Exploring evolutionary adaptations of leaf heteroblasty in subspecific taxa of Hawaiian Chenopodium oahuense

M. Bullock and J.T. Cantley, American Journal of Botany (2022).


This paper was part of Madison’s Master’s thesis and is her first first-author publication. Congratulations, Madison!


Chenopodium oahuense is a polymorphic Hawaiian endemic plant inhabiting several xeric habitats. Considerable variability in leaf morphology has made comprehensively describing its diversity difficult. Chenopodium oahuense subsp. ilioense is differentiated from C. oahuense subsp. oahuense by smaller, less-lobed, succulent leaves, smaller seeds, and prostrate to scandent habit. The lacking quantification of leaf shape, succulence, and previously unknown heteroblastic leaf transition in C. oahuense subsp. ilioense complicates the morphological boundaries separating subspecies.


This study used landmark analyses, elliptical Fourier descriptors (EFDs), and traditional shape descriptors measured from 1585 greenhouse-grown plant leaves collected over 18 weeks. Principal component analyses visualized correlations in leaf shape, and linear discriminant analyses predicted classifications, either subspecific or heteroblastic.


Identity determination and heteroblastic change visualization were limited in landmark analyses. On the basis of EFDs and shape descriptors, C. oahuense subsp. ilioense was determined to be morphologically differentiated from C. oahuense subsp. oahuense with the Pu’u Ka Pele population as intermediate. The EFDs depicted heteroblastic change, predominantly in lobing. All analyses were restricted in correctly attributing a leaf to the week collected. Shape descriptors generally represented significant heteroblastic change over the growth period.


These analyses support significant differentiation between the subspecies, particularly from shape descriptors. Furthermore, we quantified the morphological intermediacy of the Pu’u Ka Pele population. Results suggest this population could be the result of incomplete lineage sorting or a recent hybridization of the two subspecies. Hawaiian Chenopodium is a polymorphic lineage notable for future research in adaptive radiations, phenotypic plasticity, and heteroblasty.