A gastric evacuation model for three predatory gadoids and implications of using pooled field data of stomach contents to estimate food rations

The general applicability of the square root model to describe gastric evacuation in predatory gadoids independently of meal size was verified through experiments on whiting Merlangius merlangus , saithe Pollachius virens and cod Gadus morhua fed different fish and crustacean prey. Gastric evacuation rate was related by a negative power function to energy density of fish prey over an extended range from 3·4 to 11·5 kJ g^-1 of the effector variable. Gastric evacuation of crustacean prey seemed to depend on the characteristics of their exoskeleton. Gastric evacuation of mixed meals composed of fish prey with different energy densities could be described as a function of the overall energy density. The evacuation rate of each of the prey could be described directly by their share of the stomach content. A full gastric evacuation model including predator size, temperature and prey energy density was established for whiting, saithe and cod. It was demonstrated that estimates of food rations might be severely biased by use of mean values for prey composition and total mass of stomach contents.     

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

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

How feeding performance and energy intake change with a small increase in the body size of the three-spined stickleback

Changes in the foraging behaviour due to variation in the body size of the three-spined stickleback Gasterosteus aculeatus were investigated. All sizes of fish had a high probability of attacking prey whenever encountered. The probability of eating the prey increased with the size of the fish, as the larger fish had larger jaws and a greater stomach capacity. Therefore, as fish increased in size there was an increase in the probability of successful prey capture. The level of satiation did not have an effect on the prey handling time, which is contrary to other studies and is probably a result of the large prey sizes. The physical size of the prey meant that the handling times were long regardless of the motivational level of the fish. The larger fish took in more energy and at a faster rate, although the time to reach satiation was similar for all fish sizes. The advantage that large fish appear to have in successfully gaining large prey is negated by their greater metabolic requirement. The changes in feeding performance induced by small increases in body size could have important consequences for intraspecific competition, habitat Use and risk of predation.     

DIET OF WEDDELL SEALS LEPTONYCHOTES WEDDELLII AT VESTKAPP, EASTERN WEDDELL SEA (ANTARCTICA), IN RELATION TO LOCAL FOOD SUPPLY

Thirteen Weddell seals ( Leptonychotes weddellii ) were collected at Vestkapp, eastern Weddell Sea coast, in austral spring 1986. All stomachs contained partially digested food. The mean wet weight of stomach contents was 7.5 kg, 3.3% of the. mean body weight of the collected seals. Twelve fish species and three cephalopod species were identified from 372 left otoliths and 25 lower beaks, representing 58.4% of 679 total prey items obtained. Composition by number of total prey was: Chionodraco myersi (15.8%), Trematomus eulepidotus (10.0%), Pagetopsis maculatus (9.7%), Racovitzia glacialis (9.6%) and Cryodraco antarcticus (4.1%). Otoliths of the seven other fish species and beaks of the three cephalopod species together represented 9.1% of total prey numbers. The pooled wet weights calculated from 13 prey species (regressions for two octopod species were not available) amounted to 43.5 kg food mass and represented 44.7% of the combined food mass in all stomachs. Composition by mass was: C. myersi (44.5%), T. eulepidotus (19.8%), squid Psychroteuthis glacialis (8.5%), P. maculates (7.9%), C. antarcticus (7.1%) and R. glacialis (6.2%). The remaining 7 fish species together represented 5.8% by mass. Temporal variation in food availability was apparent. Midwater fish Pleuragramma antarcticum was the staple food of Weddell seals from the same area during the 1985 summer, whereas it was absent in the samples taken in spring 1986. Estimates of fish biomass from net hauls demonstrate a highly variable availability of pelagic food resources for top predators in the Vestkapp area.     

Effects of temperature, prey type and prey size on gastric evacuation in small cod and whiting

Gastric lavage was used to investigate the effects of temperature, prey type and prey size on gastric evacuation in small cod Gadus morhua and whiting Merlangius merlangus . The fish were fed to satiation and subsequently the stomach contents were sampled to determine the rate at which food was evacuated. Satiation meal size was positively related to temperature and differed between prey types. The gastric evacuation rate (GER) also tended to increase with temperature and varied with prey type. GER at temperatures of 11.3–12.7° C averaged 1.5–1.7 times higher than at 6–9.5° C. There was no significant difference (P>0.05) in the evacuation of lugworm Arenicola marina , sandeel Ammodytes spp., and herring Clupea harengus , but the GER of brown shrimp Crangon vulgaris was much slower (P<0.05). No significant changes in GER were observed when fish were fed on three different size groups of either herring or brown shrimp. In whiting, there was no significant difference in the GER of individual herring or brown shrimp when they were fed as single species meals or incorporated in meals containing a mixture of prey species.     

Rates of gastric evacuation in piscivorous brown trout, Salmo trutta

SUMMARY. 1. The dry weight of food remaining in the stomachs of piscivorous trout decreased exponentially with time. Gastric evacuation rates increased exponentially with increasing temperature but were unaffected by predator size, meal size or type of fish prey.
2. Mathematical models were developed to estimate both the rate and time for the gastric evacuation of different meal sizes (expressed as dry weight), and were applicable to piscivorous trout of different sizes (length range 10–32 cm) feeding on trout fry or sticklebacks at different temperatures (range 5–18°C).
3. The wet weight of food in the stomachs also decreased exponentially with time, but evacuation rates both increased with temperature and decreased with increasing meal size; the latter relationship occurred because relative rates of water loss from a meal also decreased with increasing meal size. Use of wet or dry weights can therefore lead to different conclusions about the effect of meal size on evacuation rates.
4. When piscivorous trout were fed three consecutive meals of varying size, the models predicted the total dry weight of food left in the stomach, but not the weight remaining for each individual meal. Interactions between meals led to an increase in evacuation rates for meals consumed early in the series and a decrease in evacuation rates for later meals.
5. Evacuation rates for piscivorous trout were compared with those for trout feeding on invertebrates in an earlier study, and were close to those for caddis larvae as prey, higher than those for mealworms and lower than those for a variety of invertebrate prey. Although a great deal is now known about the daily food intake and growth rates of trout feeding on invertebrates, there is little comparable information for piscivorous trout.     

Some observations of the effects of body weight, temperature, meal size and quality on gastric emptying time in the turbot, Scophthalmus maximus (L.) using radiography

The gastric emptying time (G.E.T.) in turbot was investigated using X-radiography and was found to decrease with temperature. Small fish processed a given ration, expressed as percent body weight, faster than large fish (G.E.T. was found to be proportional to (fish weight)^0.364). Large meals in a given fish were processed at a faster rate than small meals. Gastric emptying rate (G.E.R.) was found to be proportional to (meal size g)^0.613 at 8° C and (meal size g)^0.788 at 19° C. These exponents are in agreement with a recently proposed model relating G.E.T. and G.E.R. to meal size (Fänge & Grove, 1978). Large fish emptied a meal of given absolute size from the stomach at a faster rate (g h⁻¹) than small fish. Experimental meals diluted with kaolin were evacuated in significantly less time than a control diet, suggesting that turbot may adjust feeding rates when food quality varies.     

Constraints on prey size selection by the three-spined stickleback: energy requirements and the capacity and fullness of the gut

An experiment was designed to study how gut fullness and encounter with 5-mm Asellus aquaticus influenced acceptance or rejection of less profitable 8-mm Asellus . 45-mm sticklebacks were found to always accept 5-mm prey whereas 8-mm prey were accepted with an initial probability of about 0.9. This probability decreased as the gut filled. Fish of differing sizes and sex had similar daily energy intakes per unit body size, however the acceptance of 8-mm prey was related to fish size. Whenever a fish orientated to a prey it was followed by pursuit and manipulation independently of prey size. The decision to accept or reject prey occurred after one manipulation, a criterion that was more variable for the larger prey. For one feeding session per day the total energy intake was almost constant despite the changing combination of prey sizes eaten. The fish ate prey with long handling times if the energetic contents of the stomach had not reached 450 J. Calculations were made of how many of each millimetre prey size group would satisfy the 450 J demand and how long the estimated number would take to handle. This showed that the best option is to consume 5-mm prey if given the choice.     

Gutsy genetics: identification of digested piscine prey items in the stomach contents of sympatric native and introduced warmwater catfishes via DNA barcoding

A major focus of ecology is understanding trophic relationships and energy flows in natural systems, associated food web dynamics and changes in food webs due to introduced species. Predator-prey interactions are often assessed by examining stomach contents. However, partially digested remains may be difficult to accurately identify by traditional visual analysis. Here we evaluate the effectiveness of DNA barcoding to identify digested piscine prey remains in invasive Blue Catfish Ictalurus furcatus, non-native, but established Channel Catfish Ictalurus punctatus and native White Catfish Ameiurus catus from Chesapeake Bay, USA. Stomach contents were examined and piscine prey items were scored as lightly digested, moderately digested or severely digested. A 652 base pair region of the cytochrome c oxidase subunit I (COI-5P) mitochondrial DNA gene was sequenced for each prey item. Edited barcode sequences were compared to locally-caught and validated reference sequences in BOLD (Barcode of Life Database). A large majority of prey items were sufficiently digested to limit morphological identification (9.4 % to species and an additional 12.1 % to family). However, overall barcoding success was high (90.3 %) with little difference among the digestion classifications. Combining morphological and genetic identifications, we classified 91.6 % of fish prey items to species. Twenty-three fish species were identified, including species undergoing active restoration efforts (e.g., Alosa spp.) and commercially important species, e.g., Striped Bass Morone saxatilis, White Perch Morone americana, American Eel Anguilla rostrata and Menhaden Brevoortia tyrannus. We found DNA barcoding highly successful at identifying all but the most heavily degraded prey items and to be an efficient and effective method for obtaining diet information to strengthen the resolution of trophic analyses including diet comparisons among sympatric native and non-native predators.     

双低菜粕高水平替代饲料鱼粉对大黄鱼潜在风险的评估:生长、健康和营养价值

研究从生长、健康和营养价值方面评估了高水平的双低菜粕替代饲料鱼粉对大黄鱼潜在的危害。在鱼粉含量60%的基础饲料（FM）上按照质量分数用双低菜粕分别替代15%（CM15）、30%（CM30）、60%（CM60）和100%（CM100）的鱼粉,配制成5种实验饲料。每种饲料投喂5个网箱的大黄鱼[初重（135.38±1.02）g],即每个处理5个重复,进行12周的养殖实验。结果表明,当双低菜粕替代水平在15%和30%时,大黄鱼的生长及饲料系数并没有受到显著性的影响。然而,当替代水平高于30%时,大黄鱼的末重和特定生长率均显著降低,而饲料系数显著升高（P < 0.05）。当替代水平达到100%时,大黄鱼摄食率达到最高值而肥满度达到最低值（P < 0.05）。在组织形态方面,大黄鱼摄食双低菜粕替代的饲料后肠道绒毛的弯曲程度减少并且排列更加不规则,而肝细胞则呈现出圆形空泡状并伴随着细胞核的偏移。对大黄鱼骨骼进行X-射线扫描发现,摄食双低菜粕的大黄鱼椎体和头部出现了畸形。在营养价值方面,双低菜粕替代鱼粉并未显著影响大黄鱼背肌的脂肪含量、蛋白含量和氨基酸组成,然而脂肪酸组成受到了显著影响,即N-6系列脂肪酸含量显著升高,而DHA与EPA含量显著降低（P < 0.05）。根据欧洲食品安全局（EFSA）的相关标准,这些营养价值的变化并没有影响大黄鱼作为健康食品的功能。由此可见,高水平（60%和100%）的双低菜粕替代鱼粉对大黄鱼的负面影响主要表现为降低大黄鱼的生长性能、改变肠道和肝脏组织形态,以及影响大黄鱼的骨骼健康。然而,双低菜粕替代鱼粉养殖大黄鱼的肌肉仍然符合人类的膳食要求。因此,双低菜粕替代鱼粉并没有影响大黄鱼作为食用鱼的营养价值。     

Diet and feeding periodicity of Cox's gudgeon Gobiomorphus coxii (Krefft) in a south-eastern Australian stream

A 24 h study of stomach contents and simultaneous sampling of the drift and benthos was used to describe the diet and assess the prey selectivity of Cox's gudgeon, Gobiomorphus coxii , within a small coastal stream in south-eastern Australia. Prey items consisted entirely of macroinvertebrates with most being ephemeropterans and trichopterans. The greatest number and largest volume of macroinvertebrates were in the stomachs of fish collected just after sunset through until several hours after sunrise. Both the number of empty stomachs and proportion of digested material in the stomach contents peaked at mid-day. Overall, it was concluded that G. coxii was a nocturnal feeder that specialized on benthos, although some evidence of opportunistic prey selection was apparent. Gobiomorphus coxii was also size selective, preferring relatively large prey from both the drift and benthos.     

Prey selection by young trout fry (Salmo trutta)

Factors influencing the choice of invertebrate food by early feeding trout are sought, special attention being given to the dimensions of the prey. The stomach contents of fish of similar age feeding on lake plankton and drifting stream fauna are compared. Little consistent preference was displayed for the smaller planktonic organisms but prey size was of importance in the distribution of stream benthos among predators of different lengths. The width of the prey was of greater significance to selection than the mean volume or the area of the largest surface of the food items. When the fish were in obvious competition for food the correlation between predator length and all prey dimensions improved.     

The use of otoliths as indicators of Little Penguin Eudyptula minor diet

The validity of using otoliths from stomach contents quantitatively to determine the number and size of fish consumed was tested on Little Penguins. They were fed different meal sizes of known number and size of fish and the stomach contents were recovered after various time intervals. There were no differences in estimates of original fish size when calculated from otolith length or weight. Rate of digestion of otoliths tended to decrease with increased meal size but increased with time after ingestion. Digestion of otoliths proceeds rapidly and, if ignored, estimates of numbers of fish consumed and of original fish size can be significantly underestimated. This problem can be partially solved by inspection of otolith condition and restricting calculations of fish size to otoliths unaffected by digestion. Many factors introduce variations into rate of otolith degradation and further species-specific studies are required before appropriate correction factors can be applied.     

The use of otoliths as indicators of Little Penguin Eudyptula minor diet

The validity of using otoliths from stomach contents quantitatively to determine the number and size of fish consumed was tested on Little Penguins. They were fed different meal sizes of known number and size of fish and the stomach contents were recovered after various time intervals. There were no differences in estimates of original fish size when calculated from otolith length or weight. Rate of digestion of otoliths tended to decrease with increased meal size but increased with time after ingestion. Digestion of otoliths proceeds rapidly and, if ignored, estimates of numbers of fish consumed and of original fish size can be significantly underestimated. This problem can be partially solved by inspection of otolith condition and restricting calculations of fish size to otoliths unaffected by digestion. Many factors introduce variations into rate of otolith degradation and further species-specific studies are required before appropriate correction factors can be applied.     

Feeding behaviour of free-ranging penguins determined by oesophageal temperature

Sea birds play a major role in marine food webs, and it is important to determine when and how much they feed at sea. A major advance has been made by using the drop in stomach temperature after ingestion of ectothermic prey. This method is less sensitive when birds eat small prey or when the stomach is full. Moreover, in diving birds, independently of food ingestion, there are fluctuations in the lower abdominal temperature during the dives. Using oesophageal temperature, we present here a new method for detecting the timing of prey ingestion in free-ranging sea birds, and, to our knowledge, report the first data obtained on king penguins (Aptenodytes patagonicus). In birds ashore, which were hand-fed 2-15 g pieces of fish, all meal ingestions were detected with a sensor in the upper oesophagus. Detection was poorer with sensors at increasing distances from the beak. At sea, slow temperature drops in the upper oesophagus and stomach characterized a diving effect per se. For the upper oesophagus only, abrupt temperature variations were superimposed, therefore indicating prey ingestions. We determined the depths at which these occurred. Combining the changes in oesophageal temperatures of marine predators with their diving pattern opens new perspectives for understanding their foraging strategy, and, after validation with concurrent applications of classical techniques of prey survey, for assessing the distribution of their prey.     

Estimating size dependent diet,advection, diffusion,catchability and mortality applying a normal distribution model to spreading of released turbot

In a field experiment, breed turbot of an average length between 7–25 cm was released into a small bay. During the following week; diffusion rate, mortality and catchability of the released fish was estimated daily, by using a normal distribution model. In this model the size of the fish was incorporated in such a way, that it was possible to retrieve information about how each of the estimated parameters depended on the fish size. In addition, the length of the prey items found in the stomach was measured and compared to the prey length available in the habitat. Turbot was found to adjust to eat the available prey fast, after 3 days approximately 100% had detectable signs of prey in their stomach.     

Vertebrate diets derived from trophically transmitted fish parasites in the Bothnian Bay

Parasites that are transmitted through predator–prey interactions may be used as indicators of trophic relationships between organisms. Yet, they are rarely used as such in the construction of topological (predator–prey) food webs. We constructed food webs of vertebrate trophic interactions using observed diet alone, trophically transmitted parasites alone, and the combination of the two based on data from 31 species of fish from the Bothnian Bay, Finland. The fish food web contained 530 links derived from observed diet, 724 links inferred from parasitism, and 1,058 links calculated from a combination of both stomach contents and parasites. This sub-web constructed from stomach contents had a mean of 17.1 links per fish species, while that using parasites had 23.4 links per fish. Combining the two diet indicators yielded 34.1 links per fish species, illustrating the complementarity of the two methods. Mean number of prey species per fish species was 12.5 using observed diet items, 15.8 using parasites, and 24.5 using both measures together. Mean number of predators per fish species was 7.4 using observed diet, 11.7 using parasites and 15.0 using both. A positive correlation was found between the mean number of parasites and the number of prey taxa in the diet among the fishes. Omnivorous fish had the highest diversity of both parasite species and prey items, while benthophagous fish had among the lowest. Mean total abundance and mean total prevalence of parasites correlated positively with fish size, with piscivores being the largest with the highest abundance and prevalence, while planktivores and benthivores had the lowest. Trophically transmitted parasites may be used to help construct vertebrate sub-webs and derive information about food web processes. Parasites alone provided equivalent if not more information than observed diet. However, resolution is improved by using parasites and observed diet together.     

The influence of meal size on prey DNA detectability in piscivorous birds

Molecular methods allow noninvasive assessment of vertebrate predator–prey systems at high taxonomic resolution by examining dietary samples such as faeces and pellets. To facilitate the interpretation of field‐derived data, feeding trials, investigating the impacts of biological, methodological and environmental factors on prey DNA detection, have been conducted. The effect of meal size, however, has not yet been explicitly considered for vertebrate consumers. Moreover, different noninvasively obtained sample types remain to be compared in such experiments. Here, we present a feeding trial on abundant piscivorous birds, Great Cormorants (Phalacrocorax carbo), to assess meal size effects on postfeeding prey DNA detection success. Faeces and pellets were sampled twice a day after the feed of large (350–540 g), medium (190–345 g) and small (15–170 g) fish meals contributing either a large (>79%) or small (<38%) share to the daily consumption. Samples were examined for prey DNA and fish hard parts. Molecular analysis of faeces revealed that both large meal size and share had a significantly positive effect on prey DNA detection rate postfeeding. Furthermore, large meals were detectable for a significantly longer time span with a detection limit at ~76 hr and a 50% detection probability at ~32 hr postfeeding. In pellets, molecular methods reliably identified the meal consumed the previous day, which was not possible via morphological analysis or when examining individual faeces. The less reliable prey DNA detection of small meals or meal shares in faeces signifies the importance of large numbers of dietary samples to obtain reliable trophic data.     

The influence of prey size and abundance, and individual phenotype on prey choice by the three-spined stickleback, Gasterosteus aculeatm L.

Prey selection behaviour of three-spined sticklebacks, Gasterosteus aculeatus L., was studied in two experiments. Where possible, the experimental apparatus satisfied the assumptions of the simplest optimal diet model (the basic prey model); prey were presented sequentially, the fish could not search for and handle prey at the same time, and net energy gain, handling time and encounter rate were fixed. Experiment 1 presented fish with a range of Asellus sizes so that pursuit ( p ) and handling ( h ) time could be related to prey size. Published energy values of Asellus together with pursuit and handling times were used to calculate E /( p+h ) for Asellus measuring 3,4,5,6,7 and 9 mm. Pursuit times did not differ with prey size but handling times did. E /( p+h ) was very variable particularly at the larger prey sizes. Experiment 2 presented fish with two sequences of prey differing in the encounter rate with the most profitable prey sizes. Fish did not select the diet predicted by the basic prey model tending to always ignore the largest prey even when net energy gain would have been maximized by including them in the diet. Further analysis showed that the probability of a prey size being taken was a function of prey size, fish stomach fullness and encounter rate. It is concluded that the basic prey model is too simple to capture the behaviour of the fish. One of its main faults is that the changing state of the fish through the feeding bout is ignored.