Invertebrate prey and predatory behaviour of the omnivorous African tortoise Kinixys spekii

Invertebrates form an important part of the diet in the omnivorous African tortoises of the genus Kinixys. Millipede prey of Kinixys spekii in Zimbabwe had a mean volume of 0.9 ml, and made up 64.7% of invertebrate food by volume; beetles made up most of the remainder and had a mean volume of 1.4 ml. The mean mass of invertebrate prey was 0.19% of tortoise mass, a similar value of relative prey mass to many insectivorous lizards. Tortoises preferentially attacked moving millipedes at or just behind the head, but predatory behaviour was otherwise unspecialized, with tortoises killing millipedes by ingesting them in pieces. Handling times of millipede prey varied significantly with relative prey mass, defensive behaviour, and direction of ingestion. The profitability (mass intake/handling time) of millipedes was maximal at a relative prey mass of 0.2%; the basis on which prey are selected is discussed.     

Nestling provisioning in water pipits (Anthus spinoletta): do parents go for specific nutrients or profitable prey?

In this study, we investigated whether free-living insectivorous water pipits (Anthus spinoletta) choose prey according to biochemical quality as measured by protein, lipid, carbohydrate, energy and water contents and/or according to profitability as measured by density, size and catchability. Food preference – expressed in relation to availability – is estimated for 22 arthropod taxa (families and orders). Uni- and multivariate statistics detected no relationships between food preference and nutrient contents, but revealed that more larger prey items are fed to nestlings than smaller ones, both for all prey taken together and within individual taxa. Furthermore, slow-flying arthropods, which are easier to catch, were usually preferred over walking and fast-flying ones. Combined with results from previous studies on the effects of vegetation, prey density and catchability on search times and energy intake, these findings suggest that water pipits select their prey primarily to maximize profitability, i.e. energy intake per unit time. Qualitative traits seem to be important only for specific taxa. For example, toxins or poor digestibility may be responsible for the avoidance of heteropterans, beetles and ants and for feeding the nestlings fewer tipulids than expected at high tipulid densities. Received: 18 January 1999 / Accepted: 14 June 1999     

Prey selection by a stonefly: the influence of hunger and prey size

Summary The influences of hunger and prey size on prey selection by the stonefly Hesperoperla pacifica (Perlidae) were explored in the laboratory by observing behavioral responses toward ten prey taxa and three nonprey taxa. Patterns of behavior were consistent with most assumptions and predictions of optimal foraging theory predicting sizebased prey selection by pursuing predators. Handling time appeared to increase as an exponential function of prey mass, and prey profitability (mg/s) was highest for small and intermediate-sized prey. Fasted stoneflies consumed a wide range of prey sizes, whereas well-fed stoneflies concentrated their attacks on intermediate-sized prey. Responses of H. pacifica to nonprey taxa, however, suggest that prey recognition and selection are not based on size alone.     

Feeding time strategies of the fringe-toed lizard,Uma inornata,during breeding and non-breeding seasons

Summary I examined the foraging behavior during the breeding and non-breeding seasons, May and July 1986, of the fringe-toed lizard Uma inornata (Iguanidae). During the breeding season males differ from females in their diet and in their foraging time strategy, males exhibiting time minimization and females energy maximization. In May, plant associated foods were selectively eaten. Males concentrated on flowers, a readily available quick energy food, which reduced foraging time and increased time for reproductive activities. Time budgets indicate that males spend over twice as much time in the open and in movement in May than do females. Females at this time restrict their activities to the cover of perennial bushes, and feed primarily on plant foods (flowers and arthropods). Energy maximization appears to be maintained by both sexes in the non-breeding season when food resources diminished to one-half of those in the breeding season. The lizards were less selective in their July feeding habits, broadening their diets to include ground-dwelling arthropods and foliage. Predation by these lizards follows a wait-ambush mode of foraging.     

Trophic ecology of the Southern Grey Shrike (Lanius meridionalis) in insular environments: the influence of altitude and seasonality

The seasonal diet and prey selection of the Southern Grey Shrike (Lanius meridionalis) was studied in two different insular habitats: shrub environments of the Canary Islands in coastal and high mountain zones. We measured, in each season, food availability and prey size in order to determine prey size selection of shrikes along an altitudinal gradient. Moreover, we compared the diet patterns observed with those documented on the continent, to determine if Southern Grey Shrikes in the islands’ high mountain zone (which has a continental climate) showed seasonal diet variation similar to those in northern continental areas. We analysed a total of 1,139 shrike pellets collected in 1 year and identified 10,179 prey items. Numerically arthropods (91%), and in terms of biomass lizards (70%) were the main prey consumed by the shrikes. The proportions of the main prey items differed significantly between seasons and habitats. Diet in the coastal areas was less variable than in the high mountain zone. The greater seasonal climatic variation in the high mountain zone was associated with diet patterns similar to those found in some northern continental areas, such as the Iberian Peninsula and southern France. Finally, shrikes selected the largest prey in the high mountain habitat. This suggests that foraging behaviour in this species is related to climatic conditions, as the biggest and most profitable prey were consumed in the most harsh habitats.     

Estimation of food intake and ingested energy in Daubenton’s bats (Myotis daubentonii) during pregnancy and spermatogenesis

We studied food intake of and estimated ingested energy in female and male Myotis daubentonii during the periods of pregnancy (period 1, 8 May–4 June) and of intense spermatogenetic activity (period 2, 24 July–22 August) over 8 years (1996–2003) in central Germany. We used radiotelemetry to determine the time spent foraging and marked animals with chemiluminescent light-sticks to determine prey attack rates. Body length, body mass, moisture content, and caloric content of chironomids, the main prey of Daubenton’s bats, were measured to estimate the nightly food intake and, in consequence, energy intake. Pregnant females spent significantly more time foraging than males during period 1 and females during the post-lactation period. In contrast, male foraged longer during the period of highest spermatogenetic activity than during late spring and also significantly longer than post-lactating females. Based on a mean number of 8.3 prey attacks per minute, the time spent foraging, and a capture success rate of either 50 or 92%, calculated intake values with a feeding rate of 7.6 insects per minute (=92% capture success) were more consistent with literature data for other insectivorous bats than that of values calculated on the basis of a capture success rate of 50%. In the high capture-success model, calculated insect intake of female bats was 8.0 g during pregnancy and 4.9 g per day during post-lactation, providing 5.0 and 3.0 kJ of ingested energy per gram body mass per day. Calculated intake of male bats was 3.6 g insects per day during late spring and 8.0 g during period of intensive spermatogenesis, providing 2.6 and 5.7 kJ of ingested energy per gram body mass.     

Pushed for time or saving on fuel: fine-scale energy budgets shed light on currencies in a diving bird

Animals may forage using different currencies depending on whether time minimization or energy maximization is more pertinent at the time. Assessment of net energy acquisition requires detailed information on instantaneous activity-specific power use, which varies according to animal performance, being influenced, for example, by speed and prey loading, and which has not been measured before in wild animals. We used a new proxy for instantaneous energy expenditure (overall dynamic body acceleration), to quantify foraging effort in a model species, the imperial shag Phalacrocorax atriceps, during diving. Power costs varied nonlinearly with depth exploited owing to depth-related buoyancy. Consequently, solutions for maximizing the gross rate of gain and energetic efficiency differed for dives to any given depth. Dive effort in free-ranging imperial shags measured during the breeding season was consistent with a strategy to maximize the gross rate of energy gain. We suggest that the divergence of time and energy costs with dive depth has implications for the measurement of dive efficiency across diverse diving taxa.     

Island/mainland body size differences in Australian varanid lizards

Island varanids seem to be an exception to the rule that territorial vertebrate taxa often become gigantic relative to mainland relatives when on islands, whereas non-territorial species become dwarfed (Case 1978). However, no systematic island/mainland studies have examined the empirical size trends in this group of carnivorous lizards. We perform such an analysis for the Australian region and critically evaluate various selective agents that might be responsible for size changes in several island populations. Insular gigantism occurs at least four times among the island populations examined. The magnitude of size change is positively correlated to prey abundance on the islands (as indirectly measured through a condition index of the lizards, essentially a measure of how fat they arc) and the size of prey: islands with large prey have large varanids and vice versa. Since the island population with the largest size change, the Reevesby Varanus rosenbergi, was introduced less than 100 years ago, these size changes can be quite rapid. This might indicate that selective coefficients are strong; however, we can not exclude the possibility that these size differences have no genetic component and simply reflect environmental differences in growth rate and shifts in age structure between island and mainland locations.     

Prey size selection and cannibalistic behaviour of juvenile barramundi Lates calcarifer

This study assessed the cannibalistic behaviour of juvenile barramundi Lates calcarifer and examined the relationship between prey size selection and energy gain of cannibals. Prey handling time and capture success by cannibals were used to estimate the ratio of energy gain to energy cost in prey selection. Cannibals selected smaller prey despite its capability of ingesting larger prey individuals. In behavioural analysis, prey handling time significantly increased with prey size, but it was not significantly affected by cannibal size. Conversely, capture success significantly decreased with the increase of both prey and cannibal sizes. The profitability indices showed that the smaller prey provides the most energy return for cannibals of all size classes. These results indicate that L. calcarifer cannibals select smaller prey for more profitable return. The behavioural analysis, however, indicates that L. calcarifer cannibals attack prey of all size at a similar rate but ingest smaller prey more often, suggesting that prey size selection is passively orientated rather than at the predator's choice. The increase of prey escape ability and morphological constraint contribute to the reduction of intracohort cannibalism as fish grow larger. This study contributes to the understanding of intracohort cannibalism and development of strategies to reduce fish cannibalistic mortalities.     

The influence of predation risk on diet selectivity: A theoretical analysis

Studies that have examined the effect of experimental increases in predation risk on diet selectivity have shown both decreased and increased diet selectivity. A possible explanation for these disparate results emerges from an examination of the prey sets used in these studies, which differed in the relationship between the values of risk components associated with the capture of different prey types (‘danger’) and their profitabilities. When less profitable prey were more dangerous, selectivity increased with predation risk. When prey were equally dangerous, selectivity did not change. Finally, when the more profitable prey were also more dangerous, selectivity decreased with risk. Here, we examine theoretically the influence of a forager's estimate of the probability that a predator is present (φ) on the selection of diets from prey sets with varying danger–profitability relationships. A dynamic programming model is used to determine the maximum attack time (or distance) for each of two types of prey, differing in their energetic content, for a range of forager energy state and φ levels. The diets which would result if foragers attacked prey according to the rules provided by the dynamic model are then determined. The model results indicate that the prey danger–profitability relationship determines the diet selectivity response to φ, confirming that variation in this relationship could be responsible for the range of experimental results. This revised version was published online in July 2006 with corrections to the Cover Date.     

Temporal Variation in Japanese Macaque Bodily Mass

We investigated the bodily mass of infants and postpartum adult females longitudinally in provisioned free-ranging Japanese macaque (Macaca fuscata) troops of the Takasakiyama Natural Zoo, Oita, Japan. We investigated whether the rate of mass change in postpartum adult females and infants is more strongly associated with season (or calendar date) as opposed to the length of time since delivery/birth. Focal adult females' postpartum mass ranged between 7 and 10 kg for a year after delivery. For postpartum adult females, calender date (season) influenced the rate of mass change more than the length of time since delivery did. Rate of mass change in postpartum adult females was high in October (autumn) and April and May (spring) and low between December and February (winter). This may reflect seasonality in energy and protein intake from natural foods. Infant mass increased steadily from birth. Mass of infants varied between individuals, some infants reached 2 kg by about 240 days of age, and others by about 400 days of age. It was unclear, however, whether the rate of mass change in infants was influenced more by age or calendar date (season). Differences in trends between adult females and infants may reflect, to some degree, differences in sources of energy intake, i.e. solid foods for adult females and suckling and solid foods for infants.     

Dynamic changes in prey choice by stickleback during simultaneous encounter with large prey

When a three-spined stickleback Gasterosteus aculeatus encountered prey simultaneously the probability of hanging and the median pursuit time were greater than when prey were encountered sequentially. During simultaneous prey encounter fish did not choose to attack the more profitable prey but instead the nearer prey was handled first except when the difference between the two prey sizes was large. No difference was found in the level of total energetic intake by the fish regardless of prey size pairing. Fish that handled and ate the first prey of a pair in <5 s attacked the second prey with a high probability of success, demonstrating an opportunistic feeding strategy. Importantly however, the fish did not choose to maximize long term energy intake rate by eating both prey, but rather short-term considerations over the course of feeding took precedence. With an empty stomach, the probability of a fish eating ( P _eat) the first prey handled was high regardless of prey size. As stomach fullness increased, the P _eat the first prey handled decreased if it was the larger prey. Hence, the fish were unselective when the stomach was empty but thereafter there was a shift in preference towards the smaller prey. The decision of which prey to attack and eat appeared to be based on short-term energy considerations and the level of stomach fullness. This study demonstrates that feeding on a short-term scale is a crucial factor to take account of when analysing fish feeding during simultaneous prey encounter.     

Functional and ecological relevance of intraspecific variation in body size and shape in the lizard Podarcis melisellensis (Lacertidae)

Within populations, individual animals may vary considerably in morphology and ecology. The degree to which variation in morphology is related to ecological variation within a population remains largely unexplored. We investigated whether variation in body size and shape among sexes and age classes of the lizard Podarcis melisellensis translates in differential whole-animal performance (sprint speed, bite force), escape and prey attack behaviour in the field, microhabitat use and diet. Male and female adult lizards differed significantly in body size and head and limb proportions. These morphological differences were reflected in differences in bite strength, but not in sprint speed. Accordingly, field measurements of escape behaviour and prey attack speed did not differ between the sexes, but males ate larger, harder and faster prey than females. In addition to differences in body size, juveniles diverged from adults in relative limb and head dimensions. These shape differences may explain the relatively high sprint and bite capacities of juvenile lizards. Ontogenetic variation in morphology and performance is strongly reflected in the behaviour and ecology in the field, with juveniles differing from adults in aspects of their microhabitat use, escape behaviour and diet.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 251–264.     

Changing handling times during feeding and consequences for prey size selection of 0+ zooplanktivorous fish

The interrelationship of fish size, prey size and handling time within a 15-min feeding period was studied in three size groups of 0 + roach, Rutilus rutilus, and bleak, Alburnus alburnus. Four size classes of cladoceran prey were used to measure changes in feeding rate and handling time from initial rapid feeding to sustained feeding. Observed differences in increase of handling time between prey size classes led to a change in the prey profitability ranking of those size classes within the first 2 min of the experiments. A 1-min feeding period is interpreted as reflecting an intermediate motivational status between extreme hunger and satiation. The use of average handling times for this period revealed a substantial change in prey profitability estimates compared to previous studies which used handling times based on short-term (a few seconds up to 1 min) feeding. It is not the largest prey items a fish can handle and swallow that are most profitable, but prey of intermediate size. By this approach a closer fit between expectations derived from optimal foraging theory and empirical data on prey size selection of 0 + zooplanktivorous fish is qualitatively achieved. Optimal prey size was found to be close the mouth gape width in small fish of 15 mm standard length, decreasing to 50% of mouth gape width in fish of 40 mm standard length.     

Evaluation of the sample size used for the rapid bioassessment of rivers using macroinvertebrates

Experiments were designed to investigate selective predation by medium (40–55 mm carapace width: CW) and large (55–70 mm CW) Carcinus maenas when feeding on four bivalves of contrasting shell morphology. Size-selection was examined by presenting individual crabs with a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas and Cerastoderma edule. Medium-sized crabs preferred mussels 5–15 mm shell length (maximum shell dimension: SL) and cockles 5–10 mm SL, whereas large crabs preferred mussels 15–25 mm and cockles 10–20 mm SL. Crabs generally showed no preference for any particular size of either oyster species. Species-selection was examined by presenting individual crabs with paired combinations of the four bivalves in various proportions. When offered mussels and oysters simultaneously, both size categories of crabs consistently selected mussels, and food choice was independent of prey relative abundance. By contrast, C. maenas selected mussels and cockles as expected by the frequency in which each size category of crab encountered the preferred size ranges of prey. Crab preference clearly paralleled the rank order of prey profitability, which in turn was mainly determined by prey biomass, suggesting that active selection takes place at some point of the predation cycle. Experiments with epoxy resin models showed that initial reluctance of crabs to attack oysters was not associated with the ultimate energy reward. Moreover, they suggest that foraging decisions are partly based on evaluations of overall prey shape and volume, and that the minimum dimension of the shell constitutes an important feature which crabs recognise and associate with prey value.     

Sexual and seasonal variation in the diet and foraging behaviour of a sexually dimorphic carnivore, the honey badger (Mellivora capensis)

The honey badger, or ratel, Mellivora capensis has not been well studied despite its extensive distribution. As part of the first detailed study, visual observations of nine habituated free-living individuals (five females, four males) were used to investigate seasonal, annual and sexual differences in diet and foraging behaviour. Theory predicts that generalist predators 'switch' between alternative prey species depending on which prey species are currently most abundant, and diet breadth expands in response to decreased availability of preferred food types. There were significant seasonal differences in the consumption of eight prey categories related to changes in prey availability but no seasonal differences in food intake per kg of body mass. As predicted, the cold-dry season diet was characterized by low species richness and low foraging yield but high dietary diversity, while the reverse was true in the hot-dry and hot-wet seasons. In accordance with these predictions, results suggest that the honey badger maintains its intake level by food switching and by varying dietary breadth. Despite marked sexual size dimorphism, male and female honey badgers showed no intersexual differences in prey size, digging success, daily food intake per unit body weight or foraging behaviour. Results do not support the hypothesis that size dimorphism is primarily an adaptation to reduce intersexual competition for food.     

Size-scaling of zooplankton foraging in Arctic charr

Prey capture rate (number of prey s⁻¹) and the mode of feeding of Arctic charr Salvelinus alpinus were studied by performing foraging experiments with two sizes (1·1 and 1·8 mm) of Daphnia longispina prey. Arctic charr were particulate feeders at all densities tested. Adjusted for the effect of prey density, the capture rate showed a hump-shaped relationship with Arctic charr size for both sizes of D. longispina . Estimated attack rates ( a ) also tended to show a hump-shaped relationship with fish size. The estimated size-scaling exponent of the attack rate function, however, was relatively small, implying small changes in attack rate over fish sizes. Simultaneous estimations of a and handling time were used in combination with published data on fish metabolism and dry mass rations of prey to estimate maintenance resource density of prey as a function of Arctic charr mass. Maintenance resource densities increased monotonically with Arctic charr size, and rapidly as optimum fish size relative to attack rate on prey was passed.     

The effects of fire on the frillneck lizard (Chlamydosaurus kingii) in northern Australia

Abstract A population of frillneck lizards, Chlamydosaurus kingii, was monitored by radio telemetry and mark-recapture techniques between April 1991 and April 1994, as part of a landscape-scale fire experiment, in Kakadu National Park, Northern Territory. The study aimed to investigate both the short- and longer-term effects of fire on a lizard species in a tropical savanna where fires are frequent and often annual. Frillneck lizards are able to survive fires that occur in the first few months of the dry season by remaining perched in trees. A high level of mortality (29%) occurred during late dry-season fires, along with changes in their behavioural response to fire: sheltering in either larger trees or hollow termite mounds. Food is more accessible after fires due to the removal of ground vegetation. This is reflected in greater volume and diversity of prey in stomach contents after fires. This increase is more pronounced after late dry-season fires, possibly due to increased accessibility of prey caused by more complete vegetation removal. Frillneck lizards show an overall preference for trees with a dense canopy cover located in an area with a low density of grass. Fire has an effect on this relationship. Frillneck lizards in habitat unburnt for a number of years tend to perch in trees with a smaller canopy, whereas lizards in annually burnt habitat perch in trees with a dense canopy. Volume and composition of lizard stomach contents was broadly similar among fire treatments over a 2 year period, although termites were more predominant in stomach contents of lizards in unburnt habitat. Wet-season body condition is lower in lizards from unburnt habitat, although the reason for this is unclear. These results demonstrate the importance of different fire intensities and regimes on the ecology of a lizard species in a tropical savanna.     

The ecological cost of morphological specialization: feeding by a fossorial lizard

Summary Head size and shape of reptiles may reflect selection for multiple uses. For example, sexual selection for large head size may enhance feeding efficiency. In contrast, morphological characteristics of the heads of fossorial reptiles suggests that fossoriality may have evolved at the expense of reduced effectiveness in feeding. Our research focused on the question: Does a fossorial lizard feed less effectively than a non-fossorial lizard? To answer this question, we measured the time, number of bites, and oxygen consumption by sand-swimming (Chalcides ocellatus) and epigeal (Eumeces inexpectatus) skinks feeding on crickets. These lizard species were similar in mass, but different in body form: Chalcides had longer bodies and smaller heads than Eumeces. For lizards of the same mass, Chalcides were unable to eat prey as large as those eaten by Eumeces, Chalcides took longer to eat prey of the same size than did Eumeces, and the aerobic energy cost of eating crickets of the same relative size (cricket mass/lizard mass) tended to be greater for Chalcides than for Eumeces. The ecologically relevant costs of feeding appear to be the upper limit to the size of prey and the time of feeding. Both costs would restrict the energy intake per unit time of Chalcides. Moreover, given the same energy requirements and prey community, Chalcides would have to feed more often and would take longer to feed than would Eumeces. Both factors would increase the exposure of Chalcides to predators relative to that of Eumeces. To reduce the risk of predation, Chalcides would have to reduce energy intake or fulfill its energy requirements with relatively small prey, or both. These conclusions are potentially confounded in two ways. The first is that male Eumeces have relatively large heads as a result of sexual selection. Thus, the differences we observed between Chalcides and Eumeces (most of our specimens were males) could have been the result of reduced costs of feeding for Eumeces due to sexual selection and not the result of enhanced costs of feeding for Chalcides. A more likely explanation is that differences in the costs of feeding observed between these species reflect adaptations for fossoriality by Chalcides and sexual selection on Eumeces. Our results may also be confounded because we compared laboratory reared Chalcides with field captured Eumeces. Any deterimental effects of captivity on the vigor of Chalcides would increase their costs of feeding relative to those of Eumeces. Although short-term captivity is not associated with changes in the metabolic capcity of lizards, effects of long-term captivitiy are unknown.     

Adaptive significance of food income in European snakes: body size is related to prey energetics

Feeding strategies and diet patterns have been extensively investigated in vertebrates and, more specifically, in snakes. Although it has been hypothesized that prey species may differ in terms of energy content, almost no theoretical or practical study has been carried out to determine actual nutritional values of the common prey types of wild snakes. Our model taxa were a selection of widely distributed and well known European snake species, which have all been studied in depth: approximately 76% of their diet is composed of mammals, reptiles, and insects. We therefore selected a single model species for each of these categories and proceeded with the analyses. Nutritional values were determined using a standard procedure: lizards and mice were richer in proteins than insects (crickets); insects and mice were richer in lipids than lizards, and mice and crickets have a higher energy content than lizards; lizards were rich in ashes. We then applied our experimental results to a selected sample of European terrestrial snakes (11 populations, ten species, seven genera, two families) characterized by different body size (50–160 cm total length) and reproductive strategies (oviparous versus viviparous), aiming to correlate these parameters with patterns of energy income. A direct relationship was found between body mass/body length ratio (BCI, body condition index) and meal energetics: the higher the BCI, the higher was the metabolic requirement, whereas BCI was independent of species or of reproductive system effect. Large‐sized snakes thus need a highly diversified and more energy‐rich diet than smaller snakes, supporting previous hypotheses. The simple applicability of this method could be of valuable support in further comparative research work, reducing experimental costs and stimulating further ecological, behavioural, and, possibly, phylogenetic comparisons. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 307–317.