Lessons from the fish: a multi‐species analysis reveals common processes underlying similar species‐genetic diversity correlations

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Spatial genetic differentiation correlates with species assemblage turnover across tropical reef fish lineages

Aim

Evaluating the similarity of diversity patterns across micro- to macroevolutionary scales in natural communities, such as species–genetic diversity correlations (SGDCs), may inform on processes shaping community assembly. However, whether SGDCs not only hold across communities but also across lineages has never been explored so far. Here we investigated SGDCs across co-distributed taxa for different spatial components (α, β, γ), and formally tested the influence of dispersal traits on β-SGDCs.

Location

Western Indian Ocean.

Time period

2016–2017.

Major taxa studied

Tropical reef fish species with contrasting dispersal traits.

Methods

Using double-digest restriction-site associated DNA sequencing (ddRADseq) Single Nucleotide Polymorphism data for 20 tropical reef fishes and distribution data of 2,446 species belonging to 12 families, we analysed the correlations between within-species genetic diversity and within-family species diversity (i.e., lineage diversity) for the three spatial components (α, β, γ-SGDCs). We then related the strength of β-SGDCs per species to proxies of larval dispersal abilities.

Results

We detected positive and significant lineage-based SGDC only for the β component, that is, the families showing the greatest level of species turnover among sites contain the species with the greatest levels of genetic differentiation. We showed that the Monsoon Drift mainly explained the β-diversity patterns at both intraspecific and interspecific levels. Higher β-SGDCs were found for species with short pelagic larval duration and weak larval swimming capacity.

Main conclusions

Our study reveals a strong correlation between genetic and species β-diversity, a result explained by the presence of a ‘soft’ barrier and mediated by larval dispersal processes. This suggests that vicariance and dispersal limitation are major processes shaping β-diversity patterns from microevolutionary to macroevolutionary scales in tropical reef fishes.     

Lessons from the macroinvertebrates: species‐genetic diversity correlations highlight important dissimilar relationships

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Functional diversity and trait–environment relationships of stream fish assemblages in a large tropical catchment

1. The species composition of stream fish assemblages changes across the longitudinal fluvial gradient of large river basins. These changes may reflect both zonation in species distributions and environmental filtering of fish traits as stream environments change from the uplands to the lowlands of large catchments. Previous research has shown that taxonomic diversity generally increases in larger, lowland streams, and the River Continuum Concept, the River Habitat Template and other frameworks have provided expectations for what functional groups of fishes should predominate in certain stream types. However, studies addressing the functional trait composition of fish assemblages across large regions are lacking, particularly in tropical river basins. 2. We examined functional trait–environment relationships and functional diversity of stream fish assemblages in the Río Grijalva Basin in southern Mexico. Traits linked to feeding, locomotion and life history strategy were measured in fishes from streams throughout the catchment, from highland headwaters to broad, lowland streams. Relationships between functional traits and environmental variables at local and landscape scales were examined using multivariate ordination, and the convex hull volume of trait space occupied by fish assemblages was calculated as a measure of functional diversity. 3. Although there were a few exceptions, functional diversity of assemblages increased with species richness along the gradient from uplands to lowlands within the Grijalva Basin. Traits related to swimming, habitat preference and food resource use were associated with both local (e.g. substratum type, pool availability) and landscape‐scale (e.g. forest cover) environmental variables. 4. Along with taxonomic structure and diversity, the functional composition of fish assemblages changed across the longitudinal fluvial gradient of the basin. Trait–environment relationships documented in this study partially confirmed theoretical expectations and revealed patterns that may help in developing a better understanding of general functional responses of fish assemblages to environmental change.     

Revealing the pathways by which agricultural land‐use affects stream fish communities in South Brazilian grasslands

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Genetic and community similarities are correlated in endemic-rich springs of the northern Chihuahuan Desert

Aim Species diversity and genetic diversity within a taxon are intrinsic parts of global biodiversity. These two levels of biodiversity can show strong correlation due to a variety of reasons (i.e. parallel processes affecting both communities and populations, genotypes of a numerically or functionally dominant species affecting community composition, a species assemblage selecting for a particular genotype by affecting its selection regime). We examined correlations between species and genetic biodiversity in four isolated endemic‐rich spring systems in a hot desert and their potential link to environmental variables and physical isolation. Location Chihuahuan Desert spring systems in the Pecos River basin of New Mexico and Texas, USA. Methods We compared species richness of fish and benthic macroinvertebrate communities to within‐population allelic richness of amphipods (monophyletic Gammarus spp.) and Pecos gambusia (Gambusia nobilis) using Mantel tests. We also compared pairwise community similarities with pairwise genetic identities of populations among the same groups. We tested correlations among diversity, similarity and environmental variables after controlling for the effects of spatial distances using partial Mantel tests. We partitioned genetic and species diversity into three spatial scales (i.e. individual springs, individual spring systems, the entire region) using AMOVA and partition . Results We found strong correlations between invertebrate species richness and mosquitofish allelic richness. We found even stronger correlations of amphipod and gambusia genetic identities with fish and invertebrate community similarities; these were best explained by geographic distance rather than abiotic environmental factors. Most of the taxa and communities exhibited the largest proportion of diversity at the regional level. Main conclusions Our results suggest that drift and migration are the mechanisms that best explain our observations, and although α‐diversity among genes and species may not be strongly correlated, the pattern of species and allelic complementarity among these groups seems to be concordant at the regional level.     

Regional and longitudinal patterns of fish community structure in the Seine River basin, France

The fish communities of 371 sites from 4 natural regions of the Seine River basin were studied. The sites were located from small to medium size rivers (catchment area : 5 to 3895 km²). We examined the differences between local communities according to river size (estimated by catchment area) and region. In the Seine River basin, fish communities follow a general organisation rule: total species richness increases with river size and importance of limnophilic species versus rheophilic ones increases from upstream to downstream. However, fish communities show differences of total species richness, species richness of reproductive groups and species composition between the four natural regions of the basin. Particularly, river size and regional organisation of environmental factors interact on species composition of communities and several regional patterns of longitudinal changes of fish communities are identified. The origin and range of regional differences of fish communities are discussed according to historical and environmental factors.     

Species diversity and genetic diversity: parallel processes and correlated patterns

Species diversity and genetic diversity may be correlated as a result of processes acting in parallel at the two levels. However, no theories predict the conditions under which different relationships between species diversity and genetic diversity might arise and therefore when one level of diversity may be predicted using the other. I used simulation models to investigate the parallel influence of locality area, immigration rate, and environmental heterogeneity on species diversity and genetic diversity. The most common pattern was moderate to strong positive species-genetic diversity correlations (SGDCs). Such correlations may be driven by any one of the three locality characteristics examined, but important exceptions and patterns emerged. Genetic diversity and species diversity were more weakly correlated when genetic diversity was measured for rare versus common species. Environmental heterogeneity not only imposes spatially varying selection on populations and communities but also causes changes in species' population sizes and therefore genetic diversity; these interacting processes can create positive, negative, or unimodal relationships of genetic diversity with species diversity. When species are considered as part of multispecies communities, predictions from single-species models of genetic diversity apply in some instances (effects of area and immigration) but often not in others (effects of environmental heterogeneity).     

Importance of scale,land‐use,and stream network properties for riparian plant communities along an urban gradient

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Environmental drivers of α-diversity patterns in monsoonal tropical stream fish assemblages: a case study from tributaries of Narmada basin,India

Environmental factors, such as local physico-chemical water parameters, habitat-structural conditions, geography and human disturbances can have a profound impact on the structure and composition of biotic communities. This study investigates the relationship of these factors with fish diversity in medium to small streams in Central India and identifies the key determinants of species richness and diversity in habitats increasingly facing impacts of urban development. Data on fish diversity on eighteen sites, representative of undisturbed and disturbed aquatic habitats, located in the Narmada river basin were collected across different seasons over two years. Seasonal patterns of alpha diversity for this region were analysed and the effect of environmental drivers on seasonal patterns of alpha diversity were assessed. Analyses of species data revealed significant differences in diversity patterns across seasons for both species richness and Shannon diversity. Results of generalized linear mixed models showed that along with disturbance, altitude and substrate heterogeneity, dissolved oxygen, pH and stream width proved to be important environmental variables that predict species richness. Further, variations in pH, dissolved oxygen, temperature, stream width, stream depth and stream width heterogeneity were found to also predict Shannon diversity index. Our study revealed that seasonal changes across the year play a crucial role in shaping diversity patterns. This study is relevant as an important step in identifying roles of various ecological factors driving fish diversity in the region and towards developing long term management plans for critical catchment areas of major rivers in tropical systems.     

Hydrological and environmental variables outperform spatial factors in structuring species,trait composition,and beta diversity of pelagic algae

There has been increasing interest in algae‐based bioassessment, particularly, trait‐based approaches are increasingly suggested. However, the main drivers, especially the contribution of hydrological variables, of species composition, trait composition, and beta diversity of algae communities are less studied. To link species and trait composition to multiple factors (i.e., hydrological variables, local environmental variables, and spatial factors) that potentially control species occurrence/abundance and to determine their relative roles in shaping species composition, trait composition, and beta diversities of pelagic algae communities, samples were collected from a German lowland catchment, where a well‐proven ecohydrological modeling enabled to predict long‐term discharges at each sampling site. Both trait and species composition showed significant correlations with hydrological, environmental, and spatial variables, and variation partitioning revealed that the hydrological and local environmental variables outperformed spatial variables. A higher variation of trait composition (57.0%) than species composition (37.5%) could be explained by abiotic factors. Mantel tests showed that both species and trait‐based beta diversities were mostly related to hydrological and environmental heterogeneity with hydrological contributing more than environmental variables, while purely spatial impact was less important. Our findings revealed the relative importance of hydrological variables in shaping pelagic algae community and their spatial patterns of beta diversities, emphasizing the need to include hydrological variables in long‐term biomonitoring campaigns and biodiversity conservation or restoration. A key implication for biodiversity conservation was that maintaining the instream flow regime and keeping various habitats among rivers are of vital importance. However, further investigations at multispatial and temporal scales are greatly needed.     

Genetic variation of loci potentially under selection confounds species–genetic diversity correlations in a fragmented habitat

Positive species–genetic diversity correlations (SGDCs) are often thought to result from the parallel influence of neutral processes on genetic and species diversity. Yet, confounding effects of non‐neutral mechanisms have not been explored. Here, we investigate the impact of non‐neutral genetic diversity on SGDCs in high Andean wetlands. We compare correlations between plant species diversity and genetic diversity (GD) calculated with and without loci potentially under selection (outlier loci). The study system includes 2188 specimens from five species (three common aquatic macroinvertebrate and two dominant plant species) that were genotyped for 396 amplified fragment length polymorphism loci. We also appraise the importance of neutral processes on SGDCs by investigating the influence of habitat fragmentation features. Significant positive SGDCs were detected for all five species (mean SGDC = 0.52 ± 0.05). While only a few outlier loci were detected in each species, they resulted in significant decreases in GD and in SGDCs. This supports the hypothesis that neutral processes drive species–genetic diversity relationships in high Andean wetlands. Unexpectedly, the effects on genetic diversity GD of the habitat fragmentation characteristics in this study increased with the presence of outlier loci in two species. Overall, our results reveal pitfalls in using habitat features to infer processes driving SGDCs and show that a few loci potentially under selection are enough to cause a significant downward bias in SGDC. Investigating confounding effects of outlier loci thus represents a useful approach to evidence the contribution of neutral processes on species–genetic diversity relationships.     

Agriculture erases climate constraints on soil nematode communities across large spatial scales

Anthropogenic conversion of natural to agricultural land reduces aboveground biodiversity. Yet, the overall consequences of land‐use changes on belowground biodiversity at large scales remain insufficiently explored. Furthermore, the effects of conversion on different organism groups are usually determined at the taxonomic level, while an integrated investigation that includes functional and phylogenetic levels is rare and absent for belowground organisms. Here, we studied the Earth's most abundant metazoa—nematodes—to examine the effects of conversion from natural to agricultural habitats on soil biodiversity across a large spatial scale. To this aim, we investigated the diversity and composition of nematode communities at the taxonomic, functional, and phylogenetic level in 16 assemblage pairs (32 sites in total with 16 in each habitat type) in mainland China. While the overall alpha and beta diversity did not differ between natural and agricultural systems, all three alpha diversity facets decreased with latitude in natural habitats. Both alpha and beta diversity levels were driven by climatic differences in natural habitats, while none of the diversity levels changed in agricultural systems. This indicates that land conversion affects soil biodiversity in a geographically dependent manner and that agriculture could erase climatic constraints on soil biodiversity at such a scale. Additionally, the functional composition of nematode communities was more dissimilar in agricultural than in natural habitats, while the phylogenetic composition was more similar, indicating that changes among different biodiversity facets are asynchronous. Our study deepens the understanding of land‐use effects on soil nematode diversity across large spatial scales. Moreover, the detected asynchrony of taxonomic, functional, and phylogenetic diversity highlights the necessity to monitor multiple facets of soil biodiversity in ecological studies such as those investigating environmental changes.     

Variation in habitat connectivity generates positive correlations between species and genetic diversity in a metacommunity

An increasing number of studies are simultaneously investigating species diversity (SD) and genetic diversity (GD) in the same systems, looking for ‘species– genetic diversity correlations’ (SGDCs). From negative to positive SGDCs have been reported, but studies have generally not quantified the processes underlying these correlations. They were also mostly conducted at large biogeographical scales or in recently degraded habitats. Such correlations have not been looked for in natural networks of connected habitat fragments (metacommunities), and the underlying processes remain elusive in most systems. We investigated these issues by studying freshwater snails in a pond network in Guadeloupe (Lesser Antilles). We recorded SD and habitat characteristics in 232 ponds and assessed GD in 75 populations of two species. Strongly significant and positive SGDCs were detected in both species. Based on a decomposition of SGDC as a function of variance–covariance of habitat characteristics, we showed that connectivity (opportunity of water flow between a site and the nearest watershed during the rainy season) has the strongest contribution on SGDCs. More connective sites received both more alleles and more species through immigration resulting in both higher GD and higher SD. Other habitat characteristics did not contribute, or contributed negatively, to SGDCs. This is true of the desiccation frequency of ponds during the dry season, presumably because species markedly differ in their ability to tolerate desiccation. Our study shows that variation in environmental characteristics of habitat patches can promote SGDCs at metacommunity scale when the studied species respond homogeneously to these environmental characteristics.     

云南省把边江流域不同土地利用方式对大型底栖动物群落的影响

研究土地利用对大型底栖动物群落的影响,对流域土地利用规划修编和河流生态修复等有重要意义。以红河上游的把边江流域为研究区,调查大型底栖动物群落现状,计算37个样点上游集水区土地利用组分、配置、多样性和水文距离4个方面20个土地利用指标,分析不同土地利用方式对大型底栖动物群落的影响。本次调查共鉴定出25个大型底栖动物分类单元,其中水生昆虫最多,占分类单元总数的72%;摄食功能群以捕食者（PR）、刮食者（SC）和直接收集者（GC）为主。把边江流域城镇用地和林地对大型底栖动物群落的影响占主导地位,而耕地对大型底栖动物群落的影响较小。影响把边江流域大型底栖动物群落的关键土地利用指标为城镇用地面积比（3PLA）、城镇用地最大斑块指数（3LPI）、城镇用地聚合指数（3AI）、城镇用地水流长度（3FLOW）、林地最大斑块指数（1LPI）和林地形状指数（1LSI）。土地利用组分、配置和水文距离是影响把边江流域大型底栖动物群落的主要方面,而土地利用多样性对大型底栖动物群落的影响较小。大型底栖动物物种密度和物种多样性与3PLA、3LPI、3AI、3FLOW和1LSI呈负相关,与1LPI呈正相关。1LPI与浮游目（如小蜉科、扁蜉科）为代表的清洁物种呈现较明显的正相关关系。城镇用地指标与SC和SH（撕食者）密度具有显著的负相关关系,与GC密度有较弱的正相关关系,而与其他功能群没有明显的相关关系。SC和SH对水质和环境条件敏感,而GC对人类干扰耐受性较强。1LPI与SC和FC（滤食收集者密度）正相关,而与PR和GC密度负相关。SC和FC对生境的完整性要求较高,而GC和PR对人为干扰和较为破碎的生境更为适应。一方面,随着城镇用地的规模、边界复杂性和聚集程度增加,研究区大型底栖动物密度和多样性降低;另一方面,随着林地完整性降低,大型底栖动物物种均匀度降低。土地利用与大型底栖动物物种多样性和功能群多样性的关系不完全一致,1LSI和3FLOW与物种多样性负相关,而与功能群多样性呈正相关关系,土地利用对物种-功能群-群落不同尺度上的影响差异还需要更深入的研究。     

Response diversity,nonnative species,and disassembly rules buffer freshwater ecosystem processes from anthropogenic change

Integrating knowledge of environmental degradation, biodiversity change, and ecosystem processes across large spatial scales remains a key challenge to illuminating the resilience of earth's systems. There is now a growing realization that the manner in which communities will respond to anthropogenic impacts will ultimately control the ecosystem consequences. Here, we examine the response of freshwater fishes and their nutrient excretion – a key ecosystem process that can control aquatic productivity – to human land development across the contiguous United States. By linking a continental‐scale dataset of 533 fish species from 8100 stream locations with species functional traits, nutrient excretion, and land remote sensing, we present four key findings. First, we provide the first geographic footprint of nutrient excretion by freshwater fishes across the United States and reveal distinct local‐ and continental‐scale heterogeneity in community excretion rates. Second, fish species exhibited substantial response diversity in their sensitivity to land development; for native species, the more tolerant species were also the species contributing greater ecosystem function in terms of nutrient excretion. Third, by modeling increased land‐use change and resultant shifts in fish community composition, land development is estimated to decrease fish nutrient excretion in the majority (63%) of ecoregions. Fourth, the loss of nutrient excretion would be 28% greater if biodiversity loss was random or 84% greater if there were no nonnative species. Thus, ecosystem processes are sensitive to increased anthropogenic degradation but biotic communities provide multiple pathways for resistance and this resistance varies across space.     

Threshold responses of riverine fish communities to land use conversion across regions of the world

The growing human enterprise has sparked greater interest in identifying ecological thresholds in land use conversion beyond which populations or communities demonstrate abrupt nonlinear or substantive change in species composition. Such knowledge remains fundamental to understanding ecosystem resilience to environmental degradation and informing land use planning into the future. Confronting this challenge has been largely limited to inferring thresholds in univariate metrics of species richness and indices of biotic integrity and has largely ignored how land use legacies of the past may shape community responses of today. By leveraging data for 13,069 riverine sites from temperate, subtropical, and boreal climate zones on four continents, we characterize patterns of community change along diverse gradients of urbanization and agricultural land use, and identity threshold values beyond which significant alterations in species composition exists. Our results demonstrate the apparent universality by which freshwater fish communities are sensitive to even low levels of watershed urbanization (range of threshold values: 1%–12%), but consistently higher (and more variable) levels of agricultural development (2%–37%). We demonstrated that fish community compositional thresholds occurred, in general, at lower levels of watershed urbanization and agriculture when compared to threshold responses in species richness. This supports the notion that aggregated taxon‐specific responses may better reflect the complexity of assemblage responses to land use development. We further revealed that the ghost of land use past plays an important role in moderating how current‐day fish communities respond to land use intensification. Subbasins of the United States experiencing greater rates of past land use change demonstrated higher current‐day thresholds. Threshold responses of community composition, such as those identified in our study, illustrate the need for globally coordinated efforts to prioritize country‐specific management and policy initiatives that ensure that freshwater fish diversity is not inevitably lost in the future.     

Drawing ecological inferences from coincident patterns of population‐ and community‐level biodiversity

Biodiversity is comprised of genetic and phenotypic variation among individual organisms, which might belong to the same species or to different species. Spatial patterns of biodiversity are of central interest in ecology and evolution for several reasons: to identify general patterns in nature (e.g. species–area relationships, latitudinal gradients), to inform conservation priorities (e.g. identifying hotspots, prioritizing management efforts) and to draw inferences about processes, historical or otherwise (e.g. adaptation, the centre of origin of particular clades). There are long traditions in ecology and evolutionary biology of examining spatial patterns of biodiversity among species (i.e. in multispecies communities) and within species, respectively, and there has been a recent surge of interest in studying these two types of pattern simultaneously. The idea is that examining both levels of diversity can materially advance the above‐stated goals and perhaps lead to entirely novel lines of inquiry. Here, we review two broad categories of approach to merging studies of inter‐ and intraspecific variation: (i) the study of phenotypic trait variation along environmental gradients and (ii) the study of relationships between patterns of molecular genetic variation within species and patterns of distribution and diversity across species. For the latter, we report a new meta‐analysis in which we find that correlations between species diversity and genetic diversity are generally positive and significantly stronger in studies with discrete sampling units (e.g. islands, lakes, forest fragments) than in studies with nondiscrete sampling units (e.g. equal‐area study plots). For each topic, we summarize the current state of knowledge and key future directions.     

Order and disorder in the river continuum: the contribution of continuity and connectivity to floodplain meadow biodiversity

Aim Aspects of connectivity and continuity operating in the River Meuse were analysed for their contribution to the biodiversity of the floodplain. From this analysis of the diversity and composition of the meadow communities, we aimed to derive effective biodiversity conservation strategies. Location The River Meuse is one of the larger rivers in the European Western Plains ecoregion. The alluvial plains of the river have a long history of cultivation, and for these plains the floodplain meadow vegetation is a highly appreciated and valuable nature conservation asset. Method We sampled floodplain meadows from 400 km of the six geomorphic reaches of the middle to lower course of the River Meuse. For each, 50 vascular plant relevés were recorded, representing the spectrum of floodplain meadow communities of that reach. Beta diversity was calculated to quantify similarity in species pools between the reaches. A dissimilarity formula was used to determine the turnover between the reaches, and these dissimilarities were compared with a Mantel test to detect whether species composition of the floodplain meadows exhibited connectivity and continuity between the reaches. Species richness for the floodplain vegetation data of the reaches was compared with data for riparian invertebrate communities. The vegetation data sets were ordinated using detrended correspondence analysis (DCA) to reveal patterns in the floodplain meadow species composition, and the DCA axes were related to plant functional groups and population strategies. The axis scores of the species and plots were linked to river and plant species traits. Results We did not observe an overall continuity trend in similarity, nor one in diversity in a downstream direction. Lateral connectivity was highlighted by the dissimilarity between the reaches and in the influxes of species from adjacent ecoregions. The DCA ordination showed statistically significant separations between reaches and between the plant functional groups. The second DCA axis related to the longitudinal gradient of the river, whereas the first axis showed stronger correlations with river and plant species traits. We termed this axis the ‘disorder axis’. Plant invaders and avoiders are located at the left side of the disorder axis, whereas the true river‐adapted categories of resisters and endurers are at the right extremity. Contributions to the disorder were identified in terms of connectivity with adjacent ecoregions and the physical disturbance regime of natural and anthropogenic perturbations, resulting in community changes between the reaches. Main conclusions We concluded that a single overall strategy for biodiversity conservation of the river or its floodplains is not feasible. Strategies can, however, be derived for separate river reaches based on functional groups in the communities, the disorder characteristics of the reach, and the influence of surrounding ecoregions.     

城镇化背景下深圳典型流域鱼类群落特征及驱动因子

理解城镇的快速发展对河流鱼类群落结构的影响,是城镇河流科学管理和生物多样性保护的关键基础。本研究于2019年丰水期（8月）和枯水期（11月）,选取我国城镇化典型城市-深圳域内两个处于不同城镇化程度的代表性流域,应用多重统计方法比较分析了流域间鱼类群落结构的差异,并探讨了驱动鱼类群落变异的关键环境要素。结果发现,城镇化程度高的观澜河流域其鱼类种类组成、优势类群、生物多样性指数与城镇化程度低的坪山河流域有明显差别。 具体表现为：城镇化程度高的流域土著敏感种类如异鱲、吸鳅等几近消失,优势类群为外来入侵耐受种类,其物种多样性显著低于城镇化程度低的流域（P<0.05）。同时,外来鱼类在城镇河段其数量占比平均达92.5%,广泛分布于深圳城镇河流中。在环境因素方面,城镇化程度高的观澜河流域水体理化指标总氮、总磷、氨氮、化学需氧量、生化需氧量、高锰酸盐指数均显著性高于城镇化程度低的坪山河流域（P<0.05）。基于Bray-Curtis距离的冗余分析显示：城镇用地占比和总氮是影响观澜河和坪山河流域鱼类群落差异的主要因素。城镇化进程中河流生境的改变已影响到土著鱼类的生物多样性。因此,推动以恢复土著鱼类生物多样性的河流生态治理与保护是今后水生态目标管理的重要方向。