Morphometric analysis of Chilean blue whales and implications for their taxonomy

In the Southern Hemisphere, blue whales are currently divided into two subspecies, Antarctic blue whales (Balaenoptera musculus intermedia) and pygmy blue whales (B. m. brevicauda), but there is some debate about whether Chilean blue whales should also be considered as a separate subspecies. Here, we provide novel morphometric data to directly address this taxonomic question from a biological survey of 60 blue whales taken during the 1965/1966 Chilean whaling season. The data show that maximum body length and mean body length of both sexually mature females and males for Chilean blue whales are intermediate between pygmy and Antarctic blue whales; and that fluke-anus lengths of Chilean blue whales are significantly different from pygmy blue whales, but not necessarily from Antarctic blue whales. There is also some support from the data that snout-eye measurements are different among all three groups. These data provide further confirmation that Chilean blue whales are a distinct population requiring separate management from other blue whale populations, and are also consistent with suggestions that Chilean blue whales are not the same subspecies as pygmy blue whales.     

Tempo and mode of evolutionary divergence in modern and Holocene Vancouver Island marmots (Marmota vancouverensis) (Mammalia, Rodentia)

Marmota vancouverensis is the only insular species among the 14 species of marmots. The evolutionary history of this species is unresolved. Although M. vancouverensis is strongly differentiated in osteological and other morphological characters, its low genetic divergence suggests recent evolution from an ancestral continental species. We used geometric morphometric techniques to assess the morphology of hemimandibles from 239 modern M. vancouverensis , Marmota caligata , Marmota flaviventris , Marmota olympus and 30 Holocene (9435–735 cal. yr bp) subfossil M. vancouverensis . Our results confirm that the mandible of M. vancouverensis is strongly differentiated in shape from continental marmot species, but similar in size to its mainland sister species M. caligata . Temporal variation in size and shape over the past 2500 years among allochronic samples of M. vancouverensis was minimal suggesting that the morphological divergence of this species occurred in a period of rapid change following its isolation from mainland populations in the late Pleistocene. Selection pressures associated with environmental changes and founder effects and genetic drift resulting from population bottlenecks created by population declines and habitat fragmentation are hypothesized as factors contributing to the morphological divergence of this species.     

Geometric morphometrics of mandibles for dietary differentiation of Bovidae (Mammalia: Artiodactyla)

The mammalian family Bovidae has been widely studied in ecomorphological research, with important applications to paleoecological and paleohabitat reconstructions. Most studies of bovid craniomandibular features in relation to diet have used linear measurements. In this study, we conduct landmark-based geometric-morphometric analyses to evaluate whether different dietary groups can be distinguished by mandibular morphology. Our analysis includes data for 100 species of extant bovids, covering all bovid tribes and 2 dietary classifications. For the first classification with 3 feeding categories, we found that browsers (including frugivores), mixed feeders, and grazers are moderately well separated using mandibular shape. A finer dietary classification (frugivore, browser, browser–grazer intermediate, generalist, variable grazer, and obligate grazer) proved to be more useful for differentiating dietary extremes (frugivores and obligate grazers) but performed equally or less well for other groups. Notably, frugivorous bovids, which belong in tribe Cephalophini, have a distinct mandibular shape that is readily distinguished from all other dietary groups, yielding a 100% correct classification rate from jackknife cross-validation. The main differences in mandibular shape found among dietary groups are related to the functional needs of species during forage prehension and mastication. Compared with browsers, both frugivores and grazers have mandibles that are adapted for higher biomechanical demand of chewing. Additionally, frugivore mandibles are adapted for selective cropping. Our results call for more work on the feeding ecology and functional morphology of frugivores and offer an approach for reconstructing the diet of extinct bovids.     

Functional plasticity of the human humerus: Shape,rigidity, and muscular entheses

The relationship between the mechanical loading undergone by a bone and its form has been widely assumed as a premise in studies aiming to reconstruct behavioral patterns from skeletal remains. Nevertheless, this relationship is complex due to the existence of many factors affecting bone structure and form, and further research combining structural and shape characteristics is needed. Using two‐block PLS, which is a test to analyze the covariance between two sets of variables, we aim to investigate the relationship between upper‐limb entheseal changes, cross‐sectional properties, and contour shape of the humeral diaphysis. Our results show that individuals with strongly marked entheseal changes have increased diaphyseal rigidities. Bending rigidities are mainly related to entheseal changes of muscles that cross the shoulder. Moreover, the entheseal changes of muscles that participate in the rotation of the arm are related to mediolaterally flatter and ventrodorsally broader humeral shapes in the mid‐proximal diaphysis. In turn, this diaphyseal shape is related to diaphyseal rigidity, especially to bending loadings. The shape of the diaphysis of the rest of the humerus does not covary either with rigidity or with entheseal changes. The results indicate that large muscular scars, such as those found in the mid‐proximal diaphyses, seem to be related to diaphyseal shape, whereas this relationship is not seen for areas with less direct influences of powerful muscles. Am J Phys Anthropol 150:609–617, 2013. © 2013 Wiley Periodicals, Inc.     

Ecological preference between generalist and specialist rodents: spatial and environmental correlates of phenotypic variation

Different ecological preferences among species may result in differences in response to similar environmental variation. To test this hypothesis, we assessed the patterns of skull and mandible size and shape variation in three Sigmodontinae mice from agroecosystems of central Argentina with increasing degree of specialization: Calomys musculinus, Akodon azarae and Oxymycterus rufus. Spatial patterns in size and shape were analysed after controlling for allometry and sexual dimorphism using a total of 697 specimens. We then evaluated the covariation between shape, climatic and environmental variables and assessed the contribution of distinct climatic and environmental variables to phenotypic variability. Oxymycterus rufus displayed a marked spatial structure, and there was a high correlation between shape, climatic and environmental variables in this species. Climatic and environmental variables had a moderate effect on the phenotype of A. azarae, and were not correlated with morphological variation in C. musculinus. Our study highlights the difference in phenotypic responses to spatial and environmental gradients across coexisting species, specialist species displaying a more marked spatial structure in morphology than generalist species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 112 , 180–203.     

Shape analysis of symmetric structures: quantifying variation among individuals and asymmetry

Abstract.— Morphometric studies often consider parts with internal left-right symmetry, for instance, the vertebrate skull. This type of symmetry is called object symmetry and is distinguished from matching symmetry, in which two separate structures exist as mirror images of each other, one on each body side. We explain a method for partitioning the total shape variation of landmark configurations with object symmetry into components of symmetric variation among individuals and asymmetry. This method is based on the Procrustes superimposition of the original and a reflected copy of each landmark configuration and is compatible with the two-factor ANOVA model customary in studies of fluctuating asymmetry. We show a fully multivariate framework for testing the effects in the two-factor model with MANOVA statistics, which also applies to shapes with matching symmetry. We apply the new methods in a small case study of pharyngeal jaws of the Neotropical cichlid fish Amphilophus citrinellus . The analysis revealed that the symmetric component of variation in the pharyngeal jaws is dominated by the contrast between two alternative trophic morphs in this species and that there is subtle but statistically significant directional asymmetry. Finally, we provide some general recommendations for morphometric studies of symmetric shapes.     

Post‐weaning ontogeny of the mandible in fossorial water voles: ecological and evolutionary implications

Ventura, J. and Casado‐Cruz, M. 2011. Post‐weaning ontogeny of the mandible in fossorial water voles: ecological and evolutionary implications. —Acta Zoologica (Stockholm) 92 : 12–20. Geometric morphometrics was applied to the mandible of fossorial water voles (Arvicola terrestris monticola) to determine size and shape variations in this structure during post‐weaning ontogeny. The sample consisted of collection specimens obtained in the Aran Valley (Spain), which were grouped into six age classes. Mandible size and shape did not differ significantly between sexes, but between age classes. Mandible size accounted significantly for the shape variation. After the size‐related differences were removed, the mandible shape did not show significant sexual dimorphism but differences by age remained significant. The main shape changes occur between the third and tenth weeks of life and are related to the shift from suckling to a herbivorous diet. Although mandible shape was less remodelled after that age, an appreciable variation also occurs during adulthood. Age‐related changes lead to enhancing the digging potential of the mandible, which in adults becomes a robust structure with an increased surface and stronger crests for muscle insertion. As part of the mandible shape variation was not related to the size‐dependent adjustment and diet does not vary significantly between juvenile and adult voles, shape changes that occur during adulthood can be related to the mechanical stress derived from digging activities.     

Morphological evolution in the mandible of spiny rats, genus Trinomys (Rodentia: Echimyidae)

The development and evolution of the rodent mandible have been studied in depth in recent years. The mandible is a complex structure because it consists of six morphogenetic components formed by different condensations of mesenchymal cells. Using recent techniques for the geometric analysis of shape, we have combined developmental information with a powerful quantification of shape variation and an independent estimate of phylogeny (molecular data) to assess the evolutionary patterns of shape change in mandibles of the rodent genus Trinomys . In general, the major trends in shape variation did not agree with the expected phylogenetic pattern. However, for small-scale morphological differences, one species ( T. yonenagae ) was responsible for the lack of association between morphology and molecular divergence. This species is genetically similar to but morphologically different from other Trinomys . The coronoid process was considered to be the most conservative morphogenetic component in the mandible.     

Evolutionary persistence of phenotypic integration: influence of developmental and functional relationships on complex trait evolution

Examination of historical persistence of integration patterns provides an important insight into understanding the origin and evolution of complex traits. Specifically, the distinct effects of developmental and functional integration on the evolution of complex traits are often overlooked. Because patterns of functional integration are commonly shaped by selection exerted by the external environment, whereas patterns of developmental integration can be determined by relatively environment-independent selection for developmental homeostasis, examination of historical persistence of morphological integration patterns among species should reveal the relative importance of current selection in the evolution of complex traits. We compared historical persistence of integration patterns produced by current developmental versus ecological requirements by examining the evolution of complex mandibular structures in nine species of soricid shrews. We found that, irrespective of phylogenetic relatedness of species, patterns of developmental and functional integration were highly concordant, suggesting that strong selection for developmental homeostasis favors concordant channeling of both internal and external variation. Overall, our results suggest that divergence in mandible shape among species closely follows variation in functional demands and ecological requirements regardless of phylogenetic relatedness among species.     

Cranial and mandibular morphology of Middle Pleistocene cave bears (Ursus deningeri): implications for diet and evolution

ABSTRACT

Deninger’s bears (Ursus deningeri) have been studied less frequently than Ursus spelaeus s.l. Our objective is to present, for the first time, an analysis of the skull shape of U. deningeri.

Bear crania and mandibles were digitised with a Microscribe or CT-scanned and the surface models subsequently landmarked. The landmarks were chosen based on a compromise between functional morphology and sample size.

Results show that U. deningeri and U. spelaeus mandibles display very similar morphologies and allometric trajectories, both to each other and to Ailuropoda melanoleuca. It is inferred that masticatory adaptations to a herbivorous diet were already present in the Middle Pleistocene. U. deningeri displays a cranial morphology that is similar to that of U. spelaeus when comparing all species, but U. deningeri has a relatively narrower and dorsoventrally lower zygomatic arch than U. spelaeus, although the masticatory signal is less strong in the skull.

We observe intraspecific differences between different populations of U. deningeri, which could parallel the genetic diversity found in U. spelaeus. The intraspecific differences found within U. deningeri may be temporal and/or geographical in nature and could be related to the evolution of the Late Pleistocene cave bear, but this hypothesis remains to be tested.     

Pectoral girdle and forelimb variation in extant Crocodylia: the coracoid–humerus pair as an evolutionary module

To date, all statements about evolutionary morphological transformation in Crocodylia have essentially been based on qualitative observations. In the present study, we assessed the morphological variation and covariation (integration) between the scapula, coracoid, humerus, radius, and ulna of 15 species of Crocodylidae, Alligatoridae, and Gavialis + Tomistoma using three‐dimensional geometric morphometrics. The results obtained reveal that the variation of elements within species (intraspecific) is large. However, despite this variability, variation across species (interspecific) is mainly concentrated in two dimensions where the disparity is constrained: ‘robusticity’ and ‘twist’ (forelimbs) and ‘robusticity’ and ‘flexion’ (pectoral girdle). Robusticity (first dimension of variation) embodies a set of correlated geometrical features such as the broadening of the girdle heads and blades, or the enlargement of proximal and distal bone ends. The twist is related to the proximal and/or distal epiphyses in the forelimb elements, and flexion of the scapula and coracoid blades comprises the second dimension of variation. In all crocodylians, forelimb integration is characterized by the strong correlations of a humerus–ulna–radius triad and by a radius–ulna pair, thus forming a tight forelimb module. Unexpectedly, we found that the humerus and coracoid form the most integrated pair, whereas the scapula is a more variable and relatively independent element. The integration pattern of the humerus–coracoid pair distinguishes a relatively robust configuration in alligatorids from that of the remainder groups. The patterns of variation and integration shared by all the analyzed species have been interpreted as an inherited factor, suggesting that developmental and functional requirements would have interacted in the acquisition of a semi‐aquatic and versatile locomotion at the Crocodylia node at least 65 Mya. Our findings highlight the need to incorporate the humerus–coracoid pair in biodynamic and biomechanical studies. © 2012 The Linnean Society of London     

Craniometric variation of some Mediterranean and Atlantic populations of Stenella coeruleoalba (Mammalia,Delphinidae): A three‐dimensional geometric morphometric analysis

The variability of cranial features of Atlantic and Mediterranean samples of Stenella coeruleoalba was examined using a three‐dimensional geometric morphometric approach. Data were collected on 79 skulls from the upper and middle Mediterranean Sea, the Atlantic French coasts, and Scotland. Three‐dimensional x, y, and z coordinates of 27 landmarks were recorded on each left half skull using a Microscribe 3‐D digitizer. All configurations were rotated, centered, and scaled, and residuals from the mean configuration were analyzed through multivariate analyses of variance. Mahalanobis distances among populations were used to evaluate phenetic relationships. Consensus configurations were compared to visualize shape differences among samples. Analyses revealed significant differences among populations, a clear distinction of the Scottish coasts dolphins from the other samples, and a closer relationship of the dolphins from the French coasts to the Mediterranean populations than to the Scottish one. Shape differences are mainly concentrated in the rostral and in the occipital regions of the skull. Phylogenetic and adaptive factors were invoked as possible causes of the variation patterns.     

Shape and size variation: Growth and development of the dusky grouper (Epinephelus marginatus Lowe, 1834)

Demersal fishes have complex life cycles that involve an ontogenetic change in morphology, physiology, and behavior, as their pelagic larval stages colonize benthic habitats. The developmental transition between larvae and juveniles leads to very complex processes of morphogenesis and differentiation. These processes primarily determine changes in external morphology, which is shaped by selective pressures to optimize performance for basic activities such as swimming, escape from predators, and feeding. Fishes have provided fertile grounds for ecomorphological investigations throughout ontogeny, as the role of changing morphology in inducing ontogenetic niche shifts is not always clear. In this framework, some studies have demonstrated that certain species undergo gradual changes, whereas other species experience threshold effects in their ecomorphological relationships during ontogeny. In this study, the intraspecific allometry of the dusky grouper was examined. Geometric morphometric tools were used to quantify shape changes through the development, and a modular approach was also applied to analyze the pattern of covariation between three distinct blocks (head, trunk, and tail). For this purpose, a two‐block Partial Least Square was computed. This method reveals that the pattern of changes in the overall body shape is the result of the modularized changes of these blocks. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Evolutionary strategies of odontocete brain development

The development of the brain was studied in several species of toothed whales (harbour porpoise, spotted dolphin, narwhal, and sperm whale). As embryos, odontocetes show the general mammalian bauplan. The olfactory bulb, lacking in adult toothed whales, forms in embryos but then disappears in early fetal development. In contrast, the terminal nerve persists and shows a great increase in neuron number. Some components of the limbic system are reduced while others grow to become relatively large. The pyramidal tract is inconspicuous. The auditory system and the extrapyramidal system show rapid size increase. In the sperm whale, growth of the telencephalic hemispheres relative to the size of the brain as a whole (telencephalization) and expansion of the cortical areas (neocorticalization) are greater than in other mammals. Increases in the diameters of the cranial nerves seem to be correlated with their presumed functional roles in the postnatal animal.     

Mandibular shape variation in mainland and insular hylobatids

Although hylobatids are the most speciose of the living apes, their morphological interspecies and intraspecies variation remains poorly understood. Here, we assess mandibular shape variation in two species of Hylobates, white-handed (Hylobates lar) and black-handed (Hylobates agilis) gibbons. Using 71 three-dimensional landmarks to quantify mandibular shape, interspecies and intraspecies variation and geographic patterns of mandibular shape are examined in a mixed sex sample of adult H. lar and H. agilis through generalized Procrustes analysis, Procrustes analysis of variance, and principal components analysis. We find that relative to H. agilis, H. lar exhibits a higher amount of variation in mandibular shape. Both species demonstrate similar allometric patterns in mandibular shape. We also highlight a geographic pattern in mandibular shape variation. Compared to mainland hylobatids, insular hylobatids have relatively lower, more posteriorly oriented, and anteroposteriorly wider mandibular condyles, with an increased distance between the condyles and the coronoid processes. This geographic pattern could reflect differences in functional demands on the mandible during mastication and/or could be driven by factors often associated with evolutionary pressures of island populations relative to mainland populations. The findings of this study highlight how little is known about Hylobates morphological variation and how important this is for using Hylobates to help interpret the primate fossil record. Understanding interspecific and intraspecific variation in extant primates is vital to interpreting variation in the primate fossil record.     

Divergent in shape and convergent in function: Adaptive evolution of the mandible in Sub‐Antarctic mice

Convergent evolution in similar environments constitutes strong evidence of adaptive evolution. Transported with people around the world, house mice colonized even remote areas, such as Sub‐Antarctic islands. There, they returned to a feral way of life, shifting towards a diet enriched in terrestrial macroinvertebrates. Here, we test the hypothesis that this triggered convergent evolution of the mandible, a morphological character involved in food consumption. Mandible shape from four Sub‐Antarctic islands was compared to phylogeny, tracing the history of colonization, and climatic conditions. Mandible shape was primarily influenced by phylogenetic history, thus discarding the hypothesis of convergent evolution. The biomechanical properties of the jaw were then investigated. Incisor in‐lever and temporalis out‐lever suggested an increase in the velocity of incisor biting, in agreement with observations on various carnivorous and insectivorous rodents. The mechanical advantage related to incisor biting also revealed an increased functional performance in Sub‐Antarctic populations, and appears to be an adaptation to catch prey more efficiently. The amount of change involved was larger than expected for a plastic response, suggesting microevolutionary processes were evolved. This study thus denotes some degree of adaptive convergent evolution related to changes in habitat‐related changes in dietary items in Sub‐Antarctic mice, but only regarding simple, functionally relevant aspects of mandible morphology.