A computer‐activated food delivery system for dispensing natural prey items among complex arrays of patches

Experimental studies of the foraging ecology of drift‐feeding fishes have been limited in scope because of the lack of systems for dispensing suitable prey items to a number of patches with complex and controlled spatio‐temporal patterns of delivery rate. Here, tests are reported of a computer‐controlled feeding system that performs this requirement with considerable flexibility.     

Prey selection by juvenile cyprinids from running water

SUMMARY. 1. The juveniles of dace ( Leuciscus leuciscus (L.)) and roach ( Rtuihts rtuihis (L.)) and yearling minnows ( Phoxtnus phoxinus (L.)) exhibited generally similar patterns of feeding behaviour and preferences for particular prey types.
2. Factors affecting prey selection included predator and prey size, prey activity and environmental conditions.
3. All three species preferred prey sizes approximately 0.6x their maximum gape.
4. Prey motion was essential to induce attack, and prey were usually selected in proportion to their relative activities.
5. Specialization on one prey type was observed in still water, and was always directed at the most active prey type. In flowing water no specialization was observed, and size selection was suppressed in the juvenile fish. Yearling minnows, however, were size selective in both still and flowing water.
6. The presence of macrophyte cover did not significantly change the pattern of prey selection, but did reduce the predation rale.
7. These results are discussed in the context of previous studies of prey selection and optimal foraging.     

Balancing predation risk,social interference,and foraging opportunities in backswimmers,<Emphasis Type="Italic"> Notonecta maculata</Emphasis>

Loss of foraging opportunities and intraspecific competition for prey may be important costs of using refuges, because a hiding animal is unable to use or defend its foraging area from conspecific intrusions. Thus, animals should balance antipredator demands with other requirements in deciding when to come out from a refuge after a predators unsuccessful attack. Observations on foraging and social interactions of backswimmers Notonecta maculata suggest that foraging may be costly in terms of intraspecific agonistic interactions. When prey density is low, increasing the probability of finding a prey may require active exploration of a larger area, but this also increases the probability of encountering a competitor. After simulated exposure to predators, unfed bugs resumed feeding positions after a significantly shorter hiding period than recently fed bugs. We hypothesized that hiding time may also be reduced by recent interactions with conspecific competitors, due to an increased perceived need to defend feeding opportunities. Thus, when a predator attack occurred immediately after an agonistic conspecific interaction, backswimmers resumed feeding positions more quickly, and closer to the original position from which they were disturbed, suggesting short-term defense of particular positions. We conclude that when foraging, backswimmers balance the benefits of finding prey with the costs of predation risk and social interference in deciding their foraging strategy.Communicated by P.K. McGregor     

Environmental enrichment and prior experience of live prey improve foraging behaviour in hatchery‐reared Atlantic salmon

Atlantic salmon salmo salar L. parr were reared for 3 months under standard hatchery conditions or in a structurally enriched tank (containing plants, rocks and novel objects). Half of each of these fish had prior exposure to live prey in the form of live bloodworm while the other half were fed hatchery‐pellets. After 12 days all fish were tested on a novel live prey item (brine shrimp). A significant interaction between the two factors (prior exposure to live prey and rearing condition) revealed that foraging performance was only enhanced in fish that had been reared in a complex environment and exposed to live prey. It appears that the ability to generalize from one live prey type to another is only enhanced in fish that had been reared in an enriched environment. The findings support the assertion that the provision of enriched environments in combination with exposure to live prey prior to release may significantly improve the post‐release survival rates of hatchery‐reared fishes. As both the environmental enrichment and the prior foraging experience procedures were comparatively simple, the provision of such pre‐release experiences are likely to prove cost effective to hatcheries.     

Foraging capacities and effects of competitive release on ontogenetic diet shift in bream, Abramis brama

Bream (Abramis brama) undergo ontogenetic diet shift from zooplankton to benthic macroinvertebrates, but the switching size may be highly variable. To unravel under what conditions bream are pelagic versus benthic foragers, we experimentally determined size‐dependent foraging capacities on three prey types from the planktivory and benthivory niche; zooplankton, visible and buried macroinvertebrates. From these data we derived predictions of size‐dependent diet preferences from estimates of prey value and competitive ability, and tested these predictions on diet data from the field. Planktivorous foraging capacity described a hump‐shaped relationship with bream length that peaked for small bream of 67 mm total length. Benthivory capacity increased with increasing bream size, irrespective if benthic prey were visible on the sediment surface or buried in the sediment. From the experimental data and relationships of metabolic demand we calculated minimum resource requirement for maintenance (MRR) for each of the prey categories used in experiments. MRR increased with bream size for both zooplankton and visible chironomids, but decreased with bream size for buried chironomids, suggesting that intermediate sized bream (120–300 mm) may be competitively sandwiched between small and large bream that are more competitive planktivores and benthivores, respectively. Prey value estimates and competitive abilities qualitatively predicted diet shift in a bream population being released from competition. Competitive release did not change the diet of the largest size‐class feeding on an optimal diet of benthic invertebrates both before and after competitive release. However, profound diet shifts towards benthic macroinvertebrates were recorded for intermediate size‐classes that fed on a suboptimal diet prior to competitive release. Thus, laboratory estimates of size‐dependent foraging capacity of bream in planktivorous and benthivorous feeding niches provided useful information on size‐specific competitive ability, and successfully predicted diet preference in the field.     

Variability in diet and foraging behaviour between sexes and ploidy forms of the hybridogenetic Squalius alburnoides complex (Cyprinidae) in the Guadiana River basin, Portugal

Samples of the Squalius alburnoides complex, a hybridogenetic Iberian minnow composed of diploid, triploid and, less commonly, tetraploid forms, were collected from a tributary of the Ardila River (Guadiana River basin) between February 1999 and January 2001. Seasonal variation in diet was evident, and was probably linked to prey availability. Distinct foraging behaviours between ploidy forms were found towards several prey items, suggesting that diploid adult males fed mostly near the surface, whereas diploid adult females tended to feed near the bottom and submerged vegetation. Triploid females exhibited an intermediate foraging behaviour, although there was greater affinity towards diploid male feeding behaviour. Diploid males which, in contrast to diploid and triploid females, have non‐hybrid genomes in the Guadiana drainage, exhibited a higher specialization for food. Despite considerable dietary overlap, there appeared to be spatial segregation of feeding niches between the three forms, especially during dry periods when prey availability was lower, which may be a strategy for diminishing competition for food. Therefore, considering asexual generalist and specialist hypotheses, it appears that the different ploidy levels are generalist, opportunistic feeders that partition the resources when limited.     

The effects of turbidity and an invasive species on foraging success of rosyside dace (Clinostomus funduloides)

1.  Habitat degradation and biological invasions are important threats to fish diversity worldwide. We experimentally examined the effects of turbidity, velocity and intra- and interspecific competition on prey capture location, reactive distance and prey capture success of native rosyside dace ( Clinostomus funduloides ) and invasive yellowfin shiners ( Notropis lutipinnis ) in Coweeta Creek, North Carolina, U.S.A.
2.  Increased turbidity and velocity produced significant decreases in the number of prey captured forward of the fish's location. It is possible that this represents an increase in the amount of energy expended per prey captured.
3.  We used Akaike's Information Criterion (AIC) to evaluate competing explanatory models for reactive distance (10 generalised linear models, GLM) and prey capture success (9 generalised linear mixed models, GLMM).
4.  Reactive distance decreased by 12% with an increase from 2 to 4 conspecifics, whereas a 10 NTU increase in turbidity reduced reactive distance by 9%. Capture success was affected by velocity, dominance and competition, and varied among species. A 6 cm s⁻¹ increase in velocity produced a 28% decline in capture probability; however, dominant fish were 3.2 times more likely to capture a prey item than non-dominant fish. Yellowfin shiners only were 0.62 times as likely to capture a prey item as rosyside dace. Both intra- and interspecific competition reduced capture probability, and fish in high density intraspecific or interspecific trials were 0.46 times and 0.44 times as likely to capture prey, respectively, as fish in two fish intraspecific trials.
5.  These results suggest behavioural variables are as important as physical factors in determining reactive distance and capture probability by these minnows.     

Age-specific differences in the winter foraging strategies of rooks Corvus frugilegus

Summary Foraging efficiency and intraspecific competition were compared between wild adult and immature rooks Corvus frugilegus with respect to flock size. Behavioural time budgets, and observations of prey selection and prey energetic values revealed that adult rooks in large flocks (> 50 individuals) consumed smaller, less profitable prey, but allocated more time to feeding and fed at a faster rate and with greater success than adults in small flocks. By contrast, immature rooks in flocks of more than 30 individuals allocated proportionally less time to feeding, fed at a lower rate and fed with no increase in success rate than when foraging in smaller flocks. Agonistic encounters and the avoidance of adults by immature rooks appeared responsible for such inefficient foraging. Hence immature rooks showed a preference for smaller flocks (< 50 individuals) with low adult: immature ratios while adults preferred larger flocks (> 50 individuals). We discuss the possible influence of competitive disadvantages on immature rook distribution, flock composition and post-natal dispersal.     

Morphological and trophic divergence of lake and stream minnows (Phoxinus phoxinus)

Phenotypic divergence in response to divergent natural selection between environments is a common phenomenon in species of freshwater fishes. Intraspecific differentiation is often pronounced between individuals inhabiting lakes versus stream habitats. The different hydrodynamic regimes in the contrasting habitats may promote a variation of body shape, but this could be intertwined with morphological adaptations to a specific foraging mode. Herein, I studied the divergence pattern of the European minnow (Phoxinus phoxinus), a common freshwater fish that has received little attention despite its large distribution. In many Scandinavian mountain lakes, European minnows are considered as being invasive and were found to pose threats to the native fish populations due to resource competition. Minnows were recently found to show phenotypic adaptations in lake versus stream habitats, but the question remained if this divergence pattern is related to differences in resource use. I therefore studied the patterns of minnow divergence in morphology (i.e., using geometric morphometrics) and trophic niches (i.e., using stomach content analyses) in the lake Ånnsjön and its tributaries to link the changes in body morphology to the feeding on specific resources. Lake minnows showed a strong reliance on benthic Cladocera and a more streamlined body shape with a more upward facing snout, whereas stream minnows fed on macroinvertebrates (larvae and adults) to a higher degree and had a deeper body with a snout that was pointed down. Correlations showed a significant relationship of the proportion of macroinvertebrates in the gut and morphological features present in the stream minnows. The results of this study highlight the habitat‐specific divergence pattern in morphology and resource use in this ubiquitous freshwater fish. Consequently, interspecific interactions of invasive minnows and the native fish population could differ in the respective food webs and resource competition could target different native fish species in the contrasting habitats.     

Foraging in corallivorous butterflyfish varies with wave exposure

Understanding the foraging patterns of reef fishes is crucial for determining patterns of resource use and the sensitivity of species to environmental change. While changes in prey availability and interspecific competition have been linked to patterns of prey selection, body condition, and survival in coral reef fishes, rarely has the influence of abiotic environmental conditions on foraging been considered. We used underwater digital video to explore how prey availability and wave exposure influence the behavioural time budgets and prey selectivity of four species of obligate coral-feeding butterflyfishes. All four species displayed high selectivity towards live hard corals, both in terms of time invested and frequency of searching and feeding events. However, our novel analysis revealed that such selectivity was sensitive to wave exposure in some species, despite there being no significant differences in the availability of each prey category across exposures. In most cases, these obligate corallivores increased their selectivity towards their most favoured prey types at sites of high wave exposure. This suggests there are costs to foraging under different wave environments that can shape the foraging patterns of butterflyfishes in concert with other conditions such as prey availability, interspecific competition, and territoriality. Given that energy acquisition is crucial to the survival and fitness of fishes, we highlight how such environmental forcing of foraging behaviour may influence the ecological response of species to the ubiquitous and highly variable wave climates of shallow coral reefs.     

Disentangling interference competition from exploitative competition in a crab–bivalve system using a novel experimental approach

In predator–prey relationships such as those between crabs and their bivalve prey, interference competition is a topic of intense investigation as it can have profound consequences on the dynamics of both predator and prey populations. However in laboratory experiments – also those on crab–bivalve systems – workers never adequately disentangled interference competition from exploitative competition, as prey depletion was never compensated. Hitherto, experimental studies on crab–bivalve systems lack direct behavioural observations and have provided only indirect and thus inconclusive evidence of interference competition. We studied interference competition in adult male shore crabs Carcinus maenas that foraged on blue mussels Mytilus edulis. We developed a novel type of experimental tank to replenish each consumed mussel, and thus to keep prey levels constant. We conducted two experiments in which we varied number of crabs (1, 2, 4) and number of mussels (first experiment: 4, 8, 16, 32; second experiment: 8, 32, 128) and directly observed the foraging behaviour of crabs (foraging area=0.25 m²). In the first experiment, feeding rates decreased with increasing crab density only at mussel density 16 because both search time and time spent in agonistic interactions increased. At other mussel densities, variation in crab density did not affect feeding rates, possibly because of low statistical power and the narrow range of mussel densities offered. In the second experiment feeding rates decreased with increasing crab density because crabs spent more time in agonistic interactions and handling their prey. Feeding rates increased with increasing mussel density. Overall, crabs spent on average 14–18% of their foraging time in agonistic behaviours, while on three out of 64 occasions feeding rates decreased because mussels were stolen (kleptoparasitism). Concluding, we have shown that interference competition occurs in absence of prey depletion, while conducting direct behavioural observations aid to identify the behavioural processes that underlie interference competition.     

Optimal foraging and predator-prey dynamics, II.

In this paper we consider one-predator-two-prey population dynamics described by a control system. We study and compare conditions for permanence of the system for three types of predator feeding behaviors: (i) specialized feeding on the more profitable prey type, (ii) generalized feeding on both prey types, and (iii) optimal foraging behavior. We show that the region of parameter space leading to permanence for optimal foraging behavior is smaller than that for specialized behavior, but larger than that for generalized behavior. This suggests that optimal foraging behavior of predators may promote coexistence in predator-prey systems. We also study the effects of the above three feeding behaviors on apparent competition between the two prey types.     

Novel species interactions and environmental conditions reduce foraging competency at the temperate range edge of a range-extending coral reef fish

Poleward range extensions of coral reef species can reshuffle temperate communities by generating competitive interactions that did not exist previously. However, novel environmental conditions and locally adapted native temperate species may slow tropical invasions by reducing the ability of invaders to access local resources (e.g. food and shelter). We test this hypothesis on wild marine fish in a climate warming hotspot using a field experiment encompassing artificial prey release. We evaluated seven behaviours associated with foraging and aggressive interactions in a common range-extending coral reef fish (Abudefduf vaigiensis) and a co-shoaling temperate fish (Microcanthus strigatus) along a latitudinal temperature gradient (730 km) in SE Australia. We found that the coral reef fish had reduced foraging performance (i.e. slower prey perception, slower prey inspection, decreased prey intake, increased distance to prey) in their novel temperate range than in their subtropical range. Furthermore, higher abundance of temperate fishes was associated with increased retreat behaviour by coral reef fish (i.e. withdrawal from foraging on released prey), independent of latitude. Where their ranges overlapped, temperate fish showed higher foraging and aggression than coral reef fish. Our findings suggest that lower foraging performance of tropical fish at their leading range edge is driven by the combined effect of environmental factors (e.g. lower seawater temperature and/or unfamiliarity with novel conditions in their extended temperate ranges) and biological factors (e.g. increased abundance and larger body sizes of local temperate fishes). Whilst a future increase in ocean warming is expected to alleviate current foraging limitations in coral reef fishes at leading range edges, under current warming native temperate fishes at their trailing edges appear able to slow the range extension of coral reef fishes into temperate ecosystems by limiting their access to resources.

     

Intraspecific interference influences the use of prey hotspots

Resource clumping (prey hotspots) and intraspecific competition may interact to influence predator foraging behaviour. Optimal foraging theory suggests that predators should concentrate most of their foraging activity on prey hotspots, but this prediction has received limited empirical support. On the other hand, if prey concentration at hotspots is high enough to allow its use by several individuals, increased competition may impose constraints on foraging decisions of conspecifics, resulting in a temporal segregation in the use of shared resources. We investigated how artificial prey hotspots influence foraging behaviour in the Iberian lynx Lynx pardinus, and examined variations in the use of prey hotspots by individuals with different competitive abilities. All lynx, irrespective of their position in the competitive hierarchy, focused their activity on prey hotspots. Supplemented lynx concentrated higher proportions of active time on 100 m circular areas around hotspots than non‐supplemented lynx did on any area of the same size. However, we found evidence for a dynamic temporal segregation, where inferior competitors may actively avoid the use of prey hotspots when dominants were present. Dynamic temporal segregation may operate in solitary and territorial predators as a mechanism to facilitate coexistence of conspecifics in the absence of opportunities for spatial segregation when they use extremely clumped resources.     

外来入侵植物的根系觅养行为研究进展

土壤养分分布具有高度空间异质性, 植物的根系觅养行为是其对土壤养分异质性的一种适应。不同植物为了适应养分异质性会产生不同的根系觅养行为, 通过调整自身的根系觅养范围、觅养精度和觅养速度来更好地吸收利用土壤中的养分。外来植物与本地植物的竞争是决定其成功入侵的重要因素, 土壤养分等环境因素会影响它们之间的竞争关系。近年来, 外来入侵植物的觅养行为逐渐受到人们的关注, 关于入侵植物根系觅养行为的研究成果陆续出现: (1)总体来看, 外来入侵植物具有较强的根系觅养能力, 但根系觅养范围与觅养精度之间的权衡关系还不确定; (2)营养异质性会影响入侵植物与本地植物之间的竞争, 反过来, 二者之间的竞争也会影响根系觅养行为对营养异质性的响应; (3)丛枝菌根真菌(arbuscular mycorrhizal fungi, AMF)能够提高入侵植物的根系觅养能力, 外来植物入侵能够改变入侵植物对AMF的偏好性, 形成AMF对入侵的正反馈作用, 而本地植物与AMF的相互作用也会影响入侵植物的竞争力。未来还应加强营养异质环境下种间竞争和AMF共生对入侵植物根系觅养行为的影响机制研究, 以及全球变化背景下入侵植物根系觅养行为的变化与机制方面的研究, 可以更深入地认识外来植物的觅养行为在其成功入侵中的作用, 并为利用营养调控来防控入侵植物提供理论依据。     

Effects of Individual and Group Characteristics on Feeding Behaviors in Wild Leontopithecus rosalia

Studies have linked variation in feeding and foraging success to variation in survival and reproductive success, which makes exploring influences on feeding invaluable. In the current study, we quantified energy contents of foods consumed by wild golden lion tamarins (Leontopithecus rosalia, GLT) and feeding behaviors of 34 GLT from March 1998 to March 1999. Our objective was to test predictions regarding effects of characteristics of the 1) individual, 2) group, 3) environment, and 4) other behaviors on 3 feeding behaviors: feeding on plant matter, searching for prey, and feeding on prey. We hypothesized that environmental characteristics, e.g., resource availability, in addition to group characteristics e.g., group size, would influence feeding on plant matter, because several individuals in a group often consume fruit in the same fruit tree. We hypothesized that environmental characteristics and individual characteristics, e.g., age, would influence searching for and consuming prey because the individual often searches for and consumes prey while it is alone at a substrate. We used SAS mixed models to determine the relative influence of these characteristics on the feeding behaviors. We found that group characteristics more significantly influenced feeding on plant matter, while individual characteristics more significantly influenced searching for prey. The results emphasize the distinctly different influences of individual and group characteristics on feeding. That influences other than competition may affect feeding on plant matter warrants further exploration.     

Feeding habits of the exotic black bullhead Ameiurus melas (Rafinesque) in the Iberian Peninsula: first evidence of direct predation on native fish species

The feeding ecology of the exotic invasive black bullhead Ameiurus melas was conducted in the Iberian Peninsula for the first time. Dietary analysis based on the stomach contents of individuals caught in several Iberian basins was carried out as a first step to evaluate its potential threat for the native Iberian ichthyofauna. Aquatic macroinvertebrates (mainly Chironomidae) dominated the black bullhead's diet in all size-classes and sites, irrespective of natural riverine or artificial lentic habitats. Secondary prey items were responsible for the observed between-sites (microcrustaceans in artificial lentic habitat; oligochaeta and caddisfly larvae in natural riverine habitats) and ontogenetic diet differences (from microcrustaceans to larger prey). These diet variations were also detected in trophic diversity values and feeding strategy plots. Black bullheads consumed plant material, terrestrial prey and co-occurring fish species (native or exotic) and thus they could be considered as generalist or opportunistic, foraging on the most abundant and available prey. There was no positive relationship between black bullhead size (total length) and fish prey size, probably indicating piscivory on dead or dying vulnerable fishes as well as predation on smaller-sized active fishes. The results showed that the black bullhead could negatively affect native Iberian ichthyofauna throughout direct predation and competition. Aspects of potential conservation and management implications of fishes resulting from the undesirable presence of the black bullhead in Iberian water bodies are discussed.     

Effects of Individual and Group Characteristics on Feeding Behaviors in Wild <Emphasis Type="Italic">Leontopithecus rosalia</Emphasis>

Studies have linked variation in feeding and foraging success to variation in survival and reproductive success, which makes exploring influences on feeding invaluable. In the current study, we quantified energy contents of foods consumed by wild golden lion tamarins (Leontopithecus rosalia, GLT) and feeding behaviors of 34 GLT from March 1998 to March 1999. Our objective was to test predictions regarding effects of characteristics of the 1) individual, 2) group, 3) environment, and 4) other behaviors on 3 feeding behaviors: feeding on plant matter, searching for prey, and feeding on prey. We hypothesized that environmental characteristics, e.g., resource availability, in addition to group characteristics e.g., group size, would influence feeding on plant matter, because several individuals in a group often consume fruit in the same fruit tree. We hypothesized that environmental characteristics and individual characteristics, e.g., age, would influence searching for and consuming prey because the individual often searches for and consumes prey while it is alone at a substrate. We used SAS mixed models to determine the relative influence of these characteristics on the feeding behaviors. We found that group characteristics more significantly influenced feeding on plant matter, while individual characteristics more significantly influenced searching for prey. The results emphasize the distinctly different influences of individual and group characteristics on feeding. That influences other than competition may affect feeding on plant matter warrants further exploration. An erratum to this article can be found at     

Does diet in Celtic Sea fishes reflect prey availability?

Feeding preferences of Celtic Sea fishes were investigated using a database of stomach content records, collected between 1977 and 1994. The diet of cod Gadus morhua , hake Merluccius merluccius , megrim Lepidorhombus whiffiagonis , whiting Merlangius merlangus and saithe Pollachius virens changed markedly as the animals grew larger, and although large predators generally chose larger bodied prey, the variability of prey sizes consumed also increased. Large predators continued to select small, low value, benthic prey ( e.g . Callionymus spp. and Trisopterus spp.) which were easier to catch, rather than larger, more energy lucrative pelagic prey ( e.g . mackerel Scomber scombrus ), even though these pelagic prey‐fishes were nearly always available and were often very abundant. Stock estimates of the International Council for the Exploration of the Sea and U.K. groundfish survey catches were used as indices of prey abundance. Blue‐whiting Micromesistius poutassou and other small pelagic fishes ( Argentina spp. and clupeoids) were identified as being particularly important, and were consumed by some predators more often than would be expected given the abundance of these prey in the environment. There was no evidence for density‐dependent feeding by predators on mackerel and only hake exhibited density‐dependent feeding on horse‐mackerel. Hake, cod and megrim consumed more blue‐whiting when this prey was at higher abundance in the environment. In choosing what prey to consume, predators must balance costs and benefits, considering the quality of prey and the energy expended during search, capture and handling.     

Is larger better in sit-and-wait predators? Competitive superiority in Hydra

Contrary to a generalisation arising from many studies, a larger body size is not always the key to competitive superiority amongst animals. An analysis of competition between pairs of Hydra oligactis, Hydra vulgaris and Hydra circumcincta, that simultaneously encountered a single prey item, showed that competitive success in these sessile predators depended on species and clone in inter- and intraspecific competition, respectively. H. oligactis appeared to be competitively superior, even to the larger H. vulgaris individuals. Phenotypic traits important for prey capture, such as the fraction of the nematocyst that penetrates the prey (penetrants), were positively related to success in intraspecific competition. Body size appeared to be a positive key factor in determining foraging success in the competition between pairs of conspecifics from single or different clones. In contrast to the results of the intraspecific competition, body size was not significantly related to the foraging success of competing heterospecifics.