生物多样性 ›› 2016, Vol. 24 ›› Issue (9): 979-999. DOI: 10.17520/biods.2016203
出版日期:
2016-09-20
发布日期:
2016-10-09
通讯作者:
洪德元
基金资助:
Online:
2016-09-20
Published:
2016-10-09
Contact:
Hong De-Yuan
摘要:
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。
洪德元 (2016) 生物多样性事业需要科学、可操作的物种概念. 生物多样性, 24, 979-999. DOI: 10.17520/biods.2016203.
De-Yuan Hong (2016) Biodiversity pursuits need a scientific and operative species concept. Biodiversity Science, 24, 979-999. DOI: 10.17520/biods.2016203.
物种概念 Species concept | |
---|---|
1 | 无性繁殖物种 Agamospecies |
2 | 自衍征物种 Autapomorphic species |
3 | 生物学物种 Biospecies |
4 | 支序物种 Cladospecies |
5 | 内聚物种 Cohesion species |
6 | 掠夺物种 Compilospecies |
7 | 复合物种 Composite species |
8 | 生态学物种 Ecospecies |
9 | 进化物种 Evolutionary species |
10 | 进化显著单元 Evolutionary significant unit |
11 | 血缘和谐物种 Genealogical concordance species |
12 | 基因物种 Genic species |
13 | 遗传学物种 Genetic species |
14 | 基因型簇 Genotypic cluster |
15 | 亨氏物种 Hennigian species |
16 | 节点间物种 Internodal species |
17 | 涵盖范围最小分类单元 Least inclusive taxonomic unit (LITUS) |
18 | 形态学物种 Morphospecies |
19 | 无维物种 Non-dimensional species |
20 | 假物种 Nothospecies |
21 | 系统发生分类群物种 Phylogenetic taxon species |
22 | 表征物种 Phenospecies |
23 | 识别物种 Recognition species |
24 | 生殖竞争物种 Reproductive competition species |
25 | 接替物种 Successional (successive) species |
26 | 分类学物种 Taxonomic species |
表1 Wilkins (2006, 2009)列举的26个物种概念
物种概念 Species concept | |
---|---|
1 | 无性繁殖物种 Agamospecies |
2 | 自衍征物种 Autapomorphic species |
3 | 生物学物种 Biospecies |
4 | 支序物种 Cladospecies |
5 | 内聚物种 Cohesion species |
6 | 掠夺物种 Compilospecies |
7 | 复合物种 Composite species |
8 | 生态学物种 Ecospecies |
9 | 进化物种 Evolutionary species |
10 | 进化显著单元 Evolutionary significant unit |
11 | 血缘和谐物种 Genealogical concordance species |
12 | 基因物种 Genic species |
13 | 遗传学物种 Genetic species |
14 | 基因型簇 Genotypic cluster |
15 | 亨氏物种 Hennigian species |
16 | 节点间物种 Internodal species |
17 | 涵盖范围最小分类单元 Least inclusive taxonomic unit (LITUS) |
18 | 形态学物种 Morphospecies |
19 | 无维物种 Non-dimensional species |
20 | 假物种 Nothospecies |
21 | 系统发生分类群物种 Phylogenetic taxon species |
22 | 表征物种 Phenospecies |
23 | 识别物种 Recognition species |
24 | 生殖竞争物种 Reproductive competition species |
25 | 接替物种 Successional (successive) species |
26 | 分类学物种 Taxonomic species |
图15 Paeonia arietina (a-d, 均被柔毛)和P. mascula (e, f,两处均无毛)的茎表面和萼片背面的扫描电镜照片, 示被毛的差异。a, b (H02216)和c, d (H02217)均采自土耳其的Sivas; e, f: H01004, 采自法国的Chadieu。a, c, e: 茎表面; b, d, f: 萼片背面。
图16 Paeonia arietina (P. mascula subsp. arietina) (a, b)和P. mascula (c, d)两个类群的根形态。a: H02204, 采自土耳其的Ida山; b: H02224, 采自土耳其的Giresun; c: H02226, 采自希腊的Euboea; d: H02203, 采自土耳其的Canakkale。(洪德元摄)
图19 Paeonia obovata中花色、叶姿态、花丝和花药颜色的变异。a: 长白山, 花红色, 但叶展开; b: 河南卢氏县, 花瓣白色, 但周边粉色, 花丝下部紫色, 但上部白色; c: 日本爱媛县; d: 河南卢氏县; e: 四川卧龙; f: 吉林安图县。这里也包括了“P. japonica”的特征。
《中国植物志》27卷 ( | ||
---|---|---|
P. delavayi Franch.: 云南丽江等 P. lutea Delavay ex Franch.: 云南、西藏东南部 P. potaninii Kom. var. potaninii: 四川西部、云南永宁 var. trollioides (= P. trollioidess Stapf ex Stern): 云南德钦、西藏东南部 ……………………………… P. lutea var. ludlowii Stern & G. Taylor (1951): 西藏东南部 | P. delavayi: 四川木里、西藏察瓦龙、云南 P. lutea: 云南 P. potaninii var. potaninii: 四川木里、云南丽江 var. trollioides: 四川木里、云南丽江 ……………………………… | P. delavayi var. delavayi: 云南西北部、 四川西南部、西藏东南部 var. lutea: 云南、四川西南部、西藏东南部 var. angustiloba: 四川西部 ………………………………… |
表2 滇牡丹复合群(Paeonia delavayi Franch. complex) 1980年以前的分类处理
《中国植物志》27卷 ( | ||
---|---|---|
P. delavayi Franch.: 云南丽江等 P. lutea Delavay ex Franch.: 云南、西藏东南部 P. potaninii Kom. var. potaninii: 四川西部、云南永宁 var. trollioides (= P. trollioidess Stapf ex Stern): 云南德钦、西藏东南部 ……………………………… P. lutea var. ludlowii Stern & G. Taylor (1951): 西藏东南部 | P. delavayi: 四川木里、西藏察瓦龙、云南 P. lutea: 云南 P. potaninii var. potaninii: 四川木里、云南丽江 var. trollioides: 四川木里、云南丽江 ……………………………… | P. delavayi var. delavayi: 云南西北部、 四川西南部、西藏东南部 var. lutea: 云南、四川西南部、西藏东南部 var. angustiloba: 四川西部 ………………………………… |
图22 西藏大花黄牡丹(Paeonia ludlowii)与滇牡丹(P. delavayi)地下部分形态比较。a: 西藏的P. lutea var. ludlowii Stern & G. Taylor (1951) (P. delavayi var. lutea的一部分), 根渐细, 无地下走茎; b, c: 滇牡丹(P. delavayi) (包括“黄牡丹” P. delavayi var. lutea)根块状加粗, 具地下走茎。
图23 西藏大花黄牡丹(Paeonia ludlowii)与滇牡丹(P. delavayi)习性的比较。a: 西藏的P. lutea var. ludlowii Stern & G. Taylor (1951) (P. delavayi var. lutea的一部分)种子繁殖, 茎丛生; b: 滇牡丹(P. delavayi)(包括“黄牡丹” P. delavayi var. lutea)克隆繁殖为主, 茎散生。
图26 滇牡丹复合群(Paeonia delavayi Franch. complex)中4个分类群的模式类型。a: 滇牡丹(P. delavayi Franch.); b: 黄牡丹(P. lutea Delavay ex Franch. = P. delavayi var. lutea); c: 银莲保氏牡丹(P. potaninii var. trollioides (Stapf ex Stern) Stern); d: 保氏牡丹(P. potaninii Kom., 1921)。(洪德元摄)
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