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.     

The organization of plurisegmental mechanosensitive interneurons in the central nervous system of the wandering spider Cupiennius salei

Summary In spiders the bulk of the central nervous system (CNS) consists of fused segmental ganglia traversed by longitudinal tracts, which have precise relationships with sensory neuropils and which contain the fibers of large plurisegmental interneurons. The responses of these interneurons to various mechanical stimuli were studied electrophysiologically, and their unilateral or bilateral structure was revealed by intracellular staining. Unilateral interneurons visit all the neuromeres on one side of the CNS. They receive mechanosensory input either from a single leg or from all ipsilateral legs via sensory neurons that invade leg neuromeres and project into specific longitudinal tracts. The anatomical organization of unilateral interneurons suggests that their axons impart their information to all ipsilateral leg neuromeres. Bilateral interneurons are of two kinds, symmetric and asymmetric neurons. The latter respond to stimulation of all legs on one side of the body, having their dendrites amongst sensory tracts of the same side of the CNS. Anatomical evidence suggests that their terminals invade all four contralateral leg neuromeres. Bilaterally symmetrical plurisegmental interneurons have dendritic arborizations in both halves of the fused ventral ganglia. They respond to the stimulation of any of the 8 legs. A third class of cells, the ascending neurons have unilateral or bilateral dendritic arborizations in the fused ventral ganglia and show blebbed axons in postero-ventral regions of the brain. Their response characteristics are similar to those of other plurisegmental interneurons. Descending neurons have opposite structural polarity, arising in the brain and terminating in segmental regions of the fused ventral ganglia. Descending neurons show strong responses to visual stimulation. Approximately 50% of all the recorded neurons respond exclusively to stimulation of a single type of mechanoreceptor (either tactile hairs, or trichobothria, or slit sensilla), while the rest respond to stimulation of a variety of sensilla. However, these functional differences are not obviously reflected by the anatomy. The functional significance of plurisegmental interneurons is discussed with respect to sensory convergence and the coordination of motor output to the legs. A comparison between the response properties of certain plurisegmental interneurons and their parent longitudinal tracts suggests that the tracts themselves do not reflect a modality-specific organization.Abbreviations BPI bilateral plurisegmental interneuron - CNS central nervous system - FVG fused ventral ganglia - LT longitudinal tract - PI plurisegmental interneuron - PSTH peristimulus timehistogram - UPI unilateral plurisegmental interneuron     

Fine structure of the anterior median eyes of the funnel-web spider Agelena labyrinthica (Araneae: Agelenidae)

Only few electron microscopic studies exist on the structure of the main eyes (anterior median eyes, AME) of web spiders. The present paper provides details on the anatomy of the AME in the funnel-web spider Agelena labyrinthica. The retina consists of two separate regions with differently arranged photoreceptor cells. Its central part has sensory cells with rhabdomeres on 2, 3, or 4 sides, whereas those of the ventral retina have only two rhabdomeres on opposite sides. In addition, the rhabdomeres of the ventral retina are arranged in a specific way: Whereas in the most ventral part they form long tangential rows, those towards the center are detached and are arranged radially. All sensory cells are wrapped by unpigmented pigment cell processes. In agelenid spiders the axons of the sensory cells exit from the middle of the cell body; their fine structure and course through the eye cup is described in detail. In the central part of the retina efferent nerve fibres were found forming synapses along the distal region of the receptor cells. A muscle is attached laterally to each eye cup that allows mainly rotational movements of the eyes. The optical performance (image resolution) of these main eyes with relatively few visual cells is discussed.     

Intraguild interactions between spiders and ants and top-down control in a grassland food web

In most terrestrial ecosystems ants (Formicidae) as eusocial insects and spiders (Araneida) as solitary trappers and hunters are key predators. To study the role of predation by these generalist predators in a dry grassland, we manipulated densities of ants and spiders (natural and low density) in a two-factorial field experiment using fenced plots. The experiment revealed strong intraguild interactions between ants and spiders. Higher densities of ants negatively affected the abundance and biomass of web-building spiders. The density of Linyphiidae was threefold higher in plots without ant colonies. The abundance of Formica cunicularia workers was significantly higher in spider-removal plots. Also, population size of springtails (Collembola) was negatively affected by the presence of wandering spiders. Ants reduced the density of Lepidoptera larvae. In contrast, the abundance of coccids (Ortheziidae) was positively correlated with densities of ants. To gain a better understanding of the position of spiders, ants and other dominant invertebrate groups in the studied food web and important trophic links, we used a stable isotope analysis (¹⁵N and ¹³C). Adult wandering spiders were more enriched in ¹⁵N relative to ¹⁴N than juveniles, indicating a shift to predatory prey groups. Juvenile wandering and web-building spiders showed δ¹⁵N ratios just one trophic level above those of Collembola, and they had similar δ¹³C values, indicating that Collembola are an important prey group for ground living spiders. The effects of spiders demonstrated in the field experiment support this result. We conclude that the food resource of spiders in our study system is largely based on the detrital food web and that their effects on herbivores are weak. The effects of ants are not clear-cut and include predation as well as mutualism with herbivores. Within this diverse predator guild, intraguild interactions are important structuring forces.     

The allometry of CNS size and consequences of miniaturization in orb-weaving and cleptoparasitic spiders

Allometric studies of the gross neuroanatomy of adults from nine species of spiders from six web-weaving families (Orbicularia), and nymphs from six of these species, show that very small spiders resemble other small animals in having disproportionately larger central nervous systems (CNSs) relative to body mass when compared with large-bodied forms. Small spiderlings and minute adult spiders have similar relative CNS volumes. The relatively large CNS of a very small spider occupies up to 78% of the cephalothorax volume. The CNSs of very small spiders extend into their coxae, occupying as much as 26% of the profile area of the coxae of an Anapisona simoni spiderling (body mass < 0.005 mg). Such modifications occur both in species with minute adults, and in tiny spiderlings of species with large-bodied adults. In at least one such species, Leucauge mariana, the CNS of the spiderling extends into a prominent ventral bulge of the sternum. Tiny spiders also have reduced neuronal cell body diameters. The adults of nearly all orbicularian spiders weave prey capture webs, as do the spiderlings, beginning with second instar nymphs. Comparable allometric relations occur in adults of both orb-weaving and cleptoparasitic species, indicating that this behavioral difference is not reflected in differences in gross CNS allometry.     

应用RAPD技术对蜘蛛系统演化的初步研究

利用随机扩增多态ＤＮＡ 技术检测了３ 类不同蜘蛛的系统发生关系．用１２ 个随机引物对各实验蜘蛛的基因组ＤＮＡ 进行扩增,选择其中扩增谱带清晰的８ 个引物进行分析并计算不同类蜘蛛间的遗传距离．结果表明：所有８ 个引物获得的ＲＡＰＤ谱带均表现为不同程度的多态性;地穴型蜘蛛与结网型蜘蛛间的遗传距离及结网型与游猎型之间的遗传距离,均比地穴型与游猎型之间的遗传距离近,体现了蜘蛛由地穴→结网→游猎的系统演化进程．这一结论与根据古生物学、胚胎学、形态学及生态学等得出的结论一致,从而进一步从ＤＮＡ 分子水平上为蜘蛛系统演化提供了新的证据．     

Distribution of the spider community in the olive grove agroecosystem (Portugal): potential bioindicators

1. Spiders are successful natural enemies in different crops. Research on the role of spiders as natural enemies within agroecosystems needs to address the distribution of their communities across different spatial scales, as well as the dominant species.
2. We studied the spatial distribution of the community of spiders of the olive agroecosystem in Northeastern Portugal and investigated potential species that could be used as bioindicators of agricultural management in the olive grove.
3. We found nine functional groups (ambushers, foliage runner hunters, ground hunters, orb‐web builders, sheet web builders, sensing web‐builders, space web builders, stalkers and wandering sheet/tangle weavers) encompassing a community that changed significantly across the horizontal and vertical gradient.
4. We propose Thanatus vulgaris Simon, 1870 as potential bioindicator for the ground of the olive grove central area and Ozyptila pauxilla (Simon, 1870) for the ground of the olive grove peripheral area.
5. Adjacent shrubland areas could play an important role in biological control of pests, allowing the exchange of species and individuals with the olive crop. The role of agrobiont species as indicators of agricultural managements deserves further investigation towards the enhancement of the effectiveness of spiders within low‐impact crop management in arable landscapes.

     

Consequences of sexual size dimorphism on energetics and locomotor performance of Grammostola rosea (Araneae; Teraphosidae)

Most male spiders are smaller than females; during sexual maturity, males change their behaviour, abandoning their web or nest to seek out receptive females actively, whereas females stalk prey near their web or nest and tend not to move away from it. Considering this behavioural difference to be associated with increased locomotor activity at maturity, it may be hypothesized that males will have traits that increase locomotor performance. The present study examines the kinetics and energetics of the movements of the mygalomorph spider Grammostola rosea Walckenaer, a large spider with sexual size dimorphism. It is found that males have a higher maximum aerobic speed, average speed, distance travelled and critical angle of climbing than females, indicating better performance. Males also have lower costs of transport than females. These results support the hypothesis that sexual dimorphism in wandering spiders with active males, which are characterized by smaller body size and longer legs than the larger and more static females, is associated with low transport cost, high velocity and better locomotor performance.     

Competition for a limited space in kleptoparasitic Argyrodes spiders revealed by field experiments

Most kleptoparasitic Argyrodes spiders rely exclusively on host spider webs for obtaining their food. Because their densities occasionally reach high levels within a restricted area, competitive interactions may be important for determining the density of these unique spiders. Here I used two Argyrodes species commonly found on webs of the large orb-web spider Nephila clavata to clarify whether inter- and intraspecific competition influences abundance and within-web distribution by using observational data and field experiment. Removing Argyrodes flavescens from the host webs induced a remarkably high immigration of that species while density on control webs remained almost at the same level, which is evidence for strong intraspecific competition. Larger individuals of A. flavescens were located more frequently at the capture area of the host webs where it is easy to access prey ignored by the host spider, and spiders immigrating into webs from which that species had been removed were smaller in body size, suggesting interference competition for space among conspecific kleptoparasites. Argyrodes bonadea increased in number on webs from which A. flavescens had been removed, and the increase was correlated with the number of A. flavescens removed. This finding is evidence for interspecific competition that is rarely reported in spiders. A multiple regression model including numbers of a conspecific parasite as well as web and body sizes of the host spider could not detect competitive interactions between species, suggesting the importance of experimental approaches. Received: May 22, 2000 / Accepted: December 1, 2000     

The phylogenetic placement of Psechridae within Entelegynae and the convergent origin of orb‐like spider webs

Evolutionary convergence of phenotypic traits provides evidence for their functional success. The origin of the orb web was a critical event in the diversification of spiders that facilitated a spectacular radiation of approximately 12 000 species and promoted the evolution of novel web types. How the orb web evolved from ancestral web types, and how many times orb‐like architectures evolved in spiders, has been debated for a long time. The little known spider genus Fecenia (Psechridae) constructs a web that resembles the archetypical orb web, but morphological data suggest that Psechridae (Psechrus + Fecenia) does not belong in Orbiculariae, the ‘true orb weavers’, but to the ‘retrolateral tibial apophysis (RTA) clade’ consisting mostly of wandering spiders, but also including spiders building less regular webs. Yet, the data are sparse and no molecular phylogenetic study has estimated Fecenia's exact position in the tree of life. Adding new data to sequences pulled from GenBank, we reconstruct a phylogeny of Entelegynae and phylogenetically test the monophyly and placement of Psechridae, and in doing so, the alternative hypotheses of monophyletic origin of the orb web and the pseudo‐orb versus their independent origins, a potentially spectacular case of behavioural convergence. We also discuss the implications of our results for Entelegynae systematics. Our results firmly place a monophyletic Psechridae within the RTA clade, phylogenetically distant from true orb weavers. The architectural similarities of the orb and the pseudo‐orb are therefore clearly convergent, as also suggested by detailed comparisons of these two web types, as well as the spiders' web‐building behaviours and ontogenetic development. The convergence of Fecenia webs with true orbs provides a remarkable opportunity to investigate how these complex sets of traits may have interacted during the evolution of the orb.     

Structure,composition and mechanical properties of the silk fibres of the egg case of the Joro spider, <Emphasis Type="Italic">Nephila clavata</Emphasis> (Araneae,Nephilidae)

The silk egg case and orb web of spiders are elaborate structures that are assembled from a number of components. We analysed the structure, the amino acid and fibre compositions, and the tensile properties of the silk fibres of the egg case of Nephila clavata. SEM shows that the outer and inner covers of the egg case consist of thick, medium and thin silk fibres. The silk fibres of the outer cover of the egg case are probably produced by the major and minor ampullate glands. The silk fibres of the inner cover of the egg case from cylindrical glands appears to be distinct from the silk fibres of the major ampullate glands based on their micro-morphology, mole percent amino acid composition and types, and tensile behaviour and properties. Collectively, our investigations show that N. clavata uses silk fibres from relatively few glands in varying combinations to achieve different physical and chemical properties (e.g., color, diameter, morphology and amino acid composition) and functional and mechanical properties in the different layers of the egg case.     

Contrasting Patch Residence Strategy in Two Species of Sit‐and‐Wait Foragers Under the Same Environment: A Constraint by Life History?

The present study explored the significance of life history constraints on patch residence strategy by using two congeneric web spider species living in the same habitat. Nephila maculata had a large body size but had a shorter developmental period compared with N. clavata, indicating that N. maculata should have a greater foraging efficiency to reach maturity and reproduce. Residence time at web‐sites in N. maculata was shorter than that in N. clavata, irrespective of the season. However, supplementation of food to N. maculata increased residence time, suggesting that it searches web‐sites with higher prey intake. Investment of web materials, an important trait influencing web relocation frequency, was not greater in N. maculata. In addition, microhabitat and prey size did not differ significantly after controlling for the effect of body size. Because N. maculata needs to attain a large body size in a shorter period of time, this species appears to take a risk of moving patches to seek high quality web‐sites.     

The allometry of the arcuate body in the postembryonic development of the giant house spider <Emphasis Type="Italic">Eratigena atrica</Emphasis>

The brain of arachnids contains a special neuropil area called the arcuate body (AB), whose function has been widely discussed. Its growth and proportion in the brain volume during postembryogenesis have been investigated only in several spider species. Our allometric study is aimed at determining to what extent the development of the AB in Eratigena atrica, a spider with unique biology and behaviour, is similar to the development of this body in other species. We put forward a hypothesis of allometric growth of this body in relation to the volume of the central nervous system (CNS) and its neuropil as well as in relation to the volume of the brain and its neuropil. The analysis of paraffin embedded, H?+?E stained histological preparations confirmed our hypothesis. The AB developed more slowly than the CNS and the neuropil of both the brain and the CNS. In contrast, it exhibited positive allometry in relation to the volume of the brain. This body increased more than nine times within the postembryonic development. Its proportion in the brain volume varied; the lowest was recorded in larvae and nymphs I; then, it increased in nymphs VI and decreased to 2.93% in nymphs X. We conclude that in Eratigena atrica, the AB develops differently that in orb-weaver and wandering spiders. There is no universal model of the AB development, although in adult spiders, regardless of their behaviour, the proportion of this area in the brain volume is similar.     

The influence of food supply on foraging behaviour in a desert spider

We tested the alternative hypotheses that foraging effort will increase (energy maximizer model) or decrease (due to increased costs or risks) when food supply increased, using a Namib desert burrowing spider, Seothyra henscheli (Eresidae), which feeds mainly on ants. The web of S. henscheli has a simple geometrical configuration, comprising a horizontal mat on the sand surface, with a variable number of lobes lined with sticky silk. The sticky silk is renewed daily after being covered by wind-blown sand. In a field experiment, we supplemented the spiders' natural prey with one ant on each day that spiders had active webs and determined the response to an increase in prey. We compared the foraging activity and web geometry of prey-supplemented spiders to non-supplemented controls. We compared the same parameters in fooddeprived and supplemented spiders in captivity. The results support the costs of foraging hypothesis. Supplemented spiders reduced their foraging activity and web dimensions. They moulted at least once and grew rapidly, more than doubling their mass in 6 weeks. By contrast, food-deprived spiders increased foraging effort by enlarging the diameter of the capture web. We suggest that digestive constraints prevented supplemented spiders from fully utilizing the available prey. By reducing foraging activities on the surface, spiders in a prey-rich habitat can reduce the risk of predation. However, early maturation resulting from a higher growth rate provides no advantage to S. henscheli owing to the fact that the timing of mating and dispersal are fixed by climatic factors (wind and temperature). Instead, large female body size will increase fitness by increasing the investiment in young during the period of extended maternal care.     

Immunocytochemical localization of GAD isoforms in the CNS ganglia of the cobweb spider,Achaearanea tepidariorum

Glutamic acid decarboxylase (GAD) is an enzyme that catalyzes the decarboxylation of glutamate to γ‐aminobutyric acid (GABA) and CO₂. It has been discovered that the GAD has a restricted tissue distribution and it is highly expressed in the cytoplasm of GABAergic neurons in the CNS where GABA is used as a neurotransmitter. We have examined the microstructure of ganglionic neurons and nerves arising from the CNS and describe here the immunocytochemical localization of GAD isoforms to reveal the ecophysiological significance of GABA for the web‐building spider's behavior. In the CNS of the cobweb spider, Achaearanea tepidariorum, immunocytochemical localization of GAD isoforms can be detected in the neurons and neuropiles of the optic lobes. In addition, GAD‐like immunoreactive cell bodies are observed at the intrinsic cell bodies near the central body and the symmetric cell clusters of the protocerebrum. However, the fibrous masses within the protocerebral ganglion are not labeled at all. Based on its interconnection with other regions of the CNS, our findings suggest that the central body in the web‐building spider may act as an association center as well as a visual center.     

Regional patterns in the invasion success of <Emphasis Type="Italic">Cheiracanthium</Emphasis> spiders (Miturgidae) in vineyard ecosystems

Invasions have often been linked to reduced biodiversity, but the role of non-native species in the decline of native species is ambiguous. In a 2003 survey of four California vineyard regions, exotic spiders (Cheiracanthium spp.) were more dominant in vineyards with lower spider species diversity and reduced spider abundance. There was no evidence for the role of species interactions in the invasion of Cheiracanthium spiders, however, as native spiders from the same feeding guild were most abundant in regions with high Cheiracanthium levels. Comparison with a survey conducted 10 years earlier indicated that the recent invader C. mildei simply represented an addition to the spider community, with no apparent change in proportions of the congener C. inclusum. Invasion success is discussed with respect to agricultural habitat, as results suggest that disturbed conditions in many vineyards may favor Cheiracanthium spp. and native wandering spiders while decreasing overall spider diversity.     

Body size determines the outcome of competition for webs among alien and native sheetweb spiders (Araneae: Linyphiidae)

Abstract 1. Biotic invasions are one of the most important reasons for changes in biodiversity. The alien sheetweb spider Mermessus trilobatus (Araneae: Linyphiidae) has become abundant in large parts of Central Europe within the past three decades. Its invasion might negatively influence native spiders, for instance via competition for webs. 2. Laboratory experiments were developed to test if M. trilobatus is competitively superior to native spiders. The alien M. trilobatus and five native sheetweb spiders (Erigone dentipalpis, E. atra, Gnathonarium dentatum, Dicymbium nigrum and Micrargus herbigradus) were compared with respect to their success to take over occupied webs from E. dentipalpis. 3. The rate of web takeover or defence was determined by body size, whereby individuals with a wider thorax invaded webs more successfully. After taking body size into account, the frequency of defence or web takeover did not differ between species. In 13% of all confrontations, predation against generally smaller opponents was recorded. Contrary to the predictions, raising the web value with food resources did not raise the effort expended on web defence but reduced predation by the web owner. 4. The current study does not indicate that the invasion of the relatively small‐sized M. trilobatus is facilitated by strong competitiveness. Nevertheless, M. trilobatus may displace smaller‐sized immature specimens and thereby threaten native spiders. Still, other reasons are likely to underlie the success of M. trilobatus in Europe, such as rapid reproduction or release from natural enemies.     

Neuroarchitecture of the arcuate body in the brain of the spider Cupiennius salei (Araneae, Chelicerata) revealed by allatostatin-, proctolin-, and CCAP-immunocytochemistry and its evolutionary implications

Here we describe the neuronal organization of the arcuate body in the brain of the wandering spider Cupiennius salei. The internal anatomy of this major brain center is analyzed in detail based on allatostatin-, proctolin-, and crustacean cardioactive peptide (CCAP)-immunohistochemistry. Prominent neuronal features are demonstrated in graphic reconstructions. The stainings revealed that the neuroarchitecture of the arcuate body is characterized by several distinct layers some of which comprise nerve terminals that are organized in columnar, palisade-like arrays. The anatomy of the spider's arcuate body exhibits similarities as well as differences when compared to the central complex in the protocerebrum of the Tetraconata. Arguments for and against a possible homology of the arcuate body of the Chelicerata and the central complex of the Tetraconata and their consequences for the understanding of arthropod brain evolution are discussed.     

Relating body size to the role of aquatic subsidies for the riparian spider <Emphasis Type="Italic">Nephila clavata</Emphasis>

We examined the relationship between body size of the riparian spider Nephila clavata and the contribution of allochthonous (aquatic insects) and autochthonous (terrestrial insects) sources to its diet using stable isotope analysis. During the study period from July to September, the body size of the females increased remarkably (about 60-fold) but that of males remained small. The biomass of both aquatic and terrestrial insects trapped on the spider webs increased with spider size, with the biomass of the former ranging between 30 and 70% of that of the terrestrial insects. The average relative contribution of aquatic insects to the diet of the spiders, calculated from δ¹³C values, was 40–50% in spiders in the early juvenile and juvenile stages, 35% in adult males and 4% in adult females. There was a significant negative relationship between the relative contribution of aquatic insects and body size of the female spiders. We conclude that aquatic insects might be an important seasonal dietary subsidy for small spiders and that these allochthonous subsidies may facilitate the growth of riparian spiders, which may in turn enable the spiders to feed on larger prey.     

Male-male competition and mating success in the orb-web spider,Nephila clavata,with reference to temporal factors

Seasonal occurrence patterns of adults of both sexes, intensity of male-male interactions, and mating success in the spider,Nephila clavata, were examined in the field. Adult males began to attend female webs about 2 weeks before female maturation. Large adult males were abundant in the early breeding season, but small males increased later in the season. From the distribution of males among female webs and size relationship of males within a web, male-male interactions seemed to be more intense when most females were still subadult. This was verified by a field experiment in which males were artificially introduced to female webs that were attended by other males. It was found that the probability of introduced males remaining on subadult female webs was lower than that on adult webs. As mating occurred mostly in the period shortly after the female final molt and first male sperm precedence was known in all spiders reported so far, intense male-male competition on subadult female webs seemed to be reasonable. Male longevity had an important influence on the mating success of males with just-molted females. Mating success was also affected by the relative body size of males present in a given period. Since larger males occupied the position closest to females within a web and stayed there longer, relative body size appeared to influence mating success through male-male competition. Female body size at maturation declined with time; hence, males that attained sexual maturity earlier had the advantage of mating with larger and more fecund females. Therefore, early maturation as well as larger size seem to be two important trairs influencing the reproductive success of males.