Tharosaurus indicus Bajpai &amp; Datta &amp; Pandey &amp; Ghosh &amp; Kumar &amp; Bhattacharya 2023, gen. et sp. nov.
Creators: name: Bajpai, Sunil
name: Datta, Debajit
name: Pandey, Pragya
name: Ghosh, Triparna
name: Kumar, Krishna
name: Bhattacharya, Debasish
Publisher: name: Zenodo
Year: 2023
Subjects: Biodiversity
Taxonomy
Animalia
Dicraeosauridae
Tharosaurus
Tharosaurus indicus

description: <i>Tharosaurus indicus</i> gen. et sp. nov. 1 Department of Earth Sciences, Indian Institute of Technology, Roorkee, Uttarakhand 247667, India. 2 Geological Survey of India, Raipur, Chhattisgarh 492010, India. 3 Geological Survey of India, Jaipur, Rajasthan 302004, India. 4 Central Head Quarters, Geological Survey of India, Kolkata, West Bengal 700091, India. <i> *</i> email: sunil.bajpai@es.iitr.ac.in; debajitdatta.pd@es.iitr.ac.in; debajitdatta9@gmail.com <i>Etymology.</i> Generic name is a combination of <i>Tharo</i>, referring to the ‘Thar desert’ of western India where the type specimen was found, and <i>saurus</i>, which is derived from the Greek word ‘sauros’ meaning lizard; specific name is for the country of origin i.e., India. <i>Holotype.</i> RWR-241(A–K); partial middle/posterior cervical vertebra; middle/posterior cervical anterior condyle and right prezygapophyses; partial anterior dorsal neural arch; middle/posterior neural spines; anterior dorsal rib; partial anterior and middle caudal vertebrae (Figs. 2, 3, 4, 5; Supplementary Table 1). <i>Horizon and locality.</i> Fort Member of the Jaisalmer Formation (early–middle Bathonian), Jethwai village, District Jaisalmer, Rajasthan state, western India. <i>Diagnosis. Tharosaurus</i> exhibits a unique combination of the following characters: middle/posterior cervical centroprezygapophyseal lamina divided into lateral and medial branches, the latter connecting with the intraprezygapophyseal lamina (shared with all dicraeosaurids); middle/posterior cervical centropostzygapophyseal fossa elliptical and bordered laterally by pillar-like centropostzygapophyseal lamina (shared with the dicraeosaurids <i>Lingwulong</i>, <i>Brachytrachelopan</i>, and <i>Pilmatueia</i>); deep bifurcation of cervical neural arch extending up to the dorsal margin of the neural canal (shared with the dicraeosaurids <i>Amargasaurus</i> and <i>Pilmatueia</i>); paired fossae on the ventral surface of the middle/posterior cervical centrum separated by a mid-line keel (shared with all dicraeosaurids, barring <i>Smitanosaurus</i>, <i>Kaatedocus</i> and <i>Suuwassea</i>, and the diplodocids— <i>Barosaurus</i> and <i>Dinheirosaurus</i>); paired ventral fossae extending up to the posterior margin of centrum; anterior condyle of middle/posterior cervical more rugose compared to the rest of the centrum (shared with the dicraeosaurid <i>Kaatedocus</i>); bifid middle/posterior cervical neural spine (shared with Flagellicaudata); divided lateral fossa/ pleuroceol on cervical centra (shared with Flagellicaudata); lateral fossa divided into deep posterior and shallow anterior half by weak ridge on the ventral surface of the fossa; prominent lateroventral flanges on middle/posterior cervical centrum (shared with Flagellicaudata); middle/posterior cervical vertebrae with pre-epipophysis (shared with Flagellicaudata); cordiform cross-section of anterior caudal centra (shared with Flagellicaudata); middle/posterior dorsal transverse process laterally directed (shared with Diplodocidae); middle caudal centra articular surface with flat ventral margin (shared with Diplodocoidea); ventral surface of mid-caudal centrum with deep median fossa. Autapomorphies: triangular ventrolateral projections lying below posterior cotyle of middle/posterior cervical centrum visible in posterior view; ventral mid-line keel on middle/posterior cervical centrum bifurcating posteriorly but not meeting the lateroventral flanges; lateral pleuroceol on anterior caudal centra. <b>Description.</b> The cervical vertebrae are represented by five partial specimens found in association (Supplementary Table 1), including the anterior condyle, posterior half of a vertebra and three right prezygapophyses (Figs. 2, 3). The anterior condyle is U-shaped with an opisthocoelous condition as in other eusauropods 22, 23 (Fig. 2A), and is mediolaterally wider than dorsoventrally tall [RWR-241-A, centrum articular surface height (cH)/centrum articular surface width (cW)= 0.7; Supplementary Table 2, Supplementary Fig. 2]. In this respect, <i>Tharosaurus</i> is similar to the middle/posterior cervicals of several flagellicaudatans: <i>Suuwassea</i> 24 (cH/cW = 0.7), <i>Lingwulong</i> 13 (cH/cW =0.7), <i>Pilmatueia</i> 8 (CV7/8, cH/cW = 0.9), <i>Amargasaurus</i> 5 (CV8, cH/cW = 0.8), <i>Apatosaurus louisae</i> 25 (CV9, cH/cW = 0.7) and <i>Diplodocus carnegii</i> 26 (CV11, cH/cW = 0.9). The anterior condyle is markedly more rugose than the other vertebral elements, as in <i>Kaatedocus</i> 27 (CV14). The centrum shows strong anterior constriction with the lateral surfaces excavated by large fossae, similar to many flagellicaudatans 8, 13, 24, 26 (Fig. 2B–E). The fossa is partially preserved with much of the dorsal rim and medial wall broken off. The preserved portion suggests the fossa was elliptical, being elongated and dorsoventrally compressed (Fig. 2B–C). The dorsal margin of the fossa is rod-like and extends laterally beyond the ventral margin. The latter is robust, rounded and thickest near the posterior end of the centrum, accentuating the depth of the fossa. Anteriorly it becomes flush with the lateral surface of the centrum (Fig. 2B–C). Beneath the ventral margin of the fossa, the lateral surface of the centrum is markedly concave and flares posteriorly. The fossa is restricted to the posterior half of the centrum and includes two distinct halves: a short and shallow anterior half and a longer and deeper posterior half. The two halves are demarcated by a low U-shaped ridge on the ventral surface of the fossa, medial to the ventral margin and extending slightly onto the medial wall of the fossa. This contrasts with the dicraeosaurids <i>Amargasaurus</i>, <i>Dicraeosaurus</i>, <i>Bajadasaurus</i>, and <i>Pilmatueia</i> where the fossa is undivided and shallow 5, 7, 8. In the middle cervicals of <i>Lingwulong</i> and <i>Suuwassea</i>, however, the lateral fossa is deep and extends nearly along the entire length of the centrum 13, 24. Furthermore, the lateral fossa in <i>Tharosaurus</i> is succeeded by a posteroventral fossa (Fig. 2D,E), as seen in the mid-cervicals of <i>Amargasaurus</i>, <i>Apatosaurus</i>, <i>Bajadasaurus</i>, and <i>Pilmatueia</i> 26, 28. The ventral surface of the centrum (Fig. 2F–G) accommodates paired longitudinal depressions/fossae. Such depressions are common in diplodocids and most dicraeosaurids, barring <i>Bajadasaurus</i> which only bears a longitudinalkeel 7, 8, 13, 26. The fossae are asymmetrical, with the right fossa mediolaterally wider, and extend up to the posterior margin of the centrum where they flare out. However, in <i>Dicraeosaurus</i>, <i>Lingwulong</i>, and <i>Pilmatueia</i> the fossae are symmetrical and strongly expressed anteriorly, but do not reach the posterior margin of the centrum 8, 13, 26. The fossae in <i>Tharosaurus</i> are flanked laterally by prominent lateroventral flanges and separated by a sharp midline keel (Fig. 2F,G). The keel shows lef lateral convexity and remains prominent throughout its preserved length. It extends nearly up to the posterior margin of the fossae and bifurcates into two short ridges. The latter, however, do not reach up to the lateroventral flanges. An accessory ridge, present only on the right ventral surface, extends posterolaterally from the midline keel. The presence of a ventral keel is shared with a diverse array of flagellicaudatans including most dicraeosaurids (<i>Lingwulong</i>, <i>Dicraeosaurus</i>, <i>Pilmatueia</i>, <i>Bajadasaurus</i> and <i>Brachytrachelopan</i>) 7 – 9, 13 and some later-diverging diplodocids (<i>Barosaurus lentus</i>, <i>Diplodocus carnegii</i>, <i>Dinheirosaurus</i>) 23, 26. It is, however, absent in some putative plesiomorphic dicraeosaurids (<i>Suuwassea</i> and <i>Smitanosaurus</i>) 24, 29, barring <i>Kaatedocus</i> 27 which is the earliest-diverging dicraeosaurid. The disposition and morphology of this midline keel differentiate <i>Tharosaurus</i> from <i>Lingwulong</i>, <i>Dicraeosaurus</i>, <i>Brachytrachelopan</i> and <i>Barosaurus</i> where the keel is prominent anteriorly 4, 8, 9, 26. In <i>Dinheirosaurus</i> the keel is restricted to the posterior part of the centrum and does not extend anteriorly as in <i>Tharosaurus</i>, whereas in <i>Pilmatueia</i> the keel forks anteriorly and posteriorly with the bifurcation point placed slightly anterior to the mid-length of the centrum. The posterior cotyle of the centrum (RWR-241-B, cH/cW = 0.6, Fig. 2H,I, Supplementary Table 2) is strongly concave and mediolaterally wider than dorsoventrally tall, a feature seen in the middle cervicals of many sauropods including <i>Lingwulong</i> 13 (cH/cW =0.7), <i>Apatosaurus louisae</i> 25 (cH/cW= 0.7) and <i>Gleamopus</i> 30 (cH/cW= 0.9), and the posterior cervicals of <i>Pilmatueia</i> 8 (cH/cW = 0.6) and <i>Suuwassea</i> 24 (cH/cW = 0.7). The posterior cotyle is cordiform in outline with a weakly concave dorsal margin immediately ventral to the neural canal as in <i>Lingwulong</i> and <i>Pilmatueia</i> 8, 13. <i>Tharosaurus</i>, however, can be distinguished by the presence of posteriorly facing triangular facets along the ventrolateral margin of the posterior articular surface. The neural arch is partially preserved and includes the right prezygapophysis and the region surrounding the neural canal (Figs. 2B–K, 3, Supplementary Table 1). The anterior exit of the canal, although partial, appears oval, but the posterior exit is complete and elliptical, being transversely wider than dorsoventrally high. In <i>Pilmatueia</i> this exit is triangular, whereas in <i>Suuwassea</i> and <i>Lingwulong</i> it is circular–subcircular 8, 13, 24. The prezygapophysis is anterodorsally directed with the articular facet strongly convex transversely and longitudinally (Fig. 3A–G). The latter feature is shared with middle and posterior cervicals of the flagellicaudatans <i>Diplodocus carnegii</i> (CV8) and <i>Kaatedocus</i> (CV10) 26 – 29. The prezygapophyseal articular surface is oblong with the posterior border offset from the surrounding dorsal surface of the prezygapophysis by a transverse sulcus (Fig. 3C,D,G), similar to the posterior cervicals of <i>Kaatedocus</i> 27. Immediately ventral to the articular surface, the lateral surface of the prezygapophysis bears an anterodorsally-posteroventrally oriented pre-epipophysis (sensu Tschopp et al. 26; Fig. 3A,B). The latter is dorsoventrally compressed and succeeded ventrally by a shallow fossa. Such a ridge is also reported in the middle and posterior cervicals of <i>Kaatedocus</i> 27 (CV7–10), <i>Apatosaurus louisae</i> 26 (CV11) and <i>Suuwassea</i> 24, 26. The prezygapophysis is supported ventrally by a dorsoventrally high and robust centroprezygapophyseal lamina (cprl, Fig. 3C–F). The lamina is divided into a larger and medially concave lateral branch connecting with the prezygapophysis (lcprl) and a smaller medial branch joining the intraprezygapophyseal lamina (mcprl). The two halves of the cprl accommodate a triangular and deep centroprezygapophyseal fossa (Fig. 3C,D) which borders the neural canal dorsolaterally and is roofed by a sheet-like intraprezygapophyseal lamina. While a divided cprl is common in Flagellicaudata 1, in dicraeosaurids the medial branch of the cprl connects with the intraprezygapophyseal lamina and is listed as a synapomorphy of Dicraeosauridae (sensu Whitlock and Wilson Mantilla 29). Unlike in <i>Tharosaurus</i>, however, both branches of the cprl join the prezygapophysis in diplodocids 1, 26, 29, 31. Nonetheless, the anterior surface of the lcprl immediately ventral to the prezygapophyseal articular surface bears a depression giving the impression of a split cprl, where both branches connect to the prezygapophysis, akin to diplodocids. Further scrutiny shows this depression to be almost indistinct and unlike the morphology of the bifurcated crpl of diplodocids 26, 32. Furthermore, in diplodocids the apex of this bifurcation is ventrally directed in contrast to the incipient depression in <i>Tharosaurus</i> which is broad and dorsally convex. Therefore, the expression of the divided cprl in the new Indian taxon is closer to that of dicraeosaurids and supports its taxonomic allocation. The posterior exit of the neural canal is flanked dorsolaterally by two elliptical centropostzygapophyseal fossae [cpof(e)] which are bordered laterally by robust pillar-like centropostzygapophyseal laminae [cpol(e), Fig. 2H,I]. The cpof(e) are roofed by thin intrapostzygapophyseal laminae. The arrangement of the vertebral laminae and fossae in <i>Tharosaurus</i> is comparable with the condition in <i>Lingwulong</i>, <i>Brachytrachelopan</i>, <i>Dicraeosaurus</i>, <i>Amargasaurus</i> and <i>Pilmatueia</i> 4, 5, 8, 9, 13. However, in the latter two taxa, the cpof(e) are triangular. Furthermore, as in <i>Amargasaurus</i> 5, 8, the neural arch in <i>Tharosaurus</i> is devoid of a median tubercle. The preserved neural arch above the neural canal shows deep bifurcation which descends to the roof of the canal and encloses an anteroposteriorly extensive passage of uniform width (Fig. 2H–K; Supplementary Fig. 3). The broken ends of the neural arch dorsal to the cpof(e) suggest the presence of strongly divergent postzygapophyses. These features are reminiscent of the deeply bifurcated neural arch-spine complex of middle to cervicodorsal vertebrae of <i>Amargasaurus</i> and <i>Pilmatueia</i> 5, 8, 28. Moreover, the surface of the neural arch within this passage is finished (as seen in dorsal view, Fig. 2J,K), and bears rugose scars along the mid-line which possibly represent ligament scars. The scars largely occupy the posterior half of the passage and bordered laterally by sub-parallel striations. Rugose tuberosity on the floor of bifurcated neural arch-spine complexes of <i>Apatosaurus ajax</i> and <i>Dicraeosaurus</i> have been interpreted as ligament scars 33. Although <i>Tharosaurus</i> lacks the tuberosity, a feature shared with <i>Amargasaurus</i> 8, these scars attest to the presence of bifid neural spines in <i>Tharosaurus</i>. Similar bifid neural spines are listed as a synapomorphy of Flagellicaudata in previous studies (e.g., Wilson 1). The specimens (RWR-241-A–E) likely represent the 6th/8th cervical based on comparisons with the middle cervicals of <i>Lingwulong</i> and <i>Bajadasaurus</i>, and the middle–posterior cervicals of <i>Dicraeosaurus</i>, <i>Suuwassea</i>, <i>Amargasaurus</i>, <i>Pilmatueia</i> and <i>Kaatedocus</i> 5, 8, 13, 24, 27. Three isolated specimens including a partial neural arch and two partial neural arch-spine complexes are referable to the dorsal vertebral series (Fig. 4A–L, Supplementary Table 1). The neural arch comprises a stout, sub-horizontal transverse process with an anteroposteriorly broad and sigmoidal diapophysis (Fig. 4A–F). Anteriorly, the transverse process bears a broad and moderately deep prezygapophyseal centrodiapophyseal fossa (Fig. 4A,B), whereas its posterior surface bears a thin, mediodorsally-lateroventrally directed lamina, possibly representing the postzygodiapophyseal lamina (Fig. 4C,D). A deep centrodiapophyseal fossa (cdf) positioned ventral to the transverse process is bordered by an anteroventrally directed anterior centrodiapophyseal lamina and a near-vertical posterior centrodiapophyseal lamina (Fig. 4E,F). The specimen is tentatively assigned to the anterior dorsals based on similarity in overall morphology including laminae and fossae configuration with those of the flagellicaudatans <i>Apatosaurus louisae</i>, <i>Dinheirosaurus</i>, <i>Lingwulong</i> and <i>Amargasaurus cazaui</i> 5, 13, 23, 25, 26. The neural arch-spine complex is strongly compressed anteroposteriorly and expanded transversely, suggesting a more posterior position in the dorsal vertebral series (sensu McPhee et al. 34). The arch comprises a transversely broad and subtriangular spinoprezygapophyseal fossa (Fig. 4G,H). The transverse process is laterally directed, with a vertically oriented distal end, similar to all diplodocids 23, 35. This laterally oriented transverse process can be considered a local autapomorphy of <i>Tharosaurus</i> unlike a dorsolaterally oriented transverse process in other dicraeosaurids 5, 9, 13, 36. This feature may alternatively represent a symplesiomorphy, although Mannion et al. 23 listed the dorsolaterally projecting diapophysis in the dorsal vertebrae of the diplodocid <i>Dinheirosaurus</i> as a local autapomorphy. The neural spine is transversely expanded and non-bifid and the disposition of the lateral spinal margins suggests dorsal flaring in anterior/posterior views (Fig. 4G–J). This suggests affinity with the middle/posterior dorsal vertebrae because non-bifurcated neural spines appear from the 6th/7th dorsal of dicraeosaurids, as seen in <i>Lingwulong</i> and <i>Brachytrachelopan</i> 9, 13. Such flaring of the lateral spinal margins is also known from the middle and posterior dorsals of dicraeosaurids 4, 13, 35. In lateral view, the anterior and posterior borders of the neural spine remain subparallel (Fig. 4K,L), unlike the sub-triangular spinal lateral profile characteristic of many titanosauriforms 34, 37. The neural spine bears a sharp prespinal lamina placed along the longitudinal midline on the anterior surface (Fig. 4G,H). The prespinal lamina does not extend up to the base of the spine and terminates dorsal to the spinoprezygapophyseal fossa. This lamina, however, extends down to the base of the neural spine in the posterior dorsals of <i>Amargasaurus</i> and <i>Dicraeosaurus</i> 5, 36. Only a small part of the spinoprezygapophyseal lamina is preserved along the lateral margin of the anterior spinal surface (Fig. 4G,H). The spinodiapophyseal lamina is robust and broad, descending gently from the neural spine to the transverse process (Fig. 4G,H,K,L). The posterior surface of the spine preserves only the right spinopostzygapophyseal lamina (Fig. 4I,J). It is robust and well-rounded, extending medially from the lateral margin of the posterior spinal surface towards the midline. In lateral view, the anterior surface of the spinopostzygapophyseal lamina unites with the posterior surface of the spinodiapophyseal lamina as seen in the posterior dorsals of <i>Brachytrachelopan</i> 9 (Fig. 4K,L), and these laminae enclose the postzygapophyseal spinodiapophyseal fossa. Thus, these vertebral specimens likely belong to the middle/posterior dorsals. An isolated, nearly complete rib (Fig. 4M,N) is identified as the right anterior dorsal rib, based on similarity with that of <i>Galeamopus pabsti</i> 30. The specimen is Y-shaped, flares out proximally, and becomes dorsoventrally skewed distally. The rib lacks pneumatic openings as in diplodocoids but contrasts with those of titanosauriforms 38. The capitulum is elliptical, anteroposteriorly compressed, and directed anterodorsally. It is longer than the tuberculum and bears prominent striations on the anterior surface (Fig. 4M). The tuberculum is short and curved towards the capitulum. The tuberculum and capitulum enclose a broad U-shaped space. A robust and rounded ridge occupies the anterior surface of the rib distal to the capitulum and tuberculum (Fig. 4M). The ridge extends distally and gradually becomes flush with the surface. The ventral margin is sigmoidal with the portion immediately distal to the tuberculum strongly convex. The dorsal margin is concave, and its posterior surface is largely flat and featureless, except for the strongly rugose tuberculum. The collection includes two caudal vertebrae (Supplementary Table 1). One of these, a partial ce
description: Published as part of <i>Bajpai, Sunil, Datta, Debajit, Pandey, Pragya, Ghosh, Triparna, Kumar, Krishna &amp; Bhattacharya, Debasish, 2023, Fossils of the oldest diplodocoid dinosaur suggest India was a major centre for neosauropod radiation, pp. 1-15 in Scientific Reports 13 (1)</i> on pages 1-8, DOI: 10.1038/s41598-023-39759-2, http://zenodo.org/record/8268346
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