The Effect of Digestive Capacity on the Intake Rate of Toxic and Non-Toxic Prey in an Ecological Context

Digestive capacity often limits food intake rate in animals. Many species can flexibly adjust digestive organ mass, enabling them to increase intake rate in times of increased energy requirement and/or scarcity of high-quality prey. However, some prey species are defended by secondary compounds, thereby forcing a toxin limitation on the forager’s intake rate, a constraint that potentially cannot be alleviated by enlarging digestive capacity. Hence, physiological flexibility may have a differential effect on intake of different prey types, and consequently on dietary preferences. We tested this effect in red knots (Calidris canutus canutus), medium-sized migratory shorebirds that feed on hard-shelled, usually mollusc, prey. Because they ingest their prey whole and crush the shell in their gizzard, the intake rate of red knots is generally constrained by digestive capacity. However, one of their main prey, the bivalve Loripes lucinalis, imposes a toxin constraint due to its symbiosis with sulphide-oxidizing bacteria. We manipulated gizzard sizes of red knots through prolonged exposure to hard-shelled or soft foods. We then measured maximum intake rates of toxic Loripes versus a non-toxic bivalve, Dosinia isocardia. We found that intake of Dosinia exponentially increased with gizzard mass, confirming earlier results with non-toxic prey, whereas intake of Loripes was independent of gizzard mass. Using linear programming, we show that this leads to markedly different expected diet preferences in red knots that try to maximize energy intake rate with a small versus a large gizzard. Intra- and inter-individual variation in digestive capacity is found in many animal species. Hence, the here proposed functional link with individual differences in foraging decisions may be general. We emphasize the potential relevance of individual variation in physiology when studying trophic interactions.     

Anti-apostatic selection by wild birds on quasi-natural morphs of the land snail Cepaea hortensis: a generalised linear mixed models approach

Anti-apostatic selection occurs when predators feed disproportionately on rare prey, a process that leads to a decrease in variation within prey populations. Most of the evidence for this phenomenon comes from experiments using artificial prey items distinguished by simple artificial colour differences. We describe an experiment performed at 10 different sites in which we tested whether selection by wild birds is anti-apostatic when presented with high density populations of shells of the polymorphic land snail, Cepaea hortensis , stuffed with pastry. At each site we presented two sorts of populations in sequence: 45 yellow unbanded with 5 yellow banded, and 5 yellow unbanded with 45 yellow banded. Selection was measured using an analysis based on generalised linear mixed models (GLMMs) that has broad applicability to all resource selection studies where extraneous variance is a problem. Using this analytic method, we found that birds altered their behaviour to produce significant anti-apostatic selection in direct response to changes in frequency of the quasi-natural Cepaea morphs.     

Viewing distance affects how the presence of inedible models influence the benefit of masquerade

Masquerading prey closely resemble inedible objects found in the same locality. These animals gain protection from their predators by causing their predators to misclassify them as the inedible ‘models’ that they appear to resemble. We recently demonstrated that predators are more likely to misclassify masquerading prey as their models when masqueraders are viewed in isolation from their models than when they are viewed simultaneously with examples of their models. Using domestic chicks (Gallus gallus domesticus) as predators and the twig-mimicking caterpillars of the Early Thorn Moth (Selenia dentaria) as prey, we tested whether this effect was influenced by the relative orientations of models and masqueraders; and the distance from which models and masqueraders could be viewed simultaneously. We found no effect of orientation, but that the cost to masqueraders of being viewed simultaneously with an example of the model declined as the distance between the model and masquerader increased. These results are interpreted in terms of animal cognition, and their implications for the evolutionary ecology of masquerade.     

Incompletely informed shorebirds that face a digestive constraint maximize net energy gain when exploiting patches

Foragers that feed on hidden prey are uncertain about the intake rate they can achieve as they enter a patch. However, foraging success can inform them, especially if they have prior knowledge about the patch quality distribution in their environment. We experimentally tested whether and how red knots (Calidris canutus) use such information and whether their patch-leaving decisions maximized their long-term net energy intake rate. The results suggest that the birds combined patch sample information with prior knowledge by making use of the potential value assessment rule. We reject five alternative leaving rules. The potential encounter rate that the birds choose as their critical departure threshold maximized their foraging gain ratio (a modified form of efficiency) while foraging. The high experimental intake rates were constrained by rate of digestion. Under such conditions, maximization of the foraging gain ratio during foraging maximizes net intake rate during total time (foraging time plus digestive breaks). We conclude that molluscivore red knots, in the face of a digestive constraint, are able to combine prior environmental knowledge about patch quality with patch sample information to obtain the highest possible net intake over total time.     

Carrying capacity models should not use fixed prey density thresholds: a plea for using more tools of behavioural ecology

Earlier studies have developed models of carrying capacity to predict the number of animals a certain area can support. These models assume that resources are not renewed after consumption ('standing stock' models), and that the initial number of prey and the rate of prey consumption determine the time a population of foragers can live in an area. Within such areas, foragers give up feeding at a sub-site or patch when intake rates no longer cover energy expenditure. To improve the success rate of the models' predictions, we here change the existing rate-maximising models into fitness-maximising models, and include dynamics in the availability of patches. These new (conceptual) models show that the approaches used so far may over- as well as underestimate carrying capacity. We review empirical studies that have aimed to estimate carrying capacity, and discuss how concepts have been confused. We make explicit suggestions on how to proceed in predicting carrying capacities in future studies.     

Isolating the cow-specific part of residual energy intake in lactating dairy cows using random regressions

The ability to properly assess and accurately phenotype true differences in feed efficiency among dairy cows is key to the development of breeding programs for improving feed efficiency. The variability among individuals in feed efficiency is commonly characterised by the residual intake approach. Residual feed intake is represented by the residuals of a linear regression of intake on the corresponding quantities of the biological functions that consume (or release) energy. However, the residuals include both, model fitting and measurement errors as well as any variability in cow efficiency. The objective of this study was to isolate the individual animal variability in feed efficiency from the residual component. Two separate models were fitted, in one the standard residual energy intake (REI) was calculated as the residual of a multiple linear regression of lactation average net energy intake (NEI) on lactation average milk energy output, average metabolic BW, as well as lactation loss and gain of body condition score. In the other, a linear mixed model was used to simultaneously fit fixed linear regressions and random cow levels on the biological traits and intercept using fortnight repeated measures for the variables. This method split the predicted NEI in two parts: one quantifying the population mean intercept and coefficients, and one quantifying cow-specific deviations in the intercept and coefficients. The cow-specific part of predicted NEI was assumed to isolate true differences in feed efficiency among cows. NEI and associated energy expenditure phenotypes were available for the first 17 fortnights of lactation from 119 Holstein cows; all fed a constant energy-rich diet. Mixed models fitting cow-specific intercept and coefficients to different combinations of the aforementioned energy expenditure traits, calculated on a fortnightly basis, were compared. The variance of REI estimated with the lactation average model represented only 8% of the variance of measured NEI. Among all compared mixed models, the variance of the cow-specific part of predicted NEI represented between 53% and 59% of the variance of REI estimated from the lactation average model or between 4% and 5% of the variance of measured NEI. The remaining 41% to 47% of the variance of REI estimated with the lactation average model may therefore reflect model fitting errors or measurement errors. In conclusion, the use of a mixed model framework with cow-specific random regressions seems to be a promising method to isolate the cow-specific component of REI in dairy cows.     

The costs of becoming nocturnal: feeding efficiency in relation to light intensity in juvenile Atlantic Salmon

1. Most animals are active by day or by night, but not both; juvenile salmonids are unusual in that they switch from being predominantly diurnal for most of the year to being nocturnal in winter. They are visual foragers, and adaptations for high visual acuity at daytime light intensities are generally incompatible with sensitive night vision. Here we test whether juvenile Atlantic Salmon Salmo salar are able to maintain their efficiency of prey capture when switching between diurnal and nocturnal foraging.
2. By testing the ability of the fish to acquire drifting food items under a range of manipulated light intensities, we show that the foraging efficiency of juvenile salmon is high at light intensities down to those equivalent to dawn or dusk, but drops markedly at lower levels of illumination: even under the best night condition (full moon and clear sky), the feeding efficiency is only 35% of their diurnal efficiency, and fish will usually be feeding at less than 10% (whenever the moon is not full, skies are overcast or when in the shade of bankside trees). Fish were unable to feed on drifting prey when in complete darkness.
3. The ability of juvenile salmon to detect prey under different light intensities is similar to that of other planktivorous or drift-feeding species of fish; they thus appear to have no special adaptations for nocturnal foraging.
4. While winter drift abundance is slightly higher by night than by day, the difference is not enough to compensate for the loss in foraging efficiency. We suggest that juvenile salmon can nonetheless switch to nocturnal foraging in winter because their food requirements are low, many individuals adopting a strategy in which intake is suppressed to the minimum that ensures survival.     

Elimination of oxalate by fat sand rats (Psammomys obesus): wild and laboratory-bred animals compared

Wild fat sand rats (Psammomys obesus) can feed exclusively on plants containing much oxalate, but little calcium; oxalate intake may exceed 300 mg/d, while calcium intake is approximately 30 mg/day. By contrast, for generations, laboratory bred P. obesus have been fed a low-oxalate (<100 mg/day), high-calcium (approximately 150 mg/day) rodent chow. We compared oxalate intake and excretion between wild and laboratory-bred animals, both fed the natural high-oxalate diet, to determine whether these different dietary histories are reflected in the animal's ability to eliminate dietary oxalate. Since both wild and laboratory-bred P. obesus harbor intestinal oxalate-degrading bacteria, we predicted that their oxalate intake and excretion would be similar. Indeed, we found no significant differences in oxalate intake or excretion between the groups fed either saltbush or alfalfa (p>0.05). However, due to the differences in dietary calcium intake between the two diets, in both groups only part (23-25%) of the ingested oxalate was excreted when the animals were fed the oxalate-rich saltbush, yet most (87-90%) was excreted when feeding on calcium-rich alfalfa. Thus, even after generations of feeding on a commercial low-oxalate diet, fat sand rats maintain intestinal oxalate-degrading bacteria that appear to increase in number and activity when presented with their natural diet.     

Better the devil you know: avian predators find variation in prey toxicity aversive

Toxic prey that signal their defences to predators using conspicuous warning signals are called ‘aposematic’. Predators learn about the toxic content of aposematic prey and reduce their attacks on them. However, through regulating their toxin intake, predators will include aposematic prey in their diets when the benefits of gaining the nutrients they contain outweigh the costs of ingesting the prey''s toxins. Predators face a problem when managing their toxin intake: prey sharing the same warning signal often vary in their toxicities. Given that predators should avoid uncertainty when managing their toxin intake, we tested whether European starlings (Sturnus vulgaris) preferred to eat fixed-defence prey (where all prey contained a 2% quinine solution) to mixed-defence prey (where half the prey contained a 4% quinine solution and the other half contained only water). Our results support the idea that predators should be more ‘risk-averse’ when foraging on variably defended prey and suggest that variation in toxicity levels could be a form of defence.     

Optimal Bayesian foraging policies and prey population dynamics-some comments on Rodriguez-Girones and Vasquez

In this paper we show the density-dependent harvest rates of optimal Bayesian foragers exploiting prey occurring with clumped spatial distribution. Rodríguez-Gironés and Vásquez (1997) recently treated the issue, but they used a patch-leaving rule (current value assessment rule) that is not optimal for the case described here. An optimal Bayesian forager exploiting prey whose distribution follows the negative binomial distribution should leave a patch when the potential (and not instantaneous) gain rate in that patch equals the best long-term gain rate in the environment (potential value assessment rule). It follows that the instantaneous gain rate at which the patches are abandoned is an increasing function of the time spent searching in the patch. It also follows that the proportion of prey harvested in a patch is an increasing sigmoidal function of the number of prey initially present. In this paper we vary several parameters of the model to evaluate the effects on the forager's intake rate, the proportion of prey harvested per patch, and the prey's average mortality rate in the environment. In each case, we study an intake rate maximizing forager's optimal response to the parameter changes. For the potential value assessment rule we find that at a higher average prey density in the environment, a lower proportion of the prey is taken in a patch with a given initial prey density. The proportion of prey taken in a patch of a given prey density also decreases when the variance of the prey density distribution is increased and if the travel time between patches is reduced. We also evaluate the effect of using predation minimization, rather than rate maximization, as the currency. Then a higher proportion of the prey is taken for each given initial prey density. This is related to the assumption that traveling between patches is the most risky activity. Compared to the optimal potential value assessment rule, the current value assessment rule performs worse, in terms of long-term intake rate achieved. The difference in performance is amplified when prey density is high or highly aggregated. These results pertain to the foraging patch spatial scale and may have consequences for the spatial distribution of prey in the environment.     

Tolerance of understory plants subject to herbivory by roe deer

Classical foraging theory states that animals feeding in a patchy environment can maximise their long term prey capture rates by quitting food patches when they have depleted prey to a certain threshold level. Theory suggests that social foragers may be better able to do this if all individuals in a group have access to the prey capture information of all other group members. This will allow all foragers to make a more accurate estimation of the patch quality over time and hence enable them to quit patches closer to the optimal prey threshold level. We develop a model to examine the foraging efficiency of three strategies that could be used by a cohesive foraging group to initiate quitting a patch, where foragers do not use such information, and compare these with a fourth strategy in which foragers use public information of all prey capture events made by the group. We carried out simulations in six different prey environments, in which we varied the mean number of prey per patch and the variance of prey number between patches. Groups sharing public information were able to consistently quit patches close to the optimal prey threshold level, and obtained constant prey capture rates, in groups of all sizes. In contrast all groups not sharing public information quit patches progressively earlier than the optimal prey threshold value, and experienced decreasing prey capture rates, as group size increased. This is more apparent as the variance in prey number between patches increases. Thus in a patchy environment, where uncertainty is high, although public information use does not increase the foraging efficiency of groups over that of a lone forager, it certainly offers benefits over groups which do not, and particularly where group size is large.     

Gluttonous predators: how to estimate prey size when there are too many prey

Prey size is an important factor in food consumption. In studies of feeding ecology, prey items are usually measured individually using calipers or ocular micrometers. Among amphibians and reptiles, there are species that feed on large numbers of small prey items (e.g. ants, termites). This high intake makes it difficult to estimate prey size consumed by these animals. We addressed this problem by developing and evaluating a procedure for subsampling the stomach contents of such predators in order to estimate prey size. Specifically, we developed a protocol based on a bootstrap procedure to obtain a subsample with a precision error of at the most 5%, with a confidence level of at least 95%. This guideline should reduce the sampling effort and facilitate future studies on the feeding habits of amphibians and reptiles, and also provide a means of obtaining precise estimates of prey size.     

Repeatability of traits for characterizing feed intake patterns in dairy goats: a basis for phenotyping in the precision farming context

In ruminants, feeding behaviour variables are parameters involved in feed efficiency that show variation among individuals. This study aimed to evaluate during the first two production cycles in ruminants the repeatability of feed intake pattern, which is an important aspect of feeding behaviour. Thirty-five dairy goats from Alpine or Saanen breeds were housed in individual pens at four periods (end of first gestation, middle of first and second lactations and middle of second gestation which is also the end of first lactation) and fed a total mixed ration (TMR) ad libitum. Individual cumulative dry matter intake (DMI) was automatically measured every 2 min during the last 4 days of each period. Feed intake pattern was characterized by several measures related to the quantity of feed eaten or to the rate of intake during the 15 h following the afternoon feed delivery. Two main methods were used: modelling cumulative DMI evolution by an exponential model or by a segmentation-clustering method. The goat ability to sort against dietary fibre was also evaluated. There was a very good repeatability of the aggregate measures between days within a period for a given goat estimated by the day effect within breed and goat, tested on the residual variance (P > 0.95). The correlations between periods were the highest between the second and either the third or fourth periods. With increasing age, goats sorted more against the fibrous part of the TMR and increased their initial rate of intake. Alpine goats ate more slowly than Saanen goats but ate during a longer duration. Principal component analysis (PCA) was performed on all the aggregate measures of feed intake patterns. The factor score plots generated by the PCA highlighted the opposition between the different measures of feed intake patterns and the sorting behaviour. The projection of the animals on the scoring plots showed a breed effect and that there was a continuum for the feed intake pattern of goats. In conclusion, this study showed that the feed intake pattern was highly repeatable for an animal in a given period and between periods. This means that phenotyping goats in a younger age might be of interest, either to select them on feeding behaviour and choose preferentially the slow eaters or to adapt the quantity offered and restrict feed delivery to the fast eaters in order to increase feed efficiency and welfare by limiting the occurrence of acidosis, for example.     

Optimal Bayesian Foraging Policies and Prey Population Dynamics—Some Comments on Rodríguez-Gironés and Vásquez

In this paper we show the density-dependent harvest rates of optimal Bayesian foragers exploiting prey occurring with clumped spatial distribution. Rodríguez-Gironés and Vásquez (1997) recently treated the issue, but they used a patch-leaving rule (current value assessment rule) that is not optimal for the case described here. An optimal Bayesian forager exploiting prey whose distribution follows the negative binomial distribution should leave a patch when the potential (and not instantaneous) gain rate in that patch equals the best long-term gain rate in the environment (potential value assessment rule). It follows that the instantaneous gain rate at which the patches are abandoned is an increasing function of the time spent searching in the patch. It also follows that the proportion of prey harvested in a patch is an increasing sigmoidal function of the number of prey initially present. In this paper we vary several parameters of the model to evaluate the effects on the forager's intake rate, the proportion of prey harvested per patch, and the prey's average mortality rate in the environment. In each case, we study an intake rate maximizing forager's optimal response to the parameter changes. For the potential value assessment rule we find that at a higher average prey density in the environment, a lower proportion of the prey is taken in a patch with a given initial prey density. The proportion of prey taken in a patch of a given prey density also decreases when the variance of the prey density distribution is increased and if the travel time between patches is reduced. We also evaluate the effect of using predation minimization, rather than rate maximization, as the currency. Then a higher proportion of the prey is taken for each given initial prey density. This is related to the assumption that traveling between patches is the most risky activity. Compared to the optimal potential value assessment rule, the current value assessment rule performs worse, in terms of long-term intake rate achieved. The difference in performance is amplified when prey density is high or highly aggregated. These results pertain to the foraging patch spatial scale and may have consequences for the spatial distribution of prey in the environment.     

Do cattle egrets gain information from conspecifics when foraging?

Summary We examined whether individual cattle egrets (Bubulcus ibis) base their decisions of where to forage, and how long to stay in a patch, on the behavior of other flock members. Cattle egrets commonly forage in flocks associated with cattle and capture prey at higher rates when they do not share a cow with another egret. Foraging egrets provide cues of the location of prey and their success in capturing prey. Therefore, there is the possibility of information transfer between egrets in a flock. We predicted that egrets should only move to occupied patches when the resident was capturing enough prey that it is profitable for the invader to share the patch or take over the patch. However, egrets did not seem to decide where to forage based on neighbors' rates of energy intake, but rather on the presence or absence of conspecifics in a patch. We also predicted that an egret should remain in a patch until its rate of energy intake dropped to or below the average rate for other egrets within the flock. However, egrets that were foraging more efficiently than the average rate for the flock switched patches sooner than less efficient foragers. Egrets did not appear to increase foraging success by gaining information on patch quality from neighbors.     

Flexible search tactics and efficient foraging in saltatory searching animals

Summary Foraging is one of the most important endeavors undertaken by animals, and it has been studied intensively from both mechanistic-empirical and optimal foraging perspectives. Planktivorous fish make excellent study organisms for foraging studies because they feed frequently and in a relatively simple environment. Most optimal foraging studies of planktivorous fish have focused, either on diet choice or habitat selection and have assumed that these animals used a cruise search foraging strategy. We have recently recognized that white crappie do not use a cruise search strategy (swimming continuously and searching constantly) while foraging on zooplankton but move in a stop and go pattern, searching only while paused. We have termed thissaltatory search. Many other animals move in a stop and go pattern while foraging, but none have been shown to search only while paused. Not only do white crappie search in a saltatory manner but the components of the search cycle change when feeding on prey of different size. When feeding on large prey these fish move further and faster after an unsuccessful search than when feeding on small prey. The fish also pause for a shorter period to search when feeding on large prey. To evaluate the efficiency of these alterations in the search cycle, a net energy gain simulation model was developed. The model computes the likelihood of locating 1 or 2 different size classes of zooplankton prey as a function of the volume of water scanned. The volume of new water searched is dependent upon the dimensions of the search volume and the length of the run. Energy costs for each component of the search cycle, and energy gained from the different sized prey, were assessed. The model predicts that short runs produce maximum net energy gains when crappie feed on small prey but predicts net energy gains will be maximized with longer runs when crappie feed on large prey or a mixed assemblage of large and small prey. There is an optimal run length due to high energy costs of unsuccessful search when runs are short and reveal little new water, and high energy costs of long runs when runs are lengthy. The model predicts that if the greater search times observed when crappie feed on small prey are assessed when they feed on a mixed diet of small and large prey, net energy gained is less than if small prey are deleted from the diet. We believe the model has considerable generality. Many animals are observed to move in a saltatory manner while foraging and some are thought to search only while stationary. Some birds and lizards are, known to modify the search cycle in a manner similar to white crappie.Components of the search cycle and dimensions of the location space SST (sec) Successful search time — the average time stationary prior to a pursuit - USST (sec) Unsuccessful search time — the average time stationary prior to a run - PT (sec) Pursuit time-PL/SS — the time to pursue prey at a given distance away. It is calculated by dividing the pursuit distance by swim speed - RT (sec) Run time-RL/SS — the time to complete a run of a given length. It is calculated by dividing the run length, by swim speed - PL (cm) Pursuit length-distance moved to attack prey - RL (cm) Run length-distance moved between consecutive searches - SS (cm/sec) Swim speed — the speed of movement during a pursuit or run - LS (l) Location space — the area or volume within which prey are located. In the case of white crappie the search space is shaped like a pie wedge with the fish positioned at the apex of the wedge - LA (^o) Location angle—the angle of the wedge-shaped search space - LH (cm) Location height—the height of the wedge-shaped search space - LD (cm) Location distance—the length of long axis of the wedge-shaped search space. Components of the location probability model RND Random number-random number generated through BASICA - SV (l) Search volume—the volume of water actually searched after one run of given length - SV_MAX (l) Maximum search volume—the greatest search volume that can be based upon LA, LH, LD and unaffected by the previous search - SV_R (l) Search volume researched—that volume of SV_MAX that is researched where RLo Search volume unsearched—that volume of SV_MAX not previously searched - AD (#/1) Absolute density—the density of zooplankton prey in numbers per liter - VD (#) Visual density—the number of zooplankton prey in the search volume - LP (%) Location probability—the probability that one or more prey are in the search volume Components of the net energy gain model NEG (cal/sec) Net energy gain-total calories ingested, less total calories used, divided by total time. - E _e (cal) Energy expended on the search cycle - E _i (cal) Energy intake - e _p (cal) Energy content of a given individual prey - P _i Total number of prey ingested - e _r (cal) Energy expended while searching - e _s (cal) Energy expended while swimming - T _t (sec) Total time-time expended to eat a given number of prey     

What determines prey selection in owls? Roles of prey traits,prey class,environmental variables,and taxonomic specialization

Ecological theory suggests that prey size should increase with predator size, but this trend may be masked by other factors affecting prey selection, such as environmental constraints or specific prey preferences of predator species. Owls are an ideal case study for exploring how predator body size affects prey selection in the presence of other factors due to the ease of analyzing their diets from owl pellets and their widespread distributions, allowing interspecific comparisons between variable habitats. Here, we analyze various dimensions of prey resource selection among owls, including prey size, taxonomy (i.e., whether or not particular taxa are favored regardless of their size), and prey traits (movement type, social structure, activity pattern, and diet). We collected pellets of five sympatric owl species (Athene noctua, Tyto alba, Asio otus, Strix aluco, and Bubo bubo) from 78 sites across the Mediterranean Levant. Prey intake was compared between sites, with various environmental variables and owl species as predictors of abundance. Despite significant environmental impacts on prey intake, some key patterns emerge among owl species studied. Owls select prey by predator body size: Larger owls tend to feed on wider ranges of prey sizes, leading to higher means. In addition, guild members show both specialization and generalism in terms of prey taxa, sometimes in contrast with the expectations of the predator–prey body size hypothesis. Our results suggest that while predator body size is an important factor in prey selection, taxon specialization by predator species also has considerable impact.     

Intake rates and the functional response in shorebirds (Charadriiformes) eating macro-invertebrates

As field determinations take much effort, it would be useful to be able to predict easily the coefficients describing the functional response of free-living predators, the function relating food intake rate to the abundance of food organisms in the environment. As a means easily to parameterise an individual-based model of shorebird Charadriiformes populations, we attempted this for shorebirds eating macro-invertebrates. Intake rate is measured as the ash-free dry mass (AFDM) per second of active foraging; i.e. excluding time spent on digestive pauses and other activities, such as preening. The present and previous studies show that the general shape of the functional response in shorebirds eating approximately the same size of prey across the full range of prey density is a decelerating rise to a plateau, thus approximating the Holling type II ('disc equation') formulation. But field studies confirmed that the asymptote was not set by handling time, as assumed by the disc equation, because only about half the foraging time was spent in successfully or unsuccessfully attacking and handling prey, the rest being devoted to searching.A review of 30 functional responses showed that intake rate in free-living shorebirds varied independently of prey density over a wide range, with the asymptote being reached at very low prey densities (<150/m-2). Accordingly, most of the many studies of shorebird intake rate have probably been conducted at or near the asymptote of the functional response, suggesting that equations that predict intake rate should also predict the asymptote.A multivariate analysis of 468 'spot' estimates of intake rates from 26 shorebirds identified ten variables, representing prey and shorebird characteristics, that accounted for 81% of the variance in logarithm-transformed intake rate. But four-variables accounted for almost as much (77.3%), these being bird size, prey size, whether the bird was an oystercatcher Haematopus ostralegus eating mussels Mytilus edulis, or breeding. The four variable equation under-predicted, on average, the observed 30 estimates of the asymptote by 11.6%, but this discrepancy was reduced to 0.2% when two suspect estimates from one early study in the 1960s were removed. The equation therefore predicted the observed asymptote very successfully in 93% of cases. We conclude that the asymptote can be reliably predicted from just four easily measured variables. Indeed, if the birds are not breeding and are not oystercatchers eating mussels, reliable predictions can be obtained using just two variables, bird and prey sizes. A multivariate analysis of 23 estimates of the half-asymptote constant suggested they were smaller when prey were small but greater when the birds were large, especially in oystercatchers. The resulting equation could be used to predict the half-asymptote constant, but its predictive power has yet to be tested. As well as predicting the asymptote of the functional response, the equations will enable research workers engaged in many areas of shorebird ecology and behaviour to estimate intake rate without the need for conventional time-consuming field studies, including species for which it has not yet proved possible to measure intake rate in the field.     

Lack of an osmotic constraint on intake rate of the eastern curlew Numenius madagascariensis

Rates of food intake in animals consuming abundant prey can be constrained by the rates of digestion or excretion of ingested substances, such as salt, particularly so in the animals that regularly migrate between freshwater and saltwater environments. We tested this hypothesis in a long-distance migrant shorebird, the eastern curlew Numenius madagascariensis (suborder Charadrii), foraging on intertidal decapods in eastern Australia. We predicted that if food intake rates are constrained osmotically, individuals with access to freshwater and less saline prey (FW group) would have higher rates of food and water intake than individuals with seawater-only access (SW group). Food intake rates did not differ between the FW and SW groups (0.14 g ash-free dry mass min⁻¹), nor did the water influx rates (0.75 g  min⁻¹). Salt intake rates were lower at FW sites (19.3 versus 23.3 mg NaCl min⁻¹) and overall they were similar to those of marine birds. Food intake rate in the eastern curlew appeared limited by digestive rather than by osmoregulatory capacity.     

The costs of carnivory

Mammalian carnivores fall into two broad dietary groups: smaller carnivores (<20 kg) that feed on very small prey (invertebrates and small vertebrates) and larger carnivores (>20 kg) that specialize in feeding on large vertebrates. We develop a model that predicts the mass-related energy budgets and limits of carnivore size within these groups. We show that the transition from small to large prey can be predicted by the maximization of net energy gain; larger carnivores achieve a higher net gain rate by concentrating on large prey. However, because it requires more energy to pursue and subdue large prey, this leads to a 2-fold step increase in energy expenditure, as well as increased intake. Across all species, energy expenditure and intake both follow a three-fourths scaling with body mass. However, when each dietary group is considered individually they both display a shallower scaling. This suggests that carnivores at the upper limits of each group are constrained by intake and adopt energy conserving strategies to counter this. Given predictions of expenditure and estimates of intake, we predict a maximum carnivore mass of approximately a ton, consistent with the largest extinct species. Our approach provides a framework for understanding carnivore energetics, size, and extinction dynamics.