Global patterns and predictors of tropical reef fish species richness

In the marine realm, the tropics host an extraordinary diversity of taxa but the drivers underlying the global distribution of marine organisms are still under scrutiny and we still lack an accurate global predictive model. Using a spatial database for 6336 tropical reef fishes, we attempted to predict species richness according to geometric, biogeographical and environmental explanatory variables. In particular, we aimed to evaluate and disentangle the predictive performances of temperature, habitat area, connectivity, mid‐domain effect and biogeographical region on reef fish species richness. We used boosted regression trees, a flexible machine‐learning technique, to build our predictive model and structural equation modeling to test for potential ‘mediation effects’ among predictors. Our model proved to be accurate, explaining 80% of the total deviance in fish richness using a cross‐validated procedure. Coral reef area and biogeographical region were the primary predictors of reef fish species richness, followed by coast length, connectivity, mid‐domain effect and sea surface temperature, with interactions between the region and other predictors. Important indirect effects of water temperature on reef fish richness, mediated by coral reef area, were also identified. The relationship between environmental predictors and species richness varied markedly among biogeographical regions. Our analysis revealed that a few easily accessible variables can accurately predict reef fish species richness. They also highlight concerns regarding ongoing environmental declines, with region‐specific responses to variation in environmental conditions predicting a variable response to anthropogenic impacts.     

Human impacts on the species-area relationship in reef fish assemblages

The relationship between species richness and area is one of the oldest, most recognized patterns in ecology. Here we provide empirical evidence for strong impacts of fisheries exploitation on the slope of the species–area relationship (SAR). Using comparative field surveys of fish on protected and exploited reefs in three oceans and the Mediterranean Sea, we show that exploitation consistently depresses the slope of the SAR for both power-law and exponential models. The magnitude of change appears to be proportional to fishing intensity. Results are independent of taxonomic resolution and robust across coral and rocky reefs, sampling protocols and statistical methods. Changes in species richness, relative abundance and patch occupancy all appear to contribute to this pattern. We conclude that exploitation pressure impacts the fundamental scaling of biodiversity as well as the species richness and spatial distribution patterns of reef fish. We propose that species–area curves can be sensitive indicators of community-level changes in biodiversity, and may be useful in quantifying the human imprint on reef biodiversity, and potentially elsewhere.     

Regional and environmental effects on the species richness of mammal assemblages

Aim Variation in species richness has been related to (1) environmental conditions (water, energy and habitat characteristics) and (2) regional differences (contingent historical events and regional particularities that result in differences between regional faunas acting at broad extents). Whereas climatic factors have been widely studied, the effects of regional differences are less often quantified. This work aims to characterize global trends in the species richness of mammal assemblages with respect to both current and historical influences. Location All terrestrial biogeographical realms except Antarctica. Methods Species richness in checklists from 224 sites distributed worldwide were investigated by partitioning the variation between a general set of habitat/climate factors, biogeographical regions, and their overlaps. Additional analyses studied the specific overlaps of region, water and energy. Data were also divided according to area to determine if the strength of these effects varies according to the size of sites. Results Environmental effects explained 38% of richness variation across all sites, whereas environmentally independent regional effects explained 11% and the overlap between region and environment explained 13%. Results were similar when only larger sites (between 1000 km² and 10,000 km²) were considered. However, the importance of the overlap between region and all environmental variables was greater in smaller sites (between 100 km² and 1000 km²). In contrast, the specific importance of water and energy variables and their overlap with region was greater in larger sites. The strength of the independent effect of region remained almost invariant regardless of the size of the sites studied. Main conclusions The relationship between species richness and climate varies with scale and among regions. Although environmental variables are the strongest correlates of richness, the unique history and physiographic characteristics of a region produce differences between the richness of mammal assemblages and their response to environmental gradients. The importance of environmental variables varies with scale: climatic gradients are more important at coarse grain (larger sites), possibly as a result of their effects on species ranges, whereas habitat type is more important at the smaller sites, where the importance of ecological interactions increases. Therefore, regional differences and the scale at which richness is measured should be taken into account when evaluating species richness–energy hypotheses.     

Regional and global climate risks for reef corals: Incorporating species-specific vulnerability and exposure to climate hazards

Climate change is driving rapid and widespread erosion of the environmental conditions that formerly supported species persistence. Existing projections of climate change typically focus on forecasts of acute environmental anomalies and global extinction risks. The current projections also frequently consider all species within a broad taxonomic group together without differentiating species-specific patterns. Consequently, we still know little about the explicit dimensions of climate risk (i.e., species-specific vulnerability, exposure and hazard) that are vital for predicting future biodiversity responses (e.g., adaptation, migration) and developing management and conservation strategies. Here, we use reef corals as model organisms (n = 741 species) to project the extent of regional and global climate risks of marine organisms into the future. We characterise species-specific vulnerability based on the global geographic range and historical environmental conditions (1900–1994) of each coral species within their ranges, and quantify the projected exposure to climate hazard beyond the historical conditions as climate risk. We show that many coral species will experience a complete loss of pre-modern climate analogs at the regional scale and across their entire distributional ranges, and such exposure to hazardous conditions are predicted to pose substantial regional and global climate risks to reef corals. Although high-latitude regions may provide climate refugia for some tropical corals until the mid-21st century, they will not become a universal haven for all corals. Notably, high-latitude specialists and species with small geographic ranges remain particularly vulnerable as they tend to possess limited capacities to avoid climate risks (e.g., via adaptive and migratory responses). Predicted climate risks are amplified substantially under the SSP5-8.5 compared with the SSP1-2.6 scenario, highlighting the need for stringent emission controls. Our projections of both regional and global climate risks offer unique opportunities to facilitate climate action at spatial scales relevant to conservation and management.     

Local ecological impacts of regional biodiversity on reef fish assemblages

Aim We examined comparative data for cryptobenthic reef fishes to determine how variation in regional species richness relates to local species richness, abundance, and taxonomic and trophic composition, and to test whether systems with higher species richness exhibit finer habitat partitioning. Locations Lizard Island, Great Barrier Reef (GBR), Australia; Bahía de Loreto, Gulf of California (GoC), Mexico. Methods Cryptobenthic reef fish assemblages from four habitats (coral heads, rubble, and horizontal and vertical surfaces of boulders) were collected using clove oil. Differences in density, species richness and biomass were examined between regions and among habitats. Habitat associations were identified for each habitat/location based on multivariate ordination, and the statistical significance of patterns was tested using analysis of similarity (ANOSIM). In addition, the trophic group composition of the assemblages for both regions was examined. Results A total of 91 species in 20 families were recorded (GBR, 66 species; GoC, 25 species). Total and habitat species richness were higher on the GBR, whereas biomass was higher in the GoC. No difference in fish density between regions was found. Habitat division among assemblages was greater in the depauperate GoC. Only coral head associations proved to be distinctive on the GBR, whereas three sample groups were found in the GoC (coral heads, horizontal boulders and vertical boulders/rubble). Trophic composition in the two regions was markedly different, with omnivores dominating the GBR fauna and planktivores the GoC. Main conclusions A positive regional–local relationship in fish diversity was found between regions, but fish abundance in both regions remained similar. Contrary to expectations, habitat partitioning, at a community level, was greater in the depauperate GoC. Differences in trophic composition and patterns of habitat use appear to reflect the disparate history of the regions, whereas patterns of abundance may reflect the influence of fundamental relationships between size and abundance in communities. This study highlights the potential of reef faunas to conform to universal numerical trends while maintaining an ability to respond ecologically to local/evolutionary influences. The GoC fauna appears to be exceptionally vulnerable to natural and anthropogenic disturbance owing to the high numerical dominance of habitat‐specific species and to the limited potential for functional redundancy within the system.     

Global scale patterns of fish species richness in rivers

Explanations of spatial and temporal variation in species richness is a central theme in community ecology Until recently, most research has focused on small-scale phenomena, often emphasizing on local environmental factors and, thus, poorly reflecting large-scale processes that organize species richness In this paper, we analyze variations in species richness of indigeneous freshwater fish on a worldwide scale We show that factors related to species-area and species-energy theories statistically explain most of the variation in freshwater fish species richness across continents Historical events supposed to influence present distributions offish are of little assistence in explaining variations in fish species richness at the global scale Our model, which uses easily measured factors, should also be of practical value to aquatic conservation biology and natural resource management     

Regional species richness,hydrological characteristics and the local species richness of assemblages of North American stream fishes

1. Local assemblage structure, from a deterministic perspective, is presumably dictated by the regional species pool as well as regulated by local factors. We examined the relationships of the regional species pool and local hydrological characteristics to local species richness of North American freshwater fishes using data sets collected during the National Water Quality Assessment program conducted by the United States Geological Survey. 2. We predicted that local species richness is ultimately constrained by the composition of the regional species pool and further associated with local hydrological factors. Moreover, we predicted that variation in local species richness within major families can be explained by different combinations of hydrological characteristics that represent lineage‐specific responses to the environment. 3. Daily discharge and regional and local species richness data were assembled from 41 stream localities across the United States. Multiple stepwise regressions were conducted to predict local species richness, based on regional species richness, mean discharge and hydrological characteristics quantified by nine variables characterising flow variability. Species richness at each site was calculated for the entire assemblage as well as within the four most species‐rich families in the data set (Catostomidae, Centrarchidae, Cyprinidae and Percidae). 4. Local species richness was best predicted by a combination of regional species richness and discharge magnitude when all species were considered. Regional species richness was a significant explanatory variable of local species richness for three of four families (Catostomidae, Centrarchidae, Cyprinidae), but not for Percidae. Local richness in Centrarchidae and Cyprinidae was positively correlated with temporal flow variability as well as high and low flow duration, respectively, while richness in Catostomidae and Percidae tended to be associated with discharge volume. In addition, local species richness for three of the four major families was positively correlated with species richness of the other families in the assemblage, potentially suggesting the influence of local habitat quality and heterogeneity. 5. Results suggest the importance of the combined influences of the regional species pool and local hydrological characteristics on local richness in freshwater fishes, with variation in richness within each family predicted by different characteristics of flow regimes.     

Exploitation-related reef fish species richness depletion in the epicenter of marine biodiversity

The central Visayan region of the Philippines historically has the highest concentration of coral reef fishes than any other large marine area in the world. This well-supported biogeographic phenomenon is contradicted by recent transect observations on coral reefs that indicates that the Visayan region and the southern Philippine Sea region have the lowest species richness in the Philippines. The Visayan region has unusually low counts of species typically exploited in fisheries and the aquarium trade. This evidence, coupled with numerous reports of intense fishing and habitat degradation and subsequent species declines at local scales suggests that this exploitation is having a cumulative effect on the overall species richness of the Visayan region. Successes in Marine Protected Areas in this region in increasing species richness at local scales suggests that improved management of these protected areas coupled with much more intensive fisheries management will be key to reviving a healthy biodiversity in the Visayas.     

Predators reduce abundance and species richness of coral reef fish recruits via non-selective predation

Predators have important effects on coral reef fish populations, but their effects on community structure have only recently been investigated and are not yet well understood. Here, the effect of predation on the diversity and abundance of young coral reef fishes was experimentally examined in Moorea, French Polynesia. Effects of predators were quantified by monitoring recruitment of fishes onto standardized patch reefs in predator-exclosure cages or uncaged reefs. At the end of the 54-day experiment, recruits were 74% less abundant on reefs exposed to predators than on caged ones, and species richness was 42% lower on reefs exposed to predators. Effects of predators varied somewhat among families, however, rarefaction analysis indicated that predators foraged non-selectively among species. These results indicate that predation can alter diversity of reef fish communities by indiscriminately reducing the abundance of fishes soon after settlement, thereby reducing the number of species present on reefs.     

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.     

Assessing lake typologies and indicator fish species for Italian natural lakes using past fish richness and assemblages

In order to establish a fish-based typology of Italian lakes and identify possible reference and indicator fish species for each lake type, we analysed historical data on fish assemblages of all Italian natural lakes >0.5 km² from the period prior to the major decline in water quality in the 1950s. General linear regression models showed the ecoregion and lake altitude being the best predictors of fish species richness. The number of species was significantly higher in the Alpine than in the Mediterranean ecoregion. Among Alpine lakes, the number of fish species increased significantly with lake volume whilst decreased with altitude. In the Mediterranean lakes, none of the selected parameters was significant. Cluster analysis of fish assemblages (presence/absence) divided the lakes of the Alpine and Mediterranean ecoregions into four and two types, respectively. Pike (Esox lucius), rudd (Scardinius erythrophthalmus) and tench (Tinca tinca) were the main indicator species for the small and mostly shallow lakes in both the Alpine (Type 1) and Mediterranean (Type 6) ecoregions, minnow (Phoxinus phoxinus) for the alpine high altitude lakes (Type 2) and landlocked shad (Alosa fallax lacustris), European whitefish (Coregonus lavaretus) and burbot (Lota lota) for the large and very deep alpine lakes (Type 4). The European whitefish was the only indicator species for the deep Mediterranean lakes (Type 5). These species and associated fish assemblages may be useful indicators in future assessments of the ecological status of Italian lakes according to the European Directives (2000/60/EC and 2008/105/EC).     

Interactions between coral restoration and fish assemblages: implications for reef management

Corals create complex reef structures that provide both habitat and food for many fish species. Because of numerous natural and anthropogenic threats, many coral reefs are currently being degraded, endangering the fish assemblages they support. Coral reef restoration, an active ecological management tool, may help reverse some of the current trends in reef degradation through the transplantation of stony corals. Although restoration techniques have been extensively reviewed in relation to coral survival, our understanding of the effects of adding live coral cover and complexity on fishes is in its infancy with a lack of scientifically validated research. This study reviews the limited data on reef restoration and fish assemblages, and complements this with the more extensive understanding of complex interactions between natural reefs and fishes and how this might inform restoration efforts. It also discusses which key fish species or functional groups may promote, facilitate or inhibit restoration efforts and, in turn, how restoration efforts can be optimised to enhance coral fish assemblages. By highlighting critical knowledge gaps in relation to fishes and restoration interactions, the study aims to stimulate research into the role of reef fishes in restoration projects. A greater understanding of the functional roles of reef fishes would also help inform whether restoration projects can return fish assemblages to their natural compositions or whether alternative species compositions develop, and over what timeframe. Although alleviation of local and global reef stressors remains a priority, reef restoration is an important tool; an increased understanding of the interactions between replanted corals and the fishes they support is critical for ensuring its success for people and nature.     

Relationships between structural complexity,coral traits,and reef fish assemblages

With the ongoing loss of coral cover and the associated flattening of reef architecture, understanding the links between coral habitat and reef fishes is of critical importance. Here, we investigate whether considering coral traits and functional diversity provides new insights into the relationship between structural complexity and reef fish communities, and whether coral traits and community composition can predict structural complexity. Across 157 sites in Seychelles, Maldives, the Chagos Archipelago, and Australia’s Great Barrier Reef, we find that structural complexity and reef zone are the strongest and most consistent predictors of reef fish abundance, biomass, species richness, and trophic structure. However, coral traits, diversity, and life histories provided additional predictive power for models of reef fish assemblages, and were key drivers of structural complexity. Our findings highlight that reef complexity relies on living corals—with different traits and life histories—continuing to build carbonate skeletons, and that these nuanced relationships between coral assemblages and habitat complexity can affect the structure of reef fish assemblages. Seascape-level estimates of structural complexity are rapid and cost effective with important implications for the structure and function of fish assemblages, and should be incorporated into monitoring programs.     

Climate‐niche factor analysis: a spatial approach to quantifying species vulnerability to climate change

Climate change vulnerability assessments are an important tool for understanding the threat that climate change poses to species and populations, but do not generally yield insight into the spatial variation in vulnerability throughout a species’ habitat. We demonstrate how to adapt the method of ecological‐niche factor analysis (ENFA) to objectively quantify aspects of species sensitivity to climate change. We then expand ENFA to quantify aspects of exposure and vulnerability to climate change as well, using future projections of global climate models. This approach provides spatially‐explicit insight into geographic patterns of vulnerability, relies only on readily‐available spatial data, is suitable for a wide range of species and habitats, and invites comparison between different species. We apply our methods to a case study of two species of montane mammals, the American pika Ochotona princeps and the yellow‐bellied marmot Marmota flaviventris.     

Trait-based species richness: ecology and macroevolution

Understanding the origins of species richness patterns is a fundamental goal in ecology and evolutionary biology. Much research has focused on explaining two kinds of species richness patterns: (i) spatial species richness patterns (e.g. the latitudinal diversity gradient), and (ii) clade-based species richness patterns (e.g. the predominance of angiosperm species among plants). Here, I highlight a third kind of richness pattern: trait-based species richness (e.g. the number of species with each state of a character, such as diet or body size). Trait-based richness patterns are relevant to many topics in ecology and evolution, from ecosystem function to adaptive radiation to the paradox of sex. Although many studies have described particular trait-based richness patterns, the origins of these patterns remain far less understood, and trait-based richness has not been emphasised as a general category of richness patterns. Here, I describe a conceptual framework for how trait-based richness patterns arise compared to other richness patterns. A systematic review suggests that trait-based richness patterns are most often explained by when each state originates within a group (i.e. older states generally have higher richness), and not by differences in transition rates among states or faster diversification of species with certain states. This latter result contrasts with the widespread emphasis on diversification rates in species-richness research. I show that many recent studies of spatial richness patterns are actually studies of trait-based richness patterns, potentially confounding the causes of these patterns. Finally, I describe a plethora of unanswered questions related to trait-based richness patterns.     

Reconstructing past species assemblages reveals the changing patterns and drivers of extinction through time

Predicting future species extinctions from patterns of past extinctions or current threat status relies on the assumption that the taxonomic and biological selectivity of extinction is consistent through time. If the driving forces of extinction change through time, this assumption may be unrealistic. Testing the consistency of extinction patterns between the past and the present has been difficult, because the phylogenetically explicit methods used to model present-day extinction risk typically cannot be applied to the data from the fossil record. However, the detailed historical and fossil records of the New Zealand avifauna provide a unique opportunity to reconstruct a complete, large faunal assemblage for different periods in the past. Using the first complete phylogeny of all known native New Zealand bird species, both extant and extinct, we show how the taxonomic and phylogenetic selectivity of extinction, and biological correlates of extinction, change from the pre-human period through Polynesian and European occupation, to the present. These changes can be explained both by changes in primary threatening processes, and by the operation of extinction filter effects. The variable patterns of extinction through time may confound attempts to identify risk factors that apply across time periods, and to infer future species declines from past extinction patterns and current threat status.     

6个耐旱树种木质部结构与栓塞脆弱性的关系

木质部栓塞脆弱性对干旱响应的研究已成为全球气候变化背景下的热点和重要内容。该文以6个耐旱树种刺槐(Robinia pseudoacacia)、沙棘(Hippophae rhamnoides)、榆树(Ulmus pumila)、元宝枫(Acer truncatum)、旱柳(Salix matsudana)、榛(Corylus heterophylla)为研究对象, 采用Cochard Cavitron离心机技术建立木质部栓塞脆弱曲线, 计算木质部栓塞脆弱性, 利用染色法、硅胶注射法等测定木质部导管直径、导管内径跨度、导管连接度、导管密度、导管长度和木质部密度, 探究木质部结构与栓塞脆弱性的关系, 区分6个耐旱树种木质部结构在抗栓塞性上的差异, 以期建立6个耐旱树种在木质部结构方面的抗栓塞性指标。结果表明: 6个耐旱树种木质部栓塞脆弱性大小为刺槐>榆树>沙棘>旱柳>元宝枫>榛, 其中, 刺槐、沙棘和榆树的栓塞脆弱曲线为“r”形, 而元宝枫、旱柳和榛的栓塞脆弱曲线为“s”形, 脆弱曲线为“r”形的树种与脆弱曲线为“s”形的树种栓塞脆弱性差异极显著(p < 0.01)。线性分析表明: 木质部结构影响各树种的栓塞脆弱性, 其中, 木质部密度影响最大(t = 0.702), 导管直径次之(t = 0.532), 导管长度影响最小(t = 0.010)。     

Within and between day variability in coral reef fish assemblages: Implications for fish community surveys

Coral reef ecosystems are under increasing anthropogenic pressures making it ever more important to monitor changes in fish communities to implement appropriate management. In contrast to long-term spatial and temporal variation which has been extensively documented, little work has been carried out to identify variability in fish assemblages on short time scales, with few studies testing patterns of fish assemblages between and within days. Here we investigated the diurnal changes in species richness, relative abundance and assemblage composition in a shallow coral reef fish community in Egypt. To do so, a section of coral reef was filmed during the morning (0600 h), midday (1000 and 1400 h) and afternoon (1800 h) over eleven days. Dusk (0600 h) and dawn samples (1800 h) showed higher species richness compared to late morning (1000 h) and mid-day samples (1400 h) and borderline significantly higher numbers of total individuals, likely associated with feeding activity and predator avoidance. Assemblage composition varied across days and time-of-day, showing greater variability during dusk and dawn associated with a transition between day-time and night-time assemblages. Our results have implications for designing coral reef fish surveys, emphasising that short-term changes in fish communities should be considered when designing experiments to monitor fish assemblages over time. Where possible, we suggest increasing replication within sites and time scales or randomising data within a specific time window at all sites, looking to exclude dusk and dawn.