群落构建研究的新进展:进化和生态相结合的群落谱系结构研究

群落如何构建足群落生态学中的重要问题.群落谱系结构研究将物种间的亲缘进化关系运用到群落生态学研究中,利用物种的系统发育状况推测历史因素对现有群落的影响,为推断影响群落组成的生态学机制提供了有效方法.群落谱系结构的研究方法是首先建立可代表群落物种库的超级系统进化树,然后计算群落内物种间的谱系距离,最后通过统计方法检测其与随机模型下的谱系距离是否有显著差异来获得谱系结构(如谱系聚集、谱系发散),从而揭示群落构建中的关键生态过程(如生境过滤、竞争作用).群落谱系结构与空间尺度、分类群尺度、时间尺度等不同研究尺度有关.在小的空间尺度下,随着分类群尺度降低、树木年龄级增大,群落谱系结构从聚集逐渐转为发散;而随群落空间尺度的增大,谱系趋向于聚集.谱系结构受到环境因素影响,因此分析集合群落下的谱系可以揭示区域生态过程的影响.另外,群落谱系结构研究还有助于探讨中性理论、密度制约假说等生态学理论,并预测干扰作用下的群落演化趋势.在利用谱系结构深入探讨群落构建成因时,需要基于生态特征和环境变量共同分析,同时考虑小尺度局域过程(群落的微环境或群落内种间相互作用等)和大尺度区域过程(地史过程和物种形成等),并可结合生态控制实验,以确认群落构建的关键因素.在研究方法和手段上,今后需要注重通过选择合适的基因片段建立系统树,然后通过生态特征来加以校正,以更准确地反映物种间的亲缘距离.另外,获得谱系树后还需要寻找更加合理的统计模型和指数,增加统计分析和解决问题的能力.     

后河自然保护区珍稀植物群落谱系结构的时空变化

群落中所有物种对的亲缘关系程度构成群落的谱系结构。研究群落的谱系结构可以有效地揭示环境过滤、竞争排斥、中性过程三大生态过程对群落构建的相对重要性。本文以湖北后河自然保护区1hm。固定样地的珍稀植物群落为研究对象,分别研究了5m×5m、10m×10m、20m×20m、30m×30m、40m×40m、50m×50m6个空间尺度和在10m×10m、20m×20m、30m×30m、40m×40m、50m×50m5个空间尺度下5cm≤DBH〈10cm、10cm≤DBH〈15cm、15cm≤DBH〈20cm、DBH≥20cm4个径级对亚热带常绿落叶阔叶混交林群落谱系结构的影响。结果发现：随着空间尺度的增加,谱系结构逐渐趋向聚集。随着径级的增大,群落谱系聚集程度越来越小。但当DBHI〉20cm时,群落谱系聚集度较高。在25m。的尺度上群落谱系结构表现为随机,而在100m^2及更大的尺度上群落谱系结构都表现为聚集,说明在亚热带常绿阔叶落叶混交林的群落构建过程中,小尺度上中性过程可能起主导作用,而大尺度上生态位过程可能更重要。     

不同尺度上环境因子对常绿阔叶林群落的谱系结构的影响

群落的谱系结构是反映作用于群落组成的各种生态过程的综合指标。通过研究群落的谱系结构, 能够有效地推断形成群落谱系结构的生态过程。该文从环境因子(海拔、地形、pH、土壤湿度和土壤元素等)对群落谱系结构的影响这一视角出发, 采用圆形随机取样, 在半径为5、25、50、75和100 m 5个尺度上, 深入研究了古田山24 hm²永久样地群落的谱系结构。研究发现: 古田山样地在所有研究尺度上(半径 > 5 m)都表现为谱系聚集, 随着尺度的增加, 群落的聚集程度呈现出先增加后下降的格局。采用多元线性回归模型分析环境因子对群落谱系结构的影响时发现: 随着尺度的增加, 环境因子对群落谱系结构的影响逐渐增强; 在小尺度上, 环境因子对谱系结构没有显著影响, 这可能是由于小尺度上近缘种之间较强的竞争排斥作用所致。当尺度达到100 m半径时, 高海拔区域表现出谱系发散格局, 而低海拔区域保持谱系聚集, 这可能是由于古田山样地海拔较高的地带生境异质性较强和较大的干扰所致。该研究在所有的尺度上都表现出显著的谱系聚集, 与中性理论所预测的群落是物种的随机组合、无谱系结构相反, 因而为生态位理论提供了证据。     

不同空间尺度西藏色季拉山急尖长苞冷杉群落谱系结构特征

基于西藏色季拉山4 hm²亚高山暗针叶林固定样地调查数据,分析了急尖长苞冷杉群落内6个径级(1～2、2～4、4～7、7～11、11～16、>16 cm)的植物在5个空间尺度(5 m×5 m、10 m×10 m、20 m×20 m、50 m×50 m、100 m×100 m)上的谱系结构。结果表明: 不同空间尺度上,该群落的净谱系亲缘关系指数(NRI)和净最近种间亲缘关系指数(NTI)值均大于零,整体上表现为谱系聚集。在小空间尺度(5 m×5 m)上,谱系结构聚集强度最大。在所有空间尺度上,群落内小径级个体表现为谱系聚集,随着径级的增大(DBH>7 cm),逐渐表现为谱系发散,且发散程度随径级增加而增大。小径级个体的谱系聚集强度随空间尺度而增大;而大径级个体(DBH>7 cm)的谱系发散程度随空间尺度增加而增大。小径级群落的生境过滤和大径级群落的竞争排斥作用可能是驱动该区域急尖长苞冷杉群落构建的主要生态学过程。     

山西霍山植物群落谱系结构的空间格局

以山西霍山七里峪植物群落为研究对象,用R语言分别对9个海拔梯度(1 200~2 000m)、5个空间尺度(100、400、900、1 600和2 500m2)和5个空间尺度下划分的6个径级(Ⅰ.DBH5cm、Ⅱ.5cm≤DBH10cm、Ⅲ.10cm≤DBH15cm、Ⅳ.15cm≤DBH20cm、Ⅴ.20cm≤DBH25cm、Ⅵ.DBH≥25cm)的植物群落谱系结构及其在不同研究方向上的变化规律进行分析,探讨其群落构建成因。结果显示:(1)在所有海拔梯度上,霍山七里峪植物群落都表现出一定的谱系结构,在不同海拔梯度上,群落谱系结构表现出一定的差异性,且在中低海拔群落的谱系结构是发散的,在较高海拔群落的谱系结构是聚集的。(2)5个空间尺度下研究区植物群落都具有谱系结构,且随着空间尺度的增加,谱系聚集程度越来越大。(3)不同径级下的群落都有一定的谱系结构,随着径级的增大,谱系聚集程度逐渐降低,趋向于发散结构,但在Ⅵ径级时群落聚集程度较Ⅳ、Ⅴ径级的高。研究表明,山西霍山七里峪植物群落在不同海拔梯度、不同空间尺度和不同径级下都表现出显著的谱系结构,与中性理论所预测的谱系随机并不相符,证明生态位理论在霍山植物群落构建过程中的作用更加重要。     

放牧干扰梯度下川西亚高山植物群落的组合机理

为了阐明放牧干扰对川西亚高山区域植物群落的组合过程以及群落结构的影响, 研究了放牧干扰梯度下的功能群均匀度和群落谱系结构的变化趋势。结果显示: 在干扰较轻的阔叶林与针叶林样地, 部分样方的功能群均匀度显著高于无效模型, 随着干扰梯度的增强, 功能群均匀度呈线性下降, 样方平均值从0.930降至0.840, 其高于无效模型的次数也逐渐降低, 干扰程度较大的草甸中出现部分样方的功能群均匀度显著低于无效模型。随着干扰程度的增强, 群落的谱系结构指数也呈逐渐上升趋势, 净关联指数平均值由-0.634逐渐增加至2.360, 邻近类群指数由-0.158上升至2.179。草甸与低矮灌丛受干扰较为严重, 其大部分样方的谱系结构指数显著高于随机群落, 表明干扰群落的谱系结构呈聚集分布。功能群均匀度与谱系结构的变化趋势一致, 表明生境筛滤效应与种间竞争作用的平衡决定着群落的组合过程。干扰降低了竞争作用, 促进了少数耐干扰功能群的优势地位, 造成功能群均匀度下降, 同时通过生境筛滤作用, 使群落的谱系结构呈现出聚集分布; 而未干扰的群落中由于竞争作用的效应, 功能群均匀度较高, 谱系结构也更加分散。研究区域植物群落的功能群均匀度与物种丰富度呈负相关, 表明物种间特别是相似物种间的竞争限制了群落的物种多样性。研究结果说明, 生态位分化和物种间的相互竞争在物种共存与群落组合中具有重要作用。     

边缘效应的空间尺度与测度

综述了边缘效应的空间尺度类型以及在不同尺度上的测度方法.基于大量的研究整合,认为边缘效应空间尺度的划分,可以根据空间尺度的不同以及边缘效应形成和维持因素,分为大中小3个尺度类型,即大尺度的生物群区交错带、中尺度的景观类型之间的生态交错带和小尺度的斑块(生态系统)之间的群落交错区.大尺度主要是以植被气候带为标志的生物群区间的边缘效应,这种地带性的交错区主要受大气环境条件的影响.中尺度类型主要包括城乡交错带、林草交错带、农牧交错带等类型,是不同生态系统要素的空间交接地带,在物质能量等相互流动的作用下变得更为复杂.小尺度水平上是指斑块之间的交错所形成的边缘效应,受小地形等微环境条件及生物非生物等因子的制约,研究主要集中在群落边缘、林窗边缘和林线交错带等方面.对边缘效应测度的定量化研究有助于更加深入理解边缘效应.在大尺度水平上,边缘效应测度的研究主要是应用数量生态学等方法,研究不同气候带之间界线的划分及其物种分布的梯度规律性.中尺度水平上应用景观生态学的3S技术等方法,侧重于研究交错带的动态变化趋势及位置宽度的判定.小尺度水平上通过对距离边缘的长度,各群落中种群的数量、结构、多样性等定量指标的测定来构建测度公式,从而对边缘效应的强度进行量化,并反映边缘对群落的正负效应.总体上看,主要集中于中小尺度上,未来应该强化大尺度边缘效应测度的研究.     

山西灵空山刺五加种群空间分布格局及种间空间关联性

以山西灵空山自然保护区刺五加(Acanthopanax senticosus(Rupr.Maxim.)Harms)种群为对象,采用空间点格局方法中的Ripley’s K函数,对刺五加种群不同龄级的空间分布格局及其与主要灌木物种间的空间关联性进行研究,并对种群空间分布格局进行可视化解析。结果显示,刺五加种群中幼年个体数量多,径级结构呈金字塔型,为增长型种群,能够实现持续更新;刺五加种群的径级Ⅰ在小尺度上呈聚集分布,随着尺度增大出现随机分布;径级Ⅱ、Ⅲ、Ⅳ在全尺度上呈聚集分布,径级Ⅴ、Ⅵ在小尺度上为聚集分布,随着龄级和空间尺度的增大,刺五加种群聚集强度渐弱,呈现随机分布;刺五加种内关联在全尺度上呈正相关,但与群落中主要灌木在小尺度上呈不相关或负相关,在大尺度上呈负相关。研究表明刺五加的生长条件较为适宜,并可以通过相互之间的庇护作用,减弱种内竞争,但与其他树种间存在强烈竞争,生存现状较差,在群落中处于劣势。     

不同程度人为干扰对古田山森林群落谱系结构的影响

群落谱系结构包含群落发育的历史信息,能从新的角度反映群落形成的生态过程.作者在浙江古田山自然保护区亚热带常绿阔叶林中选择了人为干扰强度不同的4种群落类型,以20 m×20 m为研究尺度探讨了不同干扰程度对群落谱系结构的影响.结果表明,人工林(类型Ⅰ)谱系结构发散;但间伐林(类型Ⅱ)、自然恢复林(类型Ⅲ)以及自然老龄林(类型Ⅳ)谱系结构聚集,且以类型Ⅱ和Ⅳ聚集度最高.进一步分析不同径级谱系结构发现,在中小径级(DBH≤5 cm和5 cm10 cm时,除了类型Ⅳ,其他3种群落都是谱系结构发散,这反映了在恢复早期种子扩散对这些林型群落构建影响较大:而皆伐后的演替和间伐增加了群落生境异质性,生境过滤作用增强,使类型Ⅱ、Ⅲ群落中小径级谱系结构表现为聚集;类型Ⅳ群落不同径级谱系结构均表现为聚集,可能与其稳定的生境过滤作用有关.     

浙江省典型天然次生林主要树种空间分布格局及其关联性

松阔混交林和常绿阔叶林是亚热带地区天然次生林代表性的森林类型,通过研究主要树种分布变动趋势,可以有效地指导森林经营措施。在该地区4个1hm~2典型样地,在0—30m尺度范围内综合分析了主要树种的空间分布格局及种间关联性,并对同一树种分发育阶段在不同森林群落中的种群空间分布格局进行了比较,以探讨亚热带地区天然次生林群落空间格局形成和种群维持机制。样地1和2为松阔混交林,其中样地1的马尾松密度较低,样地3和4为常绿阔叶林。研究结果表明:(1)以完全随机模型为零假设时,样地1的主要种群在小尺度(10 m)呈聚集分布,随尺度增加呈随机分布;样地2—4的主要种群在所有尺度呈聚集分布,随尺度增加聚集强度逐渐减弱;以异质泊松模型为零假设时,4个样地的主要种群在大部分尺度呈随机分布;(2)青冈和苦槠的小树(5.0 cm≤DBH10.0 cm)在4样地的所有尺度以聚集分布为主,大树(DBH≥10.0 cm)在松阔混交林样地呈随机分布趋势,但在常绿阔叶林样地青冈在0—20 m尺度、苦槠在所有尺度呈聚集分布;(3)松阔混交林中建群种马尾松和其他树种的种间关联性,在样地1的小尺度为负相关,随尺度增加为不相关,在样地2的所有尺度为负相关;常绿阔叶林中建群种青冈和其他树种的种间关联性在在样地3的小尺度为负相关,随尺度增加为不相关,在样地4的所有尺度为负相关;所有伴生树种的种间关联性以负相关为主。结果说明,种群空间分布格局及其关联性随群落结构和空间尺度的不同而出现变化,在松阔混交林和常绿阔叶林群落格局形成中除了扩散限制和生境异质性以外,密度制约机制在松阔混交林中发挥了重要作用,而在常绿阔叶林中其作用随着树木生活史阶段的提高而减弱。     

Foreign exploration for Scirtothrips perseae Nakahara (Thysanoptera: Thripidae) and associated natural enemies on avocado (Persea americana Miller)

Scirtothrips perseae Nakahara was discovered attacking avocados in California, USA, in 1996. Host plant surveys in California indicated that S. perseae has a highly restricted host range with larvae being found only on avocados, while adults were collected from 11 different plant species. As part of a management program for this pest, a “classical” biological control program was initiated and foreign exploration was conducted to delineate the home range of S. perseae, to survey for associated natural enemies and inventory other species of phytophagous thrips on avocados grown in Mexico, Guatemala, Costa Rica, the Dominican Republic, Trinidad, and Brazil. Foreign exploration efforts indicate that S. perseae occurs on avocados grown at high altitudes (>1500 m) from Uruapan in Mexico south to areas around Guatemala City in Guatemala. In Costa Rica, S. perseae is replaced by an undescribed congener as the dominant phytophagous thrips on avocados grown at high altitudes (>1300 m). No species of Scirtothrips were found on avocados in the Dominican Republic, Trinidad, or Brazil. In total, 2136 phytophagous thrips were collected and identified, representing over 47 identified species from at least 19 genera. The significance of these species records is discussed. Of collected material 4% were potential thrips biological control agents. Natural enemies were dominated by six genera of predatory thrips (Aeolothrips, Aleurodothrips, Franklinothrips, Leptothrips, Scolothrips, and Karnyothrips). One genus each of parasitoid (Ceranisus) and predatory mite (Balaustium) were found. Based on the results of our sampling techniques, prospects for the importation of thrips natural enemies for use in a “classical” biological control program in California against S. perseae are not promising.     

Molecular phylogeny of Phebalium (Rutaceae: Boronieae) and related genera based on the nrDNA regions ITS 1+2

Parsimony analyses of the internal transcribed spacer regions of nuclear ribosomal DNA (ITS 1 & ITS 2) for 38 taxa sampled from the Phebalium group (Rutaceae: Boronieae) and two outgroups confirm that, with the exception of Phebalium sensu stricto and Rhadinothamnus, six of the currently recognised genera within the group are monophyletic. The data indicate that Phebaliums. str. is paraphyletic with respect to Microcybe, and Rhadinothamnus is paraphyletic with respect to Chorilaena. Rhadinothamnus and Chorilaena together are the sister group to Nematolepis. Drummondita, included as an outgroup taxon, clustered within the ingroup as sister to Muiriantha and related to Asterolasia.The phylogeny suggests that the evolution of major clades within a number of these genera (e.g. Phebalium) relates to vicariance events between eastern and south-western Australia. Leionema is an eastern genus, with the most basal taxon being the morphologically distinct Leionema ellipticum from northern Queensland. Leionema also includes one species from New Zealand, but this species (as with some others) proved difficult to sequence and its phylogenetic position remains unknown. Taxonomic changes at the generic level are recommended.The authors wish to thank Paul G.Wilson, PERTH, for advice and discussion, and Paul Forster, BRI, for collecting and providing material of Leionema ellipticum. The project was supported by a Melbourne University Postgraduate Award (to BM), the Australian Biological Resources Study (ABRS), Australian Systematic Botany Society and Wolf Den (Australia) Investments.     

Entwicklungsgeschichte und Ultrastruktur von Pollenkitt und Exine bei nahe verwandten entomophilen und anemophilen Angiospermensippen derAlismataceae,Liliaceae, Juncaceae,Cyperaceae, Poaceae undAraceae

Some closely related members of the monocotyledonous familiesAlismataceae, Liliaceae, Juncaceae, Cyperaceae, Poaceae andAraceae with variable modes of pollination (insect- and wind-pollination) were studied in relation to the ultrastructure of pollenkitt and exine (amount, consistency and distribution of pollenkitt on the surface of pollen grains). The character syndromes of pollen cementing in entomophilous, anemophilous and intermediate (ambophilous or amphiphilous) monocotyledons are the same in principal as in dicotyledons. Comparing present with former results one can summarize: 1) The pollenkitt is always produced in the same manner by the anther tapetum in all angiosperm sub-classes. 2) The variable stickiness of entomophilous and anemophilous pollen always depends on the particular distribution and consistency of the pollenkitt, but not its amount on the pollen surface. 3) The mostly dry and powdery pollen of anemophilous plants always contains a variable amount of inactive pollenkitt in its exine cavities. 4) A step-by step change of the pollen cementing syndrome can be observed from entomophily towards anemophily. 5) From the omnipresence of pollenkitt in all wind-pollinated angiosperms studied one can conclude that the ancestors of anemophilous angiosperms probably have been zoophilous (i.e. entomophilous) throughout.

     

Taxonomic status of the species of the green algal genusNeochloris

Komárek has recently reviewed the various species assigned to the green algal genusNeochloris Starr (Chlorococcales, Chlorococcaceae) and removed those with uninucleate vegetative cells to a new genus,Ettlia. Watanabe & Floyd, unaware ofKomárek's work, also reviewed the species ofNeochloris and distributed them among three genera—Neochloris, Chlorococcopsis gen. nov., andParietochloris gen. nov.—on the basis of details of the covering of the zoospore and the arrangement of the basal bodies of the flagellar apparatus. This paper reconciles these two treatments and makes additional recommendations at the ranks of genus, family, order, and class.     

New species and new combinations in Peruvian Orchidaceae

seventeen new species and combinations are proposed in the generaChondrorhyncha, Cischweinfia, Cochlioda, Eloyella, Encyclia, Kefersteinia, Koellensteinia, Macroclinium, Rodriguezia, Solenidiopsis, andStenia. All new species are illustrated. A key is provided for 2-flowered species ofMacroclinium, PeruvianSigmatostalix, and PeruvianStenia. Solenidium (Solenidiopsis) peruvianum Schltr. is lectotypified.     

安徽牯牛降的丝孢菌Ⅰ

本文报导作者采自安徽枯牛降自然保护区的18种丝孢菌,分属于5个属,其中有3个新种:牛皮冻生尾孢(Cercospora paederiicola),山鸡椒假尾饱(Pseudocercospora litseae-cubebae),鸡血藤生假尾孢(P. millettiicola)和2个中国新纪录。文中对新种进行了描述及绘图,新记录种作了简要说明。研究的标本保存在中国科学院微生物研究所真菌标本室(HMAS)。     

Changes of dung beetle communities from rainforests towards agroforestry systems and annual cultures in Sulawesi (Indonesia)

Little is known about how tropical land-use systems contribute to the conservation of functionally important insect groups, including dung beetles. In a study at the margin of Lore Lindu National Park (a biodiversity hotspot in Central Sulawesi, Indonesia) dung-beetle communities were sampled in natural forest, young secondary forest, agroforestry systems (cacao plantations with shade trees) and annual cultures (maize fields), each with four replicates (n = 16 sites). At each site we used 10 pitfall traps, baited with cattle dung, along a 100 m transect for six 3-day periods. The number of trapped specimens and species richness at the natural forest sites was higher than in all land-use systems, which did not significantly differ. Each land-use system contained, on average, 75% of the species richness of the natural forest, thereby indicating their importance for conservation. However, a two-dimensional scaling plot based on NESS indices (m = 6) indicated distinct dung beetle communities for both forest types, while agroforestry systems and annual cultures exhibited a pronounced overlap. Mean body size of dung beetles was not significantly influenced by land-use intensity. Five of the six most abundant dung beetle species were recorded in all habitats, whereas the abundance of five other species was significantly related to habitat type. Mean local abundance and number of occupied sites were closely correlated, further indicating little habitat specialisation. The low dung beetle diversity (total of 18 recorded species) may be due to the absence of larger mammals in Sulawesi during historical times, even though Sulawesi is the largest island of Wallacea. In conclusion, the dung beetle fauna of the lower montane forest zone in Central Sulawesi appears to be relatively robust to man-made habitat changes and the majority of species did not exhibit strong habitat preferences.     

Die GattungKarschia — Bindeglied zwischen bitunicaten Ascomyceten und lecanoralen Flechtenpilzen?

The genusKarschia, in the earlier sense, including saprophytes and parasites on lichens, has been thought to be a non-lichenized parallel genus of the lichen genusBuellia. Modern workers included it on the one hand inBuellia, on the other hand combined it with bitunicate ascomycetes. It is now proved thatKarschia is heterogeneous and contains but superficially similar members both of the genusBuellia of theLecanorales and of typical or masked bitunicateAscomycetes. Therefore, it can not be regarded as a link betweenLecanorales andDothideales. The type species ofKarschia belongs to theDothideales.