Estimation of roe deerCapreolus capreolus and mouflonOvis aries densities, abundance and habitat use in a mountainous Mediterranean area

The abundance, density, and habitat use of roe deerCapreolus capreolus (Linnaeus, 1758) and mouflonOvis aries Linnaeus, 1758 were studied in a confined Mediterranean area in Greece with a dung survey based on the faecal accumulation rate (FAR) technique. Estimated density was modelled with generalized additive models using altitude, habitat type, and slope as potential covariates. Model selection among the set of candidate models was conducted based on their generalized cross-validation score. Roe deer had an estimated mean density of 13.9 ind./km² and the best model included slope and habitat type as covariates. The mean density of mouflon in the study area was 22.1 ind./km² and the best model used altitude and habitat type as covariates. For both species, the highest densities were encountered in abandoned cultivations and glades, followed by conifer forests, while the lowest densities were observed in maquis. However, use of open habitats by mouflon was much greater than it was for roe deer. The strong preference of mouflon (a grazer species) for open habitats that were abundant with grasses probably reflected food availability and contrasted with the more diverse habitat use by roe deer (a selective browser).     

A meta-analysis of predation risk effects on pollinator behaviour

Flower-visiting animals are constantly under predation risk when foraging and hence might be expected to evolve behavioural adaptations to avoid predators. We reviewed the available published and unpublished data to assess the overall effects of predators on pollinator behaviour and to examine sources of variation in these effects. The results of our meta-analysis showed that predation risk significantly decreased flower visitation rates (by 36%) and time spent on flowers (by 51%) by pollinators. The strength of the predator effects depended neither on predator taxa and foraging mode (sit-and-wait or active hunters) nor on pollinator lifestyle (social vs. solitary). However, predator effects differed among pollinator taxa: predator presence reduced flower visitation rates and time spent on flowers by Squamata, Lepidoptera and Hymenoptera, but not by Diptera. Furthermore, larger pollinators showed weaker responses to predation risk, probably because they are more difficult to capture. Presence of live crab spiders on flowers had weaker effects on pollinator behaviour than presence of dead or artificial crab spiders or other objects (e.g. dead bees, spheres), suggesting that predator crypsis may be effective to some extent. These results add to a growing consensus on the importance of considering both predator and pollinator characteristics from a community perspective.     

Elevated predation risk changes mating behaviour and courtship in a fiddler crab

The fiddler crab, Uca beebei, lives in individually defended burrows, in mixed-sex colonies on intertidal mud flats. Avian predation is common, especially of crabs unable to escape into burrows. Mating pairs form in two ways. Females either mate on the surface at their burrow entrance (''surface mating'') or leave their own burrow and sequentially enter and leave (''sample'') courting males'' burrows, before staying in one to mate underground (''burrow mating''). We tested whether perceived predation risk affects the relative frequency of these mating modes. We first observed mating under natural levels of predation during one biweekly, semi-lunar cycle. We then experimentally increased the perceived predation risk by attracting grackles (Quiscalus mexicanus) to each half of the study site in two successive biweekly cycles. In each experimental cycle, crabs were significantly less likely to mate on the side with more birds. Moreover, on the side with elevated predation risk, the number of females leaving burrows to sample was greatly reduced relative to the number of females that surface-mated. Males waved less and built fewer mud pillars, which attract females, when birds were present. We discuss several plausible proximate explanations for these results and the effect of changes in predation regime on sexual selection.     

Feeding behaviour in a population of domestic fowls in the wild

In 1976 the diet of adult feral fowls on a Scottish island consisted mainly of grass from January to July and oats from August to December. Juveniles ate many invertebrates in their first two months of life, but otherwise their diet was similar to that of the adults. In contrast to the non-breeding adults, broods and their dams showed two distinct types of feeding behaviour, called “sporadic” and “intensive” feeding. These were directed towards scattered and concentrated food sources, and the type shown depended mainly on the dam's behaviour. The significance of these and other aspects of feeding behaviour is discussed.     

Predator behaviour and predation risk in the heterogeneous Arctic environment

1. Habitat heterogeneity and predator behaviour can strongly affect predator-prey interactions but these factors are rarely considered simultaneously, especially when systems encompass multiple predators and prey. 2. In the Arctic, greater snow geese Anser caerulescens atlanticus L. nest in two structurally different habitats: wetlands that form intricate networks of water channels, and mesic tundra where such obstacles are absent. In this heterogeneous environment, goose eggs are exposed to two types of predators: the arctic fox Vulpes lagopus L. and a diversity of avian predators. We hypothesized that, contrary to birds, the hunting ability of foxes would be impaired by the structurally complex wetland habitat, resulting in a lower predation risk for goose eggs. 3. In addition, lemmings, the main prey of foxes, show strong population cycles. We thus further examined how their fluctuations influenced the interaction between habitat heterogeneity and fox predation on goose eggs. 4. An experimental approach with artificial nests suggested that foxes were faster than avian predators to find unattended goose nests in mesic tundra whereas the reverse was true in wetlands. Foxes spent 3.5 times more time between consecutive attacks on real goose nests in wetlands than in mesic tundra. Their attacks on goose nests were also half as successful in wetlands than in mesic tundra whereas no difference was found for avian predators. 5. Nesting success in wetlands (65%) was higher than in mesic tundra (56%) but the difference between habitats increased during lemming crashes (15%) compared to other phases of the cycle (5%). Nests located at the edge of wetland patches were also less successful than central ones, suggesting a gradient in accessibility of goose nests in wetlands for foxes. 6. Our study shows that the structural complexity of wetlands decreases predation risk from foxes but not avian predators in arctic-nesting birds. Our results also demonstrate that cyclic lemming populations indirectly alter the spatial distribution of productive nests due to a complex interaction between habitat structure, prey-switching and foraging success of foxes.     

Fine-scale population structure, inbreeding risk and avoidance in a wild insect population

The ecological and evolutionary importance of fine-scale genetic structure within populations is increasingly appreciated. However, available data are largely restricted to wild vertebrates and eusocial insects. In addition, there is the expectation that most insects tend to have such large- and high-density populations and are so mobile that they are unlikely to face inbreeding risks through fine-scale population structuring. This has made the growing body of evidence for inbreeding avoidance in insects and its implication in mating systems evolution somewhat enigmatic. We present a 4-year study of a natural population of field crickets. Using detailed video monitoring combined with genotyping, we track the movement of all adults within the population and investigate genetic structure at a fine scale. We find some evidence for relatives being found in closer proximity, both across generations and within a single breeding season. Whilst incestuous matings are not avoided, population inbreeding is low, suggesting that mating is close to random and the limited fine-scale structure does not create significant inbreeding risk. Hence, there is little evidence for selective pressures associated with the evolution of inbreeding avoidance mechanisms in a closely related species.     

Effects of predation risk on population variation in adult size in a stream-dwelling isopod

I used a combination of laboratory experiments and field surveys to examine the role that population-specific predation risk may play in shaping the life history strategy of a stream-dwelling isopod Lirceus fontinalis. Two focal populations were identified that were exposed to different predator types. The first population was exposed to larvae of the streamside salamander (Ambystoma barbouri) and the second to banded sculpin (Cottus carolinae). A laboratory experiment, in which different size classes of prey were offered simultaneously to individual predators, revealed that L. fontinalis suffered greatest mortality risk at small sizes with A. barbouri. Alternatively, with C. carolinae the risk of mortality was independent of size. Life history theory predicts that L. fontinalis from populations exposed to the gape-limited salamander larvae should be larger at maturity relative to individuals from populations exposed to C. carolinae. Field surveys on the two focal populations both within 1 year and across 4 years supported this prediction. Four other populations, two exposed to streamside salamander larvae and two to fish, provided additional support for the prediction. I concluded that L. fontinalis exhibited an adaptive response in size at maturity in response to population-specific predation risk. I then used gut content assays of the major predators to assess whether the population-specific life history strategies adopted by L. fontinalis were successful in avoiding predation.     

Variation in predation risk and vole feeding behaviour: a field test of the risk allocation hypothesis

Many prey animals experience temporal variation in the risk of predation and therefore face the problem of allocating their time between antipredator efforts and other activities like feeding and breeding. We investigated time allocation of prey animals that balanced predation risk and feeding opportunities. The predation risk allocation hypothesis predicts that animals should forage more in low- than in high-risk situations and that this difference should increase with an increasing attack ratio (i.e. difference between low- and high-risk situations) and proportion of time spent at high risk. To test these predictions we conducted a field test using bank voles (Clethrionomys glareolus) as a prey and the least weasel (Mustela nivalis nivalis) as a predator. The temporal pattern and intensity of predation risk were manipulated in large outdoor enclosures and the foraging effort and patch use of voles were measured by recording giving-up densities. We did not observe any variation in feeding effort due to changes in the level of risk or the proportion of time spent under high-risk conditions. The only significant effect was found when the attack ratio was altered: the foraging effort of voles was higher in the treatment with a low attack ratio than in the treatment with a high attack ratio. Thus the results did not support the predation risk allocation hypothesis and we question the applicability of the hypothesis to our study system. We argue that the deviation between the observed pattern of feeding behaviour of bank voles and that predicted by the predation risk allocation hypothesis was mostly due to the inability of voles to accurately assess the changes in the level of risk. However, we also emphasise the difficulties of testing hypotheses under outdoor conditions and with mammals capable of flexible behavioural patterns.     

Assessing predation risk: optimal behaviour and rules of thumb

We look at a simple model in which an animal makes behavioural decisions over time in an environment in which all parameters are known to the animal except predation risk. In the model there is a trade-off between gaining information about predation risk and anti-predator behaviour. All predator attacks lead to death for the prey, so that the prey learns about predation risk by virtue of the fact that it is still alive. We show that it is not usually optimal to behave as if the current unbiased estimate of the predation risk is its true value. We consider two different ways to model reproduction; in the first scenario the animal reproduces throughout its life until it dies, and in the second scenario expected reproductive success depends on the level of energy reserves the animal has gained by some point in time. For both of these scenarios we find results on the form of the optimal strategy and give numerical examples which compare optimal behaviour with behaviour under simple rules of thumb. The numerical examples suggest that the value of the optimal strategy over the rules of thumb is greatest when there is little current information about predation risk, learning is not too costly in terms of predation, and it is energetically advantageous to learn about predation. We find that for the model and parameters investigated, a very simple rule of thumb such as 'use the best constant control' performs well.     

Threat-sensitive anti-intraguild predation behaviour: maternal strategies to reduce offspring predation risk in mites

Predation is a major selective force for the evolution of behavioural characteristics of prey. Predation among consumers competing for food is termed intraguild predation (IGP). From the perspective of individual prey, IGP differs from classical predation in the likelihood of occurrence because IG prey is usually more rarely encountered and less profitable because it is more difficult to handle than classical prey. It is not known whether IGP is a sufficiently strong force to evolve interspecific threat sensitivity in antipredation behaviours, as is known from classical predation, and if so whether such behaviours are innate or learned. We examined interspecific threat sensitivity in antipredation in a guild of predatory mite species differing in adaptation to the shared spider mite prey (i.e. Phytoseiulus persimilis, Neoseiulus californicus and Amblyseius andersoni). We first ranked the players in this guild according to the IGP risk posed to each other: A. andersoni was the strongest IG predator; P. persimilis was the weakest. Then, we assessed the influence of relative IGP risk and experience on maternal strategies to reduce offspring IGP risk: A. andersoni was insensitive to IGP risk. Threat sensitivity in oviposition site selection was induced by experience in P. persimilis but occurred independently of experience in N. californicus. Irrespective of experience, P. persimilis laid fewer eggs in choice situations with the high- rather than low-risk IG predator. Our study suggests that, similar to classical predation, IGP may select for sophisticated innate and learned interspecific threat-sensitive antipredation responses. We argue that such responses may promote the coexistence of IG predators and prey.     

Clouded leopard predation on a wild juvenile siamang

I report the first documented case of a wild siamang killed by a clouded leopard. The event, involving a juvenile male (weighing 3.7 kg), occurred at dawn, near a known sleeping tree in a lowland tropical forest in southern Sumatra (Indonesia). Examination of the siamang's body revealed that the first 2 cervical vertebrae were crushed and the face was partially eaten. This attack highlights the potential importance of predation as an evolutionary pressure for hylobatids, and poses some questions about the antipredator adaptations of this population of siamang.     

鼠类对山杏种子存活和萌发的影响

山杏（Prunus armeniaca)是广布于北京山区阳坡的矮林或灌丛,易生存在土壤贫脊、干旱、或严重退化的山坡、裸地,对于保护生态环境,减少水土流失及促进山区经济发展均具有重要作用。有关鼠类在山杏更新中作用的研究较少,本研究主要是确定影响山杏更新的鼠类种类、鼠类对山杏种子的取食压力、人工埋藏种子及植被因素对种子存活和萌发的影响。于1997年10月3－4日,采用夹捕法同时调查取食花生和山杏种子的鼠类种类及种群密度。于1997年9月24－25日,通过在地表放置山桔种子,调查在鼠类取食下,地表种子消失的速度。于1997年10月20日,作种子埋藏实验,调查种子存活及萌发情况。于1997年10月26日,进行扣网实验,研究网内、外山杏种子存活及萌发情况。结果表明：大林姬鼠（Apodemus speciosus)、社鼠（Rattus confucianus),黑线姬鼠（Apodemus agrarius)为取食山杏种子的主要鼠种种类;与取食花生相比,社鼠更偏爱山杏种子,大林姬鼠对山杏种子喜食性略低,黑线姬鼠对二者喜食程度相近;山杏种子放置地表后1、10、20、30d的消失率分别为13.06%,64.46%,90.70%,96.69%;扣网实验说明,山杏在地表的萌发率极低,生长也慢;而埋入土层内可明显提高萌发率和生长率（若除去鼠类的取食,萌发率还要高）;人工埋藏实验表明,将山杏种子埋入土层5cm后,能够有效地减少鼠类对种子的取食;通过分析山杏种子萌发与植被的关系,发现山杏易在开阔、阳光充分的草丛中存活和生长,而不易在阴闭的灌丛下存活和生长。     

Predator occurrence and perceived predation risk determine grouping behavior in guanaco (Lama guanicoe)

Grouping behavior of social ungulates may depend on both predator occurrence and perceived predation risk associated with habitat structure, reproductive state, and density of conspecifics. Over 3 years, we studied grouping behavior of guanaco (Lama guanicoe) families in Chilean Patagonia during the birthing season and determined their response to variation in predator occurrence and perceived predation risk (habitat structure, calf/adult rate, and density of conspecifics). We considered the effect of two predators, puma (Puma concolor) and culpeo fox (Lycalopex culpaeus). We measured two common (family group size and vigilance) and one novel (family group cohesion) behavioral responses of guanaco. Our results show that guanaco family groups adapted their grouping behavior to both predator occurrence and perceived predation risk. Larger family groups were found in open habitats and areas with high puma occurrence, while guanacos stayed in small family groups in areas with high shrub cover or low visibility. Group cohesion increased in areas with higher occurrence of pumas and culpeo foxes, and also increased in smaller family groups and in areas with low guanaco density. Vigilance (number of vigilant adults) was mainly related to group size and visibility, increasing in areas with low visibility, while residual vigilance (vigilance after removing the group‐size effect) did not vary with the explanatory variables examined. Our results suggest that a mix of predator occurrence and perceived predation risk influences guanaco grouping behavior and highlights the importance of evaluating different antipredator responses together and considering all predator species in studies aimed at understanding ungulate behavior.     

Effect of predation risk on selectivity in heteromyid rodents

Variations in predation risk affect the costs of foraging and may therefore warrant different foraging decisions. One class of models ("higher requisite profit") predicts that foragers should become more selective when predation risk increases, as low-profitability items that do not cover the increased costs are dropped from the diet. An alternative class of models ("reduced finickiness") predicts that foragers should become less selective when predation risk increases, because selectivity requires more extensive assessment and/or search behaviour, prolonging exposure to risk. We assessed the selectivity of foraging heteromyid rodents (Merriam's kangaroo rats, Dipodomys merriami, and pocket mice, Chaetodipus spp.) by comparing differences in "giving up densities" (GUD: the quantity of cryptic food left in a patch by animals for whom the diminishing marginal gains from foraging have dropped below the threshold for continued search) for foods of different value as a measure of selectivity in patches varying in predation risk. Data collected over two field seasons revealed that heteromyids were more selective when predation risk was highest; away from the protective cover of shrubs during the full moon. These findings support the predictions of higher requisite profit models.     

Selection and microevolution of coat pattern are cryptic in a wild population of sheep

Understanding the maintenance of genetic variation in natural populations is a core aim of evolutionary genetics. Insight can be gained by quantifying selection at the level of the genotype, as opposed to the phenotype. Here, we show that in a natural population of Soay sheep which is polymorphic for coat pattern, recessive genetic variants at the causal gene, agouti signalling protein (ASIP) are associated with reduced lifetime fitness. This was due primarily to a reduction in juvenile survival of uniformly coloured (self-type) sheep, which are homozygous recessive, and occurs despite significantly higher reproductive success in surviving self-type adults. Consistent with their relatively low fitness, we show that the frequency of self-type individuals has declined from 1985 to 2008. Remarkably though, the frequency of the underlying self-allele has increased, because the frequency of heterozygous individuals (who harbour the majority of all self-alleles) has increased. Indeed, the ratio of observed/expected heterozygous individuals has increased during the study, such that there is now a significant excess of heterozygotyes. By employing gene-dropping simulations, we show that microevolutionary trends in the frequency and excess of ASIP heterozygotes are too pronounced to be caused by genetic drift. Studying this polymorphism at the level of phenotype rather than underlying genotype would have failed to detect cryptic fitness differences. We would also have been unable to rule out genetic drift as an evolutionary force driving genetic change. This highlights the importance of resolving the underlying genetic basis of phenotypic variation in explaining evolutionary dynamics.     

Spatial distribution of intertidal fishes: a pattern dependent on body size and predation risk?

Traditionally, heterogeneity levels have been associated with refuge availability and, consequently, with security areas for prey. Hence, there is assumed to be a positive correlation between the spatial distribution of potential victims and habitat heterogeneity levels. Among fish, the magnitude of predator-prey interaction has been associated with their size ratio, which includes body size as a central variable for individual predation risk. To date, the importance of body size and its interaction with spatial heterogeneity in relation to fish spatial distribution has been scarcely considered, being this the main objective of the present study. Based on a semi-experimental design, defined by the level of spatial heterogeneity, censuses in 24 artificial pools were conducted for 2 years at the rocky intertidal zone. This study focused on carnivorous fish assemblages residing off the rocky shore of central Chile. The results indicated a counterintuitive response of fish to spatial heterogeneity, with a higher abundance of small fish in open areas. Apparently, this differential distribution could be related to the size ratio between prey and predator (size-dependent susceptibility to predation), which is a hypothesis supported by decreases in the co-occurrence of fish with increasing size similarity.     

Antipredatory behaviour in lizards: interactions between group size and predation risk

Group size effects on antipredatory behaviour are well documented in numerous animals, but little is known about how the level of predation risk influences this process. We tested the hypothesis that group size and level of risk interact to affect the levels of antipredatory behaviour in the group-living sun skink, Lampropholis delicata. We controlled the size of lizard groups (N=1, 2, 4, 8 or 12 females) and altered predation risk by providing either a basking tile covered with chemical cues from a predator (high risk) or one without scent (low risk). The time allocated to individual antipredatory behaviour decreased significantly with increasing group size. The relation between group size and time allocated to individual antipredatory behaviour was nonlinear and asymptotic, and did not change under low and high risks of predation. However, group size and predation risk interacted to affect significantly the time that lizards allocated to antipredatory behaviour. When the overall risk from predators was high, individual responsiveness decreased strongly as group size became larger. In contrast, when the overall risk from predators was low, individual responsiveness decreased weakly as group size became larger. Consequently, the time that lizards allocated to antipredatory behaviour under different risks of predation converged as group size increased.     

Interacting effects of predation risk and food availability on larval anuran behaviour and development

Age and size at metamorphosis are two important fitness components in species with complex life cycles. In anurans, metamorphic traits show remarkable phenotypic plasticity, especially in response to changes in growth conditions. It is also possible that the perception of risk directly determines changes in larval period and the size of metamorphs. This study examines how the perception of predation risk affects the timing of and size at metamorphosis in common frogs (Rana temporaria). I raised tadpoles at two risk levels (fish-conditioned water or unconditioned water) crossed with the availability or lack of food at night (all tadpoles had food available in the day). Tadpoles reacted to chemical cues from predatory fish by decreasing activity. A novel behavioural result was a predation×food interaction effect on refuge use, which also accounted for most of the predator main effect: predation risk only caused increased refuge use in the night-starved treatment. Despite these behavioural modifications, the perception of predation risk did not affect growth rate and mass at metamorphosis in a simple way: the effects of food regime on growth and size at metamorphosis were dependent on the level of predation risk as revealed by significant predation×food interaction effects. Tadpoles who had food withheld at night metamorphosed at the smallest size, suggesting a negative relationship between size at metamorphosis and refuge use. Tadpoles raised in fish-conditioned water had longer larval periods than those in unconditioned water, but these differences were significant only if food was available at night. These results conflict with the hypotheses that tadpoles should reduce their larval period or growth rates (and hence metamorphose at a smaller size) as the risk of predation increases. In contrast to predation risk, food availability strongly affected the length of the larval period: night-starved tadpoles metamorphosed relatively early with or without fish stimulus. Thus, early metamorphosis resulted from periods of low food availability, but not from a heightened ”perceived risk” of predation. This example counters the hypothesis of acceleration of the developmental rate (which shortens the time to metamorphosis) as a mechanism to escape a risky environment. Received: 18 August 1999 / Accepted: 10 January 2000