异涡虫属
异涡虫属 | |
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2017年新发现的日本异涡虫(Xenoturbella-japonica)[6] | |
科学分类 | |
界: | 动物界 Animalia |
门: | 异无腔动物门 Xenacoelomorpha |
亚门: | 异涡虫亚门 Xenoturbellida Bourlat et al., 2006[4][5] |
科: | 异涡虫科 Xenoturbellidae Westblad, 1949[1] |
属: | 异涡虫属 Xenoturbella Westblad, 1949[1][2] |
模式种 | |
博克异涡虫 Xenoturbella bocki | |
其他种 | |
异涡虫属(拉丁名:Xenoturbella)是一类生活在海底的小型无脊椎动物,身体构造极为简单,现被单独归至异无腔动物门下的异涡虫亚门(Xenoturbellida),已得到公认的物种仅有六个[7],分布于东北大西洋的欧洲近岸海域以及北太平洋的墨西哥、美国和日本近岸海域。
1915年,瑞典动物学家西克斯滕·博克(瑞典文:Sixten Bock)在瑞典西海岸采集到已知的第一种异涡虫的标本,但直到1949年才被瑞典动物学家埃纳尔·韦斯特布拉德(Einar Westblad)正式描述,韦斯特布拉德为纪念西克斯滕·博克,将该物种命名为博克异涡虫(Xenoturbella bocki)[1]。
分类简史
异涡虫类自被发现以来,它们在动物界中的系统发生位置便一直被认为是个未解之谜。
早期的 DNA 分析表明异涡虫与软体动物关系密切[8],但这可能是由于它们以软体动物的卵为主食,从而导致大量软体动物的 DNA 混入其体内[9]。
2006年的相关研究将异涡虫属单独置于一门,即异涡动物门(Xenoturbellida),作为后口动物的演化支暨步带动物的姐妹群[4],后续研究证实了该门是后口动物中的基底类群[10][11]或是步带动物姐妹群[12]的观点。然而,一些形态学特征(例如上皮细胞的纤毛结构)显示,异涡虫和另一类存在疑问的动物——无腔动物有着紧密的关联[13]。对异涡虫胚胎阶段的研究也显示它们为直接发育,无幼虫阶段,与无腔动物的发育模式近似[14]。基于数百种蛋白质的分子研究最终揭示了一个由异涡虫和无腔动物组成的单系群[15][12][16],被命名为异无腔动物门(Xenacoelomorpha)[12],原本的异涡动物门随之降级为亚门。
异无腔动物的单系起源很快便被确认,但其系统发生位置是处于两侧对称动物基部还是后口动物基部一直未有答案,直到2016年,随着基因和分类单元采样的增加,两项新的研究再次将异涡虫作为异无腔动物门下无腔动物的姐妹群,并将异无腔动物门作为肾管动物(包括原口动物和后口动物)的姐妹群,亦即两侧对称动物中最为基础的类群[17][18]。
异涡虫属在动物界中的系统发生位置[17] | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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下级分类
- 博克异涡虫 Xenoturbella bocki (Westblad, 1949)
=韦斯特布拉德异涡虫 X. westbladi (Israelsson, 1999):博克异涡虫的异名[19] - X. churro Rouse, Wilson, Carvajal & Vrijenhoek, 2016[17][20]
- X. hollandorum Rouse, Wilson, Carvajal & Vrijenhoek, 2016
- X. monstrosa Rouse, Wilson, Carvajal & Vrijenhoek, 2016
- X. profunda Rouse, Wilson, Carvajal & Vrijenhoek, 2016
- 日本异涡虫 X. japonica Nakano et al., 2017[6][21]
其中博克异涡虫(韦斯特布拉德异涡虫)发现于瑞典、苏格兰、冰岛和挪威的北海沿岸,以及克罗地亚的亚得里亚海沿岸[22];日本异涡虫发现于日本的西北太平洋沿岸[6];其馀四种异涡虫发现于墨西哥的加利福尼亚湾和美国的蒙特雷湾[17]。
六种异涡虫分为两个演化支,分别栖息于浅海和深海:博克异涡虫与 X. hollandorum 体型较小,身长约4厘米,生存于水深650米以内的海域,二者与身长略超过5厘米、已知栖息深度不到560米的日本异涡虫共同组成一个演化支;其他三个种 X. churro、X. monstrosa 与 X. profunda 体型较大,身长达到或超过10厘米,生存于1700–3700米以下的深水中[17],三者组成另一个演化支。
异涡虫属的分支图,基于线粒体 DNA 和蛋白质序列构建[17][6] | |||||||||||||||||||||||||||||||||||||||
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形态特征
异涡虫的身体呈两侧对称,内部结构极为简单,除平衡囊以外几乎没有真正的器官[22][23],也缺乏呼吸系统、循环系统、排泄系统和性腺,但可产生配子;嘴是腹中孔,通往胃腔,无肛门[22][23],排泄口与摄入食物的开口为同一个[24]。
参考文献
- ^ 1.0 1.1 1.2 1.3 Westblad, E. Xenoturbella bocki n. g., n. sp., a peculiar, primitive Turbellarian type. Arkiv för Zoologi. 1949, 1: 3–29.
- ^ Zhang, Zhi-Qiang. Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa (Magnolia Press). 2011-12-23, 3148: 1–237 [2021-06-16]. doi:10.11646/zootaxa.3148.1.1. (原始内容存档于2015-12-22).
- ^ Xenoturbella bocki Westblad, 1949. WoRMS. 2021 [2021-08-10]. (原始内容存档于2011-02-16).
- ^ 4.0 4.1 Bourlat, S.J.; Juliusdottir, T.; Lowe, C.J.; Freeman, R.; Aronowicz, J.; Kirschner, M.; Lander, E.S.; Thorndyke, M.; Nakano, H.; Kohn, A.B.; Heyland, A.; Moroz, L.L.; Copley, R.R. & Telford, M.J. Deuterostome phylogeny reveals monophyletic chordates and the new phylum Xenoturbellida (PDF). Nature. 2006, 444 (7115): 85–88 [2021-06-16]. Bibcode:2006Natur.444...85B. PMID 17051155. S2CID 4366885. doi:10.1038/nature05241. (原始内容存档 (PDF)于2021-06-24).
- ^ Tyler, S.; Artois, T.; Schilling, S.; Hooge, M. & Bush, L.F. (编). Xenoturbellida. World List of turbellarian worms: Acoelomorpha, Catenulida, Rhabditophora. 2006–2021 [2021-08-10]. (原始内容存档于2021-08-10) –通过WoRMS.
- ^ 6.0 6.1 6.2 6.3 Nakano, Hiroaki; Miyazawa, Hideyuki; Maeno, Akiteru; Shiroishi, Toshihiko; Kakui, Keiichi; Koyanagi, Ryo; Kanda, Miyuki; Satoh, Noriyuki; Omori, Akihito & Kohtsuka, Hisanori. A new species of Xenoturbella from the western Pacific Ocean and the evolution of Xenoturbella. BMC Evolutionary Biology. 2017, 17 (1): 245. PMC 5733810 . doi:10.1186/s12862-017-1080-2 .
- ^ 7.0 7.1 Tyler, S.; Artois, T.; Schilling, S.; Hooge, M. & Bush, L.F. (编). Xenoturbella Westblad, 1949. World List of turbellarian worms: Acoelomorpha, Catenulida, Rhabditophora. 2006–2021 [2021-08-10]. (原始内容存档于2021-03-23) –通过WoRMS.
- ^ Norén, Michael & Jondelius, Ulf. Xenoturbella 's molluscan relatives. Nature. 1997, 390 (6655): 31–32. Bibcode:1997Natur.390...31N. S2CID 4426393. doi:10.1038/36242.
- ^ Bourlat, Sarah J.; Nielsen, Claus; Lockyer, Anne E.; Littlewood, D. Timothy J. & Telford, Maximilian J. Xenoturbella is a deuterostome that eats molluscs. Nature. 2003, 424 (6951): 925–928. Bibcode:2003Natur.424..925B. PMID 12931184. S2CID 4413357. doi:10.1038/nature01851.
- ^ Perseke, Marleen; Hankeln, Thomas; Weich, Bettina; Fritzsch, Guido; Stadler, Peter F.; Israelsson, Olle; Bernhard, Detlef & Schlegel, Martin. The mitochondrial DNA of Xenoturbella bocki: genomic architecture and phylogenetic analysis. Theory in Biosciences. 2007-05-10, 126 (1): 35–42. PMID 18087755. S2CID 17065867. doi:10.1007/s12064-007-0007-7.
- ^ Telford, M.J. Xenoturbellida: the fourth deuterostome phylum and the diet of worms. Genesis. 2008, 46 (11): 580–586. PMID 18821586. S2CID 5379221. doi:10.1002/dvg.20414 .
- ^ 12.0 12.1 12.2 Philippe, H.; Brinkmann, H.; Copley, R.R.; Moroz, L.L.; Nakano, H.; Poustka, A.J.; Wallberg, A.; Peterson, K.J. & Telford, M.J. Acoelomorph flatworms are deuterostomes related to Xenoturbella. Nature. 2011, 470 (7333): 255–258. Bibcode:2011Natur.470..255P. PMC 4025995 . PMID 21307940. doi:10.1038/nature09676.
- ^ Lundin, K. The epidermal ciliary rootlets of Xenoturbella bocki (Xenoturbellida) revisited: new support for a possible kinship with the Acoelomorpha (Platyhelminthes). Zoologica Scripta. 1998, 27 (3): 263–270. S2CID 85324766. doi:10.1111/j.1463-6409.1998.tb00440.x.
- ^ Nakano, H.; Lundin, K.; Bourlat, S.J.; Telford, M.J.; Funch, P.; Nyengaard, J.R.; Obst, M. & Thorndyke, M.C. Xenoturbella bocki exhibits direct development with similarities to Acoelomorpha. Nature Communications. 2013, 4: 1537. Bibcode:2013NatCo...4.1537N. PMC 3586728 . PMID 23443565. doi:10.1038/ncomms2556 .
- ^ Hejnol, A.; Obst, M.; Stamatakis, A.; Ott, M.; Rouse, G.W.; Edgecombe, G.D.; et al. Assessing the root of bilaterian animals with scalable phylogenomic methods. Proceedings of the Royal Society, Series B. 2009, 276 (1677): 4261–4270 [2021-06-21]. doi:10.1098/rspb.2009.0896. (原始内容存档于2021-05-05).
- ^ Edgecombe, G.D.; Giribet, G.; Dunn, C.W.; Hejnol, A.; Kristensen, R.M.; Neves, R.C.; Rouse, G.W.; Worsaae, K. & Sørensen, M.V. Higher-level metazoan relationships: Recent progress and remaining questions. Organisms Diversity & Evolution. 2011, 11 (2): 151–172. S2CID 32169826. doi:10.1007/s13127-011-0044-4.
- ^ 17.0 17.1 17.2 17.3 17.4 17.5 Rouse, Greg W.; Wilson, Nerida G.; Carvajal, Jose I. & Vrijenhoek, Robert C. New deep-sea species of Xenoturbella and the position of Xenacoelomorpha. Nature. 2016-02-03, 530 (7588): 94–97. Bibcode:2016Natur.530...94R. PMID 26842060. S2CID 3870574. doi:10.1038/nature16545.
- ^ Cannon, Johanna T.; Vellutini, Bruno C.; Smith III, Julian; Ronquist, Frederik; Jondelius, Ulf & Hejnol, Andreas. Xenacoelomorpha is the sister group to Nephrozoa. Nature. 2016-02-03, 530 (7588): 89–93 [2021-06-21]. Bibcode:2016Natur.530...89C. PMID 26842059. S2CID 205247296. doi:10.1038/nature16520. (原始内容存档于2021-04-10).
- ^ Tyler, S.; Artois, T.; Schilling, S.; Hooge, M. & Bush, L.F. (编). Xenoturbella westbladi Israelsson, 1999. World List of turbellarian worms: Acoelomorpha, Catenulida, Rhabditophora. 2006–2021 [2021-08-10]. (原始内容存档于2021-03-21) –通过WoRMS.
- ^ Newly discovered deep-sea worms, including one named 'churro,' could shed light on animal evolution. Los Angeles Times. 2016-02-05 [2020-02-13]. (原始内容存档于2021-06-28).
- ^ Starr, Michelle. These Deep Sea Worms Without Butts Likely Haven't Evolved For Millions of Years. ScienceAlert. [2020-02-13]. (原始内容存档于2021-06-24).
- ^ 22.0 22.1 22.2 Israelsson, Olle. New light on the enigmatic Xenoturbella (phylum uncertain): ontogeny and phylogeny. Proceedings of the Royal Society B: Biological Sciences. 1999, 266 (1421): 835–841. PMC 1689910 . doi:10.1098/rspb.1999.0713.
- ^ 23.0 23.1 Giribet, Gonzalo & Edgecombe, Gregory D. The Invertebrate Tree of Life. Princeton, NJ: Princeton University Press. 2020-03-03: 88–89 [2021-06-16]. ISBN 978-0-691-17025-1. (原始内容存档于2021-07-07).
- ^ McCafferty, Georgia. Deep-sea 'purple sock' provides clues to early life. CNN. 2016-02-04. (原始内容存档于2016-02-05).
延伸阅读
- Haszprunar, G.; Rieger, R.M. & Schuchert, P. "Extant 'Problematica' within or near the Metazoa". Simonetta, A.M. & Conway Morris, S (编). The Early Evolution of Metazoa and the Significance of Problematic Taxa. Cambridge, UK: Oxford University Press. 1991: 99–105.
- Kjeldsen, K. U.; Obst, M.; Nakano, H.; Funch, P. & Schramm, A. Two Types of Endosymbiotic Bacteria in the Enigmatic Marine Worm Xenoturbella bocki. Applied and Environmental Microbiology. 2010, 76 (8): 2657–2662. PMC 2849209 . PMID 20139320. doi:10.1128/aem.01092-09 .