Oldest true orb-weaving spider (Araneae: Araneidae)

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

A phylogenetic analysis of the orb-weaving spider family Araneidae (Arachnida, Araneae)

We present the first cladistic analysis focused at the tribal and subfamily level of the orb-weaving spider family Araneidae. The data matrix of 82 characters scored for 57 arancid genera of 6 subfamilies and 19 tribes (and 13 genera from 8 outgroup families) resulted in 16 slightly different, most parsimonious trees. Successive weighting corroborated 62 of the 66 informative nodes on these cladograms; one is recommended as the 'working' araneid phylogcny. The sister group of Araneidae is all other Araneoidea. Araneidae comprises two major clades: the subfamily Araneinae, and the 'argiopoid' clade, which includes all other subfamilies and most tribes (((Gasteracanthinae, Caerostreae), (((Micratheninae, Xylcthreae), Eruyosaccus ), (Eurycorminae, Arciinae)), Cyrlarachninae), ((Argiopinae, Cyrtophorinae), Arachnureae)). Cyrtarachneae and Mastophoreae are united in a new subfamily, Cyr-tarachninae. The spiny orb-weavers alone (Gasteracanthinae and Micratheninae) are not monophyletic. The mimetid subfamily Arciinae and the 'tetragnathid' genus Zygiella are araneids, but .Nephila (and other tetragnathids) are not. On the preferred tree, web decorations (stabilimenta) evolved 9 times within 15 genera, and were lost once. The use of silk to subdue prey evolved once in cribellate and four times in ecribillate orb weavers. Sexual size dimorphism evolved once in nephilines, twice in araneids, and reverted to monomorphism five times. Evolution in other genitalic and somatic characters is also assessed; behavioral and spinneret features arc most consistent (male genitalia, leg and prosomal features least consistent) on the phylogeny.     

How to avoid becoming a prey: Predatory encounters between an orb-weaving spider,Araneus pinguis (Karsch) (Araneae: Araneidae) and flying insects

Insects flying into the web of an orb-weaving spiderAraneus pinguis (Karsch) and their avoidance of (pre-hitting process) and escapes from (post-hitting process) the web were examined by direct observation under natural and semi-natural conditions. In the pre-hitting process, mobile insects such as Brachycera, Lepidoptera and Hymenoptera showed a low hitting ratio (number of insects hitting/number of insects flying within 1 m³ space around the web-site) because of active web avoidance and flying activity in layers lower or higher than those in which the webs are usually laid. In contrast, less mobile insects like Heteroptera, Coleoptera and Homoptera showed a high hitting ratio. In the post-hitting process, Brachycera, Lepidoptera and some Nematocera frequently escaped without being detained by the web. Many Orthoptera and Hymenoptera escaped without any sign of detection by the spider. Coleoptera frequently escaped during the spider's attack. Small insects from the Homoptera, Nematocera and Hymenoptera rarely escaped from the web, but were not immediately attacked. Mean escape time of insects was correlated significantly with capture success of the spider. Overall most of the escapes occurred in the early phases of the predation process. This indicates that escapes are unlikely to result in heavy loss of time and energy expenditure due to unsuccessful predation. Escape patterns of insects seem to be related to their mobility.     

Anti-predator defences of Argiope appensa (Araneae, Araneidae), a tropical orb-weaving spider

Anti-predator defence behaviour of Argiope appensa (Fuesslin) (Araneae, Araneidae) was studied in the laboratory. The most frequent response of adults and large juveniles of A. appensa to disturbance was pumping, a behaviour during which this web-building spider moved its body rapidly up and down with its legs remaining on the silk. When disturbed, small juveniles differed from adult females and larger juveniles by often dropping from the web instead of pumping. Argiope appensa sometimes put its web betweeen itself and stimuli from potential predators by shuttling from one side of the hub to the other. Argiope appensa occasionally tugged on the web but this behaviour appeared to be primarily a component of prey-catching sequences instead of defence. Experiments were carried out to determine the types of stimuli that elicited pumping. Lightly touching the spider or its web, forcefully hitting the web, and air movement elicited pumping but there was no evidence that chemical stimuli from potential predators were important.     

Prey analysis of four species of tropical orb-weaving spiders (Araneae: Araneidae) and a comparison with araneids of the temperate zone

Summary The actual prey in the orb webs of four araneid spiders (Nephila clavipes, Eriophora fuliginea, Argiope argentata, and A. savignyi) and the relative abundance of their potential prey (pitfall traps, yellow traps, and sweep-netting) was investigated over 1 year at different locations in Panama. The relative abundance of insects and spiders depends on seasonal fluctuations (Fig. 2) which are reflected by corresponding variations in the effectiveness of the webs. The main prey groups are Nematocera (50%–68%), winged Formicoidea (6%–15%) and Hymenoptera, Coleoptera, and Brachycera (4%–10% each) (Fig. 4-6). The remaining 10%–17% of the prey comes from up to 26 other groups (Table 2). Differences in prey size and prey composition between the spider species are small (Fig. 7). Most prey items are 1–2 mm long: only a few insects exceed 30 mm body length (Figs. 9–12). Relative to the available prey, some groups (e.g. Nematocera, Aphidoidea, Psocoptera) are caught selectively, while other groups (e.g. Heteroptera, Coleoptera, Brachycera, Orthoptera) are underrepresented in the prey spectrum and obviously avoid orb webs (Table 7). The differences in prey composition between araneids of the tropics and of the temperate zone are discussed (Table 8) and compared to those recorded in other studies (Table 9, 10). Most of these report large numbers of big prey items (Odonata, Lepidoptera, wasps/bees). It is pointed out that those studies do not take into account the total available prey in a spider's web but only that part which the spider selects from the web (mainly according to size). The importance of small prey items even for large spiders is explained and an obvious lack of niche partitioning among coexisting araneids is discussed (Table 11).     

Genital damage in the orb-web spider Argiope bruennichi (Araneae: Araneidae) increases paternity success

The morphology of male genitalia often suggests functions besidessperm transfer that may have evolved under natural or sexualselection. In several species of sexually cannibalistic spiders,males damage their paired genitalia during mating, limitingthem to one copulation per pedipalp. Using a triple-mating experiment,we tested if genital damage in the orb-web spider Argiope bruennichiincreases male fitness either through facilitating his escapefrom an aggressive female or by obstructing the female's inseminationducts against future copulation attempts from other males. Wefound no survival advantage for males damaging their pedipalps;however, copulations into a previously used insemination ductwere significantly shorter when the previous male had left partsof his genitalia inside the insemination duct. Because copulationduration determines paternity in this species, our result suggeststhat male genital damage in A. bruennichi is sexually selected.By breaking off parts of their intromittent organs inside avirgin female, males can reduce sperm competition and therebyincrease their paternity success.     

Previous experience and site tenacity in the orb spider Nephila (Araneae,Araneidae)

Summary The tenacity of the orb spider Nephila clavipes to a web site was studied in the laboratory. No differences were found between the giving-up-times and the site tenacity of spiders reared in the laboratory or those caught in the field, nor between spiders raised under a poor or a richt diet. The animals left sites at random and seemed to ignore experiences gained at previous sites.     

Feeding ecology of the orb-weaving spiderArgiope aurantia [Araneae: Araneidae] in a cotton agroecosystem

Twenty females of the orb-weaving spiderArgiope aurantia Lucas were introduced into a cotton field in east Texas in order to study the feeding ecology of this spider. In the 24 h after the release of these spiders in the cotton field, one had moved over a distance of 53 m. The released spiders spun webs with an average diameter of 33.5 cm with the hub an average of 39 cm above the ground. The diet ofA. aurantia was diverse which characterizes this species as a food generalist. Major food components were aphids (30%), Diptera (26.8%), grasshoppers (17.9%), and Hymenoptera (12.6%). The spiders' prey length ranged from 0.4 to 47 mm (mean =7.7±0.83 mm). Adult females ofA. aurantia have the potential to kill prey of up to ca. 200% of their own size. However, two-thirds of the prey items had a length of <3 mm, while only 25% of the prey items had a length of ≥20 mm.A. aurantia was found to be a predator of the cotton fleahopper (about 1% of the spiders' diet), which is a key pest of cotton.      

Web-building management in an orb-weaving spider, Zygiella x-notata: influence of prey and conspecifics

According to optimal foraging theory, spiders should adapt their web building to environmental variations. Until now, there was no data on the influence of simultaneous information coming from different environmental factors on web building behaviour. Under laboratory conditions, we studied the behaviour of Zygiella x-notata in the presence of prey, conspecifics, or both simultaneously. There was a stimulating effect of prey, but web building was not affected by the presence of conspecifics. When spiders and prey were present simultaneously, the effect was similar to that of prey alone; it seemed that there was no interactive influence of both factors. We discussed about the use of environmental information by spiders in foraging behaviour.     

Colony size and parasitoid load in two species of colonial Metepeira spiders from Mexico (Araneae: Araneidae)

Summary While a number of advantages may result from group living, it may also lead to increased levels of attack by parasites because groups may be easier to find. This leads to the prediction that levels of parasitism should increase with colony size. We test this prediction by comparing colony size and parasitoid load for two species of colonial orb-weaving spiders from Mexico, Metepeira (undesc. sp., tentatively named atascadero) and Metepeira incrassata, which exhibit contrasting levels of social organization and utilize different habitats. For M. atascadero, which occurs solitarily or in small groups in desert/mesquite grassland habitat, rates of egg-sac parasitism fluctuate widely from year to year, and are closely tied to spider egg output. There is no relationship between colony size and rate of parasitism. For colonial M. incrassata, which occur in tropical rain forest/agricultural habitat, rates of parasitism are relatively constant from year to year. However, there is a positive relationship between colony size and rate of parasitism in this species. These differences are discussed with regard to the stability of the two habitats, prey availability, and the foraging behavior of the respective parasitoids.     

Copulatory mechanism in a sexually cannibalistic spider with genital mutilation (Araneae: Araneidae: Argiope bruennichi)

Genitalia are among the fastest evolving morphological traits as evidenced by their common function as diagnostic traits in species identification. Even though the main function of genitalia is the successful transfer of spermatozoa, the presence of diverse structures that are obviously not necessary for this suggests that genitalia are a target of sexual selection. The male genitalia of many spider species are extremely complex and equipped with numerous sclerites, plates and spines whose functions are largely unknown. Selection on male genitalia may be particularly strong in sexually cannibalistic spiders, where mating success of males is restricted to a single female. We investigated the copulatory mechanism of the sexually cannibalistic orb weaving spider Argiope bruennichi by shock freezing mating pairs and revealed a complicated interaction between the appendices and sclerites that make up the male gonopods (paired pedipalps). The plate that covers the female genital opening (scape) is secured between two appendices of the male genital bulb, while three sclerites that bear the sperm duct are unfolded and extended into the female copulatory opening. During copulation, females attack and cannibalise the male and males mutilate their genitalia in about 80% of cases. Our study demonstrates that (i) genital coupling is largely accomplished on the external part of the female genitalia, (ii) that the mechanism requires an interaction between several non-sperm-transferring structures and (iii) that there are two predetermined breaking points in the male genitalia. Further comparative work on the genus Argiope will test if the copulatory mechanism with genital mutilation indeed is an adaptation to sexual cannibalism or if cannibalism is a female counter adaptation to male monopolisation through genital plugging.     

Sublethal effects of two neurotoxican insecticides on Araneus pratensis (Araneae: Araneidae)

Spiders are important predators of several agricultural pests and they play an important role as indicators of ecosystem disturb. In Argentina, soybean crop has increased from the introduction of transgenic soybean resistant to glyphosate. This expansion produced an increase in the use of conventional and non-selective pesticides to control soybean pests. The objective of this work was to evaluate the side effects of subletal concentrations of two neurotoxican insecticides with a different mode of action: endosulfan (Glex, 35%, 25 mg/l a.i.) and spinosad (Tracer, 48%, 30 and 3 mg/l a.i) on Araneus pratensis. The insecticides were applied by ingestion of the treated prey (Musca domestica), and the effects on mortality, prey consumption, web building, mating, ootheca construction and fecundity were determined. Spinosad (30 mg/l a.i.) produced higher mortality than endosulfan (25 mg/l a.i.). Tremors and non-coordinated movements were observed in this treatment. The prey consumption was significantly reduced by the two insecticides (approximately 40% lower than control). The spider web building was significantly affected by the two insecticides, but spinosad had a greater effect. Though mating was not affected by both pesticides, abnormal oothecas and dehydrated eggs were observed. This work reports that sublethal concentrations representing approximately from 25 to 2.5% of the maximum field recommended concentrations (105 and 120 mg/l a.i., respectively) showed negative effects on A. pratensis. The consequences of these effects on role of A. pratensis as a natural mortality factor of soybean pests are discussed.     

A new combination expands the range of the African araneid spider Singafrotypa (Araneae, Araneidae)

Study of the syntype of Larinioides subinermis, a species known from Ethiopia only, revealed that it actually belongs to Singafrotypa Benoit, 1962. We redescribe Singafrotypa subinermis (Caporiacco, 1940), comb. n., and provide a key to females of four species belonging to Singafrotypa. A distribution map for all species is provided.     

Sexual cannibalism, competition, and size dimorphism in the orb-weaving spider Nephila plumipes Latreille (Araneae: Araneoidea)

The degree and direction of sexual dimorphism varies widely,but in several taxa of orb-weaving spiders, including Nephila,males may be less than one-tenth the size of females. This differenceis commonly attributed to selection through precopulation sexualcannibalism: females may either fail to detect very small males,or ignore them as potential prey items. However, there is oftenthe potential for male-male competition in these species becauseseveral males can be found on the web of a single female. Weinvestigated experimentally the effects of sexual cannibalismand male-male competition on male body size and hence sexualdimorphism in the Australian golden orb-weaver (Nephila plumipes).Small males were less likely to be detected and cannibalizedthan larger males. However, larger males excluded small malesfrom the central hub of the web, where mating takes place. Theconflicting effects of sexual cannibalism and male-male competitionmay be responsible for the relatively large variation in malebody size in this species.     

State-dependent prey type preferences of a kleptoparasitic spider Argyrodes flavescens (Araneae: Theridiidae)

Spiders from the theridiid genus Argyrodes exhibit considerable variation in foraging tactics. However, little is known about the conditions under which Argyrodes spiders switch foraging tactics. Argyrodes flavescens (Pickard-Cambridge) is commonly found in the webs of another spider Nephila pilipes (Fabricius) in Singapore. In this study, a series of prey-choice tests were conducted for A. flavescens , both in the presence and absence of N. pilipes , to investigate the state-dependent prey type preference of A. flavescens . It was found that, in the absence of N. pilipes , well-fed A. flavescens took houseflies more than fruit flies, but starved A. flavescens took more fruit flies than houseflies. Whether N. pilipes spiders were present or absent, both well-fed and starved A. flavescens preferred living prey and rarely took wrapped prey of any kind. When well fed, A. flavescens rarely took mealworms. However, when starved, A. flavescens tended to take freshly captured prey, and also tended to feed together with N. pilipes on a housefly or mealworm captured by N. pilipes . Whether A. flavescens were absent or present, both well-fed and starved N. pilipes took mealworm larvae more often than they took houseflies, and they never attacked fruit flies. This is the first study to show that Argyrodes spiders alter their foraging tactics depending on hunger level, prey type, or the presence of the host. In doing so, Argyrodes spiders may maximize their energy gain and minimize predation risk in different circumstances.     

Effects of gravity on temporary spiral construction byLeucauge mariana (Araneae: Araneidae)

Temporary spirals (TSP) in non-horizontal webs ofLeucauge mariana (Key-serling) showed consistent up-down asymmetries in initiation sites and spacing between loops that were not present in horizontal webs, even when possible effects of other web asymmetries were controlled for by changing the web's orientation just as TSP construction began. Cues from the web itself also affect spacing, since patterns of TSP placement differed between webs spun entirely at a given angle (e.g. 45°), and webs whose orientations were changed to that angle (e.g. 45°) just as TSP construction began. Spaces between successive TSP loops increased both above and below the hub, except for those of the outermost loop.