古脑翼龙属

古脑翼龙属; 化石时期：; 托阿尔阶中期; ~179.17–178.07 Ma PreЄ Є O S D C P T J K Pg N ▼
	骨骼材料
科学分类
界：	动物界 Animalia
门：	脊索动物门 Chordata
纲：	蜥形纲 Sauropsida
目：	†翼龍目 Pterosauria
演化支：	†长爪翼龙类 Macronychoptera
演化支：	†新翼龙类 Novialoidea
演化支：	†短四跖骨类 Breviquartossa
演化支：	†翼手龙形态类 Pterodactylomorpha
属：	†古脑翼龙属 Allkaruen; Codorniú et al. 2016
模式種
	†盐湖古脑翼龙; Allkaruen koi; Codorniú et al. 2016

古脑翼龙（属名：Allkaruen，意为“古代大脑”）是“喙嘴翼龙亚目”翼龙的一个属，化石发现于阿根廷早侏罗世沥青峡谷组（英语：Cañadon Asfalto Formation）。^[2]属下包括单一物种盐湖古脑翼龙（Allkaruen koi）。

描述

如CT扫描所示，古脑翼龙脑壳拥有一套介于喙嘴翼龙等较原始翼龙及古魔翼龙等较衍生翼手龙亚目翼龙之间的独特特征。这些特征包括内耳前半规管的位置、^[3]枕骨与枕髁（英语：Occipital condyles）的相对方位、绒球（英语：Flocculus）侧缘与大脑半球的相对位置及大脑长度与后脑高度的比例。^[2]此外额骨与侧半规管的相对方位更类似喙嘴翼龙而非古魔翼龙，视叶位置像翼手龙亚目般低于前脑。^[2]^[3]这一独特组合表明翼手龙亚目脑颅解剖形态是经镶嵌进化（英语：Mosaic evolution）而来。

古魔翼龙顶骨很长，占额骨长度的60%；额骨本身又圆又宽，且存在大量气腔（英语：Skeletal pneumaticity）。下颌长度约为颅骨保存部分的3.5倍，颌骨尖部向上弯曲。齿骨前半部分生有四或五个齿槽（英语：Dental alveolus），后半部分则为一沟漕所取代；该组合在翼龙中相当独特。^[2]

古脑翼龙是种小型翼龙类。正模标本颅骨完全融合，可能是具成年个体。^[2]

发现与命名

古魔翼龙正模标本由一个脑壳（MPEF-PV 3613）、一个下颌骨（MPEF-PV 3609）及一节颈椎（MPEF-PV 3615）组成；一节类似的颈椎（MPEF-PV 3616）亦归入该属。^[2]这些材料于2000年在阿根廷丘布特省神鹰山（英语：Cerro Cóndor）的沥青峡谷组（英语：Cañadon Asfalto Formation）发现，该地层可追溯至托阿尔阶中晚期，时间为179.17 ± 0.12至178.07 ± 0.21百万年前（尽管较早估计显示为巴通阶最早期）。^[4]^[5]其它材料整体上关节连接，由大量肢骨及另一块下颌骨组成，同样发现于该地点。^[2]^[6]

属名Allkaruen源于特维尔切语（英语：Tehuelche language），取自all（脑）及karuen（古代的）。种名koi源自特维尔切语“湖泊”，暗指模式产地曾经可能是一处盐湖。^[2]^[7]

分类

2016年，根据达尔文翼龙描述的数据集进行了两项系统发育分析，以确定古脑翼龙的系统发育位置。^[8]一项分析仅限于正模标本脑壳，另一项还包括正模下颌骨及颈椎。两者一致将古脑翼龙恢复为单窗孔类的姐妹群，360个最简树的共识如下所示。^[2]

短四跖骨类 Breviquartossa

矛颌翼龙 Dorygnathus

	岛翼龙 Nesodactylus

	喙嘴翼龙 Rhamphorhynchus

	天王翼龙 Cacibupteryx

翼手喙龙 Pterorhynchus

	索德斯龙 Sordes

	船颌翼龙 Scaphognathus

古脑翼龙 Allkaruen

单窗孔类 Monofenestrata

	达尔文翼龙 Darwinopterus

	悟空翼龙 Wukongopterus

翼手龙亚目 Pterodactyloidea

在2024年迷雾翼龙描述中，马丁-西韦斯通等人在系统发育分析中发现了合理证据支持古脑翼龙代表单窗孔类中达尔文翼龙类的已知最早成员。作者进一步将达尔文翼龙类重新定义为由所有更接近包含盐湖古脑翼龙及斯氏尖头翼龙的演化支而非风神翼龙的所有分类单元组成。结果如以下分支图所示：^[9]

单窗孔类 Monofenestrata

达尔文翼龙类 Darwinoptera

	长城翼龙 Changchengopterus

	鲲鹏翼龙 Kunpengopterus

	悟空翼龙 Wukongopterus

	达尔文翼龙 Darwinopterus

	藏龙 Kryptodrakon

	捻船头翼龙 Cuspicephalus

	古脑翼龙 Allkaruen

	迷雾翼龙 Ceoptera

翼手龙亚目 Pterodactyloidea

喙手龙 'Rhamphodactylus'

翼手龙 Pterodactylus

	Derived pterodactyloids

参考资料

^ Fantasia, A.; Föllmi, K. B.; Adatte, T.; Spangenberg, J. E.; Schoene, B.; Barker, R. T.; Scasso, R. A. Late Toarcian continental palaeoenvironmental conditions: An example from the Canadon Asfalto Formation in southern Argentina. Gondwana Research. 2021, 89 (1): 47–65 [27 August 2021]. Bibcode:2021GondR..89...47F. S2CID 225120452. doi:10.1016/j.gr.2020.10.001.
^ ^2.00 ^2.01 ^2.02 ^2.03 ^2.04 ^2.05 ^2.06 ^2.07 ^2.08 ^2.09 ^2.10 Codorniú, L.; Carabajal, A.P.; Pol, D.; Unwin, D.; Rauhut, O.W.M. A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium. PeerJ. 2016, 4: e2311. PMC 5012331 . PMID 27635315. doi:10.7717/peerj.2311 .
^ ^3.0 ^3.1 Witmer, L.M.; Chatterjee, S.; Franzosa, J.; Rowe, T. Neuroanatomy of flying reptiles and implications for flight, posture and behaviour (PDF). Nature. 2003, 425 (6961): 950–953. Bibcode:2003Natur.425..950W. PMID 14586467. S2CID 4431861. doi:10.1038/nature02048.
^ Pol, D.; Gomez, K.; Holwerda, F. M.; Rauhut, O. W.; Carballido, J. L. Sauropods from the Early Jurassic of South America and the Radiation of Eusauropoda. South American Sauropodomorph Dinosaurs. 2022, 1 (1): 131–163 [2 May 2022]. doi:10.1007/978-3-030-95959-3_4.
^ Cúneoa, R.; Ramezani, J.; Scasso, R.; Pol, D.; Escapa, I.; Zavattieri, A.M.; Bowring, S.A. High-precision U–Pb geochronology and a new chronostratigraphy for the Cañadón Asfalto Basin, Chubut, central Patagonia: Implications for terrestrial faunal and floral evolution in Jurassic. Gondwana Research. 2013, 24 (3–4): 1267–1275. Bibcode:2013GondR..24.1267C. doi:10.1016/j.gr.2013.01.010. hdl:11336/78351 .
^ Codorniú, L.; Rauhut, O.W.M.; Pol, D. Osteological features of Middle Jurassic pterosaurs from Patagonia (Argentina). Acta Geoscientica Sinica. 2010, 31 (suppl. 1): 12–13.
^ Cabaleri, N.G.; Armella, C.; Silva Nieto, D.G. Saline paleolake of the Cañadón Asfalto Formation (Middle-Upper Jurassic), Cerro Cóndor, Chubut province (Patagonia), Argentina. Facies. 2005, 51 (1): 350–364. Bibcode:2005Faci...51..350C. S2CID 129090656. doi:10.1007/s10347-004-0042-5.
^ Lu, J.; Unwin, D.M.; Jin, X.; Liu, Y.; Ji, Q. Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proceedings of the Royal Society B: Biological Sciences. 2010, 277 (1680): 383–389 [2009]. PMC 2842655 . PMID 19828548. doi:10.1098/rspb.2009.1603.
^ Martin-Silverstone, Elizabeth; Unwin, David M.; Cuff, Andrew R.; Brown, Emily E.; Allington-Jones, Lu; Barrett, Paul M. A new pterosaur from the Middle Jurassic of Skye, Scotland and the early diversification of flying reptile. Journal of Vertebrate Paleontology. 2024-02-05. ISSN 0272-4634. doi:10.1080/02724634.2023.2298741  （英语）.

[1] Fantasia, A.; Föllmi, K. B.; Adatte, T.; Spangenberg, J. E.; Schoene, B.; Barker, R. T.; Scasso, R. A. Late Toarcian continental palaeoenvironmental conditions: An example from the Canadon Asfalto Formation in southern Argentina. Gondwana Research. 2021, 89 (1): 47–65 [27 August 2021]. Bibcode:2021GondR..89...47F. S2CID 225120452. doi:10.1016/j.gr.2020.10.001.

[codorniu2016-2] 2.00 ^2.01 ^2.02 ^2.03 ^2.04 ^2.05 ^2.06 ^2.07 ^2.08 ^2.09 ^2.10 Codorniú, L.; Carabajal, A.P.; Pol, D.; Unwin, D.; Rauhut, O.W.M. A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium. PeerJ. 2016, 4: e2311. PMC 5012331 . PMID 27635315. doi:10.7717/peerj.2311 .

[witmer2003-3] 3.0 ^3.1 Witmer, L.M.; Chatterjee, S.; Franzosa, J.; Rowe, T. Neuroanatomy of flying reptiles and implications for flight, posture and behaviour (PDF). Nature. 2003, 425 (6961): 950–953. Bibcode:2003Natur.425..950W. PMID 14586467. S2CID 4431861. doi:10.1038/nature02048.

[4] Pol, D.; Gomez, K.; Holwerda, F. M.; Rauhut, O. W.; Carballido, J. L. Sauropods from the Early Jurassic of South America and the Radiation of Eusauropoda. South American Sauropodomorph Dinosaurs. 2022, 1 (1): 131–163 [2 May 2022]. doi:10.1007/978-3-030-95959-3_4.

[cuneoaetal2013-5] Cúneoa, R.; Ramezani, J.; Scasso, R.; Pol, D.; Escapa, I.; Zavattieri, A.M.; Bowring, S.A. High-precision U–Pb geochronology and a new chronostratigraphy for the Cañadón Asfalto Basin, Chubut, central Patagonia: Implications for terrestrial faunal and floral evolution in Jurassic. Gondwana Research. 2013, 24 (3–4): 1267–1275. Bibcode:2013GondR..24.1267C. doi:10.1016/j.gr.2013.01.010. hdl:11336/78351 .

[codorniuetal2010-6] Codorniú, L.; Rauhut, O.W.M.; Pol, D. Osteological features of Middle Jurassic pterosaurs from Patagonia (Argentina). Acta Geoscientica Sinica. 2010, 31 (suppl. 1): 12–13.

[cabaleri2005-7] Cabaleri, N.G.; Armella, C.; Silva Nieto, D.G. Saline paleolake of the Cañadón Asfalto Formation (Middle-Upper Jurassic), Cerro Cóndor, Chubut province (Patagonia), Argentina. Facies. 2005, 51 (1): 350–364. Bibcode:2005Faci...51..350C. S2CID 129090656. doi:10.1007/s10347-004-0042-5.

[luetal2009-8] Lu, J.; Unwin, D.M.; Jin, X.; Liu, Y.; Ji, Q. Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proceedings of the Royal Society B: Biological Sciences. 2010, 277 (1680): 383–389 [2009]. PMC 2842655 . PMID 19828548. doi:10.1098/rspb.2009.1603.

[Ceoptera-9] Martin-Silverstone, Elizabeth; Unwin, David M.; Cuff, Andrew R.; Brown, Emily E.; Allington-Jones, Lu; Barrett, Paul M. A new pterosaur from the Middle Jurassic of Skye, Scotland and the early diversification of flying reptile. Journal of Vertebrate Paleontology. 2024-02-05. ISSN 0272-4634. doi:10.1080/02724634.2023.2298741  （英语）.

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