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Journal of Vertebrate Paleontology

ISSN: 0272-4634 (Print) 1937-2809 (Online) Journal homepage: http://www.tandfonline.com/loi/ujvp20

First () from Early () of Oregon, U.S.A.

Gregory J. Retallack, Jessica M. Theodor, Edward B. Davis, Samantha S. B. Hopkins & Paul Z. Barrett

To cite this article: Gregory J. Retallack, Jessica M. Theodor, Edward B. Davis, Samantha S. B. Hopkins & Paul Z. Barrett (2018): First dinosaur (Ornithopoda) from (Albian) of Oregon, U.S.A., Journal of Vertebrate Paleontology, DOI: 10.1080/02724634.2018.1486847 To link to this article: https://doi.org/10.1080/02724634.2018.1486847

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Published online: 15 Oct 2018.

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Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=ujvp20 Journal of Vertebrate Paleontology e1486847 (5 pages) # by the Society of Vertebrate Paleontology DOI: 10.1080/02724634.2018.1486847

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FIRST DINOSAUR (ORNITHOPODA) FROM EARLY CRETACEOUS (ALBIAN) OF OREGON, U.S.A.

GREGORY J. RETALLACK,*,1 JESSICA M. THEODOR,2 EDWARD B. DAVIS,1 SAMANTHA S. B. HOPKINS,1 and PAUL Z. BARRETT1 1Department of Earth Sciences, University of Oregon, 1275 E. 13th Avenue, Eugene, Oregon 97403, U.S.A, [email protected]; 2Department of Biological Sciences, University of Calgary, 2500 University Drive NW, Calgary, Alberta, T2N 1N4, Canada

SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Citation for this article: Retallack, G. J., J. M. Theodor, E. B. Davis, S. S. B. Hopkins, and P. Z. Barrett. 2018. First dino- saur (Ornithopoda) from Early Cretaceous (Albian) of Oregon, U.S.A. Journal of Vertebrate Paleontology. DOI: 10.1080/ 02724634.2018.1486847.

A bone discovered in 2015 by GJR is the first diag- (Wembley, Alberta), Museum of the Rockies (Bozeman, nostic nonavian dinosaur fossil from Oregon, a state whose ), and State Museum (Salt Lake City, Utah) Mesozoic rocks are mostly marine (Orr and Orr, 2012). This dis- were used to guide literature search for taxonomic placement covery is novel evidence of Cretaceous terrestrial environments of the Mitchell phalanx. A photogrammetric three-dimensional and faunas in Oregon. The pedal phalanx was recovered from the (3D) image of the bone as a PDF using Agisoft PhotoScan is mid–Cretaceous (Albian–Cenomanian) Hudspeth Shale and is available as Supplementary Data 1. here identified informally as the ‘Mitchell ornithopod.’ This new ornithopod fills a geographic gap between the Early Cretaceous ‘ ’ () Ono ornithopod of northern California (Hilton et al., SYSTEMATIC PALEONTOLOGY 1997) and the () ‘Sucia Island theropod’ of Washington (Peecook and Sidor, 2015), as one of DINOSAURIA Owen, 1842 the few nonavian dinosaur specimens known from marine rocks Seeley, 1887 of the Pacific Northwest. Other Pacific coastal nonavian from southeast Alaska to Baja California reviewed by Peecock ORNITHOPODA Marsh, 1881 and Sidor (2015) include a possible Oregon hadrosaur sacrum gen et sp. indet. from the Late Cretaceous (Maastrichtian) Cape Sebastian Sandstone (Orr and Orr, 2012), but our examination of this unprepared specimen cannot yet determine whether it was a dino- Referred Material—UOF118600, a pedal phalanx. saur, marine , or plesiosaur. Age and Occurrence—The locality, UO13418 Mitchell, is a Institutional Abbreviations—BM, British Museum, now gentle shale slope below low cliffs of lowest Hudspeth Formation Natural History Museum, London, U.K.; IRB, Institut Royal on public lands administered by the Bureau of Land Management   des Sciences Naturelles de Belgique, Brussels, Belgium; JRF, near Mitchell, Oregon, at roughly 44.6 N, 120.2 W.Thelocalityis Judith River Foundation, Malta, Montana, U.S.A.; MOR, in the Hudspeth Shale at a stratigraphic level of 374.9 m in the Museum of the Rockies, Bozeman, Montana, U.S.A.; UMM, nearby measured section (Fig. 1) of Wilkinson and Oles (1968). University of Manchester Museum, Manchester, U.K.; UO, The biostratigraphic age is late early Albian, in the Brewericeras Condon Collection, University of Oregon Museum of Natural hulenense zone (Jones et al., 1965). Associated include the History, Eugene, Oregon, U.S.A. ammonites Brewericeras hulenense (UOF118598, UOF118606), Leconteites lecontei (UOF118597), Cleoniceras jonesi (UOF118596A–B), Anisoceras merriami (UOF118599), snail METHODS Amberleya dilleri (UOF118601), bivalves Lithophaga carolinensis (UOF118602), Entolium utukokense (UOF118603), Aucellina A single bone regarded as a dinosaur phalanx (UOF118600) dowlingi (UOF118604), an ichthyosaur centrum was collected from near Mitchell, Oregon (locality UO13418), (UOF118595A–B), and wood (UOF118605). Many of along with an ichthyosaur centrum and marine invertebrates – these ammonites are endemic North American , but (UOF11895 F118606) and cataloged into the Condon cosmopolitan ammonites also are found near the Mitchell locality, Collection of the Museum of Natural and Cultural History of which is stratigraphically above the local first appearance of the University of Oregon (paleo.uoregon.edu). Examination of Mortoniceras inflatum and below the local first appearance of specimens from the Museum of the North (Fairbanks, Alaska), Mortoniceras fallax,or102.5–103.0 Ma (Lehmann, 2015). Precise Royal Tyrrell Museum (Drumheller, Alberta), Currie Museum coordinates and further locality details are on file in the Prineville Office of the Bureau of Land Management and the Museum of *Corresponding author. Natural and Cultural History of the University of Oregon.

Published online 15 Oct 2018 Retallack et al.—Cretaceous ornithopod from Oregon (e1486847-2)

FIGURE 1. Stratigraphic location of the Mitchell ornithopod within a reconstructed geological section of the Hudspeth Formation (from Wilkinson and Oles, 1968), showing local ammonite zones (Jones et al., 1965) and first appearances of important ammonites, which are available in the Condon Collection of the Museum of Natural and Cultural History, University of Oregon (paleo.uoregon.edu).

Description—The putative dinosaur phalanx (Fig. 2) is wider phalanx with collateral pits is illustrated here in Figure 2J–L. mediolaterally (47.6 mm) than proximodistally (30.3 mm), with Generally, similar short phalanges with lateral rimmed basins like a moderately concave proximal end. The distal end has a the Mitchell bone are found in (Norman, 1980:figs. weakly saddle-shaped, convex condyle curved anteriorly 13 71–72), (Brown, 1916:fig. 14), from vertical. A mediolateral sulcus on the dorsal surface is (Zheng et al., 2011:figs. 1–3, 5–6), (Brown, continuous to the lateral surfaces, with a shallow basal ridge 1913:fig. 7), (Fig. 2G–I; Prieto-Marquez, dividing it from a short plantar sulcus below. This shallow, 2007:fig. 6.7), and (Maryanska and Osmolska, rimmed, lateral basin is better preserved in other dinosaur pha- 1984:pl. 23, figs. 11a–b). The Mitchell ornithopod (Fig. 2A–F)is langes (Fig. 2G) than the worn and broken Mitchell specimens regarded as a third pedal phalanx of the central digit (III) of the (Fig. 2C, F) and is in the position where deep collateral pits right pes of an iguanodont or hadrosaur dinosaur, by comparison are seen on many other dinosaur phalanges (Fig. 2 L, O). A with these other ornithopods (Fig. 2I; Brown, 1913, 1916;Norman, basal rim to the lateral basins also distinguishes the Mitchell 1980;MaryanskaandOsmolska, 1984; Zheng et al., 2011). phalanx from dinosaur phalanges with a girdling mediolateral Ornithopods are common in Early Cretaceous rocks of sulcus (Fig. 2M–O). Utah and , but at 102.5–103 Ma (late Albian); our phal- The maximum distal width is 47.6 mm, and the maximum anx falls within a gap in the North American dinosaur record proximal width is at least 46.6 mm. Exact measurement is com- between the Aptian–Albian and the Cenomanian promised by postmortem erosion of the cortex, especially (McDonald et al., 2012). where the museum number has been placed (Fig. 2). Taxonomic and Anatomical Position—The lateral rimmed basin of the dorsal sulcus is distinct from a completely girdling sul- SIZE ESTIMATE OF WHOLE cus or waist in other large dinosaur phalanges that are shorter than wide in sauropods (Bonnan, 2005:figs. 16.8–16.9; D’Emic et al., The Mitchell ornithopod phalanx is robust, flattened, and 2011:figs. 1–2), stegosaurs (Maidment et al., 2015:figs. 76–77), wide, thus being a load-bearing subunguligrade toe bone from nodosaurs (Carpenter et al., 1995:fig. 16), ankylosaurs (Carpenter, an animal of substantial size (Moreno et al., 2007). Although 1984:fig. 5; Currie et al., 2011:figs. 9–10), and ceratopsians (Brown, mass cannot be precisely reconstructed without more complete 1914:fig. 19; Chinnery and Horner, 2007:fig. 4P). A ceratopsian material, comparison with six Cretaceous ornithopods known phalanx with girdling sulcus is illustrated here in Figure 2M–O. from complete skeletons (Hulke, 1882: BM39460; Norman, The shallow end to the dorsal sulcus is distinct from collateral pits 1980: IRB1534, 1536, 1561; Seebacher, 2001: BM5764, and a narrow waist, found in phalanges of theropods (White et al., IRB1535; Tennant, 2013: UMM12275; Tweet et al., 2008: 2016:fig. 2), pachycephalosaurs (Perle et al., 1982:pl.44, figs. 4–7; JRF115) gives a general idea. The Mitchell ornithopod was Sues and Galton, 1987:text-fig. 14), and small ornithopods such as larger than Tenontosaurus tilleti and smaller than Iguanodon (Hulke, 1882:pl.79,fig.4;Galton,1971:fig. 1A) bernissartensis, but the Ono ornithopod was a much smaller and Tenontosaurus (Tennant, 2013:fig. 31). An ornithopod animal, closer in size to Hypsilophodon foxi (Hilton et al., Retallack et al.—Cretaceous ornithopod from Oregon (e1486847-3)

FIGURE 2. Pedal phalanx of Mitchell ornithopod, in standard views (A–F); comparable phalanx of Brachylophosaurus canadensis (G–H, arrow in I; MOR1070-7-31-99-281E); and distinct phalanges with collateral pits and girdling sulci of Tenontosaurus tilletti (J–L; MOR787) and Einiosaurus procurvi- cornis (M–O; MOR956-32-33), respectively. In all images, plantar surface is down, dorsal is up, proximal is left, and distal is right. Scale bars equal 10 mm (A–H, J–O) and 50 mm (I). (A 3D digital image of the Mitchell ornithopod is also available as Supplementary Data 1.)

1997). Closest in size to the Mitchell ornithopod is a phalanx TAPHONOMY (MOR1071B-7-99460.0) 47.3 mm wide on distal plantar edge – The Mitchell ornithopod was found with a diverse assem- and 20.6 mm long on left plantar edge (Fig. 2G I) of the Late blage of marine molluscs and ichthyosaur centrum in a dark Cretaceous hadrosaur Brachylophosaurus canadensis (Prieto- gray shale with siderite nodules. The Hudspeth Shale also Marquez, 2007), best known from the ca. 5.1 m long subadult ‘ ’ has yielded a variety of echinoids, starfish, crabs, shrimp, fish mummy Leonardo (Murphy et al., 2007; Tweet et al., 2008). bones, shark teeth, a plesiosaur, and the pterosaur, Iguanodon Body mass of a similarly proportioned ornithopod, ‘Bennettazhia’ oregonensis (Welton, 1972; Vega et al., 2010; mantelli , 5.1 m long, has been estimated at 678 kg by Averianov, 2012; Orr and Orr, 2012). The nodular dark Seebacher (2001). shales of outer marine continental shelf environments are interbedded with conglomerates of alluvial plains, fan deltas, shingle beaches, and submarine canyons of a rocky coast PALEOECOLOGY (Dorsey and Lenegan, 2007). As the only dinosaur bone Sparse plant material found in the Hudspeth Shale offers found in the Mitchell area in 150 of prospecting there insight into the terrestrial environment in which the Mitchell (Orr and Orr, 2012), the Mitchell ornithopod is a rare speci- ornithopod lived. Fossil conifer wood was found with the fossil men of a terrestrial animal, likely preserved by floating out and is common in the Hudspeth Formation (Wilkinson and to sea and disarticulated with decay. Similarly, the Ono orni- Oles, 1968). Other fossil plants from the Hudspeth Formation thopod from marine shales of California is a single lower in UO collections include conifer shoots of Sequoia condita limb with many missing (Hilton et al., 1997). Only a sin- (F118840) and Elatocladus montanensis (F118838, F118841, gle set of sauropod remains is known from the Early F120581), fragments of Nathorstia sp. cf. N. firma Posidonienschiefer of Holzmaden, southern Germany, which (F120569), horsetails Equisetites sp. cf. E. burchardtii (F120635, has been quarried for tiles and fossils since the 12th century F120661), and indeterminate or cycadeoid scales (Wild, 1978; Etter and Tang, 2002). Dinosaurs in marine (F120030A–B). No fossil angiosperms have yet been found, rocks are very rare, but the Mitchell ornithopod offers a glimpse into terrestrial environments poorly represented in and this flora predates the late Albian advent of convincing ’ angiosperm fossils in California (Retallack and Dilcher, 1986). Oregon s Mesozoic record. Paleomagnetic studies of the Hudspeth Shale reveal a paleo- latitude of 39.2 ± 4.8N (Housen and Dorsey, 2005), more like  ACKNOWLEDGMENTS modern Mendocino, California (39.3 N), than Mitchell, Oregon (45.6N). Vegetation in which the Mitchell ornithopod We thank E. W. Woodard and J. Zancanella of the Bureau lived may have been comparable to the modern coast redwood of Land Management Prineville Office for permissions and (Sequoia sempervirens) forests of northern California (Barbour help in the field. J. B. Scannella demonstrated phalanges in et al., 2007). the Museum of the Rockies. Useful advice also came from P. Retallack et al.—Cretaceous ornithopod from Oregon (e1486847-4)

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