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. 2014 Sep;225(3):271-97.
doi: 10.1111/joa.12216. Epub 2014 Jul 12.

Complete forelimb myology of the basal theropod dinosaur Tawa hallae based on a novel robust muscle reconstruction method

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Complete forelimb myology of the basal theropod dinosaur Tawa hallae based on a novel robust muscle reconstruction method

Sara H Burch. J Anat. 2014 Sep.

Abstract

The forelimbs of nonavian theropod dinosaurs have been the subject of considerable study and speculation due to their varied morphology and role in the evolution of flight. Although many studies on the functional morphology of a limb require an understanding of its musculature, comparatively little is known about the forelimb myology of theropods and other bipedal dinosaurs. Previous phylogenetically based myological reconstructions have been limited to the shoulder, restricting their utility in analyses of whole-limb function. The antebrachial and manual musculature in particular have remained largely unstudied due to uncertain muscular homologies in archosaurs. Through analysis of the musculature of extant taxa in a robust statistical framework, this study presents new hypotheses of homology for the distal limb musculature of archosaurs and provides the first complete reconstruction of dinosaurian forelimb musculature, including the antebrachial and intrinsic manual muscles. Data on the forelimb myology of a broad sample of extant birds, crocodylians, lizards, and turtles were analyzed using maximum likelihood ancestral state reconstruction and examined together with the osteology of the early theropod Tawa hallae from the Late Triassic of New Mexico to formulate a complete plesiomorphic myology for the theropod forelimb. Comparisons with previous reconstructions show that the shoulder musculature of basal theropods is more similar to that of basal ornithischians and sauropodomorphs than to that of dromaeosaurids. Greater development of the supracoracoideus and deltoideus musculature in theropods over other bipedal dinosaurs correlates with stronger movements of the forelimb at the shoulder and an emphasis on apprehension of relatively large prey. This emphasis is further supported by the morphology of the antebrachium and the intrinsic manual musculature, which exhibit a high degree of excursion and a robust morphology well-suited for powerful digital flexion. The forelimb myology of Tawa established here helps infer the ancestral conformation of the forelimb musculature and the osteological correlates of major muscle groups in early theropods. These data are critical for investigations addressing questions relating to the evolution of specialized forelimb function across Theropoda.

Keywords: Archosauria; Theropoda; functional morphology; myology; phylogenetic inference.

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Figures

Figure 1
Figure 1
Consensus phylogeny of all extant taxa used in this analysis, based on the recent phylogenies of Livezey & Zusi (2007), Jetz et al. (2012), Conrad (2008), and Sterli (2010).
Figure 2
Figure 2
Myological reconstruction of the scapulocoracoid of Tawa hallae in lateral (A) and medial (B) views. Proposed muscle origins are indicated in red, proposed insertions are indicated in blue. BB, Biceps brachii; CB, Coracobrachialis; DC, Deltoideus clavicularis; DS, Deltoideus scapularis; LS, Levator scapulae; RH, Rhomboideus; SBC, Subcoracoideus; SBS, Subscapularis; SC, Supracoracoideus; SCA, Supracoracoideus accessorius; SHA, Scapulohumeralis anterior; SHP, Scapulohumeralis posterior; SP, Serratus profundus; SS, Serratus superficialis; TBS, Triceps brachii scapularis; TR, Trapezius. Scale bar: 5 cm.
Figure 3
Figure 3
Myological reconstruction of the humerus of Tawa hallae in lateral (A), posterior (B), anterior (C), and medial (D) views. Proposed muscle origins are indicated in red, proposed insertions are indicated in blue. AN, Anconeus; AR, Abductor radialis; BB, Biceps brachii; BR, Brachialis; CB, Coracobrachialis; DC, Deltoideus clavicularis; DS, Deltoideus scapularis; EA, Epitrochleoanconeus; ECR, Extensor carpi radialis; ECU, Extensor carpi ulnaris; EDL, Extensor digitorum longus; FCU, Flexor carpi ulnaris; FDLS, Flexor digitorum longus superficialis; HR, Humeroradialis; LD, Latissimus dorsi; P, Pectoralis; PA, Pronator accessorius; PT, Pronator teres; SBC, Subcoracoideus; SBS, Subscapularis; SC, Supracoracoideus; SCA, Supracoracoideus accessorius; SHA, Scapulohumeralis anterior; SHP, Scapulohumeralis posterior; SU, Supinator; TBL, Triceps brachii longus; TBM, Triceps brachii medialis. Scale bar: 5 cm.
Figure 4
Figure 4
Myological reconstruction of the antebrachium of Tawa hallae in anterior (A), posterior (B), lateral (C), and medial (D) views. Proposed muscle origins are indicated in red, proposed insertions are indicated in blue. AN, Anconeus; APL, Abductor pollicis longus; AR, Abductor radialis; BB, Biceps brachii; BR, Brachialis; EA, Epitrochleoanconeus; FDLP, Flexor digitorum longus profundus; HR, Humeroradialis; PA, Pronator accessorius; PQ, Pronator quadratus; PT, Pronator teres; R, Radius; SU, Supinator; TB, Triceps brachii; U, Ulna. Scale bar: 5 cm.
Figure 5
Figure 5
Myological reconstruction of the carpus and manus of Tawa hallae in dorsal (A) and ventral (B) views. Proposed muscle origins are indicated in red, proposed insertions are indicated in blue. APB, Abductor pollicis brevis; APL, Abductor pollicis longus; ADM, Abductor digiti minimi; ECR, Extensor carpi radialis; ECU, Extensor carpi ulnaris; EDB, Extensor digitorum brevis; EDBP, Extensor digitorum brevis profundus; EDBS, Extensor digitorum brevis superficialis; EDL, Extensor digitorum longus; FCU, Flexor carpi ulnaris; FDBP, Flexor digitorum brevis profundus; FDBS, Flexor digitorum brevis superficialis; FDL, Flexor digitorum longus; I, Digit I; II, Digit II; III, Digit III; IV, Digit IV; PQ, Pronator quadratus. Scale bar: 5 cm.
Figure 6
Figure 6
Myological reconstruction of the articulated pectoral girdle and forelimb of Tawa hallae in left lateral view with selected muscle attachment sites labeled. Proposed muscle origins are indicated in red, proposed insertions are indicated in blue. For muscle abbreviations see Figs 5 and text. Not to scale.
Figure 7
Figure 7
Comparison of published myological reconstructions of the shoulder in a generalized basal ornithischian (A, adapted from Maidment & Barrett, 2011), the basal sauropodomorph Saturnalia (B, adapted from Langer et al. 2007), the basal theropod Tawa (C), and the dromaeosaurid Saurornitholestes (D, adapted from Jasinoski et al. and TMP 88.121.39). Muscles are labeled on Tawa and represented in the same color on other taxa. AE, antebrachial extensors; BB, Biceps brachii; CB, Coracobrachialis; DC, Deltoideus clavicularis; DS, Deltoideus scapularis; HR, Humeroradialis; LD, Latissimus dorsi; SC, Supracoracoideus; SHA, Scapulohumeralis anterior; SHP, Scapulohumeralis posterior; TBM, Triceps brachii medialis; TBS, Triceps brachii caput scapulare. Scale bars: 5 cm.

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