埃笠姆鯡目

埃笠姆鯡目; 化石時期：豪特里維期——漸新世，130–28 Ma PreЄ Є O S D C P T J K Pg N
	具齒雙棱鯡（Diplomystus dentatus）的化石
科學分類
界：	動物界 Animalia
門：	脊索動物門 Chordata
綱：	條鰭魚綱 Actinopteri
亞綱：	新鰭亞綱 Neopterygii
下綱：	真骨下綱 Teleostei
高群：	骨舌魚高群 Osteoglossocephalai
總群：	鯡頭魚總群 Clupeocephala
群：	骨鯡群 Otomorpha
亞群：	鯡形亞群 Clupei
總目：	鯡形總目 Clupeomorpha
目：	†埃笠姆鯡目 Ellimmichthyiformes; Grande, 1985
屬
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埃笠姆鯡目（學名：Ellimmichthyiformes）是一類已滅絕的硬骨魚，屬於鯡形總目。它們生存於早白堊世至漸新世（約1.3億至2800萬年前），北美洲、南美洲、歐洲、亞洲和非洲都發現了它們的化石。

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描述

埃笠姆鯡目與現生沙丁魚和大西洋鯡非常相似，它們實際上是鯡形目的姐妹群。但和沙丁魚和大西洋鯡相比，埃笠姆鯡目的身體通常高而扁，有着許多具有特殊形態的背側和腹側骨質盾形鱗片，這些鱗片以類似於翅膀的形狀橫向擴寬，這使得它們的外觀幾乎是矩形的。埃笠姆鯡目通常很小，大多數物種的體長不超過10厘米，但雙棱鯡屬等少數類群的體長可達65厘米。

由於存在非常長的腹部鱗片，一些埃笠姆鯡類具有特別高且側扁的體形，例如菱鯡屬和富鯡屬。與鯡形目一樣，埃笠姆鯡目在每個發育階段的第一個聽覺中樞也具有第二個融合的耳緣，即一個前外陰窩和帶有外殼的背鱗。阿爾米百鯡屬等較古老和特化程度較低的埃笠姆鯡類與麗黃鯡屬（Ornateglum）和弗雷鯡屬（Foreyclupea）等特化程度小的白堊紀基底鯡形目非常相似；逐漸地，埃笠姆鯡屬、似埃笠姆鯡屬、副鯡屬、托內特鯡屬、突尼斯鯡屬、長盾鯡屬、三鱯鯡屬和肘鯡屬等更衍化的埃笠姆鯡類發展出不同的身體形態和鱗片覆蓋區域。索賓鯡屬是一類基底埃笠姆鯡類，長有非常長的胸鰭和非常高的背鰭。

分類

蘭斯·格蘭德（Lance Grande）於1982年設立了埃笠姆鯡目，當時該目包含埃笠姆鯡屬和雙棱鯡屬。^[1]^[2]隨後，生存於白堊紀、古新世和始新世的其它許多鯡形總目魚類也被歸入埃笠姆鯡目。^[3]^[4]^[5]^[6]^[7]^[8]^[9]^[10]長盾棘鯡屬、哨敗鯡屬等是目前已知最古老的埃笠姆鯡類，生存年代可以追溯到早白堊世豪特里維期，但一般認為它們已經是相當衍生的埃笠姆鯡類，因此最早的埃笠姆鯡類能追溯到更早的時代。^[11]目前已知生存年代最晚的埃笠姆鯡類是桂鯡屬，其化石發現於中國廣西壯族自治區崇左市寧明縣和南寧市邕寧區的漸新世地層。^[12]

埃笠姆鯡目內的系統分類學尚不明確，在最近的研究中，具甲鯡屬等以往被認為是基底埃笠姆鯡類的屬已被確定為相對較為衍化的屬。^[10]不管怎麼說，索賓鯡科（或索賓鯡屬和生存於古新世的腹鯡屬）似乎是埃笠姆鯡目中最原始的成員，而在副鯡科內，從雙棱鯡屬到阿爾米百鯡屬可能越來越衍生，該科也包括更衍化的埃笠姆鯡類。始新埃笠姆鯡屬是目前已知的最後一類海生埃笠姆鯡類，其化石發現於意大利蒙特波卡化石庫始新世地層。^[13]

下級分類：

†具甲鯡屬 Armigatus
†腹鯡屬 Gasteroclupea
†桂鯡屬 Guiclupea Chen, Chang, Wu & Liao, 2021
- †Guiclupea superstes ^[14]
†索賓鯡科 Sorbinichthyidae
- †索賓鯡屬 Sorbinichthys Bannikov & Bacchia, 2000
- †Sorbinichthys elusivo Bannikov & Bacchia, 2000
- †雙棱鯡屬 Diplomystus
†副鯡科 Paraclupeidae
- †肘鯡屬 Codoichthys
- †似埃笠姆鯡屬 Ellimma
- †埃笠姆鯡屬 Ellimmichthys
- †始新埃笠姆鯡屬 Eoellimmichthys
- †哨敗鯡屬 Ezkutuberezi
- †副鯡屬 Paraclupea
- †菱鯡屬 Rhombichthys
- †長盾鯡屬 Scutatoclupea
- †長盾棘鯡屬 Scutatuspinosus
- †托內特鯡屬 Thorectichthys
  - †Thorectichthys fideli^[15]
- †三鱯鯡屬 Triplomystus
  - †Triplomystus applegatei^[16]
  - †Triplomystus noorae Forey et al., 2003
  - †Triplomystus oligoscutatus Forey et al., 2003
- †突尼斯鯡屬 Tunisiaclupea
- †富鯡屬 Tycheroichthys

古生態學

埃笠姆鯡目是一類在河流、湖泊以及海洋中都有分布的掠食性魚類，一些埃笠姆鯡類以微小的無脊椎動物為食，而另一些埃笠姆鯡類動物（如雙棱鯡屬）則以其他魚類為食，一些腹部區域內保存有其它魚類的化石證明了這一點。背側和腹側的鱗片能夠有效保護埃笠姆鯡類免受大型掠食者的襲擊。

參考資料

[1]
Grande, L. 1982. A revision of the fossil genus †Diplomystus, with comments on the interrelationships of clupeomorph fishes. American Museum Novitates 2728:1–34.
[2]
Grande, L. 1985. Recent and fossil clupeomorph fishes with materials for revision of the subgroups of clupeoids. Bulletin of the American Museum of Natural History 181:231–372.
[3]
Chang, M. M., and J. G. Maisey. 2003. Redescription of †Ellimma branneri and †Diplomystus shengliensis, and relationships of some basal clupeomorphs. American Museum Novitates 3404:1–35.
[4]
Alvarado-Ortega, J., E. Ovalles-Damian, and G. Arratia. 2008. A review of the interrelationships of the order Ellimmichthyiformes (Teleostei: Clupeomorpha); pp. 257–278 in G. Arratia, H.-P. Schultze and M. V. H. Wilson (eds.), Mesozoic Fishes 4—Homology and Phylogeny. Verlag Dr. Friedrich Pfeil, Munich, Germany.
[5]
B. Khalloufi, R. Zaragüeta-Bagils, and H. Lelièvre. 2010. Rhombichthys intoccabilis, gen. et sp. nov. (Ellimmichthyiformes, Clupeomorpha, Teleostei), from the Cenomanian (Upper Cretaceous) of Ein Yabrud, Middle East: anatomical description and phylogenetic implications. Journal of Vertebrate Paleontology 30(1):57-67
[6]
Murray, A. M., and M. V. H. Wilson. 2013. Two new paraclupeid fishes (Clupeomorpha: Ellimmichthyiformes) from the Upper Cretaceous of Morocco; pp. 267–290 in G. Arratia, H.-P. Schultze, and M. V. H. Wilson (eds.), Mesozoic Fishes 5—Global Diversity and Evolution. Verlag Dr. Friedrich Pfeil, Munich.
[7]
Murray, A. M., O. Vernygora, S. Japundẑic, J. Radovcic, M. V. H. Wilson, D. Bardack, and T. Grande. 2016. Relationships of the species of Armigatus (Clupeomorpha, Ellimmichthyiformes) and the description of a new species from the Cretaceous of Dalmatia, Croatia. Journal of Vertebrate Paleontology. doi: 10.1080/02724634.2017.1226851.
[8]
de Figueiredo, F. J., and D. R. M. Ribeiro. 2016. Relationships of †Codoichthys carnavalii Santos, 1994 (Teleostei, Clupeomorpha, †Ellimmichthyiformes) from the Late Aptian of S~ao Luıs-Grajau Basin, Ne Brazil. Anais da Academia Brasileira de Ciencias 88: 1277–1307.
[9]
Marramà, G., and G. Carnevale. 2017. The relationships of Gasteroclupea branisai Signeux, 1964, a freshwater double-armored herring (Clupeomorpha, Ellimmichthyiformes) from the Late Cretaceous–Paleocene of South America. Historical Biology 29:904–917.
[10]
Kamel Boukhalfa, Feixiang Wu, Walid Ben Ali & Gengyu Fang (2019): A new paraclupeid fish (Clupeomorpha: Ellimmichthyiformes) from the Lower Cretaceous Sidi Aich Formation of southern Tunisia: phylogenetic and paleobiogeographic implications, Journal of Vertebrate Paleontology, DOI: 10.1080/02724634.2018.1529675
[11]
de Figueiredo, F. J., and D. R. M. Ribeiro. 2017. Reassessment and Relationships of †Scutatuspinosus itapagipensis (Teleostei, Clupeomorpha, †Ellimmichthyiformes) from the Neocomian of Reconcavo Basin, Northeastern Brazil. Anais da Academia Brasileira de Ciencias 89:799–823.
[12]
Chen, G., Chang, M. M., Wu, F., & Liao, X. (2021). Guiclupea superstes, gen. et sp. nov., the youngest ellimmichthyiform (clupeomorph) fish to date from the Oligocene of South China. PeerJ, 9, e11418. https://doi.org/10.7717/peerj.11418
[13]
Giuseppe Marramà; Alexandre F. Bannikov; Jürgen Kriwet; Giorgio Carnevale (2018). "An Eocene paraclupeid fish (Teleostei, Ellimmichthyiformes) from Bolca, Italy: the youngest marine record of double‐armoured herrings". Papers in Palaeontology. 5 (1): 83–98. doi:10.1002/spp2.1230. PMC 6392134. PMID 30854219.
[14]
Gengjiao Chen; Mee-mann Chang; Feixiang Wu; Xiaowen Liao; Guiclupea superstes, gen. et sp. nov., the youngest ellimmichthyiform (clupeomorph) fish to date from the Oligocene of South China
[15]
Thorectichthys (Ellimmichthyiformes) in America, from the Cenomanian deposits of the Cintalapa Formation, Chiapas, southeastern Mexico. Sciencedirect. [2024-06-16]. （原始內容存檔於2024-06-16）.
[16]
Jesús Alvarado-Ortega & Ernesto Ovalles-Damián (2008) Triplomystus applegatei, sp. nov. (Teleostei: Ellimmichthyiformes), a rare 「triple armored herring」 from El Espinal Quarry (Early Cretaceous), Chiapas, southeastern Mexico, Journal of Vertebrate Paleontology, 28:1, 53-60, DOI: 10.1671/0272-4634(2008)28[53:TASNTE]2.0.CO;2

[1] [1]
Grande, L. 1982. A revision of the fossil genus †Diplomystus, with comments on the interrelationships of clupeomorph fishes. American Museum Novitates 2728:1–34.

[2] [2]
Grande, L. 1985. Recent and fossil clupeomorph fishes with materials for revision of the subgroups of clupeoids. Bulletin of the American Museum of Natural History 181:231–372.

[3] [3]
Chang, M. M., and J. G. Maisey. 2003. Redescription of †Ellimma branneri and †Diplomystus shengliensis, and relationships of some basal clupeomorphs. American Museum Novitates 3404:1–35.

[4] [4]
Alvarado-Ortega, J., E. Ovalles-Damian, and G. Arratia. 2008. A review of the interrelationships of the order Ellimmichthyiformes (Teleostei: Clupeomorpha); pp. 257–278 in G. Arratia, H.-P. Schultze and M. V. H. Wilson (eds.), Mesozoic Fishes 4—Homology and Phylogeny. Verlag Dr. Friedrich Pfeil, Munich, Germany.

[5] [5]
B. Khalloufi, R. Zaragüeta-Bagils, and H. Lelièvre. 2010. Rhombichthys intoccabilis, gen. et sp. nov. (Ellimmichthyiformes, Clupeomorpha, Teleostei), from the Cenomanian (Upper Cretaceous) of Ein Yabrud, Middle East: anatomical description and phylogenetic implications. Journal of Vertebrate Paleontology 30(1):57-67

[6] [6]
Murray, A. M., and M. V. H. Wilson. 2013. Two new paraclupeid fishes (Clupeomorpha: Ellimmichthyiformes) from the Upper Cretaceous of Morocco; pp. 267–290 in G. Arratia, H.-P. Schultze, and M. V. H. Wilson (eds.), Mesozoic Fishes 5—Global Diversity and Evolution. Verlag Dr. Friedrich Pfeil, Munich.

[7] [7]
Murray, A. M., O. Vernygora, S. Japundẑic, J. Radovcic, M. V. H. Wilson, D. Bardack, and T. Grande. 2016. Relationships of the species of Armigatus (Clupeomorpha, Ellimmichthyiformes) and the description of a new species from the Cretaceous of Dalmatia, Croatia. Journal of Vertebrate Paleontology. doi: 10.1080/02724634.2017.1226851.

[8] [8]
de Figueiredo, F. J., and D. R. M. Ribeiro. 2016. Relationships of †Codoichthys carnavalii Santos, 1994 (Teleostei, Clupeomorpha, †Ellimmichthyiformes) from the Late Aptian of S~ao Luıs-Grajau Basin, Ne Brazil. Anais da Academia Brasileira de Ciencias 88: 1277–1307.

[9] [9]
Marramà, G., and G. Carnevale. 2017. The relationships of Gasteroclupea branisai Signeux, 1964, a freshwater double-armored herring (Clupeomorpha, Ellimmichthyiformes) from the Late Cretaceous–Paleocene of South America. Historical Biology 29:904–917.

[未命名-20230318190650-10] [10]
Kamel Boukhalfa, Feixiang Wu, Walid Ben Ali & Gengyu Fang (2019): A new paraclupeid fish (Clupeomorpha: Ellimmichthyiformes) from the Lower Cretaceous Sidi Aich Formation of southern Tunisia: phylogenetic and paleobiogeographic implications, Journal of Vertebrate Paleontology, DOI: 10.1080/02724634.2018.1529675

[11] [11]
de Figueiredo, F. J., and D. R. M. Ribeiro. 2017. Reassessment and Relationships of †Scutatuspinosus itapagipensis (Teleostei, Clupeomorpha, †Ellimmichthyiformes) from the Neocomian of Reconcavo Basin, Northeastern Brazil. Anais da Academia Brasileira de Ciencias 89:799–823.

[12] [12]
Chen, G., Chang, M. M., Wu, F., & Liao, X. (2021). Guiclupea superstes, gen. et sp. nov., the youngest ellimmichthyiform (clupeomorph) fish to date from the Oligocene of South China. PeerJ, 9, e11418. https://doi.org/10.7717/peerj.11418

[13] [13]
Giuseppe Marramà; Alexandre F. Bannikov; Jürgen Kriwet; Giorgio Carnevale (2018). "An Eocene paraclupeid fish (Teleostei, Ellimmichthyiformes) from Bolca, Italy: the youngest marine record of double‐armoured herrings". Papers in Palaeontology. 5 (1): 83–98. doi:10.1002/spp2.1230. PMC 6392134. PMID 30854219.

[14] [14]
Gengjiao Chen; Mee-mann Chang; Feixiang Wu; Xiaowen Liao; Guiclupea superstes, gen. et sp. nov., the youngest ellimmichthyiform (clupeomorph) fish to date from the Oligocene of South China

[15] [15]
Thorectichthys (Ellimmichthyiformes) in America, from the Cenomanian deposits of the Cintalapa Formation, Chiapas, southeastern Mexico. Sciencedirect. [2024-06-16]. （原始內容存檔於2024-06-16）.

[16] [16]
Jesús Alvarado-Ortega & Ernesto Ovalles-Damián (2008) Triplomystus applegatei, sp. nov. (Teleostei: Ellimmichthyiformes), a rare 「triple armored herring」 from El Espinal Quarry (Early Cretaceous), Chiapas, southeastern Mexico, Journal of Vertebrate Paleontology, 28:1, 53-60, DOI: 10.1671/0272-4634(2008)28[53:TASNTE]2.0.CO;2

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