The relationship between habitat structure and fish faunas on New Zealand reefs

Reef fish abundances were sampled at 11 shallow reef localities extending over 1000 km of coastline in northern New Zealand. Sampling was restricted to the 4–10-m depth stratum and included six coastal and five island localities. These were either coralline reef flats dominated by echinoids, or algal reefs with high densities of laminarian and fucoid algae. Reefs dominated by macroscopic algae supported large numbers of small fishes, mainly labrids, and few large benthic-feeding fishes. Echinoid-dominated reefs supported a different fish fauna with more large benthic-feeding species. Additional sampling of echinoiddominated reefs and algal stands in deeper water provided confirmation of these findings. A second sampling programme was carried out at a series of eight sites within a single locality covering 5 km of coastline. These spanned a moderate exposure gradient and ranged from algal dominated reefs to typical coralline reef flats with high densities of grazing invertebrates. The relationship between habitat structure and reef fish species composition and size frequency was similar to that of the large-scale sampling programme. Thirdly, observations on reef fish foraging and feeding patterns within a single reef site suggested that larger benthic-feeding reef fishes were less likely to feed within macroscopic algal stands. Experimental reductions of grazing invertebrates designed to produce brown algal stands on echinoid-dominated reef flats supported these observations. Larger individuals capable of removing echinoids and grazing gastropods did not frequent or feed in laminarian and fucoid algal stands. This pattern is discernible at several spatial scales. Our conclusion is that the type of shallow reef habitat, echinoid- as opposed to algal-dominated, will have an important rôle in determining the associated reef fish fauna.     

Multi-scale recruitment patterns and effects on local population size of a temperate reef fish

Recruitment of the temperate reef fish Coris julis was studied across the Azores Archipelago (central North Atlantic), over four consecutive recruitment seasons and at three spatial scales: between islands (separated by 100s of km), sites within islands (separated by 10s of km) and transects within sites (separated by 10s of m). At the largest scale ( i.e . between islands) spatial recruitment patterns were highly variable, suggesting the influence of stochastic processes. Recruitment was spatially consistent within islands, even though magnitude was unpredictable between years, indicating that processes at meso-scales are probably more deterministic. Recruits settled randomly at the transect scale, probably reflecting habitat homogeneity. It was proposed that large and island-scale patterns reflect larval availability, driven by physical and biological processes occurring in the plankton. No evidence was found for a density-dependent relationship between newly settled and 2 week settled C. julis nor between cumulative recruitment and young-of-the-year. It appears that adult density is limited by larval supply (pre-settlement regulation) at low recruitment sites, and determined by post-settlement, density-dependent processes at high recruitment sites. This work is one of few to investigate multiple spatial and temporal scales of recruitment for a coastal fish species inhabiting isolated, temperate oceanic islands and hence, provides a novel comparison to the many studies of recruitment on coral reefs and other, more connected systems.     

Fine-scale temporal variation in recruitment of a temperate demersal fish: the importance of settlement versus post-settlement loss

In order to understand variability in recruitment to populations of benthic and demersal marine species, it is critical to distinguish between the contributions due to variations in larval settlement versus those caused by post-settlement mortality. In this study, fine-scale (1–2 days) temporal changes in recruit abundance were followed through an entire settlement season in a temperate demersal fish in order to determine 1) how dynamic the process of recruitment is on a daily scale, 2) whether settlement and post-settlement mortality are influenced by habitat structure and conspecific density, and 3) how the relationship between settlement and recruitment changes over time. Settlement is considered to be the arrival of new individuals from the pelagic habitat, and recruitment is defined as the number of individuals surviving arbitrary periods of time after settlement. Replicate standardized habitat units were placed in 2 spatial configurations (clumped and randomly dispersed) and monitored visually for cunner (Tautogolabrus adspersus) settlement and recruitment every 1–2 days throughout the settlement season. The process of recruitment in T. adspersus was highly variable at a fine temporal scale. Changes in the numbers of recruits present on habitat units were due to both settlement of new individuals and mortality of animals previously recruited. The relative importance of these two processes appeared to change from day to day. The magnitude of the change in recruit number did not differ between the clumped and random habitats. However, post-settlement loss was significantly greater on randomly dispersed than clumped habitats. During several sampling dates, the extent of the change in recruit abundance was correlated with the density of resident conspecifics; however, on other dates no such relationship appeared to exist. Despite the presence of significant relationships between the change in recruit number and density, there was no evidence of either density-dependent mortality or settlement. Initially, there was a strong relationship between settlement and recruitment; however, this relationship weakened over time. Within 2 months after the cessation of settlement, post-settlement loss was greater than 99%, and no correlation remained between recruitment and the initial pattern of settlement. The results of this study demonstrate that the spatial arrangement of the habitat affects the rate and intensity of post-settlement loss. Counter to much current thinking, this study suggests that in order to understand the population ecology of reef fishes, knowledge of what habitats new recruits use and how mortality varies with structural aspects of the habitats is essential.     

Variation in the effects of larval history on juvenile performance of a temperate reef fish

For organisms with complex life cycles, the transition between life stages can act as a significant demographic and selective bottleneck. Variation in developmental and growth rates among individuals present in one stage (e.g. larvae), due to initial differences in parental input and/or environmental conditions experienced, can propagate to future stages (e.g. juveniles), and such ‘carry‐over effects’ can shape fitness and phenotypic distributions within a population. However, variation in the strength of carry‐over effects between life stages and the intensity of selective mortality acting on intrinsic variation, and how these might be mediated by environmental variability in natural systems, is poorly known. Here, we evaluate variation in the strength to which larval growth histories can mediate juvenile performance (growth and survival), for a reef fish (Forsterygion lapillum) common to rocky reefs of New Zealand. We used otoliths to reconstruct demographic histories of recently settled fish that were sampled across cohorts, sites and microhabitats. We quantified sources of variation in the strength of carry‐over effects and selective mortality that operate on larval growth histories. We found overall that individuals that grew fast as larvae tended to experience proportional growth advantages as juveniles. However, the strength of growth advantages being maintained into the juvenile period varied among cohorts, sites and microhabitats. Specifically, a stronger growth advantage was found on some microhabitats (e.g. mixed stands of macroalgae) relative to others (e.g. monocultures of Carpophyllum maschalocarpum) for some cohorts and sites only. For other cohorts and sites, the degree of coupling between larval and juvenile growth rates was either indistinguishable between microhabitats or else not evident. Similarly, the intensity of growth‐based selective mortality varied among cohorts, sites and microhabitats: for the cohort and site where carry‐over effects differed between microhabitats, we also observed difference in the intensity to which fish with rapid larval growth rates were favoured. Overall, our results highlight how this spatial and temporal patchiness in extrinsic factors can interact with intrinsic variation of recruiting individuals to have a major influence on the resulting distribution of juveniles and their phenotypic traits.     

The influence of habitat characteristics and conspecifics on attraction and survival of coral reef fish juveniles

In marine species with a pelagic larval stage, search behavior and selection of a suitable reef habitat can maximize the settlement success of recently settled juveniles and their subsequent performance (growth and survival of juveniles). Our objective was to test this hypothesis for a single target coral reef fish species (Chromis viridis) at Moorea Island. C. viridis settle on living coral colonies of Porites rus already populated with conspecifics. In the present study (conducted in experimental cages), we found that: 1) mortality rate of recently settled juveniles of C. viridis was lower in the settlement habitat (living coral colonies of P. rus) than in other habitats having physical structure different from those of P. rus colonies; 2) C. viridis juveniles preferentially colonized coral heads of P. rus with conspecifics present rather than uninhabited coral heads and they also preferentially colonized uninhabited coral heads rather than coral heads with heterospecifics; 3) mortality rate of C. viridis juveniles did not vary with the presence or absence of conspecifics or heterospecifics on P. rus colonies. Overall, the study allows us to highlight that site selection by juveniles for habitat containing conspecifics does not benefit their short term mortality rates, suggesting that in the short term at least, site selection has little importance.     

枸杞岛岩礁生境主要鱼类的食物组成及食物竞争

基于2009年3月-2010年2月枸杞岛沿岸岩礁生境渔业资源调查数据,对岩礁生境中的3种定居性鱼类（褐菖鲉、大泷六线鱼和斑头六线鱼）和3种非定居性鱼类（花鲈、黄姑鱼和小黄鱼）的食物组成及食物关系进行了研究.结果表明： 麦秆虫、钩虾、褐菖鲉幼鱼、鳀鱼和中国毛虾等是6种鱼类的主要饵料生物,且饵料组成随季节而变化.3种定居性鱼类中,大泷六线鱼和斑头六线鱼在秋季食物竞争激烈;3种非定居性鱼类中,黄姑鱼和小黄鱼在夏季食物竞争激烈;非定居性鱼类和定居性鱼类之间,黄姑鱼分别与褐菖鲉（秋季）和大泷六线鱼（冬季）食物竞争激烈.黄姑鱼是引起岩礁生境中6种主要鱼类食物竞争的关键鱼种.枸杞岛岩礁生境中6种主要鱼类的摄食活动对褐菖鲉幼鱼的资源量有一定影响.     

Effects of habitat,wave exposure,and marine protected area status on coral reef fish assemblages in the Hawaiian archipelago

The relationships between fish assemblages, their associated habitat, and degree of protection from fishing were evaluated over a broad spatial scale throughout the main Hawaiian islands. Most fish assemblage characteristics showed positive responses to protection whether it was physical (e.g. habitat complexity), biological (e.g. coral cover growth forms), or human-induced (e.g. marine reserves). Fish biomass was lowest in areas of direct wave exposure and highest in areas partially sheltered from swells. Higher values for fish species richness, number of individuals, biomass, and diversity were observed in locations with higher substrate complexity. Areas completely protected from fishing had distinct fish assemblages with higher standing stock and diversity than areas where fishing was permitted or areas that were partially protected from fishing. Locations influenced by customary stewardship harbored fish biomass that was equal to or greater than that of no-take protected areas. Marine protected areas in the main Hawaiian islands with high habitat complexity, moderate wave disturbance, a high percentage of branching and/or lobate coral coupled with legal protection from fishing pressure had higher values for most fish assemblage characteristics.     

A process‐based model supports an association between dispersal and the prevalence of species traits in tropical reef fish assemblages

Habitat dynamics interacting with species dispersal abilities could generate gradients in species diversity and prevalence of species traits when the latter are associated with species dispersal potential. Using a process‐based model of diversification constrained by a dispersal parameter, we simulated the interplay between reef habitat dynamics during the past 140 million years and dispersal, shaping lineage diversification history and assemblage composition globally. The emerging patterns from the simulations were compared to current prevalence of species traits related to dispersal for 6315 tropical reef fish species. We found a significant spatial congruence between the prevalence of simulated low dispersal values and areas with a large proportion of species characterized by small adult body size, narrow home range mobility behaviour, pelagic larval duration shorter than 21 days and diurnal activity. Species characterized by such traits were found predominantly in the Indo‐Australian Archipelago and the Caribbean Sea. Furthermore, the frequency distribution of the dispersal parameter was found to match empirical distributions for body size, PLD and home range mobility behaviour. Also, the dispersal parameter in the simulations was associated to diversification rates and resulted in trait frequency matching empirical distributions. Overall, our findings suggest that past habitat dynamics, in conjunction with dispersal processes, influenced diversification in tropical reef fishes, which may explain the present‐day geography of species traits.     

Microhabitat association and temporal stability in reef fish assemblages on massive <Emphasis Type="Italic">Porites</Emphasis> microatolls

A massive Porites microatoll generally has three types of microhabitat at the top, side, and base of the microatoll. The purpose of the present study was to analyze microhabitat associations of reef fish on microatolls to determine whether habitat characteristics play an important role in the structuring of reef fish assemblages in a patchy habitat. We also investigated temporal stability of reef fish assemblage structures over a period of 17 months to determine whether fish assemblage structures vary in a random manner. The results of correspondence analysis indicated species-specific habitat associations for pomacentrids (five species) and labrids (seven species). The degree of temporal stability of fish assemblage structures, calculated by Piankas index, was relatively high in a large-sized microatoll (0.503–0.831: 3.6m in diameter), in which microhabitat associations of fishes were clearly observed. The present study suggests that a microhabitat association is one of the important factors responsible for organization of reef fish assemblages in a microatoll.     

Spatial and temporal variation of fish assemblages in a subtropical small stream of the Huangshan Mountain

Spatial and temporal variation of fish assemblages were investigated seasonally from May 2007 to February 2008 across 11 study sites in a subtropical small stream, the Puxi Stream, of the Huangshan Mountain. Along the longitudinal gradient from headwater to downstream, fish species richness and abundance increased gradually, but then decreased significantly at the lower reaches. The highest species richness and abundance were observed in August and the lowest in February. Based on analysis of similarities (...     

Spatial patterns of self‐recruitment of a coral reef fish in relation to island‐scale retention mechanisms

Oceanographic features influence the transport and delivery of marine larvae, and physical retention mechanisms, such as eddies, can enhance self‐recruitment (i.e. the return of larvae to their natal population). Knowledge of exact locations of hatching (origin) and settlement (arrival) of larvae of reef animals provides a means to compare observed patterns of self‐recruitment ‘connectivity’ with those expected from water circulation patterns. Using parentage inference based on multiple sampling years in Moorea, French Polynesia, we describe spatial and temporal variation in self‐recruitment of the anemonefish Amphiprion chrysopterus, evaluate the consistency of net dispersal distances of self‐recruits against the null expectation of passive particle dispersal and test the hypothesis that larvae originating in certain reef habitats (lagoons and passes) would be retained and thus more likely to self‐recruit than those originating on the outer (fore) reef. Estimates of known self‐recruitment were consistent across the sampling years (~25–27% of sampled recruits). For most (88%) of these self‐recruits, the net distance between hatching and settlement locations was within the maximum dispersal distance expected for a neutrally buoyant passive particle based on the longest duration of the larval dispersive phase and the average direction and speed of current flow around Moorea. Furthermore, a parent of a given body size on the outer (fore) reef of Moorea was less likely to produce self‐recruits than those in passes. Our findings show that even a simple dispersal model based on net average flow and direction of alongshore currents can provide insight into landscape‐scale retention patterns of reef fishes.     

Distribution and abundance of labrids in northeastern New Zealand: the relationship between depth,exposure and pectoral fin aspect ratio

Physical factors influencing the distribution and abundance of seven common labrid fishes were examined over four rocky reef locations in northeastern New Zealand. Depth and exposure for each species (both within and among sexes) were related to pectoral fin aspect ratio. Each of the four locations (two mainland and two island) displayed distinct labrid assemblages, which were consistent over time, likely due to the influence of the East Auckland Current. There was a consistent depth-related trend for most species, regardless of location. Several species also showed a sex related depth difference. There was also a trend for some species to be associated with certain levels of wave exposure. For most species, the relationship between pectoral fin aspect ratio and the above physical variables was not as strongly evident in this temperate assemblage as has been previously found in tropical reef fish systems. Although some species did follow the predicted shifts in fin aspect ratio with depth and/or exposure, the observed trends were unrelated to fin aspect ratio for many other species. These findings suggest that wave exposure may not be as important for labrids on northeastern New Zealand reefs as it may be in tropical coral reefs systems. The lower fin aspect ratios for New Zealand labrids, compared to tropical labrids, suggest that New Zealand labrids represent a subset of the total pectoral fin diversity in the Labridae. Consequently, the potential for distinct trends in fin aspect ratio and physical variables to be evident may be reduced.     

Spatial and temporal variation in fish assemblages of drying stream pools: The role of abiotic and biotic factors

Droughts and summer drying create unusual temporary aquatic habitats in the form of isolated pools in many small streams around the world. To examine spatial and temporal variation in fish community structure of drying stream pools, their relation to abiotic environmental variables, and associations among species, fish were sampled during summer 1995 and 1996 from pools of four streams in the Ozark mountains, Arkansas, USA. Redundancy analysis of physical-chemical variables showed significant differences among stream sites, but no significant difference between years or stream site by year interaction. Stream sites separated consistently along axes one (habitat heterogeneity) and two (temperature/canopy cover) in both years. Redundancy analysis of fish species-size class densities showed a significant stream site by year interaction. Groupings of stream sites based on fish assemblages were not well explained by physical-chemical variables measured at the pool scale, but were related to location within the drainage basin, and these groupings differed between years. There were 27 (15.8%) and 10 (5.8%) significant associations found among fish species-size classes in 1995 and 1996, respectively, and all but two significant associations in 1995 were positive. Pool depth, habitat heterogeneity, pool size and dissolved oxygen/canopy cover were important local abiotic factors depending on response variables examined. In both years, large fish total density, large central stoneroller density (80 mm TL), and small sunfish (<80 mm TL) density were positively related to pool depth. Otherwise, there was no consistent relationship between physical-chemical variables and dependent variables (fish density and species richness) within a year or between years for a given dependent variable. These results support the hypothesis that local abiotic factors are important in structuring fish assemblages in harsh environments, but the importance of those factors varies temporally, and regional influences appear to override local abiotic conditions as factors structuring fish assemblages in drying stream pools. Predation by terrestrial vertebrates may also be an important factor structuring these fish assemblages that has been largely overlooked.     

Spatial variation in the effects of size and age on reproductive dynamics of common coral trout Plectropomus leopardus

The effects of size and age on reproductive dynamics of common coral trout Plectropomus leopardus populations were compared between coral reefs open or closed (no‐take marine reserves) to fishing and among four geographic regions of the Great Barrier Reef (GBR), Australia. The specific reproductive metrics investigated were the sex ratio, the proportion of vitellogenic females and the spawning fraction of local populations. Sex ratios became increasingly male biased with length and age, as expected for a protogyne, but were more male biased in southern regions of the GBR (Mackay and Storm Cay) than in northern regions (Lizard Island and Townsville) across all lengths and ages. The proportion of vitellogenic females also increased with length and age. Female P. leopardus were capable of daily spawning during the spawning season, but on average spawned every 4·3 days. Mature females spawned most frequently on Townsville reserve reefs (every 2·3 days) and Lizard Island fished reefs (every 3·2 days). Females on Mackay reefs open to fishing showed no evidence of spawning over 4 years of sampling, while females on reserve reefs spawned only once every 2–3 months. No effect of length on spawning frequency was detected. Spawning frequency increased with age on Lizard Island fished reefs, declined with age on Storm Cay fished reefs, and declined with age on reserve reefs in all regions. It is hypothesized that the variation in P. leopardus sex ratios and spawning frequency among GBR regions is primarily driven by water temperature, while no‐take management zones influence spawning frequency depending on the region in which the reserve is located. Male bias and lack of spawning activity on southern GBR, where densities of adult P. leopardus are highest, suggest that recruits may be supplied from central or northern GBR. Significant regional variation in reproductive traits suggests that a regional approach to management of P. leopardus is appropriate and highlights the need for considering spatial variation in reproduction where reserves are used as fishery or conservation management tools.     

Similar regional effects among local habitats on the structure of tropical reef fish and coral communities

Aim We examined data on corals and reef fishes to determine how particular local habitat types contribute to variation in community structure across regions covering gradients in species richness and how consistent this was over time. Location Great Barrier Reef (GBR), Australia. Methods We compared large‐scale (1300 km), long‐term (11 years) data on fishes and corals that were collected annually at fixed sites in three habitats (inshore, mid‐shelf and outer‐shelf reefs) and six regions (latitudinal sectors) along a gradient of regional species richness in both communities. We used canonical approaches to partition variation in community structure (sites × species abundance data matrices) into components associated with habitat, region and time and Procrustes analyses to assess the degree of concordance between coral and fish community structure. Results Remarkably similar patterns emerged for both fish and coral communities occupying the same sites. Reefs that had similar coral communities also had similar fish communities. The fraction of the community data that could be explained by regional effects, independent of pure habitat effects, was similar in both fish (33%) and coral (36.9%) communities. Pure habitat effects were slightly greater in the fish (31.3%) than in the coral (20.1%) community. Time explained relatively little variation (fish = 7.9%, corals = 9.6%) compared with these two spatial factors. Conclusions Our results indicate either that fish and coral communities were structured in similar ways by processes associated with region, habitat and time, or that the variation in fish community structure tracked variation associated with the coral communities at these sites and thereby reflects an indirect link between the environment and the structure of fish communities mediated by corals. Irrespective of the causes of such commonality, we demonstrate that community structure, not just species richness, can be related to both habitat differences and regional setting simultaneously.     

Spatial structure of fish assemblages in a tropical estuarine lagoon: combining multivariate and geostatistical techniques

The spatial structure and seasonal changes of estuarine fish assemblages in the Ciénaga Grande de Santa Marta (CGSM) were analysed based on four seasonal comprehensive surveys conducted in 1993-1994 and 1997. Geostatistical and multivariate techniques were used to: (a) determine seasonal changes in spatial distribution of the species richness, and (b) identify assemblages of estuarine fish and their relation to abiotic factors. Potential biotic interactions affecting the assemblage structure were also explored. A total of 11075 individuals representing 39 species were collected, with Eugerres plumieri, Diapterus rhombeus, Micropogonias furnieri, Mugil incilis, Cathorops spixii, Elops saurus and Anchovia clupeoides as dominant species between seasons. Spatial distribution of fish richness differed between rainy and dry seasons in each year, whereas species mapping showed spatial patchiness in 1993-1994 and gradients during 1997. Strong evidence of species saturation was found in all seasons, suggesting biotic interactions limiting species richness at a threshold density of ca. 50 ind./5000 m². Marine, marine-estuarine and freshwater species were classified in each season according to their capability to cope with salinity fluctuations. Associations defined by functional feeding guilds were also identified. Empirical and statistic evidence showed that fish assemblages differed between seasons within each year, and each assemblage was always dominated by a small number of species, notably E. plumieri in both years 1993-1994 and 1997. Between-season differences in fish assemblage structure in the CGSM seem to be driven by abiotic factors; however, evidence of species saturation could suggest the existence of density-dependent factors operating together.     

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.     

Phylogeography of colour polymorphism in the coral reef fish Pseudochromis fuscus, from Papua New Guinea and the Great Barrier Reef

Body colour has played a significant role in the evolution of coral reef fishes, but the phylogenetic level at which colour variation is expressed and the evolutionary processes driving the development and persistence of different colour patterns are often poorly understood. The aim of this study was to examine the genetic relationships between multiple colour morphs of Pseudochromis fuscus (family Pseudochromidae), both within and among geographic locations. Pseudochromis fuscus is currently described as a single species, but exhibits at least six discrete colour morphs throughout its range. In this study, P. fuscus from Papua New Guinea (PNG) and the Great Barrier Reef (GBR), Australia, formed three genetically distinct clades based on mitochondrial DNA (control region) sequence data: (1) yellow and brown morphs from the GBR and southern PNG, as well as an orange morph from southern PNG; (2) a pink morph from southern PNG; and (3) all three morphs (pink, orange and grey) found in Kimbe Bay, northern PNG. The three groups showed deep levels of divergence (d=14.6–25.4%), suggesting that P. fuscus is a complex of polychromatic species, rather than a single widespread species with many different colour morphs. Population genetic analyses indicate that the three clades have experienced unique evolutionary histories, possibly from differential effects of sea level fluctuations, barriers to gene flow and historical biogeography.