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Evidence for Niche Partitioning Among Ground-Height Browsing Sauropods from the Upper Jurassic Morrison Formation of North America

Evidence for Niche Partitioning Among Ground-Height Browsing Sauropods from the Upper Jurassic Morrison Formation of North America

GEOLOGY OF THE INTERMOUNTAIN WEST an open-access journal of the Geological Association ISSN 2380-7601 Volume 5 2018

EVIDENCE FOR NICHE PARTITIONING AMONG GROUND-HEIGHT BROWSING SAUROPODS FROM THE UPPER OF NORTH AMERICA

Julia B. McHugh

Theme Issue An Ecosystem We Thought We Knew— The Emerging Complexities of the Morrison Formation SOCIETY OF Annual Meeting, October 26 – 29, 2016 Grand America Hotel Salt Lake City, Utah, USA

© 2018 Utah Geological Association. All rights reserved. For permission to copy and distribute, see the following page or visit the UGA website at www.utahgeology.org for information. Email inquiries to [email protected]. GEOLOGY OF THE INTERMOUNTAIN WEST an open-access journal of the Utah Geological Association ISSN 2380-7601

Volume 5 2018

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i GEOLOGY OF THE INTERMOUNTAIN WEST an open-access journal of the Utah Geological Association

Volume 5 2018

Evidence for Niche Partitioning Among Ground-Height Browsing Sauropods from the Upper Jurassic Morrison Formation of North America

Julia B. McHugh1 1Museums of Western Colorado, P.O. Box 20000, Grand Junction, CO 81502; [email protected]

ABSTRACT Two tooth-bearing snout fragments from a diplodocid sauropod from the Brushy Basin Member of the Morrison Formation (Upper Jurassic) excavated from the Mygatt-Moore Quarry in Rabbit Valley, Colora- do are described. The Mygatt-Moore Quarry has produced thousands of vertebrate from the Brushy Basin Member, with the diplodocid cf. louisae and the tetanuran fragilis dominat- ing the assemblage. Additionally, remains of another diplodocid, sp., have been found near the quarry within Rabbit Valley. Both specimens in this study preserve eight teeth per alveolar position, as observed through broken surfaces at the gross anatomical level and also through computed tomography (CT) scans. This is inconsistent with the genusDiplodocus sp., which has been previously shown to have a maximum of six teeth per alveolus. The presence of eight replacement teeth per alveolus has previously only been reported in the rebbachisaurid taqueti, which has been interpreted to have occupied a similar ground-height browsing feeding strategy to both Diplodocus and Apatosaurus. This is the first report of this of high-count replacement teeth in a diplodocid sauropod from the Morrison Formation. The high number of replacement teeth in a close relative to the contemporaneous Diplodocus provides evidence for niche partitioning among the contemporary ground-height browsing diplodocid sauropods of the Period in North America.

INTRODUCTION would have had a substantial impact on the available biomass in order to support populations of these two Apatosaurus and Diplodocus are contemporary sau- genera. Ecological niche partitioning in the Upper Ju- ropod genera found within the Upper Jurassic Morri- rassic Morrison Formation’s landscape has traditional- son Formation (Tschopp and others, 2015). These gen- ly been proposed to circumvent this problem (Turner era share a close phylogenetic history, as well as head and Peterson, 2004). However, empirical evidence sup- and neck morphologies that allow for similar biome- porting this partitioning among coeval genera, partic- chanical adaptions to available vegetation zones (Ste- ularly between adult Apatosaurus and Diplodocus, has vens and Parrish, 1999, 2005; Upchurch, 2000; Wilson, been limited (Fiorillo, 1998; Whitlock, 2011: D’Emic 2005; Whitlock, 2010; Stevens, 2013; Woodruff, 2017). and others, 2013). Here, I describe new specimens that The presence of such similar, large-bodied shed light on this ecological problem: two diplodocid

Citation for this article. McHugh, J.B., 2018, Evidence for niche partitioning among ground-height browsing sauropods from the Upper Jurassic Morrison Formation of North America: Geology of the Intermountain West, v. 5, p. 95–103. © 2018 Utah Geological Association. All rights reserved. For permission to use, copy, or distribute see the preceeding page or the UGA website, www.utahgeology.org, for information. Email inquiries to [email protected]. 95 Evidence for Niche Partitioning Among Ground-Height Browsing Sauropods from the Upper Jurassic Morrison Formation of North America McHugh, J.B. snout fragments containing numerous teeth ( The refining of these broad ecological groups was of Western Colorado specimens MWC 6002 and MWC attempted using snout shape morphometrics (Whit- 8430) from the Brushy Basin Member of the Morri- lock, 2011). Snout shapes were quantified for six di- son Formation excavated from Mygatt-Moore Quarry plodocoid genera (Apatosaurus sp., sp., (MMQ) in Rabbit Valley, western Colorado (figure 1). Diplodocus sp., Nigersaurus taqueti, emilieae, The MMQ is located within the McInnis Canyons and africana) using the ratio of the depth to National Conservation Area and is co-managed by the and breadth in the upper jaw, the premaxillary-maxil- Museums of Western Colorado (MWC) and the Bureau lary index, and the divergence angle of the premaxillae of Land Management (BLM). Excavations at the MMQ (Whitlock, 2011). Whereas some taxa were found to be for over thirty by crews from the MWC, as well as mathematically distinct from each other; unfortunately, the Dinamation International Society, have facilitated these metrics were unable to separate the broad snouted the recovery of over 5000 vertebrate fossils, including Apatosaurus and Diplodocus into disparate ecological specimens of the bony fish “Hulettia” hawesi groups (Whitlock, 2011). (Kirkland, 1998) and Morrolepis schaefferi (Kirkland, Tooth microwear has also been used to distinguish 1998), and the ankylosaur Mymoorapelta maysi (Kirk- among ecological resource use in sauropod land and Carpenter, 1994). However, material (Fiorillo, 1998; Whitlock, 2011). However, while tooth from the tetanuran Allosaurus fragilis and the diplodo- wear and microwear have been useful in differentiat- cid Apatosaurus cf. louisae dominate the assemblage ing feeding strategies between co-occurring taxa like (Foster and others, 2007, 2018). Two diplodocid taxa sp. and Diplodocus sp. (Fiorillo, 1998; are positively identified from the Rabbit Valley area: Di- Whitlock, 2011), suitable teeth from Apatosaurus sp. for plodocus sp. and Apatosaurus cf. louisae. Of these two, this type of analysis have been lacking (Whitlock, 2011). only Apatosaurus cf. louisae has been recovered within Although shed teeth are relatively common for preda- the MMQ. Other sauropods recovered from the quar- tors like Allosaurus fragilis, especially at the MMQ, shed ry include the macronarian, Camarasaurus sp., as well teeth from of Apatosaurus are less common and as several taxonomically ambiguous caudal vertebrae, are often difficult to differentiate from thoseDiplodo of - which are identified as Diplodocinae indet. (Foster and cus. Additional dental material is required to assess the others, 2018). microwear in the Apatosaurus, and for this type Sauropod dinosaurs include the largest terrestrial of analysis to be useful in partitioning the ecological herbivores that the Earth has ever known. Understand- niches of Apatosaurus and Diplodocus. ing their biology and interactions with plant biomass and Observations of modern herbivores show that eco- other sauropods within ecosystems has been logical niche partitioning is common among similar the subject of much research (Barrett and Upchurch, taxa within an environmental landscape (i.e., Bovidae), 1995; Stevens and Parrish, 1999, 2005; Upchurch, 2000; and that their dental and jaw morphologies are also Upchurch and Barrett, 2000; Hummel and others, 2008; correlated to this partitioning (Spencer, 1995). Howev- Yates and others, 2010; Gee, 2011; Tütken, 2011). Sau- er, with considerable morphological and biomechanical ropods have been broadly divided into “high browsing” overlap in sauropod dinosaurs between established eco- and “ground height” browsing ecological types based logical types, estimation of niche partitioning at lower upon snout shape and neck position (Whitlock, 2011), taxonomic levels has proven difficult (Fiorillo, 1998; but estimations of neck biomechanics indicate con- Whitlock, 2011). Tooth replacement may be an illu- siderable overlap in the range of feeding between dis- minating piece of evidence in fully assessing the niche parate types of sauropods (Stevens and Parrish, 1999, partitioning among sauropods. Since variable hardness 2005). Additionally, co-occurring genera within these within food sources can tend to create inconsistent wear broad categories, such as Apatosaurus and Diplodocus, on teeth, a higher level of tooth replacement are known from the Morrison Formation (Tschopp and is often associated with tougher vegetation. others, 2015). Tooth replacement rates among

Geology of the Intermountain West 96 2018 Volume 5 Evidence for Niche Partitioning Among Ground-Height Browsing Sauropods from the Upper Jurassic Morrison Formation of North America McHugh, J.B.

Figure 1. Map of Mesa County in western Colorado showing the location of the Mygatt-Moore Quarry. Insert shows exca- vations at the quarry during 2014.

have previously been described in Camarasaurus sp., physiological investment in individual teeth rather than Diplodocus sp., Dicraeosaurus hansemanni, and Niger- a biological investment in a lot of teeth. I present here saurus taqueti (Sereno and others, 2005; D’Emic and the first description of tooth replacement in a non-Di- others, 2013; Schwarz and others, 2015). The number plodocus diplodocid from the Morrison Formation that of teeth per alveolus within the premaxillae (including provides empirical evidence for an ecological partition both replacement and erupted tooth positions) vary between this and Diplodocus sp. among taxa, with four reported in Camarasaurus sp., six in Diplodocus sp., five to six in Dicraeosaurus han- semanni, and eight in Nigersaurus taqueti (Sereno and GEOLOGICAL SETTING others, 2005; D’Emic and others, 2013; Schwarz and others, 2015). D’Emic and others (2013) suggest the The fossil-bearing horizon at the MMQ is a 1- to lower number of teeth in Camarasaurus sp. is correlat- 2-m-thick unit exposed within laminated to medi- ed to the relative increase in tooth volume, and thus a um-bedded gray silty mudstone of the Brushy Basin

Geology of the Intermountain West 97 2018 Volume 5 Evidence for Niche Partitioning Among Ground-Height Browsing Sauropods from the Upper Jurassic Morrison Formation of North America McHugh, J.B. Member of the Morrison Formation (Kirkland and single alveolus. The mesial suture between the premax- Carpenter, 1994; Foster, 2003; Foster and others, 2007, illa and maxilla is poorly preserved and the ventrodistal 2018; Foster and Hunt-Foster, 2011). Radiometric anal- margin is incomplete. Therefore, it is doubtful that all ysis of ash-fall zircons from the quarry have returned maxillary alveoli are present in the specimen, and alve- an age of 152.18 ±0.29 Ma, which places the quarry on olar positions are instead referred to as mx1-mx8 and – boundary during the Late not to their exact maxillary placement (figure 2). Jurassic Period (Trujillo and others, 2014). The number of replacement teeth per alveolus de- The MMQ is interpreted to preserve an ecosystem creases from medial to distal. The mesial-most four with abundant vegetation and a high water table, but alveolar positions (mx1-mx4) each preserve seven to with standing water at the surface occurring only sea- eight replacement teeth, similar to the numbers pre- sonally (Foster, 2003; Trujillo and others, 2014; Foster served in specimen MWC 8430. The next two alveoli and others, 2018). The site preserves a -dom- (mx5-mx6) preserve six and five replacement teeth, re- inated assemblage with abundant plant material, with spectively, whereas the distal-most positions preserve crocodylomorph, , and invertebrate remains be- only four and three replacement teeth (mx7, mx8). This ing rare (Foster 2007; Foster and others, 2018). Previ- reduction in the number of replacement teeth reflects ous taphonomic work has shown the site represents an the decreasing volume in maxillary space distally. Tooth autochthonous assemblage within an attritional deposit replacement positions vi-viii are labial-linugally com- in an overbank setting with few articulated specimens pressed. Erupted crown morphology and the lingual and no preferred orientation of skeletal elements (Fos- inflection at the separation of root and crown are both ter and others, 2018). Despite the lack of transport evi- distinguishable by position v and become more pro- dent in the assemblage, fossils often preserve sur- nounced towards the erupted tooth position (i) (figures face modifications (e.g., breakage, abrasion, and feeding 3 and 4). traces) that suggest a reworked death assemblage by subsequent living individuals, possibly through feeding MWC 8430—Diplodocid Premaxilla Fragment and trampling (Foster and others, 2018). Specimen MWC 8430 is a well-preserved, isolated DESCRIPTION premaxilla fragment that preserves four sets of replace- ment teeth, each representing a single alveolus in the Both specimens in this study were excavated from upper jaw. However, lack of preservation on the exte- the gray mudstone of the Brushy Basin Member of the rior surface has not preserved the alveoli themselves. Morrison Formation at the MMQ using hand-quarry- The lingual face of the premaxilla is preserved as a thin, ing techniques. Both specimens were collected as iso- trapezoidal fragment with a smooth surface morpholo- lated elements with no additional associated cranial or gy. The fragmental nature of specimen MWC 8430 al- postcranial material. These specimens were mechani- lows for determination of lingual/labial and superior/ cally prepared at the MWC fossil preparation lab, re- inferior orientations, but not mesial/distal and it pro- paired, and stabilized using polyvinyl acetate dissolved hibits assignment to precise tooth row placement. The in acetone, as well as cyanoacrylate (PaleoBONDTM 750) alveolar positions are therefore referred to as px1-px4 adhesives, and then imaged using two-dimensional (2- (figure 5). D) photography and a 64-slice computed tomography The maximum number of replacement teeth (eight) (CT) scanner at Colorado Canyons Hospital in Fruita, Colorado. in an individual set are found within px1, with px2 and px3 each preserving seven teeth, and px4 with four teeth. Teeth in the replacement rows incrementally MWC 6002—Diplodocid Maxilla increase in length towards the erupted tooth position. Specimen MWC 6002 is a partial maxilla that pre- Tooth replacement positions vi-viii are labial-lingual- serves eight sets of teeth, with each set representing a ly compressed. Erupted crown morphology is distin-

Geology of the Intermountain West 98 2018 Volume 5 Evidence for Niche Partitioning Among Ground-Height Browsing Sauropods from the Upper Jurassic Morrison Formation of North America McHugh, J.B.

Figure 2. Diplodocid maxilla (spec- imen MWC 6002). (A) dorsal view. (B) ventral view. Scale bar equal 10 cm.

guishable by position v and becomes more pronounced tooth positions per alveolus in the premaxilla, including towards the erupted tooth position (i). Also visible at the erupted tooth (D’Emic and others, 2013; Schwarz, position v is the lingual inflection at the separation of 2015). The newly described specimens MWC 6002 and root and crown (figures 5 and 6). MWC 8430, although incomplete, preserve a maximum of eight teeth (both replacement teeth and erupted DISCUSSION tooth) per alveolus. The premaxilla exhibits the max- Despite the lack of association with more diagnostic imum number of replacement teeth, as supported by material, some taxonomic assessments for specimens the mesial increase in the number of tooth replacement MWC 6002 and MWC 8430 are possible. These speci- position seen in the maxilla, specimen MWC 6002. mens both bear the characteristic peg-like teeth of Di- However, this mesial increase in tooth replacement po- plodocoidea. However, they are not consistent with the sitions may not be characteristic of all sauropods, for known pattern of tooth replacement in Diplodocus sp. example in the titanosaur zdanskyi, the max- or the African taxon Dicraeosaurus hansemanni which imum number of tooth replacement positions is located have been shown to support no more than five to six within the maxilla and not the premaxilla (Poropat and

Geology of the Intermountain West 99 2018 Volume 5 Evidence for Niche Partitioning Among Ground-Height Browsing Sauropods from the Upper Jurassic Morrison Formation of North America McHugh, J.B.

Figure 3. CT scan of specimen MWC 6002 in and superior Figure 4. CT scan of specimen MWC 6002 in and mesial view. Alveolar positions are indicated with mx1-mx8 labels. view. Roman numerals indicate tooth positions for the mx2 alveolus.

Kear, 2013). A higher number of replacement teeth than the co- Minimally, specimens MWC 6002 and MWC 8430 eval Diplodocus sp. indicates that these large-bodied, can be assigned as a non-Diplodocus diplodocid. The “ground height” browsing herbivores were orally pro- only other known diplodocid taxa from the Rabbit Val- cessing different types of vegetation, withApatosaurus ley consists of the very abundant Apatosaurus cf. louisae sp. potentially adapted for tougher, more abrasive veg- and the ambiguous vertebrae not positively identified etation than Diplodocus sp. The Morrison Formation is beyond Diplodocinae indet. The MMQ in Rabbit Val- interpreted as representing a predominantly riparian ley has been explored in scientific earnest for more than landscape with limited botanical biomass (Foster, 2003; three decades, producing thousands of fossils, none of Engelmann and others, 2004; Rees and others, 2004; which were positively assigned to any diplodocid taxon Turner and Peterson, 2004; Hummel and others, 2008; other than Apatosaurus cf. louisae. And although the Gee, 2011). This partitioning of resources between large possibility of future diplodocoid discoveries at the site herbivores would have been critical for survival in this cannot be discounted, given the present data available, limiting environment, and may have been the key to the it is most parsimonious for specimens MWC 6002 and success and diversity of the sauropods in the Late Juras- MWC 8430 to be tentatively referred to Apatosaurus sp., sic Period. pending the recovery of more diagnosable cranial ma- terial. CONCLUSIONS Despite a closer phylogenetic, geographic, and stratigraphic relationship with Diplodocus sp., the spec- Populations of co-occurring herbivores in modern imens in this study that are referred to Apatosaurus sp. landscapes often partition plant biomass resources to are more similar to the tooth replacement counts seen facilitate survival in competitive environments. Using in the Cretaceous African form Nigersaurus taqueti. modern analogs, paleontologists have long hypothe-

Geology of the Intermountain West 100 2018 Volume 5 Evidence for Niche Partitioning Among Ground-Height Browsing Sauropods from the Upper Jurassic Morrison Formation of North America McHugh, J.B.

Figure 5. Specimen MWC 8430 in (A) ?mesial, (B) ?distal, (C) labial, (D) lingual, (E) superior, and (F) inferior views. Roman numerals indicate tooth posi- tions for an individual alveolus. All scale bars equal 1 cm.

Figure 6. CT scan of specimen MWC 8430 in and inferior view. Roman numerals indicate tooth positions for individual alveoli. Alveolar positions are indicated with px1-px4 labels.

Geology of the Intermountain West 101 2018 Volume 5 Evidence for Niche Partitioning Among Ground-Height Browsing Sauropods from the Upper Jurassic Morrison Formation of North America McHugh, J.B. sized similar niche partitioning among large sauropod Engelmann, G.F., Chure, D.J., and Fiorillo, A.R., 2004, The implica- dinosaurs in the Mesozoic. New specimens from the tions of a dry climate for the paleoecology of the fauna of the MMQ in Rabbit Valley, Colorado, reveal higher counts Upper Jurassic Morrison Formation: Sedimentary Geology, v. 167, no. 3-4, p. 297–308. of tooth replacement in Apatosaurus sp. than in Diplod- ocus sp. This indicates that species of Apatosaurus may Fiorillo, A.R., 1998, Dental micro wear patterns of the sauropod di- nosaurs camarasaurus and diplodocus—evidence for resource have been selectively feeding on tougher vegetation partitioning in the Late Jurassic of North America: Historical (e.g., and ) than species of Diplodocus Biology, v. 13, no. 1, p. 1–16. and provides the strong empirical evidence for niche Foster, J.R., 2003, Paleoecological analysis of the vertebrate fauna partitioning among these coeval and morphologically of the Morrison Formation (Upper Jurassic), Rocky Mountain similar taxa. region, USA: Museum of Natural History and Science Bulletin 23, 95 p. ACKNOWLEDGMENTS Foster, J.R., Hunt, R.K., and King, L.R., 2007, Taphonomy of the Mygatt-Moore Quarry, a large dinosaur bonebed in the Up- Thank you to Rob Gay (Colorado Canyons Associ- per Jurassic Morrison Formation of western Colorado [abs.] ation), Chris Racay (Museums of Western Colorado), Geological Society of America Abstracts with Programs, v. 39, John Foster (Museum of Moab), ReBecca Hunt-Foster no. 6, p. 400. (Bureau of Land Management, Canyon Country Dis- Foster, J.R., and Hunt-Foster, R.K., 2011, New occurrences of trict), and Michael D’Emic (Adelphi University) for dinosaur skin of two types (? and Dinosauria in- det.) from the Late Jurassic of North America (Mygatt-Moore their helpful discussions on this project. I thank Doro- Quarry, Morrison Formation): Journal of Vertebrate Paleon- thy Stewart and the volunteer corps at Dinosaur Journey tology, v. 31, no. 3, p. 717–721. Museum, as well as Dinosaur Journey field coordina- Foster, J.R., Hunt-Foster, R.K., Gorman, M.A., II, Trujillo, K.C., tor Chris Racay and field assistant Mitchell Lukens for Suarez, C.A., McHugh, J.B., Peterson, J.E., Warnock, J.P., and assistance in recovery and preparation of specimens. I Schoenstein, H.E., 2018, Paleontology, taphonomy, and sed- sincerely thank the Bureau of Land Management Grand imentology of the Mygatt-Moore Quarry, a large dinosaur Junction Field Office for support and site access, as well bonebed in the Morrison Formation, western Colorado—im- plications for Upper Jurassic dinosaur preservation modes: as Dr. Michael Neste, M.D. (Family Health West) and Geology of the Intermountain West, v. 5, p. 23–89. the Colorado Canyons Hospital in Fruita for the use of Gee, C.T., 2011, Dietary options for the sauropod dinosaurs from CT imaging facilities. Thank you to Stephen K. Poro- an integrated botanical and paleobotanical perspective, in pat (Swinburne University of Technology) and Femke Klein, N., Remes, R., Gee, C.T., and Sander, P.M., editors, Bi- Holwerda (Bayerische Staatssammlung für Paläontolo- ology of the sauropod dinosaurs—understanding the life of gie und Geologie) for their thoughtful comments on an giants: Bloomington, Indiana University Press, p. 34–56. earlier version of this manuscript, and I also thank Cary Hummel, J., Gee, C.T., Südekum, K-H., Sander, P.M., Nogge, G., Woodruff (Great Plains Dinosaur Museum), John Fos- and Clauss, M., 2008, In vitro digestibility of and gymno- ter (Museum of Moab), and Daniela Schwarz (Museum sperm foliage—implications for sauropod feeding ecology and diet selection: Proceedings of the Royal Society of London B: für Naturkunde) for their time and comments on this Biological Sciences, v. 275, no. 1638, p. 1015–1021. manuscript. Kirkland, J.I., 1998, Morrison fishes: Modern Geology, v. 22, p. 503–533. REFERENCES Kirkland, J.I., and Carpenter, K., 1994, North America’s first Barrett, P., and Upchurch, P., 1995, Sauropod feeding mecha- pre-Cretaceous ankylosaur (Dinosauria) from the Upper Ju- nisms—their bearing on palaeoecology, in Sun, A., and Wang, rassic Morrison Formation of western Colorado: Brigham Y., editors, Proceeding 6th symposium, Mesozoic terrestrial Young Univerity Geological Studies, v. 40, p. 25–42. ecosystems and biotas: Beijing, China Ocean Press, p. 107–110. Poropat, S.F., and Kear, B.P., 2013, Photographic and three-di- D’Emic, M.D., Whitlock, J.A., Smith, K.M., Fisher, D.C., and Wil- mensional reconstruction of the holotype of Euhelopus son, J.A., 2013, Evolution of high tooth replacement rates in zdanskyi with description of additional cranial elements: PLoS sauropod dinosaurs: PLoS ONE, v. 8, no. 7, p. e69235. ONE, v. 8, no. 11, p. e79932.

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