CNRS, UMR 5554 Institut des Sciences de l'Evolution (Université de Montpellier| CNRS IRD| EPHE), Place Eugène Bataillon, 34095 Montpellier, France.
Department of Biodiversity and Conservation, Real Jardín Botánico, CSIC, Plaza de Murillo, 2, 28014 Madrid, Spain.
Syst Biol. 2018 Nov 1;67(6):940-964. doi: 10.1093/sysbio/syy009.
In macroevolution, the Red Queen (RQ) model posits that biodiversity dynamics depend mainly on species-intrinsic biotic factors such as interactions among species or life-history traits, while the Court Jester (CJ) model states that extrinsic environmental abiotic factors have a stronger role. Until recently, a lack of relevant methodological approaches has prevented the unraveling of contributions from these 2 types of factors to the evolutionary history of a lineage. Herein, we take advantage of the rapid development of new macroevolution models that tie diversification rates to changes in paleoenvironmental (extrinsic) and/or biotic (intrinsic) factors. We inferred a robust and fully-sampled species-level phylogeny, as well as divergence times and ancestral geographic ranges, and related these to the radiation of Apollo butterflies (Parnassiinae) using both extant (molecular) and extinct (fossil/morphological) evidence. We tested whether their diversification dynamics are better explained by an RQ or CJ hypothesis, by assessing whether speciation and extinction were mediated by diversity-dependence (niche filling) and clade-dependent host-plant association (RQ) or by large-scale continuous changes in extrinsic factors such as climate or geology (CJ). For the RQ hypothesis, we found significant differences in speciation rates associated with different host-plants but detected no sign of diversity-dependence. For CJ, the role of Himalayan-Tibetan building was substantial for biogeography but not a driver of high speciation, while positive dependence between warm climate and speciation/extinction was supported by continuously varying maximum-likelihood models. We find that rather than a single factor, the joint effect of multiple factors (biogeography, species traits, environmental drivers, and mass extinction) is responsible for current diversity patterns and that the same factor might act differently across clades, emphasizing the notion of opportunity. This study confirms the importance of the confluence of several factors rather than single explanations in modeling diversification within lineages.
在宏观进化中，“红皇后”（Red Queen）模型假定生物多样性动态主要取决于物种内在的生物因素，如物种间的相互作用或生活史特征，而“宫廷弄臣”（Court Jester）模型则指出外在环境非生物因素具有更强的作用。直到最近，由于缺乏相关的方法学方法，一直无法揭示这两种因素对谱系进化历史的贡献。在这里，我们利用将多样化率与古环境（外在）和/或生物（内在）因素变化联系起来的新宏观进化模型的快速发展。我们推断了一个稳健且全面采样的种级系统发育，以及分化时间和祖先地理范围，并利用现生物种（分子）和已灭绝物种（化石/形态）证据，将其与阿波罗蝴蝶（Parnassiinae）的辐射联系起来。我们通过评估物种形成和灭绝是否由多样性依赖（生态位填补）和分支依赖的宿主植物关联（红皇后）或由气候或地质等大尺度连续变化（宫廷弄臣）介导，来检验它们的多样化动态是否可以更好地用红皇后或宫廷弄臣假设来解释。对于红皇后假说，我们发现与不同的宿主植物相关的物种形成率存在显著差异，但未检测到多样性依赖的迹象。对于宫廷弄臣假说，喜马拉雅-青藏高原的构建对于生物地理学具有重要作用，但不是高物种形成的驱动因素，而温暖气候与物种形成/灭绝之间的正相关性则得到了连续变化的最大似然模型的支持。我们发现，多个因素（生物地理学、物种特征、环境驱动因素和大规模灭绝）的共同作用而不是单一因素，导致了当前的多样性模式，并且同一因素在不同的分支中可能会有不同的作用，强调了机会的概念。本研究证实了在模型化谱系内多样化时，几个因素的共同作用而不是单一解释的重要性。
