Extreme drought pushes stream invertebrate communities over functional thresholds

Functional traits are increasingly being used to predict extinction risks and range shifts under long‐term climate change scenarios, but have rarely been used to study vulnerability to extreme climatic events, such as supraseasonal droughts. In streams, drought intensification can cross thresholds of habitat loss, where marginal changes in environmental conditions trigger disproportionate biotic responses. However, these thresholds have been studied only from a structural perspective, and the existence of functional nonlinearity remains unknown. We explored trends in invertebrate community functional traits along a gradient of drought intensity, simulated over 18 months, using mesocosms analogous to lowland headwater streams. We modelled the responses of 16 traits based on a priori predictions of trait filtering by drought, and also examined the responses of trait profile groups (TPGs) identified via hierarchical cluster analysis. As responses to drought intensification were both linear and nonlinear, generalized additive models (GAMs) were chosen to model response curves, with the slopes of fitted splines used to detect functional thresholds during drought. Drought triggered significant responses in 12 (75%) of the a priori‐selected traits. Behavioural traits describing movement (dispersal, locomotion) and diet were sensitive to moderate‐intensity drought, as channels fragmented into isolated pools. By comparison, morphological and physiological traits showed little response until surface water was lost, at which point we observed sudden shifts in body size, respiration mode and thermal tolerance. Responses varied widely among TPGs, ranging from population collapses of non‐aerial dispersers as channels fragmented to irruptions of small, eurythermic dietary generalists upon extreme dewatering. Our study demonstrates for the first time that relatively small changes in drought intensity can trigger disproportionately large functional shifts in stream communities, suggesting that traits‐based approaches could be particularly useful for diagnosing catastrophic ecological responses to global change.     

人为干扰对溪流鱼类功能多样性及其纵向梯度格局的影响

溪流鱼类多样性沿着河流纵向梯度的空间分布规律已得到大量报道, 但这些研究大多聚焦基于物种组成的分类α多样性, 而有关分类β多样性和功能多样性的纵向梯度分布规律及其对人类干扰的响应研究较少。本文以青弋江上游3条人为干扰程度不同的河源溪流为研究区域, 比较研究了人为干扰对溪流鱼类功能α和β多样性及其纵向梯度分布格局的影响。结果显示, 人类干扰改变了河源溪流鱼类功能多样性的纵向梯度格局——由线性变化变为二项式分布。此外, 我们发现, 人为干扰导致土著种被本地入侵种取代, 且较强的土地利用和水污染排放可能增大环境的不连续性, 而群落周转和嵌套变化往往取决于环境的变化。尽管功能β多样性由嵌套成分主导, 但周转成分占比相对于人为干扰较小的溪流而言明显增加。人为干扰显著改变了受干扰溪流鱼类的物种组成和功能多样性, 且功能多样性的纵向梯度格局在不同的多样性指标上存在差异。本研究强调, 在评估人为干扰下多样性的变化时, 需要从多方面考虑, 包括空间尺度和多样性指标等。     

Structural and functional responses of invertebrate communities to climate change and flow regulation in alpine catchments

Understanding and predicting how biological communities respond to climate change is critical for assessing biodiversity vulnerability and guiding conservation efforts. Glacier‐ and snow‐fed rivers are one of the most sensitive ecosystems to climate change, and can provide early warning of wider‐scale changes. These rivers are frequently used for hydropower production but there is minimal understanding of how biological communities are influenced by climate change in a context of flow regulation. This study sheds light on this issue by disentangling structural (water temperature preference, taxonomic composition, alpha, beta and gamma diversities) and functional (functional traits, diversity, richness, evenness, dispersion and redundancy) effects of climate change in interaction with flow regulation in the Alps. For this, we compared environmental and aquatic invertebrate data collected in the 1970s and 2010s in regulated and unregulated alpine catchments. We hypothesized a replacement of cold‐adapted species by warming‐tolerant ones, high temporal and spatial turnover in taxa and trait composition, along with reduced taxonomic and functional diversities in consequence of climate change. We expected communities in regulated rivers to respond more drastically due to additive or synergistic effects between flow regulation and climate change. We found divergent structural but convergent functional responses between free‐flowing and regulated catchments. Although cold‐adapted taxa decreased in both of them, greater colonization and spread of thermophilic species was found in the free‐flowing one, resulting in higher spatial and temporal turnover. Since the 1970s, taxonomic diversity increased in the free flowing but decreased in the regulated catchment due to biotic homogenization. Colonization by taxa with new functional strategies (i.e. multivoltine taxa with small body size, resistance forms, aerial dispersion and reproduction by clutches) increased functional diversity but decreased functional redundancy through time. These functional changes could jeopardize the ability of aquatic communities facing intensification of ongoing climate change or new anthropogenic disturbances.     

Flow intermittency alters longitudinal patterns of invertebrate diversity and assemblage composition in an arid‐land stream network

1. Temporary streams comprise a large proportion of the total length of most stream networks, and the great majority of arid‐land stream networks, so it is important to understand their contribution to biotic diversity at both local and landscape scales. 2. In late winter 2010, we sampled invertebrate assemblages in 12 reaches of a large arid‐land stream network (including perennial and intermittent headwaters, intermittent middle reaches and perennial rivers) in south‐east Arizona, U.S.A. Intermittent reaches had then been flowing for c. 60 days, following a dry period of more than 450 days. We sampled a subset of the perennial study reaches three more times between 2009 and 2011. Since intermittent reaches were dry during these additional sampling periods, we used assemblage data from two other intermittent streams in the study network (sampled in 2004–05 and 2010) to explore interannual variability in intermittent stream assemblage composition. 3. Invertebrate richness was lowest in intermittent reaches, despite their often being connected to species‐rich perennial reaches. The assemblages of these intermittent reaches were not simply a subset of the species in perennial streams, but rather were dominated by a suite of stoneflies, blackflies and midges with adaptations to intermittency (e.g. egg and/or larval diapause). On average, 86% of individuals in these samples were specialists or exclusive to intermittent streams. Predators were 7–14 times more abundant in perennial than in intermittent reaches. 4. Despite being separated by long distances (12–25 km) and having very different physical characteristics, the assemblages of perennial headwaters and rivers were more similar to one another than to intervening intermittent reaches, emphasising the prime importance of local hydrology in this system. 5. The duration and recurrence intervals of dry periods, and the relative importance of dispersal from perennial refuges, probably influence the magnitude of biological differences between neighbouring perennial and temporary streams. Although perennial headwaters supported the highest diversity of invertebrates, intermittent reaches supported a number of unique or locally rare species and as such contribute to regional species diversity and should be included in conservation planning.     

Impacts of climatic stability on the structural and functional aspects of macroinvertebrate communities after severe drought

1. Unpredictable, extreme climatic events (e.g. droughts) can potentially destabilize aquatic communities. From 1998 to 2002, southwestern Georgia, U.S.A., experienced the third worst drought of the last 100 years, leading to loss of surface flow in many small streams. We sampled macroinvertebrates, flow and water chemistry in small headwater streams from 2001 to 2007 in two adjacent coastal plain streams of contrasting headwater type (wetland and groundwater‐fed seep) following resumption of flow. 2. Decreasing water temperature, conductivity and nutrient concentrations within the first 2 years of the study indicated flushing of the streambed associated with increased discharge. Invertebrate community composition became less variable over time and during wetter years, reflecting water chemistry, hydrological and climate conditions. 3. A core set of species appeared immediately following breaking of the drought in both streams, reflecting a shared species pool. These species exhibited resilience traits, including short life cycles and resistance to desiccation, which allowed for rapid recovery from disturbance. Such species, which were small‐bodied, sclerotized and abundant in the drift, were then replaced as flows increased by those that were larger, soft‐bodied and rare in drift, suggesting a more stable and less ephemeral habitat. 4. Hydrologic regime and long‐term precipitation indices were strongly correlated with invertebrate community and trait structure. Long‐term data allowed for better interpretation of the effects of infrequent disturbances on aquatic ecosystems. Additionally, long‐term precipitation indices (i.e. 48‐month standardized precipitation index) can indicate the likelihood of a return to drought, allowing for the collection of pre‐disturbance data.     

Invertebrate assemblages of pools in arid‐land streams have high functional redundancy and are resistant to severe drying

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Drought survival strategies,dispersal potential and persistence of invertebrate species in an intermittent stream landscape

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Drought ecohydrology alters the structure and function of benthic invertebrate communities in mountain streams

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Climate extremes are associated with invertebrate taxonomic and functional composition in mountain lakes

Climate change is expected to increase climate variability and the occurrence of extreme climatic events, with potentially devastating effects on aquatic ecosystems. However, little is known about the role of climate extremes in structuring aquatic communities or the interplay between climate and local abiotic and biotic factors. Here, we examine the relative influence of climate and local abiotic and biotic conditions on biodiversity and community structure in lake invertebrates. We sampled aquatic invertebrates and measured environmental variables in 19 lakes throughout California, USA, to test hypotheses of the relationship between climate, local biotic and environmental conditions, and the taxonomic and functional structure of aquatic invertebrate communities. We found that, while local biotic and abiotic factors such as habitat availability and conductivity were the most consistent predictors of alpha diversity, extreme climate conditions such as maximum summer temperature and dry‐season precipitation were most often associated with multivariate taxonomic and functional composition. Specifically, sites with high maximum temperatures and low dry‐season precipitation housed communities containing high abundances of large predatory taxa. Furthermore, both climate dissimilarity and abiotic dissimilarity determined taxonomic turnover among sites (beta diversity). These findings suggest that while local‐scale environmental variables may predict alpha diversity, climatic variability is important to consider when projecting broad‐scale aquatic community responses to the extreme temperature and precipitation events that are expected for much of the world during the next century.     

Drought intensification alters the composition,body size,and trophic structure of invertebrate assemblages in a stream mesocosm experiment

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A community and functional comparison of coral and reef fish assemblages between four decades of coastal urbanisation and thermal stress

1. Urbanized coral reefs experience anthropogenic disturbances caused by coastal development, pollution, and nutrient runoff, resulting in turbid, marginal conditions in which only certain species can persist. Mortality effects are exacerbated by increasingly regular thermal stress events, leading to shifts towards novel communities dominated by habitat generalists and species with low structural complexity.
2. There is limited data on the turnover processes that occur due to this convergence of anthropogenic stressors, and how novel urban ecosystems are structured both at the community and functional levels. As such, it is unclear how they will respond to future disturbance events.
3. Here, we examine the patterns of coral reef community change and determine whether ecosystem functions provided by specialist species are lost post‐disturbance. We present a comparison of community and functional trait‐based changes for scleractinian coral genera and reef fish species assemblages subject to coastal development, coastal modification, and mass bleaching between two time periods, 1975–1976 and 2018, in Nakagusuku Bay, Okinawa, Japan.
4. We observed an increase in fish habitat generalists, a dominance shift from branching to massive/sub‐massive corals and increasing site‐based coral genera richness between years. Fish and coral communities significantly reassembled, but functional trait‐based multivariate space remained constant, indicating a turnover of species with similar traits. A compression of coral habitat occurred, with shallow (<5 m) and deep (>8 m) coral genera shifting towards the mid‐depths (5–8 m).
5. We show that although reef species assemblages altered post disturbance, new communities retained similar ecosystem functions. This result could be linked to the stressors experienced by urban reefs, which reflect those that will occur at an increasing frequency globally in the near future. Yet, even after shifts to disturbed communities, these fully functioning reef systems may maintain high conservation value.

     

Fish communities diverge in species but converge in traits over three decades of warming

Describing the spatial and temporal dynamics of communities is essential for understanding the impacts of global environmental change on biodiversity and ecosystem functioning. Trait‐based approaches can provide better insight than species‐based (i.e. taxonomic) approaches into community assembly and ecosystem functioning, but comparing species and trait dynamics may reveal important patterns for understanding community responses to environmental change. Here, we used a 33‐year database of fish monitoring to compare the spatio‐temporal dynamics of taxonomic and trait structure in North Sea fish communities. We found that the majority of variation in both taxonomic and trait structure was explained by a pronounced spatial gradient, with distinct communities in the southern and northern North Sea related to depth, sea surface temperature, salinity and bed shear stress. Both taxonomic and trait structure changed significantly over time; however taxonomically, communities in the south and north diverged towards different species, becoming more dissimilar over time, yet they converged towards the same traits regardless of species differences. In particular, communities shifted towards smaller, faster growing species with higher thermal preferences and pelagic water column position. Although taxonomic structure changed over time, its spatial distribution remained relatively stable, whereas in trait structure, the southern zone of the North Sea shifted northward and expanded, leading to homogenization. Our findings suggest that global environmental change, notably climate warming, will lead to convergence towards traits more adapted for novel environments regardless of species composition.     

Exploring the homogeneity of terrestrial subterranean communities at a local spatial scale

1. Although caves are generally perceived as isolated habitats, at the local scale, they are often interconnected via a network of fissures in the bedrock. Accordingly, caves in close proximity are expected to host the same, or very similar, animal communities. 2. We explored the extent to which subterranean arthropod communities are homogenous at a local spatial scale of less than 1 km², along with which cave-specific environmental features result in a departure from the expected homogeneous pattern. We approached this question by studying richness and turnover in terrestrial invertebrate communities of 27 caves in a small karst massif in the Western Italian Alps. 3. Specialised subterranean species were homogeneously distributed among caves and were not influenced by seasonality. The only factor driving their distribution was the distance from the cave entrance, with deeper caves yielding a greater diversity of species. 4. Significant spatio-temporal turnover in species not specialised to subterranean life was observed. In summer, there was a significant homogenisation of the community and a more even distribution of species among sites; in winter, these species were missing or found exclusively at greater depths, where environmental conditions were more stable. Furthermore, caves at lower elevations yielded, on average, a greater diversity and abundance of these species. 5. This study demonstrated that the theoretical expectation of no turnover in community composition in caves in close proximity is not always met. Turnover can be mostly attributed to seasonal patterns and sampling depth; thus, our findings have implications for planning sampling and monitoring activities in caves.     

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。     

Context‐dependent functional dispersion across similar ranges of trait space covered by intertidal rocky shore communities

Functional diversity is intimately linked with community assembly processes, but its large‐scale patterns of variation are often not well understood. Here, we investigated the spatiotemporal changes in multiple trait dimensions (“trait space”) along vertical intertidal environmental stress gradients and across a landscape scale. We predicted that the range of the trait space covered by local assemblages (i.e., functional richness) and the dispersion in trait abundances (i.e., functional dispersion) should increase from high‐ to low‐intertidal elevations, due to the decreasing influence of environmental filtering. The abundance of macrobenthic algae and invertebrates was estimated at four rocky shores spanning ca. 200 km of the coast over a 36‐month period. Functional richness and dispersion were contrasted against matrix‐swap models to remove any confounding effect of species richness on functional diversity. Random‐slope models showed that functional richness and dispersion significantly increased from high‐ to low‐intertidal heights, demonstrating that under harsh environmental conditions, the assemblages comprised similar abundances of functionally similar species (i.e., trait convergence), while that under milder conditions, the assemblages encompassed differing abundances of functionally dissimilar species (i.e., trait divergence). According to the Akaike information criteria, the relationship between local environmental stress and functional richness was persistent across sites and sampling times, while functional dispersion varied significantly. Environmental filtering therefore has persistent effects on the range of trait space covered by these assemblages, but context‐dependent effects on the abundances of trait combinations within such range. Our results further suggest that natural and/or anthropogenic factors might have significant effects on the relative abundance of functional traits, despite that no trait addition or extinction is detected.