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The Biogeography and Ecology of the Cretaceous Non-Avian Dinosaurs of Appalachia

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The Biogeography and Ecology of the Cretaceous Non-Avian Dinosaurs of Appalachia Palaeontologia Electronica palaeo-electronica.org The biogeography and ecology of the Cretaceous non-avian dinosaurs of Appalachia Chase D. Brownstein ABSTRACT The Cenomanian to Maastrichtian of the Late Cretaceous saw the flooding of the interior of North America by the Western Interior Seaway, which created the eastern landmass of Appalachia and the western landmass of Laramidia. Though Appalachian dinosaur faunas are poorly known, they are nevertheless important for understanding Cretaceous dinosaur paleobiogeography and ecology. In order to better track the vicariance of eastern and western North American dinosaur faunas over the duration of the Cretaceous, the former were compared with the latter from the Aptian to Maastrich- tian Stages of the Late Cretaceous using several similarity indices. The data gathered from biogeographic similarity indices suggest that an almost completely homogenous North American dinosaur fauna found in the Early Cretaceous experienced significant vicariance, splitting into a Laramidian fauna differentiated by the presence of ceratop- sids, pachycephalosaurids, saurolophids, lambeosaurines, ankylosaurids, therizino- saurids, and troodontids and an Appalachian fauna characterized by the lack of the aforementioned groups and the presence of non-hadrosaurid hadrosauroids, massive hadrosauroids, basal hadrosaurids, leptoceratopsians, “intermediate”-grade tyranno- sauroids, and nodosaurids between the Cenomanian and Campanian, with these two faunas later experiencing limited dispersal after the disappearance of the Western Inte- rior Seaway from the American Interior during the Maastrichtian. Dinosaur provincial- ism and ecology on Appalachia are also investigated and discussed. Though the fossil record of dinosaurs for parts of the Cretaceous is poor throughout North America and in the eastern portion of the continent especially, the analyses herein nevertheless allow for a greater glimpse at dinosaur biogeography and ecology in Appalachia and in North America generally during the time. Chase D. Brownstein. Research Associate, Stamford Museum & Nature Center, Stamford, CT. USA. [email protected] Keywords: paleobiogeography; paleoecology; Appalachia; Cretaceous; Dinosauria Submission: 5 July 2017 Acceptance: 17 January 2018 Brownstein, Chase D. 2018. The biogeography and ecology of the Cretaceous non-avian dinosaurs of Appalachia. Palaeontologia Electronica 21.1.5A 1-56. https://doi.org/10.26879/801 palaeo-electronica.org/content/2018/2123-appalachia-biogeography Copyright: February 2018 Society for Vertebrate Paleontology. 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. creativecommons.org/licenses/by/4.0/ BROWNSTEIN: APPALACHIA BIOGEOGRAPHY INTRODUCTION The recent discoveries of a leptoceratopsian from the Campanian Tar Heel Formation of North During the Albian to Cenomanian stages of Carolina (Longrich, 2016), the description of the the Late Cretaceous, the interior of North America dinosaurs Eotrachodon orientalis from the Santo- was flooded by a shallow sea called the Western nian-Campanian Mooreville Chalk Formation Interior Seaway (e.g., Russell, 1995; Roberts and (Prieto-Marquez et al., 2016a) and Appalachiosau- Kirschbaum, 1995). The creation of the seaway rus montgomeriensis from the Campanian Demop- caused the formation of a long, slender landmass olis Chalk Formation (Carr et al., 2005), and the known as Laramidia to the west and the wider, description of new dinosaur remains from microfos- more rectangular Appalachia to the east (e.g., sil sites like Ellisdale and Stokes Quarry (e.g., Gal- Schwimmer, 2002; Sampson et al., 2010), having lagher, 1993; Denton et al., 2011; Schwimmer, appreciable consequences in the evolution of 2015) have largely increased the non-avian dino- North American dinosaurs. The former of these two saur diversity of Appalachia. The dinosaur fauna of landmasses apparently experienced a heightened Appalachia itself was apparently dominated by rel- level of non-avian dinosaur diversification during ict forms isolated by the Western Interior Seaway the Campanian (Sampson et al., 2010; Loewen et (Schwimmer et al., 1993). However, the develop- al., 2013), when a high level of local dinosaur ment of Appalachian faunas and their eventual endemism partnered with regional differences in mixing with Laramidian ones during the Maastrich- non-avian dinosaur faunas may have occurred tian (Schwimmer et al., 1993; Carr et al., 2005) is there (Lehman, 1997; Sampson et al., 2010; Loe- poorly understood. wen et al., 2013). The cause for this rapid diversifi- Here, I review and statistically compare fau- cation of non-avian dinosaurs and other nas known from the Aptian through Maastrichtian vertebrates on Laramidia has been hypothesized of eastern North America with those known from as a consequence of climate variability leading to the west to better illustrate the vicariance of Appa- floral variability, of orogenesis or orogenic activity, lachian and Laramidian non-avian dinosaur faunas and of transgressions and regressions of the West- after the creation of the Western Interior Seaway. ern Interior Seaway (Horner et al., 1992; Lehman, This analysis provides a temporal framework for 1997; Sampson et al., 2010; Loewen et al., 2013). the evolution of non-avian dinosaur faunas on There has been some disagreement regarding the Appalachia. Additionally, non-avian dinosaur pro- existence of faunal provinces on this continent vincialism during the Coniacian, Santonian, and (e.g., Larson and Vavrek, 2010; Lucas et al., 2016). Campanian Stages of the Cretaceous was tested using several biogeographic similarity indices and Unfortunately, the terrestrial fauna of Appala- discussed, as were ecological implications regard- chia is not well-sampled, and only a limited number ing competition between predatory dinosaurs and of dinosaur taxa are known due to lack of Creta- crocodyliforms in Appalachian ecosystems. ceous-age terrestrial sediments in eastern North Because the vast majority of Appalachian dinosaur America (Baird and Horner, 1977; Baird and Gal- remains occur in marine deposits and Laramidian ton, 1981; Baird, 1986; Schwimmer, 1986; King et dinosaurs are comparatively well-known from ter- al., 1988; Gates et al., 2012). Preservation bias restrial ones, depositional bias was surely a factor thus exists against any articulated specimens of that may have skewed the results of the statistical terrestrial animals, including non-avian dinosaurs analyses conducted (e.g., Schwimmer, 1997) and (e.g., Schwimmer et al., 1993; Schwimmer, 1997). is discussed below. This analysis is important for Schwimmer (1997) suggested that, antithetic to the being the first major one to compare the non-avian hypotheses of Russell (1995), the known dinosaurs dinosaur faunas of Appalachia with Laramidian from Appalachia may represent a fair selection of ones statistically. Specific depositional compari- non-avian dinosaur groups endemic to the conti- sons between Appalachian and Laramidian faunas nent. Schwimmer (2002) also suggested that a analyzed are noted in the results section. lack of theropod diversity or abundance on Appala- chia may have also been caused by competition MATERIALS AND METHODS from the massive crocodylian Deinosuchus rugo- sus, a species abundant in eastern North America Permits with bite marks on both theropod and ornithopod No permits were required for the described bones attributed to it (Schwimmer, 1997; Galla- study, which complied with all relevant regulations. gher, 1993, 1995; Schwimmer 2002). Photographs of specimens from the Arundel Clay 2 PALAEO-ELECTRONICA.ORG referred to herein were supplied by Thomas Jors- Formation. Cenomanian Appalachian faunas were tad of the National Museum of Natural History. compared with the upper Cedar Mountain Forma- Photographs of the specimens figured herein from tion (Mussentuchit Member), Chandler Formation, the Yale Peabody Museum were provided by and Dunvegan Formation. Turonian Appalachian Jamie Henderson. faunas and the non-avian dinosaur faunas of the Moreno Hill, Frontier, and Matanuska formations Institutional Abbreviations were compared. Santonian Appalachian faunas The following abbreviations for museum col- were compared with those of the Milk River Forma- lections are used in the manuscript: USNM V/PAL: tion. Campanian Appalachian faunas were com- United States National Museum (Smithsonian), pared to those of the Wahweap, Kaiparowits, and Washington, DC, USA; UAM(1): University of Kirtland formations representing southern Campa- Arkansas at Fayettville, Fayettville, AK, USA; YPM nian Laramidian faunas and with the Oldman, VPPU: vertebrate paleontology collections, Yale Dinosaur Park, and Judith River formations repre- Peabody Museum, New Haven, CT, USA; RMM/ senting northern Campanian Laramidian faunas, MCWSC: McWane Science Center, Birmingham, whereas the Maastrichtian Appalachian faunas Alabama, USA; AMNH FARB: American Museum were compared with those of the Javelina, Hell of Natural History (fossil amphibian, bird, and rep- Creek, Lance, and Horseshoe Canyon formations. tile collections), New York, NY, USA; ANSP: Acad- The Laramidian (western North American) faunas emy of Natural Sciences at Drexel University, used for comparisons are among the most well Philadelphia, PA, USA; AUMP: Auburn University known (e.g., Kirkland et al.,
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