Mass coral bleaching causes biotic homogenization of reef fish assemblages

Global climate change is altering community composition across many ecosystems due to nonrandom species turnover, typically characterized by the loss of specialist species and increasing similarity of biological communities across spatial scales. As anthropogenic disturbances continue to alter species composition globally, there is a growing need to identify how species responses influence the establishment of distinct assemblages, such that management actions may be appropriately assigned. Here, we use trait‐based analyses to compare temporal changes in five complementary indices of reef fish assemblage structure among six taxonomically distinct coral reef habitats exposed to a system‐wide thermal stress event. Our results revealed increased taxonomic and functional similarity of previously distinct reef fish assemblages following mass coral bleaching, with changes characterized by subtle, but significant, shifts toward predominance of small‐bodied, algal‐farming habitat generalists. Furthermore, while the taxonomic or functional richness of fish assemblages did not change across all habitats, an increase in functional originality indicated an overall loss of functional redundancy. We also found that prebleaching coral composition better predicted changes in fish assemblage structure than the magnitude of coral loss. These results emphasize how measures of alpha diversity can mask important changes in the structure and functioning of ecosystems as assemblages reorganize. Our findings also highlight the role of coral species composition in structuring communities and influencing the diversity of responses of reef fishes to disturbance. As new coral species configurations emerge, their desirability will hinge upon the composition of associated species and their capacity to maintain key ecological processes in spite of ongoing disturbances.     

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.     

Four decades of functional community change reveals gradual trends and low interlinkage across trophic groups in a large marine ecosystem

The rate at which biological diversity is altered on both land and in the sea, makes temporal community development a critical and fundamental part of understanding global change. With advancements in trait‐based approaches, the focus on the impact of temporal change has shifted towards its potential effects on the functioning of the ecosystems. Our mechanistic understanding of and ability to predict community change is still impeded by the lack of knowledge in long‐term functional dynamics that span several trophic levels. To address this, we assessed species richness and multiple dimensions of functional diversity and dynamics of two interacting key organism groups in the marine food web: fish and zoobenthos. We utilized unique time series‐data spanning four decades, from three environmentally distinct coastal areas in the Baltic Sea, and assembled trait information on six traits per organism group covering aspects of feeding, living habit, reproduction and life history. We identified gradual long‐term trends, rather than abrupt changes in functional diversity (trait richness, evenness, dispersion) trait turnover, and overall multi‐trait community composition. The linkage between fish and zoobenthic functional community change, in terms of correlation in long‐term trends, was weak, with timing of changes being area and trophic group specific. Developments of fish and zoobenthos traits, particularly size (increase in small size for both groups) and feeding habits (e.g. increase in generalist feeding for fish and scavenging or predation for zoobenthos), suggest changes in trophic pathways. We summarize our findings by highlighting three key aspects for understanding functional change across trophic groups: (a) decoupling of species from trait richness, (b) decoupling of richness from density and (c) determining of turnover and multi‐trait dynamics. We therefore argue for quantifying change in multiple functional measures to help assessments of biodiversity change move beyond taxonomy and single trophic groups.     

Coral bleaching, reef fish community phase shifts and the resilience of coral reefs

The 1998 global coral bleaching event was the largest recorded historical disturbance of coral reefs and resulted in extensive habitat loss. Annual censuses of reef fish community structure over a 12-year period spanning the bleaching event revealed a marked phase shift from a prebleach to postbleach assemblage. Surprisingly, we found that the bleaching event had no detectable effect on the abundance, diversity or species richness of a local cryptobenthic reef fish community. Furthermore, there is no evidence of regeneration even after 5–35 generations of these short-lived species. These results have significant implications for our understanding of the response of coral reef ecosystems to global warming and highlight the importance of selecting appropriate criteria for evaluating reef resilience.     

Co-occurrence of herbivorous fish functional groups correlates with enhanced coral reef benthic state

Aim

Biodiversity loss is impacting essential ecosystem functions and services across the globe. Recently, our interest in the benefits of biodiversity for ecosystem function has shifted focus from measurements of species richness to functional diversity and composition. However, the additional importance of other community characteristics, such as species evenness and co-occurrence, for diversity-driven ecosystem function is less known. We used herbivorous coral reef fish as a model system to investigate how co-occurrence of different functional groups, rather than purely functional diversity, within an assemblage might affect the coral reef benthic state.

Location

Western Atlantic.

Time period

2007–2017.

Major taxa studied

Herbivorous reef fish.

Methods

We analysed benthic and fish assemblage data from 601 sites across 12 countries in the western Atlantic. Using diversity–interaction models, we investigated how the composition and relative abundances of reef fish functional groups were correlated with benthic cover and estimates of coral calcification rates. We used statistical interactions to explore the importance of co-occurrence of herbivorous fish functional groups for the coral reef benthic state.

Results

We found that co-occurrence of herbivorous fish functional groups, in addition to functional diversity, was correlated with reduced algal cover and increased coral accretion. Moreover, pairwise statistical interactions between functional groups were significantly correlated with an improvement in the coral reef benthic state.

Main conclusions

Our results support the idea that functional group co-occurrence, in addition to functional diversity, within herbivorous fish offers additional benefits to the coral reef benthic state. We identify farming damselfish and excavating parrotfish as potential key determinants of the coral reef benthic state and highlight that co-occurrence of cropping and scraping herbivores might promote coral accretion. Our findings support the argument that protecting herbivore abundance without regard to the species and functional groups present is not enough to preserve coral reef health and that fine-scale community composition must be considered.     

Interactions between herbivorous fish guilds and their influence on algal succession on a coastal coral reef

Herbivory is an important mechanism affecting algal succession, particularly on coral reefs where the relationship between algae and corals is largely controlled by herbivores. However, different functional groups of herbivores may have contrasting effects on succession, which may explain different trajectories of coral reef recovery after disturbance. Here, the effects of different herbivore groups (roving herbivores = foragers and territorial damselfish = farmers) were isolated by a multi-factorial experiment carried out on a coastal coral reef with high macroalgal cover, high farmer densities and relatively low forager abundance. The effects of foragers and farmers were distinguished by monitoring algal succession on settlement tiles placed inside and outside exclusion cages, with orthogonal treatments established inside and outside damselfish territories (with appropriate cage controls). Within 12 months, algal assemblages on ungrazed tiles inside exclusion cages proceeded rapidly from fine filamentous turfs, to corticated algae, to tough erect (e.g. Amphiroa spp.) and foliose (e.g. Peyssonnellidae) calcified algae. Farmers had a dramatic impact on succession, essentially arresting the development of the algal community at a point where it was dominated by palatable filamentous algae of the genus Polysiphonia. Fleshy macroalgae such as Sargassum spp. were excluded from farmer territories. In contrast, foragers did not suppress fleshy macroalgae, but rather, appeared to decelerate succession and promote a relatively diverse assemblage. In contrast to forager-dominated reefs, farmer territories did not appear to function solely as forager exclusion areas or promote algal diversity as a result of intermediate grazing pressure. The relatively strong effects of farmers observed here may represent a future scenario for coral reefs that are increasingly subject to overfishing of large grazing fishes.     

Observed and projected functional reorganization of riverine fish assemblages from global change

Climate and land-use/land-cover change (“global change”) are restructuring biodiversity, globally. Broadly, environmental conditions are expected to become warmer, potentially drier (particularly in arid regions), and more anthropogenically developed in the future, with spatiotemporally complex effects on ecological communities. We used functional traits to inform Chesapeake Bay Watershed fish responses to future climate and land-use scenarios (2030, 2060, and 2090). We modeled the future habitat suitability of focal species representative of key trait axes (substrate, flow, temperature, reproduction, and trophic) and used functional and phylogenetic metrics to assess variable assemblage responses across physiographic regions and habitat sizes (headwaters through large rivers). Our focal species analysis projected future habitat suitability gains for carnivorous species with preferences for warm water, pool habitats, and fine or vegetated substrates. At the assemblage level, models projected decreasing habitat suitability for cold-water, rheophilic, and lithophilic individuals but increasing suitability for carnivores in the future across all regions. Projected responses of functional and phylogenetic diversity and redundancy differed among regions. Lowland regions were projected to become less functionally and phylogenetically diverse and more redundant while upland regions (and smaller habitat sizes) were projected to become more diverse and less redundant. Next, we assessed how these model-projected assemblage changes 2005–2030 related to observed time-series trends (1999–2016). Halfway through the initial projecting period (2005–2030), we found observed trends broadly followed modeled patterns of increasing proportions of carnivorous and lithophilic individuals in lowland regions but showed opposing patterns for functional and phylogenetic metrics. Leveraging observed and predicted analyses simultaneously helps elucidate the instances and causes of discrepancies between model predictions and ongoing observed changes. Collectively, results highlight the complexity of global change impacts across broad landscapes that likely relate to differences in assemblages' intrinsic sensitivities and external exposure to stressors.     

河流鱼类分类群和功能群的纵向梯度格局——以新安江流域为例

确定鱼类群落的空间格局是保护和管理河流鱼类多样性的基础。尽管河流鱼类分类群（基于物种组成）的纵向梯度格局已得到大量报道,但其功能群（基于功能特征）的空间格局研究较少。以皖南山区新安江为研究流域,沿其"正源-下游"梯度共设置27个调查样点,分别于2017年5月和10月完成2次调查取样,着重研究了鱼类分类群和功能群结构的纵向梯度格局及其形成机制。共采集鱼类44种,可分为5个运动功能群和4个营养功能群,构成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均随河流级别显著变化,但两者均无显著的季节变化;根据相似性百分比分析,由1级至3级河流,数量优势物种和功能群的空间变化主要呈嵌套格局,而由3级至5级河流其变化主要呈周转格局。方差分解结果显示,局域栖息地、陆地景观和支流空间位置3类解释变量对分类群和功能群空间变化的解释率分别为33.6%和38.5%,其中,分类群受局域栖息地和支流空间位置变量的显著影响,而功能群受局域栖息地和陆地景观变量的显著影响。研究表明,沿着新安江的"上游-下游"纵向梯度,鱼类分类群和功能群的空间格局基本一致,但两者的形成机制不同：分类群的纵向梯度变化受环境过滤和扩散过程的联合影响,而功能群则主要受环境过滤影响。     

基于土地利用变化的热带植物群落功能性状与土壤质量的关系

为了明确热带天然林转变为橡胶林和槟榔后土壤质量变化,揭示土地利用变化下植物群落功能性状对土壤质量影响。在海南中部山区,以原始林(PF)、次生林(SF)、槟榔(Areca catechu)林(AP)、纯橡胶(Hevea brasiliensis)林(RP)和橡胶益智(Alpinia oxyphylla)林(RAP)为对象,探索天然林退化后土壤性质和质量变化,分析了植物群落功能性状(凋落物量、郁闭度、根长密度、细根密度和比根长)对土壤质量影响。结果表明:1)与原始林相比,其他土地利用类型凋落物量、根长密度、细根密度、土壤总孔隙度、最大持水量、土壤有机碳和总氮显著降低,土壤容重显著增加。人工林土壤碱解氮明显降低,但总磷、总钾和缓效钾明显升高(P0.05)。2)与原始林相比,次生林、槟榔林、纯橡胶林和橡胶益智林土壤质量指数分别降低63.4%、85.8%、81.2%和84.1%,随原始林、次生林和人工林梯度土地利用强度的增加,土壤质量显著降低(P0.05),但人工林间土壤质量无显著差异。3)凋落物量、郁闭度、根长密度和细根密度均与土壤质量指数显著正相关(P0.05),细根密度对土壤质量的直接影响效应最大,凋落物间接影响效应最大。天然林转变为橡胶和槟榔林显著改变土壤性质和质量,群落性状细根密度和凋落物可较好解释土壤质量变化,强化人工林林下植被和凋落物管理有利于土壤质量改善。     

Intercontinental trends in functional and phylogenetic structure of stream fish assemblages

Understanding of community assembly has been improved by phylogenetic and trait‐based approaches, yet there is little consensus regarding the relative importance of alternative mechanisms and few studies have been done at large geographic and phylogenetic scales. Here, we use phylogenetic and trait dispersion approaches to determine the relative contribution of limiting similarity and environmental filtering to community assembly of stream fishes at an intercontinental scale. We sampled stream fishes from five zoogeographic regions. Analysis of traits associated with habitat use, feeding, or both resulted in more occurrences of trait underdispersion than overdispersion regardless of spatial scale or species pool. Our results suggest that environmental filtering and, to a lesser extent, species interactions were important mechanisms of community assembly for fishes inhabiting small, low‐gradient streams in all five regions. However, a large proportion of the trait dispersion values were no different from random. This suggests that stochastic factors or opposing assembly mechanisms also influenced stream fish assemblages and their trait dispersion patterns. Local assemblages tended to have lower functional diversity in microhabitats with high water velocity, shallow water depth, and homogeneous substrates lacking structural complexity, lending support for the stress‐dominance hypothesis. A high prevalence of functional underdispersion coupled with phylogenetic underdispersion could reflect phylogenetic niche conservatism and/or stabilizing selection. These findings imply that environmental filtering of stream fish assemblages is not only deterministic, but also influences assemblage structure in a fairly consistent manner worldwide.     

Species’ thermal ranges predict changes in reef fish community structure during 8 years of extreme temperature variation

Aim

To assess whether observed thermal bounds in species’ latitudinal ranges (i.e., realized thermal niches) can be used to predict patterns of occurrence and abundance changes observed during a marine heatwave, relative to other important life history and functional traits.

Location

Rottnest Island, Western Australia.

Methods

A time series of standardized quantitative surveys of reef fishes spanning 8 years of pronounced ocean temperature change is used to test whether accurate predictions on shifts in species occupancy and abundance are possible using species traits.

Results

Species‐level responses in occurrence and abundance were closely related to the mid‐point of their realized thermal niche, more so than body size, range size or trophic level. Most of the species that disappeared from survey counts during the heatwave were characterized by geographic ranges that did not extend to latitudes with temperatures equivalent to the ocean temperature peak during the heatwave. We thus find support for the hypothesis that current distribution limits are set directly or indirectly by temperature and are highly responsive to ocean temperature variability.

Main conclusions

Our study shows that reef fish community structure can change very quickly when exposed to extreme thermal anomalies, in directions predicted from the realized thermal niche of the species present. Such predictions can thus identify species that will be most responsive to changing ocean climate. Continued warming, coupled with periodic extreme heat events, may lead to the loss of ecosystem services and ecological functions, as mobile species relocate to more hospitable climes, while less mobile species may head towards extinction.

     

Characterising functional trait diversity and trait–environment relationships in fish assemblages of boreal lakes

1. Characterisation of biodiversity is typically based on taxonomic approaches, while much less is known about other related aspects. Functional trait diversity is one such component of biodiversity that has not been addressed rigorously in ecological research until recently. We tested the congruence between taxonomic‐ and trait‐based approaches, and examined how spatial configuration, local abiotic environmental factors and biotic effects interact to influence taxonomic‐ and trait‐based characterisation of freshwater fish assemblages. 2. Fish assemblage data were compiled for 124 lakes in southern Finland. Variance partitioning in both linear regression analyses and redundancy analysis was used to quantify the relative contribution of spatial and environmental variables to taxonomic and functional trait diversity and structure. Additionally, a null model analysis was used to test for the potential effects of interspecific segregation and biotic interactions on the co‐occurrence of species. 3. The species pool was relatively poor. However, trait‐based classification of species indicated that most species belonged to unique functional entities, which suggested low redundancy in species composition. Correlation analysis indicated a very strong relationship between species richness (SR) and the number of unique trait combinations (UTC). Ecoregion‐level heterogeneity in SR and UTC were well represented in a relatively small group of randomly selected lakes (c. 30 lakes). Multiple regressions indicated moderate roles for abiotic environmental variables (i.e. lake surface area, depth, total phosphorous, colour and pH) in determining SR, UTC and the distribution of single trait categories, whereas geographical location was not generally influential. 4. Redundancy analysis revealed similar patterns to those of diversity analyses for taxonomic and associated trait‐based structure, emphasising the effect of abiotic environmental variables and the negligible effect of geographical position. 5. Co‐occurrence analysis indicated significant checkerboard distribution at the whole assemblage level, but interspecific segregation proved to be of relatively minor importance in the constrained analyses, where species pair combinations within trait category groups were evaluated. 6. Our results suggest that taxonomic‐ and trait‐based patterns of boreal lake fish assemblages are strongly interrelated. Environmental filtering through the effects of local abiotic variables seems to have the most prominent role in determining trait‐based assemblage patterns among lakes, which may also be secondarily shaped by biotic interactions. 7. From the applied perspective, it may not necessarily matter whether traditional taxonomic or more novel trait‐based approaches are used in characterising spatial patterns in boreal fish assemblages. However, trait‐based approaches may provide complementary information which cannot be directly revealed by taxonomic data.     

盐城滨海湿地植物群落种间格局与竞争的关系研究

盐城滨海湿地植物群落种间格局与竞争关系发生变化,导致湿地景观结构与格局发生变化。以江苏盐城国家级珍禽自然保护区核心区为典型研究区,利用1992、1997、2000、2011、2017年TM影像作为数据源,结合GIS技术,运用景观生态学方法分析了滨海湿地植物群落种间格局与竞争关系的时空变化特征。结果表明:(1)1992—2017年,植物群落结构变化呈现出芦苇群落和互花米草群落持续扩张的特征,其面积百分比到2017年分别达到37.06%和34.50%;碱蓬群落整体上呈现持续萎缩的特征,到2017年仅占5.09%。(2)1992—2017年,香农多样性指数(SHDI)和边缘密度(ED)值分别先上升至1.482和14.391,再逐渐下降至1.283和11.967,最大斑块指数(LPI)和聚集度指数(AI)先增大再减小,均反映了植物群落竞争关系从单一化走向多元化再到单一化的趋势。(3)1992—2017年,芦苇-碱蓬群落之间竞争的空间位置先向海方向推进,后向陆方面后退;碱蓬-互花米草群落竞争的位置先向海、再向陆方向、再向海方面演进;芦苇-碱蓬-互花米草群落种间竞争的位置由区域的南部向中部演进。研...     

Arthropod community diversity and trophic structure: a comparison between extremes of plant stress

Abstract.  1. Previous studies have shown that plant stress and plant vigour impact the preference and performance of many insect species. Global climate-change scenarios suggest that some regions such as continental interiors may become increasingly subject to severe drought. In combination, these two issues suggest that drought-driven plant stress may impact insect communities on a landscape scale. While there have been many population studies relating plant stress to the life history of individual herbivore species, far less is known about how plant stress affects entire communities.
2. To study the effect of plant stress on arthropod communities, arthropods were sampled from the canopies of pinyon pines ( Pinus edulis ) growing at sites with a history of chronically high environmental stress (e.g. lower water potentials, soil moisture, and reduced growth rates), and those growing under more favourable conditions. Sampling in these environments yielded >59 000 arthropods, representing 287 species from 14 orders and 80 families, and revealed three major community patterns.
3. First, chronic stress significantly altered community composition. Second, trees growing under high stress supported about 1/10th the number of arthropods, and roughly half the species as trees growing under more favourable conditions. Third, of the 33 abundant herbivore species that exhibited a significantly skewed distribution towards either high- or low-stress trees, 73% were skewed with higher numbers on low-stress trees.
4. The pattern of potentially reduced arthropod diversity and abundance on stressed pines observed in this study may further compound the loss of species resulting from the recent, landscape-scale drought-induced mortality of pines in the southwestern USA.     

海岸草本植物细根、叶片功能性状及其与土壤因子的关系

植物根、叶是对环境敏感性最高的器官,探究根叶功能性状之间的相互关系以及对环境因子的响应,有助于揭示植物对资源的利用状况及其对环境的适生策略。为探讨沿海植物的适应策略,该文以平潭岛砂质海岸草本植物为研究对象,由海及陆设置了3个距离梯度,选取6个叶功能性状和5个细根功能性状指标,分析海岸植物叶片与细根功能性状及其对土壤因子的响应。结果表明:(1)根、叶功能性状变异系数幅度在潮间带最小,在距高海潮线30～60 m的梯度上最大。单叶面积、叶磷含量、根平均直径、根组织密度、根磷含量随着由海到陆的距离增加呈上升趋势; 叶干物质含量、叶组织密度、比根长、比根面积呈下降趋势。(2)植物通过性状组合,在生长与防御间进行资源分配的权衡,表现在叶性状间、根性状间以及根-叶性状间具不同程度的相关性。其中,在地上-地下对应性状中,叶厚与根平均直径、叶磷含量与根磷含量呈极显著正相关; 而比叶面积与比根面积、比根长,叶组织密度与根组织密度均未表现出显著的相关性。(3)土壤因子对海岸植物功能性状变化的解释度为52.05%,其中土壤含盐量的影响最大,其次是土壤含水量、电导率、pH值。总体而言,在恶劣的海岸环境下,由海向陆土壤含盐量、电导率、含水量及pH值逐渐下降,整体为低磷高盐碱,植物表现为不同的生存策略:距海近的植物采取“叶片资源保守型、根系资源获取型”策略; 距海远的植物则采取“叶片资源获取型、根系资源保守型”。该研究结果为了解海岸草本植物对环境梯度变化的响应机制和适应性提供了一定参考价值,同时也利于通过分析土壤等环境特性按梯度筛选栽种适宜的物种,促进海岸植物的恢复和保护。