Prey size,prey abundance,and temperature as correlates of growth in stream populations of cutthroat trout

Growth and maximum size of stream fishes can be highly variable across populations. For salmonid fishes in streams, individuals from populations confined to headwater streams often exhibit small size at maturity in comparison to populations with access to main-stem rivers. Differences in prey size, prey availability, and metabolic constraints based on temperature may explain patterns of maximum size and growth. In this study, cutthroat trout from headwater stream populations that were isolated above a waterfall were compared to individuals from populations in similar sized streams without a movement barrier and from large main-stem rivers. Cutthroat trout from smaller streams with or without a movement barrier were significantly smaller at a given age than fish from main-stem rivers, where individuals were able to achieve a much larger maximum size. Comparisons of invertebrate drift abundance and size in the three types of streams revealed that drift size did not differ between stream categories, but was highest per volume of water in large main-stem rivers. Across all stream types, prey abundance declined from summer to fall. Temperature declined over the course of the season in a similar manner across all stream types, but remained relatively high later in the season in main-stem river habitats. Prey availability and temperature conditions in main-stem rivers may provide more optimal growing conditions for fish as individuals increase in size and become constrained by prey availability and temperature conditions in small streams. Maintaining connectivity between small spawning and rearing tributary streams and main-stem river habitats may be critical in maintaining large-bodied populations of stream salmonids.     

A PRELIMINARY INVESTIGATION INTO PREY SPECIES PREFERENCE BY THE TROPICAL GASTROPODS NATICA MACULOSA LAMARCK AND THAIS CARINIFERA (LAMARCK)

A preliminary assessment of the prey-species preference of theMalaysian intertidal mudflat gastropods Natica maculosa Lamarckand Thais carinifera (Lamarck) has been made using some commonpotential prey species. The results are assessed in terms ofthe likely benefits and costs of attacking particular prey speciesand, in the case of the two most important prey species, theassessment is supported by details of energy yield. For N. maculosaonly, a comparative assessment of growth when fed on differentprey species is presented. The consequences of observed preferencesfor previous estimates of potential energy flow from a populationof the bivalve Anadara granosa (L.) through populations of thetwo predators, are considered. ^*Present address: Welsh Water Authority, Tremains House, CoychurchRd, Bridgend, CF31 2AR (Received 3 February 1982;     

Ecological and social determinants of group size in transient killer whales

Most analyses of the relationship between group size and foodintake of social carnivores have shown a discrepancy betweenthe group size that maximizes energy intake and that which ismost frequently observed. Around southern Vancouver Island,British Columbia, killer whales of the so-called transient formforage in small groups, and appear to prey exclusively on marinemammals. Between 1986 and 1993, in approximately 434 h of observationson transient killer whales, we observed 138 attacks on fivespecies of marine mammals. Harbor seals were most frequentlyattacked (130 occasions), and the observed average energy intakerate was more than sufficient for the whale's energetic needs.Energy intake varied with group size, with groups of three havingthe highest energy intake rate per individual. While groupsof three were most frequently encountered, the group size experiencedby an average individual in the population (i.e., typical groupsize) is larger than three. However, comparisons between observedand expected group sizes should utilize only groups engagedin the behavior of interest. The typical size of groups consistingonly of adult and subadult whales that were engaged primarilyin foraging activities confirms that these individuals are foundin groups that are consistent with the maximization of energyintake hypothesis. Larger groups may form for (1) the occasionalhunting of prey other than harbor seals, for which the optimalforaging group size is probably larger than three; and (2) theprotection of calves and other social functions. Key words:dispersal, foraging, group hunting, harbor seals, killer whales,optimal group size, social structure. [Behav Ecol 7: 408-416(1996)]     

Foraging efficiency of juvenile bluegill, Lepomis macrochirus, among different vegetated habitats

Optimal foraging theory is devoted to understanding how organisms maximize net energy gain. However, both the theory and empirical studies lack critical components, such as effects of environmental variables across habitats. We addressed the hypothesis that energetic returns of juvenile bluegill are affected by environmental variables characteristic of the vegetated habitats. Predicted optimal diet breadths were calculated and compared to prey items eaten by juvenile bluegill to determine if bluegill were foraging to maximize energetic gain. Differences in habitat profitability among vegetated sites were determined by comparing predictions of maximized energetic return rates (cals^-1) with prey contents of bluegill stomachs. Sizes of most prey items eaten by juvenile bluegill throughout the vegetated sites were smaller than the predicted optimal diet breadths. However, inclusion of smaller prey items in the diet did not seem to affect rate of energetic gain. Energetic return rates were maximized at the 1.5 and 2mm prey size classes and declined only slightly with inclusion of smaller prey sizes. Predicted energetic return rates and average mass in bluegill stomachs were related negatively. Average mass in bluegill stomachs also was associated negatively with Elodea canadensis stem densities and percent of light transfer, suggesting that foraging efficiency of bluegill decreased as plant density and percent of light increased. Results of our research indicate that maximization of energetic return rates is dependent upon availability of prey sizes that contribute to optimal foraging. Thus, determination of those habitats that provide the highest availability of benthic invertebrate prey with the least interference by stems is critical. Enhanced foraging capabilities can promote recruitment, faster growth, better body condition and survival.     

Optimal body size in bumblebees

Summary It is hypothesized that the body size of a bumblebee will be that size which maximizes its average net rate of energy intake while collecting nectar. A mathematical model is developed with the result that the net rate of energy intake of a nectar-collecting bumblebee is expressed as a function of the body size of the bumblebee. From this model the body size which maximizes the net rate of energy intake (i.e., optimal body size) is found (as the solution of an implicit equation). In this situation the advantage of large size is that larger bumblebees fly faster and hence take less flight time than smaller bumblebees. The disadvantage of larger size is greater energetic costs.The parameters of the model are estimated using data obtained from the foraging behavior of bumblebees on monkshood (Aconitum columbianum). The optimal body size is then calculated for workers of Bombus appositus which obtained almost all their nectar from monkshood. The observed and expected (i.e., optimal) body size are found to be close and not significantly different.The model also predicts that, from the bumblebee's point of view, there should be a positive correlation between the size of the bumblebee and the average amount of nectar obtained per flower. Evidence of this correlation is presented and the possible significance of the correlation from the plant's point of view is discussed. A possible extension of the model to general relationships between predator body size, prey size and prey density is discussed.     

Prey size, prey nutrition, and food handling by shrews of different body sizes

We tested some predictions relating metabolic constraints offoraging behavior and prey selection by comparing food handlingand utilization in four sympatric shrew species: Sorex minutus(mean body mass = 3.0 g), S. araneus (8.0 g), Neomys anomalus(10.0 g), and N. fodiens (14.4 g). Live fly larvae, mealwormlarvae, and aquatic arthropods were offered to shrews as smallprey (body mass <0.1 g). Live earthworms, snails, and smallfish were offered as large prey (>0.3 g). The larvae werethe high-nutrition food (>8 kJ/g), and the other prey werethe low-nutrition food (<4 kJ/g). The smallest shrew, S.minutus, utilized (ate + hoarded) <30% of offered food,and the other species utilized >48% of food. The largerthe shrew, the more prey it ate per capita. However, highlyenergetic insect larvae composed 75% of food utilized by S.minutus and only >40% of the food utilized by the other species. Thus, inverse relationships appeared between shrewbody mass and mass-specific food mass utilization and betweenshrew body mass and mass-specific food energy utilization:the largest shrew, N. fodiens, utilized the least food massand the least energy quantity per 1 g of its body mass. Also,the proportion of food hoarded by shrews decreased with increase in size of shrew. With the exception of S. araneus, the sizeof prey hoarded by the shrews was significantly larger thanthe size of prey eaten. Tiny S. minutus hoarded and ate smallerprey items than the other shrews, and large N. fodiens hoardedlarger prey than the other shrews.     

Maximum body size among insular Komodo dragon populations covaries with large prey density

This study documents variation in maximum body size of Komodo dragons ( Varanus komodoensis ) among the four extant island populations in Komodo National Park and compares an indirect measure of deer density, the major prey item for large dragons, to differences in maximum body size among islands. The largest 15% of dragons from the large islands of Komodo and Rinca were significantly longer and heavier than the largest 15% of dragons on the small islands of Gili Motang and Nusa Kode. There was a 33% difference in snout vent length (SVL) between dragons found on Komodo and those found on Gili Motang, with mass varying by more than four-fold. Density of deer pellet groups between islands ranged from 5.86±0.75 groups per transect on Gili Motang to 20.73±1.02 groups per transect on Komodo Island. Maximal dragon SVL and mass was highly positively correlated with this index of deer density. Low prey density on the two small islands could constrain body size via energetic constraints. At present we can not deduce if insular body size variation has arisen through genotypic or phenotypic mechanisms.     

Food, Fat, and Feathers

The annual cycle of the White-throated Sparrow, Zonotrichiaalbicollis, is reviewed with brief references to facets of nutritionaland energetic importance (1) illustrating the life of a smallbird in a varying environment, and (2) showing that certainannual events in field populations can be compared with similarmanifestations in caged individuals. Data from captives areemployed in discussions of energetic variations related to (1)food, the source of nutritive input, (2) fat, the major formof caloric storage in birds, and (3) caloric expenditure. Metabolizableenergy is partitioned by phase of the annual cycle into existenceenergy, including the costs of thermoregulation, and productiveenergy, including expenditures for nocturnal activity (Zugunruhe)and molt. Costs of vernal migration in field birds are comparedwith costs of nocturnal activity in captives to show that energeticestimates in each situation are compatible. This conclusionis supported by a metabolic estimate made for field birds thatis within 6% of the estimated metabolism of captives under similarconditions. Data and statistics from seven additional speciesof buntings are used to examine several bioenergetic principlesfor homoiotherms. (1) Minimal metabolism measured by energybalance methods is proportional to the 0.7 power of body weightbut is higher than standard or resting metabolism measured bygaseous methods. (2) Metabolized energy is inversely relatedto ambient temperature below 25°C, the estimated ad libitumcritical temperature. (3) Heat production and loss are proportionatelyhigher in summer-acclimatized birds below the ad libitum criticaltemperature due to reduced insulation. Two summary plots, relatingtemperature, metabolizable energy, and body weight are given.Directions for future research in the study of avian nutritionare suggested.     

Foraging among cannibals and kleptoparasites: effects of prey size on pike behavior

The northern pike (Esox lucius) is an important and selective piscivorethat chooses smaller prey than predicted from energy / timebudgets. In a laboratory experiment, we investigated pike predatorybehavior to explain this selectivity. Northern pike feedingon different prey sizes in aquaria were observed when foragingalone, when in the presence of chemical cues from similar-sizedor larger conspecifics, and when in the presence of conspecifics thatwere allowed to interact with the focal pike. The results showthat prey handling time increases with prey size and that theduration of manipulating and handling prey inflicts a risk ofexposure to cannibals and kleptoparasites on the pike. Therefore,the risk of falling victim to cannibals or kleptoparasites increaseswith prey size. Attracting and experiencing intraspecific interactorscan be regarded as major fitness costs. Chemical cues from foragingconspecifics have only minor effects on pike foraging behavior.Furthermore, the ability to strike and swallow prey head first improvespike predatory performance because failing to do so increases handlingtime. Our findings emphasize the increasing potential costswith large prey and explain previous contradictory suggestionson the underlying mechanisms of behavior, selectivity, and trophiceffects of northern pike predation.     

An energetic correlate between colony size and foraging effort in seabirds, an example of the Adélie penguin Pygoscelis adeliae

Central-place foraging seabirds alter the availability of their prey around colonies, forming a "halo" of reduced prey access that ultimately constrains population size. This has been indicated indirectly by an inverse correlation between colony size and reproductive success, numbers of conspecifics at other colonies within foraging range, foraging effort (i.e. trip duration), diet quality and colony growth rate. Although ultimately mediated by density dependence relative to food through intraspecific exploitative or interference competition, the proximate mechanism involved has yet to be elucidated. Herein, we show that Adélie penguin Pygoscelis adeliae colony size positively correlates to foraging trip duration and metabolic rate, that the metabolic rate while foraging may be approaching an energetic ceiling for birds at the largest colonies, and that total energy expended increases with trip duration although uncompensated by increased mass gain. We propose that a competition-induced reduction in prey availability results in higher energy expenditure for birds foraging in the halo around large colonies, and that to escape the halo a bird must increase its foraging distance. Ultimately, the total energetic cost of a trip determines the maximum successful trip distance, as on longer trips food acquired is used more for self maintenance than for chick provisioning. When the net cost of foraging trips becomes too high, with chicks receiving insufficient food, chick survival suffers and subsequent colony growth is limited. Though the existence of energetic studies of the same species at multiple colonies is rare, because foraging metabolic rate increases with colony size in at least two other seabird species, we suggest that an energetic constraint to colony size may generally apply to other seabirds.     

Effect of energetic constraints on distribution and winter survival of weasel males

1. The absolute energy needs of small animals are generally lower than those of larger animals. This should drive higher mortality of larger animals, when the environmental conditions deteriorate. However, demonstration of the effect of energy constraints on survivals proved difficult, because the range of body mass within species is generally too small to produce enough variation for studying such an effect. An opportunity for an intraspecific study comes from weasels inhabiting the Bia?owie?a Forest (north-eastern Poland), which are characterized by a threefold variation in body mass. 2. We assumed that in summer larger weasel males are favoured by sexual selection, because they are more successful when competing for mates. We then tested whether they suffer higher mortality in winter, because they have difficulty finding sufficient food to satisfy their energy needs and/or because the additional foraging time would result in increased exposure to predation. 3. We measured daily energy expenditures (DEE) of overwintering weasel males using the doubly labelled water (DLW) technique. We constructed an energetic model predicting how individuals of different size are able to balance their energy budgets feeding on large and small prey while minimizing time spent hunting, thereby reducing their own exposure to predation. 4. The range of body mass in overwintering weasels predicted by our model corresponded very well with the distribution of prey body mass in three different habitats within our study area. Larger individuals were able to compensate for higher food requirements by using habitats with larger prey species than those available to smaller male weasels. This effectively offset the expected negative association between body mass and winter survival predicted from considerations of energy balance. 5. Our results show how energetic constraints affect body mass and spatial segregation of a species at the intra-specific level not only across large geographical ranges, but also within a relatively small area.     

Food web structure and body size: trophic position and resource acquisition

Ulrich Brose Body size is recognized as an important determinant of trophic structure as it affects individual energetic demands, population density, and the interaction between potential prey and predators. However, its relationship with trophic position remains unclear. It has been hypothesized that a positive relationship between body size and trophic position would be associated to some particular trophic structures, which would allow larger organisms to satisfy their energetic demand and sustain viable population sizes at higher trophic positions, where fewer resources are available. To test this hypothesis, we analyzed the diet of 619 killifishes from four species (Austrolebias cheradophilus, A. luteoflammulatus, A. viarius and Cynopoecilus melanotaenia), collected in temporary ponds occurring in the grasslands of Rocha, Uruguay. Trophic position, diet richness, number of energy sources, and evenness were estimated for 20 size classes, formed by consecutive groups of 31 individuals. Gape limitation and preference for the larger available prey were evaluated as explanations for observed patterns with an individual based model (IBM). In agreement with the hypothesis, killifishes presented a strong positive relationship between trophic position and body size (R²=0.86), associated with a trophic structure that could allow larger organisms to have access to more energy from the environment. This was reflected in a positive relationship between body size and 1) prey richness, 2) number of basal energy sources (i.e. plants, detritus, phytoplankton and terrestrial prey), and 3) evenness in prey use. IBM results showed that changes in trophic structure with body size are well explained by gape limitation, but not by size preferences. Our results suggest that the fulfilment of the greater energetic demands of larger organism will depend on community diversity, which typically increases with ecosystem size, indicating a novel connection between area, diversity, body size, and food chain length.     

Inferring prey size variation from mandible acceleration in northern elephant seals

Prey size is an important factor for predators as it affects prey quality (energy content) and hence total energy gain. However, it remains challenging to obtain information about prey size from free‐ranging marine predators. Here, we developed a method that estimates prey size using mandible acceleration in captive northern elephant seals and then applied it to 34 free‐ranging seals. In captive seals, the number of feeding‐related acceleration signals were positively related to prey size category (<15 cm). In free‐ranging seals, smaller number of acceleration signals occurred frequently in both mesopelagic (200–1,000 m) and bathypelagic layers (>1,000 m), suggesting that seals foraged mainly on smaller prey (possibly <15 cm). However, the quantity of larger acceleration signals increased in the bathypelagic layers, suggesting that seals were more likely to forage on larger prey (>15 cm) at deeper depths. These results suggest that seals might compensate for higher energetic costs of deeper‐diving by targeting larger prey. Although our study has practical limitations (e.g., calibrating prey size in captive conditions), our method allows concurrent inference of prey size and foraging behavior, being potentially useful to investigate how predators adjust their behavior in response to the changes in the foraging environment.     

Phenotypic plasticity and sexual dimorphism in size at post-juvenile metamorphosis: common-garden rearing of an intertidal gastropod with determinate growth

Proximate factors of the intraspecific variation in molluscan shell morphology have long received attention in biology. The intertidal gastropod Monetaria annulus (Mollusca; Gastropoda; Cypraeidae) is particularly suitable for the study of variation in body size, because this species is a determinate grower in the sense that soft-body size shows no further increase after the juvenile stage. Cross-sectional field surveys on post-juvenile individuals have indicated that the mean body size varies widely among populations and is larger in females than in males within populations. To examine whether these patterns are due to genetic differences, we conducted a common-garden rearing experiment with juvenile individuals collected from two populations on Okinawa Island. After adjusting for among-individual differences in initial degree of development, statistical analyses revealed that this species exhibits female-biased sexual size dimorphism mediated by a longer development time rather than by faster growth rates in females. Although wild individuals show a remarkable size difference between populations, no size difference was found between the populations in the individuals reared in a common-garden condition. This result suggests that the among-population size difference does not have a genetic basis and is caused by phenotypic plasticity based on environmental heterogeneity among habitats.     

ONTOGENETIC SHIFT IN PREY SIZE SELECTION AND ITS RELATION TO PROFITABILITY IN NUCELLA LAPILLUS

According to the energy maximization premise an animal shouldchoose its diet to maximize its net energy gain per unit handlingtime (E/T_b). Previous studies on adult marine gastropods haveshown this index of profitability to be a monotonicaUy increasingfunction of prey size, yet they select smaller, apparently sub-optimalprey. Laboratory experiments were used to investigate the profitabilityof different sized mussels, as prey, to juvenile Nucella lapillususing two separate criteria. First the ability to promote growthand second the more usual energy gain per unit handling time. The major difference between the two measures of prey valuewas that E/T_h predicted an optimum prey size larger than thegrowth rate model. Selection experiments were used to comparepreferences exhibited by developing Nucella with the spectraof profitability defined from the two models. In general thewhelks' choice of prey size was best predicted by the growthrate model of prey value. Field studies illustrated a functionalrelationship between size of mussel eaten and size of whelk,the mean size of mussel eaten again conformed more closely tomaximization of growth rate. The energetic return in the E/T_h model is generally taken asthe amount of food ingested, however gross growth efficiencywas found to be a decreasing function of ingestion rate. Sinceingestion rate was an increasing function of prey size, theE/T_h model overestimated the value of large prey by ignoringthe efficiency with which food is used by the predator.     

Habitat-specific clutch size and cost of incubation in eiders reconsidered

The energetic incubation constraint hypothesis (EICH) for clutch size states that birds breeding in poor habitat may free up resources for future reproduction by laying a smaller clutch. The eider (Somateria mollissima) is considered a candidate for supporting this hypothesis. Clutch size is smaller in exposed nests, presumably because of faster heat loss and higher incubation cost, and, hence, smaller optimal clutch size. However, an alternative explanation is partial predation: the first egg(s) are left unattended and vulnerable to predation, which may disproportionately affect exposed nests, so clutch size may be underestimated. We experimentally investigated whether predation on first-laid eggs in eiders depends on nest cover. We then re-evaluated how nesting habitat affects clutch size and incubation costs based on long-term data, accounting for confounding effects between habitat and individual quality. We also experimentally assessed adult survival costs of nesting in sheltered nests. The risk of egg predation in experimental nests decreased with cover. Confounding between individual and habitat quality is unlikely, as clutch size was also smaller in open nests within individuals, and early and late breeders had similar nest cover characteristics. A trade-off between clutch and female safety may explain nest cover variation, as the risk of female capture by us, mimicking predation on adults, increased with nest cover. Nest habitat had no effect on female hatching weight or weight loss, while lower temperature during incubation had an unanticipated positive relationship with hatching weight. There were no indications of elevated costs of incubating larger clutches, while clutch size and colony size were positively correlated, a pattern not predicted by the ‘energetic incubation constraint’ hypothesis. Differential partial clutch predation thus offers the more parsimonious explanation for clutch size variation among habitats in eiders, highlighting the need for caution when analysing fecundity and associated life-history parameters when habitat-specific rates of clutch predation occur.     

Carnivore body size: Ecological and taxonomic correlates

Summary Variation in body size (weight) is examined across the order Carnivora in relation to taxonomy (phylogeny), latitude, habitat, zonation, activity cycle, diet, prey size, and prey diversity. Significant differences in body weight are observed with respect to family membership. Some of these differences may be explained by phylogenetic history and/or dietary effects. Body weight is not correlated with habitat, zonation, activity cycle or latitudinal gradients. Significant differences in body weight are found among insectivorous, herbivorous and carnivorous species, and some of these differences may relate to energetic constraints. Among predatory carnivores, prey size and diversity increases with body weight. The adaptive significance, both intra- and inter-specifically, of prey characteristics (size, availability, diversity) and carnivore body weight qualities (strength, endurance, hunting technique) is discussed.     

Habitat size thresholds for predators: Why damselflies only occur in large bromeliads

Predators are often more sensitive to habitat size than their prey and frequently occur in only the largest habitats. Four explanations have been proposed for this pattern: (a) Small habitats do not have enough energy to support higher trophic levels; (b) small habitats are less likely to contain particular prey required by specialist predators; (c) small habitats are risky for predators with slow life histories or large body sizes; and (d) small habitats are numerically unlikely to be colonized by regionally rare species, such as predators. We critically examine these four hypotheses in relation to the predatory damselfly larva, Mecistogaster modesta Selys. (Pseudostigmatidae), which occurs almost exclusively in bromeliads > 100ml in capacity. We synthesize multiple years of survey data and three manipulative experiments from the Área de Conservación Guanacaste, Costa Rica, to conclude that damselflies do not occur in small bromeliads due to their higher risk of desiccation—not because of energetic limitation, trophic specialization, risk of terrestrial predation, or pure numerical effects. These results suggest that recent and predicted declines in precipitation in northwestern Costa Rica may further restrict bromeliad occupancy by damselflies, with cascading consequences for the rest of the aquatic food web.     

Body size,energetic and foraging mode of raptors in central Chile

Summary An inferential analysis of the foraging mode (opportunist or mediate by prey selection) of a taxonomic assemblage of raptors in central Chile was conducted. The analysis of energetic aspects such as daily requirements of predators and energy supplied by the preys, with estimations of prey relative abundances and their incidence in the diet of raptors, led us to conclude that contrarly to the opportunistic hunting mode suggested by others authors, these predators apparently present prey selection (on a biomass basis). This phenomenon is particularly evident in raptors of small body size.