Testing the Role of the Red Queen and Court Jester as Drivers of the Macroevolution of Apollo Butterflies
Résumé
In macroevolution, the Red Queen (RQ) model posits that biodiversity dynamics depend mainly on speciesintrinsic
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 maximumlikelihood
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.