Environmental determinants of amphibian and reptile species richness in China

Understanding the factors that regulate geographical variation in species richness has been one of the fundamental questions in ecology for decades, but our knowledge of the cause of geographical variation in species richness remains poor. This is particularly true for herpetofaunas (including amphibians and reptiles). Here, using correlation and regression analyses, we examine the relationship of herpetofaunal species richness in 245 localities across China with 30 environmental factors, which include nearly all major environmental factors that are considered to explain broad-scale species richness gradients in such theories as ambient energy, water–energy dynamics, productivity, habitat heterogeneity, and climatic stability. We found that the species richness of amphibians and reptiles is moderately to strongly correlated with most of the environmental variables examined, and that the best fit models, which include explanatory variables of temperature, precipitation, net primary productivity, minimum elevation, and range in elevation, explain ca 70% the variance in species richness for both amphibians and reptiles after accounting for sample area. Although water and temperature are important explanatory variables to both amphibians and reptiles, water variables explain more variance in amphibian species richness than in reptile species richness whereas temperature variables explain more variance in reptile species richness than in amphibian species richness, which is consistent with different physiological requirements of the two groups of organisms.     

Species richness estimation and determinants of species detectability in butterfly monitoring programmes

Abstract 1. Species richness is the most widely used biodiversity index, but can be hard to measure. Many species remain undetected, hence raw species counts will often underestimate true species richness. In contrast, capture–recapture methods estimate true species richness and correct for imperfect and varying detectability. 2. Detectability is a crucial quantity that provides the link between a species count and true species richness. For insects, it has hardly ever been estimated, although this is required for the interpretation of species counts. 3. In the Swiss butterfly monitoring programme about 100 transect routes are surveyed seven times a year using a highly standardised protocol. In July 2003, control observers made two additional surveys on 38 transects. Data from these 38 quadrats were analysed to see whether currently available capture–recapture models can provide quadrat‐specific estimates of species richness, and to estimate species detectability in relation to transect, observer, survey, region, and abundance. 4. Species richness over the entire season cannot be estimated using current capture–recapture methods. The species pool was open, preventing use of closed population models, and detectability varied by species, preventing use of current open population models. Assuming a closed species pool during two mid‐season (July) surveys, a Jackknife capture–recapture method was used that accounts for heterogeneity to estimate mean detectability and species richness. 5. In every case, more species were present than were counted. Mean species detectability was 0.61 (SE 0.01) with significant differences between observers (range 0.37–0.83). Species‐specific detection at time t+ 1 was then modelled for those species seen at t for three mid‐season surveys. Detectability averaged 0.50 (range 0.17–0.81) for individual species and 0.65, 0.44, and 0.42 for surveys. Abundant species were detected more easily, although this relationship explained only 5% of variation in species detectability. 6. These are important, although not entirely unexpected, results for species richness estimation of short‐lived animals. Raw counts of species may be misleading species richness indicators unless many surveys are conducted. Monitoring programmes should be calibrated, i.e. the assumption of constant detectability over dimensions of interest needs to be tested. The development of capture–recapture or similar models that can cope with both open populations and heterogeneous species detectability to estimate species richness should be a research priority.     

Multiple environmental determinants of regional species richness and effects of geographic range size

Understanding patterns of species richness at broad geographic extents remains one of the most challenging yet necessary scientific goals of our time. Many hypotheses have been proposed to account for spatial variation in species richness; among them, environmental determinants have played a central role. In this study, we use data on regional bat species richness in the New World to study redundancy and complementarity of three environmental hypotheses: energy, heterogeneity and seasonality. We accomplish this by partitioning variation in species richness among components associated with unique and combined effects of variables from each hypotheses, and by partitioning the overall richness gradient into gradients of species with varying breadths of geographic distribution. These three environmental hypotheses explain most variation in the species richness gradient of all bats, but do not account for all positive spatial autocorrelation at short distances. Although environmental predictors are highly redundant, energy and seasonality explain different and complementary fractions of variation in species richness of all bats. On the other hand, heterogeneity variables contribute little to explain this gradient. However, results change dramatically when richness is estimated for groups of species with different sizes of geographic distribution. First, the amount of variation explained by environment decreases with a decrease in range size; this suggests that richness gradients of small‐ranged species can not be explained as easily as those of broadly distributed species, as has been implied by analyses that do not consider differences in range size among species. Second, the relative contribution of environmental predictors to explained variation also changes with change in range size. Seasonality and energy are good predictors of species with broad distributions, but they loose almost all explanatory power for richness of species with small ranges. In contrast, heterogeneity, which is a relatively poor predictor of richness of species with large ranges, becomes the main predictor of richness gradients of species with restricted distributions. This suggests that range size is a different dimension on which heterogeneity and other environmental characteristics are complementary to each other. Our results suggest that determinants of species richness gradients might be complex, or at least more complex than many studies have previously suggested.     

Fine‐scale determinants of butterfly species richness and composition in a mountain region

Aim Global patterns of species richness are often considered to depend primarily on climate. We aimed to determine how topography and land cover affect species richness and composition at finer scales. Location Sierra de Guadarrama (central Iberian Peninsula). Methods We sampled the butterfly fauna of 180 locations (89 in 2004, 91 in 2005) at 600–2300 m elevation in a region of 10800 km². We recorded environmental variables at 100‐m resolution using GIS, and derived generalized linear models for species density (number of species per unit area) and expected richness (number of species standardized to number of individuals) based on variables of topoclimate (elevation and insolation) or land cover (vegetation type, geology and hydrology), or both (combined). We evaluated the models against independent data from the alternative study year. We also tested for differences in species composition among sites and years using constrained ordination (canonical correspondence analysis), and used variation partitioning analyses to quantify the independent and combined roles of topoclimate and land cover. Results Topoclimatic, land cover and combined models were significantly related to observed species density and expected richness. Topoclimatic and combined models outperformed models based on land cover variables, showing a humped elevational diversity gradient. Both topoclimate and land cover made significant contributions to models of species composition. Main conclusions Topoclimatic factors may dominate species richness patterns in regions with pronounced elevational gradients, as long as large areas of natural habitat remain. In contrast, both topoclimate and land cover may have important effects on species composition. Biodiversity conservation in mountainous regions therefore requires protection and management of natural habitats over a wide range of topoclimatic conditions, which may assist in facilitating range shifts and alleviating declines in species richness related to climate change.     

Spatial patterns and determinants of Moraceae richness in China

Understanding large-scale patterns of biodiversity and their drivers remains central in ecology. Many hypotheses have been proposed, including hydrothermal dynamic hypothesis, tropical niche conservatism hypothesis, Janzen’s hypothesis and a combination model containing energy, water, seasonality and habitat heterogeneity. Yet, their relative contributions to groups with different lifeforms and range sizes remain controversial, which have limited our ability to understand the general mechanisms underlying species richness patterns. Here we evaluated how lifeforms and species range sizes influenced the relative contributions of these three hypotheses to species richness patterns of a tropical family Moraceae. The distribution data of Moraceae species at a spatial resolution of 50km ×50 km and their lifeforms (i.e. shrubs, small trees and large trees) were compiled. The species richness patterns were estimated for the entire family, different life forms and species with different range sizes separately. The effects of environmental variables on species richness were analyzed, and relative contributions of different hypotheses were evaluated across life forms and species range size groups. The species richness patterns were consistent across different species groups and the species richness was the highest in Sichuan, Guangzhou and Hainan provinces, making these provinces the hotspots of this family. Climate seasonality is the primary factor in determining richness variation of Moraceae. The best combination model gave the largest explanatory power for Moraceae species richness across each group of range size and life forms followed by the hydrothermal dynamic hypothesis, Janzen’s hypothesis and tropical niche conservatism hypothesis. All these models has a large shared effects but a low independent effect (< 5%), except rare species. These findings suggest unique patterns and mechanisms underlying rare species richness and provide a theoretical basis for protection of the Moraceae species in China.     

Species richness patterns and the determinants of larch forests in China

Larch forests are important for species diversity, as well as soil and water conservation in mountain regions. In this study, we determined large-scale patterns of species richness in larch forests and identified the factors that drive these patterns. We found that larch forest species richness was high in southern China and low in northern China, and that patterns of species richness along an elevational gradient depend on larch forest type. In addition, we found that patterns of species richness in larch forests are best explained by contemporary climatic factors. Specifically, mean annual temperature and annual potential evapotranspiration were the most important factors for species richness of tree and shrub layers, while mean temperature of the coldest quarter and anomaly of annual precipitation from the Last Glacial Maximum to the present were the most important for that of herb layer and the whole community. Community structural factors, especially stand density, are also associated with the species richness of larch forests. Our findings that species richness in China''s larch forests is mainly affected by energy availability and cold conditions support the ambient energy hypothesis and the freezing tolerance hypothesis.     

中国蚂蚁丰富度地理分布格局及其与环境因子的关系

物种丰富度分布格局及其形成机制的研究对于生物多样性保护具有重要意义。为了了解中国蚂蚁物种丰富度分布格局,利用中国省级尺度蚂蚁物种分布数据和环境信息,结合GIS和数理统计方法,探讨蚂蚁物种丰富度的地理分布格局与环境因子之间的关系。研究结果表明:(1)蚂蚁丰富度随纬度增加呈逐渐递减趋势,但缺乏显著的经度梯度。丰富度最高的地区主要集中在南方省份,我国北方、西北干旱区和青藏高原北部地区丰富度较低;(2)简单线性回归分析表明,能量、水分和季节性因素中,影响蚂蚁物种丰富度最强的因子分别为最冷月均温(TEMmin)(R2adj=0.532)、年均降水量(PREC)(R2adj=0.376)和年温度变化范围(TEMvar)(R2adj=0.539),而单个生境异质性因子对蚂蚁物种丰富度的影响均不显著;(3)最优模型由年均温(TEM)、海拔变化范围(ELErange)和年温度变化范围(TEMvar)组成,能够解释68.4%的蚂蚁丰富度地理分异。鉴于海拔变化范围更多地反映与温度相关的生境异质性,因此温度是限制中国蚂蚁分布的最重要因素。另外,分析结果还表明,海南、贵州、江西、四川、安徽和山西等6省蚂蚁区系调查最不充分,是未来发现蚂蚁新分布的热点地区。     

Water–energy balance and the geographic pattern of species richness of western Palearctic butterflies

Abstract. 1. Using two sources of data to estimate butterfly species richness, the potential influences of 11 environmental variables on the richness gradient of butterflies in western/central Europe and northern Africa were examined with multiple regression and spatial autocorrelation analysis. A measure of water–energy balance, actual evapotranspiration, explained 79% of the variance in butterfly species richness using data derived from range maps, and 72% of the variance using data derived from grid‐based distribution maps. All other variables explained less than 4% of the variance in the regression models and differed depending on the data source. 2. The spatial analysis indicated that actual evapotranspiration successfully removed most of the spatial autocorrelation in both richness data sets at all spatial scales, confirming the ability of the model to account for the spatial pattern in butterfly richness. 3. Plant species richness, a rarely tested variable hypothesised to be an important determinant of herbivore diversity, was weakly associated with butterfly richness, suggesting that it has little or no direct influence on butterfly richness. 4. A historical variable, the length of time that areas have been exposed for recolonisation after the retreat of the ice sheet following the last ice age, was also not associated with richness patterns, indicating that butterfly richness is in equilibrium with contemporary climate. 5. It was not possible to confirm a result reported for Canadian butterflies that land cover diversity is a strong predictor of butterfly richness, possibly because of methodological differences in the studies, differences in the range of climates found in Canada and the western Palearctic, or because of the highly modified landscape characteristic of Europe. 6. Water–energy balance offers a parsimonious explanation for the butterfly richness gradient in this region, operating partially indirectly via effects on plant productivity and partially directly via physiological effects on butterflies, and this conclusion is robust to differences in the types of distribution maps used to estimate richness patterns.     

中国不同地理区域鸟兽物种丰富度的相关性

生物类群之间物种丰富度的相关性研究是当前物种多样性研究中的热点问题之一,目前,中国尚无相关的研究报道。我们收集了中国三种区域类型：动物地理亚区、行政区和保护区的鸟兽名录,分析了行政区与保护区、动物地理区和经纬度带中鸟兽物种数比值及其相关性。 结果表明：不同区域、动物地理区和经纬度带中鸟兽物种数都显著相关。保护区尺度鸟兽物种 数的相关系数为0.818和动物地理区中的华北区为0.768,其他所有区域和地理区域的鸟兽物 种数的相关系数都高于0.850。因此,鸟兽物种数的相关关系在一定程度上具有预测价值。我们发现不同区域鸟兽物种数比值无显著性差异;但是,不同区域间鸟兽物种数 比值差异显著。该比值在中国呈中间低四周高的分布趋势,其中东北地区最高。我们还利用历史累积调查数据与非历史累积调查数据进行了鸟兽物种数比值及其相关性分析,发现利用累积数据计算的相关性低于非累积数据计算的相关性,但利用累积数据计算的鸟兽物 种数比值高于非累积数据计算的比值。最后,探讨了为什么鸟类与兽类的物种数目会相关。我们根据物种－面积公式,S=CA^Z,导出了两个生物类群物种丰富度的相关关 系式。利用全国不同区域数据拟合,得到Z₁/Z₂＝0.913,Z₁/Z₂接近于1。于是 ,C₁/C2可视为近似等于R_am。本研究可推广到其他不同生物类群物种。物种数量的相关关系为快速评估区域的物种多样性提供了一条途径。     

Environmental controls on butterfly occurrence and species richness in Israel: The importance of temperature over rainfall

Butterflies are considered important indicators representing the state of biodiversity and key ecosystem functions, but their use as bioindicators requires a better understanding of how their observed response is linked to environmental factors. Moreover, better understanding how butterfly faunas vary with climate and land cover may be useful to estimate the potential impacts of various drivers, including climate change, botanical succession, grazing, and afforestation. It is particularly important to establish which species of butterflies are sensitive to each environmental driver.The study took place in Israel, including the West Bank and Golan Heights.To develop a robust and systematic approach for identifying how butterfly faunas vary with the environment, we analyzed the occurrence of 73 species and the abundance of 24 species from Israeli Butterfly Monitoring Scheme (BMS‐IL) data. We used regional generalized additive models to quantify butterfly abundance, and generalized linear latent variable models and generalized linear models to quantify the impact of temperature, rainfall, soil type, and habitat on individual species and on the species community.Species richness was higher for cooler transects, and also for hilly and mountainous transects in the Mediterranean region (rendzina and Terra rossa soils) compared with the coastal plain (Hamra soil) and semiarid northern Jordan Vale (loessial sierozem soil). Species occurrence was better explained by temperature (negative correlation) than precipitation, while for abundance the opposite pattern was found. Soil type and habitat were insignificant drivers of occurrence and abundance.Butterfly faunas responded very strongly to temperature, even when accounting for other environmental factors. We expect that some butterfly species will disappear from marginal sites with global warming, and a large proportion will become rarer as the region becomes increasingly arid.     

Synchrony of butterfly populations across species' geographic ranges

Understanding the mechanisms by which global climate change and habitat loss impact upon biodiversity is essential in order to mitigate any negative impacts. One such impact may be changes to population synchrony (defined as correlated fluctuations in the density of separate populations). It is well established that synchrony depends on both dispersal ability and correlated environmental conditions, for example shared climate. However, what is not clear is whether differences in habitat or position within a species' range also mediate synchrony. Since synchronous metapopulations are thought to be more extinction‐prone, establishing the drivers of synchrony has clear conservation implications. Using three butterfly species (Maniola jurtina, Pyronia tithonus and Aphantopus hyperantus) we investigated the effects of habitat similarity and range position on population synchrony, after accounting for the effects of distance and climate. Range position was present in all minimum adequate models, though non‐significant using Mantel randomization tests in one case. We show that M. jurtina and P. tithonus synchrony is not consistent across species' ranges, with marginal populations showing more synchronous dynamics. Increased climatic constraints on marginal populations, leading to a narrower range of suitable microhabitats may be responsible for this, which is supported by the result that habitat similarity between sites was also positively correlated with population synchrony. As the landscape becomes increasingly homogeneous, overall population synchrony may be expected to rise. We conclude that habitat modification and climate change have the capacity to drive changes in population synchrony that could make species more vulnerable to extinction.     

Identifying recorder-induced geographic bias in an Iberian butterfly database

A database with comprehensive butterfly faunistic information from the Iberian Peninsula and the Balearic Islands was used to estimate inventory completeness as well as the environmental, spatial, and land-use effects on sampling intensities, on a 50×50 km UTM grid. The degree of sampling effort was assessed by means of accumulation curves based on the Clench function. Using the General Linear Model regression procedure, the effects of 22 variables on the estimated sampling efforts were assessed. This combination of methods is proposed as a preliminary step in biodiversity studies, in order to evaluate not only the degree of geographic coverage of existing faunistic data, but also the amount and nature of the bias on the faunistic work done throughout the last two centuries. The degree of spatial effects on the data was greater than the effects of environmental or land-use variables, although the latter two proved to be locally relevant. The results confirm previous findings that collecting is often skewed by relatively simple factors that affect collector activity, such as accessibility and attractiveness of sampling sites. With regard to Iberian and Balearic butterflies, adequate inventories on the scale investigated may probably suffice for further studies of the diversity of this insect group. Additionally, the results enabled us to develop general guide lines for the design of further faunistic work in the area.     

国裸子植物物种丰富度空间格局与多样性中心

中国拥有世界上最丰富的裸子植物区系,对理解全球裸子植物分布变化与系统演化具有重要意义.我们利用中国天然分布的202种裸子植物的水平和垂直分布信息获得物种分布区范围,探讨了中国裸子植物在科、属、种水平的分布特点.总体上,中国裸子植物物种丰富度南高北低,山地裸子植物丰富度较高,平原和高原相对贫乏;随分类阶元变高,丰富度高值区域面积逐渐扩大,高值中心逐渐南移.占中国陆地面积5%的裸子植物最丰富区域内分布了85%的中国自然分布的裸子植物物种.我们将这些区域划分为6个裸子植物多样性中心:(1)东喜马拉雅-横断山脉-秦岭,(2)滇黔桂-南岭,(3)华中山地,(4)黄山-武夷山脉,(5)海南岛南部山地,(6)长白山(甑峰山附近).各中心裸子植物区系之间的特点和联系反映了各自地理位置的差异和空间距离的隔离作用,其中横断山脉地区是中国裸子植物最重要的分化中心.     

Environmental correlates of mammal species richness in South America: effects of spatial structure, taxonomy and geographic range

Although some consensus exists regarding the positive synergism between energy and heterogeneity in increasing species diversity, the role of environmental variability remains controversial. We examine how these factors interact to explain spatial variation in mammal species richness in South America. After taking into account the effects of spatial autocorrelation and area, elevation variability and energy mainly drive spatial variation in mammal species richness. The effect of environmental variability is less important. When different taxonomic groups of mammals are analyzed separately, three ways emerge whereby energy and heterogeneity interact to promote species richness. Heterogeneity may have no effect on species richness, habitat heterogeneity and energy availability contribute independently to species richness, or heterogeneity increases in importance with an increase in energy availability. The partition of species into range size quartiles shows that habitat heterogeneity and temporal instability in the resource supply account for the species richness pattern in the narrowest- ranging species. Habitat heterogeneity is significant also for intermediate ranging species but not for the widest-ranging species. Energy alone drives the species richness pattern in the latter species. The interplay between ecology and biogeographic history may ultimately explain these differences given that narrow- and wide-ranging species show distinct biogeographic patterns, and different taxonomic groups also unequally represent them.     

Comparisons of butterfly richness and abundance measures in prairie and barrens

Transect surveys of adult butterflies were conducted along fixed routes at 27 study sites grouped into three subregions of tallgrass prairie and one subregion of pine-oak barrens in the midwestern USA. Within subregion, each site was visited the same number of times over 5–7 years on similar dates with similar weather. For each site, five indices of species richness and/or abundance were calculated both for total butterflies and for specialist species primarily restricted to native herbaceous vegetation. These indices were then analysed as to how much they agreed or conflicted in site ranking and how site rankings based on total butterflies compared to those based on specialists. Variation in site ranking by different indices was relatively low. Mean site rank by specialist indices covaried significantly with mean site rank by indices for total butterflies. Numerous studies have shown that on a regional scale, areas of higher richness for all species in a taxonomic group are different habitat types (based on amount of canopy or degree of degradation) from hotspots for that taxon's subset of species of conservation concern (endemics or specialists). But in this study, within a habitat type, site rankings based on total butterflies significantly tended to agree with site rankings based on specialist butterflies. This suggests that site prioritization and management favouring specialist butterflies would also favour the overall butterfly fauna possible in the same habitat.     

Local versus landscape determinants of butterfly movement behaviors

Large-scale patterns of animal distributions and abundances may be determined by mechanisms that act at local or landscape scales. We studied the movement behaviors of four species of bottomland butterflies in a natural setting to examine the determinants of movement behavior across different scales. We tested the relative importance of three landscape attributes: drainage slope, boundary type, and stream proximity, and local habitat attributes related to food plants and plants that influence habitat structure. Across species, we tested the relative importance of organism size and habitat specificity to explain response variation. In general, butterfly responses to landscape features were more universal than responses to local features. Specifically, results from this study showed that drainage slope did not influence movement behaviors but boundary type, stream proximity, and host plant abundance all influenced movement patterns. Responses to local features varied by species and often complemented landscape effects on movement. Responses to all features were not related to butterfly size, but did vary in accordance with butterfly host plant specificity. These behaviors help to explain landscape-level variation in population distribution among species.     

西南地区壳斗科物种丰富度和特有性分布格局模拟及其环境解释

如何准确地模拟物种宏观丰富度格局和特有性中心是生物多样性保护工作的重点,也是生物地理学的热点话题.西南地区是我国壳斗科植物最丰富的地区之一,但物种多样性格局及环境驱动机制尚不清楚.本研究基于西南地区161种壳斗科植物7258个分布点位数据,利用点格局法和物种分布模型两种方式构建了物种丰富度、加权特有性指数和校正加权特有...     

Landscape composition and habitat area affects butterfly species richness in semi-natural grasslands

During the last 50 years, the distribution and abundance of many European butterfly species associated with semi-natural grasslands have declined. This may be the result of deteriorating habitat quality, but habitat loss, resulting in decreasing area and increasing isolation of remaining habitat, is also predicted to result in reduced species richness. To investigate the effects of habitat loss on species richness, we surveyed butterflies in semi-natural grasslands of similar quality and structure, but situated in landscapes of different habitat composition. Using spatially explicit habitat data, we selected one large (6–10 ha) and one small (0.5–2 ha) grassland site (pasture) in each of 24 non-overlapping 28.2 km² landscapes belonging to three categories differing in the proportion of the area that consisted of semi-natural grasslands. After controlling for local habitat quality, species richness was higher in grassland sites situated in landscapes consisting of a high proportion of grasslands. Species richness was also higher in larger grassland sites, and this effect was more pronounced for sedentary than for mobile species. However, the number of species for a given area did not differ between large and small grasslands. There was also a significant relationship between butterfly species richness and habitat quality in the form of vegetation height and abundance of flowers. In contrast, butterfly density was not related to landscape composition or grassland size. When species respond differently to habitat area or landscape composition this leads to effects on community structure, and nestedness analysis showed that depauperate communities were subsets of richer ones. Both grassland area and landscape composition may have contributed to this pattern, implying that small habitat fragments and landscapes with low proportions of habitat are both likely to mainly contain common generalist species. Based on these results, conservation efforts should aim at preserving landscapes with high proportions of the focal habitat.