並系群
並系(英文:paraphyly 或 paraphyletic)又称並源[1],是系统发生学上常用的概念,指某一生物类群包含了一个共同祖先及其部分後代,但未包含其所有後代。这种由同一祖先演化而来的部分後代所组成的类群,被称为並系群[2](英文:paraphyletic group;中国大陆又译作偏系群[3],臺灣又译作側系群[4]或旁系群[5])。通常並系群包含了与祖先相比性状变化不大的成员,而变化更大的成员则被排除在外[6]。根据生物分类学界普遍认同的支序系統學观点,任何合理有效的自然分類群都应为单系群(包含一个祖先及其所有後代[7]),因此並系群被视作不完整的、人为设定的非自然类群[8]。
与並系群和单系群相关的概念还有多系群,又称複系群,是指不具有最近共同祖先的生物类群,即一群来源於不同祖先的生物被人为归类在一起,同样不被视作自然合理的分类群[2][8]。並系群与多系群的产生,大部分为传统认知的局限性所致,在未修正为单系群之前,通常被学界视作无效的分类。与多系群一般作拆分处理相对,並系群一般会进行合并处理。随着生物分类研究的不断深入和遗传学技术的不断完善,许多过去遗留的並系群已被合并调整为单系群。
词源
在英语和其他一些使用拉丁字母书写的语言中,“並系”(英文:paraphyly 或 paraphyletic)这一术语源自两个古希腊文单词:前缀“παρά”(拉丁转写:pará;意为旁边、附近)+词根“φῦλον”(phûlon;属,种)[9][10],指某生物类群与其唯一共同祖先的其他所有後代相互分离、未被归于一类的情况。
与之对应的术语“單系”(英文:monophyly 或 monophyletic),指某生物类群由其唯一共同祖先的所有後代组成。词根与“並系”同源,前缀来自古希腊文“μόνος”(mónos;单独的,唯一的)[9][10]。
另一相关术语“多系”(英文:polyphyly 或 polyphyletic),指某生物类群出自多个祖先,即内部成员各自有着不同的起源。词根也与“並系”同源,前缀来自古希腊文“πολύς”(polús;多,很多)[9][10]。
范例
並系群中一个典型的例子,是传统分类法所定义的爬行纲。古生物学和遗传学证据均明确显示,鸟类为恐龙的後裔,并且是鳄鱼现存關係最近的旁亲。爬行纲的传统定义包含了鸟类的祖先(恐龙),但却未包含该祖先的後裔(鸟类),因此有部分学者使用蜥形纲来取代传统的爬行纲,将鸟类划入其中,以构成一个完整的单系群[11]。
现生羊膜動物的分化關係如下所示,传统意义上的爬行纲为並系群,只有当其包含鸟类(与蜥形纲同义)时,方能成为合理的单系群[12]。
|
爬行纲 Reptilia |
另一个广为人知的並系群是旧时定义的偶蹄目。遗传学研究表明,鲸豚类动物是偶蹄目的演化支,而在传统分类中,鲸豚类被单列为鲸目,並不屬於偶蹄动物,这意味着传统的偶蹄目实为不合理的並系群[13]。因此,现行分类将鲸目降级为鲸下目,归入偶蹄目之下,以形成完整的单系群[14][15]。
|
传统的偶蹄目 |
除上述两个例子之外,生物分类学上其他的主要並系群如下:
並系群名稱 | 應包含而未包含的演化支 | 對應的单系群 | 參考文獻 |
---|---|---|---|
原核生物(Prokaryota) | 真核生物(Eukaryota) | 生物 | [16] |
原生生物(Protista) | 植物、动物、真菌 | 真核生物 | [17] |
轮藻门(Charophyta) | 有胚植物(Embryophyta) | 链型植物(Streptophyta) | [18] |
苔藓植物 | 维管植物(Tracheophyta) | 有胚植物 | [19][18] |
裸子植物(Gymnospermae) | 被子植物(Angiospermae) | 种子植物(Spermatophyta) | [20] |
双子叶植物 | 單子葉植物 | 被子植物 | [21] |
无脊椎动物 | 脊椎动物(Vertebrata) | 动物界(Animalia) | [22] |
海绵 | 真後生動物(Eumetazoa) | 动物界 | [23][24] |
辐射对称动物(Radiata) | 两侧对称动物(Bilateria) | 真後生動物 | [25] |
珊瑚 | 水母亞門(Medusozoa)、黏體動物亞門(Myxozoa) | 刺胞動物門(Cnidaria) | [26][27] |
水母 | 软水母亚纲(Hydroidolina) | 水母亞門 | [28][29][30] |
蠕虫 | 不固定,因使用情况而异 | 腎管動物(Nephrozoa) | [31][32] |
环神经动物(Cycloneuralia) | 泛节肢动物(Panarthropoda) | 蛻皮動物(Ecdysozoa) | [33][34] |
甲殼亞門(Crustacea) | 六足亞門(Hexapoda) | 泛甲殼動物(Pancrustacea) | [35][36] |
蛾 | 蝴蝶 | 鱗翅目(Lepidoptera) | [37] |
广腰亚目(Symphyta) | 细腰亚目(Apocrita) | 膜翅目(Hymenoptera) | [38] |
黄蜂 | 蚂蚁(蟻總科 Formicoidea)、蜜蜂(花蜂類 Anthophila) | 细腰亚目 | [39] |
胡蜂總科(Vespoidea) | 蟻總科、蜜蜂总科(Apoidea) | 真针尾类(Euaculeata) | [39] |
扁蟲動物(Platyzoa) | 冠輪動物(Lophotrochozoa)、中生动物(Mesozoa) | 螺旋動物(Spiralia) | [40] |
輪蟲動物門(Rotifera) | 棘头动物门(Acanthocephala) | 合皮動物(Syndermata) | [41][42] |
鹦鹉螺亚纲(Nautiloidea) | 菊石亚纲(Ammonoidea)、蛸亞綱(Coleoidea) | 头足纲(Cephalopoda) | [43] |
魚綱(Pisces) | 四足動物(Tetrapoda) | 脊椎动物 | [38] |
蜥蜴亞目(Lacertilia) | 蛇亚目(Serpentes)、蚓蜥类(Amphisbaenia) | 有鱗目(Squamata) | [44] |
盤龍目(Pelycosaur) | 獸孔目(Therapsida) | 合弓綱(Synapsida) | [11] |
斜沟齿兽亚目(Plagiaulacida) | 土鼠亞目(Cimolodonta)、北美俊兽科(Arginbaataridae) | 多瘤齒獸目(Multituberculata) | [45] |
古鯨類(Archaeoceti) | 新鯨類(Neoceti) | 鯨類(Cetacea) | [46] |
原猿类(Prosimii) | 類人猿下目(Simiiformes) | 灵长目(Primates) | [47] |
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[目前认为,拥有两个子叶是开花植物类群的祖先特徵,而不是任何类群内部的衍徵。“双子叶植物”……是並系的……] - ^ Agassiz, Louis. Essay on Classification. Mineola, New York: Dover Publications. 2013: 115. ISBN 978-0-486-15135-9. OCLC 868969147. (原始内容存档于2021-09-09).
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Symphyta and Apocrita have long been considered as suborders of Hymenoptera but since recognition of the paraphyletic nature of the Symphyta (Köningsmann 1977, Rasnitsyn 1988) and the advent of cladistic methods the subordinal classification should be avoided. Likewise the woodwasps are thought to be non-monophyletic, forming a grade that is ancestral relative to Apocrita and Orussidae. The traditional hymenopteran classification is faulty, by cladistic criteria,in the same way as pre-cladistic vertebrate classifications in which groups sharing plesiomorphic characterswere recognized as natural, e.g., fishes were once grouped together as 'Pisces', which excluded tetrapods.
[广腰亚目和细腰亚目长期以来一直被认为是膜翅目的两个亚目,但由於认识到广腰亚目的並系性(Köningsmann 1977,Rasnitsyn 1988),加上支序學分類法的出现,这样的亚目分类应當避免。同样的,树蜂被认为是非單系的,形成了一个相对於细腰亚目和尾蜂科的祖先级别。传统的膜翅目分类是错误的,根据支序学的标准,这种情况与支序学出现之前的脊椎动物分类相同,那时认为具有共同祖徵的类群是自然的,例如鱼类曾被统一归为“鱼纲”,将四足动物排除在外。] - ^ 跳转到: 39.0 39.1 Johnson, Brian R.; Borowiec, Marek L.; Chiu, Joanna C.; Lee, Ernest K.; Atallah, Joel; Ward, Philip S. Phylogenomics Resolves Evolutionary Relationships among Ants, Bees, and Wasps (PDF). Current Biology. 2013, 23 (20): 2058–2062. PMID 24094856. S2CID 230835 . doi:10.1016/j.cub.2013.08.050 .
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外部链接
- 维基共享资源上的相關多媒體資源:並系群
- Funk, D.J.; Omland, K.E. Species-Level Paraphyly and Polyphyly: Frequency, Causes, and Consequences, with Insights from Animal Mitochondrial DNA. Annual Review of Ecology, Evolution, and Systematics. 2003, 34: 397–423. JSTOR 30033781. S2CID 33951905 . doi:10.1146/annurev.ecolsys.34.011802.132421.