Differential digestion and evacuation rates of prey in a warm-temperate grouper, gag Mycteroperca microlepis (Goode & Bean)

Prey-specific gastric evacuation rates and digestion state indices were modelled for gag Mycteroperca microlepis , a large warm-temperate grouper, consuming meals of either baitfish (scaled sardine Harengula jaguana ) or crab (purple swimmer crab Portunus gibbesii ). Power exponential models best fit the wet and dry mass gastric evacuation data and the average digestion indices over post-prandial time (PPT), regardless of prey type or gag size (Adjusted R² ≥ 0·79). Gag mass ( M ) or total length ( L _T) incorporated into an expanded power exponential model, along with exponential scalars, resulted in highly predictive ( R² ≥ 0·87) gastric evacuation and average digestion state models. The expanded power exponential models fit to the baitfish and crab wet mass gastric evacuation data differed significantly (Kimura's likelihood ratio test (LRT), both P < 0·001). Gag consuming crab showed a digestive lag period of at least 4 h (wet mass) and took a longer time to complete digestion relative to gag consuming baitfish. Gag, as well as many other warm-temperate and tropical groupers, consume a mixture of fish and crab prey and they will therefore require the development of a consumption model that incorporates mixed-prey gastric evacuation models.     

A surface-dependent gastric evacuation model for fish

A gastric evacuation curve expresses how fast prey disappear from the stomach, and empirical models are used generally for the relationship between weight of prey remaining ( W_t ) and time ( t ) after a meal. Unfortunately, empirical models are likely to have restricted applicability because their parameters often represent limited biological mechanisms. This paper develops a simple digestion model. The simplest form of the model has four parameters; the digestion velocity (expressing enzymatic breakdown of prey), prey length, initial prey radius and the density of the prey. Two more parameters are included in an extended version; a time-delay before digestion starts and environmental temperature. The approach is based on the assumption that prey digestion is a surface process in that digestive enzymes attack progressively deeper into a prey of known size and shape so that the average digestion rate is proportional to the prey radius r (m). This process is characterized by the digestion velocity d_s (m s ^–1). Unknown parameters are estimated with uncertainty using the maximum likelihood technique. Model evaluation using published data sets demonstrated that the new model is flexible. Prey geometry is incorporated into the model and temperature effects upon gastric evacuation are linked directly to the digestion velocity.     

The effects of temperature and meal size on the rate of gastric evacuation in perch (Perca fluviatilis) fed on fish larvae

SUMMARY. The rate of gastric evacuation in perch (Perca fluviatilis) (89–170 mm length) fed on fish larvae was studied at temperatures between 12.0 and 21.7°C. Gastric evacuation rates were usually described by an exponential function. The instantaneous rate of gastric evacuation ( R ) was constant for a large number of different meal sizes. At higher food rations, a lag phase in digestion was found during the first part of digestion, and this ration size was smaller for smaller perch (89–110 mm) than for bigger perch (120–170 mm). Below these larger meal sizes, gastric evacuation was similar for the different size classes studied. The relation between R and temperature was described by an exponential function. The effects of meal size, number of food items, fish size and temperature on the rate of gastric evacuation are discussed.     

Gastric evacuation of horse mackerel. II. The effects of different prey types on the evacuation model

Gastric evacuation experiments were performed on horse mackerel Trachurus trachurus. Discrete meals of different sizes of krill Meganyctiphanes norvegicus, mysids Neomysis integer , brown shrimp Crangon crangon , and herring Clupea harengus were fed to test the effect of prey type on the gastric evacuation. A general evacuation model without meal size as a variable was fitted to the data on dry weights by means of non-linear regression. With the exception of mysid evacuation the estimated parameters for curvature, predator weight effect, and temperature effect of the general evacuation model were very similar for all other food types and also confirmed the results obtained with smelt Osmerus eperlanus. In contrast the evacuation of mysids showed a strong exponential curvature, rather the effect of prey size than of prey type. The prey specific evacuation constants of the model revealed a linear relationship to the energy content. A general consumption model for field estimates is presented that can be used for prey types that have not yet been tested in the experiments, provided the energy content is known.     

Gastric evacuation in little skate

The effects of prey type and prey preparation on the mathematical forms and rates describing gastric evacuation in little skate Raja erinacea were examined. Linear and square–root models best described the gastric evacuation of whole, thin–shelled krill Meganyctiphanes norvegica , clam feet/muscle Spisula solidissima/Placopecten magellanicus , polychaetes Glycera spp. and sand lance Ammodytes dubius . Evacuation of krill and clams was faster than polychaetes and sand lance. A logistic model best described the evacuation data of thick–shelled benthic shrimp Crangon septemspinosa/Palaemonetes spp. Cut polychaetes Nereis spp. were digested at an exponential rate and were evacuated faster than would be predicted based upon comparison with live polychaetes and previously published evacuation–temperature relationships. The results of this study suggest that a single equation does not describe the evacuation process for all prey, and that whole prey should be used if laboratory–derived rates of gastric evacuation are to reflect what might occur in wild fishes.     

Offshore insular variation in the diet of the Taiwanese bamboo viper Trimeresurus stejnegeri (Schmidt)

Dietary data were ascertained for 229 T. stejnegeri (snout vent length >300mm) from 36 localities throughout the main island of Taiwan and the outlying Orchid (Lanyu) and Green (Ludau) Islands. Twenty nine percent of the snakes were devoid of any prey, and of the snakes containing prey, 43% of the cases were unidentifiable. This relatively large proportion of unidentifiable prey items (observed in the hindgut) may reflect either rapid digestion of amphibian prey and/or rapid venting of feces as an evolutionary adaptation to arboreal life. Trimeresurus stejnegeri appears euryphagous, taking primarily amphibians, but additionally reptilian, mammalian and insect prey. There was no discrepancy in prey composition based on comparisons of where the prey item was recorded in the digestive tract. No sexual variation in diet composition was evident, although males were more likely to contain prey than females, indicating the utilisation of different foraging strategies on similar sympatric prey items. Variation in diet composition was observed between mainland Taiwan and offshore islands, which is most likely the result of differences in prey availability.     

Gastric evacuation in mackerel: the effects of meal size, prey type and temperature

Gastric evacuation experiments, performed with groups of Atlantic mackerel Scomber scombrus and three different prey species: sprat Sprattus sprattus , sandeel Ammodytes marinus and krill Meganectyphanes norwegicus , revealed a strong temperature effect on gastric evacuation ( Q ₁₀=4.l) and a curvilinear evacuation with a shape intermediate between the square root and the exponential model. From the experiments with krill, where no internal tagging was possible, the medians and upper quartiles of the stomach content distributions were analysed instead of individual data. From these experiments the evacuation rate of krill was 60% higher than that of sandeel. An additional simulation study confirmed that the use of medians results in more accurate estimates of the evacuation rates when compared to the arithmetic means.     

Pacific walrus (Odobenus rosmarus divergens):Differential prey digestion and diet

Stomach content data from 798 Pacific walruses ( Odobenus rosmarus divergens ) collected during 1952–1991 were analyzed using a method that evaluates the stage of digestion of prey remains. Non-molluscan prey taxa were not well represented in previous interpretations of walrus diet due to digestion biases. Stomach contents least affected by digestion (fresh stomachs) contained more prey taxa than stomachs of an unknown or more digested state. Bivalves, gastropods, and polychaete worms were the most frequent prey items in both the Bering and Chukchi seas, although bivalves occurred more frequently in stomachs from the Bering Sea and gastropods occurred more frequently in stomachs from the Chukchi Sea. Male and female walruses consumed essentially the same prey when in the same location. Using only fresh stomachs collected between 1975 and 1985, there was no significant difference between the proportion that contained mostly bivalves and the proportion that contained non-bivalve prey items. Earlier interpretations of a change in walrus diet in this period compared to the prior two decades may have been due to digestion as well as sampling biases. Current climatic changes may affect walrus's access to diverse, productive shallow water feeding areas.     

Gastric evacuation rates and daily ration of piscivorous coho salmon, Oncorhynchus kisutch Walbaum

Effects of temperature and meal size on gastric evacuation rates of juvenile coho salmon, Oncorhynchus kisutch , consuming sockeye salmon, O. nerka , fry were examined and used in the estimation of daily meal, daily ration and number of fry consumed by coho in Chignik Lake, Alaska. Evacuation of fry by coho was best described by a negative exponential model (average R² = 0.93). A square root model also provided a good fit (average R² = 0·93), but the y-intercepts deviated more from the expected value than did the y-intercepts of the exponential model. The effect of temperature ( T , 5–13° C) and meal size (MS, 0·166–0·367 g) on the exponential evacuation rate (r_e, h^-1) could be described as
In the lake, coho fed continuously during the 24-h period in early June 1986 and 1987. Estimates of daily meal and ration of coho calculated by the Eggers method and the geometric mean of prey weight ranged from 0·224 to 0·435 g (2.1–4.4% body wt) depending on location and year. The Elliott & Persson method provided similar estimates of food consumption, whereas estimates based on the Pennington method and square root evacuation of prey differed from the exponential models. Sockeye fry represented 93% of the total prey weight. The average number of sockeye fry consumed per coho per 24 h, based on the arithmetic mean of prey weight, was 3·0–3·9 fry.     

Gastric evacuation of mixed stomach contents in predatory gadoids: an expanded application of the square root model to estimate food rations

A mechanistic gastric evacuation model of mixed stomach contents was established. Its simple generic nature should prove useful for studying predator‐prey interactions in fish communities. Only prey lengths were required as additional input. Surface dependent digestive processes were considered to act on equal fractions of individual prey‐cylinder surfaces. Primary and interactive effects of size, energy density and resistance to digestion of individual prey in a stomach were described by the model. Model predictions of results from experiments on gastric evacuation of meals composed of different prey types demonstrated the capability of this model, as opposed to previously applied model principles, to predict evacuation of mixed stomach contents involving the three above‐mentioned prey characteristics. The study furthermore illustrated that estimates of food ration might be severely biased by using improperly formulated effects of prey characters on gastric evacuation, and demonstrated that the new model holds the potential to predict food ration and diet composition for wild populations of predatory fishes.     

Evolutionarily stable kleptoparasitism: consequences of different prey types

We present two elaborations of the model of Broom and Ruxtonthat found evolutionarily stable kleptoparasitic strategiesfor foragers. These elaborations relax the assumption that thedistribution of times required to handle discovered food itemsis exponential. These changes increase the complexity of themodel but represent a significant improvement in biologicalrealism. In one elaboration, handling takes a fixed interval,t_h, at the end of which the whole value of the food item isobtained. We liken this to peeling then consuming a small orange.The other elaboration also assumes that handling takes a fixedinterval, t_h, but this time the reward from the food item isextracted continuously throughout the handling period. We likenthis to eating an apple. Both models predict that increasingfood density, the ease with which food items can be discovered,or the length of aggressive contests all act to make kleptoparasitismless common. The difference between the evolutionarily stablestrategy solutions of the apple and orange models provides aclear prediction of our theory. When prey items require handlingbefore yielding a lump sum at the end, then kleptoparasiticattacks will be focused on prey items near the end of theirhandling period. However, if prey items yield reward continuouslyduring handling, then attacks should be biased toward newlydiscovered food items. Another key difference between the modelpredictions is that kleptoparasitism increases with foragerdensity in the apple model, but decreases in the orange model.     

Unexpected effects of prey dimensions and morphologies on the size selective feeding by two bacterivorous flagellates (Ochromonas sp. and Spumella sp.)

Current models on protistan size-selective feeding assume that contact probability is the factor that largely explains observed food preferences. Contact probability is generally expected to be positively correlated with prey size and therefore to explain observed food selection for larger prey items. We critically tested these basic assumptions on size-selective feeding using the interception-feeding chrysomonad nanoflagellates Ochromonas sp. and Spumella sp. Mechanisms of differential feeding were studied during distinct stages of the selection process (i.e. contact probability, capture efficiency, ingestion efficiency, and differential digestion) by means of high-resolution video microscopy. Food selection was investigated using a mixture of microspheres ranging from 0.3-2.2 microm in diam., as well as a mixed bacterial community. In contrast to current model assumptions, the contact probability was highest for microspheres of intermediate size (0.9-1.2 microm), but was not generally positively correlated with prey size over the whole prey size range. Capture and ingestion also proved to be involved in size selection: these patterns were also independent of the food concentration (p = 0.968 for Ochromonas, p = 0.971 for Spumella). Even though the capture rate was significantly higher for attached flagellates than for swimming flagellates (p < 0.001), size selectivity was not affected (p > 0.05). Our results indicate that: (i) size selection is not actively regulated by these flagellates, but is a passive process; (ii) contact probability is not generally positively correlated with prey size, but shows a maximum for intermediate-sized prey in the prey size spectrum of 0.3-2.2 microm; and (iii) selection steps other than contact probability are crucial for size selection and should be integrated in models on size selection.     

Predator size and prey size–gut capacity ratios determine kill frequency and carcass production in terrestrial carnivorous mammals

Carnivore kill frequency is a fundamental part of predator–prey interactions, which are important shapers of ecosystems. Current field kill frequency data are rare and existing models are insufficiently adapted to carnivore functional groups. We developed a kill frequency model accounting for carnivore mass, prey mass, pack size, partial consumption of prey and carnivore gut capacity. Two main carnivore functional groups, small prey‐feeders versus large prey‐feeders, were established based on the relationship between stomach capacity (C) and pack corrected prey mass (iMprey). Although the majority of small prey‐feeders is below, and of large prey‐feeders above a body mass of 10–20 kg, both occur across the whole body size spectrum, indicating that the dichotomy is rather linked to body size‐related ecology than physiology. The model predicts a negative relationship between predator size and kill frequency for large prey‐feeders. However, for small prey‐feeders, this negative relationship was absent. When comparing carnivore prey requirements to estimated stomach capacity, small carnivores may have to eat to their full capacity repeatedly per day, requiring fast digestion and gut clearance. Large carnivores do not necessarily have to eat to full gastric capacity per day, or do not need to eat every day, which in turn reduces kill frequencies or drives other ecological processes such as scavenging, kleptoparasitism, and partial carcass consumption. Where ecological conditions allow, large prey‐feeding appears attractive for carnivores, which can thus reduce activities related to hunting. This is particularly so for large carnivores, who can achieve distinct reductions in hunting activity due to their relatively large gut capacity.     

The response of gastric pH and motility to fasting and feeding in free swimming blacktip reef sharks, Carcharhinus melanopterus

In many fish and reptiles, gastric digestion is responsible for the complete breakdown of prey items into semi-liquid chyme. The responses of the stomach to feeding and to periods of fasting are, however, unknown for many lower vertebrates. We inserted data loggers into the stomachs of free-swimming captive adult blacktip reef sharks (Carcharhinus melanopterus) to quantify gastric pH, motility and temperature during fasting and following ingestion of food. Gastric acid secretion was continuous, even during long periods of fasting, with a mean pH of 1.66 ± 0.40 (± 1 SD) when the stomach was empty. Stomach contractions were greater following meals of mackerel than for those of squid. Gastric motility following feeding on mackerel, was positively influenced by ambient temperature, and followed a quadratic relationship with meal size, with maximum motility occurring after meals of 0.8-1.0% body weight. Diel changes in gastric motility were apparent, and were most likely caused by diel changes in ambient temperature. Gastric digestion in blacktip reef sharks is affected by both biotic and abiotic variables. We hypothesize that behavioral strategies adopted by sharks in the field may be an attempt to optimize digestion by selecting for appropriate environmental conditions.     

Influences of food particle size and dietary energy content on patterns of gastric evacuation in fish: test of a physiological model of gastric emptying

The evacuation of food from the stomach is probably not a continuous smooth process but may occur in a pulse-like (step-wise) fashion. It is hypothesised that the pattern of emptying is influenced by feedback signals from receptors located in the upper intestinal tract and by factors affecting the rate of physical/chemical breakdown of the food particles. Mathematical functions used for describing evacuation patterns imply that emptying is continuous and they are, therefore, at best, only approximations. It was predicted that different mathematical expressions would give best fit to empirical data from gastric evacuation studies dependent upon the experimental conditions employed. An analysis of published data supported the predictions and revealed that an exponential function best described the evacuation of small, easily digested prey items, but a linear expression gave the best fit to data of the emptying of large food items. It is suggested that differences in surface-to-volume ratios between large and small food particles and the friabilities of different food types are important in determining the pattern of emptying. Whilst dietary energy content also appears to be an important factor in the governing of gastric evacuation, it appears that emptying pattern in fish species is rather less sensitive to changes in dietary composition than is gastric emptying in mammals.     

Diel feeding habits of juveniles of Mullus surmuletus (Linneo 1758) in the lagoon of the Stagnone di Marsala (Western Sicily, Italy)

Diet composition, feeding rhythm, gastric evacuation rate and daily ration were investigated in juvenile Mullus surmuletus (Linneo 1758). Fish were collected in the lagoon of the Stagnone di Marsala in western Sicily, in July 1995, during a 24 h sampling period. Copepoda, Polychaeta, Amphipoda and Tanaidacea were shown to be the most frequent prey items. The feeding index values showed two different daily feeding times. A unimodal trend in the daily rhythm of food consumption was derived, with a peak in feeding between 1200 and 2000 h. Gastric evacuation in juvenile M. surmuletus is best described by an exponential model, with a gastric evacuation rate R  = 0.66 g h⁻¹ ( r = 0.88) ( T  = 24.45 ± 0.64°C). The amount of food consumed daily, calculated according to the Elliott and Persson model, was 0.079 g, equal to about 8% of the average dry weight of the fish.     

Influence of predation risk on diet selection: a simple example in the grey squirrel

Current models of the optimal diet are special cases of a more general (and complex) model which incorporates the effects of predation risk on diet selection; this follows from an assumption implicit in current models that all prey items are eaten where they are encountered. Relaxing this assumption so that a forager might carry a prey item to protective cover for consumption leads to the conclusion that the value of a prey item is a function of its distance to cover as well as its energy content and handling time. Such considerations can significantly alter the outcome of diet selection relative to that expected from simple diet theory. We found that grey squirrels (Sciurus carolinensis) may reject more energetically profitable, but small food items in favour of locating larger, less energetically profitable items that can be carried to protective cover for consumption without greatly sacrificing foraging efficiency. The squirrel's tendency to reject a more profitable item is a function of its distance from cover and the size of the less profitable items. Such behaviour is inconsistent with predictions of current diet models, but is consistent with our qualitative predictions based on a previously determined predation-risk-foraging-efficiency trade-off in the grey squirrel.     

Optimal time to emerge from refuge

Factors affecting emergence by prey that enter refuges when approached by predators have been studied intensively, but only two theoretical models predict how long prey should remain in a refuge before emerging. We argue that prey can make better decisions than allowed by one model; the other model describes cases in which predators wait for prey to emerge. We present optimality models that permit prey to select a time to emerge that maximizes fitness. When in a refuge, a prey cannot obtain benefits outside; emerging too soon can be catastrophic, but delaying emergence entails loss of fitness. If predators resume foraging quickly rather than engaging in strategic waiting games, current theory suggests that prey emerge when the costs of remaining in a refuge and of emerging are equal. However, prey often can do better by emerging at the time maximizing fitness rather than when benefits equal costs (i.e. when prey break even). Optimal emergence time depends on initial fitness, benefits lost by remaining in refuge, and the decay rate of predation risk. Benefits lost if a prey is killed are modelled separately from benefits that contribute to lifetime fitness, even if the prey is killed (individual reproduction, altruism). Fitness of prey emerging at the optimal emergence time may be greater than, equal to or less than initial fitness. Break-even and optimality models base predictions on the opposing effects of risk and loss of benefits. Thus, many empirically verified predictions are identical at the ordinal level although differing quantitatively. Optimality models provide novel testable predictions for the effects of initial fitness, benefits, and, for ectotherms, the rate of cooling in refuge. They predict earlier emergence for equal retainable benefits than for those lost upon death.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 375–382.