Synopsis Fish species in many families and different trophic levels forage by following fishes and other animals. This interspecific foraging association was examined at an oceanic archipelago in the tropical West Atlantic. We recorded 27 reef fish species, two invertebrate species, and one turtle species playing the nuclear role, and 26 reef fish species acting as followers. The puddingwife wrasse following the spotted goatfish was the commonest foraging association recorded. The spotted goatfish was the nuclear fish that attracted the largest number of follower species (68% of the total number of follower species). The coney and the Noronha wrasse were the follower species that associated with the largest number of nuclear species (63 and 55% of the total number). About 20% of the reef fish species recorded in the archipelago engages in interspecific foraging associations. Substratum disturbance is a strong predictor for a fish displaying the nuclear role in the association, whereas the follower role may be predicted by carnivory. Nuclear species are diverse both in morphology and behaviour, and the nuclear role may be played either by fishes or other marine animals from invertebrates to turtles. Followers, on the other hand, comprise fishes only, which tend to display a more uniform feeding behaviour.
We studied the reef fish assemblage of eight reefs within the oceanic archipelago of Fernando de Noronha, off northeastern Brazil. In a total of 91 belt transects (20×2 m) we recorded 60 species from 28 families. The 25 most abundant species accounted for about 98% of all fish recorded in this study and most of these species are widely distributed in the Western Atlantic. The majority of fish counted were planktivores (37.0%), followed by mobile invertebrate feeders (28.5%), territorial herbivores (11.3%), roving herbivores (10.5%), omnivores (7.1%), macrocarnivores (6.5%) and sessile invertebrate feeders (0.03%). In terms of biomass, roving herbivores were the most representative (41.8%), followed by mobile invertebrate feeders (19.9%), macrocarnivores (14.3%), omnivores (14.0%), piscivores (8.3%), planktivores (1.4%), territorial herbivores (0.3%), and sessile invertebrate feeders (0.03%). Overall, density and biomass of fishes were positively correlated with coral cover and depth, and negatively correlated with wave exposure. These relationships are probably a response to the habitat complexity provided by the higher coral cover in deeper reefs (>10 m) of the archipelago or to the lower water turbulence below 10 m deep. Carnivores and mobile invertebrate feeders were mainly influenced by depth and non-consolidated substratum, planktivores and omnivores by wave exposure and herbivores by algal cover. Although our results suggest that habitat characteristics may play a role in determining the distribution of some fish species, we also found several habitat generalists, suggesting that the community is dominated by versatile species.
The influence of a substratum-disturbing forager, the spotted goatfish Pseudupeneus maculatus on the assemblage of its escorting, opportunistic-feeding fishes was examined at Fernando de Noronha Archipelago (tropical west Atlantic). Followers attracted to spotted goatfish foraging singly differed from followers of spotted goatfish foraging in groups in several characteristics. The larger the nuclear fish group, the greater the species richness and number of individuals of followers. Moreover, groups of foraging spotted goatfish attracted herbivores, not recorded for spotted goatfish foraging singly. The size of follower individuals increased with the size and the number of foraging spotted goatfish. The zoobenthivorous habits of the spotted goatfish and its ability to disturb a variety of soft substrata render it an important nuclear fish for several follower species of the reef fish assemblage at Fernando de Noronha.
Among reef fishes, it is common for "follower" individuals to accompany "nuclear" species and to feed on prey uncovered by their foraging. In this study, I examine the cues used by followers to find nuclear fish. A model of a ubiquitous nuclear fish was maintained immobile or moved to disturb the substratum and the number of fish species and individuals attracted was compared to control treatments. The results showed that: 1) bottom disturbance was the strongest attraction factor for follower reef fishes; 2) visual features of the nuclear also attracted follower reef fishes; 3) there was no evidence of an interaction between bottom disturbance and nuclear fish appearance in the attraction of followers, supporting the idea that both factors independently elicit following behaviour by reef fishes.
The goatfishes (Mullidae) include about 50 bottom-foraging fish species. The foraging activity of the yellow goatfish, Mulloidichthys martinicus, and the spotted goatfish, Pseudupeneus maculatus, was studied comparatively at Fernando de Noronha Archipelago, off coast of Northeast Brazil tropical West Atlantic. Pseudupeneus maculatus fed over a larger variety of substrate types, had lower feeding rate, roamed more per given time, spent less time in a feeding event, and displayed a more diverse repertoire of feeding modes than M. martinicus. The differences in the foraging activity and behaviour between the two species possibly minimize a potential resource overlap, as already recorded for other sympatric mullids. Pseudupeneus maculatus had lower feeding rate most likely because it feeds on larger items, and roamed over greater distance per time. Possibly this is because it foraged over a greater variety of substrate distributed over a larger area than that used by M. martinicus. Notwithstanding the overall morphological and behavioural similarity of goatfishes in general, they do differ in their substrate preferences and foraging activity, which indicates that these fishes should not be simply considered generalized bottom foragers.
The banded butterflyfish (Chaetodon striatus) from the tropical and subtropical western Atlantic is a territorial, diurnal forager on benthic invertebrates. It is usually seen moving singly or in pairs, a few meters above the sea floor. We studied the foraging activity of C. striatus on rocky reefs in southeastern Brazil. This fish spent about 11 h and 30 min per day on feeding activities, and preferred colonies of non-scleratinian anthozoans over sandy and rocky substrata while foraging. The lowest feeding rates were recorded in the early morning and late afternoon, but we found no further differences between feeding rates throughout the day. We also found no differences between the feeding rates of paired and single individuals.
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