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.     

Redescription of Natalaspis leptocarpi n.comb. (Homoptera: Diaspididae)

Abstract

Cyllobelus rufopictus? a salticid species from equatorial Africa, has a large and complex repertoire of displays. These spiders live on and near the ground in habitats of short grass, and build opaque silk nests on dead leaves. Each male uses one of three different mating tactics depending on the female’s maturity and location (communicatory versatility). Type 1 courtship, involving special movements and postures of the legs, palps, and body, occurs if the female is an adult away from the nest; apparently this type of courtship is vision-dependent. If the male encounters an adult female inside her nest, he uses Type 2 courtship, which consists of movements by the male that cause the silk to vibrate. If the female is a subadult inside her nest, the male initially uses Type 2 courtship, then builds a second chamber on the female’s nest, and cohabits until she moults and matures. Other displays occur during male-male and female-female interactions. Biting and pinching are distinctive Type 2 courtship displays that have not been described previously for salticids, and juddering is the most distinctive Type 1 display.     

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.     

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 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).     

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.     

Courtship versatility in the jumping spider, Phidippus johnsoni (Araneae: Salticidae)

A given male of P. johnsoni has at his disposal two alternative types of courtship. Each consists of distinctly different motor patterns and involves different sensory channels. If the male encounters a female outside her nest, he uses type 1 courtship. Type 2 courtship occurs if the male encounters a female inside her nest. Spiders do not respond to visual stimuli (house flies) under red light. The fact that type 2, but not type 1, courtship occurs when spiders are observed under red light is taken as evidence that type 1 is vision dependent, whereas type 2 is not vision dependent. Selection factors favouring the evolution of courtship versatility are discussed.     

Predator-prey interactions between web-invading jumping spiders and two species of tropical web-building pholcid spiders, Psilochorus sphaeroides and Smeringopus pallidus

Psilochorus sphaeroides from Queensland, Australia and Smeringopus pallidus from Sri Lanka are long-legged, web-building pholcid spiders with a special defence behaviour, whirling. The efficiency of whirling as a defence against web-invading jumping spiders (Salticidae) was examined in the laboratory. Three salticid species were used in these tests- Euryattus sp., Portia fimbriata and Portia labiata. Euryattus leapt into webs, but Portia fimbriata and Portia labiata walked slowly into webs and practised aggressive mimicry. Portia fimbriata was more consistent at using aggressive mimicry and more efficient at capturing Psilochorus sphaeroides and Smeringopus pallidus than was Portia labiata . Both species of Portia were more efficient at catching pholcids than was Euryattus. Portia , especially Portia fimbriata , was less inclined than Euryattus to stimulate pholcids to whirl. In an experiment in which pholcids were artificially induced to whirl whenever a salticid was near, salticids never captured pholcids, providing additional evidence that whirling is an effective defence of Psilochorus sphaeroides and Smeringopus pallidus against web-invaders. Results from this study are compared to those from a study of another pholcid that whirls, Pholcus phalangioides .     

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.     

The biology of ant-like jumping spiders: intraspecific interactions of Myrmarachne lupata (Araneae, Salticidae)

Myrmarachne lupata is an ant-like salticid in which males have very large chelicerae. The display repertoire of this species is unusually large and complex for a salticid spider. Each individual male uses one of three different mating tactics depending on the female's maturity and location. With adult females outside nests type 1 courtship occurs which seems to be a form of visual communication and includes specialized movements and postures of the legs, palps and body. With adult females inside nests, males use type 2 courtship, which seems to be a form of non-visual communication and consists primarily of probing with the legs on the silk; males mate with receptive females inside the nests. With subadult females, males first use type 2 courtship then spin an adjacent silken chamber and cohabit. After she moults and matures, mating occurs inside the nest. Vacant nests of conspecific females, but not those of another sympatric salticid species, elicit courtship behaviour from males. During male-male interactions, embracing occurs with the large chelicerae spread apart. Females and subadults also display, and different displays occur in interactions depending on the sex/age classes of the spiders involved. Despite the unusual morphology of these spiders, their individual displays are similar to those of more typical salticids. During copulation males stand beside the female instead of over or on her as occurs with typical salticids.     

How cross-modality effects during intraspecific interactions of jumping spiders differ depending on whether a female-choice or mutual-choice mating system is adopted

An important prediction from game theory is that the value of a resource influences the level to which conflict escalates. Here we use jumping spiders (Salticidae) to consider this prediction in the context of species adopting different mating systems (‘female mate-choice’ and ‘mutual mate-choice’). Our experiments are designed for determining whether the odour of conspecific females, more than the odour of heterospecific females, primes males to escalate conflict with a potential same-sex rival and also whether the odour of conspecific males, more than the odour of heterospecific males, primes females to escalate conflict with a potential same-sex rival. Four species were studied: Evarcha culicivora, a species in which mutual mate-choice is pronounced, and Portia fimbriata, Portia africana, and Jacksonoides queenslandicus, more conventional salticids in which female mate-choice and male–male competition appear to be dominant. Our hypothesis is that, for all four species, there is strong competition between males for access to females and that, for E. culicivora, but not for the other three species, there is also strong competition between females for access to males. Our findings are consistent with this hypothesis, as we show that, although the odour of conspecific females primes escalation of vision-based male–male conflict for all four species, E. culicivora is the only species for which there is evidence of odour from conspecific males priming the escalation of female–female conflict.     

Predator–Prey Interactions Between Aggressive-Mimic Jumping Spiders (Salticidae) and Araneophagic Spitting Spiders (Scytodidae) from the Philippines

In Los Baños in the Philippines (Laguna, Luzon), Scytodes sp. indet. is a web-building spitting spider (Scytodidae) that preys primarily on jumping spiders (Salticidae) and Portia labiata is an aggressive-mimic jumping spider that preys especially frequently on Scytodes. Tactics by which three species of Portia (P. africana, P. fimbriata, and P. labiata) and, for Portia labiata, three disjunct populations (Sri Lanka and, in the Philippines, Sagada and Los Baños) capture this especially dangerous prey are compared. Local adaptation to Scytodes by the Los Baños P. labiata is discussed. The Los Baños P. labiata uniquely made consistent use of tactics (soft plucking with palps and signal–detour–leap sequences) that were apparently responsible for greater prey-capture success and more effective avoidance of being spat on. Inter- and intraspecific differences were evident despite having used Portia that were reared in the laboratory with no prioir experience with scytodids.     

Reliance on trial and error signal derivation by <Emphasis Type="Italic">Portia africana</Emphasis>, an araneophagic jumping spider from East Africa

All species from the jumping spider (Salticidae) genus Portia appear to be predators that specialize at preying on other spiders by invading webs and, through aggressive mimicry, gaining dynamic fine control over the resident spider’s behavior. From previous research, there is evidence that P. fimbriata, P. labiata and P. schultzi derive signals by trial and error. Here, we demonstrate that P. africana is another species that uses a trial and error, or generate and test, algorithm when deriving the aggressive-mimicry signals that will be appropriate in different predator–prey encounters. We discuss the implications of these new findings and the findings from previous work in order to understand the selection factors that drive the evolution of flexibility in aggressive-mimicry strategies.     

The natural history of Myrmarachne melanotarsa,a social ant‐mimicking jumping spider

Abstract

Myrmarachne melanotarsa, an ant‐like jumping spider (Salticidae) from East Africa, is an accurate mimic of Crematogaster sp. and associates unusually closely with its models. M. melanotarsa is remarkable in that it forms dense aggregations and builds large nest complexes (numerous individually‐occupied nests connected to each other by silk). Other salticids (Pseudicius spp., Menemerus spp.) live with M. melanotarsa in the same nest complex. These aggregations, which can exceed 50 conspecific individuals per colony, are considerably larger than those few previously described, and seem to have primarily a protective function. We provide baseline information on the natural history of M. melanotarsa, paying particular attention to predatory behaviour and association with Crematogaster sp., and fit this within current theory on the function of sociality in spiders. Other unusual behaviour of M. melanotarsa includes “mouthing”, in which the spider opens and closes its chelicerae while pressing its mouthparts against nest silk. We investigated the role of prior presence of Crematogaster sp. on nest silk in eliciting this previously unreported behaviour.     

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.     

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.     

A specialized araneophagic predator's short‐term nutrient utilization depends on the macronutrient content of prey rather than on prey taxonomic affiliation

A specialist predator that has a specialized diet, prey‐specific prey‐capture behaviour and a preference for a particular type of prey may or may not be specialized metabolically. Previous studies have shown that jumping spiders of the genus Portia prey on other spiders using prey‐specific prey‐capture behaviour, prefer spiders as prey to insects and gain long‐term benefits in terms of higher survival and growth rates on spider diets than on insect diets. However, it is unclear whether there are substances uniquely present in spiders on which Portia depends, or, alternatively, spiders and insects all contain more or less the same nutrients but the relative amounts of these substances are such that Portia perform better on a spider diet. These questions are addressed by testing the hypothesis that prey specialization includes metabolic adaptations that allow Portia an enhanced nutrient extraction or nutrient utilization efficiency when feeding on spider prey compared with insect prey. Three groups of Portia quei Zabka are fed either their preferred spider prey or one of two types of flies (Drosophila melanogaster Meigen) that differ in nitrogen and lipid content. Portia quei shows a higher feeding rate of high‐protein flies than of high‐lipid flies and spiders but, after 5 days of feeding, there is no significant difference in growth between treatments, and the diets lead to significant changes in the macronutrient composition of P. quei as a result of variable extraction and utilization of the prey. The short‐term utilization of spider prey is similar to that of high‐lipid flies and both differ in several respects from the utilization of high‐protein flies. Thus, the short‐term nutrient utilization is better explained by prey macronutrient content than by whether the prey is a spider or not. The results suggest that spider prey may have a more optimal macronutrient composition for P. quei and that P. quei does not depend on spider‐specific substances.     

The Influence of Pedipalp Autotomy on the Courtship and Mating Behavior of Pardosa milvina (Araneae: Lycosidae)

Male Pardosa milvina wolf spiders use their pedipalps both for copulation and courtship. Pedipalp loss is significantly more common among adult males compared to females. We measured the courtship and mating effects associated with the loss of one or both pedipalps among adult male P. milvina. Pedipalp loss significantly reduced courtship intensity, but had no influence on mounting success. Intact males were less likely to be cannibalized and suffered fewer predatory attacks by females than autotomized males. Loss of the left pedipalp resulted in significantly less intense courtship, higher female aggression levels, and delayed onset of courtship whereas loss of the right pedipalp minimally affected male and female behavior relative to intact males. Pedipalp displays may function in reducing female aggression rather than increasing female receptivity.     

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.