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.     

Prey-capture techniques and prey preferences of Corythalia canosa and Pystira orbiculata, ant-eating jumping spiders (Araneae, Salticidae)

Corythalia canosa from Florida is an unusual salticid because it is known to eat ants. This species' specialized behaviour for catching ants is described in detail for the first time and compared to its behaviour for catching other insects. Pystira orbiculata from Queensland is shown to be another ant-eating salticid, although its behaviour for catching ants seems less specialized than that of C. canosa. Three different types of tests of prey preference were carried out. In each type of test C. canosa and P. orbiculata took ants in preference to other insects. Another species of salticid, Trite planiceps from New Zealand, failed to eat ants in these tests, although T. planiceps often attacked then released the ants. Corythalia canosa's and P. orbiculata's preference for ants, and their prey-specific predatory behaviour for catching ants, are shown not to depend on prior experience with ants.     

Conditional use of honest signaling by a Batesian mimic

Jumping spiders (Salticidae) usually avoid ants, but some specieswithin this family single out ants as preferred prey, whileothers (especially the species in the genus Myrmarachne) areBatesian mimics of ants. Field records show that ant-eatingsalticids sometimes prey on Myrmarachne, suggesting that theunwanted attention of predators that specialize on the modelmay be an important, but poorly understood, cost of Batesianmimicry. By staging encounters in the laboratory between livingant-eating salticids and Myrmarachne, we determined that ant-eatingsalticids attack Myrmarachne. However, when Myrmarachne detectsa stalking ant-eating salticid early enough, it adopts a distinctivedisplay posture (legs almost fully extended, elevated 45°,and held out to the side 45°), and this usually deters thepredator. When Myrmarachne detects an ant-eating salticid beforestalking begins, Myrmarachne makes preemptive displays thatappear to inhibit the initiation of stalking. Using immobilelures made from dead Myrmarachne that were either in a displayposture or a nondisplay posture, we ascertained that specificallythe display posture of Myrmarachne deters the initiation ofstalking (ant-eating salticids stalked nondisplaying more oftenthan displaying lures). In another experiment, we ascertainedthat it is specifically the interjection of display posturethat deters stalking. When ant-eating salticids that had alreadybegun stalking experienced lures that switched from a nondisplayto a display posture, they stopped stalking. Although the unwantedattentions of its models' predators may be, for Myrmarachne,a hidden cost of Batesian mimicry, Myrmarachne appears to havean effective defense against these predators.     

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.     

Bugs with backpacks deter vision-guided predation by jumping spiders

As a case study of how insects use masks as a defence against vision-guided predators, an experimental study was carried out using Acanthaspis petax , a reduviid bug ('ant bug') that covers itself with a 'mask', or 'backpack', made from carcasses of its preferred prey (ants), and three salticid spider species, Hyllus sp., Plexippus sp. and Thyene sp., salticids being predators with exceptionally acute vision. The ant bugs and the salticids were from the Lake Victoria region of East Africa. In each test, a salticid was presented with a live bug or a lure made from a dead bug, with the mask removed ('naked') or intact ('masked'). Salticids made predatory responses to naked bugs significantly more often than to masked bugs. These findings suggest that salticids readily identify naked bugs as prey, but fail to identify masked bugs as prey.     

Prey classification by an araneophagic ant-like jumping spider (Araneae: Salticidae)

What to attack is one of the most basic decisions predators must make, and these decisions are reliant upon the predator's sensory and cognitive capacity. Active choice of spiders as preferred prey, or araneophagy, has evolved in several distantly related spider families, including jumping spiders (Salticidae), but has never been demonstrated in ant-like jumping spiders. We used prey-choice tests with motionless lures to investigate prey-choice behaviour in Myrmarachne melanotarsa , an East African ant-like salticid that normally lives in aggregations and often associates with other spider species. We show that M . melanotarsa chooses spiders as prey in preference to insects and, furthermore, discriminates between different types of spiders. Myrmarachne melanotarsa 's preferred prey were juvenile hersiliids and its second most preferred were other salticids. To date, all documented examples of araneophagic salticids have been from the basal subfamily Spartaeinae. Myrmarachne melanotarsa is the first non-spartaeine and also the first ant-like salticid for which araneophagy has been demonstrated.     

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.     

Aggressive use of Batesian mimicry by an ant-like jumping spider

Batesian and aggressive mimicry are united by deceit: Batesian mimics deceive predators and aggressive mimics deceive prey. This distinction is blurred by Myrmarachne melanotarsa, an ant-like jumping spider (Salticidae). Besides often preying on salticids, ants are well defended against most salticids that might target them as potential prey. Earlier studies have shown that salticids identify ants by their distinctive appearance and avoid them. They also avoid ant-like salticids from the genus Myrmarachne. Myrmarachne melanotarsa is an unusual species from this genus because it typically preys on the eggs and juveniles of ant-averse salticid species. The hypothesis considered here is that, for M. melanotarsa, the distinction between Batesian and aggressive mimicry is blurred. We tested this by placing female Menemerus sp. and their associated hatchling within visual range of M. melanotarsa, its model, and various non-ant-like arthropods. Menemerus is an ant-averse salticid species. When seeing ants or ant mimics, Menemerus females abandoned their broods more frequently than when seeing non-ant-like arthropods or in control tests (no arthropods visible), as predicted by our hypothesis that resembling ants functions as a predatory ploy.     

Innate aversion to ants (Hymenoptera: Formicidae) and ant mimics: experimental findings from mantises (Mantodea)

Field data suggest that ants may be important predators of mantises which, in turn, may be important predators of jumping spiders (Salticidae). Using a tropical fauna from the Philippines as a case study, the reactions of mantises to ants, myrmecomorphic salticids (i.e. jumping spiders that resemble ants) and ordinary salticids (i.e. jumping spiders that do not resemble ants) were investigated in the laboratory. Three mantis species ( Loxomantis sp., Orthodera sp., and Statilia sp.) were tested with ten ant species, five species of Myrmarachne (i.e. myrmecomorphic salticids), and 23 ordinary salticid species. Two categories of the myrmecomorphic salticids were recognized: (1) 'typical Myrmarachne ' (four species with a strong resemblance to ants) and (2) Myrmarachne bakeri (a species with less strong resemblance to ants). Ants readily killed mantises in the laboratory, confirming that, for the mantises studied, ants are dangerous. In alternate-day testing, the mantises routinely preyed on the ordinary salticids, but avoided ants. The mantises reacted to myrmecomorphic salticids similarly to how they reacted to ants (i.e. myrmecomorphic salticids appear to be, for mantises, Batesian mimics of ants). Although myrmecomorphic salticids were rarely eaten, M . bakeri was eaten more often than typical Myrmarachne . Because the mantises had no prior experience with ants, ant mimics or ordinary salticids, our findings suggest that mantises have an innate aversion to attacking ants and that this aversion is generalized to myrmecomorphic salticids even in the absence of prior experience with ants. © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 23–32.     

Long-duration whirling of Pholcus phalangioides (Araneae, Pholcidae) is specifically elicited by Salticid spiders

Long-duration whirling (gyrating of the body during several hours a day) was shown by the pholcid spider Pholcus phalangioides to salticid spiders and hardly ever to predatory spiders from eight other families in laboratory arenas. Long-duration whirling has not been reported so far, in contrast to short-duration whirling lasting less than a few minutes. Long-duration whirling may have the anti-predatory function of disturbing continual visual fixation of prey in attacking salticids, in contrast to short-duration whirling that has been demonstrated to favour survival of pholcids in the presence of all sorts of predatory spiders.     

The meaning of predatory specialization as illustrated by Aelurillus m-nigrum, an ant-eating jumping spider (Araneae: Salticidae) from Azerbaijan

The distinctions between a predator's diet, its prey-choice behaviour and its preference are illustrated in a study of Aelurillus m-nigrum Kulczyn'ski, a salticid spider from Azerbaijan. The natural diet of A. m-nigrum was determined from records of individuals feeding in the field (N=58). Ten arthropod orders were represented. Nine were from the class Insecta (Coleoptera, Collembola, Diptera, Heteroptera, Homoptera, Hymenoptera, Lepidoptera, Orthoptera, Psocoptera) and one from the class Arachnida (Araneae). Of 50 insects among the prey, 21 (42%) were Hymenoptera, with ants (family Formicidae) alone accounting for 31% of all prey records. Although the majority (69%) of the natural prey were not ants, results from prey-choice testing in the laboratory implied that A. m-nigrum preferred ants as prey. However, this preference was evident only when the testing environment included sand and a small stone. Our findings illustrate the importance of not conflating the concept of a predator's preference with the concept of a predator's natural diet and illustrate that physical features of a predator's habitat may be an important factor in influencing how strongly preference is expressed.     

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.     

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.     

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.     

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 Pholcus phalangioides (Araneae, Pholcidae): predatory versatility, araneophagy and aggressive mimicry

Predatory versatility occurs in Pholcus phalangioides (Fuesslin). In addition to building prey-catching space webs, P. phalangioides invades webs of other spiders and feeds on the occupants. It acts as an aggressive mimic by performing specialized vibratory behaviours to which the prey-spider responds as it normally would to its own prey. Prey (spiders and insects) is attacked by wrapping. Prey that trips over lines at the edge of a web of P. phalangioides , but fails to enter the web, is successfully attacked: P. phalangioides leans out of its web to throw silk over the prey, keeping as few as two legs on the silk. However, P. phalangioides does not attack prey that is completely away from webs. Occasionally, P. phalangioides feeds on eggs of other spiders and on ensnared insects it encounters in alien webs. Experimental evidence indicates that vision is of little or no importance in the predatory behaviour of P. phalangioides . Although P. phalangioides invades diverse types of webs, in addition to using its own web, its efficiency as a predator varies with web-type. It is most efficient as a predator of spiders and, especially, insects on its own web, and least efficient as a predator of amaurobiids on their cribellate sheet webs. Sensory, locomotory and other factors which influence differential predatory efficiency are discussed. The behaviour of P. phalangioides is compared to that of Portia , an araneophagic web-invading salticid, and the results of this study are discussed in relation to hypotheses concerning salticid evolution.     

Vision-based innate aversion to ants and ant mimics

Innate vision-based aversions to model and mimic were investigatedusing a mimicry system in which the models were ants (Formicidae),and both the mimics and the predators were jumping spiders (Salticidae).Jumping spiders are a large group of predatory invertebratesthat usually prey opportunistically on prey of similar size.We used 12 representative species from this group, the "ordinarysalticids" as predators. The mimics considered belonged to anothergroup, salticids that resemble ants. A choice arena containingan empty chamber and a stimulus chamber was used for testingpredator responses to a variety of dead arthropods (ants, antmimics, and an array of non–ant-like species) mountedin a lifelike posture. When presented with visual cues fromarthropods other than ants or ant-like salticids, naive predatorschose the empty chamber no more often than the stimulus chamber.However, when visual cues were from ants or from ant-like salticids,ordinary salticids chose the empty chamber significantly moreoften than the stimulus chamber. These findings suggest learningby the predator is not necessary in order for ant-like salticidsto gain Batesian mimicry advantages.     

Compound mimicry and trading predators by the males of sexually dimorphic Batesian mimics

Sexual dimorphism is pronounced in Myrmarachne, a large genus of ant-like jumping spiders (Salticidae) and one of the major animal groups in which Batesian mimicry of ants has evolved. Although adult females and juveniles of both sexes are distinctly ant-like in appearance, Myrmarachne males have elongated chelicerae that might appear to detract from their resemblance to ants. Experimental findings suggest that the Myrmarachne male's solution is to adopt compound mimicry (i.e. the male's model seems to be not simply an ant worker but a combination of an ant and something carried in the ant's mandibles: an "encumbered ant"). By becoming a mimic of a particular subset of worker ants, Myrmarachne males may have retained their Batesian-mimicry defence against ant-averse predators, but at the price of receiving the unwanted attention of predators for which encumbered ants are preferred prey. Two salticid species were used as predators in the experiments. Portia fimbriata is known to choose other salticids as preferred prey and to avoid unencumbered ants and their mimics (Myrmarachne females). In experiments reported here, P. fimbriata avoided encumbered ants and Myrmarachne males. Ants are the preferred prey of Chalcotropis gulosus. In our experiments, C. gulosus chose safer encumbered ants in preference to more dangerous unencumbered ants, chose Myrmarachne males more often than Myrmarachne females and showed no evidence of distinguishing between Myrmarachne males and encumbered ants. The cost of reconciling sexual dimorphism with Batesian mimicry appears to be that Myrmarachne males attract the unwanted attention of specialist predators of their compound model.     

Evarcha culicivora chooses blood-fed Anopheles mosquitoes but other East African jumping spiders do not

Previous research using computer animation and lures made from dead prey has demonstrated that the East African salticid Evarcha culicivora Wesolowska & Jackson (Araneae: Salticidae) feeds indirectly on vertebrate blood by actively choosing blood-carrying female mosquitoes as prey, and also that it singles out mosquitoes of the genus Anopheles (Diptera: Culicidae) by preference. Here, we demonstrate that E. culicivora's preference is expressed when the species is tested with living prey and that it is unique to E. culicivora. As an alternative hypothesis, we considered the possibility that the preference for blood-fed female anopheline mosquitoes might be widespread in East African salticids. When live-prey choice tests were carried out in 19 additional species, there were no instances in which blood-carrying mosquitoes were chosen significantly more often than other prey. Combined with the findings of previous work, these results suggest that it is possible that specialized predators play a role in the biological control of disease vectors.     

Salticid predation as one potential driving force of ant mimicry in jumping spiders

Many spiders possess myrmecomorphy, and species of the jumping spider genus Myrmarachne exhibit nearly perfect ant mimicry. Most salticids are diurnal predators with unusually high visual acuity that prey on various arthropods, including conspecifics. In this study, we tested whether predation pressure from large jumping spiders is one possible driving force of perfect ant mimicry in jumping spiders. The results showed that small non-ant-mimicking jumping spiders were readily treated as prey by large ones (no matter whether heterospecific or conspecific) and suffered high attack and mortality rates. The size difference between small and large jumping spiders significantly affected the outcomes of predatory interactions between them: the smaller the juvenile jumping spiders, the higher the predation risk from large ones. The attack and mortality rates of ant-mimicking jumping spiders were significantly lower than those of non-ant-mimicking jumping spiders, indicating that a resemblance to ants could provide protection against salticid predation. However, results of multivariate behavioural analyses showed that the responses of large jumping spiders to ants and ant-mimicking salticids differed significantly. Results of this study indicate that predation pressure from large jumping spiders might be one selection force driving the evolution of nearly perfect myrmecomorphy in spiders and other arthropods.