The Afromontane and Afroalpine areas constitute some of the main biodiversity hotspots of Africa. They are particularly rich in plant endemics, but the biogeographic origins and evolutionary processes leading to this outstanding diversity are poorly understood. We performed phylogenomic and biogeographic analyses of one of the most species-rich plant genera in these mountains, Helichrysum (Compositae-Gnaphalieae). Most previous studies have focused on Afroalpine elements of Eurasian origin, and the southern African origin of Helichrysum provides an interesting counterexample. We obtained a comprehensive nuclear dataset from 304 species (≈50% of the genus) using target-enrichment with the Compositae1061 probe set. Summary-coalescent and concatenation approaches combined with paralog recovery yielded congruent, well-resolved phylogenies. Ancestral range estimations revealed that Helichrysum originated in arid southern Africa, whereas the southern African grasslands were the source of most lineages that dispersed within and outside Africa. Colonization of the tropical Afromontane and Afroalpine areas occurred repeatedly throughout the Miocene-Pliocene. This timing coincides with mountain uplift and the onset of glacial cycles, which together may have facilitated both speciation and intermountain gene flow, contributing to the evolution of the Afroalpine flora.

Botanic Institute of Barcelona CSIC Ajuntament de Barcelona Pg Migdia s n ES 08038 Barcelona Spain

Department of Biological Sciences Center for Biodiversity University of Memphis Memphis TN 38152 USA

Department of Botany and Plant Physiology and Plant DNA Biobank DNA National Bank University of Salamanca Edificio 1 D i Espejo St ES 37007 Salamanca Spain

Department of Botany Faculty of Science Charles University Prague Benátská 2 CZ 12801 Prague Czech Republic

Department of Botany Swedish Museum of Natural History P O Box 50007 SE 104 05 Stockholm Sweden

Department of Plant Biology and Biodiversity Management Addis Ababa University Addis Ababa P O Box 3434 Ethiopia

Foundational Biodiversity Science Kirstenbosch Research Centre South African National Biodiversity Institute Private Bag X7 Newlands Cape Town 7735 South Africa

Herbarium of the Parc Botanique et Zoologique of Tsimbazaza Antananarivo 3G9G V6C Madagascar

Institute of Botany Academy of Sciences of the Czech Republic CZ 25243 Průhonice Czech Republic

Meise Botanic Garden Nieuwelaan 38 BE 1860 Meise Belgium

Natural History Museum University of Oslo P O Box 1172 NO 0318 Oslo Norway

School of Animal Plant and Environmental Sciences University of the Witwatersrand Private Bag 3 Johannesburg 2050 South Africa

Systematics and Evolution of Vascular Plants Associated Unit to CSIC by IBB Department of Animal Biology Plant Biology and Ecology Faculty of Biosciences Autonomous University of Barcelona ES 08193 Bellaterra Spain

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