Multiple Roles of Tail Display by the Curly-Tailed Lizard Leiocephalus carinatus: Pursuit Deterrent and Deflective Roles of a Social Signal

The same display may be used in different contexts to convey different messages, or may have other, non-signaling functions. Several lines of evidence suggest that vertical tail curling, a previously documented social display in the lizard Leiocephalus carinatus , has antipredatory functions that may include pursuit deterrence and deflection of attacks from the body to the tail, which can be autotomized. An antipredatory role of tail curling is suggested by its more frequent occurrence when a predator is approaching than moving away, its greater frequency and intensity when a lizard is approached by a predator than when it moves spontaneously, and its greater frequency when the predator approaches directly rather than on a path bypassing the lizard. Evidence is presented contradicting use of tail curling for flash concealment or as an alarm signal to conspecifics. A pursuit-deterrent function of tail curling is suggested by its (1) more frequent use by lizards close to a refuge than those further from a refuge, (2) greater frequency during direct approaches by predators, and (3) much greater frequency when a predator is far enough away for pursuit to be deterred than when the predator is close enough to pose a high risk of capture. Lizards fled into a refuge without tail curling when the predator was very close, but often stopped outside a refuge while displaying the curled tail when the predator was farther away. Tail curling also may deflect attacks to the autotomizable tail, as suggested by its occurrence during spontaneous movements when no predator is approaching and by the high frequency of completely uncurled tails among individuals under bushes. The role of the tail in autotomy may facilitate evolution of pursuit-deterrent signals involving the tail.     

Effect of sex and bright coloration on survival and predator‐induced wing damage in an aposematic lantern fly with startle display

1. Aposematic coloration in prey promotes its survival by conspicuously advertising unpalatability to predators. Although classical examples of aposematic signals involve constant presentation of a signal at a distance, some animals suddenly display warning colours only when they are attacked. 2. Characteristics of body parts suddenly displayed, such as conspicuous coloration or eyespot pattern, may increase the survival of the prey by startling the predator, and/or by signalling unpalatability to the predators at the moment of attack. 3. The adaptive value of such colour patterns suddenly displayed by unpalatable prey has not been studied. We experimentally blackened the red patch in the conspicuous red–white–black hindwing pattern displayed by an unpalatable insect Lycorma delicatula White (Hemiptera: Fulgoridae) in response to predator's attack. 4. There was no evidence that the presence of the red patch increased prey survival over several weeks. We hypothesise that predators generalised from the red–white–black patches on the hindwings of unpalatable L. delicatula to any similar wing display as a signal of unpalatability. Because a higher proportion of males than females stay put at their resting sites, displaying their wings in response to repeated attacks by predators, wing damage was more frequent in males than in females. 5. To our knowledge, this is the first experimental test of an adaptive role of aposematic signals presented by unpalatable prey during sudden displays triggered by direct predatory attack.     

Sexual Differences in the Behavioral Response of Túngara Frogs, Physalaemus pustulosus, to Cues Associated with Increased Predation Risk

Engaging in mating behaviors usually increases exposure to predators for both males and females. Anti‐predator strategies during reproduction may have important fitness consequences for prey. Previous studies have shown that individuals of several species adjust their reproductive behavior according to their assessment of predation risk, but few studies have explored potential sexual differences in these strategies. In this study, we investigate whether the acoustic cues associated with predatory attacks or those associated with predators themselves affect the mating behavior of female and male túngara frogs, Physalaemus pustulosus. We compared the responses of females approaching a mate and those of calling males when exposed to mating calls associated with sounds representing increased hazard. When presented with mating calls that differed only in whether or not they were followed by a predation‐related sound, females preferentially approached the call without predation‐related sounds. In contrast to females, calling males showed greater vocal response to calls associated with increased risk than to a call by itself. We found significant differences in the responses of females and males to several sounds associated with increased hazard. Females behaved more cautiously than males, suggesting that the sexes balance the risk of predation and the cost of cautious mating strategies differently.     

Anti-Predator Signals as Advertisements: Evidence in White-Throated Magpie-Jays

Calls and displays elicited by predators usually function as alarms or to inform predators of their detection. However, predator encounters may afford some individuals the opportunity to demonstrate quality or signal their availability. Here, I report on a class of vocal signals produced in predator-elicited displays that share many characteristics with sexually selected song. White-throated magpie-jays ( Calocitta formosa ) display at low-threat predators while producing 'loud display calls' (LDCs). I use this term because the calls occur primarily in two display contexts (see below) though occasionally in other contexts as well. Such calls and displays are primarily produced by males, and also occur in one other context, at dawn. Playback experiments showed that despite being elicited by predators, males were more likely than females to respond to LDCs, and more likely to respond when their mate was fertile. Over 134 different call types were produced in over 200 displays by 34 males; the largest minimum repertoire size was 67. Presentations of taxidermic raptor mounts elicited some LDCs, but fewer calls and lower diversity than at dawn or in predator approach displays. The male bias and high diversity suggest that LDCs are an outcome of intersexual selection, while their elicitation by predators suggests an alarm function. I propose that male magpie-jays use predator encounters as opportunities to advertise their presence and availability as mates; they use LDCs as songs. Such a communication system seems to have been favored by the unusual social system of magpie-jays, in which female groups defend territories and males have little opportunity to defend resources for mate attraction, forcing them to advertise when females are paying the most attention, during predator encounters.     

Flight songs of Dusky Flycatchers: a response to bird‐hunting raptors?

ABSTRACT.   Flight-song displays usually include distinctive vocalizations and behaviors that are probably energetically expensive and also likely to attract the attention of aerial predators. We observed flight songs performed by Dusky Flycatchers ( Empidonax oberholseri ) in two breeding populations. During 15 yr at Tioga Pass, California, we observed the display twice and, during 9 yr at Steamboat Mountain, British Columbia, we observed 21 flight-song displays. These displays were generally observed later in the breeding season, suggesting that they were not used for courtship, mate attraction, or territory defense. However, for 83% of these displays, a raptor was observed to be present, usually either an American Kestrel ( Falco sparverius ) or a Sharp-shinned Hawk ( Accipiter striatus ). We hypothesize that these displays, like stotting or other pursuit-deterrent signals, may be directed at small raptors that represent a greater threat to Dusky Flycatchers and might deter attack by signaling awareness. However, because only males were observed performing flight-song displays, it is also possible that these displays serve to alert mates and offspring about the presence of an aerial predator and reduce their vulnerability to attack.     

Managing predators: The influence of kangaroo rat antipredator displays on sidewinder rattlesnake hunting behavior

Upon sensing predators in their vicinity, many prey species perform antipredator displays that are thought to provide information to the predator that deters it from attacking (predator‐deterrent signals). These displays can be complex, incorporating a variety of signaling elements as well as direct physical harassment of the predator. Although the display behaviors in these communication systems are often well characterized, evidence of the efficacy of these displays in deterring predators is limited due to the challenges associated with studying free‐ranging predators. Here, we examine how the anti‐snake signals of the desert kangaroo rat (Dipodomys deserti) influence the ambush hunting behaviors of sidewinder rattlesnakes (Crotalus cerastes). We found that, although desert kangaroo rats incorporate a number of signal elements into their antipredator display, only sand kicking behavior was a significant factor in motivating sidewinder rattlesnakes to cease hunting: high rates of sand kicking led to early abandonment of ambush coils. These results indicate that anti‐snake displays of small mammals may be especially effective at mitigating the threat posed by rattlesnakes when those displays incorporate physical harassment as well as signaling.     

Locking out predators by silk,a new counterattack behaviour in a social spider mite

1. A type of arms race that includes predation, counterattacks and cross‐counterattacks occurs between the phytophagous mite Stigmaeopsis nanjingensis (Ma et Yuan), which lives in self‐woven nests and exhibits cooperative sociality, and its specialised phytoseiid mite predator, Typhlodromus bambusae Ehara. 2. First, the efficiency of the S. nanjingensis (prey) counterattacking T. bambusae (predator) was observed. The prey females frequently locked the immature predators out of their nests using silk web, and the predators subsequently died of starvation. Furthermore, the prey males often killed immature T. bambusae mites after they invaded the nests. 3. This reversal of roles in the predator–prey system was then re‐reversed (returned to a normal state) by the behaviour of T. bambusae females. Immature predators could maintain their predacious natures due to the presence of attending adult females, which are able to cope with the prey counterattack behaviours.     

Ground squirrel tail-flag displays alter both predatory strike and ambush site selection behaviours of rattlesnakes

Many species approach, inspect and signal towards their predators. These behaviours are often interpreted as predator-deterrent signals-honest signals that indicate to a predator that continued hunting is likely to be futile. However, many of these putative predator-deterrent signals are given when no predator is present, and it remains unclear if and why such signals deter predators. We examined the effects of one such signal, the tail-flag display of California ground squirrels, which is frequently given both during and outside direct encounters with northern Pacific rattlesnakes. We video-recorded and quantified the ambush foraging responses of rattlesnakes to tail-flagging displays from ground squirrels. We found that tail-flagging deterred snakes from striking squirrels, most likely by advertising squirrel vigilance (i.e. readiness to dodge a snake strike). We also found that tail-flagging by adult squirrels increased the likelihood that snakes would leave their ambush site, apparently by elevating the vigilance of nearby squirrels which reduces the profitability of the ambush site. Our results provide some of the first empirical evidence of the mechanisms by which a prey display, although frequently given in the absence of a predator, may still deter predators during encounters.     

Increased predation risk while mate guarding as a cost of reproduction for male broad-headed skinks (Eumeces laticeps)

We investigated antipredatory costs associated with mate guarding as potential costs of reproduction for male broad-headed skinks. Mate guarding by male lizards may increase fitness by preventing loss of fertilizations of the guarded female's eggs to other males, but it may have several costs. In addition to lost opportunities to search for additional females, risk of injury while fighting other males, and energetic expenditures while following females and fighting, guarding males might suffer increased risk of predation and reduction of opportunities to forage. We studied potential antipredatory costs of mate guarding by simulating predators searching for and approaching pairs of lizards in the field. Among pairs of lizards in close proximity to each other, males were detected before females 10 times more frequently than females were detected before males, and females fled before males much more frequently than males fled before females when pairs were approached, leaving the males exposed to the predator. After one or both lizards fled, males frequently followed females by scanning visually and scent trailing, exposing themselves to the predator while the female hid. Females never followed males. The implications of these findings for antipredatory costs of mate guarding are discussed. Electronic Publication     

Pursuit-deterrence revisited

Pursuit-deterrent signals - signals used by prey that apparently convince predators not to pursue them - were discovered 15 years ago, but their existence continues to rest on shaky empirical evidence. First, pursuit-deterrent signals are usually inferred by eliminating competing hypotheses rather than testing predictions derived from the pursuit-deterrent hypothesis directly. Second, the strength of selection pressures maintaining such signals in prey populations are unknown because behavioral ecologists infrequently observe natural predation attempts. Third, the nature of information passing between prey and predators is open to misinterpretation because measures are rarely taken to separate signals that advertise perception of the predator from those that advertise perception and the prey's condition.     

Predator response to the coloured eyespots and defensive posture of Colombian four-eyed frogs

Deimatic displays, where sudden changes in prey appearance elicit aversive predator reactions, have been suggested to occur in many taxa. These (often only putative) displays frequently involve different components that may also serve antipredator functions via other mechanisms (e.g., mimicry, warning signalling, body inflation). The Colombian four-eyed frog, Pleurodema brachyops, has been suggested to gain protection against predation through putative deimatic displays where they inflate and elevate the posterior part of their body revealing eye-like colour markings. We exposed stationary artificial frogs to wild predators to test whether the two components (eyespot/colour markings, defensive posture) of their putative deimatic display, and their combination, provide protection from predation without the sudden change in appearance. We did not detect any obvious additive effect of defensive posture and eyespots/colour markings on predation risk, but found a marginally significant trend for model frogs in the resting posture to be less attacked when displaying eyespots/colour markings than when they were not, suggesting that the presence of colour markings/eyespots may provide some protection on its own. Additionally, we found that models in a resting posture were overall more frequently attacked on the head than models in a defensive posture, indicating that a defensive posture alone could help redirect predator attacks to non-vital parts of the body. The trends found in our study suggest that the different components of P. brachyops' coloration may serve different functions during a deimatic display, but further research is needed to elucidate the role of each component when accompanied by sudden prey movement.     

Tail flicking in the black redstart (<Emphasis Type="Italic">Phoenicurus ochruros</Emphasis>) and distance to cover

Tail flicking is a common behavior in many bird species, but its function is often unknown. Apart from intraspecific communication, tail flicking could be used during predator–prey communication, e.g., as a signal of prey vigilance or quality. We studied this behavior in the black redstart (Phoenicurus ochruros), a species that frequently shows tail flicking and is prone to attacks by ambushing predators that hide in cover. Hence, cover might be perceived as dangerous by this species. We hypothesized that flicking should increase with decreasing distance to cover. We counted the number of tail flicks of individuals and measured their distance to the nearest cover for an ambushing predator. We found that distance to cover had a significant effect on tail flicking behavior, as flicking increased with decreasing distance, but found no difference in flicking frequency between adults and juveniles or between sexes. Consequently, tail flicking is unlikely to signal submission or to be sexually selected in the black redstart. Since tail flicking also occurred in the absence of predators, we consider tail flicking in black redstarts to display vigilance and to be directed towards ambushing predators.     

Sex‐Specific Response of Pardosa milvina (Araneae: Lycosidae) to Experience with a Chemotactile Predation Cue

Predators frequently leave behind chemical information (i.e., semiochemicals such as pheromones or kairomones) that can be detected by their prey and used to avoid areas where predators are likely present. Prey that have interacted indirectly with predators via chemical information thus may gain insight into their risk of being consumed that naïve individuals lack. Pardosa milvina (Araneae: Lycosidae) is a chemosensitive wolf spider that shows adaptive responses to chemotactile cues deposited by the larger wolf spider Tigrosa helluo. We raised offspring from P. milvina to examine the effect of experience with a predation cue on activity, foraging, and antipredator behavior. Spiders differed in activity and foraging behavior across ontogeny and between sexes, but there was no effect of experience with a predation cue. However, a sex‐specific effect of experience was found in antipredator behavior. Male spiders, but not females, used experience with a predator cue to increase their survival in the presence of a live predator. Specifically, naïve males were attacked sooner than experienced males, indicating that prior exposure to predator cues can modify Pardosa antipredator behavior. Intersexual differences in how spiders respond to experience with a predation cue likely reflect the risk of predation faced by males and females in nature.     

Differential prey use by male and female cougars in Washington

Male and female predators are often assumed to have the same effects on prey. Because of differences in body size and behavior, however, male and female predators may use different species, sexes, and ages of prey, which could have important implications for wildlife conservation and management. We tested for differential prey use by male and female cougars (Puma concolor) from 2003 to 2008 in Washington State. We predicted that male cougars would kill a greater proportion of larger and older prey (i.e., adult elk [Cervus elaphus]), whereas females would kill smaller and younger prey (i.e., elk calves, mule deer [Odocoileus hemionus]). We marked cougars with Global Positioning System (GPS) radio collars and investigated 436 predation sites. We located prey remains at 345 sites from 9 male and 9 female cougars. We detected 184 mule deer, 142 elk, and 17 remains from 4 other species. We used log-linear modeling to detect differences in species and age of prey killed among cougar reproductive classes. Solitary females and females with dependent offspring killed more mule deer than elk (143 vs. 83, P < 0.01), whereas males killed more elk than mule deer (59 vs. 41, P < 0.01). Proportionately, males killed 4 times more adult elk than did females (24% vs. 6% of kills) and females killed 2 times more adult mule deer than did males (26% vs. 15% of kills). Managers should consider the effects of sex of predator in conservation and management of ungulates, particularly when managing for sensitive species. © 2011 The Wildlife Society.     

Aggregation of whelks, Buccinum undatum, near feeding predators: the role of reproductive requirements

In the Mingan Islands, northern Gulf of St Lawrence (eastern Canada), the whelk Buccinum undatum displays a strong escape response to its predator, the asteroid Leptasterias polaris, nevertheless large sexually mature individuals occasionally approach feeding L. polaris to obtain food. In this study, we investigated the hypothesis that reproductive requirements increase the tendency of sexually mature whelks to approach feeding asteroids. Prior to egg laying, females (which invest more energy than males into the production of reproductive structures) represented 72% of the adult whelks that approached feeding L. polaris, but only 36% of the adults randomly collected from the study area. Furthermore, females that were attracted to feeding asteroids had smaller reproductive organs (after accounting for body size) than females randomly collected from the study area. Similarly, prior to egg laying, females fed longer and ingested more food than males when tested in the presence of L. polaris in the laboratory. After egg laying, however, females and males displayed a similar tendency to feed in the presence of a predator, both in the field and in the laboratory. Predator-impact indices, computed by contrasting the feeding activity of whelks in the absence and presence of a predator, indicated that females (but not males) responded more boldly to predators prior to than after egg laying, despite a general decrease in feeding activity at that period. Taken together, our observations indicate that the tendency of adult whelks to approach feeding predators is influenced by potential reproductive gains. Because such gains are presumably more directly linked to a given feeding opportunity in sexually active individuals, whelks may be selected to display increased levels of boldness towards predators with the onset of sexual maturity. Thus, potential reproductive benefits may partly explain the size-dependent tendency of whelks to approach feeding asteroids. Copyright 2001 The Association for the Study of Animal Behaviour.     

Parasitism-mediated prey selectivity in laboratory conditions and implications for biological control

In agroecosystems, parasitoids and predators may exert top-down regulation and predators for different reasons may avoid or give preference to parasitised prey, i.e., become an intraguild predator. The success of pest suppression with multiple natural enemies depends essentially on predator–prey dynamics and how this is affected by the interplay between predation and parasitism. We conducted a simple laboratory experiment to test whether predators distinguished parasitised prey from non-parasitised prey and to study how parasitism influenced predation. We used a host-parasitoid system, Spodoptera frugiperda and one of its generalist parasitoids, Campoletis flavicincta, and included two predators, the stinkbug Podisus nigrispinus and the earwig Euborellia annulipes. In the experiment, predators were offered a choice between non-parasitised and parasitised larvae. We observed how long it took for the predator to attack a larva, which prey was attacked first, and whether predators opted to consume the other prey after their initial attack. Our results suggest that, in general, female predators are less selective than males and predators are more likely to consume non-parasitised prey with this likelihood being directly proportional to the time taken until the first prey attack. We used statistical models to show that males opted to consume the other prey with a significantly higher probability if they attacked a parasitised larva first, while females did so with the same probability irrespective of which one they attacked first. These results highlight the importance of studies on predator–parasitoid interactions, as well as on coexistence mechanisms in agroecosystems. When parasitism mediates predator choice so that intraguild predation is avoided, natural enemy populations may be larger, thus increasing the probability of more successful biological control.     

Fishing spiders, green sunfish, and a stream-dwelling water strider: male-female conflict and prey responses to single versus multiple predator environments

Many studies have experimentally addressed the effects of a particular predator species on prey behavior. In nature, however, prey frequently face multiple species of predators that often vary in their predatory mode and in their level of predation risk. Relatively few studies have considered prey responses under these complex conditions. In Kentucky, the stream-dwelling water strider (Aquariusremigis) coexists with many potentially dangerous predators, two of which are the green sunfish (Lepomiscyanellus) and the fishing spider (Dolomedesvittatus). Green sunfish occupy stream pools and attack water striders from below. In contrast, fishing spiders hunt along stream shorelines where they perch on overhanging vegetation or rocks and attack water striders near shore. We compared how A. remigis individuals respond to these two very different predators in pools with one or both predators. The presence of sunfish in pools had strong effects on male water strider behavior, including increased use of three types of refuge from sunfish (riffles, climbing out of the water, sitting on the water but at the edges of pools), decreased activity and a decreased number of aggressive males on the water. Spiders also influenced water strider behavior; male water striders avoided spiders by shifting away from the edges of pools. Comparisons of the effects of the two predator species showed that in general, antipredator responses by male water striders were stronger in pools with fish alone than in those with spiders alone. In the presence of both predators, male water strider behavior (microhabitat use and activity) was generally similar to behavior in the presence of fish alone. In contrast, female water striders showed no significant response to the presence of sunfish, and little response to the presence of spiders. This lack of response could be because females spent much of their time in refuges even in the absence of predators (apparently hiding from harassment by males). Both spiders and fish caused decreases in water strider mating activity. The presence of fish reduced both the number of matings per pool (mating frequency), and mean mating durations. Spiders induced a decrease in mean mating duration, but not in mating frequency. The largest reductions in mating activity occurred in pools with both predators present. Pools with either spiders or fish alone suffered 15–20% water strider mortality during our experiment (versus no mortality in predator-free pools). Extant theory suggests that when prey face conflicting microhabitat responses to two predators (as in this study), the predators should have facilitative effects on predation rates (i.e., prey that avoid one predator are often killed by the other and vice versa). Mortality rates in pools with both predators present, however, were not significantly different from that predicted by a null model of multiple predator effects. The lack of predator facilitation can be explained by the compensatory reductions in water strider activity and mating activity in the presence of both predators. Received: 26 August 1996 / Accepted: 12 June 1998     

Frequency of tail loss reflects variation in predation levels,predator efficiency,and the behaviour of three populations of brown anoles

We investigated two predictions regarding the incidence of tail regeneration in lizards for three populations of brown anoles exposed to varying predation levels from the same predator (cats). Firstly although inefficient predators are likely to increase the incidence of regenerated tails (i.e. lizards can escape through tail autotomy), highly efficient predators will kill and eat the lizard and thus leave no evidence of autotomy. At the site with no cats, only 4% of anoles demonstrated signs of tail regeneration. This value was not significantly different from the site where feral cats (i.e. ‘efficient’ predators that would capture prey to eat, as supported by behavioural observation) were present (7%). By contrast, 25% of anoles present at the site with pet cats (well‐fed domesticated cats that caught and played with anoles, i.e. were ‘inefficient’ predators) exhibited regenerated tails. Secondly, more obvious lizards are more susceptible to predation attempts. Supporting this hypothesis, our data indicate a higher incidence of regenerated tails (28%) was recorded amongst adult males (which are territorial, occupying exposed positions) compared to females and subadult males (17%) or juveniles (1%). In conclusion, the behaviour of both the predator and the lizard influences the frequency of regenerated tails in brown anoles. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 648–656.