Fish community changes after minimum flow increase: testing quantitative predictions in the Rhône River at Pierre-Bénite, France

1. Many aspects of the flow regime influence the structure of stream communities, among which the minimum discharge left in rivers has received particular attention. However, instream habitat models predicting the ecological impacts of discharge management often lack biological validation and spatial generality, particularly for large rivers with many fish species. 2. The minimum flow at Pierre‐Bénite, a reach of the Rhône river bypassed by artificial channels, was increased from 10 to 100 m³ s^?1 in August 2000 (natural mean discharge 1030 m³ s^?1), resulting in a fivefold increase in average velocity at minimum flow. Fish were electrofished in several habitat units on 12 surveys between 1995 and 2004. 3. Principal components analysis revealed a significant change in the relative abundance of fish species. The relative abundance of species preferring fast‐flowing and/or deep microhabitats increased from two‐ to fourfold after minimum flow increase. A change in community structure confirmed independent quantitative predictions of an instream habitat model. This change was significantly linked to minimum flow increase, but not to any other environmental variables describing high flows or temperature at key periods of fish life cycle. The rapidity of the fish response compared with the lifespan of individual species can be explained by a differential response of specific size classes. 4. The fish community at Pierre‐Bénite is in a transitional stage and only continued monitoring will indicate if the observed shift in community structure is perennial. We expect that our case study will be compared with other predictive tests of the impacts of flow restoration in large rivers, in the Rhône catchment and elsewhere.     

Relationships between life-history strategies of European freshwater fish species and their habitat preferences

1. Focusing on the current environmental characteristics, the ‘habitat template’ theory proposes that life‐history strategies summarise how evolution has shaped species to cope with the temporal and spatial variability of their present environment. The hierarchical ‘landscape filters’ concept predicts that the distribution of species reflects their specific traits that allow them to pass through multiple habitat filters. Together, these theories showed the importance of identifying the functional relationships of species to selective habitat forces in order to predict the organisation and response of communities to the environment. 2. We test the relationships between life‐history traits of European freshwater fish species’ and their habitat preferences, to detect the strategies adopted by fish to cope with their current habitat. For this purpose, we use published data on species traits and habitat preferences (microhabitat hydraulics, temperature and oxygen level). We use multivariate analyses to classify fish species’ strategies and test the relationships between strategies and habitat preferences. 3. We identified a continuum of life‐history patterns between two extremes, with opportunistic and periodic species. Our study supports the idea that microhabitat hydraulics plays a more important role as a template for these species ecological strategies than temperature and oxygen level. Indeed, riffle habitats may select opportunistic species whereas weak relationships are found between species strategies and both their temperature and oxygen level preferences. In addition, the ratio between mortality and growth rate (dimensionless trait), reflecting a trade‐off between growth and survival, varied among species according to the use of their hydraulic habitat, with species using deep habitats exhibiting the highest values. 4. These general relationships between hydraulics and traits may be of importance in community ecology to develop predictive models to understand how fish communities change with the hydraulic environment.     

Associations between wasp communities and forest structure: Do strong local patterns hold across landscapes?

Abstract Forest structure and habitat complexity have been used extensively to predict the distribution and abundance of insect assemblages in forest ecosystems. We tested empirically derived predictions of strong, consistent relationships between wasp assemblages and habitat complexity, using both field assessments and vegetation indices from remote sensing as measures of habitat complexity. Wasp samples from 26 paired ‘high and low’ complexity sites in two forests approximately 70 km apart, were compared with normalized difference vegetation indices (NDVIs) derived from multispectral videography of the survey sites. We describe a strong unequivocal link between habitat complexity and wasp communities, the patterns holding over coarse and fine landscape scales. NDVIs were also excellent predictors of habitat complexity and hence wasp community patterns. Sites with greater NDVIs consistently supported a greater abundance and species richness, and a different composition of wasps to sites with low NDVIs. Using vegetation indices from remote sensing to gauge habitat complexity has significant potential for ecosystem modelling and rapid biodiversity assessment.     

Simple predictions of instream habitat model outputs for fish habitat guilds in large streams

1. In the context of a generalised modification of hydraulic conditions in medium to large streams, modelling the impacts of stream regulation on fish communities in multiple streams is an important challenge for basic and applied freshwater ecology. Conventional instream habitat models such as PHABSIM link a hydraulic model with preference curves for various species to estimate habitat value changes with discharge in stream reaches. Despite world‐wide applications, they have been scarcely used in multiple sites with multiple species. 2. We assigned 21 size classes of European fish species to four habitat guilds (cluster analysis grouping size classes with comparable microhabitat preference curves). Then, we ran a conventional instream habitat model on 28 French stream reaches belonging to the `barbel zone', to estimate habitat values versus discharge curves for the 21 size classes. We summarised the outputs as mean habitat values for guilds, and tested if they were predictable from average characteristics of reaches (discharge, depth, width, particle size). 3. As was obtained elsewhere for populations, habitat values for guilds were strongly related to average, dimensionless characteristics of reaches. The Reynolds number of reaches, equivalent to a discharge per width unit, reflected most of the discharge‐dependent changes in habitat values (within reaches). In particular, habitat values of species preferring bank (respectively midstream) microhabitats decreased (respectively increased) with increasing Reynolds number. The Froude number at median discharge was the major predictor of reach‐dependent but discharge‐independent variations in habitat values. Habitat values of species preferring riffle versus pool or bank microhabitats were higher in reaches with high Froude numbers. These relationships were consistent with existing knowledge on the different species. 4. Such results suggest that the input variables required to estimate habitat values for fish communities can be greatly simplified, as illustrated by a general estimation of the sensitivity of species preferring midstream habitats to discharge changes in any reach. Cost‐efficient alternatives to conventional instream habitat models should facilitate their validation in multiple sites, a point that remains critical in instream habitat modelling of fish communities.     

Greater Sage-Grouse Nesting Habitat: The Importance of Managing at Multiple Scales

Abstract: Considering habitat selection at multiple scales is essential to fully understand habitat requirements and management needs for wildlife species of concern. We used a hierarchical information-theoretic approach and variance decomposition techniques to analyze habitat selection using local-scale habitat variables measured in the field and landscape-scale variables derived with a Geographic Information System (GIS) for nesting greater sage-grouse (Centrocercus urophasianus) in the Powder River Basin (PRB), Montana and Wyoming, USA, 2003–2007. We investigated relationships between habitat features that can and cannot be mapped in a GIS to provide insights into interpretation of landscape-scale—only GIS models. We produced models of habitat selection at both local and landscape scales and across scales, yet multiscale models had overwhelming statistical and biological support. Variance decomposition showed that local-scale measures explained the most pure variation (50%) in sage-grouse nesting-habitat selection. Landscape-scale features explained 20% of pure variation and shared 30% with local-scale features. Both local- and landscape-scale habitat features are important in sage-grouse nesting-habitat selection because each scale explained both pure and shared variation. Our landscape-scale model was accurate in predicting priority landscapes where sage-grouse nests would occur and is, therefore, useful in providing landscape context for management decisions. It accurately predicted locations of independent sage-grouse nests (validation R² = 0.99) and showed good discriminatory ability with >90% of nests located within only 40% of the study area. Our landscape-scale model also accurately predicted independent lek locations. We estimated twice the amount of predicted nesting habitat within 3 km of leks compared to random locations in the PRB. Likewise we estimated 1.8 times more predicted nesting habitat within 10 km of leks compared to random locations. These results support predictions of the hotspot theory of lek placement. Local-scale habitat variables that cannot currently be mapped in a GIS strongly influence sage-grouse nest-site selection, but only within priority nesting habitats defined at the landscape scale. Our results indicate that habitat treatments for nesting sage-grouse applied in areas with an unsuitable landscape context are unlikely to achieve desired conservation results.     

Habitat amount hypothesis and passive sampling explain mammal species composition in Amazonian river islands

Nested structures of species assemblages have been frequently associated with patch size and isolation, leading to the conclusion that colonization–extinction dynamics drives nestedness. The ‘passive sampling’ model states that the regional abundance of species randomly determines their occurrence in patches. The ‘habitat amount hypothesis’ also challenges patch size and isolation effects, arguing that they occur because of a ‘sample area effect’. Here, we (a) ask whether the structure of the mammal assemblages of fluvial islands shows a nested pattern, (b) test whether species’ regional abundance predicts species’ occurrence on islands, and (c) ask whether habitat amount in the landscape and matrix resistance to biological flow predict the islands’ species composition. We quantified nestedness and tested its significance using null models. We used a regression model to analyze whether a species’ relative regional abundance predicts its incidence on islands. We accessed islands’ species composition by an NMDS ordination and used multiple regression to evaluate how species composition responds to habitat amount and matrix resistance. The degree of nestedness did not differ from that expected by the passive sampling hypothesis. Likewise, species’ regional abundance predicted its occurrence on islands. Habitat amount successfully predicted the species composition on islands, whereas matrix resistance did not. We suggest the application of habitat amount hypothesis for predicting species composition in other patchy systems. Although the island biogeography perspective has dominated the literature, we suggest that the passive sampling perspective is more appropriate for explaining the assemblages’ structure in this and other non‐equilibrium patch systems. Abstract in Portuguese is available with online material.     

Simple predictions of instream habitat model outputs for target fish populations

1. Bottom-up approaches based on individual behaviour can help to identify key variables influencing populations at larger scales. Instream habitat models have been developed to predict the consequences, for populations in stream reaches, of fish preferences for particular hydraulic conditions observed at the scale of individuals. Conventional instream habitat models (e.g. PHABSIM) predict habitat values for species or life stages in reaches, and their changes with discharge. Despite their worldwide use, they have been subject to continuing criticism and have been mainly limited to site-specific case studies.
2. We ran conventional instream habitat models in 58 French stream reaches dominated by brown trout. Using non-linear mixed effect models, we demonstrated that the outputs of instream habitat models (habitat values for three trout life stages and five other species) are predictable from average characteristics of reaches (discharge, depth, width and bed particle size).
3. Our models closely reflect variations in habitat values within-reaches (with discharge) and between-reaches. Within-reach changes are linked to the Reynolds number of reaches, while between-reach changes depend mainly on the Froude number at median daily discharge. These two dimensionless variables combine discharge, mean depth and mean width of reaches. Independent model validations showed robust model predictions that are consistent with studies of habitat values for brown trout made in larger streams from western North America.
4. Our results contribute to identifying the main hydraulic variables governing estimates of fish habitat values. They should facilitate habitat studies in multiple streams, at the basin or larger scales, while reducing their cost. They should enhance the biological validation of habitat model predictions, which remains critical.     

Patterns of fish community composition along a river affected by agricultural and urban disturbance in south-central Chile

Patterns of fish community composition in a south-central Chile river were investigated along the altitudinal-spatial and environmental gradient and as a function of anthropogenic factors. The spatial pattern of fish communities in different biocoenotic zones of the Chillan River is influenced by both natural factors such a hydrologic features, habitat, and feeding types, and also by water quality variables which can reduce the diversity and abundance of sensitive species. A principal component analysis incorporating both water quality parameters and biomarker responses of representative fish species was used to evaluate the status of fish communities along the spatial gradient of the stream. The abundance and diversity of the fish community changed from a low in the upper reaches where the low pollution-tolerant species such as salmonid dominated, to a reduced diversity in the lower reaches of the river where tolerant browser species such as cypriniformes dominated. Even though the spatial pattern of fish community structure is similar to that found for the Chilean Rivers, the structure of these communities is highly influenced by human disturbance, particularly along the lower reaches of the river. Handling editor: C. Sturmbauer     

Habitat associations of imperilled fishes after conservation intervention in the Cape Fold Ecoregion,South Africa

The aim of this study was to examine the habitat associations of a native cyprinid community of the recovering Rondegat River in the Cape Fold Ecoregion of South Africa as part of a long-term native fish abundance monitoring project. Relative abundance data were extracted from underwater video camera footage across the longitudinal gradient of the river in three sampling instances. Using multivariate methods the authors assessed community composition with respect to habitat, its overlap with a protected area and species-specific abiotic predictors of relative abundance. Distance from the uppermost site in the river was the most significant predictor of species abundance, indicating spatial segregation and varying overlap between species. The protected status of sites in the upper reaches, vegetated substrates and the size of individual sites were the most impactful on the relative abundance of the endangered fiery redfin Pseudobarbus phlegethon. The results of this study indicate that underwater video monitoring is an effective and low-cost approach that can inform conservation recommendations. Reducing agricultural runoff and sedimentation in the lower reaches may be useful further interventions to maintain key habitats of submerged vegetation.     

Influence of habitat heterogeneity on the distribution of larval Pacific lamprey (Lampetra tridentata) at two spatial scales

1. Spatial patterns in channel morphology and substratum composition at small (1–10 metres) and large scales (1–10 kilometres) were analysed to determine the influence of habitat heterogeneity on the distribution and abundance of larval lamprey. 2. We used a nested sampling design and multiple logistic regression to evaluate spatial heterogeneity in the abundance of larval Pacific lamprey, Lampetra tridentata, and habitat in 30 sites (each composed of twelve 1‐m² quadrat samples) distributed throughout a 55‐km section of the Middle Fork John Day River, OR, U.SA. Statistical models predicting the relative abundance of larvae both among sites (large scale) and among samples (small scale) were ranked using Akaike's Information Criterion (AIC) to identify the ‘best approximating’ models from a set of a priori candidate models determined from the literature on larval lamprey habitat associations. 3. Stream habitat variables predicted patterns in larval abundance but played different roles at different spatial scales. The abundance of larvae at large scales was positively associated with water depth and open riparian canopy, whereas patchiness in larval occurrence at small scales was associated with low water velocity, channel‐unit morphology (pool habitats), and the availability of habitat suitable for burrowing. 4. Habitat variables explained variation in larval abundance at large and small scales, but locational factors, such as longitudinal position (river km) and sample location within the channel unit, explained additional variation in the logistic regression model. The results emphasise the need for spatially explicit analysis, both in examining fish habitat relationships and in developing conservation plans for declining fish populations.     

Influence of stream geomorphic condition on fish communities in Vermont, U.S.A.

1. Evaluations of stream geomorphic condition may increase our understanding of the composite effects of human‐induced habitat change on fish communities. Using systematic sampling of 44 reaches spread across 26 rivers in Vermont from 2002 through 2004, we tested the hypothesis that stream reaches in reference geomorphic condition would support fish assemblages that differed in diversity and productivity from fish communities found in reaches of poorer geomorphic condition. 2. At each study reach, we sampled the fish community, identified the morphological unit according to common stream classification systems and then evaluated the extent of deviation from reference geomorphic condition using a regionally adapted geomorphic assessment methodology. 3. We used principal component analysis (PCA) and linear regression to build exploratory models linking stream geomorphic condition to fish community characteristics. 4. Our results suggest that geomorphic condition significantly influences fish community diversity, productivity and condition. Geomorphic condition was a significant factor in all of our fish community models. In conjunction with additional reach characteristics, geomorphic condition explained up to 31% of the total variance observed in models for species diversity of fish communities, 44% of the variance in assemblage biomass and 45% of the variance in a regional index of biotic integrity. 5. Our work builds on single‐species evidence that geomorphic characteristics represent important local‐scale fish‐habitat variables, showing that stream geomorphic condition is a dominant factor affecting entire fish communities. Our results enhance our understanding of the hierarchy of factors that influences fish community diversity and organisation and support the use of geomorphic condition assessments in stream management.     

The Structures and Dynamics of Fish Communities in an Okinawan Coral Reef: Effects of Coral-based Habitat Structures at Sites with Rocky and Sandy Sea Bottoms

Although there have been many studies on ecological factors responsible for the organization of reef fish communities, most of the studies have focused on isolated habitats. However, findings from isolated habitats cannot necessarily be applied to fish communities in other habitats (e.g., a continuous habitat). In this study, therefore, we examined the structures of fish communities (abundance, species richness and species composition), and the dynamics of fish communities (seasonal changes in abundance, species richness and species composition) over a 2-year period in two different habitats (continuous habitat and isolated habitat) in an Okinawan coral reef. We established eight permanent quadrats (8m × 8m) on a rocky reef flat (continuous habitat) and rock reef patches surrounded by a sandy sea bottom (isolated habitat). The abundance and species richness of fishes such as pomacentrids, labrids, chaetodontids and acanthurids were greater in quadrats located in the continuous habitat, whereas those of blenniids, gobiids and mullids were greater in quadrats located in the isolated habitat. This caused marked differences between the fish community structure at the two sites. Seasonal and annual changes in fish community structure were relatively small at the continuous habitat site (>0.7 similarity based on C index) but were large at the isolated habitat site (C < 0.4), indicating that the fish community structure was relatively stable at the continuous habitat site but unstable at the isolated habitat site throughout the study period. Spatial differences between the fish community structures in the four quadrats at the continuous habitat site were small (C > 0.65 in most sites), but these differences were large at the isolated habitat site (C < 0.4). Our findings suggest that habitat structure (spatial arrangement of habitats) affects both spatial and seasonal differences in the reef fish community structure. The results also suggest that the main mechanisms underlying organization of reef fish communities in continuous and isolated habitats are different.     

Stream communities across a rural–urban landscape gradient

Rapid urbanization throughout the world is expected to cause extensive loss of biodiversity in the upcoming decades. Disturbances associated with urbanization frequently operate over multiple spatial scales such that local species extirpations have been attributed both to localized habitat degradation and to regional changes in land use. Urbanization also may shape stream communities by restricting species dispersal within and among stream reaches. In this patch-dynamics view, anthropogenic disturbances and isolation jointly reduce stream biodiversity in urbanizing landscapes. We evaluated predictions of stream invertebrate community composition and abundance based on variation in environmental conditions at five distinct spatial scales: stream habitats, reaches, riparian corridors and watersheds and their spatial location within the larger three-river basin. Despite strong associations between biodiversity loss and human density in this study, local stream habitat and stream reach conditions were poor predictors of community patterns. Instead, local community diversity and abundance were more accurately predicted by riparian vegetation and watershed landscape structure. Spatial coordinates associated with instream distances provided better predictions of stream communities than any of the environmental data sets. Together, results suggest that urbanization in the study region was associated with reduced stream invertebrate diversity through the alteration of landscape vegetation structure and patch connectivity. These findings suggest that maintaining and restoring watershed vegetation corridors in urban landscapes will aid efforts to conserve freshwater biodiversity.     

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.     

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.     

Theoretical habitat templets, species traits, and species richness: fish in the Upper Rhône River and its floodplain

• 1 Recent developments in ecological theory concerned with habitat templets, species assemblages, and life history traits were examined for the riverine fish communities of the Upper Rhône River, France, in the context of spatial–temporal habitat variability. Relationships among species traits, habitat utilization of species, the relationship between species traits and habitat utilization, and trends of species traits and species richness in the spatial–temporal variability of the habitat types were analysed.
• 2 Relationships among twelve species traits, and utilization of eight habitats were examined for twenty-five fish species using correspondence analysis; the relationship between species traits and habitat utilization was investigated by co-inertia analysis.
• 3 Positive relationships among species traits were observed for size, fecundity, and the number of reproductive cycles per individual. However, species were not well differentiated according to the habitat utilization, except for habitats rarely connected with the main channel (i.e. two types of oxbow lakes).
• 4 No significant relationship was found between species traits and habitat utilization, nor for either species traits or species richness when examined in the framework of spatial–temporal habitat variability. Only two species traits corresponded (with slight trends) to predictions in a river habitat templet: (i) the number of descendants per reproductive cycle increased along with temporal variability; and (ii) the number of reproductive cycles per individual was either low or high at low temporal variability and intermediate at elevated temporal variability.
• 5 The discrepancy between the predictions of the river habitat templet as well as of the patch dynamics concept and the results observed for the fish in the Upper Rhône was explained in terms of scale problems, the evolutionary ecology of the European fish fauna, and the history of the Rhône River.

     

Using landscape history to predict biodiversity patterns in fragmented landscapes

Landscape ecology plays a vital role in understanding the impacts of land‐use change on biodiversity, but it is not a predictive discipline, lacking theoretical models that quantitatively predict biodiversity patterns from first principles. Here, we draw heavily on ideas from phylogenetics to fill this gap, basing our approach on the insight that habitat fragments have a shared history. We develop a landscape ‘terrageny’, which represents the historical spatial separation of habitat fragments in the same way that a phylogeny represents evolutionary divergence among species. Combining a random sampling model with a terrageny generates numerical predictions about the expected proportion of species shared between any two fragments, the locations of locally endemic species, and the number of species that have been driven locally extinct. The model predicts that community similarity declines with terragenetic distance, and that local endemics are more likely to be found in terragenetically distinctive fragments than in large fragments. We derive equations to quantify the variance around predictions, and show that ignoring the spatial structure of fragmented landscapes leads to over‐estimates of local extinction rates at the landscape scale. We argue that ignoring the shared history of habitat fragments limits our ability to understand biodiversity changes in human‐modified landscapes.     

马鞍列岛岩礁生境鱼类群落结构时空格局

基于2009年马鞍列岛潮下带岩礁生境的多网目三层组合刺网逐月调查数据,对鱼类群落月相和季度间的变化、区域尺度上的空间差异和群落的稳态进行了探讨,应用优势种相对丰度和生物量、定居性鱼类和洄游种的季节动态、非度量多维标度(nMDS)和生物量丰度曲线(ABC)分析方法对群落时空格局进行了分析。结果显示:季节性洄游种集中出现在夏秋季,但对岩礁生境的利用表现出不同区域的强度差异;小黄鱼Larimichthys polyactis和黄姑鱼Nibea albiflora周年利用岩礁生境,但强度有别,尤其在冬季;定居性鱼类褐菖鲉Sebastiscus marmoratus、斑头鱼Agrammus agrammus和大泷六线鱼Hexagrammos otakii在春末夏初的种群密度最高,同样表现出某些或大部分月份的区域差异;而河口种中国花鲈Lateolabrax maculatus则更多地选择秋冬季出现在岩礁生境。多元分析结果揭示了当地岩礁生境鱼类群落格局上显著的季节和区域差异,这是定居性鱼类对岩礁生境利用的阶段性变化和区域差异、结合洄游种季节迁移和选择性分布等因素作用下共同形成的格局。ABC分析进一步发现,丰富的鱼类生态类型形成了夏季岩礁生境更为稳定的群落状态,而冬季相反;同时各季度A区的群落干扰明显强过B区。研究表明,产卵季节海域西北部的岩礁生境很可能起着主要的产卵场功能,而东南部区域各季度皆侧重于幼鱼育肥场的功能表达。丰富的种类区系加之优势种的季节性交替出现,共同塑造了岩礁生境极具动态的鱼类群落格局,对维持岛礁海域鱼类多样性以及局部区域生态系统的稳定性起着非常重要的作用。