Phylogeny, biogeography and adaptive radiation of Pachycladon (Brassicaceae) in the mountains of South Island, New Zealand

Aim To report analyses and propose hypotheses of adaptive radiation that explain distributional patterns of the alpine genus Pachycladon Hook.f. - a morphologically diverse genus from New Zealand closely related to Arabidopsis thaliana. Location South Island, New Zealand. Methods Morphological and nrDNA ITS sequence phylogenies were generated for Pachycladon. An analysis is presented of species distributional patterns and attributes. Results Phylogenetic analyses of morphological characters and nrDNA ITS sequence data were found to be congruent in supporting three New Zealand clades for Pachycladon. Monophyletic groups identified within the genus are geographically distinct and are associated with different geological parent materials. Distribution maps, latitude and altitude range, and data on geological parent material are presented for the nine named and one unnamed species of Pachycladon from New Zealand. Main conclusions (a) Panbiogeographic hypotheses accounting for the origin and present-day distribution of Pachycladon in New Zealand are not supported.
(b) Species diversity and distributions of Pachycladon are explained by a Late Tertiary-Quaternary adaptive radiation associated with increasing specialization to geological substrates. Pachycladon cheesemanii Heenan & A.D.Mitch. is morphologically similar to the closest overseas relatives. It is a geological generalist and has wide latitudinal and altitudinal ranges, and we suggest it resembles the ancestral form of the genus in New Zealand. Pachycladon novae-zelandiae (Hook.f.) Hook.f. and P. wallii (Carse) Heenan & A.D.Mitch. are a southern South Island group that predominantly occurs on Haast Schist, are polycarpic, have lobed leaves, and lateral inflorescences. Pachycladon enysii (Cheeseman) Heenan & A.D.Mitch., P. fastigiata (Hook.f.) Heenan & A.D.Mitch., and P. stellata (Allan) Heenan & A.D.Mitch. are restricted to greywacke in the eastern South Island, and are facultatively monocarpic, have serrate leaves, and stout terminal inflorescenes. (c) Present distributions of Pachycladon species may relate to Pleistocene climate change. Pachycladon enysii reaches the highest altitude of New Zealand species of Pachycladon and is most common in the Southern Alps in Canterbury. We propose that this species survived on nunataks at the height of the last glaciation. In contrast, P. fastigiata grows at a lower altitude and is absent from the high mountains of the Southern Alps. We suggest it was extirpated from this area during the last glaciation.