Parasite distribution on client reef fish determines cleaner fish foraging patterns

Recent evidence suggests that cleaner fish Labroides dimidiatus effectively control parasite densities on client reef fish that actively visit them to have parasites and dead or infected tissue removed. These findings support the hypothesis that clients benefit from cleaning, However, they do not show how cleaners reduce the parasite load of their clients. Cleaners could selectively feed on parasites or parasite removal could be a side product of cleaners foraging indifferently on the client surface, resulting in the removal of healthy mucus and scales also. To investigate cleaner fish foraging behaviour, we infected individuals of the surgeon fish Ctenochaetus striatus, with parasitic monogeneans on one body side, while the other body side was parasite free. We then allowed these clients to interact with L, dimidiatus. We found that the duration of interactions depended on parasite load, and that cleaners spent both more time and took more bites per time unit on the infected than on the uninfected side, Our data thus support the idea that parasite abundance determines food patch quality for cleaners. The overall outcome of cleaning interactions is thus likely to benefit the clients.

Learning of foraging skills by fish

This chapter outlines the relationships between a number of key factors that influence learning and memory, and illustrates them by reference to studies on the foraging behaviour of fish. Learning can lead to significant improvements in foraging performance in only a few exposures, and at least some fish species are capable of adjusting their foraging strategy as patterns of patch profitability change. There is also evidence that the memory window for prey varies between fish species, and that this may be a function of environmental predictability. Convergence between behavioural ecology and comparative psychology offers promise in terms of developing more mechanistically realistic foraging models and explaining apparently 'suboptimal' patterns of behaviour. Foraging decisions involve the interplay between several distinct systems of learning and memory, including those that relate to habitat, food patches, prey types, conspecifics and predators. Fish biologists, therefore, face an interesting challenge in developing integrated accounts of fish foraging that explain how cognitive sophistication can help individual animals to deal with the complexity of the ecological context.

Pescadores y aves marinas compitiendo por el mismo recurso: estrategias de forrajeo, interacciones y consecuencias

Con el examen conjunto de datos de seguimiento satelital de la flota pesquera y de marcaje electrónico de aves marinas, se analizan las interacciones entre la actividad pesquera y el forrajeo de las aves en periodo de reproducción. Se evidencia que las aves pueden mitigar la competencia con la pesca hasta cierto punto, forrajeando más lejos o quedándose más tiempo en el mar. Sin embargo, las aves en reproducción enfrentan a la vez un alto requerimiento energético para alimentar los pichones, y viajes de forrajeo limitados en distancia y tiempo por la necesidad de atender el nido. Para optimizar el éxito reproductivo de las aves marinas se recomienda: (1) Estimar, con la ayuda de modelos ecotróficos, qué cantidad de anchoveta sería conveniente ‘reservar’ para la alimentación de las aves; (2) Establecer zonas temporalmente cerradas a la pesca, alrededor de las principales colonias en los meses de reproducción de las aves. Los rangos máximos de forrajeo observados en guanayes y piqueros sugieren un radio de 50 a 100 km alrededor de las colonias, lo cual permitiría asegurar el forrajeo de estas especies en periodo de reproducción y así favorecer la sostenibilidad de sus poblaciones.

The moving gardens: Reef fishes grazing, cleaning, and following green turtles in SW Atlantic

Reef fishes may associate with marine turtles and graze on their shells, or clean their head, neck and flippers. on a reef flat at Fernando de Noronha Archipelago, SW Atlantic, we recorded green turtles (Chelonia mydas) grazed, cleaned and followed by reef fishes. The green turtle seeks specific sites on the reef and pose there for the grazers and/or cleaners. Fishes recorded associated to green turtles included omnivorous and herbivorous reef species such as the dam-selfish Abudefduf saxatilis and the surgeonfishes Acanthurus chirurgus and A. coeruleus. The turtle is followed by the wrasse Thalassoma noronhanum only while engaged in foraging bouts on benthic algae. Following behaviour is a previously unrecorded feeding association between turtles and fishes.

Food web structure in a tropical stream ecosystem

This study investigated the structure and properties of a tropical stream food web in a small spatial scale, characterizing its planktonic, epiphytic and benthic compartments. The study was carried out in the Potreirinho Creek, a second-order stream located in the south-east of Brazil. Some attributes of the three subwebs and of the conglomerate food web, composed by the trophic links of the three compartments plus the fish species, were determined. Among compartments, the food webs showed considerable variation in structure. The epiphytic food web was consistently more complex than the planktonic and benthic webs. The values of number of species, number of links and maximum food chain length were significantly higher in the epiphytic compartment than in the other two. Otherwise, the connectance was significantly lower in epiphyton. The significant differences of most food web parameters were determined by the increase in the number of trophic species, represented mainly by basal and intermediate species. High species richness, detritus-based system and high degree of omnivory characterized the stream food web studied. The aquatic macrophytes probably provide a substratum more stable and structurally complex than the sediment. We suggest that the greater species richness and trophic complexity in the epiphytic subweb might be due to the higher degree of habitat complexity supported by macrophyte substrate. Despite differences observed in the structure of the three subwebs, they are highly connected by trophic interactions, mainly by fishes. The high degree of fish omnivory associated with their movements at different spatial scales suggests that these animals have a significant role in the food web dynamic of Potreirinho Creek. This interface between macrophytes and the interconnections resultant from fish foraging, diluted the compartmentalization of the Potreirinho food web.

Greater Anis (Crotophaga major) Commensal Foraging with Freshwater Fish in the Pantanal Floodplain, Brazil

Foraging associations between birds and other groups of animals have been widely reported in the literature. I report the first observation of a foraging tactic involving a flock of Greater Ani (Crotophaga major), which deliberately followed fish along an artificial ditch in the Pantanal wetlands, feeding on animals flushed by the movement of the vegetation on the ditch banks. Further observations of the feeding behavior and foraging tactics of Greater Anis are necessary to ascertain if this type of behavior is a frequent event or merely sporadic. Received 22 June 2010. Accepted 11 October 2010.

A Foraging Cost of Migration for a Partially Migratory Cyprinid Fish

Migration has evolved as a strategy to maximise individual fitness in response to seasonally changing ecological and environmental conditions. However, migration can also incur costs, and quantifying these costs can provide important clues to the ultimate ecological forces that underpin migratory behaviour. A key emerging model to explain migration in many systems posits that migration is driven by seasonal changes to a predation/growth potential (p/g) trade-off that a wide range of animals face. In this study we assess a key assumption of this model for a common cyprinid partial migrant, the roach Rutilus rutilus, which migrates from shallow lakes to streams during winter. By sampling fish from stream and lake habitats in the autumn and spring and measuring their stomach fullness and diet composition, we tested if migrating roach pay a cost of reduced foraging when migrating. Resident fish had fuller stomachs containing more high quality prey items than migrant fish. Hence, we document a feeding cost to migration in roach, which adds additional support for the validity of the p/g model of migration in freshwater systems.

Foraging by the omnivorous crab Pachygrapsus transversus affects the structure of assemblages on sub-tropical rocky shores

The roles of herbivory and predation in determining the structure and diversity of communities have been tested across most intertidal systems. In contrast, the importance of omnivorous consumers remains untested in many rocky shore communities. We tested the role of a small omnivorous crab in an intertidal food web on rocky shores of the sub-tropical southwest Atlantic. Exclusion of the grapsid crab Pachygrapsus transversus in the field resulted in significant changes in the abundance of functional groups in the sublittoral fringe of sheltered shores, where the dominant cover changed from a suite of macroalgae to an assemblage of filter-feeding animals (ascidians, sponges, mussels). In contrast, limpets, whelks, large crabs and fish did not significantly affect community composition of the assemblage. To examine the omnivorous feeding pattern of P. transversus, we did laboratory experiments to test its foraging behaviour among animal and algal groups. The crab showed selective behaviour, preferring invertebrate groups to macroalgae, and opportunistic behaviour among types of prey within those major groups. According to our results, the role of slow-moving and large fast-moving consumers is apparently negligible compared to the effect of an omnivorous consumer. P. transversus plays an important role in determining the intertidal community composition on these subtropical rocky shores, causing changes in the balance of functional groups and controlling invasive species.

Effects of algal turbidity on foraging and antipredator behaviour of the three-spined stickleback (Gasterosteus aculeatus)

The aim of this thesis was to examine how aquatic organisms, such as fish, behave in an altered environmental condition. Many species of fish use vision as their primary tool to gain information about their surrounding environment. The visual conditions of aquatic habitats are often altered as a result of anthropogenic disturbance, such as eutrophication that initiates algal turbidity. In general, turbidity reduces the visibility and can be hypothesized to have an influence on the behaviour of fish. I used the three-spined stickleback (Gasterosteus aculeatus) as a model species and conducted four studies in the laboratory to test how algal turbidity affects its behaviour. In this thesis, two major behavioural aspects are discussed. The first is antipredator behaviour. In study I, the combined effects of turbidity and shoot density on habitat choice (shelter vs open) behaviour was tested on a group of sticklebacks (20 fish) in the presence and absence of piscivorous perch (Perca fluviatilis). In study II, I examined the behavioural responses of feeding sticklebacks when they were exposed to the sudden appearance of an avian predator (the silhouette of a common tern, Sterna hirundo). The study was done in turbid and clear water using three different groups sizes (1, 3 and 6 fish). The second aspect is foraging behaviour. Study III & IV focused on the effects of algal turbidity on the foraging performance of sticklebacks. In study III, I conducted two separate experiments to examine the effects of turbidity on prey consumption and prey choice of sticklebacks. In this experiment turbidity levels and the proportion of large and small prey (Daphnia spp.) were manipulated. In study IV, I studied whether a group of six sticklebacks can distribute themselves according to food input at two feeding stations in a way that provided each fish with the same amount of food in clear and turbid water. I also observed whether the fish can follow changes in resource distribution between the foraging patches. My results indicate an overall influence of algal turbidity on the antipredator and foraging behaviour of sticklebacks. In the presence of a potential predator, the use of the sheltered habitat was more pronounced at higher turbidity. Besides this, sticklebacks reduced their activity levels with predator presence at higher turbidity and shoot density levels, suggesting a possible antipredator adaptation to avoid a predator. When exposed to a sudden appearance of an avian predator, sticklebacks showed a weaker antipredator response in turbid water, which suggests that turbidity degrades the risk assessment capabilities of sticklebacks. I found an effect of group size but not turbidity in the proportion of sticklebacks that fled to the shelter area, which indicates that sticklebacks are able to communicate among group members at the experimental turbidity levels. I found an overall negative effect of turbidity on food intake. Both turbidity and changes in the proportion of prey sizes played a significant role in a stickleback’s prey selection. At lower turbidity levels (clear <1 and 5 NTU) sticklebacks showed preferences for large prey, whereas in more turbid conditions and when the proportion of large to small prey increased sticklebacks became increasingly random in their prey selection. Finally, my results showed that groups of sticklebacks disperse themselves between feeding stations according to the reward ratios following the predictions of the ideal free distribution theory. However, they took a significantly longer time to reach the equilibrium distribution in turbid water than in clear water. In addition, they showed a slower response to changes in resource distribution in a turbid environment. These findings suggest that turbidity interferes with the information transfer among group foragers. It is important to understand that aquatic animals are often exposed to a degraded environment. The findings of this thesis suggest that algal turbidity negatively affects their behavioural performance. The results also shed light on the underlying behavioural strategies of sticklebacks in turbid conditions that might help them adapt to an altered environmental situation and increase their survival. In conclusion, I hold that although algal turbidity has detrimental effects on the antipredator and foraging behaviour of sticklebacks, their behavioural adjustment might help them adapt to a changing environment.

Foraging ecology and parental behaviour in the common tern (Sterna hirundo)

One component of successful parenting is related to efficiency in foraging behaviour. The relationships among chick feeding, the size and type of food package, and length of parental foraging trips has not been well studied in seabirds. In addition, relatively few data have been collected on the activities of seabirds when foraging away from the nest site. The objectives of this study were: (1) to contrast productivity, feeding rate, and attendance patterns of individuals carrying a novel transmitter with a control group of birds; (2) to use radio-telemetry to assess the variability in foraging locations within and between individual male Common Terns; (3) to determine the seasonal variation in chick diet; (4) to determine for each transmittered bird, the relationships among the foraging patterns, parental behaviour, and seasonal reproductive success. The study took place over two years (1990-91) on a concrete breakwater 1 km offshore on Lake Erie near Port Colbome, Ontario. Ten pairs of terns in 1990 and 12 pairs in 1991 were radio-tracked by boat or car during the chick rearing stage. Concurrent behavioural observations documented the time each sex spent foraging or at the nest. The frequency and prey species composition of feeds to chicks were also recorded. The transmitters had negligible effects on the feeding frequency and brood attendance patterns of transmitter carrying birds. Peak nesting transmittered birds in 1990 and 1991 exhibited some inter-individual variability in foraging locations, however intraindividual variability was low. Birds foraged primarily to the west and northwest of the colony. Late nesters exhibited greater inter-individual variability, however intra-individual variability remained low for most birds. Neither group demonstrated sufficient variability to support the regular use of this colony as an "information centre". Individual transmittered birds had unique and predictable foraging patterns, and corresponding differences in feeding frequencies and brood attendance patterns, yet productivity was essentially equal between nests due to the impact and importance of stochastic events. Individuals that were recaptured in 1991 exhibited very similar foraging patterns to 1990, suggesting littie variability between years. Conservation of foraging patterns between years may have potential implications for mate choice decisions in future breeding seasons. Prey species delivered to chicks differed between morning and evening for peak and late nesters in 1990, but not 1991. Peak nesters in 1990 fed significantiy more Rainbow Smelt fOsmerus mordM) than Emerald Shiner (Notropis atherinoidesV this trend was reversed for late nesters who also fed large numbers of unidentified larval fish. No significant differences were found in 1991. Seasonal changes in prey species delivered to chicks is believed to be attributable to the temperature tolerances of the smelt and shiners, and the presence of large schools of larval fish during the late nesting season.

Trends in fish prey capture by common terns following zebra mussel invasion : do terns track fish species abundance? /

The Common Tern (Sterna hirundo) is a ground nesting colonial seabird. Terns rely primarily on small prey fishes which they obtain through plunge diving for their survival as well as the survival of their offspring during the breeding season. The zebra mussel (Dreissena polymorpha) is a small bivalve mollusk that invaded North American waters in the late 1980's. Through its suspension feeding, the zebra mussel has the ability to alter the entire aquatic ecosystem, ultimately leading to a reduction in pelagic organisms including small prey fish. The objective of the study was to determine what (if any) indirect effects the invasion of the zebra mussel has had on fish prey captured by terns. The study took place in two separate two-year periods, 1990-91 and 1995-96 on a concrete breakwall off the north shore of Lake Erie near Port Colborne, Ontario. Daily nest checks revealed clutch initiation dates, egg-laying chronology, hatching success and morphological egg characteristics (length and breadth). Behavioural observations included time each sex spent in attendance with its brood, the frequency of feeding chicks and the prey species composition and size fed to chicks as well as to females (courtship feeding). Egg sizes did not differ between study periods, nor did feeding rates to chicks, suggesting that food was not a limiting resource. Terns spent less time with their broods (more time foraging) in the 1995-96 period. However, they also had significantly larger broods and fledged more offspring. The time of each individual foraging trip decreased, suggesting that fish were easier to obtain in 1995 and 1996. Lastly, kleptoparasitism rates decreased, suggesting that the costs of foraging (time, energy) actually decreased as fewer birds adopted this strategy to compensate for what I assumed to be a lack of available food (fish). The only significant difference between the periods of 1990, 1991 and 1995, 1996 was a change in diet. Terns delivered significantly fewer rainbow smelt and more emerald shiner in 1995 and 1996. However, the average size of fish delivered did not change. Thus, there was little impact on prey captured by Common Terns in Lake Erie since the invasion of the zebra mussel.

Foraging ecology and parental care of common terns (Sterna hirundo) nesting in Windermere Basin, Lake Ontario

The relationships among chick feeding, size and type of prey item, and foraging time away from the brood have not been well studied in seabirds. This study investigated spatial and temporal patterns of foraging and chick-provisioning among 23 radio-tagged male common terns nesting at Hamilton Harbour, Lake Ontario during 1991 and 1992. Telemetry data were collected concurrently with behavioural observations from an elevated blind. Terns fitted with transmitters did not differ from controls with respect to either brood attendance, patterns of chick mortality, species and size distributions of prey delivered to offspring, or chick-provisioning rates. There was a clear separation of parental roles: males were primarily responsible for feeding chicks while females allocated more time to brood attendance. The prey species most commonly delivered to chicks by adults were rainbow smelt (Osmerus mordax) and alewife (A/osa pseudoharengus), followed in importance by larval fish, emerald shiner (Notropis antherinoides), salmonids, and fathead minnows (Pimepha/es prome/as). The relative proportions of various fish speCies delivered to chicks by males differed over the course of each breeding season, and there was also much variability in species composition of prey between years. Sizes of prey delivered to chicks also differed between sampling periods. The modal size of fish brought to chicks during Peak 1991 was 1.5 bill lengths, while the majority of prey in Late 1991 were small larval fish. The reverse trend occurred in 1992 when small fish were delivered to chicks predominantly during the Peak nesting period. During periods when predominantly small fish were delivered to chicks, the foraging activity of radio-tagged males was concentrated within a two kilometer radius of the colony. The observed variation in prey composition and foraging locations during the study likely reflects temporal variation in the availability of prey in the vicinity of the colony. Males delivered fish to chicks at a constant rate, while females 4 increased their feeding frequency over the first six to ten brood days. The mean length of fish delivered to chicks by adults increased significantly with increasing chick age. As a group, within each nesting period, transmittered males either foraged predominantly in the same directional bearing (north during Peak 1991, south during Late 1992), or concentrated foraging activity in the immediate vicinity of the colony (Late 1991, Peak 1992). However, individual radio-tagged males exhibited unique and predictable foraging patterns, often favouring specific locations within these areas and differing in their secondary foraging patterns. Overall, the Lake Ontario shoreline between NCB Bay" (3.5 km south of colony) and the lift bridge canal (4 km north of colony) was the foraging area used most frequently by radiotagged males during the chick-rearing period. Foraging patterns of transmittered males at Windermere Basin are similar to patterns of peak-nesting common terns, but differ from those of late-nesters, at a nearby colony (Port Colborne, Lake Erie). Differences between the foraging patterns of late-nesting terns at these colonies likely reflect differences in annual patterns of fish availability between the two locations. No relationship was found between foraging proficiency of adults and survival of offspring. Stochastic factors, such as predation by black-crowned nightherons (Nycticorax nycticorax) and adverse weather conditions during the early stages of chick rearing, may be more important determinants of common tern breeding success than parental quality or fish availability.

Nuclear-follower foraging associations among Characiformes fishes and Potamotrygonidae rays in clean waters environments of Teles Pires and Xingu rivers basins, Midwest Brazil

Por meio de observações sub e supra-aquáticas foram registradas associações alimentares do tipo nuclear-seguidor entre três espécies de peixes characiformes - Chalceus epakros, Hemiodus semitaeniatus e Hemiodus unimaculatus - e uma espécie de raia de água doce - Potamotrygon orbignyi - nas bacias dos rios Teles Pires e Xingu, no Centro-Oeste do Brasil. Os peixes teleósteos foram observados seguindo as raias quando estas revolviam o substrato à procura de invertebrados, formando discretas nuvens de sedimento. Essas situações atraíram os peixes que se aproximaram das raias para se alimentar de pequenas presas e outros tipos de alimentos expostos desta forma. Esse é um típico exemplo de relação comensal onde um participante é beneficiado enquanto o outro não é prejudicado e representa o segundo registro na literatura de associação alimentar do tipo nuclear-seguidor entre raias potamotrigonídeas e peixes teleósteos, demonstrando o potencial de estudos naturalísticos para a descoberta de novas interações envolvendo espécies de peixes de água doce.

Foraging by the omnivorous crab Pachygrapsus transversus affects the structure of assemblages on sub-tropical rocky shores

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)