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 .     

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.     

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.     

Cues for web invasion and aggressive mimicry signalling in Portia (Araneae, Salticidae)

Portia is a web-invading araneophagic spider that uses aggressive mimicry to deceive its prey. The present paper is a first step toward clarifying experimentally the cues that govern Portia's decisions of whether to enter a web, whether to make signals once in a web, and whether to persist at signalling once started. The following conclusions are supported: cues from seeing a web elicit web entry, but volatile chemical cues from webs of prey spiders are not important; seeing a spider in a web increases Portia's inclination to enter the web; after web entry, cues from webs of prey spiders are sufficient to elicit signalling behaviour, even in the absence of other cues coming directly from the prey spider; seeing a prey spider or detecting vibrations on the web make Portia more prone to signal, but volatile chemical cues from prey spiders are not important; once Portia is on a web and signalling, seeing a moving spider and detecting vibrations on the web encourage Portia to persist in signalling; on the basis of visual cues alone, Portia can distinguish between quiescent spiders, insects and eggsacs.     

Anti-predator defences of a web-building spider, Holocnemus pluchei (Araneae, Pholcidae)

Anti-predator defence behaviour of Holocnemus pluchei (Araneae, Pholcidae) was studied in the laboratory and the field. Bouncing, in which this web-building spider moves its body rapidly up and down while keeping its legs on the silk, was the most frequent response to disturbance. Less often, H. pluchei left the web when disturbed. Experiments were carried out to determine the types of stimuli that elicited bouncing and leaving the web: tactile stimulation (striking the spider with a brush) and air movement (blowing on the spider) elicited bouncing and leaving the web, but there was no evidence that chemical stimuli from potential predators were important. Tactile stimulation elicited stronger responses than air movement. There were no evident differences in how H. pluchei responded to tactile stimulation in the laboratory and the field.     

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.     

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.     

Cobweb management and control of the spider Holocnemus pluchei (Araneae: Pholcidae) on buildings

The spider Holocnemus pluchei (Scopoli) (Araneae: Pholcidae) is a nonnative species commonly found in the southwestern United States. In urban areas around homes and other structures, it is often the dominant spider species on outside surfaces; requests for control of spiders and their cobwebs are common for the commercial pest management industry. We tested two physical (brushing and vacuuming) and two low-impact chemical (permethrin and botanical insecticide) spray control methods for H. pluchei on outbuildings on the University of California-Riverside campus. Forty sites containing at least two H. pluchei spiders along a 5-m section of eave or overhang were chosen for treatment. Treatment was applied within 2 d of a pretreatment census. The number of spiders per section was counted at 2, 4, 8, 12, 16, and 52 wk posttreatment. Untreated sections showed an increase in spiders through summer and a decrease in autumn. Sites treated with permethrin significantly reduced spiders >95% and had strong residual effect throughout the remainder of the sampling period. Compared with untreated controls, the three other treatments did not significantly reduce spiders; these three were similar throughout the season and were intermediate between untreated and permethrin-treated sites. In comparison with the before and 2-wk posttreatments, spiders at the untreated sites showed no difference in body sizes, whereas those in the nonpermethrin treatments showed a decrease, indicating that larger spiders were eliminated. We also tested H. pluchei spiders in the laboratory against the two insecticides used in the field studies to determine the minimum lethal concentrations for controlling H. pluchei.     

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.     

Hatching responses of subsocial spitting spiders to predation risk

The carrying of eggs often renders parents vulnerable to predators due to increased conspicuousness or decreased mobility. Nonetheless, egg-carrying parents can escape from the predators to which they are vulnerable. Previous studies have demonstrated heavy predation by spider-eating jumping spiders (Portia labiata) on egg-carrying spitting spider (Scytodes pallida) females, but little predation on eggless females. If the timing of hatching is phenotypically plastic, then both S. pallida females and their eggs could reduce the risk of predation by hatching early. Hence, this study examines the hatching responses of S. pallida to chemical cues from P. labiata, both in the laboratory and in the field, and addresses the following questions. (i) Do cues from predatory P. labiata influence the hatching traits of S. pallida? (ii) Are the olfactory cues from predators sufficient for predator detection by S. pallida ? (iii) Are hatching responses to predatory P. labiata controlled by egg-carrying S. pallida females, or directly by their embryos? The study provides evidence of hatching as a life-history switch point, which shows an adaptive plasticity in response to predation risk in egg-carrying S. pallida. Egg-carrying S. pallida females, but not unattended eggs, adjust egg-hatching time (the interval between oviposition and hatching) in response to the threat of predation on both the female and her eggs by P. labiata. In the presence of P. labiata, eggs that are carried by females hatch sooner; the hatchlings of these eggs are therefore smaller than hatchlings born in the absence of P. labiata. Chemical cues that are released from the draglines of P. labiata are sufficient to elicit changes in the egg-hatching traits of S. pallida. Hatching early in response to this predator may benefit both females and their offspring. To my knowledge, this is the first direct experimental study to demonstrate predator-induced hatching plasticity in spiders and, in particular, in animals with parental care.     

Signals and Signal Choices made by the Araneophagic Jumping Spider Portia fimbriata while Hunting the Orb-Weaving Web spiders Zygiella x-notata and Zosis geniculatus

Portia fimbriata is a web-invading araneophagic jumping spider (Salticidae). The use of signal-generating behaviours is characteristic of how P. fimbriata captures its prey, with three basic categories of signal-generating behaviours being prevalent when the prey spider is in an orb web. The predatory behaviour of P. fimbriata has been referred to as aggressive mimicry, but no previous studies have provided details concerning the characteristics of P. fimbriata 's signals. We attempt to determine the model signals for P. fimbriata 's 'aggressive mimicry' signals. Using laser Doppler vibrometer and the orb webs of Zygiella x-notata and Zosis geniculatus , P. fimbriata 's signals are compared with signals from other sources. Each of P. fimbriata 's three categories of behaviour makes a signal that resembles one of three signals from other sources: prey of the web spider (insects) ensnared in the capture zone of the web, prey making faint contact with the periphery of the web and large-scale disturbance of the web (jarring the spider's cage). Experimental evidence from testing P. fimbriata with two sizes of lure made from Zosis (dead, mounted in a lifelike posture in standard-size orb web) clarifies P. fimbriata 's signal-use strategy: (1) when the resident spider is small, begin by simulating signals from an insect ensnared in the capture zone (attempt to lure in the resident spider); (2) when the resident spider is large, start by simulating signals from an insect brushing against the periphery of the web (keep the resident spider out in the web, but avoid provoking from it a full-scale predatory attack); (3) when walking in the resident spider's web, regardless of the resident spider's size, step toward the spider while making a signal that simulates a large-scale disturbance of the web (mask footsteps with a self-made vibratory smokescreen).     

Convergent evolution of eye ultrastructure and divergent evolution of vision-mediated predatory behaviour in jumping spiders

All jumping spiders have unique, complex eyes with exceptional spatial acuity and some of the most elaborate vision-guided predatory strategies ever documented for any animal of their size. However, it is only recently that phylogenetic techniques have been used to reconstruct the relationships and key evolutionary events within the Salticidae. Here, we used data for 35 species and six genes (4.8 kb) for reconstructing the phylogenetic relationships between Spartaeinae, Lyssomaninae and Salticoida. We document a remarkable case of morphological convergence of eye ultrastructure in two clades with divergent predatory behaviour. We, furthermore, find evidence for a stepwise, gradual evolution of a complex predatory strategy. Divergent predatory behaviour ranges from cursorial hunting to building prey-catching webs and araneophagy with web invasion and aggressive mimicry. Web invasion and aggressive mimicry evolved once from an ancestral spartaeine that was already araneophagic and had no difficulty entering webs due to glue immunity. Web invasion and aggressive mimicry was lost once, in Paracyrba, which has replaced one highly specialized predation strategy with another (hunting mosquitoes). In contrast to the evolution of divergent behaviour, eyes with similarly high spatial acuity and ultrastructural design evolved convergently in the Salticoida and in Portia.     

Masquerading predators deceive prey by aggressively mimicking bird droppings in a crab spider

In aggressive mimicry, a predator accesses prey by mimicking the appearance and/or behavior of a harmless or beneficial model in order to avoid being correctly identified by its prey. The crab spider genus Phrynarachne is often cited as a textbook example of masquerading as bird droppings (BDs) in order to avoid predation. However, Phrynarachne spiders may also aggressively mimic BDs in order to deceive potential prey. To date, there is no experimental evidence to support aggressive mimicry in masquerading crab spiders; therefore, we performed a field survey, a manipulative field experiment, and visual modeling to test this hypothesis using Phrynarachne ceylonica. We compared prey-attraction rates among BDs, spiders, and control empty leaves in the field. We found that although all prey combined and agromyzid dipterans, in particular, were attracted to BDs at a higher rate than to spiders, other dipterans and hymenopterans were attracted to BDs at a similar rate as to spiders. Both spiders and BDs attracted insects at a significantly higher rate than did control leaves. As predicted, prey was attracted to experimentally blackened or whitened spiders significantly less frequently than to unmanipulated spiders. Finally, visual modeling suggested that spiders and BDs can be detected by dipterans and hymenopterans against background leaves, but they are indistinguishable from each other. Taken together, our results suggest that insects lured by spiders may misidentify them as BDs, and bird-dropping masquerading may serve as aggressive mimicry in addition to predator avoidance in P. ceylonica.     

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.     

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 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.     

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.     

Cues by which suspended-leaf nests of Euryattus (Araneae: Salticidae) females are recognized by conspecific males and by an aggressive-mimic salticid, Portia fimbriata

Females of Euryattus sp. indet, a salticid from Queensland, suspend rolled-up leaves for nests. Euryattus males respond to conspecific females in nests with vibratory courtship. Portia fimbriata , a sympatric salticid that preys on Euryattus , responds to Euryattus females' nests by mimicking the courtship of Euryattus males. In the laboratory, cues by which potential mates (conspecific males) and potential predators ( P. fimbriata females) recognize the nests of Euryattus females are investigated experimentally. Both the position of the leaf in relation to the substratum and the number of threads between the leaf and the substratum appear to be important cues by which both the Euryattus males and P. fimbriata females find nests. In addition, chemical cues from Euryattus females influence conspecific males, but there is no evidence that chemical cues are important to the predator.     

Draglines and Assessment of Fighting Ability in Cannibalistic Jumping Spiders

The frequency of injury and death during female-female aggression varies in the jumping spider genus Portia, with interactions being more violent (likely to end in death or injury of one of the combatants) in P. labiata (from Sri Lanka) than in another two species (P. fimbriata from Australia and P. schultzi from Kenya). To investigate the role of draglines in the assessment of fighting ability, two types of tests were carried out: 1) dragline discrimination and 2) mirror image response (Portia's reaction to mirror images is comparable to interaction with conspecific rivals). For both types of testing, triplets of equal-size conspecific females were used: one female (the test spider) was exposed to draglines of two equal-size conspecific females they had not encountered before (donor spiders). The fighting abilities of donor spiders were determined directly by staging intraspecific contests between them. In dragline-discrimination tests (spider placed in petri dish containing draglines from two conspecific females with different fighting ability), females of P. labiata, but not the other two species, avoided draglines of the superior fighter (i.e., they spent the majority of their time on draglines of donor spiders with lesser fighting ability). For mirror-image testing, the test spider was placed in a petri dish containing a mirror and draglines. Each test spider was tested on two successive days, with donor draglines in the two tests coming from conspecific females with different fighting ability. In these tests, females of P. labiata (but not the other two species) spent less time embracing (each spider pressing its forelegs, palps and front of body against the other spider) and more time in a part of the petri dish where view of the mirror was obstructed when on the draglines of donor spiders with greater fighting ability than when on the other conspecific's draglines. Findings from this study suggest that P. labiata females use signpost cues associated with draglines to assess the relative fighting abilities of unknown opponents.