Varying responses of herbivorous and invertebrate-feeding fishes to macroalgal reduction on a coral reef

 The consequences of macroalgal overgrowth on reef fishes and means to reverse this condition have been little explored. An experimental reduction of macroalgae was conducted at a site in the Watamu Marine National Park in Kenya, where a documented increase in macroalgal cover has occurred over the last nine years. In four experimental 10 m by 10 m plots, macroalgae were greatly reduced (fleshy algal cover reduced by 84%) by scrubbing and shearing, while four similar plots acted as controls. The numerical abundance in all fish groups except wrasses and macroalgal-feeding parrotfishes (species in the genera Calotomus and Leptoscarus) increased in experimental algal reduction plots. Algal (Sargassum) and seagrass (Thalassia) assays, susceptible to scraping and excavating parrotfishes, were bitten more frequently in the algal reduction plots one month after the manipulation. Further, surgeonfish (Acanthurus leucosternon and A. nigrofuscus) foraging intensity increased in these algal reduction plots. The abundance of triggerfishes increased significantly in experimental plots relative to control plots, but densities remained low, and an index of sea urchin predation using tethered juvenile and adult Echinometra mathaei showed no differences between treatments following macroalgal reduction. Dominance of reefs by macrofleshy algae appears to reduce the abundance of fishes, mostly herbivores and their rates of herbivory, but also other groups such as predators of invertebrates (triggerfishes, butterflyfishes and angelfishes). Accepted: 2 February 1999     

Hyperspectral discrimination of coral reef benthic communities in the western Caribbean

Determining a subset of wavelengths that best discriminates reef benthic habitats and their associated communities is essential for the development of remote sensing techniques to monitor them. This study measured spectral reflectance from 17 species of western Caribbean reef biota including coral, algae, seagrasses, and sediments, as well as healthy and diseased coral. It sought to extend the spectral library of reef-associated species found in the literature and to test the spectral discrimination of a hierarchy of habitats, community groups, and species. We compared results from hyperspectral reflectance and derivative datasets to those simulated for the three visible multispectral wavebands of the IKONOS sensor. The best discriminating subset of wavelengths was identified by multivariate stepwise selection procedure (discriminant function analysis). Best discrimination at all levels was obtained using the derivative dataset based on 6–15 non-contiguous wavebands depending on the level of the classification, followed by the hyperspectral reflectance dataset which was based on as few as 2–4 non-contiguous wavebands. IKONOS wavebands performed worst. The best discriminating subset of wavelengths in the three classification resolutions, and particularly those of the medium resolution, was in agreement with those identified by Hochberg and Atkinson (2003) and Hochberg et al. (2003) for reef communities worldwide. At all levels of classification, reflectance wavebands selected by the analysis were similar to those reported in recent studies carried out elsewhere, confirming their applicability in different biogeographical regions. However the greater accuracies achieved using the derivative datasets suggests that hyperspectral data is required for the most accurate classification of reef biotic systems.     

Coral reef fish communities: a compromise view

Synopsis The fishes associated with coral reefs offer excellent opportunities for the study of the factors that determine the species composition of complex, highly interactive communities. Amenable to sampling and direct observations many patch reefs are small enough to be studied as entire units and yet diverse enough to include a wide range of interactions.Coral reef fishes appear to be highly specialized in morphology, color, behavior and life cycles and yet colonization experiments and repeated censusing have shown a surprisingly high variation in the fishes that are associated in similar habitats or in the same habitat at different times. This has led to two different views: (i) the chaos view that holds that the species composition is due to random factors and chance colonization, and (ii) the order view that resource sharing adaptations determine which species can live together.This paper reviews some of the obvious adaptations of reef fishes. An examination of the maximum sizes reached by infaunal reef dwellers shows that the largest individual of each species differs by a constant proportion from the next larger and next smaller species. This suggests growth limitations by interspecies competition. A hypothetical model showing how this might work is offered and it is shown that as long as there are more species than can be accommodated in the community at any one time the number of combinations is great enough to give the appearance of randomness even though the individual species may have precise environmental requirements.This paper forms part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Hellman, ed.).     

Spectral discrimination of coral reef benthic communities

Effective identification and mapping of coral reef benthic communities using high-spatial and -spectral resolution digital imaging spectrometry requires that the different communities are distinguishable by their spectral reflectance characteristics. In Kaneohe Bay, Oahu, Hawaii, USA, we collected in situ a total of 247 spectral reflectances of three coral species (Montipora capitata, Porites compressa, Porites lobata), five algal species (Dictyosphaeria cavernosa, Gracilaria salicornia, Halimeda sp., Porolithon sp., Sargassum echinocarpum) and three sand benthic communities (fine-grained carbonate sand, sand mixed with coral rubble, coral rubble). Major reflectance features were identified by peaks in fourth derivative reflectance spectra of coral (at 573, 604, 652, 675 nm), algae (at 556, 601, 649 nm) and sand (at 416, 448, 585, 652, 696 nm). Stepwise wavelength selection and linear discriminant function analysis revealed that spectral separation of the communities is possible with as few as four non-contiguous wavebands. These linear discriminant functions were applied to an airborne hyperspectral image of a patch reef in Kaneohe Bay. The results demonstrate the ability of spectral reflectance characteristics, determined in situ, to discriminate the three basic benthic community types: coral, algae and sand. Accepted: 12 January 2000     

Assessing the magnitude of intra- and interspecific competition in two coral reef fishes

Spatial associations of seven herbivore species in the Kruger National Park, South Africa, are analyzed using a new technique, Correlative Coherence Analysis (CoCA). CoCA is a generalization of the concept of correlation to more than two sequences of numbers. Prior information on the feeding ecology and metabolic requirements of these species is used to contrast spatial scales at which hypothesized guild aggregation or competition occurs. These hypotheses are tested using 13 years of aerial census data collected during the dry season. Our results are consistent with the hypothesis that distributions of large and small species of the same feeding type (i.e., grazers and browsers) overlap in potentially resource-rich areas, but have lower similarity values across all areas because the higher tolerance of large species for low quality foods results in a more even spatial distribution of large species compared to small species.     

Response of triassic reef coral communities to sea-level fluctuations, storms and sedimentation: Evidence from a spectacular outcrop (Adnet, Austria)

Summary The Upper Rhaetian coral limestone of Adnet, southeast of Salzburg Austria has been repeatedly referred to as one of the most spectacular examples of an ancient ‘autochthonous’ coral reef structure. The ‘Tropfbruch’ quarry is probably the best outcrop for interpreting the distributional patterns of biotic successions and communities of a late Triassic patch reef. Our study is based on the interpretation of a) outcrop photographs, b) reef maps resulting from quadrat transects, and c) the analysis of quantitative data describing the distribution and frequency of reef organisms and sediment. A new methodological approach (combination of reef mapping and photo-transects) is used to obtain quantitative field data which can be compared in greater detail with data from modern coral reefs investigated by corresponding quantitative surveys. Three unconformities and three well-defined ‘reef growth stages’ reflecting the vertical and lateral development of the reef structure were differrentiated using transects: Stage 1, representing the reef growth optimum, is characterized by laterally differentiated coral reef knobs with corals in growth position. Criteria supporting this interpretation are the extraordinary size of the corals, their preservation in situ and the great thickness of this interval. The massive coralPamiroseris grew under higher energy conditions at the rim of the reef knob, whereas branchingRetiophyllia colonies preferred less agitated water in the center. Vertical changes are reflected by an increase in frequency of the dasycladacean algaDiplopora adnetensis and by the decreasing size ofRetiophyllia. These sedimentological and biological criteria together with the unconformity above indicate a fall in the sea level as a major control mechanism. Stage 2, separated from stage 1 by an unconformity caused by partial subaerial exposure and karstification, is characterized by vertically stacked coral successions with diverse reef debris. Facies heterogeneity is reflected by differences in the diversity, taphonomy and packing density of reef-building organisms as well as by differences in sediment input from the platform. Water depths and accommodation space were lower, therefore minor sea level fluctuations had a stronger effect on the biotic composition. The high percentage of coral debris and corals reworked by storms and the increase in the input of platform sediment led to a reduction of reef growth. Stage 3, again separated at the base by an unconformity associated with karstification, is characterized by bioclastic sediments with isolated reefbuilders forming a level-bottom community. The distribution of different coral morphotypes suggests that sea level fluctuations were not the only controlling factor. Variations in the substrate were caused by differences in the input of platform sediment. The three-step development seen in Adnet documents the response of low-diverse coral associations to variations caused by small-scale sea level changes, storm activity and sedimentation. The vertical changes in reef community structures correspond to a sequence of ‘allogenic replacements’. The Adnet reef structure should not be regarded as a general model of Alpine Upper Rhaetian reefs, because of the particular setting of the patch reef. Only the ‘capping beds’ of the Upper Rhaetian Reef Limestone of the Steinplatte exhibit criteria similar to Adnet. Potential modern analogues of features seen in the coral communities of Adnet are the internal structure of theRetiophyllia thickets, the key role of branching corals within the communities, the scattered distribution and low and even diversity of corals subsequent to breaks in settlement, segration patterns of corals indicating ‘contact avoidance’, toppling of large coral colonies by intensive boring, and decreasing coral coverage from deeper and sheltered settings to more shallower water depths.     

Global assessment of the status of coral reef herbivorous fishes: evidence for fishing effects

On coral reefs, herbivorous fishes consume benthic primary producers and regulate competition between fleshy algae and reef-building corals. Many of these species are also important fishery targets, yet little is known about their global status. Using a large-scale synthesis of peer-reviewed and unpublished data, we examine variability in abundance and biomass of herbivorous reef fishes and explore evidence for fishing impacts globally and within regions. We show that biomass is more than twice as high in locations not accessible to fisheries relative to fisheries-accessible locations. Although there are large biogeographic differences in total biomass, the effects of fishing are consistent in nearly all regions. We also show that exposure to fishing alters the structure of the herbivore community by disproportionately reducing biomass of large-bodied functional groups (scraper/excavators, browsers, grazer/detritivores), while increasing biomass and abundance of territorial algal-farming damselfishes (Pomacentridae). The browser functional group that consumes macroalgae and can help to prevent coral–macroalgal phase shifts appears to be most susceptible to fishing. This fishing down the herbivore guild probably alters the effectiveness of these fishes in regulating algal abundance on reefs. Finally, data from remote and unfished locations provide important baselines for setting management and conservation targets for this important group of fishes.     

Behavioral thermoregulatory abilities of tropical coral reef fishes: a comparison with temperate freshwater and marine fishes

Synopsis Eight species in six different families of tropical marine reef fishes from the Indo-West Pacific region (Naso lituratus, Zebrasoma.flavescens, Balistes fuscus, B. vidua, Forcipiger longirostris, Echidna zebra, Cromileptes altivelis, Canthigaster jactator) were tested for ability to thermoregulate behaviorally in electronic shuttleboxes. All of these species preferred mean temperatures between 20 and 30°C, but differed considerably in thermoregulatory precision. All species avoided lethal high or low temperatures (i.e., they did not die during the tests), and some species thermoregulated as precisely as temperate species. Some temperate species prefer higher temperatures (above 30°C) than do these tropical reef species.     

Patterns and trends in coral reef macroalgae browsing: a review of browsing herbivorous fishes of the Indo-Pacific

Browsing fishes have been identified as an important component of coral reef resilience, because in contrast to other herbivorous fishes they are able to feed on established macroalgae. Climate change and local anthropogenic impacts have contributed to phase shifts in many coral reefs from coral to macroalgae dominance, and recent research suggests the potential ability of browsers to reverse such phase shifts. However, there is high variation among studies and some contradicting findings exist. Here, we review the relevant literature to assemble a list of species currently known to contribute to browsing in the Indo-Pacific. Furthermore, we identify spatial and temporal patterns, outline factors influencing browsing, and discuss the probability of phase shift reversal. We formulate research recommendations addressing the identified gaps in knowledge about the interactions of browsing fishes and their environment. To date, 37 species of fishes have been observed consuming macroalgae in the Indo-Pacific. The most important groups are the family Siganidae, the subfamily Scarinae (Labridae), and the subfamily Nasinae (Acanthuridae). Browsing species vary between studies depending on location, season and macroalgae species examined. Several influencing factors, such as structural complexity, palatability of macroalgae and ecosystem connectivity have been suggested to cause these discrepancies. The most promising avenues for future research are the effect of structural complexity, the importance of mobile link species and influences of food availability on the selectivity of browsing species. Increasing our knowledge in these fields will provide a better basis for successful management strategies directed at increasing the resilience of coral reefs.     

The adaptive significance of larval dispersal in coral reef fishes

Synopsis Coral reef fishes almost universally disperse over relatively great distances during a pelagic larval phase. Barlow (1981) suggested that this dispersal is adaptive because adult fishes inhabit a patchy, uncertain environment. This reiterated an older idea that the random extinction of local populations necessarily favours dispersal, since ultimately all populations of non-dispersers will disappear. Whereas this view is based on adult survival, we emphasize a less frequent view that substantial larval dispersal may be adaptive when offspring experience patchy and unpredictable survival in the pelagic habitat. We do not address the question of why these animals ‘broadcast’ rather than ‘brood’, but suggest that species committed to pelagic offspring will be under selection to disperse siblings to spread the risk of failure among members of a cohort. Our arguments are supported by a heuristic computer simulation.     

Water flow controls distribution and feeding behavior of two co-occurring coral reef fishes: I. Field measurements

The interaction of flowing water with reef topography creates a continuum of flow microhabitats that can alter species distributions directly via transport of organisms or propagules, or indirectly by modulating the availability of critical resources. To examine how water flow affects the distribution and feeding performance of two species of planktivorous tube blennies (Chaenopsidae), flow speed and turbulence were measured within the feeding areas of Acanthemblemaria spinosa and A. aspera at three sites within Glover’s Reef, Belize. Although co-occurring, A. spinosa occupies topographically high locations (e.g., upright coral skeletons) while A. aspera occupies topographically low shelters in the coral pavement. Boundary layer theory predicts that A. spinosa should experience higher flow (and a higher flux of planktonic food) relative to A. aspera; however, complex topography and oscillatory flow require that this prediction is tested directly in the field. Within each site, the flow experienced by A. spinosa was, indeed, faster and more turbulent than that experienced by A. aspera at site-specific intermediate wave heights. When waves were small, gentle velocity gradients produced similar flows for the two species. When waves were high, flow was uniformly fast through the water column due to thinning of the benthic boundary layer. Plankton availability was similar for the species, with the exception of a greater abundance of harpacticoid copepods at the shelters of A. aspera. Quantitative behavioral observations suggest that the foraging strategies employed by the two fishes exploit the prevailing hydrodynamic conditions. For example, A. spinosa, the stronger swimmer of the two, attacks nearly 100% of the time in the water column where it can exploit the higher flux of plankton associated with faster flows, while A. aspera attacks primarily toward the reef surface where currents are likely to be slower and it can exploit more abundant benthic prey. Communicated by Biology Editor Dr. Mark McCormick     

Nocturnal relocation of adult and juvenile coral reef fishes in response to reef noise

Juvenile and adult reef fishes often undergo migration, ontogenic habitat shifts, and nocturnal foraging movements. The orientation cues used for these behaviours are largely unknown. In this study, the use of sound as an orientation cue guiding the nocturnal movements of adult and juvenile reef fishes at Lizard Island, Great Barrier Reef was examined. The first experiment compared the movements of fishes to small patch reefs where reef noise was broadcast, with those to silent reefs. No significant responses were found in the 79 adults that were collected, but the 166 juveniles collected showed an increased diversity each morning on the reefs with broadcast noise, and significantly greater numbers of juveniles from three taxa (Apogonidae, Gobiidae and Pinguipedidae) were collected from reefs with broadcast noise. The second experiment compared the movement of adult and juvenile fishes to reefs broadcasting high (>570 Hz), or low (<570 Hz) frequency reef noise, or to silent reefs. Of the 122 adults collected, the highest diversity was seen at the low frequency reefs; and adults from two families (Gobiidae and Blenniidae) preferred these reefs. A similar trend was observed in the 372 juveniles collected, with higher diversity at the reefs with low frequency noises. This preference was seen in the juvenile apogonids; however, juvenile gobiids were attracted to both high and low sound treatments equally, and juvenile stage Acanthuridae preferred the high frequency noises. This evidence that juvenile and adult reef fishes orientate with respect to the soundscape raises important issues for management, conservation and the protection of sound cues used in natural behaviour.     

The role of microhabitat preference and social organization in determining the spatial distribution of a coral reef fish

Both habitat preferences and social organization can influence the spatial distributions of individuals. We explored effects of individual behavior and social organization on distributions of arc-eye hawkfish (Paracirrhites arcatus) in lagoons of French Polynesia. Analysis of habitat selectivity data obtained during surveys revealed that the most highly preferred microhabitat of arc-eye hawkfish was large Pocillopora coral with an open branching morphology. However, such corals were rare and most hawkfish occupied smaller, less preferred Pocillopora. Indeed, total abundance of Pocillopora explained nearly two thirds of the lagoon-wide variation in abundance of hawkfish and the derived relationship between the numbers of hawkfish and Pocillopora predicted 86% of the spatial variation in hawkfish abundance during subsequent surveys. In contrast, large, open-branched Pocillopora explained little of the spatial variation in abundance of hawkfish. Individual behavior and social organization significantly impacted use of the most highly preferred Pocillopora. During a colonization experiment set up outside hawkfish home ranges, all colonizers resided on the most highly preferred corals. Following addition of the most highly preferred Pocillopora corals to areas occupied by hawkfish, only the largest, socially dominant individuals obtained access to added corals, spending significantly more time and increasing both aggressive acts and prey attacks from these substrates. These results illustrate the importance of understanding the modulating effects of social behavior on habitat use and explain why most hawkfish individuals do not occupy their most preferred microhabitat type.     

The behavioral ecology of three Indian Ocean surgeonfishes (Acanthurus lineatus,A. leucosternon and Zebrasoma scopas): their feeding strategies,and social and mating systems

Synopsis The relationship between the morphology, feeding strategies and social and mating systems of three surgeonfishes was investigated. Adults of each defend feeding territories, intra-and interspecifically. The largest species, because of its morphological limitation, relies on food that has to be defended against many other species. It forms large colonies in which fishes singly defend small territories containing high standing crop algal mats. Colony formation is a mechanism by which the efficiency and effectiveness of interspecific territory defense is increased. The smallest species, because of its morphological adaptations, is able to rely most on food that other species cannot efficiently exploit. It forms pairs that defend large territories containing a thin algal mat. It is restricted to the poorest quality habitat by the aggressive activities of more dominant species. The third species, which also forms pairs, has an intermediate feeding strategy. The local coexistence of these three and other surgeonfishes results from a combination of (i) their partitioning both habitat and food resources, and (ii) the populations of two of the most dominant species apparently being below the carrying capacity. Territoriality and the absence of parental care facilitates pair formation in surgeonfishes. Permanently territorial species usually form pairs. The colonial species does not form pairs because the colonial habit facilitates interference of males in each other's spawnings.     

The evolution of complex brains and behaviors in African cichlid fishes

In this review, I explore the effects of both social organization and the physical environment, specifically habitat complexity, on the brains and behavior of highly visual African cichlid fishes, drawing on examples from primates and birds where appropriate. In closely related fishes from the monophyletic Ectodinii clade of Lake Tanganyika, both forces influence cichlid brains and behavior. Considering social influences first, visual acuity differs with respect to social organization (monogamy versus polygyny). Both the telencephalon and amygdalar homologue, area Dm, are larger in monogamous species. Monogamous species are found to have more vasotocin-immunoreactive cells in the preoptic area of the brain. Habitat complexity also influences brain and behavior in these fishes. Total brain size, telencephalic and cerebellar size are positively correlated with habitat complexity. Visual acuity and spatial memory are enhanced in cichlids living in more complex environments. However habitat complexity and social forces affect cichlid brains differently. Taken together, our field data and plasticity data suggest that some of the species-specific neural effects of habitat complexity could be the consequence of the corresponding social correlates. Environmental forces, however, exert a broader effect on brain structures than social ones do, suggesting allometric expansion of the brain structures in concert with brain size and/or co-evolution of these structures [Current Zoology 56 (1): 144-156 2010].     

Parasites of recruiting coral reef fish larvae in New Caledonia

Recruiting coral reef fish larvae from 38 species and 19 families from New Caledonia were examined for parasites. We found 13 parasite species (Platyhelminthes: Monogenea, Cestoda and Trematoda) but no acanthocephalan, crustacean or nematode parasites. Over 23% of individual fish were infected. Didymozoid metacercariae were the most abundant parasites. We conclude that most of the parasites are pelagic species that become ‘lost’ once the fish larvae have recruited to the reef. Larval coral reef fish probably contribute little to the dispersal of the parasites of the adult fish so that parasite dispersal is more difficult than that of the fish themselves.     

Ocean current variability and the spawning season of Hawaiian reef fishes

Synopsis The spawning patterns of four Hawaiian reef fishes with similar reproductive habits, but different biogeographic distributions were studied from September 1980 to October 1981. Two species are Hawaiian endemics [Centropyge potteri (Pomacanthidae) and Chaetodon multicinctus (Chaetodontidae)] and the other two have pan-tropical Pacific distributions [Ctenochaetus strigosus and Zebrasoma flavescens (Acanthuridae)]. All showed increased spawning activity from January to July, the prevalent pattern among a majority of coastal marine fishes in Hawaii. Environmental correlatives to the period of peak reproduction include cycles of (a) daylight length and temperature which probably function as proximate cues and (b) seasonal variations in ocean current patterns which may ultimately affect survival of larvae and dispersal. Peak reproduction takes place during months when (1) mesoscale eddies most likely occur, and (2) the probability is greatest of drifting objects remaining near the islands.     

The structure of a nearshore fish community of Western Australia: diel patterns and the habitat role of limestone reefs

Synopsis Extensive limestone reefs are a characteristic feature of much of the coastline of Western Australia, and potentially represent a major habitat feature influencing the structure of the coastal fish community. The structure and temporal dynamics of the fish fauna and its relationships to nearshore patch reefs and surrounding habitat near Dongara, Western Australia, were examined using (1) diel gill-netting and (2) quantitative rotenone sampling of enclosed areas of substratum. Long-term and day-to-day variability of the fauna was low. Dominant species of gill-net collections were either associated with reefs or occurred in similar abundances at both reefs and surrounding sand/seagrass flats. The overall abundance, number of species and biomass of netted fishes was higher around reefs. Rotenone collections of the more sedentary species showed a similar pattern, but suggested, however, that a simple reef versus surrounding sand and seagrass habitat comparison is complicated by the canopy-forming seagrass Amphibolis that occurs on reef tops. Time of day had an important effect on overall fish abundance and number of species, with peaks occurring at crepuscular periods. This reflected dusk and dawn activity peaks of a dominant species rather than overlapping activities of many diurnal and nocturnal species. Diel switches between reef-edge habitat and surrounding sand/seagrass flats were uncommon despite expectations (based on literature examples) that patch reefs would function primarily as sheltering habitats and surrounding non-reef areas act as foraging habitat. High catches at reef-edge sites suggest that the majority of fishes forage on or near limestone patch reefs. Fish densities of around 0.8 individuals per m^-2 of bottom on these Western Australian reefs are relatively high in comparison to visual census estimates obtained for temperate reef systems in South Australia and New Zealand, but similar to those obtained using comparable netting methods in temperate Australian seagrass systems.