Species richness and diversity in different functional groups across environmental stress gradients: a model for marine rocky shores

We present a model predicting how the species richness and diversity within benthic functional groups should vary across the full environmental stress gradient across which a regional biota from marine rocky shores can occur. Built upon previous models, our model makes predictions for sessile species (macroalgae and filter feeders), herbivores, and carnivores. We tested some of its predictions by surveying vertical (intertidal elevation) and horizontal (wave exposure and ice scour) stress gradients in northern Nova Scotia, Canada. Because of harsh winter conditions, these coasts only depict approximately intermediate‐to‐high yearly levels of stress that the cold‐temperate, rocky intertidal biota from the northwestern Atlantic can experience. The observed trends matched predictions for sessile species in 75% of the studied gradients, and showed a moderate agreement for herbivores and carnivores only when they were combined as mobile consumers. Agreement meant that both richness and diversity increased from the most stressful to the most benign habitats that can be found in northern Nova Scotia. Also as predicted, sessile species generally showed a faster rate of increase in richness than mobile consumers. Our model also predicted a higher overall richness for sessile species than for mobile consumers, which was true by a factor of 3. Therefore, our model may constitute a useful tool to understanding community composition as a function of abiotic stress, which may in turn facilitate studies on community functioning. Model predictions for lower stress ranges could be tested on more southern shores where the same regional biota occurs.     

An environmental stress model correctly predicts unimodal trends in overall species richness and diversity along intertidal elevation gradients

Environmental stress is a major factor structuring communities. An environmental stress model (ESM) predicts that overall species richness and diversity should follow a unimodal trend along the full stress gradient along which assemblages from a regional biota can occur (not to be confused with the intermediate disturbance hypothesis, which makes predictions only for basal species along an intermediate-to-high stress range). Past studies could only provide partial support for ESM predictions because of the limited stress range surveyed or a low sampling resolution. In this study, we measured overall species richness and diversity (considering all seaweeds and invertebrates) along the intertidal elevation gradient on two wave-sheltered rocky shores from Helgoland Island, on the NE Atlantic coast. In intertidal habitats, tides cause a pronounced gradient of increasing stress from low to high elevations. We surveyed up to nine contiguous elevation zones between the lowest intertidal elevation (low stress) and the high intertidal boundary (high stress). Nonlinear regression analyses revealed that overall species richness and diversity followed unimodal trends across elevations on the two studied shores. Therefore, our study suggests that the ESM might constitute a useful tool to predict local richness and diversity as a function of environmental stress. Performing tests on other systems (marine as well as terrestrial) should help to refine the model.     

Species richness along multiple gradients: testing a general multivariate model in oak savannas

Behavioral responses of prey to their predators can critically alter community dynamics. Whether or not a prey responds, clearly depends on the effectiveness of that response. The effectiveness on the other hand is predicted to depend on predator behavior. Actively searching predators can render the behavioral responses in their prey ineffective. Nevertheless, most studies investigating the optimal reaction of prey treated predators as immobile elements of the environment. I experimentally manipulated activity of poolfrog ( Rana lessonae ) tadpoles by keeping them at low and high food levels, and exposed them to three species of invertebrate predators ( Aeshna cyanea, Anax imperator , and Dytiscus marginalis ), whose activity also was manipulated through different food levels. Satiated, less active predators were more likely to kill hungry, more active tadpoles, but hungry predators killed hungry and satiated tadpoles about equally often. This result suggests that reducing their activity is a more effective strategy for tadpoles if the predators themselves are less active. On the other hand, against hungry, highly motivated predators, the behavioral avoidance strategies were essentially ineffective. Antipredator behavior is generally thought to stabilize the dynamics of predator–prey systems. The results presented here, however, suggest that the community dynamical consequences of antipredator behavior also critically depend on decisions made by predators.     

Contrasting patterns of lichen functional diversity and species richness across an elevation gradient

Major environmental gradients co‐vary with elevation and have been a longstanding natural tool allowing ecologists to study global diversity patterns at smaller scales, and to make predictions about the consequences of climate change. These analyses have traditionally studied taxonomic diversity, but new functional diversity approaches may provide a deeper understanding of the ecological mechanisms driving species assembly. We examined lichen taxonomic and functional diversity patterns on 195 plots (200 m²) together with forest structure along an elevational gradient of 1000 m in a temperate low mountain range (Bohemian Forest, Germany). Along this elevation gradient temperature decreased and precipitation increased, two macroclimatic variables critical for lichens. Elevation was more important than forest structure in driving taxonomic and functional diversity. While species richness increased with elevation, functional diversity decreased and revealed that community patterns shift with elevation from random to clustered, reflecting selection for key shared traits. Higher elevations favored species with a complex growth form (which takes advantage of high moisture) and asexual reproductive mode (facilitating establishment under low temperature conditions). Our analysis highlights the need to examine alternative forms of diversity and opens the avenue for community predictions about climate change. For a regional scenario with increasing temperature and decreasing availability of moisture, we expect a loss of specialized species with a complex growth form and those with vegetative organs at higher elevations in low mountain ranges in Europe.     

渔山岛岩石相潮间带大型底栖动物物种多样性

为系统研究渔山岛潮间带大型底栖动物群落结构与生物多样性,作者于2009年3月至2010年1月在该岛潮间带布设5条断面进行了4个季节的取样,调查大型底栖动物种类组成、生物量和丰度,并分析了多样性状况.结果显示:共鉴定出大型底栖动物100种;Shannon-Wiener多样性指数(H')、Pielou均匀度指数(J)、Ma...     

Trends in abundance of rocky intertidal seaweeds and filter feeders across gradients of elevation,wave exposure,and ice scour in eastern Canada

On NW Atlantic rocky shores, the main basal organisms in intertidal communities are seaweeds (Ascophyllum nodosum, Fucus spp. and Chondrus crispus) and filter feeders (barnacles, Semibalanus balanoides, and mussels, Mytilus spp.). Their ecology has been extensively studied in New England (United States), but knowledge gaps exist for northern shores, which are subjected to stronger environmental stress. Therefore, we studied the above organisms on Canadian shores. We quantified the summer abundance of these seaweeds and filter feeders across full vertical (intertidal elevation) and horizontal (wave exposure and winter ice scour) environmental gradients on the Gulf of St. Lawrence and open Atlantic coasts of Nova Scotia. At the regional scale along the open Atlantic coast, seaweeds showed similar abundances in Nova Scotia than values reported for New England. However, both filter feeders were considerably less abundant in Nova Scotia. At the local scale in Nova Scotia, intense winter ice scour (which only occurs on the Gulf of St. Lawrence coast) was associated with a very low abundance of all species except barnacles. Spatial trends in Nova Scotia were similar to patterns known for certain species elsewhere, such as A. nodosum being almost restricted to sheltered habitats, regardless of elevation, and C. crispus being almost restricted to low elevations, regardless of exposure. Other trends were, however, characteristic of Nova Scotia, such as C. crispus being frequent at low elevations in exposed habitats, unlike in New England, where mussels often predominate there because of competitive advantages. In Nova Scotia, mussels were always restricted to cracks and crevices, unlike in New England, where they form extensive intertidal beds on exposed shores. The direct effects of increased environmental stress and indirect effects through altered interspecific interactions might explain the regional differences in local species distribution, which will require experimental validation. Handling editor: K. Martens     

Physiological community ecology: variation in metabolic activity of ecologically important rocky intertidal invertebrates along environmental gradients

Rocky intertidal invertebrates live in heterogeneous habitatscharacterized by steep gradients in wave activity, tidal flux,temperature, food quality and food availability. These environmentalfactors impact metabolic activity via changes in energy inputand stress-induced alteration of energetic demands. For keystonespecies, small environmentally induced shifts in metabolic activitymay lead to disproportionately large impacts on community structurevia changes in growth or survival of these key species. Herewe use biochemical indicators to assess how natural differencesin wave exposure, temperature and food availability may affectmetabolic activity of mussels, barnacles, whelks and sea starsliving at rocky intertidal sites with different physical andoceanographic characteristics. We show that oxygen consumptionrate is correlated with the activity of key metabolic enzymes(e.g., citrate synthase and malate dehydrogenase) for some intertidalspecies, and concentrations of these enzymes in certain tissuesare lower for starved individuals than for those that are wellfed. We also show that the ratio of RNA to DNA (an index ofprotein synthetic capacity) is highly variable in nature andcorrelates with short-term changes in food availability. Wealso observed striking patterns in enzyme activity and RNA/DNAin nature, which are related to differences in rocky intertidalcommunity structure. Differences among species and habitatsare most pronounced in summer and are linked to high nearshoreproductivity at sites favored by suspension feeders and to exposureto stressful low-tide air temperatures in areas of low wavesplash. These studies illustrate the great promise of usingbiochemical indicators to test ecological models, which predictchanges in community structure along environmental gradients.Our results also suggest that biochemical indices must be carefullyvalidated with laboratory studies, so that the indicator selectedis likely to respond to the environmental variables of interest.     

Species richness, abundance, rarity and environmental gradients in coastal barren vegetation

Coastal barrens in Nova Scotia are heathlands characterised by short, predominantly ericaceous vegetation, sparse tree cover, exposed bedrock, pockets of Sphagnum bog, and stressful climatic conditions. Although coastal barrens are prominent in the physical and cultural landscape, they are largely unprotected. We selected six barrens along the Atlantic coast, and surveyed 20 1-m² plots at each barren for vascular plants, macrolichens, mosses and environmental factors. We recorded 173 species (105 vascular, 41 macrolichen, 27 moss), including six provincially rare vascular species found predominantly in nearshore areas with high levels of substrate salt and nutrients, variable substrate depth, and short vegetation. Although vascular plant and moss richness were similarly correlated with vegetation height, substrate depth, organic matter content, and rock exposure, there were no clear correlations between vascular plant, macrolichen and moss richness across all sites. Vascular plant rarity and species richness were not correlated, but had inverse relationships with key environmental gradients. Tailoring conservation efforts to protect areas of high richness may thus mean that rare species are missed, and vice versa. Ordination and ANOSIM show that barrens vegetation differs widely among sites; therefore, protecting any singular coastal barren will not protect the entire range of vegetation communities and species in this heathland type. Conservation planning should emphasize protecting environmental gradients correlated with richness, rarity and plant community structure, including substrate depth and moisture, and vegetation height. Additionally, protected areas should include a coastal-inland gradient and a diversity of substrate types, including exposed rock and trees.     

Species richness and taxonomic distinctness of lake macrophytes along environmental gradients in two continents

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Nematode distributions as spatial null models for macroinvertebrate species richness across environmental gradients: A case from mountain lakes

Nematode species are widely tolerant of environmental conditions and disperse passively. Therefore, the species richness distribution in this group might largely depend on the topological distribution of the habitats and main aerial and aquatic dispersal pathways connecting them. If so, the nematode species richness distributions may serve as null models for evaluating that of other groups more affected by environmental gradients. We investigated this hypothesis in lakes across an altitudinal gradient in the Pyrenees. We compared the altitudinal distribution, environmental tolerance, and species richness, of nematodes with that of three other invertebrate groups collected during the same sampling: oligochaetes, chironomids, and nonchironomid insects. We tested the altitudinal bias in distributions with t‐tests and the significance of narrow‐ranging altitudinal distributions with randomizations. We compared results between groups with Fisher's exact tests. We then explored the influence of environmental factors on species assemblages in all groups with redundancy analysis (RDA), using 28 environmental variables. And, finally, we analyzed species richness patterns across altitude with simple linear and quadratic regressions. Nematode species were rarely biased from random distributions (5% of species) in contrast with other groups (35%, 47%, and 50%, respectively). The altitudinal bias most often shifted toward low altitudes (85% of biased species). Nematodes showed a lower portion of narrow‐ranging species than any other group, and differed significantly from nonchironomid insects (10% and 43%, respectively). Environmental variables barely explained nematode assemblages (RDA adjusted R² = 0.02), in contrast with other groups (0.13, 0.19 and 0.24). Despite these substantial differences in the response to environmental factors, species richness across altitude was unimodal, peaking at mid elevations, in all groups. This similarity indicates that the spatial distribution of lakes across altitude is a primary driver of invertebrate richness. Provided that nematodes are ubiquitous, their distribution offers potential null models to investigate species richness across environmental gradients in other ecosystem types and biogeographic regions.     

The structure of Mediterranean rocky reef ecosystems across environmental and human gradients, and conservation implications

Historical exploitation of the Mediterranean Sea and the absence of rigorous baselines makes it difficult to evaluate the current health of the marine ecosystems and the efficacy of conservation actions at the ecosystem level. Here we establish the first current baseline and gradient of ecosystem structure of nearshore rocky reefs at the Mediterranean scale. We conducted underwater surveys in 14 marine protected areas and 18 open access sites across the Mediterranean, and across a 31-fold range of fish biomass (from 3.8 to 118 g m(-2)). Our data showed remarkable variation in the structure of rocky reef ecosystems. Multivariate analysis showed three alternative community states: (1) large fish biomass and reefs dominated by non-canopy algae, (2) lower fish biomass but abundant native algal canopies and suspension feeders, and (3) low fish biomass and extensive barrens, with areas covered by turf algae. Our results suggest that the healthiest shallow rocky reef ecosystems in the Mediterranean have both large fish and algal biomass. Protection level and primary production were the only variables significantly correlated to community biomass structure. Fish biomass was significantly larger in well-enforced no-take marine reserves, but there were no significant differences between multi-use marine protected areas (which allow some fishing) and open access areas at the regional scale. The gradients reported here represent a trajectory of degradation that can be used to assess the health of any similar habitat in the Mediterranean, and to evaluate the efficacy of marine protected areas.     

Ferry wakes increase seaweed richness and abundance in a sheltered rocky intertidal habitat

Because hydrodynamic regimes influence community assemblages, commercial ferry traffic can directly affect neighboring marine ecosystems by altering water movement. One of the largest ferry fleets in the world traverses the calm, protected waters of the Canadian Gulf Islands (British Columbia). To address the effects of ferry wakes on rocky marine intertidal communities, we examined community assemblages in sites impacted by ferry wakes (N?=?5) relative to geographically similar control sites not directly exposed to ferry wakes (N?=?6). Intertidal communities were significantly different between wake-influenced and control sites. Further analyses revealed that community level differences resulted from differences in seaweed assemblages, while invertebrate assemblages were similar. Sites exposed to ferry traffic displayed significantly greater overall seaweed abundance and seaweed species richness. Nitrate and nitrite concentrations, salinity, fetch, and tidal zonation were not significantly different between wake-impacted and control sites. However, dissolution blocks revealed that wake-impacted sites experienced increased overall water movement. Furthermore, block dissolution was negatively associated with distance from nearest ferry route and not fetch. Although dissolution block cannot disentangle effects of increased flow versus waves resulting from ferry wakes, we conclude that increased overall water movement from frequent and proximate ferry traffic stimulates primary production in rocky intertidal marine seaweeds by ameliorating mass transfer limitation.     

欧亚大陆东部毛茛科植物多样性格局及主导因子

毛茛科是真双子叶植物的基部类群之一, 包含多种药用植物, 具有较高的保护价值, 但关于毛茛科物种多样性和谱系多样性大尺度格局及其影响因子的研究还比较匮乏, 特别是以较高分辨率分布数据为基础的物种多样性格局研究尚未见报道。本文旨在: (1)建立欧亚大陆东部毛茛科植物分布数据库, 估算不同生活型物种多样性和谱系多样性格局, 并探究格局的形成机制。(2)分析毛茛科物种多样性和谱系多样性的相关关系, 确定多样性热点地区, 为毛茛科保护规划提供依据。根据中国、哈萨克斯坦、吉尔吉斯斯坦、塔吉克斯坦、土库曼斯坦、乌兹别克斯坦、蒙古和俄罗斯等国家的区域和地方植物志, 建立了“欧亚大陆东部地区毛茛科物种分布数据库”。该数据库包含了欧亚大陆东部地区1,688种毛茛科物种的分布数据, 空间分辨率为100 km × 100 km。在此基础上, 估算了毛茛科全部及不同生活型的物种多样性和谱系多样性格局, 并利用广义线性模型和等级方差分离方法分析了毛茛科物种和谱系多样性格局与环境因子的关系。最后比较了物种多样性和谱系多样性的相关关系, 确定了毛茛科的古热点地区。结果显示: (1)欧亚大陆东部毛茛科植物物种和谱系多样性均呈明显的纬度格局, 且在山区具有较高的多样性。(2)毛茛科植物物种和谱系多样性受现代气候、地形异质性和末次冰期以来的气候变化的共同影响, 但不同影响因子的相对贡献率在物种和谱系多样性及不同生活型之间差异显著。(3)中高纬度地区的谱系多样性高于给定物种数的预期, 是毛茛科的古热点地区, 在毛茛科保护规划中应受到重视。     

Productivity and species richness across an environmental gradient in a fire-dependent ecosystem

The fire-dependent longleaf pine-wiregrass (Pinus palustris Mill.-Aristida beyrichiana Trin. & Rupr.) savannas of the southeastern United States provide a unique opportunity to examine the relationship between productivity and species richness in a natural ecosystem because of the extremely high number of species and their range across a wide ecological amplitude (sandhills to edges of wetlands). We used a natural gradient to examine how plant species richness and plant community structure vary with standing crop biomass (which in this system is proportional to annual net productivity) as a function of soil moisture and nitrogen mineralization rates in a frequently burned longleaf pine-wiregrass savanna. Highest ground cover biomass and highest species richness were found at the same position along the gradient, the wet-mesic sites. Relative differences in species richness among site types were independent of scale, ranging from 0.01 m(2) to 100 m(2). Nitrogen availability was negatively correlated with species richness. Dominance of wiregrass (in terms of biomass) was consistent across the gradient and not correlated with species richness. Regardless of site type, the community structure of the savannas was characterized by many perennial species with infrequent occurrences, a factor in the low temporal heterogeneity (percent similarity between seasons and years) and high within-site spatial heterogeneity (percent dissimilarity of vegetation composition). The coexistence of numerous species is likely due to the high frequency of fire that removes competing hardwood vegetation and litter and to the suite of fire-adapted perennial species that, once established, are able to persist. Our results suggest that soil moisture is an important factor regulating both the number of species present and community production within the defined gradient of this study.     

天津湿地蛾类丰富度和多样性及其环境评价

天津海拔较低,在华北属湿地资源比较丰富的地区之一,是亚太地区鸟类迁徙路线的一个重要环节,也是白枕鹤、丹顶鹤等濒危鸟类迁徙过程中的一个重要的停歇地.为了研究清楚天津湿地昆虫的种类及其动态规律,为湿地的研究和保护利用服务,选取了鸭淀、团泊洼、北大港水库、七里海和独流减河大港段为样地以诱集夜间活动的昆虫为主,对天津湿地蛾类的群落生态学进行了研究,取得如下进展:(1)首次全面系统地研究了天津湿地蛾类的多样性,确定天津湿地有蛾类22科186属259种.其中以夜蛾科最多(58属85种),草螟科次之(39属60种),卷蛾科(16属28种)、螟蛾科(20属23种)、尺蛾科(14属15种)的种类也较多,生物多样性丰富.(2)所诱集的蛾类种类及个体数,以微软的Excel列表、计算及作图,进行蛾类群落的种-多度曲线、多样性指数、均匀度与多样性指数的关系以及蛾类群落的聚类(群落系数、欧氏距离)和排序(极点排序、主分量分析)等研究分析,结果表明:鸭淀和七里海的种-多度曲线表现为对数正态分布模型,而团泊洼、北大港水库和独流减河大港段的种-多度曲线多变,但总的趋势比较接近于对数级数模型,即生态位优先占领假说.Shannon-Wiener多样性指数(H′)、种间相遇机率(PIE)、物种丰富度(lnS)均为七里海＞团泊洼＞鸭淀＞北大港水库＞独流减河大港段,而均匀度(E)则为七里海＞鸭淀＞团泊洼＞北大港水库＞独流减河大港段.群落的聚类和排序表明蛾类群落的相似性依次为"七里海-团泊洼-鸭淀-北大港水库-独流减河大港段".通过研究分析得出七里海与鸭淀的湿地环境质量较好,但有退化的趋势.团泊洼、北大港水库、独流减河大港段的湿地环境不稳定,退化严重.并分析了造成这些结果的原因,提出了合理的保护和利用的建议.     

Contrasting response of native and alien plant species richness to environmental energy and human impact along alpine elevation gradients

Aim We tested whether the species–energy and species–human relationships vary between native and both naturalized and casual alien species richness when other environmental variables had been taken into account. Location Trento Province, a region (c. 6200 km²) on the southern border of the European Alps (Italy), subdivided into 156 contiguous (c. 37.5 km²) cells and ranging in elevation from 66 to 3769 m. Methods Data were separated into three subsets, representing richness of natives, naturalized aliens and casual aliens and separately related to temperature, human population and various environmental correlates of plant species diversity. We applied ordinary least squares and simultaneous autoregressive regressions to identify potential contrasting responses of the three plant status subsets and hierarchical partitioning to evaluate the relative importance of the predictor variables. Results Variation in alien plant species richness along the region was almost entirely explained by temperature and human population density. The relationships were positive but strongly curvilinear. Native species richness was less strongly related to either factor but was positively related to the presence of calcareous bedrock. Native species richness had a decelerating positive relationship with temperature (R²= 55%), whereas naturalized and casual aliens had a positive accelerating relationship explaining 86% and 62% of the variation in richness, respectively. Native species richness had a positive decelerating relationship with population density (R²= 42%), whilst both alien subsets had a positive accelerating relationship. Main conclusions Alien species richness was higher in areas with the most rich and diverse assemblages of native species. Areas at high altitudes are not especially prone to alien invasion due to energy constraints, low propagule pressure and disturbance, even considering a potential increased in temperature. Thus, if we consider future environmental change, we should expect a stronger response of aliens than natives in the currently warm, urbanized, low‐altitude areas than in cold, high‐altitude areas where human population density is low.