The most basal ankylosaurine dinosaur from the Albian-Cenomanian of China, with implications for the evolution of the tail club - PubMed (original) (raw)

The most basal ankylosaurine dinosaur from the Albian-Cenomanian of China, with implications for the evolution of the tail club

Wenjie Zheng et al. Sci Rep. 2018.

Abstract

The tail club knob is a highly specialized structure thought to characterize a subgroup of the ankylosaurine ankylosaurians, and the oldest documented tail club knob in the fossil record occurred in the Campanian ankylosaurine Pinacosaurus. Here we report a new ankylosaurid Jinyunpelta sinensis, gen. et sp. nov., from the Albian-Cenomanian Liangtoutang Formation, Jinyun County, Zhejiang, China. This is the first definitive and the best preserved ankylosaurid dinosaur ever found in southern China. Jinyunpelta possesses unique cranial features differs from other ankylosaurs including two paranasal apertures level with and posterior to the external naris, a triangular fossa on the anterodorsal edge of the maxilla, an antorbital fossa in the junction between the maxilla, lacrimal and jugal, and an anterior process of the prearticular that lies ventral to the splenial. Our phylogenetic analysis suggests Jinyunpelta as the most basal ankylosaurine dinosaur. Jinyunpelta has a tail club with interlocking caudal vertebrae and a well-developed tail club knob, it represents the oldest and the most basal ankylosaurian known to have a well-developed tail club knob. The new discovery thus demonstrates that a large and highly modified tail club evolved at the base of the ankylosaurine ankylosaurs at least about 100 million years ago.

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Conflict of interest statement

The authors declare no competing interests.

Figures

Figure 1

Figure 1

Locality and geological maps pertaining to the fossil locality. (a) The map of China showing Zhejiang Province. (b) The map of Zhejiang showing the Jinyun county; (c) map of Jinyun County with the fossil locality (marked by a star); (d) Geological map of Huzhen and Shuhong basins in Jinyun County showing type locality of Jinyunpelta sinensis (marked by a star), (a,b) and (c) Jin, et al., (d) after Lawver, et al..

Figure 2

Figure 2

The skull and mandible of Jinyunpelta sinensis holotype ZMNH M8960. Photograph and line drawing of the skull and mandible in dorsal (a), ventral (b), and anterior (c) views. Abbreviations: an, angular; ar, articular; b, paranasal aperture B; bo, basioccipital; c1, paranasal aperture C1; c2, paranasal aperture C2; d, dentary; en, external naris; exo, exoccipital; f, frontal; fen, fenestra; l, lacrimal; mso, middle supraorbital; mx, maxilla; mf, maxillary fossa; na, nasal; o, orbit; oss, ossicle; pd, predentary; pmx, premaxilla; po, postorbital; pra, prearticular; prf, prefrontal; q, quadrate; qj, quadratojugal; qjh, quadratojugal horn; sa, surangular; sno, supranarial ornamentation; spl, splenial; sq, squamosal.

Figure 3

Figure 3

The skull and mandible of Jinyunpelta sinensis holotype ZMNH M8960. Photograph and line drawing of the skull and mandible in left lateral (a), left ventrolateral (b), right lateral (c) and right ventrolateral (d) views, and enlarged view of the right narial portion (e). Abbreviations: an, angular; ar, articular; b, paranasal aperture B; bo, basioccipital; c1, paranasal aperture C1; c2, paranasal aperture C2; d, dentary; en, external naris; exo, exoccipital; f, frontal; fen, fenestra; l, lacrimal; mso, middle supraorbital; mx, maxilla; mf, maxillary fossa; na, nasal; o, orbit; qjh, quadratojugal horn; oss, ossicle; pd, predentary; pmx, premaxilla; po, postorbital; pra, prearticular; prf, prefrontal; q, quadrate; qj, quadratojugal; qjh, quadratojugal horn; sa, surangular; sno, supranarial ornamentation; spl, splenial; sq, squamosal.

Figure 4

Figure 4

The vertebrae of Jinyunpelta sinensis holotype ZMNH M8960. The axis in anterior (a), left lateral (b), right lateral (c), posterior (d), dorsal (e) and ventral (f) views; the postaxial cervical vertebra in anterior (g), left lateral (h), posterior (i), right lateral (j), ventral (k) and dorsal (l) views; the dorsal vertebra in right lateral (m), anterior (n), left lateral (o), posterior (p), dorsal (q) and ventral (r) views. Abbreviations: c, centrum; di, diapophysis, na, neural arch; nc, neural canal; ns, neural spine; od, odontoid process; pa, parapophysis; poz, postzygapophysis; prz, prezygapophysis; tp, transverse process.

Figure 5

Figure 5

The sacral vertebrae and left ilium of Jinyunpelta sinensis holotype ZMNH M8960 in dorsal (a) and ventral (b) views. Abbreviations: ac, acetabulum; il, ilium; po, plate osteoderm; oss, ossicle; r, rib; sv, sacral vertebra.

Figure 6

Figure 6

The tail club Jinyunpelta sinensis paratype ZMNH M8963 in dorsal (a) and ventral (b) views. Abbreviations: ch, chevron; maj os, major osteoderm of the tail club knob; min os, minor osteoderm of the tail club knob; ns, neural spine; os, osteoderm; ot, ossified tendon; prz, prezygapophyses.

Figure 7

Figure 7

Forelimb of Jinyunpelta sinensis holotype ZMNH M8960. The left scapula in medial (a), lateral (b), dorsal (c) and ventral (d) views; The right humerus in anterior (e) and posterior (f) views; the right manus in dorsal view (g). Abbreviations: acr, acromion; bf, bicipital fossa; dpc, deltopectoral crest; gl, glenoid; glf, glenoid fossa; hh, humeral head; it, internal tuberosity; mc, metacarpal; medr, medial ridge; mtlc, enthesis of M. triceps longus caudalis.

Figure 8

Figure 8

Hind limb of Jinyunpelta sinensis. Left ischium of ZMNH M8963 in lateral (a), anterior (b), medial (c) and posterior (d) views; the right ischium of ZMNH M8960 in lateral (e), anterior (f), medial (g) and posterior (h) views; the left femur of ZMNH M8960 in anterior (i), lateral (j), posterior (k) and medial (l) views; the left tibia and fibula of ZMNH M8963 in anterior (m), lateral (n), posterior (o) and medial (p) views. Abbreviations: 4th, fourth trochanter; ac, acetabulum; fl, fibula; g, scar for the M. gastrocnemius; gt, greater trochanter; lc, lateral condyle; mc, medial condyle; pp, pubic peduncle; s, scar; t, tibia.

Figure 9

Figure 9

The derivative strict reduced consensus tree produced by phylogenetic analysis. Aletopelta, Paw Paw scuteling, Sauroplites, and ‘_Zhejiangosaurus_’ were pruned a posteriori to improve resolution. Values above nodes represent bootstrap proportions. Values beneath nodes indicate Bremer support.

Figure 10

Figure 10

Temporal calibration of the simplified derivative strict reduced consensus tree produced by phylogenetic analysis. Taxa in red text have a tail club knob. The geologic numerical ages and coloring follow International Chronostratigraphic Chart 2017/02.

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