帽儿山红松人工林鞘翅目成虫群落小尺度空间异质性变化特征

空间异质性是群落空间格局形成的基础,也是物种共存的重要前提。基于地统计空间分析方法,研究20 m×20 m空间尺度上帽儿山红松人工林鞘翅目成虫群落及物种的空间自相关性及异质性分布格局。结果显示:5次调查共捕获鞘翅目成虫10科、42种、2721只个体,群落组成具有中等或较强的空间变异性和明显的时间变异性;Moran's I系数表明群落及部分物种具有显著的空间自相关性,各调查月份接近甚至超过一半的物种不存在空间自相关性;群落及部分物种表现为异质性特征并在特定空间尺度内形成集群,且这种空间分异多由结构性因素或结构性因素和随机性因素共同调控;物种之间表现为具有复杂正的或负的空间作用关系,这种空间关联性的形成主要是结构性因素或随机性因素单一调控的结果,但简单Mantel检验并未发现物种之间存在显著的空间关联性;群落的空间格局梯度分布特征较明显,随着季节的变化(夏天到秋天)聚集物种数量的斑块复杂程度降低。本文旨在揭示鞘翅目成虫空间异质性的变化特征,为鞘翅目成虫群落物种构建机制研究奠定基础。     

丰林典型阔叶红松林地表鞘翅目成虫空间异质性及其与环境因子的空间关联性

土壤动物群落空间异质性及其与环境因子的空间作用关系, 是揭示土壤生态系统格局与过程及生物多样性维持机制的重要基础。作者于2015年生长季节(8月)、寒冷季节(10月)在丰林典型阔叶红松林动态监测样地内, 采用陷阱法调查地表鞘翅目成虫群落, 基于地统计空间分析方法, 揭示步甲科和隐翅虫科群落个体数和物种数及优势种的空间格局, 并分析这些空间格局与土壤含水量和地形因子的空间关联性。两次采样共捕获步甲科成虫26种617只, 隐翅虫科19种222只。8月群落个体数和物种数表现为中等变异, 10月为强变异, 群落组成在两个月间具有显著差异。生长季节(8月)和寒冷季节(10月)步甲科和隐翅虫科群落多表现为中等的空间自相关性, 空间分异由随机性因素和结构性因素共同决定。单个物种的个体数多具有中等的空间异质性特征, 且其空间分异主要由随机性因素和结构性因素共同调控。生长季节群落的个体数、物种数和优势种个体数多形成斑块和孔隙镶嵌分布的空间格局。物种之间及物种与环境因子之间多为复杂的空间关联性, 这些关联性主要受到结构性因素或随机性因素的单一调控。典范对应分析(canonical correspondence analysis, CCA)结果表明, 8月土壤含水量对步甲科和隐翅虫科物种分布影响显著, 10月凹凸度对步甲科分布影响显著, 海拔对隐翅虫科分布具有显著影响。本研究表明地表步甲科和隐翅虫科在生长季节形成明显的空间格局而在寒冷季节空间格局不明显, 为不同尺度地表土壤动物空间异质性和生物多样性维持机制研究提供了理论基础。     

三江平原农田地表和地下土壤螨类丰富度与环境因子的空间关联性

地表和地下土壤动物群落空间格局及其与环境因子的空间作用关系,是揭示地表-地下生态系统格局与过程及生物多样性维持机制的重要基础。于2011年在三江平原农田生态系统,在50m×50m的空间尺度内,基于地统计空间分析方法,揭示地表和地下土壤螨群落及不同螨类物种丰富度的空间格局,并分析这种空间格局与土壤含水量、土壤p H值及大豆株高空间格局的空间关联性。半方差函数和普通克里格插值表明,8月份地表、地下和10月份地下螨群落及这些群落内大部分螨类物种在特定空间尺度内形成集群,表现为空间异质性特征,且这种空间分异多由结构性因素或结构性因素和随机性因素共同调控。交叉方差函数表明,土壤螨群落和不同螨类物种的空间格局与环境因子的空间格局在多种尺度上表现出复杂的空间关联性(正的或负的)。但简单Mantel检验仅发现8月份地表中气门亚目未定种1(Mesostigmata unidentified sp.1)和大豆株高存在明显的正的空间关联性。研究结果表明地下螨群落和生长季节的地表螨群落具有明显的空间异质性结构,地表和地下螨群落及大多数螨类物种丰富度与土壤含水量、土壤p H值及大豆株高的空间关联性并不显著。促进地表-地下生态系统土壤动物群落空间格局研究,为地表-地下格局与过程研究奠定基础。     

贺兰山冲积扇荒漠草地拟步甲群落小尺度空间格局动态

为探讨贺兰山冲积扇荒漠草地拟步甲群落小尺度空间格局动态特征,本研究于2019年5—10月每月采用陷阱法调查200 m×200 m空间尺度的拟步甲昆虫群落,基于地统计分析方法,研究拟步甲昆虫群落的空间格局,并分析空间格局与地形因子的关系。6次调查共捕获拟步甲科成虫7属10种1086只,群落组成具有较明显的空间和时间变异性。其中,拟步甲群落多样性以5月最大,7月最低。Moran I指数表明,拟步甲优势种空间自相关性具有明显的季节波动性,在5月、9月和10月表现为显著空间正相关。拟步甲群落和优势种具有较强的空间异质性,主要由结构性因素决定。普通克里金插值表明,不同季节群落空间格局梯度分布存在明显差异,夏季群落空间格局比较简单。交叉方差函数分析显示,优势种物种间大多表现为正空间关联性,且主要受结构性因素单一调控。典范对应分析(CCA)结果表明,坡度和海拔显著影响昆虫分布。本研究揭示了拟步甲昆虫群落小尺度空间异质性具有明显的季节变异,为研究贺兰山冲积扇生境地表甲虫物种多样性空间格局及维持机制奠定基础。     

阔叶红松林地表鞘翅目多样性及种间关联性

生物间相互作用被认为是群落构建机制的重要驱动力,然而目前不同空间尺度土壤动物群落物种间相互关联性仍不清楚。为了揭示小尺度空间地表鞘翅目成虫的物种组成、多样性及种间关联性,并验证是否亲缘关系较近的同属(科)物种之间多表现为负关联性,本实验于2015年8月、10月在丰林自然保护区9 hm2阔叶红松林固定监测样地,采用陷阱法捕获地表鞘翅目土壤动物。结果表明:(1)两次实验共捕获地表鞘翅目成虫1394只,隶属9科58种,其中步行虫科和隐翅虫科为优势科。10月捕获的个体数量和物种数相较于8月明显减少,群落多样性在8月和10月间具明显差异。(2)8月份,葬甲科、步行虫科和隐翅虫科群落丰富度均为显著的正关联,不同科地表鞘翅目成虫物种之间空间生态位重叠度较低,种间关系多为随机关系;同科内物种之间大多数为随机关系,极少数物种之间表现为显著的正/负的关联性。本研究表明,小尺度空间地表鞘翅目成虫群落组成和多样性在2个月份间具有明显变化,群落内物种间多为随机性关系,亲缘关系较近的物种间并未表现较多显著的负关联性。     

小兴安岭阔叶红松林地表甲虫Metacommunity格局

Metacommunity理论框架为理解生物群落的时空格局及其构建机制提供了一个强有力的方法。然而,少有研究揭示土壤动物metacommunity的格局及其构建机制,小尺度空间的研究更是少见。于2015年分别在凉水和丰林小兴安岭典型阔叶红松林长期动态监测样地内,通过空间直观定位调查监测的方法获得地表甲虫群落,揭示小尺度空间(300 m)地表甲虫metacommunity的格局,并进一步分析地形、土壤因子与这种metacommunity格局的相关性。结果表明:镶嵌型(nestedness)和随机型(random)是小尺度空间地表甲虫metacommunity的常见格局。地表甲虫的metacommunity格局具有类群依赖性,步行虫科和葬甲科多形成镶嵌型格局,而隐翅虫科则仅形成随机型格局。地表甲虫metacommunity格局具有季节波动性,在相对温暖的季节易形成显著的镶嵌型格局,而在相对寒冷的季节多为随机型格局。凉水和丰林地表甲虫的metacommunity格局没有明显差异。地形和土壤因子对地表甲虫metacommunity格局有重要的影响,该影响过程相对复杂,依类群和时间而异。本实验表明镶嵌型和随机型是小尺度空间小兴安岭典型阔叶红松林地表甲虫的常见格局,地形、土壤因子对这些格局的形成有重要的影响。     

人工红松林步行虫(Coleoptera:Carabidae)群落物种共存格局动态分析

揭示群落物种共存格局是群落生态学研究的重点内容之一,零模型的应用极大的促进了群落物种共存格局及其调控机制的进展,然而针对地下生物群落共存格局动态特征的研究并不多见。在帽儿山森林生态站的人工红松林内,通过5次调查取样基于零模型模拟分析小尺度空间(20 m×20 m)步行虫群落物种共存格局的动态特征。结果表明:(1)共捕获步行虫20种,2278只个体,其中Carabus billergi maoershanensis为所有调查季节数量最具优势且分布最广泛的物种,步行虫群落结构具明显的时间变异性;(2)2013年6月步行虫群落为集群性共存格局,而2014年8、10月为竞争性共存格局,基于目前的零模型指标和法则难以准确揭示其他月份是集群性还是竞争性的共存格局,但所有季节的群落均表现为明显的非随机性共存格局,这些共存格局的发现并不完全支持Diamond的群落构建机制理论;(3)所有调查季节均发现很少的显著物种对,基于更严格的检验表明群落中集群性物种对多于隔离性物种对,那些表现为显著的非随机性共存关系的物种对往往是群落内数量较大且分布广泛的优势和常见物种。表明非随机性共存格局可能是帽儿山人工红松林小尺度空间步行虫群落的常见格局,这种非随机性格局具一定的短期动态稳定性,但不同季节这种非随机性共存格局类型表现不同,群落内这些较少的显著物种对可能对群落物种共存格局具有一定的贡献。     

内蒙古高原针茅草原群落土壤水分和碳、氮分布的小尺度空间异质性

用地统计学的方法,研究比较了内蒙古高原4类地带性针茅草原群落,贝加尔针茅（Stipa baicalensis)群落,大针茅（S.grandis)群落,克氏针茅（S.krylovii)群落和小针茅（S.kelemenzii)群落0-20cm土壤水分和碳,氮的小尺度空间异质性特征,结果表明;4类群落土壤水分,有机碳和全氮均表现出显著的小尺度空间结构特征。自相关尺度为1.91m-10.81m,结构性方差占样本方差的35.31%-99.74%。从贝加尔针茅群落到小针茅群落空间自相关的尺度逐渐增大,纹理有逐渐变粗的趋势,土壤水分,碳和氮的小尺度空间格局共同作用于群落的生态学过程,即土壤水分格局→植物种群格局（基本斑块的大小）→土壤碳空间格局→土壤氮空间格局,同时,由于生态学过程的反馈作用,土壤氮空间格局→种群格局→土壤水分格局,土壤属性空间自相关尺度的改变可能是导致群落演替的驱动力,草原退化可能与土壤异质性尺度的改变相关。     

关帝山云杉天然更新与土壤有效氮素异质性的空间关联性

在山西关帝山华北次生林分布区选择云杉针叶林和云杉、杨、桦混交林为研究样地,应用地统计学理论和格局分析方法,分析了该地区云杉天然更新的空间格局及结构特征;运用地统计分析中的变异函数分析方法定量分析了样地土壤氮素的空间异质性特征.采用GIS叠加分析和统计分析方法,定量评价云杉更新苗分布格局和土壤有效氮素空间异质性之间的空间关联程度,以了解森林土壤氮营养生境的异质性对云杉幼苗更新的影响.结果表明：1）在云杉针叶林中,云杉更新苗分布呈空间斑块状格局,且表现出自相关空间变异特征;在云杉、杨、桦混交林中,云杉更新苗空间分布格局呈集聚分布特征,但表现为随机性的空间变异.2）云杉针叶林土壤氮素含量相对低,有效氮含量空间异质性特征明显,空间结构性变异所占比例较大;混交林土壤氮素含量相对高,有效氮呈随机性变异特征.3）在土壤氮素含量相对低的云杉针叶林中,云杉更新苗分布格局与土壤有效氮异质性之间存在较明显的空间关联性,在土壤NH₄⁺-N含量相对高的斑块中,云杉更新苗发生数量多.在土壤氮素含量相对高的混交林,云杉更新苗空间分布与土壤有效氮异质性之间的空间关联性不明显.     

土壤动物群落空间格局和构建机制研究进展

群落空间格局和构建机制一直是生态学研究的核心内容。在生物多样性严重丧失的背景下, 揭示群落空间格局及其构建机制, 有助于深刻理解生物多样性丧失的原因, 更有助于应对生物多样性保护等重大生态环境问题。然而, 陆地生态系统的研究多集中于地上生物群落, 对地下生态系统, 尤其是土壤动物空间格局和构建机制的研究尚不充分。事实上, 土壤动物多样性是全球生物多样性的关键组成之一, 是地下生态系统结构和功能维持的重要部分。对土壤动物空间格局和构建机制的研究, 能明确不同空间尺度条件下土壤动物多样性的维持机制。土壤动物群落常在多种空间尺度形成复杂的空间分布格局, 因此, 本文首先介绍了不同空间尺度主要土壤动物群落的空间自相关性特征, 阐述了土壤动物群落斑块和孔隙镶嵌分布的复杂空间格局。继而阐明这种空间格局主要受生物间作用、环境过滤和随机扩散的调控, 并说明这三个过程对土壤动物群落的调控能力和作用方式。作者提出, 这三个过程仍是今后土壤动物群落空间格局和构建机制研究的重点内容, 需要进一步加强以土壤动物为研究对象的群落构建理论的验证和发展。我国土壤动物群落空间格局和构建机制起步较晚, 希望本文能够促进我国土壤动物生态学相关领域的研究。     

Characterizing the spatial structures of riparian plant communities in the lower reaches of the Heihe River in China using geostatistical techniques

Geostatistical techniques are used to evaluate spatial characteristics of riparian plant communities that were mapped within a portion of the lower reaches of the Heihe River. The spatial structures of different plant communities were characterized using nugget, range and sill parameters of spherical or exponential model variograms. Model variograms revealed different plant communities with distinctive spatial properties that were quantified effectively by the parameters used in these models. Model variograms were fit to experimental variograms calculated from diameter at breast height and plant cover data collected for three 2,000 m × 5 m, nine 500 m × 5 m, and three 200 m × 2 m transects. The 2,000 m × 5 m transects were oriented perpendicular to the river and the other transects were located in such a way as to sample communities dominated by different plants. Riparian plant communities in the lower reaches of the Heihe River show patterns of patchiness, and the plant communities are mainly dominated by mature Populus euphratica, young P. euphratica, Tamarix chinensis, and Sophora alopecuroides. The maximum model variogram range for all communities was 55 m, suggesting that transects must be at least this long in order to be able to classify them unambiguously into recognizable communities. Experimental variograms were calculated for all plants in each of four mapped plant communities. The results show that the T. chinensis and S. alopecuroides dominated plant communities have distinctively large ranges, and the spatial structures of P. euphratica dominated plant communities show multiscale change. In the 2,000 m × 5 m transects, the spatial structures of riparian plant communities show a distinct change at the scale of 430 m, because this scale also is the scale of vegetation patchiness turnover, and at scales of more than 430 m the spatial structure of the community increases significantly. These findings confirm the potential of using high-resolution remote sensing data and geostatistics for determining the vegetation community structures of riparian plant communities.     

Spatial pattern analysis of entomopathogenic and other free‐living nematodes at landscape scales

Spatial patterns of entomopathogenic and other free‐living nematode populations were analyzed at the landscape scale. Free‐living bacterial feeder populations can be classified into trophic groups based on their functional and life history characteristics. Differences in life history traits were hypothesized to result in different spatial structures of populations. Spatial autocorrelations (Moran's I index) and exponential variogram models were calculated and estimated for each trophic group, including entomopathogenic nematodes, and comparisons were made among them. Spatial autocorrelation results showed that the spatial structure of entomopathogenic nematode populations is similar to that of relatively r‐selected free‐living bacterial feeders, which share several life history traits with entomopathogenic nematodes.     

庐山香果树毛竹混交林空间格局研究

为了解庐山毛竹入侵对香果树的影响,采用Ripley’sK(t)函数分析香果树毛竹混交林群落空间分布格局和种间关联关系。结果表明:香果树幼树在较小的尺度上呈聚集分布,较大尺度上随机分布直至均匀分布;香果树成年树在较小的尺度上呈随机分布,较大尺度上聚集分布;毛竹在所有的研究尺度上为显著聚集分布,且随着尺度的增大,聚集强度增加。毛竹与香果树幼树在较小的尺度上没有明显关联性,在较大的尺度上呈显著负相关;毛竹与香果树成年树在一系列尺度上均表现为显著负相关。根据空间关联性分析结果,应从幼龄起对生长有香果树的森林群落进行必要的管理,伐除毛竹及一些影响较大的其它乔木树种,挖除地下竹鞭,为香果树提供良好的生长环境,进而恢复其种群的天然合理配置。否则,香果树这一珍贵的植物资源在庐山将有灭绝的危险。     

黑龙江省不同纬度梯度农田大型土壤动物群落分布特征

沿着由高到低的纬度梯度,分别在塔河、带岭、帽儿山的农田生境选择研究样地,采用手捡法调查不同纬度农田生态系统大型土壤动物的群落组成、水平结构、垂直结构和多样性特征,并运用双变量相关分析及典范对应分析阐明土壤动物群落和环境因子的相互关系,旨在揭示黑龙江省不同纬度梯度农田大型土壤动物群落分布特征及其影响因素。调查共捕获大型土壤动物35类2339只,隶属于2门6纲14目35科。其中线蚓科(Enchytraeidae)、正蚓科(Lumbricidae)、步甲科(Carabida)、隐翅虫科(Staphylinidae)为优势类群,其个体数占总个体数的58.84%;常见类群为蚁科(Formicidae)、蜘蛛目(Araneida)和地蜈蚣目(Geophilomorpha)等8类,其个体数占总个体数的32.79%。结果表明:(1)水平分布上:大型土壤动物个体密度和丰富度(即类群数)的水平分布均表现为帽儿山带岭塔河,单因素方差分析表明大型土壤动物的个体密度和丰富度在不同纬度地区无显著差异。Shannon-wiener多样性指数(H')和Pielou均匀度指数(E)均为帽儿山带岭塔河;Simpson优势度指数(C)表现为塔河和带岭最高,帽儿山最少;Margalef丰富度指数(D)则是塔河最多,其次为帽儿山和带岭。(2)垂直分布上:3个不同纬度样地的农田土壤动物个体密度和丰富度在同垂直层次间不存在明显差异(P0.05)。除帽儿山5—10 cm土层土壤动物类群数量较表层增加之外,其他样地大型土壤动物个体密度和丰富度均随土层深度的增加而逐渐降低,表聚性明显。(3)与土壤环境因子关系上:双变量相关分析表明,不同纬度农田大型土壤动物的类群数、个体密度、多样性指数、均匀度指数和优势度指数与土壤p H值、有机质、全氮、全磷和速效磷均没有显著的相关性;典范对应分析(CCA)进一步表明,优势类群和常见类群对环境因子具有较强的适应能力,广泛的分布在3个样地内。研究表明,农田生态系统大型土壤动物的类群数、个体密度和多样性指数随纬度梯度的增加先增加后减少,但不同纬度间均无显著性差异。不同土壤动物类群受到环境变量的影响程度不同,在局地尺度环境因子对土壤动物的影响不容忽视。研究为区域尺度农田生态系统土壤动物空间格局和生物多样性维持机制研究奠定基础。     

Bumblebees experience landscapes at different spatial scales: possible implications for coexistence

Coexistence in bumblebee communities has largely been investigated at local spatial scales. However, local resource partitioning does not fully explain the species diversity of bumblebee communities. Theoretical studies provide new evidence that partitioning of space can promote species coexistence, when species interact with their environment at different spatial scales. If bumblebee species possess specific foraging ranges, different spatial resource utilisation patterns might operate as an additional mechanism of coexistence in bumblebee communities. We investigated the effects of the landscape-wide availability of different resources (mass flowering crops and semi-natural habitats) on the local densities of four bumblebee species at 12 spatial scales (landscape sectors with 250–3,000 m radius) to indirectly identify the spatial scales at which the bumblebees perceive their environment. The densities of all bumblebee species were enhanced in landscapes with high proportions of mass flowering crops (mainly oilseed rape). We found the strongest effects for Bombus terrestris agg. and Bombus lapidarius at large spatial scales, implying foraging distances of 3,000 and 2,750 m, respectively. The densities of Bombus pascuorum were most strongly influenced at a medium spatial scale (1,000 m), and of Bombus pratorum (with marginal significance) at a small spatial scale (250 m). The estimated foraging ranges tended to be related to body and colony sizes, indicating that larger species travel over larger distances than smaller species, presumably enabling them to build up larger colonies through a better exploitation of food resources. We conclude that coexistence in bumblebee communities could potentially be mediated by species-specific differences in the spatial resource utilisation patterns, which should be considered in conservation schemes.     

Spatial patterns of species and plant traits in response to 20 years of grazing exclusion in subalpine grassland communities

Question: Does long‐term grazing exclusion affect spatial patterns of canopy height, plant species and traits in subalpine grassland communities? Are spatial patterns of species and traits similarly affected by grazing exclusion? Are changes in spatial patterns of species associated with changes in species abundances? Location: Subalpine grasslands, Vercors and Oisans Mountains, Alps (France). Methods: Spatial sampling of vegetation and measurements of plant traits were carried out within nine pairs of grazed and ungrazed 10 m × 10‐m plots in three species‐rich communities with different productivities. We estimated within‐plot spatial patterns of canopy height, species and aggregated trait values by measuring the extent (or patch size) and intensity of spatial dependence with Moran's I. Abundance‐weighted averages for species patch size and intensity of spatial dependence were calculated across all species per plot and across species per life form. Such measures derived from analysis of spatial dependence were considered spatial traits. Results: Response of spatial patterns to grazing exclusion was only detected in patch size, whereas intensity of spatial dependence was not affected. Changes in spatial patterns were community‐dependent because spatial traits based on patch size of canopy height and species increased following grazing exclusion only in the less productive community. Unexpectedly, changes in spatial patterns of species did not support changes in spatial patterns of trait values. Abundances and patch sizes of several life forms were significantly affected by grazing exclusion. However, at the scale investigated, changes in abundance of life forms did not correspond to changes in their spatial patterns and vice versa. Conclusion: In species‐rich communities, grazing alters spatial spread of species (i.e. patch size) rather than intra‐specific aggregation (i.e. intensity of spatial dependence). Results revealed possible mechanisms of species spatial reorganisation that are independent of abundance variation. Therefore, it is important to consider changes in spatial patterns in addition to changes in mean values of vegetation features when assessing impacts of grazing management.     

Spatial patterns and associations in a Quercus‐Betula forest in northern China

Question: Are species‐specific regeneration strategies and competition the dominant processes facilitating species coexistence in a Quercus liaotungensis dominated temperate deciduous forest? Location: Dongling Mountains, North China, 1300 m a.s.l. Methods: Ripley's K‐function was used to characterize the spatial patterns and spatial associations of two dominant tree species, Quercus liaotungensis and Betula dahurica, and a common subcanopy species, Acer mono, at different growth stages (adult, sapling, seedling). Results: Seedlings, saplings and adults of all three species exhibited clumped distributions at most spatial scales. Quercus seedlings and saplings were positively associated with conspecific adult trees and spatially independent of dead trees suggesting that seed dispersal and vegetative regeneration influenced the spatial patterning of Quercus trees. Betula seedlings and saplings were positively associated with both live and dead trees of conspecific adults at small scales (<5 m) but negatively associated with live and dead trees of other species indicating sprouting as an important mechanism of reproduction. Saplings of Acer had a strong spatial dependence on the distribution of conspecific adult trees indicating its limited seed dispersal range. Negative associations between adult trees of Betula and Quercus demonstrated interspecific competition at local scales (<5 m). Conclusions: Different regeneration strategies among the three species play an important role in regulating their spatial distribution patterns, while competition between individuals of Betula and Quercus at the adult stage also contributes to spatial patterning of these communities. The recruitment limitations of Betula and Quercus may affect the persistence of these species and the long‐term dynamics of the forest.