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The Early Jurassic Ornithischian Dinosaurian Ichnogenus Anomoepus

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The Early Jurassic Ornithischian Dinosaurian Ichnogenus Anomoepus 19 The Early Jurassic Ornithischian Dinosaurian Ichnogenus Anomoepus Paul E. Olsen and Emma C. Rainforth nomoepus is an Early Jurassic footprint genus and 19.2). Because skeletons of dinosaur feet were not produced by a relatively small, gracile orni- known at the time, he naturally attributed the foot- A thischian dinosaur. It has a pentadactyl ma- prints to birds. By 1848, however, he recognized that nus and a tetradactyl pes, but only three pedal digits some of the birdlike tracks were associated with im- normally impressed while the animal was walking. The pressions of five-fingered manus, and he gave the name ichnogenus is diagnosed by having the metatarsal- Anomoepus, meaning “unlike foot,” to these birdlike phalangeal pad of digit IV of the pes lying nearly in line with the axis of pedal digit III in walking traces, in combination with a pentadactyl manus. It has a pro- portionally shorter digit III than grallatorid (theropod) tracks, but based on osteometric analysis, Anomoepus, like grallatorids, shows a relatively shorter digit III in larger specimens. Anomoepus is characteristically bi- pedal, but there are quadrupedal trackways and less common sitting traces. The ichnogenus is known from eastern and western North America, Europe, and southern Africa. On the basis of a detailed review of classic and new material, we recognize only the type ichnospecies Anomoepus scambus within eastern North America. Anomoepus is known from many hundreds of specimens, some with remarkable preservation, showing many hitherto unrecognized details of squa- mation and behavior. . Pangea at approximately 200 Ma, showing the In 1836, Edward Hitchcock described the first of what areas producing Anomoepus discussed in this chapter: 1, Newark we now recognize as dinosaur tracks from Early Juras- Supergroup, eastern North America; 2, Karoo basin; 3, Poland; sic Newark Supergroup rift strata of the Connecticut 4, Colorado Plateau. The boxed area is shown in detail in figure Valley (Hartford and Deerfield basins) (figures 19.1 19.2A. (Modified from Kent et al. 1995) Early Jurassic Ornithischian Dinosaurian Ichnogenus Anomoepus • 315 . Newark Supergroup and localities for Anomoepus.(A) Newark Supergroup, showing the locations of the maps shown in B, C, and D (rectangles); basins: 1, Fundy; 2, Hartford and Deerfield; 3, Newark; 4, Gettysburg; 5, Culpeper; 6, Taylorsville; 7, Richmond; 8, Farmville and associated basins; 9, Dan River; 10, Deep River. (B) Northern Newark basin: 1, Roseland Quarry; 2, Vreeland Quarry; 3, Shrump Quarry. (C) Deerfield basin and the northern three- quarters of the Hartford basin: 4, Portland brownstone quarries; 5, Wethersfield Cove; 6, Di- nosaur Footprint Reservation; 7, Dickinson Quarry; 8, Moody Homestead; 9, Montague City; 10, localities near the Connecticut River near Turners Falls (from west to east: near basalt in Turners Falls; “Turners Falls”; old ferry landing, Gill; Howland’s Farm, Gill; Field’s Orchard, Gill; Lily Pond Quarry, Gill). (D) North-central part of Minas subbasin, Fundy basin, Five Islands region: 11, McKay Head; 12, Blue Sac. Details of localities for specific specimens are in the caption for figure 19.3 and in table 19.1. 316 • Paul E. Olsen and Emma C. Rainforth tracks with hands. Despite having been named more preserved material allows the trackmaker to be rec- than 150 years ago and widely cited, the type material ognized as an early ornithischian and reveals details of of the species included in the genus has never been locomotory and other aspects of behavior that have attributed or described properly, nor has the wealth of not been demonstrated by the analysis of ornithischian remarkably detailed material been appreciated. Here skeletal material alone. we describe and figure the type material of all eastern North American species along with key additional ma- Material and Methods terials from other localities, placing them all in strati- graphic and temporal context (figure 19.3). We also Virtually all classic Newark Supergroup taxa have a compare the eastern North American forms with very confused history, resulting in a nomenclatural material from varied locales showing the global distri- quagmire that has remained to the present day (Olsen, bution of the ichnotaxon. The large amount of well- Smith, and McDonald 1998). Indeed, characteristic of . Stratigraphic sections of the Deerfield, Hartford, Newark, and Fundy basins, expressed on a common age scale for the Jurassic age strata, showing the position of the localities discussed in the text; A, Rockaway River, Boonton, New Jersey; B, Roseland Quarry, Roseland, New Jersey, and Vreeland Quarry, Towaco, New Jersey; C, Shrump Quarry, Pleasantdale (Roseland), New Jersey; D, Wethersfield Cove, Wethersfield, Massachusetts; E, Portland brownstone quarries, Portland, Connecticut; F, Moody Homestead and Dickinson Quarry, South Hadley, Massachusetts; G, Dinosaur Footprint Reservation, Holyoke, Massachusetts; H, Lily Pond Quarry, Gill, Massachusetts; I-J-K, Field’s Orchard and Howland’s Farm, Gill, Massachusetts, and Montague City on the old Boston Road; L, old ferry landing, Gill, Massachusetts; M, general “Turners Falls” localities; N, Turners Falls, near basalt, Turners Falls, Massachusetts; O, McKay Head and Blue Sac, Cumberland County, Nova Scotia. Note the difference in thickness scale between the basin sections and the change in timescales within the basin sections. The wavy line indicates unconformity, and the gap at H indicates possible duration of hiatus. (Stratigraphy based on Olsen 1997) Early Jurassic Ornithischian Dinosaurian Ichnogenus Anomoepus • 317 the classic Connecticut Valley taxa is massive confusion of homonyms and of objective and subjective syno- nyms that previously rendered the taxonomy indeci- pherable. Anomoepus is no exception. There fortu- nately is enough information in the early published literature and on specimen labels (including those carved into the specimen) to permit a massive stream- lining of nomenclature. The principal basis for this chapter is the Hitchcock Ichnological Collection at the Pratt Museum of Am- herst College (AC; Amherst, Massachusetts), consist- ing of natural molds and casts of footprints, mostly from the Deerfield and Hartford basins of the Newark . Supergroup. One Hartford basin track described here Definitions of measurements used in the text and in figure 19.19, based on the composite Anomoepus scambus is in the collection of the Montshire Museum of Sci- in figure 19.5 and the reconstructed skeleton in figure 19.33. ence (MM; Norwich, Vermont), consisting of the rem- nants of the collection of the Museum of Dartmouth College (Hanover, New Hampshire). Additional tracks Specimens in the Hitchcock Ichnological Collection are described from other parts of the Newark Super- presently are numbered according to the catalog of group, including the Newark and Fundy basins, and Edward Hitchcock’s son Charles H. Hitchcock (1865). these specimens reside in the collections of the Yale However, as described by Olsen, Smith, and McDonald Peabody Museum (YPM; New Haven, Connecticut), (1998), the specimen numbers were not consistent the American Museum of Natural History (AMNH; across E. Hitchcock’s publications. Numbers used New York City), Dinosaur State Park (DP; Rocky prior to 1858 are Arabic numerals usually preceded by Hill, Connecticut), Rutgers University (RU; New a “No.” carved into the rock (prefixed herein by “old Brunswick, New Jersey), the New Jersey State Museum no.”). Where the number used by E. Hitchcock (1858) (NJSM; Trenton, New Jersey), the Fundy Geological differs from that used by C. H. Hitchcock (1865), the Museum (FGM; Parrsboro, Nova Scotia), and the pri- former is distinguished herein by “1858” after the spec- vate collection of Eldon George (EG; Parrsboro, Nova imen number. In the numbering systems of both 1858 Scotia). Specimens in the Museum of Mount Holyoke and 1865, the specimen numbers were based on their College (MHM; South Hadley, Massachusetts) were location within the displays and consist of a fraction destroyed by fire in the early twentieth century. with the numerator corresponding to a wall, table, or Analog and digital photographic and drawing case and with the denominator indicating the speci- methods are the same as those described in Olsen, men number in that specific area. These different Smith, and McDonald (1998). Composite drawings are numbering schemes are important because they are shown as impressions of the right pes (figures 19.4 and the only unambiguous tool allowing individual speci- 19.5), prepared by digitally superimposing drawings of mens to be traced though the Hitchcocks’ tortuous successive tracks (with the opposite tracks reversed). nomenclatural maze in the absence of explicit draw- The elements of the tracks thought to be least affected ings or descriptions. by the processes of impression and preservation are emphasized. Conventions for quantitative measure- Osteometric Methods ments are shown in figure 19.4, as described by Olsen, Baird (1957) argued that the characters of tracks most Smith, and McDonald (1998). We follow the rules and useful for classification are those that reflect the oste- guidelines of the fourth edition of the International ology of the trackmaker, and this approach has proved Code of Zoological Nomenclature (ICZN; International very fruitful. The most important track landmarks lo- Commission on Zoological Nomenclature 1999) as ap- calizing osteological features are the pads of the digits, plied to ichnotaxa. Only citations in the
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