Food habits, occurrence, and population structure of the bat ray, Myliobatis californica, in Humboldt Bay, California

The bat ray, Myliobatis californica, is the most common, large predatory fish in Humboldt Bay, California. To prevent bat ray predation on cultured oyster beds, much effort has focused on reducing their population. A 13 month study was conducted in Humboldt Bay to examine the rays' use of the bay, population structure, and feeding ecology. Bat rays are seasonally found in Humboldt Bay during the spring, summer, and early fall months. Adult rays were most abundant during the summer months, while numbers of juveniles increased through the summer and early fall, indicating that Humboldt Bay is an important pupping and nursery ground. Very few rays were found in the bay during the colder months of the year. This decline in abundance is attributed to cooler water temperatures (< 10° C). The stomach contents from 503 bat rays were examined. Overall, clams were the predominant prey item. The index of importance and Shannon-Weiner diversity index indicated that food habits of bat rays change with increasing size. Larger prey items and a wider variety of prey items were consumed as rays increased in size. Dietary importance of small clams and Crangon shrimp decreased with increasing ray size, while the importance of larger prey items (e.g. large clams, Cancer crabs, blue mud shrimp, echiuran worms) increased. Predation on oysters was rare. Differences between the diets of male and female rays of similar sizes were also observed.     

DIET AND FORAGING BEHAVIOR OF SEA OTTERS IN SOUTHEAST ALASKA

Abstract: Direct observations of feeding sea otters ( Enhydra lutris ) at 11 sites in southeast Alaska showed infaunal clams to be the primary prey utilized by otters throughout the region. Foraging dive times associated with clam and sea urchin prey were significantly longer than those for more easily captured prey (crabs and mussels). Dive times and surface intervals were also generally correlated with water depth or apparent difficulty in obtaining buried prey. Male otters, which fed more extensively on clams than females, made significantly longer foraging dives than females. Foraging success remained high, even at sites where prey numbers were found to be very low during a related study. The very deeply burrowing geoduck clam ( Panope abrupta ), while common at several otter feeding sites, was rarely captured by otters. These results, combined with those of a companion study on prey numbers, indicate that butter clams ( Saxidomus giganteus ) account for the majority of the sea otter diet in southeast Alaska, and that sea urchins may represent relatively short-term prey in comparison to infaunal bivalves in regions where both prey types co-exist. Furthermore, the importance of butter clams in the sea otter diet and the tendency for this bivalve to retain chronically high levels of paralytic shellfish poisoning toxins in southeast Alaska increases the probability that toxic phytoplankton blooms influence sea otter distribution in this region.     

Recognizing biotic breakage of the hard clam, Mercenaria mercenaria caused by the stone crab, Menippe mercenaria: An experimental taphonomic approach

The ability to assign lethal traces left on prey to particular durophagous predators enhances our understanding of predation pressure in the fossil record. To determine whether stone crabs (Menippe mercenaria Say 1818) leave diagnostic traces in the act of feeding on hard clams (Mercenaria mercenaria Linnaeus 1758), live clams were offered to crabs in laboratory aquaria over several months and the fragments produced during predation were examined for diagnostic breakage patterns. These fragments were then compared both macroscopically and using scanning electron microscopy to the fracture patterns produced by tumbling clams in a rock tumbler which simulated breakage during transport in the surf zone, and crushing clams using an Instron which simulated breakage resulting from sediment compaction. Fossil specimens of Mercenaria mercenaria were also examined to determine whether the criteria for recognizing predation traces generated experimentally could be recognized. While not all acts of predation produce diagnostic traces, when larger fragments (greater than 50% shell remaining) are produced during feeding, predatory-diagnostic breakage ranges from 70 to 80%. Macroscopic breakage patterns generated during the predation experiments were also present in fossil specimens. Damage caused by abiotic mechanisms (tumbling and crushing) is highly unlikely to be confused with damage produced by this predator.     

Cloning and localization of MCdef, a defensin from Manila clams (Ruditapes philippinarum)

A defensin-like peptide was previously detected in hemocytes of Manila clams (Ruditapes philippinarum). In the current study, we cloned and characterized this defensin, designated MCdef. Cloning produced a full-length gene sequence of 201 bp predicted to encode a 66-amino-acid precursor protein maturing to a 44-amino-acid residue. Amino acid sequence analysis showed that MCdef is similar to defensins from marine mollusks and ticks. Phylogenetic analysis suggested that MCdef is closely related to defensins from Mytilus galloprovincialis (Mediterranean mussel) and Crassostrea gigas (Pacific cupped oyster). The three-dimensional structure of MCdef was modeled using the solution structure of C. gigas defensin as a template. With the exception of three variable loop areas, the modeled structure of MCdef was identical to that of C. gigas defensin. MCdef antiserum was raised against a synthetic MCdef peptide and verified by Western blotting using recombinant MCdef. RT-PCR analysis demonstrated high levels of MCdef mRNA in hemocytes and adductor, foot, gill, mantle, palp, and siphon tissues of Vibrio tapetis-infected Manila clams, whereas in V. tapetis-uninfected Manila clams, the level of MCdef mRNA was low in adductor, palp, and siphon tissues and even lower in the other tested tissues. Immunohistochemical analysis revealed high MCdef expression was detected in the gill, the mantle, and the digestive tubules of the diverticulum of V. tapetis-infected Manila clams. Minimum inhibitory concentration (MIC) of the purified rMCdef was determined. MCdef showed highest activity against Streptococcus iniae and Staphylococcus aureus.     

As good as dead? Sublethal predation facilitates lethal predation on an intertidal clam

Although ecologists have speculated that sublethal predation can impact prey dynamics, consequences of these predator effects have seldom been experimentally tested. In soft‐sediment marine communities, fishes crop extended feeding siphons of buried clams, potentially causing clams to reduce their burial depth, thereby enhancing their susceptibility to excavating lethal predators. We simulated cropping of the confamilial clams, Protothaca staminea and Venerupis philippinarum, by removing the top 40% of siphons, which caused each species to burrow 33–50% shallower than conspecifics with intact siphons. To examine subsequent consequences of reduced burial depth, we exposed cropped and intact clams to natural levels of predation in the field. Because of a naturally longer siphon, Protothaca, even after cropping, remained at relatively safe burial depths. In contrast, siphon cropping nearly doubled the mortality rate of Venerupis. Thus, while sublethal predation facilitates lethal predation, this linkage depends on specific life history characteristics, even among ecologically similar species.     

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.     

Methods of estimating marine mammal diets: A review of validation experiments and sources of bias and uncertainty

Diet estimation in marine mammals relies on indirect methods including recovery of prey hard parts from stomachs and feces, quantitative fatty acid signature analysis (QFASA), stable isotope mixing models, and identification of prey DNA in stomach contents and feces. Experimental evidence (9 species/13 studies) shows that digestion strongly influences the proportion and size of otoliths that can be recovered in feces. Number correction factors (NCF) and digestion coefficients have been experimentally determined to reduce the biases in fecal analysis. Correction factors and coefficients have not been determined for diet estimated from stomach contents. QFASA estimates which prey species and amounts must have been eaten to account for the fatty acid composition of the predator. Experimental studies on mammals and seabirds (9 species/10 studies) indicate that accurate estimates of diet can be determined using QFASA. Stable isotope mixing models provide rather coarse taxonomic resolution of diet composition. Prey DNA analysis shows promise as a method to estimate the species composition of diet, but further development and testing is needed to validate its use. Obtaining a representative sample from marine mammal populations is a significant challenge. Therefore, the use of complementary methods is recommended to obtain the most informative results.     

Effect of Experience on Prey Species Selection by the Bivalve Feeder Halla Okudai (Polychaeta: Lysaretidae)

We report on the effects of previous foraging experience on prey-selection by the bivalve feeding polychaete Halla okudai, including whether there was evidence of frequency-dependent predation. Three separate batches of H. okudai were maintained for 30 days on clams, mussels, or oysters, before being offered a choice among these three prey. Initially individuals from all three treatments consumed more clams than mussels, and no oysters. As the number of clams was depleted the polychaetes shifted their diet to include a greater proportion of mussels, but even after 20 days oysters were only eaten by polychaetes that had been previously acclimated to them. Nevertheless, polychaetes from each treatment inspected significantly more of the prey species to which they had initially been acclimated, suggesting that previous experience may increase the likelihood of certain prey being detected. When individuals of H. okudai were repeatedly offered the same prey species, handling time did not decrease (and therefore prey profitability did not increase) with experience, which may be because H. okudai paralyses its prey with toxic mucus. Since repeated experience of the same prey species gives no advantage in terms of reduced handling time, we suggest this may be why this species does not show frequency-dependent prey-selection.     

Effect of Experience on Prey Species Selection by the Bivalve Feeder Halla Okudai (Polychaeta: Lysaretidae)

We report on the effects of previous foraging experience on prey-selection by the bivalve feeding polychaete Halla okudai, including whether there was evidence of frequency-dependent predation. Three separate batches of H. okudai were maintained for 30 days on clams, mussels, or oysters, before being offered a choice among these three prey. Initially individuals from all three treatments consumed more clams than mussels, and no oysters. As the number of clams was depleted the polychaetes shifted their diet to include a greater proportion of mussels, but even after 20 days oysters were only eaten by polychaetes that had been previously acclimated to them. Nevertheless, polychaetes from each treatment inspected significantly more of the prey species to which they had initially been acclimated, suggesting that previous experience may increase the likelihood of certain prey being detected. When individuals of H. okudai were repeatedly offered the same prey species, handling time did not decrease (and therefore prey profitability did not increase) with experience, which may be because H. okudai paralyses its prey with toxic mucus. Since repeated experience of the same prey species gives no advantage in terms of reduced handling time, we suggest this may be why this species does not show frequency-dependent prey-selection.     

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.     

The effect of benthic prey abundance and size on red knot (Calidris canutus) distribution at an alternative migratory stopover site on the US Atlantic Coast

A population decline of the western Atlantic red knot (Calidris canutus rufa) has been linked to food limitation during the spring migratory stopover in Delaware Bay, USA. The stopover ecology at potential alternative sites has received little attention. We studied factors affecting red knot habitat selection and flock size at a coastal stopover site in Virginia in 2006–2007. The most common potential prey items were coquina clams (Donax variabilis) and crustaceans. Red knot foraging sites had more clams and crustaceans than unused sites in 2006. Prey abundance increased during the 2007 stopover period and remained high after the red knot peak. Red knot flock size in 2007 increased with mean clam shell length, and probability of flock presence decreased with increasing distance from night use locations. Our results suggest that red knots preferred coquina clams and that these clams were not depleted during the stopover period in 2007. Thus prey abundance did not appear to be a population-limiting factor at this coastal stopover site in Virginia in that year. Protection of coastal sites outside of Delaware Bay, many of which have been altered by human development, would likely benefit red knot population recovery, as they can apparently provide abundant food resources during at least some years.     

崂山湾人工鱼礁区星康吉鳗摄食生态及食物网结构

根据2015年4月至2017年1月于崂山湾人工鱼礁区地笼网和延绳钓捕获的279尾星康吉鳗样本,从胃含物组成、食性类型、摄食等级、营养生态位和营养级等方面对其摄食生态进行研究,同时结合海区许氏平鲉、大泷六线鱼、斑头鱼、褐菖鲉、花鲈等9种鱼类的胃含物分析结果,构建人工鱼礁区鱼类关键种的简化食物网模型.食性研究结果表明:星康吉鳗共摄食7类30余种饵料,虾类是其最主要的饵料类群,其次为鱼类和头足类,大泷六线鱼、方氏云鳚、鹰爪虾、玉筋鱼和日本鼓虾等是其优势饵料.星康吉鳗的饵料生物组成随肛长和季节发生显著变化.四季均以鱼类和虾类为主,春季胃含物中包括头足类,秋季包括头足类和蟹类,冬季亦有蟹类出现.肛长≤120 mm的星康吉鳗主要摄食鱼卵和鹰爪虾,120~130 mm肛长组主要摄食玉筋鱼和日本鼓虾,肛长>130 mm的星康吉鳗主要摄食大泷六线鱼和方氏云鳚.其摄食强度也随季节和肛长而变化,空胃率的季节性差异显著,平均胃饱满系数的季节性差异不显著,不同肛长组的空胃率和平均胃饱满系数均不存在显著差异.人工鱼礁区简化食物网结构显示:鱼类关键种的营养级均在3级以上,星康吉鳗的营养级为4.636,处于海区食物网的最顶端.虾类、蟹类、端足类和软体动物等是鱼类关键种的主要饵料,甲壳类、方氏云鳚、大泷六线鱼和玉筋鱼是高营养级鱼类花鲈和星康吉鳗的主要饵料.     

Activity, feeding behaviour and diet of Ogcocephalus vespertilio in southern west Atlantic

The Brazilian batfish Ogcocephalus vespertilio is a nocturnal and early morning predator on bottom invertebrates, staying stationary and hidden in rock holes, crevices or among bottom rocks during the day. While active, it searches for prey walking along the bottom with the help of its specialized paired fins, with the illicium protracted and oscillating or exploring the substrate. The prey are either snapped up from the bottom, after visual detection, or dug out with use of the mouth and rostrum. It feeds on crustaceans (hermit crabs, true crabs, shrimps, amphipods, porcelain crabs, isopods and mysid shrimps), molluscs (snails, sea slugs and clams), polychaete worms (mostly Errantia) and echinoderms (sea urchins and brittle stars).     

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.     

Control of foraging behavior of individuals within an ecosystem context: the clam Macoma balthica and interactions between competition and siphon cropping

Macoma balthica (L.) is a common clam of the estuarine seafloor, belonging to an important group of invertebrates possessing the capacity to choose between the two fundamental modes of feeding available, using its siphon to inhale either suspended food particles from the water or food particles deposited on the sediment surface. Field experiments demonstrate that intraspecific competition, effects of other competing benthic invertebrates, and complex interactions between competition and partial predation (siphon cropping by fishes) modify the foraging behavior of Macoma. When protected by caging from siphon nipping by fishes, Macoma demonstrated greater siphon regeneration at lower density, indicating the importance of competition for limited resources. In the absence of siphon croppers, these same clams also exhibited more deposit feeding at the lower density either because of improved ability to deposit-feed with longer siphons or because deposited foods become more rapidly depleted than suspended foods on local spatial scales. Addition of siphon-nipping fishes caused greater reductions in siphon size of clams at lower density, presumably because the intensity of nipping per clam was greater where clam targets were fewer and because deposit feeding, which was more intense at lower densities, confers a greater risk of cropping from greater siphon extension and activity than characterize suspension feeding. Deposit feeding by Macoma was reduced in the presence of siphon croppers at both high and low density of clams, but the intensity of deposit-feeding activity at low density was substantially higher than predicted by additive effects of clam density and cropping. This suggests operation of a balancing strategy in Macoma whereby it is accepting greater risks of partial predation when rewards of greater food harvest are larger. The surprising failure to adopt a risk-averse stratery may be explained by the non-lethal nature of partial predation, which renders siphon loss an energetic penalty replacable through regeneration. The presence of a bed of suspension-feeding Rangia cuneata also altered foraging behavior of Macoma by inducing a switch to more intense deposit feeding, in response either to the documented near-bottom depletion of suspended foods or to likely enhanced biodeposition from feces and pseudofeces. The induction of greater deposit feeding by the presence of this competing suspension feeder led to greater siphon losses during exposure to croppers because Macoma was practicing more risky feeding behavior. This enhanced loss of siphon tissues to croppers in the presence of the suspension-feeding Rangia induced an interaction between the effects of siphon croppers and Rangia, such that Macoma exhibited a larger switch away from deposit feeding in the presence of siphon croppers when Rangia were also present. Clearly, the foraging decisions made by individuals can only be understood in a broad holistic context of population, community, and ecosystem processes.     

Bioaccumulation and depuration of enteroviruses by the soft-shelled clam, Mya arenaria.

Low levels of feces-associated natural virus, simulating virus numbers estimated to exist in moderately polluted shellfish-growing waters, were used to evaluate the effectiveness of depuration as a virus depletion procedure in soft-shell clams. Depuration effectiveness depended upon the numbers of virus bioaccumulated and whether virus was solids associated. Virus uptake was greatest when viruses were solids associated and pollution levels were equivalent or greater than those likely to be found in grossly polluted growing waters. Virtually all bioaccumulated feces-associated natural virus was deposited within either the hepatopancreas or siphon tissues. Viruses usually were eliminated within a 24- to 48-h depuration period. Dependence upon depuration of clams to elimate health hazards of virus etiology involved a risk factor not measureable in the study. The greatest reduction of health risks would come from the routine depuration of clams harvested from growing waters of good sanitary quality.     

Bioaccumulation and depuration of enteroviruses by the soft-shelled clam, Mya arenaria.

Low levels of feces-associated natural virus, simulating virus numbers estimated to exist in moderately polluted shellfish-growing waters, were used to evaluate the effectiveness of depuration as a virus depletion procedure in soft-shell clams. Depuration effectiveness depended upon the numbers of virus bioaccumulated and whether virus was solids associated. Virus uptake was greatest when viruses were solids associated and pollution levels were equivalent or greater than those likely to be found in grossly polluted growing waters. Virtually all bioaccumulated feces-associated natural virus was deposited within either the hepatopancreas or siphon tissues. Viruses usually were eliminated within a 24- to 48-h depuration period. Dependence upon depuration of clams to elimate health hazards of virus etiology involved a risk factor not measureable in the study. The greatest reduction of health risks would come from the routine depuration of clams harvested from growing waters of good sanitary quality.     

Dietary shifts of sympatric buteos during a prey decline

Summary Diets of nesting Red-tailed Hawks (Buteo jamaicensis) and Ferruginous Hawks (Buteo regalis) were sampled before and after a decline in the hawks' principal prey species. Diets of pairs that shared their foraging ranges with interspecifics were contrasted with those of pairs whose home ranges did not overlap with interspecifics. Current theory predicts that diets should diverge during prey shortages and that overlap should be especially reduced in ranges shared with interspecifics. Species composition of the two hawks' diets diverged during the prey shortage, but the divergence was most pronounced in hawks that did not share foraging ranges with interspecifics. In contrast to predictions, the two species converged on similar-sized prey during the prey shortage. Available data on differences in habitat composition and prey availability at the sample sites did not explain the deviations from the predicted response. Although our findings on diet shifts differ from those of most other studies, the implications are consistent with observations of others who have studied assemblages of mammal-feeding raptors. We conclude that diet composition of the hawks we studied was not directly affected by interspecific competition and that shifts in diet overlap during prey shortages do not necessarily imply that interspecific competition is occurring.