古虫动物

古虫动物; 化石时期： 525–505 Ma PreЄ Є O S D C P T J K Pg N 寒武纪早期至中期
	;
科学分类
界：	动物界 Animalia;
超门：	后口动物总门 Deuterostomia;
门：	†古虫动物门 Vetulicolia 或; 脊索动物门 Chordata;
亚门：	（†古虫动物亚门 Vetulicolia）; Shu et al., 2001
纲
	月燕虫属 Shenzianyuloma; 古虫纲 Vetulicolida; 异形虫纲 Heteromorphida; ？云南虫纲 Yunnanozoa;

古虫动物（学名：Vetulicolia）是一类生存于寒武纪的海生动物，也是最早期的后口动物之一^[2]，传统分类上被划分为后口动物总门的古虫动物门，由中国古生物学家舒德干等人于2001年建立^[3]，一些较近期的研究则将其归类为脊索动物门的古虫动物亚门^[1]。此类动物包括十余个物种，其中大部分的化石发现于中国云南省，另外在美国、加拿大、格陵兰与澳洲坎加鲁岛亦有其分布。

生理描述

古虫动物基本特征是躯体分节，并由前体和后体两部分组成。前体为消化道的前段（咽部），口开于前体前端，背区和腹区由5对鳃囊构造组成的鳃区所分隔；后体(尾部)为消化道的后段（即肠部），肛门末位。绝大多数古虫类动物的后体由7节甚至更多的体节构成。有些种类的后体有不同程度的硬化。

古虫动物都具有背区和腹区的分化、5对鳃囊和鳃裂构造等基本身体构造，根据后体构造不同，可分为古虫纲（Vetulicolida Chen & Zhou, 1997）和异形虫纲（Heteromorphida Shu, 2005）。

研究及分类历史

澄江动物群的古虫属（Vetulicola）早在 1987 年就被描述，由于其分节特征，长期被认为是节肢动物。但随着云南虫类及地大动物、俞元动物化石的发现，大量特征显示了古虫动物是一类低等后口动物。2001年，舒德干教授建立了古虫动物门，并认为古虫动物是后口动物的早期分支。此外，大部分学者也相信云南虫类（包括云南虫和海口虫）可能与古虫动物有紧密的关系。

分类

以下分类系统以中国古生物学家陈爱林等人于2018年的研究为基础^[4]，并加入其后发表的物种：

古虫动物 Vetulicolians
- 古虫纲 Vetulicolida Chen & Zhou, 1997^[5]
  - 古虫目 Vetulicolata Hou & Bergström, 1997
    - 古虫科 Vetulicolidae Hou et al., 1999
      - 古虫属 Vetulicola Hou, 1987^[6]
        长方形古虫 Vetulicola rectangulata Luo & Hou, 1999
        
        楔形古虫 Vetulicola cuneata Hou, 1987^[6]
        
        岗头村古虫 Vetulicola gangtoucunensis Luo et al., 2005^[7]
        
        串珠古虫 Vetulicola monile Aldridge, et al., 2007^[8]
        
        龙宝山古虫 Vetulicola longbaoshanensis Yang et al., 2010^[9]
      - 古卵形虫属 Ooedigera Vinther et al., 2011^[10]
        皮氏古卵形虫 Ooedigera peeli Vinther et al., 2011^[10]
      - 北大动物属 Beidazoon Shu, 2005^[11]
        美丽北大虫 Beidazoon venustum Shu, 2005^[11]
        （= 丘疹古虫 Bullivetula variola Aldridge, et al., 2007^[8]）
    - 地大动物科 Didazoonidae Shu & Han, 2001^[3]
      - 地大动物属 Didazoon Shu & Han, 2001^[3]
        郝氏地大虫 Didazoon haoae Shu & Han, 2001^[3]
      - 岛泳虫属 Nesonektris García-Bellido et al., 2014^[12]
        奥德里奇岛泳虫 Nesonektris aldridgei García-Bellido et al., 2014^[12]
      - 圆口虫属 Pomatrum Luo & Hu, 1999^[13]
        具腹圆口虫 Pomatrum ventralis Luo & Hu, 1999^[13]
        （= 王冠西大虫 Xidazoon stephanus Shu, Conway Morris & Zhang, 1999^[14]）
      - 俞元虫属 Yuyuanozoon Chen et al., 2003^[15]
        宏大俞元虫 Yuyuanozoon magnificissimi Chen et al., 2003^[15]
- 异形虫纲 Heteromorphida Shu, 2005^[11]
  - 班府虫目 Banffiata Aldridge et al., 2007^[8]
    - 斑府虫科 Banffiidae Caron, 2006
      - 斑府虫属 Banffia Walcott, 1911
        伸缩斑府虫 Banffia constricta Walcott, 1911^[16]
        
        宽尾斑府虫 Banffia episoma Conway Morris & Selden, 2015^[17]
      - 异形虫属 Heteromorphus Luo & Hu, 1999^[13]
        困惑异形虫 Heteromorphus confusus (Chen & Zhou, 1997)^[5]
        （= 困惑斑府虫 Banffia confusa Chen & Zhou, 1997^[5]）
        （= 长尾异形虫 Heteromorphus longicaudatus Luo & Hu, 1999^[13]）
        
        "Form A" （未命名种）Shu, 2005^[11]
      - 斯金氏虫属 Skeemella Briggs et al., 2005^[18]
        杆状斯金氏虫 Skeemella clavula Briggs et al., 2005^[18]
- 纲未定
  - 目未定
    - 科未定
      - 月燕虫属 Shenzianyuloma McMenamin, 2019^[19]
        云南月燕虫 Shenzianyuloma yunnanense McMenamin, 2019^[19]
- 其他可能的古虫动物
  - ?云南虫目 Yunnanozoa Dzik, 1995^[20]
    - 云南虫科 Yunnanozoonidae Dzik, 1995^[20]
      - 云南虫属 Yunnanozoon Hou, Ramsköld & Bergström, 1991
        铅色云南虫 Yunnanozoon lividum Hou, Ramsköld & Bergström, 1991
      - 海口虫属 Haikouella Chen, Huang & Li, 1999^[21]
        矛形海口虫 Haikouella lanceolata Chen, Huang & Li, 1999^[21]
        
        尖山海口虫 Haikouella jianshanensis Shu et al., 2003^[22]

参考文献

^ ^1.0 ^1.1 García-Bellido, Diego C; Paterson, John R. A new vetulicolian from Australia and its bearing on the chordate affinities of an enigmatic Cambrian group. BMC Evolutionary Biology. 2014, 14: 214. PMC 4203957 . PMID 25273382. doi:10.1186/s12862-014-0214-z.
^ 舒德干. 寒武纪大爆发与动物树的成型. 地球科学与环境学报. 2009, 31 (2): 111–134. ISSN 1672-6561.
^ ^3.0 ^3.1 ^3.2 ^3.3 Shu, D.-G., Conway Morris, S., Han, J., Chen, L., Zhang, X.-L., Zhang, Z.-F., Liu, H.-Q., Li, Y., & Liu, J.-N. "Primitive Deuterostomes from the Chengjiang Lagerstätte (Lower Cambrian, China)" (PDF). Nature. 2001, 414: 419–424. doi:10.1038/35106514.
^ Li, Y.; Williams, M.; Gabbott, S.; Chen, A. "The enigmatic metazoan Yuyuanozoon magnificissimi from the early Cambrian Chengjiang Biota, Yunnan Province, South China". Journal of Paleontology. 2018, 92 (6): 1081–1091. doi:10.1017/jpa.2018.18.
^ ^5.0 ^5.1 ^5.2 J. Y. Chen, Y. N. Cheng, H. V. Iten.eds. "The Cambrian Explosion and the Fossil Record" 10. Taichung, Taiwan: Bulletin of the National Museum of Natural Science. 1997.
^ ^6.0 ^6.1 侯先光. "云南澄江早寒武世大型双瓣壳节肢动物" Early Cambrian large bivalved arthropods from Chengjiang, eastern Yunnan. 古生物学报. 1987, 26 (3): pp.286–298, pls. 1–6.
^ Luo Huilin, Fu Xiaoping, Hu Shixue, Li Yong, Chen, Liangzhong, You Ting, Liu Qi. "New Vetulicoliids from the Lower Cambrian Guanshan Fauna, Kunming". Acta Geologica Sinica - English Edition. 2005, 79 (1): 1–6. doi:10.1111/j.1755-6724.2005.tb00860.x.
^ ^8.0 ^8.1 ^8.2 R. J. Aldridge, Xian-guang Hou, David J. Siveter, Derek J. Siveter, S. E. Gabbott. "The Systematics and phylogenetic relationships of vetulicolians". Palaeontology. 2007, 50 (1): pp.140–142. pl.4: figs 1–5 [2014-06-24]. doi:10.1111/j.1475-4983.2006.00606.x. （原始内容存档于2015-09-07）.
^ 杨杰, 侯先光, 丛培允, 董渭, 张艳霞, 罗茂斌. "昆明地区下寒武统沧浪铺组古虫动物新发现" A new vetulicoliid from lower Cambrian, Kunming, Yunnan. 古生物学报. 2010, 49 (1): 54–63. ISSN 0001-6616.
^ ^10.0 ^10.1 J. Vinther, M. P .Smith, D. A. T. Harper. "Vetulicolians from the Lower Cambrian Sirius Passet Lagerstätte, North Greenland, and the polarity of morphological characters in basal deuterostomes". Palaeontology. 2011, 54 (3): 711–719. doi:10.1111/j.1475-4983.2011.01034.x.
^ ^11.0 ^11.1 ^11.2 ^11.3 舒德干. "再论古虫动物门" On the Phylum Vetulicolia. 科学通报. 2005, 50 (19): 2114–2126, (en) 50 (20): 2342–2354 [2014-06-19]. （原始内容存档于2014-09-08）.
^ ^12.0 ^12.1 García-Bellido, Diego C.; Paterson, John R. "A new vetulicolian from Australia and its bearing on the chordate affinities of an enigmatic Cambrian group". BMC Evolutionary Biology. 2014, 12: 214, figs.1–6. doi:10.1186/s12862-014-0214-z.
^ ^13.0 ^13.1 ^13.2 ^13.3 罗惠麟, 胡世学, 陈良忠, 张世山, 陶永和. "昆明地区早寒武世澄江动物群" Early Cambrian Chengjiang fauna from Kunming region. 云南科技出版社. 1999. ISBN 7-5416-1322-3.
^ Shu, D.-G., Conway Morris, S., Zhang, X.-L., Chen, L., Li, Y. & Han, J. "A pipiscid-like fossil from the Lower Cambrian of south China" (PDF). Nature. 1999, 400: 746–749 [2014-06-24]. doi:10.1038/23445. （原始内容 (PDF)存档于2014-08-10）.
^ ^15.0 ^15.1 Chen Ailin, Feng Hongzhen, Zhu Maoyan, Ma Dongsheng, Li Ming. "A New Vetulicolian from the Early Cambrian Chengjiang Fauna in Yunnan of China". Acta Geologica Sinica - English Edition. 2003, 77 (3): 281–287. doi:10.1111/j.1755-6724.2003.tb00742.x.
^ Caron, (2005-2006), Bannfia constricta, a putative vetulicolid from the Middle Cambrian Burgess Shales [1] （页面存档备份，存于互联网档案馆）
^ Conway Morris, S.; Halgedahl, S. L.; Selden, P.; Jarrard, R. D. "Rare primitive deuterostomes from the Cambrian (Series 3) of Utah" (PDF). Journal of Paleontology. 2015, 89: 631–636 [2017-07-30]. doi:10.1017/jpa.2015.40. （原始内容存档 (PDF)于2017-07-30）.
^ ^18.0 ^18.1 D. E. G. Briggs, B. S. Lieberman, S. L. Halgedahl & R. Jarrard. "A new metazoan from the Middle Cambrian of Utah and the nature of the Vetulicolia". Palaeontology. 2005, 48 (4): 681–686. doi:10.1111/j.1475-4983.2005.00489.x.
^ ^19.0 ^19.1 McMenamin, M. A. S. Cambrian Chordates and Vetulicolians. Geosciences. 2019, 9 (8): 354 [2020-05-05]. Bibcode:2019Geosc...9..354M. ISSN 2076-3263. doi:10.3390/geosciences9080354. （原始内容存档于2020-09-17）.
^ ^20.0 ^20.1 Dzik, Jerzy. "Yunnanozoon and the ancestry of chordates". Acta Palaeontologica Polonica. 1995, 40 (4): 341–360.
^ ^21.0 ^21.1 Y.-J. Chen, D.-Y. Huang, C.-W. Li. "An early Cambrian craniate-like chordatea". Nature. 1999, 402: 518–522. doi:10.1038/990080.
^ D.-G. Shu, S. Conway Morris, J. Han, Z.-F. Zhang, K. Yasui, P. Janvier, L. Chen, X.-L. Zhang, J.-N. Liu, Y. Li & H.-Q. Liu. "Head and backbone of the Early Cambrian vertebrate Haikouichthys" (PDF). Nature. 2003, 421: 526–529. doi:10.1111/j.1475-4983.2005.00489.x. ^{[永久失效链接]}

延伸阅读

陈均远. "动物世界的黎明". 江苏科学技术出版社. 2004: pp.309–318, 323–343. figs.493–509, 515–544. ISBN 7-5345-3797-5.
欧强, 舒德干. "后口动物咽鳃裂的起源和演化". 自然杂志. 2014年2月, 36 (1): 14–23 [2014-06-19]. ISSN 0253-9608. （原始内容存档于2016-03-05）.

外部链接

[Paterson2014-1] 1.0 ^1.1 García-Bellido, Diego C; Paterson, John R. A new vetulicolian from Australia and its bearing on the chordate affinities of an enigmatic Cambrian group. BMC Evolutionary Biology. 2014, 14: 214. PMC 4203957 . PMID 25273382. doi:10.1186/s12862-014-0214-z.

[2] 舒德干. 寒武纪大爆发与动物树的成型. 地球科学与环境学报. 2009, 31 (2): 111–134. ISSN 1672-6561.

[Shu2001-3] 3.0 ^3.1 ^3.2 ^3.3 Shu, D.-G., Conway Morris, S., Han, J., Chen, L., Zhang, X.-L., Zhang, Z.-F., Liu, H.-Q., Li, Y., & Liu, J.-N. "Primitive Deuterostomes from the Chengjiang Lagerstätte (Lower Cambrian, China)" (PDF). Nature. 2001, 414: 419–424. doi:10.1038/35106514.

[Li-4] Li, Y.; Williams, M.; Gabbott, S.; Chen, A. "The enigmatic metazoan Yuyuanozoon magnificissimi from the early Cambrian Chengjiang Biota, Yunnan Province, South China". Journal of Paleontology. 2018, 92 (6): 1081–1091. doi:10.1017/jpa.2018.18.

[Chen1997-5] 5.0 ^5.1 ^5.2 J. Y. Chen, Y. N. Cheng, H. V. Iten.eds. "The Cambrian Explosion and the Fossil Record" 10. Taichung, Taiwan: Bulletin of the National Museum of Natural Science. 1997.

[Hou1987-6] 6.0 ^6.1 侯先光. "云南澄江早寒武世大型双瓣壳节肢动物" Early Cambrian large bivalved arthropods from Chengjiang, eastern Yunnan. 古生物学报. 1987, 26 (3): pp.286–298, pls. 1–6.

[7] Luo Huilin, Fu Xiaoping, Hu Shixue, Li Yong, Chen, Liangzhong, You Ting, Liu Qi. "New Vetulicoliids from the Lower Cambrian Guanshan Fauna, Kunming". Acta Geologica Sinica - English Edition. 2005, 79 (1): 1–6. doi:10.1111/j.1755-6724.2005.tb00860.x.

[Aldridge-8] 8.0 ^8.1 ^8.2 R. J. Aldridge, Xian-guang Hou, David J. Siveter, Derek J. Siveter, S. E. Gabbott. "The Systematics and phylogenetic relationships of vetulicolians". Palaeontology. 2007, 50 (1): pp.140–142. pl.4: figs 1–5 [2014-06-24]. doi:10.1111/j.1475-4983.2006.00606.x. （原始内容存档于2015-09-07）.

[9] 杨杰, 侯先光, 丛培允, 董渭, 张艳霞, 罗茂斌. "昆明地区下寒武统沧浪铺组古虫动物新发现" A new vetulicoliid from lower Cambrian, Kunming, Yunnan. 古生物学报. 2010, 49 (1): 54–63. ISSN 0001-6616.

[Vinther-10] 10.0 ^10.1 J. Vinther, M. P .Smith, D. A. T. Harper. "Vetulicolians from the Lower Cambrian Sirius Passet Lagerstätte, North Greenland, and the polarity of morphological characters in basal deuterostomes". Palaeontology. 2011, 54 (3): 711–719. doi:10.1111/j.1475-4983.2011.01034.x.

[Shu2005-11] 11.0 ^11.1 ^11.2 ^11.3 舒德干. "再论古虫动物门" On the Phylum Vetulicolia. 科学通报. 2005, 50 (19): 2114–2126, (en) 50 (20): 2342–2354 [2014-06-19]. （原始内容存档于2014-09-08）.

[García2014-12] 12.0 ^12.1 García-Bellido, Diego C.; Paterson, John R. "A new vetulicolian from Australia and its bearing on the chordate affinities of an enigmatic Cambrian group". BMC Evolutionary Biology. 2014, 12: 214, figs.1–6. doi:10.1186/s12862-014-0214-z.

[Luo-13] 13.0 ^13.1 ^13.2 ^13.3 罗惠麟, 胡世学, 陈良忠, 张世山, 陶永和. "昆明地区早寒武世澄江动物群" Early Cambrian Chengjiang fauna from Kunming region. 云南科技出版社. 1999. ISBN 7-5416-1322-3.

[Xidazoon-14] Shu, D.-G., Conway Morris, S., Zhang, X.-L., Chen, L., Li, Y. & Han, J. "A pipiscid-like fossil from the Lower Cambrian of south China" (PDF). Nature. 1999, 400: 746–749 [2014-06-24]. doi:10.1038/23445. （原始内容 (PDF)存档于2014-08-10）.

[Chen2003-15] 15.0 ^15.1 Chen Ailin, Feng Hongzhen, Zhu Maoyan, Ma Dongsheng, Li Ming. "A New Vetulicolian from the Early Cambrian Chengjiang Fauna in Yunnan of China". Acta Geologica Sinica - English Edition. 2003, 77 (3): 281–287. doi:10.1111/j.1755-6724.2003.tb00742.x.

[16] Caron, (2005-2006), Bannfia constricta, a putative vetulicolid from the Middle Cambrian Burgess Shales [1] （页面存档备份，存于互联网档案馆）

[17] Conway Morris, S.; Halgedahl, S. L.; Selden, P.; Jarrard, R. D. "Rare primitive deuterostomes from the Cambrian (Series 3) of Utah" (PDF). Journal of Paleontology. 2015, 89: 631–636 [2017-07-30]. doi:10.1017/jpa.2015.40. （原始内容存档 (PDF)于2017-07-30）.

[Briggs-18] 18.0 ^18.1 D. E. G. Briggs, B. S. Lieberman, S. L. Halgedahl & R. Jarrard. "A new metazoan from the Middle Cambrian of Utah and the nature of the Vetulicolia". Palaeontology. 2005, 48 (4): 681–686. doi:10.1111/j.1475-4983.2005.00489.x.

[McMenamin-19] 19.0 ^19.1 McMenamin, M. A. S. Cambrian Chordates and Vetulicolians. Geosciences. 2019, 9 (8): 354 [2020-05-05]. Bibcode:2019Geosc...9..354M. ISSN 2076-3263. doi:10.3390/geosciences9080354. （原始内容存档于2020-09-17）.

[Dzik-20] 20.0 ^20.1 Dzik, Jerzy. "Yunnanozoon and the ancestry of chordates". Acta Palaeontologica Polonica. 1995, 40 (4): 341–360.

[Chen1999-21] 21.0 ^21.1 Y.-J. Chen, D.-Y. Huang, C.-W. Li. "An early Cambrian craniate-like chordatea". Nature. 1999, 402: 518–522. doi:10.1038/990080.

[22] D.-G. Shu, S. Conway Morris, J. Han, Z.-F. Zhang, K. Yasui, P. Janvier, L. Chen, X.-L. Zhang, J.-N. Liu, Y. Li & H.-Q. Liu. "Head and backbone of the Early Cambrian vertebrate Haikouichthys" (PDF). Nature. 2003, 421: 526–529. doi:10.1111/j.1475-4983.2005.00489.x. ^{[永久失效链接]}

[1]

[2]

[3]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[14]

[15]

[16]

[17]

[18]

[19]

[20]

[21]

[22]