A New Specimen of the Sauropodomorph Dinosaur Ignavusaurus Rachelis from the Early Jurassic of Lesotho
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A new specimen of the sauropodomorph dinosaur Ignavusaurus rachelis from the Early Jurassic of Lesotho § Ewan H. Bodenham1 & Paul M. Barrett2,3* 1Department of Genetics, Evolution and Environment, University College London, Darwin Building, Gower Street, London WC1E 6BT, U.K. 2Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD, U.K. 3Evolutionary Studies Institute, University of the Witwatersrand, Johannesburg, Private Bag 3, WITS, 2050 South Africa Received 30 January 2020. Accepted 10 November2020 The upper Elliot Formation (?Rhaetian–Sinemurian) of South Africa and Lesotho has yielded a rich fauna of non-avian dinosaurs, which has generally been considered to be dominated by the massopodan sauropodomorph Massospondylus carinatus. However, re-evaluation of the abundant sauropodomorph collections from this unit suggests that the species-richness of upper Elliot sites has been underestimated. Here, we describe a series of cervical and dorsal vertebrae collected from Likhoele Mountain, Lesotho, which are referred to the rare upper Elliot sauropodomorph taxon Ignavusaurus rachelis. This material represents only the second-known specimen of this taxon, extending its geographic range, and underscores the value of undertaking detailed re-assessments of neglected historical collections. Keywords: Sauropodomorpha, Massospondylidae, upper Elliot Formation, taxonomy. Palaeontologia africana 2020. ©2020 Ewan H. Bodenham & Paul M. Barrett. This is an open-access article published under the Creative Commons Attribution 4.0 Unported License (CC BY4.0). To view a copy of the license, please visit http://creativecommons.org/licenses/by/4.0/. This license permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. The article is permanently archived at: https://hdl.handle.net/10539/30351 INTRODUCTION assessed more rigorously to demonstrate their taxonomic The Stormberg Group (Late Triassic–Early Jurassic) of affinities. Moreover, large historical collections of sauro- the main Karoo Basin has yielded abundant dinosaur podomorph material from the upper Stormberg Group, remains from numerous localities in South Africa and which have generally been referred to Massospondylus Lesotho (e.g. Kitching & Raath 1984; Knoll 2004, 2005). spp. solely on the basis of provenance, now require exten- Diverse faunas have been recovered from both the Upper sive re-evaluation in order to build a more accurate Triassic lower Elliot Formation (Norian–Rhaetian) and the picture of the species-richness and palaeoecology of Upper Triassic–Lower Jurassic (?Rhaetian–Pliensbachian) upper Stormberg dinosaur faunas (see also Barrett et al. upper Elliot and Clarens formations (e.g. Knoll 2004, 2005; 2019). Viglietti et al. 2020a,b). In each case, sauropodomorph One such collection is housed at the Natural History dinosaurs are the numerically dominant tetrapod taxa Museum, London, which includes specimens acquired (Kitching & Raath 1984). in the late nineteenth century (e.g. the holotype of Until recently, almost all of the sauropodomorph dino- ‘Massospondylus browni’: Seeley 1895). However, the bulk saur specimens recovered from the upper part of the of this material was collected by a series of Anglo-South Stormberg Group were assumed to be referable to the African expeditions to South Africa and Lesotho in the massospondylid Massospondylus carinatus (e.g. Kitching & 1960s that were led jointly by either the Natural History Raath 1984; Knoll 2005). However, taxonomic re-apprais- Museum/ Birkbeck College or University College London als, new discoveries and stratigraphic revisions have (UCL), both in collaboration with the Iziko South African demonstrated that the upper Elliot and Clarens forma- Museum (e.g. Crompton 1968). Although many of the tions hosted a much more diverse sauropodomorph ornithischian dinosaurs collected by these expeditions fauna (McPhee et al. 2017). In addition to M. carinatus, this were described relatively promptly (e.g. Thulborn included at least one small-bodied ?non-massopodan 1970, 1972, 1974; Santa-Luca et al. 1976), the majority (Arcusaurus pereirabdalorum), three other likely masso- of the sauropodomorph material recovered remains spondylids or putative massospondylids (Ignavusaurus undescribed and unprepared (but see Barrett & Yates rachelis, M. kaalae and Ngwevu intloko), and four sauro- 2006). Here, we report a new specimen of the rare upper podiforms (Aardonyx celestae, Antetonitrus ingenipes, Elliot Formation sauropodomorph Ignavusaurus rachelis Ledumahadi mafube and Pulanesaura eocollum) (Yates that was collected during one of the UCL expeditions & Kitching 2003; Barrett 2009; Knoll 2010; Yates to Lesotho in July–August 1968, which was led by the et al. 2010, 2011; McPhee et al. 2015, 2018; Chapelle late Prof. Kenneth Kermack. This represents only the et al. 2019). Consequently, new discoveries can no longer second-known specimen of Ignavusaurus and under- be assumed to represent Massospondylus but must be scores the importance of unlocking the information *Author for correspondence. E-mail: [email protected] stored in historical museum collections. Palaeontologia africana 54: 48–55 — ISSN 2410-4418 [Palaeontol. afr.] Online only Permanently archived on the 11th of December 2020 at the University of the Witwatersrand, Johannesburg, South Africa The article is permanently archived at: https://hdl.handle.net/10539/30351 48 ISSN 2410-4418 Palaeont. afr. (2020) 54: 48–55 SYSTEMATIC PALAEONTOLOGY Table 1. Measurements for NHMUK PV R37375. All measurements are in mm. Measurements marked with an asterisk are minima due to Dinosauria Owen, 1842 breakage. No measurements were possible for the cervical vertebra; in Saurischia Seeley, 1887 (sensu Baron et al. 2017) other cases extensive breakage or the presence of matrix and/or overlap- ping elements prevented measurement. Sauropodomorpha von Huene, 1932 ?Massospondylidae von Huene, 1914 (sensu Sereno 1998) ACH ACW CL NSH NSL NSW PCW Ignavusaurus rachelis Knoll, 2010, Figs 1 & 2 Cervicodorsal 1 – – – 29.8 22.5 19.34 – Material. NHMUK PV R37375, an articulated series of Cervicodorsal 2 – – – 23.1 23.1 15.2 – five neural arches (representing one cervical, two Dorsal 1 – – – 24.8 28.9 9.1 – Dorsal 2 27.8* 24.1 52.8* – – – 18.9* cervicodorsals and two dorsals), also including one Isolated neural arch – – – 15.1* 29.5* 8.1 – centrum (forming part of the posterior-most articulated vertebra), and one isolated partial ?dorsal neural arch. This specimen was collected on the 1968 UCL expedition section. The lack of parapophyses on the more complete and was originally given the field number ‘C.10’ in K.A. third vertebra in the series suggests that these two verte- Kermack’s unpublished field notes (held at NHMUK) and brae are at the neck-trunk junction and are considered to identified as ‘dinosaur vertebrae’ therein. be ‘cervicodorsals’ herein. The fourth and fifth vertebrae Locality and horizon. Likhoele Mountain, near Mafeteng, have parapophyses that are situated on the neural arch, Mafeteng District, Lesotho, from a quarry named ‘Dino- as well as mediolaterally flattened neural spines, so are saur exposure’ (K.A. Kermack, unpubl. field notes). No identified as dorsals, as is the isolated partial neural stratigraphic information is recorded in the field notes. arch. Neural arch terminology follows Wilson (1999) and However, a continuous section of upper Stormberg Wilson et al. (2011). Group sediments is exposed at this site, including the The cervical neural arch (vertebra 1 in the series) lacks its lower Elliot, upper Elliot and Clarens formations (Sciscio neural spine, prezygapophyses, most of the transverse & Bordy 2016), leaving open the possibility that any of processes and the ventral parts of the neural arch pedicels these units could have been the source of this specimen. (Figs 1–3). Although the prezygapophyses and neural An elevation of ‘6,680 feet’ (2035 m) was recorded for the spine are damaged, there are broken surfaces indicating quarry (K.A. Kermack, unpubl. field notes), which would that spinoprezygadiapopyseal laminae (SPRLs) were place the source of this material within the Clarens Forma- present, framing a deep prespinal fossa. In lateral view, tion, as the upper Elliot/Clarens formation contact is at the postzygapophyses are oriented horizontally and are 1967 m (Sciscio 2016). However, the Clarens Formation at extremely elongate, so it is likely that they would have Likhoele Mountain forms sheer vertical cliffs and it is overhung the posterior margin of the centrum (Figs 1A,C, unlikely that the specimen could have been recovered 2A,C). In dorsal view,the postzygapophyses taper posteri- from this part of the section; moreover, the red, orly to a rounded apex, diverging from each another at an fine-grained sandstone encasing the specimen is identical angle of approximately 30°. A blunt, ridge-like, low to that occurring at several levels within the upper Elliot epipophysis is present, which terminates anterior to the Formation at Likhoele Mountain (Sciscio & Bordy 2016; posterior tip of the postzygapophysis (Figs 1A,C, 2A,C). L. Sciscio, pers. comm.). Consequently, we regard the Breakage prevents determination of the size or shape of source horizon for NHMUK PV R37375 as the upper Elliot the neural spine or the presence/absence of spinopost- Formation, which is of ?Rhaetian–Sinemurian age (Sciscio zygapophyseal laminae (SPOLs). The portion of the et al. 2017; Bordy et al. 2020;