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.     

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.     

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.     

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.     

Effects of prey–spider odour on intraspecific interactions of araneophagic jumping spiders

An important prediction from game theory is that the value of a resource will influence the level to which an animal is willing to escalate during conflict with conspecific rivals. Here we use this prediction as the rationale for experiments aimed at determining whether escalation decisions made by predators are influenced by the presence of preferred prey. The predators we use are adult individuals of Portia fimbriata, a jumping-spider (Salticidae) species from Queensland (Australia). P. fimbriata is known to prefer other salticid species as prey and can detect the odour of Jacksonoides queenslandicus, this being an especially common salticid species in the Queensland habitat. Here we show that, for P. fimbriata females, the odour of J. queenslandicus primes escalation of vision-based conflict. However, for P. fimbriata males, no comparable effect was found. These findings suggest intrasexual competition for prey is more important to females than to males.     

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.     

Observations in nature of detouring behaviour by Portia fimbriata, a web-invading aggressive mimic jumping spider from Queensland

Portia fimbriata , a web-invading, araneophagic salticid that uses aggressive mimicry to deceive its prey (web-building spiders), takes indirect routes to reach its prey (i.e. it makes detours). Data are presented from 18 instances of Portia making detours to reach prey in nature, the prey being five different species of web-building spiders. Portia spent 17 min (median) per predatory sequence with the prey out of view and covered 375 mm detouringper sequence. These detours were longer and more complex than those previously recorded for salticids, and these are the first detailed records of detouring behaviour by a salticid in the field.     

Side-effects of insecticides on two erigonid spider species

The current rearing technique forErigone atra (Blackwall) andOedothorax apicatus (Blackwall) (Araneae, Erigonidae) was improved. To reduce time spent rearing on live fruit flies the spiders were kept on a culture of the Collembola speciesLepidocyrtus lanuginosus (Gmelin) (Entomobryidae). Side-effects on spiders of two pyrethroid insecticides (fenvalerate and lambda-cyhalothrin) and one carbamate insecticide (pirimicarb) were tested. Sensitivity of adults of both sexes and juveniles to insecticides and their influence on the rate of emergence of spiderlings from cocoons were investigated using topical application, spraying or residual contact. LD₅₀ values for adults ranged from 0.49 to 2.52 ng a.i./spider for lambda-cyhalothrin and from 5.75 to 98.20 ng a.i./spider for fenvalerate. Topical application also resulted in up to a week's delay of web-building. A moving laboratory spraying equipment was used to spray spiders with different insecticide dosages and water volumes. Pyrethroids sprayed onto adults in webs had stronger effects than pyrethroids sprayed onto sitting or walking spiders on the soil surface. Residual contamination caused higher mortality of spiders after contact with lambda-cyhalothrin than fenvalerate. In all tests, males were more susceptible to pyrethroids than females; this difference was related to body weight. Mortality rate was higher forE. atra than forO. apicatus. Both pyrethroids were also toxic to spiderlings. Lambda-cyhalothrin inhibited emergence ofE. atra spiderlings from cocoons. Pirimicarb was harmless to both spider species.     

Predatory and nesting behaviour of Cocalus gibbosus,a spartaeine jumping spider (Araneae: Salticidae) from Queensland

Abstract

Cocalus gibbosus was studied in the field in Queensland and in the laboratory. This is the first behavioural study of a species from the spartaeine genus Cocalus. C. gibbosus often omitted elements which are usually present in the predatory sequences of typical salticids and tended to lunge at prey from close range rather than leap from afar. Experiments showed that C. gibbosus prefers moths to other prey. In nature, C. gibbosus moulted and oviposited on silk sheets spun against tree trunks, and in the laboratory on sides of cages or blocks of wood, but this species never built an enclosing nest like typical salticids nor a large prey-catching web like some other spartaeines. C. gibbosus stalked across alien webs to catch spiders and insects, but it did not make vibratory signals. It did not stick to cribellate or ecribellate glue on alien webs. The behaviour of C. gibbosus is compared to that of other spartaeine salticids.     

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.     

Interactions between citrus psylla,Trioza erytreae (Hem. Triozidae), and spiders in an unsprayed citrus orchard in the transvaal lowveld

During a two year survey, a total of 3,054 spiders represented by 21 families were sampled in an unsprayed citrus orchard in the Transvaal Lowveld. Numerically the Salticidae was the dominant family (34.4%) followed by the Theridiidae (21.9%), Thomisidae (11.9%), Araneidae (7.9%), Clubionidae (7.0%) and the Tetragnathidae (3.7%). Eighteen species of spiders were observed to prey on citrus psylla,Trioza erytreae (Del Guercio) (Hemiptera: Triozidae), while six species trap nymphs and adults under their retreats and webs. There were significant positive correlations between the weekly psylla populations and the weekly populations of web-building spiders and wandering spiders present one to four weeks later but no significant correlation between the weekly spider populations and the weekly psylla populations present one to five weeks later. This seems to indicate that while spiders are unable to keep citrus psylla populations at acceptable low levels, they may contribute in reducing their numbers.      

蛛网结构性能及其适应性

蛛网是蜘蛛的捕食工具,蛛网的结构性能不仅影响蜘蛛的捕食效率,也关系着蜘蛛的捕食投入。在不同的内外环境条件影响下,蜘蛛会通过蛛网结构性能上的相应变化来调整捕食策略和维持网结构的稳定性。本文主要综述了蛛网的结构性能以及蜘蛛通过蛛网结构性能表现出的对环境因子的适应性。     

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.     

Impact of predation by spiders on a territorial damselfly (Odonata: Calopterygidae)

Summary Predation by orb-weaving spiders and crab spiders on the damselfly Calopteryx haemorrhoidalis was studied at a small stream in Southern France. One species of orb-weaving spider, Larinioides folium, caught 76% of the damselflies which fell prey to spiders. Displacement experiments on spiders on sections of bank and the positioning of webs in male territories show that the density and distribution of damselflies is not influenced by orbweb density or by the position of webs. Predation rates corresponded to orb-web density, but neither for sex nor for stage was there a relationship with damselfly density. Mean daily predation rates ranged between 0.9% for females and 4.1% for adult males. Predation risk to adult damselflies by orb-weaving spiders was male biased, whereas among tenerals there was no bias. Males were captured more frequently at territories near the water. Captures show a maximum at noon when territorial disputes of adult males were most frequent. After orbwebs were placed within territories predation rate of males was strongly increased. Predation risk to adult females in the direct vicinity of the stream was less than in the bank vegetation where they perch close to orbwebs. The risk of predation by crab spiders, which catch damselflies at their perching sites, was not sex-biased.     

Evidence of exploitative competition among young stages of the wolf spider Schizocosa ocreata

Summary Previous research by many investigators has demonstrated food limitation in both web-building and wandering spiders. Field experiments have tested for exploitative competition for prey in web-building, but not wandering species. As a first step to examining the question of whether spiders without webs exhibit exploitative competition, we manipulated densities of young stages of a common wolf spider, Schizocos ocreata, and measured (1) spider growth rate and (2) numbers of Collembola, a potential prey organism. Replicate populations of recently hatched S. ocreata were established in 1-m² fenced plots at four levels: 0×, 0.25×, 1× and 4× natural density. Increasing spider density had a negative effect on spider growth rate, defined as increase in weight or cephalothorax width. Early in the experiment spider density had a weak negative effect on Collembola numbers [p(F)=0.08]. Taken together, this probable response by Collembola and the clear effect of spider density on growth rate constitute the first experimental evidence of intraspecific exploitative competition for prey in a species of wandering spider. We discuss (1) the strength of this evidence given the constraints of the experiment's design, and (2) the implications of the strong convergence in spider densities that had occurred after 2.5 months.     

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.     

Aging and foraging efficiency in an orb-web spider

Aging is often associated with reduced behavioral performance such as decreased locomotion or food consumption, related to a deterioration in physiological functions. In orb-web spiders, webs are used to capture prey and aging can affect web-building behavior and web structure. Here, we investigated the effect of aging on prey capture in the orb-web spider Zygiella x-notata. The ability of adult females to capture flies was examined at different ages. The rate of prey capture did not change with age, but older spiders took more time to subdue and capture the prey. Alterations which appeared in web structure with age (increase in the number of anomalies affecting radii and capture spiral) affected prey capture behavior. Furthermore, the analysis of individual performance (carried out on 17 spiders at two different ages) showed that older females spent more time handling the prey and finding it in the web. Our results suggest that, in the laboratory, age does not affect prey capture rates but it influences prey capture behavior by affecting web structure or/and spider motor functions.     

Kleptoparasites influence foraging behaviour of the spider <Emphasis Type="Italic">Stegodyphus lineatus</Emphasis> (Araneae,Eresidae)

Prey captured by a predator may attract kleptoparasites which could significantly reduce the amount of food consumed. Stegodyphus lineatus, a cribellate spider, builds an energetically costly web. Ants raid the webs of S. lineatus to steal prey and behave as kleptoparasites. We investigated ant raids in a natural population of S. lineatus and their influence on the spider’s foraging behaviour. Considering spiders that had captured a prey, 31.2% suffered an ant raid within 24 h after the prey capture. Experimental tests showed that the response to ant raid is to delay web rebuilding and this was independent of a spider’s previous foraging success. There was a tendency for spiders that were exposed to ants to build larger webs. Neither prey-handling duration nor prey consumption was modified after exposure to ants. These results suggest that Stegodyphus lineatus adapt its web-building behaviour in response to the risk of kleptoparasitism.     

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.     

Risk Sensitivity and the Paradox of Colonial Web-Building in Spiders

Group foraging is rare in spiders, occurring only where preyavailability is high. If colonial web-building increases individualprey capture rates as shown, why does group foraging not occurmore often where prey are scarce? Risk sensitivity may explainthis paradox, as variance in prey capture is reduced in groups;risk-averse spiders should join groups only when prey exceeda threshold level. Field studies show that group foraging variesas predicted between species, between populations of a singlespecies, and between sites within a population. However, recentmodels suggest the necessity of examining variance within individualsover time rather than between individuals within populations.Additionally, mechanisms responsible for variance reductionin colonial webs may be less effective than previously assumed.New field data suggest that while prey variance over time maybe somewhat less for individual spiders in groups than for solitaries,the relationship between colonial web-building and variancein prey capture is far more complex than originally thought.The influence of risk sensitivity on reproductive success andthe evolution of colonial web-building is discussed.