Endocranial asymmetry in New World monkeys: a comparative phylogenetic analysis of morphometric data.

Brain lateralization is a widespread phenomenon although its expression across primates is still controversial due to the reduced number of species analyzed and the disparity of methods used. To gain insight into the diversification of neuroanatomical asymmetries in non-human primates we analyze the endocasts, as a proxy of external brain morphology, of a large sample of New World monkeys and test the effect of brain size, home range and group sizes in the pattern and magnitude of shape asymmetry. Digital endocasts from 26 species were obtained from MicroCT scans and a set of 3D coordinates was digitized on endocast surfaces. Results indicate that Ateles, Brachyteles, Callicebus and Cacajao tend to have a rightward frontal and a leftward occipital lobe asymmetry, whereas Aotus, Callitrichinae and Cebinae have either the opposite pattern or no directional asymmetry. Such differences in the pattern of asymmetry were associated with group and home range sizes. Conversely, its magnitude was significantly associated with brain size, with larger-brained species showing higher inter-hemispheric differences. These findings support the hypothesis that reduction in inter-hemispheric connectivity in larger brains favors the lateralization and increases the structural asymmetries, whereas the patterns of shape asymmetry might be driven by socio-ecological differences among species.

Skull diversity and evolution in miniaturized amphibians, genus Brachycephalus (Anura: Brachycephalidae).

Miniaturized amphibians of the genus Brachycephalus are phenotypically diverse. The species of Brachycephalus have bufoniform or leptodactyliform Baupläne and any of three skeletal states: nonhyperossified, hyperossified without dorsal shield, and hyperossified with dorsal shield. We integrate high-resolution microcomputed tomography, geometric morphometrics, and an estimate of molecular phylogenetic relationships to investigate skull diversity in shape and size-shape space in selected species of Brachycephalus. Skull diversity amongst species of Brachycephalus can be partitioned into shape and size-shape space according to the four conditions of skeletal states-Baupläne, namely, nonhyperossified leptodactyliform, nonhyperossified bufoniform, hyperossified bufoniform without dorsal shield, and hyperossified bufoniform with dorsal shield. Skull diversity in shape and size-shape space in nonhyperossified leptodactyliform species of Brachycephalus is markedly larger, when compared to skull diversity in species of the three other conditions of skeletal states-Baupläne. Variation in skull shape scales with size across Brachycephalus and, therefore, can be explained by allometry. Skull diversity, Baupläne, and skeletal states covary to a large extent with monophyletic lineages of Brachycephalus, as revealed by a mitochondrial DNA species tree. Nonhyperossified bufoniform species and hyperossified bufoniform species with or without dorsal shield are monophyletic lineages, as inferred from a mitochondrial DNA species tree. Nonhyperossified leptodactyliform species of Brachycephalus do not share, however, a most recent common ancestor. The nonhyperossified leptodactyliform species of Brachycephalus, due to their marked skull diversity and lack of monophyly, emerge as evolutionarily complex. Therefore, further sampling of the nonhyperossified leptodactyliform condition of skeletal states-Baupläne will be necessary to further understand the evolutionary history of Brachycephalus.

Os anfíbios miniaturizados do gênero Brachycephalus são fenotipicamente diversos. As espécies de Brachycephalus têm o plano corporal bufoniforme ou leptodactyliforme e três estados esqueléticos: não-hiperossificado, hiperossificado sem placa dorsal e hiperossificado com placa dorsal. Neste trabalho nós integramos tomografia micro-computadorizada de alta resolução, morfometria geométrica e uma estimativa de relações filogenéticas moleculares para investigar diversidade craniana nos espaços de forma e tamanho-forma em determinadas espécies de Brachycephalus. A diversidade craniana entre espécies de Brachycephalus pode ser dividida no espaço de forma e tamanho-forma segundo as quatro condições de plano corporal-esqueleto, a saber, leptodactiliforme não-hiperossificado, bufoniforme não-hiperossificado, bufoniforme hiperossificado sem placa dorsal e bufoniforme hiperossificado com placa dorsal. A diversidade craniana nos espaços de forma e tamanho-forma nas espécies de Brachycephalus leptodactiliformes não-hiperossificadas é pronunciadamente maior quando comparada àquela das espécies nas outras trcs condições de plano corporal-esqueleto. A variação na forma craniana aumenta com o tamanho craniano em Brachycephalus e, portanto, pode ser explicada por alometria. Diversidade craniana, plano corporal e estados esqueléticos covariam consideravelmente com as linhagens monofiléticas de Brachycephalus, como estimado pela filogenia mitocondrial. As espécies de Brachycephalus leptodactiliformes não-hiperossificadas e bufoniformes hiperossificadas com ou sem placa dorsal são linhagens monofiléticas, como estimado pela filogenia mitocondrial. As espécies leptodactiliformes não-hiperossificadas não compartilham, todavia, um ancestral comum mais recente. As espécies de Brachycephalus leptodactiliformes não-hiperossificadas devido a sua pronunciada diversidade e não-monofilia emergem claramente como entidades evolutivamente complexas. Por conseguinte, a amostragem adicional de populações leptodactiliformes não-hiperossificadas será necessária para uma melhor compreensão da história evolutiva do gênero Brachycephalus.

Cranial and endocranial diversity in extant and fossil atelids (Platyrrhini: Atelidae): A geometric morphometric study.

OBJECTIVES: Platyrrhines constitute a diverse clade, with the modern Atelidae exhibiting the most variation in cranial and endocast morphology. The processes responsible for this diversification are not well understood. Here, we present a geometric morphometric study describing variation in cranial and endocranial shape of 14 species of Alouatta, Ateles, Brachyteles, and Lagothrix and two extinct taxa, Cartelles and Caipora. METHODS: We examined cranial and endocranial shape variation among species using images reconstructed from CT scans and geometric morphometric techniques based on three-dimensional landmarks and semilandmarks. Principal components analyses were used to explore variation, including the Procrustes shape coordinates, summing the logarithm of the Centroid Size, the common allometric component, and residual shape components. RESULTS: Differences in endocranial shape are related to a relative increase or decrease in the volume of the neocortex region with respect to brainstem and cerebellum regions. The relative position of the brainstem varies from a posterior position in Alouatta to a more ventral position in Ateles. The shape of both the cranium and endocast of Caipora is within the observed variation of Brachyteles. Cartelles occupies the most differentiated position relative to the extant taxa, especially in regards to its endocranial shape. CONCLUSIONS: The pattern of variation in the extant species in endocranial shape is similar to the variation observed in previous cranial studies, with Alouatta as an outlier. The similarities between Caipora and Brachyteles were unexpected and intriguing given the frugivorous adaptations inferred from the fossil's dentition. Our study shows the importance of considering both extant and fossil species when studying diversification of complex traits.

Increased Susceptibility of Gracilinanus microtarsus Liver Mitochondria to Ca²âº-Induced Permeability Transition Is Associated with a More Oxidized State of NAD(P).

In addition to be the cell's powerhouse, mitochondria also contain a cell death machinery that includes highly regulated processes such as the membrane permeability transition pore (PTP) and reactive oxygen species (ROS) production. In this context, the results presented here provide evidence that liver mitochondria isolated from Gracilinanus microtarsus, a small and short life span (one year) marsupial, when compared to mice, are much more susceptible to PTP opening in association with a poor NADPH dependent antioxidant capacity. Liver mitochondria isolated from the marsupial are well coupled and take up Ca(2+) but exhibited a much lower Ca(2+) retention capacity than mouse mitochondria. Although the known PTP inhibitors cyclosporin A, ADP, and ATP significantly increased the marsupial mitochondria capacity to retain Ca(2+), their effects were much larger in mice than in marsupial mitochondria. Both fluorescence and HPLC analysis of mitochondrial nicotinamide nucleotides showed that both content and state of reduction (mainly of NADPH) were lower in the marsupial mitochondria than in mice mitochondria despite the similarity in the activity of the glutathione peroxidase/reductase system. Overall, these data suggest that PTP opening is an important event in processes of Ca(2+) signalling to cell death mediated by mitochondrial redox imbalance in G. microtarsus.

The network organization of protein interactions in the spliceosome is reproduced by the simple rules of food-web models.

The network structure of biological systems provides information on the underlying processes shaping their organization and dynamics. Here we examined the structure of the network depicting protein interactions within the spliceosome, the macromolecular complex responsible for splicing in eukaryotic cells. We show the interactions of less connected spliceosome proteins are nested subsets of the connections of the highly connected proteins. At the same time, the network has a modular structure with groups of proteins sharing similar interaction patterns. We then investigated the role of affinity and specificity in shaping the spliceosome network by adapting a probabilistic model originally designed to reproduce food webs. This food-web model was as successful in reproducing the structure of protein interactions as it is in reproducing interactions among species. The good performance of the model suggests affinity and specificity, partially determined by protein size and the timing of association to the complex, may be determining network structure. Moreover, because network models allow building ensembles of realistic networks while encompassing uncertainty they can be useful to examine the dynamics and vulnerability of intracelullar processes. Unraveling the mechanisms organizing the spliceosome interactions is important to characterize the role of individual proteins on splicing catalysis and regulation.

Pleistocene megafaunal interaction networks became more vulnerable after human arrival.

The end of the Pleistocene was marked by the extinction of almost all large land mammals worldwide except in Africa. Although the debate on Pleistocene extinctions has focused on the roles of climate change and humans, the impact of perturbations depends on properties of ecological communities, such as species composition and the organization of ecological interactions. Here, we combined palaeoecological and ecological data, food-web models and community stability analysis to investigate if differences between Pleistocene and modern mammalian assemblages help us understand why the megafauna died out in the Americas while persisting in Africa. We show Pleistocene and modern assemblages share similar network topology, but differences in richness and body size distributions made Pleistocene communities significantly more vulnerable to the effects of human arrival. The structural changes promoted by humans in Pleistocene networks would have increased the likelihood of unstable dynamics, which may favour extinction cascades in communities facing extrinsic perturbations. Our findings suggest that the basic aspects of the organization of ecological communities may have played an important role in major extinction events in the past. Knowledge of community-level properties and their consequences to dynamics may be critical to understand past and future extinctions.

Encephalization and diversification of the cranial base in platyrrhine primates.

The cranial base, composed of the midline and lateral basicranium, is a structurally important region of the skull associated with several key traits, which has been extensively studied in anthropology and primatology. In particular, most studies have focused on the association between midline cranial base flexion and relative brain size, or encephalization. However, variation in lateral basicranial morphology has been studied less thoroughly. Platyrrhines are a group of primates that experienced a major evolutionary radiation accompanied by extensive morphological diversification in Central and South America over a large temporal scale. Previous studies have also suggested that they underwent several evolutionarily independent processes of encephalization. Given these characteristics, platyrrhines present an excellent opportunity to study, on a large phylogenetic scale, the morphological correlates of primate diversification in brain size. In this study we explore the pattern of variation in basicranial morphology and its relationship with phylogenetic branching and with encephalization in platyrrhines. We quantify variation in the 3D shape of the midline and lateral basicranium and endocranial volumes in a large sample of platyrrhine species, employing high-resolution CT-scans and geometric morphometric techniques. We investigate the relationship between basicranial shape and encephalization using phylogenetic regression methods and calculate a measure of phylogenetic signal in the datasets. The results showed that phylogenetic structure is the most important dimension for understanding platyrrhine cranial base diversification; only Aotus species do not show concordance with our molecular phylogeny. Encephalization was only correlated with midline basicranial flexion, and species that exhibit convergence in their relative brain size do not display convergence in lateral basicranial shape. The evolution of basicranial variation in primates is probably more complex than previously believed, and understanding it will require further studies exploring the complex interactions between encephalization, brain shape, cranial base morphology, and ecological dimensions acting along the species divergence process.

Species tree estimation for a deep phylogenetic divergence in the New World monkeys (Primates: Platyrrhini).

The estimation of a robust phylogeny is a necessary first step in understanding the biological diversification of the platyrrhines. Although the most recent phylogenies are generally robust, they differ from one another in the relationship between Aotus and other genera as well as in the relationship between Pitheciidae and other families. Here, we used coding and non-coding sequences to infer the species tree and embedded gene trees of the platyrrhine genera using the Bayesian Markov chain Monte Carlo method for the multispecies coalescent (*BEAST) for the first time and to compared the results with those of a Bayesian concatenated phylogenetic analysis. Our species tree, based on all available sequences, shows a closer phylogenetic relationship between Atelidae and Cebidae and a closer relationship between Aotus and the Cebidae clade. The posterior probabilities are lower for these conflictive tree nodes compared to those in the concatenated analysis; this finding could be explained by some gene trees showing no concordant topologies between Aotus and the other genera. Moreover, the topology of our species tree also differs from the findings of previous molecular and morphological studies regarding the position of Aotus. The existence of discrepancies between morphological data, gene trees and the species tree is widely reported and can be related to processes such as incomplete lineage sorting or selection. Although these processes are common in species trees with low divergence, they can also occur in species trees with deep and rapid divergence. The sources of the inconsistency of morphological and molecular traits with the species tree could be a main focus of further research on platyrrhines.

Marmosa paraguayana (Marsupialia: Didelphidae) as a new host for Gracilioxyuris agilisis (Nematoda: Oxyuridae) in Brazil.

Didelphids (Didelphimorphia: Didelphidae) are a large and well-studied group of Neotropical marsupials. Although knowledge of the parasitic fauna of didelphids is still scarce, recent work has suggested that Neotropical marsupials are often hosts of pinworms. Here, we isolated oxyurid nematodes from fecal samples of Marmosa paraguayana (Marsupialia: Didelphidae) and provide a general description and measurements for male and female specimens. We concluded these specimens can be assigned to Gracilioxyuris agilisis (Ascaridida: Oxyuridae), an oxyurid recently described as a parasite of the didelphid Gracilinanus agilis (Didelphimorphia: Didelphidae). The finding of G. agilisis in a different, albeit closely related, host species strengthens the previous notion of a close association between pinworms and didelphids and contributes to the knowledge of the helminthic fauna of didelphid marsupials.

Molecular phylogenetic relationships and phenotypic diversity in miniaturized toadlets, genus Brachycephalus (Amphibia: Anura: Brachycephalidae).

Toadlets of the genus Brachycephalus are endemic to the Atlantic rainforests of southeastern and southern Brazil. The 14 species currently described have snout-vent lengths less than 18 mm and are thought to have evolved through miniaturization: an evolutionary process leading to an extremely small adult body size. Here, we present the first comprehensive phylogenetic analysis for Brachycephalus, using a multilocus approach based on two nuclear (Rag-1 and Tyr) and three mitochondrial (Cyt b, 12S, and 16S rRNA) gene regions. Phylogenetic relationships were inferred using a partitioned Bayesian analysis of concatenated sequences and the hierarchical Bayesian method (BEST) that estimates species trees based on the multispecies coalescent model. Individual gene trees showed conflict and also varied in resolution. With the exception of the mitochondrial gene tree, no gene tree was completely resolved. The concatenated gene tree was completely resolved and is identical in topology and degree of statistical support to the individual mtDNA gene tree. On the other hand, the BEST species tree showed reduced significant node support relative to the concatenate tree and recovered a basal trichotomy, although some bipartitions were significantly supported at the tips of the species tree. Comparison of the log likelihoods for the concatenated and BEST trees suggests that the method implemented in BEST explains the multilocus data for Brachycephalus better than the Bayesian analysis of concatenated data. Landmark-based geometric morphometrics revealed marked variation in cranial shape between the species of Brachycephalus. In addition, a statistically significant association was demonstrated between variation in cranial shape and genetic distances estimated from the mtDNA and nuclear loci. Notably, B. ephippium and B. garbeana that are predicted to be sister-species in the individual and concatenated gene trees and the BEST species tree share an evolutionary novelty, the hyperossified dorsal plate.

Asymptotic behavior of the scaled mutation rate estimators.

Important aspects of population evolution have been investigated using nucleotide sequences. Under the neutral Wright-Fisher model, the scaled mutation rate represents twice the average number of new mutations per generations and it is one of the key parameters in population genetics. In this study, we present various methods of estimation of this parameter, analytical studies of their asymptotic behavior as well as comparisons of the distribution's behavior of these estimators through simulations. As knowledge of the genealogy is needed to estimate the maximum likelihood estimator (MLE), an application with real data is also presented, using jackknife to correct the bias of the MLE, which can be generated by the estimation of the tree. We proved analytically that the Waterson's estimator and the MLE are asymptotically equivalent with the same rate of convergence to normality. Furthermore, we showed that the MLE has a better rate of convergence than Waterson's estimator for values of the parameter greater than one and this relationship is reversed when the parameter is less than one.

Hyperossification in miniaturized toadlets of the genus Brachycephalus (Amphibia: Anura: Brachycephalidae): Microscopic structure and macroscopic patterns of variation.

Species of the genus Brachycephalus, have a snout-vent length of less than 18 mm and are believed to have evolved through miniaturization. Brachycephalus ephippium, is particularly interesting; because its entire skull is hyperossified, and the presacral vertebrae and transverse processes are covered by a dorsal shield. We demonstrate in this paper that, at the macroscopic level, a completely hyperossified skull and dorsal shield occur only in B. ephippium, but not in B. ferruginus, B. izechsohni, B. pernix, B. pombali, B. brunneus, B. didactylus, and B. hermogenesi. An intermediate condition, in which the skull is hyperossified but a dorsal shield is absent, occurs in B. vertebralis, B. nodoterga, B. pitanga, and B. alipioi. The microscopic structure of hyperossification was examined in skulls of B. ephippium and B. pitanga, revealing a complex organization involving the presence of Sharpey fibers, which in humans are characteristic of periodontal connections.

Network analysis reveals contrasting effects of intraspecific competition on individual vs. population diets.

Optimal foraging theory predicts that individuals should become more opportunistic when intraspecific competition is high and preferred resources are scarce. This density-dependent diet shift should result in increased diet breadth for individuals as they add previously unused prey to their repertoire. As a result, the niche breadth of the population as a whole should increase. In a recent study, R. Svanbäck and D. I. Bolnick confirmed that intraspecific competition led to increased population diet breadth in threespine stickleback (Gasterosteus aculeatus). However, individual diet breadth did not expand as resource levels declined. Here, we present a new method based on complex network theory that moves beyond a simple measure of diet breadth, and we use the method to reexamine the stickleback experiment. This method reveals that the population as a whole added new types of prey as stickleback density was increased. However, whereas foraging theory predicts that niche expansion is achieved by individuals accepting new prey in addition to previously preferred prey, we found that a subset of individuals ceased to use their previously preferred prey, even though other members of their population continued to specialize on the original prey types. As a result, populations were subdivided into groups of ecologically similar individuals, with diet variation among groups reflecting phenotype-dependent changes in foraging behavior as prey density declined. These results are consistent with foraging theory if we assume that quantitative trait variation among consumers affects prey preferences, and if cognitive constraints prevent individuals from continuing to use their formerly preferred prey while adding new prey.

Vulnerability of a killer whale social network to disease outbreaks.

Emerging infectious diseases are among the main threats to conservation of biological diversity. A crucial task facing epidemiologists is to predict the vulnerability of populations of endangered animals to disease outbreaks. In this context, the network structure of social interactions within animal populations may affect disease spreading. However, endangered animal populations are often small and to investigate the dynamics of small networks is a difficult task. Using network theory, we show that the social structure of an endangered population of mammal-eating killer whales is vulnerable to disease outbreaks. This feature was found to be a consequence of the combined effects of the topology and strength of social links among individuals. Our results uncover a serious challenge for conservation of the species and its ecosystem. In addition, this study shows that the network approach can be useful to study dynamical processes in very small networks.

Interaction intimacy affects structure and coevolutionary dynamics in mutualistic networks.

The structure of mutualistic networks provides clues to processes shaping biodiversity [1-10]. Among them, interaction intimacy, the degree of biological association between partners, leads to differences in specialization patterns [4, 11] and might affect network organization [12]. Here, we investigated potential consequences of interaction intimacy for the structure and coevolution of mutualistic networks. From observed processes of selection on mutualistic interactions, it is expected that symbiotic interactions (high-interaction intimacy) will form species-poor networks characterized by compartmentalization [12, 13], whereas nonsymbiotic interactions (low intimacy) will lead to species-rich, nested networks in which there is a core of generalists and specialists often interact with generalists [3, 5, 7, 12, 14]. We demonstrated an association between interaction intimacy and structure in 19 ant-plant mutualistic networks. Through numerical simulations, we found that network structure of different forms of mutualism affects evolutionary change in distinct ways. Change in one species affects primarily one mutualistic partner in symbiotic interactions but might affect multiple partners in nonsymbiotic interactions. We hypothesize that coevolution in symbiotic interactions is characterized by frequent reciprocal changes between few partners, but coevolution in nonsymbiotic networks might show rare bursts of changes in which many species respond to evolutionary changes in a single species.

Using delta13C stable isotopes to quantify individual-level diet variation.

Individual-level diet variation can be easily quantified by gut-content analysis. However, because gut contents are a 'snapshot' of individuals' feeding habits, such cross-sectional data can be subject to sampling error and lead one to overestimate levels of diet variation. In contrast, stable isotopes reflect an individual's long-term diet, so isotope variation among individuals can be interpreted as diet variation. Nevertheless, population isotope variances alone cannot be directly compared among populations, because they depend on both the level of diet variation and the variance of prey isotope ratios. We developed a method to convert population isotope variances into a standardized index of individual specialization (WIC/TNW) that can be compared among populations, or to gut-content variation. We applied this method to diet and carbon isotope data of four species of frogs of the Brazilian savannah. Isotopes showed that gut contents provided a reliable measure of diet variation in three populations, but greatly overestimated diet variation in another population. Our method is sensitive to incomplete sampling of the prey and to among-individual variance in fractionation. Therefore, thorough sampling of prey and estimates of fractionation variance are desirable. Otherwise, the method is straightforward and provides a new tool for quantifying individual-level diet variation in natural populations that combines both gut-content and isotope data.

The dynamics of intraguild predation in Chrysomya albiceps Wied. (Diptera: Calliphoridae): interactions between instars and species under different abundances of food.

The pattern of larval interaction in blowflies confined with Chrysomya albiceps Wied. and C. rufifacies Maquart can be changed in response to the predatory behaviour of the two species to a contest-type process instead of the scramble competition that usually occurs in blowflies. Facultative predation is a frequent behaviour in C. albiceps and C. rufifacies that occurs as an alternative food source during the larval stage. In this study, we investigated the dynamics of intraguild predation by C. albiceps on other fly species in order to analyse interspecific and intraspecific survival in C. albiceps, C. megacephala and C. macellaria Fabricius. The experimental design of the study allowed us to evaluate how factors such as species, density and abundance of food influenced the survival of the calliphorid species. When C. albiceps was confined with C. megacephala or C. macellaria, only adults of C. albiceps survived at different larval densities and abundance of food. In addition, the survival of C. albiceps was higher in two-species experiments when compared to single species experiments. The implications of these results for the dynamics of C. albiceps were discussed.

Evolutionary integration and morphological diversification in complex morphological structures: mandible shape divergence in spiny rats (Rodentia, Echimyidae).

The rodent mandible has become a paradigm for studies on the development and evolution of complex morphological structures. We use a combination of geometric and multivariate morphometric methods in order to assess the correspondence between integration patterns and a priori biological models in the context of evolutionary shape divergence in the mandible of rodents of the family Echimyidae. The correlation of shape distances among operational taxonomic units (individuals, species, genera) in separate morphogenetic components allowed the construction of integration matrices among mandible components for data sets corresponding to varying levels of genetic divergence (intergeneric, interspecific, and intrapopulational). The integration matrices were associated with a priori biological (developmental, genetical, modular) models, and the maximum integration axes (singular warps) were compared with realized axes of maximum interspecific variation (relative warps). The integration pattern and intensity were not stable in data sets with different levels of genetic divergence, and the varying functional demands during the ecological radiation in the family were probably responsible for the differences in observed integration patterns. Developmental and genetic models were significantly associated with the interspecific integration patterns observed, suggesting a role for neutral evolution during the evolutionary divergence of mandible shape. However, directional and stabilizing selection were not discarded as processes responsible for the generation of interspecific integration. The choreography of the morphogenetic components in the mandible is highly flexible and the integrated groups of components can be reorganized depending on functional demands during evolutionary shape changes.

Geometric estimates of heritability in biological shape.

The recently developed geometric morphometrics methods represent an important contribution of statistics and geometry to the study of biological shapes. We propose simple protocols using shape distances that incorporate geometric techniques into linear quantitative genetic models that should provide insights into the contribution of genetics to shape variation in organisms. The geometric approaches use Procrustes distances in a curved shape space and distances in tangent spaces within and among families to estimate shape heritability. We illustrate the protocols with an example of wing shape variation in the honeybee, Apis mellifera. The heritability of overall shape variation was small, but some localized components depicting shape changes on distal wing regions showed medium to large heritabilities. The genetic variance-covariance matrix of the geometric shape variables was significantly correlated with the phenotypic shape variance-covariance matrix. A comparison of the results of geometric methods with the traditional multivariate analysis of interlandmark distances indicated that even with a larger dimensionality, the interlandmark distances were not as rich in shape information as the landmark coordinates. Quantitative genetics studies of shape should greatly benefit from the application of geometric methods.