Comparative biology of Portia africana,P. albimana,P. fimbriata,P. labiata,and P. shultzi,araneophagic, web-building jumping spiders (Araneae: Salticidae): Utilisation of webs,predatory versatility,and intraspecific interactions

Abstract

Portia is a behaviourally complex and aberrant salticid genus. The genus is of unusual importance because it is morphologically primitive. Five species were studied in nature (Australia, Kenya, Malaysia, Sri Lanka) and in the laboratory in an effort to clarify the origins of the salticids and of their unique, complex eyes. All the species of Portia studied were both web builders and cursorial. Portia was also an araneophagic web invader, and it was a highly effective predator on diverse types of alien webs. Portia was an aggressive mimic, using a complex repertoire of vibratory behaviour to deceive the host spiders on which it fed. The venom of Portia was unusually potent to other spiders; its easily autotomised legs may have helped Portia escape if attacked by its frequently dangerous prey. Portia was also kleptoparasitic and oophagic when occupying alien webs. P. fimbriata from Queensland, where cursorial salticids were superabundant, used a unique manner of stalking and capturing other salticids. The display repertoires used during intraspecific interactions were complex and varied between species. Both visual (typical of other salticids) and vibratory (typical of other web spiders) displays were used. Portia copulated both on and away from webs and frequently with the female hanging from a dragline. Males cohabited with subadult females on webs, mating after the female matured. Adult and subadult females sometimes used specialised predatory attacks against courting or mating males. Sperm induction in Portia was similar to that in other cursorial spiders. Portia mimicked detritus in shape and colour, and its slow, mechanical locomotion preserved concealment. Portia occasionally used a special defensive behaviour (wild leaping) if disturbed by a potential predator. Two types of webs were spun by all species (Type 1, small resting platforms; Type 2, large prey-capture webs). Two types of egg sacs were made, both of which were highly aberrant for a salticid. Responses of different species and both sexes of Portia were quantitatively compared for different types of prey. Many of the trends in behaviour within the genus, including quantitative differences in predatory behaviour, seemed to be related to differences in the effectiveness of the cryptic morphology of Portia in concealing the spider in its natural habitat (‘effective crypsis’). The results of the study supported, in general, Jackson & Blest’s (1982a) hypothesis of salticid evolution which, in part, proposes that salticid ancestors were web builders with poorly developed vision and that acute vision evolved in conjunction with the ancestral spiders becoming proficient as araneophagic invaders of diverse types of webs.     

The biology of Tauala lepidus,a jumping spider (Araneae: Salticidae) from Queensland: display and predatory behaviour

Abstract

The display and predatory behaviour of Tauala lepidus Wanless, an abundant salticid in north Queensland rainforests, was investigated in the laboratory and, to a lesser extent, in nature. T. lepidus leapt and walked into alien webs to catch spiders and insects, and was captured and fed on by other spiders. Females ate each other’s eggs. During intraspecific interactions, a complex repertoire of displays was used. Courtship versatility occurred, each individual male having a conditional strategy of different behaviours depending on whether the female is at or away from her nest, and whether she is adult or subadult. Yet other combinations of displays occurred during male-male and female-female interactions. Apparently, pheromones on nests and draglines of females released male courtship. Abdomen twitching, a behaviour common to the display repertoires of many salticids, was an especially complex and pervasive behaviour of T. lepidus. T. lepidus also twitched its abdomen when it contacted alien webs and preyed on other species of spiders. The behaviour of T. lepidus is compared to that of Jacksonoides queenslandica Wanless, a species from the same group (Astieae).     

The comparative study of the predatory behaviour of Myrmarachne, ant-like jumping spiders (Araneae: Salticidae)

The predatory behaviour of 31 species of Myrmarachne , ant-like salticids, was studied in the laboratory and the field. The ant-like morphology and locomotion of these spiders appears to function primarily in Batesian mimicry. No evidence was found of Myrmarachne feeding on ants. However, predatory sequences were found to differ considerably from those typical of salticids. Instead of stalking and leaping on prey, Myrmarachne lunged at prey from close range. Myrmarachne used its legs I to tap prey before lunging, another unusual behaviour for a salticid. Myrmarachne fed on a wide range of arthropod prey in nature and the laboratory, but appears to be especially efficient at catching moths. Also, Myrmarachne tends to open up, or enter into, other spiders' nests and eat other spiders' eggs. Myrmarachne males were less efficient than females, in laboratory tests, at catching various types of arthropod prey, but they appear to be as efficient as females at oophagy. Myrmarachne tend to use webs of other spiders as nest sites, but no evidence was found of Myrmarachne preying on spiders in webs. It appears that the unusual features of Myrmarachne's predatory and nesting behaviour are important in enabling these spiders to preserve their ant-like appearance.     

Out of the Frying Pan and into the Fire: a Novel Trade-Off for Batesian Mimics

A mimicry system was investigated in which the models were ants (Formicidae) and both the mimics and the predators were jumping spiders (Salticidae). By using motionless lures in simultaneous‐presentation prey‐choice tests, how the predators respond specifically to the static appearance of ants and ant mimics was determined. These findings suggest a rarely considered adaptive trade‐off for Batesian mimics of ants. Mimicry may be advantageous when it deceives ant‐averse potential predators, but disadvantageous in encounters with ant‐eating specialists. Nine myrmecophagic (ant‐eating) species (from Africa, Asia, Australia and North America) and one araneophagic (spider‐eating) species (Portia fimbriata from Queensland) were tested with ants (five species), with myrmecomorphic (ant‐like) salticids (six species of Myrmarachne) and with non‐ant‐like prey (dipterans and ordinary salticids). The araneophagic salticid chose an ordinary salticid and chose flies significantly more often than ants. Portia fimbriata also chose the ordinary salticid and chose flies significantly more often than myrmecomorphic salticids. However, there was no significant difference in how P. fimbriata responded to ants and to myrmecomorphic salticids. The myrmecophagic salticids chose ants and chose myrmecomorphic salticids significantly more often than ordinary salticids and significantly more often than flies, but myrmecophagic salticids did not respond significantly differently to myrmecomorphic salticids and ants.     

Opportunistic Egg Feeding in the Kleptoparasitic Spider Argyrodes gibbosus

Argyrodes gibbosus is a kleptoparasitic spider in the web of spiders. It generally steals freshly captured prey from the web of its host. In Sicily, Argyrodes gibbosus parasitizes webs of the spider Cyrtophora citricola, a facultative colonial species. When a C. citricola female was present in its web, Argyrodes caught small prey in the web or tried to rob prey captured by the host; in that case, we never observed successful attacks on host egg-sacs. When the host disappeared from its web, the kleptoparasite modified its foraging strategies and attacked the host egg-sacs and ate the eggs. The exploitation of this new resource could ensure rapid development for the kleptoparasite which was characterized by the presence of larger females and a higher mating rate.     

Prey preferences ofPortia fimbriata,an araneophagic,web-building jumping spider (Araneae: Salticidae) from Queensland

Portia fimbriata from Queensland, a previously studied jumping spider (Salticidae), routinely includes web-building spiders and cursorial salticids in its diet, both of these types of prey being dangerous and unusual prey for a salticid. The present paper is the first detailed study ofP. fimbriata's prey preferences. Three basic types of tests of prey preference were used, providing evidence that (1)P. fimbriata males and females prefer spiders (both web-building spiders in webs and salticids away from webs) to insects; (2)P. fimbriata males and females prefer salticids to web-building spiders; (3)P. fimbriata males and females prefer larger spiders to smaller spiders; (4) there are intersexual differences in the preferences ofP. fimbriata for prey size, females preferring larger prey and males preferring smaller prey; and (5)P. fimbriata's prey preferences are not affected by a prior period without food of 2 weeks. When preferences were tested for by using both living, active prey and dead, motionless lures, the same preferences were expressed, indicating thatP. fimbriata can distinguish among different types of prey independent of the different movement patterns of different prey.     

The biology of Euryattus sp. indet., a web-building jumping spider (Araneae, Salticidae) from Queensland: utilization of silk, predatory behaviour and intraspecific interactions

The biology of an aberrant saltioid spider, Euryattus Thorell sp. indet., is described from observations in a Queensland rain forest and the laboratory. Pronounced morphological and behavioural changes occur during post-embryological development. Juveniles spin webs, but adult females make 'suspension nests' by suspending a curled-up leaf by heavy guylines from a rock ledge or vegetation. Adult males spin neither webs nor suspension nests. Females oviposit inside their suspension nests, but if denied access to leaves for suspension, they spin and oviposit in webs similar to those spun by juveniles. The flat, papery egg sacs of Euryattus are atypical for a salticid, being more like the egg sacs of many of the Gnaphosidae. Intraspecific display behaviour has characteristics in common with typical salticids, but also includes unique features. Male courtship includes vibratory displays performed on the surface of the suspended leaf. Mating occurs inside the curled-up leaf. Males co-habit with sub-adult females in suspension nests. Females take over suspension nests of other females and eat each other's eggs. Normal locomotion, intraspecific interactions, and predatory behaviour are characterized by frequent leaping. Euryattus routinely makes long and accurate leaps on to prey, including flying insects intercepted in mid-air and spiders and insects located in alien webs. The phylogenetic implications of the unusual characteristics of Euryattus are considered.     

Observations of Portia Africana,an araneophagic jumping spider,living together and sharing prey

Abstract

Instances are documented of finding individuals of Portia africana in the field living aggregated in the webs of other spiders, in the nest complexes of other salticids, around solitary nests of other salticids, and around the nests of oecobiid spiders. Aggregation members included all active juvenile stages of P. africana, as well as adult males and females. More than one individual of P. africana sometimes fed on the same prey. Small juveniles of P. africana were more often than other stages found aggregated and more often observed feeding together. Small juveniles of P. africana surrounded the nests occupied by other salticid genera and nests occupied by oecobiid spiders. When the resident salticid or oecobiid attempted to leave or enter the nest, one of the P. africana juveniles lunged and captured it, after which other P. africana individuals sometimes joined to feed.     

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.     

Prey-capture techniques and prey preferences of Chrysilla, Natta and Siler, ant-eating jumping spiders (Araneae, Salticidae) from Kenya and Sri Lanka

The predatory behaviour of Chrysilla lauta and Sliersemiglaucus from Sri Lanka, and four species of Natta from Kenya, was studied in the laboratory for the first time. These salticids eat ants, a prey most salticids avoid. These species' specialized behaviour for catching ants is described and compared to their behaviour for catching other insects. Three different types of tests of prey preference were carried out and, in each type, ants were taken in preference to other insects. Preference for ants, and prey-specific predatory behaviour did not depend on prior experience with ants. Results from this study are discussed in relation to recent findings on other ant-eating salticids.     

Complex display behaviour of Evarcha culicivora,an East African mosquito‐eating jumping spider

Abstract

Evarcha culicivora, an East African jumping spider (Salticidae), is an unusual predator because it feeds indirectly on vertebrate blood by choosing blood‐carrying mosquitoes as preferred prey. It also associates with particular plant species, Lantana camara and Ricinus communis. Here we document this species’ exceptionally complex display repertoire. In common with many other salticids, E. culicivora exhibits pronounced courtship versatility, with males using different tactics depending on the female's location and state of maturity. However, in contrast to most other salticids that have been studied, the males and the females of E. culicivora are both active at initiating and sustaining courtship, and both sexes are cannibalistic. Contrary to the emphasis in the literature on female spiders eating males, females of E. culicivora run a higher risk of being killed by males than vice versa during courtship. E. culicivora males also differ from other salticids that have been studied by adopting pronounced copulatory courtship. Male‐female interactions of E. culicivora are especially complex when encounters are in the foliage of L. camara and R. communis.     

The Trochidae and Turbinidae of the Kermadec Ridge (Mollusca: Gastropoda)

Abstract

Taieria erebus (Gnaphosidae) was found to be a versatile predator: it captured insects both cursorially (away from webs) and kleptopar-asitically (on alien webs); it captured spiders in both the presence and absence of webs; and it also ate the eggs of host spiders (oophagy). When T. erebus invaded webs, it was as an aggressive mimic — it performed a repertoire of vibratory behaviours to lure the host spider. Although T. erebus pursued and captured spiders on diverse web-types, it was more effective as a predator when invading densely (rather than sparsely) woven cribellate and non-sticky webs, and was especially effective on non-cribellate sticky webs. Gnaphosids are traditionally referred to as hunting spiders, but T. erebus built a small prey-capture web. T. erebus also preyed on segestriid spiders, then used their webs to catch more prey, this being an unusual example of a spider using, as a tool for predation, the spinning-work of another species from an unrelated family. T. erebus used specialised behaviours to prey on nesting cursorial spiders. Prey was either grasped or stabbed; the venom of T. erebus was highly potent against spiders. Experiments indicated that vision was of little or no importance in the predatory behaviour of T. erebus. The behaviour of T. erebus is compared to that of Portia, a web-building salticid spider which is very versatile in its predatory behaviour and has acute vision. T. erebus is discussed in relation to hypotheses concerning gnaphosid and salticid evolution.     

Fish predation on Notonecta (Hemiptera): relationship between prey risk and habitat utilization

Summary Prey risk was examined in the laboratory to compare the ability of 6 Notonecta species to coexist with insectivorous fishes (Lepomis cyanellus, L. gibbosus). Because of their smaller size, lighter color, greater tendency to remain motionless in the presence of the predator, greater ability to avoid capture when attacked, and predicted lower profitability as prey, N. lunata and N. petrunkevitchi were estimated to have the lowest prey risk and greatest chance of coexisting with insectivorous fishes in nature. The 2 largest notonectids, N. irrorata and N. insulata, both highly melanistic and rapidly discovered by foraging sunfishes, were judged to be the most susceptible to predation by large fishes. The presence of vegetation as a potential refuge tended to decrease prey risk but did not significantly alter the relative risk among the prey species. Finally, the relative prey risk measured in the laboratory was consistent with the general distribution of these species in relation to fishes in local habitats. We suggest that fish predation is an important determinant of Notonecta habitat utilization patterns.     

Conditional strategies and interpopulation variation in the behaviour of jumping spiders

Abstract

Conditional strategies and interpopulation variation in the mating and predatory behaviour of salticid spiders are reviewed. A functional approach is adopted, and defended, in which specified behavioural phenotypes are accounted for, in large part, by specified selection factors. Courtship versatility, in which a male's behaviour depends on the female's maturity and location, is common in the Salticidae. If a male encounters an adult female in the open, where there is ample ambient light, he performs vision-dependent displays (Type 1 courtship) in front of her. If he encounters an adult female inside her nest, he uses different displays (Type 2 courtship) which are not vision–dependent and consist of various tugging, probing and jerking movements on the silk of the nest. These displays apparently send vibratory stimuli to the female. When a male encounters a subadult female inside her nest, he initially performs Type 2 courtship, then spins a second chamber on the nest and cohabits until the female moults and matures. A modification of optimal foraging theory has been used to examine factors that influence interpopulation variation in male courtship persistence. A study of five populations corroborated predictions from the model. Persistence appears to be related to female availability. Female availability is related to local phenology, which is, in turn, related to local climate. Complex examples of predatory versatility also have evolved in the Salticidae, especially in the genus Portia. All species of Portia studied are araneophagic spiders that invade other spiders' webs and practise aggressive mimicry. Portia fimbriata, uniquely among Portia species studied, uses specialised behaviour to prey on other salticids. Portia fimbriata and one of the salticids on which it preys, Euryattus sp., appear to be co-adapted to each other.     

Introduction of Alien Host–parasite Complexes in a Natural Environment and the Symbiota Concept

When a species is introduced in a new territory its parasites may also follow, thus forming a sort of biotic unit. We propose the term symbiota to indicate this host–parasite complex. This concept is at the basis of the present research in which 11 species of alien and native freshwater fish were collected by electrofishing in the Po river area (Northern Italy). The alien host fish Ictalurus melas and Silurus glanis were parasitized by only one monogenean species each. Lepomis gibbosus, of North American origin, hosted four monogeneans and Carassius carassius three. All these parasites should be considered as alien species, except those collected from C. carassius that should be considered in part natives and in part aliens.     

The biology of Portia fimbriata, a web-building jumping spider (Araneae, Salticidae) from Queensland: intraspecific interactions

Portia fimbriata (Doleschall) is an unusual salticid because it spins webs and uses its own webs and those of other species in predation. However, the courtship and threat displays of this species are more like those of typical, cursorial salticids than like typical web-building spiders. During male-female interactions, males perform leg-waving and leg-shaking displays, with legs I and II extended stiffly forward, while distant from the females. Before mounting they tap the females with their legs; and after mounting they tap, scrape and stroke the females' abdomens. Copulation takes place on or near the female's web or on the webs of other species. Hunched-legs displays, with legs I–III highly flexed and held to the side of the body, occur in male-male interactions. Also, during male-female and female-female interactions, females perform hunched-legs displays, strike, charge, ram, embrace, grapple with and leap at conspecifics. Sometimes they lose legs while grappling. After female-female interactions associated with maternal webs, if the resident decamps, the intruder eats the eggs left behind on a suspended leaf, spins a new egg-case over the destroyed one, and oviposits. Adult and sub-adult males co-habit in webs with sub-adult females for as long as 48 days and mate when the spiders mature.     

Foraging traits of native predators determine their vulnerability to a toxic alien prey

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Testing the Functions of Detritus Stabilimenta in Webs of Cyclosa fililineata and Cyclosa morretes (Araneae: Araneidae): Do They Attract Prey or Reduce the Risk of Predation?

Spiders of the genus Cyclosa often add prey remains and other debris to their orb‐webs. The function of silk decorations is generally associated with defense against predators or with the attraction of prey, but few studies have focused on stabilimenta containing detritus. In this study, we used artificial webs with and without the detritus stabilimenta of two species of Cyclosa to investigate whether these structures increase the number of insects intercepted. Artificial models of spiders and stabilimenta were used to compare the frequency of attacks against different shapes. We also conducted choice experiments in laboratory to determine whether detritus columns attracted Drosophila melanogaster (Diptera: Drosophilidae) and Trigona angustula (Hymenoptera: Apidae, Meliponinae) to the webs. The frequency of interception in artificial webs with a stabilimentum was similar to that of webs without such structure. The taxonomic composition and biomass of insects were also similar in both types of artificial webs. The choice experiments showed no significant tendency in attraction to webs with a stabilimentum. However, models of spiders were attacked at a higher frequency than those simulating detritus columns and silk decorations. These findings argue against the prey attraction hypothesis and suggest that the addition of stabilimenta to webs of Cyclosa could reduce the intensity of predation, possibly by disrupting the image of the spider's outline.     

Predator-prey interactions between jumping spiders (Araneae, Salticidae) and Phokus phalangioides (Araneae, Pholcidae)

Portia is a genus of specialized web-invading salticids that use aggressive mimicry. Some other salticids leap into webs to catch spiders but do not use aggressive mimicry. Pholcus phalangioides is a web-building spider with a special defensive behaviour—called whirling—in which it swings its body around in a circle while keeping its long legs on the silk. Pholcus phalangioides is preyed on by Portia and probably other salticid spiders in nature. Interactions between P. phalangioides and 13 species of salticids were studied in the laboratory to compare how effective salticids with different styles of predation were at catching the pholcids. Four species of Portia were studied and each was more efficient at catching P. phalangioides than were the other nine salticids tested. For one species—Portia fimbriata—individuals from three different populations were studied. The Queensland P. fimbriata used aggressive mimicry more consistently and were more efficient at catching P. phalangioides than were the other species of Portia and the other populations of P. fimbriata . The salticids that were the most efficient at catching pholcids were also better able to avoid setting off whirling by the pholcids. An experiment in which pholcids were artificially induced to whirl whenever the predator was near provided additional evidence that whirling is an effective defence of pholcids against predation by salticids.     

Competition between the native and the introduced hornets Vespa crabro and Vespa velutina: a comparison of potentially relevant life‐history traits

1. Invasive alien species are a major threat to biodiversity. In addition to predation and parasitism, native species might suffer from competition when invasive alien species occupy a similar ecological niche. 2. This study focused on the potential interspecific interaction between two hornets: the Asian yellow‐legged hornet, Vespa velutina, a high‐concern invasive alien species recently arrived in Europe; and the native European hornet, Vespa crabro. The two species share a similar ecological niche and V. velutina is rapidly expanding across Europe, which suggests that V. crabro might suffer from competition. 3. Under laboratory‐controlled conditions, two life‐history traits that might cause the two species to compete were investigated: (i) the ability of workers to find food sources and their flexibility in exploiting them (through individual food item choice tests and exploration assays); and (ii) the worker resistance to pathogens (through immune challenge tests). 4. The results show that trophic preference of both species highly overlaps, with a marked dietary preference for honeybees compared with other insect prey and non‐prey protein items. No differences were observed in the exploratory behaviour of both species. Finally, constitutive antibacterial activity was greater in workers of the native species than in workers of the invasive hornet. 5. This laboratory study provides a first assessment under controlled conditions of the factors affecting competition between workers of two hornet species and proposes a framework to assess, in wild contexts, the magnitude of the competition and the impact of the introduced V. velutina on the native V. crabro.