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Dinosaur Census Reveals Abundant Tyrannosaurus and Rare Ontogenetic Stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA

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Dinosaur Census Reveals Abundant Tyrannosaurus and Rare Ontogenetic Stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA Dinosaur Census Reveals Abundant Tyrannosaurus and Rare Ontogenetic Stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA John R. Horner1*, Mark B. Goodwin2, Nathan Myhrvold3 1 Museum of the Rockies, Montana State University, Bozeman, Montana, United States of America, 2 Museum of Paleontology, University of California, Berkeley, California, United States of America, 3 Intellectual Ventures, Bellevue, Washington, United States of America Abstract Background: A dinosaur census recorded during the Hell Creek Project (1999–2009) incorporates multiple lines of evidence from geography, taphohistory, stratigraphy, phylogeny and ontogeny to investigate the relative abundance of large dinosaurs preserved in the Upper Cretaceous Hell Creek Formation of northeastern Montana, USA. Overall, the dinosaur skeletal assemblages in the Hell Creek Formation (excluding lag-influenced records) consist primarily of subadult or small adult size individuals. Small juveniles and large adults are both extremely rare, whereas subadult individuals are relatively common. We propose that mature individuals of at least some dinosaur taxa either lived in a separate geographic locale analogous to younger individuals inhabiting an upland environment where sedimentation rates were relatively less, or these taxa experienced high mortality before reaching terminal size where late stage and often extreme cranial morphology is expressed. Methodology/Principal Findings: Tyrannosaurus skeletons are as abundant as Edmontosaurus, an herbivore, in the upper Hell Creek Formation and nearly twice as common in the lower third of the formation. Smaller, predatory dinosaurs (e.g., Troodon and dromaeosaurids) are primarily represented by teeth found in microvertebrate localities and their skeletons or identifiable lag specimens were conspicuously absent. This relative abundance suggests Tyrannosaurus was not a typical predator and likely benefited from much wider food choice opportunities than exclusively live prey and/or specific taxa. Tyrannosaurus adults may not have competed with Tyrannosaurus juveniles if the potential for selecting carrion increased with size during ontogeny. Conclusions/Significance: Triceratops is the most common dinosaur and isolated skulls contribute to a significant portion of this census. Associated specimens of Triceratops consisting of both cranial and postcranial elements remain relatively rare. This rarity may be explained by a historical collecting bias influenced by facies and taphonomic factors. The limited discovery of postcranial elements may also depend on how extensive a fossil quarry is expanded after a skull is collected. Citation: Horner JR, Goodwin MB, Myhrvold N (2011) Dinosaur Census Reveals Abundant Tyrannosaurus and Rare Ontogenetic Stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA. PLoS ONE 6(2): e16574. doi:10.1371/journal.pone.0016574 Editor: Peter Roopnarine, California Academy of Sciences, United States of America Received July 29, 2010; Accepted January 4, 2011; Published February 9, 2011 Copyright: ß 2011 Horner et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: This work was supported with funding by the following individuals: James Kinsey, Catherine B. Reynolds, and Homer Hickam. Intellectual Ventures, the Windway Foundation, and the Smithsonian Institution provided additional funding. The University of California Museum of Paleontology provided funding to MBG. NM is the founder and CEO of Intellectual Ventures and contributed to the study design. Homer Hickham participated as a Hell Creek Project volunteer in the field in Montana. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: Nathan Myhrvold contributed financially to the Hell Creek Project and intellectually to the design of the study. * E-mail: [email protected] Introduction Creek Formation. A dataset by Sheehan et al. [1] was used by White et al. [3] to examine the structure of the dinosaurian assemblage The Hell Creek Project (1999–2009), a collaborative, multi- with regard to taphonomy only. Russell and Manabe [4] provided a institutional field study of the Upper Cretaceous Hell Creek clearer picture of relative abundances within the dinosaur Formation, northeastern Montana, produced this dinosaur census assemblage, but failed to subdivide the Hell Creek Formation from a well-documented collection of all taxa. The overall goal of stratigraphically or to include ontogenetic perspectives in their the project was to create a comprehensive biotic foundation from analysis of previous surveys and collections. We think it is essential which paleobiological and geological hypotheses could be tested. that all temporal and spatial points of reference be considered One of the many projects included the collection of dinosaur synthetically when analyzing taxa from the fossil record – a type of specimens to test hypotheses focusing on the relative abundances unified field theory for paleontological specimens. Unified frames of and the presence or absence of various dinosaurian ontogenetic reference (UFR) include geography, taphohistory (defined here as stages. Previous Hell Creek Formation surveys [1,2] attempted to the history of the specimen from death to final disposition within statistically support particular extinction hypotheses, but offered space and time), stratigraphy, phylogeny and ontogeny. minimal information on the actual composition of the stratigraphi- In this current survey, we focused on the Upper Cretaceous Hell cally dispersed assemblages through the entire section of the Hell Creek Formation exposed around Fort Peck Reservoir in PLoS ONE | www.plosone.org 1 February 2011 | Volume 6 | Issue 2 | e16574 Dinosaur Census from the Hell Creek Formation northeastern Montana. Here, contiguous outcrops are traceable continuing evaluation of the geology and paleontology of M3 will over an area of about 1000 sq. km (Figure 1). Facies changes were be reported on in a follow-up study by the senior author and the tracked within the formation’s 90 to 100 meter thickness. Each Hell Creek Project team. confirmed skeleton was recorded and evaluated with regard to its In this study, the L3 and U3 are subdivided based on UFR. This dataset is archived in the Museum of the Rockies sedimentological and stratigraphic details confirmed in the study (MOR). area (Figure 1). The L3 basal sandstone (L3.BS) is further Here we present the relative abundance of large dinosaur taxa, subdivided into three stratigraphic horizons: lower, L3.lBS; their general ontogenetic stage within a stratigraphic and middle, L3.mBS; and upper, L3.uBS. An overlying variegated sedimentological context and offer generalized hypotheses to mudstone (L3.MS) is also subdivided into three traceable units: explain several of the more interesting patterns that have emerged. lower, L3.lMS; middle, L3.mMS; and upper, L3.uMS. Because a This dinosaur census sets the stage for future statistical analysis and dinosaur – bearing lag deposit was not identified in the lower third evaluation of the end Cretaceous faunal record and will add to (L3) of the Hell Creek Formation, we use the Jen-rex sand [7] at recent paleoecological studies of diverse dinosaur faunas and the base of M3 as a proxy for this facies in L3 because of its clades that focused on body size, habitat partitioning and living stratigraphic position along the M3–L3 contact (see Figure 2). space requirements [5] or abundance modeling [6]. The U3 is comprised of two sandstone units: the Apex The Hell Creek Project spanned 11 field seasons from 1999– sandstone (U3.AS) and the 10-meter sandstone (U3.10mS). An 2009. The first five years focused on the collection of specimens interbedded variegated mudstone (U3.ibMS) separates these two from the lower third of the formation (See Figure 1, dark orange sandstones and an upper variegated mudstone (U3.uMS) overlies area), while the remaining six years were aimed at specimen the 10-meter sandstone. U3 is capped by an overlying uppermost collection from the middle and upper strata (Figure 1, light orange coal zone (U3.CZ). area). Since several of the dinosaur specimens from the most High-resolution digital video (remote control camera mounted recent field seasons (2008–2010) remain unprepared, the data on a Bell 407 helicopter) was utilized to track stratigraphic presented here for the upper two-thirds of the formation will be horizons and facies over large distances to assure stratigraphic updated as Hell Creek Project specimens are prepared and resolution and supplement standard stratigraphic sections and available from the corresponding author and on www.museu- mapping within the study area. moftherockies.org. Census Methods Methods Census figures for the dinosaurs were determined using two different methods: (1) recording numbers of articulated or Geological Methods associated skeletons and (2) recording numbers of individuals In order to evaluate possible changes in dinosaur taxa and their based on isolated elements collected from lag deposits in the L3 relative abundance through time, the Hell Creek Formation was ( = Jen-rex sand) and U3 ( = Apex sandstone). Although more than divided into three stratigraphic units [7,8] designated L3 (lowest), 150
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