The evolutionary history of spines: a Cenozoic arms race with mammals
The role of mammal herbivory in plant evolution is largely unrecognised. Spines on stems are a common and important feature found in [~]9% of eudicot woody plant species worldwide. Spines evolved independently multiple times during the Cenozoic. The timing and extent of spiny plant diversification varied among continents, pointing towards continental rather than global drivers. Spine evolution is closely related to radiation of extant ungulates and extinct ground sloths, rather than climate variation. Diversification began in the Paleogene in herbivore species-rich Eurasia and North America, emerging later in the Neogene in species-poorer South America, Africa and Australia. Spiny lineages expanded their ecological footprint over non-spiny plants, mainly through intercontinental migrations, indicating that spines likely provided a competitive advantage with increasing, and novel, mammal herbivory pressure.