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Fig 1.

Muscular, fleshy, bone or cartilage elements of the tongue in extant archosaurs and outgroups.

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Table 1.

Extant specimens and staining protocol applied.

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Fig 2.

DiceCT imaging of the head of Alligator mississippiensis.

Individual cross-sections (A-D) from cranial to caudal region with targeted muscular and bony elements colored and labeled. Abbreviations: bh, basihyal; cb, ceratobranchial; ephc, episterno-hyoid muscular complex; lar, larynx; mbm, M. branchiomandibularis; mhy, M. hyoglossus; mge, M. genioglossus; tg, tongue.

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Fig 3.

DiceCT imaging of the head of Dromaius novaehollandiae.

Individual cross-sections (A-D) from cranial to caudal region with targeted muscular and bony elements colored and labeled. Abbreviations: lar, larynx; mbm, M. branchiomandibularis; mcg, M. ceratoglossus; mge, M. genioglossus; mhys, M. hypoglossus; msh, M. stylohyoideus and M. serpihyoideus; mtl, M. tracheolateralis.

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Fig 4.

DiceCT imaging of the head of Nothoprocta perdicaria.

Individual cross-sections (A-D) from cranial to caudal region with targeted muscular and bony elements colored and labeled. Abbreviations: lar, larynx; mbm, M. branchiomandibularis; mcg, M. ceratoglossus; mge, M. genioglossus; mhys, M. hypoglossus; msh, M. stylohyoideus and M. serpihyoideus; mtl, M. tracheolateralis; md, mandible.

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Fig 5.

DiceCT imaging of the head of Phasianus colchicus.

Individual cross-sections (A-D) from cranial to caudal region provided with targeted muscular and bony elements colored and labeled. Abbreviations: bh, basihyal; cb, ceratobranchial; ep, epibranchial; mbm, M. branchiomandibularis; mcg, M. ceratoglossus; mch, M. cricohyoideus; msh, M. stylohyoideus and M. serpihyoideus; md, mandible.

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Fig 6.

Hyoid remains in extinct archosaurs.

(A)Alligator prenasalis, pterosaurs (B)Liaoxipterus brachycephalus and (C)Ludodactylus sibbicki, (D)basal ornithischian Jeholosaurus shangyuanensis, (E)tyrannosaur Yutyrannus huali (F) Sinosauropteryx prima.

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Fig 7.

Hyoid remains in paravians.

(A) Microraptor zhaoianus, (B)Confuciusornis sp., (C)Enantiornithine sp., (D)Hongshanornis longicresta. The blue arrow indicates the ossified basihyal in Confuciusornis and Hongshanornis; it was also observed in one specimen of Microraptor. The green arrow indicates the phylogenetically earliest epibranchial [15]. See S1 Table for specimen numbers and supplemental references for published specimens.

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Table 2.

Scanning parameters for extant specimens.

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Table 2 Expand

Fig 8.

The evolution of the tongue in Archosauria.

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Fig 9.

Episterno-hyoid and M. coracohyoideus muscle in Alligator mississippiensis and proposed homologous muscles in Aythya americana.

(A)(B)Alligator mississippiensis (TMM M-12053), (C)(D)the homologous M. sternotrachealis in Aythya americana (TMM M-12682). Abbreviations: ephc, episterno-hyoid complex; epis or inter, episternum or the interclavicle; fur-furcula; mclh, M. cleidohyoideus (in Aythya); mcoh, M. coracohyoideus (in Alligator); tra, trachea.

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Fig 10.

The M. branchiomandubularis (arrows) in birds and Alligator.

(A)Rhea americana (USNM 615363), (B)Dromaius novaehollandiae (TMM M-14235), (C) Nothura maculosa (USNM 631209), (D)Megapodius pritchardii (USNM 319640), (E) Phasianus colchicus (TMM M-12000), (F)Alligator mississippiensis (TNHC specimen). The two portions of the muscle are indicated in Galliformes: ‘M. branchiomandubularis cranialis’ (mbm_a) and ‘caudalis’ (mbm_p).

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Fig 11.

M. intermandibularis cranialis in birds and Alligator.

(A)Rhea americana (USNM 615363), (B)Nothura maculosa (USNM 631209), (C)Dromaius novaehollandiae (TMM M-14235), (D)Struthio camelus (TMM M-14237), (E)Alligator mississippiensis (TNHC specimen) and (F)Phasianus colchicus (TMM M-12000). The muscle was indicated by black arrow.

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Fig 12.

Episterno-hyoid and M. coracohyoideus muscle in Alligator and proposed homologous muscles in birds.

(A)(B)Alligator mississippiensis (TMM M-12053), (C)(D)the homologous M. sternotrachealis in Aythya americana (TMM M-12682). (E)(F) show the M. tracheolateralis in Aythya americana (USNM 643741) and Rhea americana (USNM 615363) respectively; (G)(H)(I) show M. cricohyoideus in Nothura machulosa (USNM 631210), Megapodius pritchardii (USNM 319640) and Aythya americana (USNM 643741) respectively Abbreviations: ephc, episterno-hyoid complex; mch, M. cricohyoideus; mclh, M. cleidohyoideus; mtl, M. tracheolateralis, and mstt, M. sternotrachealis.

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Fig 13.

The position of the laryngeal opening in birds and Alligator.

Laryngeal opening is indicated by the red circle. (A)Alligator mississippiensis (TNHC specimen), (B)Dromaius novaehollandiae (TMM M-14236); (C)Nothura maculosa (USNM 631210), (D)Phasianus colchicums (TMM M-12001).

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