Positive indirect effects in a coral reef fish community

Indirect effects occur when two species interact through one or more intermediate species. Theoretical studies indicate that indirect interactions between two prey types that share common predators can be positive, neutral or negative. We document a positive indirect interaction between different types of prey fish on coral reefs in Australia. A high abundance of one type of prey fish (cardinalfishes: Apogonidae) resulted in higher recruitment, abundance and species richness of other prey fish. Our evidence indicates that these effects were not due to differential settlement but were instead due to differential post-settlement predation. We hypothesize that resident piscivores altered their foraging behaviour by concentrating on highly abundant cardinal-fish when they were present, leaving recruits of other species relatively unmolested. Indirect effects were evident within 48 h of settlement and persisted throughout the 42-day experiment, highlighting the importance of early post-settlement processes in these communities.     

Mass spawning of corals on a high latitude coral reef

Evidence is presented that at least 60% of the 184 species of scleractinian corals found on reefs surrounding the Houtman Abrolhos Islands (Western Australia) participate in a late summer mass spawning. These populations are thus reproductively active, despite most species being at the extreme southern limit of their latitudinal range (28° 29°S). In the present study, coral mass spawning occurred in the same month on both temperate (Houtman-Abrolhos) and tropical (Ningaloo) reefs of Western Australia, despite more than two months difference in the timing of seasonal temperture minima between the two regions. This concurrence in the month of spawning suggests that temperature does not operate as a simple direct proximate cue for seasonal spawning synchrony in these populations. Seasonal variation in photoperiod may provide a similar and more reliable signal in the two regions, and thus might be more likely to synchronize the seasonal reproductive rhythms of these corals. Also there is overlap in the nights of mass spawning on the Houtman Abrolhos and tropical reefs of Western Australia, despite significant differences in tidal phase and amplitude between the two regions. This indicates that tidal cycle does not synchronize with the night(s) of spawning on these reefs. Spawning is more likely to be synchronised by lunar cycles. The co-occurrence of the mass spawning with spring tides in Houtman Abrolhos coral populations may be evidence of a genetic legacy inherited from northern, tropical ancestors. Micro-tidal regimes in the Houtman Abrolhos region may have exerted insufficient selective pressure to counteract this legacy.     

Long-term effects of the cleaner fish Labroides dimidiatus on coral reef fish communities

Cleaning behaviour is deemed a mutualism, however the benefit of cleaning interactions to client individuals is unknown. Furthermore, mechanisms that may shift fish community structure in the presence of cleaning organisms are unclear. Here we show that on patch reefs (61–285 m²) which had all cleaner wrasse Labroides dimidiatus (Labridae) experimentally removed (1–5 adults reef⁻¹) and which were then maintained cleaner-fish free over 8.5 years, individuals of two site-attached (resident) client damselfishes (Pomacentridae) were smaller compared to those on control reefs. Furthermore, resident fishes were 37% less abundant and 23% less species rich per reef, compared to control reefs. Such changes in site-attached fish may reflect lower fish growth rates and/or survivorship. Additionally, juveniles of visitors (fish likely to move between reefs) were 65% less abundant on removal reefs suggesting cleaners may also affect recruitment. This may, in part, explain the 23% lower abundance and 33% lower species richness of visitor fishes, and 66% lower abundance of visitor herbivores (Acanthuridae) on removal reefs that we also observed. This is the first study to demonstrate a benefit of cleaning behaviour to client individuals, in the form of increased size, and to elucidate potential mechanisms leading to community-wide effects on the fish population. Many of the fish groups affected may also indirectly affect other reef organisms, thus further impacting the reef community. The large-scale effect of the presence of the relatively small and uncommon fish, Labroides dimidiadus, on other fishes is unparalleled on coral reefs.     

Temporal settlement patterns of larvae of the broadcast spawning reef coral Favites chinensis and the broadcast spawning and brooding reef coral Goniastrea aspera from Okinawa, Japan

The faviid corals, Favites chinensis and Goniastrea aspera are widely distributed in the Indo-Pacific region. Both corals are hermaphroditic broadcast spawners, but G. aspera is also known to brood planula larvae in Okinawa. This study investigated the temporal settlement patterns of planula larvae of the scleractinian corals F. chinensis and G. aspera that developed from spawned gametes, and planula release and settlement of brooded larvae of G. aspera from Okinawa, Japan. Some of the broadcast-spawned larvae of F. chinensis and G. aspera had very short pre-competency periods of 1–2 and 2–3 days after spawning, and relatively long maximum settlement-competency periods of 56–63 and 63–70 days after spawning, respectively. These pre-competency periods are among the shortest reported for larvae of broadcast spawning coral species, and appear to be negatively correlated with seawater temperature. F. chinensis larvae tended to settle rapidly with 34–39% of larvae settling in the first week after spawning, while broadcast-spawned G. aspera larvae had a slower settlement pattern with 11–15% of larvae settling in the first week after spawning. Brooded larvae of G. aspera settled more rapidly, with settlement rates of 27–31% within the first 24 h and 45–65% within the first week after the start of the experiment. The production of planula larvae with rapid settlement capabilities may enable F. chinensis and G. aspera to establish and maintain populations in shallow reef sites at Okinawa. The release of the brooded planulae for up to 2 months may explain why G. aspera is locally more dominant on shallow reefs in Okinawa than F. chinensis. On a broader scale, the longer settlement competency periods of some of the broadcast-spawned larvae of these species increase their potential for longer-distance dispersal and may partly explain the wide biogeographic distribution of these species in the Indo-Pacific region.     

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.     

Disturbance and the structure of coral reef fish communities on the reef slope

Three levels of physical disturbance were applied to corals in permanent 10x10 m quadrats along a section of fringing reef at Lizard Island on the Great Barrier Reef to investigate the response of fish assemblages. Tabular and corymbose corals were overturned and left in situ, reducing total hard coral cover from ˜55% to ˜47%, ˜43%, and ˜34%. Despite pre-existing associations with benthic cover, all fish groups examined (pomacentrids, labrids, chaetodontids, and acanthurids) were resistent to benthic disturbances at the level and scale at which they were applied. Partial Mantel's tests, in combination with partial Canonical Correspondence Analysis enabled spatial and temporal variation to be factored out from experimental effects. Most of the variation in the fish community could be assigned to spatio-temporal variables, indicating that spatial structure over the reef landscape may moderate localised disturbance effects. This study indicates that coral reef fish assemblages may be more resistant to disturbance than many correlative studies would suggest, and highlights a need for further information on levels and scales of natural habitat disturbance in order to apply a structured approach to the experimental investigation of the importance of habitat in structuring coral-reef fish assemblages.     

Size-spectra as indicators of the effects of fishing on coral reef fish assemblages

The data requirements and resources needed to develop multispecies indicators of fishing impacts are often lacking and this is particularly true for coral reef fisheries. Size-spectra, relationships between abundance and body-size class, regardless of taxonomy, can be calculated from simple sizeabundance data. Both the slope and the mid-point height of the relationship can be compared at different fishing intensities. Here, we develop size-spectra for reef fish assemblages using body size- abundance data collected by underwater visual census in each of ten fishing grounds across a known gradient of fishing intensity in the Kadavu Island group, Fiji. Slopes of the size-spectra became steeper (F_9,69=3.20, p<0.01) and the height declined (F_9,69=15.78, p<0.001) with increasing fishing intensity. Regressions of numbers of individuals per size class across grounds were negative for all size classes, although the slope was almost zero for the smallest size class. Response to exploitation of each size class category was greatest for larger fish. Steepening of the slope with increasing fishing intensity largely resulted from reductions in the relative abundance of large fish and not from the ecological release of small fish following depletion of their predators. The slope and height of the size-spectrum appear to be good indicators of fishing effects on reef fish assemblages.     

Interactive effects of interspecific competition and microhabitat on early post-settlement survival in a coral reef fish

Microhabitat type and the competition for microhabitats can each influence patterns of abundance and mortality in coral reef fish communities; however, the effect of microhabitat on the intensity and outcome of competition is not well understood. In Kimbe Bay, Papua New Guinea, surveys were used to quantify microhabitat use and selectivity in two live-coral specialist damselfishes (Pomacentridae), Chrysiptera parasema, and Dascyllus melanurus. A patch reef experiment was then conducted to test how intra- and interspecific competition interacts with two types of microhabitat to influence survival of recently settled C. parasema. Surveys demonstrated that C. parasema and D. melanurus recruits utilized similar coral microhabitats; 72% of C. parasema and 85% of D. melanurus used corymbose and bottlebrush growth forms of Acropora. One microhabitat type, Pocillopora sp. coral, was commonly used by D. melanurus but rarely by C. parasema. The patch reef experiment revealed that both microhabitat and interspecific competition influence abundance of recently settled C. parasema. Microhabitat had the strongest influence on survival of C. parasema. In the absence of interspecific competitors, ~85% of C. parasema survived for 5 days after transplantation to high-complexity bottlebrush Acropora reefs when compared to only 25% survival of Pocillopora reefs. In both microhabitats, interspecific competition with D. melanurus, but not intraspecific competition, significantly decreased the survival of C. parasema. Taken together, these results suggest that the observed distribution of C. parasema results from specialized microhabitat requirements and competition for space in those microhabitats. This study demonstrates that interspecific competition and microhabitat type can interact to influence early post-settlement survival in coral reef fishes, though, whether and how these factors influence survival will depend on the behavioural attributes and strength of habitat associations among potential competitors. Communicated by Environment Editor Prof. Rob van Woesik     

Aggressive mimicry in a coral reef fish: The prey's view

Since all forms of mimicry are based on perceptual deception, the sensory ecology of the intended receiver is of paramount importance to test the necessary precondition for mimicry to occur, that is, model‐mimic misidentification, and to gain insight in the origin and evolutionary trajectory of the signals. Here we test the potential for aggressive mimicry by a group of coral reef fishes, the color polymorphic Hypoplectrus hamlets, from the point of view of their most common prey, small epibenthic gobies and mysid shrimp. We build visual models based on the visual pigments and spatial resolution of the prey, the underwater light spectrum and color reflectances of putative models and their hamlet mimics. Our results are consistent with one mimic‐model relationship between the butter hamlet H. unicolor and its model the butterflyfish Chaetodon capistratus but do not support a second proposed mimic‐model pair between the black hamlet H. nigricans and the dusky damselfish Stegastes adustus. We discuss our results in the context of color morphs divergence in the Hypoplectrus species radiation and suggest that aggressive mimicry in H. unicolor might have originated in the context of protective (Batesian) mimicry by the hamlet from its fish predators rather than aggressive mimicry driven by its prey.     

Physiological effects of heat and cold exposure in the common reef coral Acropora millepora

Reef-forming corals are under threat globally from climate change, leading to changes in sea temperatures with both hot and cold events recorded and projected to increase in frequency and severity in the future. Tolerance to heat and cold exposure has been found to be mutually exclusive in other marine invertebrates, but it is currently unclear whether a trade-off exists between hot and cold thermal tolerance in tropical corals. This study quantified the changes in physiology in Acropora millepora from the central Great Barrier Reef subjected to three temperature treatments; sub-lethal cold, ambient and sub-lethal heat (23.0 °C, 27.0 °C and 29.5 °C, respectively). After 10 weeks, pigment content and Symbiodiniaceae density increased in cold-treated corals but decreased in heat-treated corals relative to corals at ambient conditions. Heat-treated corals gained less mass relative to both ambient and cold-treated corals. These results indicate that the physiological condition of A. millepora corals examined here improved in response to mild cold exposure compared to ambient exposure and decreased under mild heat exposure despite both these temperatures occurring in situ around 15% of the year. The energetic condition of corals in the hotter treatment was reduced compared to both ambient and cooler groups, indicating that corals may be more resilient to mild cold exposure relative to mild heat exposure. The results indicate that the corals shifted their resource allocation in response to temperature treatment, investing more energy into skeletal extension rather than maintenance. No evidence of thermal tolerance trade-offs was found, and cold thermal tolerance was not lost in more heat-tolerant individuals. An enhanced understanding of physiological responses of corals at both ends of the thermal spectrum is important for predicting the resilience of corals under projected climate change conditions.

     

Patterns of migration between feeding and spawning sites in a coral reef surgeonfish

Many coral reef fishes exhibit regular localised migrations between feeding and spawning areas, but the factors affecting these migration patterns, such as the distance, frequency and spawning site fidelity are poorly understood. The aim of this study was to investigate the patterns of migration to spawning sites of the surgeonfish, Ctenochaetus striatus (Acanthuridae). We explored relationships amongst an individual’s size and sex, the distance and frequency it migrated from its feeding area to spawning sites, fidelity to particular spawning sites and the number of individuals that aggregated to spawn. In order to achieve this, 406 C. striatus were captured and tagged on inshore reefs in Kimbe Bay (5°30′S 150°6′E), New Britain, Papua New Guinea. Tagged individuals were consistently observed within spatially discrete but overlapping feeding areas (maximum diameter averaging <13 m). The mean distance migrated was 58 m (ranging from 2 to 291 m). No tagged individuals were witnessed spawning at more than one site. Whilst most individuals (n = 88) migrated to the spawning site that was closest to their feeding areas, those that migrated to sites further away (n = 9) always spawned at sites where the number of conspecifics aggregating was larger. Neither the size nor the sex of individuals limited migration distance. However, males migrated significantly more frequently than females (on average once every 2 days vs. once every 3 days), and migration frequency was positively correlated with size in females. Migration distance did not affect the frequency with which individuals spawned. Whether patterns of migration are determined by cost-benefit optimisation, tradition, or an alternative mechanism is unknown.     

Interactive effects of ontogeny, food ration and temperature on elemental incorporation in otoliths of a coral reef fish

The potential for environmental and physiological modification of elemental incorporation in otoliths is significant and must be validated before otoliths can be used reliably to estimate water parameters over the life history of a fish. We experimentally manipulated temperature and diet quantity for juvenile, sub-adult, and adult Acanthochromis polyacanthus, a tropical damselfish of the SW Pacific. Significant interactive effects between life history stage, temperature and food quantity were observed for otolith Ba/Ca, while significant interactions between stage and food were observed for Sr/Ca. Specific growth rates were negatively correlated with D_Ba and D_Sr for juveniles and sub-adults. These interactions indicated elemental incorporation dynamics varied depending on the life history stage, suggesting variation in effects of stage-specific metabolism or reproductive status. Our results highlight complex responses of elemental incorporation to both endogenous and exogenous factors. Interpretations of life history transects across otoliths must account for these effects to avoid confounding environmental variability with ontogenetic changes in physiology.     

Impacts of the live reef fish trade on populations of coral reef fish off northern Borneo

The live reef fish trade (LRFT) is one of the greatest but least-quantified sources of fishing pressure for several species of large coral reef fish across the Indo-Pacific. For the first time we quantify the localized impact of the LRFT. We collected data from three LRFT traders in northern Borneo, which yielded information on daily fishing effort and the species and mass of all fishes sold every day by individual fishers or vessels over 2, 3 and 8 years. Total monthly catch and relative abundance (catch-per-unit-effort) declined significantly in several species, including the most valuable species the Napoleon wrasse (Cheilinus undulatus, estimated changes of -98 and -78% over 8 years in catch and relative abundance, respectively) and lower-value bluelined groupers (Plectropomus oligocanthus: -99 and -81%) and Epinephelus groupers (-89 and -32%). These severe declines were rapid, species-specific and occurred in the first 2-4 years of the dataset and are, we believe, directly attributable to the LRFT. This has crucial implications for future data collection and monitoring if population collapses in other parts of the LRFT and similar wildlife trades are to be successfully detected.     

Eleven polymorphic microsatellite loci in a coral reef fish,Pterapogon kauderni

We describe the isolation and characterization of 11 polymorphic tetranucleotide microsatellite loci from a male mouthbrooding coral reef fish, the Banggai cardinalfish Pterapogon kauderni. In a sample of 37 fish from a natural population, polymorphism at these loci ranged from two to 15 alleles, with expected heterozygosities ranging from 0.107 to 0.928, enabling high‐resolution genetic studies of this coral reef fish.     

Social transmission of behavioural traditions in a coral reef fish

Traditional behaviours involve the non-genetic transmission of social information across age classes or generations. French grunts (Haemulon flavolineatum) exhibit social traditions of daytime schooling sites and twilight migration routes. Individuals transplanted to new schooling sites and allowed to follow residents at the new sites used the new migration routes and returned to the new sites in the absence of resident fish. Control fish with no opportunity to learn showed no such directionality or return. This is the first demonstration of apparent pre-cultural behaviour in free-living fish. Our observations suggest additional classes of behaviour and taxonomic groups in which pre-cultural activities are likely to have evolved.     

Intraspecific competition in a reef coral: effects on growth and reproduction

Summary Growth rates and reproduction of a branching coral (Stylophora pistillata) were compared in the presence and in the absence of intraspecific competition. Field experiments demonstrated a significant decline in the growth rate of competing colonies compared to noncompeting control colonies; the growth rate slowed in all of the interacting individuals, irrespective of their place in the hierarchy of the intraspecific dominance or of their color morph. In case of immediate killing of the subordinate, the dominant colony grew at a normal rate. In addition to the marked decrease in the growth rate of interacting colonies, the typical symmetry shape of these colonies was changed to an abnormal growth form. The number of female gonads per polyp was significantly reduced in colonies competing intraspecifically, and the typical synchrony in reproduction among different branches of a given colony was changed and desynchronized. Again, these results did not correlate with the hierarchy of dominance. We conclude that intraspecific competition in reef corals involves great investment of energy. The ecological significance and the different pathways of this competition are discussed.     

Diurnal variability in a fish assemblage of a Bahamian coral reef

Synopsis Variation in the diurnal composition of a fish assemblage of a Bahamian coral reef was investigated by comparing visual counts of fishes taken along a 100 × 4 m wide fixed transect at four times: 0900, 1200, 1500 and 1800 hours during the summer of 1979. One sample per day was obtained at each time period over a span of 20 consecutive days. Forty-two species were recorded in these samples, with 25 occurring frequently enough to permit statistical analysis of diurnal variations in abundance. Of the 25 species compared, nearly one fourth (6 species) displayed significant variation in abundance patterns among the four time periods tested. It is suggested, because of the strong possibility of bias which might otherwise be introduced because of these variations, that repetitive quantitative visual censusing of coral reef fishes be undertaken at about the same time each day.