Abstract
This paper proposes a new methodology to quantify patterns of egg shape variation using geometric morphometrics of three-dimensional landmarks captured on digitally reconstructed eggshells and demonstrates its performance in capturing shape variation at multiple biological levels. This methodology offers unique benefits to complement established linear measurement or two-dimensional (2D) contour profiling techniques by (i) providing a more precise representation of eggshell curvature by accounting for variation across the entire surface of the egg; (ii) avoids the occurrence of correlations from combining multiple egg shape features; (iii) avoids error stemming from projecting a highly-curved three-dimensional (3D) object into 2D space; and (iv) enables integration into 3D workflows such as finite elements analysis. To demonstrate, we quantify patterns of egg shape variation and estimate morphological disparity at multiple biological levels, within and between clutches and among species of four passerine species of different lineages, using volumetric dataset obtained from micro computed tomography. The results indicate that species broadly have differently shaped eggs, but with extensive within-species variation so that all four-focal species occupy a range of shapes. Within-species variation is attributed to between-clutch differences in egg shape; within-clutch variation is surprisingly substantial. Recent comparative analyses that aim to explain shape variation among avian taxa have largely ignored potential biases due to within-species variation, or use methods limited to a narrow range of egg shapes. Through our approach, we suggest that there is appreciable variation in egg shape across clutches and that this variation needs to be accounted for in future research. The approach developed in this study to assess variation in shape is freely accessible and can be applied to any spherical-to-conical shaped object, including eggs of non-avian dinosaurs and reptiles through to other extant taxa such as poultry.
Highlights
IntroductionAvian egg shape diversity is a well-known biological phenomenon reflecting taxonomic diversity (Olsen, Cunningham & Donnelly, 1994; Stoddard et al, 2017), ranging from the nearly spherical eggs of owls (Hoyt, 1976), slightly pointed eggs of domestic fowl (Havlıcek et al, 2008), to the extremely pointed eggs of certain waders, alcids and penguins (Birkhead et al, 2017; Stoddard et al, 2017)
The first two PCs accounted for 83.8% of the total egg shape variation across species, and a biplot of the two axes provides a reasonable approximation of the egg morphospace for this study
Overall mean clutch disparity was not significantly different among species (ANOVA, F3,12 = 0.184, P = 0.906). Together these results indicate that within-species egg shape variation is due to between-clutch differences as well as within-clutch differences in egg shape
Summary
Avian egg shape diversity is a well-known biological phenomenon reflecting taxonomic diversity (Olsen, Cunningham & Donnelly, 1994; Stoddard et al, 2017), ranging from the nearly spherical eggs of owls (Hoyt, 1976), slightly pointed eggs of domestic fowl (Havlıcek et al, 2008), to the extremely pointed eggs of certain waders, alcids and penguins (Birkhead et al, 2017; Stoddard et al, 2017). Various hypotheses have been proposed to explain the evolution of specific egg shapes (Rensch, 1947; Andersson, 1978; Bain & Solomon, 1991; Attard et al, 2017; Deeming, 2017; Stoddard et al, 2017), but so far only explain a small proportion of variation in egg shape (Deeming, 2017; Stoddard et al, 2017), or are applicable to a limited number of species These findings are complicated by the use of multiple methods by different studies to evaluate and compare egg shape, with some more comprehensive in their evaluation of shape differences than others (Havlıcek et al, 2008). These variables are likely to be correlated with one another, leading to multiple difficulties in identifying the extent of shape variation between samples and selecting which indices best characterise egg shape for a given species (Narushin, 2001)
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