Optimal foraging as the criteria of prey selection by two centrarchid fishes

The nature of prey selection by two centrarchids (white crappie and bluegill) is presented as a model incorporating optimal foraging strategies. The visual field of the foraging fish as represented by the reactive distance is analysed in detail to estimate the number of prey encounters per search bout. The predicted reactive distances are compared with experimental data. The energetic cost associated with fish foraging behaviour is calculated based on the sequence of events that takes place for each prey consumed. Comparisons of the relative abundance of prey species and size categories in the stomach to the lake environment indicated that both white crappie and bluegill (length < 100 mm) strongly select prey utilising an energy optimization strategy. In most cases, the fish exclusively selected large Daphnia ignoring evasive prey types (Cyclops, Diaptomids) and small cladocera. This selectivity is the result of fish actively avoiding prey with high evasion capabilities even though they appear to be high in energetic content and having translated this into optimal selectivity through capture success rates. The energy consideration and visual system, apart from the forager's ability to capture prey, are the major determinants of prey selectivity for large-sized bluegill and white crappie still at planktivorous stages.     

Mechanisms of prey choice by planktivorous fish: perceptual constraints and rules of thumb

Visually foraging planktivorous fish feed preferentially on larger, energetically more valuable zooplankton. Laboratory studies of bluegill sunfish, Lepomis macrochirus, feeding on different size classes of Daphnia indicate that, at low prey densities, these fish generally select among prey encountered simultaneously by pursuing whichever one appears largest, i.e. projects the largest retinal image. At high prey densities, however, bluegills take fewer small Daphnia than predicted for fish foraging by this apparent size rule. The pattern of prey selection observed suggests that bluegills employ the apparent size rule, not as the sole determinant of prey choice, but only to direct their attention to an individual prey item. In this way, fish may consider prey encountered simultaneously in a sequential manner. The fish can then evaluate the actual size of the apparently largest prey item, and ignore it if it is below a minimum threshold actual size. The less-preferred small prey, however, are never completely excluded from the diets of the fish. This partial preference may be the result of perceptual constraints; fish may be able to evaluate the actual size of zooplankton prey only when they occur within the fish's binocular visual field.     

The effects of perceived competition and parasitism on the foraging behaviour of the upland bully (Eleotridae)

Prey choice by fish is subject to many constraints, some of which may interact to determine the relative preference of fish for prey with different profitabilities. The constraining effects of parasitism and perceived competition on foraging behaviour were examined in the upland bully, Gobiomorphus breviceps . In the laboratory, bullies faced with a choice of prey items of two different sizes chose the larger prey significantly more often than the smaller ones. The presence of a conspecific fish near the source of large prey significantly reduced the bullies'preference for larger prey. Neither the size of the test fish, nor the number of digenean metacercarial cysts they harboured, had any influence on their relative preference for larger prey, or on how that preference was dampened by the presence of a competitor. The threat of competition appears to be a more important constraint on prey choice in upland bullies than parasitism.     

An experimental study of body size and food size relations in crucian carp,Carassius carassius

Synopsis In dense, single-species assemblages of crucian carp, competition is intense and results in populations of stunted fish. To explore mechanisms underlying this competition, we measured handling times, return rates, and prey choice for five sizes of crucian carp feeding on six sizes of a standardized food. Handling times increased with prey size and decreased with fish size. Return rates (dry mass ingested per unit handling time) increased dramatically with fish size, and generally decreased with increasing prey size, especially for small fish. Patterns of return rates among size-classes suggested that one or more size-related shifts in feeding efficiency exist for crucian carp; combined with physiological stresses related to winter anoxia, the inability of fish to make these shifts may contribute to size structures observed in high-density populations. Comparisons of relations among fish size, prey size, and return rates for crucian carp and bluegill, Lepomis macrochirus, suggest that similar intraspecific competitive relations exist between generalist species with size-structured populations. Despite differences in return rates among prey sizes, the extent of food selectivity based on prey size exhibited by crucian carp in two types of choice trials was less than predicted. Crucian carp commonly take in several items before mechanically processing food with their pharyngeal apparatus; this multiple prey processing may contribute to disparities between observed choice patterns and those predicted based on return rates for single prey.     

Virtual plankton: A novel approach to the investigation of aquatic predator‐prey interactions

Small‐scale zooplankton swimming behaviors can affect aquatic predator‐prey interactions. Difficulties in controlling prey swimming behavior however, have restricted the ability to test hypotheses relating differences in small‐scale swimming behavior to frequency of predation by fish. We report here a Virtual Plankton (VP) system that circumvents this problem by allowing the observation of fish “preying"on computer‐generated prey images whose size, shape, color and swimming behavior can be precisely controlled. Two experiments were performed in which bluegill sunfish (Lepomis macrochirus) were given a choice of either two VP images, one of which moved twice as fast as the other, or six VP, one of which moved either faster (1.25 x, 1.5 x or 2 x ) or slower (0.5 x) than the other five. Current predator‐prey models based on encounter probabilities and prey visibility predict that moving faster increases predation risk and conversely, moving slower decreases predation risk. In agreement with existing predator‐prey models, in both experiments, fish chose faster moving VP significantly more often than their slower moving neighbors. Contrary to the predictions of existing models, in the second experiment with six VP, the rate at which fish chose a prey image moving half as fast as the five surrounding images did not differ significantly from the rate predicted by chance(l/6). These results suggest that current fish‐zooplankton predation models would benefit by the incorporation of small‐scale swimming behavior and assessments of its influence on overall prey visibility.     

The effect of shoal size on patterns of body colour segregation in mollies

Individual white and black mollies Poecilia latipinna spent significantly more time near the larger group when given the choice between two shoals of similar colouration to themselves. When given the choice between a large and a small shoal of dissimilar colouration to themselves, black test fish spent significantly more time with the larger shoal while white test fish showed no preference for either group. Both white and black mollies chose the smaller of two shoals when given the choice between a large dissimilarly coloured shoal and a small similarly coloured shoal. The results indicate that mollies actively discriminate between shoals on the basis of both body colour and shoal size. However, body colour segregation appears to have a stronger influence on shoal choice.     

Host specificity among Ancyrocephalinae (Monogenoidea) of Nebraska sunfish

This study addressed the problem of local patterns of host specificity among Ancyrocephalinae (Monogenoidea) on bass and sunfish species, when the hosts occur in different species combinations in separate ponds. One hundred fifty-three fish of the Centrarchidae, from 4 study sites in Nebraska, were collected. Host species included bluegill (Lepomis macrochirus), green sunfish (L. cyanellus), largemouth bass (Micropterus salmoides), black crappie (Pomoxis nigromaculatus), white crappie (P. annularis), and rock bass (Ambloplites rupestris). These fish occurred in different species combinations, depending on the pond sampled. Results indicated that several centrarchid species could inhabit the same pond and yet support distinct monogene communities. Clavunculus bursatus, Onchocleidus helicis, O. principalis, and Syncleithrum fusiformis were found only on largemouth bass, regardless of what other centrarchids were present in a particular pond. Haplocleidus dispar occurred on green sunfish, bluegill, largemouth bass, and black crappie, and H. furcatus occurred on both bluegill and largemouth bass. Onchocleidus cyanellus and O. ferox were found on both bluegill and green sunfish. Rock bass were present in only 1 of the 4 ponds, but were not infected with any monogenes, even though co-occurring centrarchids were often heavily infected. Largemouth bass had the most diverse ancyrocephaline communities. The degree of parasite host specificity among these monogenes was inversely related to the diversity of host species present in a particular pond. In general, the parasites were more host specific than might be inferred from the literature; parasite species did not necessarily colonize supposedly receptive host species even when the latter were present, and host relatedness was the major factor in determining whether host species shared a common parasite species.     

Prey resources before spawning influence gonadal investment of female, but not male, white crappie

In this study, an outdoor pool experiment was used to evaluate the effect of prey resources during 4 months before spawning on the gonadal investments of male and female white crappie Pomoxis annularis , a popular freshwater sportfish that exhibits erratic recruitment. Fish were assigned one of three feeding treatments: starved, fed once every 5 days (intermediate) or fed daily (high). All measurements of male testes ( i.e. wet mass, energy density and spermatocrit) were similar across treatments. Conversely, high-fed females produced larger ovaries than those of intermediate-fed and starved fish, and invested more energy in their ovaries than starved fish. Compared to pre-experiment fish, starved and intermediate-fed females appeared to increase their ovary size by relying on liver energy stores ('capital' spawning). Conversely, high-fed females increased liver and gonad mass, implying an 'income'-spawning strategy (where gonads are built from recently acquired energy). Fecundity did not differ among treatments, but high-fed fish built larger eggs than those starved. Females rarely 'skipped' spawning opportunities when prey resources were low, as only 8% of starved females and 8% of intermediate-fed females lacked vitellogenic eggs. These results suggest that limited prey resources during the months before spawning can limit ovary production, which, in turn, can limit reproductive success of white crappies.     

The ontogeny of search behavior in the white crappie,Pomoxis annularis

Synopsis Animals that forage for discrete, isolated resources are often characterized as either ambush (sit-and-wait) or cruise (active) searchers. Juvenile white crappie, Pomoxis annularis, search for zooplankton prey using a saltatory search (SS) strategy. Unlike ambush and cruise search, SS involves scanning for prey only during the brief stationary periods that punctuate repositioning movements. If prey are not found, these fish swim a short distance, stop, and scan again. In this paper, we describe the ontogeny of prey search in the white crappie and compare the search pattern that they employ with that of juveniles. White crappie larvae searched for prey throughout the search space and only during the pauses that punctuated swimming movements. Prey location distances increased with fish size, as did several other components of the predation cycle. We conclude that white crappie larvae employ a search strategy similar to that exhibited by juveniles. We emphasize that, to obtain an accurate assessment of the feeding ecology of early life history stages, the search pattern that they employ must be characterized, and its components quantified.     

Intraspecific phenotypic variation among alewife populations drives parallel phenotypic shifts in bluegill

Evolutionary diversification within consumer species may generate selection on local ecological communities, affecting prey community structure. However, the extent to which this niche construction can propagate across food webs and shape trait variation in competing species is unknown. Here, we tested whether niche construction by different life-history variants of the planktivorous fish alewife (Alosa pseudoharengus) can drive phenotypic divergence and resource use in the competing species bluegill (Lepomis macrochirus). Using a combination of common garden experiments and a comparative field study, we found that bluegill from landlocked alewife lakes grew relatively better when fed small than large zooplankton, had gill rakers better adapted for feeding on small-bodied prey and selected smaller zooplankton compared with bluegill from lakes with anadromous or no alewife. Observed shifts in bluegill foraging traits in lakes with landlocked alewife parallel those in alewife, suggesting interspecific competition leading to parallel phenotypic changes rather than to divergence (which is commonly predicted). Our findings suggest that species may be locally adapted to prey communities structured by different life-history variants of a competing dominant species.     

Diel feeding periodicity, daily ration and prey selection of a riverine population of juvenile chinook salmon, Oncorhynchus tshawytscha (Walbaum)

The diel feeding periodicity, daily ration and prey selection of juvenile chinook salmon, Oncorhynchus tshawytscha , were studied in relation to the available prey. Maximum dry weight of food intake occurred about dawn, when mayflies were the major prey, but the greatest number of freshly eaten prey occurred during the afternoon, when chironomids and terrestrial dipterans predominated. Feeding activity at night was low, with smaller mayflies comprising up to 50% of the prey. During the day the young salmon fed selectively on chironomids and the larger mayflies, while trichopterans and terrestrial taxa were under-represented in the diet. Food consumption over the 24-h period averaged 8.3% of the fish dry body weight. Prey abundance in the drift explained about 50% of the composition of the diet. Although the fish selected larger mayflies, size apparently was not a main criterion for selection because chironomids, although smaller than mayflies, were also frequently eaten. Previous dietary experience of the fish and the diel pattern of prey abundance appear to best explain the selective feeding of juvenile chinook salmon.     

Utilization of the exotic cladoceran Daphnia lumholtzi by juvenile fishes in an Illinois River floodplain lake

Daphnia lumholtzi comprises a substantial component of the zooplankton community during mid‐ to late‐summer in Lake Chautauqua, a floodplain lake along the Illinois River near Havana, Illinois. In order to quantify the utilization of D. lumholtzi by juvenile fishes, diet analyses were conducted for seven juvenile fish species collected from Lake Chautauqua during the 2001 annual drawdown period. Freshwater drum Aplodinotus grunniens and emerald shiner Notropis atherinoides demonstrated negative selectivity for D. lumholtzi relative to native zooplankton species whereas four species of fish (bluegill Lepomis macrochirus, white bass Morone chrysops, white crappie Pomoxis annularis and black crappie Pomoxis nigromaculatus) consumed substantial amounts of D. lumholtzi. Although selectivity values for D. lumholtzi varied among these fish species, positive selection for D. lumholtzi increased similarly among larger size classes of each fish species, and corresponded with ontogenetic shifts in diet. Mean body length of D. lumholtzi consumed by 20–69 mm L_T juvenile fishes ranged from 0·75 to 0·99 mm with a calculated total length range of 2·0–2·6 mm. Results from this study provide evidence that high abundances of D. lumholtzi in mid‐ to late‐summer provide an additional food source for several juvenile fish species during a time when abundances of large native cladoceran species (i.e. Daphnia) are low, and juvenile fishes are searching for larger prey associated with ontogenetic shifts from zooplankton to macroinvertebrates and fishes. Because zooplankton production is typically lower in rivers than in lakes, survivorship of juvenile fishes produced in floodplain lakes may be higher in riverine systems if they are not reliant on zooplankton as a primary food resource. Therefore, high abundances of D. lumholtzi may benefit juvenile fishes in managed floodplain lakes, such as Lake Chautauqua, by increasing growth and facilitating the transition from zooplanktivory to insectivory or piscivory.     

The Ability of Three Species of Tadpoles to Differentiate among Potential Fish Predators

The ability of prey to respond to novel predator cues may depend on the generality or specificity of the response to predator cues. We used laboratory behavioral experiments to examine the ability of tadpoles of three species of anurans (American toad, Bufo americanus ; bullfrog, Rana catesbeiana ; and green frog, R. clamitans ) to respond to the presence of two native potential predators (bluegill, Lepomis macrochirus ; and largemouth bass, Micropterus salmoides ) and one non-native potential predator (goldfish, Carassius auratus ). We also examined the effect of tadpole size on the behavioral responses of American toads and green frogs to predator cues. All three species of tadpoles responded to the presence of predator cues, although the specific responses varied among species. American toads and green frogs reduced activity in the presence of at least some fish cues, but bullfrog tadpoles did not change their activity. Bullfrogs decreased use of vegetation in the presence of some predator cues, whereas American toads and green frogs did not. American toads only responded to the presence of bluegill cues but not the other fish predator cues, whereas bullfrogs and green frogs responded more generally to the fish predators. In both American toads and green frogs, tadpole size affected behavior. For American toads, activity increased, as did the use of the vegetated side of the aquarium, in larger tadpoles. Not only did size affect American toad behavior, but it also influenced the responses of the tadpoles to predator cues. For green frogs, activity decreased in larger tadpoles. Our results suggest that behavioral responses of tadpoles to predator cues can be influenced by both the identity of the predator and the prey, as well as the size of the potential prey.     

Do juvenile gape-limited predators compensate for their small size when feeding?

When juvenile and adult animals occur syntopically, juveniles are at a distinct performance disadvantage due to their absolutely small size. Yet, optimal foraging theory predicts that juvenile predators should feed efficiently in order to compete with adults for food, and to minimize their exposure to predators. Previous authors have suggested that one way for juvenile animals to accomplish these ecological tasks is by increasing their overall feeding performance relative to adults (compensation hypothesis). Nonetheless, only a handful of studies have tested whether juvenile animals have increased feeding performance (e.g. decreased ingestion and/or handling times relative to body size) compared with adults. We tested this hypothesis by examining the ontogeny of head dimensions and feeding performance (ingestion time and number of mandibular protractions) on fish prey for broad-banded water snakes Nerodia fasciata . Individuals were fed fish scaled in a 1:1 ratio to their head width. All head dimensions scaled with significant negative allometry versus body size, and thus smaller snakes had relatively larger heads for their body size compared with larger snakes. By contrast, most head variables (except head volume) exhibited positive allometry versus head length, demonstrating that larger snakes had larger head dimensions relative to head size compared with smaller snakes. In the performance trials, smaller snakes had worse feeding performances when feeding on similarly sized fish prey (relative to their head width) compared with larger snakes. Therefore, these data show that smaller water snakes do not compensate for their size through increased feeding performance.     

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.     

Evidence for leptin expression in fishes

Tissues from bony fish were screened with anti-mouse leptin antibodies to detect the presence of the fat-regulating hormone in fishes. Low molecular-weight (16 kDa) immunoreactive bands were detected in blood, brain, heart and liver of green sunfish (Lepomis cyanellus), bluegill sunfish (Lepomis macrochirus), largemouth bass (Micropterus salmoides), white crappie (Pomonix annularis), channel catfish (Ictalurus punctatus), and rainbow trout (Oncorhynchus mykiss). To further verify that we had identified leptin, the response of fish "leptin" was measured in fed and fasted green sunfish. Fed sunfish had approximately threefold higher concentration of leptin in blood than did fasted sunfish (fed vs. fasted; 0.599 +/- 0.03 microg/microl vs. 0.196 +/- 0.04 microg/microl; P > F = 0.0001), which is consistent with mammalian models of leptin function. Brain leptin concentration is also positively correlated with percent body fat in white crappie and bluegill. Based upon electrophoretic mobility, immunoreactivity, response to fasting, and correlation with adiposity, we believe we have the first evidence for leptin expression in an ectotherm.     

Predator-prey interactions in size-structured fish communities: implications of prey growth

Predator-prey interactions among size-structured populations may be strongly influenced by factors which affect growth rates of prey. I examined the importance of prey growth in the interaction between large-mouth bass (Micropterus salmoides) and their prey, bluegill (Lepomis macrochirus), by analyzing diets and growth rates of bass in a set of seven lakes in south-central Wisconsin. Sizes of bluegill consumed by bass changed dramatically across a gradient of bluegill growth, which resulted in differing patterns of bass growth. In lakes with slow bluegill growth, small bass fed on the youngest bluegill cohort, but large bass were capable of feeding on several age classes. Consequently, bass growth rates were strongly size-dependent; small bass ate small prey and had low growth, but growth rates increased substantially with size as bass ate progressively larger prey. When bluegill had high growth rates, they quickly reached a size refuge from predation and bass of all sizes were restricted to feeding on the youngest/smallest prey. In these lakes, bass growth rates were more uniform across bass sizes. Because growth rates influence population size-distributions, variation in bluegill growth can have strong effects on the structure of bass populations. These effects could potentially feed back to further influence the interaction between predator and prey.     

Do bovids evolve hindquarter markings for anti-predation?

Conspicuous coloration in animals serves many functions such as anti-predation. Anti-predation strategies include motion dazzle and flash behavior. Motion dazzle markings can reduce the probability of being preyed on because the predators misjudge their movement. In flash behavior, prey demonstrate conspicuous cue while fleeing; the predators follow them; however, the prey hide their markings and the predators assume that the prey has vanished. To investigate whether bovids use conspicuous hindquarter markings as an anti-predatory behavior, we undertook phylogenetically controlled analyses to explore under what physiological characteristics and environmental factors bovids might have this color pattern. The results suggested that rump patches and tail markings were more prevalent in bovids living in larger-sized groups, which supports the hypothesis of intraspecific communication. Moreover, we observed the occurrence of conspicuous white hindquarter markings in bovids having smaller body size and living in larger groups, suggesting a motion dazzle function. However, the feature of facultative exposing color patterns (flash markings) was not associated with body size, which was inconsistent with predictions and implied that bovids may not adopt this as an anti-predator strategy. It was concluded that species in bovids with conspicuous white hindquarter markings adopt motion dazzle as an anti-predation strategy while fleeing and escaping from being prey on.     

Population structure and food habits of white crappie Pomoxis annularis Rafinesque in a turbid Oklahoma reservoir

We assessed population structure and food habits of white crappie Pomoxis annuluris Rafinesque in Lake Carl Blackwell, a turbid reservoir in north-central Oklahoma, U.S.A. White crappie [ n = 8549; total length ( t.l. ) mean = 147 mm, s.d. = 28 mm, range = 73–445 mm] were collected with gillnets, frame nets, and hoop nets. Proportional stock density was 2, as 97% of the fish were under 200 mm t.l . Relative stock density was 96, 2, 1, and 1 for 'stock-quality', 'quality-preferred', 'preferred-memorable', and 'memorable-trophy' lengths. Total annual mortality rate was 54% for fish older than 2 years. Growth was poor and lower than the Oklahoma-Arkansas regional averages ( P <0.01). Relative weights were 79, 86, 106, 109, and 123 for fish in the stock-quality, quality-preferred, preferred-memorable, memorable-trophy, and trophy size categories. Ephemeroptera nymphs were an important dietary component for fish 150 mm t.l. or smaller. Gizzard shad Dorosoma cepedianum (Lesueur) was a major forage item, especially for fish greater than 150 mm t.l. ( P <0.01). Empirical information on prey availability indicates that the white crappie population is forage limited and that gizzard shad are not available in appropriate sizes or sufficient numbers to sustain the predator population. Poor fish condition, slow growth, and high mortality are probably the result of inadequate forage.     

Prey size selection and cannibalistic behaviour of juvenile barramundi Lates calcarifer

This study assessed the cannibalistic behaviour of juvenile barramundi Lates calcarifer and examined the relationship between prey size selection and energy gain of cannibals. Prey handling time and capture success by cannibals were used to estimate the ratio of energy gain to energy cost in prey selection. Cannibals selected smaller prey despite its capability of ingesting larger prey individuals. In behavioural analysis, prey handling time significantly increased with prey size, but it was not significantly affected by cannibal size. Conversely, capture success significantly decreased with the increase of both prey and cannibal sizes. The profitability indices showed that the smaller prey provides the most energy return for cannibals of all size classes. These results indicate that L. calcarifer cannibals select smaller prey for more profitable return. The behavioural analysis, however, indicates that L. calcarifer cannibals attack prey of all size at a similar rate but ingest smaller prey more often, suggesting that prey size selection is passively orientated rather than at the predator's choice. The increase of prey escape ability and morphological constraint contribute to the reduction of intracohort cannibalism as fish grow larger. This study contributes to the understanding of intracohort cannibalism and development of strategies to reduce fish cannibalistic mortalities.