The costs and benefits of predator inspection behaviour in Thomson's gazelles

Summary When Thomson's gazelles (Gazella thomsoni) detect stalking predators, such as cheetahs (Acinonyx jubatus) and lions (Panthera leo), they often approach and follow the predator for up to 72 min (average 14 min). Coursing predators are rarely approached. Gazelle groups were more likely to approach cheetahs if the groups were larger, if the vegetation was low, or if the cheetahs came closer to the group. Immature gazelles were more likely to approach than adults, and a higher proportion of group members participated in inspection behaviour in small groups than in large ones. Gazelles approached closer in less risky situations: if they were in larger groups or if the vegetation was low. Inspection behaviour caused cheetahs to move further between rests and between hunting attempts. Approaching cheetahs was risky, particularly for younger gazelles (probability of being killed while inspecting a cheetah was 1 in 5000 approaches for adults and 1 in 417 approaches for half-grown/adolescent gazelles), and the risks were higher than monitoring cheetahs from a distance. The time costs of predator inspection were also considerable (less than 4.2% of daylight time budget), suggesting that the benefits must be substantial to offset these costs. The results suggested that inspection behaviour was multifunctional, causing stalking predators to move out of the vicinity, enabling gazelles to monitor the predators' movements, and providing an opportunity, particularly for younger animals, to learn about predators. By approaching, gazelles also inform predators that they have been detected and alert other gazelles to the predators' presence.     

Does information of predators influence general wariness?

Antipredator behavior includes several qualitatively distinct activities, but few studies have determined the degree to which these activities are independent. If behaviors are not independent, then the nature of the relationship would illustrate potential performance constraints. We studied crimson rosellas (Platycercus elegans) and first focused on acoustic predator discrimination. We measured time allocation before and after playback of one of three experimental treatments (peregrine falcons—Falco peregrinus, wedge-tailed eagles—Aquila audax, and crimson rosellas) to determine whether or not rosellas discriminated predators from non-predators, and specifically whether or not they discriminated large from small predators. We then focused on the decision to flee. We experimentally approached subjects and measured the distance at which they oriented to us (alert distance) and the distance at which they fled (flight initiation distance; FID). We found that rosellas could distinguish among predators; however, there was no effect on general wariness as measured by FID. These two processes of antipredator behavior may, thus, be independent.     

Making sense of predator scents: investigating the sophistication of predator assessment abilities of fathead minnows

According to the threat-sensitive predator avoidance hypothesis, selection favors prey that accurately assess the degree of threat posed by a predator and adjust their anti-predator response to match the level of risk. Many species of animals rely on chemical cues to estimate predation risk; however, the information content conveyed in these chemical signatures is not well understood. We tested the threat-sensitive predator avoidance hypothesis by determining the specificity of the information conveyed to prey in the chemical signature of their predator. We found that fathead minnows (Pimephales promelas) could determine the degree of threat posed by northern pike (Esox lucius) based on the concentration of chemical cues used. The proportion of minnows that exhibited an anti-predator response when exposed to a predator cue increased as the concentration of the pike cue used increased. More surprisingly, the prey could also distinguish large pike from small pike based on their odor alone. The minnows responded more intensely to cues of small pike than to cues of large pike. In this predator–prey system small pike likely represent a greater threat than large pike.Communicated by A. Mathis     

Innate antipredator responses of Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) depend on predator species and their diet

The ability to discriminate between more dangerous and less dangerous predators can have serious fitness advantages for fish juveniles. This is especially true for hatchery-reared fish young used for stocking, because their post-release mortality is often much higher than that of wild-born conspecifics. We tested whether two coexisting fish predators and their different diets induce innate behavioral responses in predator-naive Arctic charr (Salvelinus alpinus) young originating from an endangered hatchery-bred population used for re-introductions. We predicted the antipredator responses of charr to be stronger towards chemical cues of brown trout (Salmo trutta) and pikeperch (Stizostedion lucioperca) than towards odorless control water. More pronounced antipredator behavior was predicted in treatments with predators fed on charr than when their diet consisted of another sympatric salmonid, European grayling (Thymallus thymallus), or when they were food-deprived. The Arctic charr young showed strong antipredator responses in all brown trout treatments, whereas odors of the less likely predator pikeperch were avoided with conspecific diet only. Freezing was the most sensitive antipredator behavior, as it was completely absent in control treatments. We found considerable individual variation in the amount and strength of antipredator responses. Although almost half of the charr failed to show antipredator behavior towards the piscivores, those with the innate ability showed highly sensitive recognition of predator odors. Our results indicate that the innate antipredator behavior of the juvenile fish is already finely tuned to respond specifically to chemical cues from different fish predators and even their diets.Communicated by J. Krause     

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     

A nesting association between semi-colonial Bullock's orioles and yellow-billed magpies: evidence for the predator protection hypothesis

Mixed-species nesting associations have been reported for many birds. However, few studies have shown that one species specifically chooses to nest close to another in order to obtain a direct benefit and only two studies have managed to adequately test the predator protection hypothesis for such associations. Here we demonstrate a nesting association between Bullock's orioles (Icterus galbula bullockii) and yellow-billed magpies (Pica nuttalli). The evidence indicates that this is an active association in which, given a choice of suitable habitat areas in which to nest, orioles choose those areas that contain magpie nests. Oriole nests associated with magpie nests have a significantly reduced predation rate compared to those without associated magpie nests. This suggests that orioles prefer to nest near to magpies and that this preference has evolved due to the protection provided by neighbouring magpies. The nesting association results in the clumping of oriole nests around magpie nests, and therefore provides a possible explanation for the semi-colonial nature of Bullock's orioles. Received: 23 November 1998 / Received in revised form: 25 May 1999 / Accepted: 30 May 1999     

Less is more: social learning of predator recognition requires a low demonstrator to observer ratio in Arctic charr (Salvelinus alpinus)

Reintroduction programs for conservation purposes suffer from low post-release survival of captive-bred animals, especially so with fish reintroductions. High mortality is most often due to weakened antipredator skills, which could be enhanced through direct experience with predators and social learning. As yet, the relative efficiency of these learning methods has not been tested, and the significance of the factors determining how efficiently antipredator responses are transmitted socially from experienced (demonstrator) to predator naive (observer) individuals is largely unknown. Using hatchery-bred stock of the endangered Saimaa Arctic charr (Salvelinus alpinus), we show for the first time that: (1) contrary to theoretical predictions, acquired predator recognition was only socially transmitted from predator-experienced to predator-naive fish in groups with a smaller proportion of experienced than naive individuals, and that (2) socially acquired avoidance of the predator odors was as strong as in those fish conditioned with a combination of odors and predator exposure. We conclude that using social transmission procedures in training fish for reintroduction programs may provide considerable ecological, economical and ethical advantages.Communicated by J. Krause     

Hunger-dependent predator inspection and foraging behaviours in the threespine stickleback (Gasterosteus aculeatus) under predation risk

Individual fish commonly leave the relative safety of the shoal to approach potential predators at a distance. Not all members of a shoal are equally likely to initiate such predator inspection visits. Here, we show for the first time that the current hunger state of individual fish strongly influences their predator inspection behaviour, as well as their foraging rate, in the face of predation hazard. When all members of threespine stickleback (Gasterosteus aculeatus) test shoals were in a similar hunger state, they were equally likely to inspect a trout predator model alone and did not differ in the frequency of their inspection visits or foraging rate. However, when individual sticklebacks in a shoal differed in their hunger state, the food-deprived (i.e. hungrier) member of the shoal fed at a higher rate, was significantly more likely to initiate solitary predator inspection visits, and inspected the predator model significantly more often than its less hungry (i.e. well-fed) shoal mates. Individual fish which inspected the predator model more frequently also tended to have higher feeding rates. The results indicate that the hungrier fish in a shoal are more willing to take greater risks to inspect a potential threat at a distance, compared with their well-fed shoal mates, and suggest that they may gain a foraging benefit in doing so. If marked asymmetries in hunger state exist among members of fish shoals, then mutual cooperation during predator inspection visits may be difficult to achieve because well-fed individuals are not as likely to initiate or participate in inspection visits as are hungry individuals.Correspondence to: J.-G.J. Godin     

Social dominance in migrant and resident turkey vultures at carcasses: evidence for a despotic distribution?

We placed carcasses in three different vegetation types in the heterogeneous savannas of central Venezuela to investigate the role of social dominance in habitat use by flocking migrant and resident turkey vultures (Cathartes aura meridionalis and C. a. ruficollis). Migrants foraged primarily in savanna habitats while residents foraged almost exclusively in gallery forest. In the gallery forest residents discovered carrion first significantly more often than migrants, despite there being equal densities of residents and migrants foraging over this habitat. Because residents fed in smaller groups than migrants at carcasses they had higher feeding rates. There was also a negative relationship between group sizes of residents and migrants. The feeding rate of residents declined in response to increased group size of migrants, but group size of residents had no effect. Migrant group size also had a greater effect on resident feeding rates than king vulture presence or absence. When the effect of migrant and resident group size on feeding rates in migrants was compared, the most significant factor was migrant group size. A second analysis showed that both resident group size and presence or absence of king vultures had a significant effect on feeding rates in migrants. Rates of agonistic encounters in migrant and resident turkey vultures increased weakly in relation to group size. However, there was an increase in residents' encounter rate with migrants in relation to increased migrant group size; there was no difference in resident encounter rates with other residents in relation to resident group size. Migrants dominated residents in almost all agonistic interactions over carcasses. We suggest that savanna habitats were less attractive to residents for foraging because they held larger groups of migrants.     

米埔湿地冬季鸟类对气象及生态质量响应研究

为探讨气象及生态气象质量与湿地生态系统内鸟类变化的关系,文章以香港米埔湿地为例,利用米铺湿地1979—2009年留鸟人工观测数据、香港天文台公布气象数据和以2000—2009年4期等时间间距研究区的景观变化数据,分析湿地内留鸟数量对研究区气象及生态气象质量变化的响应。结果表明：（1）湿地留鸟数量与温度和最低能见度时数相关性显著,与降水量和月均日照时数变化相关性较小;（2）5种生态气象评价指数（EMI）与同期鸟类数量的相关性分析表明,留鸟与综合生态评价指数（ECAI）呈正相关。其中与水体指数（WDI）正相关系数最大,与土地利用退化指数（LDI）负相关系数最大,相关系数分别为0.88和-0.85;而与湿润指数（HI）和植被指数（VDI）相关性较小。研究表明,保护生态环境,调节气候变化幅度,对保护留鸟的生存空间,延续生物的生态平衡具有重要意义。     

Sensory complementation and the acquisition of predator recognition by salmonid fishes

Following disturbance, some aquatic prey species release chemicals that act as a warning cue and increase vigilance in nearby conspecifics. Such disturbance cues evoke consistent low intensity anti-predator responses. In contrast, alarm cues from injured conspecifics often evoke stronger intensity responses in prey animals. In this study, we test the sensory complement hypothesis, which suggests that multiple cues act in an additive or synergistic fashion to provide additional information for risk assessment by prey. In the first experiment, we showed that juvenile rainbow trout pre-exposed to disturbance cues respond to a given concentration of damage-released alarm cues with a higher intensity of response than the trout that were pre-exposed to cues from undisturbed conspecifics. The two cues acted in an additive fashion. In the second experiment, we demonstrated that disturbance cues alone were not enough to elicit a conditioned response to the odour of a novel predator. We also showed that while disturbance cues elicit an increase in the response of trout to alarm cues, this increase does not translate into a stronger learned response to the predator when the predator odour is paired with alarm cues. Future studies should take into account sensory complementation to avoid underestimating the responses of prey to predators.     

Migratory orientation of juvenile yellow-rumped warblers (Dendroica coronata) following stopover: sources of variation and the importance of geographic origins

Birds migrating along coastlines may be at increased risk if displacement occurs toward open-ocean. Eastern North America experiences prevailing northwesterly winds during autumn, which could compel some migrants to drift eastward. Therefore, migrants at stop-over sites along this route may be a mixture of on- and off-course individuals. We assessed whether orientation behavior of juvenile yellow-rumped warblers (Dendroica coronata) captured at a stop-over site in southern Nova Scotia was related to where they originated from (i.e., likely on/off-course). We hypothesized three scenarios after displacement: 1) continued orientation in the migratory direction selected before displacement, 2) orientation from the new location toward the previous destination, or 3) correction to regain the original pathway. Using stable isotopes, we determined that stop-over migrants originated from nearby areas (and assumed on-course) and as far away as western Hudson Bay, over 1,600 km northwest (and assumed off-course) of the site. We used video-based orientation registration cages to determine an individuals’ migratory orientation. Because numerous factors influence migratory orientation (e.g., fuel reserves, celestial cues), we simultaneously assessed the influence of body condition and cloud cover, in addition to geographic origin, on orientation behavior. Individuals that originated closer to the site tended to orient more southwesterly. Orientation directions became increasingly more west-northwesterly the further away an individual originated from the site (i.e., the more likely it was to have been displaced). The result is most consistent with scenario three: individuals from northwest origins likely respond to easterly displacement by orienting westerly to reestablish their previous migratory route.     

Cooperative breeding and immunity: a comparative study of PHA response in African birds

Cooperatively breeding birds might be expected to suffer from higher costs of parasitism than pair-breeding species because of two aspects of their ecology which should facilitate horizontal transmission and possibly select for higher parasite virulence: first, they interact regularly with more individuals than pair-breeding species, and second, these individuals are commonly close relatives that could share similar resistance alleles. This hypothesis predicts that cooperative breeders should invest relatively more in immune defence than closely related species which breed in pairs. I tested this prediction comparatively in African birds by examining the response of the immune system to the mitogenic lectin, phytoahemagglutinin (PHA response) in relation to cooperative breeding. Among 66 species, of which 18 breed cooperatively, PHA response was significantly higher in cooperatively breeding species. This association appeared not to be confounded by body size, clutch size, nest position, coloniality or similarity owing to common phylogenetic descent. These results suggest that cooperatively breeding birds may have been selected to invest more than pair-breeders in defences against parasites. If so, then additional costs of philopatry and helping behaviour might be imposed on breeders, helpers and offspring.     

A Comparison of Logging Systems and Bat Diversity in the Neotropics

Abstract:  Evaluating logging systems to determine which are most compatible with the maintenance of biodiversity is of prime importance if tropical forests are to be managed in a sustainable way. Bats are model taxa for this purpose. Two different logging systems are used in the natural forest of the Victoria-Mayaro Forest Reserve in Trinidad: open range and periodic block. Open range is a continuous harvesting system and, in common with most methods used to log tropical forests worldwide, has few harvest controls other than girth limits for selected species. Periodic block is a polycyclic system, with felling based on ecological criteria assumed to be compatible with the maintenance of biodiversity. To compare the effects of periodic block and open-range systems on biodiversity, we determined bat species richness and abundance in each system and in primary forest. We caught bats in mist nets set at ground level and in the canopy and in harp traps. In total 1959 individuals representing 38 species were captured. Species richness was similar among primary forest and logged forest habitats, although bat diversity was lower in logged forest. The distributions of bat species abundance did not differ significantly between logged forest and primary forest. We found, however, that both logging systems lead to a decrease in gleaning animalivores and an increase in frugivores. The increase in frugivores was likely the result of an increase in the abundance of bat-dispersed pioneer fruiting plants in logged forest. Bats of periodic-block-managed forest were more similar to those of primary forest than those of forest logged using the open-range system, indicating that the periodic-block system is more compatible with the maintenance of bat diversity. Our results support the suggestion that the measured use of tropical forests can largely be compatible with biodiversity conservation.     

Antipredator responses of California ground squirrels to rattlesnakes and rattling sounds: the roles of sex,reproductive parity,and offspring age in assessment and decision-making rules

California ground squirrels (Spermophilus beecheyi) and northern Pacific rattlesnakes (Crotalus viridus oreganus) have an adversarial relationship. Adults are partially protected by venom resistance and harass rattlesnakes in part to defend their more vulnerable offspring. Larger, warmer snakes are more dangerous than smaller colder snakes, and in escalated conflict squirrels could benefit from risk assessment strategies. Rattlesnakes often rattle at harassing squirrels and rattling sounds produce cues related to body size and temperature. In study 1 we played back rattling sounds from snakes that varied in dangerousness and evaluated the roles of sex and parity in squirrel risk assessment strategies. In general, squirrels tail flagged and stood bipedally more, and were slower to reapproach the playback speaker following playbacks of rattling sounds from more dangerous snakes. In comparison with males and nonmothers, mothers were most responsive to rattling sounds and more sensitive to variation in snake dangerousness. Mothers tail flagged more than males and nonmothers, and this behavior tracked variation in snake dangerousness most closely, perhaps reflecting the effects of snake size and temperature on pup vulnerability. These findings suggest that many aspects of squirrel antisnake behavior are governed by their effects on descendant kin. In study 2 we tested the effects of offspring age on mothers responses to live rattlesnakes and rattling sounds. According to the offspring value hypothesis, mothers should take more risks in defense of older offspring because they are more likely to survive to reproductive age. By contrast, under the offspring vulnerability hypothesis, older offspring are less vulnerable to predators and thus mothers should take fewer risks. Risk-taking, as measured by behaviors that bring the squirrel close to the snakes strike range, was either unaffected by or negatively correlated with offspring age. Thus, our findings suggest that whereas offspring value is unimportant in squirrel antisnake behavior, offspring vulnerability may affect maternal defense. We suggest that offspring vulnerability in mammals, in comparison with birds, may play a larger role in parental defense against predators.Communicated by P.A. BednekoffAn erratum to this article can be found at     

Learning to recognize novel predators under weakly acidic conditions: the effects of reduced pH on acquired predator recognition by juvenile rainbow trout

Summary. Recent studies have demonstrated that under weakly acidic conditions (pH 6.0), many prey fishes, including juvenile rainbow trout (Onchorhynchus mykiss), do not exhibit overt antipredator responses to conspecific chemical alarm cues. In laboratory trials, we investigated the potential effects of reduced pH on the ability of hatchery reared, predator naïve juvenile rainbow trout to acquire the recognition of a novel predator (yellow perch, Perca flavenscens). Initially, we exposed trout to the odour of a predatory yellow perch, buffered to pH 6.0 (weakly acidic) or pH 7.0 (neutral) paired with conspecific skin extracts (also buffered to pH 6.0 or 7.0) or a distilled water control. Juvenile trout exhibited significant increase in antipredator behaviour when exposed to neutral skin extract (pH 7.0). When retested 48 hours later to perch odour alone (pH 7.0), only trout initially conditioned with neutral skin extracts (pairs with either neutral or acidic perch odour) exhibited a learned recognition of perch odour as a predator risk. Those initially exposed to weakly acidic skin extract or the distilled water control did not show a learned response to predator odour. These results demonstrate that the ability to acquire the recognition of novel predators is impaired under weakly acidic conditions, as would occur in natural waterways affected by acidic precipitation.     

A trade-off between prey- and predator-induced polyphenisms in larvae of the salamander Hynobius retardatus

Organisms in natural habitats participate in complex ecological interactions that include competition, predation, and foraging. Under natural aquatic environmental conditions, amphibian larvae can simultaneously receive multiple signals from conspecifics, predators, and prey, implying that predator-induced morphological defenses can occur in prey and that prey-induced offensive morphological traits may develop in predators. Although multiple adaptive plasticity, such as inducible defenses and inducible offensive traits, can be expected to have not only ecological but also evolutionary implications, few empirical studies report on species having such plasticity. The broad-headed larval morph of Hynobius retardatus, which is induced by crowding with heterospecific anuran (Rana pirica) larvae, is a representative example of prey-induced polyphenism. The morph is one of two distinct morphs that have been identified in this species; the other is the typical morph. In this paper, we report that typical larval morphs of Hynobius can respond rapidly to a predatory environment and show conspicuous predator-induced plasticity of larval tail depth, but that broad-headed morphs cannot respond similarly to a predation threat. Our findings support the hypothesis that induction or maintenance of adaptive plasticity (e.g., predator-induced polyphenism) trades off against other adaptive plastic responses (e.g., prey-induced polyphenism). For a species to retain both an ability to forage for larger prey and an ability to more effectively resist predation makes sense in light of the range of environments that many salamander larvae experience in nature. Our results suggest that the salamander larvae clearly discriminate between cues from prey and those from predators and accurately respond to each cue; that is, they adjust their phenotype to the current environment.     

Seasonal changes in the diet and feeding behaviour of a top predator indicate a flexible response to deteriorating oceanographic conditions

Shifts in the diet of top predators can be linked to changes in environmental conditions. In this study, we tested relationships between environmental variation and seasonal changes in diet of a top predator, the grey-headed albatross Thalassarche chrysostoma, breeding at Bird Island, South Georgia in an austral summer of 1999/2000. Oceanographic conditions in that year around South Georgia were abnormal (i.e. anomalously high sea surface temperature to a relative 19-year long-term mean). The diet of grey-headed albatrosses showed high seasonal variation, shifting from cephalopods (42.9 % by mass) in late February to Antarctic krill Euphausia superba (58.3 %) in late April, and grey-headed albatrosses breeding performance was low (16.8 %). This study shows these albatrosses did not manage to find sufficient alternative prey and highlight the risk to top predators if there is an increase in the frequency or severity of food shortages in Antarctic waters.     

Escape take-off strategies in birds: the significance of protective cover

Avian escape strategy is highly dependent on the ability to fly, and the success rate of raptor attacks is reduced if the prey gets fully airborne. Therefore, when escaping from predator attacks, the initial take-off is crucial and a rapid take-off, high velocity, and high angle of ascent maximize the chance of survival. However, due to the laws of gravity, birds face a trade-off between maximizing its linear acceleration and maximizing its rate of climb when taking off. The optimal policy between velocity and angle of ascent when a bird escapes from an attacking predator might depend on the detailed nature of the predator’s attack, the proximity to cover, and the presence of conspecifics. Many small birds rely on a quick dash to protective cover. In this study, we examine how the availability of protective cover affects take-off strategy in birds. Male great tits (Parus major) were subjected to a surprise attack by a model predator either in the presence or absence of protective cover. When attacked by the predator, great tits took off and perched in the cover when it was available. Birds subjected to the predator attack in the absence of cover took off in 10° higher angle of ascent and flew faster in the start of the take-off, compared to birds that took off in the presence of cover. Thus, this study for the first time shows that a bird do trade off speed with angle of ascent in an adaptive way, depending on the presence of protective cover.