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Biggest of the Big: a Critical Re-Evaluation of the Mega-Sauropod Amphicoelias Fragillimus Cope, 1878

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Biggest of the Big: a Critical Re-Evaluation of the Mega-Sauropod Amphicoelias Fragillimus Cope, 1878 Foster, J.R. and Lucas, S. G., eds., 2006, Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin 36. 131 BIGGEST OF THE BIG: A CRITICAL RE-EVALUATION OF THE MEGA-SAUROPOD AMPHICOELIAS FRAGILLIMUS COPE, 1878 KENNETH CARPENTER Department of Earth Sciences, Denver Museum of Nature & Science, Denver, CO 80205 [email protected] Abstract—Questions about the largest dinosaur have largely ignored or downplayed a discovery made in 1878 of a 1.5 m tall neural arch. The specimen is lost, but the original description by E.D. Cope of Amphicoelias fragillimus provides enough information to reconstruct the vertebra as a posterior dorsal of a diplodocid. The estimated 2.7 m height of the dorsal suggests a skeleton 58 m long, about 9.25 m at the highest point of the back, and with a body mass of 122,400 kg. These dimensions indicate that A. fragillimus was several orders of magnitude larger than any other sauropod. A. fragillimus existed at the peak of a trend toward gigantism in sauropods that progressed since their origin in the Late Triassic. One possible cause for large body size in sauropods, based on studies of extant mammalian megaherbivores, may be due to increased gut size for more efficient digestion of low quality browse by hindgut fermentation. INTRODUCTION Discoveries during the past decade or so of very large sauropods (>30 m) have led to the question about which was the largest dinosaur. Van Valen (1969) attempted to answer that question before many of the recent discoveries of large sauropods, such as of Argentinosaurus, thus was limited in the data available for his analysis. Contenders today include the diplodocids Seismosaurus hallorum from the Upper Jurassic of New Mexico (Gillette, 1991; as Diplodocus hallorum by Lucas et al., 2004 and Lucas et al. this volume) and Supersaurus vivianae from the Upper Juras- sic of Colorado and Wyoming (Jensen, 1985; Lovelace et al., 2005), and the titanosaurids Argentinosaurus huinculensis from the Lower Cretaceous of Argentina (Bonaparte and Coria, 1993) and “Antarctosaurus” giganteus from the Upper Cretaceous of Argentina (Huene, 1929; Powell, 2003). All of these sauropods have estimated lengths of 30-35 m and estimated weights of between 30-90 tons (Mazzetta et al., 2004; Paul,1994) one possible contender, Paralatitan stromeri (Smith et al., 2001) is apparently shorter than 30 m. Intriguingly, these sauropods are only known from fragmentary remains. Based on the bone sample study in Amboseli Park by Behrensmeyer and Dechant Boaz (1980), most of the skeleton for such giant sauropods should be present. Although some of the loss is undoubt- edly due to erosion of the fossils, most of the specimens consist of disar- ticulated and scattered skeletons. Considering the great mass of the bones, this dispersal is remarkable in light of the bone dispersal studies by Voorhies (1969) and Behrensmeyer (1975). One other giant sauropod from the Upper Jurassic of North America, Amphicoelias fragillimus Cope, 1878b, is either ignored (e.g., Gillette, 1991) or only briefly mentioned (e.g., Mazzetta et al., 2004). Only Paul (1997), following preliminary work by Carpenter (1996), has discussed FIGURE 1. A. Amphicoelias fragillimus (AMNH 5777) as figured by Cope this taxon, but even this is brief. Because this taxon may indeed be the (1878b), modified with identification of features. B. Dorsal reconstructed based on largest dinosaur known, a fuller assessment is presented below. modified A. altus (extensively modified from Osborn and Mook, 1921, fig. 21. C. Dorsal of A. altus (AMNH 5764) for comparison (modified from Osborn and Mook, The holotype of A. fragillimus consists of a neural arch and spine 1921, fig. 21). C. cavities bounded by lamina within the neural arch; h, hyposphene; (Fig. 1) believed to have been from the last or second to last dorsal (D10 or l, lamina within the neural arch (outer bone eroded away exposing interior); lat D9). It was collect by Oramel Lucas near the main Camarasaurus supremus spol, lateral spinopostzygapophyseal lamina; med spol, medial localities in Garden Park, north of Cañon City, Colorado, as indicated spinopostzygopophyseal lamina; pcdl, posterior centroparapophyseal lamina; posl, by E.D. Cope’s field notes (Fig. 2). Lucas began collecting for Cope in postspinal lamina; poz, postzygapophysis; spol, spinopostzygapophyseal lamina. late 1877, and shipped the specimen to Cope during the spring or early summer of 1878 (McIntosh, 1998), who then published it in August mudstone and the specimen was probably very pale tan, tinted with a 1878. Cope visited Lucas in July, 1879 and recorded in a small note- maroon red, as are all the surviving specimens from this level. Cope book the various quarries from which Lucas had been excavating (McIn- also records “IV Immense distal end of femur near 1st broken smaller tosh, 1998; Monaco, 1998). Among the entries is “III Amphicoelias femur.” It seems very likely that this femur belonged to the same speci- fragillissimus [sic] from between the two lots” (lots refers to quarries men as the neural arch and spine because it was found in the next site or sites). Specifically, the site is southwest of the “hill” of Cope’s notes a few tens of meters away. Unfortunately, the whereabouts of both the (now referred to as Cope’s Nipple), and south of the Camarasaurus neural spine and femur are unknown as first reported by Osborn and supremus I Quarry. The matrix in the vicinity of the quarry is a reddish Mook (1921), and all attempts to locate the specimens have failed 132 DESCRIPTION It is unfortunate that the specimen of Amphicoelias fragillimus is missing, and that comparisons with other sauropods must rely on Cope’s description. Although missing, it has been assigned the catalog number AMNH 5777. The generic referral by Cope was based on his (1878b, p. 563) observation that “[i]t exhibits the general characteris- tics of the genus Amphicoelias, in the hyposphen [sic], antero-posteri- orly placed neural spine, and elevated diapophysis for the rib articula- tion. The diapophyses are compressed and supported by a superior [i.e., spinodiapophyseal lamina] and inferior [i.e., centrodiapophyseal lamina], and anterior [i.e., prezygodiapophyseal lamina] and posterior [i.e., postzygodiapophyseal lamina], thin buttresses [i.e., lamina], sepa- rated by deep cavities.” The genus Amphicoelias (as A. altus) was named by Cope in De- cember, 1877 (published in 1878a) for a mid-dorsal vertebra (D6?), a pos- terior dorsal vertebra (D10?), a pubis, and a femur (all AMNH 5764) from Quarry XII located at the north end of a long ridge (Fig. 2) about 1.5 km northwest of the main quarries. Osborn and Mook (1921) also referred a scapula, a coracoid, an ulna, and a tooth to the holotype. Quarry XII is stratigraphically the highest in the Morrison Formation in the Garden Park area, being about 15 m higher than the Camarasaurus supremus sites (Car- penter, 1998). As noted by Osborn and Mook (1921), the specimen closely resembles Diplodocus, especially in the posterior dorsal (best preserved): tall, slender neural spine, tall neural arch, well developed spinodiapophyseal laminae, and prominent posterior centroparapophyseal laminae. However, it also differs from Diplodocus having in a centrum that is as long as tall rather than taller than long, having an anteroposteriorly elongated, laterally compressed neural spine, parapophysis separate from the diapophysis, dia- mond-shaped hyposphene, pleurofossa (defined by Carpenter and Tidwell, FIGURE 2. A. E.D. Cope’s 1879 field notebook contains a map showing the location 2005, p. 94) restricted to the centrum (not extending onto the neural arch), of the Lucas brothers quarries. B. This map allowed relocation of many of the quarries. and less constricted (“pinched”) centrum. The ulna is significantly longer Amphicoelias fragillimus was found at quarry III in a string of closely spaced quarries relative to the femur length (0.6) than in Diplodocus (0.46); thus, the fore- (I-VIII). I-XIII quarries; A.f. Amphicoelias fragillimus; A.l., - Amphicoelias latus limbs were proportionally longer as well. The circular cross-section of the quarry; CS1, 2, Camarasaurus supremus 1 and 2 quarries; H.v., probable site for femur, once used to separate Amphicoelias from Diplodocus, is known to Hallopus victor Marsh (see Ague et al., 1995). Most of the quarries along the ridge occur in Diplodocus as well (V. Tidwell, personal commun.). have not yet been relocated. See McIntosh (1998) for further discussion of the Cope The species name, fragillimus, apparently reflects the fragility or map. delicateness of the specimen produced by the thinness of the numerous (McIntosh, 1998). laminae as noted by Cope (1878b, p. 563): “[in] the extreme tenuity [i.e., An attempt was made to relocate the quarry in 1994 including using thinness] of all its parts, this vertebra exceeds those of this type [i.e., ground penetrating radar. However, because the density of fossilized bone Amphicoelias altus] already described...”. These laminae not only include is the same as the encasing matrix, the technique, which relies on density the external ones connecting various structures, but also internal struts within differences, did not work. In addition, the mudstone containing lots I-IV is the neural arch (Fig. 1A). These internal struts or laminae are exposed be- nearly stripped down to the underlying sandstone. The local topography cause the missing outer cortical bone was lost, probably to erosion. The suggests that most of the strata had already been stripped off by the time delicateness of the specimen made collecting difficult, “much care was Lucas made his first dinosaur bone discovery in 1877. Thus, it is probable requisite to secure its preservation” (Cope, 1878b, p. 563), and may pro- that most of the Amphicoelias skeleton was destroyed long before the neu- vide a clue to its disappearance (as discussed below).
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