新疆下三叠统韭菜园组一新小型兽头类

doi:10.19615/j.cnki.2096-9899.240726

摘要/Abstract

摘要：

主要基于晚二叠世的头部材料，中国近年来报道了多个兽头类物种。记述了新疆韭菜园组的小型包氏兽超科(Baurioidea)新材料，包括一个头骨吻端及咬合的下颌以及两个头后骨架，并命名为迷惑韭菜园颌兽( Jiucaiyuangnathus confusus )。虽然标本为幼年个体，新种有以下鉴定特征：上颌骨侧面中部有小圆坑；泪骨与鼻骨接触；前颌骨后端在内鼻孔外侧有容纳下颌齿的窝；犁骨前部侧缘有小型三角形脊向背腹两侧延伸，组成细长的犁骨孔的侧缘；额骨有尖角延伸到鼻骨后侧部；齿骨的联合部收窄并指向前方；5枚前颌骨齿，上下犬齿缺失，前颌骨齿与上颌骨齿间有齿缺，下颌前部无齿缺，10枚上颌骨齿，12枚齿骨齿，后部齿加长，有大的主尖及小的前后副尖；寰椎神经弓与神经棘愈合，枢椎神经棘倒向后方，前乌喙骨孔位于前乌喙骨与肩胛骨间。齿系与小型包氏兽超科成员南非的 Ericiolacerta parva 及俄罗斯的 Silphedosuchus orenburgensis 类似。新的头后骨骼材料提供了难得的研究包氏兽超科成员头后骨骼的机会。

关键词: 新疆吉木萨尔, 早三叠世, 韭菜园组, 兽头类，包氏兽超科

Abstract:

Several therocephalian species, mainly represented by cranial material from the late Permian, have been reported from China in recent years. Here we describe a tiny new baurioid therocephalian, Jiucaiyuangnathus confusus gen. et sp. nov., from the Jiucaiyuan Formation, Xinjiang, China. The new taxon is represented by a partial snout with occluded partial lower jaw and two postcranial skeletons. Although juvenile in stage, the new species is diagnosed by the following features: round pit in middle of lateral surface of maxilla; lacrimal contact nasal; fossa for dentary tooth on the posterior end of the premaxilla, lateral to the anterior choana; two small vertical triangular ridges extending dorsally and ventrally on the vomerine anterior portion, and bordering a thin vomerine foramen laterally; anterior projection of the lateral part of the frontal on the nasal; symphyseal region of the dentary projected anteriorly; 5 upper premaxillary teeth, upper and lower canines absent, diastema between the last premaxillary upper incisor and first maxillary tooth present, no diastema separating anterior from posterior dentition in the mandible, 10 maxillary teeth and 12 dentary teeth, posterior postcanine expands mesiodistally, having a main large cusps and tiny anterior and posterior accessory cusps in line; neural arches of the atlas fused by the neural spine, neural spine of the axis projected posteriorly, procoracoid foramen lies between procoracoid and scapula. Features of the dentition resembles those of the small baurioid Ericiolacerta parva from South Africa and Silphedosuchus orenburgensis from Russia. The specimens provide the rare opportunity to know in detail the postcranial skeleton of baurioids.

Key words: Jimsar, Xinjiang, Early Triassic, Jiucaiyuan Formation, Therocephalia, Baurioidea

中图分类号:

Q915.864

刘俊. 新疆下三叠统韭菜园组一新小型兽头类. 古脊椎动物学报, 2024, 62(3): 201-224.

LIU Jun, Fernando ABDALA. A new small baurioid therocephalian from the Lower Triassic Jiucaiyuan Formation, Xinjiang, China. Vertebrata Palasiatica, 2024, 62(3): 201-224.

图/表 10

Fig. 1 Jiucaiyuangnathus confusus gen. et sp. nov. (IVPP V32945, holotype) from Jimsar, Xinjiang A-C. photo (A) and 3D rendering (B) of the left side of the snout in lateral and 3D rendering of the medial view of the left side of the snout (C);D. 3D rendering of the snout in posterior view showing the bones as preserved Abbreviations: afo. anteriormost foramen; cc. christa choanalis; F. frontal; L. lacrimal; l.c. lacrimal canal;M. maxilla; m. maxillary tooth; m.a. maxillary antrum; msf. maxillo-septomaxillary foramen;pdl. dorsal layer of palatine; pit. maxillary round pit; PL. palatine; PM. premaxilla;pml. medial layer of palatine; sH, sinus Highmore; SM. septomaxilla; V. vomer

Fig. 2 Snout of Jiucaiyuangnathus confusus gen. et sp. nov. (IVPP V32945, holotype) from Jimsar, Xinjiang A. photo in dorsal view; B, C. 3D rendering of the left nasal and frontal in dorsal (B) and ventral (C) views; D, E. 3D rendering of the left side of the snout in ventral (D) and dorsal (E) views Abbreviations: cc. christa choanalis; ch. choana; F. frontal; f. fossa for lower anterior tooth; L. lacrimal; lpr. lacrimal-palatine ridge; M. maxilla; m. maxillary tooth; mpf. maxilla-palatine foramen; N. nasal; p. premaxillary tooth; pdl. dorsal layer of palatine; p.f. premaxillary fossa; PF. prefrontal; PL. palatine; PM. premaxilla; pml. medial layer of palatine; pvl. ventral layer of palatine; SM. septomaxilla

Fig. 3 3D rendering of Jiucaiyuangnathus confusus gen. et sp. nov. (IVPP V32945, holotype) from Jimsar, Xinjiang A. the preserved snout in ventral view; B, C. vomer in dorsal (B) and ventral (C) views, vomer is reconstructed in the natural position in (C); D. left dentary in lateral view; E, F. mandibles in dorsal (E) and ventral (F) views Abbreviations: 1st ic. first incisor; ch. choana; D. dentary; d. dentary tooth; F. frontal; for vn. place for vomeronasal organ; M. maxilla; N. nasal; PL. palatine; PM. premaxilla; SP. splenial; V. vomer; v.f. vomerine foramen

Fig. 4 Posterior skeletons of Jiucaiyuangnathus confusus gen. et sp. nov. (IVPP V32946) from Jimsar, Xinjiang A, B. photo of the slab; C, D. 3D rendering of bones Grey and light purple are bones of each specimen. Light blue uncertain. See online version for color

Fig. 5 3D rendering of Jiucaiyuangnathus confusus gen. et sp. nov. (IVPP V32946-1) from Jimsar, Xinjiang A, B. the preserved skeleton in dorsal (A) and ventral (B) views; C. right femur in distal view; D-F. atlas and axis in anterior (D), left (E), and right (F) lateral views; G. four thoracic vertebrae in right lateral view; H, I. sacral region in lateral (H) and dorsal (I) views; J. caudal vertebrae Abbreviations: aic. atlas intercentrum; cdr. caudal rib; cdv. caudal vertebra; cr. cervical rib; FE. femur; lv. lumbar vertebra; ns. neural spine; sr. sacral rib; sv. sacral vertebra; tp. transverse process; tr. thoracic rib; tv. thoracic vertebra

Fig. 6 3D rendering of Jiucaiyuangnathus confusus gen. et sp. nov. (IVPP V32946-2) from Jimsar, Xinjiang A, B. skeleton in dorsal (A) and ventral (B) views; C. part of thoracic vertebrae in left lateral view; D. two sacral and three anterior caudal ribs; E-H. sacral ribs of (D) in dorsal (E, G) and ventral (F, H) views;I-L. two smaller sacral ribs in dorsal (I, K) and ventral (J, L) views Abbreviations: cdr. caudal rib; mt. metatarsal; sr. sacral rib

Fig. 7 3D rendering of shoulder girdle of Jiucaiyuangnathus confusus gen. et sp. nov. (IVPP V32946-2) from Jimsar, Xinjiang A-D. left scapula in anterior (A), lateral (B), posterior (C), and medial (D) views; E. partial right scapula in lateral view; F, G. left procoracoid in lateral (F) and medial (G) views Abbreviation: gl. glenoid

Fig. 8 3D rendering of pelvic girdles of Jiucaiyuangnathus confusus gen. et sp. nov. from Jimsar, Xinjiang IVPP V32946-1: A, B. left ilium in lateral (A) and medial (B) views; C, D. right ilium in lateral (C) and medial (D) views; E. relocated pubes and right ischium in ventral view; F. left pubis in dorsal view; G. right ischium in dorsal view. V32946-2: H, I. left ilium in lateral (H) and medial (I) views; J, K. right ilium in lateral (J) and medial (K) views; L-N. left ischium in ventral (L), lateral (M), and dorsal (N) views; O-Q. incomplete right pubis in dorsal (O), lateral (P), and ventral (Q) views; R. reconstructed left pelvis in lateral view based on the small individual (right ischium reversed). Scale bars equal 5 mm

Fig. 9 3D rendering of right femur of Jiucaiyuangnathus confusus gen. et sp. nov. (IVPP V32946-2) from Jimsar, Xinjiang in ventral (A), anterior (B), dorsal (C), posterior (D) and proximal (E) views Abbreviations: tr.int. internal trochanter; tr.maj. trochanter major; tr.min. trochanter minor

Fig. 10 Relationships of Baurioidea, showing the positions of Jiucaiyuangnathus when canines are coded as unknown Abbreviations: Chan. Changhsingian; Ind. Induan

参考文献 63

[1]	Abdala F, Rubidge B S, Van den Heever J A, 2008. The oldest therocephalians (Therapsida, Eutheriodontia) and the early diversification of Therapsida. Palaeontology, 51: 1011-1024
[2]	Angielczyk K D, Liu J, Sidor C A et al., 2022. The stratigraphic and geographic occurrences of Permo-Triassic tetrapods in the Bogda Mountains, NW China - implications of a new cyclostratigraphic framework and Bayesian age model. J Afr Earth Sci, 195: 104655, https://doi.org/10.1016/j.jafrearsci.2022.104655
[3]	Benoit J, Manger P R, Rubidge B S, 2016. Palaeoneurological clues to the evolution of defining mammalian soft tissue traits. Sci Rep, 6: 25604, 10.1038/srep25604 DOI URL PMID
[4]	Boonstra L D, 1954. A scaloposaurid from the Tapinocephalus -Zone. Ann Mag Nat Hist, 7: 153-157
[5]	Botha-Brink J, Modesto S P, 2007. A mixed-age classed ‘pelycosaur’ aggregation from South Africa: earliest evidence of parental care in amniotes? Proc R Soc B, 274: 2829-2834
[6]	Botha-Brink J, Modesto S P, 2011. A new skeleton of the therocephalian synapsid Olivierosuchus parringtoni from the Lower Triassic South African Karoo Basin. Palaeontology, 54: 591-606
[7]	Brink A S, 1950. Notes on a second specimen of Homodontosaurus kitchingi . South Afr J Sci, 47: 118-119
[8]	Broom R, 1901. On Ictidosuchus primaevus. Trans S Afr Philos Soc, 11: 177-184, 10.1080/21560382.1900.9525964
[9]	Broom R, 1911. On some new South African Permian reptiles. Proc Zool Soc Lond, 81: 1073-1082
[10]	Broom R, 1938. On a nearly complete therocephalian skeleton. Ann Trans Mus, 19: 257-261
[11]	Broom R, 1940. On some new genera and species of fossil reptiles from the Karroo beds of Graaff-Reinet. Ann Trans Mus, 20: 157-192
[12]	Broom R, 1948a. The skeleton of a very small therocephalian. Ann Trans Mus, 21: 39-41
[13]	Broom R, 1948b. New fossil reptile genera from the Bernard Price collection. Ann Trans Mus, 21: 187-194
[14]	Colbert E H, Kitching J W, 1981. Scaloposaurian reptiles from the Triassic of Antarctica. Am Mus Novit, 709: 1-22
[15]	Fourie H, Rubidge B S, 2007. The postcranial skeletal anatomy of the therocephalian Regisaurus (Therapsida: Regisauridae) and its utilization for biostratigraphic correlation. Palaeont Afr, 42: 1-16
[16]	Gigliotti A, Pusch L, Kammerer C F et al., 2023. Craniomandibular anatomy of the akidnognathid therocephalian Olivierosuchus parringtoni from the Early Triassic of South Africa. Palaeont Afr, 56: 142-170
[17]	Goloboff P A, Catalano S, 2016. TNT, version 1.5, with a full implementation of phylogenetic morphometrics. Cladistics, 32: 221-238, doi: 10.1111/cla.12160 PMID
[18]	Hillenius W J, 2000. Septomaxilla of nonmammalian synapsids: soft-tissue correlates and a new functional interpretation. J Morphol, 245: 29-50 PMID
[19]	Hopson J A, Barghusen H R, 1986. An analysis of therapsid relationships. In: HottonN III, MacLeanP D, RothJ J et al. eds. The Ecology and Biology of Mammal-like Reptiles. Washington D C: Smithsonian Institution Press. 83-106
[20]	Hou L H, 1979. On a new theriodont from Inner Mongolia. Vert PalAsiat, 17: 121-130
[21]	Huttenlocker A K, Abdala F, 2015. Revision of the first therocephalian, Theriognathus Owen (Therapsida: Whaitsiidae), and implications for cranial ontogeny and allometry in nonmammaliaform eutheriodonts. J Paleontol, 89: 645-664
[22]	Huttenlocker A K, Sidor C A, 2016. The first karenitid (Therapsida, Therocephalia) from the upper Permian of Gondwana and the biogeography of Permo-Triassic therocephalians. J Vert Paleont, 36: e1111897, 10.1080/02724634.2016.1111897
[23]	Huttenlocker A K, Botha J, Browning C et al., 2022. A Gulliver Scaloposaurus (Therapsida, Therocephalia) from the Katberg Formation of South Africa and its implication for Lilliput assemblages during the Early Triassic recovery. J Afr Earth Sci, 196, 10.1016/j.jafrearsci.2022.104720
[24]	Ivakhnenko M F, 2011. Permian and Triassic Therocephals (Eutherapsida) of Eastern Europe. Paleontol J, 45: 981-1144, 10.1134/s0031030111090012
[25]	Jasinoski S C, Abdala F, 2017. Aggregations and parental care in the Early Triassic basal cynodonts Galesaurus planiceps and Thrinaxodon liorhinus . PeerJ, 5: e2875, 10.7717/peerj.2875
[26]	Kemp T S, 1978. Stance and gait in the hindlimb of a therocephalian mammal-like reptile. J Zool, 186: 143-161
[27]	Kemp T S, 1979. The primitive cynodont Procynosuchus : functional anatomy of the skull and relationships. Philos Trans R Soc Lond B, 285: 73-122
[28]	Kemp T S, 1986. The skeleton of a baurioid therocephalian therapsid from the Lower Triassic ( Lystrosaurus Zone) of South Africa. J Vert Paleont, 6: 215-232
[29]	Koh T P, 1940. Santaisaurus yuani gen. et sp. nov., ein neues Reptil aus der unteren Trias von China. Bull Geol Surv China, 20: 73-92
[30]	Li Y H, 1984. On a new scaloposaurid from Inner Mongolia. Vert PalAsiat, 22: 21-28
[31]	Liu J, 2020. New chroniosuchian materials from Xinjiang, China. Vert PalAsiat, 58: 283-292
[32]	Liu J, Abdala F, 2017a. Therocephalian (Therapsida) and chroniosuchian (Reptiliomorpha) from the Permo-Triassic transitional Guodikeng Formation of the Dalongkou Section, Jimsar, Xinjiang, China. Vert PalAsiat, 55: 24-40
[33]	Liu J, Abdala F, 2017b. The tetrapod fauna of the upper Permian Naobaogou Formation of China: 1. Shiguaignathus wangi gen. et sp. nov., the first akidnognathid therocephalian from China. PeerJ, 5: e4150, 10.7717/peerj.4150
[34]	Liu J, Abdala F, 2019. The tetrapod fauna of the upper Permian Naobaogou Formation of China: 3. Jiufengia jiai gen. et sp. nov., a large akidnognathid therocephalian. PeerJ, 7: e6463, 10.7717/peerj.6463
[35]	Liu J, Abdala F, 2020. The tetrapod fauna of the upper Permian Naobaogou Formation of China: 5. Caodeyao liuyufengi gen. et sp. nov., a new peculiar therocephalian. PeerJ, 8: e9160, 10.7717/peerj.9160
[36]	Liu J, Abdala F, 2022. The emblematic South African therocephalian Euchambersia in China: a new link in the dispersal of late Permian vertebrates across Pangea. Biol Lett, 18: 20220222, 10.1098/rsbl.2022.0222
[37]	Liu J, Abdala F, 2023. Late Permian terrestrial faunal connections invigorated: the first whaitsioid therocephalian from China. Palaeont Afr, 56: 111-117
[38]	Liu J, Yang W, 2022. A gorgonopsian from the Wuttonggou Formation (Changhsingian, Permian) of Turpan Basin. Xinjiang, China. Palaeoworld, 31(3): 383-388
[39]	Liu J, Li J L, Cheng Z W, 2002. The Lystrosaurus fossils from Xinjiang and their bearing on the terrestrial Permian-Triassic boundary. Vert PalAsiat, 40: 267-275
[40]	Mendrez C H, 1973. Révision du genre Homodontosaurus Broom 1949 et problème de la survivance des Pélycosaures au Permien supérieur en Afrique australe. C R Acad Sci Paris, 276: 1821-1822
[41]	Mendrez C H, 1975. Principales variations du palais chez les thérocéphales Sud-Africains (Pristerosauria et Scaloposauria) au cours du Permien Supérieur et du Trias Inférieur. Colloque Int CNRS, 218: 379-408
[42]	Mendrez-Carroll C H, 1979. Nouvelle étude du crâne du type de Scaloposaurus constrictus Owen, 1876, spécimen jeune, Therocephalia, Scaloposauria, Scaloposauridae, de la zone à Cistecephalus (Permian supérieur) d'Afrique australe. Bull Mus Natl Hist Nat C, 3: 155-201
[43]	Meng Q, Liu J, Varricchio D J et al., 2004. Parental care in an ornithischian dinosaur. Nature, 431: 145-146, 10.1038/431145a
[44]	Pusch L C, Kammerer C F, Fröbisch J, 2024. The origin and evolution of Cynodontia (Synapsida, Therapsida): reassessment of the phylogeny and systematics of the earliest members of this clade using 3D-imaging technologies. Anat Rec, 307: 1634-1730, https://doi.org/10.1002/ar.25394
[45]	Sigogneau D, 1963. Note sur une nouvelle espece de Scaloposauridae. Palaeont Afr, 8: 13-37
[46]	Sigurdsen T, Huttenlocker A K, Modesto S P et al., 2012. Reassessment of the morphology and paleobiology of the therocephalian Tetracynodon darti (Therapsida), and the phylogenetic relationships of Baurioidea. J Vert Paleont, 32: 1113-1134, 10.1080/02724634.2012.688693
[47]	Sun A L, 1963. The Chinese kannemeyerids. Palaeont Sin, New Ser C, 17: 1-109
[48]	Sun A L, 1991. A review of Chinese therocephalian reptiles. Vert PalAsiat, 29: 85-94
[49]	Sun A L, Hou L H, 1981. Hazhenia , a new genus of Scaloposauria. Acta Palaeont Sin, 20(4): 297-310
[50]	Tatarinov L P, 2004. The postcranial skeleton of the Late Permian scaloposaurian Karenites ornamentatus (Reptilia, Theriodontia) from the Kirov Region. Paleontol J, 38: 548-555
[51]	van den Heever J A, 1994. The cranial anatomy of the early Therocephalia (Amniota: Therapsida). Ann Univ Stellenbosch, 1994: 1-59
[52]	Watson D M S, 1931. On the skeleton of a bauriamorph reptile. Proc Zool Soc Lond, 1931: 1163-1205
[53]	Yang W, Wan M, Crowley J L et al., 2021. Paleoenvironmental and paleoclimatic evolution and cyclo- and chrono-stratigraphy of upper permian-Lower Triassic fluvial-lacustrine deposits in Bogda Mountains, NW China - implications for diachronous plant evolution across the Permian-Triassic boundary. Earth Sci Rev, 222: 103741, https://doi.org/10.1016/j.earscirev.2021.103741
[54]	Young C C, 1936. On a new Chasmatosaurus from Sinkiang. Bull Geol Surv China, 15: 291-320
[55]	Young C C, 1939. Additional Dicynodontia remains from Sinkiang. Bull Geol Surv China, 19: 111-139
[56]	Young C C, 1952. On a new therocephalian from Sinkiang, China. Acta Sci Sin, 2: 152-165
[57]	Young C C, 1961. On a new cynodont from N.W. Shansi. Vert PalAsiat, 1961(2): 109-113
[58]	Young C C, 1963. Additional remains of Chasmatosaurus yuani Young from Sinkiang, China. Vert PalAsiat, 7: 215-222
[59]	Young C C, 1973a. The discovery of Prolacertilia in Jimusar, Sinkiang. Vert PalAsiat, 11: 46-48
[60]	Young C C, 1973b. Note on the labyrinthodont from the Lystrosaurus Zone in Jimusar, Sinkiang. Vert PalAsiat, 11: 50-51
[61]	Young C C, 1974. A new genus of Traversodontidae in Jiyuan, Henan. Vert PalAsiat, 12: 203-211
[62]	Yuan P L, Young C C, 1934a. On the discovery of a new dicynodon in Sinkiang. Bull Geol Surv China, 13: 563-573
[63]	Yuan P L, Young C C, 1934b. On the occurrence of Lystrosaurus in Sinkiang. Bull Geol Surv China, 13: 575-580