Juvenile coral reef fish use sound to locate habitats

There is limited knowledge of the orientation cues used by reef fish in their movement among different habitats, especially those cues used during darkness. Although acoustic cues have been found to be important for settlement-stage fish as they seek settlement habitats, only a small number of studies support the possible role of acoustic cues in the orientation of post-settled and adult reef fish. Therefore, the aim of this study was to determine whether habitat-specific acoustic cues were involved in the nocturnal movements of juvenile reef fish to small experimental patch reefs that were broadcasting sound previously recorded from different habitats (Fringing Reef, Lagoon, Silent). Juvenile fish arriving at each patch reef were caught the next morning by divers and were identified. There were a greater number of occasions when juvenile fish (from all species together) moved onto the patch reefs broadcasting Fringing Reef and Lagoon sound (43 and 38%, respectively) compared to Silent reefs (19%) (χ² = 33.5; P < 0.05). There were significantly more occasions when juvenile fish from the family Nemipteridae were attracted to the patch reefs broadcasting Lagoon sound (63%) versus those reefs broadcasting either Fringing Reef sound (31%) or Silent (6%). In contrast, there were more occasions when juveniles from the family Pomacentridae were attracted to the patch reefs broadcasting Fringing Reef sound (56%) than either Lagoon (24%) or Silent patch reefs (20%) (χ² = 19.5; P < 0.05). These results indicate that some juvenile fish use specific habitat sounds to guide their nocturnal movements. Therefore, the fish are able to not only use the directional information contained in acoustic cues, but can also interpret the content of the acoustic signals for relevant habitat information which is then used in their decision-making for orientation.     

Ontogeny of microhabitat use and two-step recruitment in a specialist reef fish, the Browncheek Blenny (Chaenopsidae)

Ecological specialization is common on coral reefs and almost certainly contributes to the high diversity of fishes and invertebrates associated with reefs. Here, the recruitment pathway of an endemic Gulf of California fish, the Browncheek Blenny, Acanthemblemaria crockeri (Teleostei: Chaenopsidae), which specializes as an adult on vacant invertebrate tests or tubes, is reported. Like most reef fishes, Browncheek Blennies have a planktonic larval stage that leaves the reef and later settles on suitable habitat as a fully developed juvenile. These blennies follow a clear, “two-step” recruitment pathway, however, and do not reside in invertebrate tests until reaching an adult body size. Individual juveniles and adults were observed for 3 min intervals in order to develop average time budgets for this species. Members of both sexes and all post-settlement life-history stages were included in the analysis. The difference in habitat use by post-settlement juveniles and adults is striking; the average juvenile spends none of its time inside a test, and the average large adult spends all of its time inside a test. Using data on intermediate-sized individuals, the behavioral change associated with invading a test was determined to be size-cued, and it occurs between 20 and 30 mm standard length. Changes in feeding and predator avoidance behaviors are also associated with the ontogenetic shift from life in the open to life in a shelter. Addition of artificial shelters demonstrated the essential role of access to this specialized resource in the population regulation of adults but not juveniles of these blennies.     

Spatio-temporal variability in fish recruitment to a coral reef (Moorea,French Polynesia)

Spatial and temporal patterns of recruitment of juvenile coral reef fishes were studied on the reefs of the high island of Moorea (Society Archipelago, French Polynesia) during the wet season (October 1988 to April 1989). Two size-classes of fishes (new-recruits and juveniles) were censused by visual counts within 24 quadrats along a transect across the lagoon. Correspondence analysis was used to calculate the hypothetical movements of the two size classes. Spatial variability was far more important than temporal variability; six main spatial communities were revealed, which could result from differences among habitats. Temporal variability in recuitment occurred only at nearshore stations; stations on the outer fringing reef and inner barrier reef displayed stable recruitment patterns.     

Highlighting ontogenetic shifts in habitat use by nocturnal coral reef fish

The lagoon of Moorea Island was characterised by 12 distinct reef zones. Visual censuses allowed us to document the spatial distributions of recently settled juveniles vs adults of 17 nocturnal fish species among the 12 reef zones. Five distinct patterns in habitat use were found: an increase in the number of reef zones used during the adult stage (four species); a decrease in the number of reef zones adults used compared to recently settled juveniles (two species); the use of different reef zones (one species); the use of same reef zones but with relative densities different (one species); and no change in habitat use (nine species). Overall, this study is the first to explore the use of space by a broad range of nocturnal fish taxa to document the patterns and determinism of habitat shifts between juvenile and adult life stages.     

Importance of live coral habitat for reef fishes

Live corals are the key habitat forming organisms on coral reefs, contributing to both biological and physical structure. Understanding the importance of corals for reef fishes is, however, restricted to a few key families of fishes, whereas it is likely that a vast number of fish species will be adversely affected by the loss of live corals. This study used data from published literature together with independent field based surveys to quantify the range of reef fish species that use live coral habitats. A total of 320 species from 39 families use live coral habitats, accounting for approximately 8 % of all reef fishes. Many of the fishes reported to use live corals are from the families Pomacentridae (68 spp.) and Gobiidae (44 spp.) and most (66 %) are either planktivores or omnivores. 126 species of fish associate with corals as juveniles, although many of these fishes have no apparent affiliation with coral as adults, suggesting an ontogenetic shift in coral reliance. Collectively, reef fishes have been reported to use at least 93 species of coral, mainly from the genus Acropora and Porities and associate predominantly with branching growth forms. Some fish associate with a single coral species, whilst others can be found on more than 20 different species of coral indicating there is considerable variation in habitat specialisation among coral associated fish species. The large number of fishes that rely on coral highlights that habitat degradation and coral loss will have significant consequences for biodiversity and productivity of reef fish assemblages.     

Strong density-dependent survival and recruitment regulate the abundance of a coral reef fish

Debate on the control of population dynamics in reef fishes has centred on whether patterns in abundance are determined by the supply of planktonic recruits, or by post-recruitment processes. Recruitment limitation implies little or no regulation of the reef-associated population, and is supported by several experimental studies that failed to detect density dependence. Previous manipulations of population density have, however, focused on juveniles, and there have been no tests for density-dependent interactions among adult reef fishes. I tested for population regulation in Coryphopterus glaucofraenum, a small, short-lived goby that is common in the Caribbean. Adult density was manipulated on artificial reefs and adults were also monitored on reefs where they varied in density naturally. Survival of adult gobies showed a strong inverse relationship with their initial density across a realistic range of densities. Individually marked gobies, however, grew at similar rates across all densities, suggesting that density-dependent survival was not associated with depressed growth, and so may result from predation or parasitism rather than from food shortage. Like adult survival, the accumulation of new recruits on reefs was also much lower at high adult densities than at low densities. Suppression of recruitment by adults may occur because adults cause either reduced larval settlement or reduced early post-settlement survival. In summary, this study has documented a previously unrecorded regulatory mechanism for reef fish populations (density-dependent adult mortality) and provided a particularly strong example of a well-established mechanism (density-dependent recruitment). In combination, these two compensatory mechanisms have the potential to strongly regulate the abundance of this species, and rule out the control of abundance by the supply of recruits.     

Temporal consistency of ontogenetic shifts in habitat use by coral reef fishes in the northernmost coral ecosystem in the world (Kudaka Island, Japan)

The prevalence of major habitat shifts in tropical fishes between juvenile and adult stages (ontogenetic shifts) in one of the northernmost coral reefs in the world (Kudaka Island, Japan) is given. The comparative analysis of spatial distribution of juveniles v . adults highlighted four ontogenetic patterns: no change in habitat use between juveniles and adults (five species), a decrease in the number of habitats used by adults compared to juveniles (three species), an increase in the number of habitats used during the adult stage (four species) and use of nursery areas by juveniles followed by extensive movements to different adult habitats (three species). The comparative analysis of fish distribution over time ( i.e. during three consecutive settlement months) showed that 84% of species had temporal consistency in ontogenetic patterns of habitat use.     

Bipartite life cycle of coral reef fishes promotes increasing shape disparity of the head skeleton during ontogeny: an example from damselfishes (Pomacentridae)

Background  

Quantitative studies of the variation of disparity during ontogeny exhibited by the radiation of coral reef fishes are lacking. Such studies dealing with the variation of disparity, i.e. the diversity of organic form, over ontogeny could be a first step in detecting evolutionary mechanisms in these fishes. The damselfishes (Pomacentridae) have a bipartite life-cycle, as do the majority of demersal coral reef fishes. During their pelagic dispersion phase, all larvae feed on planktonic prey. On the other hand, juveniles and adults associated with the coral reef environment show a higher diversity of diets. Using geometric morphometrics, we study the ontogenetic dynamic of shape disparity of different head skeletal units (neurocranium, suspensorium and opercle, mandible and premaxilla) in this fish family. We expected that larvae of different species might be relatively similar in shapes. Alternatively, specialization may become notable even in the juvenile and adult phase.     

Simple ecological trade-offs give rise to emergent cross-ecosystem distributions of a coral reef fish

Ecosystems are intricately linked by the flow of organisms across their boundaries, and such connectivity can be essential to the structure and function of the linked ecosystems. For example, many coral reef fish populations are maintained by the movement of individuals from spatially segregated juvenile habitats (i.e., nurseries, such as mangroves and seagrass beds) to areas preferred by adults. It is presumed that nursery habitats provide for faster growth (higher food availability) and/or low predation risk for juveniles, but empirical data supporting this hypothesis is surprisingly lacking for coral reef fishes. Here, we investigate potential mechanisms (growth, predation risk, and reproductive investment) that give rise to the distribution patterns of a common Caribbean reef fish species, Haemulon flavolineatum (French grunt). Adults were primarily found on coral reefs, whereas juvenile fish only occurred in non-reef habitats. Contrary to our initial expectations, analysis of length-at-age revealed that growth rates were highest on coral reefs and not within nursery habitats. Survival rates in tethering trials were 0% for small juvenile fish transplanted to coral reefs and 24-47% in the nurseries. As fish grew, survival rates on coral reefs approached those in non-reef habitats (56 vs. 77-100%, respectively). As such, predation seems to be the primary factor driving across-ecosystem distributions of this fish, and thus the primary reason why mangrove and seagrass habitats function as nursery habitat. Identifying the mechanisms that lead to such distributions is critical to develop appropriate conservation initiatives, identify essential fish habitat, and predict impacts associated with environmental change.     

SMURFs: standard monitoring units for the recruitment of temperate reef fishes

I evaluated a standard monitoring unit for the recruitment of reef fishes (SMURF) as a tool for ascertaining spatial and temporal patterns of reef fish recruitment in central California, USA. SMURFs consisted of a 1.0×0.35 m dia. cylinder of fine mesh plastic grid that contained a folded section of larger mesh plastic grid. SMURFs collected new recruits of 20 species of fish with 92% of the individuals collected from 10 species, mostly rockfish (genus Sebastes). An experiment varying depth of SMURFs in the water column (surface, mid-depth, or bottom) showed that surface SMURFs collected the greatest diversity of species and significantly greater abundance for eight species, with two species having significantly greater abundance on mid-depth SMURFs and three species having significantly greater abundance on bottom SMURFs. A comparison of cumulated recruitment from SMURFs that varied in sampling frequency (removal of new recruits every 1-3, 7, or 28 days) suggested that increasing the time between sampling caused a significant decrease in recruitment estimates for some species but not for others. To determine how well temporal patterns of recruitment to SMURFs reflected patterns to nearby reefs, I compared within season temporal patterns of recruitment to SMURFs with that at nearby reefs, estimated by visual transect surveys conducted on scuba. Temporal patterns of recruitment to SMURFs were significantly and positively related to early recruitment on reefs for one group of benthic-algal associated rockfish species when diver surveys were lagged by 30 days (r=0.87) and for another group of canopy-algal associated rockfish species when lagged by 5 days (r=0.72). SMURFs appeared to be an effective and efficient method for indexing relative rates of delivery of competent juveniles for many temperate nearshore reef fishes.     

Spatial structure of coral reef fish communities at Kudaka Island (Ryukyu Archipelago), Japan

The present study aimed to investigate the spatial structure of fish communities at juvenile and adult stages on coral reefs at Kudaka Island (Ryukyu Archipelago, Japan) and to relate spatial patterns in the structure of the fish communities to gradients in environmental variables. Diurnal visual censuses allowed us to record 2,602 juveniles belonging to 60 species and 1,543 adults belonging to 53 species from October to December 2005. The distribution of species highlighted that the juvenile community was organised into three distinct assemblages, rather than exhibiting gradual change in community structure along the cross-reef gradient. Correlations between spatial patterns of juvenile community and environmental variables revealed that the most significant factors explaining variation in community structure were coral rubble and coral slab. In contrast, the adult community was organised into one assemblage, and the most significant variation factors in community structure were depth, live coral in massive form, live coral in branched form, dead coral and sand. Overall, the present study showed that most juvenile and adult coral reef fish at Kudaka Island exhibited striking patterns in their distribution and depth and some biological factors (e.g., abundance of live coral, dead coral and coral rubble) might exert considerable influence on the distribution of fishes.     

Non‐reef habitats in a tropical seascape affect density and biomass of fishes on coral reefs

Nonreef habitats such as mangroves, seagrass, and macroalgal beds are important for foraging, spawning, and as nursery habitat for some coral reef fishes. The spatial configuration of nonreef habitats adjacent to coral reefs can therefore have a substantial influence on the distribution and composition of reef fish. We investigate how different habitats in a tropical seascape in the Philippines influence the presence, density, and biomass of coral reef fishes to understand the relative importance of different habitats across various spatial scales. A detailed seascape map generated from satellite imagery was combined with field surveys of fish and benthic habitat on coral reefs. We then compared the relative importance of local reef (within coral reef) and adjacent habitat (habitats in the surrounding seascape) variables for coral reef fishes. Overall, adjacent habitat variables were as important as local reef variables in explaining reef fish density and biomass, despite being fewer in number in final models. For adult and juvenile wrasses (Labridae), and juveniles of some parrotfish taxa (Chlorurus), adjacent habitat was more important in explaining fish density and biomass. Notably, wrasses were positively influenced by the amount of sand and macroalgae in the adjacent seascape. Adjacent habitat metrics with the highest relative importance were sand (positive), macroalgae (positive), and mangrove habitats (negative), and fish responses to these metrics were consistent across fish groups evaluated. The 500‐m spatial scale was selected most often in models for seascape variables. Local coral reef variables with the greatest importance were percent cover of live coral (positive), sand (negative), and macroalgae (mixed). Incorporating spatial metrics that describe the surrounding seascape will capture more holistic patterns of fish–habitat relationships on reefs. This is important in regions where protection of reef fish habitat is an integral part of fisheries management but where protection of nonreef habitats is often overlooked.     

Otolith growth of Springer's demoiselle, Chrysiptera springeri (Pomacentridae, Allen & Lubbock), on a protected and non-protected coral reef

The structural complexity of coral reefs is important for their function as shelter and feeding habitats for coral reef fishes, but physical disturbance by human activities often reduce complexity of the reefs by selectively destroying fragile and more complex coral species. The damselfish Springer's demoiselle Chrysiptera springeri primarily utilize complex coral heads for shelter and are hence vulnerable to human disturbance. In order to evaluate the potential effect of habitat degradation on juvenile fish growth, coral reef cover, fish age at settling and otolith growth, juvenile Springer's demoiselle was investigated on a protected and non‐protected coral reef in Darvel Bay, Borneo. The protected reef had higher coverage of complex branching corals and exhibited a more complex 3‐dimensional structure than the non‐protected reef. Springer's demoiselle settled at the same age on non‐protected and protected reefs. The growth rates of the otoliths from Springer's demoiselle were similar during the pre‐settlement period on the two reefs (manova , P > 0.05), but from age 20 to 48 days (post‐settlement period) the otolith growth rate of juveniles on the non‐protected reef was reduced compared to those from the protected reef (manova , P = 0.017). However, the differences in the otolith size, and by inference, fish size, after 48 days were small. The small effect of habitat degradation on growth is likely related to the fact that the Springer's demoiselles collected on the non‐protected reef were associated with the few remaining complex coral heads. Increased foraging‐predation tradeoffs on the non‐protected reef may decrease food intake and growth of juvenile Springer's demoiselle, but the main effect of habitat degradation on their abundance is likely to be related to lack of suitable shelter, and consequently reduced carrying capacity, on disturbed reefs.     

Spatiotemporal distribution of nocturnal coral reef fish juveniles in Moorea Island,French Polynesia

This study aimed to investigate the spatial structure of nocturnal fish communities at settlement on coral reefs in Moorea Island lagoon, French Polynesia; and the temporal consistency of habitat selection between winter (April to June 2001) and summer (November 2001). The Moorea lagoon was divided into 12 habitat zones (i.e., coral reef zones), which were distinct in terms of depth, wave exposure, and substratum composition. Nocturnal visual censuses among the 12 habitats found that the recently settled juveniles of 25 species recorded were dispatched among three communities spatially distributed according to the distance from the reef crest (reef crest, barrier reef, and fringing reef communities). This spatial communities structure of nocturnal juveniles was consistent in both winter and summer and would be explained primarily by a current gradient in Moorea lagoon (current speed was high near the reef crest and decreased towards the beach) and by the topographic characteristics of reef zones. Among the 25 species, 13 were recorded in both winter and summer. A comparison of the spatial distribution between summer and winter for 13 species that occurred during both seasons found that only 4 differed between the two seasons. For these species, habitat selection would be organized primarily by some stochastic processes such as inter- and intraspecific competition, predation, and food availability. Overall, the present study allowed us to highlight that most nocturnal coral reef fish juveniles at Moorea Island exhibited striking patterns in their distribution and current and topographic characteristics of reef zones might exert considerable influence on the distribution of fishes.     

Using passive acoustic telemetry to infer mortality events in adult herbivorous coral reef fishes

Mortality is considered to be an important factor shaping the structure of coral reef fish communities, but data on the rate and nature of mortality of adult coral reef fishes are sparse. Mortality on coral reefs is intrinsically linked with predation, with most evidence suggesting that predation is highest during crepuscular periods. We tested this hypothesis using passive acoustic telemetry data to determine the time of day of potential mortality events (PMEs) of adult herbivorous reef fishes. A total of 94 fishes were tagged with acoustic transmitters, of which 43 exhibited a PME. Furthermore, we identified five categories of PMEs based on the nature of change in acoustic signal detections from tagged fishes. The majority of PMEs were characterised by an abrupt stop in detections, possibly as a result of a large, mobile predator. Overall, mortality rates were estimated to be approximately 59 % per year using passive acoustic telemetry. The time of day of PMEs suggests that predation was highest during the day and crepuscular periods and lowest at night, offering only partial support for the crepuscular predation hypothesis. Visually oriented, diurnal and crepuscular predators appear to be more important than their nocturnal counterparts in terms of predation on adult reef fishes. By timing PMEs, passive acoustic telemetry may offer an important new tool for investigating the nature of predation on coral reefs.     

The use of artificial reefs in enhancing fish communities in Singapore

Intense development of the coastal zone in Singapore has resulted in the degradation of much of the marine ecosystem. In order to restore and enhance fish communities of denuded areas, an artificial reef consisting of a tyre reef and a concrete reef, was established in the vicinity of the southern islands of Singapore. Results from fish visual censuses after the establishment of the artificial reef indicated an increase in numbers of juveniles and adults. A total of 37 and 32 fish species were recorded over a period of 1/2 years at the concrete and tyre reefs respectively. The dominant fish families were Pomacentridae, Labridae, Chaetodontidae, Apogonidae, Gobiidae and Nemipteridae. The artificial reefs also serve as a nursery ground for some species (e.g. Neopomacentrus sp.) which are important primary consumers of algae on natural reefs. Greater numbers of target (food-important) fishes were observed at the concrete reef while the tyre reef harboured more juveniles and smallersized adults. The results indicate that the concrete modules were more effective than the tyre reef in terms of fish abundance per unit volume. Such structures can enhance the biological resources of relatively unproductive areas.     

Food ration and condition affect early survival of the coral reef damselfish, Stegastes partitus

The supply of larvae is a major determinant of population and community structure in coral reef fishes. However, spatial and temporal variation in condition (i.e. quality) of potential recruits, as well as their density (i.e. quantity), may influence survival and growth of juveniles. We conducted an experiment to test whether recent feeding history could affect growth, condition and post-recruitment survival in a Caribbean damselfish, Stegastes partitus. Fish were collected soon after settlement, and fed either low or high rations in aquaria for 7 days. Fish fed the high ration grew faster in aquaria and were in a better condition (higher total lipids and Fulton’s condition factor) at the end of the feeding period. Subsequently, we released 50 fish in 25 pairs (one fish subjected to low rations, the other to high rations) on a Bahamian coral reef and monitored survival for 10 days. Survivorship of high-ration fish was double that of low-ration fish (80 vs 40% over 10 days). However, low-ration fish that survived 10 days were of similar condition and grew at similar rates to high-ration fish, suggesting that short-term ration differences may not persist in surviving fish. Laboratory experiments showed that low-ration fish were taken by piscivorous fishes before high-ration fish, indicating that differential predation may account for survival differences. This study highlights the potential of feeding history and condition to affect the relationship between patterns of larval arrival at reefs, and subsequent juvenile and adult population densities. Received: 1 March 1999 / Accepted: 15 July 1999     

Ontogenetic shifts in the social behaviour of pairing coral reef rabbitfishes (Siganidae)

Adult rabbitfishes (Siganidae) differ from most other herbivorous coral reef fishes by forming stable pair bonds throughout their adult lives. However, little is known about the early life stages of rabbitfishes, and no quantitative evidence regarding the ontogeny of pairing behaviour exists to date. This study describes the abundance, distribution and ontogeny of social associations in juvenile rabbitfishes on the mid-shelf reefs around Lizard Island, Great Barrier Reef, Australia. Using underwater visual censuses, we surveyed an area of 5728?m² across three distinct habitat types, revealing that the abundance of juveniles varies significantly among species, size class and exposure level. Furthermore, we demonstrate a pronounced ontogenetic shift in the social associations of juveniles of Siganus doliatus and Siganus corallinus, changing from primarily solitary individuals in the smallest size class (<50?mm) to predominantly paired individuals in the larger size classes (50–100?mm and 100–150?mm). In this context, we report the presence of several mixed-species pairs of rabbitfishes, providing the first evidence for this behaviour within the family. This supports previous research, which posits that there are strong ecological drivers, rather than a solely reproductive basis, for pairing behaviour in rabbitfishes. Based on our results, we suggest that further exploration of the ecology of early life stages of herbivorous reef fishes will increase our knowledge about ecological processes on coral reefs.     

Species-specific habitat distribution of coral reef fish assemblages in relation to habitat characteristics in an Okinawan coral reef

We determined the species-specific habitat associations of coral reef fishes and environmental characteristics in an Okinawan coral reef in Japan. We focused on three families (Pomacentridae, Gobiidae and Labridae) and attempted to determine differences in habitat utilization. We selected six sites along the coast of Amitori Bay, from the entrance to the innermost part, in order to cover a wide range of habitat characteristics (exposed habitat, semi-exposed habitat and sheltered habitat). The species diversity of coral assemblages was greater at the exposed and semi-exposed habitats, whereas branching coral mostly covered the sheltered habitat. The environmental factors that determine the species-specific spatial association in fishes differed among families. Both biological characteristics (coral morphology and coral species diversity) and physical characteristics (water depth and wave exposure) affected the spatial association of pomacentrids and gobiids. In contrast, physical characteristics such as substrate complexity and water depth affected the species-specific spatial association of labrid species. Further study is needed to determine the ecological factors that regulate the species-specific habitat preference in Okinawan coral reefs.