Behavioral manipulation of the orb‐weaver spider Argiope argentata (Araneae: Araneidae) by Acrotaphus chedelae (Hymenoptera: Ichneumonidae)

We observed the first case of host‐behavioral manipulation of an orb‐weaver spider Argiope argentata induced by a parasitoid wasp of the genus Acrotaphus. The modified web is similar of those constructed by other orb weavers attacked by wasps of the close related genus Hymenoepimecis. The stick spirals and radii are absent and the web is composed of a three‐dimensional structure of non adhesive threads. The discovery of the ability to induce changes in host's web‐building behavior in Acrotaphus is indicative that this trait may be primitively present in the clade that includes the genus Hymenoepimecis.     

Vertical asymmetries in orb webs

In almost all vertical orb webs the hub is above the geometric centre and consequently, the extent of the capture area is larger below the hub than above. In addition to this vertical web‐extent asymmetry, orb webs show vertical asymmetries in number of spiral loops, mesh widths, and angles between radii. However, it was unknown whether these asymmetries are adaptations to the web‐extent asymmetry or whether they are linked to gravity in a different way than through web‐extent asymmetry. We reviewed known vertical asymmetries of orb webs, and we analysed the asymmetries of webs built by four different Cyclosa species, which show large intra‐ and inter‐specific variation in web‐extent asymmetry. We found all analysed structural asymmetries to be linked both to web‐extent asymmetry and to gravity: Larger web extents below the hub and gravity both led to more sticky‐spiral loops and to smaller angles between radii below the hub, whereas web‐extent asymmetry and gravity had opposing effects on mesh width (mean and peripheral). Independent of web‐extent asymmetry, almost all analysed webs had narrower peripheral meshes and smaller angles between radii below the hub than above. We interpret the narrow peripheral meshes along the web's lower edge as an adaptation to prevent tumbling prey from escaping, and the small angles between radii as an adaptation to prevent the sticky‐spiral lines in these narrow meshes to come into contact with each other. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 659–672.     

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.     

Ladder webs in orb‐web spiders: ontogenetic and evolutionary patterns in Nephilidae

Spider web research bridges ethology, ecology, functional morphology, material science, development, genetics, and evolution. Recent work proposes the aerial orb web as a one‐time key evolutionary innovation that has freed spider‐web architecture from substrate constraints. However, the orb has repeatedly been modified or lost within araneoid spiders. Modifications include not only sheet‐ and cobwebs, but also ladder webs, which secondarily utilize the substrate. A recent nephilid species level phylogeny suggests that the ancestral nephilid web architecture was an arboricolous ladder and that round aerial webs were derived. Because the web biology of the basalmost Clitaetra and the derived Nephila are well understood, the present study focuses on the webs of the two phylogenetically intervening genera, Herennia and Nephilengys, to establish ontogenetic and macroevolutionary patterns across the nephilid tree. We compared juvenile and adult webs of 95 Herennia multipuncta and 143 Nephilengys malabarensis for two measures of ontogenetic allometric web changes: web asymmetry quantified by the ladder index, and hub asymmetry quantified by the hub displacement index. We define a ‘ladder web’ as a vertically elongated orb exceeding twice the length over width (ladder index ≥ 2) and possessing (sub)parallel rather than round side frames. Webs in both genera allometrically grew from orbs to ladders, more so in Herennia. Such allometric web growth enables the spider to maintain its arboricolous web site. Unexpectedly, hub asymmetry only increased significantly in heavy‐bodied Nephilengys females, and not in Herennia, challenging the commonly invoked gravity hypothesis. The findings obtained in the present study support the intrageneric uniformness of nephilid webs, with Herennia etruscilla webs being identical to H. multipuncta. The nephilid web evolution suggests that the ancestor of Nephila reinvented the aerial orb web because the orb arises at a much more inclusive phylogenetic level, and all intervening nephilids retained the secondarily acquired substrate‐dependent ladder web. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 849–866.     

Mesh Width Influences Prey Retention in Spider Orb Webs

Orb‐weaving spiders depend upon the sticky capture spirals of webs to retain insects long enough to be captured. However, insects often escape from orb webs before the spiders can attack them. Therefore, the architectures of orb webs likely reflect strong selective pressure to increase retention times of insects. We experimentally increased the mesh width of one side of an orb web while maintaining the original mesh width on the other side as a control, and then tested the effect of this manipulation on the retention times of four different taxa of insects. We found evidence that increased mesh width of Argiope aurantia orb webs resulted in a general reduction in the retention times of insects. However, retention times for different taxa of insects were not predicted by any one specific morphological or flight characteristic. The influence of mesh width on the retention times of insects is very complex, but our results suggest that mesh width can act to selectively favor the capture of certain taxa of insect prey over others. This effect may help to explain both species level differences in web‐building behaviors and variation in the architectures of webs spun by individual spiders on different days.     

Low metabolic rates in primitive hunters and weaver spiders

The rates of oxygen consumption and carbon dioxide release of primitive hunters and weaver spiders, the Chilean Recluse spider Loxosceles laeta Nicolet (Araneae: Sicariidae) and the Chilean Tiger spider Scytodes globula Nicolet (Araneae: Scytodidae), are analyzed, and their relationship with body mass is studied. The results are compared with the metabolic data available for other spiders. A low metabolic rate is found both for these two species and other primitive hunters and weavers, such as spiders of the families Dysderidae and Plectreuridae. The metabolic rate of this group is lower than in nonprimitive spiders, such as the orb weavers (Araneae: Araneidae). The results reject the proposition of a general relationship for metabolic rate for all land arthropods (related to body mass) and agree with the hypothesis that metabolic rates are affected not only by sex, reproductive and developmental status, but also by ecology and life style, recognizing here, at least in the araneomorph spiders, a group having low metabolism, comprising the primitive hunters and weaver spiders, and another group comprising the higher metabolic rate web building spiders (e.g. orb weavers).     

Phylogenetic position and composition of Zygiellinae and Caerostris,with new insight into orb‐web evolution and gigantism

Orb‐weaving spiders are good objects for evolutionary research, but phylogenetic relationships among and within orb‐weaving lineages are poorly understood. Here we present the first species‐level molecular phylogeny that includes the enigmatic orb weavers ‘Zygiellidae’ and Caerostris. Zygiellidae is interesting for the evolution of the sector web, and Caerostris is noteworthy for web gigantism and extraordinary silk biomechanics. We assembled a molecular data set using mitochondrial (COI, 16S) and nuclear (H3, 18S, 28S, ITS2) gene fragments for 112 orbicularian exemplars, focusing on taxa with diverse web architecture and size. We show that ‘Zygiellidae’ contains the Holarctic Zygiella genus group (Leviellus, Parazygiella, Stroemiellus, and Zygiella) and the Australasian Phonognatha and Deliochus. As this clade is placed with Araneidae in all analyses we treat it as a subfamily, Zygiellinae. Using the new phylogeny, we show that the sector web evolved eight times, and coevolved with the silk tube retreat, but that these features are not zygielline synapomorphies. Zygiellinae, Caerostris, and some other araneids form a basal grade of araneids that differ from ‘classical’ araneids in web‐building and preying behaviour. We also confirm that Caerostris represents the most striking case of spider‐web gigantism. © 2015 The Linnean Society of London     

Manipulation of araneid spider web architecture by the polysphinctine parasitoid Zatypota picticollis (Hymenoptera: Ichneumonidae: Pimplinae)

We found that the koinobiont ectoparasitoid wasp Zatypota picticollis is exclusively associated with three orb weaving spiders Cyclosa conica, Mangora acalypha and Zilla diodia from the family Araneidae. Under the influence of the parasitoid's final instar larva the spiders built a specific web architecture, which differed considerably from the capturing orb web. Manipulated webs of C. conica and M. acalypha lacked the spiral, stabilimentum and central hub, and the radials were reduced in number. The manipulated web of Z. diodia consisted of one strong horizontally oriented thread.     

ORB WEBS IN "NON-ORB WEAVING" OGRE-FACED SPIDERS (ARANEAE: DINOPIDAE): A QUESTION OF GENEALOGY

Abstract— Observations of web spinning behavior in Costa Rican Dinopis sp. reveal the same behaviors synapomorphic for orb weavers: specifically frame, radius, non-sticky spiral construction, and sticky spiral construction, as well as more detailed motor patterns. Dinopids are therefore highly derived orb weavers, although the behavioral data do not conclusively indicate whether they are more closely related to the uloborid or araneoid orb weavers. A cladogram of dinopids, uloborids, and araneoids is presented.     

<Emphasis Type="Italic">Argiope bruennichi</Emphasis> shows a drinking-like behaviour in web hub decorations (Araneae,Araneidae)

As stationary predators, araneid spiders that lack protective retreats are especially vulnerable to abiotic influences. Species of the genus Argiope permanently remain at the hub of their orb webs and are thereby exposed to desiccating circumstances. Like other land arthropods, spiders must balance their hygric status. Beside desiccation avoidance behaviours, they can manage this balance by water gain through either liquefied prey items or direct ingestions of free water. Drinking-like behaviours are sparely documented for Araneids. We observed Argiope bruennichi ingesting accumulated water droplets from the silk-overstitched web hub, a part of the web decoration, and subsequently tested whether this behaviour is a regular feature of this species. In 50% of our observations, spiders that had been sprayed with water actively searched the hub decoration for water droplets and ingested them. The behavioural elements were very stereotypic among the tested individuals. Significantly, A. bruennichi females only searched the covered web hubs for water, even though the entire web was moistened. These data suggest that hub decorations of A. bruennichi might have an adaptive significance by helping to maintain a balance of water metabolism, adding yet another element to the spirited debate about the functional significance of web decorations.     

A new view of orb webs: multiple trap designs in a single structure

Spider orb webs are impressive for their apparently uniform geometric patterns. There are, however, consistent, substantial and taxonomically widespread periphery‐to‐hub differences in the distances between both adjacent radii and between sticky spiral lines. Radii in typical orbs were on average about 4–5 times farther apart at the outer edge than the inner edge of the area covered by sticky lines. Distances between sticky spiral loops were on average about two times larger near the outer edge than in more inner portions. This pattern in sticky spiral spacing was absent in the modified orbs of Nephila clavipes, in which distances between radii varied less. Thus, patterns in sticky spiral spacing may be related to inter‐radial spacing; there is, however, probably no single explanation for all of the different patterns of sticky spiral spacing. The patterned differences in radius and sticky spiral spacing have important consequences for understanding orb function, because the lines in a prey's immediate vicinity largely determine whether it will be stopped and then retained, and elementary physics dictates that contact with more lines will tend to increase prey being stopped and retained. Rather than being a unit trap with a single set of prey capture properties, an orb has locally different trapping properties in different sectors. Abandoning the previous typological style of discussion of ‘the’ ability of a given design to stop and retain prey promises to lead to improved understanding of orb web designs. Published 2014. This article is a U.S. Government work and is in the public domain in the USA, Biological Journal of the Linnean Society, 2014, 111 , 437–449.     

The influence of web monitoring tactics on the tracheal systems of spiders in the family Uloboridae (Arachnida,Araneida)

Summary Uloborids that spin reduced webs more actively monitor them than those that construct orb webs. Hyptiotes use both their first and fourth legs to tense their triangle-webs, whereas Miagrammopes rely principally on their first legs to monitor and jerk the threads of their irregular webs. The respiratory systems of these spiders include tracheae that extend into the prosoma, bifurcate, and enter the legs. To determine if the legs responsible for active web-monitoring tactics have more extensive tracheal supplies, the total cross sectional area has been computed of the tracheae entering the legs of mature female orb web and reduced web uloborids. Each leg's value has been divided by the cross sectional area of the tracheal trunks that enter the prosoma. These indexes reveal no significant differences between the relative tracheal supplies of the orb weavers investigated (Waitkera waitkerensis, Tangaroa beattyi, Uloborus glomosus). But the first, third, and fourth legs of H. cavatus and the first legs of M. animotus and M. pinopus have greater relative tracheal supplies than those of the three orb weaving species. Relative to leg volume, the first and fourth legs of H. cavatus have the greatest and the first legs of Miagrammopes species the next greatest tracheal supplies. When tracheal lengths are considered, these differences in potential oxygen supplies remain, showing that area differences do not simply compensate for differences in the distances over which oxygen must diffuse. These differences are leg-specific and not species-specific, and uloborids with the most extensive tracheal supplies are found in moist habitats. Thus the observed differences are best explained as adaptations to meet the greater oxygen demands of legs responsible for active web-monitoring tactics and not as adaptations to reduce respiratory water loss.     

The spinning apparatus of Polenecia producta (Araneae,Uloboridae): Structure and histochemistry

Summary The spinning apparatus of the uloborid spider Polenecia producta was studied to complete previous studies on the same family of spiders. The structure of spinnerets and spigots, under scanning electron microscopy, and the main anatomical and histochemical characteristics of the spinning glands of adult females and males are described. In addition some observations on the spinning apparatus at three successive stages of development are made. There are nine kinds of silk glands in Polenecia, i.e. one more (aciniform — B glands) than found in other uloborids. The spinning apparatus of Polenecia is, therefore, the most complex so far known. It is also more complex than that presently known of Araneoidea. The characteristics of the spinning glands of Polenecia are compared with those of other uloborids. Present knowledge of the spinning apparatus of uloborids leads to a renewed discussion of the origin of the orb web in this family and in araneids. It is concluded that these two types of orb webs emerged from independent evolutionary processes.     

Memory of distances and directions moved as cues during temporary spiral construction in the spiderLeucauge mariana (Araneae: Araneidae)

The spider Leucauge marianaprobably uses the presence of temporary spiral (TSP) lines already in place to determine sites of attachment of currently produced loops of temporary spiral, but less rigidly than has been previously supposed for orb weavers. Memory of distances and directions traveled recently to and from TSP lines is implicated by the fact that adjustments to experimental and natural discontinuities in previous TSP lines occur gradually rather than being abrupt. Distances and directions traveled along both radii and previous TSP lines correlate with relative amounts of adjustment. Body size may also be used as a reference measure, but not in the simple, inflexible way suggested by R. W. G. Hingston (A Naturalist in Himalaya,Small, Maynard, Boston, 1920). Tensions on radii are not used as cues in any simple way, and may not be used at all, since experimental changes in tensions produced effects consistent with resulting changes in thread positions but inconsistent with tension differences.     

蜘蛛位置对成功捕获猎物和球型网图案的影响

静坐在球型网的中心,蜘蛛可能遭受天敌的攻击并暴露在不利的天气条件下,如风和雨。然而,栖居于网的中心使蜘蛛比隐藏在隐蔽场所中的蜘蛛能更迅速地察觉并捕获猎物,这是因为猎物的位置仅能被位于网中心的蜘蛛所确定。对在隐蔽场所中的蜘蛛而言,提高对猎物捕获率的方式之一是尽量减少隐蔽所与网中心的距离。而且,网中心与隐蔽所之间较短的距离使蜘蛛能更迅速地逃离危险境况。我使用既在网中心、又在隐蔽场所的硬类肥蛛(Larinioides sclopetarius Clerck),来检验这两种行为如何影响对猎物的捕获成功率。隐藏在隐蔽场所中的蜘蛛更经常忽略猎物,使猎物也有比较多的逃离机会,这样,与在网中心的蜘蛛相比,猎物的损失率就更高。另外,研究了隐蔽场所的位置对球型网图案的影响。在大多数球型网中,网中心上方的区域比网下方小,丝也比较少,形成了结构不对称的网;隐蔽场所通常在网的上方。当隐蔽场所的位置在实验中被倒转时,就形成了非典型的球型网。最后,L．sclopetarius建造的网有很突出的边缘非对称性,与隐蔽场所相邻的区域面积较小,而远离隐蔽场所的区域面积较大,这也可解释为减少了隐蔽场所和网中心之间的距离[动物学报50(4)：559-565．2004]。     

Tangled in a sparse spider web: single origin of orb weavers and their spinning work unravelled by denser taxonomic sampling

In order to study the tempo and the mode of spider orb web evolution and diversification, we conducted a phylogenetic analysis using six genetic markers along with a comprehensive taxon sample. The present analyses are the first to recover the monophyly of orb-weaving spiders based solely on DNA sequence data and an extensive taxon sample. We present the first dated orb weaver phylogeny. Our results suggest that orb weavers appeared by the Middle Triassic and underwent a rapid diversification during the end of the Triassic and Early Jurassic. By the second half of the Jurassic, most of the extant orb-weaving families and web designs were already present. The processes that may have given origin to this diversification of lineages and web architectures are discussed. A combination of biotic factors, such as key innovations in web design and silk composition, as well as abiotic environmental changes, may have played important roles in the diversification of orb weavers. Our analyses also show that increased taxon sampling density in both ingroups and outgroups greatly improves phylogenetic accuracy even when extensive data are missing. This effect is particularly important when addition of character data improves gene overlap.     

Host behavior modification of Achaearanea tingo (Araneae: Theridiidae) induced by the parasitoid wasp Zatypota alborhombarta (Hymenoptera: Ichneumonidae)

Behavioral manipulation involving Zatypota (Ichneumonidae: Pimplinae) parasitoids and their spider hosts is usually associated with an increase in web complexity at the location where the parasitoid larva builds its cocoon. A higher number of web threads at this location may improve stability and provide a physical barrier against potential predators. However, we observed that parasitized individuals of Achaearanea tingo attacked by Z. alborhombarta change the three‐dimensional structure of their webs to a very simple and strong structure composed of two cables attached to the surrounding vegetation. This structure holds the curled leaf formerly used by the spider as a shelter. The parasitoid larva remains protected within this shelter after killing the host. The architectural pattern of the cocoon webs of A. tingo indicates that host manipulation is characterized by the repetition of one specific subroutine involved in web construction. Similar alterations have been previously described for cocoon webs constructed by parasitized orb‐weavers, but not for the three‐dimensional webs of theridiids.     

Forces in the spider orb web

Summary Pretensile forces were measured in individual threads of intact spider webs. In the orb web of Araneus diadematus forces decrease from mooring threads to frame threads and radii, a typical ratio being 1071. The smaller number of radii in the upper than in the lower half of the orb is paralleled by force ratios of 21 to 31. A similar difference between radii built first during web construction and radii added after completion of the frame underlines the importance of the former as part of the scaffolding. High tensions in the auxiliary spiral stabilize the radii in addition to providing a pathway for the spider when inserting the sticky spiral. Radial pretension (F) changes with spider mass (m). F/m is similar for different animals indicating an adaptation of radial forces to those resulting from spider mass. Several observations suggest tension control by the spider. When forced to anchor its web to thin flexible rods tension in the threads remains in the normal range. Tension values are similar in the webs of A. diadematus, Zygiella x-notata, Nuctenea umbratica, and Nephila clavipes indicating independence from details of web geometry. Only the mooring threads of Nephila show unusually large forces suggesting a narrower working range of tensions for the catching area than for the scaffolding.     

Building a better insect trap; An experimental investigation of prey capture in a variety of spider webs

Summary Web-building spiders (Araneae; Theridiidae, Linyphiidae, Araneidae) are catagorized as searchers because they devote a large amount of energy to the construction of the web which constitutes the search phase in the foraging sequence. In this study search energy is equated with the density of threads in a web and the effectiveness of a variety of webs in three broad catagories (tangle webs, sheet webs & orb webs) is tested in the light of current foraging theory. Within each web type there is a distinct thread density at which the number of approaching Drosophila (Diptera; Drosophilidae) that are captured is maximized (Figs. 1, 2, 3). That maximum results from a combination of factors that are a function of the density of threads in the web. The visibility of the web to an approaching Drosophila increases which acts to decrease the number of flies that enter the web (Tables 2, 3, 4). The ability of the web to detain a Drosophila that contacts it (capture efficiency) increases to an asymptote as a function of thread density (Fig. 4). These data support an assumption of many optimal foraging models that with increasing investment in search the predator receives a diminishing return.More Drosophila intercept orb webs than intercept sheet or tangle webs. In addition orb webs detain a greater proportion of the flies that contact them (Fig. 4). Sheet webs are intermediate between orb and tangle webs in their relative abilities to contact and detain Drosophila.     

The escape strategy of green lacewings from orb webs

When green lacewings (Neuroptera: Chrysopidae) fly into spider orb webs, they often simply reverse their flight direction and pull away (Table I). If a lacewing is trapped, it uses a specialized escape behavior. It first cuts away the sticky strands entangling head, feet, and antennae. If an antenna cannot be freed by tugging, it uses an antenna climb (Fig. 5A). After its body is free, the lacewing remains suspended by its hair-covered wings, which are held in a characteristic cruciform position (Fig. 5B). Orb web sticky strands adhere poorly to the hairy wings (Fig. 7), so the chrysopid may just wait until the strands slide off and it falls free. If placed in an orb web when the spider is at the web hub and ready to attack, a lacewing usually does not have time to escape (Fig. 1). When the spider is at the hub but eating, the chances of escape improve, and when the spider is away from the hub attacking other prey, nearly all lacewings in our experiment were able to escape. This finding emphasizes the importance of the spider's activity in its capture success.Paper No. 88 of the series Defense Mechanisms of Arthropods.