Measuring the functional responses of farmland birds: an example for a declining seed-feeding bunting

1. Many farmland bird species have undergone significant declines. It is important to predict the effect of agricultural change on these birds and their response to conservation measures. This requirement could be met by mechanistic models that predict population size from the optimal foraging behaviour and fates of individuals within populations. A key component of these models is the functional response, the relationship between food and competitor density and feeding rate. 2. This paper describes a method for measuring functional responses of farmland birds, and applies this method to a declining farmland bird, the corn bunting Miliaria calandra L. We derive five alternative models to predict the functional responses of farmland birds and parameterize these for corn bunting. We also assess the minimum sample sizes required to predict accurately the functional response. 3. We show that the functional response of corn bunting can be predicted accurately from a few behavioural parameters (searching rate, handling time, vigilance time) that are straightforward to measure in the field. These parameters can be measured more quickly than the alternative of measuring the functional response directly. 4. While corn bunting violated some of the assumptions of Holling's disk equation (model 1 in our study), it still provided the most accurate fit to the observed feeding rates while remaining the most statistically simple model tested. Our other models may be more applicable to other species, or corn bunting feeding in other locations. 5. Although further tests are required, our study shows how functional responses can be predicted, simplifying the development of mechanistic models of farmland bird populations.     

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.     

Shape and sources of variations of the functional response of wildfowl: an experiment with mallards, Anas platyrhynchos

Understanding the variations of the functional response of an organism, i.e. the predation rate in relation to prey density, is necessary to understand the interactions between the animal and its food supply. This has received little attention in dabbling ducks so we investigated experimentally the shape of the functional response of mallard feeding on poultry pellets, and assessed the influence of several factors such as the size of food items, sex or individual performance on this functional response. Individual differences in intake rate are of crucial importance in group or gregarious foraging species. We used two approaches of the functional response: 1) the relation between feeding rate (pellets/s) and pellet densities (pellets/m²), and 2) the relationship between instantaneous intake rate (g/s) and biomass density (g/m²). For both approaches, we found that the Type II functional response gave better estimates than a Type I linear functional response but explained only a third of the variance. Our results show that pellet size has a large effect on instantaneous intake rate. The comparison of the functional response parameters suggest that handling time per prey may not reflect the real constraints on intake rate, but that handling time per gram ingested may be more appropriate to integrate the effect of item size in the functional response. We then discuss the possible mechanisms involved. We also found individual variations in the functional response for each of the experiments, with some consistency in the hierarchy regarding feeding efficiency. We did not find any differences between males and females. Our results provide an evaluation of individual variations in intake rate in interference-free conditions, which has rarely been done, and call for more controlled experiments to allow a finer understanding of the mechanisms of food acquisition in dabbling ducks.     

Effects of acute and chronic temperature changes on the functional responses of the dogfish <Emphasis Type="Italic">Scyliorhinus canicula</Emphasis> (Linnaeus, 1758) towards amphipod prey <Emphasis Type="Italic">Echinogammarus marinus</Emphasis> (Leach, 1815)

Predation is a strong driver of population dynamics and community structure and it is essential to reliably quantify and predict predation impacts on prey populations in a changing thermal landscape. Here, we used comparative functional response analyses to assess how predator-prey interactions between dogfish and invertebrate prey change under different warming scenarios. The Functional Response Type, attack rate, handling time and maximum feeding rate estimates were calculated for Scyliorhinus canicula preying upon Echinogammarus marinus under temperatures of 11.3 °C and 16.3 °C, which represent both the potential daily variation and predicted higher summer temperatures within Strangford Lough, N. Ireland. A two x two design of “Predator Acclimated”, “Prey Acclimated”, “Both Acclimated”, and “Both Unacclimated” was implemented to test functional responses to temperature rise. Attack rate was higher at 11.3 °C than at 16.3 °C, but handling time was lower and maximum feeding rates were higher at 16.3 °C. Non-acclimated predators had similar maximum feeding rate towards non-acclimated and acclimated prey, whereas acclimated predators had significantly higher maximum feeding rates towards acclimated prey as compared to non-acclimated prey. Results suggests that the predator attack rate is decreased by increasing temperature but when both predator and prey are acclimated the shorter handling times considerably increase predator impact. The functional response of the fish changed from Type II to Type III with an increase in temperature, except when only the prey were acclimated. This change from population destabilizing Type II to more stabilizing Type III could confer protection to prey at low densities but increase the maximum feeding rate by Scyliorhinus canicula in the future. However, predator movement between different thermal regimes may maintain a Type II response, albeit with a lower maximum feeding rate. This has implications for the way the increasing population Scyliorhinus canicula in the Irish Sea may exploit valuable fisheries stocks in the future.     

Noise-induced reduction in the attack rate of a planktivorous freshwater fish revealed by functional response analysis

1. Anthropogenic noise can affect animals physically, physiologically, and behaviourally. Although individual responses to noise are well documented, the consequences in terms of community structure, species coexistence, and ecosystem functioning remain fairly unknown.
2. The impact of noise on predation has received a growing interest and alterations in trophic links are observed when animals shift from foraging to stress-related behaviours, are distracted by noise, or because of acoustic masking. However, the experimental procedures classically used to quantify predation do not inform on the potential demographic impact on prey.
3. We derived the relationship between resource use and availability (the functional response) for European minnows (Phoxinus phoxinus) feeding on dipteran larvae (Chaoborus sp.) under two noise conditions: ambient noise and ambient noise supplemented with motorboat noise. The shape and magnitude of the functional response are powerful indicators of population outcomes and predator–prey dynamics. We also recorded fish behaviour to explore some proximate determinants of altered predation.
4. For both noise conditions, fish displayed a saturating (type II) functional response whose shape depends on two parameters: attack rate and handling time. Boat noise did not affect handling time but significantly reduced attack rate, resulting in a functional response curve of the same height but with a less steep initial slope. Fish exhibited a stress-related response to noise including increased swimming distance, more social interactions, and altered spatial distribution.
5. Our study shows the usefulness of the functional response approach to study the ecological impacts of noise and illustrates how the behavioural responses of predators to noise can modify the demographic pressure on prey. It also suggests that prey availability might mediate the negative effect of noise on predation. Community outcomes are expected if the reduced consumption of the main food sources goes with the overconsumption of alternative food sources, changing the distribution pattern of interaction strengths. Predation release could also trigger a trophic cascade, propagating the effect of noise to lower trophic levels.

     

Dynamics of handling time and functional response by larvae of Chrysomya albiceps (Dipt., Calliphoridae) on different prey species

Abstract:  One way to understand the behavioural patterns exhibited by a predator in response to prey density is to evaluate its functional response. Such evaluation yields information about basic mechanisms of prey–predator dynamics, and is an essential component of prey–predator models. In this paper we analysed experimentally the functional response and the handling time spent by Chrysomya albiceps on different prey species and larval instars of blowflies. The type II functional response was observed when second instar larvae of Chrysomya megacephala and Chrysomya macellaria were consumed. The handling time spent by the predator was significantly different between instars and species. The implications of the functional response and handling time for the interaction dynamics of Brazilian Chrysomyinae species are discussed.     

Depletion models can predict shorebird distribution at different spatial scales

Predicting the impact of habitat change on populations requires an understanding of the number of animals that a given area can support. Depletion models enable predictions of the numbers of individuals an area can support from prey density and predator searching efficiency and handling time. Depletion models have been successfully employed to predict patterns of abundance over small spatial scales, but most environmental change occurs over large spatial scales. We test the ability of depletion models to predict abundance at a range of scales with black-tailed godwits, Limosa limosa islandica. From the type II functional response of godwits to their prey, we calculated the handling time and searching efficiency associated with these prey. These were incorporated in a depletion model, together with the density of available prey determined from surveys, in order to predict godwit abundance. Tests of these predictions with Wetland Bird Survey data from the British Trust for Ornithology showed significant correlations between predicted and observed densities at three scales: within mudflats, within estuaries and between estuaries. Depletion models can thus be powerful tools for predicting the population size that can be supported on sites at a range of scales. This greatly enhances our confidence in predictions of the consequences of environmental change.     

Increased searching and handling effort in tall swards lead to a Type IV functional response in small grazing herbivores

Understanding the functional response of species is important in comprehending the species’ population dynamics and the functioning of multi-species assemblages. A Type II functional response, where instantaneous intake rate increases asymptotically with sward biomass, is thought to be common in grazers. However, at tall, dense swards, food intake might decline due to mechanical limitations or if animals selectively forage on the most nutritious parts of a sward, leading to a Type IV functional response, especially for smaller herbivores. We tested the predictions that bite mass, cropping time, swallowing time and searching time increase, and bite rate decreases with increasing grass biomass for different-sized Canada geese (Branta canadensis) foraging on grass swards. Bite mass indeed showed an increasing asymptotic relationship with grass biomass. At high biomass, difficulties in handling long leaves and in locating bites were responsible for increasing cropping, swallowing, and searching times. Constant bite mass and decreasing bite rate caused the intake rate to decrease at high sward biomass after reaching an optimum, leading to a Type IV functional response. Grazer body mass affected maximum bite mass and intake rate, but did not change the shape of the functional response. As grass nutrient contents are usually highest in short swards, this Type IV functional response in geese leads to an intake rate that is maximised in these swards. The lower grass biomass at which intake rate was maximised allows resource partitioning between different-sized grazers. We argue that this Type IV functional response is of more importance than previously thought.     

Predator functional response changed by induced defenses in prey

Functional responses play a central role in the nature and stability of predator-prey population dynamics. Here we investigate how induced defenses affect predator functional responses. In experimental communities, prey (Paramecium) expressed two previously undocumented inducible defenses--a speed reduction and a width increase--in response to nonlethal exposure to predatory Stenostomum. Nonlethal exposure also changed the shape of the predator's functional response from Type II to Type III, consistent with changes in the density dependence of attack rates. Handling times were also affected by prey defenses, increasing at least sixfold. These changes show that induced changes in prey have a real defensive function. At low prey densities, induction led to lower attack success; at high prey densities, attack rates were actually higher for induced prey. However, induction increased handling times sufficiently that consumption rates of defended prey were lower than those of undefended prey. Modification of attack rate and handling time has important potential consequences for population dynamics; Type III functional responses can increase the stability of population dynamics and persistence because predation on small populations is low, allowing a relict population to survive. Simulations of a predator-prey population dynamic model revealed the stabilizing potential of the Type III response.     

The role of sense organs in the feeding behaviour of Chinese perch

Experiments were conducted to identify the roles of the individual sense organs in the feeding behaviour of Chinese perch Siniperca chuatsi by determining the consumption of natural food after selective removal or blocking of eyes, lateral lines and olfactory organs, and also by observing the behavioural response to visual, mechanical and chemical stimulation by artificial prey. Chinese perch were able to feed properly on live prey fish when either eyes or lateral lines were intact or functional, but could scarcely feed without these two senses. Chinese perch recognized its prey by vision through the perception of motion and shape, and showed a greater dependence on vision in predation when both visual and mechanical cues were available. Chemical stimulation by natural food could not elicit any feeding response in Chinese perch, and gustation was only important to the fish for the last stage of food discrimination in the oropharyngeal cavity. The sensory basis of Chinese perch in feeding is well adapted to its nocturnal stalking hunting strategy, and also explains its peculiar food habit of accepting live prey fish only and refusing dead prey fish or artificial diets.     

Consumer-food systems: why type I functional responses are exclusive to filter feeders

The functional response of a consumer is the relationship between its consumption rate and the abundance of its food. A functional response is said to be of type I if consumption rate increases linearly with food abundance up to a threshold level at which it remains constant. According to conventional wisdom, such type I responses are more frequent among filter feeders than among other consumers. However, the validity of this claim has never been tested. We review 814 functional responses from 235 studies, thereby showing that type I responses are not only exceptionally frequent among filter feeders but that they have only been reported from these consumers. These findings can be understood by considering the conditions that a consumer must fulfil in order to show a type I response. First, the handling condition: the consumer must have a negligibly small handling time (i.e. the time needed for capturing and eating a food item), or it must be able to search for and to capture food while handling other food. Second, the satiation condition: unless its gut is completely filled and gut passage time is minimal, the consumer must search for food at a maximal rate with maximal effort. It thus has to spend much time on foraging (i.e. searching for food and handling it). Our functional response review suggests that only filter feeders sometimes meet both of these conditions. This suggestion is reasonable because filter feeders typically fulfil the handling condition and can meet the satiation condition without losing time, for they are, by contrast to non-filter feeders, able simultaneously to perform foraging and non-foraging activities, such as migration or reproduction.     

外来鱼类雀鳝与本地斑鳢功能反应的比较

为评估外来鱼类雀鳝对本地鱼类资源的捕食效率及与本地肉食鱼类摄食的差异, 研究对眼斑雀鳝(Lepisosteus oculatus)捕食鲮(Cirrhinus molitorella)、广东鲂(Megalobrama terminalis)和草鱼(Ctenopharyngodon idellus)的功能反应进行了实验研究, 并与本地生态位相似肉食性鱼类斑鳢(Channa maculata)的功能反应进行了比较。研究分析了雀鳝和斑鳢的功能反应类型, 建立了功能反应曲线, 估计了资源消耗率相关的参数, 并结合功能反应模型和食物转化效率模拟了2种鱼类的种群增长趋势。结果表明: 雀鳝与斑鳢的功能反应类型均为Type-Ⅱ型; 2种捕食者的袭击率(P=0.383)及食物处理时间(P=0.663)均无显著性差异, 表明2种捕食者在单位资源消耗率上并无明显差别; 雀鳝的相对生长效率显著高于斑鳢(P<0.05), 而食物转化效率则无显著差异(P=0.132); 雀鳝的种群增长快于斑鳢且具有更高的稳定种群密度。研究为系统评估外来鱼类雀鳝的生态影响提供了基础, 采用的量化功能反应的方法可为其他外来鱼类生态影响的评估提供借鉴和参考。     

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.     

Feeding by marine fish larvae: developmental and functional responses

Synopsis The relationship between prey consumption rate and prey concentration (functional response), and its change with growth (developmental response) were examined in the laboratory for three species of marine fish larvae: bay anchovy Anchoa mitchilli (Engraulidae), sea bream Archosargus rhomboidalis (Sparidae) and lined sole Achirus lineatus (Soleidae). The major objective was to determine relative predatory abilities of the larvae by fitting feeding rate data to developmental and functional response models. Feeding success, prey capture success, attack rates, handling times and search rates were estimated. Prey consumption rates and attack rates of bay anchovy usually were highest, but at the lowest prey level (50 per liter) first-feeding sea bream larvae had the highest consumption rate. Sea bream could consume prey at near-maximum rates at prey levels lower than those required by the other species. As larvae grew, time searching per attack decreased rapidly for all species, especially at low prey levels. Handling time also decreased, but most rapidly for bay anchovy. Search rates were highest for bay anchovy and lowest for lined sole. Bay anchovy had the best apparent predation ability, but when previous results on larval growth rates, survival rates and growth efficiencies were considered, sea bream larvae were the most efficient predators and the least likely of the three species to be limited by low prey levels.     

Flexible components of functional responses

1. The functional response of predators describes the rate at which a predator consumes prey and is an important determinant of community dynamics. Despite the importance, most empirical studies have considered a limited number of models of functional response. In addition, the models often make strong assumptions about the pattern of predation processes, even though functional responses can potentially exhibit a wide variety of patterns. 2. In addition to the limited model consideration, model selections of functional response models cannot tease apart the components of predation (i.e. capture rate and handling time) when flexible traits are considered because it is always possible that many different combinations of the capture rate and handling time can lead to the same predation rate. 3. This study directly examined the capture rate and handling time of functional response in a mite community. To avoid the model selection problem, the searching and handling behaviour data were collected. The model selection was applied directly to these two components of predation data. Commonly used functional response models and models that allow for more flexible patterns were compared. 4. The results indicated that assumptions of the commonly used models were not supported by the data, and a flexible model was selected as the best model. These results suggest the need to consider a wider variety of predation patterns when characterizing a functional response. Without making a strong assumption (e.g. static handling time), model selections on functional response models cannot be used to make reliable inferences on the predation mechanisms.     

Feeding behaviour determining differential capture success of evasive prey in underyearling European perch (<Emphasis Type="Italic">Perca fluviatilis</Emphasis> L.) and roach (<Emphasis Type="Italic">Rutilus rutilus</Emphasis> (L.))

The effect of feeding behaviour on the prey capture efficiency of young-of-the-year European perch and roach was investigated in laboratory experiments using planktonic crustaceans possessing different escape abilities—Daphnia sp. and Cyclops sp. Two sets of experiments were performed. In the first set, the feeding efficiency and behaviour of 270 fish individuals were determined by stomach content analyses and video record evaluations. In the second set of experiments, analysis of attack-effort, which was evaluated as attack-distance and repeated strikes, was undertaken. Except for situations in which Daphnia was offered at high densities, the feeding efficiency of perch was significantly higher compared to roach in all other combinations of prey types and densities. Roach consumed significantly less prey compared to perch when feeding exclusively on the evasive Cyclops and when it was offered in a 1:1 ratio mixture with Daphnia. The mean swimming speed was similar in both fish species, but behavioural differences were evident during prey search and capture. Perch swam through the aquaria in short and fast movements that were interrupted by many stops. Roach exhibited rather continuous swimming that was punctuated by slowdowns instead of stops. The perch attacks were very intensive and repeated strikes occurred, particularly when feeding on evasive Cyclops. On the other hand, roach revealed strong schooling behaviour restricting the fish during inspection of the experimental aquaria. The distinct differences in feeding efficiency between perch and roach were demonstrated to be closely related to differences in their feeding behaviour. Discontinuous searching for prey, vigorous attacks, occurrence of repeated strikes and the absence of schooling increased perch prey capture efficiency, particularly when foraging on evasive copepods.     

Mechanisms of drift-feeding behavior in juvenile Chinook salmon and the role of inedible debris in a clear-water Alaskan stream

Drift-feeding fish are challenged to discriminate between prey and similar-sized particles of debris, which are ubiquitous even in clear-water streams. Spending time and energy pursuing debris mistaken as prey could affect fish growth and the fitness potential of different foraging strategies. Our goal was to determine the extent to which debris influences drift-feeding fish in clear water under low-flow conditions when the distracting effect of debris should be at a minimum. We used high-definition video to measure the reactions of drift-feeding juvenile Chinook salmon (Oncorhynchus tshawytscha) to natural debris and prey in situ in the Chena River, Alaska. Among all potential food items fish pursued, 52 % were captured and quickly expelled from the mouth, 39 % were visually inspected but not captured, and only 9 % were ingested. Foraging attempt rate was only moderately correlated with ingestion rate (Kendall’s τ?=?0.55), raising concerns about the common use of foraging attempts as a presumed index of foraging success. The total time fish spent handling debris increased linearly with foraging attempt rate and ranged between 4 and 25 % of total foraging time among observed groups. Our results help motivate a revised theoretical view of drift feeding that emphasizes prey detection and discrimination, incorporating ideas from signal detection theory and the study of visual attention in cognitive ecology. We discuss how these ideas could lead to better explanations and predictions of the spatial behavior, prey selection, and energy intake of drift-feeding fish.     

How predation by Podisus nigrispinus is influenced by developmental stage and density of its prey Alabama argillacea

The functional response of a predator to the density of its prey is affected by several factors, including the prey's developmental stage. This study evaluated the functional response of Podisus nigrispinus (Dallas) (Hemiptera: Heteroptera: Pentatomidae) females to fourth instars and pupae of Alabama argillacea (Hübner) (Lepidoptera: Noctuidae), an important pest of cotton (Gossypium hirsutum L., Malvaceae) in Brazil. The prey were exposed to the predator for 12 and 24 h, and in densities of 1, 6, 12, 18, 24, and 30 items per predator female. The predation data were subjected to polynomial regression logistic analysis to determine the type of functional response. Holling and Rogers' equations were used to estimate parameters such as attack rate and handling time. Podisus nigrispinus females showed functional response types II and III by preying on larvae and pupae, respectively. The attack rate and handling time did not differ between the 12 and 24 h exposure times. Predation rate was higher at higher larval and pupal densities; predation was highest at a density of 30 prey items per female, and it was similar at 18 and 24 prey per predator. Understanding the interaction of predators and their food resources helps to optimize biological control strategies. It also helps the decision‐making and the improvement of release techniques of P. nigrispinus in the field.     

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.     

Are digestive characteristics important contributors to the profitability of prey?

Summary In the field, Spinachia fed on four types of prey; copepods, isopods, mysids and amphipods. As fish size increased, mysids gradually succeeded amphipods as the most important food type in the diet. Prey dimensions and morphometry of the fish's mouth most accurately predicted capture efficiency for amphipods, whereas for mysids capture efficiency was determined by the prey's escape response and the fish's fast-start capability. Responses to model prey revealed the ability of fish to differentiate among contrasting prey characteristics, resulting in the adoption of appropriate predatory tactics. Amphipods were associated with a shorter gut evacuation time than mysids, although approximately equal proportions of energy were absorbed from each. Similar rations of mysids and amphipods were required to satiate fish. The lower energy content per unit dry mass of amphipods was off set by their lower water content. From pre-digestive behaviour, we predicted that mysids were more profitable than amphipods, and this was reflected in the fish's choice. Conversely, incorporating the net rate of energy uptake by the gut led us to predict that amphipods were more profitable. Although physiological constraints clearly influence the net rate of energy uptake, it appears that dietary preferences are based on pre-digestive predatory behaviour and hence on time minimisation.