植物物候变化的全气候生产要素影响机制与模型研究

物候是气候变化的重要指示指标,不仅是植被动态模型的重要参数,也是陆地碳循环模型和大气环流模式的重要参数.本文从物候变化机制和物候模拟模型两方面综述了植物物候研究最新进展,剖析了物候研究趋势,指出当前物候研究大都仅考虑单一气候因子或少数几个气候因子的影响,涉及多气候因子相互作用影响的研究仍很少,还没有开展植物全气候生产要素相互作用影响物候的研究;受物候变化机制认识与研究物种的限制,现有物候模型仍不能反映植被生长的真实性,也不能有效模拟物候变化.本文指出,全面认识物候变化的机制迫切需要考虑影响植物物候的全气候生产要素及其相互作用,为物候模型发展提供依据.本文基于植物光合作用是最主要的物候影响因子的研究结论,提出采用气候生产潜力作为植物物候变化驱动因子,不仅体现气候环境因子对植物生产的综合作用及其在植物生长全过程中的一致性,还体现了生物因子、环境因子及其相互作用以及极端天气气候事件的影响,从而实现物候的准确模拟.本文提出的未来物候研究方向,有助于丰富物候认知、提升陆地碳循环和大气环流模式模拟的准确性.     

气候和土地利用与空间结构在影响大熊猫同域分布大中型哺乳动物物种丰富度中的相对作用

气候因子和土地利用因子是影响生物多样性分布格局的两个主要驱动因素.然而,当前关于气候因子和土地利用因子对生物多样性影响的研究主要集中在物种层面上,在群落水平上对生物多样性的影响依然知之甚少.本研究以大熊猫同域分布大中型哺乳动物为研究对象,结合物种丰富度数据、气候数据、土地利用数据以及经纬度数据,构建基于不同变量组合的多...     

中国东部海岛植物功能性状及其影响因子

研究海岛植物功能性状及其影响因子,有助于理解植物在海岛环境中的适应策略。以中国东部近海7个海岛典型植物群落为研究对象,测定群落内每种植物的6个功能性状(胸径、株高、平均单叶面积、比叶面积、小枝密度和小枝含水率),比较植物功能性状在不同气候带间的差异,探讨功能性状对气候因子(年均温、年降水和年平均风速)和土壤因子(土壤含水率、土壤总碳、土壤总氮和土壤总磷)的响应关系,并分析影响这些性状的主要驱动因子。研究结果表明:(1)所测功能性状中除比叶面积外,其他性状在不同气候带间差异显著,但常绿和落叶植物的功能性状在不同气候带间的差异不同,表明不同生活型植物对环境的响应程度和适应能力不同。(2)海岛植物功能性状对气候和土壤因子均有不同程度的响应,气候因子是影响中国东部海岛植物功能性状的重要因子,其中水热条件(年均温和年降水)是主导因子,在水热资源丰富的南方海岛,植物具有较大的胸径和小枝含水率、较小的叶片和小枝密度,北方海岛反之。此外,海岛植物的小枝性状(小枝密度和小枝含水率)受年平均风速的影响显著。该研究结果可为海岛植被恢复重建中物种的选择及植被布局规划提供一定的参考。     

基于物种分布模型评价土壤因子对我国毛竹潜在分布的影响

基于单类别支持向量机方法的物种分布模型, 利用政府间气候变化专门委员会(IPCC)气候情景模式和联合国粮食与农业组织(FAO)的全球土壤数据, 模拟1981-2099年我国毛竹(Phyllostachys edulis)的潜在空间分布及变化趋势, 比较考虑土壤因子前后模拟结果的差异, 旨在探究土壤因子对毛竹潜在空间分布模拟结果的影响。结果表明, 仅以气候因子为模拟变量和同时考虑气候与土壤因子为模拟变量的毛竹潜在空间分布模拟均具有较高精度, 毛竹潜在分布区表现为面积增加并向北扩张。模拟因子重要性分析表明表征温暖程度的气候因子在毛竹潜在分布模拟中起主导作用, 而表征土壤质地和酸碱性的土壤因子以限制性作用为主。同时考虑气候与土壤因子的模拟结果具有较高的模拟效率, 且在未来气候变化情景模式下毛竹潜在分布区面积增幅与向北迁移幅度均小于仅使用气候因子的模拟, 表明土壤要素对毛竹潜在分布具有明显的限制作用, 该结果对现在的毛竹潜在分布模拟研究具有重要的补充作用。     

中国芦苇沼泽植物物种丰富度分布格局及其驱动因素

明确大尺度物种丰富度的分布格局及其影响因素一直是生态学及生物地理学领域的核心议题之一,其对于预测全球环境变化背景下生物多样性的响应和制定减少生物多样性丧失的保护方案有着重要意义。本文基于全国芦苇(Phragmites australis)沼泽野外调查数据,结合气候、地理、土壤等环境因子,探讨了中国芦苇沼泽植物物种丰富度的分布格局及其驱动机制。结果表明:(1)中国的芦苇沼泽植物物种丰富度总体表现为亚热带湿润区和温带湿润半湿润区较高,青藏高原区、温带干旱半干旱区和滨海区较低的特点;(2)芦苇沼泽植物物种丰富度与年降水、土壤有机碳、土壤总氮、纬度均呈显著正相关,而与年均温、最冷月最低温、土壤pH值、海拔之间表现为显著负相关;(3)土壤因子特别是土壤pH值是影响芦苇沼泽植物物种丰富度的最主要因素,其次分别为淹水状况、气候因子和地理因子;(4)结构方程模型结果表明土壤因子、淹水状况和气候因子直接影响植物物种丰富度,而地理因子通过调控土壤因子、淹水状况与气候因子间接影响植物物种丰富度。总体而言,中国芦苇沼泽植物物种丰富度具有空间异质性,其分布格局受到多种因素的共同影响,土壤因子是影响植物物种丰富度...     

中国水稻潜在分布及其气候特征

基于全国层次和年尺度筛选的影响中国水稻分布的潜在气候指标,结合水稻地理分布信息,利用最大熵模型和ArcGIS软件的空间分析功能,分析了中国水稻潜在分布及其气候特征.结果表明:年降水量(P)、湿润指数(MI)、稳定通过18℃持续日数( N18)和≥10℃积温(∑T10)4个因子是影响水稻分布的主导气候因子,其累积贡献百分率达97.6％.采用主导气候因子作为环境变量重建气候水稻分布关系的最大熵模型,利用重建的最大熵模型给出的中国水稻存在概率,对中国水稻潜在分布区的气候适宜等级进行了划分,并分析了各适宜区的气候特征.研究结果可为中国水稻生产布局及制定应对气候变化政策提供参考.     

不同环境类型对华北落叶松分布的潜在影响

植物分布-环境因子之间的关系是生态学、生物地理学研究中一个核心问题。采用最大熵物种分布模型,选取气候、土壤和地形3类环境类型中33个因子,对气候、气候-土壤、气候-地形、气候-土壤-地形4种环境组合对华北落叶松在河北省、山西省和内蒙古自治区的分布预测进行建模和检验。利用ArcGIS空间统计,划分华北落叶松分布适宜区,并制作分布适宜性等级图。结果表明,不同环境类型组合对华北落叶松分布影响的训练集和检验集的AUC值在0.965-0.983之间,均达到极准确的精度水平。在影响华北落叶松分布的主导因子上,气候、气候-土壤两种环境类型组合均为最热月的最高温度、温度的年较差和季节性温度变异,累计贡献率均达到74%以上,而气候-地形、气候-土壤-地形两种环境类型组合中海拔和坡度的影响最大,分别为48.8%和51.8%。在影响华北落叶松适生区（中、高、极高适宜区）面积上,气候、气候-地形两种环境类型组合差异不大,分别为102583km²和100698 km²,而气候-土壤、气候-土壤-地形两种环境类型组合影响下的华北落叶松适生区面积出现显著下降分别为57134 km²和66754 km²。最大熵模型能够很好的预测华北落叶松分布,地形因子能明显改变单一气候因子对华北落叶松分布的预测结果,虽然土壤因子对落叶松分布格局的影响不大,但在适宜性、尤其是中等以上适宜区分布上,其影响作用显著。以上结论可为华北地区的生态修复和落叶松可持续经营提供依据。     

气候与土壤对烤后烟叶类胡萝卜素和表面提取物含量的影响

选择贵州生态条件差异较大的威宁县、开阳县和天柱县进行客土田间小区试验,以云烟85为材料,探讨气候与土壤对烤后烟叶类胡萝卜素和表面提取物含量的影响.结果表明:类胡萝卜素和表面提取物含量受生态条件的影响较大,地域性差异显著,打破犁底层耕作方式无明显作用.类胡萝卜素含量有随海拔高度增加而升高的趋势,但海拔高度的影响存在阈值效应;清香型烟叶的表面提取物含量明显高于中间香型.不同生态因子的影响程度明显不同,气候是关键生态影响因子.类胡萝卜素含量气候间的变异度是土壤的2-3倍,气候对β-胡萝卜素与叶黄素含量变异的贡献率分别为74％和64％,土壤贡献率分别为9％和12％.影响较大的前3个气候因子为大田生长前期的累积日照时数和累积降水量、成熟期的相对湿度,前3个土壤因子为有效钾、pH值和有效磷.表面提取物含量气候间的变异度是土壤的3-4倍,气候对腺毛分泌物与烷烃类蜡质含量变异的贡献率分别为61％和63％,土壤贡献率分别为13％和7％.影响腺毛分泌物含量的前3个气候因子为大田生长前期的平均气温、成熟期的相对湿度和≥10℃积温,前3个土壤因子为有效钾、有效铜含量和pH值.烷烃类蜡质含量的前3个气候影响因子为大田生长前期的累积降水量和平均气温、成熟期的平均气温,前3个土壤因子为有效锰、有效磷含量和阳离子交换量.土壤因子对烤后烟叶类胡萝卜素含量部位特征的影响较气候因子突出,气候因子对表面提取物含量部位特征的影响较土壤因子明显.     

大兴安岭人为火发生影响因素及气候变化下的趋势

我国重要的北方针叶林地区大兴安岭是林火高发地区.受气候变暖影响,该地区林火发生频率将会发生显著变化.模拟人为火的发生分布与影响因素之间的关系、加强气候变化下人为火的发生分布预测,对于林火管理和减少森林碳损失具有重要作用.本文采用点格局分析方法,基于大兴安岭1967—2006年的火烧数据,建立人为火空间分布与影响因素之间的关系模型,该模型以林火发生次数为因变量,选取非生物因子(年均温和降水量、坡度、坡向和海拔)、生物因子(植被类型)和人为活动因子(距离道路距离、距离居民点距离、道路密度)共9个因子为自变量.并采用RCP 2.6和RCP 8.5气候情景数据代替当前气候情景预测2050年大兴安岭人为火的空间分布状况.结果表明: 点格局模型能够较好地模拟人为火发生分布与空间变量的关系,可以预测未来气候下人为火的发生概率.其中,气候因子对人为火的发生具有明显的控制作用,植被类型、海拔和人为活动等因子对人为火的发生也具有重要影响.林火发生预测结果表明,未来气候变化下,南部地区的林火发生概率将进一步增加,北部和沿主要道路干线附近将成为新的人为火高发区.与当前相比,2050年大兴安岭人为火的发生概率将增加72.2%～166.7%.在未来气候情景下,人为火的发生更多受气候和人为活动因素的控制.     

极小种群东北红豆杉所在群落特征及其环境解释

以东北红豆杉(Taxus cuspidata Sieb.et Zucc.)所在群落为对象,选择有东北红豆杉分布的21个样方进行群落调查,结合样方植物和环境数据,采用系统聚类、RDA分析、方差分解等方法研究东北红豆杉所在群落的特征及其主要影响因素。结果表明:(1)东北红豆杉所在群落共记录到48科90属107种植物,其物种组成丰富,可将群落划分为红松-紫椴林、红松-云杉-冷杉林、红松-槭树林、红松-风桦林4个林分类型;(2)温度、降水、林分类型和土壤pH等环境因子显著影响东北红豆杉的分布及其群落特征,整体上温度是最显著的影响因子,但在不同林型中各环境因子的主要作用不同;(3)气候因子、地形与土壤因子、生物因子三者对东北红豆杉所在群落分布的共同解释率为49.85%,地形与土壤因子的单独解释力最大(15.70%),其次为气候因子(14.96%)和生物因子(9.79%)。总之,由于自然和人为因素的干扰,东北红豆杉所处的群落类型多样,且东北红豆杉种群特征在不同的群落中各不相同,主要分布在红松-紫椴林和红松-云杉-冷杉林中,其他2种群落类型对回归保护地的选择具有很好的指示作用,应针对不同的分布群落采取不同的保护措施。     

Patterns and determinants of wood physical and mechanical properties across major tree species in China

The physical and mechanical properties of wood affect the growth and development of trees, and also act as the main criteria when determining wood usage. Our understanding on patterns and controls of wood physical and mechanical properties could provide benefits for forestry management and bases for wood application and forest tree breeding. However, current studies on wood properties mainly focus on wood density and ignore other wood physical properties. In this study, we established a comprehensive database of wood physical properties across major tree species in China. Based on this database, we explored spatial patterns and driving factors of wood properties across major tree species in China. Our results showed that(i) compared with wood density, air-dried density, tangential shrinkage coefficient and resilience provide more accuracy and higher explanation power when used as the evaluation index of wood physical properties.(ii) Among life form, climatic and edaphic variables, life form is the dominant factor shaping spatial patterns of wood physical properties, climatic factors the next, and edaphic factors have the least effects, suggesting that the effects of climatic factors on spatial variations of wood properties are indirectly induced by their effects on species distribution.     

Spatial patterns of wood traits in China are controlled by phylogeny and the environment

Aim Wood properties are related to tree physiology and mechanical stability and are influenced by both phylogeny and the environment. However, it remains unclear to what extent geographical gradients in wood traits are shaped by either phylogeny or the environment. Here we aimed to disentangle the influences of phylogeny and the environment on spatial trends in wood traits. Location China. Methods We compiled a data set of 11 wood properties for 618 tree species from 98 sampling sites in China to assess their phylogenetic and spatial patterns, and to determine how many of the spatial patterns in wood properties are attributable to the environment after correction for phylogenetic influences. Result All wood traits examined exhibited significant phylogenetic signal. The widest divergence in wood traits was observed between gymnosperms and angiosperms, Rosids and Asterids, Magnoiliids and Eudicots, and in Lamiales. For most wood traits, the majority of trait variation was observed at genus and species levels. The mechanical properties of wood showed correlated evolution with wood density. Most of the mechanical properties of wood exhibited significant latitudinal variation but limited or no altitudinal variation, and were positively correlated with mean annual precipitation based on both Pearson's correlation analysis and the phylogenetic comparative method. Correlations at family level between mean annual temperature and wood density, compression strength, cross‐section hardness, modulus of elasticity and volumetric shrinkage coefficient became significant after correction for phylogenetic influences. Main conclusions Phylogeny interacted with the environment in shaping the spatial patterns of wood traits of trees across China because most wood properties showed strong phylogenetic conservatism and thus affected environmental tolerances and distributions of tree species. Mean annual precipitation was a key environmental factor explaining the spatial patterns of wood traits. Our study provides valuable insights into the geographical patterns in productivity, distribution and ecological strategy of trees linking to wood traits.     

Relative importance of climate vs local factors in shaping the regional patterns of forest plant richness across northeast China

Northeast (NE) China covers three climatic zones and contains all the major forest types of NE Asia. We sampled 108 forest plots in six nature reserves across NE China to examine the influence of climate and local factors (canopy seasonality, successional stage, topography and forest structure) on geographic patterns of plant richness. We analyzed the relative effects of different factors at two spatial scales: the regional scale (across both latitude and altitude) and the local scale (along the altitudinal gradient within site). Our results showed that the relative importance of climate vs local factors differed remarkably depending on scale and functional group. While total and tree species richness were mainly limited by climate, herb and shrub richness was more related to local factors (especially at the local scale). In the climatic factors, heat sum was the major correlate of tree, shrub and total species richness, while herb richness was more associated with winter coldness. Precipitation was not a limiting factor for forest plant richness in NE China. Climate accounted for 34–76% of variation in richness at the regional scale, but explained only 0–44% at the local scale. Among the local factors, shrub species richness was sensitive to seasonal canopy openness, with higher richness in deciduous forests than in the evergreen needle-leaf forest. On the other hand, herb richness was sensitive to forest successional stage, with higher richness in middle- successional forests than in the early and late-sucessional forests. Local topography (aspect and position on slope) and forest structure (tree density) also showed remarkable influence on species richness. Our results suggest the importance of including local factors when examining large scale diversity gradient (especially for understory species), and the necessity of comparing diversity patterns among functional groups at different spatial scales.     

The structural characteristics and climatic and human impacts of deciduous oak forests in China

中国落叶栎林群落结构特征及气候和人类活动的影响 落叶栎林是中国温带地区和亚热带山地典型的植被类型之一。虽然已有大量关于中国落叶栎林的群落结构特征研究,但以往研究多局限于局域尺度或单个类群,而在国家尺度上的系统性研究较为缺乏。本文采用统一的调查方法获取682个天然落叶栎林样方数据,基于此数据分析了我国落叶栎林的群落结构特征及其空间格局,探究了影响其特征的气候和人为因素。研究结果表明,中国落叶栎林乔木 层的平均胸径、平均树高、平均林分密度和平均总胸高断面积分别为：13.7 cm、10.0 m、1468株/ha 和 24.3 m²/ha。落叶栎林乔木层、灌木层和草本层的平均物种丰富度分别为：6种/600 m²、10种/100 m²和 4种/1 m²。随纬度的升高,落叶栎林平均树高、林分密度、总胸高断面积、乔木层物种丰富度和灌木层物 种丰富度均呈现显著降低趋势,林分平均胸径无显著变化,而草本层物种丰富度呈现显著增加趋势。 进一步分析发现,相比于林分密度和草本层物种丰富度,气候和人类活动可以更多地解释平均胸径、平均树高、总胸高断面积、乔木层物种丰富度和灌木层物种丰富度的空间分异。此外,与降水相关的气候因子在塑造群落结构特征的空间格局方面起着更为重要的作用。总之,本研究为认识中国落叶栎林群落结构特征的生物地理格局及其对全球变化的响应提供了基础。     

Tree species distribution in temperate forests is more influenced by soil than by climate

Knowledge of the ecological requirements determining tree species distributions is a precondition for sustainable forest management. At present, the abiotic requirements and the relative importance of the different abiotic factors are still unclear for many temperate tree species. We therefore investigated the relative importance of climatic and edaphic factors for the abundance of 12 temperate tree species along environmental gradients. Our investigations are based on data from 1,075 forest stands across Switzerland including the cold‐induced tree line of all studied species and the drought‐induced range boundaries of several species. Four climatic and four edaphic predictors represented the important growth factors temperature, water supply, nutrient availability, and soil aeration. The climatic predictors were derived from the meteorological network of MeteoSwiss, and the edaphic predictors were available from soil profiles. Species cover abundances were recorded in field surveys. The explanatory power of the predictors was assessed by variation partitioning analyses with generalized linear models. For six of the 12 species, edaphic predictors were more important than climatic predictors in shaping species distribution. Over all species, abundances depended mainly on nutrient availability, followed by temperature, water supply, and soil aeration. The often co‐occurring species responded similar to these growth factors. Drought turned out to be a determinant of the lower range boundary for some species. We conclude that over all 12 studied tree species, soil properties were more important than climate variables in shaping tree species distribution. The inclusion of appropriate soil variables in species distribution models allowed to better explain species' ecological niches. Moreover, our study revealed that the ecological requirements of tree species assessed in local field studies and in experiments are valid at larger scales across Switzerland.     

不同林龄、树种落叶松人工林径向生长与气候变化的关系

研究人工林径向生长与气候变化的关系对全球气候变暖背景下人工林合理经营有着重要的意义。该文对在辽东山区广泛栽培的黄花落叶松(Larix olgensis)和日本落叶松(Larix kaempferi)人工林, 运用树木年轮气候学方法建立了辽宁草河口和湾甸子林场落叶松人工林年表, 分析了落叶松径向生长对气候变化的响应以及气候条件、树种、立地条件和林分因子(林龄、密度、蓄积量等)的相对影响程度。结果发现在影响年轮-气候关系的因素中, 气象因子的潜在蒸发散(PET)的影响力最大; 林龄、密度和蓄积量同时也具有重要的影响作用。中龄落叶松人工林径向生长主要与气温呈正相关关系, 成熟落叶松人工林径向生长主要与气温呈负相关关系; 而其他因素, 如树种、立地条件等的影响作用不大。这表明在气候变暖背景下随着林龄增加, 林分会逐渐受到气温升高导致的水分亏缺的限制, 导致明显的生长下降趋势, 因而气候变暖对成熟落叶松人工林威胁更为严重, 所以要注重对成熟林的优先保护, 同时可以预测, 随着东北地区今后气候进一步变暖, 可能将逐步影响到林龄更小的林分的生长, 因此需要进一步研究如何在落叶松人工林经营中采取科学的措施来更好地应对未来气候变化。     

Shifts in trait means and variances in North American tree assemblages: species richness patterns are loosely related to the functional space

One of the key hypothesized drivers of gradients in species richness is environmental filtering, where environmental stress limits which species from a larger species pool gain membership in a local community owing to their traits. Whereas most studies focus on small‐scale variation in functional traits along environmental gradient, the effect of large‐scale environmental filtering is less well understood. Furthermore, it has been rarely tested whether the factors that constrain the niche space limit the total number of coexisting species. We assessed the role of environmental filtering in shaping tree assemblages across North America north of Mexico by testing the hypothesis that colder, drier, or seasonal environments (stressful conditions for most plants) constrain tree trait diversity and thereby limit species richness. We assessed geographic patterns in trait filtering and their relationships to species richness pattern using a comprehensive set of tree range maps. We focused on four key plant functional traits reflecting major life history axes (maximum height, specific leaf area, seed mass, and wood density) and four climatic variables (annual mean and seasonality of temperature and precipitation). We tested for significant spatial shifts in trait means and variances using a null model approach. While we found significant shifts in mean species’ trait values at most grid cells, trait variances at most grid cells did not deviate from the null expectation. Measures of environmental harshness (cold, dry, seasonal climates) and lower species richness were weakly associated with a reduction in variance of seed mass and specific leaf area. The pattern in variance of height and wood density was, however, opposite. These findings do not support the hypothesis that more stressful conditions universally limit species and trait diversity in North America. Environmental filtering does, however, structure assemblage composition, by selecting for certain optimum trait values under a given set of conditions.     

Idiosyncratic soil-tree species associations and their relationships with drought in a monodominant Amazon forest

Monodominant forests are characterized by the strong influence of a single species on the structure and diversity of the community. In the tropics, monodominant forests are rare exceptions within the generally highly diverse tropical forest biome. Some studies have shown that tree monodominance may be a transient state caused by successional and demographic variation among species over time. Working in a Brosimum rubescens Taub. (Moraceae) monodominant forest at the southern edge of Amazonia, we tested the hypotheses that local-scale variation in intra- and interspecific spatial patterns of dominant tree species is affected by i) demographic rates of recruitment and mortality following severe droughts, ii) local variation in edaphic properties, and iii) occupation of species in the vertical layer of the forest. We quantified intra- and interspecific spatial patterns and edaphic associations of the five most abundant species using aggregation and association distance indices, and examined changes over time. We found some support for all hypotheses. Thus, intra- and interspecific spatial patterns of most species varied over time, principally after severe drought, emphasizing species-level variability and their interactions in sensitivity to this disturbance, even as B. rubescens monodominance was maintained. While positive and negative spatial associations with edaphic properties provide evidence of habitat specialization, the absence of negative spatial associations of B. rubescens with edaphic properties indicates that this species experiences little environmental restriction, and this may be one of the factors that explain its monodominance. Spatial repulsion and attraction between species in the same and in different vertical layers, respectively, indicates niche overlap and differentiation, while changes over time indicate that the relationships between species are dynamic and affected by drought disturbance.     

Multi-scale determinants of dung beetle assemblage structure across abiotic gradients of the Kalahari–Nama Karoo ecotone, South Africa

Aim Changing conditions across spatial gradients are primary determinants of biotic regions, local habitats, and distributional edges. We investigate how a climatic gradient and edaphic mosaic interact as multi‐scale drivers of spatial patterns in scarabaeine dung beetles. The patterns are tested for congruency with ecoregion and floral boundaries over the same gradient, as responses to physical factors often differ among higher taxa. Location Southern Africa and the Nama Karoo–Kalahari ecotone, Northern Cape, South Africa. Methods Data consisted of the climatic distributions of 104 species and their abundances at 223 sites in two ecoregions/floral biomes, four bioregions, and 13 vegetation units. Factor analyses determined the biogeographical composition of the species, and regional‐ to local‐scale patterns in species abundance structure. Hierarchical analysis of oblique factors determined the proportional contribution of spatial variance to patterns. One‐way anova was used to test for significant separation of patterns along factor axes. Stepwise multiple regression was used to determine correlations of five physical attributes with species richness, Shannon‐Wiener diversity, and factor loadings for the study sites. Results Four biogeographical influences overlap in the study region, although rank contribution declines from south‐west arid through north‐east savanna to widespread and south‐east highland taxa. Species abundance structure comprises five subregional patterns, two centred to the north‐east (Kalahari, Isolated Kalahari Dune) dominated by Kalahari influence, and three to the south‐west (Nama Karoo subdivisions: Bushmanland, ‘Upper’, ‘Stony Prieska’) dominated by south‐west arid influence. Kalahari deep sands are characterized especially by a warmer, moister climate, whereas the Nama Karoo mosaic of deep or stony soils is characterized especially by north‐west aridity (Bushmanland), south‐east cooler temperatures (‘Upper’), or excessively stony soils (‘Stony Prieska’). Four of the subregional patterns each comprised three localized patterns related primarily to relative stoniness, edge effects from geographical position, or incidence of rainfall. Species richness and diversity declined with decreasing rainfall and increasing stoniness. Main conclusions Climatic and edaphic factors are important multi‐scale determinants of spatial patterns in dung beetle assemblage structure, with edaphic factors becoming more important at local spatial scales. The patterns are roughly congruent with the Kalahari Savanna–Nama Karoo ecotone at the floral biome or ecoregion scale, but show limited coincidence with finer‐scale floral classification.     

Co‐occurrence patterns of trees along macro‐climatic gradients and their potential influence on the present and future distribution of Fagus sylvatica L.

Aim During recent and future climate change, shifts in large‐scale species ranges are expected due to the hypothesized major role of climatic factors in regulating species distributions. The stress‐gradient hypothesis suggests that biotic interactions may act as major constraints on species distributions under more favourable growing conditions, while climatic constraints may dominate under unfavourable conditions. We tested this hypothesis for one focal tree species having three major competitors using broad‐scale environmental data. We evaluated the variation of species co‐occurrence patterns in climate space and estimated the influence of these patterns on the distribution of the focal species for current and projected future climates. Location Europe. Methods We used ICP Forest Level 1 data as well as climatic, topographic and edaphic variables. First, correlations between the relative abundance of European beech (Fagus sylvatica) and three major competitor species (Picea abies, Pinus sylvestris and Quercus robur) were analysed in environmental space, and then projected to geographic space. Second, a sensitivity analysis was performed using generalized additive models (GAM) to evaluate where and how much the predicted F. sylvatica distribution varied under current and future climates if potential competitor species were included or excluded. We evaluated if these areas coincide with current species co‐occurrence patterns. Results Correlation analyses supported the stress‐gradient hypothesis: towards favourable growing conditions of F. sylvatica, its abundance was strongly linked to the abundance of its competitors, while this link weakened towards unfavourable growing conditions, with stronger correlations in the south and at low elevations than in the north and at high elevations. The sensitivity analysis showed a potential spatial segregation of species with changing climate and a pronounced shift of zones where co‐occurrence patterns may play a major role. Main conclusions Our results demonstrate the importance of species co‐occurrence patterns for calibrating improved species distribution models for use in projections of climate effects. The correlation approach is able to localize European areas where inclusion of biotic predictors is effective. The climate‐induced spatial segregation of the major tree species could have ecological and economic consequences.