The influence of hunger and predation risk on group cohesion in a pelagic fish, walleye pollock Theragra chalcogramma

Variation in the intensity of schooling behavior in fishes suggests that the benefits of aggregation are balanced by certain costs. We examined the proximity of group members to each other in juvenile walleye pollock, Theragra chalcogramma, under different environmental conditions. Food availability, simulated by a gradient of six ration treatments, had a major influence on group cohesion, with increasing dispersion as food level decreased. Group cohesion also decreased at night relative to daytime levels. Small juveniles (x=53 mm TL) maintained on high rations were highly responsive to the potential threat of a predator, with groups becoming more cohesive and remaining so for up to an hour after the initial threat. A chronic threat (continual presence of predators) resulted in tighter group cohesion than an acute threat (single simulated attack). Small juveniles maintained on low rations were less responsive to predation threats and recovered quickly, supporting the hypothesis that hunger induces risk-taking behavior. Large juveniles (x=149 mm TL) did not change their degree of aggregation in response to either type of predation threat. An overall plasticity in the degree of cohesiveness among group members indicates that walleye pollock are capable of gradually modifying their schooling behavior according to the environmental context.     

Nesting and post-fledging predation risk influence diel patterns of songbird fledging

Among stages of avian ontogeny, the act of nest departure or fledging is an abrupt transition into a new environment and a major leap toward independence for offspring. In altricial birds, the timing (i.e. time of day) of fledging is notable in that many species tend to fledge early in the morning. Past studies have proposed nest predation as a key factor driving birds to fledge earlier in the morning (the ‘survival hypothesis’), whereby offspring avoid peak times of nest predation that occur later in the day. A natural extension of this hypothesis is the predation of offspring post-fledging, whereby offspring are also timing their fledging with future survival prospects outside of the nest. However, few studies have investigated fledging behaviour in the context of both nesting and post-fledging predation. To help fill this knowledge gap, we investigated factors driving the timing and duration of fledging across six songbird species in the context of offspring predation: daily nest mortality, post-fledging mortality and diel patterns of nest predation risk. We found that > 60% of songbirds fledged early in the morning, whereas the peaks in nest predation risk occurred several hours post-fledging. Furthermore, species under greater risk of nest predation fledged earlier in the day and in closer succession to their siblings. Parameters of post-fledging mortality were poor predictors of fledging timing, but individuals from broods of species under higher risk of post-fledging mortality fledged in closer succession to their siblings. These results provide evidence in support of the survival hypothesis, and suggest that songbirds fledge in the morning to avoid peak times of nest predation risk that occur later in the day (~ 8 h after civil dawn). Such results corroborate past research highlighting predation on dependent offspring as a key factor driving variation in life histories across animal taxa; however, estimates of post-fledging mortality suggest that nest predation alone does not fully explain variation in fledging behaviour among species. Future research is therefore needed to investigate the contribution of other factors, such as energetics, parent–offspring conflict and diel patterns of post-fledging survival, which may help to mediate diel patterns of fledging within and among songbird species.     

Alterations in the grazing activities of cased caddisfly larvae in response to variations in predation risk and resource level

Flexible behavioral response to avoid encountering predators has never been reported in lotic, cased caddisfly larvae with low mobility. However, an earlier laboratory experiment found the growth of such a caddisfly species, Glossosoma sp., decreased in the presence of a predatory sculpin, Cottus nozawae. We conducted laboratory experiments to test whether Glossosoma responses to sculpin varied according to resource level and/or time of day. Lower periphyton biomass resulted in increased movement at any time of day, but was followed by decreased grazing success of Glossosoma in the morning when they were sampled. Although sculpins were active in the morning, evening and night but never in the afternoon, movements of Glossosoma were suppressed by sculpin stimuli only in the morning, when food intake of Glossosoma also decreased, regardless of periphyton biomass. Glossosoma reduced the risk of predation by sculpin by flexibly controlling its grazing activities during the most risky period of the day (i.e. morning). Even in the morning in the presence of sculpin, lower resource levels resulted in longer movement distances similar to those in predator-free conditions, suggesting that lower resource availability promotes predation risk in natural streams inhabited by predatory sculpin.     

Parasite and predator risk assessment: nuanced use of olfactory cues

Foraging herbivores face twin threats of predation and parasite infection, but the risk of predation has received much more attention. We evaluated, experimentally, the role of olfactory cues in predator and parasite risk assessment on the foraging behaviour of a population of marked, free-ranging, red-necked wallabies (Macropus rufogriseus). The wallabies adjusted their behaviour according to these olfactory cues. They foraged less, were more vigilant and spent less time at feeders placed in the vicinity of faeces from dogs that had consumed wallaby or kangaroo meat compared with that of dogs feeding on sheep, rabbit or possum meat. Wallabies also showed a species-specific faecal aversion by consuming less food from feeders contaminated with wallaby faeces compared with sympatric kangaroo faeces, whose gastrointestinal parasite fauna differs from that of the wallabies. Combining both parasite and predation cues in a single field experiment revealed that these risks had an additive effect, rather than the wallabies compromising their response to one risk at the expense of the other.     

Cheek pouch use, predation risk, and feeding competition in blue monkeys (Cercopithecus mitis stuhlmanni)

The adaptive function of cheek pouches in the primate subfamily Cercopithecinae remains unresolved. By analyzing the circumstances of cheek pouch use, we tested two hypotheses for the evolution of cercopithecine cheek pouches proposed in earlier studies: (1) cheek pouches reduce vulnerability to predation, and (2) cheek pouches increase feeding efficiency by reducing competition. We studied two groups of wild blue monkeys (Cercopithecus mitis stuhlmanni) in the Kakamega Forest, Kenya, conducting focal observations of feeding individuals. Monkeys were less exposed while emptying their cheek pouches than filling them, supporting the predation-avoidance hypothesis. We investigated several measures of competitive threat, but only one supported the competition-reduction hypothesis: when the nearest neighbor's rank increased, subjects were more likely to increase than to decrease cheek pouch use. Overall, our findings supported the predation-avoidance hypothesis more strongly than the competition-reduction hypothesis. We suggest that variation in cheek pouch use may reflect differing behavioral strategies used by cercopithecines to mitigate competition and predation, as well as factors such as resource size and distribution, home range size, and travel patterns.     

Ontogeny, phylogeny, and morphology in anuran larvae: morphometric analysis of cranial development and evolution in Rana tadpoles (Anura: Ranidae)

Comparative studies of chondrocranial morphology in larval anurans are typically qualitative in nature, focusing primarily on discrete variation or gross differences in the size or shape of individual structures. Detailed data on chondrocranial allometry are currently limited to only two species, Rana sylvatica and Bufo americanus. This study uses geometric morphometric and multivariate statistical analyses to examine interspecific variation in both larval chondrocranial shape and patterns of ontogenetic allometry among six species of Rana. Variation is interpreted within the context of hypothesized phylogenetic relationships among these species. Canonical variates analyses of geometric morphometric datasets indicate that species can be clearly discriminated based on chondrocranial shape, even when whole ontogenies are included in the analysis. Ordinations and cluster analyses based on chondrocranial shape data indicate the presence of three primary groupings (R. sylvatica; R. catesbeiana + R. clamitans; and R. palustris + R. pipiens + R. sphenocephala), and patterns of similarity closely reflect phylogenetic relationships. Analysis of chondrocranial allometry reveals that some patterns are conserved across all species (e.g., most measurements scale with negative allometry, those associated with the posterior palatoquadrate tend to scale with isometry or positive allometry). Ontogenetic scaling along similar allometric trajectories, lateral transpositions of individual trajectories, and variable allometric relationships all contribute to shape differences among species. Overall patterns of similarity among ontogenetic trajectories also strongly reflect phylogenetic relationships. Thus, this study demonstrates a tight link between ontogeny, phylogeny, and morphology, and highlights the importance of including both ontogenetic and phylogenetic data in studies of chondrocranial evolution in larval anurans.     

Density is more important than predation risk for predicting growth and developmental outcomes in tadpoles of spotted tree frog,Litoria spenceri (Dubois 1984)

The diverse benefits of group living include protection against predators through dilution effects and greater group vigilance. However, intraspecific aggregation can decrease developmental rates and survival in prey species. We investigated the impact on tadpole development and behaviour of the interaction between population density and predation risk. Spotted tree frog (Litoria spenceri: Hylidae, Dubois 1984) tadpoles were kept at one of three different densities (two tadpoles per litre, five tadpoles per litre or 10 tadpoles per litre) until metamorphosis in the presence or absence of predatory cues. We aimed to determine the influence of population density, predation and the interaction of both factors in determining growth rates in tadpoles. Tadpoles were measured weekly to assess growth and development and filmed to quantify differences in activity and feeding frequency between groups. Generally, tadpoles housed without predators had longer developmental periods when housed with a predator, but there was no effect on tail length or total length. There was no effect of either predation cues or density on percentage of individuals feeding or moving. Although the effects of the presence of predators alone may appear to be less than the effects of the presence of competitors, the prioritisation of competitiveness over predator avoidance may increase vulnerability of tadpoles to the lethal threat of predators. This is particularly important in species such as L. spenceri, which is at risk from introduced fish predators.     

The effects of flow on schooling Devario aequipinnatus: school structure,startle response and information transmission

To assess how flow affects school structure and threat detection, startle response rates of solitary and small groups of giant danio Devario aequipinnatus to visual looming stimuli were compared in flow and no‐flow conditions. The instantaneous position and heading of each D. aequipinnatus was extracted from high‐speed videos. Behavioural results indicate that (1) school structure is altered in flow such that D. aequipinnatus orient upstream while spanning out in a crosswise direction, (2) the probability of at least one D. aequipinnatus detecting the visual looming stimulus is higher in flow than no flow for both solitary D. aequipinnatus and groups of eight D. aequipinnatus; however, (3) the probability of three or more individuals responding is higher in no flow than in flow. These results indicate a higher probability of stimulus detection in flow but a higher probability of internal transmission of information in no flow. Finally, results were well predicted by a computational model of collective fright response that included the probability of direct detection (based on signal detection theory) and indirect detection (i.e. via interactions between group members) of threatening stimuli. This model provides a new theoretical framework for analysing the collective transfer of information among groups of fishes and other organisms.     

Behavioural activity levels and expression of stress proteins under predation risk in two damselfly species

Abstract 1. It has become apparent that predators may strongly decrease prey fitness without direct contact with the prey, as they induce the development of defence systems that limit the availability of energy for growth and reproduction. Recent studies suggest that stress proteins may help prey organisms deal with this stress. The pattern is not general, however, and little is known about species differences in physiological traits in coping with predator stress, and covariation of physiological with other antipredator traits. 2. To explore these issues, we quantified levels of constitutive and fish‐induced stress proteins (Hsp60 and Hsp70) and anti‐predator behaviours in larvae of two damselfly species that differ in lifestyle. Both stress proteins were fixed at higher levels in Erythromma najas, which has a slow lifestyle, than in Lestes sponsa, which has a fast lifestyle. Similarly, anti‐predator behaviours were fixed at safer levels in E. najas than in L. sponsa. 3. These results suggest that stress proteins may be part of anti‐predator syndromes of damselfly larvae, and there may be trait co‐specialisation between stress proteins and behavioural anti‐predator traits. Studies formally testing these hypotheses in more species may prove rewarding in advancing our understanding of the functional integration of physiological anti‐predator traits in relation to the prey’s lifestyle.     

Effects of body size, age and maturity stage on diet in a large shark: ecological and applied implications

Ontogenetic diet shifts are a widespread phenomenon among vertebrates, although their relationships with life history traits are poorly known. We analyzed the relative importance of body size, age and maturity stage as determinants of the diet of a marine top predator, the copper shark, Carcharhinus brachyurus, by examining stomach contents using a multiple-hypothesis modeling approach. Copper sharks shifted their diet as size and age increased and as they became sexually mature, incorporated larger prey as they grew, and had a discrete shift in diet with body size, with only individuals larger than ≈200 cm total length able to prey on chondrichthyans. Body size was the most important trait explaining the consumption of chondrichthyans, while age determined the consumption of pelagic teleosts. Pelagic teleosts were consumed mostly by medium-aged sharks, a result, probably, of a risk-reducing feeding strategy at young ages coupled with either a senescence-related decline in performance or a change in sensory capabilities as sharks age. Copper sharks of all sizes were able to cut prey in pieces, implying that gape limitation (i.e., the impossibility of eating prey larger than a predator’s mouth) did not play a role in producing the diet shift. Our results suggest that, contrary to the current practice of setting minimum but not maximum size limits in catches, any plan to conserve or restore the ecological function of sharks, through their predatory control of large prey, should aim to maintain the largest individuals.     

Hide, rest or die: a light-mediated diapause response in Daphnia magna to the threat of fish predation

1. In a laboratory batch culture experiment, a diapause response of Daphnia magna to a simulated threat of fish predation was tested at various light intensities, which under natural conditions determine potential vulnerability of Daphnia to visual planktivorous fish. 2. Under moderate light intensity (1.4 μmol m^?2 s^?1) that allows effective predation by fish, the proportion of females producing dormant eggs was significantly higher than under dim light conditions (0.001 μmol m^?2 s^?1) that are not favourable for visual detection of prey. Production of dormant eggs was not observed in complete darkness or in treatments missing fish kairomones, irrespective of tested light conditions. 3. The observed phenomenon is interpreted as a flexible response of prey to the conditional risk of predation assessed by Daphnia according to the presence of fish‐derived cues on the one hand and the presence of dark refugia on the other. Irrespective of the presence of fish kairomones, Daphnia may not produce resting eggs as long as a safe, dark, bottom zone is accessible.     

Functional mechanisms of an inducible defence in tadpoles: morphology and behaviour influence mortality risk from predation

In many amphibian larvae a suite of morphological and behavioural characters varies together in an induced defence against predators, but it remains unclear which features are functionally related to defence. We independently manipulated behaviour and morphology in tadpoles of Hyla versicolor and assessed their consequences for swimming performance and predator escape. Data on burst swimming showed that tadpoles which accelerated rapidly were elongate, with shallow bodies and tails. Predator escape was measured by exposing tadpoles to predators (larval Anax dragonflies or larval Ambystoma salamanders) and recording time until death. Tadpoles were first reared for 30 days in ponds containing either caged Anax or no predators; individuals responded to predators by developing large brightly coloured tails and short bodies. We placed tadpoles of both morphological phenotypes into plastic tubs, and manipulated their behaviour using food and chemical cues from predators. Mortality risk experienced by the predator‐induced phenotype was about half that of the no‐predator phenotype, and risk increased with time spent swimming. An interaction between morphology and behaviour arose because increasing activity caused higher risk for tadpoles with deep tail fins but not shallow tail fins.     

Direct and indirect cues of predation risk influence behavior and reproduction of prey: a case for acarine interactions

Little is known about the flow of chemical information fromhigher to lower levels within the animal food chain. However,this information may determine the behavior and distributionof many animals (e.g., that of potential prey) when exposedto direct and indirect cues of predation risk. We used herbivorousspider mites, Tetranychus urticae Koch (Tetranychidae) as amodel to examine the foraging and oviposition decisions thatprey make when exposed to these cues. We conducted laboratorytests to determine if the previous presence of predators (directcues) on leaf discs or the presence of injured conspecifics(indirect cues) alters the distribution of adults and eggs ofT. urticae. When given a choice, after 24 h, fewer adults and eggswere found on leaf discs that had previously contained specialistspider mite predators, Phytoseiulus persimilis Athias-Henriot (Phytoseiidae),than on discs unexposed to predators. Also, more T. urticaeemigrated from predator-exposed discs than from unexposed discs orfrom those that had previously contained nonpredatory mites(Tyrophagus putrescentiae, Acaridae). Finally, fewer T. urticaeforaged and laid eggs on predator-exposed discs or on thosewith artificially damaged conspecifics (eggs or dead adults)than on discs with intact conspecifics. Tetranychus urticaeprobably recognizes infochemicals (kairomones) from its predatorsor cues from injured spider mites and consequently avoids feedingor ovipositing in areas exposed to these cues. Recognition and avoidanceof kairomones from specialist predators by this prey are likelyto be hereditary, but avoidance of injured conspecifics maybe an adaptation to avoid predators that are not inherentlyrecognized. We discuss the behavioral and ecological implicationsof our findings.     

Contrasting migration behaviour of Daphnia pulicaria and D. galeata x hyalina, in avoidance of predation by 0+perch (Perca fluviatilis)

In spring and early summer, a small population of the large-bodied Daphnia pulicaria coexists with a much larger population of the medium-sized hybrid Daphnia galeata × hyalina in the epilimnion of Lake Maarsseveen (The Netherlands). When large shoals of juvenile perch (Perca fluviatilis) appear in the open water, both species start to migrate vertically. Since D. pulicaria has a larger body-size than D. galeata × hyalina, and is therefore competitive dominant over the hybrid, it is unlikely that both species interact via their common food resource, but they react both to fish predation. However, since they differ in size, and therefore in vulnerability for fish predation, both species adopt different strategies. The smaller bodied, and less vulnerable D. galeata × hyalina exhibits diel vertical migration ascending to the surface at dusk, and staying there during the night. In this way, benefiting from the higher temperatures of the surface layers. In contrast, the large-bodied, and more vulnerable D. pulicaria selects the deep cold hypolimnion water layers as refuge against fish predation. In this way it benefits from a safe habitat, free from fish predators, but on the other hand suffers from low water temperatures, which decrease its fitness. It is likely that the relatively higher temperature in the upper water layers serves as a proximate factor for the downward migration of D. pulicaria.     

Vigilance in ursine black and white colobus monkeys (Colobus vellerosus): an examination of the effects of conspecific threat and predation

Vigilance is thought to have evolved as an antipredator defense but, in primates, conspecific threat often better explains this behavior. We examined vigilance in one group of Colobus vellerosus inhabiting the Boabeng‐Fiema Monkey Sanctuary in Ghana. We aimed to (1) describe factors affecting vigilance in this population, and (2) examine the importance of predation avoidance and conspecific threat in explaining vigilance patterns. Because of a male takeover preceding the study, our focal group (B2) consisted of six adult males and three adult females. We collected 490 10‐min focal samples (82 hr) from all adults in the group (N=9) from July to November, 2001. To avoid predators, individuals should be more vigilant (i) with fewer neighbors, and (ii) in areas of the canopy with higher predation risks. Conspecific threats can be divided into extra‐ and intra‐group threats. Extra‐group threats should lead to higher vigilance levels (iii) during intergroup encounters, and (iv) in areas where the home range overlaps with other groups of colobus. Intra‐group threats should lead to greater vigilance (v) in the presence of neighbors and (vi) while feeding or occupying food patches (if resources are limiting). Our results best support the hypothesis that vigilance functions primarily to detect extra‐group, conspecific threats. Individuals were most vigilant during intergroup encounters and in areas of range overlap, and monthly vigilance rates were associated with monthly encounter rates. Individuals tended to scan less in proximity to familiar vs. unfamiliar group mates, suggesting that relationship quality may affect colobus vigilance. Finally, predation pressures or anthropogenic disturbances might have influenced vigilance, as individuals were more vigilant low in the canopy. However, this last result could also be due to the lower visibility because of dense foliage or to the fact that the monkeys have access to fewer escape routes when ranging lower in the canopy. Am. J. Primatol. 71:919–927, 2009. © 2009 Wiley‐Liss, Inc.