Environmental and historical imprints on beta diversity: insights from variation in rates of species turnover along gradients

A common approach for analysing geographical variation in biodiversity involves using linear models to determine the rate at which species similarity declines with geographical or environmental distance and comparing this rate among regions, taxa or communities. Implicit in this approach are weakly justified assumptions that the rate of species turnover remains constant along gradients and that this rate can therefore serve as a means to compare ecological systems. We use generalized dissimilarity modelling, a novel method that accommodates variation in rates of species turnover along gradients and between different gradients, to compare environmental and spatial controls on the floras of two regions with contrasting evolutionary and climatic histories: southwest Australia and northern Europe. We find stronger signals of climate history in the northern European flora and demonstrate that variation in rates of species turnover is persistent across regions, taxa and different gradients. Such variation may represent an important but often overlooked component of biodiversity that complicates comparisons of distance–decay relationships and underscores the importance of using methods that accommodate the curvilinear relationships expected when modelling beta diversity. Determining how rates of species turnover vary along and between gradients is relevant to understanding the sensitivity of ecological systems to environmental change.     

Warming leads to higher species turnover in a coastal ecosystem

The responses of ecological communities and ecosystems to increased rates of environmental change will be strongly influenced by variation in the diversity of community composition. Yet, our understanding of how diversity is affected by rising temperatures is inconclusive and mainly based on indirect evidence or short‐term experiments. In our study, we analyse the diversity and species turnover of benthic epilithic communities within the thermal flume of a nuclear power plant at the Swedish coast. This flume covers the range of predicted future temperature rises. Species composition was significantly different between control sites and sites with higher temperatures. We found that temperature had little effect on the number of species in three functional groups (macroinvertebrates, benthic diatoms, and macrophytes, which here comprise multicellular algae and macroscopic colonies of unicellular algae and cyanobacteria), neither at single sampling dates nor summed for the entire observation year. However, species turnover significantly increased with increasing temperature for diatoms, macrophytes and invertebrates. Different temperature regimes resulted in significantly different species composition and indicator species. Thus, increasing temperatures in the thermal flume increased temporal beta‐diversity and decreased compositional stability of communities, although observed richness did not change at any point in time. We highlight the need to investigate the consequences of such declines in compositional stability for functional stability of ecosystem processes.     

The relationship between species replacement,dissimilarity derived from nestedness,and nestedness

Aim Beta diversity can be partitioned into two components: dissimilarity due to species replacement and dissimilarity due to nestedness ( Baselga, 2010 , Global Ecology and Biogeography, 19 , 134–143). Several contributions have challenged this approach or proposed alternative frameworks. Here, I review the concepts and methods used in these recent contributions, with the aim of clarifying: (1) the rationale behind the partitioning of beta diversity into species replacement and nestedness‐resultant dissimilarity, (2) how, based on this rationale, numerators and denominators of indices have to match, and (3) how nestedness and nestedness‐resultant dissimilarity are related but different concepts. Innovation The rationale behind measures of species replacement (turnover) dictates that the number of species that are replaced between sites (numerator of the index) has to be relativized with respect to the total number of species that could potentially be replaced (denominator). However, a recently proposed partition of Jaccard dissimilarity fails to do this. In consequence, this partition underestimates the contribution of species replacement and overestimates the contribution of richness differences to total dissimilarity. I show how Jaccard dissimilarity can be partitioned into meaningful turnover and nestedness components, and extend these new indices to multiple‐site situations. Finally the concepts of nestedness and nestedness‐resultant dissimilarity are discussed. Main conclusions Nestedness should be assessed using consistent measures that depend both on paired overlap and matrix filling, e.g. NODF, whereas beta‐diversity patterns should be examined using measures that allow the total dissimilarity to be separated into the components of dissimilarity due to species replacement and dissimilarity due to nestedness. In the case of multiple‐site dissimilarity patterns, averaged pairwise indices should never be used because the mean of the pairwise values is unable to accurately reflect the multiple‐site attributes of dissimilarity.     

The relationships between local and regional species richness and spatial turnover

Aim To determine the empirical relationships between species richness and spatial turnover in species composition across spatial scales. These have remained little explored despite the fact that such relationships are fundamental to understanding spatial diversity patterns. Location South‐east Scotland. Methods Defining local species richness simply as the total number of species at a finer resolution than regional species richness and spatial turnover as turnover in species identity between any two or more areas, we determined the empirical relationships between all three, and the influence of spatial scale upon them, using data on breeding bird distributions. We estimated spatial turnover using a measure independent of species richness gradients, a fundamental feature which has been neglected in theoretical studies. Results Local species richness and spatial turnover exhibited a negative relationship, which became stronger as larger neighbourhood sizes were considered in estimating the latter. Spatial turnover and regional species richness did not show any significant relationship, suggesting that spatial species replacement occurs independently of the size of the regional species pool. Local and regional species richness only showed the expected positive relationship when the size of the local scale was relatively large in relation to the regional scale. Conclusions Explanations for the relationships between spatial turnover and local and regional species richness can be found in the spatial patterns of species commonality, gain and loss between areas.     

神农架植物物种空间周转的驱动因素

揭示区域物种组成随环境梯度的变化规律, 掌握物种多样性的周转过程及其与环境的耦合关系, 是理解物种多样性形成的生态过程的核心, 也是生物多样性保护和保护地网络构建的科学基础。神农架是众多古老、孑遗和特有植物的关键栖息地, 也是全球落叶木本植物最丰富的地区之一。然而, 我们对影响神农架植物物种周转并形成镶嵌格局的驱动因素知之甚少。本研究基于对神农架长期系统的植物群落物种调查数据, 通过广义相异模型(GDM)和偏回归分析等方法, 分析了气候差异、地理距离、人为干扰等因素对神农架植物物种周转的驱动效应。结果表明, 随着年均温(MAT)、≥ 0℃积温、≥ 10℃积温差异和地理距离增大, 植物β多样性显著增加, 而年均降水量(MAP)、距离道路距离及森林破碎化等人为干扰因素对植物β多样性无显著影响。地理距离、气候差异和人为干扰分别解释了物种相异性变异的28.75%、34.56%和12.55%, 其交互作用解释了变异的8.52%。地理距离、气候差异和人为干扰共同解释了物种相异性变异的43.47%。综上, 地理距离和温度分异是驱动神农架植物物种空间周转过程的关键因素。保护海拔梯度上植被的垂直带谱及复杂多样的地貌异质性, 是维持神农架物种周转生态过程的关键, 也是神农架国家公园建设乃至鄂西北保护地网络构建应关注的重点。     

Low species turnover of upland Amazonian birds in the absence of physical barriers

Aim

One of the oldest and most powerful ways for ecologists to explain distinct biological communities is to invoke underlying environmental differences. But in hyper-diverse systems, which often display high species richness and low species abundance, these sorts of community comparisons are especially challenging. The classic view for Amazonian birds posits that riverine barriers and habitat specialization determine local and regional community composition. We test the tacit, complementary assumption that similar bird communities should therefore permeate uniform habitat between major rivers, regardless of distance.

Location

Upland (terra firme) rainforests of central Amazonia.

Methods

We conducted intensive whole-community surveys of birds in three pairs of 100-ha plots, separated by 40–60 km. We then used dissimilarity indices, cluster analysis, and ordination to characterize differences among the six avian communities.

Results

In all, we detected 244 forest-dependent birds, with an average of 190 species (78%) per plot. Species turnover was negligible, no unique indicator species were found among plot pairs, and all documented species were already known from a complete inventory at one of the three sites.

Main Conclusions

Our study corroborates the classic biogeographical pattern and suggests that turnover contributes little to regional avian diversity within upland forests. Using a grain size of 100 ha, this implies that upland birds perceive the environment as uniform, at least over distances of ~60 km. Therefore, to maximize both local species richness and population persistence, our findings support the conservation of very large tracts of upland rainforest. Our analyses also revealed that the avifauna at Reserva Ducke, encroached by urban sprawl from the city of Manaus, shows the hallmarks of a disturbed community, with fewer vulnerable insectivores. This defaunation signals that even an enormous preserve (10 × 10 km) in lowland Amazonia is not insulated from anthropogenic degradation within the surrounding landscape.     

Patterns of species richness and turnover in endemic amphibians of the Guineo-Congolian rain forest

Aim

The African Guineo-Congolian (GC) region is a global biodiversity hotspot with high species endemism, bioclimatic heterogeneity, complex landscape features, and multiple biogeographic barriers. Bioclimatic and geographic variables influence global patterns of species richness and endemism, but their relative importance varies across taxa and regions and is poorly understood for many faunas. Here, we test the hypothesis that turnover in endemic amphibians of the GC biodiversity hotspot is influenced mainly by the geographic distance between grid cells and secondarily by rainfall- and temperature-related variables.

Location

West and Central Africa.

Major Taxa Studied

Amphibians.

Methods

We compiled species-occurrence records via field sampling, online databases, and taxonomic literature. Our study used 1205 unique georeferenced records of 222 amphibian species endemic to the GC region. Patterns of species richness were mapped onto a grid with a spatial resolution of 0.5° × 0.5°. We estimated weighted endemism and tested whether endemism was higher than the expected species richness (randomization test). We quantified species turnover using generalized dissimilarity modelling to evaluate the processes underlying observed patterns of species richness in GC endemic amphibians. We explored bioregionalization using agglomerative hierarchical clustering based on the unweighted pair group method with arithmetic averages.

Results

We identified seven areas within the lower GC region – forests in Cameroon, Gabon, Southern Nigeria, Equatorial Guinea, Republic of Congo, Democratic Republic of Congo, and Cote d'Ivoire – as having high species richness of endemic amphibians. The randomization test returned four major areas of significant weighted endemism: Nigeria-Cameroon mountains, forest regions of the Democratic Republic of Congo, Cote d'Ivoire, and Ghana. Our analysis revealed five bioregions for amphibian endemism, four of which were located within the lower Guineo-Congolian forest. Species turnover was strongly related to the geographic distance between grid cells; contributing bioclimatic variables included precipitation of the warmest quarter, mean temperature of the wettest quarter, and mean diurnal temperature range.

Main Conclusions

Our results indicate that geographic distance between grid cells is the primary determinant of turnover in GC endemic amphibians, with secondary but significant effects of rainfall- and temperature-related variables. Our study identifies key areas of endemic amphibian richness that could be prioritized for conservation actions.     

Disentangling the effects of geographic distance and environmental dissimilarity on global patterns of species turnover

Aim To distinguish the effects of geographic distance and environmental dissimilarity on global patterns of species turnover in four classes of terrestrial vertebrates (mammals, birds, reptiles and amphibians). Location Six hundred and sixty terrestrial ecoregions across the globe. Methods We calculated species turnover between each pair of ecoregions, using the Jaccard index (J). We selected seven variables to quantify environment in each ecoregion, and subjected the environmental values to a principal components analysis. For each realm, we applied multiple regression analysis relating the natural logarithm of the Jaccard index (lnJ) to geographic distance alone and in combination with differences in the environment variables measured as principal components (PC). We used partial correlations to partition variance in lnJ between unique contributions of distance and environmental PC scores, the covariation between distance and environment, and unexplained variance. To examine the latitude and species turnover relationship, we regressed lnJ on latitude with distance between ecoregions being included as a covariate. Results The natural logarithm of the Jaccard index (lnJ) decreased significantly with increasing geographic distance for all vertebrate classes in each zoogeographic realm, and the slopes of the relationships per 1000 km ranged from ?0.251 to ?1.043. With environmental differences included in the analysis, both geographic distance and environmental differences were substantial predictors of lnJ for every combination of taxon and realm. On average, the unique contribution of geographic distance to variation in species turnover between ecoregions was about 1.4 times that of the environmental differences between ecoregions. Species turnover generally decreased with increasing latitude when controlling for geographic distance. The value of lnJ for each vertebrate class was highly and positively correlated with those of the other vertebrate classes. Main conclusions Our analyses suggest that both dispersal‐based and niche‐based processes have played important roles in determining faunal similarities among vertebrate assemblages at the spatial scale examined. Furthermore, reptiles and amphibians exhibited greater distance‐independent faunal heterogeneity among ecoregions and greater turnover among ecoregions with respect to geographic and environmental distance than birds and mammals.     

环境梯度下蒙古栎群落的物种多样性特征

通过样地法研究了东北地区处于不同经度、纬度和海拔的 13个地点蒙古栎群落的物种丰富度、Gini指数、PIE指数、Shannon指数和 Pielou指数 ,利用相关和回归的统计方法分析了不同地点物种的丰富度指数、Simpson多样性指数和 Shannon多样性指数与各地所处的经度、纬度和海拔的关系。结果发现 :不仅不同地点 (较大尺度 )的物种丰富度和多样性指数均有差异 ,即使在相同的地点 (较小尺度 ) ,物种丰富度及多样性指数也有差异 ,有时还具有很大的差异 ,呈现空间异质性分布的特征 ;因为影响这些多样性指数的环境因子更加复杂 ,不仅受经度、纬度和海拔的影响 ,也受地形、群落的年龄、干扰史等多种生态因子影响。不同地点的物种丰富度与海拔和纬度都具有明显的相关性 (p<0 .0 5 ) ,物种丰富度随海拔和纬度的升高而降低 ,依据显著度的大小可以推测物种丰富度与海拔的相关性比与纬度的相关性更密切 ;蒙古栎群落不同类群的植物种的丰富度具有不同的分布格局 ,木本植物的丰富度与当地纬度具有明显的相关性 (p<0 .0 5 ) ,而与所在地的海拔没有显著的关系 (p>0 .0 5 ) ,而草本植物受海拔的影响更显著 (p<0 .0 5 ) ,而与纬度之间没有显著的关系 (p>0 .0 5 )。群落的 Gini指数、PIE指数、Shannon指数和Pielou指数未发现与海     

On the relationship between plant species diversity and genetic diversity of Plantago lanceolata (Plantaginaceae) within and between grassland communities

Aims and Methods The relationship between genetic diversity and species diversity and the underlying mechanisms are of both fundamental and applied interest. We used amplified fragment length polymorphism (AFLP) and vegetation records to investigate the association between genetic diversity of Plantago lanceolata and plant species diversity using 15 grassland communities in central Germany. We used correlation and partial correlation analyses to examine whether relationships between genetic and species diversity were direct or mediated by environmental differences between habitats.Important findings Both within- and between-population genetic diversity of P. lanceolata were significantly positively correlated with plant species diversity within and between sites. Simple and partial correlations revealed that the positive correlations indirectly resulted from the effects of abiotic habitat characteristics on plant species diversity and, via abundance, on genetic diversity of P. lanceolata. Thus, they did not reflect a direct causal relationship between plant species diversity and genetic diversity of P. lanceolata, as would have been expected based on the hypothesis of a positive relationship between plant species diversity and niche diversity.     

Spatial and environmental determinants of vascular plant species richness distribution in the Iberian Peninsula and Balearic Islands

Using an exhaustive data compilation, Iberian vascular plant species richness in 50 times 50 UTM grid cells was regressed against 24 explanatory variables (spatial, geographical, topographical, geological, climatic, land use and environmental diversity variables) using Generalized Linear Models and partial regression analysis in order to ascertain the relative contribution of primary, heterogeneous and spatially structured variables. The species richness variation accounted for by these variables is reasonably high (65% of total deviance). Little less than half of this variation is accounted for spatially structured variables. A purely spatial component of variation is hardly significant. The most significant variables are those related to altitude, and particularly maximum altitude, whose cubic response reflects the occurrence of the maximum number of species at the highest altitudes. This result highlighted the importance of Iberian mountains as hotspots of diversity and the relevance of large and small scale historical factors in contemporary plant distribution patterns. Climatic or energy-related variables contributed little, whereas geological (calcareous and acid rocks) and, to a lesser extent, environmental heterogeneity variables (land use diversity and altitude range) seem to be more important.     

Reproductive traits associated with species turnover of amphibians in Amazonia and its Andean slopes

Assembly of ecological communities is important for the conservation of ecosystems, predicting perturbation impacts, and understanding the origin and loss of biodiversity. We tested how amphibian communities are assembled by neutral and niche‐based mechanisms, such as habitat filtering. Species richness, β‐diversities, and reproductive traits of amphibians were evaluated at local scale in seven habitats at different elevation and disturbance levels in Wisui Biological Station, Morona‐Santiago, Ecuador, on the foothills of the Cordillera del Kutukú; and at regional scale using 109 localities across evergreen forests of Amazonia and its Andean slopes (0–3,900 m a.s.l.). At local scale, species composition showed strong differences among habitats, explained mainly by turnover. Reproductive modes occurred differently across habitats (e.g., prevalence of direct developers at high elevation, where breeding in ground level water disappears). At regional scale, elevation was the most important factor explaining the changes in species richness, reproductive trait occurrences, and biotic dissimilarities. Species number in all groups decreased with elevation except for those with lotic tadpoles and terrestrial reproduction stages. Seasonality, annual precipitation, and relative humidity partially explained the occurrence of some reproductive traits. Biotic dissimilarities were also mostly caused by turnover rather than nestedness and were particularly high in montane and foothill sites. Within lowlands, geographic distance explained more variability than elevation. Habitat filtering was supported by the different occurrence of reproductive traits according to elevation, water availability, and breeding microhabitats at both scales, as well as other assembly mechanisms based in biotic interactions at local scale. Human‐generated land use changes in Amazonia and its Andean slopes reduce local amphibian biodiversity by alteration of primary forests and loss of their microhabitats and the interaction network that maintains their unique amphibian assemblages with different reproductive strategies.     

云南横断山区蚤类物种多样性的地理分布趋势与重要环境因素的关系

根据云南横断山区具代表性的7个山系共9个样区山地蚤类及其宿主动物的垂直分布调查结果,并参考以往的部分调查资料,对所获数据进行统计学处理,应用不同山系样区各垂直带蚤类物种多样性的平均值作为指标,就云南横断山区蚤类物种多样性的地理分布趋势和规律,与宿主动物和重要环境因素的关系等问题进行了分析和探讨。结果显示,蚤类物种多样性的地理分布趋势是由西北→南→东→北呈递减的趋势。与变化趋势密切相关的重要因素是水湿条件,其次是经度的影响。另外还与蚤类的丰富度、地理环境、垂直带和人们的生产活动等因素有关。研究认为,形成蚤类与宿主动物间多样性地理分布趋势上的差异的主要原因是对环境的适应性不同,其中水湿条件就是很重要的因素之一。研究还发现,横断山蚤类在我国蚤类区系中是多样性最高的地区,它充分地反映了横断山区栖息地异质性对物种数的重要影响,并说明栖息地异质性对物种—面积关系起着重要作用。     

羌塘高寒草地物种多样性与生态系统多功能关系格局

传统的生物多样性-生态系统功能研究大多侧重于单一生态系统功能与物种多样性的关系,忽略了生态系统的重要价值在于其能够同时提供多种功能或服务,即生态系统的多功能性。基于藏北羌塘高寒草地样带调查数据,选取植被地上生物量、地下生物量、土壤全氮、硝态氮及铵态氮含量、土壤全磷含量、土壤有机碳储量等7个与植物生长、养分循环、土壤有机碳蓄积相关的参数来表征生态系统多功能性。采用上述参数转换为Z分数后的平均值计算多功能性指数(M)。分析了不同生物多样性指数与生态系统多功能指数的关系以及年降水量和年均温度对物种多样性和生态系统多功能性指数的影响。结果表明,物种丰富度指数与生态系统多功能性之间呈极显著的正相关关系,Shannon-wiener和Simpson物种多样性指数也与多功能性指数间呈显著的正相关,但多功能性指数与Pielou均匀度指数没有表现出明显的相关关系。物种丰富度与表征植物生长、养分循环以及土壤有机碳蓄积的生态系统功能指数间也均呈极显著的正相关关系。降水格局显著影响羌塘高原物种丰富度和生态系统多功能指数,二者均随年降雨量的增加而显著增加,但物种多样性指数并未与年降水量呈现显著相关关系。研究强调了群落物种丰富度即群落物种数量对维持生态系统多功能性的重要意义,这意味着由于人类活动导致的物种丧失可能会给藏北高寒草地生态系统多功能和生态服务带来更为严重的后果。就退化草地恢复或草地可持续管理而言,在藏北羌塘地区,本地植物种的物种丰富度恢复和维持应作为重要目标之一。     

High species diversity and turnover in granite inselberg floras highlight the need for a conservation strategy protecting many outcrops

Determining patterns of plant diversity on granite inselbergs is an important task for conservation biogeography due to mounting threats. However, beyond the tropics there are relatively few quantitative studies of floristic diversity, or consideration of these patterns and their environmental, biogeographic, and historical correlates for conservation. We sought to contribute broader understanding of global patterns of species diversity on granite inselbergs and inform biodiversity conservation in the globally significant Southwest Australian Floristic Region (SWAFR). We surveyed floristics from 16 inselbergs (478 plots) across the climate gradient of the SWAFR stratified into three major habitats on each outcrop. We recorded 1,060 species from 92 families. At the plot level, local soil and topographic variables affecting aridity were correlated with species richness in herbaceous (HO) and woody vegetation (WO) of soil‐filled depressions, but not in woody vegetation on deeper soils at the base of outcrops (WOB). At the outcrop level, bioclimatic variables affecting aridity were correlated with species richness in two habitats (WO and WOB) but, contrary to predictions from island biogeography, were not correlated with inselberg area and isolation in any of the three habitats. Species turnover in each of the three habitats was also influenced by aridity, being correlated with bioclimatic variables and with interplot geographic distance, and for HO and WO habitats with local site variables. At the outcrop level, species replacement was the dominant component of species turnover in each of the three habitats, consistent with expectations for long‐term stable landscapes. Our results therefore highlight high species diversity and turnover associated with granite outcrop flora. Hence, effective conservation strategies will need to focus on protecting multiple inselbergs across the entire climate gradient of the region.