新发现的长吻型离龙(双孔亚纲：离龙目)幼年个体——袖珍蒙山龙,兼论新离龙类的个体发育

doi:10.19615/j.cnki.2096-9899.210607

古脊椎动物学报 ›› 2021, Vol. 59 ›› Issue (3): 213-228.DOI: 10.19615/j.cnki.2096-9899.210607

新发现的长吻型离龙(双孔亚纲：离龙目)幼年个体——袖珍蒙山龙,兼论新离龙类的个体发育

袁梦¹^,²^,³, 李大庆⁴^,^*(), Daniel T. KSEPKA⁵, 易鸿宇¹^,²^,^*()

1 中国科学院古脊椎动物与古人类研究所,中国科学院脊椎动物演化与人类起源重点实验室北京 100044
2 中国科学院生物演化与环境卓越创新中心北京 100044
3 中国科学院大学北京 100049
4 甘肃农业大学古脊椎动物研究所兰州 730070
5 美国布鲁斯博物馆康涅狄格州格林尼治镇 06830

收稿日期:2021-03-15 出版日期:2021-07-20 发布日期:2021-07-20
通讯作者: * daqingli@gsau.edu.cn;
yihongyu@ivpp.ac.cn
基金资助:
中国科学院战略性先导科技专项(XDA19050102);中国科学院战略性先导科技专项(XDB18030504);中国科学院战略性先导科技专项(XDB26000000);中国科学院“率先行动”项目;国家自然科学基金青年科学基金(41702020);国家自然科学基金基础科学中心项目(41688103)

A juvenile skull of the longirostrine choristodere (Diapsida: Choristodera), Mengshanosaurus minimus gen. et sp. nov., with comments on neochoristodere ontogeny

YUAN Meng¹^,²^,³, LI Da-Qing⁴^,^*(), Daniel T. KSEPKA⁵, YI Hong-Yu¹^,²^,^*()

1 Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences Beijing 100049, China
2 CAS Center for Excellence in Life and Paleoenvironment Beijing 100044, China
3 University of Chinese Academy of Sciences Beijing 100049, China
4 Institute of Vertebrate Paleontology, Gansu Agricultural University Lanzhou 730070, China
5 Bruce Museum Greenwich, Connecticut 06830, USA

Received:2021-03-15 Published:2021-07-20 Online:2021-07-20

1. 附件.pdf(295KB)

摘要/Abstract

摘要：

离龙是一类生活在中侏罗世至中新世的半水生掠食性动物。白垩纪早期,部分离龙类演化为与现生鳄鱼形态类似的大型长吻爬行动物,称为新离龙类。报道了来自山东省下白垩统蒙阴组的一长吻型离龙新种——袖珍蒙山龙(Mengshanosaurus minimus), 正型标本头骨全长仅35 mm, 是迄今为止发现的最小新离龙类个体。根据未完全骨化的颅腔和额骨-顶骨间尚存未愈合的圆孔,推测蒙山龙正型标本为一幼年个体。系统发育分析显示,蒙山龙属于新离龙类,具有单个外鼻孔,单一鼻骨和下颞孔开放等新离龙类的共有衍征。在新离龙类内部,蒙山龙分类位置较为基干,为伊克昭龙属(Ikechosaurus)、车尔龙属(Tchoiria)、西莫多龙属(Simoedosaurus)和鳄龙属(Champsosaurus)组成的支系的姐妹群。蒙山龙区别于其他新离龙类的特征包括泪孔位于前额骨与泪骨之间,以及增大的腭面齿(宽度超过上颌齿的1/3)。蒙山龙具有长吻和紧密排列的尖利牙齿,推测离龙类幼体与现生鳄类幼体食性类似,以水生昆虫和无脊椎动物为食。现生鳄类在发育过程中,吻部的形态常发生改变,牙齿也会从尖细的形状转向圆钝;而蒙山龙的吻部和牙齿的形态与已知大体型的成体新离龙类没有明显区别,这说明相较于现生鳄类,新离龙类在发育过程中生态位的变化并不显著。

关键词: 山东新泰, 早白垩世, 蒙阴组, 新离龙类

Abstract:

Choristoderes were an important clade of semi-aquatic predators that occupied Laurasian freshwater ecosystems from the Middle Jurassic to the Miocene. During the Early Cretaceous, the neochoristodere lineage evolved large size and long snouts, converging on the body plan of modern crocodilians. Here, we describe a new longirostrine choristodere, Mengshanosaurus minimus gen. et sp. nov. from the Lower Cretaceous Mengyin Formation of Shandong Province, China. The holotype is the smallest reported neochoristodere individual, with a skull length of only 35 mm. The poorly ossified braincase, along with retention of a fontanel at the frontal-parietal suture, indicates this individual was a juvenile. Phylogenetic analyses recovered Mengshanosaurus as a neochoristodere, a placement supported by the presence of a single narial opening, fusion of the nasals, and expansion of the temporal fenestrae. In the Neochoristodera,Mengshanosaurus is sister to a clade consisting of Ikechosaurus, Tchoiria, Simoedosaurus, and Champsosaurus. It differs from other neochoristoderes in having the lacrimal foramen between the prefrontal and lacrimal, in addition to having large vomerine teeth (exceeding one-third the width of corresponding maxillary teeth). The closely arranged marginal teeth and large vomerine teeth suggest juvenile choristoderes may have fed on invertebrates and insects, similar to juveniles of modern crocodilians. However, the observation that very young neochoristoderes had similar skull proportions and marginal tooth shapes to adults, along with features suggesting a more fully aquatic ecology, suggest that neochoristoderes exhibited less pronounced ontogenetic niche shifts than modern crocodilians.

Key words: Xintai,Shandong, Early Cretaceous, Mengyin Formation, Neochoristodera

中图分类号:

Q915.864

袁梦, 李大庆, Daniel T. KSEPKA, 易鸿宇. 新发现的长吻型离龙(双孔亚纲：离龙目)幼年个体——袖珍蒙山龙,兼论新离龙类的个体发育. 古脊椎动物学报, 2021, 59(3): 213-228.

YUAN Meng, LI Da-Qing, Daniel T. KSEPKA, YI Hong-Yu. A juvenile skull of the longirostrine choristodere (Diapsida: Choristodera), Mengshanosaurus minimus gen. et sp. nov., with comments on neochoristodere ontogeny. Vertebrata Palasiatica, 2021, 59(3): 213-228.

图/表 9

Fig. 1 A new choristodere specimen from Shandong Province A. map of the fossil locality in Shandong, China; B. photos of the new specimen IVPG-T002 in dorsal (left) and ventral (right) views; C. paleogeographic map of Asia in the Early Cretaceous (not to scale), modified from Matsumoto et al. (2015), red triangle represents the fossil locality Abbreviations: EUR. Europe; INC. Indo-China; J. Japan; K. Korea; KAZ. Kazakhstan; MON. Mongolia; NCB. North China Block; SCB. South China Block; SH. Shan Thai; SIB. Siberia

Fig. 2 Holotype of Mengshanosaurus minimus (IVPG-T002) from Xintai, Shandong CT model (A, C) and line drawing (B, D) of the skull in dorsal (A, B) and ventral (C, D) views Abbreviations: bo. basioccipital; cod. mandibular condyle; ect. ectopterygoid; ex n. external naris; exo. exoccipital; f. frontal; ipt v. interpterygoid vacuity; j. jugal; la. lacrimal; m. maxilla; n. nasal; p. parietal; pbsh. parabasisphenoid; pl. palatine; pmx. premaxilla; pof. postorbitofrontal; prf. prefrontal; pt. pterygoid; q. quadrate; qj. quadratojugal; so. supraoccipital; sof. suborbital fenestra; sq. squamosal; v. vomer. Scale bars = 5 mm

Fig. 3 The lacrimal foramen of Mengshanosaurus minimus (IVPG-T002) from Xintai, Shandong A. three-dimensional reconstruction of the anterior margin of the left orbit; B. two-dimensional CT slice through the anterior margin of left orbit. The red arrows mark the lacrimal foramen Abbreviations see Fig. 2. Scale bars = 1 mm

Fig. 4 Size comparisons between the palatal and marginal teeth of Mengshanosaurus minimus (IVPG-T002) from Xintai, Shandong A. line drawing of the skull in ventral view (not to scale); B-D. the palatal teeth (white arrows) and the maxillary teeth and tooth sockets (red arrows) in the same coronal plane. Scale bars = 1 mm

Table 1 Measurements of vomerine and maxillary teeth in the same cross section (mm)

		Vomerine tooth width	Maxillary tooth width	Proportion
Mengshanosaurus minimus (IVPG-T002)	V1	0.26	0.70	0.37
	V2	0.20*	0.73	0.27
	V3	0.19*	0.73	0.26
	V4	0.22	0.64	0.34
	Average proportion			0.31
Tchoiria klauseni (IGM 1/8)	V1	1.40	9.99	0.14
	V2	1.03	9.32	0.11
	Average proportion			0.13
Champsosaurus lindoei (CMN 8920)	V1	0.65	4.25	0.15
	V2	0.77	3.54	0.22
	Average proportion			0.19
Ikechosaurus sunailinae (IVPP V 9611-3)	V1	0.38	2.40	0.16
	V2	0.33	2.30	0.15
	Average proportion			0.16

Fig. 5 Relative position between the occipital condyle and the jaw articulation among neochoristoderes A. a reconstructed skull of neochoristoderes denoting the position of the craniomandibular joint (red circle) B, C. Ikechosaurus sunailinae: B. IVPP V 9611-3, C. IVPP V 9611-1, juvenile; D. Champsosaurus lindoei (CMN 8920); E. Tchoiria namsarai (PIN 3386/1, modified from Skutschas and Efimov, 2015); F. Mengshanosaurus minimus (IVPG-T002). The dashed lines indicate the position of the craniomandibular joint, and the red arrows denote the occipital condyle

Table 2 Tooth count from several specimens of neochoristoderes

	Upper tooth row	Premaxilla	Maxilla	Lower tooth row	Mandibular symphysis	Posterior to the mandibular symphysis
Ikechosaurus sunailinae (IVPP V 9611-1)	68	4	64	~ 70	~ 22	~ 48
Ikechosaurus sunailinae (IVPP V 9611-3)	>43*	NA	>43*	>64**	NA	42
Ikechosaurus pijiagouensis (IVPP V 13283)	NA	NA	NA	~ 60	NA	NA
Tchoiria namsarai (PIN 3386/1)	>60*	NA	>60*	75	17	60
Tchoiria klauseni (IGM 1/8)	>34*	NA	>34*	NA	12	NA
Simoedosaurus lemoinei (MNHN R1935)	~45	4	~41	NA	NA	NA
Simoedosaurus dakotensis (SMMP.76.10.1)	~49	4	~45	NA	NA	NA
Champsosaurus lindoei (CMN 8920)	47	6	41	NA	NA	NA
Mengshanosaurus minimus (IVPG-T002)	42-43	4-5	38-39	45	12	33

Fig. 6 Strict consensus tree from the phylogenetic analysis, resulting in Mengshanosaurus minimus being recovered as the sister taxon to other neochoristoderes (CI=0.812, RI=0.844) Blue branches represent longirostrine neochoristoderes and red branches represent brevirostrine choristoderes Numbers above tree branches are bootstrap values over 50, and those below the branches are Bremer support values

Table 3 Skull and total length of three species of choristoderes in various developmental stages (mm)

Species	Developmental stage	Skull length	Total length
Hyphalosaurus baitaigouensis	Adult LPMC R00066¹⁾	66	1100
	Juvenile BMNHC V050¹⁾	27.0	~450
	“small free-living individual”²⁾	14.62	NA
	Embryo³⁾	7.745	NA
Mengshanosaurus minimums (IVPG-T002)	Juvenile	35	NA
Ikechosaurus pijiagouensis (IVPP V 13283)	Adult⁴⁾	275	1700

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新发现的长吻型离龙(双孔亚纲：离龙目)幼年个体——袖珍蒙山龙,兼论新离龙类的个体发育

A juvenile skull of the longirostrine choristodere (Diapsida: Choristodera), Mengshanosaurus minimus gen. et sp. nov., with comments on neochoristodere ontogeny

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