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Most morphological studies focus on adult specimens, or if developmental studies are pursued, especially in Euarthropoda, they focus on embryonic development. Araneae (spiders) is one of these groups, in consequence with understudied post-embryonic development. Here we present aspects of the post-embryonic stages of different species of Mesothelae, sister group to the remaining spiders (when fossil species are not taken into account). We used different imaging methods and measured different external morphological structures to detect possible ontogenetic changes. One structure exhibiting post-embryonic changes is the chelicera. Here the significant change occurs between the last immature stage and the adult, yet only in males. For the spinnerets, we could not detect ontogenetic changes, but instead a high variability in length and width, probably due to their lack of pivot joints between the elements. The strongest morphological change during ontogeny occurred on the sternum, which begins with a rather roundish shape in the first stage and changes to being fairly elongate in shape in the last immature stages and the adult. This specific sternum shape only occurs in adults of mesothelan spiders, while opisthothelan spiders have a broader sternum also in the adult. We discuss our results in an evolutionary context, also taking into account recent finds of fossil spiders.

Prey vary dramatically in quality, and maternal diet is generally assumed to substantially influence offspring survivorship, growth, and reproduction in spiders. Numerous studies that have tested this hypothesis have focused exclusively on parental generation or have considered relatively few fitness components of juvenile offspring. However, maternal diet may have a substantial effect on fitness performance beyond juvenile offspring. Here, we investigated the influence of one-time maternal feeding on multiple offspring fitness components, including the survival rate and growth of juvenile offspring as well as the mating and reproductive success of adult offspring in Hylyphantes graminicola , a sheetweb spider with an extremely short lifespan (∼1 month). We fed field-collected adult female spiders two different diets only once immediately before oviposition: midges ( Tendipes sp.) only (MO) or flies ( Drosophila melanogaster ) only (FO). Juvenile offspring of MO females had significantly higher survival rate, faster growth, and larger male size at maturity than FO offspring. Although maternal diet did not significantly influence mating behavior or fecundity of female offspring overall, those of MO females laid eggs earlier and their eggs also hatched earlier and had a higher hatching rate than those of FO females. Intriguingly, one-time maternal feeding was sufficient to have such an influence on offspring fitness even beyond juvenile offspring in H graminicola This one-time maternal effect may be widespread in other spiders and other invertebrates with a short lifespan.This article has an associated First Person interview with the first author of the paper., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

Winter provides many challenges for terrestrial arthropods, including low temperatures and decreased food availability. Most arthropods are dormant in the winter and resume activity when conditions are favorable, but a select few species remain active during winter. Winter activity is thought to provide a head start on spring growth and reproduction, but few studies have explicitly tested this idea or investigated tradeoffs associated with winter activity. Here, we detail biochemical changes in overwintering winter-active wolf spiders, Schizocosa stridulans, to test the hypothesis that winter activity promotes growth and energy balance. We also quantified levels of putative cryoprotectants throughout winter to test the prediction that winter activity is incompatible with biochemical adaptations for coping with extreme cold. Body mass of juveniles increased 3.5-fold across winter, providing empirical evidence that winter activity promotes growth and therefore advancement of spring reproduction. While spiders maintained protein content throughout most of the winter, lipid content decreased steadily, suggesting either a lack of available prey to maintain lipids, or more likely, an allometric shift in body composition as spiders grew larger. Carbohydrate content showed no clear seasonal trend but also tended to be higher at the beginning of the winter. Finally, we tested the hypothesis that winter activity is incompatible with cryoprotectant accumulation. However, we observed accumulation of glycerol, myo-inositol, and several other cryoprotectants, although levels were lower than those typically observed in overwintering arthropods. Together, our results indicate that winter-active wolf spiders grow during the winter, and while cryoprotectant accumulation was observed in the winter, the modest levels relative to other species could make them susceptible to extreme winter events., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

We report the discovery of the wasp Tachinobia repanda Bouček collected from egg sacs of the colonial spider Cyrtophora moluccensis (Doleschall) in Morobe Province, Papua New Guinea (PNG) by Lubin, Y.D. in 1980. This is the first record of T. repanda from egg sacs of a colonial spider. The likely host of this eulophid wasp was the larvae of a sarcophagid fly that parasitizes the egg sacs of these spiders. The 67 T. repanda collected were all females and varied little in body size. We suggest that this species is a gregarious hyperparasitoid., (© The Author(s) 2020. Published by Oxford University Press on behalf of Entomological Society of America.)

Species that are Batesian mimics during postembryonic development shift between mimetic models as they grow in size. However, it has not yet been tested whether these successive mimetic phenotypes are similarly protected from predators. Early-instar phenotypes could represent an inaccurate phenotype or an accurate phenotype because of selection from different predators. Here, we tested the hypotheses of transformational Batesian mimicry in the ant-mimicking jumping spider Leptochestes berolinensis . We quantified the mimetic accuracy of different ontogenetic stages to potential ant models by using a multitrait approach. We measured movement, body profile, body size, and coloration. Analysis revealed adults to be more accurate mimics than juveniles. Adults were similar to smaller morphs of Camponotus or Lasius ants, whereas juveniles were more similar to Lasius and Colobopsis ants. We tested whether predators, mantises, and Pisaura spiders were deceived by mimics after having experience with ant models. These predators never captured any ant or a mimic but always captured the nonmyrmecomorphic spider. We conclude that L. berolinensis is a Batesian mimic of ants undergoing transformational mimicry, with all stages being accurate mimics.

The early reports and our previous work confirmed the existence of the toxic proteinaceous components in the body of the L. tredecimguttatus newborn and adult spiders. For revealing the differences in the protein expression profiles and toxicity bases of the spiders at different developmental stages, the spiderling and adult spider proteins were comparatively analyzed using a proteomic strategy. Totals of 429 and 958 proteins were identified from the spiderlings and adult spiders, respectively, with 239 proteins being identified from both of them. Although some similarities between the spiderling and adult spider proteomes exist, there are obvious differences between the two proteomes in size, complexity, molecular weight (MW) distribution, acid-base property, and hydropathicity, etc. Gene ontology (GO) analysis demonstrates that, comparing based on the percentages of proteins, the spiderling and adult spider proteins have generally similar distribution profiles with respect to the subcellular localization, molecular function and biological process. However, there are still some differences between these two sets of proteins in some classifications of the three GO categories. For the adult spiders, latrotoxins together with other toxins and toxin-like proteins, etc. constitute their toxicity basis, whereas the toxicity of the spiderlings depends mainly on the synergistic action of atypical latrotoxins and toxin-like proteins, most of which are different from those of the adult spiders, demonstrating that the spiders at different developmental stages have largely different toxicity mechanisms., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2020 Elsevier Inc. All rights reserved.)

In the spider, determination of the dorsal-ventral body (DV) axis depends on the interplay of the dorsal morphogen encoding gene decapentaplegic (Dpp) and its antagonist, short gastrulation (sog), a gene that is involved in the correct establishment of ventral tissues. Recent work demonstrated that the forkhead domain encoding gene FoxB is involved in dorsal-ventral axis formation in spider limbs. Here, Dpp likely acts as a dorsal morphogen, and FoxB is likely in control of ventral tissues as RNAi-mediated knockdown of FoxB causes dorsalization of the limbs. In this study, we present phenotypes of FoxB knockdown that demonstrate a function in the establishment of the DV body axis. Knockdown of FoxB function leads to embryos with partially duplicated median germ bands (Duplicitas media) that are possibly the result of ectopic activation of Dpp signalling. Another class of phenotypes is characterized by unnaturally slim (dorsal-ventrally compressed) germ bands in which ventral tissue is either not formed, or is specified incorrectly, likely a result of Dpp over-activity. These results suggest that FoxB functions as an antagonist of Dpp signalling during body axis patterning, similarly as it is the case in limb development. FoxB thus represents a general player in the establishment of dorsal-ventral structures during spider ontogeny.

Due to fascinating mechanical and biological characteristics spider silk is of great interest in many research fields. Among the orb-weavers Nephila edulis is one of the species used as source for natural spider silk in laboratories. Under appropriate conditions, animals can be kept and bred easily. This manuscript gives information about the spiders' natural habitat, behavior, and breeding and compares them with the established methods and conditions within a research laboratory. Keeping conditions and methods of rearing are described in detail. Within a keeping facility with reliable supply of food, cannibalism rate is significantly reduced and spiders mate all year long. Cohabitants of the genus Steatoda are routinely found in laboratory keeping. While these small spiders do not pose a threat to Nephila edulis, cellar spiders (family Pholcidae) have to be extracted as they have been observed hunting for Nephila spiders.

Anterior patterning in animals is based on a gene regulatory network, which comprises highly conserved transcription factors like six3, pax6 and otx. More recently, foxQ2 was found to be an ancestral component of this network but its regulatory interactions showed evolutionary differences. In most animals, foxQ2 is a downstream target of six3 and knockdown leads to mild or no epidermal phenotypes. In contrast, in the red flour beetle Tribolium castaneum, foxQ2 gained a more prominent role in patterning leading to strong epidermal and brain phenotypes and being required for six3 expression. However, it has remained unclear which of these novel aspects were insect or arthropod specific. Here, we study expression and RNAi phenotype of the single foxQ2 ortholog of the spider Parasteatoda tepidariorum. We find early anterior expression similar to the one of insects. Further, we show an epidermal phenotype in the labrum similar to the insect phenotype. However, our data indicate that foxQ2 is positioned downstream of six3 like in other animals but unlike insects. Hence, the epidermal and neural pattering function of foxQ2 is ancestral for arthropods while the upstream role of foxQ2 may have evolved in the lineage leading to the insects.

Spiders are equipped with a large number of innervated cuticular specializations, which respond to various sensory stimuli. The physiological function of mechanosensory organs has been analysed in great detail in some model spider species (e.g. Cupiennius salei); however, much less is known about the distribution and function of chemosensory organs. Furthermore, our knowledge on how the sense organ pattern develops on the spider appendages is limited. Here we analyse the development of the pattern and distribution of six different external mechano- and chemosensory organs in all postembryonic stages and in adult male and female spiders of the species Parasteatoda tepidariorum. We show that except for small mechanosensory setae, external sense organs appear in fixed positions on the pedipalps and first walking legs, arranged in longitudinal rows along the proximal-distal axis or in invariable positions relative to morphological landmarks (joints, distal tarsal tip). A comparison to other Entelegynae spiders shows that these features are conserved. We hope that this study lays the foundation for future molecular analysis to address the question how this conserved pattern is generated.

The Hox gene labial (lab) governs the formation of the tritocerebral head segment in insects and spiders. However, the morphology that results from lab action is very different in the two groups. In insects, the tritocerebral segment (intercalary segment) is reduced and lacks appendages, whereas in spiders the corresponding segment (pedipalpal segment) is a proper segment including a pair of appendages (pedipalps). It is likely that this difference between lab action in insects and spiders is mediated by regulatory targets or interacting partners of lab. However, only a few such genes are known in insects and none in spiders. We have conducted a candidate gene screen in the spider Parasteatoda tepidariorum using as candidates Drosophila melanogaster genes known to (potentially) interact with lab or to be expressed in the intercalary segment. We have studied 75 P. tepidariorum genes (including previously published and duplicated genes). Only 3 of these (proboscipedia-A (pb-A) and two paralogs of extradenticle (exd)) showed differential expression between leg and pedipalp. The low success rate points to a weakness of the candidate gene approach when it is applied to lineage specific organs. The spider pedipalp has no counterpart in insects, and therefore relying on insect data apparently cannot identify larger numbers of factors implicated in its specification and formation. We argue that in these cases a de novo approach to gene discovery might be superior to the candidate gene approach.

Sexual reproduction is pervasive in animals and has led to the evolution of sexual dimorphism. In most animals, males and females show marked differences in primary and secondary sexual traits. The formation of sex-specific organs and eventually sex-specific behaviors is defined during the development of an organism. Sex determination processes have been extensively studied in a few well-established model organisms. While some key molecular regulators are conserved across animals, the initiation of sex determination is highly diverse. To reveal the mechanisms underlying the development of sexual dimorphism and to identify the evolutionary forces driving the evolution of different sexes, sex determination mechanisms must thus be studied in detail in many different animal species beyond the typical model systems. In this perspective article, we argue that spiders represent an excellent group of animals in which to study sex determination mechanisms. We show that spiders are sexually dimorphic in various morphological, behavioral, and life history traits. The availability of an increasing number of genomic and transcriptomic resources and functional tools provides a great starting point to scrutinize the extensive sexual dimorphism present in spiders on a mechanistic level. We provide an overview of the current knowledge of sex determination in spiders and propose approaches to reveal the molecular and genetic underpinnings of sexual dimorphism in these exciting animals.

Large part of the morphological diversity observed across taxa is attributed to the effect of sexual selection; and the static allometry of these structures vary largely from highly positive to negative, depending on their function, and position on the animal's body. In arthropods, information of how sexually selected contact and reaching male structures use during courtship scale on body size is scarce. We tested two complementary hypotheses: the reaching structure hypothesis and the contact-function hypothesis, in the spider Kukulcania hibernalis. We used the length of the proximal segments of the male pedipalp to test the reaching structure hypothesis, and claw features to test the contact-function hypothesis. Our results support both hypotheses. Small males have disproportionally longer pedipalps (highly negative allometry) than large males, increasing the probability of small-bodied males to inseminate even large females. We also found that both distal contact and noncontact homologous structures scaled shallow (slope < 1) on body size, but allometry was significantly shallower for contact than for noncontact distal structures, providing support to the contact-function hypothesis, and allowing teasing apart the effect of sexual selection on distal contact structures with dual functions., (© 2019 Wiley Periodicals, Inc.)

Allatoregulatory neuropeptides are multifunctional proteins that take part in the synthesis and secretion of juvenile hormones. In insects, allatostatins are inhibitors of juvenile hormone biosynthesis in the corpora allata while allatotropins, act as stimulators. By quantitative real-time PCR, we analyzed the gene expression of allatostatin A (PtASTA), allatostatin B (PtASTB), allatostatin C (PtASTC), allatotropin (PtAT) and their receptors (PtASTA-R, PtASTB-R, PtASTC-R, PtAT-R) in various tissues in different age groups of female spiders. In the presented manuscript, the presence of allatostatin A, allatostatin C, and allatotropin are reported in females of the spider P. tepidariorum. The obtained results indicated substantial differences in gene expression levels for allatoregulatory neuropeptides and their receptors in the different tissues. Additionally, the gene expression levels also varied depending on the female age. Strong expression was observed coinciding with sexual maturation in the neuroendocrine and nervous system, and to a lower extent in the digestive tissues and ovaries. Reverse trends were observed for the expression of genes encoding the receptors of these neuropeptides. In conclusion, our study is the first hint that the site of synthesis and secretion is fulfilled by similar structures as observed in other arthropods. In addition, the results of the analysis of spider physiology give evidence that the general functions like regulation of the juvenile hormone synthesis, regulation of the digestive tract and ovaries action, control of vitellogenesis process by the neuropeptides seem to be conserved among arthropods and are the milestone to future functional studies., Competing Interests: The authors have declared that no competing interests exist.

Spiders (order Araneae, class Arachnida) are an important group of predatory arthropods in terrestrial ecosystems that have been recently identified as an untapped reservoir of single-stranded (ss)DNA viruses. Specifically, spiders harbour a diversity of ssDNA viruses encoding a replication-associated protein (Rep) within a circular genome. However, little is known about the ecology of novel circular Rep-encoding ssDNA (CRESS DNA) viruses. Here we investigated two CRESS DNA viruses recently identified in spinybacked orbweavers (Gasteracantha cancriformis), namely spinybacked orbweaver circular virus (SpOrbCV) 1 and 2. SpOrbCV-1 was detected in the majority (> 65 %) of spider specimens from all life stages, including eggs, spiderlings and adults, demonstrating that this virus is active within spinybacked orbweavers. In contrast, SpOrbCV-2 was only detected in adults at a lower (36 %) prevalence. Since we also detected SpOrbCV-2 in other spider species and this virus has been reported from a dragonfly, we suggest that SpOrbCV-2 is accumulated in these predators through common insect prey. The prevalence of SpOrbCV-1 in collected specimens allowed us to design assays to characterize this virus, which represents a new group of CRESS DNA viruses, the 'circularisviruses'. To our knowledge, SpOrbCV-1 is the first example of a vertically transmitted virus in spiders, which may explain its high prevalence in spinybacked orbweavers. Since vertically transmitted viruses infecting insects (class Insecta) can manipulate their host's behaviour and physiology, future studies should investigate the ecological role of vertically transmitted viruses in spiders.

Tetragnathid spiders (Tetragnatha spp.) found in riparian habitats have recently been used as bioindicators of sediment contamination and insect-mediated contaminant flux. We investigated whether sexual dimorphism (size and behavior) influenced the female:male ratio in composite samples, stable isotope ratios (carbon [δ 13 C], nitrogen [δ 15 N]), and Hg concentrations in the southern United States. Additionally, we explored whether biomass for contaminant analysis could be preserved by using the legs of tetragnathids as a surrogate for whole-body δ 13 C and δ 15 N signatures. We found that female tetragnathids were significantly larger than male spiders and represented a larger proportion of spiders collected at all sites. However, despite the difference in size between sexes, no differences in growth dynamics, isotopic signatures (δ 13 C and δ 15 N), or mercury concentrations were observed. It was determined that the leg of a tetragnathid can accurately represent the stable isotope signature of an entire spider.

Lycosids are a diverse family of spiders distributed worldwide. Previous studies recovered some of the deeper splits of the family, but with little support. We present a broad phylogenetic analysis of the Lycosidae including a wide geographic sampling of representatives and all the subfamilies described to date. Additionally, we extend the amount of molecular data used in previous studies (28S, 12S and NADH) through the inclusion of two additional markers, the nuclear H3 and the mitochondrial COI. We estimated the divergence times through the inclusion of fossils as calibration points and used the phylogenetic hypothesis obtained to explore the evolution of particular traits associated with dispersal capabilities. We recovered most of the currently recognized subfamilies with high nodal support. Based on these results, we synonymize Piratinae and Wadicosinae with Zoicinae and Pardosinae, respectively, and revalidate the subfamily Hippasinae. We corroborated that lycosids are a family with a relatively young origin that diversified with the reduction of tropical forests and the advance of open habitats. We show that a gradual accumulation of behavioral traits associated with ambulatory dispersal made Lycosidae the most vagrant subfamily of spiders, with an impressive ability to disperse long distances which helps to explain the worldwide distribution of some very young clades, such as the members of the subfamily Lycosinae., (Copyright © 2019 Elsevier Inc. All rights reserved.)

There is growing evidence that speciation can occur between populations that are not geographically isolated. The emergence of assortative mating is believed to be critical to this process, but how assortative mating arises in diverging populations is poorly understood. The wolf spider genus Schizocosa has become a model system for studying mechanisms of assortative mating. We conducted a series of experiments to identify the factors that control mate pair formation in a Schizocosa population that includes both ornamented and nonornamented males. We show that the population also includes two previously unrecognized female phenotypes. One female phenotype mates mostly or exclusively with ornamented males, and the other mates mostly or exclusively with unornamented males. Assortative mating within these groups is maintained by differences in maturation time, microhabitat use, and female mate preference. We conclude that the population is not a single species, as previously believed, but rather an incipient species pair with multiple overlapping mechanisms of reproductive isolation. The identification of a new incipient species pair in the well-studied and rapidly speciating Schizocosa clade presents new opportunities for the study of speciation without geographic isolation.

Sox genes encode a set of highly conserved transcription factors that regulate many developmental processes. In insects, the SoxB gene Dichaete is the only Sox gene known to be involved in segmentation. To determine if similar mechanisms are used in other arthropods, we investigated the role of Sox genes during segmentation in the spider Parasteatoda tepidariorum . While Dichaete does not appear to be involved in spider segmentation, we found that the closely related Sox21b-1 gene acts as a gap gene during formation of anterior segments and is also part of the segmentation clock for development of the segment addition zone and sequential addition of opisthosomal segments. Thus, we have found that two different mechanisms of segmentation in a non-mandibulate arthropod are regulated by a SoxB gene. Our work provides new insights into the function of an important and conserved gene family, and the evolution of the regulation of segmentation in arthropods., Competing Interests: CP, AS, DL, SR, AM No competing interests declared, (© 2018, Paese et al.)

The genus Physokermes Targioni Tozzetti includes species that are distributed in the Holarctic region and feed on conifers. The recently described scale Physokermes hellenicus (Kozár and Gounari) (Hemiptera: Coccidae) is an endemic species of Greece whose host plants are fir trees of the genus Abies (Pinales: Pinaceae). It is considered as beneficial scale insect species since its honeydew secretions are exploited by honeybees leading to the production of a special honey with important physicochemical characteristics. Since there are no previous data on the natural enemies of P. hellenicus, an investigation was carried out during 2013 in forested areas of eight mountains in south and central Greece aiming to correlate the presence of P. hellenicus with certain parasitoids and predators. Seven species of Encyrtidae, Eulophidae, Pteromalidae, and Eurytomidae (Hymenoptera); five species of Anthribidae and Coccinellidae (Coleoptera); and four species of Dictinidae, Linyphiidae, and Theridiidae (Araneae) were identified. Twelve of them were identified at the species level while four at the genus level. Among them Microterys lunatus (Dalman) (Hymenoptera: Encyrtidae), Pseudorhopus testaceus (Ratzeburg) (Hymenoptera: Encyrtidae), and Anthribus fasciatus Forster (Coleoptera: Anthribidae) were the most abundant natural enemies of P. hellenicus adult female while Metaphycus unicolor Hoffer (Hymenoptera: Encyrtidae) and Trichomasthus sp. (Hymenoptera: Encyrtidae) were found to parasitize P. hellenicus male nymph. Cinetata gradata (Simon) (Araneae: Linyphiidae) is reported for first time in the Greek arachnofauna. Our results suggest that the abundance of the fir scale P. hellenicus could be affected by a complex of parasitoid and predator species of different taxa. Future long-term research on these species in relation with abiotic factors would help to understand possible fluctuation of the scale's population.

Light and transmission electron microscopy were used to study the development of new book lung lamellae in juvenile and adult spiders (Parasteatoda tepidariorum). As hypothesized earlier in a study of embryos, mesenchyme cells dispersed throughout the opisthosoma (EMT) are a likely source of precursor epithelial cells (MET) for the new lamellae. The precursor cells in juveniles and adults continue many of the complex activities observed in embryos, e.g., migration, alignment, lumen formation, thinning, elongation, and secretion of the cuticle of air channel walls and trabeculae. The apicobasal polarity of precursor cells for new channels is apparently induced by the polarity pattern of precursor cells of channels produced earlier. Thus, new air and hemolymph channels extend and continue the alternating pattern of older channels. At sites more distant from the spiracle and atrium, new channels are usually produced by the mode II process (intracellular alignment and merging of vesicles). These air channels have bridging trabeculae and are quite stable in size throughout their length. At sites closer to the spiracle and atrium, new channels may be produced by mode I (coalescence of merocrine vesicle secretion). This raises the hypothesis that structural and functional differences in mode I and II channels and differing oxygen and fluid conditions with distance from the spiracle and atrium determine the mode of formation of new channels. Observations herein support an earlier hypothesis that there is some intercellular apical/apical and basal/basal affinity among the opposed surfaces of aligned precursor cells. This results in the alternating pattern of air channels at the apical and hemolymph channels at the basal cell surfaces., (Copyright © 2018 The Author(s). Published by Elsevier Ltd.. All rights reserved.)

Embryogenesis and post-embryogenesis of spiders depend on several environmental factors including light and temperature. This study was aimed at evaluating the impact of different thermal and lighting conditions on embryonic and early post-embryonic development of Eratigena atrica. Embryos, larvae, nymphs I and II were incubated at constant temperatures of 12, 22, 25 and 32°C under three different light regimes: light, dark, light/dark. Extreme temperatures (12 and 32°C) significantly increased mortality of embryos (to 100%) and nymphs II, whereas larvae and nymphs I suffered reduced survival only at the lowest temperature. Moreover, the lowest temperature reduced the development rate of all stages. The impact of light conditions was less pronounced and more variable: constant light reduced the survival of nymphs I at lower temperatures, but increased that of larvae. Moreover, light increased the time of embryonic development and duration of nymphal stages, particularly at lower temperatures (12-22°C). Thus, the most optimal locations for spiders seem to be dark (though except larval stage) and warm (25°C) sites, where their development is fastest and mortality lowest., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Zinc finger transcription factors of the Sp6-9 group are evolutionarily conserved in all metazoans and have important functions in, e.g., limb formation and heart development. The function of Sp6-9-related genes has been studied in a number of vertebrates and invertebrates, but data from chelicerates (spiders and allies) was lacking so far. We have isolated the ortholog of Sp6-9 from the common house spider Parasteatoda tepidariorum and the cellar spider Pholcus phalangioides. We show that the Sp6-9 gene in these spider species is expressed in the developing appendages thus suggesting a conserved role in limb formation. Indeed, RNAi with Sp6-9 in P. tepidariorum leads not only to strong limb defects, but also to the loss of body segments and head defects in more strongly affected animals. Together with a new expression domain in the early embryo, these data suggest that Sp6-9 has a dual role P. tepidariorum. The early role in head and body segment formation is not known from other arthropods, but the role in limb formation is evolutionarily highly conserved.

During ontogenesis, the size of a spider body, tissues and organs increases dramatically. The aim of the study was to estimate changes in the central nervous system of postembryonic stages of Eratigena atrica and compare them with the literature data on species differing in behavioural traits. Allometric analysis involved evaluation of histological slides embedded in paraffin and stained with hematoxylin and eosin. The reduced major axis regression (RMA) was applied to find allometric relationships between the volumes of the particular parts of the body. All the measured parts of the central nervous system (CNS) were negatively allometrically related to the volume of the prosoma, showing that the increment of the CNS was lower than that of the entire body. The growth of the brain was negatively allometrically related to the growth of the CNS but the increment of the subesophageal ganglion was greater than that of the CNS, exhibiting a positive allometry. Within both these structures, the increase in neuropil volume was greater than the growth of the cortex (cell body rind). Thus, in postembryonic development, the share of the subesophageal ganglion and neuropil in the total volume of the CNS increased, whereas that of the brain and cortex decreased. The mode of the CNS development in E. atrica is similar to that observed in other arthropods, including Argiope aurantia, a spider of different ecology and behaviour., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

While molting occurs in the development of many animals, especially arthropods, post-maturity molting (PMM, organisms continue to molt after sexual maturity) has received little attention. The mechanism of molting has been studied intensively; however, the mechanism of PMM remains unknown although it is suggested to be crucial for the development of body size. In this study, we investigated factors that potentially induce PMM in the golden orb-web spider Nephila pilipes , which has the greatest degree of sexual dimorphism among terrestrial animals. We manipulated the mating history and the nutrient consumption of the females to examine whether they affect PMM. The results showed that female spiders under low nutrition were more likely to molt as adults, and mating had no significant influence on the occurrence of PMM. Moreover, spiders that underwent PMM lived longer than those that did not and their body sizes were significantly increased. Therefore, we concluded that nutritional condition rather than mating history affect PMM., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)

Generalist predators such as spiders may help mitigate the spread and impact of exotic herbivores. The lack of prey specificity and long generation times of spiders may allow them to persist when pests are scarce, and to limit the growth of pest populations before they reach damaging levels. We examined whether resident spiders are likely to play a role in maintaining populations of the invasive light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae), below outbreak levels in California. We surveyed the spider community on two E. postvittana host plants, the ornamental Australian tea tree, Leptospermum laevigatum, and the weed French broom, Genista monspessulana, to characterize spider and larval E. postvittana abundance and spider species composition throughout the year. Spider densities and species composition showed slight seasonal changes. Spiders were present during periods of high and low E. postvittana abundance. Anyphaenid hunting spiders, Anyphaena aperta Banks in Australian tea tree and Anyphaena pacifica Banks in French broom, dominated spider species composition at four of five sampled sites, and underwent only slight seasonal variation in abundance. Adult A. aperta were rare at all times of the year, suggesting that high mortality among juvenile A. aperta limits the potential of this species as a predator of E. postvittana. Nevertheless, the continued presence of spiders throughout the year indicates that the resident spider community is likely to play a key role in reducing E. postvittana populations in California., (Published by Oxford University Press on behalf of Entomological Society of America 2017. This work is written by US Government employees and is in the public domain in the US.)

Ecdysteroids play an essential role in the regulation of the molting processes of arthropods. Nuclear receptors of the spider Agelena silvatica that showed high homology with other arthropods especially in the functional domains were identified, two isoforms of ecdysone receptor (AsEcRA, AsEcRB), retinoid X receptor (AsRXR) and two isoforms of E75 (AsE75A, AsE75D). AsEcR and AsRXR mRNA did not show major changes in expression but occurred throughout the third instar nymphal stage. AsE75DBD was low or non-existent at first then showed a sudden increase from D7 to D10. On the other hand, AsE75D was expressed in the first half and decreased from D6 to D10. Ecdysteroid titers showed a peak on D6 in A. silvatica third instar nymphs. LC-MS/MS analysis of the ecdysteroid peak revealed only 20-hydroxyecdysone (20E) was present. The 20E peak on D6 and increase in AsE75DBD from D7 is likely a result of ecdysteroids binding to the heterodimer formed with constant expression of the AsEcR and AsRXR receptors. These findings indicate the mechanisms regulating molting widely conserved in insects and other arthropods also similarly function in spiders., (Copyright © 2017 Elsevier Inc. All rights reserved.)

Evarcha culicivora, an East African salticid spider, is a mosquito specialist and it is also a plant specialist, with juveniles visiting plants primarily for acquiring nectar meals and adults visiting plants primarily as mating sites. The hypothesis we consider here is that there are ontogenetic shifts in cognition-related responses by E. culicivora to plant odour. Our experiments pertain to cross-modality priming effects in three specific contexts: executing behaviour that we call the 'visual inspection of plants' (Experiment 1), adopting selective visual attention to specific visual targets (Experiment 2) and becoming prepared to respond rapidly to specific visual targets (Experiment 3). Our findings appear not to be a consequence of salient odours in general elevating E. culicivora's motivation to respond to salient visual stimuli. Instead, effects were specific to particular odours paired with particular visual targets, with the salient volatile plant compounds being caryophyllene and humulene. We found evidence that prey odour primes juveniles and adults to respond to seeing specifically prey, mate odour primes adults to respond to seeing specifically mates and plant odour primes juveniles to respond to seeing specifically flowers. However, plant odour appears to prime adults to respond to seeing specifically a mate associated with a plant., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Theraphosid spiders (tarantulas) are venomous arthropods found in most tropical and subtropical regions of the world. Tarantula venoms are a complex cocktail of toxins with potential use as pharmacological tools, drugs and bioinsecticides. Although numerous toxins have been isolated from tarantula venoms, little research has been carried out on the venom of Australian tarantulas. We therefore investigated the venom profile of the Australian theraphosid spider Phlogius crassipes and examined whether there are ontogenetic changes in venom composition. Spiders were divided into four ontogenic groups according to cephalothorax length, then the venom composition of each group was examined using gel electrophoresis and mass spectrometry. We found that the venom of P. crassipes changes continuously during development and throughout adulthood. Our data highlight the need to investigate the venom of organisms over the course of their lives to uncover and understand the changing functions of venom and the full range of toxins expressed. This in turn should lead to a deeper understanding of the organism's ecology and enhance the potential for biodiscovery.

The Southeast Asian pholcid genus Calapnita Simon, 1892 is revised, with descriptions of 17 new species, five of them in the phyllicola group (Borneo: C. lehi, C. kubah, C. bidayuh, C. bankirai; Malay Peninsula, Sumatra, Java: C. anai), 12 in the vermiformis group (Borneo: C. bario, C. bariengi, C. magaseng, C. dayak, C. lawangan, C. loksado; Sulawesi: C. bugis; Philippines: C. bohol, C. dinagat, C. mae, C. nunezae, C. maragusan). New records are listed for six of the eight previously described species. A morphological cladistic analysis supports the monophyly of Calapnita and of its two previously proposed species groups and presents several new phylogenetically informative characters. New data are presented about ultrastructure and natural history (web, egg-sac, egg parasitism).

The spider genus Oxyopes Latreille occurs throughout the world, but is insufficiently known in South America and the Caribbean. In this study, three apparently related species of this genus are reported from the region. Oxyopes salticus Hentz, 1845 is redescribed and illustrated, and extreme variation in colour pattern and in female genitalia is reported within this species. Oxyopes m-fasciatus Piza, 1938 and Oxyopes nigrolineatus Mello-Leitão, 1941 are both considered junior synonyms of O. salticus. The material examined in this study, together with information from the literature, show this species is widely distributed throughout the continental Americas and the Caribbean. Oxyopes crewi Bryant, 1948 is redescribed and illustrated, and its geographic distribution is extended to Bahamas, Cuba, Dominican Republic, Jamaica, and Saint Kitts. A new species, Oxyopes incantatus sp. nov., is described and illustrated for the first time based on specimens from the Galápagos Islands.

In this paper we propose Trebacosa brunhesi Villepoux, 2007 as a junior synonym of Trebacosa europaea Szinetár & Kan-csal, 2007 based on the examination of specimens from all the localities from where those species are known. Illustration of the type species of the genus, Trebacosa marxi (Stone, 1890) and specimens from all known localities of T. europaea are given to show both the inter- and the intraspecific differences of the genus. Scanning electron micrographs were used to illustrate the detailed structure of the female's genitalia.

Three species of the genus Allocosa Banks, 1900 from southern South America are redescribed: Allocosa alticeps (Mello-Leitão, 1944), A. brasiliensis (Petrunkevitch, 1910) and A. senex (Mello-Leitão, 1945). The female of A. senex is described for the first time and the species is revalidated. A new species, A. marindia sp. nov. from southern Uruguay and southern Brazil is described. The new species is distinguished by the flattened terminal apophysis of the male bulb and the conspicuous pointed projections on the posterior margin of the female epigynum. The species inhabits in sandy estuarine and oceanic coasts with psammophile vegetation.

Alopecosa atypica Ponomarev, 2008 is known from the holotype male collected in Western Kazakhstan. Here we synonymize it with A. cronebergi (Thorell, 1875), which was described from two syntype females from the Volga River and Daghestan. This species is redescribed in detail and its distribution is shown to be restricted to Russia and Ukraine, and previous records from Hungary are probably mistaken.

A species of the Afrotropical spider genus Afroneutria is newly described. Afroneutria tanga sp. nov is endemic to Tanzania, southeast Africa, and can be distinguished from the remaining species of the genus by the retrolateral excavation of the embolus. An identification key for all Afroneutria species is presented.

In this communication, we report important preliminary evidence for possibly the toughest egg case silk threads recorded to date spun by the hermit spider, Nephilengys cruentata (G¯=193MJm(-3)). We further elucidate that the egg case itself is woven with a specialised repeat cross-weave that when subjected to tension, drives perpendicular-to-force threads to pile. This piling of threads constrains damage to small areas and retains the architectural integrity of the surrounding egg case material. We deduce that by having ultra-tough threads coupled to a tear resistant architecture, N. cruentata is able to protect its eggs from predators with a considerable level of effectiveness., (Copyright © 2016 Elsevier B.V. All rights reserved.)

Four species of Cytaea Keyserling, 1882 are diagnosed, redescribed, and illustrated: C. laticeps (Thorell, 1878), C. nimbata (Thorell, 1881), C. rubra (Walckenaer, 1837), and C. sinuata (Doleschall, 1859).

Males of Ariadna kiwirrkurra sp. nov., collected on the 2015 Bush Blitz biodiscovery expedition to the Kiwirrkurra Indigenous Protected Area (IPA) are the first Segestriid males described for mainland Australia.

Three new species of the genus Xestaspis Simon, 1884: X. nuwaraeliya sp. nov., X. padaviya sp. nov. and X. pophami sp. nov. are described based on both sexes. Xestaspis kandy Eichenberger, 2012 is illustrated based on new material. A key and a distribution map for Sri Lankan Xestaspis is provided. Phylogenetic placement of Sri Lankan Xestaspis spp. obtained by the analysis of 43 morphological characters is presented and discussed. In all our analysis Gamasomorpha species appear paraphyletic suggesting that Xestaspis is most probably a junior synonym of Gamasomorpha. Thus, forming monophyletic group of oriental hard bodied oonopids.

Light and transmission electron microscopy were used to study the development of book lungs in embryos of the spider Parasteatoda tepidariorum. There is a bilateral cluster of temporary lamellae that form just posterior to the second opisthosomal (O2) limb buds. These lamellae are replaced by advanced embryo (AE) book lungs that continue into postembryonic stages. Results herein agree with earlier suggestions that the O2 limb buds become the AE book lungs. Each O2 limb bud merges with the ventral surface of the O2 segment, where the limb bud/book lung is internalized by covering with epidermis. A strand of tissue (entapophysis) from the epidermis at the posterior opisthosoma provides precursor cells for the book lung lamellae, and possibly entapophysis cells induce limb bud cells to align and produce lamellae. Electron micrographs show the different modes (I-III) of lumen formation. The result is a spiracle, atrium and alternating air and hemolymph channels. A hypothesis is presented for the role of precursor cell polarity in producing the planar tissue polarity of the channels. Some type of apical/apical affinity results in air channels, while basal/basal affinity results in hemolymph channels. Strong basal/basal affinity is likely as opposed cells in hemolymph channels extend basal processes that span the channel and start pillar trabeculae that continue in postembryonic stages., (Copyright © 2016 The Author. Published by Elsevier Ltd.. All rights reserved.)

Most recent studies of spider embryonic development have focused on representatives of the species-rich group of entelegyne spiders (over 80 % of all extant species). Embryogenesis in the smaller spider groups, however, is less well studied. Here, we describe the development of the germ band in the spider species Pholcus phalangioides, a representative of the haplogyne spiders that are phylogenetically the sister group of the entelegyne spiders. We show that the transition from radially symmetric embryonic anlage to the bilaterally symmetric germ band involves the accumulation of cells in the centre of the embryonic anlage (primary thickening). These cells then disperse all across the embryonic anlage. A secondary thickening of cells then appears in the centre of the embryonic anlage, and this thickening expands and forms the segment addition zone. We also confirm that the major part of the opisthosoma initially develops as a tube shaped structure, and its segments are then sequentially folded down on the yolk during inversion. This special mode of opisthosoma formation has not been reported for entelegyne spiders, but a more comprehensive sampling of this diverse group is necessary to decide whether this peculiarity is indeed lacking in the entelegyne spiders.

Two ctenid species, Anahita maolan Zhu, Chen & Song, 1999 and Ctenus yaeyamensis Yoshida, 1998 from Taiwan are redescribed in detail based on both sexes. Anahita maolan is reported from Taiwan for the first time. For both species, Kenting National Park (southernmost Taiwan) is the southernmost known locality.

Trophic specialists are expected to possess adaptations that increase the efficiency of handling preferred prey. Such adaptations may constrain the ability to utilise alternative prey. Here we tested whether the ant-eating spider Euryopis episinoides possesses metabolic specialisations with increased efficiency in utilising preferred prey and decreased efficiency in utilising alternative prey. In addition, we investigated the contribution of genetic variation via maternal effects. We reared E. episinoides spiders from the first instar on two different diets, either ants (preferred prey) or fruit flies (alternative prey). Spider survival rate and increases in body mass were significantly higher on the ant diet. The total development time did not differ between diet groups, nor did the number of egg sacs per female or the incubation period. However, the number of eggs per egg sac and hatching success were higher on the ant diet. There was a genetic variation in several offspring traits. Our data support the hypothesis that stenophagous ant-eating E. episinoides have a metabolic specialisation on ant utilisation indicated by higher efficiency in utilising ants than fruit flies. While most individuals of E. episinoides were able to capture fruit flies, only very few spiders were able to develop and reproduce on a pure fruit fly diet, suggesting the existence of within-species genetic variation regarding the tolerance to alternative prey., (Copyright © 2016 Elsevier GmbH. All rights reserved.)

The genus Tibellus Simon, 1875 currently includes 51 species (WSC 2016); it presents some taxonomic difficulties, due to morphological variability, and similarities among some congeners (Efimik 1999). Studies on the genus were undertaken by Gertsch (1933) and Schick (1965) from America, Tikader (1980) from India, Van den Berg & Dippenaar-Schoeman (1994) from the Afrotropical region, and Efimik (1999) from the East Palearctic region.

Copulatory cannibalism of male 'widow' spiders (genus Latrodectus) is a model example of the extreme effects of sexual selection, particularly in L. hasselti and L. geometricus where males typically facilitate cannibalism by females and mate only once. We show that these males can increase their reproductive success by copulating with final-instar, immature females after piercing the female's exoskeleton to access her newly developed sperm storage organs. Females retain sperm through their final moult and have similar fecundity to adult-mated females. This is an adaptive male tactic because immature mating increases insemination success relative to adult mating (which predicts higher paternity) and moreover, rarely ends in cannibalism, so males can mate again. Although successful only during a brief period before the female's final moult, males may employ this tactic when they associate with final-instar females in nature. Consistent with this, one-third of L. hasselti females collected as immatures in nature were already mated. Immature mating alters sexual selection on these otherwise monogynous males, and may explain male traits allowing facultative polygyny in Latrodectus Since male cohabitation with immature females is common among invertebrates, immature mating may be a widespread, previously unrecognized mating tactic, particularly when unmated females are of high reproductive value., (© 2016 The Author(s).)

Background: Socially cued anticipatory plasticity (SCAP) has been proposed as a widespread mechanism of adaptive life-history shifts in semelparous species with extreme male mating investment. Such mating systems evolved several times independently in spiders and male reproductive success should critically depend on timely maturation and rapid location of a receptive and, ideally, virgin female. We experimentally investigated socially cued anticipatory plasticity in two sympatric, closely related Nephila species that share many components of their mating systems, but differ in the degree to which male reproductive success depends on mating with virgin females. Juveniles of both species were reared either in the presence or absence of virgin female silk cues. We predicted strong selection on socially cued plasticity in N. fenestrata in which males follow a highly specialized terminal investment strategy, but expected a weaker plastic response in N. senegalensis in which males lost the ability to monopolize females., Results: Contrary to our predictions, N. fenestrata males presented with virgin female silk cues did not mature earlier than siblings reared isolated from such cues. Males in N. senegalensis, however, showed a significant response to female cues and matured several days earlier than control males. Plastic adjustment of maturation had no effect on male size., Conclusions: Our results indicate that a strong benefit of mating with virgins due to first male sperm priority does not necessarily promote socially cued anticipatory plasticity. We emphasize the bidirectional mode of developmental responses and suggest that this form of plasticity may not only yield benefits through accelerated maturation, but also by avoiding costs of precipitate maturation in the absence of female cues.

Six new species of the peacock spider genus Maratus Karsch, 1878 are described from Australia: M. eliasi sp. nov., M. julianneae sp. nov., M. licunxini sp. nov., M. michaelorum sp. nov. and M. ottoi sp. nov. from Queensland, and M. kiwirrkurra sp. nov. from Western  Australia. Five species groups are further documented within the genus and new records, detailed SEM and automontage images are provided for six previously described species: M. anomalus Karsch, 1878, M. chrysomelas (Simon, 1909), M. digitatus Otto & Hill, 2012, M. pavonis (Dunn, 1947), M. speciosus (O.P.-Cambridge, 1874) and M. volans (O.P.-Cambridge, 1874).

The rare ant mimicking sac spider genus Aetius was erected by O. Pickard-Cambridge in 1896 based on an unspecified number of female specimen(s) collected from Sri Lanka. The type species of the genus, A. decollatus O. Pickard-Cambridge, 1896, has been redescribed twice based on the holotype (Majumder & Tikader 1991; Deeleman-Reinhold 2001). Reimoser (1934) recorded the genus for the first time from India, who collected a male specimen from Mudumalai Tiger Reserve in Tamil Nadu State of southern India. This specimen was identified as A. decollatus, but it was never formally described and was later recognised to be a penultimate male (Dankittipakul & Singtripop 2013). Deeleman-Reinhold (2001) described the second representative of the genus, A. nocturnus, based on a single female specimen from Borneo, 105 years after the establishment of the genus. Dankittipakul & Singtripop (2013) described the male of A. nocturnus, thereby revealing the male genitalia of the genus, but the type species was still known only from the female sex.

Extracting venom from small species is usually challenging. We describe here an affordable and versatile electrical venom extractor based on the Arduino(®) Mega 2560 Board, which is designed to extract venom from arthropods and other small animals. The device includes fine tuning of stimulation time and voltage. It was used to collect venom without apparent deleterious effects, and characterized for the first time the venom of Zoropsis spinimana, a common spider in French Mediterranean regions., (Copyright © 2016 Elsevier Ltd. All rights reserved.)