Nick Milne

Structure and function in armadillo limbs (Mammalia: Xenarthra: Dasypodidae)

Armadillos are a very diverse group ranging from non-fossorial Tolypeutini through to the powerful diggers like the giant armadillo Priodontes maximus to the totally subterranean Chlamyphorini. A previous study demonstrated a close relationship between the relative length of the olecranon of the ulna (index of fossorial ability, IFA) and the fossorial ability of armadillos. This study examines a wide range of limb proportions to explore the biomechanical correlates with fossorial ability. The study demonstrates that the indices of the forelimb (brachial index, shoulder moment index and IFA) do correlate well with digging habits, but also reveals some interesting exceptions, particularly in the most fossorial and most cursorial forms. On the other hand, the hindlimb indices apparently do not correlate with digging habits, but seem to be influenced more by body size. The correlations among the forelimb indices are quite strong and positive but correlations between forelimb and hindlimb indices are negative or very low. It is apparent that there is still much to be learned about structure and function in armadillos.

Scaling of form and function in the xenarthran femur: a 100-fold increase in body mass is mitigated by repositioning of the third trochanter

How animals cope with increases in body size is a key issue in biology. Here, we consider scaling of xenarthrans, particularly how femoral form and function varies to accommodate the size range between the 3 kg armadillo and its giant relative the 300 kg glyptodont. It has already been noted that femoral morphology differs between these animals and suggested that this reflects a novel adaptation to size increase in glyptodont. We test this idea by applying a finite element analysis of coronal plane forces to femoral models of these animals, simulating the stance phase in the hind limb; where the femur is subject to bending owing to longitudinal compressive as well as abduction loads on the greater trochanter. We use these models to examine the hypothesis that muscles attaching on the third trochanter (T3) can reduce this bending in the loaded femur and that the T3 forces are more effective at reducing bending in glyptodont where the T3 is situated at the level of the knee. The analysis uses traditional finite element methods to produce strain maps and examine strains at 200 points on the femur. The coordinates of these points before and after loading are also used to carry out geometric morphometric (GM) analyses of the gross deformation of the model in different loading scenarios. The results show that longitudinal compressive and abductor muscle loading increases bending in the coronal plane, and that loads applied to the T3 reduce that bending. In the glyptodont model, the T3 loads are more effective and can more readily compensate for the bending owing to longitudinal and abductor loads. This study also demonstrates the usefulness of GM methods in interpreting the results of finite element analyses.

Adaptation in the vertebral column: a comparative study of patterns of metameric variation in mice and men

In this paper we examine metamerism in the vertebral column of certain mammals from the perspectives of development and adaptation. To this end we examine the patterns of metameric variation of dimensions of the neural (vertebral) canal, vertebral body and spinous process in man and inbred strains of mice. The data from inbred strains of mice indicate that variability in dimensions within a strain reflects the temporal ordering and nature of developmental influences on vertebral morphogenesis. Differences between strains parallel the within‐strain findings. These findings are attributed to somatic and neural influences on morphogenesis. Comparisons between mice and man indicate that these same influences can be invoked to explain and interpret the mosaic nature of vertebral column evolution. These findings lead us to conclude that different vertebral elements and levels are subject to different interactions of evolutionary and morphogenetic influences. The study of these influences and their interactions should prove fruitful in developing an understanding of the relationship between adaptation, development, growth and function in the skeleton generally.

Allometric and Group Differences in the Xenarthran Femur

The aim of this study is to analyze shape variation in the xenarthran femur to gain insights into their behavior and locomotion. Specimens of both Cingulata (armadillos and glyptodonts) and Pilosa (anteaters and sloths) were studied and within each group body mass varies by several orders of magnitude. The main focus of the analysis was allometric variation in femoral shape in the three groups studied, armadillos, glyptodonts, and pilosans. Three dimensional coordinates were recorded for 40 homologous landmarks on each of 51 xenarthran femurs. The data were analyzed by geometric morphometric methods, and form space analysis was used to identify the allometric variation in each of the three groups. Across all groups, larger specimens tended to have larger articular surfaces, more robust femora generally, and the shape of the femoral condyles was more suited to extended postures. In addition, in larger specimens the medial condyle was much larger than the lateral condyle and the third trochanter was located more distally. The larger armadillo femora had a greater trochanter located considerably proximal to the femoral head and this is thought to improve femoral extension, but in glyptodonts and pilosans the larger specimens had a greater trochanter that was far lateral to the femoral head and this is interpreted as enhancing femoral rotation.

What's wrong with this picture? an experiment in quantifying accuracy in 2D portrait drawing

Traditional portraiture aims to produce a life-like representation of an individual’s unique facial features, but there are a number of perceptual factors that may affect how an artist sees and depicts a sitter’s facial shapes. The anatomical accuracy of a portrait is predominantly ascertained subjectively, and not through quantitative methods. To see if it was possible to apply the methods of analysis more typically used to study variation in biological forms, the authors produced 30 portrait drawings based on 30 pose photographs, and analysed how the portrait drawings differed from the photographs using three different measures: visual assessment, anthropometric measures, and geometric morphometric analysis. Of the three methods, geometric morphometric analysis was found to be the far better technique to account for patterns of depiction common to one artist, and our study indicates this method may have broad research applicability to the face as it is variously represented across a range of visual media.

Discrete trait variation in Western Australian Aboriginal skulls

The non-metrical features of 125 Western Australian Aboriginal skulls in four regional samples have been analysed. An analysis of variance revealed significant differences in the trait frequencies between the sexes. The data for males and females were analysed separately by a modified Grewal-Smith technique (Green et al., 1979) . The relationships between the samples were similar for both sexes. The inland sample (east) was shown to diverge greatly from the three coastal samples (north, central and south). On the basis of the sizes of the standard deviations of the divergences for each trait in the six pairwise comparisons, the number of traits was reduced to 16 for males and 20 for females. The reduction did not alter the relationships found. Extending the analysis to include samples from coastal New South Wales and Queensland (adapted from Larnach and Macintosh, 1966 , Larnach and Macintosh, 1970 again showed east to be the most divergent of the Western Australian samples. The east sample also showed the greatest divergences from New South Wales and Queensland, between which the divergences were very small. These and other inter-relationships essentially support the craniometric and odontometric studies of similar samples by Margetts & Freedman (1977) and Freedman & Lofgren (1981) .

A cranial morphometric study of deer (Mammalia, Cervidae) from Argentina using three-dimensional landmarks.

This study examines morphological variation in the crania (n = 70) of eight cervid species from Argentina. Forty 3-dimensional landmarks were acquired on each adult cervid cranium. The data were analysed using Morphologika software. The co-ordinates were registered and scaled to remove size differences by Procrustes analysis, and then principal components analysis was applied to examine shape variation. Shape variation associated with each principal component can be visualised in the program. The first principal component correlates strongly with the centroid size of the crania and also with the body mass and height of each species. The larger species were distinguished by relatively longer snouts and relatively smaller brains. The smallerMazama andPudu species cluster closely on the first as well as the other principal components. Among the larger species, the twoHippocamelus species, which live at higher altitudes, were clearly distinguished from the lowland species,Ozotoceros bezoarticus andBlastocerus dichotomus, on the basis of cranial flexion and the orientation of the occipital region. FinallyO. bezoarticus andB. dichotomus were compared directly and small differences were noted in the orbital region. The shape data was used to produce a distance matrix and a phenogram, which we relate to some of currently accepted phylogenetic relationships of this group of cervids.

A three-dimensional geometric morphometric analysis of variation in cranial size and shape in tammar wallaby (Macropus eugenii) populations

This study uses geometric morphometric techniques to examine cranial size and shape variation in nine isolated populations of the tammar wallaby (Macropus eugenii). A set of 36 three-dimensional landmarks were digitised on 143 tammar crania from two mainland and seven island populations. While there was no evidence of island dwarfism or gigantism, cranial size increased with both increasing island size and increasing latitude. As latitude increased, the palate narrowed relative to the nasal bones, cranial flexion and nasal height increased, and the zygomatic arches spread out laterally from the cranium. Overall, the anterior nasal aperture (nares) narrowed with increasing latitude. Mean shapes were calculated for each population, and pair-wise comparisons were made; most of these were significantly different. There was a clear tendency for island populations and those with greater geographic separation to show greater shape differentiation. Thus, regional and population differences in the cranial size and shape of tammar wallabies provide examples of selection, founder effect and random genetic drift.

Curved bones: An adaptation to habitual loading

Why are long bones curved? It has long been considered a paradox that many long bones supporting mammalian bodies are curved, since this curvature results in the bone undergoing greater bending, with higher strains and so greater fracture risk under load. This study develops a theoretical model wherein the curvature is a response to bending strains imposed by the requirements of locomotion. In particular the radioulna of obligate quadrupeds is a lever operated by the triceps muscle, and the bending strains induced by the triceps muscle counter the bending resulting from longitudinal loads acting on the curved bone. Indeed the theoretical model reverses this logic and suggests that the curvature is itself a response to the predictable bending strains induced by the triceps muscle. This, in turn, results in anatomical arrangements of bone, muscle and tendon that create a simple physiological mechanism whereby the bone can resist the bending due to the action of triceps in supporting and moving the body. The model is illustrated by contrasting the behaviour of a finite element model of a llama radioulna to that of a straightened version of the same bone. The results show that longitudinal and flexor muscle forces produce bending strains that effectively counter strains due to the pull of the triceps muscle in the curved but not in the straightened model. It is concluded that the curvature of these and other curved bones adds resilience to the skeleton by acting as pre-stressed beams or strainable pre-buckled struts. It is also proposed that the cranial bending strains that result from triceps, acting on the lever that is the radioulna, can explain the development of the curvature of such bones.

Forelimb bone curvature in terrestrial and arboreal mammals

It has recently been proposed that the caudal curvature (concave caudal side) observed in the radioulna of terrestrial quadrupeds is an adaptation to the habitual action of the triceps muscle which causes cranial bending strains (compression on cranial side). The caudal curvature is proposed to be adaptive because longitudinal loading induces caudal bending strains (increased compression on the caudal side), and these opposing bending strains counteract each other leaving the radioulna less strained. If this is true for terrestrial quadrupeds, where triceps is required for habitual elbow extension, then we might expect that in arboreal species, where brachialis is habitually required to maintain elbow flexion, the radioulna should instead be cranially curved. This study measures sagittal curvature of the ulna in a range of terrestrial and arboreal primates and marsupials, and finds that their ulnae are curved in opposite directions in these two locomotor categories. This study also examines sagittal curvature in the humerus in the same species, and finds differences that can be attributed to similar adaptations: the bone is curved to counter the habitual muscle action required by the animal’s lifestyle, the difference being mainly in the distal part of the humerus, where arboreal animals tend have a cranial concavity, thought to be in response the carpal and digital muscles that pull cranially on the distal humerus.