Phylogeny, molecular sequences, fossils, biogeography, and biome occupancy are all lines of evidence that reflect the singular evolutionary history of a clade, but they are most often studied separately, by first inferring a fossil-dated molecular phylogeny, then mapping on ancestral ranges and biomes inferred from extant species. Here we jointly model the evolution of biogeographic ranges, biome affinities, and molecular sequences, while incorporating fossils to estimate a dated phylogeny for all of the 163 extant species of the woody plant clade Viburnum (Adoxaceae) that we currently recognize in our ongoing worldwide monographic treatment of the group. Our analyses indicate that while the major Viburnum lineages evolved in the Eocene, the majority of extant species originated since the Miocene. Viburnum radiated first in Asia, in warm, broad-leaved evergreen (lucidophyllous) forests. Within Asia we infer several early shifts into more tropical forests, and multiple shifts into forests that experience prolonged freezing. From Asia we infer two early movements into the New World. These two lineages probably first occupied warm temperate forests and adapted later to spreading cold climates. One of these lineages (Porphyrotinus) occupied cloud forests and moved south through the mountains of the Neotropics. Several other movements into North America took place more recently, facilitated by prior adaptations to freezing in the Old World. We also infer four disjunctions between Asia and Europe: the Tinus lineage is the oldest and probably occupied warm forests when it spread, while the other three were more recent and in cold-adapted lineages. These results variously contradict published accounts, especially the view that Viburnum radiated initially in cold forests and, accordingly, maintained vessel elements with scalariform perforations. We explored how the location and biome assignments of fossils affected our inference of ancestral areas and biome states. Our results are sensitive to, but not entirely dependent upon, the inclusion of fossil biome data. It will be critical to take advantage of all available lines of evidence to decipher events in the distant past. The joint estimation approach developed here provides cautious hope even when fossil evidence is limited.

中文翻译：

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荚果全球多样化期间地理运动和生物群落变化的联合系统发育估计

系统发育、分子序列、化石、生物地理学和生物群落占有率都是反映进化枝单一进化历史的证据线，但它们最常被单独研究，首先推断化石年代的分子系统发育，然后绘制祖先范围和从现存物种推断的生物群落。在这里，我们共同模拟了生物地理范围、生物群系亲和性和分子序列的演变，同时结合化石来估计我们目前在我们正在进行的全球专着中认识到的木本植物进化枝荚蒾科 (Adoxaceae) 的所有 163 种现存物种的过时系统发育组的治疗。我们的分析表明，虽然主要的 Viburnum 谱系在始新世进化，但大多数现存物种起源于中新世。荚蒾首先在亚洲辐射，在温暖，阔叶常绿（lucidophyllous）森林。在亚洲，我们推断出有几次早期转变为更多的热带森林，并且多次转变为经历长时间冰冻的森林。我们从亚洲推断出两个早期进入新世界的运动。这两个谱系可能首先占据了暖温带森林，后来适应了蔓延的寒冷气候。其中一个谱系（Porphyrotinus）占据了云雾林并通过新热带山脉向南移动。最近发生了其他几次进入北美的运动，这得益于先前对旧世界冻结的适应。我们还推断出亚洲和欧洲之间的四个分离：Tinus 谱系是最古老的，在它传播时可能被占领的温暖森林，而其他三个则较新，并且处于冷适应谱系中。这些结果与已发表的报道有不同的矛盾，尤其是荚蒾最初在寒冷的森林中辐射并因此保持带有鳞片状穿孔的血管元素的观点。我们探索了化石的位置和生物群落分配如何影响我们对祖先地区和生物群落状态的推断。我们的结果对包含化石生物群落数据很敏感，但并不完全依赖于。利用所有可用的证据来破译遥远过去的事件至关重要。即使化石证据有限，这里开发的联合估计方法也提供了谨慎的希望。我们探索了化石的位置和生物群落分配如何影响我们对祖先地区和生物群落状态的推断。我们的结果对包含化石生物群落数据很敏感，但并不完全依赖于。利用所有可用的证据来破译遥远过去的事件至关重要。即使化石证据有限，这里开发的联合估计方法也提供了谨慎的希望。我们探索了化石的位置和生物群落分配如何影响我们对祖先地区和生物群落状态的推断。我们的结果对包含化石生物群落数据很敏感，但并不完全依赖于。利用所有可用的证据来破译遥远过去的事件至关重要。即使化石证据有限，这里开发的联合估计方法也提供了谨慎的希望。