Anti-predator defences of Psilochorus sphaeroides and Smeringopus pallidus (Araneae, Pholcidae), tropical web-building spiders

Anti-predator defence behaviours of Psilochorus sphaeroides from Queensland and Smeringopus pallidus from Sri Lanka were studied in the laboratory. Whirling, in which these web-building pholcid spiders move their bodies rapidly around in circles, with legs remaining on the silk, is the most frequent response of adults to disturbance. Experiments were carried out to determine what types of stimuli elicited whirling. Forceful tactile and air movement stimuli elicited whirling, but there was no evidence that chemical stimuli from potential predators were important. Tactile stimulation was more effective than air movement. The behaviour of these pholcids was compared to that of a cosmopolitan pholcid, Pholcus phalangioides .     

Predator-prey interactions between web-invading jumping spiders and a web-building spider, Holocnemus pluchei (Araneae, Pholcidae)

Three species of web-invading salticid spiders, with different predatory strategies, were tested with Holocnemus pluchei in the laboratory: Portia fimbriata and Portia labiata , which practise aggressive mimicry, and Euyattus sp., which leaps from outside on to spiders in webs but does not practise aggressive mimicry. Portia was shown to be more efficient than Euryattus at catching H. pluchei. Portia fimbriata from Queensland was more consistent at using aggressive mimicry than was Portia labiata and was also more efficient at catching H. pluchei . The web-invaders that were more efficient at catching H. pluchei were also better able to avoid setting off bouncing, a special defence behaviour used by H. pluchei . An experiment, in which H. pluchei was artificially induced to bounce whenever the predator was near, provided additional evidence that bouncing is effective in defending H. pluchei against web-invaders.     

Predator-prey interactions between web-invading jumping spiders and Argiope appensa (Araneae, Araneidae), a tropical orb-weaving spider

A range of web-invading jumping spiders with different predatory strategies was tested with A. appensa in the laboratory: Mimetus maculosus (Mimetidae), Pholcus phalangioides (Pholcidae), Taieria erebus (Gnaphosidae), and 11 species of salticids. Spiders that are known to specialize at web-invading, either by leaping into webs or by walking slowly into webs and practising aggressive mimicry, captured A. appensa ; three salticid species not known to be web-invaders never did. Web-invaders that practised aggressive mimicry were more efficient than were species that only leapt into webs. Portia fimbriata from Queensland was the most consistent at using aggressive mimicry and was also the most efficient at catching A. appensa . Web-invaders that were more efficient at catching A. appensa were also better able to avoid setting off pumping, a special defence behaviour used by A. appensa . Portia fimbriata from Queensland was especially efficient at avoiding setting off pumping: P. fimbriata more consistently than other Portia made its final approach toward A. appensa by coming down from above the web on a dragline and making minimal contact with the web. An experiment, in which A. appensa was artificially induced to pump whenever the predator was near, provided additional evidence that pumping is effective in defending A. appensa against web-invaders.     

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.     

Predator-prey interactions between two amphibian species: effects of insecticide exposure

The presence of environmental contaminants may alter predator-prey interactions among aquatic species by altering activity levels of predators or prey, or by altering predator avoidance behavior. The outcome of a predatory encounter may be dependent upon whether both species are exposed to a contaminant simultaneously, or whether exposure occurs only in one of the species. In a laboratory experiment, I used the insecticide carbaryl to examine predation of southern leopard frog tadpoles (Rana sphenocephala) by adult red-spotted newts (Notophthalmus viridescens) under four conditions: both tadpoles and newts exposed, neither tadpoles nor newts exposed, and either newts or tadpoles only exposed. After one hour, exposed newts consumed half as many tadpoles as non-exposed newts. Carbaryl potentially affected newt activity enough to reduce time spent searching for prey, or may have altered the speed and coordination necessary to capture tadpoles. After six hours, non-exposed and exposed newts consumed similar numbers of tadpoles, most likely indicating recovery from exposure. After 24 h, predation rates were lowest when both newts and tadpoles were simultaneously either exposed or not exposed, and were greatest when newts and tadpoles were not exposed simultaneously. This study suggests that when tadpoles and newts are exposed to a sublethal level of a contaminant simultaneously, that predation rates do not differ from those observed under natural conditions, but exposure of either predator or prey at different times can disrupt predator-prey dynamics. This revised version was published online in August 2006 with corrections to the Cover Date.     

Cross-modality effects during male-male interactions of jumping spiders

An important prediction from game theory is that the value of a resource influences the level to which male-male conflict escalates. Earlier experimental studies have shown that the seven salticid species we study here (Bavia aericeps, Euryattus sp., Hypoblemum albovittatum, Jacksonoides queenslandicus, Marpissa marina, Portia africana and Simaetha paetula) determine by sight whether a female is a conspecific or a heterospecific and then escalate the intensity with which they interact (i.e., they adopt behaviour that is likely to put them at greater risk of injury after detecting the presence of a conspecific female). Here the earlier studies are extended by using the odour of conspecific females (experimental tests) and heterospecific females (control tests), and by presenting each male with his mirror image as well as having two males interact with each other. Findings from this study suggest that, for J. queenslandicus and P. africana, the odour of conspecific females, more than the odour of heterospecific females, primes the male to escalate conflict with a potential rival. However, this was not found for the other five species tested.     

Anti-predator defences of Pholcus phalangioides (Araneae, Pholcidae), a web-building and web-invading spider

The behaviours used by Pholcus phalangioides (Fuesslin) (Araneae, Pholcidae) to evade its predators were studied with particular attention being given to a special defence behaviour, whirling. To whirl, this long-legged web-building spider swings its body around in a circle, with its legs remaining on the silk. Experiments were carried out to determine the types of stimuli that elicited whirling. Touching the spider or its web elicited whirling, as did air movement over the spider, but there was no evidence that chemical stimuli from potential predators were important. Small juveniles differed from adult females and larger juveniles by more often dropping from the web instead of whirling when confronted by a potential predator. Besides catching prey on its own web P. phalangioides invades other spiders' webs to catch the other spiders. By whirling in alien webs, P. phalangioides could deter attacks by the resident spider, but P. phalangioides was less inclined to whirl when in an alien than when in its own web.     

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.     

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.     

Synergy between two invasive species,redback spiders and rabbits,threaten the endangered Cromwell chafer beetle

Interactions between invasive species can be difficult to predict and can result in unanticipated impacts of significance for native fauna. Here we show that introduced European rabbits (Oryctolagus cuniculus) create habitat that enables invasive redback spiders (Latrodectus hasselti Thorell, 1870) to establish and prey upon the nationally endangered, endemic Cromwell chafer beetle (Prodontria lewisii Broun, 1904). We examined the spatial relationship between rabbit holes and redback spider occurrence, recorded all prey caught in redback spider webs over a 4-month period, and tested the role rabbit holes play in providing habitat for redback spiders experimentally, by filling in rabbit holes in areas used by spiders and monitoring subsequent occupation of the areas over four months. Redback spiders predominately resided in old rabbit holes, with the highest densities of spiders coinciding with high densities of rabbit holes. Cromwell chafer beetles were commonly caught in webs. Filling in rabbit holes eliminated the presence of redback spiders at all treated sites and reduced prey capture in those areas. Conservation management to protect Cromwell chafer beetles should focus on eliminating rabbits and their holes from beetle habitat.     

Karyotype, sex determination, and meiotic chromosome behavior in two pholcid (Araneomorphae, Pholcidae) spiders: implications for karyotype evolution

There are 1,111 species of pholcid spiders, of which less than 2% have published karyotypes. Our aim in this study was to determine the karyotypes and sex determination mechanisms of two species of pholcids: Physocyclus mexicanus (Banks, 1898) and Holocnemus pluchei (Scopoli, 1763), and to observe sex chromosome behavior during meiosis. We constructed karyotypes for P. mexicanus and H. pluchei using information from both living and fixed cells. We found that P. mexicanus has a chromosome number of 2n = 15 in males and 2n = 16 in females with X0-XX sex determination, like other members of the genus Physocyclus. H. pluchei has a chromosome number of 2n = 28 in males and 2n = 28 in females with XY-XX sex determination, which is substantially different from its closest relatives. These data contribute to our knowledge of the evolution of this large and geographically ubiquitous family, and are the first evidence of XY-XX sex determination in pholcids.     

Molecular taxonomy in pholcid spiders (Pholcidae, Araneae): evaluation of species identification methods using CO1 and 16S rRNA

The identification of species using molecular characters is a promising approach in alpha taxonomy and in any discipline depending on reliable assignment of specimens. Previous studies have shown the feasibility of the method, but considerable controversy persists. In this study, we use pholcid spiders in an effort to address two main issues. First, we evaluate and calibrate molecular species (re-)identification within a closely related group of organisms by using specimens that are morphologically unambiguously either conspecific or not. Species limits hypothesized a priori based on morphology were almost universally reconstructed by both mitochondrial markers used. Second, we focus on species identification methodology in a morphology-calibrated scenario, i.e. on how to assess the quality of a dataset and of the method used to obtain distance estimates (e.g. choice of markers, alignment strategy, type of distance data). We develop a number of statistical estimators permitting the measurement and communication of the clarity of species boundaries in a dataset and discuss their benefits and drawbacks. We propose that box plots rather than histograms are the superior tool for graphically illustrating taxonomic signal and that the median is a more appropriate measure of central tendency than the mean. Applying the suggested tools to our data, we propose that in molecular species identification, indel-related alignment uncertainties may often be even advantageous (by accentuating taxonomy-relevant information) and we conclude that — at least for our dataset — 16S is better suited to taxonomy than CO1.     

Small-scale spatial pattern of web-building spiders (Araneae) in alfalfa: relationship to disturbance from cutting, prey availability, and intraguild interactions

Understanding the development of spatial patterns in generalist predators will improve our ability to incorporate them into biological control programs. We studied the small-scale spatial patterns of spider webs in alfalfa by analyzing the relationship between web locations over distances ranging from 4 to 66 cm. Using a coordinate-based spatial statistic (O-ring) and assuming a heterogeneous distribution of suitable web sites, we analyzed the impact of cutting and changes in spider abundance on web distribution. We analyzed the influence of small-scale variation in prey availability by comparing web distributions to the pattern of sticky-trap captures of Aphididae and Diptera described by a count-based spatial statistic (SADIE). Cutting of alfalfa reduced the overall density of web-building spiders but had no immediate impact on the spatial distribution of their webs. Availability of aphids was highest before the alfalfa was cut and was clumped at a scale of 66 cm. Spider webs, however, were not clumped at any scale or date. In contrast, webs were regularly distributed at smaller distances (<20 cm) immediately before and after cutting. Because cursorial and web-building spiders were most active during this period, we hypothesize that the development of small-scale regularity in web locations was driven by intraguild interactions. Our results suggest that intraguild interactions contribute to the development of small-scale spatial patterns of spider webs in alfalfa. Variation in prey availability may have more of an influence on web distribution in crops with a different vegetation structure or if patterns are studied at larger spatial scales.     

Reproductive interactions between two Australian reptile tick species

In South Australia the two tick species Amblyomma limbatum and Aponomma hydrosauri share the same common reptile host species, but have allopatric distributions which abut along a narrow parapatric boundary. Reproductive interference is a mechanism that has previously been suggested could contribute to maintaining the boundary. Populations of each species were established in pens within the range of Aponomma hydrosauri. Pens held either each species alone, or the two species together. The performance of females in those pens was monitored over 28 months. There was no indication that the proportion of attached females which mated and engorged was reduced by the presence of heterospecifics. There was no indication that the time taken to mate, engorge and detach was any longer in the presence of heterospecifics. The experiment did not support the hypothesis that reproductive interference contributes to maintaining the parapatric boundary. However, Amblyomma limbatum in the pens had a shorter season of reproductive activity, and achieved much smaller numbers of reproductive females. This may inhibit successful colonization of cooler habitats to the south of its distribution.     

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.     

Contrasting responses of web-building spiders to deer browsing among habitats and feeding guilds

We examined web-building spider species richness and abundance in forests across a deer density gradient to determine the effects of sika deer browsing on spiders among habitats and feeding guilds. Deer decreased the abundance of web-building spiders in understory vegetation but increased their abundance in the litter layer. Deer seemed to affect web-building spiders in the understory vegetation by reducing the number of sites for webs because vegetation complexity was positively correlated with spider density and negatively correlated with deer density. In contrast, the presence of vegetation just above the litter layer decreased the spider density, and deer exerted a negative effect on this vegetation, possibly resulting in an indirect positive effect on spider density. The vegetation just above the litter layer may be unsuitable as a scaffold for building webs if it is too flexible to serve as a reliable web support, and may even hinder spiders from building webs on litter. Alternatively, the negative effect of this vegetation on spiders in the litter may be as a result of reduced local prey availability under the leaves because of the reduced accessibility of aerial insects. The response to deer browsing on web-building spiders that inhabit the understory vegetation varied with feeding guild. Deer tended to affect web-invading spiders, which inhabit the webs of other spiders and steal prey, more heavily than other web-building spiders, probably because of the accumulated effects of habitat fragmentation through the trophic levels. Thus, the treatment of a particular higher-order taxon as a homogeneous group could result in misleading conclusions about the effects of mammalian herbivores.     

Interference interactions and nest usurpation between two subordinate ant species

Camponotus foreli (Emery) and Cataglyphis iberica (Emery) are two sympatric, subordinate ant species that have been found to fight in attacks that usually conclude with the death of many workers of both species and with nest abandonment by C. iberica. These harassment episodes have been observed in two different areas and over many years of study. No such attacks of C. foreli were observed in the study areas against any other ant species, nor did any other ants attack C. iberica nests, and laboratory confrontations confirmed this specificity. These attacks neither eliminated C. iberica colonies, nor distanced them from C. foreli nests. Moreover, there was no real competition for food between the species: in an experiment where all C. iberica colonies were eliminated from an area, rates of prey and liquid food collection by C. foreli nests in the exclusion zone were similar to those found in the control zone with C. iberica, and the activity rhythms of C. foreli did not change in the absence of C. iberica. The hypothesis of competition for a nest site is more consistent. Both in the laboratory and the field, the most frequent outcome of these aggressive interactions was the occupation of the C. iberica nest by C. foreli. This behavior may be advantageous for C. foreli, because it is much less skilful at excavating than C. iberica. One of the chief concerns of this study is to show that such interference interactions, typical especially of dominant, very aggressive species, are also found between subordinate, apparently nonaggressive species. Received: 20 March 1997 / Accepted: 29 September 1997