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. 2017 Oct;231(4):515-531.
doi: 10.1111/joa.12660. Epub 2017 Aug 1.

Myology of the forelimb of Majungasaurus crenatissimus (Theropoda, Abelisauridae) and the morphological consequences of extreme limb reduction

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Myology of the forelimb of Majungasaurus crenatissimus (Theropoda, Abelisauridae) and the morphological consequences of extreme limb reduction

Sara H Burch. J Anat. 2017 Oct.

Abstract

Forelimb reduction occurred independently in multiple lineages of theropod dinosaurs. Although tyrannosaurs are renowned for their tiny, two-fingered forelimbs, the degree of their reduction in length is surpassed by abelisaurids, which possess an unusual morphology distinct from that of other theropods. The forelimbs of abelisaurids are short but robust and exhibit numerous crests, tubercles, and scars that allow for inferences of muscle attachment sites. Phylogenetically based reconstructions of the musculature were used in combination with close examination of the osteology in the Malagasy abelisaurid Majungasaurus to create detailed muscle maps of the forelimbs, and patterns of the muscular and bony morphology were compared with those of extant tetrapods with reduced or vestigial limbs. The lever arms of muscles crossing the glenohumeral joint are shortened relative to the basal condition, reducing the torque of these muscles but increasing the excursion of the humerus. Fusion of the antebrachial muscles into a set of flexors and extensors is common in other tetrapods and occurred to some extent in Majungasaurus. However, the presence of tubercles on the antebrachial and manual elements of abelisaurids indicates that many of the individual distal muscles acting on the wrist and digits were retained. Majungasaurus shows some signs of the advanced stages of forelimb reduction preceding limb loss, while also exhibiting features suggesting that the forelimb was not completely functionless. The conformation of abelisaurid forelimb musculature was unique among theropods and further emphasizes the unusual morphology of the forelimbs in this clade.

Keywords: Archosauria; Theropoda; functional morphology; musculature; phylogenetic inference; vestigial structures.

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Figures

Figure 1
Figure 1
Reconstruction of articulated right scapulocoracoid and forelimb of Majungasaurus crenatissimus in lateral view. Model is composed of CT scans of FMNH PR 2836, right scapulocoracoid and humerus; UA 9860, left ulna (reversed); and FMNH PR 2836, left radius, metacarpals, and phalanges (reversed). H, humerus; MC I, metacarpal I; MC IV, metacarpal IV; P, phalanges; R, radius; SC, scapulocoracoid; U, ulna. Scale bar: 5 cm.
Figure 2
Figure 2
Myological reconstruction of the right scapulocoracoid (FMNH PR 2836) of Majungasaurus crenatissimus 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; P, pectoralis; RH, rhomboideus; SBC, subcoracoideus; SBS, subscapularis; SC, supracoracoideus; SCA, supracoracoideus accessorius; 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 Majungasaurus crenatissimus in anterior (A), medial (B), posterior (C), and lateral (D) views. Proposed muscle origins are indicated in red, proposed insertions in blue. The humerus is a composite reconstruction based on the right humerus of FMNH PR 2836 and the isolated humerus FMNH PR 2423. AN, anconeus; AR, abductor radialis; BB, biceps brachii; BR, brachialis; CB, coracobrachialis; DC, deltoideus clavicularis; DS, deltoideus scapularis; EA, epitrocheloanconeus; ECR, extensor carpi radialis; ECU, extensor carpi ulnaris; EDL, extensor digitorum longus; FCU, flexor carpi ulnaris; FDLS, flexor digitorum longus superficialis; LD, latissimus dorsi; P, pectoralis; PT, pronator teres; SBC, subcoracoideus; SBS, subscapularis; SC, supracoracoideus; SCA, supracoracoideus accessorius; 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 Majungasaurus crenatissimus in anterior (A), posterior (B), medial (C), and lateral (D) views. Proposed muscle origins are indicated in red, proposed insertions in blue. Radius is FMNH PR 2836, ulna is UA 9860 (reversed). Cross‐hatching indicates broken bone surface, shaded areas indicate matrix. AN, anconeus; APL, abuctor pollicis longus; AR, abductor radialis; BB, biceps brachii; BR, brachialis; EA, epitrocheloanconeus; ECR, extensor carpi radialis; FCU, flexor carpi ulnaris; FDLP, flexor digitorum longus profundus; PT, pronator teres; R, radius; SU, supinator; TB, triceps brachii; U, ulna. Scale bar: 5 cm.
Figure 5
Figure 5
Myological reconstruction of the manus (FMNH PR 2836) of Majungasaurus crenatissimus in dorsal (A) and ventral (B) views. Proposed muscle origins are indicated in red, proposed insertions in blue. Cross‐hatching indicates broken bone surface, shaded areas indicate matrix. APL, abductor pollicis longus; ECU, extensor carpi ulnaris; EDB, extensor digitorum brevis; EDL, extensor digitorum longus; FDB, flexor digitorum brevis; FDL, flexor digitorum longus; I, digit I; II, digit II; III, digit III; IV, digit IV. Scale bar: 1 cm.
Figure 6
Figure 6
Comparison of myological reconstructions of the shoulder in the early theropod Tawa hallae (A) and Majungasaurus crenatissimus (B). Muscles are labeled on Tawa and represented in the same color on Majungasaurus. AE, antebrachial extensors; BB, biceps brachii; CB, coracobrachialis; DC, deltoideus clavicularis; DS, deltoideus scapularis; HR, humeroradialis; LD, latissimus dorsi; LS, levator scapulae; SC, supracoracoideus; SHA, scapulohumeralis anterior; SHP, scapulohumeralis posterior; SS, serratus superficialis; TBL, triceps brachii lateralis; TBM, triceps brachii medialis; TBS, triceps brachii scapularis; TR, trapezius. Scale bars: 5 cm (for each taxon).
Figure 7
Figure 7
Patterns of manual reduction among tetrapods showing typical reduction of external digits (A–C), and the less common uniform digital reduction (D–F). All diagrams show a dorsal view of the right manus, and phalangeal formulae are given below each manus. Examples shown are the lepidosaur Hemiergis quadrilineata (A), the paleognathous bird Struthio camelus (B), the nonavian theropod Tyrannosaurus rex (C), the tortoise Testudo (D), the sauropod Diplodocus (E), and Majungasaurus crenatissimus (F). All diagrams except B, D, and E after Shapiro et al. (2007).

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