重庆云阳中侏罗统三列齿兽类化石新发现及其古地理意义<sup>&#x0002A;</sup>

doi:10.7605/gdlxb.2024.05.048

古地理学报 ›› 2025, Vol. 27 ›› Issue (2): 484-498. doi: 10.7605/gdlxb.2024.05.048

• 生物古地理学及古生态 • 上一篇下一篇

重庆云阳中侏罗统三列齿兽类化石新发现及其古地理意义^*

刘璐¹(), 魏光飚²(), 王萍³, 魏曌英³, 任纪澄⁴, 毛方园⁴()

1 国家自然博物馆,北京 100050
2 重庆市地质调查院,重庆 401122
3 重庆市地质矿产勘查开发局208水文地质工程地质队,川渝共建古生物与古环境协同演化重庆市重点实验室,重庆 400700
4 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

收稿日期:2024-01-25 修回日期:2024-03-05 出版日期:2025-04-01 发布日期:2025-04-01
作者简介:
刘璐,女,1994年生,助理研究员,主要研究方向为早期哺乳动物与植物的协同演化。 E-mail: liulu@nnhm.org.cn。魏光飚,男,1969年生,研究馆员,主要研究方向为古脊椎动物化石的形态学分类与系统演化。 E-mail: elephantfossil@aliyun.com。
基金资助:
*重庆市自然科学基金项目(编号: CSTB2022NSCQ-MSX1537)、重庆市规划和自然资源局《重庆市重要古生物化石资源保护研究》项目、重庆市市级人才计划“包干制”项目(编号: cstc2022ycjh-bgzxm0170)、国家自然科学基金(编号: 42122010,42072002)、中国科学院青年促进会项目(编号: 2019076)、北科萌芽计划(编号: BGS202201)共同资助

New discovery of tritylodontids from the Milldle Jurassic in Yunyang area,Chongqing and their palaeogeographic significance

LIU Lu¹(), WEI Guangbiao²(), WANG Ping³, WEI Zhaoying³, REN Jicheng⁴, MAO Fangyuan⁴()

1 National Natural History Museum of China,Beijing 100050,China
2 Chongqing Institute of Geological Survey,Chongqing 401122, China
3 Chongqing Key Laboratory of Paleontology and Paleoenvironment Co-evolution,No.208 Hydrogeological and Engineering Geological Team,Chongqing Bureau of Geological and Mineral Resource Exploration and Development(Sichuan-Chongqing Joint Construction), Chongqing 400700,China
4 Key Laboratory of Evolutionary Systematics of Vertebrates,Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences,Beijing 100044,China

Received:2024-01-25 Revised:2024-03-05 Online:2025-04-01 Published:2025-04-01
About author:
LIU Lu,born in 1994,is an assistant research fellow of National Natural History Museum of China,whose current research focuses on the evolution and palaeogeographic distribution of Mesozoic tritylodontids. E-mail: liulu@nnhm.org.cn. WEI Guangbiao,born in 1969,curator of Chongqing Institute of Geological Survey,is engaged in the researches on Paleontology and Stratigraphy(including paleoanthropology).E-mail: elephantfossil@aliyun.com.
Supported by:
Co-funded by the Natural Science Foundation of Chongqing,China(No. CSTB2022NSCQ-MSX1537),Chongqing Bureau of Urban Planning and Natural Resources “Chongqing important paleontological fossil resource conservation research”project,Chongqing Municipal Talent Plan “the lump-sum project-funding system”project(No. cstc2022ycjh-bgzxm0170),the National Natural Science Foundation of China(Nos. 42122010,42072002),the Youth Innovation Promotion Association,CAS(No.2019076)and BJAST Budding Talent Program(No. BGS202201)

1. 刘璐附表1.pdf(200KB)

摘要/Abstract

摘要：

三列齿兽类(tritylodontids)是一类特化的、植食性的进步下孔类,在晚三叠世—早侏罗世呈全球性分布,之后趋于衰退,在亚洲东北部地区的分布可以延续至早白垩世。三列齿兽类被认为是与哺乳动物亲缘关系较近的下孔类,可为认识哺乳动物起源提供重要证据。借助CT扫描和三维重建技术,文中报道了2件发现于重庆云阳下沙溪庙组和新田沟组的三列齿兽类头骨化石,其中一件标本保存较完整的头骨,具有颧弓宽大且腹向扩展、上颊齿短宽、上颊齿的齿尖型式2-3-3等特征,而另一件标本仅保存下颌骨。经过形态特征对比和研究,认为这2件新标本应归入似卞氏兽属,但由于标本数量有限且保存不太完整,缺乏其他特征佐证,并且三列齿兽种内差异很大,所以笔者暂将2件新标本定为似卞氏兽属的2个未定种Bienotheroides sp.1和Bienotheroides sp.2。结合已有研究成果,讨论了三列齿兽类时空分布的演化,认为在早侏罗世三列齿兽类的明显灭绝可能与图阿尔期全球变暖相关,中—晚侏罗世三列齿兽类有明显的向北迁移的趋势。上述化石的发现不仅丰富了该地区的三列齿兽类化石材料,而且为探讨该类群的分类多样性、演化分异等提供了重要支撑。

关键词: 重庆, 中侏罗统, 下沙溪庙组, 新田沟组, 三列齿兽类

Abstract:

The Tritylodontidae represent a highly specialized group of herbivorous,derived synapsids that exhibited a global distribution during the Late Triassic and Early Jurassic. Following this era,their presence declined,with the last known occurrences recorded in northeastern Asia during the Early Cretaceous. Tritylodontids are considered one of the closest clades to mammals in terms of phylogeny,making them essential for understanding mammalian origins. This paper utilizes CT scanning and 3D reconstruction techniques to describe two new tritylodontid specimens from the Middle Jurassic Xintiangou Formation and Lower Shaximiao Formation in Yunyang,Chongqing. One specimen preserves a more complete skull,characterized by broad,ventrally extended zygomatic arches,short and broad upper cheek teeth,and an upper cheek tooth cusp formula of 2-3-3. The other specimen retains only the lower jaw. Comparative analysis of morphological features indicates that both new specimens should be classified as Bienotheroides. The number of specimens is limited,and the preservation is incomplete,with no additional characteristics available as evidence. The two new specimens are tentatively designated as Bienotheroides sp.1 and Bienotheroides sp.2. Furthermore,this paper discusses the spatio-temporal evolution of tritylodontids,suggesting that the apparent extinction of Early Jurassic tritylodontids may be linked to the Toarcian global warming event. Additionally,a clear northward migration trend is observed during the Middle to Late Jurassic. The discovery of these fossils not only enriches the tristichid fossil assemblage in this region but also provides significant support for discussions regarding the taxonomic diversity and evolutionary differentiation of this group.

Key words: Chongqing, Middle Jurassic, Lower Shaximiao Formation, Xintiangou Formation, tritylodontids

刘璐, 魏光飚, 王萍, 魏曌英, 任纪澄, 毛方园. 重庆云阳中侏罗统三列齿兽类化石新发现及其古地理意义^*[J]. 古地理学报, 2025, 27(2): 484-498.

LIU Lu, WEI Guangbiao, WANG Ping, WEI Zhaoying, REN Jicheng, MAO Fangyuan. New discovery of tritylodontids from the Milldle Jurassic in Yunyang area,Chongqing and their palaeogeographic significance[J]. Journal of Palaeogeography（Chinese Edition）, 2025, 27(2): 484-498.

http://www.gdlxb.cn/CN/Y2025/V27/I2/484

图/表 9

图1 重庆云阳三列齿兽类化石采集点交通位置图

Fig.1 Maps showing the tritylodontid fossil locality in Yunyang area,Chongqing

图2 重庆云阳侏罗系岩性柱状图及化石分布位置

Fig.2 Stratigraphic column and fossil beds of the Jurassic in Yunyang area,Chongqing

图3 重庆云阳下沙溪庙组三列齿兽类似卞氏兽头骨(16YPDC-01)
A—背视图; B—腹视图; C—右侧视图; D—枕面视图。构造命名缩写同图4

Fig.3 Skull of tritylodontid Bienotheroides sp.1(16YPDC-01)from the Lower Shaximiao Formation in Yunyang area,Chongqing

图4 重庆云阳下沙溪庙组三列齿兽类似卞氏兽(16YPDC-01)头骨线条图
A—背视图; B—腹视图; C—右侧视图; D—枕面视图。缩写: a.p.p.-anterior paroccipital process前副枕骨突,BO-basioccipital基枕骨,BS-basisphenoid基蝶骨,bs.w.-basisphenoidale wing基蝶骨翼,c.ep.-cavum epiptericum上翼腔,EP-epipterygoid上翼骨,EXO-exoccipital外枕骨,e.a.m.-external auditory meatus外耳道,F-frontal额骨,f.m.-foramen magnum枕骨大孔,i.o.f.-infraorbital foramen眶下孔,J-jugal轭骨,L-lacrimal泪骨,l.f.pe.-lateral flange of petrosal岩骨侧翼,M-maxilla上颌骨,N-nasal鼻骨,ob.f.-orbital fissure眶裂,P-parietal顶骨,PAL-palatine腭骨,PAR-parasphenoid副蝶骨,PC-upper postcanines上犬齿后齿,PM-premaxilla前颌骨,p.p.f.-pterygo-paroccipital foramen翼骨—侧枕骨孔,p.p.p.-posterior paroccipital process后副枕骨突,PE-petrosal岩骨,pro.f.-prootic foramen前耳孔,PT-pterygoid翼骨,p.t.f.-post temporal foramen后颞孔,i.n.-internal naris内鼻孔,Q-quadrate方骨,q.r.ep.-quadrate ramus of epipterygoid上翼骨方骨支,SO-supraoccipital上枕骨,SQ-squamosal鳞骨,V-vomer犁骨

Fig.4 Line drawing of the skull of the tritylodontid Bienotheroides sp.1(16YPDC-01)from the Lower Shaximiao Formation in Yunyang area,Chongqing

图5 重庆云阳下沙溪庙组三列齿兽类似卞氏兽新材料(16YPDC-01)的三维重建与CT扫描图
A—头骨腹面观; B—右上颊齿齿列腹视图; C—左上颊齿齿列腹视图; D—右上颊齿齿列; E—左上颊齿齿列; 箭头指可能的未萌发齿。缩写: PC-postcanines下犬齿后齿,L1,L2,L3-舌侧列颊齿前尖、中间尖和后尖; M1,M2,M3-中间列颊齿前尖、中间尖和后尖; B1,B2-颊侧列颊齿前尖和后尖

Fig.5 3-D reconstructions and CT scan images of new materials of tritylodontid Bienotheroides sp.1(16YPDC-01)from the Lower Shaximiao Formation in Yunyang area,Chongqing

图6 重庆云阳新田沟组三列齿兽类似卞氏兽下颌骨化石(20YX-1#-0510-1)照片和线条图
缩写: I-incisor门齿,ap-angular process隅突,cp-coronoid process冠状突,fm-mental foramen颏孔,ma.fos.-masseteric fossa咬肌窝,pc-lower postcanines下犬齿后齿

Fig.6 Photograph and line drawing of lower jaw of tritylodontid Bienotheroides sp.2(20YX-1#-0510-1)from the Xintiangou Formation in Yunyang area,Chongqing

图7 重庆云阳新田沟组三列齿兽类似卞氏兽下颌骨新材料(20YX-1#-0510-1)的三维重建与CT扫描图
A—下颌骨颊侧视图; B—下颌骨舌侧视图(箭头示齿骨后骨槽);C,D—下颊齿冠面视图; E,F—下颌骨右侧视图,箭头指可能的未萌发齿

Fig.7 3-D reconstructions and CT scan images of new lower jaw materials of tritylodontid Bienotheroides sp.2(20YX-1#-0510-1) from the Xintiangou Formation in Yunyang area,Chongqing

表1 重庆云阳三列齿兽类新标本与已报道的5种似卞氏兽的形态特征对比

Table1 Morphological comparisons between new specimens of tritylodontid in Yunyang area, Chongqing and five reported species of Bienotheroides

属种	Bienotheroides wansienensis 万县似卞氏兽	Bienotheroides zigongensis 自贡似卞氏兽	Bienotheroides ultimus	Bienotheroides shartegensis	Bienotheroides xingshanensis	Bienotheroides sp.1	Bienotheroides sp.2
吻部	短而宽	极度缩短,宽	短而宽	短而宽	短而宽	—	—
上颊齿齿冠	短宽	长窄	长宽近相等	短宽	短宽	短宽	—
保存的颊齿数量	7	7	1	4	6	7	5(下颊齿)
齿尖结构	3-3-3(上颊齿);2-2(下颊齿)	2-3-3(上颊齿);2-2(下颊齿)	2-3-3(上颊齿)	2-3-3 (上颊齿)	2-3-3 (上颊齿)	2-3-3 (上颊齿)	2-2(下颊齿)
齿尖形态	齿尖退化,每个齿尖列有2个主要齿尖和其前方的一个小尖	唇侧列后齿尖显著大于前齿尖; 中间列后2个齿尖呈新月形; 前齿尖小、锥形	中间列3个齿尖,前齿尖明显缩小,舌侧列3个齿尖,前齿尖明显缩小	仅中间列的前尖明显缩小,舌侧列的前尖较为发育	中间列和舌侧列的两个前尖都缩小,L1<M1	中间列和舌侧列的两个前尖都缩小,L1>M1	唇舌侧齿列大小和形态相当,前尖大于后尖(下颊齿)
产地	重庆万州区高岭	四川自贡大山铺和新疆准噶尔盆地	新疆准噶尔盆地东部	Shar Teg(Gobi-Altai Aimag)蒙古西南部	湖北秭归盆地	重庆云阳县	重庆云阳县
文献来源	杨钟健,1982;孙艾玲,1984;孙艾玲和李雨,1985	孙艾玲,1986;孙艾玲和崔贵海,1989	Maisch et al., 2004	Watabe et al.,2007	Liu et al.,2023	本文	本文

图8 三列齿兽类的分布和网络分析(改自Liu et al., 2023)
A—已报道的三列齿兽类的化石分布图,数字代表不同属种: 1,Tritylodon;2,Bienotherium;3,Lufengia;4,Yunnanodon;5,Oligokyphus;6,Bocatherium;7,Dinnebitodon;8,Kayentatherium;9,Stereognathus;10,Bienotheroides;11,Polistodon;12,Yuanotherium;13,Shartegodon;14,Nuurtherium;15,Montirictus;16,Xenocretosuchus;17,Fossiomanus;18,Tritylodontidae indet。B—侏罗纪时期的三列齿兽类和产地的网络分析图,依据王骞和黄冰(2020)的方法,使用Gephi软件制作完成(www.Gephi.org),其中: 较大的圆圈代表不同的地理区块,较小的圆圈代表侏罗纪时期的三列齿兽类化石属,地理区块之间通过连接共享属的线连接,只出现在1个地理区块中的三列齿兽类化石属用1条线绘制在相应地理区块的外部。三列齿兽类的属种类型、时代和产地信息详见附表1

Fig.8 Distribution map and network analysis of family Tritylodontidae(modified from Liu et al., 2023)

参考文献 62

[1]	崔贵海. 1976. 云南禄丰兽孔类一新属. 古脊椎动物与古人类, 14(2): 85-90.
	[Cui G H. 1976. Yunnania,a new tritylodont genus from Lufeng,Yunnan. Vertebrata PalAsiatica, 14(2): 85-90,135]
[2]	崔贵海. 1981. 记三列齿科一新属. 古脊椎动物与古人类, 19(1): 5-10.
	[Cui G H. 1981. A new genus of Tritylodontidae. Vertebrata PalAsiatica, 19(1): 5-10]
[3]	崔贵海,孙艾玲. 1987. 三列齿类的颊齿齿根. 古脊椎动物学报, 25(4): 245-259.
	[Cui G H,Sun A L. 1987. Postcanine root system in tritylodonts. Vertebrata PalAsiatica, 25(4): 245-259]
[4]	何信禄,蔡开基. 1984. 自贡大山铺的三列齿兽类化石. 成都地质学院学报,11(S2): 33-45.
	[He X L,Cai K J. 1984. The tritylodont remains from Dashanpu,Zigong. Journal of Chengdu College of Geology,11(S2): 33-45]
[5]	李锦玲,刘俊. 2015. 中国古脊椎动物志. 第3卷. 基干下孔类、哺乳类. 第1册,基干下孔类(总第14册). 北京: 科学出版社,1-105.
	[Li J L,Liu J. 2015. Basal synapsids. In: Qiu Zhan-xiang and Li Chuan-kui(eds). Palaeovertebrata Sinica,Volume ⅢBasal synapsids and mammals,Fascicle 1(serial no. 14). Beijing: Science Press, 1-105]
[6]	孙艾玲. 1984. 四川三列齿兽类似卞氏兽(兽形类爬行动物)的头骨. 中国科学(B辑), 14(3): 257-268.
	[Sun A L. 1984. Skull morphology of the tritylodont genus Bienotehroides of Sichuan. Science China B, 14(3): 257-268]
[7]	孙艾玲. 1986. 似卞氏兽(三列齿兽类爬行动物)新材料. 古脊椎动物学报, 24: 165-170.
	[Sun A L. 1986. New material of Bienotehroides (tritylodontid reptile)from the Shaximiao Formation of Sichuan. Vertebrata PalAsiatica, 24: 165-170]
[8]	孙艾玲,崔贵海. 1989. 三列齿兽类爬行动物化石在新疆的发现. 古脊椎动物学报, 27(1): 1-8.
	[Sun A L,Cui G H. 1989. Tritylodont reptile from Xinjiang. Vertebrata PalAsiatica, 27(1): 1-8]
[9]	孙艾玲,李雨和. 1985. 四川侏罗纪三列齿兽类头后骨骼. 古脊椎动物学报, 23(2): 135-152.
	[Sun A L,Li Y H. 1985. The postcranial skeleton of the late ritylodont Bienotheroides. Vertebrata PalAsiatica, 23(2): 135-152]
[10]	王骞,黄冰. 2020. 浅谈网络分析法及其在古生物学中的应用. 古生物学报, 59(3): 380-392.
	[Wang Q,Huang B. 2020. Network analysis and its application in paleontology: a preliminary introduction. Acta Palaeontologica Sinica, 59(3): 380-392]
[11]	王永栋,付碧宏,谢小平. 2010. 四川盆地陆相三叠系与侏罗系. 合肥: 中国科学技术大学出版社,1-178.
	[Wang Y D,Fu B H,Xie X P. 2010. The terrestrial Triassic and Jurassic System in Sichuan Basin,China. Hefei: University of Science and Technology of China Press,1-178]
[12]	杨钟健. 1974. 云南禄丰兽孔类新材料. 古脊椎动物与古人类, 12: 111-114.
	[Young C C. 1974. New material of therapsids from Lufeng,Yunnan,China. Vertebrata PalAsiatica, 12: 111-114]
[13]	杨钟健. 1982. 似卞氏兽. 杨钟健文集,北京: 科学出版社,10-13.
	[Young C C. 1982. On a Bienotherium-like tritylodont from Szechuan,China. In: Selected Workd of Yang Zhongjian. Beijing: Science Press, 10-13]
[14]	余海东,张洪亮,肖明,李德亮,周羽漩,杨文光. 2021. 重庆云阳恐龙化石埋藏地新田沟组孢粉化石组合特征及意义. 矿物岩石, 41(2): 99-108.
	[Yu H D,Zhang H L,Xiao M,Li D L,Zhou Y X,Yang W G. 2021. Characteristics and significance of palynological assemblage of Xintiangou Formation at the Yunyang dinosaur burial site,Chongqing. Mineralogy and Petrology, 41(2): 99-108]
[15]	周明镇. 1962. 云南禄丰一巨大的卞氏兽类化石. 古脊椎动物与古人类, 6(4): 365-367.
	[Chow M C. 1962. A tritylodont specimen from Lufeng,Yunnan. Vertebrata PalAsiatica, 6(4): 365-367]
[16]	周明镇,胡承志. 1959. 云南禄丰三列齿科一新属. 古脊椎动物与古人类, 1(1): 7-10.
	[Chow M C,Hu C Z. 1959. A new tritylodontid from Lufeng,Yunnan. Vertebrata PalAsiatica, 1(1): 7-10]
[17]	周羽漩,胡海虔,代辉,谭超,杨文光,张洪亮. 2020. 重庆云阳新津乡中侏罗统新田沟组恐龙的发现及沉积环境. 地质通报, 39(10): 1527-1537.
	[Zhou Y X,Hu H Q,Dai H,Tan C,Yang W G,Zhang H L. 2020. The discovery of dinosaurs from the Middle Jurassic Xintiangou Formation in Xinjin Township,Yunyang,Chongqing,and its sedimentary environment. Geological Bulletin of China, 39(10): 1527-1537]
[18]	Bonaparte J F,Martinelli A G,Schultz C L,Rubert R. 2003. The sister group of mammals: small cynodonts from the Late Triassic of Southern Brazil. Revista Brasileira de Paleonología, 5: 5-27.
[19]	Bonaparte J F,Martinelli A G,Schultz C L. 2005. New information on Brasilodon and Brasilitherium(Cynodontia,Probainognathia)from the Late Trassic of southern Brazil. Revista Brasileira de Paleonología, 8: 25-46.
[20]	Clark J M,Hopson J A. 1985. Distinctive mammal-like reptile from Mexico and its bearing on the phylogeny of the Tritylodontidae. Nature, 315: 398-400.
[21]	Cope E D. 1884. The Tertiary Marsupialia. American Naturalist, 18: 686-697.
[22]	Crompton A W. 1972. Postcanine occlusion in cynodonts and tritylodonts. Bulletin of the British Museum(Natural History). Geology, 21: 30-71.
[23]	Crompton A W,Ellenberger F. 1957. On a new cynodont from the Molteno Beds and the origin of tritylodontids. Annals of the South African Museum, 44: 1-14.
[24]	Fedak T J,Sues H D,Olsen P E. 2015. First record of the tritylodontid cynodont Oligokyphus and cynodont postcranial bones from the McCoy brook formation of Nova Scotia,Canada. Canadian Journal of Earth Science, 52: 244-249.
[25]	Hoffman E A,Rowe T B. 2018. Jurassic stem-mammal perinates and the origin of mammalian reproduction and growth. Nature, 561: 104-108.
[26]	Hopson J A. 1991. Systematics of the nonmammalian Synapsida and implications for patterns of evolution in synapsids. In: Schultze H P,Trueb L(eds). Origins of the Higher Groups of Tetrapods: Controversy and Consensus. Ithaca,New York: Comstock Publishing Associates,635-693.
[27]	Hopson J A,Crompton A W. 1969. Origin of mammals. In: Dobzhansky T,Hecht M K,Steere W C(eds). Evolutionary Biology. New York: Appleton-Century-Croft,15-71.
[28]	Hopson J A,Kitching J W. 2001. A probainognathian cynodont from South Africa and the phylogeny of nonmammalian cynodonts. Bulletin Museum of Comparative Zoology, 156: 5-35.
[29]	Hu Y M,Meng J,Clark J M. 2009. A new tritylodontid from the Upper Jurassic of Xinjiang,China. Acta Palaeontologica Polonica, 54: 385-391.
[30]	Kemp T S. 1982. Mammal-like Reptiles and the Origin of Mammals. London: Academic Press, 1-363.
[31]	Kemp T S. 1983. The relationships of mammals. Zoological Journal of the Linnean Society, 77: 353-384.
[32]	Kemp T S. 2005. The Origin and Evolution of Mammals. Oxford: Oxford University Press,1-344.
[33]	Kermack D M. 1982. A new tritylodontid from the Kayenta ormation of Arizona. Zoological Journal of the Linnean Society, 76: 1-17.
[34]	Kiel S. 2017. Using network analysis to trace the evolution of biogeography through geologic time: a case study. Geology, 45: 711-714.
[35]	Kühne W G. 1956. The Liassic Therapsid Oligokyphus. London: British Museum of Natural History,1-149.
[36]	Liu J,Olsen P. 2010. The phylogenetic relationships of Eucynodontia(Amniota: Synapsida). Journal of Mammal Evolution, 17: 151-176.
[37]	Liu J W,Wang L H,Beatty B L,Zhang G,Wang T,Bi S D. 2023a. New material(Tritylodontidae,Mammaliamorpha)from the lower Jurassic of Lufeng and its implication on the taxonomy of Lufengia. Historical Biology, 35(9): 1605-1615.
[38]	Liu L,Zhou C F,Wang J S,Xue J Z. 2023b. A new tritylodontid from the Middle Jurassic Shaximiao Formation of western Hubei,China. Historical Biology, 35(8): 1391-1400.
[39]	Luo Z X. 1994. Sister-group relationships of mammals and transformations of diagnostic mammalian characters. In: Fraser N C, Sues H D(eds). In the Shadow of the Dinaosaurs-Early Mesozoic Tetrapods. Cambridge: Cambridge University Press,98-128.
[40]	Luo Z X,Sun A L. 1993. Oligokyphus (Cynodontia: Tritylodontidae)from the Lower Lufeng Formation(Lower Jurassic)of Yunnan,China. Journal of Vertebrate Paleontology, 13: 477-482.
[41]	Luo Z X,Wu X C. 1994. The small tetrapods of the Lower Lufeng Formation,Yunnan,China. In: Fraser N C, Sues H D(eds). In the Shadow of the Dinosaurs-Early Mesozoic Tetrapods. Cambridge: Cambridge University Press,251-270.
[42]	Lopatin A V,Agadjanian A K. 2008. A tritylodont(Tritylodontidae,Synapsida)from the Mesozoic of Yakutia. Doklady Biological Sciences, 419: 107-110.
[43]	Maisch M W,Matzke A T,Sun G. 2004. A new tritylodontid from the Upper Jurassic Shishugou Formation of the Junggar Basin(Xinjiang,NW China). Journal of Vertebrate Paleontology, 24: 649-656.
[44]	Mao F Y,Zhang C,Liu C Y,Meng J. 2021. Fossoriality and evolutionary development in two Cretaceous mammaliamorphs. Nature, 592: 577-582.
[45]	Matsuoka H,Kusuhashi N,Corfe I J. 2016. A new Early Cretaceous tritylodontid(Synapsida,Cynodontia,Mammaliamorpha)from the Kuwajima Formation(Tetori Group)of Central Japan. Journal of Vertebrate Paleontology, 36: e1112289.
[46]	Matsuoka H,Setoguchi T. 2000. Significance of Chinese tritylodontids(Synapsida,Cynodontia)for the systematic study of Japanese materials from the Lower Cretaceous Kuwajima Formation,Tetori Group of Shiramine,Ishikawa,Japan. Asian Primate Paleontology, 1: 161-176.
[47]	Owen R. 1884. On the skull and dentition of a Triassic mammal(Tritylodon longaevus)from South Africa. Quarterly Journal of the Geological Society of London, 40: 146-152.
[48]	Rowe T. 1988. Definition,diagnosis,and origin of Mammalia. Journal of Vertebrate Paleontology, 8: 241-264.
[49]	Ruta M,Botha-Brink J,Mitchell S A,Benton M J. 2013. The radiation of cynodonts and the ground plan of mammalian morphological diversity. Proceedings Biological Sciences, 280: 20131865.
[50]	Sidor C A,Hopson J A. 1998. Ghost lieages and “mammalness”: assessing the temporal pattern of character acquisition in the Synapsida. Paleobiology, 24: 254-273.
[51]	Simpson G G. 1928. A catalogue of the Mesozoic Mammalia in the Geological Department of the British Museum. London: Trustees of the British Museum,1-215.
[52]	Sues H D. 1985. The relationships of the Tritylodontidae(Synapsida). Zoological Journal of the Linnean Society, 85: 205-217.
[53]	Sues H D. 1986a. The skull and dentition of two tritylodontid synapsids from the Lower Jurassic of western North America. Bulletin of the Museum of Comparative Zoology, 151: 217-268.
[54]	Sues H D. 1986b. Dinnebitodon amarali,a new tritylodontid(Synapsida)from the Lower Jurassic of western North America. Journal of Paleontology, 60: 758-762.
[55]	Tatarinov L P,Matchenko E N. 1999. A find of an aberrant tritylodont(Reptilia,Cynodontia)in the Lower Cretaceous of the Kemerovo Region. Paleontological Journal, 33: 422-428.
[56]	Velazco P M,Buczek A J,Novacek M J. 2017. Two new tritylodontids(Synapsida,Cynodontia,Mammaliamorpha)from the Upper Jurassic,southwestern Mongolia. American Museum Novitates, 3874: 1-35.
[57]	Waldman M,Savage R J G. 1972. The first Jurassic mammal from Scotland. Journal of the Geological Society, 128: 119-125.
[58]	Watabe M,Tsubamoto T,Tsogtbaatar K. 2007. A new tritylodontid synapsid from Mongolia. Acta Palaeontologica Polonica, 52: 263-274.
[59]	Wible J R. 1991. Origin of Mammalia: the craniodental evidence reexamined. Journal of Vertebrate Paleontology, 11: 1-28.
[60]	Young C C. 1940. Preliminary notes on the Mesozoic mammals of Lufeng,Yunnan,China. Bulletin of the Geological Society of China, 20: 93-111.
[61]	Young C C. 1947. Mammal-like reptiles from Lufeng,Yunnan,China. Proceedings of the Zoological Society of London, 117: 537-597.
[62]	Hennig E. 1922. Die Säugerzähne des württembergischen Rhät-Lias-Boneneds. Neues Jahrb Mineral Geology and Palaeontology, 46: 181-267.

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