Trait-based metrics as bioindicators: Responses of stream fish assemblages to a gradient of environmental degradation

The development of biomonitoring tools is increasingly appealing in light of the increasing degradation of aquatic ecosystems. In this context, we investigated the responses of stream fish communities to the gradient of environmental degradation in different basins using a variety of indices based on functional diversity, functional composition, and taxonomic diversity. We used datasets from three Brazilian ecoregions. In order to describe the gradient of environmental degradation we scored streams based on local and landscape variables. The functional structure of the assemblages was described in relation to seven functional traits related to habitat use, swimming capacity, and maneuverability. We described assemblages in each ecoregion separately in relation to 20 potential indicators, grouped in four families: indices of functional diversity based on presence/absence (FD_p/a); indices of functional diversity weighted by abundance (FD_abund); indices of functional composition (mT); and, indices based on taxonomic diversity (TDiv). The relationship between the indicators and the gradient of environmental degradation were evaluated using linear regression. We found a significant interaction effect (ANOVA, p = 0.006) between group of indices and ecoregions for the performance of indicators, suggesting a context-dependent response. The indices, on an individual basis, had variable performance and consistency among ecoregions. Four mT indicators demonstrated the highest average performance and consistency. Taxonomic diversity indicators consistently had the lowest average performance, while FD_p/a and FD_abund indicators had low average performance and variable inter-regional consistency. The differential inter-regional performance of indicators was due to the differences in the lengths of the gradients of environmental degradation. Our results indicate that functional traits have greater predictive power compared to taxonomic indicators for fish responses to a gradient of environmental degradation. Although indicators of functional composition are the most promising, we emphasize that caution is needed when generalizing functional diversity indicators across ecoregions because most of them are context-dependent.     

Patterns of functional and taxonomic organization of stream fishes: inferences based on α, β, and γ diversities

The primary objective of this study was to determine whether total biodiversity (γ) is partitioned into within‐community (α) and among‐community (β) components differently for taxonomic and functional organization. I hypothesized that α diversity will contribute more to the functional organization of γ diversity and that β diversity will contribute more to the taxonomic organization of γ diversity. A secondary objective was to determine whether the relationship between taxonomic and functional diversity is scale dependent. Species abundance data was obtained from fisheries surveys conducted by the Texas Parks and Wildlife Dept that focused on least disturbed streams from 11 different ecoregions of Texas, including 62 localities from 18 drainages. Functional and taxonomic organization of assemblages was quantified with two different measures of biodiversity, including richness and the numbers equivalent of Shannon diversity. Scale‐dependent effects on these indices were assessed by multiplicatively partitioning γ into α and β components. The contribution of α and β components to γ diversity differed between functional and taxonomic organization and among different measures of biodiversity. Among‐community components were more influential in structuring the taxonomic organization of stream‐fish assemblages, whereas within‐community components were more important in structuring the functional organization of assemblages. The relationship between taxonomic and functional diversity differed between α and β components and between spatial scales. Indeed, ecological patterns not only change with spatial scale, but how they change is dependent on which aspect of biodiversity is considered.     

Local and regional determinants of stream fish assemblage structure: inferences based on taxonomic vs. functional groups

Aim  To examine the roles of local and regional environmental variables and biotic interactions in determining the structure of local stream fish assemblages, and to compare results derived from analyses based on taxonomic and functional groups.
Location  Texas, USA.
Methods  Species abundance data were compiled for 157 stream fish assemblages in several river basins across Texas. Species were condensed into functional groups based on trophic and life-history characteristics. Local and regional environmental variables were either measured at each location or determined from scale maps and public-access data bases. The original taxonomic and functional group data sets were analysed using similarity indices, null models of co-occurrence, and direct and indirect ordination techniques. Results derived from taxonomic and functional group data sets are compared.
Results  Inferences regarding the relative roles of local and larger-scale factors in determining stream fish assemblage structure differ dramatically between analyses of taxonomic and functional groups. Taxonomic analyses suggest a prominent role of regional-scale environmental factors, and local assemblages sorted according to a biogeographic pattern. Functional group analyses suggest almost equal roles of factors representative of local and larger scales, and assemblages were distinguished by a habitat template irrespective of geographic region.
Main conclusions  The structure of local stream fish assemblages is determined ultimately by factors representing multiple scales, with the relative importance of each depending on the biological unit employed (species or functional groups). We suggest that analyses using functional groups can more directly infer ecological responses to environmental variation, and therefore may provide a more fruitful avenue for developing and testing ecological theory of community organization across biogeographic scales.     

Fishes of Indiana streams: current and historic assemblage structure

We examined Indiana fish assemblages using taxonomy and ecological categories to assess temporal shifts in community structure and recent environmental relationships. Historic (1945) and recent (1996–2007) presence/absence data were compiled by subbasin and analyzed with Nonmetric Multidimensional Scaling (NMS) ordination and by species richness. Canonical Correspondence Analysis (CCA) was used to test taxonomic identity and ecological category abundance data for explanation with recent (1996–2007) environmental variables. We found a decrease in assemblage heterogeneity for recent assemblages and an increase in the number of tolerant species per subbasin. Recent Indiana streams are dominated by tolerant fishes with generalist life history strategies and low functional variation. The use of ecological categories resulted in weaker relationships with environmental variables than analyses with taxonomic identities. Analyses using taxonomy resulted in strong assemblage explanation from stream size and flow variation, while analyses using ecological categories resulted in strong assemblage explanation from habitat variation in silt substrates and flow. Analyses of recent assemblage structure using ecological categories resulted in decreased assemblage variation among subbasins than in analyses using taxonomic identities. We found that fish assemblages of Indiana streams are structured primarily by habitat complexity and have been altered during the past 50 years through multiple disturbances including fragmentation, siltation, and species introductions.     

Seasonal fluctuations in taxonomic and functional diversity in assemblages of catfishes in the Venezuelan Arauca River Floodplain

The Arauca River harbors a considerable fish biodiversity, yet it remains poorly sampled and the knowledge about its fish communities is incomplete. We studied the taxonomic and functional composition and diversity of catfish assemblages in one lotic and one lentic habitat of a segment of this floodplain during one hydrological cycle, from August 2014 to August 2015, comprising six samplings using gillnets, and identified the relationships between catfish traits and the environment based on five limnological variables. The functional structure was studied through 11 morphological measures and the species diets. We identified 31 catfish species and nine trophic groups. There were significant differences in species composition and diversity between habitats and months, but the functional structure of the catfishes was only determined by the temporal variability. The functional structure of the catfish assemblages was ordered by morphological traits related to locomotion, habitat use, and trophic ecology and these traits were correlated with the limnological variables, supporting the existence of environmental filters. Integrating the taxonomic and functional perspectives can provide a more complete picture of the ecological patterns of freshwater fish assemblages in the Neotropics and can help us predict how these assemblages might change in response to anthropogenic alterations of the environment.     

海州湾鱼类群落功能多样性的时空变化

功能多样性是联系物种、生态环境和生态系统功能的基础.从功能多样性的角度研究群落结构,有助于更好地理解生物多样性与生态系统功能之间的关系.本研究根据2011—2017年(除2012年外)春、秋季海州湾渔业资源底拖网调查数据,选取反映鱼类摄食、运动、生态适应性、繁殖行为和种群动态特征的13个功能性状,利用功能丰富度指数、功能均匀度指数、功能离散度指数和群落特征加权平均数指数(CWM),研究了海州湾鱼类群落功能多样性的季节、年际和空间的变化.结果表明: 海州湾鱼类群落功能多样性指数具有显著的季节差异,其中秋季功能丰富度指数显著高于春季,春季功能离散度指数显著高于秋季,鱼类洄游是造成功能多样性指数季节变化的主要原因;CWM分析表明,春季鱼类群落优势种为营养级较高、运动能力较强、生长系数、恢复力和脆弱性较高的冷/暖温性鱼类,秋季则相反,春、秋季鱼类群落均以产浮性卵鱼类为主;鱼类群落功能多样性指数呈现一定的年际变化,其中春、秋季各功能多样性指数在不同年际均呈波动趋势,表明海州湾鱼类群落稳定性较低;鱼类群落功能多样性指数呈现显著的空间差异,其中20 m以深远岸水域功能离散度指数较高.海州湾鱼类群落功能多样性具有明显的时空变化特征,鱼类生态位与其对资源的利用均随季节、年际和空间而变化.     

Low‐head dams induce biotic homogenization/differentiation of fish assemblages in subtropical streams

Extensive distribution of widespread species and the loss of native species driven by anthropogenic disturbances modify community similarity, resulting in a decrease or increase in community distinctiveness. Data from four basins in the Wannan Mountains, China, were used to evaluate the effects of low‐head dams on patterns of fish faunal homogenization and differentiation based on abundance data. We aimed to examine the spatial changes in taxonomic and functional similarities of fish assemblages driven by low‐head dams and to examine whether the changes in the similarity of fish assemblages differed between taxonomic and functional components. We found that low‐head dams significantly decreased the mean taxonomic similarity but increased the mean functional similarity of fish assemblages in impoundments using abundance‐based approaches, suggesting that taxonomic differentiation accompanied functional homogenization in stream fish assemblages. These results show the importance of population abundance in structuring fish faunal homogenization and differentiation at small scales, especially when the major differences among assemblages are in species abundance ranks rather than species identities. Additionally, we also found only a weak positive correlation between changes in mean taxonomic and functional similarities, and partial pairs exhibited considerable variation in patterns of fish faunal homogenization and differentiation for taxonomic and functional components. In conclusion, this study highlighted that the observed taxonomic differentiation of current fish assemblages (short‐term phenomenon) is probably an early warning sign of further homogenization in regions where native species are completely predominated and that changes in taxonomic similarity cannot be used to predict changes in functional similarity.     

青弋江鱼类分类群和功能群的α和β多样性纵向梯度格局

确定溪流鱼类多样性的时空分布格局可为鱼类多样性保护与管理提供科学基础。尽管溪流鱼类分类群多样性的纵向梯度格局已有大量报道, 但以鱼类生物学特征为基础的功能多样性研究较少。本文基于2009-2010年4个季度对青弋江1-5级溪流共15个样点的调查数据, 利用形态特征数据和食性构建了鱼类复合功能群, 研究了不同级别溪流间鱼类分类群和功能群组成及多样性的异同, 着重探讨了鱼类分类群和功能群的α和β多样性沿溪流纵向梯度的变化规律。采集到的56种鱼类可分为4个营养功能群和5个运动功能群, 共计14个“营养-运动”复合功能群。双因素交互相似性分析结果显示, 鱼类分类群和功能群组成都随河流级别显著变化, 但季节动态不显著; 双因素方差分析后发现, 鱼类分类群和功能群α、β多样性都随河流级别显著变化, 但受季节影响不显著。经回归分析, 分类群和功能群α多样性与河流级别大小呈显著的线性正相关, 但最大分类群α多样性出现于4级河流, 最大功能群α多样性在4级和5级河流间一致; 分类群和功能群β多样性与河流级别大小呈显著的二项式关系, 呈U型分布。分类群β多样性的空间变化主要取决于物种周转, 而功能群β多样性主要由嵌套所驱动。本研究表明, 沿着“上游-下游”的纵向梯度, 河流鱼类的α和β多样性的空间变化规律不同, 分类群和功能群α多样性的空间格局基本一致, 但分类群(主要是物种周转)和功能群β多样性(主要是功能嵌套)的空间变化过程的驱动机制不同。     

Spatial mismatch in morphological,ecological and phylogenetic diversity,in historical and contemporary European freshwater fish faunas

Biodiversity encompasses multiple facets, among which taxonomic, functional and phylogenetic aspects are the most often considered. Understanding how those diversity facets are distributed and what are their determinants has become a central concern in the current context of biodiversity crisis, but such multi‐faceted measures over large geographical areas are still pending. Here, we measured the congruence between the biogeographical patterns of freshwater fish morphological, ecological and phylogenetic diversity across Europe and identified the natural and anthropogenic drivers shaping those patterns. Based on freshwater fish occurrence records in 290 European river catchments, we computed richness and evenness for morphological, ecological and phylogenetic diversity using standardized effect sizes for each diversity index. We then used linear models including climatic, geo‐morphological, biotic and human‐related factors to determine the key drivers shaping freshwater fish biodiversity patterns across Europe. We found a weak spatial congruence between facets of diversity. Patterns of diversity were mainly driven by elevation range, climatic seasonality and species richness while other factors played a minor role. Finally, we found that non‐native species introductions significantly affected diversity patterns and influenced the effects of some environmental drivers. Morphological, ecological and phylogenetic diversity constitute complementary facets of fish diversity rather than surrogates, testifying that they deserve to be considered altogether to properly assess biodiversity. Although the same environmental and anthropogenic factors overall explained those diversity facets, their relative influence varied. In the current context of global change, non‐native species introductions may also lead to important reshuffling of assemblages resulting in profound changes of diversity patterns.     

A multi-faceted framework of diversity for prioritizing the conservation of fish assemblages

Floodplain waterbodies and their biodiversity are increasingly threatened by human activities. Given the limited resources available to protect them, methods to identify the most valuable areas for biodiversity conservation are urgently needed. In this study, we used freshwater fish assemblages in floodplain waterbodies to propose an innovative method for selecting priority areas based on four aspects of their diversity: taxonomic (i.e. according to species classification), functional (i.e. relationship between species and ecosystem processes), natural heritage (i.e. species threat level), and socio-economic (i.e. species interest to anglers and fishermen) diversity. To quantitatively evaluate those aspects, we selected nine indices derived either from metrics computed at the species level and then combined for each assemblage (species rarity, origin, biodiversity conservation concern, functional uniqueness, functional originality, fishing interest), or from metrics directly computed at the assemblage level (species richness, assemblage rarity, diversity of biological traits). Each of these indices belongs to one of the four aspects of diversity. A synthetic index defined as the sum of the standardized aspects of diversity was used to assess the multi-faceted diversity of fish assemblages. We also investigated whether the two main environmental gradients at the catchment (distance from the sea) and at the floodplain (lateral connectivity of the waterbodies) scales influenced the diversity of fish assemblages, and consequently their potential conservation value. Finally, we propose that the floodplain waterbodies that should be conserved as a priority are those located in the downstream part of the catchment and which have a substantial lateral connectivity with the main channel.     

Numerical abundance and biomass reveal different temporal trends of functional diversity change in tropical fish assemblages

Understanding how the biodiversity of freshwater fish assemblages changes over time is an important challenge. Until recently most emphasis has been on taxonomic diversity, but it is now clear that measures of functional diversity (FD) can shed new light on the mechanisms that underpin this temporal change. Fish biologists use different currencies, such as numerical abundance and biomass, to measure the abundance of fish species. Nonetheless, because they are not necessarily equivalent, these alternative currencies have the potential to reveal different insights into trends of FD in natural assemblages. In this study, the authors asked how conclusions about temporal trends in FD are influenced by the way in which the abundance of species has been quantified. To do this, the authors computed two informative metrics, for each currency, for 16 freshwater fish assemblages in Trinidad's Northern Range that had been surveyed repeatedly over 5 years. The authors found that numerical abundance and biomass uncover different directional trends in these assemblages for each facet of FD, and as such inform hypotheses about the ways in which these systems are being restructured. On the basis of these results, the authors concluded that a combined approach, in which both currencies are used, contributes to our understanding of the ecological processes that are involved in biodiversity change in freshwater fish assemblages.     

DER: An algorithm for comparing species diversity between assemblages

There is no single diversity index that adequately summarises species diversity, since this is a multidimensional concept and hence should be quantified using a compound statistical measure. Here, we present the DER algorithm, available as an R package on CRAN and as an RWizard application on http://www.ipez.es/RWizard. This algorithm provides tools for differentiating assemblages on the basis of five compounds of diversity: rarity, heterogeneity, evenness, taxonomic/phylogenetic diversity and functional diversity. For all the samples, the algorithm calculates a total of 39 of the indices most often used. All indices of all samples are transformed to a scale range of between 0 and 1, and the algorithm calculates the polar coordinates of all samples with all possible combinations for all five groups of indices. Thus, for each combination, an index of rarity, heterogeneity (species richness is included in this group), evenness, taxonomic/phylogenetic diversity and functional diversity is used for each group to calculate the polar coordinates of all samples. When the polar coordinates of the samples are obtained for each combination, the convex hull and the mean Euclidean distance between samples are calculated. The algorithm selects the combination of indices with the highest value of the mean between convex hull and mean Euclidean distance between samples; priority is therefore given to maximising dispersion between the samples. The polar coordinates of the selected combination are depicted using a diagram from which it is possible to determine the differences in terms of rarity, heterogeneity, evenness, taxonomic/phylogenetic diversity and functional diversity between assemblages.     

What governs the functional diversity patterns of fishes in the headwater streams of the humid forest enclaves: environmental conditions,taxonomic diversity or biotic interactions?

The relationship between functional and taxonomic diversity is a major issue in ecology. Biodiversity in aquatic environments is strongly influenced by environmental gradients that act as dispersion and niche barriers. Environmental conditions act as filters to select functional traits, but biotic interactions also play a role in assemblage structure. In headwater streams, the relationship between functional and taxonomic diversity remains unclear. In this study we investigated how environmental conditions, taxonomic diversity and biotic interactions influence the spatial distribution of traits and functional diversity in stream fish species. Standardized sampling of fish species was carried out in 50 m sections of 16 streams located in rainforest enclaves in a semiarid region of Brazil (Caatinga biome). The functional diversity indices displayed different responses to the predictor variables used. Functional richness was mainly influenced by environmental conditions, while functional evenness was mostly determined by taxonomic diversity. On the other hand, functional dispersion was explained by a combination of environmental conditions and taxonomic diversity. The spatial distribution of fish species with the same functional traits was random, indicating that biotic interactions are not a strong predictor in these ecosystems. Channel width, pH and substrate were the most important variables in the spatial distribution of the functional traits of the fish species. Our results suggest that the functional structure of fish assemblages in headwater streams depends mainly on environmental conditions and taxonomic diversity.     

Utilization of species checklist data in revealing the spatial distribution of fish diversity

Marine fish species checklists from six Chinese coastal waters were combined for the analysis of taxonomic diversity. The Genus-Family index (G-F index) ranged between 0.39 and 0.84, which generally indicated a decreasing trend with increasing latitude, with the exception of the southernmost area. Average taxonomic distinctness showed a slight increasing trend from northern to central study areas, but whether the taxonomic distinctness indices represent a latitudinal gradient of biodiversity requires further study. The multivariate analysis revealed a distinct latitudinal variation in fish assemblages. These results indicate that species checklist data are helpful in understanding the diversity distribution of fish species in the coastal zone. The potential of a species inventory should be exploited to fully understand biodiversity.     

淡水鱼类功能多样性及其研究方法

目前,群落功能多样性备受生态学界关注,被认为是能解决生态问题的一种重要途径。我国对于群落功能多样性主要集中在植物群落和微生物群落,而在鱼类群落方面的研究几乎是空白。我国鱼类资源正面临着严重威胁,包括水坝建设导致的鱼类通道受阻、水库形成造成鱼类产卵场功能消失、过度捕捞、水质恶化和富营养化加重、外来种入侵等因素,导致渔业资源急剧衰退,水生生态系统功能下降。以淡水鱼类群落为例,对鱼类功能多样性的数据获取及处理分析与评价、测定指标及计算方法与研究难点等进行综述,以期为鱼类资源保护提供新的理论依据和切入点。     

Biogeographic patterns in major marine realms: function not taxonomy unites fish assemblages in reef,seagrass and mangrove systems

We examine the effects of different biogeographic histories on assemblage composition in three major marine habitats in two biogeographically distinct marine realms. Specifically, we quantify the taxonomic and functional composition of fish assemblages that characterise coral reef, seagrass and mangrove habitats, to explore the potential effects of biogeographic history and environment on assemblage composition. The three habitats were surveyed in the Caribbean and on the Great Barrier Reef using a standardised underwater visual census method to record fish size and abundance data. The taxonomic composition of assemblages followed biogeographic expectations, with realm‐specific family‐level compositions. In marked contrast, the functional composition of assemblages separated habitats regardless of their biogeographic locations. In essence, taxonomy characterises biogeographic realms while functional groups characterise habitats. The Caribbean and Indo‐West Pacific have been separated for approximately 15 million years. The two realms have different taxonomic structures which reflect this extended separation, however, the three dominant shallow‐water marine habitats all retain distinct functional characteristics: seagrass fishes are functionally similar regardless of their taxonomic composition or biogeographic location. Likewise, for coral reefs and mangroves. The results emphasise the advantages and limitations of taxonomic vs. functional metrics in evaluating patterns. Taxonomy primarily reflects biogeographic and evolutionary history while functional characteristics may better reflect ecological constraints.     

Measuring ecosystem degradation through half a century of fish species introductions and extirpations in a large isolated lake

The introduction of exotic species and the extirpation of native species that occurred during the past two centuries have strongly modified the structure of most plant and animal assemblages across the globe. Such a biotic change is particularly marked in isolated environments such as islands or isolated lakes. Most studies reported drastic changes between before and after human disturbances, but the dynamics of change in assemblage structure through the invasion and extirpation processes are rarely reported. Here we measured the aquatic ecosystem degradation through exotic species introduction and native species extirpation experienced by Lake Erhai (China) during the last 50 years using structural, functional and taxonomic distinctness biodiversity indices. Structural diversity (species richness) did not varied monotonically along the temporal gradient, due to an opposite trend between exotic species increase and a concomitant decline of native species richness. Functional diversity displayed unclear ascending trends driven by the introduction of exotic species having distinct functional traits than natives. Taxonomic distinctness indices exhibited an increase of the average taxonomic distinctness (Δ⁺), but a decrease of the variation in taxonomic distinctness (Λ⁺) through time. Structural, functional and distinctness indices providing complementary information on ecosystem degradation, we here proposed a new multifaceted degradation index integrating these three facets of biodiversity. Such an index provided an accurate representation of the faunistic changes experienced by Lake Erhai and might constitute a comprehensive way to measure ecosystem degradation through exotic fish species introductions and native fish species extirpations.     

Ecomorphological analysis as a complementary tool to detect changes in fish communities following major perturbations in two South African estuarine systems

Ecomorphological changes as a result of natural perturbations in estuarine fish communities were investigated in two South African estuaries (Swartvlei and East Kleinemonde), both before and after the loss of aquatic macrophyte beds in these systems. The fish communities were analysed using an ecomorphological diversity index (EMI) and the results compared to a traditional index, the Shannon-Wiener diversity index. The EMI revealed that the major changes in fish community composition recorded in both estuaries were associated with quantitative variations at the species level. Both estuaries essentially lost their macrophyte beds and ended up with the same type of bottom habitat (bare sediment). In both cases the fish morphological variability decreased immediately after aquatic macrophyte loss and then increased to end above the initial value. The ecomorphological analysis appeared to be sensitive to major ecological disturbances that occurred during the study period and this was confirmed by the morphospace configuration. The results indicate that the ecomorphology of the fish community responds to habitat changes and that this change corresponds to alterations in the representation of the different feeding types. These findings therefore contribute to the measurement of morphological changes in estuarine fish assemblages as a result of habitat changes within the ecosystem and we propose that ecomorphological analyses add another dimension to the information provided by existing diversity indices in studying changing fish communities.     

Finding the Minimum Sample Richness (MSR) for multivariate analyses: implications for palaeoecology

Many techniques have been developed to estimate species richness and beta diversity. Those techniques, dependent on sampling, require abundance or presence/absence data. Palaeontological data is by nature incomplete, and presence/absence data is often the only type of data that can be used to provide an estimate of ancient biodiversity. We used a simulation approach to investigate the behaviour of commonly used similarity indices, and the reliability of classifications derived from these indices, when working with incomplete data. We drew samples, of varying number and richness, from artificial species lists, which represented original life assemblages, and calculated error rates for classifications of the parent lists and samples. Using these results, we estimated the Minimum Sample Richness (MSR) needed to achieve 95% classification accuracy. Results were compared for classifications derived from several commonly used similarity indexes (Dice, Jaccard, Simpson and Raup–Crick). MSR was similar for the Dice, Jaccard and Simpson indices. MSR for the Raup–Crick index was often much lower, suggesting that it is preferable for classifying patchy data, however the performance of this index was less stable than the other three in the simulations, which required an even lower MSR. MSR can be found for all presence/absence data from the contour graphs and equations as long as the absolute species richness and the beta diversity can be estimated.