Measuring the magnitude of morphological integration: The effect of differences in morphometric representations and the inclusion of size

The magnitude of morphological integration is a major aspect of multivariate evolution, providing a simple measure of the intensity of association between morphological traits. Studies concerned with morphological integration usually translate phenotypes into morphometric representations to quantify how different morphological elements covary. Geometric and classic morphometric representations translate biological form in different ways, raising the question if magnitudes of morphological integration estimates obtained from different morphometric representations are compatible. Here we sought to answer this question using the relative eigenvalue variance of the covariance matrix obtained for both geometric and classical representations of empirical and simulated datasets. We quantified the magnitude of morphological integration for both shape and form and compared results between representations. Furthermore, we compared integration values between shape and form to evaluate the effect of the inclusion or not of size on the quantification of the magnitude of morphological integration. Results show that the choice of morphological representation has significant impact on the integration magnitude estimate, either for shape or form. Despite this, ordination of the integration values within representations is relatively the same, allowing for similar conclusions to be reached using different methods. However, the inclusion of size in the dataset significantly changes the estimates of magnitude of morphological integration, hindering the comparison of this statistic obtained from different spaces. Morphometricians should be aware of these differences and must consider how biological hypothesis translate into predictions about integration in each particular choice of representation.Fil: de Andrade Machado, Fabio. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina. University of Massachussets; Estados UnidosFil: Hubbe, Alex. Universidade Federal da Bahia; BrasilFil: Melo, Diogo. Universidade de Sao Paulo; BrasilFil: Porto, Arthur. University of Oslo; NoruegaFil: Marroig, Gabriel. Universidade de Sao Paulo; Brasi

Late Quaternary population expansion in gray brocket deer [ Subulo gouazoubira (G. Fisher, 1814), Cervidae, Mammalia] in Brazil

There is little information about the changes in the abundance of mammalian species in South America during the late Quaternary, which is a key parameter to understanding past ecological and evolutionary dynamics. Here, we describe a conspicuous increase in the abundance of the gray brocket deer (Subulo gouazoubira G. Fisher, 1814; Mammalia) during the late Quaternary in Brazil based on paleontological evidence. We studied the material retrieved from pitfall deposits in Cuvieri Cave (Brazil) representing three time intervals (Late Pleistocene, Pleistocene/Holocene transition, and Holocene). The deposits in this cave were previously characterized in terms of facies composition, chronology, and overall taxa and taphonomy. We combined previous data with taxonomic identification at the species level for deer (morphological and ancient DNA analyses), quantification of the number of individuals for large mammals (>10 kg), taphonomic analyses, and stratigraphic distribution of specimens. Our results show that while in the Late Pleistocene gray brocket deer corresponded to up to 18% of 38 individuals, in the Holocene they represented at least 82% of 78 individuals. We interpret this change in frequency as a considerable increase in abundance in the surroundings of the cave, after excluding major taphonomical biases. The causes for this phenomenon are uncertain but could have involved a surplus of food availability and predation release due to the late Quaternary extinction event

Extant and extinct Xenarthran skull morphological analysis: evolutionary and functional inferences

Os Xenarthra representam um clado de mamíferos eutérios. Pouco se sabe sobre a evolução morfológica craniana do grupo. Esta tese iniciou os estudos relativos a esta questão com base na genética quantitativa, na morfometria e na sistemática, e teve por objetivos específicos: 1) avaliar empiricamente se as matrizes de variância e covariância fenotípica (matriz-P) dos diversos gêneros de Xenarthra estudados podem ser utilizadas como substitutas das respectivas matrizes de variância e covariância genética aditiva (matriz-G), uma vez que não existem matrizes-G estimadas para os Xenarthra, e também se elas podem ser utilizadas em estudos macroevolutivos; 2) testar se a diversificação morfológica craniana no grupo ocorreu somente através de deriva genética; e 3) compreender como a relação entre os caracteres morfológicos (módulos) e a magnitude geral de integração podem influir na evolução morfológica craniana. Além destes objetivos focados na evolução do grupo, também foi escopo desta tese inferir o hábito alimentar de taxa fósseis do final do Pleistoceno/início do Holoceno para melhorar o conhecimento sobre a ecologia de alguns grupos fósseis. O banco de dados utilizado foi composto por medidas lineares de aproximadamente 1150 espécimes adultos, representando 12 dos 14 gêneros atuais e sete dos diversos gêneros extintos de Xenarthra. Com base nesses dados, matrizes-P de variância e covariância e de correlação foram estimadas para cada gênero. Essas matrizes foram posteriormente comparadas par a par para avaliar a semelhança na estrutura das diferentes matrizes. Também a partir dessas matrizes, foram obtidas as variâncias entre e intra populações para testar se a diversificação morfológica ocorreu de acordo com a expectativa teórica de diversificação sob a ação exclusiva de deriva genética. As mesmas matrizes-P foram comparadas a diferentes matrizes teóricas de hipóteses de modularidade craniana. As matrizes teóricas expressaram a relação entre os caracteres com base no desenvolvimento e/ou desempenho de função compartilhado pelas partes do crânio. Para cada matriz-P de correlação calculou-se a magnitude geral de integração. Além disto, a dieta dos grupos extintos foi inferida através de análises de funções discriminantes a partir da relação entre forma e função dos animais atuais. Os resultados obtidos indicam que as matrizes-P dos diversos gêneros são similares entre si, o que sugere que matrizes-P podem ser utilizadas tanto como substitutas das matrizes-G quanto no contexto macroevolutivo. Os resultados obtidos refutaram a hipótese nula da diversificação morfológica craniana ocorrendo somente por deriva genética, ao menos nos níveis mais inclusivos da filogenia dos Xenarthra. Consequentemente, a seleção natural provavelmente atuou neste processo de diversificação. Os resultados também sugeriram que o crânio desse grupo está organizado em módulos, sendo os módulos mais conspícuos os relacionados à face. Além disso, foi detectada grande variação na magnitude geral de integração entre gêneros. A variação no padrão modular, mas principalmente na magnitude geral de integração, faz com que os gêneros apresentem diferenças nas possíveis capacidades de responder de forma alinhada às pressões seletivas. Por último, as análises morfofuncionais indicaram elevada diversidade de hábitos alimentares entre os Xenarthra extintosXenarthra are an eutherian mammal clade and little is known about their cranial morphological evolution. This thesis has initiated studies related to this topic and, based on quantitative genetics, morphometrics and systematics, aimed to: 1) empirically assess if the phenotypic variance and covariance matrices (P-matrix) of several genera can be used as surrogates for their respective additive genetic variance and covariance matrices (G-matrix), since G-matrices for Xenarthra are not available, and also if P-matrices can be used in macroevolutionary studies; 2) test whether the skull morphological diversification within the group occurred only through genetic drift; and 3) understand how the relationship between the traits (modules) and overall magnitude of integration may influence cranial morphological evolution. Besides these objectives focused on the evolution of the group, it was also within the scope of this thesis to infer the feeding habits of late Pleistocene/early Holocene fossil taxa to better understand the ecology of some fossil groups. The database used consist of linear measurements of approximately 1150 adult specimens, representing 12 of the 14 extant genera and seven of the several extinct genera of Xenarthra. The data gathered were used to estimate variance/covariance and correlation P-matrices for every genus. These matrices were compared between pairs of genera to evaluate the matrices\' structural similarities. Based on these matrices, within and between population variances were obtained and it was tested whether morphological diversification was in accordance to the theoretical expectation of diversification under genetic drift alone. The same matrices were compared to theoretical matrices expressing modularity hypotheses. These theoretical matrices represent the relationship among traits in reference to the shared development and/or function of different skull\'s anatomical regions (modules). For every correlation P-matrix the overall magnitude of integration was calculated. Moreover, the extinct groups\' diet was inferred through discriminant function analysis relying on the relationship between form and function of extant animals. Results indicate that P-matrices from several genera were structurally similar. This suggests that P-matrices can be used as surrogates of their G-matrices and in the macroevolutionary context. Results refuted the null hypothesis of cranial morphological diversification occurring only due to genetic drift, at least in more inclusive levels of Xenarthran phylogeny. Consequently, natural selection probably acted on this diversification process. The results also suggested that the Xenarthran skull is organized in modules, and the most conspicuous modules are in the face region. A large variation in the overall magnitude of integration among genera was detected. The variation in the modular pattern, but especially in the overall magnitude of integration, allows genera to differ in their potential capacity to respond aligned with selective pressures. Finally, morphofunctional analyses indicate a high diversity of feeding habits among extinct Xenarthr

Taxonomic, taphonomic and morphometric contextualization of the megamastofauna bone remains from Cuvieri Cave (MG), a Late Pleistocene Palaeontological Site.

O conhecimento limitado em relação à megafauna brasileira (oriundo de uma tradição acadêmica taxonomista no país) e a falta de escavações estratigraficamente controladas de sítios paleontológicos no Brasil motivou o início das escavações paleontológicas na Gruta Cuvieri, Minas Gerais, através de um método de escavação inédito no país. O presente trabalho analisou o material fóssil exumado de uma das áreas escavadas da gruta (Locus 3), enfocando os seguintes tópicos: (1) descrição e classificação taxonômica dos elementos da megafauna recuperados; (2) estudo tafonômico desse material; e (3) análise da função dos membros anteriores dos grandes mamíferos extintos exumados da gruta e de outros animais depositados em coleções de museus nacionais. Desta forma, tentou-se aumentar o conhecimento em relação à megafauna brasileira em temas pouco explorados. A análise taxonômica teve a finalidade de identificar o material encontrado para permitir o desenvolvimento dos demais tópicos do estudo. A análise tafonômica buscou subsídios em algumas assinaturas tafonômicas para auxiliar a compreensão da formação do pacote fossilífero. Já o estudo da função do membro anterior foi realizado através de morfometria geométrica e estatística multivariada na região distal do úmero de Xenarthra atuais e extintos para inferir a função do membro anterior de algumas espécies extintas, a saber: Catonyx cuvieri, Eremotherium laurilardii, Glossotherium aff. lettsomi, Nothrotherium maquinense, Ocnotherium giganteum, Holmesina paulacoutoi, Pampatherium humboldti e Valgipes bucklandii. Os resultados da análise taxonômica indicaram a presença de 3 espécimes de Tapirus terrestris, 3 de Catonyx cuvieri e 1 de Valgipes bucklandii, revelando uma elevada abundância de fósseis para o local. Através da análise tafonômica concluiu-se que a queda/pisoteamente de animais no interior do Locus 3 teve papel importante na fragmentação e dispersão dos ossos de megafauna e que, provavelmente, uma sedimentação lenta favoreceu este processo. Por último, os resultados morfo-funcionais sugerem que os tatus extintos (Pampatherium e Holmesina ), assim como a grande maioria das preguiças extintas (Catonyx, Glossotherium, Ocnotherium, Valgipes), podiam ser capazes de escavar o substrato ocasionalmente ou apenas remover pequenas quantidades de terra, mas não seriam escavadores especializados, tampouco tinham a habilidade de extensão pronunciada do membro anterior. Já Eremotherium, ao contrário das demais espécies, seria capaz justamente de estender seus braços, não tendo a habilidade de raspar o substrato. Os resultados foram inconclusivos quanto à função dos membros para Nothrotherium.The limited knowledge regarding the Brazilian megafauna (due to a mainly taxonomist tradition of brazilian paleontologists) and the lack of controlled stratigraphic excavations of paleontological sites in Brazil motivated the excavations of Cuvieri Cave, Minas Gerais, using an unprecedented method of excavation in the country, based on detailed exhumation of paleontological remains. This study examined the fossil material exhumed from one fossiliferous areas of the cave (Locus 3), focusing on the following topics: (1) taxonomic classification and description of the megafauna remains recovered, (2) taphonomic study of this material, and (3) functional analysis of large extinct mammals\' forelimbs found in the cave and from other specimens deposited in distinct national museums. The main objective of this study was to increase the knowledge regarding the Brazilian megafauna on these marginalized topics. The taxonomic analysis was intended to identify the bones found in the cave to enable the development of the other parts of the study. The taphonomic analysis were performed to assist the understanding of the formation of the fossiliferous package in Locus 3. Finally, the functional study of the forelimb was performed through morphometric geometric analysis and multivariate statistics applied to the distal region of the humerus of extant and extinct Xenarthra in order to infer the function of the forelimb of some extinct species, namely: Catonyx cuvieri, Eremotherium laurilardii, Glossotherium aff. lettsomi, Nothrotherium maquinense, Ocnotherium giganteum, Holmesina paulacoutoi, Pampatherium humboldti and Valgipes bucklandii. The results of the taxonomic analysis indicated the presence of three specimens of Tapirus terrestris, three of Catonyx cuvieri and one of Valgipes bucklandii, revealing a high density of specimens in the site. Through the taphonomic analysis it was concluded that the fall/trampling of animals within the Locus 3 was very important in the fragmentation and dispersion of the megafauna bones, and also that a potential slow sedimentation favored this process. Finally, the morpho-functional results suggest that the extinct armadillos (Pampatherium and Holmesina) as well as the majority of extinct sloths studied (Catonyx, Glossotherium, Ocnotherium, Valgipes) might have been able to dig occasionally the substrate or only remove small quantities of earth, being neither specialized burrowers nor able to pronounced extension of the forelimb. On the other hand, Eremotherium, unlike the other sloth species, was able to extend its arms, not having the ability to dig the substrate. The results were inconclusive regarding the function of Nothrotherium\'s forelimb

A large Cervidae Holocene accumulation in Eastern Brazil: an example of extreme taphonomi- cal control in a cave environment

A remarkable cervid bone accumulation occurs at a single passage (named Cervid Passage; CP) at Lapa Nova, a maze cave in eastern Brazil. CP lies away from cave entrances, is a typical pitfall passage and contains bone remains of at least 121 cervids, besides few bats, peccaries and rodents remains. There is no evidence of water (or sediment) flow at the site and in general bones lack post depositional alterations and display anatomical proximity, suggesting that the majority of the remains found inside CP (mainly cervids) are due to animals that after entering the cave got trapped in the site. Observations suggest that two entrances could have provided access to cervids (and the few other animals, besides bats), either by falling inside the cave or by entering by their own free will. Once inside the cave, the maze pattern would make route finding difficult, and of all passage intersections, only the one leading to CP would result in a non-return situation, starving the animal to death. Radiocarbon dates suggest that animal entrapment occurred during at least 5 thousand years, during the Holocene. The reasons why mainly cervids were found are unknown but they are probably related to the biology of this group coupled with the fact that caves provide several specific taphonomic processes that may account for a strong bias in bone accumulation. Indeed, the frequent occurrence of Cervidae in both the fossil and sub-fossil record in Brazilian caves may be related to an overall high faunal abundance or may suggest that these animals were especially prone to enter caves, perhaps in search of nutrients (as cave saltpetre) or water

A large cervidae Holocene accumulation in eastern Brazil: An example of extreme taphonomical control in a cave environment

A remarkable cervid bone accumulation occurs at a single passage (named Cervid Passage; CP) at Lapa Nova, a maze cave in eastern Brazil. CP lies away from cave entrances, is a typical pitfall passage and contains bone remains of at least 121 cervids, besides few bats, peccaries and rodents remains. There is no evidence of water (or sediment) flow at the site and in general bones lack post depositional alterations and display anatomical proximity, suggesting that the majority of the remains found inside CP (mainly cervids) are due to animals that after entering the cave got trapped in the site. Observations suggest that two entrances could have provided access to cervids (and the few other animals, besides bats), either by falling inside the cave or by entering by their own free will. Once inside the cave, the maze pattern would make route finding difficult, and of all passage intersections, only the one leading to CP would result in a non-return situation, starving the animal to death. Radiocarbon dates suggest that animal entrapment occurred during at least 5 thousand years, during the Holocene. The reasons why mainly cervids were found are unknown but they are probably related to the biology of this group coupled with the fact that caves provide several specific taphonomic processes that may account for a strong bias in bone accumulation. Indeed, the frequent occurrence of Cervidae in both the fossil and sub-fossil record in Brazilian caves may be related to an overall high faunal abundance or may suggest that these animals were especially prone to enter caves, perhaps in search of nutrients (as cave saltpetre) or water

New Accelerator Mass Spectrometry (AMS) Ages Suggest a Revision of the Electron Spin Resonance (ESR) Middle Holocene Dates Obtained for a Toxodon platensis (Toxodontidae, Mammalia) from Southeast Brazil

In a paper published in Applied Radiation and Isotopes, Baffa et al. (2000) reported a Middle Holocene date (~6.5 kyr BP) for a specimen of Toxodon platensis from Ribeira do Iguape, southeast Brazil, using the emergent technique electron spin resonance (ESR). Through an accelerator mass spectrometry (AMS) procedure applied on tooth collagen, we provide a new set of dates to test the accuracy of the ages generated by ESR. We obtained 2 dates more than 4500 BP older than the previous one, suggesting a minimum Late Pleistocene age for the specimen