“…Expansion of temperate biomes is thought to have provided ecological opportunities for lineages to colonize and diversify (Simpson, 1953;Donoghue, 2008;Stroud & Losos, 2016). Indeed, many temperate-adapted lineages evolved and diversified around and after Oligocene, such as drought-and cold-adapted C4 grasses (Poaceae), succulents (e.g., Aizoaceae, Cactaceae), and orchids (Orchidoideae), leading to the subsequent spread of grasslands and deserts (Spriggs et al, 2014;Palazzesi et al, 2022;Thompson et al, 2023a;Thompson et al, 2023b). In most of these cases, it is thought that lineages already possessed traits needed for colonizing a new region, and that the niche was largely conserved as lineages radiated after arrival (Wiens & Donoghue, 2004;Donoghue, 2008;Crisp et al, 2009;Donoghue & Edwards, 2014).…”