並系群
![本頁使用了標題或全文手工轉換](http://upload.wikimedia.org/wikipedia/commons/thumb/c/cd/Zh_conversion_icon_m.svg/35px-Zh_conversion_icon_m.svg.png)
![](http://upload.wikimedia.org/wikipedia/commons/thumb/d/df/Clade-grade_II.png/300px-Clade-grade_II.png)
並系(英文:paraphyly 或 paraphyletic)又稱並源[1],是系統發生學上常用的概念,指某一生物類群包含了一個共同祖先及其部分後代,但未包含其所有後代。這種由同一祖先演化而來的部分後代所組成的類群,被稱為並系群[2](英文:paraphyletic group;中國大陸又譯作偏系群[3],臺灣又譯作側系群[4]或旁系群[5])。通常並系群包含了與祖先相比性狀變化不大的成員,而變化更大的成員則被排除在外[6]。根據生物分類學界普遍認同的支序系統學觀點,任何合理有效的自然分類群都應為單系群(包含一個祖先及其所有後代[7]),因此並系群被視作不完整的、人為設定的非自然類群[8]。
與並系群和單系群相關的概念還有多系群,又稱複系群,是指不具有最近共同祖先的生物類群,即一群來源於不同祖先的生物被人為歸類在一起,同樣不被視作自然合理的分類群[2][8]。並系群與多系群的產生,大部分為傳統認知的局限性所致,在未修正為單系群之前,通常被學界視作無效的分類。與多系群一般作拆分處理相對,並系群一般會進行合併處理。隨着生物分類研究的不斷深入和遺傳學技術的不斷完善,許多過去遺留的並系群已被合併調整為單系群。
詞源
[編輯]在英語和其他一些使用拉丁字母書寫的語言中,「並系」(英文:paraphyly 或 paraphyletic)這一術語源自兩個古希臘文單詞:前綴「παρά」(拉丁轉寫:pará;意為旁邊、附近)+詞根「φῦλον」(phûlon;屬,種)[9][10],指某生物類群與其唯一共同祖先的其他所有後代相互分離、未被歸於一類的情況。
與之對應的術語「單系」(英文:monophyly 或 monophyletic),指某生物類群由其唯一共同祖先的所有後代組成。詞根與「並系」同源,前綴來自古希臘文「μόνος」(mónos;單獨的,唯一的)[9][10]。
另一相關術語「多系」(英文:polyphyly 或 polyphyletic),指某生物類群出自多個祖先,即內部成員各自有着不同的起源。詞根也與「並系」同源,前綴來自古希臘文「πολύς」(polús;多,很多)[9][10]。
範例
[編輯]並系群中一個典型的例子,是傳統分類法所定義的爬行綱。古生物學和遺傳學證據均明確顯示,鳥類為恐龍的後裔,並且是鱷魚現存關係最近的旁親。爬行綱的傳統定義包含了鳥類的祖先(恐龍),但卻未包含該祖先的後裔(鳥類),因此有部分學者使用蜥形綱來取代傳統的爬行綱,將鳥類劃入其中,以構成一個完整的單系群[11]。
現生羊膜動物的分化關係如下所示,傳統意義上的爬行綱為並系群,只有當其包含鳥類(與蜥形綱同義)時,方能成為合理的單系群[12]。
|
爬行綱 Reptilia |
另一個廣為人知的並系群是舊時定義的偶蹄目。遺傳學研究表明,鯨豚類動物是偶蹄目的演化支,而在傳統分類中,鯨豚類被單列為鯨目,並不屬於偶蹄動物,這意味着傳統的偶蹄目實為不合理的並系群[13]。因此,現行分類將鯨目降級為鯨下目,歸入偶蹄目之下,以形成完整的單系群[14][15]。
|
傳統的偶蹄目 |
除上述兩個例子之外,生物分類學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 | 對應的單系群 | 參考文獻 |
---|---|---|---|
原核生物(Prokaryota) | 真核生物(Eukaryota) | 生物 | [16] |
原生生物(Protista) | 植物、動物、真菌 | 真核生物 | [17] |
輪藻門(Charophyta) | 有胚植物(Embryophyta) | 鏈型植物(Streptophyta) | [18] |
苔蘚植物 | 維管植物(Tracheophyta) | 有胚植物 | [19][18] |
裸子植物(Gymnospermae) | 被子植物(Angiospermae) | 種子植物(Spermatophyta) | [20] |
雙子葉植物 | 單子葉植物 | 被子植物 | [21] |
無脊椎動物 | 脊椎動物(Vertebrata) | 動物界(Animalia) | [22] |
海綿 | 真後生動物(Eumetazoa) | 動物界 | [23][24] |
輻射對稱動物(Radiata) | 兩側對稱動物(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 | 軟水母亞綱(Hydroidolina) | 水母亞門 | [28][29][30] |
蠕蟲 | 不固定,因使用情況而異 | 腎管動物(Nephrozoa) | [31][32] |
環神經動物(Cycloneuralia) | 泛節肢動物(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] |
廣腰亞目(Symphyta) | 細腰亞目(Apocrita) | 膜翅目(Hymenoptera) | [38] |
黃蜂 | 螞蟻(蟻總科 Formicoidea)、蜜蜂(花蜂類 Anthophila) | 細腰亞目 | [39] |
胡蜂總科(Vespoidea) | 蟻總科、蜜蜂總科(Apoidea) | 真針尾類(Euaculeata) | [39] |
扁蟲動物(Platyzoa) | 冠輪動物(Lophotrochozoa)、中生動物(Mesozoa) | 螺旋動物(Spiralia) | [40] |
輪蟲動物門(Rotifera) | 棘頭動物門(Acanthocephala) | 合皮動物(Syndermata) | [41][42] |
鸚鵡螺亞綱(Nautiloidea) | 菊石亞綱(Ammonoidea)、蛸亞綱(Coleoidea) | 頭足綱(Cephalopoda) | [43] |
魚綱(Pisces) | 四足動物(Tetrapoda) | 脊椎動物 | [38] |
蜥蜴亞目(Lacertilia) | 蛇亞目(Serpentes)、蚓蜥類(Amphisbaenia) | 有鱗目(Squamata) | [44] |
盤龍目(Pelycosaur) | 獸孔目(Therapsida) | 合弓綱(Synapsida) | [11] |
斜溝齒獸亞目(Plagiaulacida) | 土鼠亞目(Cimolodonta)、北美俊獸科(Arginbaataridae) | 多瘤齒獸目(Multituberculata) | [45] |
古鯨類(Archaeoceti) | 新鯨類(Neoceti) | 鯨類(Cetacea) | [46] |
原猿類(Prosimii) | 類人猿下目(Simiiformes) | 靈長目(Primates) | [47] |
參考文獻
[編輯]- ^ Paraphyletic-並源[的];並系[的]. 雙語詞彙、學術名詞暨辭書資訊網. 新北: 國家教育研究院. (原始內容存檔於2021-12-30).(繁體中文)
- ^ 2.0 2.1 第二屆遺傳學名詞審定委員會 (編). 遗传学名词 (2版). 北京: 科學出版社. 2006. ISBN 978-7-03-016800-9. LCCN 2009393587. OCLC 71742122. OL 23974572M. NLC 003136637.(並系群,復系)(簡體中文)
- ^ 第二屆微生物學名詞審定委員會 (編). 微生物学名词 (2版). 北京: 科學出版社. 2012. ISBN 978-7-03-035918-6. OCLC 910353784. OL 37808101M. NLC 006772289.(偏系群)(簡體中文)
- ^ 黃啟東. 以形態、分子與珠芽發育特徵研究臺灣目賊芋的側系群分類 [Paraphyly Study of Remusatia vivipara (Roxb.) Schott (Araceae) by Morphology, Molecular and Bulbil Development Traits] (博士論文). 臺北: 國立臺灣大學生態學與演化生物學研究所. 2015. doi:10.6342/NTU.2015.00008. (原始內容存檔於2022-01-09).
- ^ Biodiversity Terms. 兩岸生物多樣性名詞翻譯查詢編修系統. 臺北: 中央研究院生物多樣性研究中心. (原始內容存檔於2022-03-11).(繁體中文)
- ^ Ridley, Mark. Evolution - A-Z - Paraphyletic group. Evolution. Blackwell Publishing. (原始內容存檔於2004-09-20).
- ^ 第二屆植物學名詞審定委員會 (編). 植物学名词 (2版). 北京: 科學出版社. 2019. ISBN 978-7-03-062178-8. OCLC 1198145621. OL 37808061M. NLC 010446494.(單系[類]群)(簡體中文)
- ^ 8.0 8.1 古生物學名詞審定委員會 (編). 古生物学名词 (2版). 北京: 科學出版社. 2009. ISBN 978-7-03-025433-7. LCCN 2010372103. OCLC 886143158. NLC 004299369.(並系[性],多系[性])(簡體中文)
- ^ 9.0 9.1 9.2 Bailly, Anatole. Abrégé du dictionnaire grec français. Paris: Hachette. 1981-01-01. ISBN 978-2-01003528-9. OCLC 461974285.(法文)
- ^ 10.0 10.1 10.2 Bailly, Anatole. Greek-french dictionary online. Tabularium.be. (原始內容存檔於2016-10-05).(法文)
- ^ 11.0 11.1 Benton, M.J. Appendix: Classification of the vertebrates. Vertebrate Palaeontology (3rd ed.). Oxford, England: Blackwell Publishing. 2005: 389–403 [2021-12-30]. ISBN 0-632-05637-1. LCCN 2003028152. OL 3681987M. NLC 001102922. (原始內容存檔於2008-10-19).
- ^ Modesto, S.P.; Anderson, J.S. The Phylogenetic Definition of Reptilia. Systematic Biology. 2004, 53 (5): 815–821. CiteSeerX 10.1.1.507.6893
. PMID 15545258. S2CID 4112111
. doi:10.1080/10635150490503026
.
- ^ O'Leary, Maureen A. The Phylogenetic Position of Cetaceans: Further Combined Data Analyses, Comparisons with the Stratigraphic Record and a Discussion of Character Optimization. American Zoologist. 2001, 41 (3): 487–506. CiteSeerX 10.1.1.555.8631
. S2CID 198156070
. doi:10.1093/icb/41.3.487
.
- ^ Groves, C.P.; Grubb, P. Ungulate Taxonomy (PDF). Baltimore, Maryland: Johns Hopkins University Press. 2011: 27–28. ISBN 978-1-4214-0093-8. LCCN 2011008168. OCLC 708357723. OL 25220152M. NLC 003158499. (原始內容存檔 (PDF)於2019-12-26).
- ^ List of marine mammal species and subspecies. MarineMammalScience.org. Society for Marine Mammalogy. 2022-05 [2022-08-09]. (原始內容存檔於2020-11-24).
- ^ Linda, Berg. Introductory Botany: Plants, People, and the Environment (2nd ed.). Belmont, CA: Thomson-Brooks/Cole. 2008: 360. ISBN 978-0495384786. LCCN 2006939406. OCLC 153914481. OL 7651605M.
- ^ Schlegel, Martin; Hülsmann, Norbert. Protists – A textbook example for a paraphyletic taxon. Organisms Diversity & Evolution. 2007, 7 (2): 166–172. ISSN 1439-6092. S2CID 84631175
. doi:10.1016/j.ode.2006.11.001
.
- ^ 18.0 18.1 Becker, B.; Marin, B. Streptophyte algae and the origin of embryophytes. Annals of Botany. 2009, 103 (7): 999–1004. JSTOR 43576186. PMC 2707909
. PMID 19273476. S2CID 7018235
. doi:10.1093/aob/mcp044
.
- ^ Cox, Cymon J.; Li, Blaise; Foster, Peter G.; Embley, T. Martin; Civáň, Peter. Conflicting Phylogenies for Early Land Plants are Caused by Composition Biases among Synonymous Substitutions. Systematic Biology. 2014, 63 (2): 272–279. PMC 3926305
. PMID 24399481. S2CID 9140429
. doi:10.1093/sysbio/syt109
.
- ^ Christenhusz, M.J.M.; Reveal, J.L.; Farjon, A.; Gardner, M.F.; Mill, R.R.; Chase, M.W. A new classification and linear sequence of extant gymnosperms (PDF). Phytotaxa. 2011, 19: 55–70. S2CID 86797396
. doi:10.11646/phytotaxa.19.1.3
. (原始內容存檔 (PDF)於2022-03-10).
- ^ Simpson, Michael George. Plant systematics. Burlington; San Diego; London: Academic Press. 2006: 139–140. ISBN 978-0-12-644460-5. OL 7329015M.
It is now thought that the possession of two cotyledons is an ancestral feature for the taxa of the flowering plants and not an apomorphy for any group within. The 'dicots' ... are paraphyletic ...
[目前認為,擁有兩個子葉是開花植物類群的祖先特徵,而不是任何類群內部的衍徵。「雙子葉植物」……是並系的……] - ^ Agassiz, Louis. Essay on Classification. Mineola, New York: Dover Publications. 2013: 115. ISBN 978-0-486-15135-9. OCLC 868969147. (原始內容存檔於2021-09-09).
- ^ Borchiellini, C.; Manuel, M.; Alivon, E.; Boury-Esnault, N.; Vacelet, J.; Le Parco, Y. Sponge paraphyly and the origin of Metazoa. Journal of Evolutionary Biology. 2001, 14 (1): 171–179. PMID 29280585. S2CID 25119754
. doi:10.1046/j.1420-9101.2001.00244.x
.
- ^ Philippe, H.; Derelle, R.; Lopez, P.; Pick, K.; Borchiellini, C.; Boury-Esnault, N.; Vacelet, J.; Renard, E.; Houliston, E.; Quéinnec, E.; Da Silva, C.; Wincker, P.; Le Guyader, H.; Leys, S.; Jackson, D.J.; Schreiber, F.; Erpenbeck, D.; Morgenstern, B.; Wörheide, G.; Manuel, M. Phylogenomics Revives Traditional Views on Deep Animal Relationships. Current Biology. 2009, 19 (8): 706–712. PMID 19345102. S2CID 15282843
. doi:10.1016/j.cub.2009.02.052
.
- ^ Martindale, Mark; Finnerty, J.R.; Henry, J.Q. The Radiata and the evolutionary origins of the bilaterian body plan. Molecular Phylogenetics and Evolution. 2002, 24 (3): 358–365. PMID 12220977. S2CID 6570344. doi:10.1016/s1055-7903(02)00208-7.
- ^ Stampar, S.N.; Maronna, M.M.; Kitahara, M.V.; Reimer, J.D.; Morandini, A.C. Fast-Evolving Mitochondrial DNA in Ceriantharia: A Reflection of Hexacorallia Paraphyly?. PLOS ONE. 2014, 9 (1): e86612. Bibcode:2014PLoSO...986612S. PMC 3903554
. PMID 24475157. S2CID 11763910
. doi:10.1371/journal.pone.0086612
.
- ^ Ruppert, Edward E.; Fox, Richard S.; Barnes, Robert D. 7 CNIDARIA
. Invertebrate zoology: a functional evolutionary approach (7th ed.). Belmont, CA: Thomson-Brooks/Cole. 2004: 132–148. ISBN 0-03-025982-7. LCCN 2003107287. OCLC 752875516. OL 7271214M.
- ^ Marques, Antonio C.; Collins, Allen G. Cladistic analysis of Medusozoa and cnidarian evolution. Invertebrate Biology. 2004, 123 (1): 23–42. S2CID 28342963
. doi:10.1111/j.1744-7410.2004.tb00139.x
.
- ^ Zou, H.; Zhang, J.; Li, W.; Wu, S.; Wang, G. Mitochondrial Genome of the Freshwater Jellyfish Craspedacusta sowerbyi and Phylogenetics of Medusozoa. PLOS ONE. 2012, 7 (12): e51465. Bibcode:2012PLoSO...751465Z. PMC 3519871
. PMID 23240028. S2CID 17907871
. doi:10.1371/journal.pone.0051465
.
- ^ Zapata, F.; Goetz, F.E.; Smith, S.A.; Howison, M.; Siebert, S.; Church, S.H.; Sanders, S.M.; Ames, C.L.; McFadden, C.S.; France, S.C.; Daly, M.; Collins, A.G.; Haddock, S.H.D.; Dunn, C.W.; Cartwright, P. Phylogenomic Analyses Support Traditional Relationships within Cnidaria. PLOS ONE. 2015, 10 (10): e0139068. Bibcode:2015PLoSO..1039068Z. PMC 4605497
. PMID 26465609. S2CID 16364776
. bioRxiv 10.1101/017632
. doi:10.1371/journal.pone.0139068
.
- ^ Fedonkin, M.A.; Simonetta, A.; Ivantsov, A.Y. New data on Kimberella, the Vendian mollusc-like organism (White sea region, Russia): palaeoecological and evolutionary implications. Vickers-Rich, Patricia; Komarower, Patricia (編). The Rise and Fall of the Ediacaran Biota. Special publications 286. London: Geological Society. 2007: 157–179. ISBN 978-1-86239233-5. LCCN 2008273095. OCLC 156823511. OL 23157129M.
- ^ Butterfield, N.J. Hooking some stem-group "worms": fossil lophotrochozoans in the Burgess Shale. BioEssays. 2006, 28 (12): 1161–1166. PMID 17120226. S2CID 29130876. doi:10.1002/bies.20507.
- ^ Dunn, C.W.; Hejnol, A.; Matus, D.Q.; Pang, K.; Browne, W.E.; Smith, S.A.; Seaver, E.; Rouse, G.W.; Obst, M.; Edgecombe, G.D.; Sørensen, M.V.; Haddock, S.H.D.; Schmidt-Rhaesa, A.; Okusu, A.; Kristensen, R.M.; Wheeler, W.C.; Martindale, M.Q. & Giribet, G. Broad phylogenomic sampling improves resolution of the animal tree of life. Nature. 2008, 452 (7188): 745–749. Bibcode:2008Natur.452..745D. PMID 18322464. S2CID 4397099
. doi:10.1038/nature06614.
- ^ Webster, Bonnie L.; Copley, Richard R.; Jenner, Ronald A.; Mackenzie-Dodds, Jacqueline A.; Bourlat, Sarah J.; Rota-Stabelli, Omar; Littlewood, D.T.J.; Telford, Maximilian J. Mitogenomics and phylogenomics reveal priapulid worms as extant models of the ancestral Ecdysozoan. Evolution & Development. 2006, 8 (6): 502–510. PMID 17073934. S2CID 22823313. doi:10.1111/j.1525-142X.2006.00123.x.
- ^ Andrew, David R. A new view of insect–crustacean relationships II. Inferences from expressed sequence tags and comparisons with neural cladistics. Arthropod Structure & Development. 2011, 40 (3): 289–302. PMID 21315832. S2CID 29360314. doi:10.1016/j.asd.2011.02.001.
- ^ Bjoern, M.; von Reumont, Ronald A.; Jenner, Matthew A.; Wills, Emiliano; Dell'Ampio, Günther; Pass, Ingo; Ebersberger, Benjamin; Meyer, Stefan; Koenemann, Thomas M. Iliffe. Pancrustacean phylogeny in the light of new phylogenomic data: support for Remipedia as the possible sister group of Hexapoda. Molecular Biology and Evolution. 2012, 29 (3): 1031–1045. CiteSeerX 10.1.1.460.4464
. PMID 22049065. S2CID 1047649
. doi:10.1093/molbev/msr270
. (原始內容存檔於2022-01-09).
- ^ Scoble, M.J. The Lepidoptera: form, function and diversity (Reprint). Oxford, England: Oxford University Press. 1995. ISBN 0-19-854952-0. OCLC 639626639. OL 7400889M.
- ^ 38.0 38.1 Sharkey, M.J. Phylogeny and classification of Hymenoptera (PDF). Zootaxa. 2007, 1668 (1): 521–548. S2CID 86772806
. doi:10.11646/zootaxa.1668.1.25. (原始內容存檔 (PDF)於2022-03-08).
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.
[廣腰亞目和細腰亞目長期以來一直被認為是膜翅目的兩個亞目,但由於認識到廣腰亞目的並系性(Köningsmann 1977,Rasnitsyn 1988),加上支序學分類法的出現,這樣的亞目分類應當避免。同樣的,樹蜂被認為是非單系的,形成了一個相對於細腰亞目和尾蜂科的祖先級別。傳統的膜翅目分類是錯誤的,根據支序學的標準,這種情況與支序學出現之前的脊椎動物分類相同,那時認為具有共同祖徵的類群是自然的,例如魚類曾被統一歸為「魚綱」,將四足動物排除在外。] - ^ 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
.
- ^ Brusca, Richard C. Ch 16 Gnathifera (PDF). RickBrusca.com. (原始內容存檔 (PDF)於2022-01-09).
- ^ Ruppert, Edward E.; Fox, Richard S.; Barnes, Robert D. 23 GNATHIFERA
. Invertebrate zoology: a functional evolutionary approach (7th ed.). Belmont, CA: Thomson-Brooks/Cole. 2004: 788–806. ISBN 0-03-025982-7. LCCN 2003107287. OCLC 752875516. OL 7271214M.
- ^ Shimek, Ronald L. Nano-Animals, Part I: Rotifers. Reefkeeping. 2006-01. (原始內容存檔於2022-03-08).
- ^ Lindgren, A.R.; Giribet, G.; Nishiguchi, M.K. A combined approach to the phylogeny of Cephalopoda (Mollusca). Cladistics. 2004, 20 (5): 454–486. PMID 34892953. S2CID 85975284
. doi:10.1111/j.1096-0031.2004.00032.x
.
- ^ Reeder, Tod W.; Townsend, Ted M.; Mulcahy, Daniel G.; Noonan, Brice P.; Wood, Perry L.; Sites, Jack W.; Wiens, John J. Integrated Analyses Resolve Conflicts over Squamate Reptile Phylogeny and Reveal Unexpected Placements for Fossil Taxa. PLOS ONE. 2015, 10 (3): e0118199. Bibcode:2015PLoSO..1018199R. PMC 4372529
. PMID 25803280. S2CID 5046152
. doi:10.1371/journal.pone.0118199
. hdl:10088/25654.
- ^ Kielan-Jaworowska, Z. & Hurum, J. Phylogeny and Systematics of Multituberculate Animals. Palaeontology. 2001, 44 (3): 389–429. doi:10.1111/1475-4983.00185
.
- ^ Thewissen, J.G.M.; Williams, E.M. The Early Radiations of Cetacea (Mammalia): Evolutionary Pattern and Developmental Correlations. Annual Review of Ecology and Systematics. 2002, 33: 73–90. JSTOR 3069257. S2CID 85720339. doi:10.1146/annurev.ecolsys.33.020602.095426.
- ^ Groves, C.P. Systematics of tarsiers and lorises. Primates. 1998, 39 (1): 13–27. S2CID 10869981. doi:10.1007/BF02557740.
外部連結
[編輯]維基共享資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.
|