Functional morphologists have extensively used measurements of performance to investigate the relationship among form, function and ecology through ontogeny. Among different measurements of performance bite force play a crucial role influencing fitness. Although, bite force has been thoroughly investigated among vertebrates, the majority of the studies on fishes have been concentrated only in small species. Consequently, this is the first study that compares the bite force performance in two large marine predators, the great barracuda (Sphyraena barracuda) and bull shark (Carcharhinus leucas). Values of posterior bite force in S. barracuda varied from 3 - 258 N for an ontogenetic series of 27 individuals (23 - 130 cm, TL). Bite force as well as the majority of the biomechanical variables that contribute to it scaled with isometry in S. barracuda. Values of posterior bite force in C. leucas varied from 170 - 5,914 N for and ontogenetic series of 16 individuals (73 - 285 cm, TL). Bite force at the most anterior bite point scaled with positive allometry as well as the majority of the subdivisions of the adductive musculature that greatly contribute to bite force. Bite force performance in this two species showed strong differences, where S. barracuda has one of the lowest relative values of bite force among fishes and C. leucas has one of the largest ones. Additionally, the scaling patterns for bite force and most of the biomechanical variables investigated in this study differed among these two species. These results suggest that predatory success may be acquired by different strategies, and that the same ecological role in a marine ecosystem may be reached by having different bite force performance.

Pocket gophers (family Geomyidae) are the dominant burrowing rodents in North America today. Their fossil record is also incredibly rich; in particular, entoptychine gophers, a diverse extinct subfamily of the Geomyidae, are known from countless teeth and jaws from Oligocene and Miocene-aged deposits of the western United States and Mexico. Their postcranial remains, however, are much rarer and little studied. Yet, they offer the opportunity to investigate the locomotion of fossil gophers, shed light on the evolution of fossoriality, and enable ecomorphological comparisons with contemporaneous rodents. We present herein a quantitative study of the cranial and postcranial remains of eight different species of entoptychine gophers as well as many contemporary rodent species. We find a range of burrowing capabilities within Entoptychinae, including semifossorial scratch-digging animals and fossorial taxa with cranial adaptations to burrowing. Our results suggest the repeated evolution of chisel-tooth digging across genera. Comparisons between entoptychine gophers and contemporaneous rodent taxa show little ecomorphological overlap and suggest that the succession of burrowing rodent taxa on the landscape may have had more to do with habitat partitioning than competition.

京都大学 / 0048 / 新制・課程博士 / 博士(人間・環境学) / 甲第20458号 / 人博第808号 / 新制||人||194(附属図書館) / 28||人博||808(吉田南総合図書館) / 京都大学大学院人間・環境学研究科相関環境学専攻 / (主査)教授 加藤 眞, 教授 市岡 孝朗, 准教授 西川 完途 / 学位規則第4条第1項該当 / Doctor of Human and Environmental Studies / Kyoto University / DGAM

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Thesis (PhD)--Stellenbosch University, 2013. / ENGLISH ABSTRACT: The phenotype of an individual has often been used as the descriminating factor in distinguishing species. However, with the advent of more precise molecular techniques, the genotype of species is increasingly being used as the preferred method in taxonomic classifications. Many taxa have recently been demonstrated to be incongruent in terms of their genetic and morphological groupings, and this may due to the influence that the environment may have on the morphological and functional aspects of a species. Selective pressures often act upon the performance of a species within a particular habitat first, and then selection for the morphological characters that allow for optimal performance occurs. Should genetically disparate species inhabit a particular environment, convergence in morphologies and performance may evolve. Historically, lizard species descriptions were based primarily on external morphologies, and thus misclassfication of species may have occurred due to mistakenly grouping species with convergent morphologies together. In the current dissertation, the links between morphology, performance capacities, diet and behaviour is explored in comparison to the environment and genetic relationships of southern African lacertid lizards. The performance capacities and associated morphological traits were expected to be more closely linked with the environment, and not closely linked with genetic relationships. To investigate these expectations, a multidisciplinary approach was taken, and genetic, morphological and performance analyses were done and compared with dietary behavioural and environmental analyses. In the first chapter, the link between habitat openness and the lizard bauplans is investigated and the presence of convergent morphologies within this group of lizards is uncovered. These convergences are shown to have resulted in misclassification of two lacertid species, and taxonomic revisions within the family are discussed. The second chapter explores the link between performance and associated morphological traits, and the dietary composition of the members of the Nucras genus. The third chapter identifies the link between the predator escape strategies employed by the members of the Meroles genus, and their morphologies and performance capacities. The fourth chapter explores the intraspecific, inter-population differences in morphologies and investigates the link between the morphological groupings and the population genetic groupings within Pedioplanis lineoocellata. The final chapter identifies whether adaptation to a novel habitat can occur over a relatively short period of time, and the morphological traits, functional aspects, and population genetic structure is investigated in conjunction with environmental analyses of vegetation and substrate between the populations of Meroles knoxii. It was concluded that the morphological and functional aspects of the southern African lacertid lizards are more closely related to the environment, particularly the microhabitat structure, than to their genetic relationships, and that future work using this group of lizards should involve a multidisplinary approach as different selective pressures are playing a role in shaping the morphologies and performance capacities of these lizards, compared to those that are acting upon the genotypes of the lizards. / AFRIKAANSE OPSOMMING: Die fenotipe van 'n individu is dikwels gebruik as die diskriminerende faktor in kenmerkende spesies. Maar, met die ontwikkeling van meer akkurate molekulêre tegnieke, word die genotipe van spesies toenemend gebruik as die voorkeur-metode in taksonomiese klassifikasie. Die onversoenbaarheid van genetiese en morfologiese eienskappe kom voor in ‘n verskeidenheid taksa, dit kan wees as gevolg van die invloed wat die omgewing het op die morfologiese en funksionele aspekte van ‘n spesie. Selektiewe druk beїnvloed dikwels doeltreffende funktionaliteit van 'n spesie in 'n bepaalde habitat eerste, en gevolglik word morfologiese karakters wat voorsiening maak vir optimale funktionaliteit geselekteer. Indien geneties uiteenlopende spesies woon in 'n bepaalde omgewing, kan konvergensie in morfologie en soortgelyke werksverrigtinge ontwikkel. Histories, is akkedis spesiesbeskrywings hoofsaaklik gebaseer op eksterne morfologieë, en kan dus misklassifikasie tot gevolg hê wat kan lei tot foutiewe taksonomie van spesies met konvergente morfologieë. In die huidige verhandeling, is die verband tussen die morfologie, werksverrigtingsvermoë, dieët en gedrag ondersoek, in vergelyking met die omgewing en die genetiese verwantskappe van Suider-Afrikaanse sandakkedisse. Die werksverrigtingsvermoë en gepaardgaande morfologiese eienskappe word verwag om te meer verband te hou met die omgewing, en dus nie in noue verband te wees met die genetiese verwantskappe nie. Om hierdie verwagtinge te ondersoek, is 'n multi-dissiplinêre benadering geneem, en genetiese, morfologiese en werksverrigtingontledings is gedoen in vergelyking met dieët, gedrags-en omgewings-ontleding. In die eerste hoofstuk, is die skakel tussen die habitat openheid en die akkedis bauplans ondersoek en die teenwoordigheid van konvergente morfologieë binne hierdie groep akkedisse word ten toon gestel. Hierdie konvergensies het gelei tot foutiewe klassifikasie van twee sandspesies, en taksonomiese hersiening binne die gesin word bespreek. Die tweede hoofstuk ondersoek die verband tussen werksverrigting en gepaardgaande morfologiese eienskappe, en die samestelling van die dieët van die lede van die Nucras genus. Die derde hoofstuk identifiseer die verband tussen die roofdier ontsnapping strategieë, morfologieë en werksverrigtingsvermoë van die Meroles genus. Die vierde hoofstuk ondersoek die intraspesifieke, inter-bevolkingsverskille in morfologieë en ondersoek die verband tussen die morfologiese groepe en die bevolking genetiese groepe binne die Pedioplanis lineoocellata spesies kompleks. Die finale hoofstuk identifiseer hoe die aanpassings na 'n nuwe habitat kan plaasvind oor 'n relatief kort tydperk, en die morfologiese eienskappe, funksionele aspekte en die bevolking genetiese struktuur word ondersoek in vergelyking met die omgewingsanalise van plantegroei en substraat tussen die bevolkings van Meroles knoxii. Die gevolgtrekking is dat die morfologiese en funksionele aspekte van die Suider- Afrikaanse sandakkedisse nader verwant is aan die omgewing, veral die mikrohabitat struktuur, as aan hul genetiese verwantskappe. Toekomstige werk op hierdie groep akkedisse moet ‘n multidisiplinêre benadering behels siende dat verskillende selektiewe drukke 'n rol speel in die vorming van die morfologie en werksverrigtingsvermoë van hierdie akkedisse, in vergelyking met selektiewe drukke wat die genotipes van die akkedisse beinvloed.

Previous research on the biomechanics of tool use has focused heavily on traits correlated with locomotion, tool manufacturing, and habitual tool use. Features like the breadth of the metacarpals, relative length of the thumb, styloid process of the third metacarpal, and the breadth of the apical tufts are skeletal features associated with the use and development of stone tools. However, there are many traits of the distal forelimb that may also be correlated directly with the development and use of tools. The purpose of this research is to analyze morphological features of the hands and compare them to features of the arm in humans, fossil Homo and the great apes to understand how the hominin distal arm functions as a mosaic in response to the use of stone tools. The results indicate a separation between tool-users and non-tool users when all distal forelimb dimensions are examined. Omo 40-19 falls closer to non-tool users when univariate plots of ulna length and breadth are examined. Ratios of hand measurements to radius length are better at polarizing the tool-users from non-tool users than are hand dimensions to ulna length ratios. These results highlight the role of the radius in stabilizing the hand during stone tool production.

Os isópodos terrestres possuem uma cutícula protetora que mantém a forma corporal, permite locomoção e comunicação com o ambiente e protege contra dessecação, infecção e predação. Assim como nos demais crustáceos, a cutícula é composta por uma matriz orgânica que é mineralizada com cálcio. A cutícula é uma estrutura versátil que reflete adaptações ambientais e a ampla distribuição geográfica do grupo. Dessa forma, a ultraestrutura e a composição da cutícula variam entre espécies. Isópodos terrestres fazem mudas frequentes ao longo da vida para crescer e/ou renovar receptores de superfície, o que resulta em reabsorção e deposição cuticular constante. Esse grupo apresenta muda bifásica e deposição de placas de cálcio nos esternitos anteriores antes da ecdise como estratégia para reciclar o cálcio corporal. Estudos relacionados à cutícula nesse grupo contemplam a ultraestrutura, a composição e a deposição bem como o efeito de alguns fatores ambientais na muda. No entanto, poucos estudos exploram as ligações entre ecomorfologia e história de vida. Essa tese visa explorar diversos aspectos relacionados à cutícula e a muda de duas espécies neotropicais de isópodos terrestres. Atlantoscia floridana e Balloniscus glaber foram usados como modelo uma vez que são frequentemente encontrados nas mesmas localidades e diferem em tipo ecomorfológico e estratégias ambientais. No Capítulo I, foram exploradas as estruturas de superfície e a ultraestrutura da cutícula das duas espécies a fim de ver como as diferenças encontradas podem ser relacionadas à história de vida de cada espécie, utilizando técnicas de microscopia. As espécies diferiram em tipo e disposição das estruturas de superfície bem como em espessura e proporção de camadas da cutícula. De maneira geral, as diferenças das estruturas de superfície estão relacionadas à seleção de microhabitat e nicho ecológico. Características cuticulares ajudam a explicar o hábito endógeno de B. glaber e epígeo de A. floridana enquanto que as diferenças em ultraestrutura são relacionadas a estratégias comportamentais e tolerância ambiental. No Capítulo II, o efeito do cálcio alimentar no ciclo de muda das espécies foi testado. Para isso, dieta experimental e análise estrutural da cutícula foram realizadas a fim de entender como diferentes concentrações de cálcio alimentar interferem na duração do ciclo de muda. Peculiaridades na ecdise em A. floridana refletem morfologia específica do animal. A duração da intramuda foi maior em B. glaber assim como a sobrevivência média nos tratamentos. A duração do ciclo de muda foi influenciada pela concentração de cálcio; uma tendência a menor duração do ciclo com o aumento da concentração de cálcio foi observada em B. glaber, enquanto em A. floridana a diferença encontrada foi apenas entre o controle sem cálcio e os demais tratamentos. Não houve efeito da dieta no grau de mineralização ou na ultraestrutura em B. glaber. Independentemente do tratamento, a maior taxa de mortalidade em laboratório parece estar relacionada com o próprio processo de ecdise, com mortalidade acumulada de 20% do início da ecdise até o início da pós-muda para ambas as espécies. No Capítulo III, a secreção da cutícula durante a pré- e pós-muda muda foi observada utilizando microscopia de transmissão. A deposição seguiu o padrão observado para outros isópodos. Entretanto, grânulos eletrondensos presentes no espaço ecdisial durante a pré-muda são provavelmente constituídos de cálcio, sugerindo a reciclagem de cálcio diretamente da cutícula velha para a nova no mesmo segmento. Esses grânulos são depositados nas escamas na nova epicutícula antes da ecdise, sugerindo a presença de cálcio na superfície cuticular das espécies. Além disso, regiões sem a ultraestrutura típica encontrada na pós-muda indicam que há modificação na exocutícula após a ecdise uma vez que a expansão e endurecimento da nova cutícula apenas após a ecdise. De maneira geral, esse trabalho não apenas trouxe novas informações sobre a estrutura cuticular de duas espécies neotropicias, mas também contribuiu para esclarecer conexões entre ecomorfologia e requerimentos biológicos de isópodos terrestres. / Terrestrial isopods have a protective cuticle that maintains body shape, allows locomotion, enables communication with the environment and protects them against desiccation, infection and predation. As in all crustaceans, their cuticle is composed of an organic matrix that is mineralized with calcium. The cuticle is highly versatile reflecting adaptations to environmental conditions and large geographical distribution of this group. Therefore, cuticle ultrastructure and composition vary among species. Terrestrial isopods molt frequently throughout their lives in order to grow and/or renew surface receptors, resulting in constant cuticular resorption and deposition. In this group, this dynamics of cuticle formation is affected by the biphasic molt and by the calcium deposition on sternal deposits prior ecdysis, strategies to recycle body calcium. Studies related to cuticle on this group include ultrastructure, composition and deposition as well as effect of some environmental factors on molting. However, few studies explore connections of ecomorphology and life history of animals. This thesis aimed to explore various aspects of cuticle structure and molting using two Neotropical species of terrestrial isopods. Atlantoscia floridana and Balloniscus glaber were used as models since they are found in the same locations while differing in ecomorphology and behavioral strategies. In Chapter I, I explored the cuticle surface structures and ultrastructure of both species to see how their differences can be related to each species life history traits, using microscopy techniques. Species differed in surface structures type and disposition, as well as cuticle thickness and layer proportion. Overall, differences in surface structure are related to microhabitat selection and ecological niche. Cuticular features further explain the endogeic habit of B. glaber and epigeic habit of A. floridana, while differences in cuticle ultrastructure relate to behavioral strategies and environmental tolerance. Next, differences on molting cycle and environmental requirements were analyzed. In Chapter II, I tested the effects of dietary calcium on the molting cycle of both species. For that, artificial diet and structural analysis of the cuticle were used to understand how different concentrations of dietary calcium interfere with molt cycle duration. Peculiarities were observed during ecdysis in A. floridana and reflect to specific morphology of the species. Intramolt duration was longer for B. glaber as well as overall survivorship in treatments. Cycle duration was influenced by calcium concentration; a trend of shorter molt cycle length with increasing calcium concentration was observed for B. glaber, while in A. floridana, only difference between control without calcium and other treatments was observed. Degree of mineralization and cuticle ultrastructure of B. glaber showed no difference between treatments. Regardless of treatment or species, higher mortality rate under lab conditions seems to be related to the process of ecdysis itself, with cumulative mortality of 20% from the beginning of ecdysis until the beginning of postmolt. In Chapter III, I used transmission electron microscopy to analyze cuticle secretion in both species during pre- and postmolt stages. Cuticle deposition during premolt followed the same pattern as other terrestrial isopods. Nonetheless, electron dense granules present on the ecdysial space during premolt are likely calcium granules, suggesting the recycling of calcium within the same segment. These granules are deposited on the scales of the new epicuticle prior ecdysis, suggesting the presence of calcium on the cuticular surface of both species. Moreover, regions without typical lamellate ultrastructure during postmolt indicate modification of the exocuticle after ecdysis since expansion and hardening must occur after ecdysis. Overall, this work not only added information on cuticular structure of two Neotropical species but also clarified connections between ecomorphology and biological requirements of terrestrial isopods.

abstract: As the junction between the head and the trunk, the neck functions in providing head stability during behaviors like feeding to facilitating head mobility during behavior like grooming and predator vigilance. Despite its importance to these vital behaviors, its form and function remain poorly understood. Fossil hominin cervical vertebrae preserve a striking diversity in form despite the commitment to orthograde bipedality. Do these differences in cervical vertebral form correspond to functional variations among our recent ancestors? This dissertation attempts to understand 1) how does the neck function in head stability and mobility 2) how do these functions relate to cervical vertebral form. Kinematic and passive range of motion studies were conducted in several species of primate to obtain measures of function which were subsequently related to skeletal form. Results show that cervical vertebral morphology does not significantly covary with differences in joint mobility. Rather, they implicate the critical role of ligaments and muscles in facilitating head mobility. Results of the kinematics study show that the neck plays a role in maintaining head stability during locomotion. However, the kinematic data do not significantly correlate with morphological variation among primate species. Given the negative results of the extant morphological analyses, it is difficult to apply them to the fossil record. As such, the functional significance of the disparate morphologies found in the hominin fossil record remain ambiguous. / Dissertation/Thesis / Doctoral Dissertation Anthropology 2019