Consistency of females’ stridulatory behaviour during inter‐sexual interactions in spiders

In a sexual context, it is expected that females base their choice of mate on the behaviours that males perform during courtship, as such behaviours are associated with the male's mate quality. Stridulation is one form of female communication in arthropods, for example, spiders. In spiders, stridulation during sexual interactions is relatively common in some groups but mainly restricted to males. In the pholcid spider Holocnemus pluchei (Pholcidae), both sexes have stridulatory organs. The aims of the present work were to: (a) determine possible differences in the frequency of occurrence of stridulation between females during inter‐sexual interactions, (b) establish female consistency in stridulation along repeated interactions and (c) analyse if female stridulation is associated with certain male behaviours during pre‐copulatory courtship and with male size. Female H. pluchei showed highly repeatable differences in their frequency of stridulation across consecutive encounters with males (ICC = 0.64). However, only a modest level of repeatability was detected in total time females spent stridulating across trials (ICC = 0.19). Females’ mean stridulatory behaviour did not change across ten consecutive trials spread across 20 days, and their behaviour was apparently unaffected by male persistence of copulatory attempted and/or size. These results imply that the frequency of female stridulatory behaviour is a trait that is highly characteristic of each individual. Finally, our work opens the door to determine whether behavioural consistency manifests in other ecological contexts and their functional implications.     

Signalling and Sexual Conflict: Female Spiders Use Stridulation to Inform Males of Sexual Receptivity

The use of acoustic signals by males during courtship and mating is well known. Nevertheless, their association with female unwillingness to mate is much less studied. In spiders, stridulation during sexual interactions is relatively common in some groups, but mainly restricted to males. In the pholcid spider Holocnemus pluchei, both sexes have stridulatory organs. The aims of the present work were (1) to establish whether female stridulation occurs during intra‐ and inter‐sexual interactions, (2) to determine whether female reproductive status affects the likelihood that she will stridulate and (3) to determine whether female stridulation is influenced by male sexual behaviour. We found that female stridulation usually occurs both during intrasexual interactions and, most frequently, during intersexual interactions. Females with more previous matings stridulated more frequently. Stridulation intensity was higher in females that did not accept new copulations compared with those that copulated. Female stridulation did not vary in elaborated and non‐elaborated courtship. Thus, females use stridulation to communicate levels of sexual receptivity. It is also possible that females use stridulation to indirectly assess male ability to persist and persuade.     

Southern African pholcid spiders: revision and cladistic analysis of Quamtana gen. nov. and Spermophora Hentz (Araneae: Pholcidae), with notes on male−female covariation

The southern African pholcid spiders previously assigned to Pholcus and Spermophora are revised and their phylogenetic position within Pholcidae is analysed cladistically. Two distinct groups are recognized: the first is restricted to the Cape region and probably correctly placed in Spermophora ; it includes Spermophora peninsulae Lawrence and four new species. The second is more widely distributed in southern and eastern South Africa, and reaches as far north as Cameroon, Congo (Democratic Republic) and Uganda; it is assigned generic status ( Quamtana gen. nov .), and includes two species transferred from Pholcus [ Q. leptopholcica (Strand), Q. ciliata (Lawrence)] as well as 23 new species. A key to the pholcid spiders of South Africa is presented. A close correlation between a male character (distance between cheliceral apophyses) and a female character (distance between pockets on epigynum) is documented in Quamtana .  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139 , 477−527.     

Colonization and diversification of the spider genus Pholcus Walckenaer, 1805 (Araneae, Pholcidae) in the Macaronesian archipelagos: evidence for long-term occupancy yet rapid recent speciation

Macaronesian archipelagos stand apart from other oceanic islands reputed as laboratories for the study of evolution by their proximity to the mainland, lack of subsidence, and steep ecological gradients. The genus Pholcus Walckenaer, 1805, commonly known as daddy-long-leg spiders, is one of the most speciose arthropod groups in the region, with 25 endemic species. In the present study, we use information from four mitochondrial genes, along with morphological data, to examine the phylogenetic relationships and diversification patterns of the genus in the region. Phylogenetic analyses support monophyly of Macaronesian Pholcus including the Moroccan species Pholcus vachoni and hence a single colonization event in the archipelagos. Madeira colonizers most likely originated from the Canaries, and a back-colonization of the nearby mainland receives further support. Estimated lineage divergence times suggest a long-time presence of Pholcus in the region, but also reveal that most present-day species are the result of recent, and probably rapid, speciation events. Diagnostic characters among Macaronesian Pholcus are confined to structures involved in copulation. Coupled with the extremely high diversification rate, the highest recorded for spiders, these copulatory characters suggest that sexual selection has played a key role in the local diversification of Pholcus in Macaronesia.     

Individual decisions and group dynamics: why pholcid spiders join and leave groups

To examine the role of individual variation in the dynamics of group formation, I conducted a mark-recapture study and a series of laboratory and field experiments with Holocnemus pluchei spiders (Araneae: Pholcidae). These spiders can either share webs or live alone, and individuals shift frequently between these strategies. Spiders' decisions were influenced by size and recent feeding success. In the laboratory, small hungry spiders introduced into a web that held a larger conspecific resident were more likely than small well-fed spiders to abandon the web and build their own web. This behaviour pattern gradually reversed as spiders grew: large hungry spiders were more likely than large well-fed spiders to stay in the shared web. When I introduced spiders into empty webs, they were more likely to stay compared with spiders tested with conspecifics. However, hunger level also influenced behaviour even when conspecifics were not present. Food-deprived spiders were more likely to abandon webs and build their own, consistent with the idea that spiders were following a win-stay/lose-shift strategy. In the field, spiders were more likely to stay in webs overnight when they were given supplemental food. In another experiment, spiders that were found building webs in cleared areas were smaller and thinner than average. Finally, I tested whether the size of the intruder or the resident affected whether spiders joined webs. Large intruders were more successful at remaining in webs than smaller intruders, although spiders of all sizes had some success in joining groups. Additional synthetic theoretical work is needed to integrate the complex processes underlying the formation and persistence of groups.     

Slow genital and genetic but rapid non-genital and ecological differentiation in a pair of spider species (Araneae,Pholcidae)

Like most animals with internal fertilization, spiders tend to have species-specific genitalia, allowing closely related species to be identified by their reproductive morphology more easily than by non-genital characters. This implies that genitalia evolve on average more rapidly than non-genital morphological traits. Here we describe two putative species of Pholcidae from Taiwan (Pholcus pingtung, n. sp.; Pholcus chengpoi, n. sp.) that differ conspicuously in their microhabitat (rocks vs. leaves), coloration, color pattern, and body proportions, but have almost indistinguishable genitalia and cytochrome oxidase I (co1) sequences. The two species have identical yet highly unusual male cheliceral modifications, strongly arguing for sister species. Despite the almost identical genitalia and co1 sequences, we treat the two ‘morphs’ as species for three reasons: (1) they are easily distinguished by several characters; no intermediate specimens were found; (2) subtle yet consistent differences in genital (uncus) shape support the idea of reproductively isolated entities beyond the more conspicuous non-genital differences; (3) each locality provided both types of microhabitat but only one of the two species, arguing against environmental plasticity or polymorphism. We conclude that probably a very recent expansion into a novel microhabitat has led to speciation and rapid ecological and non-genital differentiation, with insufficient time to accumulate significant genital and genetic differences.     

Comparative morphology of cheliceral muscles using high-resolution X-ray microcomputed-tomography in palpimanoid spiders (Araneae,Palpimanoidea)

Spiders are important predators in terrestrial ecosystems, yet we know very little about the principal feeding structures of spiders, the chelicerae, which are functionally equivalent to “jaws” or “mandibles” and are an extremely important aspect of spider biology. In particular, members of Palpimanoidea have evolved highly unusual cheliceral morphologies and functions, including high-speed, ballistic movements in mecysmaucheniid spiders, the fastest arachnid movements known thus far, and the elongated, highly maneuverable chelicerae of archaeids that use an attack-at-a-distance strategy. Here, using micro-Computed-Tomography scanning techniques, we perform a comparative study to examine cheliceral muscle morphology in six different spider specimens representing five palpimanoid families. We provide a hypothesis for homology in palpimanoid cheliceral muscles and then compare and contrast these findings with previous studies on other non-palpimanoid spiders. We document and discuss two sets of cheliceral muscles in palpimanoids that have not been previously observed in other spiders or which may represent a position shift compared to other spiders. In the palpimanoids, Palpimanus sp., Huttonia sp., and Colopea sp. showed similar cheliceral muscle anatomy. In Eriauchenius ranavalona, which has highly maneuverable chelicerae, some of the muscles have a more horizontal orientation, and there is a greater degree of cheliceral muscle divergence. In Zearchaea sp. and Aotearoa magna, some muscles have also shifted to a more horizontal orientation, and in Zearchaea sp., a species with a ballistic, high-speed predatory strike, there is a loss of cheliceral muscles. This research is a first step toward understanding cheliceral form and function across spiders.     

Karyotype, sex determination, and meiotic chromosome behavior in two pholcid (Araneomorphae, Pholcidae) spiders: implications for karyotype evolution

There are 1,111 species of pholcid spiders, of which less than 2% have published karyotypes. Our aim in this study was to determine the karyotypes and sex determination mechanisms of two species of pholcids: Physocyclus mexicanus (Banks, 1898) and Holocnemus pluchei (Scopoli, 1763), and to observe sex chromosome behavior during meiosis. We constructed karyotypes for P. mexicanus and H. pluchei using information from both living and fixed cells. We found that P. mexicanus has a chromosome number of 2n = 15 in males and 2n = 16 in females with X0-XX sex determination, like other members of the genus Physocyclus. H. pluchei has a chromosome number of 2n = 28 in males and 2n = 28 in females with XY-XX sex determination, which is substantially different from its closest relatives. These data contribute to our knowledge of the evolution of this large and geographically ubiquitous family, and are the first evidence of XY-XX sex determination in pholcids.     

Ultrastructure of spider thread anchorages

Spiders attach silken threads to substrates by means of glue-coated nanofibers (piriform silk), spun into disc-like structures. The organization and ultrastructure of this nano-composite silk are largely unknown, despite their implications for the biomechanical function and material properties of thread anchorages. In this work, the ultrastructure of silken attachment discs was studied in representatives of four spider families with Transmission Electron Microscopy to facilitate a mechanistic understanding of piriform silk function across spiders. Based on previous findings from comparative studies of piriform silk gland morphology, we hypothesized that the fibre-glue proportion of piriform silk differs in different spiders, while the composition of fibre and glue fractions is consistent. Results confirmed large differences in the relative proportion of glue with low amounts in the orb weaver Nephila senegalensis (Araneidae) and the hunting spider Cupiennius salei (Ctenidae), larger amounts in the cobweb spider Parasteatoda tepidariorum (Theridiidae) and a complete reduction of the fibrous component in the haplogyne spider Pholcus phalangioides (Pholcidae). We rejected our hypothesis that glue ultrastructure is consistent. The glue is a colloid with polymeric and fluid fractions that strongly differ in proportions and assembly. We further confirmed that in all species studied both dragline and piriform silk fibres do not make contact with the environmental substrate. Instead, adhesion is established by a thin dense skin layer of the piriform glue. These results advance our understanding of piriform silk function and the interspecific variation of its properties, which is significant for spider biology, web function and the bioengineering of silk.     

Web-invasion and araneophagy by New Zealand and Australian pholcid spiders

Abstract

Pholcus phalangioides, Pholcus ancoralis, and Psilochorus sphaeroides were observed in the field in New Zealand (first species only) and Australia (all three species) to invade the webs of other species. P. phalangioides was observed to feed on spiders in alien webs. Webs of each species were frequently contiguous with webs of other spiders and P. sphaeroides sometimes invaded neighbouring webs in pursuit of insects.     

A Parental Care-Mating Dilemma? Potential Risks for Offspring in the Pholcid Spider When Egg-Carrying Females Accept Mating

Many species face a trade-off between additional mating opportunities and the offspring benefits (viability, quality) provided by parental care. Female Holocnemus pluchei spiders must abandon their egg-sac, which they otherwise carry with their chelicerae, to copulate. This may involve risks for the offspring, such as predation and fungal infection. We assessed whether (1) males discriminate between egg-carrying females according to the egg development stage, (2) females are influenced by the egg development stage in regard to their proneness to mate, and (3) offspring are less likely to survive, due to high humidity and/or predation, in the absence of egg-carrying females. Apparently, males did not distinguish between females according to the developmental stage of the eggs they carried. However, females were more likely to mate when carrying relatively mature versus immature eggs (14–18 days and 3–6 days post-oviposition, respectively). All egg-sacs hatched successfully when guarded by egg-carrying females because this avoided both fungal infections (at high humidity) and cannibalism by conspecific spiders. Thus, H. pluchei females may face a trade-off between mating and parental care. Further research should clarify why egg-carrying females mate and how females prevent their egg-sacs from being infected by fungi.     

北京幽灵蛛的附肢自残

附肢自残是北京幽灵蛛（Pholcus beijingensis Zhu ＆ Song,1999）的习性之一。野外调查表明,约20％的个体至少失去1只步足;个体自残的发生率在2—7月间有显著的差异,同时第一对步足自残的比例最高（43．47％）;个体失去步足的比例随着年龄的增加而增加;左侧和右侧步足失去的比率接近（1．03：1）。野外调查发现,正在交配的个体与单独生活的个体相比,步足的完整性上并无显著差别。实验室研究表明,北京幽灵蛛附肢自残后没有再生现象。通过对自残幼体与非自残幼体的3龄与4龄龄期研究,发现各处理组间并无显著的差别。步足自残被认为是北京幽灵蛛在极端情况下的“避险策略”（Bet-hedging strategy）[动物学报54（6）：998-1004,2008]。     

Exploring the effect of the Cardinium endosymbiont on spiders

Spiders have recently emerged as important diversity hot spots for endosymbiotic bacteria, but the consequences of these symbiotic interactions are largely unknown. In this article, we examined the evolutionary history and effect of the intracellular bacterium Cardinium hertigii in the marbled cellar spider Holocnemus pluchei. We showed that Cardinium infection is primarily transmitted in spider populations maternally via egg cytoplasm, with 100% of the progeny from infected mothers being also infected. Examination of a co‐inherited marker, mitochondrial DNA (mtDNA), revealed that Cardinium infection is associated with a wide diversity of mtDNA haplotypes, showing that the interaction between Cardinium and H. pluchei has a long‐term evolutionary dimension and that horizontal transmission among individuals could also occur. Although Cardinium is well known to exert sex ratio distortion or cytoplasmic incompatibility in various arthropod hosts, we show, however, that Cardinium does not interact with the reproductive biology of H. pluchei. A field survey shows a clear geographical structuring of Cardinium infection, with a marked gradual variation of infection frequencies from ca. 0.80 to 0. We discuss different mechanistic and evolutionary explanations for these results as well as their consequences for spider phenotypes. Notably, we suggest that Cardinium can either behave as a neutral cytoplasmic element within H. pluchei or exhibit a context‐dependent effect, depending on the environmental conditions.     

Females of the Cellar Spider Discriminate Against Previous Mates

Mate choice for novel partners should evolve when remating with males of varying genetic quality provides females with fitness‐enhancing benefits. We investigated sequential mate choice for same or novel mating partners in females of the cellar spider Pholcus phalangioides (Pholcidae) to understand what drives female remating in this system. Pholcus phalangioides females are moderately polyandrous and show reluctance to remating, but double‐mated females benefit from a higher oviposition probability compared to single‐mated females. We exposed mated females to either their former (same male) or a novel mating partner and assessed mating success together with courtship and copulatory behaviours in both sexes. We found clear evidence for mate discrimination: females experienced three‐fold higher remating probabilities with novel males, being more often aggressive towards former males and accepting novel males faster in the second than in the first mating trial. The preference for novel males suggests that remating is driven by benefits derived from multiple partners. The low remating rates and the strong last male sperm precedence in this system suggest that mating with novel partners that represent alternative genotypes may be a means for selecting against a former mate of lower quality.     

Hyalomma dromedarii (Acari: Ixodoidea: Ixodidae): Central and peripheral nervous system anatomy

TheHyalommadromedarii central nervous system, the synganglion, is an integrated nerve mass concentrated around the esophagus and formed by fusion of a small anterodorsal supraesophageal part an a large posteroventral subesophageal part. The supraesophageal part consists of the protocerebrum including a pair of optic ganglia, a pair of cheliceral ganglia, a pair of pedipalpal ganglia, and the stomodeal pons. The subesophageal part includes four paired pedal ganglia and the complex opisthosomatic ganglion. The peripheral nervous system includes the following pairs of nerves: optic, cheliceral, pedipalpal, primary and accessory (histologically traced); also unpaired pharyngeal and recurrent nerves, four pairs of pedal nerve trunks, each with a hemal branch, and two pairs of opisthosomatic nerves. Each peripheral nerve is traced distally to the innervation site. The salivary glands are innervated anteriorly by branches of the pedipalpal nerve and medially by branches of the hemal nerves associated with the third pedal nerves.Reprint request should be sent to: Medical Zoology Department, NAMRU-3, Fleet Post Office, New York 09527, U.S.A.     

近亲幽灵蛛用于兼性群居生活的时间消耗

所谓近亲幽灵蛛（Pholcus affinis）的兼性群居生活,是指他们可以单独生活,也可以几个个体共栖于一张网上成为一组。本文使用焦点动物追踪法,研究不同大小的组（指组内个体数量多少）、不同年龄阶段的蜘蛛在组内生活中用于“消耗性”行为（移动、织网、弹跳、拨丝或种内相互作用）的时间耗费。主要结论包括：蜘蛛的年龄和组的大小影响组内成员的付出。在不同大小的组里,2～3龄幼蛛（S型）、亚成体或成体（L型）用在“消耗性”行为上所用的时间百分率没有显著性差异,但4～5龄幼蛛（M型）随着组内成员的增多而要花费更多的时间。当组内个体数量超过3个,L型和M型蜘蛛之间有着更多的相互作用,而S型蜘蛛不论组内个体多少,其个体之间相互作用都很少。研究结果同时显示,近亲幽灵蛛在不同发育阶段有不同的群居生活策略。     

Intersexual cooperation during male clasping of external female genitalia in the spider Physocyclus dugesi (Araneae, Pholcidae)

Sexual conflict may influence the shape and evolution of body structures that males use to grasp females during mating. Not only sexual coercion but also intersexual cooperation may be involved during clasping behavior. Among pholcid spiders, secondary sexual modifications of the male chelicerae, such as apophyses with spines or tooth-like processes, function to grasp the female by specific parts of her external genitalia such as grooves or apophyses of the epigynum. We analyzed how the female and the male respond when their structures for clasping are experimentally modified in the pholcid Physocyclus dugesi. We used three treatment groups for virgin females that differed in the manipulation of the epigynum apophyses (uncovered, partially covered, and fully covered by a plaster) and two groups of males (uncovered and fully covered cheliceral apophyses). We found that females are mainly cooperative to courting males not only when the female genital apophyses were experimentally covered but also when the male cheliceral apophyses were covered. The current data also indicate behavioral flexibility in males during courtship, especially when they had difficulty in genital intromission. Our experimental results, together with previous observational studies, support a modulated-cooperative scenario between the sexes for cheliceral clasping and genital intromission in pholcid spiders.     

Fine structure of the chelicera in the spider Nephila clavata

The fine structural characteristics of the biting apparatus in the orb‐web spider Nephila clavata were studied using scanning electron microscopy. The main biting apparatuses of spiders are the chelicerae and cheliceral fangs in the cephalothorax. The chelicera of N. clavata is that of the jack‐knife (folding knife) type, which is composed of two segments, and has a labidognathous form that moves at right angles to the body axis. Each chelicera bears a hinged fang that folds into a cheliceral groove. The tips of the fangs are quite sharp, and the spider's body is well adapted to driving the fangs into prey. Just below the fang, each side of the cheliceral groove is covered with a total of seven cuticular teeth (four promarginal teeth and three retromarginal teeth) in two rows. The cheliceral fang has a single aperture at the tip of the posterior surface, and the lower margin of the fang which meets the promarginal teeth is a saw‐like groove. Fine structural observation reveals that each fang has a single venom pore, and each cuticular depressive area on the cheliceral groove has two different types of surface pit. Approximately 40 to 50 spiky protrusions were counted at the cheliceral groove, to hold prey tightly.     

Cobweb management and control of the spider Holocnemus pluchei (Araneae: Pholcidae) on buildings

The spider Holocnemus pluchei (Scopoli) (Araneae: Pholcidae) is a nonnative species commonly found in the southwestern United States. In urban areas around homes and other structures, it is often the dominant spider species on outside surfaces; requests for control of spiders and their cobwebs are common for the commercial pest management industry. We tested two physical (brushing and vacuuming) and two low-impact chemical (permethrin and botanical insecticide) spray control methods for H. pluchei on outbuildings on the University of California-Riverside campus. Forty sites containing at least two H. pluchei spiders along a 5-m section of eave or overhang were chosen for treatment. Treatment was applied within 2 d of a pretreatment census. The number of spiders per section was counted at 2, 4, 8, 12, 16, and 52 wk posttreatment. Untreated sections showed an increase in spiders through summer and a decrease in autumn. Sites treated with permethrin significantly reduced spiders >95% and had strong residual effect throughout the remainder of the sampling period. Compared with untreated controls, the three other treatments did not significantly reduce spiders; these three were similar throughout the season and were intermediate between untreated and permethrin-treated sites. In comparison with the before and 2-wk posttreatments, spiders at the untreated sites showed no difference in body sizes, whereas those in the nonpermethrin treatments showed a decrease, indicating that larger spiders were eliminated. We also tested H. pluchei spiders in the laboratory against the two insecticides used in the field studies to determine the minimum lethal concentrations for controlling H. pluchei.     

Structure and anatomical relationships of the synganglion in the American dog tick,Dermacentor variabilis (Acari: Ixodiadae)

The synganglion of Dermacentor variabilis Say is a single nerve mass, condensed around the esophagus and within the periganglionic sinus of the ciculatory system. Protocerebral, cheliceral (including stomodeal bridge), and pedipalpal ganglia lie in the pre-esophageal portion of the nerve mass and bear optic, cheliceral, and pedipalpal nerves respectively. The unpaired stomodeal and the recurrent nerve which forms the hyper-esophageal ganglion arise from the stomodeal bridge. Paired primary and accessory nerves to the retrocerebral organ complex have mixed protocerebral-cheliceral origins. Pedal ganglia (including ventral olfactory lobes of pedal ganglia I) and composite opisthosomal ganglion lie in the post-esophageal nerve mass and bear pedal nerve trunks and two pairs of opisthosomal nerves respectively. Internally, the synganglion consists of cellular rind and fibrous core. A welldefined neurilemma with a laminar matrix covers nerve mass and peripheral nerves. The rind contains the somata of ganglionic neurons and ensheathing glial cells and is restricted to the synganglion mass. It is limited by two specialized glial layers, the external perineurium and internal subperineurium. Discrete glomerular formations are present within the protocerebrum and olfactory lobes. Olfactory glomeruli located in pedal ganglia I are associated with a pair of globuli cell groups. Possible physiological relationships between anatomical specializations of the synganglion, extraneural sinuses and circulating hemocytes are considered. The evolutionary significances of condensation in the stomatogastric neuropile regions and throughout the synganglion, together with the simplification and loss of glomerular formations, are discussed.