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Osteology and relationships of Revueltosaurus callenderi (Archosauria: Suchia) from the Upper Triassic (Norian) Chinle Formation of Petrified Forest National Park, Arizona, United States - PubMed

Review

. 2022 Oct;305(10):2353-2414.

doi: 10.1002/ar.24757. Epub 2021 Sep 29.

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Review

Osteology and relationships of Revueltosaurus callenderi (Archosauria: Suchia) from the Upper Triassic (Norian) Chinle Formation of Petrified Forest National Park, Arizona, United States

William G Parker et al. Anat Rec (Hoboken). 2022 Oct.

Abstract

Once known solely from dental material and thought to represent an early ornithischian dinosaur, the early-diverging pseudosuchian Revueltosaurus callenderi is described from a minimum of 12 skeletons from a monodominant bonebed in the upper part of the Chinle Formation of Arizona. This material includes nearly the entire skeleton and possesses a combination of plesiomorphic and derived character states that help clarify ingroup relationships within Pseudosuchia. A phylogenetic analysis recovers R. callenderi in a clade with Aetosauria and Acaenasuchus geoffreyi that is named Aetosauriformes. Key autapomorphies of R. callenderi include a skull that is longer than the femur, a complete carapace of dermal armor including paramedian and lateral rows, as well as ventral osteoderms, and a tail end sheathed in bone. Histology of the femur and associated osteoderms demonstrate that R. callenderi was slow growing and that the individuals from the bonebed were not young juveniles but had not ceased growing. A review of other material assigned to Revueltosaurus concludes that the genus cannot be adequately diagnosed based on the type materials of the three assigned species and that only R. callenderi can be confidently referred to Revueltosaurus.

Keywords: Aetosauriformes; Krzyzanowkisaurus; Pseudosuchia; histology; phylogeny.

© 2021 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy. This article has been contributed to by US Government employees and their work is in the public domain in the USA.

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Figures

FIGURE 1
FIGURE 1

Localities at Petrified Forest National Park containing Revueltosaurus callenderi (a) and stratigraphic distribution of those localities in the Chinle Formation (b). Stratigraphic column modified from Marsh et al. (2020). Numbers on the left side of the column are numeric ages in millions of years (Ma); those at the boundaries between members are inferred from the age models of Rasmussen et al. (2020), the smaller numbers with uncertainties are key individual U‐Pb zircon ages from the same study

FIGURE 2
FIGURE 2

Hypothesized evolutionary relationships of Revueltosaurus callenderi hypothesized by Nesbitt (, a), Butler et al. (, b), Ezcurra et al. ( supplement, c), Marsh et al. (, d), and the revised analysis in this study (e). A, Archosauria; Af, Aetosauriformes; P, Pseudosuchia; S, Suchia

FIGURE 3
FIGURE 3

Reconstruction of the skull and mandible of Revueltosaurus callenderi (a, b) based off of PEFO 34561 and PEFO 34089 and articulated skull and mandibles referred to R. callenderi (c, d, PEFO 38609) in dorsal (a, c), ventral (d), and lateral view (b). an, angular; ar, articular; bsp, basisphenoid; ch, ceratohyal; d, dentary; en, external naris; f, frontal; ift, infratemporal fenestra; j, jugal; l., left; la, lacrimal; lal, last alveolus; mx, maxilla; na, nasal; o, orbit; oo, otoccipital; pa, parietal; pf, prefrontal; pm, premaxilla; po, postorbital; pof, postfrontal; qj, quadratojugal; r., right; sa, surangular; so, supraoccipital; sp, splenial; sq, squamosal; stf, supratemporal fenestra; t, tooth. Dashed lines indicate a fault, which obscures the antorbital fenestra. Scale bars equal 5 cm

FIGURE 4
FIGURE 4

Skull elements of specimens referred to Revueltosaurus callenderi. Left nasal and premaxilla (a, b, PEFO 34561), left maxilla and lacrimal (c, d, PEFO 34274), right premaxilla and tooth (e, f, PEFO 34561), right maxilla and teeth (g, h, PEFO 34561), right frontal postfrontal, and parietal (i, j, PEFO 34561), left prefrontal (k, l), right jugal (m, n, PEFO 34561), right quadratojugal (o, p, PEFO 34561), right postorbital (q, r, PEFO 34561), right quadrate (s, t, PEFO 34561), and left squamosal (u, v, PEFO 34561) in lateral (a, c, e, g, i, k, m, q, u), medial (b, d, f, h, j, l, n, r, v), dorsal (i), ventral (j), posterior (s), and anterior view (t). a., articulates with; af, antorbital fenestra; aof, antorbital fossa; be, buccal emargination; cr, crest; ect, ectopterygoid; en, external naris; fo, fossa; gr, groove; idp, interdental plates; itf, infratemporal fenestra; j, jugal; kn, knob; la, lacrimal; mc, medial condyle; mx, maxilla; na, nasal; ofo, orbital fossa; om, orbital margin; pa, parietal; pf, prefrontal; pm, premaxilla; po, postorbital; pof, postfrontal; ppmx, palatal process of maxilla; ppo, paroccipital process of the otoccipital; pppm, palatal process of the premaxilla; q, quadrate; qf, quadrate foramen; qj, quadratojugal; r, ridge; sq, squamosal; s, sulcus; stf, supratemporal fenestra. Scale bars equal 1 cm, arrows point in anterior direction

FIGURE 5
FIGURE 5

Palatal and mandibular elements referred to Revueltosaurus callenderi (PEFO 34561). Right pterygoid (a), right ectopterygoid (b), left?palatine (c), left dentary (d), left surangular (e), left angular (f), left splenial (g), and left articular (h‐j). Ventral view (left) and dorsal view (right) is visible in a‐c and lateral view (left) and medial view (right) is visible in d‐f. Splenial (g) is in medial view. Articular is visible in lateral (h), dorsal (i), and ventral view (j). a. articulates with; an, angular; ap, anterior process; ar, articular; d, dentary; ect, ectopterygoid; emf, external mandibular fenestra; f, foramen; fo, fossa; gl, glenoid; ich, internal choana; j, jugal; lp, lateral process; mck, Meckelian canal; mp, medial process; ptg, pterygoid; r, ridge; rp, retroarticular process; sa, surangular; su, surangular; sy, symphysis. Scale bars equal 1 cm, arrows point in anterior direction

FIGURE 6
FIGURE 6

Braincase of specimens referred to Revueltosaurus callenderi (PEFO 34561). Parabasisphenoid (a‐d), right prootic (e, f), left otoccipital (g, h), basioccipital (i‐k), and reconstructed basicranium (l‐n) in lateral (a, e, k, l), medial (f), dorsal (b, i), ventral (c, j, n), anterior (g), and posterior view (d, h, m). a., articulates with; ab, alaris basisphenoidei; aip, anterior inferior process; bo, basioccipital; bpt, basipterygoid process; bt, basitubera; cp, cultriform process; crp, crista prootica; f, foramen; fm, foramen magnum; fo, fossa; hf, hypophyseal fossa; ic?, entrance of internal carotids?; ls, laterosphenoid; mcv, middle cerebral vein; mf, metotic foramen; mpr, median pharyngeal recess; mr, median ridge; np, notochoral pit; oc, occipital condyle; on, occipital neck; ot, otoccipital; pa, parietal; pbs, parabasisphenoid; pp, paroccipital process; pr, prootic; r, ridge; re, recess; so, supraoccipital; ssc, semicircular canal; ssr, subsellar recess; stg, stapedial groove; ve, vestibule; V, notch for trigeminal nerve; VI, canal for abducens nerve; VII, canal for facial nerve; VII, canal for vestibulocochlear nerve; XII, canal for hypoglossal nerve; vrop, ventral ramus of the otoccipital. Scale bars equal 1 cm, arrows point in anterior direction

FIGURE 7
FIGURE 7

Reconstruction of the skeletal anatomy of Revueltosaurus callenderi based off of PEFO 34561, PEFO 34089, and PEFO 33991 with osteoderms (a) and without osteoderms (b). Scale bars equal 10 cm

FIGURE 8
FIGURE 8

Axial skeleton of specimens referred to Revueltosaurus callenderi. Atlantal neurapophysis (a, PEFO 34561), atlantal pleurocentrum (b, PEFO 34561), atlantal intercentrum (c, PEFO 34561), axis pleurocentrum (d, PEFO 34561), anterior cervical vertebrae (e, PEFO 34833), posterior cervical vertebra (f, PEFO 34561), anterior trunk vertebra (g, PEFO 41387), last trunk vertebra (h, PEFO 34561), coossified sacral centra (i‐k, PEFO 36876), second sacral vertebra (l, PEFO 34269), anterior caudal vertebra (m, PEFO 34561), posterior caudal vertebra (n, PEFO 41387), cervical rib (o, PEFO 34561), trunk rib (p, PEFO 34561), and gastralium (q, PEFO 33847) in lateral (a, e, k, n, o, p), anterior (b), posterior (c, h), dorsal (i, l), and ventral view (j, q). Vertebrae in d, f, g, and m are shown in anterior, posterior, and lateral view from left to right. a., articulates with; ap, anterior process; ati, atlantal intercentrum; atp, atlantal pluerocentrum; axi, axial intercentrum; axp, axial pleurocentrum; ca, capitulum; ch, chevron facet; di, diapophysis; epi, epipophysis; k, keel; ns, neural spine; poz, postzygapophysis; pp, parapophysis; prz, prezygapophysis; r, ridge; sr, sacral rib; tp, transverse process; tu, tuberculum. Scale bars equal 1 cm, arrows point in anterior direction

FIGURE 9
FIGURE 9

Pectoral girdle and forelimb specimens referred to Revueltosaurus callenderi. Right scapula and coracoid (a, b, PEFO 34561), interclavicle (c, d, PEFO 34561), left humerus (e–h, PEFO 34561), right ulna (i‐l, PEFO 34561), right radius (m‐p, PEFO 34561), partial right manus (q, r, PEFO 33870), and isolated manual ungual (S, T, PEFO 34582) in lateral (a, t), medial (b), dorsal (d, q), ventral (c, r), anterior (e, j, n), posterior (f, i, m), proximal (g, k, o, s), and distal view (h, l, p). acp, acromion process; btu, biceps tubricle; cf, coracoid foramen; dc, distal carpals; dpc, deltopectoral crest; ef, ectepicondylar foramen; fo, fossa; g, glenoid; glr, glenoid rim; gr, groove; hh, humeral head; k, kink; ms, muscle scar; ol, olecranon; rat, radial tuber; rc, radial condyle; uc, ulnar condyle. Roman numerals indicate metacarpals. Scale bars equal 1 cm, arrows point in anterior direction

FIGURE 10
FIGURE 10

Pelvic girdle and hindlimb specimens referred to Revueltosaurus callenderi. Right ilium (a, PEFO 34561) and left ilium (b, PEFO 34561), right ischium (c, d, PEFO 36875), left pubis (e, f, PEFO 36875), left pubis and ischium (g, PEFO 34561), left femur (h‐k, PEFO 34561), left tibia (l‐o, PEFO 34273), and left fibula (p‐s, PEFO 34561) in lateral (a, c, e), medial (b, d, f), anterior (h, l, p), posterior (g, i, m, q), proximal (j, n, r), and distal view (k, o, s). a. articulates with; ace, acetabulum; alt, anterolateral tuber; amt, anteromedial tuber; ctf, crista tibiofibularis; ft, fourth trochanter; gr, groove; il, ilium; if, iliofibularis scar; ip, ischial peduncle; is, ischium; lc, lateral condyle; lt, lateral tuber; mc, medial condyle; of, obturator foramen; os, osdeoderm; pap, preacetabular process; pmt, posteromedial tuber; pop, postacetabular process; pu, pubis; pua, pubic apron; pup, pubic peduncle; r, ridge; sac, supraacetabuler crest; sr, sacral rib. Scale bars equal 1 cm, arrows point in anterior direction

FIGURE 11
FIGURE 11

Tarsal and metatarsal specimens referred to Revueltosaurus callenderi. Left astragalus[NS1] (a‐f PEFO 34561), left calcaneum (g‐l PEFO 34561), left? distal tarsal 4 (m‐o, PEFO 34561), right metatarsals I‐IV (p, q, PEFO 34561), and metatarsal V (p, PEFO 34269) in anterior (a, i), posterior (b, j), proximal (c, h, n), distal (d, g), lateral (e, k, m), medial (f, l, o), dorsal (p), and ventral view (q). 3, distal tarsal 3; 4, distal tarsal 4; a. articulates with; ah, anterior hollow; ca, calcaneum; ct, calcaneal tuber; p, peg; pg, posterior groove; fi, fibula; ti, tibia. Roman numerals indicate metatarsals. Scale bars equal 1 cm, arrows point in anterior direction

FIGURE 12
FIGURE 12

Dermal skeleton of specimens referred to Revueltosaurus callenderi. Articulated trunk paramedian osteoderms (a, PEFO 34561), four isolated trunk paramedian osteoderms (b, PEFO 34561), a trunk lateral osteoderm (c, PEFO 34561), three caudal paramedian osteoderms (d, PEFO 34561), articulated ventral osteoderms (e, PEFO 34269), articulated distal tail osteoderms (f, PEFO 33850; g, PEFO 38669), and articulated dorsal carapace (h, PEFO 42442/UWBM 116869). Dorsal view overlies ventral view in b, c, and d. ab, anterior bar; amp, anteromedial projection; ap, anterior projection; cs, caudal sheath; de, dorsal eminence; lr, lateral osteoderm; pm, paramedian osteoderm; ve, ventral osteoderm. Scale bars equal 1 cm, arrows point in anterior direction

FIGURE 13
FIGURE 13

Osteohistology of Revueltosaurus callenderi (a‐f, i) and Krzyzanowskisaurus hunti (g‐h, j). Left femur of R. callenderi (PEFO 33843) in posterior view (a) and transverse section (b‐d), paramedian osteoderm of R. callenderi (PEFO 33940a) in dorsal view (E) and longitudinal section (f, i), and paramedian osteoderm of K. hunti (UCMP 175058) in dorsal view (g) and longitudinal section (h, j). Dashed lines in a, e, and g indicate plane of histological section. Black arrows in d, i, and j indicate growth marks, white arrows in d indicate possible growth marks, and gray arrow in j indicates resorption line. All histological photographs are imaged in plane light except for the right side of c (purple hues), which is cross‐polarized with a quartz plate, and the left side of i (black background), which is cross‐polarized. Surrounding epoxy has been cropped out for clarity; see Morphobank Project P620 for original full resolution histological images. Specimen in g was coated with ammonium chloride sublimate for clarity. An, anterior; Do, dorsal; and Me, medial

FIGURE 14
FIGURE 14

Select elements of specimens referred to?Krzyzanowskisaurus hunti from the Blue Hills, AZ. Isolated tooth coated in ammonium chloride sublimate for clarity (a, b, UCMP 139573), left parietal (c, UCMP 195227), left postorbital (d, UCMP 195217), right quadratojugal (e, UCMP 194219), right quadrate (f, UCMP 165206), left squamosal (g, UCMP 165205), distal end of left humerus (h, UCMP 165207), trunk paramedian osteoderm (i, UCMP 175055), and caudal paramedian osteoderm (j, UCMP 175064) in lingual (a), labial (b), dorsal (c, i, j), lateral (d, e, g), and posterior view (f, h). Abbreviations: a., articulates with; ab, anterior bar; ci, “cingulum”; de, dorsal eminence; ef, ectepicondylar foramen; fo, fossa; itf, infratemporal fenestra; j, jugal; mc, medial condyle; om, orbital margin; pof, postfrontal; sq, squamosal; stf, supratemporal fenestra. Scale bars equal 1 cm, arrows point in anterior direction

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References

    1. Arcucci, A. , Previtera, E. , & Mancuso, A. C. (2019). Ecomorphology and bone microstructure of Proterochampsia from the Chañares Formation. Acta Palaeontologica Polonica, 64, 157–170.
    1. Ash, S. R. (2005). Synopsis of the Upper Triassic flora of Petrified Forest National Park and vicinity. Mesa Southwest Museum Bulletin, 9, 53–61.
    1. Atchley, S. C. , Nordt, L. C. , Dworkin, S. I. , Ramezani, J. , Parker, W. G. , Ash, S. R. , & Bowring, S. A. (2013). A linkage among Pangean tectonism, cyclic alluviation, climate change, and biologic turnover in the Late Triassic: The record from the Chinle Formation, southwestern United States. Journal of Sedimentary Research, 83, 1147–1161.
    1. Baird, I. L. (1970). The anatomy of the reptilian ear. In Gans C. & Parsons T. (Eds.), Biology of the Reptilia 2, morphology B (pp. 193–275). Academic Press.
    1. Baranyi, V. , Reichgelt, T. , Olsen, P. E. , Parker, W. G. , & Kürschner, S. (2018). Norian vegetation history and related environmental changes: New data from the Chinle Formation, Petrified Forest National Park (Arizona SW USA). Geological Society of America Bulletin, 130, 775–795.

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