Estudo Da Biomecânica Craniana de Um Rauissuquídeo A Partir de Tomografias Computadorizadas E Técnicas de Imagens Digitais em 3 Dimensões

UNIVERSIDADE FEDERAL DO RIO GRANDE DO SUL
INSTITUTO DE GEOCIÊNCIAS
PROGRAMA DE PÓS-GRADUAÇÃO EM GEOCIÊNCIAS
ESTUDO DA BIOMECÂNICA CRANIANA DE UM

RAUISSUQUÍDEO A PARTIR DE TOMOGRAFIAS
COMPUTADORIZADAS E TÉCNICAS DE IMAGENS
DIGITAIS EM 3 DIMENSÕES
ALEXANDRE LIPARINI
ORIENTADOR – Prof. Dr. Cesar Leandro Schultz
Porto Alegre, 2008

UNIVERSIDADE FEDERAL DO RIO GRANDE DO SUL
INSTITUTO DE GEOCIÊNCIAS
PROGRAMA DE PÓS-GRADUAÇÃO EM GEOCIÊNCIAS
ESTUDO DA BIOMECÂNICA CRANIANA DE UM

RAUISSUQUÍDEO A PARTIR DE TOMOGRAFIAS
COMPUTADORIZADAS E TÉCNICAS DE IMAGENS
DIGITAIS EM 3 DIMENSÕES
ALEXANDRE LIPARINI
ORIENTADOR – Prof. Dr. Cesar Leandro Schultz
BANCA EXAMINADORA:
Prof. Dr. José Eduardo Figueiredo Dornelles – Universidade Federal de Pelotas
Prof. Dr. Richard Fariña – Universidad de la Republica, Uruguai
Prof. Dr. Sergio Alex Kugland Azevedo – Museu Nacional, Rio de Janeiro
Dissertação de Mestrado apresentada

como requisito parcial para a
obtenção do Título de Mestre em
Geociências.
Porto Alegre, 2008

Liparini, Alexandre
Estudo da biomecânica craniana de um Rauissuquídeo a partir de
tomografias computadorizadas e técnicas de imagens digitais em 3
dimensões. / Alexandre Liparini. - Porto Alegre : IGEO/UFRGS,
2008.
[122 f]. il.
Dissertação (Mestrado). - Universidade Federal do Rio Grande

do Sul. Instituto de Geociências. Programa de Pós-Graduação em
Geociências. Porto Alegre, RS - BR, 2008.
1. Archosauria. 2. Cinelismo craniano. 3. Crurotarsi. 4. Triássico.

I. Título.
_____________________________
Catalogação na Publicação
Biblioteca Geociências - UFRGS
Veleida Ana Blank CRB 10/571
DEDICATÓRIA
Dedico esta dissertação aos meus pais, Agostinho A. G. Campos e

Ângela L. Campos, que sempre me apoiaram e incentivaram independente do caminho
que eu escolhesse.
Dedico este trabalho também ao meu tio Antônio C. Liparini que,

inconscientemente ou não, me ajudou a escolher o (tal) caminho, me apresentando as
curiosidades da biologia (animal/vegetal/marinha) que marcaram a minha infância e
adolescência. Assim despertou um admirável interesse por esta área e uma paixão pela
ciência e pela pesquisa.
AGRADECIMENTOS
Tenho muito a agradecer e não será nada fácil colocar todos aqui nestas
folhas sem que eu me esqueça de alguém. Por isto, este primeiro agradecimento vai, de
coração, para todos aqueles que de alguma forma entraram na minha história, de
maneira mais ou menos influente, mas não menos importante, para fazer de mim o que
sou hoje.
MUITO OBRIGADO!
À minha querida e Sempre-Linda do Cerrado, Lorena C. Fleury,

companheira de mais de quatro anos de estrada, que descobriu comigo as dificuldades e
maravilhas de voar “sozinho” em um ambiente totalmente desconhecido.
Ao meu pai, Agostinho A. G. Campos, minha mãe Ângela Liparini
Campos e à mina irmã, também muito querida, Tânia Liparini Campos, essências para o
meu crescimento (para não dizer, a minha própria existência). Além de todos os meus
familiares Campos, Liparinis, Cândidos e Fleurys.
À Adamir Cunha (Tia Preta) e toda sua generosa família que me acolheu
e ajudou muito nas primeiras semanas de estada em Porto Alegre.
Aos eternos amigos de Belo Horizonte: das antigas (Late), como o Bruno
P. Ribeiro (Brunão) e a Fernanda de A. P. Rennó (Fubá); do meio (Middle), como o
Ivan B. Campos (Sorin), Fernando B. Campos (Bozão), Pablo Saldanha, Gabriel Lana e
Paula M. Bizzotto; e da nova leva (Early), como o Ivan Seixas, Tiago C. Silva
(Tiaguinho), Felipe Almeida (Vili), Artur Veloso e Rogério Oliveira. E aos mais novos
amigos de Porto Alegre como o Daniel C. Fortier (Tatá) e a Ana Emília Quezado
(Tatinha) que são do Ceará, Luiz A. Araújo e Janine Arruda, mineiros, Ednair
Nascimento, de Rondônia, a Elizete C. Holanda, que é do Acre e o Juan C. Cisneros da
Bahia (ops!) de El Salvador.
Aos belos dançarinos e amigos do grupo Sarandeiros, que fizeram parte
da minha vida por tanto tempo e com os quais aprendi uma nova maneira de ver e sentir
o Brasil.
Ao Leonardo Morato, conterrâneo, que, além de ter me ensinado muito
em seus cursos de extensão, me repassou valiosas dicas sobre a estrutura e
funcionamento de uma instituição que eu mal conhecia.
Aos participantes da banca examinadora, José Eduardo F. Dornelles,
Richard A. Fariña e Sérgio Alex K. Azevedo, e às suplentes, Mariana B. Soares e Ana
Maria Ribeiro, que prontamente aceitaram ler e contribuir com este trabalho.
Ao meu orientador, Cesar L. Schultz, pela realização eficiente e precisa
de seu papel como orientador, podando onde deveria ser podado, mas ao mesmo tempo
permitindo que eu caminhasse sobre meus próprios pés. Agradeço-o também pela
grande quantidade de informação repassada durante as suas aulas e discussões em
campo. Aproveito também para agradecer a três outros grandes orientadores, Jorge Luis
Pesquero, Carlos Henrique C. Moreira e Jadson C. Belchior, que contribuíram muito
para o meu crescimento como pesquisador.
Ao Roberto M. Pereira (Robertinho), pelo empenho e eficiência
administrativa, facilitando significativamente a resolução de questões burocráticas
referentes à bolsa de estudo e ao curso em si.
A todos os colegas de trabalho, alunos, professores e funcionários, do
Instituto de Geociências, especialmente aos colegas do Laboratório de Paleontologia de
Vertebrados e do Departamento de Paleontologia e Estratigrafia em geral.
À Coordenação de Aperfeiçoamento de Pessoal de Nível Superior
(CAPES), pela concessão da bolsa de estudos sem a qual seria impossível me manter na
cidade de Porto Alegre, e realizar os estudos com a dedicação a qual me dispus
inicialmente.
Enfim, mais uma vez, registro que agradeço imensamente a todos vocês
que participaram, participam e, ainda os que participarão da minha história,
conseqüentemente da história desta dissertação.
Obrigado!
RESUMO
Um esqueleto parcialmente completo e semi-articulado de um rauissuquídeo,

proveniente de níveis pertencentes à Cenozona de Therapsida (Mesotriássico) da
Formação Santa Maria, foi coletado em 2003, no município de Dona Francisca, RS,
Brasil. Neste espécime, os ossos do crânio e mandíbula estavam quase todos
desarticulados e puderam ser retirados isoladamente da rocha. Tal forma de preservação
possivelmente seria uma evidência de que os ossos do crânio deste animal poderiam
apresentar mobilidades entre si. Para verificar tal hipótese, foram utilizadas como
metodologias tomografias computadorizadas e programas específicos de manipulação
digital em três dimensões com o objetivo de criar modelos animados que permitissem o
estudo das possibilidades de movimento entre os ossos do crânio deste exemplar de
maneira isolada e integrada em todo o crânio. Como resultado, vídeos animados,
simulando as possibilidades de movimento entre os ossos do crânio foram gerados e,
para cada elemento craniano tomografado, um arquivo em 3D contendo o modelo de
sua superfície foi elaborado. Este material foi preparado para publicação em uma revista
eletrônica (i.e. Palaeontologia electronica) que permite a exposição e divulgação de
conteúdos digitais animados. Os resultados apresentados neste estudo indicaram um
número maior de evidências que favorecem um tipo passivo de cinetismo craniano,
sendo elas: - movimentos restritos, porém possíveis, entre vários ossos intracranianos; e,
- a presença de contatos frouxamente ligados, incapazes de realizar movimentos amplos
ou incapazes de realizar qualquer tipo de movimento quando o crânio como um todo é
considerado. Apesar disso, mais estudos referentes à reconstrução de tecidos moles e ao
desenvolvimento ontogenético de rauissuquídeos são necessários para se testar ou
falsear a hipótese de um possível modelo de cinetismo craniano ativo.
PALAVRAS-CHAVE: Archosauria; cinetismo craniano; Crurotarsi; Triássico.

ABSTRACT
Partially disarticulated skull bones of an unidentified rauisuchid were found in Dona

Francisca, south Brazil (Therapsid Cenozone, Mesotriassic). The skull bones and
mandible of this specimen were almost all disarticulated and could be isolated from the
rock separately. This peculiar type of preservation may probably be an evidence of
possible intracranial mobility. To verify this hypothesis, computer tomography (CT)
scan and specific 3D software methodologies were used to analyse suture morphology
and articulation areas of each individual pair of adjacent bones and for an integrated
mechanical model, which considered all bones that comprises the skull of the studied
specimen. As results, illustrative 3D animations for each discussed videos that
simulated possible mobility between skull bones were generated and, a 3D surface
model of each CT scanned skull element. Illustrative 3D animations for each discussed
movements were created and 3D surface model files for each CT scanned skull element
are presented. This material was prepared to be submitted in an electronic journal (i.e.
Palaeontologia Electronica), where animated digital documents could be published. As
results, this study presented more evidences in favour of passive cranial kinesis, which
includes: -restricted, but possible mobility between a considerable number of bones;
and, -presence of loosely contacts that would not be able to do extensive movements or
would present no mobility at all when the integrated model is considered. Though, more
studies on soft tissue reconstructions and ontogenetic development of rauisuchids are
needed to testify or falsify the hypothesis of an active cranial kinesis.
KEYWORDS: Archosauria; cranial kinesis; Crurotarsi; Triassic.

LISTA DE FIGURAS E TABELAS
FIGURA 1 - Principais pontos de articulação e eixos de rotação em um crânio de

Varanus salvator (Varanidae) em vistas dorsal, ventral e lateral
direita .......................................................................................................20
FIGURA 2 - Modelo mecânico integrando os pontos de cinetismo em um lagarto

(Varanus). O crânio está representado em vista lateral esquerda
durante as fases de repouso, protração e retração ....................................21
FIGURE 1 - Location map where UFRGS PV0629T was collected............................56
FIGURE 2 - UFRGS PV0629T reconstruction ............................................................57
FIGURE 3 - Illustration indicating original position of the bones in the fossil

when collected and prepared....................................................................58
FIGURE 4 - Prepared skull bones of UFRGS PV0629T .............................................59
FIGURE 5 - 3D images of the neurocranium/skull roof of PV0629T in posterior

view, laying on its right side, as founded in the rock, with probable
deformation sequence during the fossilization process ...........................60
FIGURE 6 - 3D images of the left premaxilla in lateral view as an example of the

different quality types of each generated 3D surface model....................61
FIGURE 7 - Correlations between the anatomical planes of symmetry and

coordinate planes using the left premaxilla as an example......................62
FIGURE 8 - Images comparing original shape of the prepared

Neurocranium/skull roof in anterior, left, posterior and dorsal views .....63
FIGURE 9 - Comparison of the original and retro-deformed shapes (in lateral and
ventral views) of the left quadrate/quadrato-jugal ...................................64
FIGURE 10 - Comparison of the original and retro-deformed shapes of the left

maxilla in anterior view ...........................................................................65
FIGURE 11 - Schematic illustration of some cranial bones in left lateral view and
their surrounding contacts........................................................................66
FIGURE 12 - 3D surface model of the left premaxilla ..................................................67
FIGURE 13 - 3D animated movie of the possibilities of mobility between the left

premaxilla and maxilla in left lateral view ..............................................68

premaxilla and the mirrored right nasal in left lateral view.....................69
FIGURE 15 - 3D surface model of the left maxilla (not retro-deformed)......................70

maxilla and the mirrored right nasal ........................................................71

maxilla and lacrimal/prefrontal................................................................72
FIGURE 18 - 3D animated movie of the mobility between the left maxilla and the
left jugal ...................................................................................................73
FIGURE 19 - 3D surface model of the right nasal .........................................................74
FIGURE 20 - 3D animated movie of the possibilities of mobility between the right

nasal and the right frontal, which is integrated to the
neurocranium/skull roof...........................................................................75
FIGURE 21 - 3D surface model of the left fused lacrimal/prefrontal............................76

lacrimal/prefrontal and the left frontal .....................................................77
FIGURE 23 - 3D surface model of the left jugal and ectopterygoid..............................78

jugal and the left postorbital, which is fused to the
neurocranium/skull roof unit....................................................................79

ectopterygoid and the left pterygoid ........................................................80
FIGURE 26 - 3D surface model of the left pterygoid ....................................................81

pterygoid and the pterygoid process of the basisphenoid ........................82

pterygoid and the left quadrate ................................................................83
FIGURE 29 - 3D surface model of the left palatine.......................................................84
FIGURE 30 - 3D surface model of the left fused quadrate/quadrato-jugal (not

retro-deformed) ........................................................................................85
FIGURE 31 - 3D animated movie of the possibilities of mobility between the

quadrate and squamosal ...........................................................................86
FIGURE 32 - 3D surface model of the fused neurocranium/skull roof/right

quadrate (not retro-deformed)..................................................................87
FIGURE 33 - 3D surface model of the anterior and posterior portions of the left
mandibular ramus.....................................................................................88
FIGURE 34 - 3D animated movie of the possibilities of translational mobility

between each mandibular ramus at the mandibular symphysis ...............89
FIGURE 35 - 3D animated movie of the possibilities of rotational mobility
between mandibular rami.........................................................................90
FIGURE 36 - 3D surface model of the neurocranium/skull roof and its contact with
the nasals, lacrimals/prefrontals, jugals/ectopterygoids, quadrates/
quadrato-jugals and pterygoids ................................................................91
FIGURE 37 - 3D animated movie of the possibilities of movement of the quadrate

unit in relation to the neurocranium/skull roof ........................................92
FIGURE 38 - 3D animated movie of the possibilities of movement of the pterygoid

unit in relation to the neurocranium/skull roof ........................................93

unit integrated with a fixed quadrate unit and without considering the
nasals........................................................................................................94

unit integrated with a fixed quadrate unit and considering the
mobility of the nasals ...............................................................................95
FIGURE 41 - 3D animated movie of the possibilities of movement at the

articulation site among the premaxilla, maxilla and nasal contact,
considering the premaxilla as an isolated unit not fixed to the nasal.......96
FIGURE 42 - 3D animated movie of the possibilities of movement at the

articulation site among the premaxilla, maxilla and nasal contact,
considering the premaxilla fixed to the nasal ..........................................97
FIGURE 43 - 3D surface model of the whole reconstructed skull of the UFRGS

PV0629T specimen..................................................................................98
FIGURE 44 - 3D animated movie of model 1 - possible intracranial mobility

considering the whole integrated mechanical model, where the
lacrimal/prefrontal-jugal and premaxilla-nasal contacts would
present no mobile sutures.........................................................................99

lacrimal/prefrontal-jugal contact would present no mobility and the
premaxilla-nasal contact would have a mobile articulation site ............100

lacrimal/prefrontal-frontal contact would present mobility and the
premaxilla-nasal contact would not have a mobile suture .....................101

lacrimal/prefrontal-jugal and the premaxilla-nasal articulation sites
would be mobile.....................................................................................102
TABLE 1 - Technical CT specification for each scanned skull element ..................104
TABLE 2 - Summary of the observed bone contacts and possible movements

described for each isolated element .......................................................105
TABLE 3 - Reference list for each described isolated element movement and its
illustrative animated figure ....................................................................106
TABLE 4 - List of bone contacts that presented or could present mobility in

isolated elements and whether the mobility remained or not in each
considered mechanical model ................................................................107
SUMÁRIO
TEXTO EXPLICATIVO DA ESTRUTURA DA DISSERTAÇÃO.........................13
1. INTRODUÇÃO .........................................................................................................15
1.1 OBJETIVOS .........................................................................................................16
1.2 ESTADO DA ARTE ............................................................................................17
1.2.1 Cinetismo craniano .....................................................................................17
1.2.2 Tomografias e imagens 3D .........................................................................22
1.2.3 Rauissuquídeos ............................................................................................22
1.3 MÉTODOS ...........................................................................................................23
1.4 ANÁLISE INTEGRADORA ...............................................................................24
1.5 REFERÊNCIAS BIBLIOGRÁFICAS .................................................................27
2. CORPO PRINCIPAL DA DISSERTAÇÃO...........................................................32

2.1 ARTIGO SUBMETIDO À REVISTA PALAEONTOLOGIA ELECTRONICA. .32
2.1.1 Abstract........................................................................................................33
2.1.2 Introduction .................................................................................................33
2.1.3 Material and methods .................................................................................36
2.1.4 Results and Discussion................................................................................38
2.1.4.1 Isolated Elements ..................................................................................39
2.1.4.1.1 Premaxilla......................................................................................39
2.1.4.1.2 Maxilla...........................................................................................40
2.1.4.1.3 Nasal ..............................................................................................40
2.1.4.1.4 Lacrimal/prefontal.........................................................................41
2.1.4.1.5 Jugal ..............................................................................................41
2.1.4.1.6 Ectopterygoid.................................................................................42
2.1.4.1.7 Pterygoid........................................................................................42
2.1.4.1.8 Palatine..........................................................................................43
2.1.4.1.9 Quadrate/quadrato-jugal...............................................................43
2.1.4.1.10 Neurocranium/skull roof..............................................................43
2.1.4.1.11 Mandible ......................................................................................44
2.1.4.2 Integrated Elements...............................................................................45
2.1.4.2.1 Quadrate unit.................................................................................45
2.1.4.2.2 Pterygoid unit ................................................................................46
2.1.4.3 Functional Inferences ............................................................................47
2.1.5 Conclusions ..................................................................................................49
2.1.6 Perspectives..................................................................................................49
2.1.7 Acknowledgements......................................................................................50
2.1.8 References ....................................................................................................51
2.1.9 Figures..........................................................................................................56
2.1.10 Tables .......................................................................................................103
2.1.11 Appendix A - Activation of non still figures .........................................108
2.1.11.1 3D surface models .............................................................................108
2.1.11.2 3D animated movies..........................................................................108
APÊNDICE A - DOCUMENTOS EM MEIO ELETRÔNICO ..............................109
ANEXO A - CARTA DE SUBMISSÃO DO ARTIGO ............................................111
ANEXO B - RESUMO EXPANDIDO NA I SEMANA ACADÊMICA EM 2006.112
ANEXO C - RESUMOS NA PALEO RS EM 2006..................................................118
ANEXO D - RESUMO NA XXIII JORNADAS ARGENTINAS EM 2007...........121

13
TEXTO EXPLICATIVO DA ESTRUTURA DA DISSERTAÇÃO
Este texto tem por objetivo explicar a estrutura da dissertação adotada

pelo autor e seu orientador, a qual seguiu rigorosamente as "Normas para a
Apresentação de Dissertações de Mestrado na Forma de Artigos" formuladas pelo
Programa de Pós-Graduação em Geociências da UFRGS. Para o leitor que não estiver
familiarizado com este modelo de dissertação (diferente do modelo tradicional), foi
disponibilizado para eventual consulta o arquivo contento as normas de formatação e
estruturação exigidas pelo Programa de Pós-Graduação em Geociências (Apêndice A).
Neste modelo de dissertação, o trabalho desenvolvido pelo autor durante
o período de seu curso é feito visando a produção de pelo menos um artigo para
publicação. Desta forma, os resultados e discussões obtidos ao longo de dois anos de
pesquisa são condensados e apresentados de forma clara e sucinta atendendo aos
objetivos de um texto elaborado para publicação. Como, normalmente, os textos na
forma de artigo estão associados a assuntos específicos, torna-se necessário um texto
integrador que permita ao leitor, que não está diretamente envolvido com o tema da
pesquisa, uma contextualização ao objeto de estudo, aos objetivos da pesquisa, às
metodologias adotadas pelo autor e seu orientador, e aos principais resultados obtidos.
Inicialmente, no primeiro capítulo, é apresentada uma breve introdução
ao tema na qual os principais assuntos abordados na dissertação são apresentados de
maneira generalizada. Nesta parte, o leitor terá um primeiro contato com as questões
levantadas pelo autor e seu orientador e um apanhado geral dos principais trabalhos já
publicados relacionados ao tema cinetismo craniano. Em seguida, são apresentados os
objetivos gerais e específicos que o autor se propôs a alcançar com a elaboração do
artigo. Na seção Estado da Arte é apresentado, de maneira mais detalhada e completa,
os principais temas abordados na introdução. Nesta parte, o leitor encontrará um número
maior de referências relativas aos grupos fósseis e atuais que apresentam cinetismo
craniano, aos principais tipos de cinetismo descritos, ao uso de tomografias na
paleontologia, além dos principais trabalhos que discutem e tentam definir
filogeneticamente o grupo dos rauissuquídeos. Após uma apresentação geral dos
métodos adotados e a justificativa de sua utilização, faz-se necessária uma análise
integradora, na qual alguns dos principais resultados são apresentados e discutidos, além
14
de serem discutidas também as possibilidades de estudos futuros que poderão ser

realizados com a continuidade de tal pesquisa.
O corpo principal da dissertação, contido no segundo capítulo deste
trabalho, é composto pela versão integral do artigo elaborado pelo autor e seu orientador
sobre o tema exposto no texto integrador. Tal artigo, por apresentar também resultados
em formatos específicos para visualização em meios eletrônicos, foi submetido à revista
eletrônica Palaeontologia Electronica, que permite a inclusão de elementos com
conteúdos ativos no trabalho submetido. Desta forma, animações, simulações, arquivos
3D ou vídeos produzidos para exemplificar os tipos de movimentos descritos neste
trabalho, além de sua parte textual, poderão ser visualizados em qualquer parte do globo
em que o acesso à internet esteja disponível. Tendo em vista que se trata de um
periódico internacional, no qual seus textos são publicados em inglês, tal artigo foi
redigido e será apresentado nesta dissertação neste idioma.
Em seguida, para que os leitores desta dissertação também possam
consultar e ter acesso ao material digital produzido, foi incluído um CD-ROM como
apêndice contendo os arquivos com conteúdos ativos apresentados no corpo principal da
dissertação. Além disso, foi disponibilizada, também neste CD, uma versão digital da
dissertação, na qual as figuras estão interligadas, na forma de hyperlinks, às suas
respectivas animações e aos seus arquivos 3D. Também estão incluídas janelas de
navegação no formato html que permitem ao leitor um acesso prático e rápido à versão
digital da dissertação, bem como de maneira independente às figuras de alta resolução,
tabelas, arquivos 3D e vídeos.
Finalmente, anexos como a carta de recebimento do artigo e outras
atividades relacionadas ao tema da dissertação, realizadas pelo autor durante o período
de elaboração deste estudo, também foram incluídos.
15
1. INTRODUÇÃO
A preservação excepcional e peculiar de um espécime de rauissuquídeo,

coletado no ano de 2003 no município de Dona Francisca, RS, Brasil gerou vários
questionamentos sobre seu modo de vida, causas de sua morte e processos envolvidos
na sua fossilização. Alguns destes aspectos vêm sendo estudados, isoladamente e em
conjunto, por alunos de Mestrado e Doutorado do Setor de Paleovertebrados da
UFRGS.
A presença, neste exemplar, de um crânio quase completo, mas
representado por elementos - quase todos - desarticulados, levou às seguintes perguntas:
- “Como estavam dispostos os ossos do crânio deste animal quando em vida?”; -
“Poderiam tais elementos apresentar mobilidades entre si?”; ou ainda: - “Como esta
eventual mobilidade teria sido utilizada pelo animal?”
A utilização de princípios mecânicos para explicar a resposta de
estruturas biológicas a determinados estímulos consiste no estudo da biomecânica. Sua
aplicação à paleontologia tem se mostrado eficiente, contribuindo com novas
interpretações comportamentais e morfofuncionais de grupos extintos.
Estudos relacionados à mobilidade intracraniana, ou cinetismo craniano,
são relativamente comuns para grupos de vertebrados viventes, tais como peixes,
lagartos (Frazzetta, 1962; Metzger, 2002), cobras, aves (Bock, 1964; Meekangvan et al.,
2006), e anfíbios [e.g. cecílias (Summers & Wake, 2005)]. Alguns destes estudos
apresentam importantes aspectos funcionais, que podem relacionar hábitos de vida
específicos com tipos determinados de mobilidade entre os ossos do crânio. A
possibilidade de cinetismo craniano também tem sido considerada em estudos
paleontológicos de grupos como dinossauros terópodes (McClelland 1993 apud
Rayfield 2005a; Mazzetta et al. 1998; Currie et al. 2003; Rayfield 2004, 2005a, 2005b),
ornitópodes (Norman & Weishampel 1985), aves do Cretáceo (Buhler, Martin &
Witmer, 1988), crocodilomorfos do Mesozóico (Walker, 1990) e, inclusive,
rauissuquídeos (Chatterjee, 1985; Gower, 1999).
O grupo dos rauissuquídeos (=Rauisuchia Bonaparte, 1982, sensu
Gower, 2000), devido ao escasso e fragmentário registro de seus espécimes, não
apresenta ainda uma relação filogenética consensual e, dependendo do autor, é
considerado como sendo mono-, para- ou polifilético (Gower, 2000). Tal grupo
16
compreende animais de médio e grande porte, predadores de topo de cadeia restritos ao

período Triássico. Morfologicamente – especialmente em relação ao crânio -
assemelham-se aos dinossauros terópodes, dominantes no Jurássico e no Cretáceo,
provavelmente por terem ocupado o mesmo nicho ecológico como predadores de
grande porte. Porém, como arcossauros crurotarsais, estariam filogeneticamente mais
proximamente relacionados ao grupo dos crocodilos.
Como parâmetros de comparação em relação ao espécime aqui estudado,
foram utilizados como representantes atuais o grupo dos lagartos e o das aves, pois são
os grupos viventes mais próximos filogeneticamente aos rauissuquídeos que apresentam
algum tipo de mobilidade intracraniana. Em relação aos grupos fósseis, foram
considerados para comparação, o grupo dos dinossauros terópodes por serem
morfologicamente parecidos aos rauissuquídeos e também apresentarem evidências de
cinetismo craniano, e as espécies Postosuchus kirkpatricki e Batrachotomus
kupferzellensis que compreenderiam, possivelmente, as espécies fósseis mais
proximamente relacionadas filogeneticamente ao espécime em questão, que também
apresentariam possíveis regiões de articulação intracraniana.
1.1 OBJETIVOS
Os objetivos gerais que guiaram a elaboração do presente estudo foram:

1) reconstituir digitalmente o crânio de um espécime de
rauissuquídeo;
2) verificar se há possibilidade de movimento entre os elementos
ósseos do crânio do espécime em estudo.
Como objetivos específicos são listados:
1) testar ferramentas computacionais de manipulação de objetos
digitais em 3 dimensões para reconstituições em
paleontologia;
2) descrever detalhadamente as regiões de contato entre pares de
ossos adjacentes do crânio, indicando a possibilidade ou não
de movimento entre eles;
17
3) gerar animações que simulem os possíveis movimentos

descritos;
4) analisar a possibilidade mecânica de realização integrada dos
movimentos descritos, considerando o crânio como uma
unidade.
1.2 ESTADO DA ARTE
1.2.1 Cinetismo craniano
A expressão cinetismo craniano se refere à presença de articulações

móveis entre elementos ósseos da caixa craniana e tem sido estudada para diversos
grupos de vertebrados viventes, incluindo peixes (Thomson, 1971), anfíbios (Summers
& Wake, 2005), mamíferos (Herring, 1972 apud Metzger, 2002; Jaslow & Biewener,
1995 apud Metzger, 2002; Herring & Teng, 2000 apud Metzger, 2002), aves (Bock,
1964; Meekangvan et al., 2006) e répteis (Frazzetta, 1962; Metzger, 2002), e também
para grupos fósseis como dinossauros terópodes (McClelland 1993 apud Rayfield
2005a; Mazzetta et al. 1998; Currie et al. 2003; Rayfield 2004, 2005a, 2005b),
ornitópodes (Norman & Weishampel 1985), aves do Cretáceo (Buhler, Martin &
Witmer, 1988), crocodilomorfos do Mesozóico (Walker, 1990) e rauissuquídeos
(Chatterjee, 1985; Gower, 1999). O termo “cinetismo craniano”, em alguns casos, é
aplicado indicando apenas a presença de articulações ou de possíveis suturas móveis
entre ossos de um crânio. Porém, tal termo também pode ser utilizado com a implicação
de um sentido funcional para tal mobilidade. Desta forma, animais que potencialmente
poderiam apresentar articulações ou suturas móveis entre ossos, podem ser definidos
como acinéticos caso tal mobilidade não seja observada durante uma atividade
comportamental (e.g. alimentação, disputa intra ou interespecífica).
Na paleontologia, com a impossibilidade de se observar diretamente o
funcionamento de um crânio cinético, bem como sua composição muscular, são feitas
inferências, através de observações morfológicas dos ossos, a partir das quais os limites
de mobilidade podem ser definidos. Isto não demonstra propriamente o tipo de
18
movimento realizado pelo animal quando em vida, mas sim as possibilidades de

movimentação que poderiam ser realizadas entre esses ossos. Rayfield (2004) utiliza,
para isto, termos como “mobilidade entre suturas”, “mobilidade craniana” ou
“flexibilidade craniana”, o que dá um aspecto menos funcional e mais descritivo ao tipo
de análise.
A realização de movimentos entre ossos do crânio tem sido estudada
desde meados do século XVIII (Hérrisant, 1748 apud Buhler, Martin & Witmer, 1988;
Nitzsch, 1816-1817 apud Buhler, Martin & Witmer, 1988r; Versulys, 1910), porém de
forma isolada e pouco expressiva. A partir de meados do século XX, estudos sobre
cinetismo craniano, principalmente em aves e lagartos, vêm sendo realizados de forma
mais sistemática e significativa (Hofer 1945; Frazzetta 1962; Bock 1964). Inicialmente,
tais estudos se focavam na identificação de características anatômicas e morfológicas
dos ossos cranianos relacionadas a possíveis pontos de mobilidade. Além disso, a
descrição detalhada da musculatura que compunha o crânio e suas prováveis relações
com o cinetismo craniano também era um objeto de estudo muito pesquisado até
meados de 1970. A partir desta época, com o desenvolvimento de novas metodologias,
como a cineradiografia e a eletromiografia, uma análise mais funcional sobre cinetismo
craniano pôde ser determinada (Smith, 1982; Trockmorton & Clarke, 1981; Herrel et al.
1999; Gussekloo & Bout, 2005a, 2005b; Metzger, 2002). Tais técnicas permitiram
acompanhar, passo a passo, a seqüência de realização de determinadas atividades, como
a alimentação in vivo, sendo possível associar cada etapa da atividade à disposição de
cada osso no crânio.
Tipos específicos de cinetismo craniano para vertebrados terrestres foram
definidos principalmente para lagartos e aves. De acordo com a revisão feita por
Metzger (2002), para lagartos, os termos utilizados para descrever os principais tipos de
cinetismo encontrados são: 1) metacinético; 2) mesocinético; e, 3) estreptostílico. Tais
termos são definidos pelo tipo de movimento que realizam e os ossos envolvidos em tal
ponto de articulação. Um crânio metacinético é definido pelo deslizamento curvilíneo
entre os ossos parietal e supraoccipital, sendo o eixo de rotação orientado
transversalmente e passando pelos processos paraoccipitais dos ossos exoccipitais do
crânio. O tipo mesocinético de cinetismo craniano esta ligado à flexão e extensão
dorsoventral ao redor do eixo transversal que corre ao longo da sutura fronto-parietal. A
estreptostilia é definida como a rotação da articulação dorsal do quadrado com o osso
esquamosal e/ou supratemporal. Nesta articulação o eixo de rotação se apresenta,
19
normalmente, orientado transversalmente. Além disso, o termo anficinético é utilizado

para se referir a crânios que sejam, simultaneamente, metacinéticos e mesocinéticos
(Metzger, 2002). A Figura 1 indica os pontos de articulação e eixos de rotação dos tipos
de cinetismo apresentados acima e a Figura 2 ilustra como seriam realizados os
movimentos de um crânio cinético segundo o modelo mecânico apresentado por
Frazzetta (1962).
Já para o grupo das aves, além de poderem apresentar um crânio
estreptostílico, algumas áreas de cinetismo são também relacionadas a regiões flexíveis
capazes de encurvar-se. Os dois principais tipos de cinetismo craniano reconhecidos
para as aves atuais são: 1) procinético; e 2) rincocinético, estando ambos localizados
anteriormente às órbitas. No tipo procinético, a região de curvatura envolve os ossos
nasal e pré-maxilar, próximo ao seu contato com o osso frontal, ao longo de um único
eixo transversal. Em um crânio rincocinético, a barra dorsal - composta pelo pré-maxilar
e pela porção medial do nasal - e a porção lateral do nasal apresentam duas regiões de
curvatura distintas.
20
FIGURA 1. Principais pontos de articulação e eixos de rotação em um crânio de

Varanus salvator (Varanidae) em vistas dorsal (A), ventral (B) e lateral direita (C). As
linhas tracejadas, os círculos marcados e as articulações (2b) indicam os eixos de
cinetismo. 1, eixo mesocinético; 2a, eixo metacinético; 2b, articulação metacinética
(supraoccipital desliza anteroposteriormente na região ventral do osso parietal como
indicado pelas setas); 3 eixo estreptostílico; 4, eixo hipocinético. bo, basioccipital; bp,
processo basipterigóide; bs, basiesfenóide; ect, ectopterigóide; ep, epipterigóide; fr,
frontal; jug, jugal; l, lacrimal; max, maxilar; n, nasal; pa, parietal; pal, palatino; pf, pré-
frontal; pmx, pré-maxilar; pop, processo paraoccipital; po+pof, pós-orbital+pós-frontal;
pro, proótico; pt, pterigóide; q, quadrado; s, stapes; sep, septomaxilar; soc,
supraoccipital; sq, esquamosal; stp, supratemporal; vom, vômer. Modificado de Rieppel
(1980). © Birkhäuser, Verlag.
21
FIGURA 2. Modelo mecânico integrando os pontos de cinetismo em um lagarto

(Varanus). O crânio está representado em vista lateral esquerda durante as fases de
repouso (A), protração (B) e retração (C). Região sombreada indica o segmento
occipital do crânio. As denominações muzzle, parietal, epipterygoid, basal e
quadratic são relativas às cinco unidades do segmento maxilar Sendo elas,
respectivamente, focinho, parietal, epipterigóide, basal e quadrado. Os círculos
marcados indicam a localização dos eixos de cinetismo. 1, eixo mesocinético; 2, eixo
estreptostílico; 3, eixo hipocinético. Modificado de Frazzetta (1962). © Wiley-Liss.
A utilização de termos iguais para se referir a tipos de cinetismo craniano

encontrado em grupos distintos (i.e. aves e lagartos) não indica, necessariamente, uma
homologia entre as partes. Smith (1982) sugere que, mesmo dentro do grupo dos
lagartos, a estreptostilia provavelmente tenha surgido independentemente mais de uma
vez. Tal fato poderia indicar, também, que apesar de tais movimentos serem realizados
de maneira muito similar, poderiam estar associados a funções distintas.
22
1.2.2 Tomografias e imagens 3D
A utilização e manipulação de imagens digitais têm sido facilitadas pela

maior acessibilidade a computadores com uma maior capacidade de processamento de
dados. Com isso, aumentou também a utilização de técnicas, como a tomografia
computadorizada, capazes de extrair, dos fósseis, informações anteriormente
inacessíveis (Ketcham & Carlson, 2001). É possível, por exemplo, saber como um fóssil
está disposto no interior de uma rocha para que sua preparação seja planejada. Ou então,
regiões internas, como cavidades nasal e craniana, podem ser acessadas e estudadas sem
que o material seja danificado. Reconstituições das cavidades cranianas feitas a partir de
imagens tomografadas podem apresentar importante fonte de dados para a análise de
características fisiológicas em grupos fósseis como mostrou Rodrigues (2005). Além
disso, reconstituições de peças fósseis em 3D têm sido utilizadas para se fazer retro-
deformações (Polcyn et al., 2005), simulações de encaixes e movimentos, além da
divulgação de material via world wide web (i.e. http://digimorph.org/).
Neste trabalho, foi escolhida a utilização desta metodologia para a
obtenção das imagens em 3 dimensões pelo grau de detalhamento que esta metodologia
poderia obter das imagens, não havendo perdas significativas de informações. Além
disso, a escolha foi favorecida também pelo contato já estabelecido entre o orientador
deste trabalho e o pessoal responsável pelo setor de tomografias do Hospital de Clínicas
de Porto Alegre, que facilitou o acesso e a realização das sessões.
1.2.3 Rauissuquídeos
O grupo dos rauissuquídeos compreende um grupo de arcossauros

crurotarsais exclusivamente triássicos que apresenta registros em todos os continentes
exceto Austrália e Antártica, porém de forma esparsa e fragmentária (Gower, 2000). A
sistemática deste grupo ainda é polêmica e muito discutida (Benton & Clark, 1988;
Sereno & Arcucci, 1990; Sereno, 1991; Parrish, 1993; Juul, 1994). O termo Rauisuchia
23
(Bonaparte, 1982), ou rauissuquídeos, tem sido empregado para se referir a um grupo

composto pelos táxons incluídos nas famílias Rauisuchidae, Prestosuchidae,
Poposauridae e Chatterjeeidae, independentemente de assumi-lo ou não como um grupo
monofilético (Chatterjee, 1985; Galton, 1985; Long & Murry, 1995).
Os rauissuquídeos compuseram a fauna de predadores de topo de cadeia
durante todo o período Triássico, tendo sido, no Jurássico, substituídos pelos
dinossauros terópodes, que apresentavam um aspecto morfológico geral bastante
semelhante àqueles, possivelmente devido à convergência de hábitos (Benton, 1984).
Apesar de existirem descrições para vários grupos de rauissuquídeos (para uma revisão,
consultar Gower, 2000), poucos trabalhos tratam da possibilidade de existirem
articulações ou estruturas móveis no crânio. Apenas Chatterjee (1985) apresenta um
estudo um pouco mais aprofundado sobre cinetismo craniano em um táxon pertencente
a este grupo (i.e., Postosuchus), enquanto Gower, em 1999, discute a possibilidade de
cinetismo craniano em um rauissuquídeo do Triássico Médio do sul da Alemanha (i.e.,
Batrachotomus kupferzellensis). Gower (2000) comenta a possibilidade de existir uma
articulação entre pré-maxilar e maxilar de rauissuquídeos e sugere maiores estudos
nesse sentido, a fim de refinar as descrições sobre esta região, o que poderá contribuir
também para a resolução da filogenia deste grupo.
1.3 MÉTODOS
O material craniano do espécime UFRGS PV0629T, depositado no

Laboratório de Paleontologia de Vertebrados da UFRGS, foi preparado mecanicamente
e, posteriormente, cada elemento ósseo foi fotografado separadamente, em alta
resolução, preservando o maior número de detalhes possível. A seguir, foram
selecionados ossos para serem submetidos a tomografias computadorizadas. Uma vez
que, nem sempre, estavam presentes (ou em bom estado) os ossos correspondentes das
duas metades do crânio, foram escolhidas apenas aquelas peças mais bem preservadas
para serem tomografadas. A obtenção das imagens tomografadas só foi possível com o
apoio do Hospital de Clínicas de Porto Alegre, RS, Brasil, que gentilmente cedeu alguns
horários de utilização do aparelho e técnicos responsáveis pela manipulação do
24
equipamento. A partir das imagens obtidas, foram gerados arquivos 3D com os modelos
das superfícies de cada peça tomografada.
Utilizando tanto o material fóssil preparado, quanto as imagens obtidas
através das tomografias, foi feito um estudo anatômico e morfológico detalhado das
suturas dos ossos do crânio a fim de se identificar possíveis regiões de mobilidade.
Identificadas tais regiões, foram feitas descrições sobre os tipos de movimentos capazes
de serem realizados entre cada par de ossos isoladamente sem a preocupação com
possíveis interferências de outros ossos do crânio. Só então, após listadas todas as
possibilidades de movimentos entre os pares de osso, é que o efeito do conjunto como
um todo foi analisada integrando-se a influência de um grupo cada vez maior de ossos
até se obter um modelo completo com o crânio totalmente integrado e reconstituído.
Para a reconstituição do crânio, foi utilizado o programa computacional
Maya 2008, específico para manipulação de arquivos 3D. Tal programa permite alinhar
e colocar em escala todos os arquivos 3D gerados na tomografia, gerando um único
arquivo, composto por todos os ossos que compunham o crânio do animal em estudo.
Além disso, com a possibilidade de deformar as superfícies 3D, elementos cranianos
que se encontravam distorcidos pelo processo de fossilização foram retro-deformados
alinhando-se pontos anatômicos de referência, assim como foi feito por Polcyn et al.
(2005).
Animações, simulando os tipos de movimentos realizados entre pares de
ossos adjacentes, bem como do crânio como uma unidade mecânica integrada, também
foram feitos utilizando o software Maya 2008. Os tipos de mobilidade e seus limites
foram identificados através de observações anatômicas das regiões de sutura, barreiras
impostas por outros ossos e por comparações com mobilidades intracraniana já descritas
para outros grupos. Assim, foram criados filmes de curta duração para cada movimento
descrito, além daqueles para os diferentes modelos de cinetismo propostos para o crânio
como um todo.
1.4 ANÁLISE INTEGRADORA
Com a digitalização dos elementos mais bem preservados do exemplar

UFRGS PV0629T foi possível fazer a reconstituição digital de todo o crânio,
25
permitindo manipular e analisar a influência de cada osso sobre o conjunto como um

todo. Desta forma, foram geradas animações, simulando os possíveis tipos de
mobilidades que pares de ossos isolados poderiam ter. Em seguida, grupos de ossos
foram sendo adicionados ao modelo, gerando assim, simulações cada vez mais
completas de todo o conjunto até que modelos com possíveis tipos de mobilidades entre
todas as peças do crânio fossem considerados. Além disso, também foi possível retro-
deformar os ossos maxilar, quadrado/quadrado-jugal esquerdo e neurocrânio/teto
craniano, simulando como seria a forma original destes ossos sem os efeitos de
deformação do processo de fossilização. Como resultados também foram obtidos
arquivos em três dimensões de cada peça tomografada, passíveis de serem manipulados
pela internet sem a necessidade de programas pagos, facilitando assim o acesso a esse
tipo de informação.
Em relação aos possíveis movimentos realizados entre os pares isolados,
foram identificadas áreas nítidas de articulação nos contatos do: - pré-maxilar com o
maxilar; - maxilar com o jugal; - maxilar com o lacrimal; - pterigóide com o processo
basipterigóide do basiesfenóide; - pterigóide com o quadrado; e, - pterigóide com o
ectopterigóide.
Integrando os elementos em um modelo geral para todo o crânio, foi
observado que alguns movimentos descritos como possíveis entre pares isolados, como
por exemplo, a rotação latero-medial do quadrado com o esquamosal, não poderiam ser
realizados no modelo integrado. No entanto, possibilidades de movimentos entre outros
elementos continuaram sendo plausíveis, tais como o deslocamento antero-
dorsal/postero-ventral do pterigóide sobre o processo basipterigóide do basiesfenóide e
o deslocamento do maxilar sobre o jugal na mesma direção.
A mobilidade intracraniana pode estar relacionada a diversos fatores
funcionais, e uma característica determinante para as possíveis funções de tal
mobilidade seria o tipo de cinetismo apresentado pelo animal. Este cinetismo poderia
ser realizado de forma ativa ou passiva, dependendo da presença ou não de
determinados grupos musculares, disposição de ligamentos e tendões, entre outros
fatores. Apesar das limitações presentes para este tipo de análise (reconstituição de
tecidos moles em fósseis), há algumas alternativas que permitem um acesso a estas
informações. Com a metodologia de cladismo reverso, proposta por Witmer (1995), e
com os trabalhos de descrição de musculaturas cranianas de crocodilos e aves, seria
possível fazer um levantamento dos possíveis músculos presentes, por exemplo, no
26
rauissúquio analisado neste estudo. Desta forma, seria possível verificar se algum dos
músculos encontrados poderia desencadear a mobilidade do crânio, permitindo um
modelo de cinetismo ativo para o grupo em questão. Por outro lado, a metodologia de
análise por elementos finitos, recentemente utilizada e aplicada à paleontologia por
diversos autores (Snively & Russel, 2002; Rayfiel, 2005a, 2005b; Richmond et al.,
2005), permite examinar a distribuição de forças e resistência geral, por exemplo, de um
crânio durante uma mordida. Portanto, seria possível, também, pesquisar como se
dariam as variações na distribuição destas forças ao incluir pontos móveis ao crânio.
Assim, alívios, redistribuições de tensões e resistência, poderiam ser acompanhados em
um crânio que apresentasse um modelo passivo de cinetismo craniano.
Um estudo voltado para estas novas abordagens, só seria possível após a
identificação das possíveis regiões de articulação intracraniana de um determinado
grupo. Neste trabalho, tal objetivo foi alcançado para um espécime de rauissuquídeo, o
que motivou o presente autor a dar continuidade ao estudo e à pesquisa nesta área.
Partindo deste ponto, será possível tratar tais temas ao longo de seu doutorado, com o
propósito de obter algumas respostas sobre os aspectos funcionais de um crânio
cinético.
27
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32
2. CORPO PRINCIPAL DA DISSERTAÇÃO
2.1 ARTIGO SUBMETIDO À REVISTA PALAEONTOLOGIA ELECTRONICA.
BIOMECHANICAL SIMULATION OF INTRACRANIAL MOBILITY IN A

RAUISUCHIDAE SPECIMEN USING COMPUTER TOMOGRAPHY AND 3D
MANIPULATION TECHNICS.
KEY WORDS: Archosauria; cranial kinesis; Crurotarsi; skull suture; Triassic
Alexandre Liparini and Cesar L. Schultz

Alexandre Liparini. Instituto de Geociências, Universidade Federal do Rio Grande do
Sul, Avenida Bento Gonçalves 9500, CEP 91501-970, Porto Alegre, Rio Grande do Sul,
Brasil. alexandreliparini@yahoo.com.br
BSc in Biology at the Federal University of Minas Gerais, Brazil, with focus on
vertebrate zoology and morphometry. Presently, finishing MSc at the Federal University
of Rio Grande do Sul in Geoscience - subarea Palaeontology. Interests and research
areas: -vertebrate palaeontology; -biomechanics; -3D digital modeling; and, -computer
based simulations.
Cesar L. Schultz. Instituto de Geociências, Universidade Federal do Rio Grande do Sul,

Avenida Bento Gonçalves 9500, CEP 91501-970, Porto Alegre, Rio Grande do Sul,
Brasil. cesar.schultz@ufrgs.br
BSc in Geology at the Federal University of Rio Grande do Sul (1983), MSc in
Geoscience (1986) and PhD in Science (1991) at the same university. Currently, is an
associate professor at the Federal University of Rio Grande do Sul (UFRGS) and assists
master and doctoral students of the UFRGS Geosciences Post Graduation Program.
Works on geosciences with focus on vertebrate palaeontology and biostratigraphy,
including mainly researches on Permian and Triassic tetrapods, upper Permian and
Triassic biostratigraphy, vertebrate taphonomy and vertebrate fossils digital
reconstructions.
33
2.1.1 Abstract
Partially disarticulated skull bones of an unidentified rauisuchid were

found in Dona Francisca, south Brazil. CT scan was used to analyse possible
intracranial mobility using specific 3D software. After mechanical preparation and CT
scan, sutures morphology and articulation areas were analysed for each individual pair
of adjacent bones and for an integrated mechanical model, which considered all bones
that comprises the skull of the studied specimen. Illustrative 3D animations for each
discussed movements were created and 3D surface model files for each CT scanned
skull element are presented. Regarding the mobility of individual pairs of bones, some
patent mobile sutures or articulation areas were identified such as the premaxilla and
maxilla, maxilla and jugal, pterygoid and basisphenoid and the pterygoid and
ectopterygoid contact. Some of the described mobility for isolated pair of bones were
suppressed when the integrated model was considered, however the intracranial
mobility as a whole remained possible. This study presented more evidences in favour
of passive cranial kinesis, which includes: -restricted, but possible mobility between
some bones; and, -presence of loosely contacts that would not be able to do extensive
movements or would present no mobility at all when the integrated model is considered.
Though, more studies on soft tissue reconstructions and ontogenetic development of
rauisuchids are needed to testify or falsify the hypothesis of an active cranial kinesis.
2.1.2 Introduction
In general, the term cranial kinesis has been used to indicate any type of
intracranial mobility (Versluys 1910; Hofer 1945; Frazzetta 1962; Bock 1964).
Different types of cranial kinesis are described for living vertebrates, especially lizards
(e.g., Frazzetta 1962) and birds (e.g., Bock 1964). Initially, those works were based on
comparative morphology and anatomy, considering both the morphology of skull bones
sutures and muscles arrangement. Since the beginning of the 1980s, with the
development of new methodologies such as cineradiography and electromyography,
authors came across new evidences on cranial kinesis functionality, making possible
34
innovative interpretations on this field (Throckmorton and Clarke 1981; Smith 1982;
Herrel et al. 1999; Herrel et al. 2000; Metzger 2002; Gussekloo and Bout 2005a,
2005b).
Cranial kinesis may have raised independently many times in the history
of vertebrate evolution (Schwenk 2000; Evans 2003). Different types of cranial kinesis,
such as streptostyly and mesokinesis, may also have appeared independently and,
therefore, might have autonomous mobility and functionality one from each other. This
hypothesis was confirmed for lizards by Smith (1980) and Herrel et al. (2000) and could
also have happened in other groups, including fossil ones.
Due to the impossibility of observing direct functionality of a kinetic
skull in fossils, as well as its muscular composition, inferences are made by studying the
morphology of skull bones shapes and contacts (Rayfield 2005a). Investigation on
cranial sutures morphology permits to determinate whether the mobility of adjacent
bones at a contact is possible or not, and which movements are feasible or not (e.g.,
rotation and/or translation). Those remarks may not demonstrate what movements, if
any, the animal actually did when alive, but indicates only the possible mobility
between bones without considering ligaments and muscles attachments. Therefore,
terms such as "skull mobility", "cranial flexibility" and "intracranial mobility" will be
preferably used, to give a more descriptive aspect to the analysis, instead of "kinetic
skull" or "cranial kinesis", which would lead to precipitated functional interpretations.
Regarding fossil archosaurs, there are some bibliographies on intracranial
mobility for: -rauisuchids (e.g., Chatterjee 1985, 1991); -theropod dinosaurs
(McClelland 1993 apud Rayfield 2005a; Mazzetta et al. 1998; Currie et al. 2003;
Rayfield 2004, 2005a, 2005b); and, -ornithopod dinosaurs (Norman and Weishampel
1985).
The rauisuchids comprise a group of exclusively Triassic crurotarsian
archosaurs with global distribution, except for Australia and Antarctica, however, their
findings are often sparse and incomplete (Gower 2000). The systematic of this group is
still controversy and much discussed (Benton and Clark 1988; Sereno and Arcucci
1990; Sereno 1991; Parrish 1993; Juul 1994). The terms Rauisuchia BONAPARTE
1982, and rauisuchids, will be used in this work as reference to a group which may
include the Rauisuchidae, Prestosuchidae, Poposauridae and Chatterjeeidae families, as
used by Gower (2000), independently of considering it a monophyletic taxon or not
(Chatterjee 1985; Galton 1985; Long and Murry 1995).
35
Rauisuchids were the top predators from the Triassic period, being
replaced, in the Jurassic, by theropod dinosaurs. Both groups presented the same general
morphology, probably due to convergent habits (Benton 1984). Despite the fact that
many groups of rauisuchids were formally described (for an overview see Gower 2000),
only few studies deal with cranial sutures morphology and the possibility of intracranial
movements. For example, Chatterjee (1985) described with some detail the cranial
kinesis of Postosuchus kirkpatricki, and Gower (1999) indicated some possible areas of
mobility in a rauisuchid skull from the Middle Triassic southern Germany (i.e.,
Batrachotomus kupferzellensis). Gower (2000) pointed towards the need of more
studies in this area in order to improve detailed descriptions on sutures and bone
contacts, such as the premaxilla and maxilla contact, what would also contribute to
character determination for phylogenetic analysis of the rauisuchids.
On the other hand, more studies on theropod’s intracranial mobility have
been made (McClelland 1993; Mazzetta et al. 1998; Currie et al. 2003; Rayfield 2004,
2005a, 2005b), possibly due to the facilitated access to this material, in comparison with
rauisuchids material available, and also because of the popularity of the former group.
Nowadays, with an easier access to computers with high data processing
capacity, the use and manipulation of digital images have been facilitated. Therefore,
techniques such as computed tomography (CT), became also more accessible,
permitting to extract previously unreachable fossil information (Ketcham and Carlson
2001). Furthermore, fossil 3D reconstitutions may be used to study taphonomic
distortion (e.g., Polcyn et al. 2005), retro-deformations, bone contacts and mobility, as
well as divulgation via World Wide Web (e.g., Alcober 2001 at
http://digimorph.org/specimens/Saurosuchus_galilei/).
In this work, disarticulated skull bones from a rauisuchid (probably
related to Prestosuchus HUENE 1942, but not yet formally identified), were
individually digitalized using CT scan to completely reconstruct a digital articulated
skull. Beyond that, intracranial joints and sutures were analysed and their possible
movements studied. Based on these considerations, 3D animations were created,
simulating the possible skull flexibility.
Besides the evident skull morphological/functional implications for
rauisuchids, this analysis may also contribute to systematics and phylogenetic studies on
rauisuchids, since morphological characters of joints and sutures will be treated in the
36
perspective of kinetics. In addition, we expect to contribute with detailed morphological

cranial information of a rauisuchid specimen.
2.1.3 Material and methods
Our material consisted of cranial elements from a rauisuchid not formally

described (cf. Prestosuchus), placed in the fossil vertebrate collection of the Vertebrate
Palaeontology Laboratory, Palaeontology and Stratigraphy Department of the
Geosciences Institute from the Federal University of Rio Grande do Sul. This specimen
was found and collected in 2003, near Dona Francisca town, central region of the
federal state Rio Grande do Sul, Brazil (Figure 1), identified as UFRGS PV0629T. This
specimen was collected from Mesotriassic sediments of the Therapsid Cenozone from
the Santa Maria Formation. This material comprises an almost complete individual,
with some articulated parts (including the posterior half of the backbone, pelvic girdle
and posterior members) and disarticulated ones, such as part of the skull and pectoral
girdle (Figures 2, 3).
After mechanical preparation, the cranial elements that could be
recovered and identified include: fused neurocranium/skull roof/right prefrontal/right
quadrate/right quadratojugal (Figure 4.1), left lacrimal/prefrontal (Figure 4.2), left
quadrate/quadratojugal (Figure 4.3), left jugal/ectopterygoid (Figure 4.4), right jugal
(Figure 4.5), right ectopterygoid/pterygoid (4.6), left pterygoid (Figure 4.7), left palatine
(Figure 4.8), left and right maxillae (Figure 4.9, 4.10), nasals (Figure 4.11, 4.12) and
premaxillae (Figure 4.13, 4.14). All mandibular elements (articular, surangular, angular,
splenial and dentary), were also identified for both sides (Figure 4.15-4.18).
In general, the bones of the left side of the skull were less deformed and
better preserved, when compared to the right ones. The left nasal, right jugal, pterygoid
and articular are fragmented. Furthermore, the fused neurocranium/skull roof (including
both anterior parts of the lower temporal fenestra, formed by the postorbitals)/right
prefrontal/right quadrate/right quadratojugal were deformed as shown in Figure 5. The
ventral bar of the left lower temporal fenestra and the ascending process of the left
maxilla were also deformed during the fossilization process.
37
The methodology proposed by Polcyn et al. (2005) to test alignment and

displacement of cranial elements of a fossil snake, was used for retro-deform the
following elements: 1) fused neurocranium/skull roof/right prefrontal/right
quadrate/right quadratojugal; 2) fused left quadrate/quadratojugal; and, 3) ascending
process of the left maxilla. This methodology was applied to the 3D files of the three
mentioned skull elements, acting directly in all symmetry planes. The retro-deformation
process was used as an approach to a more realistic reconstruction of the 3D skull.
Only the best preserved side of each paired skull bones were selected for
CT scanning. Thus, the left lacrimal/prefrontal (Figure 4.2), jugal/ectopterygoid (Figure
4.4), pterygoid (Figure 4.7), palatine (Figure 4.8), maxillae (Figure 4.9), premaxillae
(Figure 4.13) and mandibular ramus (Figure 4.15, 4.16), and right nasal (Figure 4.12)
were chosen to generate the 3D files. Furthermore, the fused neurocranium/skull
roof/right prefrontal/right quadrate/right quadratojugal (Figure 4.1) and the left
quadrate/quadratojugal (Figure 4.3) were also CT scanned.
A Brilliance 16, Philips tomograph (clinical use), placed at Hospital de
Clínicas de Porto Alegre, Brazil, was used to scan the fossil elements cited above. A
total of 12 pieces were scanned and their technical data are specified in Table 1. For
each scanned element two distinct visualization masks for the material density were
created. A wider one ranging from 0 to 3.000 HU and a narrower, which ranged from
450 to 3.000 HU. Then, two distinct 3D models were created for each scanned element
with its specific mask. For the first model, smoothing, triangle reduction and higher
matrix resolution methods were used, which resulted in higher quality files. For the
second one, neither smoothing nor triangle reduction was used, as well as a lower
matrix resolution was considered, presenting files with an intermediate quality (Figure
6.1-6.4). The files with higher qualities, consequently, demand higher computer
processing and handling time when compared to the files created with intermediate
quality.
All 3D files created were dealt in Maya 2008 ® software. Using this
program, each element was orientated in the X, Y and Z axes, so that the planes of
symmetry agreed with the program coordinate planes, that is to say X-plane corresponds
to the sagital plane of symmetry, Y-plane corresponds to the horizontal plane and Z-
plane to the coronal plane (for an illustrative example see Figure 7). In addition, each
3D file was scaled so that one unit measured by the software matched one centimetre.
Furthermore, the reduce polygon tool was used to reduce 75% of the 3D mesh faces
38
(Figure 6.5, 6.6). This method was applied to each file with intermediate quality,
generating files with low quality (for comparisons see Figure 6). Simple movements
simulation, including two to three elements, were made using files with higher quality,
whereas animations with more than three elements were done with lower quality files.
Morphological observations of sutures and articulation areas were
described separately for each element and their adjacent bones. Potential mobility of
each bone contributing to the sutures or articulation areas were recorded in the same
way as did Rayfield (2005a) (see details below). This potential mobility was also
simulated using 3D animations created on Maya. Formerly, possible movements in six
degrees of freedom (three translational and three rotational) for each contiguous pair of
elements were analysed before considering the restriction imposed on suture mobility
by surrounding contacts. Then, an animation framework and elements hierarchy was
created connecting all skull bones. Movements and limitations for each bone were
established on the animation framework, which permitted analyse skull flexibility as an
integrated mechanical unit.
2.1.4 Results and Discussion
Images of the retro-deformed and original positioned: -

neurocranium/skull roof/right prefrontal/right quadrate/right quadratojugal; -fused left
quadrate/quadratojugal; and, -ascending process of the left maxilla, are presented on
Figures 8-10, respectively. Henceforward, one should consider images of isolated 3D
elements being in its original form and images or animations with two or more elements
in its retro-deformed 3D form.
For sutures morphology and intracranial movements, results and
discussion were separated into two different sections. For the first one, articular areas
for each isolated element and their surrounding contacts were pointed out and discussed.
Demonstrative 3D animations were created for each pair of elements, showing how
each proposed movement could occur. In this level, as mentioned previously, the
restrictions imposed by other contacts were not taken into account. On the other hand,
the second section considered the whole biomechanical chain, where all bones were
mutually connected. Therefore, movements that formerly where possible between two
39
isolated elements, but restricted at another point were reconsidered. It is not the aim of
these analyses to reconstruct skull musculature or possible ligaments attachment, so that
a functional analysis will be restricted and only a preliminar overview will be done.
On the first analysis, the bones that, individually, clearly presented at
least one well defined region of mobility were the premaxilla, maxilla, jugal, pterygoid,
ectopterygoid and quadrate. As no vomers were found, it was not possible to define
whether mobility with their surrounding contacts was present or not.
2.1.4.1 Isolated Elements
2.1.4.1.1 Premaxilla (Figures 11-14)
For a schematic illustration of the premaxilla and its surrounding contacts

see Figure 11 and for a 3D surface model visualization of the CT scanned bone see
Figure 12. Figures 13 and 14 are simulations of possible movements of the premaxilla
with maxilla and nasal, respectively. The premaxilla contacts the nasal, maxilla and
probably the vomer. It has a postero-medial process that, in lateral view, corresponds to
one fourth of its total length. This process articulates ventrally with the palatal process
of the maxilla permitting an anterior-posterior movement of the premaxilla. Besides this
mobility, a limited rotation over the X axis in the sagital plane was possible (Figure 13).
The lateral face of the premaxillar anterior ascending process, contacts the medial face
of the most distal part of the nasal. At this point a sliding movement in the
anteroventral-posterodorsal direction was possible. In the other hand, the suture between
the posterior ascending process of the premaxilla and the descending process of the
nasal seems to present no mobility or may rotate in the X axis in the sagital plane
(Figure 14). Due to poorly preserved boundaries, the mobility of this contact could not
be more accurately defined.
40
2.1.4.1.2 Maxilla (Figures 15-18)
The maxilla contacts anteriorly the premaxilla, dorsomedially, the nasal

and lacrimal, medially the palatine (and possibly the vomer), and posteriorly the jugal
bone. The anterior edge of the ascending process of the maxilla contacts the ventro-
lateral edge of the nasal. Along the axis of this contact, a sliding movement in the
anteroventral-posterodorsal direction and rotation of the maxilla was possible (Figure
16). The lacrimal bone articulates its distal lateral face with the medial face of the final
portion of the ascending process of the maxilla. This suture could present a sliding
mobility in an anterior-posterior direction as well as in a dorso-ventral direction as
shown in the simulation of Figure 17. Palatine’s lateral edge, from the choana to the
suborbital fenestra, contacts the medial face of the maxilla in the same line of its palatal
process. This articulation area could not be clearly interpreted, so that no mobility could
be determined such as in the posterior ascending process of the premaxilla. A conical
process at the posterior portion of the maxilla was observed and, in this work, referred
as jugal process of the maxilla (Figure 11). Jugal articulates at the dorso-lateral face of
this process, where sliding in the anterodorsal-posteroventral direction was possible
(Figure 18).
2.1.4.1.3 Nasal (Figures 19, 20)
The nasal bone contacts posteriorly the frontal, laterally lacrimal and
maxilla, and anteriorly the premaxilla. In dorsal view, the posterior part of the nasal is
“L-shaped”. This region contacts the frontal laterally and anteriorly, where the medial
contact of the nasal acts as a rotation pivot so that its posterior contact slightly slides
over the anterior edge of the frontal (Figure 20). This type of movement would be
analogous to the prokinetic mobility present in some lizards and many groups of birds
(Frazzetta 1962; Bock 1964), differently from Allosaurus fragilis, for example, that
rather presented antero-posterior translational movements than rotational ones (Rayfield
2005a). Regarding the nasal-lacrimal contact, the anterior portion of the lacrimal medial
41
border articulates with the lateral and posterior edge of the nasal. At this suture no
possible movements could be determined.
2.1.4.1.4 Lacrimal/prefontal (Figures 21, 22)
The left lacrimal and prefrontal are fused and were found isolated from
the frontal bone. At the right side the lacrimal was not preserved, but a fragment of the
prefrontal remained in contact with the frontal. The frontal-prefrontal contact at the
right side is masked by sediment and was not analysed. On the other hand, this
articulation at the left side is clearer and a rotation of the lacrimal/prefrontal over the X
axis was possible (Figure 22). The descending process of the prefrontal (or descending
pillar of the prefrontal) may form a continuous with the anterior ascending process of
the jugal forming the preorbital bar, but, for this specimen it was broken (or
disarticulated) before its fossilization (Figure 11), so that the possibility of movements
at this suture could not be determined. An interesting observation was made by Gower
(1999), who reported disarticulation and loss of the prefrontals from the frontals of
Batrachotomus kupferzellensis (SMNS 52970), suggesting a loosely contact between
those bones in both species of rauisuchids.
2.1.4.1.5 Jugal (Figures 23, 24)
Five articulation areas around the jugal were observed. At its ventral
border and medial face, it articulates with the ectopterygoid. Left jugal and
ectopterygoid are merged by sediment in a single unit, and it is not possible to
determinate whether this contact is fused or could present any mobility (Figure 23).
However, the right jugal was preserved separately, showing a fragment of the right
ectopterygoid united to the pterygoid. This indicates rather a slackly contact between
those bones (jugal, ectopterygoid and pterygoid), than a firmly one. Anteriorly, the jugal
contacts the maxilla, and the lacrimal/prefrontal bones. The anterior border of the
posterior ascending process articulates with the posterior border of the postorbital
42
descending process. At this suture, the jugal could slide along the axis of the postorbital
bar in an anteroventral-posterodorsal direction (Figure 24). The posterior part of the
jugal contacts the anterior process of the quadrato-jugal forming the lower temporal bar.
Despite the evident suture between these bones, they were fused, so that no mobility
was observed.
2.1.4.1.6 Ectopterygoid (Figure 23, 25)
The ectopterygoid beyond articulating with the jugal, contacts the

postero-ventral ramus of the pterygoid. At this articulation, the medial face of the
postero-ventral ramus of the ectopterygoid contacts, almost completely, the lateral face
of the postero-ventral ramus of the pterygoid, where movements in the anterodorsal-
posteroventral direction were possible (Figure 25).
2.1.4.1.7 Pterygoid (Figure 26-28)
The pterygoid contacts the quadrate, the basipterygoid process of the

basisphenoid, the ectopterygoid and the palatine. Medially, at the base of the quadrate
ramus of the pterygoid, a short rod-like groove can be observed, where the
basipterygoid process of the basisphenoid fits. At this joint rotation around the long axis
of this rod-like process and translation in the anterodorsal-posteroventral direction were
possible (Figure 27). Posteriorly, the quadrate ramus of the pterygoid presents two
flanges, which lateral faces articulates with the medial face of the quadrate. At this
contact, the pterygoid could slide over the quadrate face in an anterior-posterior as well
as in a dorso-ventral direction (Figure 28). Finally, almost the whole ventral face of the
anterior portion of the pterygoid seems to articulate with the dorsal face of the posterior
half of the palatine, but their contact, thus also their possible mobility, could not be
clearly defined.
43
2.1.4.1.8 Palatine (Figure 29)
In respect to the palatine, all its contacts and types of possible

movements were described above, although no clear articulation site could be identified
for this bone.
2.1.4.1.9 Quadrate/quadrato-jugal (Figure 30-32)
The left quadrate fractured before fossilization next to its articulation

with the lower jaw (Figure 11). It is fused to the quadrato-jugal, and both bones were
found disarticulated from the squamosal at the skull roof. Considering the articulation
joint with the squamosal, the quadrate/quadrato-jugal could rotate back and forwards,
and also to the right and left side around the pivot established between these two bones
(Figure 31). These movements would be restricted anteriorly and medially till it
contacts the descending process of the squamosal. This type of mobility can be
compared to the streptostylic kinesis described for lizards and birds. However, it is
different in the way that the quadrate is fused to the quadrato-jugal that forms the lower
temporal bar, which would restrict this movement. The integration of this movement to
their surrounding bones (jugal, ectopterygoid, postorbital, lacrimal/prefrontal, maxilla
and pterygoid) will be discussed later. On the other hand, at the right side, the
quadrate/quadrato-jugal remained fixed to the squamosal at the skull roof (Figure 32).
Nevertheless, as observed in the left side, the descending process of the right squamosal
would limit only an anterior and medial rotation of the quadrate.
2.1.4.1.10 Neurocranium/skull roof (Figure 32)
The neurocranium and the skull roof are fused to each other, but it is
worth to highlight that the paroccipital process of the opistotic and the supraoccipital are
44
clearly individualized. This may indicate some mobility between the neurocranium and
the skull roof, in the same way as occurs in some lizards (i.e. metakinesis).
2.1.4.1.11 Mandible (Figure 33-35)
No specific suture type was identified at the anterior portion of the

dentaries and splenials, delimitating a mandibular symphysis. This indicates a loosely
contact between each mandibular ramus, made only by ligaments and perhaps, would
have allowed each ramus to move in a partially independently manner. Slight
adjustments in the antero-posterior and dorso-ventral directions (Figure 34) and
outwards (lateral) rotation of each mandibular ramus (Figure 35) could be realized at
this contact. This mobility would compensate irregular prey forms in bilateral biting and
adjust to possible cranial kinesis. On the other hand, intramandibular kinesis, such as
described for Postosuchus kirkpatricki (Chatterjee 1985) or those described for some
theropod groups (Mazzetta et al. 1998; Sampson and Witmer 2007) between anterior
and posterior segments of the lower jaw, were not possible in the observed specimen.
Movements described above for isolated elements were summarized in
Table 2 and Table 3. Bone contacts and types of mobility encountered for each
articulation can be seen in Table 2. Table 3 relates movements described for each bone
to its animated movement simulation figures.
Similar descriptions of possibilities of mobility for isolated elements
were made elsewhere for some rauisuchids and theropods. Rayfield (2004) described
kinetic contacts between maxilla-jugal and postorbital-jugal for Tyrannosaurus rex,
where, respectively, anterodorsal-posteroventral and dorso-ventral possibilities of
mobility were observed in most analysed specimens. Rayfield concluded that the
maxilla-jugal suture would provide a tensile shock-absorbing site, reducing local
stresses. Furthermore, Rayfield (2005a) investigated the relation between stress
environment, cranial strength and sutural morphology and mobility in the skull of
Allosaurus fragilis and concluded that A. fragilis cranial sutures would be capable of
accommodating stress and strain patterns generated during biting. Gower (1999), while
describing Batrachotomus kupferzellensis, indicated potentially mobile joints between
individual skull elements. However, in his conception, consideration of the skull and
45
mandible as a whole suggests that adults of B. kupferzellensis lacked significant cranial

and mandibular kinesis.
Detailed morphological descriptions of sutures can also be used as
character determination for phylogenetical analysis. Parrish (1993) used 42 characters to
study crurotarsian phylogeny. In his list of characters, one could identify at least five
characters that were directly related to isolated skull elements mobility. For example,
character 17 stays for the posterior process of the maxilla that could be loosely or firmly
attached to the maxilla and character 39 stays for the presence or absence of the
descending process of the squamosal (Parrish 1993). In this way, it is also possible to
identify probable mobility regions in close phylogenetic related groups by observing
bone structures rather than articulation sites (e.g., presence of the descending process of
the squamosal would limit anterior movements of the quadrate).
2.1.4.2 Integrated Elements
Considering the skull as a whole, one should think of an integrated model

of the skull, where all bone elements and its individual movements were taken into
account as a single mechanical chain. At this mechanical chain, actions of one element
are reflected in other ones even if they do not contact directly. So, a bone with limited
mobility could block a sequence of movements initiated by other ones.
At first place the neurocranium/skull roof was considered as the
stationary (fixed) unit and movements of other elements were described in relation to
this one. The contact with this stationary unit occurs at 5 points, which are the nasals,
lacrimals/prefrontals, jugals, quadrates and pterygoids (Figure 36).
2.1.4.2.1 Quadrate unit (Figure37)
Considering that the quadrates are fused to the quadrate-jugals and that
these are fixed to the jugals and ectopterygoids, those bones would form a unit that
could move as a block in an anterior-posterior direction (Figure 37). However, if one
46
considers the lacrimals/prefrontals also fused to the anterior ascending bar of the jugals,
the mobility described for the lacrimals/prefrontals (rotation at the frontals) and the one
described above for the quadrate unit (antero-posterior rotation at the squamosal) would
block one another. These two movements would only be permitted if the preorbital bar
had a mobile articulation between the lacrimal and the jugal. The rotation of the
quadrate moving this unit medio-laterally was not permitted, because the pterygoid
would have to translate laterally, what was not possible due to the basipterygoid process
of the basisphenoid. Without the medio-laterally rotation of the quadrate, no lateral
rotation of each mandibular ramus, as shown in Figure 35 would be possible.
Additionally, if the lacrimal-jugal contact was fused, the quadrates would present no
mobility at all, and the movements described for the jugal-postorbital and frontal-
prefrontal (see Table 4) would not be feasible too.
2.1.4.2.2 Pterygoid unit (Figure38)
At the basipterygoid process of the basisphenoid, the medial-laterally

rotation of the pterygoids would be blocked by the ectopterygoids, since this movement
would pull the dorsal part of the quadrate inwards, which is limited to the descending
process of the squamosal. On the other hand, the anterodorsal-posteroventral translation
of the pterygoid would encounter no limitation, since this movement could also be done
at the contacts pterygoid-quadrate and pterygoid-ectopterygoid (Table 4). At the
anterior region of the pterygoid, the palatine was not fused, but no specific type of
movements could be established at this contact as well as at the contact between the
palatine and the maxilla. So, another unit was determined including the pterygoid,
palatine and maxilla (Figure 38). The anterodorsal-posteroventral translation of the
pterygoid would result in the same movement at the maxilla, which reflected at the
maxilla contacts with the jugal, lacrimal, nasal and premaxilla. Even with the fixed
model of the quadrate unit and the fused preorbital bar, the anterodorsal-posteroventral
translation of the maxillae would occur at their contacts with the jugals and lacrimals
(Figure 39). As the maxilla is also linked to the nasal and premaxilla, its influence over
those bones were also analysed. The forth and upwards movement of the maxilla would
be possible, because of the rotation of the nasal at its contact with the frontal, and, at the
47
same time, the sliding movement of the maxilla over the nasal (Figure 40). Rotation of
the maxilla at the nasal contact as described above (see Table 2), would only be possible
if the quadrate unit could rotate outwards at the squamosal contact, which was blocked
by the pterygoid. Regarding the premaxillae, they could follow the anterodorsal-
posteroventral movements of the maxillae in a wider range only if the posterior
ascending process of the premaxilla presented a rotation with the descending process of
the nasal, which is dubious. If this movement was possible, the premaxilla would rotate
at its contact with the maxilla and the anterior process of the premaxilla would slide
over the nasal most distal part as shown in Figure 41. If there was no mobility between
the nasal and premaxilla, the anterodorsal-posteroventral translation of the maxilla
would still be possible, but with a more limited range of translation (Figure 42).
Considering all the possibilities of intracranial movements discussed
above, a complete 3D surface model reconstruction of the UFRGS PV0629T skull was
created (Figure 43). According to the descriptions above, 4 models as a mechanical unit
were determined: 1) without mobility at the lacrimal-jugal and at the posterior
premaxilla-nasal contact (Figure 44); 2) without mobility at the lacrimal-jugal contact
and a mobile premaxilla-nasal contact (Figure 45); 3) with a mobile lacrimal-jugal
contact and a not mobile premaxilla-nasal contact (Figure 46); and, 4) with both
lacrimal-jugal and premaxilla-nasal mobile contact (Figure 47). All models could
potentially be kinetic and the major difference observed between these 4 models was the
possible mobility of the quadrate in models 3 and 4 (Figures 46, 47), representing an
analogue of the streptostilic movements described for lizards and birds, and an
independent movement of the premaxilla in models 2 and 4, which has no analogue
considering terrestrial vertebrates, what may be of great importance for further studies.
2.1.4.3 Functional Inferences
For Squamata extant groups, the mobility of the quadrate was related to
the increase of mechanical advantage for: 1) m. adductor mandibulae externus
(Gingerich 1971) and m. pterygoideus (Smith 1980); 2) gape increase (Patchell and
Shine 1986; MacLean 1974); 3) efficient intraoral transport of food items
(Throckmorton and Clarke 1981); and, 4) shearing of prey for processing (De Vree and
48
Gans 1994; Herrel and De Vree 1999), among others (see Metzger 2002). More
evidences are necessary to confirm the mobility of the quadrate unit of the studied
specimen and other rauisuchids, but such possibility of mobility could be associated to
specific and distinct feeding habits, which would give important insights of the living
habits of those extinct groups. To achieve this data specific functional analysis should
be done and more fossil material is needed.
The loosely contact between the premaxilla and maxilla have been also
recorded elsewhere for other rauisuchids (Parrish 1993; Gower 1999), which could also
indicate possible mobile contact. This type of mobility does not present any analogue in
extant groups and no functional implications were proposed yet. Morphofunctional
analysis would be a promissory and challenging field regarding this subject.
Independent of which model one considers, possible intracranial mobility
was observed for the studied specimen. This can reflect many different ecological habits
depending on the degree of freedom of the movements. Furthermore, the degree of
freedom of those movements will depend on other features such as ontogenetic stage,
soft tissues association with sutures and adjacent bones, and type of intracranial
mobility (i.e. active or passive kinesis).
Despite the total length size estimated for the studied specimen (≈ 6
meters), the rauisuchid analysed could be a juvenile, which sutures were not completely
developed. If this was the case, it would not prevent skull flexibility but could indicate a
decrease in cranial kinesis through ontogeny, as proposed by Walker (1990) for
Sphenosuchus acutus. However, to validate this hypothesis, more specimens in different
ontogenetic stages would be necessary to compare suture morphologies.
Studies on soft tissues rarely can be done by analysing direct evidences
on fossils, nevertheless they could leave impressions in some cases that would improve
reconstructions and biomechanical and morphofunctional analyses. Accessing soft
tissue data would help to determinate if the studied animal could have active cranial
kinesis, where specific major muscles and, therefore, their impressions should be
present (e.g. insertion and/or origin marks of m. pterigoideus). On the other hand,
loosely contacts between bones, which actually could not present considerable mobility
to work actively, is an evidence for passive intracranial mobility. Regarding the studied
specimen, it seems more probably to present passive intracranial mobility than active
muscle powered cranial kinesis, mainly because most skull bones were preserved
separately as shown in Figures 3 and 4, and some predicted isolated movements (Table
49
2) could not be realized when the integrated mechanical model was considered (Table
4), though other movements encountered no limitations even in the integrated model.
2.1.5 Conclusions
Considering isolated movements of adjacent bones and its sutures, some

patent regions of mobility could be determined. Clear mobility areas were identified
between the premaxilla and maxilla, maxilla and jugal, maxilla and lacrimal, pterygoid
and basipterygoid process of the basisphenoid, pterygoid and quadrate and at the
pterygoid and ectopterygoid contact. In other contacts, mobility was not present or may
be present but could not be accurately defined.
Four integrated models of skull mobility were created to analyse
influences of each bone in the whole mechanical chain. Each model considered different
possibilities of movements between some bones, which contacts and possible
movements could not be precisely defined (i.e. lacrimal/jugal and premaxilla/nasal
contact). At the integrated model, some possible movements described for isolated pair
of bones were suppressed, such as outward rotation of the quadrate around the antero-
posterior axis (Z axis), medio-laterally rotation of the pterygoid around the Z axis and
outward rotation of the maxilla around the oblique axis formed at its contact with the
nasal. Nevertheless other movements remain possible and intracranial mobility as a
whole may have occurred. This mobility could occur as a result to external forces
during biting (i.e. passive kinesis) or could be an active muscle powered intracranial
mobility (i.e. active kinesis). More evidences lead to passive rather than active cranial
kinesis for the studied rauisuchid specimen. However, functional analyses were not
made and remain an interesting subject for further studies.
2.1.6 Perspectives
Functional implications for intracranial mobility, independent of being

active or passive kinesis, can be elucidated with the help of methodologies that have
50
been recently applied to palaeontology. Advanced morphological studies of extant

groups, closely related to extinct groups, have contributed for the reconstruction of soft
tissue of fossil groups. For example, using the Extant Phylogenetic Bracket
methodology proposed by Witmer (1995), and cranial musculatures descriptions for
crocodiles and birds, one could reconstruct major cranial musculature groups of well
preserved rauisuchid skull material, such as the one presented in this study. Such
analysis would give clues on what possible types or models of intracranial mobility
could be present depending on the existing cranial muscles. On the other hand, finite
element analysis, recently applied to palaeontology by many authors (Rayfield et al.
2001; Snively and Russel 2002; Rayfield 2005a, 2005b; Richmond et al. 2005), may
help to elucidate questions on stress and strain distribution and overall skull resistance
of the analysed specimens. Therefore, comparisons between stress relief, strain
redistribution and overall resistance in a rigid and a flexible skull (i.e. akinetic and
passive kinetic skulls) could be done for rauisuchids and would lead to exciting and
innovative analyses for this group.
2.1.7 Acknowledgements
We would like to thank Daniel C. Fortier for imaging and computational

assistance; Lorena C. Fleury and Juan C. Cisneros for reviews and useful comments of
the manuscript; Bianca M. Mastrantonio for refined mechanical preparation of most
material and for conceding her illustrations; Teo V. de Oliveira for contributing with his
illustrations; Adolfo Bittencourt and Martin Reus for 3D modelling and 3D animation
assistance; and Coordenação de Aperfeiçoamento de Pessoal de Nível Superior
(CAPES) for financial support.
51
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56
2.1.9 Figures
FIGURE 1. Location map where UFRGS PV0629T was collected.

57
FIGURE 2. UFRGS PV0629T reconstruction. Blue paintings indicates which

elements were found. Total length estimated in 6 meters.
58
FIGURE 3. Illustration indicating original position of the bones in the fossil when
collected and prepared. Cranial elements are marked in blue. Scale of the drawing:
approximately 180 cm in length and 90 cm in width.
59
FIGURE 4. Prepared skull bones of UFRGS PV0629T. Skull roof, neurocranium and
right quadrate/quadrato-jugal (1) in dorso-posterior view. Left lacrimal (2), right
quadrate/quadrato-jugal (3), left jugal/ectopterygoid (4) and right jugal (5) in lateral
views. Right pterygoid/ectopterygoid (6) and left pterygoid (7) in dorso-lateral views.
Left palatine (8) in dorsal view. Left (9) and right (10) maxillae, left (11) and right (12)
nasals, left (13) and right (14) premaxillae, anterior (15) and posterior (16) portions of
the left mandibular ramus, and anterior (17) and posterior (18) portions of the right
mandibular ramus, all in lateral views.
60
FIGURE 5. 3D images of the neurocranium/skull roof of PV0629T in posterior view,

laying on its right side, as founded in the rock, with probable deformation sequence
during the fossilization process. Possible original shape (1) and deformed shape (2) as
removed from sediments. Arrows indicate the deformation forces direction. Scale bar
equals 7 cm.
61
FIGURE 6. 3D images of the left premaxilla in lateral view as an example of the

different quality types of each generated 3D surface model. 3D files were created based
on a narrower HU density range with high quality (1), intermediate (3), and low quality
(5). The same 3D surface model simplifications were applied for 3D files with a wider
HU density range, with a high quality 3D file (2), an intermediate (4) and a low quality
one (6). Scale bar equals 7 cm.
62
FIGURE 7. Correlations between the anatomical planes of symmetry and coordinate

planes using the left premaxilla as an example. The sagital plane (lateral (1) and medial
(2) views) is related to the X coordinate plane (red X box), the coronal plane (anterior
(3) and posterior (4) views) to the Z coordinate plane (blue Z box) and the horizontal
plane (dorsal (5) and ventral (6) views) is related to the Y coordinate plane (green Y
box). The X letter at each red axes indicates the left side, the Y letters at the green axes
indicate the dorsal (top) region, and the Z letters at the blue axes indicate the anterior
(front) region. Any of the boxes may be used as scale bars considering each side equal
to 1 cm.
63
FIGURE 8. Images comparing original shape of the prepared Neurocranium/skull

roof in anterior (1), left (3), posterior (5) and dorsal (7) views, where the points of
alignment were indicated by yellow linked dots and the retro-deformed results in
anterior (2), left (4), posterior (6) and dorsal (8) views. Coordinate axes were used as
reference lines of symmetry. Scale bar equals 7 cm.
64
FIGURE 9. Comparison of the original (in lateral (1) and ventral (3) views) and
retro-deformed shapes (in lateral (2) and ventral (4) views) of the left
quadrate/quadrato-jugal. Points of alignment were indicated with yellow linked dots.
Coordinate axes were used as reference lines of symmetry. Scale bar equals 7 cm.
65
FIGURE 10. Comparison of the original (1) and retro-deformed (2) shapes of the left
maxilla in anterior view. Points of alignment were indicated with yellow linked dots.
Coordinate axes were used as reference lines of symmetry. Scale bar equals 7 cm.
66
FIGURE 11. Schematic illustration of some cranial bones in left lateral view and their
surrounding contacts. Abbreviations: aap, anterior ascending process; ap, ascending
process; d, disarticulation; dp, descending process; Ec, ectopterygoid; f, fracture; J,
jugal; jp, jugal process; L/Prf, lacrimal/prefrontal; M, maxilla; N, nasal; pap, posterior
ascending process; pmp, posterior medial process; pp, palatal process; Prm,
premaxilla; Q, quadrate; Qj, Quadrato-jugal. Scale bar equals 7 cm.
67
FIGURE 12. 3D surface model of the left premaxilla. Initial image in left lateral view.
Click on the image to activate 3D controls (read Appendix 1 for detailed activation
information). Premaxilla total length is approximately 8.5 cm.
68
FIGURE 13. 3D animated movie of the possibilities of mobility between the left
premaxilla and maxilla in left lateral view. Arrows indicate movement direction in the
still image. Click on the image to activate the movie (read Appendix 1 for detailed
activation information). Maxilla total length is approximately 27.0 cm.
69
premaxilla and the mirrored right nasal in left lateral view. Arrows indicate movement
direction in the still image. Click on the image to activate the movie. Premaxilla total
length is approximately 8.5 cm.
70
FIGURE 15. 3D surface model of the left maxilla (not retro-deformed). Click on the
image to activate 3D controls. Initial image in left lateral view. Maxilla total length is
approximately 27.0 cm.
71
maxilla and the mirrored right nasal in oblique left lateral/anterior/dorsal view. Arrows
indicate movement direction in the still image. Click on the image to activate the movie.
Maxilla total length is approximately 27.0 cm.
72
maxilla and lacrimal/prefrontal in left lateral view. Arrows indicate movement direction
in the still image. Click on the image to activate the movie. Maxilla total length is
73
FIGURE 18. 3D animated movie of the mobility between the left maxilla and the left
jugal in left lateral view. Arrow indicates movement direction in the still image. Click
on the image to activate the movie. Maxilla total length is approximately 27.0 cm.
74
FIGURE 19. 3D surface model of the right nasal. Initial image in right lateral view.
Click on the image to activate 3D controls. Nasal total length is approximately 25.0 cm.
75
FIGURE 20. 3D animated movie of the possibilities of mobility between the right
nasal and the right frontal, which is integrated to the neurocranium/skull roof. Arrow
indicates movement direction in the still image. Left lateral view. Click on the image to
activate the movie. Nasal total length is approximately 25.0 cm.
76
FIGURE 21. 3D surface model of the left fused lacrimal/prefrontal. Initial image in
left lateral view. Click on the image to activate 3D controls. Lacrimal/prefrontal total
77
lacrimal/prefrontal and the left frontal in left lateral view. Arrow indicates movement
direction in the still image. Click on the image to activate the movie.
Lacrimal/prefrontal total length is approximately 18.0 cm.
78
FIGURE 23. 3D surface model of the left jugal and ectopterygoid. Initial image in left
lateral view. Click on the image to activate 3D controls. Jugal total length is
79
FIGURE 24. 3D animated movie of the possibilities of mobility between the left jugal
and the left postorbital, which is fused to the neurocranium/skull roof unit. Arrow
indicates movement direction in the still image. Left lateral view. Click on the image to
activate the movie. Jugal total length is approximately 15.0 cm.
80
ectopterygoid and the left pterygoid. Arrow indicates movement direction in the still
image. Right medial view. Click on the image to activate the movie. Pterygoid total
81
FIGURE 26. 3D surface model of the left pterygoid. Initial image in left lateral view.
Click on the image to activate 3D controls. Pterygoid total length is approximately 24.0
cm.
82
pterygoid and the pterygoid process of the basisphenoid in oblique medial/ventral view.
Arrows indicate movement directions in the still image. Click on the image to activate
the movie. Pterygoid visible length is approximately 18.0 cm.
83
pterygoid and the left quadrate in right medial view. Arrows indicate movement
directions in the still image. Click on the image to activate the movie. Pterygoid total
84
FIGURE 29. 3D surface model of the left palatine. Initial image in ventral view. Click
on the image to activate 3D controls. Palatine total length is approximately 25.5 cm.
85
FIGURE 30. 3D surface model of the left fused quadrate/quadrato-jugal (not retro-
deformed). Initial image in left lateral view. Click on the image to activate 3D controls.
Quadrate/quadrato-jugal total length is approximately 12.0 cm.
86
FIGURE 31. 3D animated movie of the possibilities of mobility between the quadrate
and squamosal in oblique left lateral/posterior view. Arrows indicate movement
directions in the still image. Click on the image to activate the movie.
Quadrate/quadrato-jugal total height is approximately 13.5 cm.
87
FIGURE 32. 3D surface model of the fused neurocranium/skull roof/right quadrate

(not retro-deformed). Initial image in left lateral view. Click on the image to activate 3D
controls. Neurocranium/skull roof total length is approximately 25.5 cm.
88
FIGURE 33. 3D surface model of the anterior and posterior portions of the left
mandibular ramus. Initial image in left lateral view. Click on the image to activate 3D
controls. Left mandibular ramus total length is approximately 47.5 cm.
89
FIGURE 34. 3D animated movie of the possibilities of translational mobility between

each mandibular ramus at the mandibular symphysis in oblique left
lateral/anterior/dorsal view. Arrows indicate movement directions in the still image.
Click on the image to activate the movie. Left mandibular ramus total length is
90
FIGURE 35. 3D animated movie of the possibilities of rotational mobility between

mandibular rami. Arrows indicate movement directions in the still image. Dorsal view.
Click on the image to activate the movie. Left mandibular ramus total length is
91
FIGURE 36. 3D surface model of the neurocranium/skull roof and its contact with the
nasals, lacrimals/prefrontals, jugals/ectopterygoids, quadrates/quadrato-jugals and
pterygoids. Initial image in left lateral view. Click on the image to activate 3D controls.
Total length in lateral view is approximately 46,0 cm.
92
FIGURE 37. 3D animated movie of the possibilities of movement of the quadrate unit
in relation to the neurocranium/skull roof. Arrows indicate movement directions in the
still image. Left lateral view. Click on the image to activate the movie.
Neurocranium/skull roof total length is approximately 25.5 cm.
93
FIGURE 38. 3D animated movie of the possibilities of movement of the pterygoid unit
in relation to the neurocranium/skull roof. Arrow indicates movement directions in the
still image. Left lateral view. Click on the image to activate the movie. Maxilla total
94
integrated with a fixed quadrate unit and without considering the nasals. Arrow
indicates movement directions in the still image. Left lateral view. Click on the image to
activate the movie. Maxilla total length is approximately 27.0 cm.
95
integrated with a fixed quadrate unit and considering the mobility of the nasals. Arrows
indicate movement directions in the still image. Left lateral view. Click on the image to
activate the movie. Maxilla total length is approximately 27.0 cm.
96
FIGURE 41. 3D animated movie of the possibilities of movement at the articulation

site among the premaxilla, maxilla and nasal contact, considering the premaxilla as an
isolated unit not fixed to the nasal. Arrows indicate movement directions in the still
image. Left lateral view. Click on the image to activate the movie. Maxilla total length
is approximately 27.0 cm.
97
FIGURE 42. 3D animated movie of the possibilities of movement at the articulation

site among the premaxilla, maxilla and nasal contact, considering the premaxilla fixed
to the nasal. This fixation would reduce the movement but not block it. Arrows indicate
movement directions in the still image. Left lateral view. Click on the image to activate
the movie. Maxilla total length is approximately 27.0 cm.
98
FIGURE 43. 3D surface model of the whole reconstructed skull of the UFRGS
PV0629T specimen. Initial image in left lateral view. Click on the image to activate 3D
controls. Total length in left lateral view is approximately 49.0 cm.
99
FIGURE 44. 3D animated movie of model 1 - possible intracranial mobility

considering the whole integrated mechanical model, where the lacrimal/prefrontal-jugal
and premaxilla-nasal contacts would present no mobile sutures. In the still image, the
"X" mark indicates the considered fixed regions for this model. Oblique left
lateral/anterior/dorsal view. Click on the image to activate the movie. Maxilla total
100

contact would present no mobility and the premaxilla-nasal contact would have a
mobile articulation site. For this model, circle indicates mobile sutures whether "X"
mark indicates fixed regions in the still image. Oblique left lateral/anterior/dorsal view.
Click on the image to activate the movie. Maxilla total length is approximately 27.0 cm.
101

considering the whole integrated mechanical model, where the lacrimal/prefrontal-
frontal contact would present mobility and the premaxilla-nasal contact would not have
a mobile suture. For this model, circle indicates mobile sutures whether "X" mark
indicates fixed regions in the still image. Oblique left lateral/anterior/dorsal view. Click
on the image to activate the movie. Maxilla total length is approximately 27.0 cm.
102

and the premaxilla-nasal articulation sites would be mobile. Circles in the still image
indicate the considered regions where mobility were possible for this model. Oblique
left lateral/anterior/dorsal view. Click on the image to activate the movie. Maxilla total
103
2.1.10 Tables
104
TABLE 1. Technical CT specification for each scanned skull element.

Voltage Current Slice Slice Number Resolution Pixel Orientation* Field
Thickness Increment of Size of View
Elements (V) (mAs) (mm) (mm) Slices (pxl) (cm)
Left jugal/ectopterygoid 120 375 1 1 153 512x512 0.8008 ALB 41
Left lacrimal 120 497 1 1 124 512x512 0.8008 PLT 41
Anterior left mandibular ramus 120 375 1 1 117 512x512 0.8008 PRB 41
Posterior left mandibular ramus 120 375 1 1 147 512x512 0.8008 ART 41
Left maxilla 120 497 1 1 194 512x512 0.8008 PLT 41
Right nasal 120 375 1 1 113 512x512 0.8008 ABR 41
Neurocranium/skull roof 120 281 1 1 292 512x512 0.9258 TRP 47.4
Left premaxilla 120 375 1 1 105 512x512 0.8008 BLP 41
Left pterygoid 120 469 1 0.5 352 512x512 0.6641 PLT 34
left palatine 120 375 1 1 105 512x512 0.8008 ATL 41
Left quadrate/quadrato-jugal 140 344 2 1 139 512x512 0.268 ALB 13.7
Left quadrate fragment 140 399 2 1 88 512x512 0.236 LAT 12.1
* A=anterior; P=posterior; L=left; R=right; T=top; B=botton.

Comparing to the position of a human laid on the CT bed, the first letter corresponds to the human left side, the second letter to his
dorsal plane (contacts the bed) and the last letter to his cranial position.
105
TABLE 2. Summary of the observed bone contacts and possible movements described for each isolated element.
Mandibular
Premaxilla Maxilla Nasal Lacrimal Prefrontal Frontal Postorbital Basisphenoid Squamosal Quadrate Quadrato-jugal Jugal Ectopterygoid Pterygoid Palatine Vomer Articular synphysis
Premaxilla NF Rx;Tz ? (0;Rx;T1) -- -- -- -- -- -- -- -- -- -- -- -- ? -- --
Maxilla Rx;Tz -- R1;T1 T3 -- -- -- -- -- -- -- T2 -- -- ? (NF) ? -- --
Nasal ? (0;Rx;T1) R1;T1 NF ? (NF) ? (NF) Rx -- -- -- -- -- -- -- -- -- -- -- --
Lacrimal -- T3 ? (NF) -- F -- -- -- -- -- -- ? -- -- -- -- -- --
Prefrontal -- -- ? (NF) F -- Rx -- -- -- -- -- ? -- -- -- -- -- --
Frontal -- -- Rx -- Rx -- -- -- -- -- -- -- -- -- -- -- -- --
Postorbital -- -- -- -- -- -- -- -- -- -- -- T1 -- -- -- -- -- --
Basisphenoid -- -- -- -- -- -- -- F -- -- -- -- -- R2;T2 -- -- -- --
Squamosal -- -- -- -- -- -- -- -- -- Rx;Rz ? -- -- -- -- -- -- --
Quadrate -- -- -- -- -- -- -- -- Rx;Rz -- F -- -- T3 -- -- Ry --
Quadrato-jugal -- -- -- -- -- -- -- -- ? F -- 0 -- -- -- -- -- --
Jugal -- T2 -- ? ? -- T1 -- -- -- 0 -- ? (0;Rz;Tz) -- -- -- -- --
Ectopterygoid -- -- -- -- -- -- -- -- -- -- -- ? (0;Rz;Tz) -- T2 -- -- -- --
Pterygoid -- -- -- -- -- -- -- R2;T2 -- T3 -- -- T2 NF ? (NF) -- -- --
Palatine -- ? (NF) -- -- -- -- -- -- -- -- -- -- -- ? (NF) NF ? -- --
Vomer ? ? -- -- -- -- -- -- -- -- -- -- -- -- ? ? -- --
Articular -- -- -- -- -- -- -- -- -- Ry -- -- -- -- -- -- -- --
Mandibular
synphysis -- -- -- -- -- -- -- -- -- -- -- -- -- -- -- -- -- Ry;T3
-- (not applicable) R (rotate around the axis) 1 (oblique - anteroventral-posterodorsal)

? (unknown, dubious or unassigned) T (translate along the axis - slide) 2 (oblique - anterodorsal-posteroventral)
; (and/or) x (latero-medial - x axis) 3 (all directions at the sagital plane)
F (fused) y (dorso-ventral - y axis)
NF (not fused) z (antero-posterior - z axis)
0 (no mobility)
106
TABLE 3. Reference list for each described isolated element movement and its
illustrative animated figure.
Reference Figures Type of mobility observed and bones involved
Figure 13 Rotation and translation between premaxilla and maxilla
Figure 14 Possible rotation and translation between premaxilla and nasal
Figure 16 Rotation and translation between maxilla and nasal
Figure 17 Translation between maxilla and lacrimal
Figure 18 Translation between maxilla and jugal
Figure 20 Rotation between nasal and frontal
Figure 22 Rotation between prefrontal and frontal
Figure 24 Translation beween jugal and postorbital
Figure 25 Translation between ectopterygoid and pterygoid
Figure 27 Rotation and translation between pterygoid and basisphenoid
Figure 28 Translation between pterygoid and quadrate
Figure 31 Rotation between quadrate and squamosal
Figure 34 Translation at the mandibular synphysis
Figure 35 Rotation at the mandibular synphysis
107
TABLE 4. List of bone contacts that presented or could present mobility in isolated
elements and whether the mobility remained or not in each considered mechanical
model.
Mobility
Considered bone Adjacent bone Isolated pair Model 1 Model 2 Model 3 Model 4
Premaxilla Maxilla + + + + +
Nasal (‡) ? 0 + 0 +
Vomer ? ? ? ? ?
Maxilla Premaxilla + + + + +
Nasal + 0 (R1); + (T1) 0 (R1); + (T1) 0 (R1); + (T1) 0 (R1); + (T1)
Vomer ? ? ? ? ?
Palatine ? 0 0 0 0
Lacrimal/prefrontal + + + + +
Jugal + + + + +
Nasal Premaxilla (‡) ? 0 + 0 +
Maxilla + 0 (R1); + (T1) 0 (R1); + (T1) 0 (R1); + (T1) 0 (R1); + (T1)
Lacrimal/prefrontal ? ? ? ? ?
Frontal + + + + +
Palatine Maxilla ? 0 0 0 0
Vomer ? ? ? ? ?
Pterygoid ? 0 0 0 0
Lacrimal/ Maxilla + + + + +
prefrontal Nasal ? ? ? ? ?
Jugal (‡) ? 0 0 + +
Frontal + 0 0 + +
Jugal Maxilla + + + + +
Lacrimal/prefrontal (‡) ? 0 0 + +
Postorbital + 0 0 + +
Ectopterygoid ? 0 0 0 0
Quadrate/quadrato-jugal 0 0 0 0 0
Pterygoid Palatine ? 0 0 0 0
Ectopterygoid + + + + +
Basisphenoid + 0 (R2); + (T2) 0 (R2); + (T2) 0 (R2); + (T2) 0 (R2); + (T2)
Ectopterygoid Jugal ? 0 0 0 0
Pterygoid + + + + +
Quadrate/ Jugal 0 0 0 0 0
quadrato-jugal Pterygoid + + + + +
Squamosal + 0 0 + (Rx); 0 (Rz) + (Rx); 0 (Rz)
Articular + + + + +
Mandibular Translation + + + + +
synphysis Rotation + 0 0 0 0
‡ (contacts that differentiate each model) x (latero-medial - x axis)

? (unknown, dubious or unassigned) z (antero-posterior - z axis)
0 (no mobility) 1 (oblique - anteroventral-posterodorsal)
+ (mobile) 2 (oblique - anterodorsal-posteroventral)
R (rotate around the axis)
T (translate along the axis - slide)
108
2.1.11 Appendix A - Activation of non still figures
2.1.11.1 3D surface models
Figures where a 3D surface model can be manipulated may need to have

a free 3DCT plug-in installed that can be donwloaded in the folowing website:
http://www.3dcompress.com/web/prod_view.asp. Usually, it is automatically instaled
when one tries to open a web site with an embedded 3D file (.3dc) for the first time.
Once activated, the main controls are: 1) rotation tool (pressing the left
mouse button while moving the pointer); 2) zoom tool (pressing the right mouse button
while moving the pointer for zoom in or zoom out); and, 3) hand tool to move the object
(pressing both mouse buttons while moving the pointer).
Other functions such as anatomical views are extra tools and may be
acessed by clicking at each respective link when available.
2.1.11.2 3D animated movies
Figures where short animated movies can be seen may need the free
quick time viewer software. This software can be downloaded for free at the following
website: http://www.apple.com/quicktime/download/
109
APÊNDICE A - Documentos em meio eletrônico
CD-ROM contendo os arquivos, tais como figuras, arquivos 3D e

animações, apresentados no corpo principal da dissertação, além de uma versão digital
da dissertação com os seus devidos endereços ligando cada figura ao seu respectivo
conteúdo ativo, e as normas para apresentação de dissertações de mestrado na forma de
artigo, elaborada pelo Programa de Pós-Graduação em Geociências. Esta mídia contem
também arquivos em formatos de páginas de internet (html) para facilitar a navegação
entre os documentos disponíveis no CD.
Neste CD estão incluídos:
− 01 arquivo da dissertação em formato digital;
− 01 arquivo com as normas para apresentação da dissertação;
− 47 figuras (formato jpg);
− 04 tabelas (formato html);
− 24 animações (formato mov);
− 12 arquivos contendo o código html para ativação e
visualização dos arquivos 3D (formato html);
− 12 arquivos referente às imagens em três dimensões (formato
3dc);
− 01 arquivo no formato html (index.html) para navegar pelos
documentos disponíveis no CD-ROM, e sua pasta
correspondente contendo os arquivos necessários para que a
página seja carregada corretamente.
110
111
ANEXO A - Carta de submissão do artigo

112
ANEXO B - Resumo expandido na I Semana Acadêmica em 2006

113
114
115
116
117
118
ANEXO C - Resumos na Paleo RS em 2006

119
120
121
ANEXO D - Resumo na XXIII Jornadas Argentinas em 2007

122

Estudo Da Biomecânica Craniana de Um Rauissuquídeo A Partir de Tomografias Computadorizadas E Técnicas de Imagens Digitais em 3 Dimensões

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Estudo Da Biomecânica Craniana de Um Rauissuquídeo A Partir de Tomografias Computadorizadas E Técnicas de Imagens Digitais em 3 Dimensões

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UNIVERSIDADE FEDERAL DO RIO GRANDE DO SUL

ESTUDO DA BIOMECÂNICA CRANIANA DE UM

ORIENTADOR – Prof. Dr. Cesar Leandro Schultz

Porto Alegre, 2008

ESTUDO DA BIOMECÂNICA CRANIANA DE UM

ORIENTADOR – Prof. Dr. Cesar Leandro Schultz

Dissertação de Mestrado apresentada

Porto Alegre, 2008

Dissertação (Mestrado). - Universidade Federal do Rio Grande

1. Archosauria. 2. Cinelismo craniano. 3. Crurotarsi. 4. Triássico.

Dedico esta dissertação aos meus pais, Agostinho A. G. Campos e

Dedico este trabalho também ao meu tio Antônio C. Liparini que,

À minha querida e Sempre-Linda do Cerrado, Lorena C. Fleury,

Um esqueleto parcialmente completo e semi-articulado de um rauissuquídeo,

PALAVRAS-CHAVE: Archosauria; cinetismo craniano; Crurotarsi; Triássico.

Partially disarticulated skull bones of an unidentified rauisuchid were found in Dona

KEYWORDS: Archosauria; cranial kinesis; Crurotarsi; Triassic.

FIGURA 1 - Principais pontos de articulação e eixos de rotação em um crânio de

FIGURA 2 - Modelo mecânico integrando os pontos de cinetismo em um lagarto

FIGURE 1 - Location map where UFRGS PV0629T was collected............................56

FIGURE 2 - UFRGS PV0629T reconstruction ............................................................57

FIGURE 3 - Illustration indicating original position of the bones in the fossil

FIGURE 4 - Prepared skull bones of UFRGS PV0629T .............................................59

FIGURE 5 - 3D images of the neurocranium/skull roof of PV0629T in posterior

FIGURE 6 - 3D images of the left premaxilla in lateral view as an example of the

FIGURE 7 - Correlations between the anatomical planes of symmetry and

FIGURE 8 - Images comparing original shape of the prepared

FIGURE 10 - Comparison of the original and retro-deformed shapes of the left

FIGURE 12 - 3D surface model of the left premaxilla ..................................................67

FIGURE 13 - 3D animated movie of the possibilities of mobility between the left

FIGURE 14 - 3D animated movie of the possibilities of mobility between the left

FIGURE 15 - 3D surface model of the left maxilla (not retro-deformed)......................70

FIGURE 17 - 3D animated movie of the possibilities of mobility between the left

FIGURE 19 - 3D surface model of the right nasal .........................................................74

FIGURE 20 - 3D animated movie of the possibilities of mobility between the right

FIGURE 21 - 3D surface model of the left fused lacrimal/prefrontal............................76

FIGURE 22 - 3D animated movie of the possibilities of mobility between the left

FIGURE 23 - 3D surface model of the left jugal and ectopterygoid..............................78

FIGURE 24 - 3D animated movie of the possibilities of mobility between the left

FIGURE 25 - 3D animated movie of the possibilities of mobility between the left

FIGURE 26 - 3D surface model of the left pterygoid ....................................................81

FIGURE 27 - 3D animated movie of the possibilities of mobility between the left

FIGURE 28 - 3D animated movie of the possibilities of mobility between the left

FIGURE 29 - 3D surface model of the left palatine.......................................................84

FIGURE 30 - 3D surface model of the left fused quadrate/quadrato-jugal (not

FIGURE 31 - 3D animated movie of the possibilities of mobility between the

FIGURE 32 - 3D surface model of the fused neurocranium/skull roof/right

FIGURE 34 - 3D animated movie of the possibilities of translational mobility

FIGURE 37 - 3D animated movie of the possibilities of movement of the quadrate

FIGURE 38 - 3D animated movie of the possibilities of movement of the pterygoid

FIGURE 39 - 3D animated movie of the possibilities of movement of the pterygoid

FIGURE 40 - 3D animated movie of the possibilities of movement of the pterygoid

FIGURE 41 - 3D animated movie of the possibilities of movement at the

FIGURE 42 - 3D animated movie of the possibilities of movement at the

FIGURE 43 - 3D surface model of the whole reconstructed skull of the UFRGS

FIGURE 44 - 3D animated movie of model 1 - possible intracranial mobility

FIGURE 45 - 3D animated movie of model 2 - possible intracranial mobility

FIGURE 46 - 3D animated movie of model 3 - possible intracranial mobility

FIGURE 47 - 3D animated movie of model 4 - possible intracranial mobility

TABLE 2 - Summary of the observed bone contacts and possible movements

TABLE 4 - List of bone contacts that presented or could present mobility in

TEXTO EXPLICATIVO DA ESTRUTURA DA DISSERTAÇÃO.........................13

2. CORPO PRINCIPAL DA DISSERTAÇÃO...........................................................32