The Winston Churchill Memorial Trust of Australia

Report by Dr Stephen F. Poropat

2017 Churchill Fellow

To explore the impact of continental drift and climate change on Southern Hemisphere dinosaur faunas by studying ornithopod, theropod and sauropod fossils from Argentina and Antarctica, held in various institutions across Argentina.

I understand that the Churchill Trust may publish this report, either in hard copy or on the internet or both, and consent to such publication.

I indemnify the Churchill Trust against any loss, costs or damages it may suffer arising out of any claim or proceedings made against the Trust in respect of or arising out of the publication of any report submitted to the Trust and which the Trust places on a website for access over the internet.

I also warrant that my Final Report is original and does not infringe on the copyright of any person, or contain anything which is, or the incorporation of which into the Final Report is, actionable for defamation, a breach of any privacy law or obligation, breach of confidence, contempt of court, passing-off or contravention of any other private right or of any law.

Signed:

Dr Stephen F. Poropat 26/02/2019

Keywords: dinosaur, ornithopod, sauropod, theropod, Gondwana, Argentina, Antarctica, Australia

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Replica of Talenkauen santacrucensis as found in the field, at the Museo Argentino de Ciencias Naturales in Buenos Aires

Reconstructed skeleton of Talenkauen santacrucensis on display at the Museo Argentino de Ciencias Naturales in Buenos Aires

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Executive Summary Dr Stephen F. Poropat (Ph: +61 3 9214 5152) Postdoctoral Researcher (Palaeontology) Research Associate Faculty of Science and Technology Australian Age of Dinosaurs Museum of Swinburne University of Technology Natural History Hawthorn, Victoria, Australia 3122 Winton, Queensland, Australia 4735 Email: [email protected] Email: [email protected] Aim: To explore the impact of continental drift and climate change on Southern Hemisphere dinosaur faunas by studying ornithopod, theropod and sauropod fossils from Argentina and Antarctica, held in various institutions across Argentina. Highlights: Museo Argentino de Ciencias Naturales: Loncosaurus argentinus, Morrosaurus antarcticus Museo de La Plata: Trinisaura santamartaensis Museo Municipal Carmen Funes: Anabisetia saldiviai Centro Paleontológico Lago Barreales: Macrogryphosaurus gondwanicus, Gasparinisaura cincosaltensis Museo Regional de Cinco Saltos: Gasparinisaura cincosaltensis Universidad Nacional de la Patagonia San Juan Bosco: Notohypsilophodon comodorensis, Sektensaurus sanjuanboscoi Museo Regional Provincial Padre Manuel Jesús Molina: Talenkauen santacrucensis Project Description: I developed a comprehensive understanding of South American and Antarctic non-hadrosaurid ornithopod dinosaur anatomy. I will use this information in phylogenetic analyses incorporating all known South America, Antarctic and Australian ornithopods, and these will form the basis for palaeobiogeographic analyses interpreted in the light of palaeoclimatic and palaeoenvironmental changes throughout the Cretaceous. I also studied several titanosauriform sauropods and megaraptorid theropods, with a view to conducting similar research on them. Major lessons, conclusions and dissemination of information: Loncosaurus, Macrogryphosaurus, Sektensaurus and Talenkauen show remarkable morphological uniformity. Their narrow age range implies that they constitute a short-lived radiation of medium-sized ornithopods in Argentina, which might have extended into Australia via Antarctica. Gasparinisaura, Trinisaura and Morrosaurus are all tens of millions of years younger than all known Australian ornithopods and, despite being closer in size, are evidently more distantly related. Notohypsilophodon is morphologically similar and temporally proximal to Victoria’s ornithopods, but the only known specimen is a rather fragmentary subadult. The knowledge I gained during my Churchill Fellowship will greatly enhance my research on dinosaur specimens held in the collections of the Australian Age of Dinosaurs Museum (Winton, Queensland) and Museums Victoria (Melbourne). My students and I will write scientific and popular articles using data collected during this Fellowship, with a particular focus on the phylogenetic and biogeographic implications of Argentinian, Antarctic and Australian dinosaurs. I also disseminated some of my work through a blog during my trip (https://stephenporopat.weebly.com/blog). 4

Program for Churchill Fellowship to Argentina Dates Place Institution Contact(s) Specimen(s) studied 19/10/2018 Depart Melbourne 22/10/2018– Buenos Aires, Museo Argentino de Martin Ezcurra Loncosaurus argentinus 26/10/2018 Buenos Aires Ciencias Naturales [email protected] Morrosaurus antarcticus “Bernardino Rivadavia” Chubutisaurus insignis 29/10/2018– La Plata, Museo de La Plata Marcelo Reguero Trinisaura santamartaensis 02/11/2018 Buenos Aires [email protected] Antarctopelta oliveroi 05/11/2018– Plaza Huincul, Museo Municipal Rodolfo Coria Anabisetia saldiviai 09/11/2018 Neuquén Carmen Funes [email protected] Megaraptor namunhuaiquii Murusraptor barrosaensis 13/11/2018– Lago Barreales, Centro Jorge Calvo Macrogryphosaurus 19/11/2018 Neuquén Paleontológico Lago [email protected] gondwanicus Barreales Gasparinisaura cincosaltensis Megaraptor namunhuaiquii Futalognkosaurus dukei 20/11/2018 Cipolletti, Río Museo Provincial de Ignacio Cerda Rocasaurus muniozi Negro Cipolletti “Carlos [email protected] Ameghino” Virginia Zurriaguz [email protected] 21/11/2018 Neuquén, Universidad Juan Porfiri Megaraptor namunhuaiquii Neuquén Nacional del [email protected] Comahue 22/11/2018– Cinco Saltos, Museo Regional de Ignacio Cerda Gasparinisaura 23/11/2018 Río Negro Cinco Saltos [email protected] cincosaltensis Guido Riccono [email protected] 26/11/2018– Trelew, Museo Diego Pol Patagotitan mayorum 30/11/2018 Chubut Paleontológico [email protected] Egidio Feruglio 03/12/2018– Comodoro Universidad Rubén Martínez Notohypsilophodon 07/12/2018 Rivadavia, Nacional de la [email protected] comodorensis Chubut Patagonia San Juan Marcelo Luna Sektensaurus sanjuanboscoi Bosco [email protected] Un-named megaraptorids 11/12/2018– Río Gallegos, Museo Regional Claudia Aguilar Talenkauen santacrucensis 14/12/2018 Santa Cruz Provincial Padre [email protected] Manuel Jesús Molina 18/12/2018 Return to Melbourne

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Introduction My childhood dream was to become a palaeontologist. I was utterly fascinated by dinosaurs and other prehistoric lifeforms. Moreover, I wondered what Earth was like when the dinosaurs were alive during the Mesozoic Era (251–66 million years ago), and how much our planet had changed between their first appearance in the fossil record (~230 million years ago) and the extinction event 66 million years ago that wiped out all dinosaurs other than birds. As a child, I was also keenly aware that our understanding of what Australia was like during the Mesozoic Era was limited. When I was born, only nine Mesozoic dinosaurs had been named based on bones found in Australia (there are now 23; still relatively few, but a definite improvement). Even back then, several American states, at least one Canadian province and several parts of China could boast more dinosaur species than could our entire country. I was determined to change that, and to enhance our understanding of the Australian Mesozoic. I undertook a BA/BSc at Monash University from 2003–2006, majoring in Geography and Geology with a minor in Biology. In so doing, I acquired the requisite skillset to do an Honours year in palaeontology in 2007. Throughout my undergraduate studies, I volunteered on the annual Dinosaur Dreaming digs on the Bass Coast. However, being somewhat concerned about my job prospects, I shifted away from studying fossil vertebrates and studied invertebrate microfossils instead, since they have applications in hydrocarbon exploration. The most important thing I learned during my Honours year, and during the PhD that followed (from 2008–2011), was that a job in the petroleum industry was not for me, for many reasons. However, I was at a bit of a loss as to what to do next. In 2010, nearing the end of my PhD and concerned about what was to come after, a colleague of mine, Prof. Patricia Vickers-Rich, informed me about an unusual job being offered in Sweden. The successful applicant was expected to spend part of each year in Uppsala, and part in a remote town in Queensland named Winton. Winton was familiar to me as the home of the Lark Quarry Dinosaur Stampede, and I also knew of it because three new dinosaurs from Winton were formally named and described in 2009. Intrigued, I resolved to apply for the job, and was encouraged to visit Winton as a result. This took place in late 2010. During my first week in Winton, I met David Elliott (awarded a Churchill Fellowship in 2011), the co- founder of the Australian Age of Dinosaurs Museum (AAOD hereafter), as well as Trish Sloan and George Sinapius who were, and still very much are, integral members of the AAOD team. David, Trish, George and AAOD opened my eyes regarding the palaeontological potential of central Queensland: we did indeed have dinosaurs, they were just harder to find than had previously been appreciated! I have since been up to Winton between one and five times a year, every year since 2011. I landed the job at Uppsala University (2011–2014) and started working with AAOD. When the job ended, I moved back to Australia and continued my association with AAOD, even when full-time work was difficult to find. In 2017, I was employed by Swinburne University of Technology to continue working with the AAOD, but with the Mesozoic fossil collections of Melbourne Museum at my fingertips as well. These presented quite a different kettle of (highly-derived) fish. Winton has rightly become famous for producing bones of dinosaurs called sauropods, which count among their ranks the largest land animals of all time by any measure (the tallest, longest, and 6

heaviest). All bones found in Winton Shire since 2004 have stayed in Winton at AAOD, meaning that that museum’s collection is dominated by sauropod specimens. By contrast, Melbourne Museum — which houses all of the Mesozoic fossil specimens found in Victoria — houses no sauropod remains. The most common dinosaurs by far in Victorian Mesozoic rocks are ornithopods. These are the dinosaurs that seem to get the least attention from the general public, although one group of ornithopods, the hadrosauroids (especially Parasaurolophus), gets a little bit of airtime. No hadrosauroids have been found in Australia; however, non-hadrosauroid ornithopods are the most commonly found dinosaurs in both Victoria (four named species) and New South Wales (three). One of my tasks in my job at Swinburne University of Technology is to study the ornithopod dinosaurs in the Melbourne Museum, in order to understand their evolutionary relationships and their palaeoecology. However, in order to understand their evolution and their palaeobiogeography in particular, it is necessary to broaden our horizons somewhat. The simple fact is that Australia was not an island continent during the mid-Cretaceous (~125–105 Ma), when ornithopods reigned supreme in Victoria. Instead, Australia was firmly connected to Antarctica via Tasmania. Further west, South America was also linked to Antarctica, albeit intermittently: what was, at times, a more or less contiguous land bridge between the two had a more archipelagic aspect at other times. This would have had a marked effect on the ability of terrestrial vertebrates — like dinosaurs — to disperse between South America, and the conjoined Australian– Antarctic continent. In 2016, when I and my colleagues announced the discovery of Savannasaurus elliottorum (Australia’s 20th named Mesozoic dinosaur), we also presented the results of a study aimed at elucidating their palaeobiogeographic signal; that is, what the position of our sauropods on the dinosaur family tree told us about where and when they might have originated (Poropat et al., 2016). We determined that our sauropods were very basal titanosaurs. Titanosaurs as a group were fashionably late to the Mesozoic party, only appearing, and eventually dominating, during the Cretaceous (the third and final period of the Mesozoic Era). However, as far as the fossil record has revealed, they were also the only sauropods to survive until the end of the Cretaceous. Their bones are found worldwide in Upper Cretaceous rocks and, in some places in Lower Cretaceous rocks too. Their presence in Australia in the earliest Late Cretaceous — the same time that the most basal titanosaur by definition, Andesaurus delgadoi, was living in Argentina — suggests that titanosaurs as a whole might have originated somewhere between Australia + Antarctica + South America, then dispersed across the globe. Alternatively, they might have evolved earlier and only reached Australia relatively late in the game. More research is required, and it was hinted that enhancing the sauropod analyses through the inclusion of more species — and the running of similar analyses focused on other dinosaur and tetrapod groups — would help to reveal the pattern of dispersal between Australia + Antarctica and South America throughout the Cretaceous in response to sea level, climatic and tectonic changes. To this end, I realised I needed to understand South American and Antarctic non-hadrosaurid ornithopods much better. Having now completed my Churchill Fellowship, I am pleased to report that I have studied virtually every single South American and Antarctic non-hadrosaurid ornithopod specimen ever described, as well as an ankylosaur, a few theropods, and several sauropods. Although I definitely now need a break from bones, it was amazing. 7

Acknowledgments First and foremost: to the Winston Churchill Memorial Trust, it has been an absolute privilege and an honour to have become a Churchill Fellow, and to have had the opportunity to study dinosaur fossils in Argentina. The photographs, measurements and observations that I was able to collect on this trip are utterly invaluable and will feature prominently in my and my students’ ongoing research on Australian dinosaurs and their phylogenetic relationships and palaeobiogeographic implications. Thank you for everything. To all of my hosts at the various museums and universities, I extend you my warmest regards and my sincerest thanks. Each and every one of you made me feel welcome in your institution, your town, and your country. The jobs you do — maintaining palaeontological collections, conducting and communicating scientific research, and welcoming researchers like myself with open arms, is far too often undervalued and under-appreciated. I assure you that I could not value or appreciate it more. Specifically, I would like to thank the following people at the following institutions for their hospitality: • Martin Ezcurra (Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”); • Marcelo Reguero and Julia Desojo (Museo de La Plata); • Rodolfo Coria, Guillermo Windholz, Mattia Baiano and Ludmila Coria (Museo Municipal Carmen Funes) • Jorge Calvo and Juan Mansilla (Centro Paleontológico Lago Barreales, Universidad Nacional del Comahue); • Ignacio Cerda and Virginia Zurriaguz (Museo Provincial de Cipolletti “Carlos Ameghino”); • Juan Porfiri and Domenica dos Santos (Universidad Nacional del Comahue); • Ignacio Cerda and Guido Riccono (Museo Regional de Cinco Saltos); • Diego Pol and José Carballido (Museo Paleontológico Egidio Feruglio); • Rubén Martínez and Marcelo Luna (Universidad Nacional de la Patagonia San Juan Bosco), and • Claudia Aguilar, Patricia Campan, Yael Güemes and Tiziana Bellini (Museo Regional Provincial Padre Manuel Jesús Molina). I also want to acknowledge the staff at the hotels in which I stayed, the chefs and waiters at the restaurants in which I ate, the non-technical staff at the institutions in which I studied, the pilots of the planes I caught and the drivers of the buses and taxis I rode in, and the operators at the various tourist destinations I visited. I am keenly aware of how privileged I was to be able to spend eight weeks in Argentina doing what I love doing, while all of you were working to make a living and possibly not enjoying it as much as I. To all those Argentines who welcomed me at hotels, exchanged money, booked taxis, cleaned rooms, made breakfasts, and generally made my trip to your wonderful country safe and amazing, muchas gracias. David Elliott’s (Australian Age of Dinosaurs) support throughout my career to date in vertebrate palaeontology, and his encouragement for my Churchill Fellowship application, has been immense. Thank you so much David, your immediate family (Judy, Bob and Harry), and to the “extended family” at the Australian Age of Dinosaurs Museum that you have assembled — you really are changing the face of Australian Mesozoic palaeontology. 8

To Patricia Vickers-Rich (Swinburne University of Technology) and Thomas Rich (Melbourne Museum): you were my inspiration when I was a kid, and it is still sometimes hard to believe that I work for, and with, you now. It is a dream come true. Thank you. To all of my colleagues, friends and family in Melbourne: your support throughout my eight weeks away, never mind the preceding several decades, has been massive. I love you all. Mum and Dad: you are the best parents conceivable. You have supported me throughout my life in so many ways, and have helped my aspirations become a reality. Thank you so much. Finally, spending eight weeks away from home, more or less by myself, in a country where relatively few people speak English, was always going to be lonely. When I planned my Churchill Fellowship itinerary, I had no idea what the months between finalising flight times and hotel details and actually embarking on the trip would hold for me. As it turns out, I met the woman of my dreams just weeks before I was to depart for Argentina… Amy, you were there for me every step of the journey on my Churchill Fellowship, despite being 10,000 km away. Our talks almost every morning and evening, and on your lunchbreaks on workdays, meant the world to me. Put simply, you made this trip so much more enjoyable, and the only way that it could have been more enjoyable still would have been if you were able to accompany me on it.

With a life-sized restoration of Patagotitan mayorum on the outskirts of Trelew, Chubut Province, Argentina

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Overview The primary objective of my Churchill Fellowship was to gather as much anatomical and palaeoenvironmental information as possible about the non-hadrosaurid ornithopod dinosaurs of Argentina and Antarctica. This is important to my research because one of the two fossil vertebrate collections in Australia to which I have essentially unrestricted access — the Cretaceous collection at Melbourne Museum, Museums Victoria (formerly the National Museum of Victoria) — houses abundant remains of ornithopod dinosaurs from sites along the Victorian coast. The majority of the hundreds of ornithopod specimens in the Museums Victoria collections are isolated bones. However, four partial skeletons have also been recovered since the late 1980s. Two of these have historically been attributed to Leaellynasaura amicagraphica (Rich & Rich, 1989; Rich et al., 2010): the first (NMV P185992–3) preserves an exceedingly long tail (comprising more than 70 vertebrae) in connection with a partial pelvis and hind limb (Herne et al., 2016), whereas the second (NMV P186047, a.k.a. ‘Junior’) preserves a more complete and better preserved pair of hind limbs, portions of the pelvis and the vertebrae from the hip region and the base of the tail (Gross et al., 1993; Herne et al., 2016; Poropat et al., 2018). Both of these specimens were excavated from the now famous Dinosaur Cove site in the 1980s, and the latter is significant inasmuch as it preserves evidence of the bone disease osteomyelitis in its left shin (Gross et al., 1993; Woodward et al., 2018). Several kilometres east of Dinosaur Cove, another specimen (NMV P221080) was collected in 2005 (Rich et al., 2009), and in early 2018 was described as the new species Diluvicursor pickeringi (Herne et al., 2018). The fourth and final specimen, discovered in 2010 near Inverloch and nicknamed ‘Noddy’ (because it was preserved in a nodule), remains undescribed but is probably the most exciting of the ornithopods found in Victoria to date. This is because it preserves a partial skull, a complete neck, and much of the thorax. Along with researchers based in the United Kingdom and Australia, I am going to be describing this specimen and comparing it — where possible — with other Cretaceous ornithopod specimens. Those that should show the greatest degree of similarity to ‘Noddy’ are from South America and Antarctica — hence the reason for this Churchill Fellowship. Only three partial ornithopod skeletons (Hooker et al., 1991; Coria et al., 2013; Rozadilla et al., 2016) and two isolated elements pertaining to hadrosaurs (Rich et al., 1999; Case et al., 2000) have been found in Antarctica. One of the non-hadrosaurid ornithopod specimens from Antarctica (NHMUK BAS R.2450) is reposited in the Natural History Museum in London; however, despite the fact that it was discovered in 1989 — and reported on briefly in 1991 (Hooker et al., 1991; Thomson & Hooker, 1991) — it has never been described (Barrett et al., 2014). I asked permission to see the specimen in 2017, but was denied. As such, I decided to focus on the two published specimens, both of which are held in Argentinian museums and both of which date to near the end of the Cretaceous Period. Perhaps unsurprisingly, the Argentinian ornithopod record is far better than that of Antarctica (Cruzado-Caballero, 2016). It is also substantially better than that of Australia, comprising a greater number of partial skeletons spanning a far greater age range. The majority of the record — including all of the named taxa — is restricted to the Late Cretaceous (Coria & Cambiaso, 2007; Coria, 2016). What this means is that the South American/Antarctic and Australian ornithopod records are complementary: The Australian ornithopods span the end of the Early Cretaceous and the earliest Late Cretaceous, whereas those of South America and Antarctica span the Late Cretaceous. 10

Hadrosaurs (often informally, albeit incorrectly, referred to as “duckbills”) have an extensive record in the South American Late Cretaceous (Casamiquela, 1964; Martinelli & Forasiepi, 2004; Coria, 2015), and several taxa have been named. These include Secernosaurus (Brett-Surman, 1979; Bonaparte et al., 1984; Bonaparte & Rougier, 1987; Prieto-Márquez & Salinas, 2010; Becerra et al., 2018), Willinakaqe (Powell, 1987; Juárez Valieri et al., 2010; Cruzado-Caballero & Coria, 2016) and Lapampasaurus (González Riga & Casadío, 2000; Coria et al., 2012). However, hadrosaurs are not known from Australia, and I decided not to spend any time working on them during the Fellowship. Taken together, the fossil records of Australia, Antarctica and South America have the potential to elucidate how ornithopods, the most abundant small-bodied herbivores of the time, responded to climate change and continental drift throughout the Cretaceous. By also incorporating the sauropod and theropod records, a fairly complete picture of dinosaurian response to natural climatic and geographic shifts throughout the Cretaceous in the Southern Hemisphere should be approachable. My mission was to personally observe all of the named ornithopod specimens from Antarctica (Table 1) and Argentina (Table 2) for the purposes of better learning ornithopod anatomy and variability, and for comparison with the Australian specimens (Table 3). Along the way, I also attempted to observe as many relevant theropod and sauropod specimens as possible. In this report, I will only present details of my work on the ornithopod dinosaurs. However, I also worked on the sauropods Chubutisaurus insignis (Del Corro, 1975; Salgado, 1993; Carballido et al., 2011), Futalognkosaurus dukei (Calvo et al., 2007a, b; Calvo, 2014), Rocasaurus muniozi (Salgado & Azpilicueta, 2000; García & Salgado, 2013), and Patagotitan mayorum (Carballido et al., 2017), as well as the ankylosaur Antarctopelta oliveroi in La Plata (Gasparini et al., 1987, 1996; Salgado & Gasparini, 2006). Finally, I also had the opportunity to study several megaraptorid theropod specimens in detail. In Plaza Huincul, I studied Murusraptor barrosaensis (Coria & Currie, 2016; Paulina-Carabajal & Currie, 2017), focusing mainly on the teeth. In Comodoro Rivadavia, I was given permission to study several unpublished megaraptorid specimens (Lamanna, 2004; Casal et al., 2016). However, the most important and intriguing of all the theropod specimens that I studied were the three best exemplars of Megaraptor namunhuaiquii: the holotype in Plaza Huincul (Novas, 1998), a more complete referred specimen in Lago Barreales (Calvo et al., 2004), and the juvenile specimen in Neuquén (Porfiri et al., 2014).

Table 1: Antarctic ornithopod body fossils (non Hadrosauroidea)

Identification Specimen(s) Formation Locality Age Reference(s) Morrosaurus MACN Pv Cape Lamb El Morro (‘The lower Cambiaso (2007); Reguero et antarcticus 19777 Member, Snow Naze’), James Maastrichtian al. (2013a); Reguero et al. (holotype) Hill Island Ross Island (2013b); Rozadilla et al. (2016) Iguanodontia NHMUK BAS Cape Lamb Cape Lamb, lower Hooker et al. (1991); Thomson indet. R2450 Member, Snow Vega Island Maastrichtian and Hooker (1991); Reguero et Hill Island al. (2013a, b); Barrett et al. (2014) Trinisaura MLP 08-III-1-1 Gamma Santa Marta upper Coria et al. (2013); Reguero et santamartaensis (holotype) Member, Snow Cove, James Campanian al. (2013a); Reguero et al. Hill Island Ross Island (2013b) Ornithopoda MLP 07-III-2- Gamma Santa Marta upper Coria et al. (2007b); Reguero et indet. 1–2 Member, Snow Cove, James Campanian al. (2013a); Reguero et al. Hill Island Ross Island (2013b) 11

Table 2: Argentinian ornithopod body fossils (non Hadrosauroidea)

Identification Specimen(s) Formation Locality Age Reference(s) Iguanodontia indet. MEPyG-177 Allen Salitral Ojo de Agua, middle Campanian– Coria et al. (2007a) Río Negro lower Maastrichtian Gasparinisaura MUCPv-208 Anacleto near Cinco Saltos lower Campanian Coria and Salgado (1996a); cincosaltensis (holotype), 212–218, Cemetery, Río Negro Salgado et al. (1997); Coria 225–227; MCSPv 1–3, (1999); Heredia and Salgado 111–112 (1999); Cambiaso (2007); Coria and Cambiaso (2007); Cerda (2008); Novas (2009); Cerda and Chinsamy (2012) Mahuidacursor MAU-Pv-CO-596 Bajo de la Cerro Overo, Neuquén Santonian Cruzado-Caballero et al. (2019) lipanglef (holotype) Carpa Ornithopoda indet. MAU-Pv-CO-564– Bajo de la Cerro Overo, Neuquén Santonian Cruzado-Caballero et al. (2018) 565, 569–572, 574–579 Carpa Ornithopoda indet. MAU-Pv-LE-616–8 Bajo de la La Escalonada, Santonian Jiménez-Gomis et al. (2018) Carpa Neuquén Ornithopoda indet. MPCA-AT 261 Plottier El Anfiteatro, Rio upper Coniacian–upper Salgado et al. (2009) Negro Santonian Ornithopoda indet. MAU-Pv-PH-458 Plottier Puesto Hernández upper Coniacian–upper Cruzado-Caballero et al. (2016) quarry, Neuquén Santonian Macrogryphosaurus MUCPv-321 Sierra near Mari Menuco middle–upper Coniacian Calvo et al. (2007c); Novas gondwanicus (holotype) Barrosa Lake, Neuquén (2009); Garrido (2010) Ornithopoda indet. MCF-PVPH-73, 165 Portezuelo ?Cerro Bandera, Coniacian Coria (1999) Neuquén Iguanodontia indet. MUCPv-352, etc. Portezuelo localities near Lago Coniacian Porfiri and Calvo (2002); Calvo Barreales, Neuquén and Porfiri (2003); Calvo et al. (2007a) Iguanodontia indet. MCF-PVPH-167 Portezuelo Cerro Bandera, Coniacian Coria (1999); Coria and Neuquén Cambiaso (2007) Sektensaurus UNPSJB-PV 960 + 973 Lago Colhué Lago Colhué Huapi, Coniacian–Maastrichtian Luna et al. (2003); Ibiricu et al. sanjuanboscoi + 1054 (holotype) Huapi Chubut (2010); Ibiricu et al. (2019) Ornithopoda indet. MOZ-PV 020 Huincul Agrio del Medio, upper Cenomanian– Canudo et al. (2013) Neuquén Turonian Iguanodontia indet. MCF-PVPH-479 Huincul near Plaza Huincul, upper Cenomanian– Coria et al. (2007a) Neuquén Turonian Anabisetia saldiviai MCF-PVPH-74 Cerro Cerro Bayo Mesa, Cenomanian Coria (1999); Coria and Calvo (holotype), 75–77 Lisandro Neuquén (2002); Cambiaso (2007); Coria et al. (2007a); Novas (2009) cf. Gasparinisaura MCF-PVPH-168 Cerro near Neuquén Cenomanian Coria (1999) Lisandro Notohypsilophodon UNPSJB-Pv 942 Bajo Barreal near Buen Pasto, Cenomanian Martínez (1998); Cambiaso comodorensis (holotype) Chubut (2007); Coria and Cambiaso (2007); Novas (2009); Ibiricu et al. (2014) Loncosaurus MACN [1]1629 Mata Pari-Aike, Río Sehuen, Cenomanian Ameghino (1899); Molnar argentinus (holotype) Amarilla Santa Cruz (1980a); Coria and Salgado (1996b); Cambiaso (2007); Coria and Cambiaso (2007); Novas (2009) Talenkauen MPM-10001A Mata Los Hornos Hill, Santa Cenomanian Novas et al. (2004); Cambiaso santacrucensis (holotype), 10001B Amarilla Cruz (2007); Coria and Cambiaso (2007); Novas (2009); Egerton et al. (2013)

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Table 3: Australian and New Zealand ornithopod body fossils

Identification Specimen(s) Formation Locality Age Reference(s) Ornithopoda indet. NZGS CD529 Maungataniwha Mangahouanga Campanian– Wiffen and Molnar (1989); Wiffen Sandstone Stream, New Maastrichtian (1996); Agnolin et al. (2010) Zealand Ornithopoda indet. QM F52774 Winton near Winton, uppermost Hocknull and Cook (2008) Queensland Albian– ?lowermost Turonian Weewarrasaurus LRF 3076 (holotype), 766 Griman Creek Lightning Ridge, upper Albian / Bell et al. (2018); Bell et al. (2019) pobeni New South Wales Cenomanian Fostoria LRF 3050 (holotype) Griman Creek Lightning Ridge, upper Albian / Bell et al. (2019); Bell et al. (in press) dhimbangunmal New South Wales Cenomanian Fulgurotherium NHMUK R3719 (holotype) Griman Creek Lightning Ridge, upper Albian / Huene (1932); Molnar (1980a); australe New South Wales Cenomanian Molnar and Galton (1986); Bell et al. (2019) Ornithopoda indet. QM F various; AM F Griman Creek Lightning Ridge, upper Albian / Molnar and Galton (1986); Bell et al. various; LRF 660 New South Wales Cenomanian (2018); Bell et al. (2019) Iguanodontia AM F81865, 119236, Griman Creek Lightning Ridge, upper Albian / Molnar and Galton (1986); Bell et al. indet. 119282; LRF 492, 1556 New South Wales Cenomanian (2018) Ankylopollexia LRF 267 Griman Creek Lightning Ridge, upper Albian / Bell et al. (2018); Bell et al. (2019) indet. New South Wales Cenomanian Muttaburrasaurus QM F6140 (holotype) Mackunda near Muttaburra, upper Albian Bartholomai and Molnar (1981); langdoni Queensland Molnar (1995); Molnar (1996a); Agnolin et al. (2010) Muttaburrasaurus QM F12541, F14921 Allaru Mudstone near Hughenden, upper Albian Molnar (1996a) sp. Queensland Ornithopoda indet. QM F10942 Allaru Mudstone near Hughenden, upper Albian Molnar (1980a); Molnar (1980c); Queensland Molnar (1984) Ornithopoda indet. In the United Kingdom Toolebuc Near Boulia, upper Albian Molnar (1984) Queensland Leaellynasaura NMV P185991 (holotype), Eumeralla Dinosaur Cove, upper Aptian– Rich and Rich (1988); Rich and Rich amicagraphica 185990 (?), 185992–3 (?), Victoria lower Albian (1989); Gross et al. (1993); Agnolin P186047 (?) et al. (2010); Rich et al. (2010); Woodward et al. (2011); Herne (2014); Herne et al. (2016); Poropat et al. (2018); Woodward et al. (2018) Atlascopcosaurus NMV P166409 (holotype), Eumeralla Point Lewis, upper Aptian– Rich and Rich (1989); Agnolin et al. loadsi 157390 Victoria lower Albian (2010); Herne (2014); Poropat et al. (2018) Diluvicursor NMV P221080 (holotype) Eumeralla Eric the Red West, upper Aptian– Rich et al. (2009); Herne (2014); pickeringi Victoria lower Albian Herne et al. (2018); Poropat et al. (2018) Ornithopoda indet. NMV various Eumeralla Several localities on upper Aptian– Rich and Vickers-Rich (1999); the Otway Coast, lower Albian Agnolin et al. (2010); Woodward et Victoria al. (2011); Herne (2014); Poropat et al. (2018); Woodward et al. (2018) Qantassaurus NMV P199075 (holotype), Upper Strzelecki Flat Rocks, Victoria upper Rich and Vickers-Rich (1999); intrepidus 186389, 186456, 198962, Group Barremian– Agnolin et al. (2010); Herne (2014); 199087, 208093 lower Aptian Poropat et al. (2018) Ornithopoda indet. NMV various Upper Strzelecki Several localities on upper Rich and Rich (1989); Vickers-Rich Group the Bass Coast, Barremian– et al. (1999); Agnolin et al. (2010); Victoria lower Aptian Woodward et al. (2011); Poropat et al. (2018); Woodward et al. (2018)

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Table 4: Institutional Abbreviations Abbreviation Institution AM Australian Museum, Sydney, New South Wales, Australia LRF Australian Opal Centre, Lightning Ridge, New South Wales, Australia MACN Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina MAU Museo Municipal Argentino Urquiza, Rincón de los Sauces, Neuquén, Argentina MCF-PVPH Museo Municipal Carmen Funes, Paleontología de Vertebrados, Plaza Huincul, Neuquén, Argentina MCSPv Museo Regional de Cinco Saltos, Cinco Saltos, Río Negro, Argentina MEPyG Museo Educativo de Paleontología y Geología del Instituto de Formación Docente Continua de General Roca, Río Negro, Argentina MLP Museo de La Plata, La Plata, Argentina MOZ Museo Provincial de Ciencias Naturales “Prof. Dr. Juan A. Olsacher” de Zapala, Neuquén, Argentina MPCA Museo Provincial de Cipolletti “Carlos Ameghino”, Cipolletti, Río Negro, Argentina MPEF Museo Paleontológico Egidio Feruglio, Trelew, Chubut, Argentina MPM Museo Regional Provincial Padre Manuel Jesús Molina, Río Gallegos, Argentina MUCPv Museo de Geologia y Paleontologia de la Universidad Nacional del Comahue, Neuquén, Argentina NHMUK Natural History Museum, London, United Kingdom NMV Museums Victoria (National Museum of Victoria), Melbourne, Victoria, Australia NZGS New Zealand Geological Survey Collection, Lower Hutt, New Zealand QM Queensland Museum, Brisbane, Queensland, Australia UNPSJB Universidad Nacional de la Patagonia San Juan Bosco, Comodoro Rivadavia, Argentina

Type specimen of Andesaurus delgadoi on display at Centro Paleontológico Lago Barreales, Universidad Nacional del Comahue

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Megaraptor menacing and munching on Macrogryphosaurus at the Museo Argentino de Ciencias Naturales in Buenos Aires

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Stop 1: Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires

The large unenlagiine dromaeosaurid theropod Austroraptor cabazai, on display at the Museo Argentino de Ciencias Naturales

The Museo Argentino de Ciencias Naturales in Buenos Aires is one of the oldest and largest natural history museums in Argentina. Historically, fossils from all over the country were sent to Buenos Aires for study, and then reposited there — as happened with Loncosaurus argentinus from Santa Cruz Province. However, in the last few decades there have been changes to the law regarding where fossils are to be kept in Argentina. Now, fossils are not to permanently leave the province in which they are found, and they are to be reposited in the closest museum (that is able to store fossils) to the fossil site. Relatively few dinosaur specimens have been lodged in the Museo Argentino de Ciencias Naturales in the last few decades. However, one of the few is an Antarctic ornithopod of great interest. 16

Loncosaurus argentinus Ameghino, 1899

Type femur of Loncosaurus argentinus in five different views. It was probably better to start than finish with this specimen… The ornithopod dinosaur Loncosaurus argentinus was named on the basis of a single incomplete femur (MACN 1629 or MACN 11629) from Santa Cruz Province. Although it was originally described (Ameghino, 1899) and later maintained (Huene, 1929) as a theropod, part of the reason for this was its apparent association with an undoubted theropod tooth (MACN 10985) in the original description (Ezcurra & Agnolín, 2017). However, it was subsequently realised that the tooth and the femur came from different animals, and that the femur was actually from an ornithopod (Molnar, 1980a). Following on from that realisation, Loncosaurus was fully redescribed but regarded as a nomen vanum (Coria & Salgado, 1996b), despite the recognition of an apparently unique trochanteric fossa near the fourth trochanter. More recently, it was upheld as a potentially valid ornithopod (Coria & Cambiaso, 2007), included in a phylogenetic analysis for the first time [albeit in an unpublished thesis (Cambiaso, 2007)], and, most recently, maintained as a nomen vanum (Cruzado-Caballero et al., 2018). The Loncosaurus femur provides relatively little anatomical information but is of significance nonetheless because it is possibly one of the oldest ornithopod body fossils from South America. Unfortunately, the age of the formation from which it is thought to derive — the “Mata Amarilla Formation” — is not well constrained (Varela et al., 2012). 17

None of the Australian ornithopod femora that I have observed so far possess the trochanteric fossa that is present on Loncosaurus, implying that it might only be distantly related to them. However, I will have to conduct a more thorough survey of the Australian material to ascertain whether any of them possess this feature. Before personally observing the ornithopod Talenkauen santacrucensis, which derives from the same formation that Loncosaurus is thought to have been collected from, I wondered if it, too, would possess a trochanteric fossa on its femur. Based on my observations, it certainly does — and no other ornithopod specimen that I saw during my Churchill Fellowship, or that I have read about in published papers, possesses this feature. This might imply that Loncosaurus and Talenkauen are actually one and the same. If this is formalised, the name Loncosaurus might have to take priority because it was named more than 100 years before Talenkauen.

My work station at the Museo Argentino de Ciencias Naturales, with the femur of Loncosaurus argentinus in the spotlight

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Morrosaurus antarcticus Rozadilla, Agnolin, Novas, Aranciaga Rolando, Motta, Lirio & Isasi, 2016 To date, Antarctica has only produced evidence of dinosaurs from the Jurassic and Cretaceous periods. All of the Jurassic specimens (of which I am aware) are held in institutions in the USA. However, fortunately for me, almost all of the Cretaceous-aged dinosaur specimens collected from Antarctica are housed in Argentina. At the Museo Argentino de Ciencias Naturales I was privileged to be able to study the most recently named dinosaur from Antarctica — Morrosaurus antarcticus. The only known specimen of Morrosaurus was collected by geologists Juan M. Lirio and Héctor Nuñez in the early 2000s. The first description of MACN Pv 19777 — the specimen that was to become Morrosaurus antarcticus — was produced by Andrea Cambiaso five years later and included as part of her Ph.D. dissertation (Cambiaso, 2007). However, Cambiaso never published on this specimen, and evidently did not consider it worthy of a name since she did not give it one. In 2016, Sebastian Rozadilla and colleagues decided that it was worth naming MACN Pv 19777, and they dubbed it Morrosaurus antarcticus (Rozadilla et al., 2016). When I first saw the type specimen of Morrosaurus, I was struck by how large it was (relatively speaking; large for a small ornithopod, if you will). In fact, the same was true of Loncosaurus — both it and Morrosaurus were evidently quite a bit larger than even the largest of the known Cretaceous ornithopods from Victoria. Leaellynasaura, Atlascopcosaurus, Qantassaurus and Diluvicursor were quite titchy by comparison. Talenkauen, Sektensaurus and Macrogryphosaurus were larger still. The type specimen of Morrosaurus comprises bits and pieces of a right hind leg. The shafts of the femur (thighbone) and the tibia and fibula (shinbones) are missing, but at least their top and bottom ends are preserved (only one end in the case of the fibula). By contrast, the metatarsals (the upper toe bones in the foot bound in flesh) are almost complete. I suspect that the only reason that they are incomplete is because of repeated freezing and thawing of the specimen: the part of Antarctica from which Morrosaurus derives is, in the modern day, covered by ice in winter but not summer. The metatarsals are quite distinctive: the middle one was evidently massive, whereas the flanking metatarsals are distinctly compressed, meaning that the metatarsus as a whole would have comprised a tightly-packed bundle of bones. Whether Morrosaurus possessed a rudimentary first and/or fifth toe is unknown based on the available material.

Left: reconstructed right hind limb of Morrosaurus antarcticus, viewed from the front, with proportions of missing elements based on Anabisetia saldiviai. Photos by the author 19

Morrosaurus lived during the Maastrichtian (72–66 Ma), the last stage of the Cretaceous Period (after which non- avian dinosaurs were no more). At that time, Antarctica was in much the same position on the planet as it is today, but the planet as a whole was far warmer. Consequently, the southernmost continent appears to have been ice-free all year round — at least as far as the geological record tells us. Instead, it was covered in forest, and was evidently quite hospitable for dinosaurs — remains of ornithopods, ankylosaurs, sauropods and theropods (including birds) have all been found in the Snow Hill Island Formation, the rock unit from which Morrosaurus derives. Specifically, Morrosaurus itself was found in the Cape Lamb Member at a locality called “The Naze”, more properly known as Península el Morro (from which its name derives), on James Ross Island. Intriguingly, this specimen comes from the same geological unit as the undescribed Antarctic ornithopod held in Right metatarsals of Morrosaurus antarcticus, viewed from the back the United Kingdom, which might either represent a new species of another specimen of Morrosaurus (Hooker et al., 1991; Thomson & Hooker, 1991; Reguero et al., 2013a, b; Barrett et al., 2014).

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Stop 2: Museo de La Plata, La Plata

With a glyptodont for a guide, I was surely in the right place. The Museo de La Plata is a grandiose building, with equally grandiose fossil galleries. During the late 19th Century and early 20th Century, an abundance of fossil material from all over Argentina was sent here for safekeeping and study. Consequently, historically important dinosaur holotypes — like Genyodectes, Argyrosaurus, Neuquensaurus and Amygdalodon — are held here. Recently implemented laws have stemmed the flow of fossils from other territories into the Museo de La Plata; however, since the 1980s, it has been a hub for researchers working on Antarctic fossil specimens, including the ornithopod Trinisaura and the ankylosaur My, what big teeth you have: Genyodectes serus holotype Antarctopelta. These latter were my focus…

Ground sloths & glyptodonts: Lestodon, Glossotherium × 2, Scelidotherium, Sclerocalyptus, Glyptodon, Panochthus, Doedicurus 21

Trinisaura santamartaensis Coria, Moly, Reguero, Santillana & Marenssi, 2013 In January 2008, geologists Juan Moly and Rodolfo Coria collected the fossilised skeleton of a small ornithopod in Santa Marta Cove on James Ross Island, Antarctica. When they collected it was, and remains, the smallest ornithopod recovered from Antarctica with an estimated overall length of 1.5 metres. Despite the effects of recent freeze- thaw cycles, a substantial portion of the skeleton was able to be recovered by Moly and Reguero. This then underwent preparation and conservation at Museo de La Plata, and by the early 2010s the specimen (MLP 08-III-1-1) was ready for description. Rodolfo Coria lead authored the paper (with Moly and Reguero as coauthors), published in 2013, that officially named this Antarctic ornithopod

The type specimen of Trinisaura santamartaensis in left side view, assembled Trinisaura santamartaensis (Coria from my photos of the original bones et al., 2013). The “Trini” part of the genus name honours Trinidad

Diaz, a geologist who did pioneering geological studies on the Antarctic Peninsula, whereas the species name alludes to Santa Marta Cove, where the specimen was discovered. The holotype and only known skeleton of Trinisaura includes several dorsal (thoracic), sacral (hip) and caudal (tail) vertebrae, both shoulder girdles, a bone identified as a humerus, four of the six bones in the pelvis (both ilia, and the right pubis and ischium), and several elements from the right hind leg and a few from the left one. Altogether, then, we have a good understanding of the relative size and shape of Trinisaura. More importantly, it has proven possible to identify enough significant anatomical features in this specimen to differentiate it from other ornithopods and to assess its position on the ornithopod family tree. In their 2013 paper describing the new dinosaur, Rodolfo Coria and colleagues included Trinisaura in a phylogenetic analysis to determine its position among other ornithopods. They found that it sat on a branch among ornithopods from South America, including Gasparinisaura, Talenkauen and Anabisetia. In 2016, Sebastian Rozadilla and colleagues found that Trinisaura was part of the same group as all three of these ornithopods, although they found that they formed a natural evolutionary 22

group together with Morrosaurus from Antarctica, and the Argentinian Macrogryphosaurus and Notohypsilophodon (Rozadilla et al., 2016). In 2018, Daniel Madzia and colleagues ran analyses which suggested that the same group of ornithopods was more inclusive still, since they also found that two Australian ornithopods — Atlascopcosaurus and Qantassaurus — were also part of this group, which is known as Elasmaria (Madzia et al., 2018). The fact that the Antarctic Trinisaura and Morrosaurus, and ornithopods from Argentina Tail vertebrae of Trinisaura santamartaensis, still embedded and Australia, share several anatomical features in the Antarctic rock in which they were found and seem to be more closely related to one another than to ornithopods from elsewhere, suggests that there was a radiation of these plant-eating dinosaurs in the southern supercontinent Gondwana during the Cretaceous Period. Why this was so is unclear, although it might have been related to their geographic or climatic isolation from more northern ornithopod faunas. Indeed, despite the fact that the Cretaceous ornithopod record of Africa is poor, and that India and Madagascar have yielded none to date, it is interesting to note that one of the few known African taxa — Kangnasaurus coetzeei (Haughton, 1915; Cooper, 1985) — also seems to show ties to this group. Hopefully, more fossils from the continents that formerly constituted Gondwana will be discovered and described to shed further light on this ornithopod radiation.

Articulated right pelvis of Trinisaura santamartaensis, viewed from the right hand side 23

Stop 3: Museo Carmen Funes, Plaza Huincul

Reconstructed Argentinosaurus skeleton: simply gigantic With the immense reconstructed skeleton of Giganotosaurus

Battle of the theropods: Patagonykus versus two Unenlagia Aucasaurus garridoi restored skeleton and in situ replica The Museo Municipal Carmen Funes in Plaza Huincul is probably best known for housing the type specimen of Argentinosaurus huinculensis, one of the largest sauropods — and land animals — of all time (Bonaparte & Coria, 1993). Unfortunately, during my visit, the galleries at the Museum were all closed for refurbishment. Nevertheless, curator Rodolfo Coria was kind enough to let me see the displays – and to spend a week behind the scenes! Few real fossils are on display — Argentinosaurus being a notable exception. However, behind the scenes there were so many specimens, many of them holotypes, that it was hard to know where to begin and where to stop! While in Plaza Huincul, I worked most days from 10am to 7pm without a lunch break; a surprise asado (Argentinian barbeque) on the Holding the holotype claw of Megaraptor namunhuaiquii! Tuesday with the staff was a welcome change. 24

Anabisetia saldiviai Coria & Calvo, 2002

Skeletal reconstruction of Anabisetia saldiviai at the Museo Municipal Carmen Funes, with rather elongate neck The four known specimens of Anabisetia saldiviai are all housed at the Museo Municipal Carmen Funes, and I spent several days there learning as much as I could about them. Put simply, they were absolutely amazing to work with. Most of the material is undistorted, and much of it was found articulated. The only reason that we know about Anabisetia at all is that Roberto Saldivia, a farmer from Plaza Huincul, discovered some bones on his land and reported his find to palaeontologists. If this story sounds familiar, it is because many dinosaur discoveries in Queensland, Australia are made in exactly this way — by graziers, working their land or mustering stock. The significance of the collection accrued by the Australian Age of Dinosaurs Museum in Winton since 1999 attests to the critical role that people living on the land play in enhancing our palaeontological understanding. Back in 1993, Saldivia brought a few fossilised bone fragments to the Museo Municipal Carmen Funes, where they were soon recognised as ornithopod remains. As a result of his discovery, an expedition was mounted to Cerro Bayo Mesa (30 km south of Plaza Huincul), and — with Saldivia’s help — four partial ornithopod skeletons were soon uncovered. Although the discovery was briefly mentioned six years later, and although the proposed name for the new ornithopod was leaked (Coria, 1999), almost a decade elapsed between the discovery of the new ornithopod and its official naming and description. Rodolfo Coria and Jorge Calvo finally announced Anabisetia saldiviai to the world in 2002 (Coria & Calvo, 2002). The genus name honours archaeologist Ana Biset, who helped to sort out provincial fossil legislation in Argentina, whereas the species name honours Roberto Saldivia for discovering Anabisetia. Although the original paper is quite concise and does not provide many measurements, it is quite well-illustrated. However, it includes no specimen photographs. My mission at the Museo Municipal Carmen Funes was to measure, photograph and make my own observations of Anabisetia. 25

Articulated right back foot of Anabisetia saldiviai Right hand of Anabisetia saldiviai viewed from the side The bones comprising each Anabisetia specimen were stored in big wooden boxes, with the individual fossils supported in polystyrene trays. The sole exception was a complete hind foot, maintained in the position in which it was found, which was kept in what looked like an upside-down fish tank sitting on a small trap door. One of the specimens in the trays included a tiny, nearly complete hand with five fingers. Over a period of three days I compiled a pretty comprehensive database on Anabisetia, so much so that I could just about write a complete osteology of the animal (with a little help…). One thing I did notice was that some of the “individuals” included elements that had to come from other animals. In one specimen, a complete left ilium (upper hip bone) was present, along with a second, incomplete left ilium; in another, three humeri (upper arm bones) were present. In each case, one specimen was preserved differently to the rest of the registered material, so it seemed prudent to exclude it; however, it does mean that there are more than four Anabisetia individuals known! While studying Anabisetia I realised that the proportions of its hind limbs were quite different to those of a Victorian ornithopod specimen nicknamed ‘Junior’ (NMV P186047), sometimes referred to Leaellynasaura amicagraphica. The shins of Anabisetia were only slightly longer than the thighs, whereas in ‘Junior’ the shins were about 40% longer than the thighs. This almost certainly reflects a behavioural disparity between ‘Junior’ and Anabisetia (in the parlance of some palaeos, ‘Junior’ was more cursorial: a better runner), and I made sure to take the requisite measurements of all the other ornithopods I observed to see if they were closer to Anabisetia or ‘Junior’ in this regard. Another thing I found curious was this: one specimen of Anabisetia preserved both thighbones, but these two bones looked completely different from one another despite being effectively identical in size and preservation. These differences had to be due to distortion, either before or during the fossilisation process. I think this will impact our understanding of ornithopod diversity in southeastern Australia where, on the basis of thighbones alone, it has been hypothesised that up to five different species were present (Rich & Rich, 1989). Lastly, I was intrigued to learn that all of the Anabisetia specimens had been found in close proximity to one another. Small-bodied ornithopods seem to be found clustered together fairly often: examples I can think of include Dysalotosaurus lettowvorbecki at Tendaguru in Tanzania (Janensch, 1955), the numerous Hypsilophodon foxii specimens recovered from a small area on the Isle of Wight (Galton, 1974), and the specimens attributed to Leaellynasaura amicagraphica from Dinosaur Cove in Victoria (Rich & Rich, 1989). It would be really interesting to find out why this is so. 26

Stop 4: Centro Paleontológico Lago Barreales, Lago Barreales, Neuquén

Studying the neck vertebrae of Futalognkosaurus dukei

With the gigantic left femur of Futalognkosaurus dukei Nasty looking arm and hand of Megaraptor namunhuaiquii The Centro Paleontológico Lago Barreales, which is part of the Universidad Nacional del Comahue, sits on the north shore of Lago Barreales, about 80 km northwest of Neuquén. This lake, and its near neighbour Lago Marí Menuco, are artificial: once dry basins, a deliberate partial deviation of the Neuquén River has caused them to fill with water. As a result of numerous fossil discoveries in the area — particularly at a site now known as Futalognko — the Centro Paleontológico Lago Barreales was built. This facility operates as a museum, an educational centre and a tourist attraction, and I could not help being reminded of the Australian Age of Dinosaurs Museum in Winton the whole time that I was there. To the south of the Centre lies the lake and the Futalognko fossil site itself; to the north, cliffs comprising Cretaceous sedimentary rocks. 27

Macrogryphosaurus gondwanicus Calvo, Porfiri & Novas, 2007

The type specimen of Macrogryphosaurus gondwanicus in left side view, assembled from my photos of the original bones

In May 1999 — before the Centro Paleontológico Lago Barreales was built — the Universidad Nacional del Comahue ran a geological field trip to Lago Marí Menuco, Lago Barreales’ next door neighbour. While there, the team was informed by a small boy named Rafael Moyano of a dinosaur discovery that he had made. The specimen was collected by the university, and in 2007 Jorge Calvo, Juan Porfiri and Fernando Novas described it as the new ornithopod species Macrogryphosaurus gondwanicus — the “Gondwanan big enigmatic lizard” (Calvo et al., 2007c). Originally, it was thought to have derived from the Portezuelo Formation; however, more recently it has been attributed to the Sierra Barrosa Formation (Garrido, 2010). The Macrogryphosaurus type specimen was found to be remarkable in several ways: bony plates were present on the rib cage that were only otherwise known from a few other ornithopods; the sternum had projections in three different directions; and the neck was quite long by ornithopod standards, comprising at least ten vertebrae. When I had spoken with Rodolfo Coria about my trip post-Plaza Huincul, I had mentioned that I was planning to visit Lago Barreales to see ornithopods. Rodolfo told me that the Macrogryphosaurus type specimen was so beautifully preserved that it was as if it had died yesterday. On hearing this, I could not wait to see it, so I did what was at that time the next best thing: I re-read the Macrogryphosaurus paper (Calvo et al., 2007c). Honestly, I was a bit underwhelmed; the bones looked great in the figures, but not earth-shatteringly awesome. I decided to reserve judgment until I saw the specimen for myself. Once I started studying Macrogryphosaurus firsthand, I could not agree more with Rodolfo’s assessment of it: it is amazing. Most of the bones are beautifully preserved, even though many are broken and the vertebrae between the hips have suffered minor crushing. Perhaps the best thing about the specimen is that there is so much of it: most of the neck, the entire thorax, the pelvis and a significant portion of the tail. The ribs and several dorsal vertebrae have been left in articulation, as they were found, which impeded my attempts to photograph them somewhat. Nevertheless, I was happy with the results. I spent two days, working from 9am to 9pm, on Macrogryphosaurus, annotating the original description (Calvo et al., 2007c) with my own observations. I obtained a full set of measurements and photographs of the specimen, and I now have an excellent understanding of its overall anatomy. On the whole, the vertebrae were quite accurately

Thorax of Macrogryphosaurus, showing vertebrae and ribs in situ described by Calvo et al. (2007c), and I

28 identified very few points of difference on the specimens before me. However, in my opinion, the pelvic elements were less well interpreted by Calvo et al. (2007c); I wrote several pages of notes detailing my own thoughts. As an example, the preacetabular process of the ilium was described by Calvo et al. (2007c) as being triangular in cross-section, strongly excavated medially, and slightly concave dorsally and ventrally, with the medial surface bearing a horizontal ridge for the transverse processes of the sacral vertebrae. The way I saw it, “This is a little misleading. Triangular is not really the shape: because of the degree of the development of the medial concavity, the right preacetabular process looks like an italicised “L”, then a “C” in anterior view (whereas the left looks like a horizontally flipped italicised “L”, then a “C”). Anterior to the mid- length of sacral 2, the preactabular process has one surface oriented dorsomedially–ventrolaterally and another horizontal surface projecting medially (which, as stated by Calvo et al. (2007) supports the transverse process of the sacrals — HOWEVER: the transverse process should be attached to the MEDIAL surface of it, not sitting on its dorsal surface! The sacrum has evidently been transversely compressed). Posterior to the level of the mid-length of sacral 2, the “C” shape develops: the dorsal surface becomes flat and develops a ventrally-projecting lip on its medial margin. A true lateral margin also comes into being at about the level of the pubic peduncle, and the medially-projecting lip deviates to project ventrally and merge with the pubic peduncle.” What the above suggests is that Calvo et al. (2007c) did not account for the side-to-side crushing of the pelvis and sacrum that has occurred, and described the bone as if it was undistorted. The other pelvic elements, the pubes and ischia, were hard to work with because they were broken. However, the difficulty was even greater because both of each were present, and the lefts and rights had often broken in the same places! Working out which fragments went with which others was challenging, but achievable eventually. For some reason, Calvo et al. (2007c) did not described what the distal end of the pubis looked like; I decided this Pelvis of Macrogryphosaurus viewed from the top, crushed side-to-side was important (and was glad I did when studying Talenkauen later on in the trip!), so here’s what I said: “The distal end of the pubis is shaped like a fat piece of Trivial Pursuit-style pie: the dorsal–lateral– ventral margin is broadly convex, with straight dorsomedial and ventromedial margins. The pubis articulates with the distal end of its opposite number on the dorsomedial margin.” I also decided it would be worth describing the distal end of the ischium, since little information given by Calvo et al. (2007c): “The lateral surface is flat; the medial one bears a prominent ridge which starts at the dorsal margin but migrates to the ventral one along the length of the middle third of the medial surface. This was easily observable on both ischia. Dorsal to this, at about 5/6 the length of the shaft, another medial ridge 29

develops which continues along the medial surface to the distal end. The distal ends of the ischia have small “boots” that project more ventrally than dorsally. Viewed distally, the ischia are diamond shaped. It would seem the ventromedial surfaces were the ones that articulated; the dorsomedial ones appear to have formed the cloacal channel.” The fact that I took these notes was absolutely critical later in the trip when I was studying Sektensaurus and Talenkauen: the distal end of the ischium of Sektensaurus was interpreted as a distal radius (forearm bone) by Ibiricu et al. (2019), whereas the distal ends of Talenkauen’s ischia were described as distal fibulae (shin bones) by Cambiaso (2007)! As far as I am aware, this is the first time that these elements have been identified correctly. It would not have been possible without Macrogryphosaurus. Now that I have had the privilege of studying Macrogryphosaurus, I have a much improved understanding of its anatomy. At five or six metres in total length, Macrogryphosaurus was relatively big for a non-hadrosaurid ornithopod. During my Fellowship I studied several ornithopods from Argentina (Talenkauen, Sektensaurus, Loncosaurus) that were similar in size and anatomy to Macrogryphosaurus, an could understand why they had been classified as the unique group Elasmaria. It also seemed reasonable Left and right pubes and ischia (and vertebrae) of Macrogryphosaurus to speculate that the undescribed, similarly-sized Antarctic ornithopod might be an elasmarian. However, one thing that is holding back our understanding of elasmarian evolution is the geological age of these specimens, since several are poorly constrained. For example, Talenkauen and Loncosaurus were originally thought to be from the end of the Late Cretaceous but are now thought to be from the start of it, whereas Sektensaurus’ potential age range spans more than 20 million years.

The neck of Macrogryphosaurus gondwanicus, which is more than 1 metre long despite missing the front few vertebrae

Finally, the neck of Macrogryphosaurus made me think of an east African antelope called a gerenuk. Gerenuks have much longer necks than other antelope, allowing them to browse at higher levels than most other species. They also routinely rear up on to their hind legs so that they can reach even higher into the trees — up to two metres. Perhaps Macrogryphosaurus was putting its relatively long neck to a similar use. Given that the neck of Talenkauen was similarly long, this feature — unique to elasmarians among ornithopods — might have adapted to enable acquisition of a specific foodstuff. 30

Gasparinisaura cincosaltensis Coria & Salgado, 1996: Part I

The beautiful type skull of Gasparinisaura cincosaltensis: photograph (top) and interpretive schematic (bottom) by the author

At the start of 1996, the year I finished high school, non-hadrosaurid ornithopods from South America were effectively unknown. That year, however, a paper published in Journal of Vertebrate Paleontology changed that (Coria & Salgado, 1996a). Several partial skeletons were reported, one of which preserved an almost complete skull. These specimens painted a vivid picture of an ornithopod little bigger than 31

a chicken. Rodolfo Coria and Leonardo Salgado named the new ornithopod Gasparinisaura cincosaltensis, honouring the place in which it was found (Cinco Saltos) as well as Zulma Gasparini, a world-renowned Argentinian palaeontologist. The type skull of Gasparinisaura is absolutely exquisite. To be able to see the ridges and wear facets on the teeth and the holes that facilitated the passage of blood vessels into the jaw bones was simply phenomenal. One thing I had not realised before observing the skull for myself was that the palpebral — the bone above the eye socket that might have made these small In situ forelimbs and ribs of Gasparinisaura cincosaltensis ornithopods look perpetually angry — was broken, displaced inwards, and distally incomplete. This was not evident from the published illustrations or descriptions, but might prove important in our work on ‘Noddy’. The bones preserved with the skull of Gasparinisaura were similarly spectacular, including the partial shoulder and pelvic girdles and even some complete limb bones. In their initial publication, Coria and Salgado (1996a) mentioned only two specimens of Gasparinisaura, and almost all of the illustrations they provided (none of which were photographs) were based on the skeleton with the skull. The other specimen they reported, a partial tail, was only briefly summarised. However, the following year, Salgado et al. (1997) reported the discovery of several new specimens of Gasparinisaura. They revealed that the locality from which the material was collected was situated 2.5 km southeast of the Cinco Saltos Cemetery — a juxtaposition of two graveyards of markedly disparate ages. In total, they reported twelve new Gasparinisaura specimens of varying levels of completeness: some comprised substantial portions of skeletons, others were only represented by one or two incomplete bones. Taken together, these specimens spanned quite a size range: from sub-chicken-sized individuals that were almost certainly juveniles to substantially larger exemplars. However, even the largest were still quite small by dinosaur standards! On one of the thigh bones of a particularly large Gasparinisaura individual, I observed a puncture mark that was clearly made prior to the bone being fossilised. This specimen was not illustrated by Salgado et al. (1997), but they Femur of Gasparinisaura cincosaltensis showing bite mark 32 did briefly allude to it. Their conclusion as to the origin of the puncture seems perfectly reasonable to me: they interpreted it as a crocodylomorph tooth mark. This implied that this and other Gasparinisaura individuals might have fallen victim to an aquatic predator attack as they came down to drink at a waterhole. Whatever the crocodylomorphs did not eat was eventually buried in sediment at the bottom of the water body and, ultimately, fossilised. However, it is possible that the Gasparinisaura individuals congregated around a drying waterhole, died, and were scavenged by opportunistic crocodylomorphs who scattered their carcasses. With these new specimens available, the overall anatomy and appearance of Gasparinisaura became rather well understood. Some of the new specimens included portions of the tail, bones from the upper arm and forearm, partial pelves, articulated ankles, and even in two instances a fairly complete foot. However, many of the long bones (especially those of larger individuals) were incomplete: their top and/or bottom ends were often preserved, but their shafts were almost always missing. Unfortunately, this meant that gauging the Complete, and tiny, relative proportions of the hind limbs was difficult. femur of Gasparinisaura This was one of my frustrations reading about Gasparinisaura. Based on all that I saw at Lago Barreales, it also seemed to be an insurmountable problem: few of the limb bones were complete, and few specimens preserved bones from both the fore- and hind limbs, making it difficult to get a sense of Gasparinisaura’s overall proportions. I took photographs and measurements of all the material I could lay my hands on, and resolved to do my best. I should not have worried — the specimens I studied in Cinco Saltos the week after answered most of my questions!

Partially restored back foot of Gasparinisaura In situ right shoulder girdle and humerus of Gasparinisaura 33

Stop 5: Museo Provincial de Cipolletti “Carlos Ameghino” (Cipolletti, Río Negro): 20/11/2018

Me with the left pelvic bones of Rocasaurus muniozi The restored holotype skull of Abelisaurus comahuensis

The holotype skull of the unenlagiine dromaeosaurid theropod Buitreraptor gonzalezorum. Small, sublime, and so delicate…

The Museo Provincial de Cipolletti “Carlos Ameghino” houses some amazing fossils. Most of the ornithopod material held there pertains to hadrosaurids (Juárez Valieri et al., 2010; Cruzado- Caballero, 2017); the rest is very fragmentary. However, the only known South American ankylosaur material is there too (Salgado & Coria, 1996; Coria & Salgado, 2001). I wanted to study this material to compare it with Minmi paravertebra (Molnar, 1980b; Molnar & Frey, 1987) and Kunbarrasaurus ieversi (Molnar, 1996b; Molnar & Clifford, 2000, 2001; Leahey et al., 2015) from Queensland, and isolated ankylosaur specimens from Queensland (Leahey & Salisbury, 2013) and Victoria (Barrett et al., 2010). Unfortunately, it was being worked on by a student, and I could not study it. However, there were other specimens that I wanted to see, primarily the sauropod Rocasaurus (Salgado & Azpilicueta, 2000; García & Salgado, 2013). I worked on Rocasaurus for a day in order to better understand its vertebral and pelvic anatomy, and I was also able to see several other specimens, including the sauropods Bonitasaura and Pellegrinisaurus and the theropods Abelisaurus and Buitreraptor. 34

Stop 6: Universidad Nacional del Comahue (Neuquén, Neuquén): 21/11/2018

Juvenile Megaraptor right maxilla (upper jaw bone with teeth) Juvenile Megaraptor skeletal remains including vertebrae

Cast of juvenile Megaraptor skeleton as it was found in the field, on display at Centro Paleontológico Lago Barreales

Having observed the holotype and referred specimens of Megaraptor namunhuaiquii in Plaza Huincul and Lago Barreales respectively, I made it my mission to ‘complete the set’, as it were, by seeing the referred juvenile specimen of Megaraptor described by Porfiri et al. (2014). Unlike most of the dinosaurs in the Universidad Nacional del Comahue collections, this specimen is not out at Lago Barreales — although that institution does retain a cast of the skeleton as it was found in the field, showing the position of the spinal column, skull bones, and girdle and limb elements. The real bones of this specimen are kept in the Geology Department of the Universidad Nacional del Comahue’s Neuquén City campus, so I planned to spend an entire day there studying them. Unfortunately, I was only given four hours to do so; nevertheless, I managed to take measurements and photos of all the specimens, and gained an excellent understanding of the animal’s anatomy. I remain somewhat unconvinced that this specimen truly pertains to Megaraptor, given that very few of the preserved bones overlap with the type or referred specimen. However, it is clearly a juvenile megaraptorid. 35

Stop 7: Museo Regional Cinco Saltos (Cinco Saltos, Río Negro): 22–23/11/2018

All of the known specimens of the small ornithopod Gasparinisaura — with the exception of a partial femur provisionally assigned to the genus by Coria (1999) — derive from a single locality near the cemetery of the small town of Cinco Saltos. Indeed, the type and only species of Gasparinisaura is named after the town. Much material has left Cinco Saltos, but some remains in the Museo Regional Cinco Saltos — which occupies a long disused railway station. The quaint nature of the Museo Regional Cinco Saltos building does not detract from the significance of the dinosaur specimens held there. One such specimen is the most complete skeleton of the titanosaurian sauropod Neuquensaurus ever found. Mostly, though, their collection is dominated by specimens of Gasparinisaura, and some of these are absolutely spectacular.

With a skeleton of Gasparinisaura cincosaltensis With one of the two skulls of Gasparinisaura cincosaltensis 36

Gasparinisaura cincosaltensis Coria & Salgado, 1996: Part II

In situ skeleton of Gasparinisaura cincosaltensis, showing the fore- and hind limbs, pelvis, and gastroliths (stomach stones) After observing several partial skeletons of Gasparinisaura at Centro Paleontológico Lago Barreales, you could be forgiven for thinking that I would have been ‘Gasparinisauraed out’ and that I’d never want to see another specimen. However, I knew there were more skeletons of this small ornithopod at Museo Regional Cinco Saltos: some assigned to the genus by Salgado et al. (1997), others by Cerda (2008). Using one of these specimens, I was finally able to collect a comprehensive series of limb and girdle measurements for Gasparinisaura — data which was previously unavailable. The specimen in question is phenomenal: the bones from the thorax, pelvis and all four limbs are all preserved in life position, with the knees bent, the toes curled up and the hands spread out. Despite the fact that the new specimens he announced were quite complete and well preserved, Ignacio Cerda’s main interest in Gasparinisaura was not anatomical in his 2008 paper. Instead, Cerda (2008) focused on what lay within the abdominal regions of both new specimens, as well as what had been found in association with the type specimen: gastroliths, also known as stomach stones! To find them associated with three specimens of the same species, all derived from the same locality, with no other rocks in the associated sediment, meant that their interpretation as gastroliths was unequivocal. Nevertheless, one has to wonder why Gasparinisaura needed gastroliths at all. Its teeth were clearly adapted for mastication, so did they aid in digestion? Perhaps, as a small dinosaur, Gasparinisaura needed to extract as much nutriment from its food as possible, and in order to achieve this it needed to pulverise whatever plants it was eating, well beyond what its teeth were capable of. Cerda (2008) made a strong case for the interpretation of these rock clusters as gastroliths, even going so far as to demonstrate that the ratio between the mass of the stomach stones in a Gasparinisaura individual, and the approximate mass of that individual, was comparable to that found in modern birds. The most recent paper on Gasparinisaura was published in 2012 by Ignacio Cerda and Anusuya Chinsamy. In this work, they focused on the histology of Gasparinisaura: its internal bone structure 37

(Cerda & Chinsamy, 2012). What they found, from their substantial sampling, was that Gasparinisaura grew and lived fast and — if the specimens known to date are anything to go by — often died young. Based on their observations, Gasparinisaura grew relatively rapidly until it reached ~60% of the size of the largest individual preserved. This growth did not always take place at a continuous rate, with some individuals showing evidence of brief periods of stasis. However, only after Gasparinisaura exceeded this size did growth slow down more dramatically: evidence of slower, periodically arrested growth was only seen in larger individuals. Nevertheless, it was clear that even the largest preserved individuals were still growing at their time of death. This suggested two possibilities: that Gasparinisaura’s growth was indeterminate — i.e. it never stopped growing until it died — or that no adult Gasparinisaura individuals were known. Cerda and Chinsamy (2012) suggested that the latter was more likely, and I agree. In recent years, several papers have been published on the internal bone structure of the ornithopod dinosaurs from Victoria (Chinsamy et al., 1998; Woodward et al., 2011, 2018). It will be interesting to compare and contrast the results of these studies against those obtained by Cerda and Chinsamy (2012), and also to conduct similar studies on other Australian ornithopod dinosaurs to determine how similar or different their internal bone structure is, and the implications of that.

In situ tail and pelvis of Gasparinisaura cincosaltensis. Isolated specimens show that the tail was encased in ossified tendons

Back foot of Gasparinisaura cincosaltensis. Note the curling of the toes, as in the more complete skeleton with gastroliths

38

Stop 8: Museo Paleontológico Egidio Feruglio (Trelew, Chubut): 26–30/11/2018

Me with the 2.4 metre long femur of Patagotitan mayorum The mezzanine, with bones of Patagotitan in the background Unlike almost every other institution that I visited as part of my Churchill Fellowship, the Museo Palaeontológico Egidio Feruglio in Trelew does not house any ornithopod material. However, it does hold all of the known remains of the titanosaurian sauropod Patagotitan mayorum, one of the largest dinosaurs ever discovered (Carballido et al., 2017). The significance of this titanosaur is such that it was the focus of a 2016 BBC documentary entitled Attenborough And The Giant Dinosaur, released shortly before Patagotitan was named. By coincidence, Dr Philip Mannion — a colleague from University College London, with whom I had travelled to Argentina twice before — was in Trelew during my stay. We independently studied the remains of Patagotitan, then discussed our interpretations, enabling more robust Reconstructed hind limb of Patagotitan interpretations of vague anatomical features to be made than would have been possible otherwise. My research on Patagotitan resulted in hundreds of photographs and a Word document (replete with measurements) almost twenty pages long. The anatomical information I collected on this trip will be hugely helpful to my ongoing research on Australian Cretaceous sauropods, but since the primary focus of the Churchill Fellowship was ornithopod dinosaurs, I will not detail that further here. Suffice it to say — our sauropods are quite different! 39

Stop 9: Universidad Nacional de la Patagonia, San Juan Bosco (Comodoro Rivadavia, Chubut): 3– 7/12/2018

Me with a megaraptoran theropod claw in the University lab Photographing the Notohypsilophodon femur in Comodoro The second last stop on my Argentinian trip was Comodoro Rivadavia, a rather bleak city on the southeast coast of Chubut Province. Specifically, I travelled here to visit the Geology Department of the Universidad Nacional de la Patagonia, San Juan Bosco, which houses an impressive dinosaur collection. Since the 1980s, Rubén Martínez, Marcelo Luna and Gabriel Casal have been conducting fieldwork throughout southern

Chubut Province, and their efforts have reaped Reconstructed Notohypsilophodon skeleton at top left rich rewards. On this trip I was focused on ornithopod dinosaurs, and there were two specimens that I wanted to study. One of these, Notohypsilophodon comodorensis (Cenomanian Bajo Barreal Formation), had been known to science for two decades (Martínez, 1998; Ibiricu et al., 2014). The was partially described in 2010 (Ibiricu et al., 2010), but new excavations at the site (Coniacian–Maastrichtian Lago Colhué With Marcelo Luna, fossil hunter extraordinaire — one of Huapi Formation) led to the recovery of more my hosts in the Geology Department in Comodoro Rivadavia bones. On Valentine’s Day 2019, it was named Sektensaurus sanjuanboscoi (Ibiricu et al., 2019). 40

Notohypsilophodon comodorensis Martínez, 1998

The type specimen of Notohypsilophodon comodorensis in left side view, assembled from my photos of the original bones Notohypsilophodon comodorensis is known only from a single specimen (UNPSJB-Pv 942: Martínez, 1998; Cambiaso, 2007; Ibiricu et al., 2014). However, it is significant because it is the oldest partial ornithopod skeleton known from South America, and is therefore the closest one in time to the known Australian ornithopods. During my first two days in the Department of Geology at Universidad Nacional de la Patagonia, San Juan Bosco in Comodoro Rivadavia (3–4/12/2018) I photographed and measured all of the bones of Notohypsilophodon. I also made notes on all of the bones in order to test claims made in the two papers published to date on this specimen (Martínez, 1998; Ibiricu et al., 2014). Somewhere along the line, I had forgotten that Cambiaso (2007) redescribed Notohypsilophodon in her unpublished thesis. Consequently, on my first day in Comodoro, I referred exclusively to the description of Notohypsilophodon presented by Ibiricu et al. (2014) as I studied the specimen myself. As I did so, I began to doubt the veracity of several of the unique features of Notohypsilophodon identified by Ibiricu et al. (2014). This was because many of these features appeared to be based on surfaces that were clearly broken. As an example, in describing the scapula (shoulder blade), Ibiricu et al. (2014) stated that the glenoid fossa (for the humerus) was concave. However, personal observation 41

of the specimen showed that the glenoid fossa is incomplete on its medial margin, meaning that it was not necessarily concave in life. Ibiricu et al. (2014) also described the coracoids as if the dorsal margins were complete (as far as I can tell, they aren’t), and the femur as if it was uncrushed and preserved complete distal condyles (when in my opinion it isn’t, and it doesn’t). Furthermore, they described only one of the preserved fibulae: the one they focused on is more complete than the other, but is also more damaged. Consequently, some of the unique features Ibiricu et al. (2014) identified on this bone might not be unique, since they cannot be observed on the less damaged, albeit less complete, fibula. My first day working on Notohypsilophodon happened to be ‘leg day’, meaning that part of my time was spent studying its back foot. Unfortunately, Ibiricu et al. (2014) did not describe the bones of the foot in detail, and did not work out which toes were represented. I determined that parts of both feet were present, and that neither was complete. I also worked out that one of the “toe” bones was actually a metacarpal from the hand. Back at my hotel that night, I flicked through Cambiaso’s (2007) unpublished thesis — and found The right back foot of Notohypsilophodon, with the positions of the pedal her thorough redescription of phalanges based on personal observations [and Cambiaso (2007)] Notohypsilophodon! After loosely translating her anatomical descriptions and figure captions, I compared her interpretations against mine — and was delighted to realise that we were in agreement on virtually every point, including the identification of the metacarpal. In retrospect, had I not seen the articulated feet of Anabisetia and Gasparinisaura beforehand, I might not have been able to correctly place the pedal phalanges of Notohypsilophodon. I suspect that Cambiaso took advantage of having done the same, since she also briefly described some specimens of Anabisetia in her thesis as well (Cambiaso, 2007). The long and the short of it: based on my personal observations, the unpublished description of Notohypsilophodon produced by Cambiaso (2007) is more detailed than the one published by Ibiricu et al. (2014). Furthermore, in terms of its age, morphology and size, I consider Notohypsilophodon to be the South American ornithopod most likely to share a close relationship with Australia’s ornithopods. However, the anatomical incompleteness of the only known specimen, and its subadult status (shown by the lack of fusion between the scapula and coracoid, and the incomplete coalescence of the sacral vertebrae), mean that it is going to be difficult to resolve its position on the ornithopod family tree with certainty. Nevertheless, by personally observing the specimen I feel that I have now developed a much better understanding of the features that make Notohypsilophodon unique, and those that demonstrate its relationships to other ornithopods within South America and worldwide. 42

Sektensaurus sanjuanboscoi Ibiricu, Casal, Martínez, Luna, Canale, Álvarez & González Riga, 2019

The type specimen of Sektensaurus sanjuanboscoi in left side view, assembled from my photos of the original bones

In 1993, several incomplete bones representing a fairly large ornithopod were collected from an island in Colhue Huapi Lake. A decade later, a left ilium (registered as UNPSJB-Pv 973) was briefly described in an abstract (Luna et al., 2003). Seven years after that, more of the material (registered as UNPSJB- Pv 960) was briefly described but not named (Ibiricu et al., 2010). Then, in 2017, Colhue Huapi Lake receded, enabling researchers at UNPSJB to ride motorbikes and all-terrain vehicles to the island in order to collect more fossils. They succeeded. On 14 February 2019, eight weeks after I returned from Argentina, all of these bones, which were interpreted as belonging to a single ornithopod individual, were made the type specimen of Sektensaurus sanjuanboscoi (Ibiricu et al., 2019). However, during my Fellowship, I was generously given permission to work on the then unpublished material. While working on Sektensaurus in Comodoro Rivadavia, I did not have access to the now published description by Ibiricu et al. (2019). As such, I essentially had to describe Sektensaurus as if no-one else had. I was quite glad that I was only attempting this after studying much more complete ornithopods like Anabisetia, Gasparinisaura and Macrogryphosaurus; had I not, I doubt I would have been able to interpret Sektensaurus particularly well at all. Now that the published description is out (Ibiricu et al., 2019), I have been able to compare my notes with it, and I have found several points of difference. Ibiricu et al. (2019) described one incomplete element as a left tibia; however, I worked out that this bone was in fact right metatarsal III, and that another incomplete bone (which was not described) was right metatarsal IV and articulated with it perfectly. Other foot bones were also preserved, but also were not described. Another incomplete bone, interpreted as a right radius, is in fact part of the right ischium, whereas the distal end of a humerus and the proximal end of a fibula — interpreted by Ibiricu et al. (2019) as being from the right side — are both from the left. Most importantly, perhaps, the bone interpreted as a frontal was, to my eyes, more likely a fragment of turtle carapace or plastron. 43

Stop 10: Museo Regional Provincial Padre Manuel Jesús Molina (Río Gallegos, Santa Cruz): 11– 14/12/2018 The last stop on my Argentina trip was the Museo Regional Provincial Padre Manuel Jesús Molina in Río Gallegos, not far from the southern tip of South America. Prior to my departure from Australia, I checked what December weather was like in Río Gallegos, and many sources said very windy. It certainly lived up to the billing! I was glad I brought a jumper. The museum is situated on the corner of one of the main streets in Río Gallegos. Unfortunately, the street-side façade is riddled with graffiti. Several extremely significant dinosaur fossils, like the large sauropods Drusilasaura, Dreadnoughtus and Puertasaurus, and the theropods Austrocheirus and Orkoraptor are held here. Crucially, so is the only known skeleton of the ornithopod Talenkauen — one of the specimens I was most looking forward to studying on my Churchill Fellowship. All of these dinosaur specimens were found in Santa Cruz province, shipped to Buenos Aires for study, and then returned to Río Gallegos. Many of the specimens remain in shipping crates, including virtually all of the remains of Talenkauen. The only exceptions were a few bones from the pectoral girdle and forelimb, which were on display. My interactions with the museum staff indicated that they did not realise just how amazing the Talenkauen specimen was. As I conducted my research, they realised that the entire vertebral column was preserved and would make a far more impressive display piece than the few bones situated in their galleries. As such, after my visit they completely changed their Talenkauen display. The staff publicised my visit on their Facebook page, and used it to highlight the significance of the specimens held Talenkauen at Museo Regional Provincial Padre Manuel Jésus Molina. Although ‘fossiled out’ by this stage, I enjoyed this! in their palaeontological department. 44

Talenkauen santacrucensis Novas, Cambiaso & Ambrosio, 2004

The type specimen of Talenkauen santacrucensis in left side view, assembled from my photos of the original bones

One of the dinosaurs that I was most looking forward to studying on my Churchill Fellowship was the last one I studied — the type specimen of Talenkauen santacrucensis. This magnificent skeleton (MPM-PV 10001A) was originally thought to date to the Maastrichtian (Novas et al., 2004; Cambiaso, 2007), the last stage of the Cretaceous — and the last stage of the non-avian dinosaur age. However, the rocks from which Talenkauen was extracted are now thought to be Cenomanian in age (Varela et al., 2012), meaning that they are in fact from the mid-Cretaceous (~101.5–95 Ma). Only one scientific paper beyond the original, brief description by Novas et al. (2004) has focused on Talenkauen, and that revealed that teeth from a hatchling or juvenile were preserved with the type specimen (MPM‐PV 10001B: Egerton et al., 2013). Although the discovery of Talenkauen was announced in 2004, no comprehensive description of this ornithopod has ever been published. An entire chapter of Andrea Cambiaso’s (2007) thesis was dedicated to such a description, but this was never published and is at present only available in Spanish. Consequently, Talenkauen has remained rather enigmatic, despite its skeletal completeness, its potential phylogenetic proximity to Australian (and other South American) ornithopods, and its similar geological age. I had known before travelling to Rio Gallegos that Talenkauen was fairly large (~5 metres long) as far as “small” ornithopods go, and that the specimen was quite complete. Nevertheless, I was completely unprepared for how well preserved many of the bones were — and how poorly preserved were others. The few preserved skull bones and the neck vertebrae were solid and beautiful, whereas the bones of the hind limb and pelvis were quite fragile. 45

In order to properly study Talenkauen in the week I had available, I (loosely) translated the description of Talenkauen from Andrea Cambiaso’s (2007) thesis, then studied the bones firsthand. I photographed almost every single specimen, and took measurements of everything too. The skull and limb bones were particularly informative from an anatomical perspective. However, the vertebrae were perhaps the most instructive. Ossified tendons were present along the dorsal and sacral neural spines, and the ribs were braced against each other by large, dish-like uncinate processes. Presumably, all of these Reconstructed Talenkauen skeleton on display at the Museo features increased the rigidity of the Argentino de Ciencias Naturales in Buenos Aires. thorax. Why this was necessary is unclear. There were several things I noticed about Talenkauen that did not seem to match up with Cambiaso’s (2007) description. One was her interpretation that Talenkauen had sixteen thoracic vertebrae. This seemed like a very high number, especially in light of my observations of Macrogryphosaurus at Lago Barreales. I carefully observed the vertebrae of Talenkauen and wrote the following in my notes: “I contend that the vertebra interpreted herein as dorsal vertebra 16 is in fact sacral 1, based on comparisons with Macrogryphosaurus. In addition, the dorsal vertebra labelled as “6” (which is articulated with “7”–“10”) connects to that labelled “5” (which is articulated with “4” and “3”). As such, it seems that Talenkauen, like Macrogryphosaurus, had 14 — not 16 — dorsal vertebrae” This is an important observation, because it means that Macrogryphosaurus and Talenkauen might be even more closely related than previously thought. In addition, I made the following observations: “I completely agree with Cambiaso’s (2007) interpretation of the right prepubis, based on firsthand observation of Macrogryphosaurus. The left pubis in that taxon preserves the prepubic process, and it is nigh on identical to this one — even to the point where there is very clearly a raised convexity on the lateral side. “Two elements identified as distal fibulae are in fact the distal ends of the pubes. They look just like those of Macrogryphosaurus, they are mirror images of one another, they articulate with each other on a flat, longitudinally-striated surface, and they are ventrally hooked. They do not look like the distal fibulae of any ornithopods that I have observed to date.” Thus, Talenkauen only preserves the proximal end of one fibula; the “distal” fibulae are in fact part of the pubes. In fairness to Cambiaso, who (as far as I am aware) was not able to study Macrogryphosaurus in the course of her research, it was only because I had carefully scrutinised the pelvic elements of 46

Macrogryphosaurus that I was able to recognise these elements for what they truly were. Had I not done so, I doubt I would have been able to work this out. Based on my observations, it would seem Cambiaso also erred in her assessment of Talenkauen’s metatarsals: “The preserved portions of II and III are more distal than she [Cambiaso] indicated, and they actually key into each other perfectly demonstrating that the preserved portion of II extends further proximally than that of III. IV preserves the proximal end (complete with a concavity as in Sektensaurus!), and appears to just overlap with III in terms of the preserved portion of its proximal medial articular surface.” The fact that this feature was shared with Sektensaurus suggests a close relationship between it and Talenkauen; curiously, in their paper describing Sektensaurus that was published six weeks after my return, Ibiricu et al. (2019) only ran phylogenetic analyses that did not Talenkauen left metatarsals in correct articulation include Talenkauen or Macrogryphosaurus. This means that they did not test the possibility of a close phylogenetic relationship between Sektensaurus and Talenkauen, despite discussing anatomical similarities and differences between the two taxa in their discussion, and despite the fact that they tentatively identified Sektensaurus as an elasmarian — the group which contains Talenkauen and Macrogryphosaurus. I also reassessed the bones of the back foot, identifying one toe bone (phalanx) as a claw (ungual): “I am quite confident that the element identified by Cambiaso (2007) as the proximal end of pedal phalanx I-1 is in fact the proximal end of an ungual, specifically pedal phalanx IV-5. This would suggest that the other phalanges from this pedal digit are IV-2 and IV- 3, since the ungual is far too small to articulate with the distal end of the more distal of the two.” I also studied some bones of Talenkauen that Cambiaso (2007) either missed or did not have access to. Primary among these was a bone from the tail called a chevron, which showed two interesting features: a proximally bridged haemal canal, and a blade with a subtle posterior flange. The chevrons of Macrogryphosaurus are similar. By the end of my week in Río Gallegos, I had a more comprehensive database on Talenkauen than had ever been published. This will be exceedingly useful for my The left back foot of Talenkauen in life position. future research on Australian ornithopod dinosaurs. Note the robustness of the toe bones and claws 47

Conclusions and recommendations

Over the course of my eight weeks in Argentina, I was able to study all (then) named non-hadrosaurid ornithopod taxa from South America and Antarctica (sadly, I missed Mahuidacursor, which was only named after my return). Had this been the only result of the Fellowship, I would have considered it an overwhelming success. Because of my Churchill Fellowship, I now:

• have a far better understanding of South American and Antarctic ornithopods; • understand which anatomical characters separate the various species; • appreciate how similar and how different each species is to all of the others; and • appreciate their scale, which is often hard to appreciate from a table of measurements.

However, this trip provided me with so much more than just an understanding of Gondwanan ornithopods. It enabled me to meet several of the researchers responsible for excavating, curating and describing these specimens, which I hope (and expect) will lead to collaborations in the future. In addition, I was able to study several titanosaurian sauropod and megaraptorid theropod specimens. This will greatly enhance my ongoing research on both of those groups.

Ornithopods

Having now studied seven of the eight named species of ornithopod from Argentina, it is clear to me that the type specimens of the Argentinian taxa are far better than most of the Australian ones, both in terms of quantity of bones preserved their and quality of preservation. As a result, it has been — and will continue to be — easier to place the South American species on the ornithopod family tree. Further study on the Australian taxa might enable more robust family tree placements, but what will enable to us to understand their positions thereon even better is the discovery of new, more complete specimens — like ‘Noddy’ from Victoria. Future exploration and excavation in known and new sites across Victoria, New South Wales and Queensland will improve this situation, and this is something that anyone hoping to understand Gondwanan ornithopod evolution should strive to achieve.

One thing that struck me during my travels was how similar several of Argentina’s ornithopods were to one another. Those that are classified within Elasmaria — Talenkauen, Macrogryphosaurus, and Sektensaurus — were particularly similar, and the establishment of that group for those genera seems to be fully justified. What is now of great interest to me is exactly where and for how long (geologically speaking) elasmarians lived. Talenkauen and Macrogryphosaurus are now thought to be very similar in age to each other (~100.5–93.9 million years old), making them approximately the same age as much of the rock unit known as the Winton Formation, which is the focus of annual digs by the Australian Age of Dinosaurs Museum in Winton. By contrast, Sektensaurus’ rather poorly constrained age range (~86.3–66 million years ago) does not overlap with that of either Talenkauen or Macrogryphosaurus. If Sektensaurus truly does date to somewhere within this presently understood age range, then elasmarians were rather tenacious. So, should we expect to find elasmarians in Australia? I think so, unless they were so specialised they could only eat South American plants! 48

With respect to the dietary preferences and possible environmental constraints of ornithopods, one thing really surprised me about the species I studied: very few Argentinian ornithopods are associated with abundant and diverse fossil floras. Consequently, their palaeoenvironmental surroundings are often not well understood (except Talenkauen, if it is indeed from the Mata Amarilla Formation as is now thought). This is in stark contrast to the situation in Australia, where the palaeoenvironmental settings inhabited by our dinosaurs are often better understood than the dinosaurs themselves!

In both Australia and Argentina, there appear to be significant issues with chronostratigraphic resolution. What this means is that, as presently constrained, the geological age ranges of many ornithopod taxa across Gondwana are very broad. Even if we were able to place all of the South American, Antarctic and Australian species on the ornithopod family tree (something I am working towards), any attempts to pin down the timing of origins, evolutionary radiations and extinctions are undermined when the correct age of a species is either unknown, or so broad that it is rendered essentially uninformative. Australian palaeontologists are certainly working towards overcoming this problem, and the Antarctic species seem to be quite accurately constrained in geological time. However, from what I can tell much, work remains to be done in this regard for the Argentinian taxa.

Ankylosaurs, megaraptorid theropods and titanosaurian sauropods

Having not yet studied the Australian ankylosaurs Minmi or Kunbarrasaurus in detail, it was difficult for me to determine Antarctopelta’s level of similarity or difference to them. However, having studied the Antarctopelta specimen, I now feel much better equipped to supervise a student who will work to describe a newly discovered ankylosaur vertebra (with osteoderm [armour plate]) from Victoria.

In recent years I have been involved in much research on Australovenator, Australia’s best represented megaraptorid theropod. Having now studied several Megaraptor specimens in Argentina, I was struck by its anatomical similarity to Australovenator. The only major difference was Megaraptor’s greater size (although there were several subtle anatomical differences, of course). I am confident that the two are closely related, and plan to use the data collected to improve understanding of Australovenator’s position within Megaraptoridae and in the description of new megaraptorid material from Victoria.

The sauropods I studied provided me with vast quantities of useful anatomical information. Chubutisaurus was particularly instructive, since it seems to be very similar to Diamantinasaurus and Savannasaurus. Studying Patagotitan and Futalognkosaurus was similarly informative, particularly in understanding how these beasts were able to support their vast bulks. I plan to incorporate the information derived from these specimens in my ongoing research on Australian sauropods.

Recommendations

To the ends outlined above, I would like to see more collaboration between Argentinian and Australian palaeontologists. My Churchill Fellowship has definitely opened the door to such collaborations for me personally, and I fully intend to strengthen those relationships. Many Australian–Argentinian collaborations have occurred in the last few decades, and I would hope that many will in future. 49

I had hoped that many of my hosts, and other palaeontologists that I met throughout my Fellowship, would be attending the 79th Annual Society of Vertebrate Paleontology Conference (the biggest conference in vertebrate palaeontology), which is being held later this year in Brisbane. This will be the first time the meeting has been held in the Southern Hemisphere, it will provide many attendees with the opportunity to visit Australian vertebrate palaeontology collections, and it would seem to be the perfect gateway to collaborations between Argentinian and Australian palaeontologists. However, when I asked the Argentinian palaeontologists hosting me in each institution whether or not they would be attending, almost all said no. The reason? A lack of funding. Given that Argentina’s economy had collapsed shortly before my arrival, I must admit that I did not find this that surprising. However, even had the economy been in better shape, I would still not have found it surprising, since Australian palaeontologists often find it difficult to attend overseas conferences themselves for the same reason.

Perhaps a solution would be to have virtual scientific meetings that enable South American and Australian researchers — and those from other continents or countries that were formerly part of the Gondwanan supercontinent, like Africa, Madagascar and India — to overcome the tyranny of distance between these now widely-separated landmasses? My experience with internet connectivity throughout Argentina gave me the sense that researchers there would have no trouble participating in a video conference. The language of science is English, which is great for Australian palaeontologists but not so for some of our Argentinian colleagues; many of my hosts’ English was exemplary, whereas others relished in the opportunity to practise. Virtual conferences would give Argentinian palaeontologists the chance to speak English in a congenial setting, to find like-minded colleagues who could read over manuscripts and improve the quality of their English, and to discuss the current state- of-play in Argentinian palaeontology while learning the same of Australian palaeontology.

The study of past life helps us understand how our planet has changed through time and places modern floras and faunas (and humanity) in an appropriate context. Critically, palaeontology is often the spark that ignites an interest in science in children. I saw this in every Argentinian museum I visited: schoolchildren of all ages gazed in wonder at skeletons of dinosaurs and fossil mammals, chatting excitedly with their classmates about our planet’s past denizens. Those Argentinian schoolchildren — and the children of Australia — include among their number the majority of the Southern Hemisphere’s future scientists. By instilling an appreciation for nature, for humanity’s place within the natural world, for the scale and mechanics of past environmental and evolutionary changes, and for the real risk of extinction for all species (including us), we will give them a chance to better look after their home than did previous generations. The quality of the Earth these children inherit, and their capacity to stabilise or improve it, will be impacted by scientific understanding and technology. Therefore, it is essential that as much scientific interest as possible is sparked and nurtured in children and adults alike.

To this end, palaeontology has a huge part to play. However, in order to recognise this potential, palaeontology will need more financial support. The employment of more palaeontologists in museums and universities across Australia would go some way to achieving this. 50

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Tools of the trade: tape measure, dinosaur bones, computer and brains (not pictured). Studying Sektensaurus in Comodoro

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The lengths a homesick Churchill Fellow will go to for phone reception... (photos Juan Mansilla)