Microhabitat selectivity underpins regional indicators of fish abundance and replenishment

Species with specialized resource use can display strong spatial heterogeneity in abundance according to the availability of their preferred habitats. If these preferences shift with ontogeny, then a wide range of habitats may need to be protected in order to support both adult populations and their replenishment. We explored whether microhabitat selectivity interacts with habitat availability to provide an effective suite of indicators for regional fish abundance and replenishment, using offshore rocky reefs in south-eastern Australia as a case study. We examined generalized additive mixed models (GAMMs) in a full subsets approach to infer the best predictors for adult and juvenile fish density in four diverse families (Labridae, Odacidae, Pomacentridae, Serranidae), based on rapid underwater visual surveys across transects (⿼500 m²), wave exposures (0.3⿿1 km), and sites (0.3⿿48 km). We then examined whether these regional fish-habitat models aligned with the microhabitat electivity of individuals (at scale of <1 m²). Microhabitat selection by reef fishes at the local scale underpinned the most effective habitat indicators for regional heterogeneity in fish abundance, and pointed to critical nursery habitats that support hotspots of juvenile recruitment. Strong species-habitat relationships, such as these, can be combined with broad-scale habitat mapping to assess the potential carrying capacity of focal areas, spatial management zone placements, and nursery habitats that warrant special protection. A number of emerging threats to these key habitat types indicates an urgent need for habitat-based protection and monitoring as a key part of holistic marine ecosystem conservation and management.     

Spatial predictability of juvenile fish species richness and abundance in a coral reef environment

Juvenile reef fish communities represent an essential component of coral reef ecosystems in the current focus of fish population dynamics and coral reef resilience. Juvenile fish survival depends on habitat characteristics and is, following settlement, the first determinant of the number of individuals within adult populations. The goal of this study was to provide methods for mapping juvenile fish species richness and abundance into spatial domains suitable for micro and meso-scale analysis and management decisions. Generalized Linear Models predicting juvenile fish species richness and abundance were developed according to spatial and temporal environmental variables measured from 10 m up to 10 km in the southwest lagoon of New Caledonia. The statistical model was further spatially generalized using a 1.5-m resolution, independently created, remotely sensed, habitat map. This procedure revealed that : (1) spatial factors at 10 to 100-m scale explained up to 71% of variability in juvenile species richness, (2) a small improvement (75%) was gained when a combination of environmental variables at different spatial and temporal scales was used and (3) the coupling of remotely sensed data, geographical information system tools and point-based ecological data showed that the highest species richness and abundance were predicted along a narrow margin overlapping the coral reef flat and adjacent seagrass beds. Spatially explicit models of species distribution may be relevant for the management of reef communities when strong relationships exist between faunistic and environmental variables and when models are built at appropriate scales.     

The relative influence of local to regional drivers of variation in reef fishes

In this study, fishes and habitat attributes were quantified, four times over 1 year, on three reefs within four regions encompassing a c. 6° latitudinal gradient across south-western Australia. The variability observed was partitioned at these spatio-temporal scales in relation to reef fish variables and the influence of environmental drivers quantified at local scales, i.e. at the scale of reefs (the number of small and large topographic elements, the cover of kelp, fucalean and red algae, depth and wave exposure) and at the scale of regions (mean and maximum nutrient concentrations and mean seawater temperature) with regard to the total abundance, species density, species diversity and the multivariate structure of reef fishes. Variation in reef fish species density and diversity was significant at the regional scale, whereas variation in the total abundance and assemblage structure of fishes was also significant at local scales. Spatial variation was greater than temporal variation in all cases. A systematic and gradual species turnover in assemblage structure was observed between adjacent regions across the latitudinal gradient. The cover of red algae within larger patches of brown macroalgae (a biological attribute of the reef) and the number of large topographic elements (a structural attribute of the reef) were correlated with variation observed at local scales, while seawater temperature correlated with variation at the scale of regions. In conclusion, conservation efforts on reef fishes need to incorporate processes operating at regional scales with processes that shape local reef fish communities at local scales.     

Relative Importance of Coral Cover,Habitat Complexity and Diversity in Determining the Structure of Reef Fish Communities

The structure of coral reef habitat has a pronounced influence on the diversity, composition and abundance of reef-associated fishes. However, the particular features of the habitat that are most critical are not always known. Coral habitats can vary in many characteristics, notably live coral cover, topographic complexity and coral diversity, but the relative effects of these habitat characteristics are often not distinguished. Here, we investigate the strength of the relationships between these habitat features and local fish diversity, abundance and community structure in the lagoon of Lizard Island, Great Barrier Reef. In a spatial comparison using sixty-six 2m² quadrats, fish species richness, total abundance and community structure were examined in relation to a wide range of habitat variables, including topographic complexity, habitat diversity, coral diversity, coral species richness, hard coral cover, branching coral cover and the cover of corymbose corals. Fish species richness and total abundance were strongly associated with coral species richness and cover, but only weakly associated with topographic complexity. Regression tree analysis showed that coral species richness accounted for most of the variation in fish species richness (63.6%), while hard coral cover explained more variation in total fish abundance (17.4%), than any other variable. In contrast, topographic complexity accounted for little spatial variation in reef fish assemblages. In degrading coral reef environments, the potential effects of loss of coral cover and topographic complexity are often emphasized, but these findings suggest that reduced coral biodiversity may ultimately have an equal, or greater, impact on reef-associated fish communities.     

Predation on reef-building corals: multiscale variation in the density of three corallivorous gastropods, Drupella spp.

 Feeding aggregations of three corallivorous gastropods, Drupella cornus, D. fragum and D. rugosa, have caused considerable coral damage on reefs across the Indo-West Pacific. Distribution and abundance of these three species were explored at Lizard Island, Great Barrier Reef, to determine within-reef variations in density, and spatial relationships between Drupella and their prey corals. The scales of greatest variation were between reef habitats (combinations of exposure and depth) and individual coral colonies. Density varied 12-fold among four habitats: exposed crests (2.55/m²), exposed slopes (0.22/m²), sheltered crests (0.34/m²) and sheltered slopes (2.07/m²). Species composition also varied markedly between habitats. Individuals were highly aggregated, usually forming small clusters (<10 individuals) on live coral colonies and other substrata, and occasional large aggregations of=200 to>2000. Five basic tenets for sampling Drupella are established, based on patterns of variation in density and species composition, and small-scale habitat use. Accepted: 8 February 1999     

Environmental factors influencing urchin spatial distributions on disturbed coral reefs (New Caledonia, South Pacific)

Few works have examined the relative contributions of habitat variables to the distribution of coral reef urchins. In the present study, the spatial distribution of two common urchin species (Diadema setosum and Echinometra mathaei) was studied in the fringing reefs of two urban bays in New Caledonia (South Pacific). Urchins were surveyed at 105 stations with contrasted habitat structure/anthropic disturbance levels; 32 environmental variables (water/sediment characteristics, reef structuring species) were considered. Moderate densities were generally observed at station scale (mean 0.5 individuals m^− 2). The combination of univariate and multivariate techniques highlighted patchy distributions for Diadema as well as Echinometra, with distinct species/habitat associations; environmental gradients occurring within the bays did not seem to influence the species patterns. For Diadema, the spatial variability was better explained by sediment type than by biotic cover; increasing densities occurred across habitats with larger sediment sizes and decreasing coral complexity/macrophytes cover. In contrast, the distribution of E. mathaei exhibited weak relationships with habitat variables. In coral reefs, small-scale heterogeneity may thus be responsible for most of urchins spatial variability.     

Hierarchical and scale‐dependent effects of fishing pressure and environment on the structure and size distribution of parrotfish communities

Parrotfishes are considered to have a major influence on coral reef ecosystems through grazing the benthic biota and are also primary fishery targets in the Indo‐Pacific. Consequently, the impact of human exploitation on parrotfish communities is of prime interest. As anthropogenic and environmental factors interact across spatial scales, sampling programs designed to disentangle these are required by both ecologists and resource managers. We present a multi‐scale examination of patterns in parrotfish assemblage structure, size distribution and diversity across eight oceanic islands of Micronesia. Results indicate that correlates of assemblage structure are scale‐dependent; biogeographic distributions of species and island geomorphology hierarchically influenced community patterns across islands whereas biophysical features and anthropogenic pressure influenced community assemblage structure at the within‐island scale. Species richness and phylogenetic diversity increased with greater broad‐scale habitat diversity associated with different island geomorphologies. However, within‐island patterns of abundance and biomass varied in response to biophysical factors and levels of human influence unique to particular islands. While the effect of fishing activities on community composition and phylogenetic diversity was obscured across island types, fishing pressure was the primary correlate of mean parrotfish length at all spatial scales. Despite widespread fishery‐induced pressure on Pacific coral reefs, the structuring of parrotfish communities at broad spatial scales remains a story largely dependent on habitat. Thus, we propose better incorporation of scale‐dependent habitat effects in future assessments of overexploitation on reef fish assemblages. However, strong community‐level responses within islands necessitate an improved understanding of the phylogenetic and functional consequences of altering community structure.     

Effect of simulated macroalgae on the fish assemblage associated with a temperate reef system

Increased habitat complexity is supposed to promote increased diversity, abundance and biomass. This study tested the effect of the macroalgal cover on temperate reef fishes by mimicking macroalgae on artificial reefs in NW Sicily (Mediterranean Sea). Macroalgal cover affected reef fishes in different ways and independently of intrinsic temporal trends. The fish assemblages of manipulated and control artificial reef units differed in the relative abundances of the associated species, but little in species composition. In line with studies in seagrass habitats, fishes were most abundant in reefs covered by artificial macroalgae. Three species (Boops boops, Serranus scriba and Symphodus ocellatus) exhibited consistently greater abundance on vegetated reef units than on control reef units. The total number of species and the abundance of three particular species (S. scriba, S. ocellatus and Thalassoma pavo) displayed temporal trends which were independent on short and large temporal scales. Only fish total biomass and one species (Spicara flexuosa) displayed strong effects of interaction among the experimental factors. Mechanisms to explain these findings are discussed from observational evidence on habitat use and interactions among multiple species. This study highlights that manipulative experiments involving repeated sampling of fish in artificial habitats appear to be a valid approach to study fish-habitat relationships in fluctuating environments. It is also concluded that macroalgae mimics may serve as a tool for restoring lost marine vegetated habitats when current human-induced conditions prevent the recovery of pristine macroalgal stands.     

BIODIVERSITY RESEARCH: Native‐exotic species richness relationships across spatial scales and biotic homogenization in wetland plant communities of Illinois,USA

Aim To examine native‐exotic species richness relationships across spatial scales and corresponding biotic homogenization in wetland plant communities. Location Illinois, USA. Methods We analysed the native‐exotic species richness relationship for vascular plants at three spatial scales (small, 0.25 m² of sample area; medium, 1 m² of sample area; large, 5 m² of sample area) in 103 wetlands across Illinois. At each scale, Spearman’s correlation coefficient between native and exotic richness was calculated. We also investigated the potential for biotic homogenization by comparing all species surveyed in a wetland community (from the large sample area) with the species composition in all other wetlands using paired comparisons of their Jaccard’s and Simpson’s similarity indices. Results At large and medium scales, native richness was positively correlated with exotic richness, with the strength of the correlation decreasing from the large to the medium scale; at the smallest scale, the native‐exotic richness correlation was negative. The average value for homogenization indices was 0.096 and 0.168, using Jaccard’s and Simpson’s indices, respectively, indicating that these wetland plant communities have been homogenized because of invasion by exotic species. Main Conclusions Our study demonstrated a clear shift from a positive to a negative native‐exotic species richness relationship from larger to smaller spatial scales. The negative native‐exotic richness relationship that we found is suggested to result from direct biotic interactions (competitive exclusion) between native and exotic species, whereas positive correlations likely reflect the more prominent influence of habitat heterogeneity on richness at larger scales. Our finding of homogenization at the community level extends conclusions from previous studies having found this pattern at much larger spatial scales. Furthermore, these results suggest that even while exhibiting a positive native‐exotic richness relationship, community level biotas can/are still being homogenized because of exotic species invasion.     

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.     

Retention of Habitat Complexity Minimizes Disassembly of Reef Fish Communities following Disturbance: A Large-Scale Natural Experiment

High biodiversity ecosystems are commonly associated with complex habitats. Coral reefs are highly diverse ecosystems, but are under increasing pressure from numerous stressors, many of which reduce live coral cover and habitat complexity with concomitant effects on other organisms such as reef fishes. While previous studies have highlighted the importance of habitat complexity in structuring reef fish communities, they employed gradient or meta-analyses which lacked a controlled experimental design over broad spatial scales to explicitly separate the influence of live coral cover from overall habitat complexity. Here a natural experiment using a long term (20 year), spatially extensive (∼115,000 kms²) dataset from the Great Barrier Reef revealed the fundamental importance of overall habitat complexity for reef fishes. Reductions of both live coral cover and habitat complexity had substantial impacts on fish communities compared to relatively minor impacts after major reductions in coral cover but not habitat complexity. Where habitat complexity was substantially reduced, species abundances broadly declined and a far greater number of fish species were locally extirpated, including economically important fishes. This resulted in decreased species richness and a loss of diversity within functional groups. Our results suggest that the retention of habitat complexity following disturbances can ameliorate the impacts of coral declines on reef fishes, so preserving their capacity to perform important functional roles essential to reef resilience. These results add to a growing body of evidence about the importance of habitat complexity for reef fishes, and represent the first large-scale examination of this question on the Great Barrier Reef.     

The effectiveness of different meso-scale rugosity metrics for predicting intra-habitat variation in coral-reef fish assemblages

Habitat structure is frequently an important variable affecting species’ abundances and diversity, and identifying the key aspects and spatial scales of habitat complexity is critical for understanding the ecology and conservation of a range of communities. Many coral-reef fishes are intimately linked with benthic habitat structure, and previous research has demonstrated rugosity as an important predictive variable of assemblage parameters. However, these studies typically consider rugosity at small scales, amalgamate multiple habitat features, or are semi-quantitative. This study considers meso-scale rugosity (within 51 plots of 25 m² on a Belizean forereef) generated by varying coral densities, heights, and complexities. Seven rugosity metrics were calculated for each plot, and were regressed against each of 11 fish assemblage parameters. Intra-habitat variability of each fish parameter was significantly positively or curvilinearly correlated to at least one metric of meso-scale rugosity, but the metric generating the strongest correlation varied. The abundance of small fishes, and consequently most of the assemblage statistics (e.g., total fish abundance and diversity) were best predicted by the number of tall (>50 cm) corals. The abundance of damselfishes, parrotfish biomass, the abundances of medium-sized and large fishes, and total fish biomass were curvilinearly related to mean coral height. The abundances of wrasses and surgeonfishes were most strongly correlated with the number of corals within a plot. Because coral-generated meso-scale rugosity is an important factor influencing intra-habitat variation in fishes, it should be explicitly considered when investigating fish-habitat relationships and predicting the impacts of coral mortality on ecosystem processes and services.     

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

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

Patterns of species density and productivity at different spatial scales in herbaceous plant communities

A major challenge in evaluating patterns of species richness and productivity involves acquiring data to examine these relationships empirically across a range of ecologically significant spatial scales. In this paper, we use data from herb‐dominated plant communities at six Long‐Term Ecological Research (LTER) sites to examine how the relationship between plant species density and above‐ground net primary productivity (ANPP) differs when the spatial scale of analysis is changed. We quantified this relationship at different spatial scales in which we varied the focus and extent of analysis: (1) among fields within communities, (2) among fields within biomes or biogeographic regions, and (3) among communities within biomes or biogeographic regions. We used species density (D=number of species per m²) as our measure of diversity to have a comparable index across all sites and scales. Although we expected unimodal relationships at all spatial scales, we found that spatial scale influenced the form of the relationship. At the scale of fields within different grassland communities, we detected a significant relationship at only one site (Minnesota old‐fields), and it was negative linear. When we expanded the extent of analyses to biogeographic regions (grasslands or North America), we found significant unimodal relationships in both cases. However, when we combined data to examine patterns among community types within different biogeographic regions (grassland, alpine tundra, arctic tundra, or North America), we did not detect significant relationships between species density and ANPP for any region. The results of our analyses demonstrate that the spatial scale of analysis – how data are aggregated and patterns examined – can influence the form of the relationship between species density and productivity. It also demonstrates the need for data sets from a broad spectrum of sites sampled over a range of scales for examining challenging and controversial ecological hypotheses.     

Non-Random Variability in Functional Composition of Coral Reef Fish Communities along an Environmental Gradient

Changes in the coral reef complex can affect predator-prey relationships, resource availability and niche utilisation in the associated fish community, which may be reflected in decreased stability of the functional traits present in a community. This is because particular traits may be favoured by a changing environment, or by habitat degradation. Furthermore, other traits can be selected against because degradation can relax the association between fishes and benthic habitat. We characterised six important ecological traits for fish species occurring at seven sites across a disturbed coral reef archipelago in Indonesia, where reefs have been exposed to eutrophication and destructive fishing practices for decades. Functional diversity was assessed using two complementary indices (FRic and RaoQ) and correlated to important environmental factors (live coral cover and rugosity, representing local reef health, and distance from shore, representing a cross-shelf environmental gradient). Indices were examined for both a change in their mean, as well as temporal (short-term; hours) and spatial (cross-shelf) variability, to assess whether fish-habitat association became relaxed along with habitat degradation. Furthermore, variability in individual traits was examined to identify the traits that are most affected by habitat change. Increases in the general reef health indicators, live coral cover and rugosity (correlated with distance from the mainland), were associated with decreases in the variability of functional diversity and with community-level changes in the abundance of several traits (notably home range size, maximum length, microalgae, detritus and small invertebrate feeding and reproductive turnover). A decrease in coral cover increased variability of RaoQ while rugosity and distance both inversely affected variability of FRic; however, averages for these indices did not reveal patterns associated with the environment. These results suggest that increased degradation of coral reefs is associated with increased variability in fish community functional composition resulting from selective impacts on specific traits, thereby affecting the functional response of these communities to increasing perturbations.     

A Large-scale, Hierarchical Approach for Assessing Habitat Associations of Fish Assemblages in Large Dryland Rivers

Multiple-scale assessments of fish-habitat associations are limited despite the fact that riverine fish assemblages are influenced by factors operating over a range of spatial scales. A method for assessing fish-habitat assemblages at multiple scales is proposed and tested in a large Australian dryland river, the Barwon–Darling River. Six discrete mesohabitat types (large wood, smooth bank, irregular bank, matted bank, mid-channel and deep pool) nested within 10 km long river reaches were sampled. Individual reaches were, in turn, nested within four larger geomorphological zones, previously identified along the river. Fish assemblages varied significantly between mesohabitat types and at different spatial scales. Golden perch (Macquaria ambigua), Murray cod (Maccullochella peelii peelii) and common carp (Cyprinus carpio) were strongly associated with large wood, but golden perch and Murray cod exhibited higher habitat specificity than carp. Bony herring (Nematalosa erebi) were more common in shallow edgewater habitats. At the river-scale, regional differences in the fish assemblage occurred at scales closely corresponding to geomorphological zones and these differences were associated with changes in the relative abundance of species rather than the addition or replacement of species. The proposed hierarchical framework improves the efficiency of fish surveys in large rivers by viewing meso-scale fish-habitat associations in the context of larger-scale geomorphological processes.     

Multiscale habitat associations of detrivorous blennies (Blenniidae: Salariini)

The distribution and habitat associations of detrivorous blennies on a tropical coral reef were investigated at several spatial scales and compared with other fish that feed on the epilithic algal matrix to assess density and biomass contributions of small detrivorous fishes to these assemblages. At broad spatial scales total blenny abundance and biomass were highest on the tops of reefs exposed to prevailing winds. On the finer scale of microhabitat use, all species showed a preference for non-living corals, although the type of coral utilised differed between species. The high abundance of blennies on reef tops and non-living corals may be partially related to the quality and availability of detritus in these habitats. Comparisons of total blenny abundance and biomass with other territorial detrivores found that blennies accounted for approximately 60% of this functional group's density and 21% of their biomass on exposed reef tops. Overall, territorial detrivores were found to constitute approximately 37% of the density and 26% of the biomass of the detrivorous/herbivorous fish assemblage on exposed reef tops. Small detrivorous fish therefore represent a substantial proportion of fish assemblages that feed on epilithic algae and associated detritus on coral reefs.     

Reef fish and habitat relationships in a Caribbean seascape: the importance of reef context

Marine protected area (MPA) effectiveness is contingent on understanding key ecological patterns and processes at appropriate spatial scales and may depend upon maintaining critical linkages among essential habitat patches to conserve reef-fish communities. Hypotheses were tested to investigate the importance of habitat linkages in the US Virgin Islands. As expected, reef context (the spatial pattern of surrounding habitat patches) was a strong predictor of reef fish assemblage structure. Specific relationships were functionally consistent with the ecology of the fishes of interest. For example, reefs with large amounts of seagrass nearby harbored the greatest numerical abundance of fishes, particularly mobile invertebrate feeders and the exploited fish families of Haemulidae (grunts) and Lutjanidae (snappers). Species richness for the entire fish community and within these fish groups was also strongly associated with reef context. Furthermore, reef fish mobility influenced how fishes related to reef context. Fish-habitat relationships were detected as far as 1 km from study reefs, suggesting that fish movements result in habitat encounter rates that may influence their patterns of distribution. Consequently, functional habitat connectivity of habitat patches appears important in structuring reef-fish assemblages, and suggests that landscape-scale metrics may provide insights useful to managers in the design of MPAs.