浅析物种概念的演变历史

本文是一篇关于物种概念演变的简述。生物学家用不同的方法或标准划分物种, 就形成了不同的物种概念, 如生物学物种、形态学物种、生态学物种、进化物种、系统发生或支序物种, 或它们的组合, 等等。它们都揭示了物种属性的特定侧面, 都是不同物种客观存在的真实反映, 但都无法令所有人满意。对真核生物来说, 无论它们在形态上的差别有多大, 生殖隔离(不能产生可育的后代)应该是两个群体能否真正分化成不同物种的关键, 这种隔离机制可以是地理的、行为的或其他方式; 而生殖隔离总会伴随着一些形态或遗传上的变化, 虽然这些特征可能与生殖隔离本身并无多大关系, 但往往成为分类学家或分子进化生物学家区分种的依据,对已经灭绝的化石物种来说, 生殖隔离的物种划分方式就无能为力了。如何准确定义一个物种依然充满着矛盾, 因为基于生殖隔离的物种概念不实用, 而实用的物种概念(如形态学物种)又被认为是人为的。     

生物多样性事业需要科学、可操作的物种概念

物种概念(species concept)是生物学家们持续关注的中心问题。物种概念决定物种划分, 而物种划分的合理性关系到生物多样性的研究、保护和可持续利用。本文把现有较流行的物种概念分为6类, 并对它们予以述评后指出: 虽然生物学物种概念、遗传学物种概念、进化物种概念、系统发生物种概念等从不同方面认识了物种的客观真实性和物种的本质, 但在实践中都难以操作。绝大多数物种是由分类学家划分的, 但目前所有的分类学物种概念都包含有不同程度的主观因素, 从而造成物种划分的人为性, 对生物多样性研究造成负面影响。因此, 生物多样性事业需要科学、可操作的物种概念。本文在吸收了生物学物种概念、遗传学物种概念、进化物种概念以及系统发生物种概念等的长处, 也分析了它们的不足和问题的基础上提出一个新的物种概念, 即形态-生物学物种概念。最后, 以芍药属(Paeonia)几个物种的处理为例, 说明这一新的物种概念是可操作的, 划分的物种在形态上区别分明, 易于鉴别。更重要的是, 其结果得到基于25或26个单拷贝或寡拷贝核基因DNA序列所作的系统发生分析的强有力支持。各个物种在系统发生树上形成单系和独立的谱系, 表明其间各自形成独立的基因库, 没有基因交换, 它们独立进化, 有各自的生态位和独立的分布区。因此, 利用这一新的物种概念能够达到预期目标。     

物种——老问题新看法

简要回顾了物种概念争论的历史.根据来源将这些争论划分为4个阶段:第一阶段是在达尔文《物种起源》发表前,争论的主导概念是模式种概念;第二阶段从《物种起源》发表到综合进化论期间,争论的主导概念是形态种概念;第三阶段从综合进化论到Hennig的分支分类时代,以Mayr的生物学种概念为代表从机制上阐明了物种存在的生物学原因;第四阶段,分支分类以来,以系统发育种的概念为代表.在分析物种关键问题的基础上,结合物种研究新的进展,提出物种是由特定机制所维系的具有一定属性表现的优良基因组合的保存单元.基于植物演化的特点,提出认识植物物种的3个阶段:形态识别阶段、多学科证据积累和检验阶段与优良基因组合保存机制阐明阶段.     

物种与物种多样性

本文首先讨论生物物种的科学概念和生物学本质，分析物种客观存在的自然属性和物种概念的局限性，认为物种的生物学属性和物种多样性的科学属性之间有着本质联系。物种多样性研究的实质是研究生物物种的生物学多样性。度量物种多样性程度有多种方法，但物种数目是物种多样性程度最直接、也是最基本的表达，估计物种多样性数目是当前国际上物种多样性研究的核心与热点内容。物种多样性产生的根源是物种形成，物种绝灭速率是维持物种多样性的关键因素。本文简要总结了物种形成与绝灭的基本模式和机制，通过分析生物地理区系与物种多样性研究的密切关系，说明物种的区系成份分析是物种多样性大尺度格局研究的重要内容。     

菌物分类学研究中常见的物种概念

物种是生物多样性与分类学研究的基本单元, 物种识别是生物学研究的基本问题之一。物种的划分一直以来都没有一个明确统一的标准, 这使得分类学多少带有主观的色彩, 并经常被看作艺术而不完全是科学的研究。本文简要概述了菌物分类学研究中常见的3个物种概念, 即形态学种、生物学种和系统发育学种的背景和应用现状, 并通过实例讨论了这3个物种概念的特点及应用中存在的问题, 特别是各个物种概念之间的交错, 以期为菌物分类学研究和物种概念探讨提供参考。     

生物学物种概念在化石物种鉴定中的应用问题北大核心CSCD

物种概念是生物学的核心问题,而在化石鉴定中的应用存在诸多困难和分歧。本文介绍生物学物种概念在古生物学研究中的优点、难点与必要性。近年来,生物学物种概念已被古生物学家们广泛接受,居群和种内变异的重要性越来越受到重视。文章以二叠纪牙形类和腕足类化石种等为例,探讨如何在厘定物种、建立新种和使用亚种概念时,运用生物学物种概念及其延伸出来的各个标准。文章还讨论生物学物种在不同门类中的适用情况。认为在后生动物中,生物学物种可以得到较好的验证,在化石鉴定中,要尽量识别每一个种的发育序列和充分考虑种内变异,这样鉴定的化石种具有较高的地层对比价值,应当尽量避免仅仅根据个别标本的形态特征鉴定化石物种。     

细胞水平的分类学——细胞分类学

各学科互相渗透形成众多的边缘学科,这是当代科学发展的重要特点。细胞学和细胞遗传学同分类学之间的渗透,出现了细胞水平的分类学——细胞分类学。形态-地理学标准是传统分类学的主要方法。本世纪以来,由于实验生物学的蓬勃发展,分类学曾一度遭到冷落。然而,实验对象必须正确鉴定,实验所得信息需要记录和储存在确切的信息库内;自然界又发现不少兄弟种(sibling species)它们在形态上不能或难以区别,但却有被保护的独立的基因库。因此形态-地理学的物种概念被生物学物种概念所替代。与此同时,实验生物学,特别是     

中国植物分类学中的物种问题

Nooteboom和Peter Raven都认为中国的分类学家往往持有狭隘的物种概念,而这是由于他们缺 少足够的标本,尤其是模式标本,因而不能充分研究种的变异性。Raven说这使他相信中国植物分类学 中真正的物种概念颇有模式概念的色彩。他们的说法不无道理。的确,中国的植物区系中可能有大量 的种其真实身份还有讨论的余地。根据单个性状或完全按营养器官性状来命名新种是常有的事。有些 学者甚至把发表新种作为其分类工作的主要目标。现有的、为数非常有限的关于中国植物变异式样研 究的证据证明有些“种”实际上是生态宗(Clinopodium)、地理宗(Cunninghamia和Indigofera)或呈地形 梯度变异式样的分类群(Lespedeza和Rhododendron)而已。根据表型的可塑性变化而命名的种曾经或 仍然被作为“好种”对待(Rorippa)。具各种不同叶形特性的种间杂种的分离系被命名为不同的种 (Ilex)。分类学上本来就已经很复杂的无融合生殖复合体由于新种的不断增加而变得更加复杂 (Malus)。对一个具有复杂变异式样的种的精心研究导致对25个种名的归并,其中有10个是80年代 由中国分类学家发表的(Clematoclethra)。诸如此类的例子随着研究工作的扩大而势必大量增加。传统 植物分类学主要或完全以形态性状为依据。外部形态性状具有比较容易观察和记录的优点。分类学种 概念能满足多种用途分类的需要。但如果把分类学种概念说成是完全凭分类学家个人的主观判断或甚 至偏爱而很难有是非之分就颇成问题了。现在比过去任何时候都有条件对种这个分类阶元作客观的解 释。正确的物种概念来自对植物变异式样及其分类学价值的正确和充分的理解。因此,作为标本室分 类学家,首先要研究尽可能多的标本。其次,尽可能收编来自其他方面的证据是十分可取的。这些证据 即使不是很有用的分类性状,对更好地了解或解释类群的变异式样往往具有重要的意义。分类学家可 借以发现他所寻找的分类群在表型或基因型变异上的间断。对这两种间断的合理应用将导致在许多属内种级水平的更加合理的分类。     

我国植物种级水平分类学研究刍议

对洪德元先生最近在《生物多样性》(2016年第24卷第3期)发表的《关于提高物种划分合理性的意见》一文中的部分观点进行了进一步阐述。强调我国植物确实还存在大量种级水平的分类学问题有待解决, 我国植物分类学研究在一些重要发展阶段(如系统阶段和物种生物学阶段)上存在明显缺失, 需要弥补。指出分类学发展到今天, 不宜再强调“经典分类学”和“实验分类学”之分, 应采用多学科手段解决分类学问题; 我国应加强植物分类专著水平的研究工作, 注意培养年轻一辈植物分类学专著工作者; 在分类处理中应用居群概念和统计学方法时应特别谨慎; 在系统植物学中接受物种概念的多元性是必要的, 但要向达到广义的生物学种概念努力, 不宜以有所谓的“归并派”和“细分派”之分为借口而完全主观地划分物种。     

生物多样性事业呼唤对物种概念和 物种划分标准的深度讨论

正物种概念也许是这个世界上最重要、最复杂的概念了。说它重要,是因为它是划分物种的依据,而物种划分的结果直接或者间接地影响到社会的方方面面;说它复杂,是因为人们对它的讨论在过去几百年中从来就没有停止过,而且至今尚未达成共识。近年来,随着学科的发展和社会的进步,人们对物种概念和物种划分标准的讨论进入了新的阶段。一方面,大家普遍认识到正确划分物种的必要     

Species concept and speciation

Defining and recognizing a species has been a controversial issue for a long time. To determine the variation and the limitation between species, many concepts have been proposed. When a taxonomist study a particular taxa, he/she must adopted a species concept and provide a species limitation to define this taxa. In this paper some of species concepts are discussed starting from the typological species concepts to the phylogenetic concept. Positive and negative aspects of these concepts are represented in addition to their application.     

Patterns of bryophyte diversity and rarity at a regional scale

Information on bryophyte diversity and rarity were combined withinformation on soil conditions and land use for Walloon Brabant (centralBelgium, 1091 km²) in order to investigate whichlandscape features sustain the most rare and diverse species assemblages.Presence–absence of 325 bryophyte species was recorded in 87 grid-squaresof 4 × 4 km. Species diversity was significantly correlatedwith forest cover (r = 0.71, P <0.001), sandy soils (r = 0.61, P <0.001), loamy soils, (r = –0.68,P < 0.001), and agricultural fields(r = –0.49, P < 0.001). Themost diverse grid-squares possessed up to 182 species and were characterized byat least 10% forest cover and the presence of unique micro-habitats.Grid-squares with forest cover reaching at least 10% but lacking uniquemicro-habitats contained between 90 and 130 species. Below 10% forest cover,diversity ranged between 55 and 110 species per grid-square. However, even theleast diverse cultivated areas included a significant amount of the regionallyrare species. A number of the latter are characteristic in other areas forspecific primary habitats lacking in Walloon Brabant but display an unexpectedability to disperse throughout hostile areas and colonize secondary habitats.The tendency of such species to occur in man-made habitats decreased our abilityto predict species richness and rarity from landscape features and soilconditions.     

Diversity and abundance of leafhopper species (Homoptera: Cicadellidae) among red maple clones

A survey was conducted in the 2001 growing season to examine the leafhopper diversity and abundance among trees of 17 red maple (Acer rubrum) clones. Yellow sticky traps were used to qualify and quantify the number of aerial leafhoppers from 1 May 2001 until 4 September 2001. A total of 45 species from eight different leafhopper subfamilies, for a total of 6055 individuals, were considered in this study. The mean number of leafhoppers collected, mean species richness, diversity and evenness were significantly lower on traps of trees for October Glory than for the other clones. Yet, none of the leafhopper species dominated the weekly samples. Species similarity among clones ranged from 56 to 90%. No two clones had complete similarity. Franksred and trees of a controlled cross between October Glory × Autumn Flame shared the highest degree of species similarity, while clones from PA, USA and RI, USA were the least similar. The development of new clones did not create new pest problems, but suppressed populations of damaging pests, and maintained the diversity of low abundance species.     

Species richness, diversity and biomass production of desert annuals in an ungrazed Rhanterium epapposum community over three growth seasons in Kuwait

Two blocks, each consisting of twelve 0.25 m²permanent plots, were established in a community dominated by desert annuals inan area of Kuwait protected from grazing, and community attributes wereinvestigated over three growth seasons. A linear relationship was found betweenestimated desert annual cover and biomass production. Species frequency wasassessed by the number of 100 cm² subplots occupied byaspecies in a block (block incidence). Over the three seasons,vegetation features varied considerably. The first season (spring 1997) wascharacterised by very sparse vegetation cover, low vegetation block incidence(sum of individual block incidences of each species present in a block) and lowspecies richness. The second season (spring 1998) had high vegetation cover,relatively low vegetation block incidence, but high species richness. In thethird season (spring 2000), vegetation cover was very sparse (as in 1997), butboth blocks had the highest vegetation block incidence of the three years andspecies richness was moderately high, although individual plants were verysmall. The growth period was much shorter in the very productive andspecies-rich 1998 season than in 2000. Fluctuations in biomass and speciesincidence coincided with pronounced variation in key meteorological variables,in particular the onset of the first rains, the amount of rainfall during thegrowing season and its temporal distribution. Species composition, however,appears mainly to be determined by the amount and possibly the timing of thefirst rainfalls of the season to trigger germination. There was little evidenceto suggest that particularly favourable or unfavourable microsites existed inplots for more than one vegetation period, either for individual species or fordiversity in general. Despite substantial between-season fluctuations inincidence, the relative abundances of the more frequent species to each otherremained broadly similar over the three seasons, with Plantagoboissieri being the predominant species and accounting for thelargest proportion of biomass.     

The species-area relation III. A third model for a delimited community

Summary A mathematical model of the species-area relation is described for a group of limited species. This model is a modification of that proposed earlier (Kobayashi, 1975), being assumed that the limited species are expected to occur in a habitat under consideration. The model equation gives a sigmoid species-log area curve implying that few rare species are found in a group of species. The good agreement between observation and this model is exemplified with the data of plant and arthropod communities. The implication of parameters involved are examined in connection with those of the preceding model, and the underlying ecology of the model is discussed. Contribution from the Laboratory of Applied Zoology, Yamagata University, No. 83.     

The species in primatology

Biologists of the late eighteenth and early nineteenth centuries all bandied about the term “species,” but very rarely actually said what they meant by it. Often, however, one can get inside their thinking by piecing together some of their remarks. One of the most nearly explicit‐appropriately, for the man who wrote a book called The Origin of Species – was Charles Darwin 1 : “Practically, when a naturalist can unite two forms together by others having intermediate characters, he treats the one as a variety of the other… He later translated this into evolutionary terms: “Hereafter, we shall be compelled to acknowledge that the only distinction between species and well‐marked varieties is, that the latter are known, or believed, to be connected at the present day by intermediate gradations, whereas species were formerly thus connected”^1:484‐5     

Changes in plant species diversity along a chronosequence of vegetation restoration in the humid evergreen broad-leaved forest in the Rainy Zone of West China

Plant species diversity has been recognized as one of the vital attributes for assessing vegetation restoration. Changes in the diversity may be related to different stages of succession. In this study, 54 sites of humid, evergreen, broad-leaved forest were selected in the Rainy Zone of West China. A chronosequence of the sites was used to study the successive patterns of the diversity in the forest that had undergone natural regeneration for 5 to 350 years and to test the hypothesis that the diversity is maximized in mid-succession. Data were collected simultaneously at different stages of succession, and four α-diversity indices (species richness, Margalef index, Shannon-Wiener index, Pielou Evenness index) and two β-diversity indices (Whittaker index, Sørensen’s index) were calculated for each stratum in each plot. A total of 394 vascular plant species were recorded. From the β-diversity indices, the forest succession may be divided into the early-successional stage (before 50 years), mid-successional stage (from 50 to 300 years), and late-successional stage (after 300 years). In this community, the species diversity and richness were found to be the greatest at the mid-successional stage, followed by the late- and early-successional stages. The results of regression analysis indicated that the richness and Margalef index peaked around the 175th and 165th year, respectively. Shannon-Wiener index values also appeared to follow an approximately humped pattern of succession and were maximal around the 100th year. However, the species evenness did not show any significant relationship with successional age. Our results demonstrate (1) forest restoration is a long-term process and the formation of climax forest requires at least 300 years and (2) the forest has a strong capacity for restoration. Our results also suggest Lindera limprichitii and Machilus pingii as ideal tree species for afforestation because of their wide niche.     

Microhabitat variations and seed bank-vegetation relationships in a desert wadi ecosystem

Due to extreme variability in patterns of rainfall, plant seed banks are an important component of desert habitats. Here I report on effects of standing vegetation and three different microhabitats (channel, bank and terrace) on the soil seed bank of a desert wadi ecosystem in the Eastern Desert of Egypt. A total of 450 soil samples at 45 stands were collected to represent the different wadi microhabitats. The germinable seed bank was estimated by controlled counts of seedling emergence. The floristic composition, functional properties and diversity of the soil seed bank, as well as its similarity with the standing vegetation varied among wadi microhabitats. Such variation could be attributed to differences in disturbance intensity among microhabitats (terrace < bank < channel) and variation of soil factors along the microtopographic gradient. Channel showed the highest species richness and size of soil seed bank, followed by bank and then terrace. Moreover the Shannon index of diversity of the seed bank and its similarity with standing vegetation were significantly greater in both channel and bank microhabitats than in terrace. At the level of plant functional groups, number of seeds of annuals was higher in both channel and bank than in terrace. Shrubs were more abundant in seed banks of channel compared to terrace. The size and species richness of seed bank were increased with the total plant cover, annual/perennial ratio and species richness of the standing vegetation.     

Environmental drivers of the bryophyte propagule bank and its comparison with forest-floor assemblage in Central European temperate mixed forests

Species richness, composition and abundance of the bryophyte diaspore bank of Central European temperate mixed forests were compared with the forest-floor bryophyte assemblage. The impact of environmental variables and anthropogenic disturbances, including tree species composition, stand structure, microclimate, light conditions, soil and litter properties, management history, and landscape properties, potentially influencing bryophyte diaspore bank assemblages were explored. Thirty-four, 70–100 years old mixed stands with differing tree species composition were examined in the ?rség National Park, Western Hungary. The diaspore bank was studied by soil collection and cultivation, and data were analysed by multivariate methods. Contrary to the forest-floor bryophyte assemblage, where substrate availability, tree species composition and stand structure were the most influential environmental variables, the composition and abundance of the diaspore bank was mainly affected by site conditions (microclimate, litter and soil properties). Species richness of the bryophyte diaspore bank was lower than that of the forest-floor bryophyte assemblage. Short-lived mosses (colonists, short-lived shuttles) were dominant in the diaspore bank, as opposed to the forest-floor bryophyte community, where perennial mosses dominated. In the studied forests, the importance of the bryophyte diaspore bank was relatively low in the regeneration and maintenance of the forest-floor bryophyte vegetation.     

科尔沁沙地沙漠化过程中的物种多样性

对内蒙古科尔沁地区沙漠化过程中物种多样性的变化特征进行了分析 ,依据植物种在各类型沙漠化土地的出现频率和不同沙漠化阶段各生活型植物种的组成 ,探讨了沙漠化过程中植物种绝灭和定居特点 .物种多样性指数分析表明 ,沙漠化过程是一个物种多样性衰减的过程 ,沙漠化首先导致特有种的绝灭 ,其次为稀有种和普通种 .从植物的生活型来看 ,多年生草本和灌木类植物受沙漠化影响最大 ,而 1年生草本和半灌木类植物可存活于沙漠化过程各个阶段 .在沙漠化过程中植物种的绝灭速率大于定居速率 .