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Optimizing time and resource allocation trade-offs for investment into morphological and behavioral defense.

Prey organisms are confronted with time and resource allocation trade-offs. Time allocation trade-offs partition time, for example, between foraging effort to acquire resources and behavioral defense. Resource allocation trade-offs partition the acquired resources between multiple traits, such as growth or morphological defense. We develop a mathematical model for prey organisms that comprise time and resource allocation trade-offs for multiple defense traits. Fitness is determined by growth and survival during ontogeny. We determine optimal defense strategies for environments that differ in their resource abundance, predation risk, and defense effectiveness. We compare the results with results of simplified models where single defense traits are optimized. Our results indicate that selection acts in favor of integrated traits. The selective advantage of expressing multiple defense traits is most pronounced at intermediate environmental conditions. Optimizing single traits generally leads to a more pronounced response of the defense traits, which implies that studying single traits leads to an overestimation of their response to predation. Behavioral defense and morphological defense compensate for and augment each other depending on predator densities and the effectiveness of the defense mechanisms. In the presence of time constraints, the model shows peak investment into morphological and behavioral defense at intermediate resource levels.

Animals↗

Foraging and feeding ecology of the gray wolf (Canis lupus): lessons from Yellowstone National Park, Wyoming, USA.

The foraging and feeding ecology of gray wolves is an essential component to understanding the role that top carnivores play in shaping the structure and function of terrestrial ecosystems. In Yellowstone National Park (YNP), predation studies on a highly visible, reintroduced population of wolves are increasing our understanding of this aspect of wolf ecology. Wolves in YNP feed primarily on elk, despite the presence of other ungulate species. Patterns of prey selection and kill rates in winter have varied seasonally each year from 1995 to 2004 and changed in recent years as the wolf population has become established. Wolves select elk based on their vulnerability as a result of age, sex, and season and therefore kill primarily calves, old cows, and bulls that have been weakened by winter. Summer scat analysis reveals an increased variety in diet compared with observed winter diets, including other ungulate species, rodents, and vegetation. Wolves in YNP hunt in packs and, upon a successful kill, share in the evisceration and consumption of highly nutritious organs first, followed by major muscle tissue, and eventually bone and hide. Wolves are adapted to a feast-or-famine foraging pattern, and YNP packs typically kill and consume an elk every 2-3 d. However, wolves in YNP have gone without fresh meat for several weeks by scavenging off old carcasses that consist mostly of bone and hide. As patterns of wolf density, prey density, weather, and vulnerability of prey change, in comparison with the conditions of the study period described here, we predict that there will also be significant changes in wolf predation patterns and feeding behavior.

Animals↗

Eating without hands or tongue: specialization, elaboration and the evolution of prey processing mechanisms in cartilaginous fishes.

The ability to separate edible from inedible portions of prey is integral to feeding. However, this is typically overlooked in favour of prey capture as a driving force in the evolution of vertebrate feeding mechanisms. In processing prey, cartilaginous fishes appear handicapped because they lack the pharyngeal jaws of most bony fishes and the muscular tongue and forelimbs of most tetrapods. We argue that the elaborate cranial muscles of some cartilaginous fishes allow complex prey processing in addition to their usual roles in prey capture. The ability to manipulate prey has evolved twice along different mechanical pathways. Batoid chondrichthyans (rays and relatives) use elaborate lower jaw muscles to process armored benthic prey, separating out energetically useless material. In contrast, megacarnivorous carcharhiniform and lamniform sharks use a diversity of upper jaw muscles to control the jaws while gouging, allowing for reduction of prey much larger than the gape. We suggest experimental methods to test these hypotheses empirically.

Animals↗

Rotational feeding in caecilians: putting a spin on the evolution of cranial design.

Caecilians are a poorly known group of amphibians with a highly derived skull and cranial musculature that has evolved in response to their specialized head-first burrowing lifestyle. They possess a unique jaw-closing system, which is shown to be capable of generating considerable bite forces for its head width (1.09+/-0.34 and 0.62+/-0.31 N for Schistometopum thomense and Boulengerula taitanus, respectively). However, comprehensive dietary studies indicate that there is no need for large bite forces, since most caecilians appear to be generalist predators of subterranean macrofauna. Here, we demonstrate, based on in vivo external and X-ray video recordings of animals feeding, that long-axis body rotations are used independent of prey size by these two species of caeciliid caecilians when feeding underground. Further, we show that individuals are capable of generating a substantial spinning force, which is greater than their bite force (1.35+/-0.26 and 1.02+/-0.18 N, respectively). These observations shed light on the functional and the evolutionary significance of several unique features of the cranial design in derived caecilians; spinning may allow the individuals to judge prey size and subsequently reduce oversized prey within gape limits.

Amphibians↗

Low-frequency echolocation enables the bat Tadarida teniotis to feed on tympanate insects.

The European free-tailed bat, Tadarida teniotis, forages in uncluttered airspace by using intense narrow-band echolocation calls with low frequency (11-12 kHz), and feeds on relatively large flying insects, mainly (90% by volume) of the tympanate orders Lepidoptera and Neuroptera. The use of low-frequency echolocation calls without strong harmonics appears to be a specialization for long-range detection of large, tympanate insects, which are less well represented in the diet of most other aerial-hawking bats. The results provide evidence in support of the allotonic frequency hypothesis, i.e. that use of echolocation calls with frequencies above or below the best hearing of tympanate insects is an adaptation to increase the availability of these insects.

Animals↗

Nest predation increases with parental activity: separating nest site and parental activity effects.

Alexander Skutch hypothesized that increased parental activity can increase the risk of nest predation. We tested this hypothesis using ten open-nesting bird species in Arizona, USA. Parental activity was greater during the nestling than incubation stage because parents visited the nest frequently to feed their young during the nestling stage. However, nest predation did not generally increase with parental activity between nesting stages across the ten study species. Previous investigators have found similar results. We tested whether nest site effects might yield higher predation during incubation because the most obvious sites are depredated most rapidly. We conducted experiments using nest sites from the previous year to remove parental activity. Our results showed that nest sites have highly repeatable effects on nest predation risk; poor nest sites incurred rapid predation and caused predation rates to be greater during the incubation than nestling stage. This pattern also was exhibited in a bird species with similar (i.e. controlled) parental activity between nesting stages. Once nest site effects are taken into account, nest predation shows a strong proximate increase with parental activity during the nestling stage within and across species. Parental activity and nest sites exert antagonistic influences on current estimates of nest predation between nesting stages and both must be considered in order to understand current patterns of nest predation, which is an important source of natural selection.

Animals↗

The effects of temporal variation in predation risk on anti-predator behaviour: an empirical test using marine snails.

Foraging animals must often balance the conflicting demands of finding food and avoiding predators. Temporal variation in predation risk is expected to influence how animals allocate time to these behaviours. Counterintuitively, the proportion of time spent foraging during both high- and low-risk periods should increase with increasing time exposed to high risk. We tested this prediction using intertidal marine snails (Littorina spp.) that were exposed to temporal variation in perceived predation risk from crabs (Cancer productus and Cancer magister). Our results were consistent with those predicted for high-risk, but not low-risk, periods. During high-risk periods, a greater number of snails foraged (versus those that left the water or remained in their shells) as time at high perceived risk increased. For low-risk periods, there was no relationship between the number of snails foraging and time at high risk. This might be due to snails in all treatments foraging maximally in the low-risk periods. As a consequence, the difference in the number of snails foraging between high- and low-risk periods decreased with increasing time subject to high risk. These results indicate that the commonly used protocol of exposing foragers to a single pulse of heightened risk might tend to overestimate their typical investment in anti-predator behaviour.

Adaptation, Biological↗

A division of labour with role specialization in group-hunting bottlenose dolphins (Tursiops truncatus) off Cedar Key, Florida.

Individual role specialization during group hunting is extremely rare in mammals. Observations on two groups of bottlenose dolphins (Tursiops truncatus) in Cedar Key, Florida revealed distinctive behavioural roles during group feeding. In each group, one individual was consistently the 'driver', herding the fishes in a circle toward the remaining 'barrier' dolphins. Aerial fish-capture rates differed between groups, as well as between the driver and barrier dolphins, in one group but not in the other. These differences between the two groups may reflect differences in group stability or in prey school size.

Animals↗

A theoretical investigation of the effect of predators on foraging behaviour and energy reserves.

Data show that when small birds are exposed to a model of a predator, their body mass may either increase or decrease. Although attempts have been made to explain the data using previous models, these models are based on a constant level of predation and hence are not appropriate for making predictions about the response of a bird to the sight of a predator. We have developed a novel model that includes encounters between a bird and potential predators. We show that, depending on the biology of the predator, optimal body mass may either increase or decrease. The model also makes predictions about the foraging behaviour of the bird after it has seen a predator.

Animals↗

Do unprofitable prey evolve traits that profitable prey find difficult to exploit?

Prey that are unprofitable to attack (for example, those containing noxious chemicals) are often conspicuously patterned and move in a slower and more predictable manner than species lacking these defences. Contemporary theories suggest these traits have evolved as warning signals because they can facilitate both associative and discriminative avoidance learning in predators. However, it is unclear why these particular traits and not others have tended to evolve in unprofitable prey. Here we show using a signal detection model that unprofitable prey will evolve conspicuous colours and patterns partly because these characteristics cannot readily evolve in profitable prey without close mimicry. The stability of this signal is maintained through the costs of dishonesty in profitable prey. Indeed, unprofitable prey will sometimes evolve a conspicuous form to reduce mimetic parasitism, even in the unlikely event that this form can be more closely mimicked. This is one of the first mathematical models of the evolution of warning signals to allow for the possibility of mimicry, yet our analyses suggest it may offer a general explanation as to why warning signals take the form that they do. Warning signals and mimicry may therefore be more closely related than is currently supposed.

Animals↗

The influence of prey experience on movement pattern preference in Salamandra salamandra (L.).

Groups of juvenile Salamandra salamandra were raised selectively with different types of prey. The influence of these conditions on the movement pattern preference was studied during the first six months after metamorphosis. Responses to a black horizontal bar moving continuously or stepwise at step frequencies between 0.25 and 8 steps/s were tested. The differently raised groups showed significant differences in their respective preference for stimulus movement patterns which corresponded with the movement pattern of their food. At the same time, superimposed upon this learning process, a general increase in the relative efficiency of continuous stimulus movement with regard to stepwise stimulus movement could be observed.

Animals↗

The role of early sensory experience in the prey catching responses of Salamandra salamandra to stationary prey.

Two groups of newly metamorphosed Salamandra salamandra (L.) were raised with living and with stationary prey, respectively. One year later, the group which had experience exclusively with stationary prey was significantly better in responding to stationary prey than the group which only had experience with moving prey. This was the case both under light and dark conditions. The experiments show that prey-catching behavior in salamanders can be considerably modified by experience.

Animals↗

Neonatal lamb losses due to feral pig predation.

An investigation into causes of low lamb marking percentages was made on a property in north-west New South Wales from 1971 to 1975. Investigations revealed that from 11 to 70% of the ewes in lamb were losing all their lambs. Observations suggested that feral pig predation was a factor in the perinatal loss. In 1975, 2 groups of ewes were placed in adjoining paddocks prior to lambing. Feral pigs were excluded from one paddock for most of lambing by means of an electric fence. In this paddock, 117% of lambs were marked compared with 80% in the adjoining paddock. It was estimated that in 1975 over 600 lambs were killed by feral pigs from 1,422 ewes lambing in paddocks with feral pigs. The problems involved in the diagnosis of feral pig predation are discussed.

Animal Population Groups↗

Extreme reversed sexual dichromatism in a bird without sex role reversal.

Brilliant plumage is typical of male birds, reflecting differential enhancement of male traits when females are the limiting sex. Brighter females are thought to evolve exclusively in response to sex role reversal. The striking reversed plumage dichromatism of Eclectus roratus parrots does not fit this pattern. We quantify plumage color in this species and show that very different selection pressures are acting on males and females. Male plumage reflects a compromise between the conflicting requirements for camouflage from predators while foraging and conspicuousness during display. Females are liberated from the need for camouflage but compete for rare nest hollows.

Adaptation, Biological↗