Foraging plasticity in obligate corallivorous Melon butterflyfish across three recently bleached reefs

Because obligate corallivorous butterflyfish feed exclusively on coral polyps, they are particularly sensitive to changes in coral cover and its spatial distribution. To understand how such differences in coral cover influence obligate corallivores, we examined the densities and foraging behavior of Melon butterflyfish Chaetodon trifasciatus across three reefs in the Lakshadweep archipelago. These reefs suffered differential bleaching mortality after the 2010 El Niño Southern Oscillation, resulting in wide variation in coral cover and community composition. Despite these differences, C. trifasciatus were able to persist at similar densities across reefs. However, our analysis of high‐resolution video recordings of multiple focal fish revealed that time budgets, bite rates, and diet selectivity differed significantly. Fish in resource‐poor reefs spent more time moving between coral patches and less time foraging than ones in relatively resource‐rich reefs. We also found that fish in resource‐poor reefs had higher bite rates and were less selective in their foraging. Our results provide novel insights into how obligate corallivores cope with even large differences in resource availability. At a time when we are rapidly losing corals to repeated climate‐induced bleaching events, this flexibility may represent a critical mechanism that enables persistence of obligate corallivores in resource‐poor reefs, even if it does not guarantee longer‐term survival.     

Effects of different disturbance types on butterflyfish communities of Australia’s Great Barrier Reef

The effects of disturbances on coral reef fishes have been extensively documented but most studies have relied on opportunistic sampling following single events. Few studies have the spatial and temporal extent to directly compare the effects of multiple disturbances over a large geographic scale. Here, benthic communities and butterflyfishes on 47 reefs of the Great Barrier Reef were surveyed annually to examine their responses to physical disturbances (cyclones and storms) and/or biological disturbances (bleaching, outbreaks of crown-of-thorns starfish and white syndrome disease). The effects on benthic and butterflyfish communities varied among reefs depending on the structure and geographical setting of each community, on the size and type of disturbance, and on the disturbance history of that reef. There was considerable variability in the response of butterflyfishes to different disturbances: physical disturbances (occurring with or without biological disturbances) produced substantial declines in abundance, whilst biological disturbances occurring on their own did not. Butterflyfishes with the narrowest feeding preferences, such as obligate corallivores, were always the species most affected. The response of generalist feeders varied with the extent of damage. Wholesale changes to the butterflyfish community were only recorded where structural complexity of reefs was drastically reduced. The observed effects of disturbances on butterflyfishes coupled with predictions of increased frequency and intensity of disturbances sound a dire warning for the future of butterflyfish communities in particular and reef fish communities in general.     

Territoriality of the omnivorous butterflyfish, Chaetodon auriga (Chaetodontidae)

Butterflyfish research has focused on corallivorous species, with little attention paid to the other, equally abundant, omnivorous and planktivorous species. This study sought to explore the nature of ranging behaviour of the omnivorous butterflyfish, Chaetodon auriga, across two geographic regions. Pairs of C. auriga defended non-overlapping territories, ranging in area from 97.44 m² to 2281.64 m². Mean territory area varied among study sites while remaining consistent between geographic regions. Most of the variation in territory area was explained by variation in butterflyfish density, while damselfish density, total fish density, and percent sand cover inside territories were also correlated with territory area. Benthic assemblages of territories did not mirror study sites, suggesting that territory establishment is non-random. At all study sites, Chaetodon auriga exhibited significant selection in its patterns of habitat resource use. These results highlight the importance of research on generalist butterflyfishes as damage to coral reef ecosystems continues to threaten their specialist counterparts.     

Toxic coral gobies reduce the feeding rate of a corallivorous butterflyfish on Acropora corals

The obligate coral-dwelling gobiid genus Gobiodon inhabits Acropora corals and has developed various physiological, morphological and ethological adaptations towards this life habit. While the advantages of this coral-fish association are well documented for Gobiodon, possible fitness-increasing factors for the host coral are unknown. This study examines the influence of coral-dwelling gobies on the feeding behaviour of obligate corallivorous butterflyfishes. In an aquarium experiment using video observation, the corallivorous butterflyfish Chaetodon austriacus fed significantly less on corals inhabited by two Gobiodon species compared to unoccupied coral colonies of similar size. The more agonistic species G. histrio, which mostly displayed directed movements towards butterflyfishes, decreased butterflyfish bite rate by 62–98 % compared to uninhabited colonies. For Gobiodon sp. 3, which mostly displayed undirected movements in response to visits by C. austriacus, bite rate reduction was 64–68 %. The scale-less skin of Gobiodon spp. is covered by mucus that is toxic and multi-functional by reducing predation as well as affecting parasite attachment. A choice flume experiment suggests that the highly diluted skin mucus of Gobiodon spp. also functions as a corallivore repellent. This study demonstrates that Gobiodon spp. exhibit resource defence against coral-feeding butterflyfishes and also that coral colonies without resident Gobiodon suffer higher predation rates. Although the genus Gobiodon is probably a facultative corallivore, this study shows that by reducing predation on inhabited colonies by other fishes, these obligate coral-dwellers either compensate for their own fitness-decreasing impact on host colonies or live in a mutualistic association with them.     

An analysis of fish‐habitat associations on disturbed coral reefs: chaetodontid fishes in New Caledonia

A concordance analysis was used to study the simultaneous influence of several environmental data sets on chaetodontid (butterflyfish) distributions. This multivariate and multitable method enabled the correlation of three types of benthic characteristics (mineral substratum, coverage of structural species and large echinoderms) with butterflyfish abundances in two bays of the urban centre of Nouméa (New Caledonia). The first concordance axis was related to a gradient in the coverage of branching corals. This disturbance gradient compared damaged reef areas dominated by long‐spined sea urchins to areas with an extensive coverage of branching corals. The abundance of corallivorous chaetodontids was related to this gradient, supporting the view of corals as a food and shelter source for these fishes. The second concordance axis was interpreted as a gradient of heterogeneity in the coverage of benthic life‐forms. The abundance of omnivorous chaetodontids was related to this gradient. Thus, the concordance axes defined two key components of habitat structure that were related to the entire fish community structure.     

Coral mortality versus structural collapse as drivers of corallivorous butterflyfish decline

As climate change increasingly threatens biodiversity, identifying specific drivers of species loss as well as the attributes of species most vulnerable to climatic disturbances is a key challenge to ecologists and conservationists. Here we assess the effects of coral loss versus declines in structural complexity on obligate and facultative coral feeding butterflyfishes on coral reefs in the central and western Indian Ocean. In the inner Seychelles, the abundance of the obligate coral feeding group declined markedly in response to live coral mortality (r ² = 0.48), but showed no further decline with respect to erosion of the physical matrix of the reef. Conversely, the facultative feeding group showed no decline in response to live coral loss, reflecting their feeding versatility; however they did decline in response to structural erosion of the reef framework (r ² = 0.26). There were no significant changes in either obligate or facultative corallivore abundances at a reference location (Chagos archipelago), highlighting that butterflyfish populations are stable in the absence of habitat disturbance. While specialised coral dependant fishes are highly vulnerable to coral loss caused by climate-induced coral bleaching, the structural collapse of dead coral colonies may have significant, but more variable, impacts across a wide range of fishes. If conservation and mitigation planning are to be effective, there is a clear need to better understand the mechanisms of reef structural collapse and the dynamics of system recovery following large-scale disturbance.     

Great Barrier Reef butterflyfish community structure: the role of shelf position and benthic community type

The extent to which fish communities are structured by spatial variability in coral reef habitats versus stochastic processes (such as larval supply) is very important in predicting responses to sustained and ongoing habitat degradation. In this study, butterflyfish and benthic communities were surveyed annually over 15 years on 47 reefs (spanning 12° of latitude) of the Great Barrier Reef (GBR). Spatial autocorrelation in the structure of butterflyfish communities versus key differences in reef habitats was investigated to assess the extent to which the structure of these fish communities is influenced by habitat conditions. Benthic communities on each of the 47 reefs were broadly categorised as either: 1. Poritidae/Alcyoniidae, 2. mixed taxa, 3. soft coral or 4. Acropora-dominated habitats. These habitat types most reflected increases in water clarity and wave exposure, moving across the GBR shelf from coastal to outer-shelf environments. In turn, each habitat type also supported very distinct butterflyfish communities. Hard coral feeders were always the dominant butterflyfish species in each community type. However, the numerically dominant species changed according to habitat type, representing spatial replacement of species across the shelf. This study reveals clear and consistent differences in the structure of fish communities among reefs associated with marked differences in habitat structure.     

Corallivory in tubelip wrasses: diet,feeding and trophic importance

This paper describes a 2 month study of the patterns of abundance, feeding pressure, diet and feeding selectivity in corallivorous tubelip wrasses (Labridae), rarely studied, yet widespread and abundant group of corallivores on Indo‐Pacific coral reefs. The relative abundance and feeding pressure of corallivorous wrasses and butterflyfishes (Chaetodontidae) in Kimbe Bay, Papua New Guinea, were compared. Overall, tubelip wrasses were more than twice as abundant as corallivorous butterflyfishes and accounted for three times as many feeding bites on corals. The three most abundant tubelip wrasses (yellowtail tubelip Diproctacanthus xanthurus, Allen's tubelip Labropsis alleni and the tubelip wrasse Labrichthys unilineatus) were all obligate corallivores taking > 97% of bites from the surface of live corals. Labropsis alleni and D. xanthurus were highly selective, consuming preferred prey species in proportions significantly higher than expected given their availability. In contrast, L. unilineatus was fairly non‐selective and consumed most corals in direct accordance with their availability. As coral predators, tubelip wrasses are highly comparable to coral‐feeding butterflyfishes in the coral species consumed, range of dietary specialization and their reliance on live coral. Tubelip wrasses, however, may supersede butterflyfishes as the predominant corallivorous family in some Indo‐Pacific locations, and coral‐feeding tubelip wrasses are likely to be severely affected by coral decline.     

Specialist corallivores dominate butterflyfish assemblages in coral‐dominated reef habitats

This study examined the dietary habits and functional composition of butterflyfishes in the Chagos Archipelago, central Indian Ocean. Eighteen species of butterflyfishes were recorded in Chagos, including six obligate corallivores (Chaetodon bennetti, Chaetodon guttatissimus, Chaetodon meyeri, Chaetodon trifascialis, Chaetodon trifasciatus and Chaetodon zanzibarensis), five facultative corallivores (Chaetodon auriga, Chaetodon falcula, Chaetodon interruptus, Chaetodon kleinii and Chaetodon madagaskariensis), two non‐corallivores (Chaetodon lunula and Chaetodon xanthocephalus) and a further five species (Chaetodon citrinellus, Chaetodon lineolatus, Heimitaurichthys zoster, Heniochus monoceros and Forcipiger flavissimus), for which local dietary habits were not studied. There were marked differences in the abundance of butterflyfishes among sites and between reef zones, mostly associated with variation in abundance of scleractinian corals. Obligate coral‐feeding species (mostly C. trifascialis) dominated across all sites. This study suggests that coral feeding and high levels of dietary specialization contribute to high population‐level fitness among coral reef butterflyfishes. Despite being more vulnerable to habitat disturbances and coral loss, it appears likely that specialist coral‐feeding butterflyfishes are also much more resilient to occasional disturbances, and therefore dominate in a wide range of coral reef habitats.     

Mode of development and interspecific avian brood parasitism

Avian interspecific brood parasites differ considerably in theircommitment to parasitism; 87 species are obligate brood parasites,whereas 35 species are known to be facultative brood parasites.This variation is strongly related to mode of development. Obligateparasitism is found almost exclusively in altricial species,whereas facultative interspecific parasitism is predominantin precocial birds. We propose that the association betweenmode of development and form of parasitism reflects a fundamentaldifference between altricial and precocial birds in the relativebenefits of emancipation from parental care after laying. Weargue that altricial brood parasites obtain such a large increasein realized fecundity by avoiding the costs of parental carethat obligate parasitism is favored over facultative parasitism.In contrast, precocial brood parasites gain relatively littlein terms of increased fecundity via obligate parasitism, andmuch of this increase could potentially be gained by facultativeparasitism. Thus, obligate interspecific brood parasitism willnot be favored in precocial birds. Three factors influence thisdifference between altricial and precocial species: (1) altricialbirds have relatively more energy and nutrients with which tolay additional eggs, (2) altricial birds can produce more eggsfor the same amount of energy and nutrients, and (3) altricialbirds realize a greater relative gain in fecundity for eachadditional egg laid. We suggest further that facultative interspecificparasitism in birds may originate simply through a carry overof intraspecific parasitism; 29 of 33 facultative interspecificparasites also parasitize conspecifics. Facultative parasitismof other species would provide a greater range of potentialhost nests and could be maintained as an evolutionarily stableend point by the same mechanisms that maintain intraspecificbrood parasitism. [Behav Ecol 1991;2:309–318]     

Community structure and biogeography of shore fishes in the Gulf of Aqaba,Red Sea

Shore fish community structure off the Jordanian Red Sea coast was determined on fringing coral reefs and in a seagrass-dominated bay at 6 m and 12 m depths. A total of 198 fish species belonging to 121 genera and 43 families was recorded. Labridae and Pomacentridae dominated the ichthyofauna in terms of species richness and Pomacentridae were most abundant. Neither diversity nor species richness was correlated to depth. The abundance of fishes was higher at the deep reef slope, due to schooling planktivorous fishes. At 12 m depth abundance of fishes at the seagrass-dominated site was higher than on the coral reefs. Multivariate analysis demonstrated a strong influence on the fish assemblages by depth and benthic habitat. Fish species richness was positively correlated with hard substrate cover and habitat diversity. Abundance of corallivores was positively linked with live hard coral cover. The assemblages of fishes were different on the shallow reef slope, deep reef slope and seagrass meadows. An analysis of the fish fauna showed that the Gulf of Aqaba harbours a higher species richness than previously reported. The comparison with fish communities on other reefs around the Arabian Peninsula and Indian Ocean supported the recognition of an Arabian subprovince within the Indian Ocean. The affinity of the Arabian Gulf ichthyofauna to the Red Sea is not clear. Received in revised form: 2 November 2001 Electronic Publication     

Tropical Fishes Dominate Temperate Reef Fish Communities within Western Japan

Climate change is resulting in rapid poleward shifts in the geographical distribution of tropical and subtropical fish species. We can expect that such range shifts are likely to be limited by species-specific resource requirements, with temperate rocky reefs potentially lacking a range of settlement substrates or specific dietary components important in structuring the settlement and success of tropical and subtropical fish species. We examined the importance of resource use in structuring the distribution patterns of range shifting tropical and subtropical fishes, comparing this with resident temperate fish species within western Japan (Tosa Bay); the abundance, diversity, size class, functional structure and latitudinal range of reef fishes utilizing both coral reef and adjacent rocky reef habitat were quantified over a 2 year period (2008–2010). This region has undergone rapid poleward expansion of reef-building corals in response to increasing coastal water temperatures, and forms one of the global hotspots for rapid coastal changes. Despite the temperate latitude surveyed (33°N, 133°E) the fish assemblage was both numerically, and in terms of richness, dominated by tropical fishes. Such tropical faunal dominance was apparent within both coral, and rocky reef habitats. The size structure of the assemblage suggested that a relatively large number of tropical species are overwintering within both coral and rocky habitats, with a subset of these species being potentially reproductively active. The relatively high abundance and richness of tropical species with obligate associations with live coral resources (i.e., obligate corallivores) shows that this region holds the most well developed temperate-located tropical fish fauna globally. We argue that future tropicalisation of the fish fauna in western Japan, associated with increasing coral habitat development and reported increasing shifts in coastal water temperatures, may have considerable positive economic impacts to the local tourism industry and bring qualitative changes to both local and regional fisheries resources.     

Relationships between butterflyfish (Chaetodontidae) feeding rates and coral consumption on the Great Barrier Reef

This study explored differences in the feeding rate among 20 species of coral reef butterflyfishes (Chaetodontidae) from Lizard Island, Great Barrier Reef. Feeding rate, measured as bites per minute (b.p.m.), varied between 2.98 ± 0.65 and 12.29 ± 0.27 (mean ± SE) according to species and was positively related to the proportional consumption of coral (r ² = 0.40, n = 20, P < 0.01), independent of phylogeny (standardised independent contrasts r ² = 0.29, n = 19, P < 0.05). All species fed actively throughout the day, with obligate corallivores having a higher feeding rate at all times than either facultative corallivores or non-corallivores. The feeding rate of the obligate corallivores was also highest during the middle of the day. For eight of the species for which data was available, there was a positive correlation between bite rate and competitive dominance (r = 0.71, P < 0.05). Chaetodon ephippium was the only species for which the feeding rate of pairs was higher than for solitary individuals.     

Small-scale environmental variation influences whether coral-dwelling fish promote or impede coral growth

Mutualistic symbioses are ubiquitous in nature and facilitate high biodiversity and productivity of ecosystems by enhancing the efficiency of energy and nutrient use within ecological communities. For example, small groups of fish that inhabit coral colonies in reef ecosystems potentially enhance coral growth through defense from coral predators, aeration of coral tissue and nutrient provisioning. This study examines whether the prevalence and consequences of fish-coral interactions vary among sites with different environmental conditions in a coral reef lagoon, using the humbug damselfish Dascyllus aruanus and its preferred coral host Pocillopora damicornis as a study system. Using a field experiment, we tested the site-specific effects of D. aruanus on coral growth, and show that the cost-benefit ratio for corals hosting fish varies with local environmental variation. Results of this study also demonstrate that fish prefer to inhabit coral colonies with particular branch-spacing characteristics, and that the local abundance of D. aruanus influences the proportion of coral colonies within a site that are occupied by fish rather than increasing the number of fish per colony. We also show that corals consistently benefit from hosting D. aruanus via defense from predation by corallivorous butterflyfish, regardless of local environmental conditions. These findings highlight the need to consider the potential for multiple scale- and state-dependent interaction effects when examining the ecology of fish-coral associations. We suggest that fluctuating cost-benefit ratios for species interactions may contribute to the maintenance of different colony phenotypes within coral populations.     

Draft genome of an iconic Red Sea reef fish,the blacktail butterflyfish (Chaetodon austriacus): current status and its characteristics

Butterflyfish are among the most iconic of the coral reef fishes and represent a model system to study general questions of biogeography, evolution and population genetics. We assembled and annotated the genome sequence of the blacktail butterflyfish (Chaetodon austriacus), an Arabian region endemic species that is reliant on coral reefs for food and shelter. Using available bony fish (superclass Osteichthyes) genomes as a reference, a total of 28 926 high‐quality protein‐coding genes were predicted from 13 967 assembled scaffolds. The quality and completeness of the draft genome of C. austriacus suggest that it has the potential to serve as a resource for studies on the co‐evolution of reef fish adaptations to the unique Red Sea environment, as well as a comparison of gene sequences between closely related congeneric species of butterflyfish distributed more broadly across the tropical Indo‐Pacific.     

Habitat preferences of a corallivorous reef fish: predation risk versus food quality

Many animals preferentially select a habitat from a range of those potentially available. However, the consequences of these preferences for distribution and abundance, and the underlying basis of habitat preferences are often unknown. The present study, conducted at Great Keppel Island, Australia, examined how distribution and abundance of an obligate corallivorous filefish, Oxymonacanthus longirostris, relates to coral architecture and diversity. The main drivers of the distribution and abundance of O. longirostris among reefs were coral species richness and availability of branching coral. Feeding territories had a higher percentage of Acropora coral than surrounding habitat. In addition, feeding territories had a higher percentage of the structurally important branching coral, Acropora nobilis, and a primary prey species, Acropora millepora. A series of pair-wise choice experiments in which both structural complexity and coral tissue quality were independently manipulated showed that habitat choice was primarily based on structural complexity and shelter characteristics. In addition, the choice for the preferred coral (A. nobilis) was stronger in the presence of a piscivorous fish. These results indicate that species-diverse coral habitats, which provide sufficient structural complexity along with nutritionally important prey, are essential for population persistence of this small, corallivorous reef fish.     

The influence of coral reef benthic condition on associated fish assemblages

Accumulative disturbances can erode a coral reef's resilience, often leading to replacement of scleractinian corals by macroalgae or other non-coral organisms. These degraded reef systems have been mostly described based on changes in the composition of the reef benthos, and there is little understanding of how such changes are influenced by, and in turn influence, other components of the reef ecosystem. This study investigated the spatial variation in benthic communities on fringing reefs around the inner Seychelles islands. Specifically, relationships between benthic composition and the underlying substrata, as well as the associated fish assemblages were assessed. High variability in benthic composition was found among reefs, with a gradient from high coral cover (up to 58%) and high structural complexity to high macroalgae cover (up to 95%) and low structural complexity at the extremes. This gradient was associated with declining species richness of fishes, reduced diversity of fish functional groups, and lower abundance of corallivorous fishes. There were no reciprocal increases in herbivorous fish abundances, and relationships with other fish functional groups and total fish abundance were weak. Reefs grouping at the extremes of complex coral habitats or low-complexity macroalgal habitats displayed markedly different fish communities, with only two species of benthic invertebrate feeding fishes in greater abundance in the macroalgal habitat. These results have negative implications for the continuation of many coral reef ecosystem processes and services if more reefs shift to extreme degraded conditions dominated by macroalgae.     

Prehistoric fisheries in the Caribbean

We studied faunal remains from archaeological sites on five Caribbean islands, each with an early (1,850-1,280 years B.P.) and late (1,415-560 years B.P.) occupation. On each of these islands (Puerto Rico, St. Thomas, St. Martin, Saba, and Nevis), the mean size of reef fishes in the faunal remains declined from the early to the late occupation. The large samples from sites on St. Thomas and Nevis allowed examination of the size distribution of individual taxa. Samples of obligate reef fishes (Scaridae, Acanthuridae, Lutjanidae, and Serranidae) showed large reductions in size between the early and late occupations. Samples of facultative reef fishes (Carangidae and Clupeidae) showed little change in size frequency distribution. The percentage of estimated reef fish biomass in the total aquatic faunal record sharply declined in the samples from four of the islands, while on Nevis there was a slight increase. The mean trophic level of reef fishes declined from the early to the late occupations on each island. Together these patterns suggest that populations of reef fishes adjacent to occupation sites on these islands were heavily exploited in prehistoric times. Such exploitation resulted in shifts in size structure and species composition among the reef fish fauna. On some islands the decline in reef fish resources corresponded with a shift towards greater exploitation of pelagic species.     

Kenyan coral reef lagoon fish: effects of fishing,substrate complexity,and sea urchins

Population density, number of species, diversity, and species-area relationships of fish species in eight common coral reef-associated families were studied in three marine parks receiving total protection from fishing, four sites with unregulated fishing, and one reef which recently received protection from fishing (referred to as a transition reef). Data on coral cover, reef topographic complexity, and sea urchin abundance were collected and correlated with fish abundance and species richness. The most striking result of this survey is a consistent and large reduction in the population density and species richness of 5 families (surgeonfish, triggerfish, butterflyfish, angelfish, and parrotfish). Poor recovery of parrotfish in the transition reef, relative to other fish families, is interpreted as evidence for competitive exclusion of parrotfish by sea urchins. Reef substrate complexity is significantly associated with fish abundance and diversity, but data suggest different responses for protected versus fished reefs, protected reefs having higher species richness and numbers of individuals than unprotected reefs for the same reef complexity. Sea urchin abundance is negatively associated with numbers of fish and fish species but the interrelationship between sea urchins, substrate complexity, coral cover, and management make it difficult to attribute a set percent of variance to each factor-although fishing versus no fishing appears to be the strongest variable in predicting numbers of individuals and species of fish, and their community similarity. Localized species extirpation is evident for many species on fished reefs (for the sampled area of 1.0 ha). Fifty-two of 110 species found on protected reefs were not found on unprotected reefs.     

Diet of combtooth blennies (Blenniidae) in Kochi and Okinawa,Japan

Combtooth blennies (Blenniidae) have a wide array of diets, including coral polyps, fish scales/mucus/rays, and detritus. However, there is limited knowledge about the diet of some species. We described the diet of 30 blenny species from subtidal, intertidal, or supralittoral habitats in Kochi (warm temperate) and Okinawa (tropical), Japan. Cluster analysis of dietary overlap grouped blennies into seven feeding groups: omnivores, herbivores, detritivores, molluscivores, corallivores, worm-like invertebrate feeders, and fish mucus/scale/ray feeders. The largest cluster, detritivores, contained 17 blenny species from all habitat zones and both climate zones. Our findings suggest that closely related species fill similar feeding niches and detritus is an important component of many blenny diets.