Abstract

Core Goodeniaceae is a clade of ~330 species primarily distributed in Australia. Considerable variation in flower morphology exists within this group and we aim to use geometric morphometrics to characterize this variation across the two major subclades: Scaevola sensu lato (s.l.) and Goodenia s.l., the latter of which was hypothesized to exhibit greater variability in floral symmetry form. We test the hypothesis that floral morphological variation can be adequately characterized by our morphometric approach, and that discrete groups of floral symmetry morphologies exist, which broadly correlate with subjectively determined groups. From 335 images of 44 species in the Core Goodeniaceae, two principal components were computed that describe >98% of variation in all datasets. Increasing values of PC1 ventralize the dorsal petals (increasing the angle between them), whereas increasing values of PC2 primarily ventralize the lateral petals (decreasing the angle between them). Manipulation of these two morphological “axes” alone was sufficient to recreate any of the general floral symmetry patterns in the Core Goodeniaceae. Goodenia s.l. exhibits greater variance than Scaevola s.l. in PC1 and PC2, and has a significantly lower mean value for PC1. Clustering clearly separates fan-flowers (with dorsal petals at least 120° separated) from the others, whereas the distinction between pseudo-radial and bilabiate clusters is less clear and may form a continuum rather than two distinct groups. Transitioning from the average fan-flower to the average non-fan-flower is described almost exclusively by PC1, whereas PC2 partially describes the transition between bilabiate and pseudo-radial morphologies. Our geometric morphometric method accurately models Core Goodeniaceae floral symmetry diversity.

Highlights

  • Categorization of morphological forms is an important component of comparative biology

  • A textbook example is the adaptive radiation in Darwin’s finches. Their beaks exhibit multiple discrete forms correlating with food sources, the overall shape of the beak is dictated by modulation of two pathways: Bmp4 for depth, and Calmodulin for length [3, 4], operating along an axis of shape variation discerned by geometric morphometrics [5]

  • principle components (PC) values were mapped back onto floral morphology using the eigenvectors of the residual covariance matrix

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Summary

Introduction

Categorization of morphological forms is an important component of comparative biology. Methods that provide objective, quantitative alternatives to subjective categorization are becoming important, when categories are not immediately obvious or variation appears continuous. Quantifying these traits provides mathematical leverage to studies of variation within populations, and to comparative character evolution and gene expression studies among species to help identify the key molecular targets of adaptive evolution. A textbook example is the adaptive radiation in Darwin’s finches Their beaks exhibit multiple discrete forms correlating with food sources, the overall shape of the beak is dictated by modulation of two pathways: Bmp for depth, and Calmodulin for length [3, 4], operating along an axis of shape variation discerned by geometric morphometrics [5]. Studies that have shifted from binary or discrete characters to continuous characters have opened our eyes to previously unseen patterns, as in the example of the punctuated adaptive diversification of the Aquilegia nectar spur in response to different pollinators (e.g., [6, 7])

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