Abstract
In macroevolution, the Red Queen (RQ) model posits that biodiversity dynamics depend mainly on speciesintrinsicbiotic factors such as interactions among species or life-history traits, while the Court Jester (CJ) model states thatextrinsic environmental abiotic factors have a stronger role. Until recently, a lack of relevant methodological approaches hasprevented 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 changesin paleoenvironmental (extrinsic) and/or biotic (intrinsic) factors. We inferred a robust and fully-sampled species-levelphylogeny, as well as divergence times and ancestral geographic ranges, and related these to the radiation of Apollobutterflies (Parnassiinae) using both extant (molecular) and extinct (fossil/morphological) evidence.We tested whether theirdiversification dynamics are better explained by an RQ or CJ hypothesis, by assessing whether speciation and extinctionwere mediated by diversity-dependence (niche filling) and clade-dependent host-plant association (RQ) or by large-scalecontinuous changes in extrinsic factors such as climate or geology (CJ). For the RQ hypothesis, we found significantdifferences in speciation rates associated with different host-plants but detected no sign of diversity-dependence. ForCJ, the role of Himalayan–Tibetan building was substantial for biogeography but not a driver of high speciation, whilepositive dependence between warm climate and speciation/extinction was supported by continuously varying maximumlikelihoodmodels. 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 mightact differently across clades, emphasizing the notion of opportunity. This study confirms the importance of the confluenceof several factors rather than single explanations in modeling diversification within lineages.