古蟲動物

古蟲動物; 化石時期： 525–505 Ma PreЄ Є O S D C P T J K Pg N 寒武紀早期至中期
	;
科學分類
界：	動物界 Animalia;
超門：	後口動物總門 Deuterostomia;
門：	†古蟲動物門 Vetulicolia 或; 脊索動物門 Chordata;
亞門：	（†古蟲動物亞門 Vetulicolia）; Shu et al., 2001
綱
	月燕蟲屬 Shenzianyuloma; 古蟲綱 Vetulicolida; 異形蟲綱 Heteromorphida; ？雲南蟲綱 Yunnanozoa;

古蟲動物（學名：Vetulicolia）是一類生存於寒武紀的海生動物，也是最早期的後口動物之一^[2]，傳統分類上被劃分為後口動物總門的古蟲動物門，由中國古生物學家舒德干等人於2001年建立^[3]，一些較近期的研究則將其歸類為脊索動物門的古蟲動物亞門^[1]。此類動物包括十餘個物種，其中大部分的化石發現於中國雲南省，另外在美國、加拿大、格陵蘭與澳洲坎加魯島亦有其分布。

生理描述

古蟲動物基本特徵是軀體分節，並由前體和後體兩部分組成。前體為消化道的前段（咽部），口開於前體前端，背區和腹區由5對鰓囊構造組成的鰓區所分隔；後體(尾部)為消化道的後段（即腸部），肛門末位。絕大多數古蟲類動物的後體由7節甚至更多的體節構成。有些種類的後體有不同程度的硬化。

古蟲動物都具有背區和腹區的分化、5對鰓囊和鰓裂構造等基本身體構造，根據後體構造不同，可分為古蟲綱（Vetulicolida Chen & Zhou, 1997）和異形蟲綱（Heteromorphida Shu, 2005）。

研究及分類歷史

澄江動物群的古蟲屬（Vetulicola）早在 1987 年就被描述，由於其分節特徵，長期被認為是節肢動物。但隨著雲南蟲類及地大動物、俞元動物化石的發現，大量特徵顯示了古蟲動物是一類低等後口動物。2001年，舒德干教授建立了古蟲動物門，並認為古蟲動物是後口動物的早期分支。此外，大部分學者也相信雲南蟲類（包括雲南蟲和海口蟲）可能與古蟲動物有緊密的關係。

分類

以下分類系統以中國古生物學家陳愛林等人於2018年的研究為基礎^[4]，並加入其後發表的物種：

古蟲動物 Vetulicolians
- 古蟲綱 Vetulicolida Chen & Zhou, 1997^[5]
  - 古蟲目 Vetulicolata Hou & Bergström, 1997
    - 古蟲科 Vetulicolidae Hou et al., 1999
      - 古蟲屬 Vetulicola Hou, 1987^[6]
        長方形古蟲 Vetulicola rectangulata Luo & Hou, 1999
        
        楔形古蟲 Vetulicola cuneata Hou, 1987^[6]
        
        崗頭村古蟲 Vetulicola gangtoucunensis Luo et al., 2005^[7]
        
        串珠古蟲 Vetulicola monile Aldridge, et al., 2007^[8]
        
        龍寶山古蟲 Vetulicola longbaoshanensis Yang et al., 2010^[9]
      - 古卵形蟲屬 Ooedigera Vinther et al., 2011^[10]
        皮氏古卵形蟲 Ooedigera peeli Vinther et al., 2011^[10]
      - 北大動物屬 Beidazoon Shu, 2005^[11]
        美麗北大蟲 Beidazoon venustum Shu, 2005^[11]
        （= 丘疹古蟲 Bullivetula variola Aldridge, et al., 2007^[8]）
    - 地大動物科 Didazoonidae Shu & Han, 2001^[3]
      - 地大動物屬 Didazoon Shu & Han, 2001^[3]
        郝氏地大蟲 Didazoon haoae Shu & Han, 2001^[3]
      - 島泳蟲屬 Nesonektris García-Bellido et al., 2014^[12]
        奧德里奇島泳蟲 Nesonektris aldridgei García-Bellido et al., 2014^[12]
      - 圓口蟲屬 Pomatrum Luo & Hu, 1999^[13]
        具腹圓口蟲 Pomatrum ventralis Luo & Hu, 1999^[13]
        （= 王冠西大蟲 Xidazoon stephanus Shu, Conway Morris & Zhang, 1999^[14]）
      - 俞元蟲屬 Yuyuanozoon Chen et al., 2003^[15]
        宏大俞元蟲 Yuyuanozoon magnificissimi Chen et al., 2003^[15]
- 異形蟲綱 Heteromorphida Shu, 2005^[11]
  - 班府蟲目 Banffiata Aldridge et al., 2007^[8]
    - 斑府蟲科 Banffiidae Caron, 2006
      - 斑府蟲屬 Banffia Walcott, 1911
        伸縮斑府蟲 Banffia constricta Walcott, 1911^[16]
        
        寬尾斑府蟲 Banffia episoma Conway Morris & Selden, 2015^[17]
      - 異形蟲屬 Heteromorphus Luo & Hu, 1999^[13]
        困惑異形蟲 Heteromorphus confusus (Chen & Zhou, 1997)^[5]
        （= 困惑斑府蟲 Banffia confusa Chen & Zhou, 1997^[5]）
        （= 長尾異形蟲 Heteromorphus longicaudatus Luo & Hu, 1999^[13]）
        
        "Form A" （未命名種）Shu, 2005^[11]
      - 斯金氏蟲屬 Skeemella Briggs et al., 2005^[18]
        桿狀斯金氏蟲 Skeemella clavula Briggs et al., 2005^[18]
- 綱未定
  - 目未定
    - 科未定
      - 月燕蟲屬 Shenzianyuloma McMenamin, 2019^[19]
        雲南月燕蟲 Shenzianyuloma yunnanense McMenamin, 2019^[19]
- 其他可能的古蟲動物
  - ?雲南蟲目 Yunnanozoa Dzik, 1995^[20]
    - 雲南蟲科 Yunnanozoonidae Dzik, 1995^[20]
      - 雲南蟲屬 Yunnanozoon Hou, Ramsköld & Bergström, 1991
        鉛色雲南蟲 Yunnanozoon lividum Hou, Ramsköld & Bergström, 1991
      - 海口蟲屬 Haikouella Chen, Huang & Li, 1999^[21]
        矛形海口蟲 Haikouella lanceolata Chen, Huang & Li, 1999^[21]
        
        尖山海口蟲 Haikouella jianshanensis Shu et al., 2003^[22]

參考文獻

^ ^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. 地球科學與環境學報. 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. 古生物學報. 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. （原始內容存檔於2015-09-07）.
^ 楊傑, 侯先光, 叢培允, 董渭, 張艷霞, 羅茂斌. "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. 古生物學報. 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. 科學通報. 2005, 50 (19): 2114–2126, (en) 50 (20): 2342–2354 [2014-06-19]. （原始內容存檔於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 羅惠麟, 胡世學, 陳良忠, 張世山, 陶永和. "昆明地区早寒武世澄江动物群" Early Cambrian Chengjiang fauna from Kunming region. 雲南科技出版社. 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. （原始內容 (PDF)存檔於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] （頁面存檔備份，存於網際網路檔案館）
^ 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. （原始內容存檔 (PDF)於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. （原始內容存檔於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. ^{[永久失效連結]}

延伸閱讀

陳均遠. "动物世界的黎明". 江蘇科學技術出版社. 2004: pp.309–318, 323–343. figs.493–509, 515–544. ISBN 7-5345-3797-5.
歐強, 舒德干. "后口动物咽鳃裂的起源和演化". 自然雜誌. 2014年2月, 36 (1): 14–23 [2014-06-19]. ISSN 0253-9608. （原始內容存檔於2016-03-05）.

外部連結

[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. 地球科學與環境學報. 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. 古生物學報. 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. （原始內容存檔於2015-09-07）.

[9] 楊傑, 侯先光, 叢培允, 董渭, 張艷霞, 羅茂斌. "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. 古生物學報. 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. 科學通報. 2005, 50 (19): 2114–2126, (en) 50 (20): 2342–2354 [2014-06-19]. （原始內容存檔於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 羅惠麟, 胡世學, 陳良忠, 張世山, 陶永和. "昆明地区早寒武世澄江动物群" Early Cambrian Chengjiang fauna from Kunming region. 雲南科技出版社. 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. （原始內容 (PDF)存檔於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] （頁面存檔備份，存於網際網路檔案館）

[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. （原始內容存檔 (PDF)於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. （原始內容存檔於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. ^{[永久失效連結]}

[2]

[3]

[1]

[4]

[5]

[6]

[7]

[8]

[9]

[10]

[11]

[12]

[13]

[14]

[15]

[16]

[17]

[18]

[19]

[20]

[21]

[22]