群体的快速多样化通常与利用生态机会和/或关键创新的演变有关。然而,这些非生物和生物因素的相互作用如何与生物多样化相关,在实证研究中很少记录,特别是对于居住在旱地的生物。Fumarioideae是罂粟科中最大的亚科,主要分布在北半球的温带地区。这里,我们使用了一个核(ITS)和六个质体(rbcL,atpB,matK,rps16,trnL-F,和trnG)DNA序列,以研究该亚家族多样化的时空模式和潜在的相关因素。我们首先介绍了迄今为止最全面的Fumarioideae系统发育分析。我们的综合分子测年和生物地理分析表明,在上白垩纪期间,Fumarioideae的最新共同祖先开始在亚洲多样化,然后在新生代多次散布出亚洲。特别是,我们发现了两个独立的扩散事件,从欧亚到新世晚期的东非,这表明阿拉伯半岛可能是新世晚期欧亚大陆和东非之间重要的交流走廊。在Fumarioideae中,在两组中检测到物种形成率增加,延胡索和奋马科。延胡索在42Ma的皇冠组首次经历了多样化的爆发,从上世纪中期开始,多元化进一步加速。在这两个时期,延胡索进化了不同的生活史类型,这可能促进了源自北半球广泛造山运动以及亚洲内部荒漠化的各种栖息地的定殖。Fumariinae在15Ma时经历了一阵多元化,这在时间上与欧亚大陆中部日益加剧的干旱化相吻合,但明显落后于栖息地(从潮湿到干旱)和生活史(从常年到年度)以及从亚洲到欧洲的扩展,这表明Fumariinae物种可能已经通过获取年度生活史而预先适应了入侵欧洲干旱栖息地。我们的研究提供了一个经验案例,该案例记录了适应前对旱地生物多样化的重要性,并强调了非生物和生物因素在促进植物多样化方面的协同作用。
Rapid diversification of a group is often associated with exploiting an ecological opportunity and/or the evolution of a key innovation. However, how the interplay of such abiotic and biotic factors correlates with organismal diversification has been rarely documented in empirical studies, especially for organisms inhabiting drylands. Fumarioideae is the largest subfamily in Papaveraceae and is mainly distributed in temperate regions of the Northern Hemisphere. Here, we used one nuclear (ITS) and six plastid (rbcL, atpB, matK, rps16, trnL-F, and trnG) DNA sequences to investigate the spatio-temporal patterns of diversification and potential related factors of this subfamily. We first present the most comprehensive phylogenetic analysis of Fumarioideae to date. The results of our integrated molecular dating and biogeographic analyses indicate that the most recent common ancestor of Fumarioideae started to diversify in Asia during the Upper Cretaceous, and then dispersed multiple times out of Asia in the Cenozoic. In particular, we discover two independent dispersal events from Eurasia to East Africa in the late Miocene, suggesting that the Arabian Peninsula might be an important exchange corridor between Eurasia and East Africa in the late Miocene. Within the Fumarioideae, increased speciation rates were detected in two groups, Corydalis and Fumariinae. Corydalis first experienced a burst of diversification in its crown group at ∼ 42 Ma, and further accelerated diversification from the mid-Miocene onwards. During these two periods, Corydalis had evolved diverse life history types, which could have facilitated the colonization of diverse habitats originating from extensive orogenesis in the Northern Hemisphere as well as Asian interior desertification. Fumariinae underwent a burst of diversification at ∼ 15 Ma, which temporally coincides with the increasing aridification in central Eurasia, but is markedly posterior to the shifts in habitat (from moist to arid) and in life history (from perennial to annual) and to range expansion from Asia to Europe, suggesting that Fumariinae species may have been pre-adapted to invade European arid habitats by the acquisition of annual life history. Our study provides an empirical case that documents the importance of pre-adaptation on organismal diversification in drylands and highlights the significant roles of the synergy of abiotic and biotic factors in promoting plant diversification.