The neck of flying archosaurs (Amniota, Reptilia) : description, reconstruction and biomechanics

Detalhes bibliográficos
Autor(a) principal: Oliveira, Richard Santos Buchmann de
Data de Publicação: 2022
Tipo de documento: Tese
Idioma: por
Título da fonte: Repositório Institucional da Universidade Federal do Espírito Santo (riUfes)
Texto Completo: http://repositorio.ufes.br/handle/10/16409
Resumo: Pterosaurs and birds are winged archosaurs that originated in the Mesozoic. Flight allowed representatives of both clades to explore previously vacant niches, and the differentiation of forelimbs into wings made the neck of these animals an active, functional limb used during foraging. Presently, birds have diversified and have necks of varying lengths, which are adapted to different life habits. However, the lack of extant descendants of pterosaurs creates gaps in the knowledge regarding the biology of this group, including its cervical anatomy and biomechanics. Here, we identify and describe the vertebral arrangement and influence of cervical soft tissues on the neck position at rest of extant birds. We then apply this data to establish and infer the neck position at rest and associated cervical soft tissues, and quantify the likely movements performed by the neck musculature of pterosaurs. For that end, we dissected the necks of specimens representing sixteen species of extant birds and used computed tomography scans of the cervical series of the pterosaurs Anhanguera piscator, Azhdarcho lancicollis and Rhamphorhynchus muensteri with three-dimensional preservation of the largest number of vertebral elements. We inferred the cervical muscles forces for pterosaurs by multiplying the thickness of the widest point of the muscle and its stress value. We found that the thickness of the intervertebral cartilage of extant birds varies along the neck and that excluding it can distort reconstructions of the neck of extinct animals. We also recognized sixteen muscles associated with the neck of extant birds. The vertebrae of extant archosaurs and pterosaurs showed evolutionary convergences that allowed us to reconstruct synovial cartilages in joints and ligaments in the latter. According to the angulation of the cervical vertebrae, the pterosaur neck probably had a slightly sinuous shape when in a rest position. Furthermore, we inferred thirteen cervical muscles in pterosaurs. We conclude that in pterosaurs the musculature responsible for the dorsoventral movements of the skull and neck was probably more robust and stronger, as in extant birds, and that less robust muscles were associated with stabilizing the neck or performing additional force for cervical movements
id UFES_ac167eadb7f67eaddf3255e25611770b
oai_identifier_str oai:repositorio.ufes.br:10/16409
network_acronym_str UFES
network_name_str Repositório Institucional da Universidade Federal do Espírito Santo (riUfes)
repository_id_str 2108
spelling Silva, Taissa Rodrigues Marques dahttps://orcid.org/0000-0001-7918-1358hhttp://lattes.cnpq.br/9562316044920852Oliveira, Richard Santos Buchmann dehttps://orcid.org/0000-0003-4428-9224http://lattes.cnpq.br/9978598888603093Nunes, Fabiana Rodrigues Costahttps://orcid.org/0000-0003-3596-0143http://lattes.cnpq.br/8396814023400187Montefeltro, Felipe Chinaglia https://orcid.org/0000-0001-6519-8546http://lattes.cnpq.br/9313332827151714Campos, Alexandre Liparinihttps://orcid.org/0000-0002-9967-2311http://lattes.cnpq.br/3860946535384466Pinheiro, Felipe Lima https://orcid.org/0000-0003-3354-914Xhttp://lattes.cnpq.br/15898745772256042024-05-30T01:40:58Z2024-05-30T01:40:58Z2022-11-29Pterosaurs and birds are winged archosaurs that originated in the Mesozoic. Flight allowed representatives of both clades to explore previously vacant niches, and the differentiation of forelimbs into wings made the neck of these animals an active, functional limb used during foraging. Presently, birds have diversified and have necks of varying lengths, which are adapted to different life habits. However, the lack of extant descendants of pterosaurs creates gaps in the knowledge regarding the biology of this group, including its cervical anatomy and biomechanics. Here, we identify and describe the vertebral arrangement and influence of cervical soft tissues on the neck position at rest of extant birds. We then apply this data to establish and infer the neck position at rest and associated cervical soft tissues, and quantify the likely movements performed by the neck musculature of pterosaurs. For that end, we dissected the necks of specimens representing sixteen species of extant birds and used computed tomography scans of the cervical series of the pterosaurs Anhanguera piscator, Azhdarcho lancicollis and Rhamphorhynchus muensteri with three-dimensional preservation of the largest number of vertebral elements. We inferred the cervical muscles forces for pterosaurs by multiplying the thickness of the widest point of the muscle and its stress value. We found that the thickness of the intervertebral cartilage of extant birds varies along the neck and that excluding it can distort reconstructions of the neck of extinct animals. We also recognized sixteen muscles associated with the neck of extant birds. The vertebrae of extant archosaurs and pterosaurs showed evolutionary convergences that allowed us to reconstruct synovial cartilages in joints and ligaments in the latter. According to the angulation of the cervical vertebrae, the pterosaur neck probably had a slightly sinuous shape when in a rest position. Furthermore, we inferred thirteen cervical muscles in pterosaurs. We conclude that in pterosaurs the musculature responsible for the dorsoventral movements of the skull and neck was probably more robust and stronger, as in extant birds, and that less robust muscles were associated with stabilizing the neck or performing additional force for cervical movementsPterossauros e aves são arcossauros alados que se originaram no Mesozóico. O voo permitiu que representantes de ambos os clados explorassem nichos anteriormente vagos, e a diferenciação dos membros anteriores em asas tornou o pescoço desses animais um membro ativo e funcional usado durante o forrageamento. Atualmente, as aves se diversificaram e possuem pescoços de comprimentos variados, adaptados a diferentes hábitos de vida. No entanto, a falta de descendentes existentes de pterossauros cria lacunas no conhecimento sobre a biologia desse grupo, incluindo sua anatomia cervical e biomecânica. Aqui, identificamos e descrevemos a disposição vertebral e a influência dos tecidos moles cervicais na posição do pescoço em repouso de aves viventes. Em seguida, aplicamos esses dados para estabelecer e inferir a posição do pescoço em repouso, reconstruir os tecidos moles cervicais e quantificar os prováveis movimentos realizados pela musculatura do pescoço dos pterossauros. Para tanto, dissecamos os pescoços de espécimes representando dezesseis espécies de aves viventes e utilizamos tomografias computadorizadas da série cervical dos pterossauros Anhanguera piscator, Azhdarcho lancicollis e Rhamphorhynchus muensteri com preservação tridimensional do maior número de elementos vertebrais. Inferimos as forças dos músculos cervicais de pterossauros multiplicando a espessura do ponto mais largo do músculo e seu valor de tensão. Descobrimos que a espessura da cartilagem intervertebral de aves viventes varia ao longo do pescoço e que a excluir pode distorcer as reconstruções do pescoço de animais extintos. Também reconhecemos dezesseis músculos associados ao pescoço de aves viventes. As vértebras de arcossauros viventes e pterossauros mostraram convergências evolutivas que nos permitiram reconstruir cartilagens sinoviais nas articulações e ligamentos neste último. De acordo com a angulação das vértebras cervicais, o pescoço dos pterossauros provavelmente tinha uma forma levemente sinuosa quando em posição de repouso. Além disso, inferimos treze músculos cervicais em pterossauros. Concluímos que em pterossauros a musculatura responsável pelos movimentos dorsoventrais do crânio e pescoço era provavelmente mais robusta e forte, como nas aves viventes, e que os músculos menos robustos estavam associados à estabilização do pescoço ou à realização de força adicional para os movimentos cervicaisConselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)Texthttp://repositorio.ufes.br/handle/10/16409porUniversidade Federal do Espírito SantoDoutorado em Biologia AnimalPrograma de Pós-Graduação em Ciências Biológicas (Biologia Animal)UFESBRCentro de Ciências Humanas e Naturaishttps://creativecommons.org/licenses/by/4.0/info:eu-repo/semantics/openAccessZoologiaPaleozoologiaAnatomiaVértebras cervicaisPterosauriaAvesAequorlitornithesMúsculos cervicaisLigamentos cervicaisCervical vertebraePterosauriaBirdsAequolitornithesCervical musclesCervical ligamentsThe neck of flying archosaurs (Amniota, Reptilia) : description, reconstruction and biomechanicsinfo:eu-repo/semantics/publishedVersioninfo:eu-repo/semantics/doctoralThesisreponame:Repositório Institucional da Universidade Federal do Espírito Santo (riUfes)instname:Universidade Federal do Espírito Santo (UFES)instacron:UFESrichardbuchmann@gmail.comORIGINALRichardSantosBuchmanndeOliveira-2022-teseapplication/pdf6234569http://repositorio.ufes.br/bitstreams/c6601972-ac14-4fcf-add3-cc9c9ce1f681/download0c798bc1a28508e6e83d5324cc61dd9fMD5110/164092024-09-25 17:13:57.007https://creativecommons.org/licenses/by/4.0/open accessoai:repositorio.ufes.br:10/16409http://repositorio.ufes.brRepositório InstitucionalPUBhttp://repositorio.ufes.br/oai/requestopendoar:21082024-10-15T17:54:43.481406Repositório Institucional da Universidade Federal do Espírito Santo (riUfes) - Universidade Federal do Espírito Santo (UFES)false
dc.title.none.fl_str_mv The neck of flying archosaurs (Amniota, Reptilia) : description, reconstruction and biomechanics
title The neck of flying archosaurs (Amniota, Reptilia) : description, reconstruction and biomechanics
spellingShingle The neck of flying archosaurs (Amniota, Reptilia) : description, reconstruction and biomechanics
Oliveira, Richard Santos Buchmann de
Zoologia
Paleozoologia
Anatomia
Vértebras cervicais
Pterosauria
Aves
Aequorlitornithes
Músculos cervicais
Ligamentos cervicais
Cervical vertebrae
Pterosauria
Birds
Aequolitornithes
Cervical muscles
Cervical ligaments
title_short The neck of flying archosaurs (Amniota, Reptilia) : description, reconstruction and biomechanics
title_full The neck of flying archosaurs (Amniota, Reptilia) : description, reconstruction and biomechanics
title_fullStr The neck of flying archosaurs (Amniota, Reptilia) : description, reconstruction and biomechanics
title_full_unstemmed The neck of flying archosaurs (Amniota, Reptilia) : description, reconstruction and biomechanics
title_sort The neck of flying archosaurs (Amniota, Reptilia) : description, reconstruction and biomechanics
author Oliveira, Richard Santos Buchmann de
author_facet Oliveira, Richard Santos Buchmann de
author_role author
dc.contributor.authorID.none.fl_str_mv https://orcid.org/0000-0003-4428-9224
dc.contributor.authorLattes.none.fl_str_mv http://lattes.cnpq.br/9978598888603093
dc.contributor.advisor1.fl_str_mv Silva, Taissa Rodrigues Marques da
dc.contributor.advisor1ID.fl_str_mv https://orcid.org/0000-0001-7918-1358
dc.contributor.advisor1Lattes.fl_str_mv hhttp://lattes.cnpq.br/9562316044920852
dc.contributor.author.fl_str_mv Oliveira, Richard Santos Buchmann de
dc.contributor.referee1.fl_str_mv Nunes, Fabiana Rodrigues Costa
dc.contributor.referee1ID.fl_str_mv https://orcid.org/0000-0003-3596-0143
dc.contributor.referee1Lattes.fl_str_mv http://lattes.cnpq.br/8396814023400187
dc.contributor.referee2.fl_str_mv Montefeltro, Felipe Chinaglia
dc.contributor.referee2ID.fl_str_mv https://orcid.org/0000-0001-6519-8546
dc.contributor.referee2Lattes.fl_str_mv http://lattes.cnpq.br/9313332827151714
dc.contributor.referee3.fl_str_mv Campos, Alexandre Liparini
dc.contributor.referee3ID.fl_str_mv https://orcid.org/0000-0002-9967-2311
dc.contributor.referee3Lattes.fl_str_mv http://lattes.cnpq.br/3860946535384466
dc.contributor.referee4.fl_str_mv Pinheiro, Felipe Lima
dc.contributor.referee4ID.fl_str_mv https://orcid.org/0000-0003-3354-914X
dc.contributor.referee4Lattes.fl_str_mv http://lattes.cnpq.br/1589874577225604
contributor_str_mv Silva, Taissa Rodrigues Marques da
Nunes, Fabiana Rodrigues Costa
Montefeltro, Felipe Chinaglia
Campos, Alexandre Liparini
Pinheiro, Felipe Lima
dc.subject.cnpq.fl_str_mv Zoologia
topic Zoologia
Paleozoologia
Anatomia
Vértebras cervicais
Pterosauria
Aves
Aequorlitornithes
Músculos cervicais
Ligamentos cervicais
Cervical vertebrae
Pterosauria
Birds
Aequolitornithes
Cervical muscles
Cervical ligaments
dc.subject.por.fl_str_mv Paleozoologia
Anatomia
Vértebras cervicais
Pterosauria
Aves
Aequorlitornithes
Músculos cervicais
Ligamentos cervicais
Cervical vertebrae
Pterosauria
Birds
Aequolitornithes
Cervical muscles
Cervical ligaments
description Pterosaurs and birds are winged archosaurs that originated in the Mesozoic. Flight allowed representatives of both clades to explore previously vacant niches, and the differentiation of forelimbs into wings made the neck of these animals an active, functional limb used during foraging. Presently, birds have diversified and have necks of varying lengths, which are adapted to different life habits. However, the lack of extant descendants of pterosaurs creates gaps in the knowledge regarding the biology of this group, including its cervical anatomy and biomechanics. Here, we identify and describe the vertebral arrangement and influence of cervical soft tissues on the neck position at rest of extant birds. We then apply this data to establish and infer the neck position at rest and associated cervical soft tissues, and quantify the likely movements performed by the neck musculature of pterosaurs. For that end, we dissected the necks of specimens representing sixteen species of extant birds and used computed tomography scans of the cervical series of the pterosaurs Anhanguera piscator, Azhdarcho lancicollis and Rhamphorhynchus muensteri with three-dimensional preservation of the largest number of vertebral elements. We inferred the cervical muscles forces for pterosaurs by multiplying the thickness of the widest point of the muscle and its stress value. We found that the thickness of the intervertebral cartilage of extant birds varies along the neck and that excluding it can distort reconstructions of the neck of extinct animals. We also recognized sixteen muscles associated with the neck of extant birds. The vertebrae of extant archosaurs and pterosaurs showed evolutionary convergences that allowed us to reconstruct synovial cartilages in joints and ligaments in the latter. According to the angulation of the cervical vertebrae, the pterosaur neck probably had a slightly sinuous shape when in a rest position. Furthermore, we inferred thirteen cervical muscles in pterosaurs. We conclude that in pterosaurs the musculature responsible for the dorsoventral movements of the skull and neck was probably more robust and stronger, as in extant birds, and that less robust muscles were associated with stabilizing the neck or performing additional force for cervical movements
publishDate 2022
dc.date.issued.fl_str_mv 2022-11-29
dc.date.accessioned.fl_str_mv 2024-05-30T01:40:58Z
dc.date.available.fl_str_mv 2024-05-30T01:40:58Z
dc.type.status.fl_str_mv info:eu-repo/semantics/publishedVersion
dc.type.driver.fl_str_mv info:eu-repo/semantics/doctoralThesis
format doctoralThesis
status_str publishedVersion
dc.identifier.uri.fl_str_mv http://repositorio.ufes.br/handle/10/16409
url http://repositorio.ufes.br/handle/10/16409
dc.language.iso.fl_str_mv por
language por
dc.rights.driver.fl_str_mv https://creativecommons.org/licenses/by/4.0/
info:eu-repo/semantics/openAccess
rights_invalid_str_mv https://creativecommons.org/licenses/by/4.0/
eu_rights_str_mv openAccess
dc.format.none.fl_str_mv Text
dc.publisher.none.fl_str_mv Universidade Federal do Espírito Santo
Doutorado em Biologia Animal
dc.publisher.program.fl_str_mv Programa de Pós-Graduação em Ciências Biológicas (Biologia Animal)
dc.publisher.initials.fl_str_mv UFES
dc.publisher.country.fl_str_mv BR
dc.publisher.department.fl_str_mv Centro de Ciências Humanas e Naturais
publisher.none.fl_str_mv Universidade Federal do Espírito Santo
Doutorado em Biologia Animal
dc.source.none.fl_str_mv reponame:Repositório Institucional da Universidade Federal do Espírito Santo (riUfes)
instname:Universidade Federal do Espírito Santo (UFES)
instacron:UFES
instname_str Universidade Federal do Espírito Santo (UFES)
instacron_str UFES
institution UFES
reponame_str Repositório Institucional da Universidade Federal do Espírito Santo (riUfes)
collection Repositório Institucional da Universidade Federal do Espírito Santo (riUfes)
bitstream.url.fl_str_mv http://repositorio.ufes.br/bitstreams/c6601972-ac14-4fcf-add3-cc9c9ce1f681/download
bitstream.checksum.fl_str_mv 0c798bc1a28508e6e83d5324cc61dd9f
bitstream.checksumAlgorithm.fl_str_mv MD5
repository.name.fl_str_mv Repositório Institucional da Universidade Federal do Espírito Santo (riUfes) - Universidade Federal do Espírito Santo (UFES)
repository.mail.fl_str_mv
_version_ 1813022523637366784