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Article Redescription And Journal of Vertebrate Paleontology 29(1):58–79, March 2009 # 2009 by the Society of Vertebrate Paleontology ARTICLE REDESCRIPTION AND PHYLOGENETIC RELATIONSHIPS OF DOSWELLIA KALTENBACHI (DIAPSIDA: ARCHOSAURIFORMES) FROM THE UPPER TRIASSIC OF VIRGINIA DAVID DILKES*,1 and HANS-DIETER SUES2 1Department of Biology & Microbiology, University of Wisconsin Oshkosh, Oshkosh, Wisconsin 54901, U.S.A., [email protected]; 2National Museum of Natural History, Smithsonian Institution, MRC 106, P. O. Box 37012, Washington, DC 20013-7012, U.S.A. ABSTRACT—A detailed redescription of the Late Triassic archosauromorph reptile Doswellia kaltenbachi Weems, 1980 from the Poor Farm Member of the Falling Creek Formation in the Taylorsville basin (Newark Supergroup) in Virginia is presented based upon additional preparation of the holotype. The euryapsid skull has a distinctive occiput with a prominent supraoccipital process that is flanked by posterior “horn-like” projections of the squamosals. Postfrontals, tabulars, and postparietals are absent. Plesiomorphic features of the palate and braincase include a plate-like horizontal parabasisphenoid, a pair of foramina for the internal carotid arteries on the ventral surface of the basisphenoid, and two fields of teeth on the palatal surface of the pterygoid. A sharp angle along the cervical and anterior dorsal ribs clearly separates the dorsal and lateral sides of the neck and anterior thoracic region. The posterior thoracic region has shorter ribs that project laterally with only a slight curvature. The ilium has a laterally deflected blade with numerous deep grooves along its distal edge. The laterally extensive set of osteoderms includes a nuchal element that is composed of several interlocking osteoderms that lack the arrangement in distinct transverse rows that characterizes the remainder of the osteoderms. A phylogenetic analysis of basal archosauriforms incorporates new data of Doswellia and the taxa Turfanosuchus, Yonghesuchus, and Qianosuchus that have not previously been combined in a single study. Results include a sister-group relationship between Doswellia and proterochampsids, placement of Qianosuchus as a crurotarsan archosaur, and Yonghesuchus and Turfanosuchus as successive sister taxa to Archosauria. INTRODUCTION scales. To date, tetrapods are represented only by unidentified archosaurian teeth and Doswellia (Weems, 1980). The initial breakup of the supercontinent Pangaea during the Based on palynological data, Cornet and Olsen (1990) Triassic commenced with rifting along the future long axis of the argued for a Carnian age for the fluviolacustrine Taylorsville Atlantic Ocean, from Greenland to Mexico (Olsen et al., 1989). basin sequence 1 (TVB1), which includes the unit from which This led to the formation of a long chain of rift basins in present- Doswellia is known. Huber et al. (1993) and Lucas and Huber day eastern North America. These basins became filled with (2003) assigned Doswellia to their Sanfordian Land Vertebrate thousands of meters of early Mesozoic sediments and, in many Faunachron (LVF) for eastern North America. Huber et al. cases, basalt flows that are collectively referred to as the Newark (1993) considered the Sanfordian LVF late Carnian (Tuvalian) in Supergroup. Although the sedimentary strata were long consid- age, but Lucas and Hunt (2003) extended the possible age range ered largely devoid of tetrapod skeletal remains, fieldwork in for this faunachron to ?early to late Carnian (Julian to early recent decades, especially in the southern rift basins, has Tuvalian) and dated the Falling Creek Formation as ?Julian. revealed a considerable diversity of Late Triassic amphibians, Long and Murry (1995) described and illustrated a number of reptiles, and synapsids. vertebrae and osteoderms as well as what appears to be a dis- Among the Triassic tetrapods described to date from the New- articulated carapace from the Dockum Group of Howard Coun- ark Supergroup, the distinctive, heavily armored reptile Doswel- ty, Texas, and referred them to Doswellia kaltenbachi. This lia kaltenbachi Weems, 1980 is noteworthy. Although referral appears justified because the Texas and Virginia speci- represented by much of the skeleton, the phylogenetic relation- mens share features such as a large articular facet and a dorsal ships of this taxon have remained uncertain because it combines eminence with many small pits on each osteoderm and elongat- a considerable number of autapomorphies with plesiomorphic ed, ventrally concave diapophyses on the dorsal vertebrae. To features and most of its appendicular skeleton is still unknown. date, the remains from the Dockum Group represent the only The type material of Doswellia kaltenbachi was collected from known record of Doswellia outside the Taylorsville basin. siltstones of the Upper Triassic (Carnian) Poor Farm Member or Additional preparation of the material of Doswellia kaltenba- the upper portion of the Falling Creek Formation (Doswell chi described by Weems (1980) has revealed new anatomical Group) in the Taylorsville basin of the Newark Supergroup in features of this distinctive reptile. In this paper we redescribe eastern Virginia (LeTourneau, 2003). The Poor Farm Member Doswellia in detail and provide a revised diagnosis. We also comprises gray, gray-green, dark gray, and black sandstone, silt- present a new phylogenetic analysis to assess the phylogenetic stone, calcareous sandstone and siltstone, and thin coaly layers position of Doswellia among basal archosauriforms and to test (LeTourneau, 2003). Its dark-gray and black siltstones have current hypotheses of archosauriform interrelationships. yielded well-preserved remains of a diverse flora (Cornet and Institutional Abbreviations—USNM, National Museum of Olsen, 1990), abundant casts of unionid shells, and isolated fish Natural History (formerly United States National Museum), Smithsonian Institution, Washington, DC. Anatomical Abbreviations—ac, rim for articular capsule of *Corresponding author. joint; af, articular facet; an, angular; ap, anterior projection; ar, 58 DILKES AND SUES—DOSWELLIA 59 articular; ax, axis; bo, basioccipital; bpt, basipterygoid process; and including at least five longitudinal rows on each side of bt, basal tuber of basioccipital; cap, capitulum; cpr, cultriform vertebral column in posterior dorsal region; ilium with laterally process; de, dorsal eminence; df, distal facet; dia, diapophysis; deflected dorsal blade. Doswellia is also distinguished by the ec, ectopterygoid; eo, exoccipital; f, frontal; fc, rim for fibrous following unique combination of features: prominent occipital capsule of joint; for, foramen; ica, foramen for cerebral branch peg of supraoccipital that projects over dorsal rim of foramen of internal carotid artery; i. tub., ischial tuberosity; j, jugal; mf, magnum; euryapsid construction of temporal region with en- metotic foramen; mx, maxilla; of, obturator foramen; op, larged jugal below supratemporal fenestra; absence of postpar- opisthotic; or. marg., orbital margin; p, parietal; pbs, parabasi- ietals, tabulars, and postfrontals; small elliptical supratemporal sphenoid; pal, palatine; para, parapophysis; po, postorbital; pt, fenestra that does not reach occipital margin; squamosals with pterygoid; p. tub., pubic tuberosity; q, quadrate; qj, quadratoju- posteriorly directed “horn-like” processes; elongate convex gal; rtp, retroarticular process; sa, surangular; so, supraoccipital; dorsal end of quadrate that fits into elongate ventral groove on sq, squamosal; st.f., supratemporal fenestra; su.an., suture for squamosal; step between the flat skull roof and temporal region; angular; su.ar., suture for articular; su.d., suture for dentary; su. absence of lateral mandibular fenestra; teeth with slender, coni- ec., suture for ectopterygoid; su.l., suture for lacrimal; su.m., cal crowns lacking carinae; three sacral ribs, the first derived suture for maxilla; su.po., suture for postorbital; su.pra., suture from dorsal region; pair of oval articular facets at distal tips of for prearticular; su.qj., suture for quadratojugal; su.sa., suture for first two caudal ribs. surangular; su.sq., suture for squamosal; tp, tooth puncture; tub, tuberculum; vrop, ventral ramus of opisthotic; vt, crista ventro- DESCRIPTION lateralis; XII, foramen for N. hypoglossus (XII). Skull The known cranial remains of Doswellia consist of the postor- SYSTEMATIC PALEONTOLOGY bital portion of a skull associated with a left angular, both sur- angulars, and a right articular (USNM 214823) and an isolated DIAPSIDA Osborn, 1903 right jugal (USNM 437574) from the Doswell locality. Weems ARCHOSAUROMORPHA Huene, 1946 (1980) reconstructed a long and tapering rostrum for Doswellia DOSWELLIIDAE Weems, 1980 based on a referred left dentary (USNM 186989) from the Ash- DOSWELLIA KALTENBACHI Weems, 1980 land locality. The dentary was found with isolated vertebrae and (Figs. 1–15) osteoderms that conform to those of the holotype of Doswellia Holotype—USNM 244214, “axial skeleton from seventh cer- kaltenbachi, and the teeth preserved in alveoli 3 and 4 closely vical through fifth caudal, scattered more posterior caudals, resemble a tooth in the left maxilla of USNM 214823 in the associated ribs; pelvis complete except for left ischium; clavicle, possession of slender, conical crowns without carinae. interclavicle; dorsal and lateral armor badly shattered except for As preserved, the postorbital region of USNM 214823 is broad an articulated patch from the posterior region” (Weems,
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