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Feeding and vigilance behaviour of breeding ostriches (Struthio camelus) in a farming environment in Britain.

1. Vigilance and feeding behaviour of male and female adult breeding ostriches were recorded to determine feeding and scanning bout lengths, a time budget and the pattern of vigilance immediately after food was provided. 2. Males were more vigilant and fed for shorter periods than females immediately after concentrate food was delivered but not throughout the whole day. 3. Most interscan periods of males were below 40 s with a maximum of 90 s compared with most interscan periods of females lasting less than 70 s with a maximum of 160 s. 4. Gender differences in behaviour are attributed to increased male vigilance for predators and/or conspecifics, and increased female feeding required for egg production and greater opportunity to feed because of male vigilance.

Animals↗

High reproductive rates result in high predation risks: a mechanism promoting the coexistence of competing prey in spatially structured populations.

I tested the hypothesis that spatial structure provides a trade-off between reproduction and predation risk and thereby facilitates predator-mediated coexistence of competing prey species. I compared a cellular automata model to a mean-field model of two prey species and their common predator. In the mean-field model, the prey species with the higher reproductive rate (the superior competitor) always outcompeted the other species (the inferior competitor), both in the presence of and the absence of the predator. In the cellular automata model, both prey species, which differed only in their reproductive rates, coexisted for a long time in the presence of their common predator at intermediate levels of predation. At low predation rates, the superior competitor dominated, while high predation rates favored the inferior competitor. This discrepancy in the results of the different models was due to a trade-off that spontaneously emerged in spatially structured populations; that is, the more clustered distribution of the superior competitor made it more susceptible to predation. In addition, coexistence of competing prey species declined with increasing dispersal ranges of either prey or predator, which suggests that the trade-off that results from spatial structure becomes less important as either prey or predator disperse over a broader range.

Animals↗

How localized consumption stabilizes predator-prey systems with finite frequency of mixing.

Predator-prey theory began with aspatial models that assumed organisms interacted as if they were "well-mixed" particles that obey the laws of mass action, but it has become clear that both the spatial and individual nature of many organisms can change how the dynamics of such systems function. Here I examine how localized consumption of prey by predators changes the dynamics of predator-prey systems; I use an individual-based simulation of the Rosenzweig-MacArthur model in implicit space and its mean-field approximation. In combination with limited movement, localized consumption makes the predator-prey dynamics more stable than the comparable "well-mixed" Rosenzweig-MacArthur model. Using a spatial correlation, one can directly compare a simplified version of the individual-based model with the Rosenzweig-MacArthur model. While this comparison allows the changes in the dynamics to be captured by the "well-mixed" Rosenzweig-MacArthur model, the parameters of the functional response are now dependent on the movement parameters, and so the functional response must be estimated statistically from the dynamics of the individual-based model. Yet this implies that aspatial models may work in a scale-specific fashion for spatial systems. Unlike many recent spatial models, the localized consumption and limited movement in the model presented here cannot produce coherent spatial patterns and do not depend on a patchy structure, as found in metapopulation models. Instead, the individual nature of the interactions creates a diffusion-limited reaction, which appears closer to a form of ephemeral refuge.

Animals↗

Increasing search rate over time may cause a slower than expected increase in prey encounter rate with increasing prey density.

A previous experiment with birds searching for caterpillars in an aviary demonstrated a highly counterintuitive result, that the rate at which a forager encounters prey does not increase linearly with prey density. Here, I demonstrate that if search rate increases over time then this can produce exactly the observed type of behaviour. Further, I argue that declining perception of predation risk over time in the absence of reinforcement, coupled with a trade-off between anti-predator vigilance and searching ability (both widely reported in field and laboratory studies), could generate such a change in search rate over time. Hence, if my hypothesis is correct, the previous experimental results could have considerable generality, and invite reconsideration of our mathematical descriptions of predator-prey interactions.

Animals↗

Spatial processes can determine the relationship between prey encounter rate and prey density.

Theoretical models frequently assume that the rate at which a searching predator encounters prey increases linearly with prey density. In a recent experiment using great tits searching for winter moth caterpillars, the time to find the first prey item did not decline as quickly with density as the standard theory assumes. Using a spatial simulation model, we show that prey aggregation and/or spatially correlated searching behaviour by the predator can generate a range of relationships, including results that are qualitatively similar to those found in the great tit experiment. We suggest that further experiments are required to determine whether the explanation proposed here is correct, and that theoretical work is needed to determine how this behaviour is likely to influence the ecological and evolutionary dynamics of predator-prey communities.

Animals↗

Trends in the maternal investment of harbour porpoises are uncoupled from the dynamics of their primary prey.

Harbour porpoises in the Bay of Fundy and Gulf of Maine feed primarily on Atlantic herring. Herring stocks have undergone dramatic fluctuations in abundance over the past three decades due to changes in fishing intensity. In order to understand the effects of these changes in prey abundance on the patterns of maternal investment, I examined interdecadal variation in the size of porpoise calves measured in August prior to weaning. Female harbour porpoises exhibited significant variation in maternal investment between 1970 and 1999. During the 1980s, females consumed more herring and produced larger calves. Surprisingly, however, this increased maternal investment occurred during the period of lowest prey abundance, perhaps because the herring stock assessment does not reflect the availability or quality of prey to female porpoises.

Adipose Tissue↗

Good foragers can also be good at detecting predators.

The degree to which foraging and vigilance are mutually exclusive is crucial to understanding the management of the predation and starvation risk trade-off in animals. We tested whether wild-caught captive chaffinches that feed at a higher rate do so at the expense of their speed in responding to a model sparrowhawk flying nearby, and whether consistently good foragers will therefore tend to respond more slowly on average. First, we confirmed that the time taken to respond to the approaching predator depended on the rate of scanning: as head-up rate increased so chaffinches responded more quickly. However, against predictions, as peck rate increased so head-up rate increased and mean length of head-up and head-down periods decreased. Head-up rate was probably dependent on peck rate because almost every time a seed was found, a bird raised its head to handle it. Therefore chaffinches with higher peck rates responded more quickly. Individual chaffinches showed consistent durations of both their head-down and head-up periods and, therefore, individuals that were good foragers were also good detectors of predators. In relation to the broad range of species that have a similar foraging mode to chaffinches, our results have two major implications for predation/starvation risk trade-offs: (i) feeding rate can determine vigilance scanning patterns; and (ii) the best foragers can also be the best at detecting predators. We discuss how our results can be explained in mechanistic terms relating to fundamental differences in how the probabilities of detecting food rather than a predator are affected by time. In addition, our results offer a plausible explanation for the widely observed effect that vigilance continues to decline with group size even when there is no further benefit to reducing vigilance.

Animals↗

Risk allocation and competition in foraging groups: reversed effects of competition if group size varies under risk of predation.

Animals often feed more quickly when in larger groups. This group-size effect is often explained by safety advantages for groups but an alternative explanation is that animals feed faster in larger groups because of greater scramble competition for limited food. We show that predation risk enhances the group-size effect if groups vary in size. By contrast, competition leads to the group-size effect only when individuals feed in groups of constant size. When individuals feed in groups that vary in size, the best strategy for dealing with competition is to feed intensely when in smaller groups and feed little when in larger (more competitive) groups. In all situations, the effects of competition interact with the effects of predation risk in a simple multiplicative way. Our results suggest that scramble competition is not a general explanation for the group-size effect on vigilance in situations where group size changes relatively rapidly.

Animals↗

Caste evolution and ecology: a special worker for novel prey.

Individual specialization underpins the division of labour within ant societies, but only in a small minority do morphological specialists, or physical castes, exist in the workforce. The genetic conditions that allow such castes to evolve are well understood, but the ecological pressures that select for them are not. We provide compelling evidence that the task of transporting novel prey selected for an exaggerated transport caste, or 'submajor', in the army ant Eciton burchellii. This species is the only Eciton that preys upon large arthropods as well as ants, the ancestral prey type, and by comparing load-transport among Eciton species and within E. burchellii, we show that this mixed diet significantly constrains transport efficiency. Crucially, however, we also show that E. burchellii submajors are highly specialized on transporting non-ant prey, and we demonstrate experientially that it is specifically this prey type that constrains prey-transport efficiency. Our study also suggests that phylogenetic constraints associated with the Eciton lifestyle intensified selection for the exaggerated submajor of E. burchellii. Thus, we propose that a novel task may only select for a special caste when phylogenetic constraints preclude the evolution of alternative solutions. This identifies a new and potentially general scenario for the evolution of physical castes.

Analysis of Variance↗

Hydrodynamic image formation by the peripheral lateral line system of the Lake Michigan mottled sculpin, Cottus bairdi.

Lake Michigan mottled sculpin (Cottus bairdi) have a lateral-line-mediated prey-capture behaviour that consists of an initial orientation towards the prey, a sequence of approach movements, and a final strike at the prey. This unconditioned behaviour can be elicited from blinded sculpin in the laboratory by both real and artificial (vibrating sphere) prey. In order to visualize what Lake Michigan mottled sculpin might perceive through their lateral line when approaching prey, we have combined anatomical, neurophysiological, behavioural and computational modelling techniques to produce three-dimensional maps of how excitation patterns along the lateral line sensory surface change as sculpin approach a vibrating sphere. Changes in the excitation patterns and the information they contain about source location are consistent with behavioural performance, including the approach pathways taken by sculpin to the sphere, the maximum distances at which approaches can be elicited, distances from which strikes are launched, and strike success. Information content is generally higher for laterally located sources than for frontally located sources and this may explain exceptional performance (e.g. successful strikes from unusually long distances) in response to lateral sources and poor performance (e.g. unsuccessful strikes) to frontal sources.

Animals↗

Behavioural and ecological consequences of limited attention.

Ecological research in the past few decades has shown that most animals acquire and respond adaptively to information that affects survival and reproduction. At the same time, neurobiological studies have established that the rate of information processing by the brain is much lower than the rate at which information is encountered in the environment, and that attentional mechanisms enable the brain to focus only on the most essential information at any given time. Recent integration of the ecological and neurobiological approaches helps us to understand key behaviours with broad ecological and evolutionary implications. Specifically, current data indicate that limited attention affects diet choice and constrains animals' ability simultaneously to feed and attend to predators. Recent experiments also suggest that limited attention influences social interactions, courtship and mating behaviour.

Animals↗

Evolution of handling time can destroy the coexistence of cycling predators.

Several consumers (predators) with Holling type II functional response may robustly coexist even if they utilize the same resource (prey), provided that the population exhibits nonequilibrium dynamics and the handling time of predators is sufficiently different. We investigate the evolution of handling time and, in particular, its effect on coexistence. Longer handling time is costly in terms of lost foraging time, but allows more nutrients to be extracted from a captured prey individual. Assuming a hyperbolically saturating relationship between handling time and the number of new predators produced per prey consumed, we obtain three results: (i) There is a globally evolutionarily stable handling time; (ii) At most two predator strategies can coexist in this model; (iii) When two predators coexist, a mutant with intermediate handling time can always invade. This implies that there is no evolutionarily stable coexistence, and the evolution of handling time eventually leads to a single evolutionarily stable predator. These results are proven analytically and are valid for arbitrary (not only small) mutations; they however depend on the relationship between handling time and offspring production and on the assumption that predators differ only in their prey handling strategy.

Animals↗

Reduced competition and altered feeding behavior among marine snails after a mass extinction.

Extinction may alter competitive interactions among surviving species, affecting their subsequent recovery and evolution, but these processes remain poorly understood. Analysis of predation traces produced by shell-drilling muricid snails on bivalve prey reveals that species interactions were substantially different before and after a Plio-Pleistocene mass extinction in the western Atlantic. Muricids edge- and wall-drilled their prey in the Pliocene, but Pleistocene and Recent snails attacked prey only through the shell wall. Experiments with living animals suggest that intense competition induces muricid snails to attack shell edges. Pliocene predators, therefore, probably competed for resources more intensely than their post-extinction counterparts.

Animals↗

Behavioral regulation of the milieu interne in man and rat.

In regulating the internal homeostatic environment mammals, by necessity, employ behavioral strategies that differ from the tactics used in coping with contingencies in the external environment. When an animal consumes a meal, the palatability of that meal is automatically adjusted in accordance with the ultimate internal effects of that meal. If the meal causes toxicosis, the animal acquires an aversion for the taste of the meal; conversely, if recuperation follows ingestion of the meal, the taste of that meal is enhanced. Unlike the learning that occurs when externally referred visual and auditory signals are followed by punishment in the form of peripheral pain or reward in the form of food in the mouth, conditioning to the homeostatic effects of food can occur in a single trial and rarely requires more than three to five trials, even though the ultimate effects of the meal are delayed for hours. Paradoxically, the animal need not be aware of the ultimate internal effect in the same sense that it is aware of external contingencies. For example, an aversion can be acquired even if the animal is unconscious when the agent of illness is administered. Thus, the way in which food-effects are stored in memory may be fundamentally different from the way in which memories of specific time-space strategies devised for external contingencies are stored. This separation of function is indicated by limbic lesions which disrupt conditioning to a buzzer that is followed by shock and facilitate conditioning to a taste that is followed by illness. Operationally speaking, one can describe both aversion conditioning and buzzer-shock conditioning in the spacetime associationistic terms of classical conditioning. However, psychologically speaking, one must realize that in aversion conditioning the animal does not act as if it were acquiring an "if-then" strategy. It acts as if a hedonic shift, or a change in the incentive value of the flavor were taking place. Such hedonic shifts are critical in regulation of the internal milieu. When an animal is in need of calories, food tends to be more palatable; as the caloric deficit is restored, food becomes less palatable. If the animal's body temperature is below optimum, a warm stimulus applied to the skin is pleasant. When body temperature is too high, the converse is true. In this way, homeostatic states monitored by internal receptors produce changes in the incentive values of external stimuli sensed by the peripheral receptors, and guide feeding behavior. In mammals at least, the gustatory system, which provides sensory control of feeding, sends fibers to the nucleus solitarius. This brainstem relay station also receives fibers from the viscera and the internal monitors of the area postrema. Ascending fibers bifurcate at the level of the pons and project toward the feeding areas of the hypothalamus and the cortex. The olfactory system which primarily projects to the limbic system does not play a primary role in adjusting food incentives. Rather, it plays a secondary role in the activation of feeding, as do other external sensory systems. This specialized conditioning mechanism, which specifically adjusts gustatory hedonic values through delayed visceral feedback, is widespread among animals, including man and rat. These two species are remarkably similar in their thresholds and preferences for gustatory stimuli. The behavioral similarities are based on the animals' having similar gustatory systems, similar convergence of gustatory and internal afferents to the nucleus solitarius, and similar midbrain regulatory mechanisms. Thus, it is not surprising that the feeding of obese rats with internal hypothalamic damage resembles the feeding of obese human beings insensitive to the internal signs of this caloric state. Obviously, man has a highly specialized form of symbolic communication and the rat does not, yet man's cognitive specialization does not prevent him from developing aversions to food consumed before illness even when he knows that his illness was not caused by food (43).

Animals↗