The holly genus, Ilex L., in the monogeneric Aquifoliaceae, is the largest woody dioecious genus (>664 spp.), with a near‐cosmopolitan distribution in mesic environments. We constructed a phylogeny based on two nuclear genes, representing 177 species spread across the geographical range, and dated using macrofossil records. The five main clades had a common ancestor in the early Eocene, much earlier than previously suggested. Ilex originated in subtropical Asia and extant clades colonized South America by 30 Ma, North America by 23 Ma, Australia by 8 Ma, Europe by 6 Ma, and Africa by 4 Ma. South and North America were colonized multiple times. Ilex also reached Hawaii (10 Ma) and other oceanic islands. Macrofossil and pollen records show the genus has tracked mesic climates through time and space, and had a wider distribution before late Miocene global cooling. Our phylogeny provides a framework for studies in comparative ecology and evolution.

中文翻译：

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冬青的系统发育和生物地理学（冬青属，Aquifoliaceae）

冬青属 Ilex L. 属于单属 Aquifoliaceae，是最大的木本雌雄异株属 (>664 spp.)，在中等环境中具有近乎世界性的分布。我们基于两个核基因构建了系统发育，代表分布在地理范围内的 177 种物种，并使用大型化石记录确定年代。这五个主要进化枝在始新世早期有一个共同的祖先，比以前认为的要早得多。冬青起源于亚热带亚洲，现存的进化枝在南美洲殖民了 30 马，北美是 23 马，澳大利亚是 8 马，欧洲是 6 马，非洲是 4 马。南美洲和北美洲曾多次被殖民。冬青还到达了夏威夷（10 Ma）和其他海洋岛屿。宏观化石和花粉记录显示该属已经通过时间和空间追踪了中等气候，并且在晚中新世全球变冷之前分布更广。我们的系统发育为比较生态学和进化研究提供了一个框架。