Sound production during competitive feeding in the grey gurnard

The acoustic repertoire of captive grey gurnard Eutrigla gurnardus during competitive feeding consisted of three types of sound: knocks, grunts and growls. Knocks were audible as a single sound, whereas grunts and growls were perceived as longer, pulsed sounds to the human ear. Typically, knocks were composed of 1–2 pulses, grunts of 4–8 pulses and growls >10 pulses. Growls were longer and had shorter pulse periods than grunts. All sound types had peak frequencies of c . 500 Hz. The sequences of behaviours observed during feeding interactions suggest that grey gurnard obtain food both by scramble and contest tactics. Competing fish emitted knocks mainly while grasping a food item and also during other non‐agonistic behaviour, suggesting that knock production may reflect a state of feeding arousal but could also serve as a warning of the forager's presence to nearby competitors. Grunts were mainly emitted during frontal displays, which were the most frequent behavioural act preceding grasps, suggesting that they may play a role in deterring other fish from gaining access to disputed food items.     

Ontogeny of Acoustic and Feeding Behaviour in the Grey Gurnard, Eutrigla gurnardus

Although sound production in teleost fish is often associated with territorial behaviour, little is known of fish acoustic behaviour in other agonistic contexts such as competitive feeding and how it changes during ontogeny. The grey gurnard, Eutrigla gurnardus, frequently emits knock and grunt sounds during competitive feeding and seems to adopt both contest and scramble tactics under defensible resource conditions. Here we examine, for the first time, the effect of fish size on sound production and agonistic behaviour during competitive feeding. We have made sound (alone) and video (synchronized image and sound) recordings of grey gurnards during competitive feeding interactions. Experimental fish ranged from small juveniles to large adults and were grouped in four size classes: 10–15, 15–20, 25–30 and 30–40 cm in total length. We show that, in this species, both sound production and feeding behaviour change with fish size. Sound production rate decreased in larger fish. Sound duration, pulse duration and the number of pulses increased whereas the peak frequency decreased with fish size, in both sound types (knocks and grunts). Interaction rate and the frequency of agonistic behaviour decreased with increasing fish size during competitive feeding sessions. The proportion of feeding interactions accompanied by sound production was similar in all size classes. However, the proportion of interactions accompanied by knocks (less aggressive sounds) and by grunts (more aggressive) increased and decreased with fish size, respectively. Taken together, these results suggest that smaller grey gurnards compete for food by contest tactics whereas larger specimens predominantly scramble for food, probably because body size gives an advantage in locating, capturing and handling prey. We further suggest that sounds emitted during feeding may potentially give information on the motivation and ability of the individual to compete for food resources.     

Feeding of grey gurnard Eutrigla gurnardus (Triglidae) in the area of the Rockall Seamount

Specific features of feeding of grey gurnard Eutrigla gurnardus depending on vertical distribution and habitat conditions in the area of the Rockall Seamount in 2000 were considered. The intensity of feeding of grey gurnard from April to September decreased by several times. The main food of this species in spring are euphausiids and that in autumn are fish. Males feed slightly more intensively than females. The intensity of feeding, distribution, and behavior of grey gurnard in different seasons and at different depths near bottom and in the pelagial noticeably differ and depend on the composition of distribution and numbers of dominant food items (euphausiids, sand eel Ammodytes marinus, fish juveniles).     

Foraging on patchily distributed prey by a cichlid fish (Teleostei,Cichlidae): A test of the ideal free distribution theory

Cichlid fish (Aequidens curviceps) distributed themselves and allocated their foraging time between two drift food patches in close approximation to the patch profitability ratio, as predicted by the ideal free distribution theory. The fish thereby achieved similar average feeding rates in the two patches, in two of three patch profitability ratio experiments. However, one major assumption of the ideal free model was violated, since individual fish differed in their competitive abilities for limited food resources, which resulted in unequal payoffs among individuals within each patch. Individual variation in feeding rates, and thus in competitive ability, was not related to despotism, but perhaps rather to individual differences in perceptual ability and in the ability to learn which patch was currently the more profitable. The strategy used by the fish to assess patch profitability included sampling available patches. However, individual fish switched (sampled) patches with varying frequency. Sampling had an associated cost, since high-frequency switchers had lower feeding rates on average than low-frequency switchers. Differences in foraging strategy among the fish therefore contributed to the observed in-equality in individual payoffs within patches.     

Networks of prey specialization in an Arctic monomorphic seabird

Generalist predator populations are sometimes made up of individuals that specialize on particular prey items. To examine specialization in thick‐billed murres Uria lomvia during self‐feeding we obtained stomach contents and muscle stable isotope values for 213 birds feeding close to five colonies in the Canadian Arctic. Adults were less specialized during self‐feeding than during chick‐provisioning. Nonetheless, particular specialists clustered together within the foraging network. While sexes showed similar levels of specialization, individuals of the same sex clustered together within the foraging network. The significant degree of clustering regardless of sex showed that individuals specializing on one prey item tend to also specialize on another, although network topology varied from colony to colony. Adult muscle stable isotope values correlated with the stable isotope values of the prey found in stomachs, at least at the one colony with relevant prey data, suggesting that specializations are maintained over time. Degree of specialization increased with niche width across the five colonies, but similarity in gastro‐intestinal and bill morphology was independent of dietary similarity. Thus, although individual specialization is thought to play a key role in sympatric speciation through trophic specialization, we found no support for an association between morphology and foraging patterns in our species. We conclude that self‐feeding murres show clustered dietary specialization, and that specialization is highest where diet is most diverse.     

Water depth selection, daily feeding routines and diets of waterbirds in coastal lagoons in Ghana

Water depth requirements, diet, feeding styles and diurnal activity patterns are described for waterbirds using two brackish water lagoon systems in coastal Ghana, the Songor and Keta Lagoons. We project the habitat and activity data on a guild structure defined on the basis of individual feeding style and the sensory mechanism used to detect food. A total of 3199 flocks containing 118,648 individuals of 36 different waterbird species were examined during October-November 1994. Feeding habitats varied from dry mudflats to wet mud and shallow water of not more than 20 cm. The depth of water selected by waterbirds for foraging (but not for roosting) was correlated with tarsus length. Foraging birds exhibited a wide range of feeding styles using visual and/or tactile means for detecting prey: pecking, probing, stabbing, sweeping and ploughing, sometimes feeding singly, communally or socially in loose or dense flocks. Prey items taken ranged from seeds of Widgeongrass Ruppia maritima to invertebrates (mainly polychaetes, molluscs and crabs) and fish, mainly juvenile Tilapia. The daytime was spent on two main activities, feeding and roosting, with a small fraction of the time (average of 10% for 25 species) spent on comfort activities. The waterbirds exhibited either a circadian (most waders, except Common Sandpipers Actitis hypoleucos and Turnstones Arenaria interpres) or a diurnal foraging activity pattern (herons and terns), with no purely nocturnal species. Some species fed throughout the day, others showed peak foraging at various times of the day. The proportion of time spent foraging was related to guild (highest in visual and tactile surfaceforaging waders) and was negatively correlated with the size of the species. We conclude that the observed patterns in the use of the 24-h day by waterbirds for foraging are not species specific but vary depending on conditions on the feeding grounds. Nocturnal foraging is a normal and a regular strategy used by waterbirds to obtain enough food to fulfill their energetic requirements, so that irrespective of the sensory mechanism used to detect prey and the conditions prevailing on the feeding grounds, waterbirds forage day and night as dictated by their energetic needs. Water depth appears to be the key environmental factor controlling the availability of food for the waterbirds in the Ghanaian lagoons.     

Foraging and habitat use by Nycteris grandis (Chiroptera: Nycteridae) in Zimbabwe

This study describes the foraging strategies, foraging areas, and roosts used by four female radio-tagged Nycteris grandis along the Zambezi River in Mana Pools National Park in Zimbabwe. The bats exhibited two foraging strategies, sometimes hunting from perches, other times flying continuously close to the ground. Some individuals used one strategy more than the other. During the study, the bats fed mainly on frogs, selecting only Ptychadena anchietae ; they also took crickets, moths and cicadas. Each bat usually consumed two frogs each night and patterns of occupation of feeding perches suggest that their foraging efficiency was between 21 and 36%. The marked bats roosted in an old water tower or in a hollow Acacia albida. Four of five marked females were nursing young during this study.     

Foraging innovation in the guppy

When novel behaviour patterns spread through animal populations, typically one animal will initiate the diffusion. It is not known whether such 'innovators' are particularly creative individuals, individuals exposed to the appropriate environmental contingencies, or individuals in a particular motivational state. We describe three experiments that investigated the factors influencing foraging innovation in the guppy Poecilia reticulata. We exposed small laboratory populations of fish to novel foraging tasks, which involved exploration and problem solving to locate a novel food source. Experiments 1 and 2 found that (1) females were more likely to innovate than males, (2) food-deprived fish were more likely to innovate than nonfood-deprived subjects, and (3) smaller fish were more likely to innovate than larger fish. We suggest that the sex difference may reflect parental investment asymmetries in males and females. Experiment 3 found that past innovators were more likely to innovate than past noninnovators. Collectively, the results suggest that differences in foraging innovation in guppies are best accounted for by differences in motivational state, but, in addition, guppies may vary in their predisposition to innovate. Copyright 1999 The Association for the Study of Animal Behaviour.     

Shifting the balance between foraging and predator avoidance: the importance of food distribution for a schooling pelagic forager

It is widely held that when predator avoidance conflicts with other activities, such as feeding, avoidance of predators often takes precedence. In this study, we examine how predation risk and food distribution interact to influence the schooling behavior and swimming speed of foraging juvenile walleye pollock, Theragra chalcogramma. Fish were acclimated to either spatially and temporally clumped, or spatially and temporally dispersed food for 3 weeks. Fish were then monitored while feeding in the absence and presence of predatory sablefish, Anoplopoma fimbria. Fish foraging for clumped food swam rapidly in a loose school when predators were absent, but swam more slowly and adopted more cohesive schooling in the presence of predators, trading-off foraging opportunity for decreased vulnerability to predators. Fish foraging for dispersed food swam about slowly and did not engage in cohesive schooling in either the absence or presence of predators. These fish accepted greater predation risk in order to continue foraging, suggesting that the cost of schooling, in terms of decreased foraging opportunity, was greater when food was dispersed than when it was clumped. This lower responsiveness to predators among fish receiving dispersed food demonstrates that predator avoidance does not always take precedence over other activities, but rather, that a balance is maintained between predator avoidance and feeding, which shifts as food distribution changes.     

Feeding ecology of wintering terns in Guinea-Bissau

We studied the feeding ecology of Little Terns Sterna albifrons , Sandwich Terns S. sandvicensis and Royal Terns S. maxima in the Archipélago dos Bijagós (11°40'N, 15°45'W) in Guinea-Bissau (West Africa) during the winter of 1992/1993. More than 95% of all prey taken by these terns were roundfish, ranging in weight from 0.3 to 40 g. Birds usually fed alone, but sometimes they were observed feeding in mixed-species flocks consisting of 15–200 individuals. Capture rate ( n fish per hour foraging) in these flocks was higher than that of solitary birds. However, smaller fish were caught by birds foraging in flocks, so food intake rate (g/h) did not differ between solitary and flock-feeding birds. The relationships between foraging behaviour of the three tern species and abiotic factors, such as time, tide and water clarity, have been investigated. Capture rate of Royal Terns increased with water clarity. For Little Terns and Sandwich Terns, food intake rate was lower in the most turbid waters compared to clearer waters. There was very little foraging activity during high tide. For Little Terns and Royal Terns, food intake rate was about twice as high during receding and low tides as during an incoming tide. Food intake rate averaged 8 g/h in Little Terns, 60 g/h in Sandwich Terns and 45 g/h in Royal Terns. With a rough model, we estimate the maximum rate of daily energy expenditure of terns wintering in the tropics at 3 × BMR (defined as energy expenditure of inactive bird at thermoneutrality in a post-absorptive state during the resting phase of the daily cycle). From an energetic viewpoint, wintering Sandwich Terns in Guinea-Bissau seem to have an easy living.     

Vocalisation Repertoire of Female Bluefin Gurnard (Chelidonichthys kumu) in Captivity: Sound Structure,Context and Vocal Activity

Fish vocalisation is often a major component of underwater soundscapes. Therefore, interpretation of these soundscapes requires an understanding of the vocalisation characteristics of common soniferous fish species. This study of captive female bluefin gurnard, Chelidonichthys kumu, aims to formally characterise their vocalisation sounds and daily pattern of sound production. Four types of sound were produced and characterised, twice as many as previously reported in this species. These sounds fit two aural categories; grunt and growl, the mean peak frequencies for which ranged between 129 to 215 Hz. This species vocalized throughout the 24 hour period at an average rate of (18.5 ± 2.0 sounds fish^-1 h^-1) with an increase in vocalization rate at dawn and dusk. Competitive feeding did not elevate vocalisation as has been found in other gurnard species. Bluefin gurnard are common in coastal waters of New Zealand, Australia and Japan and, given their vocalization rate, are likely to be significant contributors to ambient underwater soundscape in these areas.     

Food choice causes interrupted feeding in the generalist grasshopper Schistocerca americana: further evidence for inefficient decision-making

Individual grasshoppers were observed foraging on either single food plant species or binary mixtures of them. Individuals ate slightly less on the mixtures, but grew faster on them. In the mixtures, individuals fed regularly from both plant species, typically eating both within a single meal. Feeding bouts were longer on the preferred plant species in the mixture. The pattern of foraging behavior was markedly different in mixtures and single food treatments. In mixtures, although the total length of time engaged in ingestion over the day was slightly less than in single food treatments, there were relatively more short feeding bouts and more gaps between feeding bouts within meals, so that the time spent in activities associated with the food was markedly longer on mixtures. This is discussed in terms of efficiency and the problem of choice that faces individual generalists. The efficiency factor and its impact on vigilance is added to the other costs of being a polyphagous herbivore.     

Can a minority of informed leaders determine the foraging movements of a fish shoal?

There is no information on whether the daily foraging movements of fish shoals are the result of chance, the collective will of all shoalmates, or the leadership of a few individuals. This study tested the latter possibility. Shoals of 12 golden shiners, Notemigonus crysoleucas, were trained to expect food around midday in one of the brightly lit corners of their tank. They displayed daily food-anticipatory activity by leaving the shady area of their tank and spending more and more time in the food corner up to the normal time of feeding. Past this normal time they remained in the shade, even on test days when no food was delivered. Most of these experienced individuals were then replaced by na?ve ones. The resulting ratio of experienced:na?ve fish could be 5:7, 3:9 or 1:11. On their own, na?ve individuals would normally spend the whole day in the shade, but in all tests the experienced individual(s) were able to entrain these more numerous na?ve fish out of the shade and into the brightly lit food corner at the right time of day. Entrainment was stronger in the 5:7 than in the 1:11 experiment. The test shoals never split up and were always led by the same fish, presumably the experienced individuals. These results indicate that in a strongly gregarious species, such as the golden shiner, a minority of informed individuals can lead a shoal to food, either through social facilitation of foraging movements or by eliciting following behaviour. Copyright 2000 The Association for the Study of Animal Behaviour.     

Enhancing knowledge on low-value fishing species: the distinct reproductive strategy of two gurnard species

The depletion and overexploitation of several fish stock demands for a valorisation of non-target and discarded species. Nonetheless, such species are often poorly studied, and information on their biological parameters must be gathered for effective population management. For 1 year, the reproductive strategy of the piper gurnard Trigla lyra and the red gurnard Chelidonichthys cuculus was studied by monthly samples obtained from commercial boats operating on western Portuguese coast. Both species showed a biased sex ratio towards females, especially for larger length classes. Length at first maturity could be estimated only for red gurnard (22.1 and 19.9 cm for females and males, respectively) because all piper gurnard individuals caught were mature. Piper gurnard showed determinate fecundity and a short spawning season, from November to February with a peak in January, whereas red gurnard showed indeterminate fecundity and a wide spawning season, from late December to May. The relative annual fecundity estimated for red gurnard (1893 ± 728 oocytes × g⁻¹ eviscerated weight [EW]) was higher than the one estimated for piper gurnard (1018 ± 250 oocytes×g⁻¹ EW). Although important information for understanding the species dynamics is presented in this study, additional information on other life-history parameters and of species landings is required.     

Within-colony feeding selectivity by a corallivorous reef fish: foraging to maximize reward?

Foraging theory predicts that individuals should choose a prey that maximizes energy rewards relative to the energy expended to access, capture, and consume the prey. However, the relative roles of differences in the nutritive value of foods and costs associated with differences in prey accessibility are not always clear. Coral‐feeding fishes are known to be highly selective feeders on particular coral genera or species and even different parts of individual coral colonies. The absence of strong correlations between the nutritional value of corals and prey preferences suggests other factors such as polyp accessibility may be important. Here, we investigated within‐colony feeding selectivity by the corallivorous filefish, Oxymonacanthus longirostris, and if prey accessibility determines foraging patterns. After confirming that this fish primarily feeds on coral polyps, we examined whether fish show a preference for different parts of a common branching coral, Acropora nobilis, both in the field and in the laboratory experiments with simulated corals. We then experimentally tested whether nonuniform patterns of feeding on preferred coral species reflect structural differences between polyps. We found that O. longirostris exhibits nonuniform patterns of foraging in the field, selectively feeding midway along branches. On simulated corals, fish replicated this pattern when food accessibility was equal along the branch. However, when food access varied, fish consistently modified their foraging behavior, preferring to feed where food was most accessible. When foraging patterns were compared with coral morphology, fish preferred larger polyps and less skeletal protection. Our results highlight that patterns of interspecific and intraspecific selectivity can reflect coral morphology, with fish preferring corals or parts of coral colonies with structural characteristics that increase prey accessibility.     

Optimal foraging theory predicts effects of environmental enrichment in a group of adult golden lion tamarins

The success of environmental enrichment programs in effecting specific changes in the behavior of captive animals has not always been uniform. Separate studies demonstrated both an increase in food competition and a decrease in food competition among captive group-living primates upon introduction of foraging devices. The objectives of this study were to measure the effects of variation in resource distribution and availability on food competition in a group of captive adult golden lion tamarins (Leontopithecus rosalia). The resource variables chosen were suggested from optimal foraging theory. The energy invested to obtain an item influenced food transfer and aggressive behaviors while food abundance did not. All individuals obtained an equivalent number of items over the course of the experiment from the foraging device, even though some tamarins obtained most of their food rewards directly from the device while others received their food rewards primarily through food transfer from other group members. Because the monkeys appeared highly motivated to obtain food from the test apparatus and did not habituate to it, the foraging device used in this experiment could be used as regular environmental enrichment for golden lion tamarins. One way to circumvent potentially unacceptable rates of aggression, with this or any feeding protocol that increases foraging task complexity and search time, may be to provide more than one foraging device per group. Zoo Biol 17:231–244, 1998. © 1998 Wiley-Liss, Inc.     

Swimbladder volume and body density in an armoured benthic fish, the streaked gurnard

Body density of streaked gurnards Trigloporus lastoviza is anomalously low (mean 1·043 g ml⁻¹) for a benthic fish and overlaps with that of pelagic fish such as cod and lumpfish. It is clear that the heavy armour of gurnards is offset by buoyancy provided by the swimbladder. Swimbladder volume is a mean 3·88% of gurnard volume. Evidence is given to indicate that this is quite large for a marine teleost: data are also presented to show that the '5% of fish volume for marine fish, 7% for freshwater fish' rule for swimbladder volumes is unreliable.     

Tracing the life history of red gurnard (Aspitrigla cuculus) using validated otolith annual rings

Back calculation, marginal increment analysis and modal progression analysis were used to trace growth processes of the red gurnard, Aspitrigla cuculus L., in the Mediterranean Sea. Three false rings appeared regularly on otoliths and were hypothesized to refer to the ontogeny on the basis of the study of the recruitment and reproduction processes of the species. The first two rings were laid down during the first pelagic period and the transformation to the bottom mode of life, respectively. The third ring corresponded to the first spawning at the end of the second year of life. The marginal increment analysis demonstrated that one opaque and one translucent zone is formed each year, the translucent ring corresponding to the period of slow growth in the winter. Temperature and salinity of Mediterranean waters below the thermocline where red gurnard live do not undergo seasonal variations. Therefore, traditional interpretation of the discontinuity patterns on otoliths related to the decrease of water temperature should be rejected for Mediterranean fish distributed on the deep shelf and slope. Discontinuities in otolith structure may be related to seasonal variation in feeding intensity as a consequence of decreases in the benthic mysids biomass and time available to find food due to reduced daylight. Future research should focus on the trophic ecology of the demersal fishes and the seasonal fluctuations of the prey resources during the year in order to assess the role of the food supply with regard to otolith growth patterns in the Mediterranean Sea.     

The costs of becoming nocturnal: feeding efficiency in relation to light intensity in juvenile Atlantic Salmon

1. Most animals are active by day or by night, but not both; juvenile salmonids are unusual in that they switch from being predominantly diurnal for most of the year to being nocturnal in winter. They are visual foragers, and adaptations for high visual acuity at daytime light intensities are generally incompatible with sensitive night vision. Here we test whether juvenile Atlantic Salmon Salmo salar are able to maintain their efficiency of prey capture when switching between diurnal and nocturnal foraging.
2. By testing the ability of the fish to acquire drifting food items under a range of manipulated light intensities, we show that the foraging efficiency of juvenile salmon is high at light intensities down to those equivalent to dawn or dusk, but drops markedly at lower levels of illumination: even under the best night condition (full moon and clear sky), the feeding efficiency is only 35% of their diurnal efficiency, and fish will usually be feeding at less than 10% (whenever the moon is not full, skies are overcast or when in the shade of bankside trees). Fish were unable to feed on drifting prey when in complete darkness.
3. The ability of juvenile salmon to detect prey under different light intensities is similar to that of other planktivorous or drift-feeding species of fish; they thus appear to have no special adaptations for nocturnal foraging.
4. While winter drift abundance is slightly higher by night than by day, the difference is not enough to compensate for the loss in foraging efficiency. We suggest that juvenile salmon can nonetheless switch to nocturnal foraging in winter because their food requirements are low, many individuals adopting a strategy in which intake is suppressed to the minimum that ensures survival.     

Facilitation and Dominance in a Schooling Predator: Foraging Behavior of Florida Pompano,Trachinotus carolinus

Presumably an individual’s risk of predation is reduced by group membership and this ‘safety in numbers’ concept has been readily applied to investigations of schooling prey; however, foraging in groups may also be beneficial. We tested the hypothesis that, when feeding in groups, foraging of a coastal fish (Florida Pompano, Trachinotus carolinus) on a benthic prey source would be facilitated (i.e. fish feeding in groups will consume more prey items). Although this question has been addressed for other fish species, it has not been previously addressed for Florida Pompano, a fish known to exhibit schooling behavior and that is used for aquaculture, where understanding the feeding ecology is important for healthy and efficient grow-out. In this experiment, juvenile Florida Pompano were offered a fixed number of coquina clams (Donax spp.) for one hour either in a group or as individuals. The following day they were tested in the opposite configuration. Fish in groups achieved greater consumption (average of 26 clams consumed by the entire group) than the individuals comprising the group (average of 14 clams consumed [sum of clams consumed by all individuals of the group]). Fish in groups also had fewer unsuccessful foraging attempts (2.75 compared to 4.75 hr^-1) and tended to have a shorter latency until the first feeding activity. Our results suggest fish in groups were more comfortable feeding and more successful in their feeding attempts. Interestingly, the consumption benefit of group foraging was not shared by all – not all fish within a group consumed equal numbers of clams. Taken together, the results support our hypothesis that foraging in a group provides facilitation, but the short-term benefits are not equally shared by all individuals.