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FIGURE 4 in Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)

FIGURE 4. Comparison of palatal and mandibular bones: Albertosaurus libratus vomer (A) in lateral view, ectopterygoid (ROM 1247, C) in rostroventral view, growth series of surangulars (ROM 1247, E; CMN 2120, F) in lateral view, angular (ROM 1247, H) in lateral view, and growth series of prearticulars (ROM 1247, J; CMN 2120, K) in medial view. Daspletosaurus torosus vomer (CMN 8506, B) in lateral view, ectopterygoid (CMN 8506, D) in rostroventral view, surangular (CMN 8506, G) in lateral view, angular (CMN 8506, I) in lateral view, and prearticular (CMN 8506, L, M) in medial view. Arrows indicate features discussed in text. Bones have been reversed to face right when required. Scale bar equals 50 mm.

FIGURE 3 in Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)

FIGURE 3. Growth series of Albertosaurus libratus craniofacial bones: lacrimals in lateral view (TMP 86.144.1, A; ROM 1247 B; AMNH 5336, C, D; CMN 2120, E); jugals in lateral view (ROM 1247, I; AMNH 5336, J; CMN 2120, K); and postorbitals in lateral view (TMP 86.144.1, M; AMNH 5664, N; AMNH 5336, O). Craniofacial bones of Daspletosaurus torosus'. lacrimals in lateral view (CMN 8506, F; TMP 85.62.1, G; CMN 11594, H); jugal in lateral view (FMNH PR308, L); and postorbitals in lateral view (CMN 11594, P; FMNH PR308, Q). Arrows indicate features discussed in text. Bones have been reversed to face right when required. Scale bar equals 50 mm; J is not to scale.

FIGURE 2 in Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)

FIGURE 2. Growth series of Albertosaurus libratus craniofacial bones: nasals in dorsal view (TMP 86.144.1, B; ROM 1247, C); maxillae in lateral (ROM 1247, E; AMNH 5336, F) and medial (CMN 12063, J; ROM 1247, I) views. Craniofacial bones of Daspletosaurus torosus'. premaxilla (CMN 8506, A); nasals (CMN 8506, D) in dorsal view; maxilla in lateral (CMN 8506, G; AMNH 5346, H) and medial (CMN 8506, K) views. Arrows indicate features discussed in text. Bones have been reversed to face right when required. Scale bar equals 50 mm; F is not to scale.

FIGURE 1 in Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)

FIGURE 1. Relative completeness of the skull and jaws of FMNH PR308 (Daspletosaurus torosus) in left lateral (A) and right lateral (B) views. Skull length is 1,050 mm.

FIGURE 8 in Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)

FIGURE 8. Comparison of Stage 1 (CMNH 7541) and Stage 4 (AMNH 5027, MOR 555) skulls of Tyrannosaurus rex in palatal (A, B), dorsal (C, D), lateral (E, F), caudal (G, H), and rostral (I, J) views. Numbered labels indicate T. rex autapomorphies: (1) nasal processes of the premaxillae tightly appressed throughout their entire length; (2) restricted exposure of the jugal within the antorbital fenestra; (3) antorbital fossa reaches the nasal suture caudodorsally; (4) transversely broad jugal pneumatic recess; (5) elongate frontal sagittal crest; (6) strongly divergent and short basal tubers; (7) rostroventrally-oriented caudal occipital plate; (8) shallow subcondylar recess; (9) rostroventrally deep basisphenoid plate and rostrocaudally-restricted basisphenoid recess; (10) inflated ectopterygoid; (11) strongly convex rostral plate of the surangular; (12) transversely narrow snout and broad temporal region relative to other tyrannosaurids; and (13) deep mandible relative to other tyrannosaurids.

Figure 22. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 22. A, origins and insertions of mm. intercristales (top, light-fill shapes), mm. intertransversarii (dark-fill shapes) and an interpretation of dorsal, rib head origins (large light-filled shapes) of m. iliocostalis capitis, of Tyrannosaurus rex (AMNH 5027). B, moment arms for lateroflexion by mm. intertransversarii on C6–C8 of Tyrannosaurus rex (BHI 3033, ventral view).

Figure 23. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 23. A, topological appearance of m. rectus capitis ventralis (anterior muscles) and m. iliocostalis cervicis (posteroventrally originating muscles) on anterior axial skeleton of Tyrannosaurus rex (BHI 3033), ventral view. The moment arm for lateral flexion by m. iliocostalis cervicis is superimposed. B, m. r.c.v.: origins of m. rectus capitis ventralis from ventral spinous processes of Tyrannosaurus rex (BHI 3033) with arrow showing course of the muscle. m. il. cerv.: origin from ventral centra and insertions onto ventral fascia of cervical ribs of m. iliocostalis cervicis, with arrow showing course of the muscle. C, insertion of m. rectus capitis ventralis onto basioccipital of Daspletosaurus torosus (CMN 8506; the specimen is incomplete and the image partly mirrored), showing moment arms. Because the tyrannosaurid occiput slopes anteroventrally, the ventroflexion moment arm is somewhat longer than depicted in this view. D, strength of functional inference for head ventroflexion by m. rectus capitis ventralis in Tyrannosaurus rex. E, strength of functional inference for neck lateroflexion by m. iliocostalis cervicis of Tyrannosaurus rex.

Figure 20. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 20. A, origin scar of m. splenius capitis from C2 of Tyrannosaurus rex (BHI 3033), in anterodorsal view. B, area of possible insertions of m. splenius capitis (medial part) on the occiput of Daspletosaurus torosus (CMN 8506; the specimen is incomplete and the image partly mirrored), with moment arms shown for dorsiflexion and lateroflexion. C, insertions of m. longus colli dorsalis/m. transversospinalis cervicis onto posterior and dorsal surfaces of epipophyses, from C2 to C5. The most prominent insertion is a posteriorly concave scar on the C2 epipophysis. D, centres of rotation (white circles) and moment arms (lines) for insertions of m. longus colli dorsalis/m. transversospinalis cervicis, on cervical vertebrae of Tyrannosaurus rex (BHI 3033). Centres of rotation are estimated to be at positions similar to those Selbie, Thomson & Richmond (1993) determined for intervertebral flexion in cats.

Figure 21. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 21. A, origin scars of m. longissimus capitis superficialis (C7–D1 parapophyses) and m. longissimus capitis profundus (C6–C3 parapophyses) of Tyrannosaurus rex (AMNH 5027). B, short moment arm of m. longissimus capitis superficialis for neck plus head dorsiflexion. C, paroccipital process insertions and moment arms of m. longissimus capitis superficialis on Daspletosaurus torosus (CMN 8506). D, basioccipital insertions and moment arms of m. longissimus capitis profundus on Daspletosaurus torosus (CMN 8506; the specimen is incomplete and the images are partly mirrored).

Figure 16 in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 16. Schematic diagram of major neck muscles of Tyrannosaurus rex in dorsal view. A–C represent successively deeper layers, and other conventions are as in Figure 15. M. complexus inserts dorsally on the squamosal, and m. iliocostalis capitis inserts along the ventral edge of the paroccipital process. M. longissimus capitis superficialis inserts between these on the lateral edge of the paroccipital process. In C a probable lateral part of m. spinalis capitis is signified by a '?'.

Figure 11. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 11. A, origins (dark shapes) and insertions (lighter outlined shapes) of mm. intertransversarii aponeuroses on posterior cervical vertebrae of Struthio camelus. Origins are from anterior faces of lateral tubercles, and insertions are onto posterior projections of the lateral tubercles. Arrows represent lines of action whereby insertions are drawn towards the origins to effect intervertebral lateroflexion. B, origins (dark) and insertions (lighter) of mm. inclusii on posterior cervical vertebrae of Struthio camelus. Origins are from anterior faces of the costal processes, and insertions are onto the lateral and dorsolateral tubercles. Arrows represent lines of action whereby insertions are drawn towards the origins. C, origins (dark-filled shapes) and insertions (light-filled shapes) of lateral portions of mm. intertransversarii in Caiman crocodylus. Arrows represent lines of action from origin to insertion, by which the muscles would lateroflex the anterior vertebra of each pair relative to the posterior one.

Figure 9. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 9. A, insertions of (dark grey) m. longus colli dorsalis pars cranialis onto anterior cervical epipophyses of Haliaeetus leucocephalus. B, schematic origin (light grey) of m. longus colli dorsalis pars cranialis, from cervicodorsal region of Struthio camelus. C, schematic origins (dark grey) of m. longus colli dorsalis pars cranialis from epipophyses (processes dorsales) of posterior cervicals in Struthio camelus. Slips from these origins coalasce with the main belly of the muscle, as demarcated by the light-shaded lines. The main belly of the muscle continues dorsally. D, schematic origin and insertions of m. longus colli dorsalis pars caudalis. This muscle subsystem originates from the cervicodorsal region and sends multiple bellies to insert on posterior cervical epipophyses. E, origins of m. longissimus cervicis/m interarticulares of Caiman crocodylus. The gradient-filled slips represent association of origins with the fascia surrounding the muscles. F, anteriormost insertions (dark grey) of m. longissimus cervicis/m. interarticulares, and m. transversospinalis cervicis, of Caiman crocodylus.

Figure 10. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 10. A, several muscles of Pelicanus occidentalis, disseted in lateral view. Abbreviations are as in the main text. Several slips of m. rectus capitis dorsalis (m. r.c.d.) converge ventrally, towards tendinous insertions on the basioccipital tuberosities. M. complexus has an unusual lateral, tendinous insertion. B, posterolateral view of m. longissimus capitis superficialis (outlined in dark grey) of Alligator mississippiensis. The posterior origin and the insertion are tendinous. M. iliocostallis capitis (m. il. cap.) is depicted, and m. constrictor colli has not been dissected away. The neck is slightly dorsiflexed in this view.

Figure 12. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 12. A, origins of m. rectus capitis lateralis (m. r.c.l.) and rectus capitis ventralis (m. r.c.v.) of Corvus brachyrhynchos, from C2 prosessus ventralis. B, all origins of m. r.c.v. from anterior cervicals of another specimen of Corvus brachyrhynchos, and its m. rectus capitis lateralis origin from C2. Both images are ventrolateral views. C, ventrolateral view of m. rectus capitis ventralis (with light outline) of Falco columbarius.

Figure 8. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 8. A, lateral view of m. longus colli dorsalis pars cranialis (m. l.c.d. cranialis), m. longus colli dorsalis pars caudalis (m. l.c.d. caudalis) and mm. intertransversarii (mm. intertrans.) of Cygnus columbianus. Note posterior slips contributing to m. l.c.d. cranialis, ventrally inserting heads of m. l.c.d. caudalis and multiple tendinous divisions of all mm. intertrans. B, left m. transversospinalis cervicis (m. trans. cerv., outlined in grey) of Caiman crocodylus, inserts by a white tendon onto C1. M. spinocapitis posticus (m. sp. cap. post.) and m. longissimus capitis superficialis (m. long. cap. sup., with part of surrounding fascia left on) are also outlined in grey. Superficial muscles have been removed.

Figure 7. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 7. A, dorsal view of m. biventer cervicis (m. biv. c., dark outline) and m. longus colli dorsalis pars cranialis of Aquila chrysaetos. Note the tendon intervening between anterior and posterior bellies of m. biv. c. M. complexus has been removed. B, dorsal view of m. transversospinalis capitis (m. trans. cap., dark outline) of Alligator mississippiensis. Medial and lateral portions of m. trans. cap. in Alligator are not distinguished here. C, lateral view of m. trans. cap. of Alligator mississippiensis (dark outline). D, m. complexus (dark outline) and m. rectus capitis lateralis (light outline) of Pelicanus occidentalis in lateral view. E, m. complexus of Pelicanus occidentalis, outlined in dark grey on the right. This is a dorsal view, with anterior towards the top. F, dorsal view of dissected Alligator mississippiensis, with m. epistrpheo-capitis lateralis, m. spinocapitis posticus and m. transversospinalis cervicis outlined in grey. M. transversospinalis capitis has been removed. G, m. splenius capitis of Anas platyrhynchos, from C2 to the occiput, outlined in grey, in posterior view. M. complexus and m. biventer cervicis have been removed. H, m. epistropheo-capitis medialis/m. altoïdius capitis of Caiman crocodylus, outlined in grey. M. transversospinalis capitis and m. epistropheo-capitis lateralis have been removed.

Figure 5 in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 5. Attachments of muscles inserting on the occiput of birds. A, posterior cervical vertebrae of Strutio camelus, showing osteological origin of posterior belly of m. biventer cervicis. B, anterior cervical vertebrae of Haliaeetus leucocephalus, with origins of m. complexus, m. splenius capitis and m. rectus capitis dorsalis (outlined). M. complexus originates from the epipophyses dorsally (as in Haliaeetus leucocephalus), and sometimes the lateral tubercles ventrally. For adjoining origins of m. complexus and m. rectus capitis dorsalis from the lateral tubercles, the latter is the anterior of each pair. C, occiput of Struthio camelus, depicting all insertions. In Struthio camelus and many other birds m. splenius capitis lateralis inserts laterally onto the occiput, but in other birds part of m. complexus inserts here.

Figure 4 in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 4. Comparison of major sites of muscle attachment on the necks of tyrannosaurids versus crocodilians and birds. A, cervical series of Caiman crocodylus (above; C1–C9) and Tyrannosaurus rex (below; C1–C10), depicting similarities of posterior transverse process morpohology. B, cervical series of Asio flamaeus (above) and Tyrannosaurus rex (below), depicting morphological similarities of epipophysis and the C2 neural spine. In all three groups the anterior transverse processes are smaller than their posterior counterparts (to show this clearly the C2 cerivical rib of T. rex is not pictured). Cervical rib morphology differs markedly among these archosaurs. The specimens are scaled to similar lengths from C1 to C9. The Tyrannosaurus rex is a composite reconstruction of C1 from Osborn (1905), BHI 3033 (C2) and AMNH 5027 (the remaining bones).

Figure 3 in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 3. Flow chart for extant behavioural interpolation, which enables inference of muscle-modulated behaviours in extinct animals. The certainty of behavioural inference in an extinct taxon is inversely related to the behaviour's specificity. Behaviour can be inferred by phylogenetic bracketing if similarity of muscle function is established in extant clades by kinematic and physiological considerations, and if these functions correlate with similar behaviours in the extant groups.

Figure 19. A in Functional morphology of neck musculature in the Tyrannosauridae (Dinosauria, Theropoda) as determined via a hierarchical inferential approach

Figure 19. A, origins of m. transversospinalis capitis (C2–C9), m. complexus (C2–C5) and m. splenius capitis from C2 and possibly C3, of Tyrannosaurus rex (BHI 3033). B, rugose scarring of m. transversospinalis capitis insertion on parietals of Tyrannosaurus rex (AMNH 5029). C, insertion of m. transversospinalis capitis onto parietals of Daspletosaurus torosus (CMN 8506; the specimen is incomplete and the image partly mirrored), with moment arms for lateral and dorsiflexion. D, insertions and moment arms for m. complexus (two dorsal) and m. iliocostalis capitis (ventral) on occiput of Daspletosaurus torosus (CMN 8506; the specimen is incomplete and the image partly mirrored for clarity).