Prey selection by molluscivorous cichlids foraging on a schistosomiasis vector snail,Biomphalaria glabrata

Summary This paper considers prey size selection by four molluscivorous cichlids feeding on the intermediate host snail of Schistosoma parasites, Biomphalaria glabrata. Haplochromis ishmaeli obtains its prey by crushing the snails between the pharyngeal jaws, whereas H. xenognathus, H. sauvagei and Macropleurodus bicolor apply both pharyngeal crushing and oral shelling. The fishes crushed significantly more snails with the highest reward in biomass per second of crushing. Oral shelling occurred far less often than pharyngeal crushing. Encounter rates with prey showed significant variations between different size classes of prey. The fish have no overall knowledge of snail availability in a tank. The probability that a snail will be eaten at encounter, calculated from the number encountered and the number eaten, reflects the prey size preference of the fish. Those snails with the highest biomass/crushing-time ratio had the highest probability of being crushed; observed and predicted prey size preferences corresponded well. Although for oral shelling the potential reward in biomass per second is of the same magnitude as for crushing, the probability of successful shelling is very low. Apparently the fish prefer prey with lowest risks.     

Prosobranch mollusc species- and size-preferences of Sargochromis codringtonii (Cichlidae) in Lake Kariba,Zimbabwe

From the stomach contents of 28 wild-caught Sargochromis codringtonii from Lake Kariba the size and species of snails consumed were estimated from opercula found. An energetic cost/benefit ratio of the various sizes of the most frequent prey, Bellamya capillata, was calculated. The size of snails consumed increased significantly with fish size, with large fish showing individual size preferences. Ingested snails were of sizes with relatively low energetic cost/benefit ratios.     

Home-field advantage: native signal crayfish (Pacifastacus leniusculus) out consume newly introduced crayfishes for invasive Chinese mystery snail (Bellamya chinensis)

The introduction of non-indigenous plants, animals and pathogens is one of today’s most pressing environmental challenges. Freshwater ecologists are challenged to predict the potential consequences of species invasions because many ecosystems increasingly support novel assemblages of native and non-native species that are likely to interact in complex ways. In this study we evaluated how native signal crayfish (Pacifastacus leniusculus) and non-native red swamp crayfish (Procambarus clarkii) and northern crayfish (Orconectes virilis) utilize a novel prey resource: the non-native Chinese mystery snail (Bellamya chinensis). All species are widespread in the United States, as well as globally, and recent surveys have discovered them co-occurring in lakes of Washington State. A series of mesocosm experiments revealed that crayfish are able to consume B. chinensis, despite the snail’s large size, thick outer shell and trapdoor defense behaviour. Crayfish exhibited size-selective predation whereby consumption levels decreased with increasing snail size; a common pattern among decapod predators. Comparison of prey profitability curves—defined as the yield of food (weight of snail tissue) per second of feeding time (the time taken to crack the shell and consume the contents)—suggests that small and very large snails may represent the most profitable prey choice. By contrast, previous studies have reported the opposite pattern for crayfish consumption on thin-shelled snails. For all snail size classes, we found that native P. leniusculus and invasive O. virilis consumed greater numbers of snails than invasive P. clarkii. Moreover, P. leniusculus consistently handled and consumed snails at a faster pace compared to both invasive crayfishes across the range of snail sizes examined in our study. These results suggest not only that B. chinensis is a suitable food source for crayfish, but also that native P. leniusculus may ultimately out-consume invasive crayfishes for this new prey resource.     

Experimental tumbling of <Emphasis Type="Italic">Dreissena polymorpha</Emphasis>: implications for recognizing durophagous predation in the fossil record

Shell damage left by predators constitutes an important source of information on predator–prey interactions. However, recognition of the origins of shell damage can often be controversial, and needs to be assessed cautiously. More specifically, differentiation between predation- and abiotic-induced shell damage remains challenging. Here, we show the results of tumbling experiments using a bivalve species Dreissena polymorpha in order to determine rates and patterns of shell damage induced by physical forces in high-energy conditions. It is demonstrated that, in contrast to durophagous fish and crab predation, abiotic-induced fragmentation and damage are typically characterized by the presence of distinct abrasive scratches and wear scars on the surface of shell fragments. Furthermore, fragmented shells typically reveal a wide size distribution, and a different degree of sphericity and roundness resulting from abrasion. Importantly, large shell fragments commonly display smooth edges. These data suggest that durophagous predation, which typically induces fragmentation into large and angular shell fragments bearing no wear scars, can be reliably recognized both in present-day environments and in the fossil record.     

Integration of an invasive consumer into an estuarine food web: direct and indirect effects of the New Zealand mud snail

Introduced species interact both directly and indirectly with native species. We examine interactions between the introduced New Zealand mud snail (Potamopyrgus antipodarum) and native estuarine invertebrates and predators through experiments and field studies. A widely held management concern is that when P. antipodarum, which has low nutritional value, becomes abundant, it replaces nutritious prey in fish diets. We tested two key components of this view: (1) that fish consume, but get little direct nutritional value from P. antipodarum; and (2) that P. antipodarum has an indirect negative effect on fish by reducing the energy derived from native prey. We also examined predation by the native signal crayfish, Pacifastacus leniusculus. Laboratory feeding trials showed that both crayfish and fish consume P. antipodarum, a direct effect. Crayfish consumed and successfully digested higher numbers of snails than did fish [Pacific staghorn sculpin (Leptocottus armatus), three spine stickleback (Gasterosteus aculeatus), and juvenile starry flounder (Platicthys stellatus)]. P. antipodarum occurred at low frequencies in the stomachs of wild-caught fish. More interesting were the indirect effects of this invader, which ran counter to predictions. P. antipodarum presence was associated with no change or an increase in the amount of energy derived from native prey by predators. The presence of P. antipodarum also led to increased consumption of and preference for the native amphipod Americorophium salmonis over the native isopod Gnorimosphaeroma insulare. This is an example of short-term, asymmetric, apparent competition, in which the presence of one prey species (snails) increases predation on another prey species (the amphipod).     

Biomechanical limits to ecological performance: mollusc-crushing by the Caribbean hogfish, Lachnolaimus maximus (Labridae)

Functional limitations on feeding ability were investigated in the mollusc-crushing Caribbean hogfish, Lachnolaimus maximus (Labridae). Two constraints were proposed to limit the maximum size prey L. maximus can eat: pharyngeal jaw gape and crushing force. These factors yield diflferent quantitative expectations for the relationship between fish size and maximum prey size. Their relative importance for predation on a frequently consumed gastropod ( Cerithium litteratum ) was investigated in laboratory performance tests designed to determine the largest snails fish could eat. Cerithium predation was found to be force limited rather than gape limited. The importance of this functional constraint in determining the largest Cerithium consumed by wild fish was examined by comparing hogfish feeding capability, as determined by the performance tests, to the maximum size snails found in the stomach contents of field-collected fish. Crushing ability appears to limit Cerithium predation in natural fish populations. The utility of performance testing for determining the functional and ecological importance of morphology is discussed.     

Feeding behavior of black carp Mylopharyngodon piceus (Pisces: Cyprinidae) on fry of other fish species and trematode transmitting snail species

Fish raised in aquaculture ponds may get infected with fishborne zoonotic trematodes (FZT) during the nursing stage. Freshwater snails serve as intermediate hosts for FZT and we wanted to explore the possibility of controlling snails by stocking nursery ponds with a few juvenile specimens of the mollusc-eating fish, black carp (Mylopharyngodon piceus). Obviously, the risk that black carp might also prey on the juvenile fishes in nursery ponds should first be assessed. Laboratory trials showed that all size classes of juvenile black carp consumed fry of common carp (Cyprinus carpio) even when offered snails as food; the odds of survival of fry from tanks with medium sized and large black carp was 5.6% and 39.9%, respectively of that of fry in tanks with small sized black carp. Since the large black carp also consumed fewer snails than medium sized fish, we believe that large specimens were stressed in the experimental aquaria. Under semi-field conditions, presence of the black carp had no effect on survival of fry of Oreochromis niloticus and C. carpio both in the absence and presence of snails as alternative food. The black carp consumed most snails offered with the exception of some of the large snails. We conclude that under field conditions, predation by black carp on fish fry is minimal and field trials in nursery ponds are warranted. Due to the risks that black carp pose to native imperiled snails and other molluscs, trials should be restricted to ponds within the fish’s native or existing range.     

Independent and combined effects of multiple predators across ontogeny of a dominant grazer

Ecosystems host multiple coexisting predator species whose interactions may strengthen or weaken top–down control of grazers. Grazer populations often exhibit size‐structure, but the nature of multiple predator effects on suppression of size‐structured prey has seldom been explicitly considered. In a southeastern US salt‐marsh, we used both field (additive design) and mesocosm (additive‐substitutive design) experiments to test the independent and combined effects of two species of predatory crab on the survival and predator‐avoidance behavior (i.e. a non‐consumptive effect) of both juveniles and adults of a dominant grazing snail. Results showed: 1) juvenile snails were more vulnerable to predation; 2) consumptive impacts of predators were hierarchically nested, i.e. the larger predator consumed both juvenile and adult snails, while the smaller‐bodied predator consumed only juvenile snails; 3) there were no emergent multiple predator effects on snail consumption; and 4) non‐consumptive effects differed from consumptive effects, with only the large predator inducing predator‐avoidance behavior of individuals within either snail ontogenetic class. The smaller predator therefore played a functionally redundant trophic role across the prey classes considered, augmenting and potentially stabilizing trophic regulation of juvenile snails. Meanwhile, the larger predator played a complementary and functionally unique role by both expanding the size‐spectrum of prey trophic regulation and non‐consumptively altering prey behavior. While our study suggests that nestedness of consumptive interactions determined by predator and prey body sizes may allow prediction of the functional redundancy of particular predator species, it also shows that traits beyond predator body size (e.g. habitat domain) may be required to predict potentially cascading non‐consumptive effects. Future studies of multiple predators (and predator biodiversity) should continue to strive towards greater realism by incorporating not only size‐structured prey, but also other aspects of resource and environmental heterogeneity typical of natural ecosystems.     

Ontogenetic variations in the type and size of prey consumed by juvenile coho,Oncorhynchus kisutch,and chinook,O. tshawytscha,salmon

Synopsis Stomach contents of juvenile coho,Oncorhynchus kisutch, and chinook,O. tshawytscha, salmon collected in purse seines off the coast of Washington and Oregon were examined for variations related to predator size. There was a general trend toward increasing consumption of fish with increasing body size, due mainly to the increase in northern anchovy biomass consumed by the larger salmon. Most of the major prey taxa showed significant differences among the size classes examined for both salmon species. There was a direct relationship between predator and prey size for both coho and chinook, but considerable variation was found in prey length consumed within each size class. Prey width did not provide as good a fit as prey length for either species. In general, coho consumed larger fish prey in relation to their body length than chinook but there were substantial differences by month or year of collection.     

Dragonfly predators influence biomass and density of pond snails

Studies in lakes show that fish and crayfish predators play an important role in determining the abundance of freshwater snails. In contrast, there are few studies of snails and their predators in shallow ponds and marshes. Ponds often lack fish and crayfish but have abundant insect populations. Here we present the results of field surveys, laboratory foraging trials, and an outdoor mesocosm experiment, testing the hypothesis that insects are important predators of pulmonate snails. In laboratory foraging trials, conducted with ten species of insects, most insect taxa consumed snails, and larval dragonflies were especially effective predators. The field surveys showed that dragonflies constitute the majority of the insect biomass in fishless ponds. More focused foraging trials evaluated the ability of the dragonflies Anax junius and Pantala hymenaea to prey upon different sizes and species of pulmonate snails (Helisoma trivolvis, Physa acuta, and Stagnicola elodes). Anax junius consumed all three species up to the maximum size tested. Pantala hymenaea consumed snails with a shell height of 3 mm and smaller, but did not kill larger snails. P. acuta were more vulnerable to predators than were H. trivolvis or S. elodes. In the mesocosm experiment, conducted with predator treatments of A. junius, P. hymenaea, and the hemipteran Belostoma flumineum, insect predators had a pronounced negative effect on snail biomass and density. A. junius and B. flumineum reduced biomass and density to a similar degree, and both reduced biomass more than did P. hymenaea. Predators did not have a strong effect on species composition. A model suggested that A. junius and P. hymenaea have the largest effects on snail biomass in the field. Given that both pulmonate snails and dragonfly nymphs are widespread and abundant in marshes and ponds, snail assemblages in these water bodies are likely regulated in large part by odonate predation.     

Feeding selectivity of four species of sympatric stickleback in brackish-water habitats in eastern Canada

Although sympatric populations of sticklebacks occur commonly along the north-eastern coast of North America, very few studies have considered the interspecific relationships within such populations. This paper investigates prey selectivity, feeding behaviour and potential prey availability within sympatric populations from five brackish-water sites in New Brunswick. Differences between the diets of species from different sites were related to differences in the range of prey available. Fish collected from sites with abundant aquatic vegetation consumed greater numbers of prey than those from vegetation-poor sites. The proportions of prey types eaten by both adults and juveniles reflected prey species abundance at the different sites; thus shifts in diet over time were found to be related to changes in the abundance of potential prey and not to shifts in prey preference on the part of the fish. The size of prey consumed was related to mouth size; adult and juvenile Gasterosteus wheatlandi , having the smallest mouths, consumed the smallest prey. Differences in prey size detected in the four species studied were due to differences in the types of prey consumed. Laboratory experiments on feeding behaviour showed that adult Apeltes quadracus and Pungitius pungitius were more efficient at capturing benthic prey than were Gasterosteus aculeatus and G. wheatlandi. Competition for food is thought not to occur in these populations, because of the abundance of potential prey and the morphological constraints on feeding behaviour.     

Parasites and salinity: costly tradeoffs in a threatened species

Parasites and environmental conditions can have direct and indirect effects on individuals. We explore the relationship between salinity and parasites in an endemic New Mexico State threatened fish, the White Sands pupfish (Cyprinodon tularosa). Spatial variation in salinity limits the distribution of the endemic springsnail (Juturnia tularosae) within Salt Creek, a small desert stream. The springsnail is the presumed intermediate host for trematodes that infect the White Sands pupfish, and trematode prevalence and intensity in pupfish are positively associated with the springsnail. Salinity and parasites both have negative impacts on pupfish, but in areas of high salinity, pupfish can effectively escape parasites. Pupfish trematodes were absent from sites lacking snails. At the upstream site, the absence of parasites and lower variance in salinity were correlated with larger pupfish that were in better condition than pupfish at either the middle or lower sites. Springsnails were present in the middle section, an area with moderate salinity, and all pupfish had trematodes (median abundance 847 trematodes/fish). Lipid levels and condition were lowest in fish from the middle site. Additionally, fewer older fish indicated an increased mortality rate. At the lower site, springsnails were absent due to high salinity; pupfish trematode abundance was much lower (six trematodes/fish), and fish condition was intermediate. An additional experiment revealed that snail activity and survival were significantly reduced at high salinities commonly present at the lower site. Although both high salinity and parasites significantly affect pupfish, parasites might be more detrimental.     

Patterns of shell penetration by Chorus giganteus juveniles (Gastropoda: Muricidae) on the mussel Semimytilus algosus

Patterns are described for shell penetration by the sublittoral muricid snail Chorus giganteus during predatory attacks on the mussel Semimytilus algosus. Location, form and size of shell penetrations were observed in relation to the size of the predator. The results suggested that positions of the perforations on the mussel shells were related to size of the attacking snail. Smaller snails perforated areas near the shell ligament and in the central zone of the shell, while larger snails more frequently attacked shell borders, principally on the ventral side. These observations may be related to: (a) changes in the process of manipulation of the prey during development of the foot and the shell tooth of the predator, (b) changes in internal structure of the snails related to the shell perforation mechanism, or (c) learned behavior acquired experientially by the snails during early growth. Although in other studies of muricid penetration patterns larger boreholes made in shells of the prey were positively correlated with increasing predator size, this relation did not appear to hold with C. giganteus, as larger specimens often made relatively small shell perforations. Areas of boreholes made in the mussel shells by this snail varied from 0.01 to 1.1 mm(2), and were unusually variable in size and shape, especially when compared with literature results on bores characteristic of other muricid species.     

Evolution of bite performance in turtles

Abstract Among vertebrates, there is often a tight correlation between variation in cranial morphology and diet. Yet, the relationships between morphological characteristics and feeding performance are usually only inferred from biomechanical models. Here, we empirically test whether differences in body dimensions are correlated with bite performance and trophic ecology for a large number of turtle species. A comparative phylogenetic analysis indicates that turtles with carnivorous and durophagous diets are capable of biting harder than species with other diets. This pattern is consistent with the hypothesis that an evolutionary increase in bite performance has allowed certain turtles to consume harder or larger prey. Changes in carapace length tend to be associated with proportional changes in linear head dimensions (no shape change). However, maximum bite force tends to change in proportion to length cubed, rather than length squared, implying that changes in body size are associated with changes in the design of the jaw apparatus. After the effect of body size is accounted for in the analysis, only changes in head height are significantly correlated with changes in bite force. Additionally, our data suggest that the ability to bite hard might trade off with the ability to feed on fast agile prey. Rather than being the direct result of conflicting biomechanical or physiological demands for force and speed, this trade‐off may be mediated through the constraints imposed by the need to retract the head into the shell for defensive purposes.     

Predation of Japanese dace, Tribolodon hakonensis, by largemouth bass, Micropterus salmoides, in experimental aquaria

 To test the size range of prey fish that largemouth bass, Micropterus salmoides, can successfully consume, live Japanese dace, Tribolodon hakonensis, were given as prey fish to individual largemouth bass in aquaria. The ratio of maximum standard length (SL) of the Japanese dace consumed by largemouth bass was 46–69% of bass SL. The maximum length of Japanese dace consumed did not differ significantly between largemouth bass and smallmouth bass (M. dolomieu) previously studied, although largemouth bass have relatively larger mouth sizes than smallmouth bass. Largemouth bass occasionally injured and killed Japanese dace larger than the limit that could be consumed.     

The effects of predation by the mummichog, Fundulus heteroclitus (L.), on the abundance and distribution of the salt marsh snail, Melampus bidentatus (Say)

Enclosure and exclosure experiments were conducted in Canary Creek marsh to examine how predation by a killifish, Fundulus heteroclitus (L.), affects the abundance and size distribution of the salt marsh snail, Melampus bidentatus (Say). Enclosures (7.6 × 19.7 m) were stocked with Fundulus heteroclitus at densities of one-half normal, normal, twice normal, and four times normal density. Fish exclusion pens were also built. In both years of study, the mean density of snails increased significantly in pens where fish were excluded or their density reduced. During the same period in each year, the density of snails in pens containing higher than normal fish density fell by ≈50%.

Fish density also affected the size distribution of snails within pens. In both years, mean shell length of snails in the pen with the highest density of fish was significantly greater, and mean shell length of snails within fish exclusion pens was significantly lower than in all other treatments. Gape size limitation of F. heteroclitus causes selective predation on small snails and apparently is responsible for the difference in mean shell length among treatments.

Density and size distribution measurements of the natural Melampus bidentatus population in Canary Creek marsh were also taken. Larger snails were found to occur in the low marsh zone, whereas smaller snails occurred in the high marsh zone. Since the high marsh area is flooded less often than the low marsh, and the grass types found in the high marsh zone are known to afford protection from fish predation, the distribution of snails in the marsh is consistent with the idea that fish predation is an important factor influencing the distribution of snails in Canary Creek marsh.     

LABORATORY DIGESTION OF PREY AND INTERPRETATION OF WALRUS STOMACH CONTENTS

A new approach for evaluating the potential biases of walrus ( Odobenus rosmarus ) diet data derived from stomach contents was examined based on the rates at which different prey types were digested. In this study controlled digestion experiments on polychaetes, echiurid and sipunculid worms, clams, snails, and crabs demonstrated that these prey items did not remain equally identifiable during digestion. Polychaetes, echiurids, and sipunculids were the least persistent prey. All worms became unidentifiable during the six-hour digestion trials. Over 50% of the clams maintained their diagnostic tissues ( i. e. , foot and/or siphon). Clam viscera did not survive hour 2. Snails and crustaceans were the most persistent prey. Without a consideration of the state of digestion, a stomach sample may not accurately reflect the species composition and size of prey consumed. Volume is not a reliable measure of a prey's relative importance, as the diagnostic fragments of invertebrate prey in a stomach vary greatly in physical composition. Walruses probably consume all of the soft tissues of clams, not solely the foot and siphon.     

Heterochrony, maternal effects, and phenotypic variation among sympatric pupfishes

Variation in ontogeny can produce phenotypic variation both within and among species. I investigated whether changes in timing and rate of growth were a source of phenotypic variation in a putative incipient species group of pupfish (Cyprinodon spp.). On San Salvador Island, Bahamas, sympatric forms of pupfish differ in morphology but show only partial reproductive isolation in the laboratory. Offspring from two forms and two geographical areas and their hybrids were bred in the laboratory, and ontogenetic trajectories of their feeding morphology were followed until maturity. In the Bahamian pupfish the two forms grow along similar size but not shape trajectories. Two heterochronic parameters, onset and rate of growth, alter shape trajectories in the Bahamian pupfish. Similar forms from different geographical areas (Florida and the Bahamas) grow along parallel shape trajectories, differing only in one heterochronic parameter, the onset shape. Hybrids within and between the pupfish forms produced intermediate feeding morphologies that were influenced by their maternal phenotype, suggesting that maternal effects may be a source of phenotypic variation in shape that can persist to maturity. In Cyprinodon, small changes in multiple heterochronic parameters translate into large phenotypic differences in feeding morphology.     

Killing them softly: Ontogeny of jaw mechanics and stiffness in mollusk-feeding freshwater stingrays

Durophagous predators consume hard-shelled prey such as bivalves, gastropods, and large crustaceans, typically by crushing the mineralized exoskeleton. This is costly from the point of view of the bite forces involved, handling times, and the stresses inflicted on the predator's skeleton. It is not uncommon for durophagous taxa to display an ontogenetic shift from softer to harder prey items, implying that it is relatively difficult for smaller animals to consume shelled prey. Batoid fishes (rays, skates, sawfishes, and guitarfishes) have independently evolved durophagy multiple times, despite the challenges associated with crushing prey harder than their own cartilaginous skeleton. Potamotrygon leopoldi is a durophagous freshwater ray endemic to the Xingu River in Brazil, with a jaw morphology superficially similar to its distant durophagous marine relatives, eagle rays (e.g., Aetomylaeus, Aetobatus). We used second moment of area as a proxy for the ability to resist bending and analyzed the arrangement of the mineralized skeleton of the jaw of P. leopoldi over ontogeny using data from computed tomography (CT) scans. The jaws of P. leopoldi do not resist bending nearly as well as other durophagous elasmobranchs, and the jaws are stiffest nearest the joints rather than beneath the dentition. While second moment has similar material distribution over ontogeny, mineralization of the jaws under the teeth increases with age. Neonate rays have low jaw stiffness and poor mineralization, suggesting that P. leopoldi may not feed on hard-shelled prey early in life. These differences in the shape, stiffness and mineralization of the jaws of P. leopoldi compared to its durophagous relatives show there are several solutions to the problem of crushing shelled prey with a compliant skeleton.     

The potential for intracohort cannibalism in age-0 walleye pollock,Theragra chalcogramma,as determined under laboratory conditions

Synopsis In laboratory experiments, we tested the capability of larger age-0 walleye pollock to consume smaller members of their cohort. In separate aquaria, 81 pairs of juveniles covering a wide range of size differences (total lengths differing by 12 to 61 mm) were held and monitored over a 4 day period. Complete consumption, in which a smaller fish was swallowed whole by a larger fish, occurred 11% of the time. In 36% of the pairs, attacks by the larger fish resulted in mortality of the smaller fish. The mouth width:body depth ratio between the larger and smaller fish of a pair differed significantly depending on whether the smaller fish survived, was killed but not consumed, or was ingested whole by the larger fish. Cannibalistic individuals could consume fish close to the maximum size physically possible under gape limitation; at this size the length of the cannibal was approximately 1.7 times the length of the prey. Length-frequency distributions of age-0 pollock in field concentrations suggested that, at least in some geographical areas, potential cannibals and prey commonly co-occur. Unsuccessful predatory attacks by larger individuals may have additional detrimental effects on smaller pollock in natural populations.