面积、温度及分布区限制对物种丰富度海拔格局的影响: 以秦岭太白山为例

 物种丰富度的分布格局及其形成机制是生态学研究的热点。以往的研究主要描述丰富度的格局, 而对其形成机制研究较少, 且主要集中于探讨单个因子或过程的影响。物种丰富度同时受到多个因子和过程的综合作用, 面积、温度及物种分布区限制被认为是控制山地物种丰富度海拔格局的主要因素, 三者同时沿海拔梯度而变化, 同时作用于丰富度的海拔格局。幂函数种-面积关系(SAR)、生态学代谢理论(MTE)及中域效应假说(MDE)分别基于以上3个因素, 从机制上解释了物种丰富度 的海拔格局。探讨这些假说的相对影响对研究物种丰富度的大尺度格局及其形成机制具有重要意义。方差分离方法有利于分解不同因素的影响, 为此, 该文以秦岭太白山的植物物种丰富度为例, 采用方差分离和逐步回归方法, 分析了SAR、MTE及MDE对物种丰富度海拔格局的影响。结果表明, 太白山的植物物种丰富度沿海拔梯度呈单峰分布格局, 但丰富度峰值存在类群差异; 对太白山所有植物物种丰富度的垂直格局而言, SAR、MTE及MDE分别解释了其物种丰富度随海拔变化的66.4%、19.8%和37.9%, 共同解释了84.6%, 在消除其他因素的影响后, SAR和MTE的独立影响较高(分别为25.5%和17.7%), 而MDE的独立影响不显著; 分类群研究则发现, 苔藓植物丰富度的海拔格局主要受MDE的影响, 蕨类植物丰富度的海拔格局同时受到SAR、MTE以及MDE的影响, 而种子植物物种丰富度的海拔格局主要受SAR和MTE影响。     

基于等面积高度带划分的贺兰山维管植物物种丰富度的海拔分布格局

山地植物物种丰富度海拔分布格局是生物多样性研究的热点之一。以往研究中一般将山体划分为等海拔间距的高度带, 以分析物种丰富度的垂直格局, 其缺陷在于因各高度带面积不相等而可比性下降。为消除面积不相等的影响, 作者利用数字高程数据(DEM, Digital Elevation Model)在地理信息系统(GIS)工具支持下, 尝试将贺兰山(海拔范围1,300–3,500 m)划分为等面积的数个高度带, 从而分析其物种丰富度的海拔格局。结果表明: (1) 贺兰山物种丰富度呈现为单峰式海拔格局, 峰值出现在海拔2,000 m附近。(2) 逐步回归分析显示, 坡度异质性是解释物种丰富度海拔分布格局的最优因子。高度带的坡度异质性越大, 意味着地形的起伏变化越大, 反映出生境类型越趋多样化, 从而可维持多个物种的共存。(3) 贺兰山植物物种丰富度在海拔2,000 m 附近达到峰值, 可能与植被演变历史、气候条件、地形复杂度、生态过渡带和中间膨胀效应的共同影响有关。(4) 对山体进行等面积划带, 可直接消除面积不相等带来的影响, 与等间距划带的方法相比, 尤其在物种海拔分布信息准确度较高时更具优势。     

长江三峡库区物种多样性的垂直分布格局:气候、几何限制、面积及地形异质性的影响

为了验证生物多样性地理格局的几个重要假说,即种-面积关系、水分-能量动态假说、几何限制(中域效应)假说和生境异质性假说,作者以长江三峡库区维管植物物种丰富度沿海拔梯度的分布格局为例,采用多元回归和方差分解方法,研究了面积、气候、几何限制、地形异质性对多样性垂直格局的独立影响和协同作用,及其对各植物类群(不同分布宽度、不同分布区类型和不同生长型)影响的差异.结果表明,三峡库区各种植物类群的物种丰富度随着海拔上升均呈先升后降的单峰格局.水分-能量动态假说对多样性格局有很强的解释能力,其总的解释力(＞93％)明显高于其他所有解释机制.但对于很多植物类群而言,水分和能量的解释力中有很大一部分属于几何限制、面积及地形异质性等因素的协同作用.几何限制对分布宽度大的物种的多样性格局解释力很强,但对分布宽度小的物种作用很小;面积自身对物种丰富度解释力较强,但在考虑了其他环境因素的影响时,仅对少数植物类群有解释力;地形异质性自身对多样性的解释能力很弱,但在多元回归模型中起着必要的作用.综合来看,水分-能量动态是解释三峡库区植物多样性垂直格局的最重要的机制.几何限制的作用随着物种分布宽度减小而递减;地形异质性虽然对多样性垂直格局的影响较弱,但也是一种必要的补充解释机制;由于面积与气候、几何限制等因素存在强烈的共线性,面积对植物多样性垂直格局的相对作用大小还需要进一步的系统比较研究.     

西双版纳种子植物物种多样性的垂直格局及机制

物种多样性海拔分布格局及其形成机制的研究是生物地理学和宏观生态学的重要议题之一。本文利用西双版纳植物专著资料, 结合高分辨率的地形和气候等数据, 探讨了面积、边界限制和现代气候对西双版纳野生种子植物物种丰富度及物种密度海拔分布格局的影响。结果表明: (1)物种丰富度呈单峰分布格局, 面积(81.9%)、边界限制(17.5%)和气候(60.0-69.3%)都不同程度地解释了物种丰富度的单峰格局; (2)利用幂函数种-面积关系计算的物种密度沿海拔大致呈减小的分布趋势, 气候的解释率降低为32.6-40.6%, 与边界限制无显著相关关系; (3)利用等面积高度带划分得到的物种密度沿海拔呈单峰变化趋势, 物种密度与边界限制无显著相关性, 但气候对物种密度的解释率为81.6-89.9%。研究结果有助于准确全面地理解物种多样性的海拔分布格局及其成因机制, 为西双版纳生物多样性保护提供理论支撑和实践指导。     

白水江自然保护区植物物种多样性的垂直格局: 面积、气候、边界限制的解释

采用线性回归模型和方差分离方法,分析面积、气候、边界限制对白水江自然保护区植物物种垂直分布格局的相对作用和独立作用.结果表明:白水江自然保护区植物物种及不同类群的物种多样性随海拔的升高呈单峰分布格局,峰值分布偏向低海拔段.白水江自然保护区植物物种的垂直分布格局是由各种因素协同作用的结果.水分能量动态假设对白水江自然保护区植物物种多样性垂直格局的解释力最强,面积是影响物种多样性格局的第二重要因子.边界限制效应可作为对白水江自然保护区植物物种多样性垂直格局解释的补充机制.边界限制对不同种域的物种解释力存在较大差异,随着物种分布宽度的增加,边界限制的解释力逐渐增加.     

高黎贡山种子植物物种丰富度沿海拔梯度的变化

物种丰富度沿海拔梯度的分布格局成为生物多样性研究的热点。为探讨中尺度区域物种丰富度沿海拔梯度的分布,本文以高黎贡山为研究对象,利用该地区的地方植物志资料,结合通过GIS生成的区域数字高程模型(DEM)数据,分析了该区域全部种子植物和乔木、灌木、草本三种生活型种子植物物种丰富度的垂直分布格局以及物种密度沿海拔梯度的变化特征。结果表明：(1)全部种子植物和不同生活型植物物种丰富度随着海拔的升高呈现先增加后减小的趋势,最大值出现在海拔1500—2000m的范围;(2)物种密度与海拔也呈现单峰曲线关系;(3)物种丰富度和物种密度分布格局的形成主要受海拔所反映的水、热状况组合以及物种分布的边界影响。     

物种丰富度垂直分布格局及影响机制

物种丰富度分布格局是一定地域内物种丰富度沿三维空间的立体分布,包括物种丰富度在经度、纬度和垂直梯度(海拔高度和海水深度)三个维度上的空间分异。近年来物种多样性的垂直分布格局与机制研究得到了生物地理学家和生态学家的重视。物种丰富度的垂直分布格局存在多种类型,但随海拔增加而物种数减少的单调递减模型和中海拔物种丰富度最高的单峰模型较为常见。目前在机制研究中验证较多的是气候稳定性、生物因子(种间相互作用)、能量、生境异质性、干扰、进化时间、物种分化速率、面积、中域效应(mid-domain effect)、生态位保守性(niche conservatism)等假说和机制。物种丰富度的分布格局是多方面因素综合作用的结果;由于地理、地形、气候、地质演化历史、物种库和进化历史、物种分化速率、干扰等差异,在不同地区存在着特别的物种丰富度空间分布格局和机制;处于同一地区的不同类群的物种也因进化扩散历史和生态适应能力不同而呈现多样化的分布格局。因此,对不同地区和类群的物种丰富度格局和机制进行研究应具体分析后才能得到可信结论。     

物种丰富度的垂直分布格局及其形成机制

物种丰富度海拔梯度格局及其形成机制一直是宏观生态学研究的重要议题。物种丰富度沿海拔梯度呈4种分布格局,其中单峰分布格局最为普遍。人们提出各种假设从不同角度解释物种丰富度的海拔梯度格局,这些假设主要以气候、空间效应、物种间相互作用及其进化历史为出发点,提出的假设主要有水分-能量动态假设、生物多样性代谢理论、生产力假设、种-面积关系、中域效应、栖息地异质性假设及静态进化模型。本文系统介绍了物种丰富度的海拔格局及影响其格局的生态过程。目前,物种丰富度的海拔格局主要集中在现代气候对物种丰富度海拔格局的形成过程的作用,但这些现代气候的参数之间具有显著共线性,难以分辨具体的某种环境因子对其的绝对贡献。结合现代气候和物种的谱系进化过程,系统比较不同海拔区域物种的系统发育特征有助于进一步理解物种丰富度的海拔分布格局及其成因机制,同时有助于理解现代环境的波动对植被群落的影响。     

拉萨河流域亏组山植物物种丰富度和群落特征研究

为研究草地植物群落、物种丰富度及其分布格局与影响因子之间的关系,该文以拉萨河流域林周县卡孜乡亏组山为研究地点,对山体垂直样带(3900~5100 m)植物群落特征、植物物种丰富度与各影响因子之间的关系进行了研究。研究区域共设置了13个样带(每隔100 m设一个样带),每样带设置0.5 m×0.5 m的5个样方进行植被调查,运用主成分分析(PCA)和双向指示种分析法(TWINSPAN)对植物群落进行排序和分类,运用冗余分析法(RDA)对群落及其分布格局与影响因子之间的关系进行分析,对植物物种丰富度与各因子之间的关系进行了回归分析。结果表明:该山体植物可分为3个群落类型,含7个群丛;影响区域植物群落物种组成和分布格局的主要环境因子为海拔,其次是坡度;物种丰富度与海拔、分种盖度呈单峰关系,与总盖度、坡度、地上生物量呈正相关。该研究结果为区域植物物种多样性和山地植物资源的保护和可持续利用提供了科学依据。     

伊犁河谷山地植物群落物种多样性分布格局及环境解释

结合对新疆伊犁河两岸科古琴山南坡(河谷北坡)和乌孙山北坡(河谷南坡) 94个样地的调查资料, 采用DCCA (detrended canonical correspondence analysis)排序法, 分析了物种多样性指数与环境因子之间的关系, 运用广义可加模型拟合植物群落总体多样性指数对海拔梯度的响应曲线, 探讨了伊犁河谷山地植物群落物种多样性的垂直分布格局。结果表明: 在调查的94个样地中, 共出现259种植物, 其中, 草本植物的种类极其丰富, 多达235种, 木本植物的种类极其有限; 垂直结构完整的植物群落具有较高的多样性指数; 河谷北坡植物群落物种多样性的分布格局受海拔、坡度、坡向以及土壤全氮、全钾、有机质、含水量等环境因子的影响较大, 而在河谷南坡, 物种多样性分布格局主要受坡度、海拔、有效磷含量和土壤含水量等环境因子的影响; 在河谷北坡, 植物群落的Patrick丰富度指数与Shannon-Wiener指数与海拔呈明显的双峰曲线关系, Simpson指数与Pielou均匀度指数呈不对称的单峰格局, 而河谷南坡的物种多样性指数随海拔均呈双峰格局, 尽管Patrick丰富度指数不甚明显。山地植物群落物种多样性的垂直分布格局是由海拔为主的多种环境因子综合作用的结果。     

Climate and history explain the species richness peak at mid-elevation for Schizothorax fishes (Cypriniformes: Cyprinidae) distributed in the Tibetan Plateau and its adjacent regions

Aim  We studied elevational species richness patterns of Schizothorax fishes and identified the roles of ecological and evolutionary factors in shaping the patterns of elevational diversity.
Location  The Tibetan Plateau and its adjacent regions.
Methods  We assembled distribution and altitude data for all Schizothorax species using the literature. We merged ecological and evolutionary approaches to test the relationships between species richness and ecological factors (climate, area, the mid-domain effect) or evolutionary factors (diversification rates and time of colonization).
Results  We found that species richness of Schizothorax fishes peaked at mid-elevations. Rainfall, area, the mid-domain effect and diversification rate were weak predictors of the richness pattern. Temperature showed a nonlinear relationship with species richness. Temperature and time of colonization were the most important variables in explaining the elevational diversity pattern.
Main conclusion  Our findings indicate that the time-for-speciation effect and niche conservatism play important roles in variation of species richness.     

Elevational patterns of frog species richness and endemic richness in the Hengduan Mountains, China: geometric constraints, area and climate effects

We studied frog biodiversity along an elevational gradient in the Hengduan Mountains, China. Endemic and non-endemic elevational diversity patterns were examined individually. Competing hypotheses were also tested for these patterns. Species richness of total frogs, endemics and non-endemics peaked at mid-elevations. The peak in endemic species richness was at higher elevations than the maxima of total species richness. Endemic species richness followed the mid-domain model predictions, and showed a nonlinear relationship with temperature. Water and energy were the most important variables in explaining elevational patterns of non-endemic species richness. A suite of interacting climatic and geometric factors best explained total species richness patterns along the elevational gradient. We suggest that the mid-domain effect was an important factor to explain elevational richness patterns, especially in regions with high endemism.     

Elevational gradients, area and tropical island diversity: an example from the palms of New Guinea

The factors causing spatial variation in species richness remain poorly known. In this study, factors affecting species richness of palms (Palmae/Arecaceae) were studied along the elevational gradient of New Guinea. Interpolated elevational ranges were calculated from a database of all known collections for 145 species in 32 genera. The amount of land area at different elevations greatly affects the species richness gradient. If assessed in equal-elevation bands species richness appears to decline monotonically, but when assessed in equal-area bands species richness shows a pronounced mid-elevation peak, due to the large proportion of lowlands in New Guinea. By randomising species ranges within the total elevational gradient for palms and accounting for area, we found the mid-elevation peak to be consistent with a mid-domain effect caused by the upper and lower limits to palm distribution. Our study illustrates the importance of accounting for area in macroecological studies of richness gradients and introduces a novel yet simple method for doing this through the use of equal-area bands. Together, the effect of area and the mid-domain effect explain the majority of variation in species richness of New Guinea palms. We support calls for the multivariate assessment of the mid-domain effect on an equal footing with other potential explanations of species richness.     

Geographical patterns in species richness of the benthic polychaetes in the continental shelf of the Gulf of California, Mexican Pacific

The present study is the first attempt to describe meso-scale patterns in the species richness of polychaetes along the Gulf of California, which stretches from about 23°N to 31°N. We examine herein the spatial changes in species distribution and explore the overlapping of species’ ranges towards the centre of the Gulf, to test whether the mid-domain effect (MDE) could explain an expected mid-domain peak in species richness. The faunal composition and the latitudinal range of 244 species of polychaetes recorded along the continental shelf of the Gulf of California were analysed in latitude bands of 1°. The species composition changes around the Gulf’s archipelago (~29°N), and the highest values of species richness are found at the 25° (197 species) and 26° (193 species) of latitude. Although the species richness pattern could be described by a parabolic shape, the regional trend was not strongly consistent with the peak of diversity at 27°N (176–191 species) predicted by the mid-domain effect: the random sorting of species’ ranges within spatial domain does not explain satisfactorily the geographical patterns of diversity. Nevertheless, a partial contribution of MDE to these natural patterns of diversity could be detected, and the increase in species richness towards middle latitudes was basically determined by species with distribution ranges larger than 6°. The low level of significance between the empirical species richness pattern and the mid-domain model prediction for polychaetes in the Gulf does not restrict their use as a model for exploring the randomness of the diversity patterns.     

Altitudinal patterns of seed plant richness in the Gaoligong Mountains, south-east Tibet, China

Elevational patterns of species richness and their underlying mechanisms have long been a controversial issue in biodiversity and biogeographical research, and several hypotheses have been proposed in the past decades. Local and regional studies have suggested that area and geometric constraint are two of major factors affecting the elevational pattern of species richness. In this study, using data of seed plants and their distribution ranges and a Digital Elevation Model data set, we explored altitudinal patterns of seed plant richness and quantified the effects of area and the mid-domain effect (MDE) on the richness patterns in a high mountain area, Gaoligong Mountains (ranging from 215 m to 5791 m a.s.l.) located in south-eastern Tibet, China. The results showed that richness and density (richness/log-transformed area) of seed plants at species, genus, and family levels all showed hump-shaped patterns along the altitudinal gradient. The altitudinal changes in richness of species with three different range sizes (< 500 m, 500–1500 m, and > 1500 m), species of different plant life-forms (trees, shrubs, and herbs), and endemic species further confirmed this finding. Analysis of Generalized Linear Model depicted that although the area of each elevational band was always in high correlation with the species richness, the MDE could explain 84.9%, 33.8%, 83.8%, and 84.5% of the total variation in richness for all species and the three species groups with different range sizes, respectively. This suggests that the MDE significantly influences the patterns of species richness and is likely be stronger for broad-ranged species than for narrow-ranged ones in the Gaoligong Mountains.     

Evaluating environmental and geometrical constraints on endemic vertebrates of the semiarid Caatinga (Brazil)

Most hypotheses explaining species diversity are related to the current and/or past environment, but explanations that rely on geometrical constraints, such as the mid-domain effect, have been recently proposed. Under this type of model, more species are expected in the centre of a domain due to the extent of occurrence of the species being constrained by the domain's geometry. Here we evaluated the proportion of the explanatory power of space, the current environment, historical vicariance events and geometric constraints on the richness of the Caatinga's terrestrial endemic vertebrates using a partial regression approach. The current environment had the highest proportion of explanatory power for the variance of the Caatinga's endemic species, in its shared effect, explaining approximately 47%. On the other hand the geometric constraints model (GCM) had the highest proportion in its single explanatory power with 15% of explanatory power. The amount of explanation by GCM changes when analyzing richness of endemics calculated from species with different range sizes. Despite the fact that GCM is the main driver of the Caatinga's endemic species, in its single effect, this does not necessarily indicate that geometric constraints and random dispersal within species ranges drive richness. The GCM model underestimates richness in the centre of the Caatinga, suggesting that other ecological and evolutionary processes may help explain the overall pattern of decreasing richness towards the borders of the biome.     

河西走廊水生植物多样性格局、群落特征及影响因素

水生植物是湿地生态系统重要组成部分,研究水生植物多样性分布格局及其影响因素对地区水生植物资源保护具有重要意义。通过野外调查并结合气候等环境因素,研究了河西走廊主要水生植物群落类型、数量特征、水生植物多样性分布格局及影响因素,并对中域效应假说进行了验证。研究结果表明:(1)河西走廊地区共有水生植物29科42属84种,群落的聚类分析可将河西走廊水生植物群落划分为15个主要群落类型;(2)河西走廊水生植物群落类型主要受到水温、海拔、经纬度等环境因子影响,群落物种多样性指数与盐度以及溶解性固体总量呈显著性相关;(3)河西走廊水生植物多样性空间格局呈现出"∩"型的单峰格局,中域效应模型能较好地解释该地区水生植物多样性水平的纬度格局及海拔垂直分布格局,对该区域水生植物物种丰富度在纬度和海拔梯度上的变异解释率分别为57.56%、63.5%。分析表明,河西走廊水生植物物种丰富度格局由几何(边界)限制和随机过程及其他未知因素共同控制,且几何(边界)限制和随机过程贡献率较大;同时本研究中未考虑的环境异质性、气候、人为干扰等因素也对河西走廊水生植物多样性空间分布产生重要影响。     

Three-dimensional mid-domain predictions: geometric constraints in North American amphibian, bird, mammal and tree species richness patterns

The "mid-domain effect" (MDE) has received much attention as a candidate explanation for patterns in species richness over large geographic areas. Mid-domain models generate a central peak in richness when species ranges are placed randomly within a bounded geographic area (i.e. the domain). Until now, domain limits have been described mostly in one-dimension, usually latitude or elevation, and only occasionally in two-dimensions. Here we test 1-D, 2-D and, for the first time, 3-D mid-domain models and assess the effects of geometric constraints on species richness in North American amphibian, bird, mammal and tree species. Using spatially lagged simultaneous autoregressive models, empirical richness was predicted quite well by the mid-domain predictions and the spatial autoregressive term (45–92% R²). However, our results show that empirical species richness peaks do deviate from those of the MDE predictions in 3 dimensions. Variation explained (R²) by MDE predictions generally increased with increasing mean range size of the different biotic groups (from amphibian, to tree, mammal and finally bird data), and decreased with increasing dimensions being accounted for in the models. The results suggest geometric constraints alone can explain much of the variation in species richness with elevation, specifically with respect to the larger-range taxa, birds and mammals. Our analysis addresses many of the recent methodological criticisms directed at studies testing the MDE, and our results support the hypothesis that species diversity patterns are influenced by geometric constraints.