Evidence supporting an evolutionary trade-off between material properties and architectural design in Anolis lizard long bones.

In biology, "many-to-one mapping" occurs when multiple morphological forms can meet a particular functional demand. Knowledge of this mapping is crucial for understanding how selection on performance shapes the evolution of morphological diversity. Past research has focused primarily on the potential for geometrically alternative morphological designs to produce equivalent performance outcomes. Here, we ask whether the material properties of biological tissues hold similar potential. Through a phylogenetic comparative study of Anolis lizards, we show that the architectural design and mineral density of the femur trade off in a many-to-one functional system, yielding a morphospace featuring parallel isolines in size-relative bending strength. Anole femur evolution has largely tracked a narrow band of strength isolines over phylogenetic timescales, suggesting that geometry and mineral content shape the course of macroevolution through compensatory effects on performance. Despite this conserved evolutionary relationship, insular and continental species evolve strong bones differently, likely reflecting underlying ecological differences. Mainland anoles, which exhibit fast-paced life histories, typically have femora with lower mineralization and thinner walls than island species, which exhibit the opposite strategy. Together, our results reveal an overlooked dimension in the relationship between form and function, expanding our understanding of how many-to-one mapping can shape patterns of phenotypic diversity.

Ovariectomized rat model and shape variation in the bony labyrinth.

Postmenopausal osteoporosis is a serious concern in aging individuals, but has not been explored for its potential to alter the shape of the inner ear by way of increased remodeling in the otic capsule. The otic capsule, or bony labyrinth, is thought to experience uniquely limited remodeling after development due to high levels of osteoprotegerin. On this basis, despite the widespread remodeling that accompanies osteoporosis, we hypothesize that both the shape and volume of the semicircular canals will resist such changes. To test this hypothesis, we conducted three-dimensional geometric morphometric shape analysis on microcomputed tomographic data collected on the semicircular canals of an ovariectomized (OVX) rat model. A Procrustes ANOVA found no statistically significant differences in shape between surgery and sham groups, and morphological disparity testing likewise found no differences in shape variation. Univariate testing found no differences in semicircular volume between OVX and control groups. The range of variation in the OVX group, however, is greater than in the sham group but this difference does not reach statistical significance, perhaps because of a combination of small effect size and low sample size. This finding suggests that labyrinthine shape remains a tool for assessing phylogeny and function in the fossil record, but that it is possible that osteoporosis may be contributing to intraspecific shape variation in the bony labyrinth. This effect warrants further exploration at a microstructural level with continued focus on variables related to remodeling.

Ecomorphological associations of scapulocoracoid form in Greater Antillean Anolis lizards.

External morphological metrics have featured prominently in comparative studies examining the morphological convergence that characterizes anoline ecomorphs. To what degree the appendicular-skeletal morphology of Greater Antillean island Anolis lizards tracks their diversity and ecological adaptation, however, remains relatively unexplored. Here we employ computed tomographic scanning techniques to visualize in situ the scapulocoracoid of ecomorph representatives (trunk-ground, trunk-crown, crown-giant, twig) from three islands (Jamaica, Hispaniola, and Puerto Rico), and compare its three-dimensional geometry using qualitative-descriptive and quantitative-morphometric techniques. In contrast to our previous, similarly-conducted study of the pelvic girdle of these same species, the form of the scapulocoracoid varies markedly both within and between species, with much of the variation relating to phylogenetic relationship, specimen size, and assigned ecomorph category. Morphometric variation that correlates with size and/or phylogenetic signal varies between species and cannot be eliminated from the data set without markedly reducing its overall variability. The discovered patterns of skeletal variation are consistent with the demands of locomotor mechanics imposed by the structural configuration of the microhabitat of these ecomorphs. Most pertinently the ecomorphs differ in the anteroposterior length of the coracoid, the dorsoventral height of the scapulocoracoid, the dorsoventral height of the scapula in relation to the height of the suprascapula, and the relative positioning of the borders of the scapulocoracoid fenestra. In the examined ecomorph categories these skeletal differences likely relate to microhabitat usage by permitting different degrees of tilting and displacement of the scapulocoracoid in the parasagittal plane and influencing the sizes of muscle origins and the vectors of their actions. These differences relate to the amount of humeral adduction applied during its protraction, and to the structural stability of the shoulder girdle during acrobatic maneuvers, thus influencing the perch diameter that can be effectively negotiated, a critical factor in the microhabitat structure of Anolis ecomorphs.

What is bred in the bone: Ecomorphological associations of pelvic girdle form in greater Antillean Anolis lizards.

Ecological niche partitioning of Anolis lizards of the Greater Antillean islands has been the focus of many comparative studies, and much is known about external morphological convergence that characterizes anole ecomorphs. Their internal anatomy, however, has rarely been explored in an ecomorphological context, and it remains unknown to what degree skeletal morphology tracks the diversity and ecological adaptation of these lizards. Herein, we employ CT scanning techniques to visualise the skeleton of the pelvic girdle in situ, and 3D geometric morphometrics to compare the form of the ilium, ischium, and pubis within and between ecomorphs. We examine 26 species of anoles representing four ecomorphs (trunk-ground, trunk-crown, crown-giant, twig) from three islands (Jamaica, Hispaniola, and Puerto Rico). The subtle variations in pelvic girdle morphology discovered are directly associable with all three parameters that we set out to focus on: phylogenetic relationship, specimen size, and assigned ecomorph category. Morphometric variation that correlates with size and/or phylogenetic signal varies between species and cannot be eliminated from the data set without markedly reducing its overall variability. The discovered patterns of skeletal variation are consistent with the demands of locomotor mechanics pertinent to the structural configuration of the microhabitat of three of the four ecomorphs, with the fourth having no discernible distinctive features. This manifests itself chiefly in the relative anteroposterior extent and anteroventral inclination of the ilium and pubis, which differ between ecomorphs and are postulated to reflect optimization of the direction of muscle vectors of the femoral protractors and retractors. Our investigation of the form of the pelvic girdle of anoles allows us to generalize our findings to entire ecomorph categories within a broad phylogenetic and biogeographic context. Differences in the form and configuration of the postcranial skeleton are directly related to ecological patterns.

Points on the curve: An analysis of methods for assessing the shape of vertebrate claws.

The form of amniote claws has been extensively investigated, often with inferences about ecological association being drawn from studies of their geometry. Various methods have been used to quantify differences in the geometry of claws, but rarely have the underlying assumptions of such methods been addressed. Here, we use one set of bird claws and apply six methods (five that have been previously used, and a new one) that are tasked with comparing their shape. In doing so, we compare the (1) ability of these methods to represent the shape of the claw; (2) validity of the assumptions made about underlying claw geometry; (3) their ability to be applied unambiguously; and (4) their ability to differentiate between predetermined functional clusters. We find that of the six methods considered only the geometric morphometric approach reveals differences in the shapes of bird claws. Our comparison shows that geometry-based methods can provide a general estimate of the degree of curvature of claw arcs, but are unable to differentiate between shapes. Of all of the geometry-based approaches, we conclude that the adjusted version of the Zani (2000) method is the most useful because it can be applied without ambiguity, and provides a reliable estimate of claw curvature. The three landmarks that define that method (tip and base of the claw arc, plus the intersection between said claw arc and a line drawn perpendicular from the midpoint of tip and claw base) do not all bear biological significance, but relatively clearly circumscribe the length-to-height ratio of the claw, which relates to its curvature. Overall, our comparisons reveal that the shape of avian claws does not differ significantly between climbing and perching birds, and that the utilization of preordained functional clusters in comparative data analysis can hinder the discovery of meaningful differences in claw shape. J. Morphol. 278:150-169, 2017. © 2016 Wiley Periodicals,Inc.

Geometric morphometric analysis of the breast-shoulder apparatus of lizards: a test case using Jamaican anoles (Squamata: Dactyloidae).

The breast-shoulder apparatus (BSA) is a structurally and kinematically complex region of lizards. Compared with the pelvic region it has received little attention, even though its morphological variation is known to be extensive. This variability has seldom been the focus of functional explanation, possibly because the BSA has been difficult to explore as a composite entity. In this study we apply geometric morphometric techniques to the analysis of the BSA in an attempt to more fully understand its configuration in relation to differential use in locomotion. Our approach centers upon the Jamaican radiation of anoline lizards (genus Norops) as a tractable, small monophyletic assemblage consisting of species representing several ecomorphs. We hypothesized that the different species and ecomorphs would exhibit variation in the configuration of the BSA. Our findings indicate that this is so, and is expressed in the component parts of the BSA, although it is subtle except for Norops valencienni (twig ecomorph), which differs greatly in morphology (and behavior) from its island congeners. We further found similarities in the BSA of N. grahami, N. opalinus (both trunk-crown ecomorphs), and N. garmani (crown giant). These outcomes are promising for associating morphology with ecomorphological specialization and for furthering our understanding of the adaptive response of the BSA to demands on the locomotor system.