格兰道鱼龙属

格兰道鱼龙属; 化石时期：晚侏罗世，启莫里期至提通期 PreЄ Є O S D C P T J K Pg N
	朱氏格兰道鱼龙复原图
科学分类
界：	动物界 Animalia
门：	脊索动物门 Chordata
纲：	爬行綱 Reptilia
目：	†鱼龙目 Ichthyosauria
科：	†大眼鱼龙科 Ophthalmosauridae
属：	†格兰道鱼龙属 Grendelius; McGowan, 1976
种
	†亚氏格兰道鱼龙 G. alekseevi ; (Arkangelsky, 2001) [原为奥茨舍夫鱼龙]; †噬咬格兰道鱼龙 G. mordax ; McGowan, 1976（模式种）; †伪斯基泰格兰道鱼龙 G. pseudoscythicus ; (Efimov, 1998) [原为奥茨舍夫鱼龙]; †朱氏格兰道鱼龙 G. zhuravlevi ; (Arkangelsky, 1998);
異名
	奥茨舍夫鱼龙 Otschevia ; Efimov, 1998

格兰道鱼龙属（学名：Grendelius）是大眼鱼龙科扁鳍鱼龙亚科鱼龙的一个属，生存于晚侏罗世（启莫里期至提通期）的英国和欧洲俄罗斯。它是种长约4米（13英尺）的中型鱼龙类。^[1]

分类

模式种噬咬格兰道鱼龙（Grendelius mordax）是于1976年根据CAMSM J.68516即英国启莫里黏土组（英语：Kimmeridge Clay）发现的完整颅骨及相关颅后材料所描述，^[2]后来被重新归入扁鳍鱼龙，因为当时认为二者差异过小，不足以建立新属甚至新种。^[3]^[4]

叶菲莫夫于1998年根据俄罗斯伏尔加地区（英语：Volga region）乌里扬诺夫斯克州伪斯基泰菊石带（Pseudoscythia Zone，晚侏罗世晚提通阶）发现的单件标本（正模标本）命名了鱼龙类新属伪斯基泰奥茨舍夫鱼龙（Otschevia pseudoscythica）。^[5]后来阿尔坎盖尔斯基于1998年从俄罗斯萨拉托夫提通阶岩层中描述了朱氏扁鳍鱼龙（Brachypterygius zhuravlevi），随后将其更名为朱氏奥茨舍夫鱼龙（Otschevia zhuravlevi）。^[6]^[7]迈施与马茨凯于2000年提出这两个俄罗斯分类单元互为异名，并将新组合伪斯基泰扁鳍鱼龙（B. pseudoscythicus）归入扁鳍鱼龙。^[8]格兰道鱼龙与奥茨舍夫鱼龙因此成为扁鳍鱼龙的次异名。阿尔坎盖尔斯基另于2001命名了亚氏奥茨舍夫鱼龙（Otschevia alekseevi），同样来自俄罗斯晚的侏罗世地层。^[9]麦高文与藻谷（2003年）认为这些俄罗斯分类单元均为极端扁鳍鱼龙（B. extremus）的次异名，^[4]迈施（2010年）则持保留意见，并将噬咬扁鳍鱼龙（B. mordax）视为独立物种。^[10]2015年发表的一篇论文发现格兰道鱼龙完全不同于扁鳍鱼龙模式种，在扁鳍鱼龙亚科中更为衍生，且与扁鳍鱼龙的关系近于短鳍鱼龙。奥茨舍夫鱼龙则被视为格兰道鱼龙的次异名。^[1]

以下分支图来自泽维科夫与雅各（2020年）的分析，显示了本属在大眼鱼龙科中可能的系统发育位置。^[11]

大眼鱼龙类 Ophthalmosauria

大眼鱼龙亚科 Ophthalmosaurinae

密接实泳鱼龙 Acamptonectes densus

始祖莫勒斯鱼龙 Mollesaurus periallus

	泅泳大眼鱼龙 Ophthalmosaurus natans

	艾森尼大眼鱼龙 Ophthalmosaurus icenicus

尼氏真加鱼龙 Gengasaurus nicosiai

亚氏小鳍鱼龙 Nannopterygius yasykovi

	包齿小鳍鱼龙 Nannopterygius enthekiodon

	萨茹列夫小鳍鱼龙 Nannopterygius saveljeviensis

	北方小鳍鱼龙 Nannopterygius borealis

伏尔加关节鳍鱼龙 Arthropterygius volgensis

隆氏关节鳍鱼龙 Arthropterygius lundi

南海关节鳍鱼龙 Arthropterygius thalassonotus

	霍氏关节鳍鱼龙 Arthropterygius hoybergeti

	克氏关节鳍鱼龙 Arthropterygius chrisorum

扁鳍鱼龙亚科 Platypterygiinae

极端短鳍鱼龙 Brachypterygius extremus

细椎阿戈尔鱼龙 Aegirosaurus leptospondylus

垂直穆伊斯卡鱼龙 Muiscasaurus catheti

星纹列宁鱼龙 Leninia stellans

独特巧泳鱼龙 Sveltonectes insolitus

油砂矿阿萨巴斯卡鱼龙 Athabascasaurus bitumineus

美洲扁鳍鱼龙 Platypterygius americanus

	卡氏水鱼龙 Acuetzpalin carranzai

	萨奇卡扁鳍鱼龙 Platypterygius sachicarum

	波氏礁灵鱼龙 Caypullisaurus bonapartei

	噬咬格兰道鱼龙 Grendelius mordax

	亚氏格兰道鱼龙 Grendelius alekseevi

	伪斯基泰格兰道鱼龙 Grendelius pseudoscythicus

	朱氏格兰道鱼龙 Grendelius zhuravlevi

凯氏温多雷鱼龙 Undorosaurus kielanae

	尼氏温多雷鱼龙 Undorosaurus nessovi

	戈罗季谢温多雷鱼龙 Undorosaurus gorodischensis

南方扁鳍鱼龙 Platypterygius australis

	贝登普卢托尼亚鱼龙 Plutoniosaurus bedengensis

	比氏辛比尔斯克鱼龙 Simbirskiasaurus birjukovi

	哈茨山扁鳍鱼龙 Platypterygius hercynicus

	西氏锡斯特龙鱼龙 Sisteronia seeleyi

	平趾扁鳍鱼龙 Platypterygius platydactylus

	林氏慈母椎龙 Maiaspondylus lindoei

参见

鱼龙列表
鱼龙类研究历史（英语：Timeline of ichthyosaur research）

参考资料

^ ^1.0 ^1.1 Zverkov, N. G.; Arkhangelsky, M. S.; Stenshin, I. M. A review of Russian Upper Jurassic ichthyosaurs with an intermedium/humeral contact. Reassessing Grendelius McGowan, 1976. Proceedings of the Zoological Institute. 2015, 318 (4): 558–588. doi:10.31610/trudyzin/2015.319.4.558.
^ McGowan C. 1976. The description and phenetic relation-ships of a new ichthyosaur genus from the Upper Jurassic of England. Canadian Journal of Earth Sciences 13: 668–683.
^ McGowan, C. 1997. The taxonomic status of Grendelius mordax: a preliminary report. Journal of Vertebrate Paleontology, 17, 428–430. doi:10.1080/02724634.1997.10010986
^ ^4.0 ^4.1 McGowan, C. & Motani, R. Ichthyopterygia. In Sues, H.-D. (ed.) Handbook of Paleoherpetology, vol. 8. Verlag Dr. Friedrich Pfeil, Munich, 175 pp., 19 pls.
^ Efimov VM. 1998. An Ichthyosaur, Otschevia pseudoscythica gen. et sp. nov. from the Upper Jurassic Strata of the Ulyanovsk Region (Volga Region). Paleontologicheskii Zhurnal 32 (2): 187-191.
^ Arkhangelsky, M. S. 1998. On the ichthyosaurian fossils from the Volgian stage of the Saratov Region. Paleontologicheskii Zhurnal, 1998, 87–91.
^ Arkhangelsky, M. S. 2000. On the ichthyosaur Otschevia from the Volgian Stage of the Volga Region. Paleontological Journal, 34, 549–552.
^ Maisch MW, Matzke AT. 2000. The Ichthyosauria. Stuttgarter Beiträge zur Naturkunde, Serie B (Geologie und Paläontologie) 298: 1-159
^ Arkhangelsky, M. S. 2001. On a new ichthyosaur of the genus Otschevia from the Volgian Stage of the Volga Region near Ulyanovsk. Paleontological Journal, 35, 629–635.
^ Michael W. Maisch. Phylogeny, systematics, and origin of the Ichthyosauria – the state of the art (PDF). Palaeodiversity. 2010, 3: 151–214.
^ Nikolay G. Zverkov & Megan L. Jacobs. Revision of Nannopterygius (Ichthyosauria: Ophthalmosauridae): reappraisal of the 'inaccessible' holotype resolves a taxonomic tangle and reveals an obscure ophthalmosaurid lineage with a wide distribution. Zoological Journal of the Linnean Society. 2021, 191 (1): 228–275 [2020]. doi:10.1093/zoolinnean/zlaa028.

[ZAS2015-1] 1.0 ^1.1 Zverkov, N. G.; Arkhangelsky, M. S.; Stenshin, I. M. A review of Russian Upper Jurassic ichthyosaurs with an intermedium/humeral contact. Reassessing Grendelius McGowan, 1976. Proceedings of the Zoological Institute. 2015, 318 (4): 558–588. doi:10.31610/trudyzin/2015.319.4.558.

[2] McGowan C. 1976. The description and phenetic relation-ships of a new ichthyosaur genus from the Upper Jurassic of England. Canadian Journal of Earth Sciences 13: 668–683.

[3] McGowan, C. 1997. The taxonomic status of Grendelius mordax: a preliminary report. Journal of Vertebrate Paleontology, 17, 428–430. doi:10.1080/02724634.1997.10010986

[MM03-4] 4.0 ^4.1 McGowan, C. & Motani, R. Ichthyopterygia. In Sues, H.-D. (ed.) Handbook of Paleoherpetology, vol. 8. Verlag Dr. Friedrich Pfeil, Munich, 175 pp., 19 pls.

[Efimov,_1998-5] Efimov VM. 1998. An Ichthyosaur, Otschevia pseudoscythica gen. et sp. nov. from the Upper Jurassic Strata of the Ulyanovsk Region (Volga Region). Paleontologicheskii Zhurnal 32 (2): 187-191.

[Arkhangelsky,_1998-6] Arkhangelsky, M. S. 1998. On the ichthyosaurian fossils from the Volgian stage of the Saratov Region. Paleontologicheskii Zhurnal, 1998, 87–91.

[Arkhangelsky,_2000-7] Arkhangelsky, M. S. 2000. On the ichthyosaur Otschevia from the Volgian Stage of the Volga Region. Paleontological Journal, 34, 549–552.

[Maisch_&_Matzke,_2000-8] Maisch MW, Matzke AT. 2000. The Ichthyosauria. Stuttgarter Beiträge zur Naturkunde, Serie B (Geologie und Paläontologie) 298: 1-159

[Arkhangelsky,_2001-9] Arkhangelsky, M. S. 2001. On a new ichthyosaur of the genus Otschevia from the Volgian Stage of the Volga Region near Ulyanovsk. Paleontological Journal, 35, 629–635.

[Maisch2010-10] Michael W. Maisch. Phylogeny, systematics, and origin of the Ichthyosauria – the state of the art (PDF). Palaeodiversity. 2010, 3: 151–214.

[Zverkov2020-11] Nikolay G. Zverkov & Megan L. Jacobs. Revision of Nannopterygius (Ichthyosauria: Ophthalmosauridae): reappraisal of the 'inaccessible' holotype resolves a taxonomic tangle and reveals an obscure ophthalmosaurid lineage with a wide distribution. Zoological Journal of the Linnean Society. 2021, 191 (1): 228–275 [2020]. doi:10.1093/zoolinnean/zlaa028.

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