Morphology of Funiusaurus luanchuanensis (Squamata, Polyglyphanodontia) based on new material

doi:10.19615/j.cnki.2096-9899.260814

Abstract

Abstract:

Polyglyphanodontia is a successful clade of squamates thriving during the Late Cretaceous. They were widely distributed in Asia, North America and possibly some other regions, and are highly diverse in taxonomy, morphology and ecology. Although the clade is better known to have many large herbivores, quite a few small insectivorous species were described from all the main localities where polyglyphanodontian fossils were found. Funiusaurus luanchuanensis from the Upper Cretaceous Qiupa Formation of Henan Province, China is one of them. Previously, it was described based only on a nearly complete but relatively fragmentary skull which therefore provided limited morphological information. Herein we add a great amount of anatomical data of this species based on high-resolution computerized tomography data of a new completely preserved skull and several other cranial and postcranial specimens, especially of the palate, braincase, vertebral column and limbs. The new data of Funiusaurus will help to build a more comprehensive phylogenetic data matrix of Polyglyphanodontia in the future, and further clarify the evolutionary history of body size or diet shift within the clade.

Key words: Luanchuan, Henan, Qiupa Formation, Late Cretaceous, Funiusaurus, Polyglyphanodontia

摘要：

多凿齿蜥类是一类繁盛于晚白垩世的有鳞类，广泛分布于亚洲、北美洲以及其他地区，在属种、形态和生态上都表现出高度多样性。虽然多凿齿蜥类以拥有众多大型植食性代表而闻名，但它的主要化石地点也产出了不少小型食虫物种，中国河南上白垩统秋扒组的栾川伏牛蜥(Funiusaurus luanchuanensis)就是其中之一。伏牛蜥之前仅有一个虽近完整但较为破碎的头骨，因此提供的形态学信息有限。基于一件新发现于河南栾川的完整头骨的高精度CT数据和其他几件头骨和头后骨骼标本，补充了伏牛蜥的大量解剖学数据，尤其是腭区、脑颅、脊柱以及四肢部分。这些新数据将有助于建立一个更完善的多凿齿蜥类形态矩阵，从而深入讨论多凿齿蜥类的体型大小和食性转变等演化历史。

关键词: 河南栾川, 秋扒组, 晚白垩世, 伏牛蜥, 多凿齿蜥类

CLC Number:

Q915.864

DONG Li-Ping, LI Jie, CHANG Fei, HAN Ye, LIU Di, WANG Yuan. Morphology of Funiusaurus luanchuanensis (Squamata, Polyglyphanodontia) based on new material. Vertebrata Palasiatica, 2026, 64(4): 372-395.

董丽萍, 李杰, 常飞, 韩晔, 刘迪, 王原. 2026, 64(4): 372-395, 栾川伏牛蜥(有鳞类，多凿齿蜥类)新材料的形态学研究. 古脊椎动物学报.

Figures/Tables 6

Fig. 1 The newly referred specimen of Funiusaurus luanchuanensis (HGM L2021-1-8) from the Upper Cretaceous Qiupa Formation of Luanchuan, Henan, China A. photograph of the specimen in dorsolateral view; B. 3D rendering of the specimen in the same view Abbreviations: an. angular; ar. articular; bo. basioccipital; bs. basisphenoid; co. coronoid; de. dentary; ept. epipterygoid; fr. frontal; ju. jugal; l. left; la. lacrimal; ma. maxilla; na. nasal; ot-op. otic-occipital; pa. parietal; pal. palatine; pm. premaxilla; pocc. paroccipital process; pof?. possible postfrontal; prf. prefrontal; pt. pterygoid; qu. quadrate; r. right; san. surangular; sm. septomaxilla; so. supraoccipital

Fig. 2 3D rendering of isolated bones in Funiusaurus luanchuanensis (HGM L2021-1-8) from the Upper Cretaceous Qiupa Formation of Luanchuan, Henan, China A. right maxilla in labial (A1), medial (A2) and dorsal (A3) views and one isolated maxillary tooth (A4) showing the tricuspid nature; the red box in A1 shows where the isolated tooth is from; B. premaxilla in right lateral view; C. frontal in dorsal (C1), left lateral (C2) and ventral (C3) views; D. quadrate in posterior view; E. left jugal in lateral (E1) and medial (E2) views; F. the palate in ventral view (F1), and the vomer (F2), palatine (F3) and pterygoid (F4) in dorsal view; G. anterior portion of the right dentary in labial (G1) and lingual (G2) views; H. left mandible in labial (H1) and lingual (H2) views Abbreviations: a.i.a.f. anterior interior alveolar foramen; a.s.f. anterior surangular foramen; an. angular; ar.co. articular condyle; bs.ft. basisphenoid facet; ce.co. cephalic condyle; co. coronoid; cr.cr. crista cranii; de. dentary; ecpt. ectopterygoid; ecpt.ft. ectopterygoid facet; fa.pr. facial process; fo.col. fossa columellae; io.ca. infraorbital canal; ju.ft. jugal facet; la.ft. lacrimal facet; ma.ft. maxilla facet; man.co. mandible condyle; med.rd. medial ridge; na.ft. nasal facet; na.pr. nasal process; p.m.f. posterior mylohyoid foramen; p.pr. posterior process; p.s.f. posterior surangular foramen; pa.f. parietal foramen; pa.lp. parietal lappet; pal. palatine; pal.ft. palatine facet; pal.sf. palatal shelf; pal.ss. palatine sinus; par. prearticular; pob.rm. postorbital ramus; pof.ft. postfrontal facet; prf.ft. prefrontal facet; pt. pterygoid; pt.lp. pterygoid lappet; pt.t. pterygoid teeth; rap. retroarticular process; rd. ridge; sac. opening of superior alveolar canal; san. surangular; sd.rd. subdental ridge; sm.ft. septomaxilla facet; spl. splenial; sob.rm. suborbital ramus; sym. symphysis; ty.cr. tympanic crest; vo. vomer; vo.f. vomerine foramen

Fig. 3 3D rendering of the braincase in Funiusaurus luanchuanensis (HGM L2021-1-8), from the Upper Cretaceous Qiupa Formation of Luanchuan, Henan, China, in posterior (A) and left lateral (B) views, and restored basisphenoid and basioccipital in anterior (C), dorsal (D) and ventral (E) views Abbreviations: al.pr. alar process; b.tb. basal tubercle; bpt.pr. basipterygoid process; CN6. opening of cranial nerve abducens; ds. dorsum sellae; f.mg. foramen magnum; f.o. foramen ovalis; icf. internal carotid foramen; lrst. lateral opening of resessus scalae tympani; oc.re. occipital recess; pocc. paroccipital process;psp. parasphenoid; so. supraoccipital

Fig. 4 Additional cranial anatomy of Funiusaurus luanchuanensis from the Upper Cretaceous Qiupa Formation of Luanchuan, Henan, China A. HGM L2021-1-8: 3D rendering of the skull in ventral view after removing the mandibles; B. HGM 1137-2: the middle section of a left mandible in lingual view; C. HGM L08-15: a partial skull in lateral view;D. HGM L2021-1-9: a right dentary in labial view (D1) and a left dentary in lingual view (D2), arrows in D1 point to the mental foramina; E. HGM L2021-1-2: the skull in lateroposterior view Abbreviations: a.i.a.f. anterior interior alveolar foramen; an. angular; ar. articular; at.n?. possible atlas neural arch; bc. braincase; bo. basioccipital; bs. basisphenoid; co. coronoid; co.ft. coronoid facet; de. dentary; ecpt. ectopterygoid; fr. frontal; i.m.f. inferior myhyoid foramen; ju. jugal; l. left; la. lacrimal; ma. maxilla; Mk. Meckelian canal; pa. parietal; pal. palatine; pm. premaxilla; pocc. paroccipital process; pof?. possible postfrontal; prf. prefrontal; pt. pterygoid; qu. quadrate; r. right; san. surangular; scl. scleral ossicles; sd.rd. subdental ridge; sm. septomaxilla; spl. splenial; spl.ft. splenial facet; st?. possible supratemporal bone; sym. symphysis; vo. vomer

Fig. 5 A referred specimen of Funiusaurus luanchuanensis (HGM L2021-1-7) from the Upper Cretaceous Qiupa Formation of Luanchuan, Henan, China A. the photo of the specimen with rectangles indicating the positions of other subfigures; B. the sacral and anterior caudal vertebral section with the right hind limb in different views (B1-3); C. the articulated dorsal vertebrae in ventrolateral view; D. four cervical or dorsal vertebrae in lateral view; E. the partial skull in ventral view. Abbreviations: ac. astragalocalcaneum; con. condyle; cV. caudal vertebra; de. dentary; dt. distal tarsal; dV. dorsal vertebra; fe. femur; fi. fibula; hm. haemal arch; isc. ischium; ma. maxilla; mt. metatarsal; ns. neural spine; pm. premaxilla; pu. pubis; sV. sacral vertebra; syn. synapophysis; ti. tibia; tp. transverse process; ul. ulna; zyg. zygopophysis

Table 1 Polyglyphanodontian taxa with at least a nearly complete or partial skull preserved: the skull and/or mandible length (when skull not available) of the largest specimen, the tooth type and their diet (mm)

Region	Taxa	Specimen	Skull length	Mandible length	Tooth type (most complex)	Diet	Reference
North America	Polyglyphanodon sternbergi	USNM 15477	81.5		IV	Herbivorous	Gilmore, 1940
Ganzhou	Xenodontolacerta fangi	CVEB20180206	~107		II	Omnivorous	Jiang et al., 2026
Ganzhou	Yechilacerta yingliangia	YLSNHM01791	~105		I	Herbivorous	Xing et al., 2023
Ganzhou	Tianyusaurus sp.	NHMG8502	87.4		I	Herbivorous	Mo et al., 2010
Ganzhou	Caninosaurus ganzhouensis	CUGW VH110	~30		III	Insectivorous	Wang et al., 2025
Gobi	Gilmoreteius ferrugenous	AMNH 6520	~115		I	Herbivorous	Gilmore, 1943a
Gobi	Gilmoreteius chulsanensis	ZPAL MgR-I/18	80		I	Herbivorous	Sulimski, 1975
Gobi	Cherminsaurus kozlowskii	ZPAL MgR- III/24	65		I	Herbivorous	Sulimski, 1975
Gobi	Erdenetesaurus robinsonae	ZPAL MgR- III/19	55		I	Herbivorous	Sulimski, 1975
Gobi	Darchansaurus estesi	ZPAL MgR- III/6	56		I	Herbivorous	Sulimski, 1975
Gobi	Tuberocephalosaurus pompabilis	PIN No. 3124/328	85	75	I	Herbivorous	Alifanov, 2000
Gobi	Aprisaurus bidentatus	PIN No. 3142/302	~55		I	Herbivorous	Alifanov, 2000
Gobi	Campedosaurus sulcularis	PIN No. 3142/341	~50.5		I	Herbivorous	Alifanov, 2000
Gobi	Gobinatus arenosus	IGM 3/126	35.2		III, unicuspid	Insectivorous	Gao and Norell, 2000
Gobi	Tchingisaurus multivagus	IGM 3/129	34		III, tricuspid	Insectivorous	Gao and Norell, 2000
Gobi	Pyramicephalosaurs cherminicus	IGM 3/130	~20.9		III, tricuspid	Insectivorous	Gao and Norell, 2000
Gobi	Conicodontosaurus djadochtaensis	IVPP V10051	~25		III, unicuspid	Insectivorous	Gilmore, 1943b
Gobi	Mongolochamops reshetovi	PIN No. 3142/304	~43		III, tricuspid	Insectivorous	Alifanov, 1988
Gobi	Altanteius facilis	PIN No. 3142/306	~39		III, tricuspid	Insectivorous	Alifanov, 1993
Gobi	Barungoia vasta	PIN No. 4487/2	~24	~23	III, unicuspid	Insectivorous	Alifanov, 1993
Gobi	Gurvansaurus potissimus	PIN No. 3143/104		35	III, unicuspid	Insectivorous	Alifanov, 1993
Gobi	Parameiva oculea	PIN no. 3142/310		~20	III, unicuspid	Insectivorous	Alifanov, 1993
Gobi	Prodenteia ministra	PIN No. 3142/324		~18	III, unicuspid	Insectivorous	Alifanov, 1993
Gobi	Dzhadochtosaurus giganteus	PIN No. 3143/103	69	53	III, unicuspid	Insectivorous	Alifanov, 1993
Gobi	Cyclurasia multidentata	PIN No. 3142/336	~35		III, ?	Insectivorous	Alifanov, 2000
Gobi	Adamisaurus magnidentatus	IGM 3/117	32.2		III, unicuspid	Insectivorous	Gao and Norell, 2000
Qiupa	Funiusaurus luanchuanensis	HGM 41HIII-114	~30		III, tricuspid	Insectivorous	Xu et al., 2014

Region	Taxa	Specimen	Skull length	Mandible length	Tooth type (most complex)	Diet	Reference
North America	Polyglyphanodon sternbergi	USNM 15477	81.5		IV	Herbivorous	Gilmore, 1940
Ganzhou	Xenodontolacerta fangi	CVEB20180206	~107		II	Omnivorous	Jiang et al., 2026
Ganzhou	Yechilacerta yingliangia	YLSNHM01791	~105		I	Herbivorous	Xing et al., 2023
Ganzhou	Tianyusaurus sp.	NHMG8502	87.4		I	Herbivorous	Mo et al., 2010
Ganzhou	Caninosaurus ganzhouensis	CUGW VH110	~30		III	Insectivorous	Wang et al., 2025
Gobi	Gilmoreteius ferrugenous	AMNH 6520	~115		I	Herbivorous	Gilmore, 1943a
Gobi	Gilmoreteius chulsanensis	ZPAL MgR-I/18	80		I	Herbivorous	Sulimski, 1975
Gobi	Cherminsaurus kozlowskii	ZPAL MgR- III/24	65		I	Herbivorous	Sulimski, 1975
Gobi	Erdenetesaurus robinsonae	ZPAL MgR- III/19	55		I	Herbivorous	Sulimski, 1975
Gobi	Darchansaurus estesi	ZPAL MgR- III/6	56		I	Herbivorous	Sulimski, 1975
Gobi	Tuberocephalosaurus pompabilis	PIN No. 3124/328	85	75	I	Herbivorous	Alifanov, 2000
Gobi	Aprisaurus bidentatus	PIN No. 3142/302	~55		I	Herbivorous	Alifanov, 2000
Gobi	Campedosaurus sulcularis	PIN No. 3142/341	~50.5		I	Herbivorous	Alifanov, 2000
Gobi	Gobinatus arenosus	IGM 3/126	35.2		III, unicuspid	Insectivorous	Gao and Norell, 2000
Gobi	Tchingisaurus multivagus	IGM 3/129	34		III, tricuspid	Insectivorous	Gao and Norell, 2000
Gobi	Pyramicephalosaurs cherminicus	IGM 3/130	~20.9		III, tricuspid	Insectivorous	Gao and Norell, 2000
Gobi	Conicodontosaurus djadochtaensis	IVPP V10051	~25		III, unicuspid	Insectivorous	Gilmore, 1943b
Gobi	Mongolochamops reshetovi	PIN No. 3142/304	~43		III, tricuspid	Insectivorous	Alifanov, 1988
Gobi	Altanteius facilis	PIN No. 3142/306	~39		III, tricuspid	Insectivorous	Alifanov, 1993
Gobi	Barungoia vasta	PIN No. 4487/2	~24	~23	III, unicuspid	Insectivorous	Alifanov, 1993
Gobi	Gurvansaurus potissimus	PIN No. 3143/104		35	III, unicuspid	Insectivorous	Alifanov, 1993
Gobi	Parameiva oculea	PIN no. 3142/310		~20	III, unicuspid	Insectivorous	Alifanov, 1993
Gobi	Prodenteia ministra	PIN No. 3142/324		~18	III, unicuspid	Insectivorous	Alifanov, 1993
Gobi	Dzhadochtosaurus giganteus	PIN No. 3143/103	69	53	III, unicuspid	Insectivorous	Alifanov, 1993
Gobi	Cyclurasia multidentata	PIN No. 3142/336	~35		III, ?	Insectivorous	Alifanov, 2000
Gobi	Adamisaurus magnidentatus	IGM 3/117	32.2		III, unicuspid	Insectivorous	Gao and Norell, 2000
Qiupa	Funiusaurus luanchuanensis	HGM 41HIII-114	~30		III, tricuspid	Insectivorous	Xu et al., 2014

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