Predator detection and dilution as benefits of associations between yellow mongooses and Cape ground squirrels

Associations among organisms are thought to form because the benefits, such as increased foraging efficiency or decreased risk of predation, outweigh any costs, such as resource competition. Though many interspecific associations have been described for closely related mammals, few studies have examined the associations between mammals in different orders. The yellow mongoose (Cynictus pencillata), a carnivore, and the Cape ground squirrel (Xerus inauris), a rodent, co-occur in arid and semi-arid South Africa where they share sleeping burrows, predators, a similar body size, and the capability to emit alarm calls in response to predators. To investigate enhanced predator avoidance as a potential benefit explaining the persistence of this association, we assessed individual mongoose vigilance alone and with squirrels or other mongooses, and with varying interspecific group size, using field observations. We also tested for responses to conspecific and heterospecific alarm calls in both study species using playback experiments. The proportion of time mongoose individuals spent vigilant decreased in the presence of squirrels or other mongooses and was negatively correlated with interspecific group size; a similar pattern was previously shown for conspecific groups of Cape ground squirrels. These results are predicted by both the dilution and collective detection hypotheses. In addition, hetero- and conspecific alarm calls elicited vigilance responses in both species. These results suggest that both species can benefit from the collective detection and dilution arising from their interspecific association and that this interspecific association could be mutualistic.     

Interspecific associations of Cape ground squirrels with two mongoose species: benefit or cost?

Mixed-species associations have been described in many vertebrate species, but few behavioral studies have investigated associations between species from different mammalian orders. Cape ground squirrels (Xerus inauris) are highly social rodents that inhabit burrows with two species of mongoose, but the benefits of these interspecific associations to ground squirrels remain unresolved. We compared the behavior of squirrels while solitary, with conspecifics, and in the presence of suricates (Suricata suricatta) and yellow mongooses (Cynictis pencillatus). Squirrels spent less time alert and more time feeding when suricates were present, but increased vigilance in the presence of yellow mongooses. In a series of mobbing trials with a puff adder (Bitis arietans), a common predator of all three species, Cape ground squirrels were the most active in mobbing the snake. Our results suggest that Cape ground squirrels benefit from associating with suricates, but not necessarily with yellow mongooses. Both mongoose species benefit from the burrowing activities of the squirrels for thermoregulation and escape from predators, and a suite of other organisms may similarly benefit from the habitat modifications by Cape ground squirrels, suggesting they could be considered ecosystem engineers of the arid and semi-arid regions of southern Africa. Thus, the association between Cape ground squirrels and suricates appears mutually beneficial, whereas yellow mongooses may merely be commensals of the squirrels.     

Dwarf mongoose and hornbill mutualism in the Taru desert,Kenya

Summary Dwarf mongooses in the Taru desert region of Kenya form foraging communities with a variety of endemic bird species, especially hornbills. The prey spectra of the mongooses and hornbills overlap almost completely. For the other bird species forming the foraging community only partial overlap exists. The association between the birds and mongooses is actively sought by both parties. The birds wait in tress around the termite mound where the monogooses are sleeping for them to emerge and the mongooses delay their foraging departure if no birds are present. There is a positive relationship between the number of mongooses in the group and the number of birds accompanying them. A true mutualism only exists between the mongooses and the two hornbill species Tockus deckeni and T. flavirostris since their presence or arrival affects the subsequent start of foraging. These two hornbill species have also been observed to influence the start of foraging actively by means of two behaviour patterns termed chivvying and waking. Both the mongooses and birds are exposed to a high predator pressure from raptors with an overlap in the birds of prey predating the various species. This predator pressure is counteracted behaviourally by the mongooses by means of an altruistic behaviour pattern, guarding. Both mongooses and birds warn vocally and flee when a raptor is sighted. The mongooses modify their guarding behaviour to compensate for the warning behaviour of the birds in two ways: (a) fewer mongooses guard when large numbers of birds are present and vice versa, (b) the frequency of the mongooses' intraspecific warning calls is significantly reduced in cases where birds are present in comparison with those where they are absent. The birds also sight and respond to the raptor first on significantly more occasions than the mongooses. In addition, the birds also warn for raptor species which do not predate them but which are mongoose predators, not, however, for raptors which are not mongoose predators. This mutualistic association with its high degree of compensatory behaviour by both parties appears to be unique for free-living vertebrates and has its closest parallel in the trophobiosis described for ants and aphids.     

All clear? Meerkats attend to contextual information in close calls to coordinate vigilance

Socio-demographic factors, such as group size and their effect on predation vulnerability, have, in addition to intrinsic factors, dominated as explanations when attempting to understand animal vigilance behaviour. It is generally assumed that animals evaluate these external factors visually; however, many socially foraging species adopt a foraging technique that directly compromises the visual system. In these instances, such species may instead rely more on the acoustical medium to assess their relative risk and guide their subsequent anti-predator behaviour. We addressed this question in the socially foraging meerkat (Suricata suricatta). Meerkats forage with their head down, but at the same time frequently produce close calls (‘Foraging’ close calls). Close calls are also produced just after an individual has briefly scanned the surrounding environment for predators (‘Guarding’ close calls). Here, we firstly show that these Guarding and Foraging close call variants are in fact acoustically distinct and secondly subjects are less vigilant (in terms of frequency and time) when exposed to Guarding close call playbacks than when they hear Foraging close calls. We argue that this is the first evidence for socially foraging animals using the information encoded within calls, the main adaptive function of which is unrelated to immediate predator encounters, to coordinate their vigilance behaviour. In addition, these results provide new insights into the potential cognitive mechanisms underlying anti-predator behaviour and suggest meerkats may be capable of signalling to group members the ‘absence’ of predatory threat. If we are to fully understand the complexities underlying the coordination of animal anti-predator behaviour, we encourage future studies to take these additional auditory and cognitive dimensions into account.     

Within-group spatial position and vigilance: a role also for competition? The case of impalas (<Emphasis Type="Italic">Aepyceros melampus</Emphasis>) with a controlled food supply

Theory predicts that individuals at the periphery of a group should be at higher risk than their more central conspecifics since they would be the first to be encountered by an approaching terrestrial predator. As a result, it is expected that peripheral individuals display higher vigilance levels. However, the role of conspecifics in this “edge effect” may have been previously overlooked, and taking into account the possible role of within-group competition is needed. Vigilance behavior in relation to within-group spatial position was studied in impalas (Aepyceros melampus) feeding on standardized patches. We also controlled for food distribution in order to accurately define a “central” as opposed to a “peripheral” position. Our data clearly supported an edge effect, with peripheral individuals spending more time vigilant than their central conspecifics. Data on social interactions suggest that it was easier for a foraging individual to defend its feeding patch with its head lowered, and that more interactions occurred at the center of the group. Together, these results indicate that central foragers may reduce their vigilance rates in response to increased competition. Disentangling how the effects of competition and predation risk contribute to the edge effect requires further investigations.     

Information transfer and gain in flocks: the effects of quality and quantity of social information at different neighbour distances

We assessed experimentally how the quality and quantity of social information affected foraging decisions of starlings (Sturnus vulgaris) at different neighbour distances, and how individuals gained social information as a function of head position. Our experimental set up comprised three bottomless enclosures, each housing one individual placed on a line at different distances. The birds in the extreme enclosures were labelled senders and the one in the centre receiver. We manipulated the foraging opportunities of senders (enhanced, natural, no-foraging), and recorded the behaviour of the receiver. In the first experiment, receivers responded to the condition of senders. Their searching rate and food intake increased when senders foraged in enhanced conditions, and decreased in no-foraging conditions, in relation to natural conditions. Scanning was oriented more in the direction of conspecifics when senders behaviour departed from normal. In the second experiment, responses were dose dependent: receivers increased their searching rate and orientated their gaze more towards conspecifics with the number of senders foraging in enhanced food conditions. In no-foraging conditions, receivers decreased their searching and intake rates with the number of senders, but no variation was found in scanning towards conspecifics. Differences in foraging and scanning behaviour between enhanced and no-foraging conditions were much lower when neighbours were separated farther. Overall, information transfer within starling flocks affects individual foraging and scanning behaviour, with receivers monitoring and copying senders behaviour mainly when neighbours are close. Information transfer may be related to predation information (responding to the vigilance of conspecifics) and foraging information (responding to the feeding success of conspecifics). Both sources of information, balanced by neighbour distance, may simultaneously affect the behaviour of individuals in natural conditions.Communicated by H. Kokko     

Group-living in cliff swallows as an advantage in avoiding predators

Summary Cliff swallows (Hirundo pyrrhonota) in SW Nebraska, USA, nest in colonies and associate in groups away from their colonies. The degree to which group-living in this species affords advantages in the avoiding of predators was examined. The distance from the colony at which a snake predator was detected increased with colony size. In flocks away from the colonies, group vigilance increased, but the time that each individual spent vigilant decreased, with flock size. As a result, birds in large flocks had more time for preening and mud-gathering. Cliff swallows did not effectively mob predators and thus were unable to deter predators regardless of group size. Nesting within each colony was highly synchronous, but when the effects of ectoparasites on nesting success were removed, individuals nesting during the peak breeding period were no more successful than those nesting before or after the peak. This suggests that swamping of predators is unlikely in cliff swallow colonies. Nests at the edges of colonies were more likely to be preyed upon than nests nearer the center, suggesting that colonial nesting conferred some selfish herd benefits. Overall reproductive success did not vary with colony size. While cliff swallows receive some anti-predator benefits by living in groups, the avoidance of predators is probably not a major selective force for the evolution of coloniality in this species.     

Individual variation in the relationship between vigilance and group size in eastern grey kangaroos

The mean vigilance of animals in a group often decreases as their group size increases, yet nothing is known about whether there is individual variability in this relationship in species that change group sizes frequently, such as those that exhibit fission–fusion social systems. We investigated variability in the relationship between group size and vigilance in the eastern grey kangaroo (Macropus giganteus) by testing whether all individuals showed decreased vigilance with increased group size, as has been commonly assumed. We carried out both behavioural observations of entire groups of kangaroos and focal observations of individually recognised wild female kangaroos. As in other studies, we found a collective group-size effect on vigilance; however, individuals varied in their vigilance patterns. The majority (57%) of the identified individual kangaroos did not show significant group-size effects for any of the recorded measures of vigilance. The females that did not show a negative group-size effect were, on average, more vigilant than those females that did show a group-size effect, but this difference was not significant. We propose that some females exhibit higher levels of social vigilance than others, and that this social vigilance increases with group size, cancelling out any group-size effect on anti-predator vigilance for those females. Our results therefore suggest that only some prey individuals may gain anti-predator benefits by reducing their time spent scanning when in larger groups. The large amount of variation that we found in the vigilance behaviour of individual kangaroos highlights the importance of collecting and analysing vigilance data at the individual level, which requires individual recognition.     

Flexible bat echolocation: the influence of individual,habitat and conspecifics on sonar signal design

Acoustic signals which are used in animal communication must carry a variety of information and are therefore highly flexible. Echolocation has probably such functions and could prove as flexible. Measurable variabitlity can indicate flexibility in a behaviour. To quantify variability in bat sonar and relate to behavioural and environmental factors, I recorded echolocation calls of Euderma maculatum, Eptesicus fuscus, Lasiurus borealis and L. cinereus while the bats hunted in their natural habitat. I analysed 3390 search phase calls emitted by 16 known and 16 unknown individuals foraging in different environmental and behvioural situations. All four species used mainly multiharmonic signals that showed considerable intra- and inter-individual variability in the five signal variables I analysed (call duration, call interval, highest and lowest frequency and frequency with maximum energy) and also in the shape of the sonagram. A nested multivariate analysis of variance identified the influences of individual, hunting site, close conspecifics and of each observation on the frequency with maximum energy in the calls, and on other variables measured. Individual bats differed in multiple comparisons, most often in the main call frequency and least often in call interval. In a discriminant function analysis with resubstitution, 56–76% of a species' calls were assigned to the correct individual. Distinct individual call patterns were recorded in special situations in all species and the size of foraging areas in forested areas influenced temporal and spectral call structure. Echolocation behaviour was influenced by the presence of conspecifics. When bats were hunting together, call duration decreased and call interval increased in all species, but spectral effects were less pronounced. The role of morphometric differences as the source of individually distinct vocalizations is discussed. I also examined signal adaptations to long range echolocation and the influence of obstacle distance on echolocation call design. My results allow to discuss the problems of echo recognition and jamming avoidance in vespertilionid bats.     

Group size and foraging efficiency in yellow baboons

Summary I studied the foraging behaviour of adults in three different-sized groups of yellow baboons (Papio cynocephalus) at Amboseli National Park in Kenya to assess the relationship between group size and foraging efficiency in this species. Study groups ranged in size from 8 to 44 members; within each group, I collected feeding data for the dominant adult male, the highest ranking pregnant female, and the highest ranking female with a young infant. There were no significant differences between groups during the study in either the mean estimated energy value of the food ingested per day for each individual (385±27 kJ kg^-1 day^-1) or in the estimated energy expended to obtain that food (114±3 kJ kg^-1 day^-1). Mean foraging efficiency ratios, which reflect net energy gain per unit of foraging time, also did not vary as a function of the size of the group in which the baboons were living. There was substantial variation between days in the efficiency ratios of all animals; this was the result of large differences in energy intake rather than in the energy expended during foraging itself. The members of the smallest group spent on the average only one-half as much time feeding each day as did individuals in the two larger groups. However, they obtained almost as much energy while foraging, primarily because their rate of food intake while actually eating tended to be higher than the rate in the other groups. The baboons in the small group were observed closer to trees that they could climb to escape ground predators, and they also were more likely to sit in locations elevated above the ground while resting. Such differences would be expected if the members of the small group were less able to detect approaching predators than individuals that lived in the larger groups. The results of this study suggest that predator detection or avoidance, rather than increased foraging efficiency, may be the primary benefit of living in larger groups in this population.     

Threat-sensitive foraging by larval threespine sticklebacks (Gasterosteus aculeatus)

Summary The threat-sensitive predator avoidance hypothesis predicts that prey can assess the relative threat posed by a predator and adjust their behaviour to reflect the magnitude of the threat. We tested the ability of larval threespine sticklebacks to adjust their foraging in the presence of predators by exposing them to conspecific predators of various sizes and recording their foraging and predator avoidance behaviours. Larvae (<30 days post-hatch) displayed predator escape behaviours only towards attacking predators. At 3 weeks post-hatch larvae approached the predator after fleeing, a behaviour which may be the precursor to predator inspection. Larvae reduced foraging and spent less time in the proximity of large and medium-sized predators compared to small predators. The reduction in foraging was negatively correlated to the predator/larva size ratio, indicating that larvae increased their foraging as they increased in size relative to the predator. We conclude that larval sticklebacks can assess the threat of predation early in their ontogeny and adjust their behaviour accordingly.Correspondence to: J.A. Brown     

Vigilance for predators: detection and dilution effects

Summary Grouped individuals are less vigilant for predators than solitary conspecifics because (1) grouping increases the likelihood of predator detection (detection effect) and (2) grouping makes it less likely that any given individual will be preyed upon (dilution effect). However, many models of vigilance behaviour consider only the detection effect, and the interaction of the two effects has been insufficiently considered. I present two models of vigilance behaviour and test them using data on the vigilance of elk, Cervus elaphus. The first model, based on the detection effect alone, is implicit in many published formulations of the relationship between vigilance and group size. Although it predicts the direction of the relationship between vigilance and group size, it provides a poor explanation of the form of the relationship. The second model incorporates both detection and dilution effects. Regression analysis on this security model indicates that the model provides a good prediction of both the direction and the form of the relationship between vigilance and group size, explaining 69% of the variance in vigilance frequency. The security model demonstrates that both detection and dilution are important in determining the frequency of vigilance behaviour but that the relative importance of these two effects changes across group size, with detection providing relatively less benefit as group size increases. However, even when groups are large, individuals should exhibit at least some vigilance because although dilution alone provides much protection from predation, even a low level of vigilance greatly increases the likelihood that an individual will survive repeated predation attempts.     

Social foraging in stingless bees: how colonies of Melipona fasciata choose among nectar sources

In an experimental set-up, a colony of the stingless bee Melipona fasciata demonstrated its ability to choose the better of two nectar sources. This colony pattern was a result of the following individual behavioural decisions: continue foraging, abandon the feeder, restart foraging and initiate foraging. Only very rarely did individuals switch from one feeder to the other. With the first combination of a rich (2.7 M) and a poor (0.8 M) feeder M. fasciata behaved differently from Apis mellifera. Recruitment occurred to both feeders and the poor feeder was not abandoned completely. When the poor feeder was set to 0.4 M, M. fasciata abandoned the poor feeder rapidly and allocated more foragers to the rich feeder. These patterns were similar to those reported for A. mellifera with the first combination of feeders. Over a sequence of 4 days, experienced bees increasingly determined the colony patterns, and the major function of communication between workers became the reactivation of experienced foragers. The foragers modulated their behaviour not only according to the profitability of the feeder, but also according to previous experience with profitability switches. Thus, experience and communication together regulated colony foraging behaviour. These findings and the results of studies with honeybees suggest that M. fasciata and honeybees use similar decision-making mechanisms and only partly different tools. Received: 21 December 1998 / Accepted: 5 January 1999     

Ontogenetic changes in social behaviour in the forest tent caterpillar,<Emphasis Type="Italic"> Malacosoma disstria</Emphasis>

Many animals, including gregarious caterpillars, begin life in groups and become increasingly solitary as they grow larger. These ontogenetic changes in social behaviour suggest that the costs and benefits of grouping change with increasing individual size. Forest tent caterpillar (Malacosoma disstria) colonies exhibit complex social behaviour during the early larval instars, using pheromone trails to move together between temporary bivouacs and feeding sites, and break up as the caterpillars grow. We demonstrate changes in individual responses to cues from conspecifics that explain changes in aggregation during caterpillar development. We used Markov chain analysis to test the influence of pheromone trails and colony-mates on an individual caterpillars tendency to switch between quiescence, searching, walking and spinning. Pheromone-laden silk trails increased the tendency to begin locomotion, whereas colony-mates increased switching from activity to quiescence. Trails also influenced the form taken by locomotor behaviour, and promoted directed walking over searching. Social cues thus increase the efficiency of individual locomotion. Younger larvae were more quiescent and more reluctant to walk in the absence of trails than were older insects. An increase in independent locomotion as the larvae grow provides a mechanism to explain colony break-up and points to an ontogenetic shift in the internal processes driving behaviour. Scaling relationships suggest that many of the benefits associated with group-living in caterpillars decrease as individuals grow larger, providing an adaptive explanation for observed ontogenetic changes in social behaviour.Communicated by N. Wedell     

Testing a stochastic version of the Beddington–DeAngelis functional response in foraging shore crabs

Current behaviour-based interference models assume that the predator population is infinitely large and that interference is weak. While the realism of the first assumption is questionable, the second assumption conflicts with the purpose of interference models. Here, we tested a recently developed stochastic version of the Beddington–DeAngelis functional response—which applies to a finite predator population without assuming weak interference—against experimental data of shore crabs (Carcinus maenas) foraging on mussels (Mytilus edulis). We present an approximate maximum likelihood procedure for parameter estimation when only one focal individual is observed, and introduce ‘correction factors’ that capture the average behaviour of the competing but unobserved individuals. We used the method to estimate shore crab handling time, interaction time, and searching rates for prey and competitor. Especially the searching rates were sensitive to variation in prey and competitor density. Incorporating constant parameter values in the model and comparing observed and predicted feeding rates revealed that the predictive power of the model is high. Our stochastic version of the Beddington–DeAngelis model better reflects reality than current interference models and is also amenable for modelling effects of interference on predator distributions.     

Female foraging and male vigilance in white-tailed ptarmigan (Lagopus leucurus): opportunism or behavioural coordination?

For several species of birds, high rates of male vigilance are correlated with high rates of female foraging. This relationship is thought to ultimately result in higher reproductive success for females paired with highly vigilant males. However, previous research has not examined the behavioural mechanism that produces the correlation between male vigilance and rates of female foraging. Foraging females may take advantage of vigilance that males are using for other purposes. Alternatively, the purpose of male vigilance may be to increase females' ability to forage. We examined these alternatives by testing whether vigilance preceded or followed bouts of female foraging more often than would occur by chance alone, using simultaneous behaviour observations of pre-incubation pairs of white-tailed ptarmigan (Lagopus leucurus). Our results indicate that each member of a pair may influence the behaviour of the other. Females were more likely to initiate foraging bouts after males became vigilant than if their mate remained non-vigilant. Moreover, non-vigilant males were more likely to become vigilant if their mate was foraging than if she was engaged in some other activity. Despite the possibility that a sexual conflict exists as each member of a pair attempts to maximize its fitness, both sexes behave as though a major role of male vigilance is to enhance female foraging opportunities. Received: 3 May 1999 / Received in revised form: 14 June 1999 / Accepted: 16 June 1999     

<Emphasis Type="Italic">Octopus insularis</Emphasis> (Octopodidae), evidences of a specialized predator and a time-minimizing hunter

Shallow-water octopuses have been reported as major predators of motile species in benthonic marine communities, capturing their prey by different foraging techniques. This study assessed for the first time the feeding ecology, foraging behavior, and defensive strategy during foraging, including the use of body patterns, to construct a general octopus foraging strategy in a shallow water-reef system. Octopus insularis was studied in situ using visual observations and video recordings. The diet included at least 55 species of crustaceans (70%), bivalves (17.5%), and gastropods (12.5%); however, only four species accounted for half of the occurrences: the small crabs Pitho sp. (26.8%) and Mithrax forceps (23.9%), the bivalve Lima lima (5.3%), and the gastropod Pisania pusio (4.9%). Poke and crawl were most frequent foraging behaviors observed in the video recordings. The foraging behaviors were associated with environmental variables and octopus body size. The sequences of foraging behavior showed characteristics of a tactile saltatory searching predator, as well as a visual opportunist. Body patterns showed a relationship with foraging behavior, habitat variables, and octopus body size. Mottle was the most frequent pattern, especially during poke and crawl, in shallower depths. Dorsal light–ventral blue green was more frequent during swimming at mid-water, and Blotch was the normal pattern during web-over by large animals. The large proportion of two species of small crabs in den remains, the intense search for food during short hunting trips, and the intense use of cryptic body patterns during foraging trips, suggest that this species is a ‘time-minimizing’ forager instead of a ‘rate-maximizer’.     

Vocal mediation of foraging competition in the cooperatively breeding green woodhoopoe (<Emphasis Type="Italic">Phoeniculus purpureus</Emphasis>)

Many group-living species produce frequent vocalisations when foraging, but the function of these food-associated calls is often difficult to divine. I investigated the kek call of the cooperatively breeding green woodhoopoe (Phoeniculus purpureus), a species in which individuals have preferred foraging techniques dependent on their bill size. Individuals called at a greater rate (1) in foraging compared to non-foraging situations, and (2) in groups containing potential foraging competitors (i.e. individuals that foraged using the same preferred techniques). I therefore asked whether the kek call is used to recruit conspecific foragers or whether it acts as a vocal signal of foraging niche and mediates foraging competition. Foragers that were vocalising were no more likely to be approached than those that were silent, and individuals gained no foraging advantage from the close proximity of another group member. Thus, keks are unlikely to be used to recruit conspecifics. Instead, they appear to regulate spacing between potential foraging competitors. Although an individual forager was equally likely to be closely approached by all other group members, it increased its calling rate only in response to potential foraging competitors. This increase in calling rate resulted in the approaching individual moving away, thus maintaining some separation between individuals that forage in the same way. Maintenance of such spacing is important because the success rate of an individual decreased when a foraging competitor was close by.Communicated by M. Leonard     

Social facilitation of exploration in mosquitofish (<Emphasis Type="Italic">Gambusia holbrooki</Emphasis>)

The social environment can exert a powerful influence on the expression of an individual’s behaviour patterns. For example, social facilitation occurs where individuals are more likely to express a given behaviour, or express it a greater rate, in the presence of conspecifics. Social facilitation is partly driven by an individual’s perception of risk, which is a function both of the size of its social group and the information that it gathers relating to predator activity and risk. Here I tested the effects of social group size (one, two, four, eight or 16 fish) and the presence of ‘social’, ‘predation’ or ‘neutral’ chemical cues (derived respectively from live conspecifics, injured conspecifics or a blank water control) on the exploratory behaviour of juvenile mosquitofish (Gambusia holbrooki) in a novel environment. Focal fish in larger groups explored a greater proportion of the arena during the course of the experiment, demonstrating social facilitation of exploration. After 4 h in the arena, focal fish in all group sizes showed significantly reduced swimming activity, suggesting that the initial faster swimming activity of fish on entry to the arena may be in response to the motivation to explore. The presence of predation cues in the environment had the effect of reducing exploratory behaviour across groups in a novel environment, whereas social cues had no effect on exploratory behaviour. Taken as a whole, these results suggest that there is a high degree of context dependency in the expression of exploratory behaviour, with a strong influence of both the presence of conspecifics and cues relating to potential danger.     

Different antipredator behaviour in two anuran tadpoles: effects of predator diet

Recent investigations have indicated that animals are able to use chemical cues of predators to assess the magnitude of predation risk. One possible source of such cues is predator diet. Chemical cues may also be important in the development of antipredator behaviour, especially in animals that possess chemical alarm substances. Tadpoles of the common toad (Bufo bufo) are unpalatable to most vertebrate predators and have an alarm substance. Tadpoles of the common frog (Rana temporaria) lack both these characters. We experimentally studied how predator diet, previous experience of predators and body size affect antipredator behaviour in these two tadpole species. Late-instar larvae of the dragonfly Aeshna juncea were used as predators. The dragonfly larvae were fed a diet exclusively of insects, R. temporaria tadpoles or B. bufo tadpoles. R. temporaria tadpoles modified their behaviour according to the perceived predation risk. Depending on predator diet, the tadpoles responded with weak antipredatory behaviour (triggered by insect-fed predators) or strong behaviour (triggered by tadpole-fed predators) with distinct spatial avoidance and lowered activity level. The behaviour of B. bufo in predator diet treatments was indistinguishable from that in the control treatment. This lack of antipredator behaviour is probably related to the effective post-encounter defenses and more intense competitive regime experienced by B. bufo. The behaviour of both tadpole species was dependent on body size, but this was not related to predator treatments. Our results also indicate that antipredator behaviour is largely innate in tadpoles of both species and is not modified by a brief exposure to predators. Received: 22 August 1996 / Accepted after revision: 31 January 1997