Fabien L Condamine
email: fabien.condamine@gmail.com

Rémi Allio

Eliette L Reboud

Julian R Dupuis

Emmanuel F A Toussaint

Nathan Mazet

Shao-Ji Hu

Delano S Lewis

Krushnamegh Kunte

Adam M Cotton

Felix A H Sperling

† Affiliations

A comprehensive phylogeny and revised taxonomy illuminate the origin and diversification of the global radiation of Papilio (Lepidoptera: Papilionidae)

Keywords: Butterflies, Historical biogeography, Macroevolution, Paleotropics, Systematics, Taxonomy

niveau recherche, publiés ou non, émanant des établissements d'enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

Introduction

Insects are indisputably the most diverse terrestrial eukaryotic clade, with over a million insect species described and many more awaiting discovery or formal description [START_REF] Stork | How many species of insects and other terrestrial arthropods are there on Earth[END_REF]. This staggering species diversity is unevenly distributed across the insect tree of life, with many clades comprising substantially more species than their sister clades [START_REF] Grimaldi | Evolution of the Insects[END_REF][START_REF] Engel | Insect evolution[END_REF]. For instance, within ants (Formicidae), some genera such as Camponotus and Pheidole have over 1,500 and 1,100 species, respectively [START_REF] Hölldobler | The Ants[END_REF], while their corresponding sister clades have less than 700 and 200 species, respectively (Economo et al., 2018). Within flies (Diptera), the most well-known case is the genus Drosophila, which has more than 1600 species [START_REF] Brake | Drosophilidae (Diptera)[END_REF][START_REF] O'grady | Phylogeny of the genus Drosophila[END_REF], which is sister to a clade composed of seven genera including about 600 species [START_REF] Wiegmann | Episodic radiations in the fly tree of life[END_REF][START_REF] O'grady | Phylogeny of the genus Drosophila[END_REF]. Such genera are difficult to resolve phylogenetically and manage taxonomically. The genus rank is widely used in molecular, morphological and paleontological systematics. However, species-rich genera that are difficult to delimit can also be treated using the subgenus rank, rendering their relationships less complicated while allowing flexibility to taxonomic ranks that are less frequently employed [START_REF] Winston | Describing Species: Practical Taxonomic Procedure for Biologists[END_REF][START_REF] Teta | The usage of subgenera in mammalian taxonomy[END_REF].

In Lepidoptera, swallowtail butterflies (Papilionidae) of the genus Papilio Linnaeus, 1758 include more than 200 species and represent more than one third of all Papilionidae (Fig. 1), which has about 600 recognized species (Zakharov et al., 2004a;Haüser et al., 2005;[START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF]Nakae, 2021a). As one of the most well-known and broadly studied groups of insects, Papilio swallowtails are recognized as model organisms in evolutionary biology, ecology, genomics, and conservation biology (e.g. [START_REF] Collins | Threatened Swallowtail Butterflies of the World[END_REF][START_REF] Scriber | Swallowtail Butterflies: Their Ecology and Evolutionary Biology[END_REF][START_REF] Kunte | The diversity and evolution of Batesian mimicry in Papilio swallowtails butterflies[END_REF][START_REF] Kunte | doublesex is a mimicry supergene[END_REF][START_REF] Dupuis | Repeated reticulate evolution in North American Papilio machaon group swallowtail butterflies[END_REF]. Yet the phylogeny of Papilio is far from being resolved despite numerous studies [START_REF] Ae | The phylogeny of some Papilio based on interspecific hybridization data[END_REF][START_REF] Hancock | Classification of the Papilionidae (Lepidoptera): a phylogenetic approach[END_REF][START_REF] Igarashi | The classification of the Papilionidae mainly based on the morphology of their immature stages[END_REF][START_REF] Miller | Phylogenetic studies in the Papilioninae (Lepidoptera: Papilionidae)[END_REF][START_REF] Tyler | Swallowtail Butterflies of the Americas: A Study in Biological Dynamics[END_REF][START_REF] Scriber | Swallowtail Butterflies: Their Ecology and Evolutionary Biology[END_REF][START_REF] Aubert | Molecular phylogeny of swallowtail butterflies of the tribe Papilionini (Papilionidae, Lepidoptera)[END_REF][START_REF] Caterino | Papilio phylogeny based on mitochondrial cytochrome oxidase I and II genes[END_REF][START_REF] Reed | Interaction of process partitions in phylogenetic analysis: an example from the swallowtail butterfly genus Papilio[END_REF][START_REF] Yagi | Phylogeny of Japanese papilionid butterflies inferred from nucleotide sequences of the mitochondrial ND5 gene[END_REF][START_REF] Caterino | A partitioned likelihood analysis of swallowtail butterfly phylogeny (Lepidoptera: Papilionidae)[END_REF]Zakharov et al., 2004a;Condamine et al., 2013a;[START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF][START_REF] Wu | Phylogeny and historical biogeography of Asian Pterourus butterflies (Lepidoptera: Papilionidae): A case of intercontinental dispersal from North America to East Asia[END_REF][START_REF] Owens | The latitudinal diversity gradient in New World swallowtail butterflies is caused by contrasting patterns of out-of-and into-thetropics dispersal[END_REF][START_REF] Owens | Comparative phylogenetics of Papilio butterfly wing shape and size demonstrates independent hindwing and forewing evolution[END_REF]. Ecological and evolutionary hypotheses on their diversification rely on a comprehensive and strongly supported phylogeny as well as accurate age estimates for significant phylogenetic events like host-plant shifts or mimicry evolution.

The first classifications of Papilio species relied on morphological characters (summarized in Zakharov et al., 2004a: table 1). [START_REF] Munroe | The classification of the Papilionidae (Lepidoptera)[END_REF] divided Papilio into five sections but did not designate them as subgenera because they lacked a simple diagnosis with adult characters. Using an explicitly cladistic estimation of relationships within Papilio, Hancock (1983) recognized six genera (Chilasa, Eleppone, Heraclides, Papilio, Princeps, and Pterourus) based on phylogenetic evidence and inferred evolutionary antiquity, but this phylogeny also suffered from lack of character justification. Another classification was proposed by [START_REF] Igarashi | The classification of the Papilionidae mainly based on the morphology of their immature stages[END_REF] based on the morphology of immature stages, but this work did not represent all of Hancock's genera. [START_REF] Igarashi | The classification of the Papilionidae mainly based on the morphology of their immature stages[END_REF] nonetheless recognized seven genera (Achillides, Agehana, Chilasa, Euchenor, Menelaides, Papilio, and Pterourus), with numerous discrepancies between his treatment and that of [START_REF] Hancock | Classification of the Papilionidae (Lepidoptera): a phylogenetic approach[END_REF]. Hancock's classification was criticized by [START_REF] Miller | Phylogenetic studies in the Papilioninae (Lepidoptera: Papilionidae)[END_REF], who did not consider elevation of Papilio subdivisions to the genus level to be justified. However, one group within Papilio was elevated in a widely available checklist of swallowtail butterflies (Haüser et al., 2005), where Chilasa was treated as a distinct genus. It has been challenging to apply the rules of the PhyloCode in designating subgroups within Papilio to generic status as it has proven difficult to find morphological synapomorphies as required by the PhyloCode (Cantino and de Queiroz, 2020). The sole use of genetic data to delimit taxa has not been adequately addressed by the PhyloCode.

Due to the limitations of traditional morphological approaches and with the development of new molecular systematic approaches, the classification of Papilio has received significant attention in the last three decades. Relationships among species within the P. machaon and P. glaucus-troilus species groups were studied based on allozyme variation [START_REF] Sperling | Evolution of the Papilio machaon species group in western Canada[END_REF]Hagen and Scriber, 1991). Restriction fragment length polymorphism of mitochondrial DNA was used to compare taxa within the same species groups in later studies [START_REF] Sperling | Mitochondrial DNA phylogeny, speciation, and host-plant coevolution of Papilio butterflies[END_REF](Sperling, , 1993a(Sperling, , 1993b;;Sperling andHarrison, 1994, Tyler et al., 1994). Phylogenetic relationships within Papilio have also been analyzed using DNA sequences of a variety of genes, but these studies have been confined to single species groups or local geographic areas (e.g. [START_REF] Vane-Wright | Evolution of the mimetic African swallowtail butterfly Papilio dardanus: Molecular data confirm relationships with P. phorcas and P. constantinus[END_REF][START_REF] Yagi | Phylogeny of Japanese papilionid butterflies inferred from nucleotide sequences of the mitochondrial ND5 gene[END_REF]Zakharov et al., 2004b;Condamine et al., 2013;[START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF][START_REF] Owens | The latitudinal diversity gradient in New World swallowtail butterflies is caused by contrasting patterns of out-of-and into-thetropics dispersal[END_REF][START_REF] Owens | Comparative phylogenetics of Papilio butterfly wing shape and size demonstrates independent hindwing and forewing evolution[END_REF][START_REF] Dupuis | Phylogenomic test of mitochondrial clues to archaic ancestors in a group of hybridizing swallowtail butterflies[END_REF][START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF] or have included limited sampling across Papilio subdivisions (e.g. [START_REF] Aubert | Molecular phylogeny of swallowtail butterflies of the tribe Papilionini (Papilionidae, Lepidoptera)[END_REF][START_REF] Caterino | Papilio phylogeny based on mitochondrial cytochrome oxidase I and II genes[END_REF][START_REF] Reed | Interaction of process partitions in phylogenetic analysis: an example from the swallowtail butterfly genus Papilio[END_REF]Zakharov et al., 2004a). Although the two latest molecular phylogenetic studies of Papilionidae [START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF][START_REF] Allio | Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants[END_REF] substantially increased the fraction of sampled species, they did not provide a sufficient resolution to assess the fine-scale taxonomic delimitation within the genus. Yet, a number of systematic enigmas remain in Papilio, starting with the number of valid species to consider in the genus.

Here we establish a list of valid Papilio species based on previous molecular studies and build a reliable and comprehensive time-calibrated species-level phylogeny for the genus Papilio using seven gene fragments for ca. 80% of the total species diversity. Our study aims at establishing a reference phylogenetic framework to evaluate both subgeneric monophyly and species relationships within Papilio, for the enigmatic and long-debated placement of several groups and species. The phylogeny includes species that are key for testing biogeographic hypotheses (e.g. P. anactus, P. benguetana), evolution of mimicry (e.g. Chilasa, P. nobilis), and host-plant associations (e.g. P. alexanor), as well as species that are rare and insufficiently studied (e.g. P. antimachus, P. himeros), endangered (P. buddha, P. chikae) or recently described (e.g. the Fijian P. natewa). The recent discovery of P. natewa [START_REF] Tennent | A remarkable new swallowtail butterfly from Fiji (Lepidoptera, Papilionidae)[END_REF] may clarify the systematic position of P. anactus, presently placed in a monotypic subgenus (Eleppone) but with an unstable phylogenetic position [START_REF] Hancock | The systematic position of Papilio anactus Macleay (Lepidoptera: Papilionidae)[END_REF]Zakharov et al., 2004a;[START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF]. Indeed, external morphology and genitalia of P. natewa suggest a phylogenetic affinity with P. anactus [START_REF] Tennent | A remarkable new swallowtail butterfly from Fiji (Lepidoptera, Papilionidae)[END_REF], but this remains to be tested.

Without knowledge of its larva, the enigmatic African giant swallowtail, P. antimachus, has often been placed in species groups of two distinct Papilio subgenera (Druryia or Princeps), and the subgenus Druryia has never been studied with a molecular approach. The position of the nobilis species-group also remains poorly resolved, wavering between the iconic phorcas and hesperus species-groups [START_REF] Munroe | The classification of the Papilionidae (Lepidoptera)[END_REF]Hancock, 1983, respectively). Yet, its placement has important implications for the understanding of mimicry evolution in the phorcas group [START_REF] Vane-Wright | Evolution of the mimetic African swallowtail butterfly Papilio dardanus: Molecular data confirm relationships with P. phorcas and P. constantinus[END_REF] and evolution of iridescent wings (common origin or convergence with the subgenus Achillides). Including the giant blue swallowtail, P. zalmoxis, in the phylogeny could help solve this puzzle. Moreover, the monophyly and rank of the danaine-and moth-mimicking Chilasa are uncertain. [START_REF] Munroe | The classification of the Papilionidae (Lepidoptera)[END_REF] split its members among two Papilio subunits, but [START_REF] Hancock | Classification of the Papilionidae (Lepidoptera): a phylogenetic approach[END_REF] placed them together in a single genus, considered to be the sister taxon of Eleppone. Finally, we also revisited the placement of P. alexanor. The relationships of this odd European Apiaceae feeder have been examined several times [START_REF] Aubert | Molecular phylogeny of swallowtail butterflies of the tribe Papilionini (Papilionidae, Lepidoptera)[END_REF][START_REF] Caterino | Papilio phylogeny based on mitochondrial cytochrome oxidase I and II genes[END_REF][START_REF] Reed | Interaction of process partitions in phylogenetic analysis: an example from the swallowtail butterfly genus Papilio[END_REF]Zakharov et al., 2004a), nearly reaching a consensus that P. alexanor is sister to a clade comprising Pterourus and Chilasa (Zakharov et al., 2004a;[START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF]. However, no strong resolution has been obtained and this position has been questioned with phylogenomic analyses [START_REF] Allio | Whole genome shotgun phylogenomics resolves the pattern and timing of swallowtail butterfly evolution[END_REF], albeit with less-than-ideal sampling density. We not only resolve these systematic conflicts and uncertainties, but also provide a phylogenetic framework to infer an evolutionary timescale for Papilio, estimate historical biogeography, and investigate heterogeneity of diversification dynamics.

Materials and methods

Taxon sampling and molecular data

We first established a taxonomic working list of Papilio species (Table 1). This species list combines several previous studies (Zakharov et al., 2004a(Zakharov et al., , 2004b;;[START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF]Condamine et al., , 2013aCondamine et al., , 2013b;;[START_REF] Shiraiwa | A new Heraclides swallowtail (Lepidoptera, Papilionidae) from North America is recognized by the pattern on its neck[END_REF][START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF][START_REF] Wu | Phylogeny and historical biogeography of Asian Pterourus butterflies (Lepidoptera: Papilionidae): A case of intercontinental dispersal from North America to East Asia[END_REF][START_REF] Owens | The latitudinal diversity gradient in New World swallowtail butterflies is caused by contrasting patterns of out-of-and into-thetropics dispersal[END_REF][START_REF] Owens | Comparative phylogenetics of Papilio butterfly wing shape and size demonstrates independent hindwing and forewing evolution[END_REF][START_REF] Allio | Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants[END_REF][START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF]. Like all other working lists [START_REF] Garnett | Principles for creating a single authoritative list of the world's species[END_REF], we acknowledge that this species list remains provisional and is likely to evolve through time (i.e. species are hypotheses, [START_REF] Pante | Species are hypotheses: Avoid connectivity assessments based on pillars of sand[END_REF]. Indeed, Papilio butterflies will inevitably continue to receive attention as a model clade, and we expect many of the species' boundaries to be tested with new genomic data and evolutionary models in further studies, which are very likely to eventually modify the taxonomic list (e.g. [START_REF] Kunte | Sex chromosome mosaicism and hybrid speciation among tiger swallowtail butterflies[END_REF]Dupuis and Sperling, 2022). From the 235 species that are currently recognized in the new taxonomic list presented here for the genus Papilio, we sampled 184 species representing 78.3% of the total diversity (Table 1). However, the total number of species will evolve with further systematic studies, especially of the African clades that are currently poorly represented in our analysis (Fig. 2).

We added 18 outgroup species representing several swallowtail genera to root the Papilio tree and provide deeper relationships for secondary calibration of nodes based on previous dated studies [START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF][START_REF] Allio | Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants[END_REF]. The outgroups included: [START_REF]Changes to subspecies nomenclature resulting from literature review and this study. 1462 1463 Species New subspecies combinations Papilio (Achillides) orsippus Godman & Salvin[END_REF] 7) the single species of the subfamily Baroniinae, Baronia brevicornis, which is the sister lineage to all Papilionidae and was used as root of the phylogenetic tree [START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF][START_REF] Allio | Whole genome shotgun phylogenomics resolves the pattern and timing of swallowtail butterfly evolution[END_REF]. Overall, the molecular dataset comprised 202 species (184 ingroups and 18 outgroups).

We assembled a supermatrix dataset with data extracted from Sanger sequencing and genome sequencing available on GenBank as of September 2021. The molecular data mainly came from previous studies (e.g. Zakharov et al., 2004aZakharov et al., , 2004b;;[START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF]Condamine et al., , 2013aCondamine et al., , 2013b;;[START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF][START_REF] Wu | Phylogeny and historical biogeography of Asian Pterourus butterflies (Lepidoptera: Papilionidae): A case of intercontinental dispersal from North America to East Asia[END_REF][START_REF] Owens | The latitudinal diversity gradient in New World swallowtail butterflies is caused by contrasting patterns of out-of-and into-thetropics dispersal[END_REF][START_REF] Owens | Comparative phylogenetics of Papilio butterfly wing shape and size demonstrates independent hindwing and forewing evolution[END_REF][START_REF] Allio | Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants[END_REF][START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF]. We used five mitochondrial gene fragments (COI, COII, ND1, ND5 and rRNA 16S) and two nuclear gene fragments (EF-1a and Wg), chosen based on their availability among Papilio species. We aligned the DNA sequences for the rRNA 16S using MAFFT 7.110 [START_REF] Katoh | MAFFT multiple sequence alignment software version 7: improvements in performance and usability[END_REF] with the E-INS-i algorithm, while we aligned the coding genes using MACSE 2.00 [START_REF] Ranwez | MACSE: Multiple Alignment of Coding SEquences accounting for frameshifts and stop codons[END_REF] with the alignSequences subprogram and default options. All the resulting alignments were checked for codon stops and eventually refined by eye with Mesquite 3.7 [START_REF] Maddison | Mesquite: a modular system for evolutionary analysis[END_REF]. All gene alignments were concatenated into a nucleotide supermatrix, which is available in FigShare (Data 1: https://figshare.com/s/a32288db7e6429714a62).

Inferring phylogenetic relationships

We performed both maximum likelihood (ML) and Bayesian Inference (BI) to reconstruct phylogenetic relationships. Although phylogenomic approaches have used amino acids on Papilionidae at the genus level [START_REF] Allio | Whole genome shotgun phylogenomics resolves the pattern and timing of swallowtail butterfly evolution[END_REF], all our analyses relied on nucleotides for the tree inference of Papilio because the molecular dataset is limited to seven gene fragments and converting it to amino acids will result in a smaller dataset with limited phylogenetic information given the species diversity of the focal clade. ML inference was implemented with IQ-TREE 2.1.2 [START_REF] Minh | IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era[END_REF] using ModelFinder to select the best-fit partition scheme and the best-fitting substitution model for each partition (-m MFP+MERGE option, Chernomor et al., 2016;[START_REF] Kalyaanamoorthy | ModelFinder: fast model selection for accurate phylogenetic estimates[END_REF]. For IQ-TREE analyses, we estimated the most likely tree with 100 separate ML searches, which after initial model optimization on a parsimony tree used 100 random tree topologies as starting trees for each search. As recommended, we optimized ML searches to avoid local optima by [START_REF]Changes to subspecies nomenclature resulting from literature review and this study. 1462 1463 Species New subspecies combinations Papilio (Achillides) orsippus Godman & Salvin[END_REF] increasing the number of unsuccessful iterations before stopping tree optimization to 500 (-nstop 500 option), and (2) decreasing the perturbation strength for randomized NNI to 0.2 (-pers 0.2 option). Statistical reliability of the ML tree was evaluated with 100 non-parametric bootstraps under the optimal partitioned model to obtain ML bootstrap percentages (BPPART). To compare branch supports, a second ML analysis with IQ-TREE was carried out under the same conditions but with 2,000 ultrafast bootstraps (UFBSPART; [START_REF] Hoang | UFBoot2: improving the ultrafast bootstrap approximation[END_REF]. BS values and UFBS values were considered strong when higher than 70% and 95%, respectively. We also assessed branch support using Shimodaira-Hasegawa approximate likelihood ratio test (SH-aLRT; [START_REF] Guindon | New algorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3.0[END_REF], with SH-aLRT values above 80% considered as strong support for a clade. BI analyses were performed using MrBayes 3.2.7a [START_REF] Ronquist | MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space[END_REF]. Because ModelFinder includes more substitution models than BEAST or MrBayes, the most likely partitioning scheme and substitution models can be altered, the molecular dataset was analyzed with PartitionFinder 2.1.1 [START_REF] Lanfear | PartitionFinder 2: new methods for selecting partitioned models of evolution for molecular and morphological phylogenetic analyses[END_REF] to estimate the best partition scheme with an initial subset of 19 possible partitions (all three codon positions for coding sequences and noncoding genes treated separately). Partitions and corresponding optimal substitution models were searched using the greedy algorithm, the mrbayes set of models, and the Bayesian Information Criterion (BIC) to compare the fit of different models. We used reversible-jump Markov chain Monte Carlo (rjMCMC) to sample the entire space of possible models instead of using the a priori substitution models recovered by PartitionFinder [START_REF] Huelsenbeck | Bayesian phylogenetic model selection using reversible jump Markov chain Monte Carlo[END_REF].

MrBayes analyses were performed with two separate runs, a random starting tree, and eight rjMCMC (one cold and seven incrementally heated, temp=0.1) running for 30 million generations with tree sampling occurring every 3,000 generations (resulting in 10,000 trees) to calculate the clade posterior probabilities (PPPART). We also specified (1) a uniform prior probability of phylogenies (i.e. all possible trees are considered a priori equally probable), and

(2) a uniform prior probability distribution on branch lengths. Convergence of the Bayesian runs was ensured by checking the average deviation of split frequencies (ADSF), the potential scale reduction factor (PSRF) values, the effective sample size (ESS) of all parameters, and by plotting the log-likelihood of the samples against the number of generations in Tracer 1.7.1 [START_REF] Rambaut | Posterior summarization in Bayesian phylogenetics using Tracer 1.7[END_REF]. The runs had to have values of ADSF approaching zero, PSRF close to 1.0 and ESS above 200 to assume convergence. All trees that predated the time needed to reach a log-likelihood plateau were discarded as burn-in, and the remaining samples were used to generate a 50% majority rule consensus tree (option halfcompat). Branch support was estimated and PPPART≥0.95 was considered to indicate strong support for a given clade [START_REF] Douady | Comparison of Bayesian and maximum likelihood bootstrap measures of phylogenetic reliability[END_REF]. All MrBayes analyses were performed on the computer cluster CIPRES Science Gateway [START_REF] Miller | A RESTful API for access to phylogenetic tools via the CIPRES science gateway[END_REF], using BEAGLE [START_REF] Ayres | BEAGLE: an application programming interface and high-performance computing library for statistical phylogenetics[END_REF] with default parameters. BI was also conducted using PhyloBayes MPI 1.8 [START_REF] Lartillot | PhyloBayes MPI: phylogenetic reconstruction with infinite mixtures of profiles in a parallel environment[END_REF] under the CAT-GTR-Γ4 mixture model [START_REF] Lartillot | A Bayesian mixture model for across-site heterogeneities in the amino-acid replacement process[END_REF], which has proven to perform well on large molecular datasets (e.g. [START_REF] Allio | Whole genome shotgun phylogenomics resolves the pattern and timing of swallowtail butterfly evolution[END_REF]. The analyses were conducted on the nucleotide dataset. For each analysis, two independent MCMC starting from a random tree were run at least 10,000 cycles, with trees and associated model parameters being sampled every 10 cycles.

The initial 2,000 trees sampled in each MCMC run were discarded as burn-in after checking for convergence in both likelihood and model parameters (tracecomp subprogram), and clade posterior probability (bpcomp subprogram). We checked the mean (meandiff) discrepancy observed across all bipartitions, considering that meandiff < 0.01 indicates convergence. The 50% majority-rule Bayesian consensus tree and the associated posterior probabilities (PPCAT) were then computed from the remaining trees using bpcomp.

Testing topology hypotheses

We conducted tests of topology hypotheses with MrBayes when phylogenetic analyses did not recover the same relationships as those from previous studies (see in Zakharov et al., 2004a;[START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF][START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF]. To do so, we conducted topology hypotheses by enforcing the clade (species, genera) to be sister to another clade or to be monophyletic in a case of non-monophyly. Bayesian inferences were run with the exact same priors and parameters (see above) but including constraints and a model to compute the marginal likelihood estimate (MLE) to obtain the likelihood score of each constrained topology, which was then compared to the score of an unconstrained topology.

We used stepping-stone sampling [START_REF] Xie | Improving marginal likelihood estimation for Bayesian phylogenetic model selection[END_REF] to estimate the marginal likelihood of the specified topology and the unconstrained topology. Stepping-stone sampling (SS) is considerably more accurate than the harmonic mean of the likelihoods from a standard MCMC run, and has been shown to be more efficient than the thermodynamic integration [START_REF] Baele | Accurate model selection of relaxed molecular clocks in Bayesian phylogenetics[END_REF]. SS estimates the model likelihood by sampling a series of distributions that represent different mixtures of posterior distribution and prior distribution [START_REF] Xie | Improving marginal likelihood estimation for Bayesian phylogenetic model selection[END_REF]. To obtain an adequate sample from most of the steps in the algorithm, we used 100 steps with 300,000 generations each, for a total of 30 million generations. To monitor convergence during each step, we set the diagnostics frequency to once every 1,000 generations. The MLE obtained with the SS in MrBayes were used to calculate the Bayes factors (BF). The logarithm of the BF is the difference in the logarithms of the marginal model likelihoods. The BF is calculated by subtracting the MLE of the constrained topology and the MLE of the unconstrained topology.

We considered BF values >10 favoring one model over another as very significant (Nylander et al., 2004;Brown and Lemmon, 2007).

Estimation of divergence times

We first tested the hypothesis of a molecular clock with PATHd8 [START_REF] Britton | Estimating divergence times in large phylogenetic trees[END_REF]. Since a strict molecular clock was not supported for 72.7% of the nodes in this dataset at P<0.05, a Bayesian relaxed-clock approach considering rate variations across lineages was employed to estimate divergence times [START_REF] Drummond | Relaxed phylogenetics and dating with confidence[END_REF]. Bayesian MCMC analyses implemented in BEAST 1.10.4 (Suchard et al., 2018) were performed to approximate the posterior distribution of rates and divergence times and infer their credibility intervals.

Molecular dating analyses can be sensitive to several parameters or priors [START_REF] Alfaro | The posterior and the prior in Bayesian phylogenetics[END_REF][START_REF] Drummond | Bayesian evolutionary analysis with BEAST[END_REF]. For instance the prior governing the branching process [START_REF] Condamine | Origin and diversification of living cycads: A cautionary tale on the impact of the branching process prior on Bayesian molecular dating[END_REF] or the number of molecular clocks [START_REF] Angelis | An evaluation of different partitioning strategies for Bayesian estimation of species divergence times[END_REF]Condamine et al., 2018a) may impact estimates of divergence times. Accordingly, we conducted analyses with the following non-default settings and priors: (1) the molecular dataset was partitioned according to the best-fit scheme recovered by PartitionFinder, (2) substitution models were set based on ModelFinder estimation in IQ-TREE (see above), (3) a birth-death tree prior was set for the branching process, and (4) one clock model was set for the mitochondrial partitions, and one clock for the nuclear partitions. Finally, we let the tree topology be estimated by BEAST (classic operator mix) but enforced the monophyly of subfamilies, tribes, and genus Papilio following the results of phylogenomic analyses [START_REF] Allio | Whole genome shotgun phylogenomics resolves the pattern and timing of swallowtail butterfly evolution[END_REF]. Bayesian relaxed clocks were set up with an uncorrelated lognormal distribution clock model, with the mean set to a uniform prior between 0 and 0.5 (starting value=0.1), and an exponential prior (lambda=0.333) for the standard deviation. The birth-death process [START_REF] Gernhard | The conditioned reconstructed process[END_REF]) is a more realistic branching tree prior than a Yule model for species-level divergence times analysis. The birth-death process was set using the following uniform priors: the mean growth rate ranged between 0 and 1 with a starting value at 0.1 births per lineage per million years, and the relative death rate ranged between 0 and 1 deaths per lineage per million years (starting value=0.5). We performed four independent BEAST runs (different seeds) for 50 million generations of MCMC each and with a sampling frequency of 5,000 generations.

We discarded the first 10% of generations as burn-in and checked for convergence using Tracer (ESS>200). We combined the four runs using LogCombiner. Using TreeAnnotator, we computed the maximum clade credibility tree with median ages and the 95% credibility intervals (CI) at each node. The xml files are available in FigShare (Data 2: https://figshare.com/s/a32288db7e6429714a62).

Fossil and secondary calibrations

To explore the effect of fossil calibrations versus secondary calibrations on estimating divergence times, we designed two calibration sets. All node calibrations were assigned a uniform prior distribution with hard bounds (Yang and Rannala, 2006).

First, we used three unambiguous fossil calibrations assigned to Papilionidae, two of which are Parnassiinae (de Jong, 2017). The first is †Thaites ruminiana [START_REF] Scudder | Fossil butterflies[END_REF], a compression fossil from limestone in the Niveau du gypse d'Aix Formation of France (Aix-en-Provence, Bouches-du-Rhône) within the Chattian (23.03-28.1 Ma) of the late Oligocene [START_REF] Sohn | An annotated catalog of fossil and subfossil Lepidoptera (Insecta: Holometabola) of the world[END_REF]. †Thaites is sister to Parnassiini, and occasionally sister to Luehdorfiini + Zerynthiini (Condamine et al., 2018b). Thus we constrained the crown age of Parnassiinae with a uniform distribution bounded by a minimum age of 23.03 Ma. The second is †Doritites bosniaskii [START_REF] Rebel | Doritites bosniaskii. Sitzungsberichte der akademie der wissenschaften[END_REF], an exoskeleton and compression fossil from Italy (Tuscany) from the Messinian (5.33-7.25 Ma, late Miocene;[START_REF] Sohn | An annotated catalog of fossil and subfossil Lepidoptera (Insecta: Holometabola) of the world[END_REF]. †Doritites is sister to Archon (Luehdorfiini, Condamine et al., 2018b), in agreement with [START_REF] Carpenter | Treatise on Invertebrate Paleontology, Part R, Arthropoda 3-4[END_REF]. The crown of Luehdorfiini was thus constrained for divergence time estimation using a uniform distribution bounded with 5.33 Ma. Third is the genus †Praepapilio with two fossil species †P. colorado and †P. gracilis [START_REF] Durden | Butterflies from the middle Eocene: the earliest occurrence of fossil Papilionidae[END_REF] found in the Green River Formation (Colorado, U.S.A.). The age of †Praepapilio was used to constrain the crown age of Papilionidae with a uniform distribution bounded by a minimum age of 47.8 Ma since the Green River Formation encompasses a 5 million-year period between ~48.5 and 53.5 Ma, which falls within the Ypresian (47.8-56 Ma) in the early Eocene (Smith et al., 2003;[START_REF] De Jong | Estimating time and space in the evolution of the Lepidoptera[END_REF]. These three fossil calibrations have been used in previous studies (e.g. [START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF]Condamine et al., , 2013aCondamine et al., , 2018aCondamine et al., , 2018b;;[START_REF] Allio | Whole genome shotgun phylogenomics resolves the pattern and timing of swallowtail butterfly evolution[END_REF]. Uniform distributions of the fossil calibrations were bounded with a maximum age of 150 Ma, a conservative age congruent with the major radiation of angiosperms [START_REF] Magallón | A metacalibrated time-tree documents the early rise of flowering plant phylogenetic diversity[END_REF][START_REF] Foster | Evaluating the impact of genomic data and priors on Bayesian estimates of the angiosperm evolutionary timescale[END_REF][START_REF] Li | Origin of angiosperms and the puzzle of the Jurassic gap[END_REF].

Second, we relied on secondary calibrations with uniform priors applied to the nodes where we set the fossil calibrations plus nodes that are in common between our phylogeny and the phylogeny of Papilionidae. We retrieved the 95% CI from recent estimations of divergence times of the family [START_REF] Allio | Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants[END_REF] as follows: (1) the crown of Papilionidae (root of the tree) between 47.8 and 70.9 Ma, (2) the crown of Parnassiinae between 29.9 and 58.9 Ma, (3) the crown of Luehdorfiini between 13.9 and 35.3 Ma, (4) the crown of Papilioninae between 34.4 and 62.9 Ma, (5) the crown of Leptocircini between 26.6 and 49.9 Ma, (6) the crown of Papilionini + Troidini between 30.8 and 56.1 Ma, (7) the crown of Troidini between 26.9 and 50.4 Ma, and (8) the crown of Papilionini between 27.5 and 50.9 Ma [START_REF] Allio | Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants[END_REF].

Inference of historical biogeography

We estimated the ancestral ranges of origin and geographic range evolution for Papilio using the ML approach of dispersal-extinction-cladogenesis (DEC, Ree and Smith, 2008) as implemented in the DEC eXtended version (DECX, Beeravolu and Condamine, 2016; available at: https://github.com/champost/DECX). To infer the biogeographic history of a clade, DEC requires a time-calibrated tree, the current distribution of each species for a set of geographic areas, and a time-stratified geographic model that is represented by connectivity and dispersal scalar matrices for specified time intervals spanning the entire evolutionary history of the group. DECX allows classical vicariance as a cladogenetic event by using temporally flexible constraints on the connectivity between any two given areas following the movement of landmasses and dispersal opportunity over time. DECX can analyze phylogenies on the order of several thousand tips with numerous areas [START_REF] Rolland | The contribution of temperature and continental fragmentation to amphibian diversification[END_REF]Kawahara et al., 2023). DECX does not incorporate the founder-event speciation (+J parameter) because of concerns with statistical validity of model choice among DEC-derived models [START_REF] Ree | Conceptual and statistical problems with the DEC+ J model of founder-event speciation and its comparison with DEC via model selection[END_REF]. Also, founder-event speciation often leads to inferences that are decoupled from time, with null or extremely low extinction rates, an effect of the model favoring cladogenetic events over anagenetic events [START_REF] Ree | Conceptual and statistical problems with the DEC+ J model of founder-event speciation and its comparison with DEC via model selection[END_REF], which makes it inadequate for reconstructing the history of ancient groups with widespread distributions.

The geographic distribution for all 235 Papilio species was categorized by coding the presence or the absence of each species in the following areas: (1) West Palearctic, defined as Europe west of the Urals and the part of Asia west of this north-south line, (2) East Palearctic, defined as everywhere considered Palearctic east of the Urals, above 3,000 m in the Himalayas and north of Sichuan in China, (3) West Nearctic, defined as North America west of the Rocky Mountains, (4) East Nearctic, defined as North America east of the Rocky Mountains, (5) Central America, defined as from the northern border of Mexico southwards to the border between Panama and Colombia, also including the Caribbean islands except Trinidad and Tobago, (6) South America, defined as all countries from Colombia to Argentina and including Trinidad and Tobago, (7) Africa, defined as the whole of the African continent and Arabian Peninsula but excluding the islands in the Indian Ocean, (8) Madagascar, defined as the island of Madagascar and all other Indian Ocean islands in the vicinity, (9) India, defined as the area below 3,000 m from NW Pakistan to the border with Myanmar, (10) Indonesia and Wallacea, defined as Myanmar, SE Asia, southern China, western Indonesia to Lydekker's Line; including the Lesser Sunda Islands but excluding Timor, Wetar and associated islands, which are Australasian in origin, and (11) Australasia, defined as everywhere east of Lydekker's Line but including Timor, Wetar and small nearby islands. Species which only marginally enter an area were excluded. We used data available in the literature (e.g. [START_REF] Collins | Threatened Swallowtail Butterflies of the World[END_REF][START_REF] Tyler | Swallowtail Butterflies of the Americas: A Study in Biological Dynamics[END_REF][START_REF] Scriber | Swallowtail Butterflies: Their Ecology and Evolutionary Biology[END_REF]. The geographic distribution of all species and the species sampled in this study is presented in Fig. 2. The resulting matrix of species distribution for Papilio is available in Table S1.

A time-stratified geographic model was built using connectivity matrices that consider paleogeographic changes through time with time slices indicating the possibility or not for a species to colonize a new area (Beeravolu and Condamine, 2016). Based on paleogeographic reconstructions (e.g. Scotese, 2004;Blakey, 2008;Seton et al., 2012), we created a connectivity matrix to represent major changes in tectonic conditions that may have affected the distribution of these butterflies. We specified constraints on area connectivity by coding 0 if any two areas are not connected or 1 if they are connected at a given period. We did not add dispersal matrices because setting the values for dispersal rates between regions through time is highly subjective, and it has been shown that dispersal probability categories had minor effects on ancestral state estimation (Chacon and Renner, 2014). Therefore, we assumed a dispersal scalar matrix with equal dispersal rates between areas through time. Biogeographic ranges larger than four areas in size were disallowed as valid biogeographic states if they were not subsets of the terminal species ranges; widespread ranges comprising areas that have never been geographically connected were also removed.

We estimated the most likely ancestral states at each node by performing a DEC analysis under an ML procedure as implemented in DECX (Beeravolu and Condamine, 2016) using both species distribution matrix and connectivity matrices. The files for reproducing the analyses are available in FigShare (Data 3: https://figshare.com/s/a32288db7e6429714a62).

Investigating heterogeneity of diversification rates

To provide an assessment of diversification rates through time, we used the ML approach of [START_REF] Morlon | Reconciling molecular phylogenies with the fossil record[END_REF], tested and automated in [START_REF] Mazet | Estimating clade-specific diversification rates and palaeodiversity dynamics from reconstructed phylogenies[END_REF] and implemented in the Rpackage RPANDA 2.0 [START_REF] Morlon | RPANDA: an R package for macroevolutionary analyses on phylogenetic trees[END_REF]. This method aims at capturing heterogeneity of diversification by allowing preselected subclades to follow different birth-death models from the deeper pruned tree (the backbone). For both subclades and backbones (different backbones are tested because of the different combination of shifts), speciation and/or extinction rates can change exponentially through time: λ(t)= λ0e αt with λ0 denoting speciation at present, α the trend of rate variation of speciation through time t. Extinction rate can exceed speciation, meaning that diversification rates can be negative [START_REF] Morlon | Reconciling molecular phylogenies with the fossil record[END_REF], which results in a declining paleodiversity dynamic that can be expected for the backbone because of the higher proportion of long branches after isolating recently-originated subclades.

Following [START_REF] Mazet | Estimating clade-specific diversification rates and palaeodiversity dynamics from reconstructed phylogenies[END_REF], for the analysis of Papilio, we first computed the clades' sampling fractions based on our revised taxonomy with the get.sampling.fraction function. We

Results and Discussion

Global phylogeny of Papilio

Partitioned phylogenetic analyses with IQ-TREE and MrBayes provided almost identical phylogenetic trees, differing in branch length estimates (Fig. 3; Figs S1, S2 for IQ-TREE). The genus Papilio was always recovered as monophyletic with maximal (PPCAT=1, PPPART=1, BPPART=100, UFBSPART=100) branch support (Table 3), which was expected since molecular data have never found it non-monophyletic (e.g. [START_REF] Aubert | Molecular phylogeny of swallowtail butterflies of the tribe Papilionini (Papilionidae, Lepidoptera)[END_REF]Zakharov et al., 2004a;[START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF][START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF][START_REF] Owens | The latitudinal diversity gradient in New World swallowtail butterflies is caused by contrasting patterns of out-of-and into-thetropics dispersal[END_REF][START_REF] Owens | Comparative phylogenetics of Papilio butterfly wing shape and size demonstrates independent hindwing and forewing evolution[END_REF][START_REF] Allio | Whole genome shotgun phylogenomics resolves the pattern and timing of swallowtail butterfly evolution[END_REF]. The Bayesian inference with MrBayes converged well (ADSF=0.007293, average PSRF for parameter values=1.000, and ESS>>200 for all parameters). Bayesian analyses reconstructed a robust phylogeny with 69.4% of the nodes recovered with strong support within Papilio (nodes with PPPART≥0.95, Fig. 3). Robustness was slightly lower for ML analysis with non-parametric bootstrap: 67.4% of branches in the tree were strongly supported (nodes with BPPART≥70, Fig. S1). In comparison, IQ-TREE analysis with UFBSPART recovered the most robust phylogeny with 82.5% of branches supported by UFBSPART≥95 (Fig. S2). PhyloBayes analysis under the mixture (CAT-GTR-Γ4) model ran during 11,350 cycles and converged well (meandiff=0.00814). The topology (Fig. S3) was very similar to traditional partitioned analyses with IQ-TREE and MrBayes, except within the Old World Papilio (see below). Overall, branch support in PhyloBayes was also like IQ-TREE and MrBayes with 66.7% of branches having PPCAT≥0.95 (see Fig. S4 for a comparison of branch support across all analyses).

All phylogenetic analyses reconstructed a backbone topology with three main clades (Clades 1 to 3 on Fig. 3, UFBSPART=100). The species relationships within Heraclides largely correspond to previous works [START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF][START_REF] Owens | The latitudinal diversity gradient in New World swallowtail butterflies is caused by contrasting patterns of out-of-and into-thetropics dispersal[END_REF][START_REF] Owens | Comparative phylogenetics of Papilio butterfly wing shape and size demonstrates independent hindwing and forewing evolution[END_REF].

New World Papilio sensu stricto (i.e. subgenera Heraclides and Pterourus) were not inferred as monophyletic in all analyses. We always recovered the American subgenus Pterourus within a clade including Asian subgenera Chilasa and 'Agehana' (Clade 2 on Fig.

3) with strong branch support (PPCAT=0.99, PPPART=1, BPPART=91, and UFBSPART=100). In addition, 'Agehana' was internal to subgenus Pterourus in all analyses (Table 3). This topology agrees with some phylogenetic work [START_REF] Wu | Phylogeny and historical biogeography of Asian Pterourus butterflies (Lepidoptera: Papilionidae): A case of intercontinental dispersal from North America to East Asia[END_REF] but contrasts with others (Zakharov et al., 2004a;[START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF][START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF], which proposed that Heraclides is sister to Pterourus + Chilasa + Alexanoria. The substantial increase of taxon sampling may account for the difference between topologies: we sampled 64 species belonging to these subgenera, while Zakharov et al. (2004a) and [START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF] analyzed only 21 species (including P. alexanor). Bayesian topology tests provided strong support (BF>6) for the non-monophyly of the New World Papilio clade (sensu Zakharov et al., 2004a;[START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF][START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF], and decisive support (BF>10) for the non-monophyly of Pterourus sensu lato (Table 2). These new phylogenetic arrangements have important biogeographic implications (see below).

Papilio alexanor, a mysterious Eurasian lineage that is notoriously difficult to place within Papilio (Zakharov et al., 2004a;[START_REF] Wu | Phylogeny and historical biogeography of Asian Pterourus butterflies (Lepidoptera: Papilionidae): A case of intercontinental dispersal from North America to East Asia[END_REF], was sister to the Old World clade including all remaining species, with moderate to high branch support (PPCAT=1, PPPART=0.99, BPPART=58, and UFBSPART=85). The Bayesian topology test constraining P. alexanor to be sister to the clade Pterourus + Chilasa + 'Agehana' (Zakharov et al., 2004a;[START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF] yielded a worse MLE than the unconstrained topology with strong support (BF>10, Table 2). Our unconstrained topology agrees with a recent but sparsely sampled phylogenomic study [START_REF] Allio | Whole genome shotgun phylogenomics resolves the pattern and timing of swallowtail butterfly evolution[END_REF], suggesting that the phylogenetic position of P. alexanor is reaching a consensus.

The Old World Papilio clade (Clade 3 on Fig. 3) includes the highest species diversity of the genus. After the divergence with P. alexanor, partitioned phylogenetic analyses recovered a series of five main clades (Clades 3a to 3e on Fig. 3) sustained by short internal branches leading to different subgenera with moderate to high branch support (Table 3). These internal branches were the source of discrepancy between the Bayesian analyses (under both the mixture model and the partitioned analyses) and ML analyses. The main discrepancy was that Clades 3a and 3b were recovered as sisters in ML analyses and weakly supported (BPPART=23, and UFBSPART=58; Figs S1-2) but not recovered in Bayesian analyses (Fig. 3, Fig. S3). Despite a substantially increased taxon sampling compared to previous studies (37% of total species diversity in [START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF] versus 78% in this study), these results suggest that the systematic backbone of Papilio is still not definitely resolved, which impedes a higherlevel systematic revision of the genus. We also acknowledge the limited size of the Sangerbased dataset and phylogenomic studies may illuminate the early evolutionary history of Papilio in future studies to provide a definitive resolution of subgeneric classification.

Clade 3a was always composed of the subgenus Euchenor [START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF] including two New Guinean species (Papilio euchenor and P. depilis), subtended by a long branch, which was systematically recovered as sister to a clade comprising species of subgenus Druryia (including the antimachus and dardanus species-groups) with moderate to strong branch support (Clade 3a on Fig. 3; PPCAT=0.82, PPPART=0.99, BPPART=74, and UFBSPART=99).

We still have poor knowledge on subgenus Druryia because existing phylogenies only included a handful of species. Here we found the nireus, oribazus, and zalmoxis species groups in a separate clade sister to Papilio nobilis with strong branch support (PPCAT=1, PPPART=1, BPPART=76, UFBSPART=98). This clade has recently been ranked as the subgenus Nireopapilio [START_REF] Cotton | Replacement name for Eques Kirby, 1896 (Lepidoptera: Papilionidae)[END_REF]. A Bayesian topology test provided decisive support (BF>10) for non-monophyly of subgenus Druryia (Table 2), therefore lending support to the validity of subgenus Nireopapilio. However, it is important to remain cautious because future studies with more comprehensive sampling could recover subgenus Druryia as monophyletic, notably through the inclusion of the zenobia species-group not sampled in this or previous studies.

Papilio natewa, the latest described Papilio species [START_REF] Tennent | A remarkable new swallowtail butterfly from Fiji (Lepidoptera, Papilionidae)[END_REF], was always sister to the monotypic subgenus Eleppone, with maximal branch support in all analyses (Fig. 3). This indicates that Papilio natewa can be placed in the subgenus Eleppone, together with P. anactus from eastern Australia. The subgenus Eleppone was always found to be sister to a strongly supported clade of five Indonesian species, the demolion species group, which was previously considered to belong to the subgenus Menelaides but now assigned to the subgenus Araminta. Although often recovered, this sister relationship was not highly supported (Clade 3b on Fig. 3; PPPART=0.72, BPPART=50, and UFBSPART=93).

3, Table

The next clade (Clades 3c + 3d + 3e on Fig. 3) included species classified in six subgenera and was generally strongly supported in all analyses (PPCAT=0.99, PPPART=1, BPPART=72, and UFBSPART=99). The first to branch off this clade were subgenus Sinoprinceps (Papilio xuthus and P. benguetana) and subgenus Papilio (machaon species-group) found to be sisters (Clade 3c on Fig. 3) in all analyses with maximal branch support (except BPPART=97).

These two subgenera were both recovered as monophyletic with maximal branch supports. The remaining subgenera comprised Nireopapilio + Achillides + Princeps + Menelaides (Clade 3d + Clade 3e on Fig. 3) and formed a strongly supported clade (PPCAT=0.99, PPPART=1, BPPART=60, and UFBSPART=99). The species Papilio nobilis (usually ranked in subgenus Princeps, nobilis group) was always found as sister to Nireopapilio (PPCAT=1, PPPART=1, BPPART=76, and UFBSPART=98; Clade 3d on Fig. 3), and the latter contained the species of the nireus, oribazus and zalmoxis groups. The subgenus Achillides was strongly supported as monophyletic in all analyses (PPCAT=1, PPPART=1, BPPART=85, and UFBSPART=99), and was often found as sister to the clade Princeps + Menelaides with moderate nodal support (Clade 3e on Fig. 3). Within Achillides, the species relationships largely agree with the study of Condamine et al. (2013b), but we added two endangered species (P. buddha and P. chikae; [START_REF] Collins | Threatened Swallowtail Butterflies of the World[END_REF]. Endemic to the Western Ghats biodiversity hotspot of Southern India, P. buddha was always recovered in the palinurus group (P. daedalus and P. palinurus)

from Southeast Asia with strong support (PPCAT=1, PPPART=1, BPPART=100, and UFBSPART=100), but its sister relationship with P. daedalus was weakly supported (PPCAT=0.8, PPPART=0.72, BPPART=64, and UFBSPART=62). Such a relationship is interesting to study in terms of wing morphological evolution between the palinurus group and the other Achillides endemic to India, P. crino. Papilio chikae is endemic to the Philippines (North Luzon) and was always sister to P. hermeli as expected [START_REF] Cabusas | Cytochrome c oxidase subunit I and II (COI and COII) sequences support recognition of Papilio chikae Igarashi and Papilio hermeli Nuyda as distinct species[END_REF], also endemic to the Philippines (North Mindoro) with maximal support in all analyses. They were together nested in the bianor group with maximal branch support as previously found (Condamine et al., 2013b). Subgenus

Princeps, containing five species of the demoleus species group, constituted a strongly supported clade with maximal branch support, which was found to be sister to subgenus Menelaides (PPCAT=0.93, PPPART=1, BPPART=70, and UFBSPART=99). A Bayesian topology test rejected the hypothesis of a monophyletic origin for all species usually placed in subgenus

Princeps like the dardanus species-group (BF>10, Table 2). Finally, the species comprising the subgenus Menelaides formed a solid monophyletic group with robust branch support in all analyses (PPCAT=1, PPPART=0.99, BPPART=99, and UFBSPART=100). The species relationships within Menelaides largely agree with the recent phylogenetic study of [START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF].

We constrained subgenera Menelaides and Araminta (species previously included in Menelaides) to form a single clade, but the Bayesian analysis confirmed the non-monophyly of such an artificially inclusive Menelaides with decisive support (BF>10, Table 2).

Origin of Papilio and subgeneric diversification

The four independent Bayesian runs of the two dating analyses converged well (ESS>>200 for most of the parameters) and were therefore combined. The analyses yielded almost identical estimates of divergence times with less than 0.5 million years of difference for all nodes, regardless of the calibrations used: fossils or secondary (Fig. 4, Table 3, see Figs S5-6 for the chronograms resulting from the BEAST analyses). The dating analyses estimated that Papilio originated in the Oligocene ca. 30.06 Ma (95% CI=20.66-47.42 Ma) with the three-fossilcalibrations analysis, and ca. 29.95 Ma (95% CI=23.97-37.34 Ma) with the eight-secondarycalibrations analysis. These results indicate that fossil and secondary calibrations can provide similar and consistent results in age estimates, which has not always been the case [START_REF] Sauquet | Testing the impact of calibration on molecular divergence times using a fossil-rich group: the case of Nothofagus (Fagales)[END_REF]. For the subsequent analyses (biogeography and diversification), we selected the maximum clade credibility tree with median ages estimated with a Bayesian uncorrelated lognormal method calibrated with fossils (Fig. 4).

We estimated that the origin of subgenera ranges from the early Miocene (e.g. Chilasa, Heraclides, Pterourus), middle Miocene (e.g. Achillides), to the late Miocene (e.g. Araminta, Menelaides, Papilio) (Fig. 4, Table 3). These age estimates are slightly younger than previous analyses with age differences ranging from ca. Condamine et al., 2013). We argue that these slight age discrepancies mostly come from the differences in taxon sampling resulting in new phylogenetic placements (e.g. P. alexanor) and potentially from practices in dating techniques both influencing divergence time estimates.

Northern origin and dynamic dispersal into the tropics

DEC analyses recovered a region including West Nearctic, Central America, East Palearctic, and Sundaland as the most likely ancestral geographic origin (relative probability=0.467), when Asia and North America were connected by the Bering land bridge in the Oligocene (Fig. 4).

The second best ancestral area was composed of West Nearctic, Central America, and East Palearctic (relative probability=0.215). Hence, a Northern (Laurasian) origin is preferred over a Southern (Gondwanan) origin. This is not unexpected given the age of the genus and the results of previous biogeographic analyses of the genus (e.g. [START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF]Condamine et al., , 2013;;[START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF][START_REF] Wu | Phylogeny and historical biogeography of Asian Pterourus butterflies (Lepidoptera: Papilionidae): A case of intercontinental dispersal from North America to East Asia[END_REF], although some studies estimated an older age and Gondwanan origin (Zakharov et al., 2004a). The DEC analyses indicate a dynamic biogeographic history with numerous dispersal events inferred (n=64) in comparison to vicariance events (n=22). Dispersal events were mostly southward (n=32) from a northern origin (or dispersal into the tropics, [START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF][START_REF] Rolland | Dispersal is a major driver of the latitudinal diversity gradient of Carnivora[END_REF] than northward (n=17, when including northwestward dispersals) from the equator (or dispersal out of the tropics, [START_REF] Jablonski | Out of the tropics: evolutionary dynamics of the latitudinal diversity gradient[END_REF]. Northward dispersals are only estimated in the last 10 million years. Excluding vicariance events involving areas around the Bering Strait (n=2), we found more vicariance in the Old World (n=13) than in the New World (n=7). We also estimated numerous local extirpation events (n=51), which tend to be more numerous in the Old World subgenera (n=27) than in the New World subgenera (n=15), likely due to the Miocene fragmentation of the Boreotropical forest in the Holarctic [START_REF] Pound | Global vegetation dynamics and latitudinal temperature gradients during the Mid to Late Miocene (15.97-5.33 Ma)[END_REF].

The Paleotropics as a biogeographic crossroad

In the non-monophyletic New World Papilio, subgenus Heraclides originated in Central America (very likely including Caribbean Islands; [START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF] We found the Paleotropics to be an important ancestral area for the historical biogeography of Papilio. It is striking that, within the Old World clade (Clade 3), all the main nodes of the backbone (seven in total out of 26, Table 3) are estimated to originate within the Paleotropics in the early Miocene (23 to 15 Ma, Fig. 4). During this time period, Africa, West and East Palearctic, and Sundaland were mostly covered by tropical-adapted or paratropical forests providing an almost continuous tropical habitat between these large regions, which likely facilitated biotic movements within the Paleotropics (e.g. [START_REF] Ziegler | Tracing the tropics across land and sea: Permian to present[END_REF][START_REF] Morley | Cretaceous and Tertiary climate change and the past distribution of megathermal rainforests[END_REF]. However, starting after the middle Miocene climatic optimum [START_REF] Steinthorsdottir | The Miocene: The future of the past[END_REF], the global cooling of the Cenozoic fostered the geographic contraction of the tropical belt toward the equator that disappeared from higher latitudes [START_REF] Pound | Global vegetation dynamics and latitudinal temperature gradients during the Mid to Late Miocene (15.97-5.33 Ma)[END_REF]. Our biogeographic estimates indicated widespread geographic extinctions in the Paleotropics, especially in the Holarctic, and at least seven different lineages were able to disperse southwards to track their preferred macroclimatic conditions, supposedly tropical climates [START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF]. Like crematogastrine ants [START_REF] Blaimer | Paleotropical diversification dominates the evolution of the hyperdiverse ant tribe Crematogastrini (Hymenoptera: Formicidae)[END_REF], we show that the Paleotropics had a central role in the origin and evolution of Papilio. The vast and stable ecological opportunity offered by the tropical rainforests likely explained the into-the-tropics dispersal trend to track tropical contractions as climate cooled down toward the present. This supports the hypothesis that potentially many clades, particularly inhabitants of Boreotropical floras, were likely extirpated from the Holarctic and persist today in more southern tropical locations [START_REF] Meseguer | Reconstructing deep-time palaeoclimate legacies in the clusioid Malpighiales unveils their role in the evolution and extinction of the boreotropical flora[END_REF].

Tempo and mode of Papilio diversification

The diversification analyses investigating rate heterogeneity across Papilio revealed four best equally-likely scenarios of diversification (ΔAICc≤2, Table S2), which refutes the hypothesis of a single diversification rate for the whole genus (∆AICc=14.06; Table S2). S3).

Interestingly, after isolating the subclades that significantly diversify at different rates, two patterns emerge from the backbone (remaining lineages) of best combinations. For the two first best combinations, the backbone is explained by a model with a decreasing speciation rate over time without extinction (Fig. 5B), while the backbone in the two other scenarios follows a model with the same decreasing speciation rate and includes a constant extinction rate (Fig. S7B). In the latter case, net diversification rate becomes negative in the Pliocene onward (ca. 5

Ma), leading to a waxing-waning pattern of diversification (Fig. S7C). This short diversity decline near the present did not affect the trend of the global diversity dynamics because it happens when subclades are already diversifying, thus compensating the decline (Fig. S7B,C,D). The diversity dynamics of these two different patterns in the backbone (decline or not) are similar except with a faster accumulation of lineages during the early Miocene for the scenarios with a decline (Fig. 5, Fig. S7).

These four scenarios highlight common features related to adaptive radiation. First, all scenarios agree to show a two-step diversification history. The first phase of the Papilio radiation proceeded at a high net diversification (ranging from 0.224 to 0.426 events/Myr/lineage depending on the scenario), while the second period of diversification is supported by subclade dynamics having lower speciation rates than at the origin of Papilio (e.g. 0.122 for Heraclides, 0.178 for Papilio), except for Menelaides (0.386 events/Myr/lineage).

The first radiation phase reaches an equilibrium before a short declining phase in half of the best scenarios. For the two other scenarios, the backbones have lower speciation rates at the origin but also contain more lineages (only three or two shifts). These results agree with the general tendency for diversification to slow down as evolution proceeds, considered as one of the most pervasive macroevolutionary principles [START_REF] Rabosky | Ecological limits and diversification rate: alternative paradigms to explain the variation in species richness among clades and regions[END_REF][START_REF] Morlon | Inferring the dynamics of diversification: a coalescent approach[END_REF][START_REF] Moen | Why does diversification slow down?[END_REF][START_REF] Condamine | Assessing the causes of diversification slowdowns: temperature-dependent and diversity-dependent models receive equivalent support[END_REF]. Such diversification slowdowns have often been interpreted as the effect of competition for resources or niche availability [START_REF] Rabosky | Ecological limits and diversification rate: alternative paradigms to explain the variation in species richness among clades and regions[END_REF] or the role of past environmental changes [START_REF] Condamine | Assessing the causes of diversification slowdowns: temperature-dependent and diversity-dependent models receive equivalent support[END_REF]. It is thus possible that the diversity of some Papilio lineages can be at equilibrium or limited by ecological resources, although these lineages are still expanding toward the present but at a slower pace than in the past, supporting the 'damped increase' hypothesis [START_REF] Cornell | Is regional species diversity bounded or unbounded?[END_REF].

Clade-specific rates of diversification can further explain differences in clades' species richness. Although the number of clade shifts varies from one scenario to another, this pattern of radiating subclades within the genus Papilio seems to be supported by a strong signal (Table S2). Clade-heterogeneous diversification has been unveiled across Papilionidae as a whole [START_REF] Condamine | What caused the latitudinal gradient of species diversity in swallowtail butterflies?[END_REF][START_REF] Allio | Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants[END_REF] or for other swallowtail clades (Condamine et al., 2018b), which has been then linked to species' traits or environments. In the case of Papilio, the two most important shifts in diversification are recovered for subgenera Achillides and

Menelaides. These two subclades are island-dwelling groups in the Indo-Malayan and Australasian Archipelagos (Condamine et al., 2013;[START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF], a region where species diversity is the highest at the global scale for this genus (Fig. 2). Their diversification proceeded through repeated island colonizations since the middle Miocene leading to rampant allopatric speciation across the entire archipelago (Condamine et al., 2013;[START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF]. In addition, the subgenus Menelaides shows an early-burst pattern with high speciation rate when it originated (0.386 events/Myr/lineage), followed by a decrease toward the present (0.145 events/Myr/lineage, Fig. 5). Besides their island distribution, Menelaides are wellknown for their Batesian mimicry of aposematic and toxic troidine swallowtails [START_REF] Kunte | The diversity and evolution of Batesian mimicry in Papilio swallowtails butterflies[END_REF][START_REF] Palmer | A shared genetic basis of mimicry across swallowtail butterflies points to ancestral co-option of doublesex[END_REF][START_REF] Kizhakke | Mimicry promotes morphological diversification but not speciation in mimetic swallowtail butterflies[END_REF], which may have spurred their high speciation rates. The subgenus Menelaides diversified faster than any other subgenus, and yet is the most recent clade with subgenus Papilio (Table S3). The latter is often found as a significant shift (Fig. 5), which may be linked to the host-plant shift on Apiaceae [START_REF] Allio | Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants[END_REF] and/or to the glaciation cycles initiated in the late Pliocene [START_REF] Dupuis | Phylogenomic test of mitochondrial clues to archaic ancestors in a group of hybridizing swallowtail butterflies[END_REF].

We did not recover a model incorporating extinction rate in any subclade, which might be artifactual due to the difficulty of estimating extinction rates from phylogenies of extant species alone, or this may be biologically realistic if the nascent subclades did not have enough time to experience extinction to be detectable (e.g. [START_REF] Morlon | Reconciling molecular phylogenies with the fossil record[END_REF]. Assuming these estimates are not biased, this may suggest that subclades of Papilio evolved under a 'museum model of diversity' with a very low extinction rate. Altogether, while their geographic ranges contracted toward the equator (Fig. 4), the genus sustained high rates of species diversification thanks to multiple speciation shifts leading to a global increase of species accumulation toward the present (Fig. 5). This suggests that the tropics are not only the evolutionary source of the Papilio diversity but have also played an important role in mitigating their extinction.

Should more than one genus be recognized within Papilio?

The rank of genus is the most visible rung in the Linnaean hierarchy. It forms part of every species name, indicating shared relationships when there are multiple species in a genus, or unusual distinctness of a species when it constitutes a monotypic genus. Other than monophyly (evidenced by one or more synapomorphies), there are no objective criteria for determining genus boundaries, and even monophyly does not determine how high up on its tree a branch should be cut. Further, when subclades within a genus are elevated to the rank of genus, the information provided by the name about closer relationships is balanced by the lost information about the broader relationships of the group. Nonetheless, numerous attempts have been made

to provide general guidelines for recognizing a genus [START_REF] Talavera | Establishing criteria for higherlevel classification using molecular data: the systematics of Polyommatus blue butterflies (Lepidoptera, Lycaenidae)[END_REF][START_REF] Dorchin | Phylogeny, new generic-level classification, and historical biogeography of the Eucera complex (Hymenoptera: Apidae)[END_REF][START_REF] Sigward | How big is a genus? Towards a nomothetic systematics[END_REF][START_REF] Zhang | Genomes reveal drastic and recurrent phenotypic divergence in firetip skipper butterflies (Hesperiidae: Pyrrhopyginae)[END_REF][START_REF] Nakahara | Description of a new genus and species for a common and widespread Amazonian satyrine butterfly (Lepidoptera: Nymphalidae: Satyrinae: Satyrini)[END_REF], with criteria including the compactness of a group, its distinctness from other such groups, the number of species it contains, its estimated age of divergence, comparability to related genera, consistency with established usage of a name, and degree of confidence in the assessment of its phylogenetic relationships [START_REF] Ashlock | Principles of Systematic Zoology[END_REF]. Balancing and prioritizing potential conflicts among these criteria, while at the same time conveying as much information as possible about relationships between species to non-expert end-users of the names, remains one of the most subjective aspects of systematics.

It is clear that taxonomic stability is a primary aim, if not the primary aim, of the International Code of Zoological Nomenclature (ICZN, 1999(ICZN, , 2012)). The genus Papilio sensu lato has been stable and unambiguously defined by morphology since [START_REF] Munroe | The classification of the Papilionidae (Lepidoptera)[END_REF], and later supported by molecular evidence (e.g. [START_REF] Aubert | Molecular phylogeny of swallowtail butterflies of the tribe Papilionini (Papilionidae, Lepidoptera)[END_REF][START_REF] Caterino | Papilio phylogeny based on mitochondrial cytochrome oxidase I and II genes[END_REF]Zakharov et al., 2004a), although the clades within that genus have continued to be volatile [START_REF] Hancock | Classification of the Papilionidae (Lepidoptera): a phylogenetic approach[END_REF][START_REF] Miller | Phylogenetic studies in the Papilioninae (Lepidoptera: Papilionidae)[END_REF]Zakharov et al., 2004a). Even Chilasa and 'Agehana ', which Hauser et al. (2005) treated as distinct, have continued to move back and forth with each new phylogeny (Zakharov et al., 2004a;[START_REF] Wu | Phylogeny and historical biogeography of Asian Pterourus butterflies (Lepidoptera: Papilionidae): A case of intercontinental dispersal from North America to East Asia[END_REF]. Consequently, we have chosen here to retain the use of the genus Papilio in its broad sense, pending a more rigorous and objective integration of the multiple criteria that may be used to determine the boundary of a genus. For widely recognized and previously stable taxonomic groups, we consider it to be in the best interests of systematists to be conservative in proposing new name changes for anything other than well supported cases of non-monophyly. In our experience, continued name changes for high-profile taxa will consistently elicit frustration and disrespect from the larger community of biologists, conservationists, and the general public. At the same time, if names reflect current knowledge, then advances based on new evidence or even different weighting of evidence will inevitably result in name changes. Fortunately, a solution to this conflict is available through the use of subgeneric names, which allow systematists to refer clearly to their refined taxonomic concepts without putting the burden of increased confusion from name changes onto the users of these species and genus names in other fields.

Conclusion

Building a comprehensive species-level dated phylogeny for a globally distributed insect clade is challenging. We addressed this challenge with the genus Papilio, which currently includes 235 species worldwide. Our study assembles ~80% of the species diversity in a molecular supermatrix and provides a generally robust time-calibrated phylogenetic reference for Papilio.

This new tree confirms previous results such as the early-diverging position of New World

Papilio and the sister relationships between some subgenera and unveils novel relationships like the placement of P. alexanor and the non-monophyly of several subgenera. This phylogenetic framework provides the foundation for a systematic revision, but we urge specialists to refrain from changing the status of subgenera to the rank of genus. Deep nodes within Papilio remain poorly resolved and some subgenera are still poorly sampled, especially in Africa. Future studies with denser taxon sampling and full genomic data will undoubtedly provide a more accurate phylogeny for Papilio and justify a thorough systematic revision. Here we used this dated phylogeny to assess the evolutionary history of Papilio, with estimated divergence times that place its origin in the Oligocene (~30 Ma). Biogeographic analyses suggest a Beringian origin followed by southward dispersals into the tropics, with a pivotal role of the Paleotropics as a biogeographic crossroad for the worldwide colonization of Papilio.

Formal taxonomic changes

Our results confirm the status of 'Agehana' (two species, P. elwesi and P. maraho) within subgenus Pterourus as proposed by [START_REF] Wu | Phylogeny and historical biogeography of Asian Pterourus butterflies (Lepidoptera: Papilionidae): A case of intercontinental dispersal from North America to East Asia[END_REF]. Papilio zalmoxis Hewitson, 1864 is transferred from subgenus Druryia to subgenus Nireopapilio. This finding is also supported morphologically by [START_REF] Huxley | The coloration of Papilio zalmoxis and P. antimachus, and the discovery of Tyndall blue in butterflies[END_REF], who showed that the wing scale structure of P. zalmoxis is identical to P. bromius (valid name: P. chrapkowskoides Storace, 1952), both also containing blue-fluorescent pigment. [START_REF] Zhang | Tracing the origin and evolution of supergene mimicry in butterflies[END_REF][START_REF] Deshmukh | Reproductive barriers and genomic hotspots of adaptation during allopatric species divergence[END_REF].

Specific separation between the mainland Asian taxa and those in the Sundaic islands was also found in the memnon group, but the status of taxa within this group is shown to be further complicated by several taxa traditionally considered as separate species being conspecific with related species [START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF]. The larger memnon group then split into two clades, one containing the bootes and alcmenor species groups and the other P. 1 Table 2. Bayesian tests of topology hypotheses using the stepping-stone sampling (SS). The topology was constrained with previously inferred relationships (Zakharov et al., 2004) that were not recovered in our best topology. The SS estimated the marginal likelihood for each topology and was compared to the marginal likelihood of the best topology (unconstrained). The marginal likelihood is used to calculate Bayes factors to select the topology for the dataset. Results for BF are as follows: non-significant (0<BF<2), positive support (2<BF<6), strong support (6<BF<10), and very strong support (BF>10).

Hypothesis tests on various systematic positions

  two species of the genus Meandrusa (M. payeni and M. sciron) to represent the sister genus of Papilio and define the crown of tribe Papilionini, (2) five species of the tribe Troidini (Battus philenor, Ornithoptera priamus, Parides photinus, Pharmacophagus antenor and Troides helena), which is sister to Papilionini, (3) two species of the tribe Leptocircini (Graphium sarpedon and Lamproptera meges), which is sister to all other Papilioninae, (4) two species of the tribe Parnassiini (Hypermnestra helios and Parnassius apollo), (5) four species of the tribe Zerynthiini (Allancastria louristana, Bhutanitis mansfieldi, Sericinus montela and Zerynthia polyxena), (6) two species of the tribe Luehdorfiini (Archon apollinaris and Luehdorfia puziloi), and (

  specified the nine following subclades: the subgenera Achillides,Chilasa, Druryia, Heraclides, Menelaides, Nireopapilio, Papilio, Pterourus, and the monophyletic group of Lauraceae feeders regrouping Chilasa and Pterourus (Apiaceae and Rutaceae feeders are not monophyletic). We then estimated the number of possible shifts to be tested in combination with corresponding backbones using the get.comb.shift function. The selection of subclades created 319 combinations of subclade(s)/backbone that were compared to the cladehomogeneous birth-death model. We designed and fitted five diversification models to each clade and backbone with the shift.estimate function that determines the best fitting model and most likely combination of shifts. The five models are: (1) a Yule model, where speciation is constant and extinction is null (BCST); (2) a constant birth-death model, where speciation and extinction rates are constant (BCST_DCST); (3) a variable speciation rate model without extinction (BVAR); (4) a variable speciation rate model with constant extinction (BVAR_BCST); and (5) a rate-constant speciation and variable extinction rate model (BCST_BVAR). Diversification rates are defined backward in time such as a positive dependency parameter (α for speciation and β for extinction) reflects a slowdown of rates towards the present. Finally, based on the most likely rate estimates and best combination of shifts, we estimated the diversity dynamics of each clade and resulting backbone recovered in the best shift combination using the apply_prob_dtt and paleodiv function. The files figshare.com/s/a32288db7e6429714a62).

  and clade Chilasa + Pterourus originated in a region comprising the West Nearctic, Central America, East Palearctic, and Sundaland. The clade of Old World Papilio originated in East Palearctic with P. alexanor being sister to all remaining Old World Papilio, which soon after their divergence colonized the Paleotropics through West and East Palearctic + Sundaland and extended to Africa (see shaded area on Fig. 4).

  These shiftconfiguration scenarios include three shifts located at subgenera Achillides, Menelaides and Papilio for the first best scenario (Fig. 5A), two shifts (Achillides and Menelaides) for the second best scenario, five shifts (Achillides, Heraclides, Menelaides, Papilio and Pterourus) for the third best scenario (Fig. S7A), and four shifts (Achillides, Heraclides, Menelaides and Papilio) for the fourth best scenario. Subgenera Achillides and Menelaides are always found as significant shifts. All subclades are better explained by a pure-birth model (BCST), except subgenus Menelaides that follows a model with a decreasing speciation rate through time (BVAR; Fig. 5B, Fig. S7B, Table

Fig. 1 .Fig. 2 .

 12 Fig. 1. Illustration of swallowtail butterfly diversity in the genus Papilio, showing several subgenera that have been widely used since Munroe (1961). Specimens are not uniformly scaled. Pictures from Fabien L. Condamine.

Fig. 3 .

 3 Fig. 3. Bayesian molecular phylogeny of Papilio. The phylogeny was inferred with MrBayes using a traditional partitioning strategy and a reversible-jump MCMC approach for selecting the best fitting substitution models. Posterior probabilities ≥0.95 are indicated at nodes with filled circles. The red dashed rectangle indicates phylogenetic uncertainties between methods. Existing subgeneric classification is shown with colored rectangles delineating subgenera. The outgroups are removed. Asterisks indicate species illustrated on the right. Pictures from Fabien L. Condamine.

Fig. 4 .

 4 Fig. 4. Dated phylogeny and historical biogeography of Papilio. The maximum clade credibility tree shows median ages estimated with a Bayesian uncorrelated lognormal method calibrated with fossils (see Figs S5-7 for additional results). The bottom-left corner map represents the global paleogeography about 25 Ma with continents delimited into 11 areas. Colored areas on the map correspond to colored squares for each node, representing inferred ancestral area(s) with the DEC model, and colored circles at tips, representing present-day distributions. The outgroups are removed. Asterisks indicate species illustrated on the right. Pictures from Fabien L. Condamine.

Fig. 5 .

 5 Fig. 5. Diversification shifts and diversity dynamics estimated for Papilio as estimated by the best-fitting combination of shifts. A) The phylogeny of Papilio with shifts highlighted in colors and best models in parenthesis. Red dots correspond to all tested nodes. B) The evolution of diversification rates through time for the backbone and all subclades that are found as significant shifts. C) Diversity dynamics for the backbone and subclade trees as estimated with the probabilistic approach (dotted line represents the confidence interval of diversity estimates for the backbone). For the sake of clarity, confidence intervals of diversity estimates for subclades are not represented. D) Global diversity dynamics of Papilio obtained by summing all the diversity dynamics for the backbone and subclade trees. Pli.=Pliocene, Q=Quaternary, Myrs=million years.

Table 3 )

 3 . Subgenus Heraclides (Clade 1) was sister to all remaining Papilio, which is composed of a clade mostly including the New World Papilio (Clade 2, subgenera Chilasa and Pterourus) and another clade comprising the Old World Papilio (Clade 3, the remaining subgenera). Although this topology was consistent across methods, branch support for the large clade of Papilio excluding the subgenus Heraclides was not robust (PPCAT=0.75, PPPART=0.98, BPPART=55, and UFBSPART=90). Subgenus Heraclides was always monophyletic with high branch support (PPCAT=1, PPPART=0.96, BPPART=75, and

   Ma) compared to our previous estimation of 22.1 Ma (95% CI=18.0-26.9 Ma;Lewis et al.,

	2015) or 21.9 Ma (95% CI=17.5-26.4 Ma; Wu et al., 2015); and (2) subgenus Achillides
	estimated at 17.1 Ma (95% CI=11.45-26.88 Ma) whereas we previously recovered a 2-million-
	year older age at 19.3 Ma (95% CI=16.4-21.8 Ma;
	1 to 2 million years younger, but it is important
	to note that 95% CIs for these nodes strongly overlap with earlier studies. For instance, we
	estimated the origin of the clade 'Agehana' + Chilasa + Pterourus at 22.73 Ma (95% CI=15.36-

35.9 Ma), whereas we previously estimated this age at 20.9 Ma (95% CI=16.9-25.6 Ma;

[START_REF] Lewis | Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails[END_REF]

or 19.7 Ma (95% CI=17.2-22.9 Ma,

[START_REF] Owens | The latitudinal diversity gradient in New World swallowtail butterflies is caused by contrasting patterns of out-of-and into-thetropics dispersal[END_REF][START_REF] Owens | Comparative phylogenetics of Papilio butterfly wing shape and size demonstrates independent hindwing and forewing evolution[END_REF]

but we have a similar estimation to

[START_REF] Wu | Phylogeny and historical biogeography of Asian Pterourus butterflies (Lepidoptera: Papilionidae): A case of intercontinental dispersal from North America to East Asia[END_REF]

who found an age of 22.63 Ma (95% CI=18.93-26.58 Ma). Slight differences are found in (1) subgenus

  Two taxa in subgenus Druryia are recognized as separate species to Papilio dardanus[START_REF] Tyler | Swallowtail Butterflies of the Americas: A Study in Biological Dynamics[END_REF] placedPapilio victorinus Doubleday, 1844 from Mexico within Pterourus menatius(Hübner, [1819]), but we consider this Central American taxon as specifically distinct from the South American Papilio menatius[START_REF] Owens | The latitudinal diversity gradient in New World swallowtail butterflies is caused by contrasting patterns of out-of-and into-thetropics dispersal[END_REF][START_REF] Owens | Comparative phylogenetics of Papilio butterfly wing shape and size demonstrates independent hindwing and forewing evolution[END_REF], and we reinstate Papilio victorinus Doubleday, 1844 as a separate species (stat.In subgenus Papilio two North American species are confirmed as separate from P. machaon Linnaeus, 1758, namelyPapilio kahli F. & R. Chermock, 1937 (stat. nov.) and Papilio bairdii Edwards, 1866 (stat. rev.). Papilio bairdii includes two subspecies, oregoniaEdwards, 1876 and dodi McDunnough, 1939 (comb. nov.) as well as the nominate subspecies.We found two commonly recurrent patterns in several Asian groups, with speciation between taxa from mainland southern Asia and Sundaland, and between taxa in the Bismarck Archipelago from those in New Guinea, as detailed below.Within subgenus Achillides, five taxa, three with more than one subspecies, are separated at species level based on molecular results fromCondamine et al. (2013b) and Cabusas et al. (2020). Papilio polyctor Boisduval, 1836 (stat. rev.) from northern Pakistan and NW India is separated from Papilio bianor Cramer, 1777, and Papilio hermeli Nuyda, 1992 (stat. rev.) from Mindoro, Philippines is reinstated as a separate species to P. chikae Igarashi, 1965 from Luzon. Papilio daedalus C. Felder & R. Felder, 1861 (stat. rev.), with subspecies angustatus Staudinger, 1888, is confirmed as a separate species to Papilio palinurus Fabricius, 1787. Hiura and Alagar (1971) first separated P. daedalus from P. palinurus based on morphology, but Page and Treadaway (2003a) placed them as conspecific based on perceived similarity of genitalia. Molecular phylogenies clearly show they are separate species, and all subspecies of the two species can easily be separated based on the position of the postdiscal green hindwing band in relation to the anal eyespot. In P. palinurus the lowest point of the band is always above the eyespot, whereas in P. daedalus the band meets the eyespot. The Papilio ulysses group was found to consist of three separate species, Papilio ulysses Linnaeus, 1758 (South Moluccas, New Guinea and most associated islands, and Australia), Papilio telegonus C. Felder & R. Felder, 1860 (stat. rev.) from Northern Moluccas, and Papilio orsippus Godman & Salvin, 1888 (stat. rev.) from the Bismarck Archipelago and Solomon Islands. Nakae (2021) separated Papilio arjuna Horsfield, 1828, found on Sumatra and Java, from mainland Asian P. paris based on the findings of Condamine et al. (2013b), which is confirmed here due to paraphyly with P. karna. That study of Achillides also suggested that P. maackii and P. syfanius Papilio euchenor Guérin-Méneville, 1830 and Papilio depilis Rothschild, 1895 (stat. rev.).Jordan, in Rothschild (1895), stated that forewing scale structure in these two taxa are different but refrained from treating them as separate species. Jordan (1896) then separated the two species, citing several morphological differences including genitalia. Jordan (1908-09) without explanation reunited P. depilis with P. euchenor, which has been followed by all authors until now. This specific separation of Bismarck Archipelago taxa from New Guinea taxa mirrors the separation of P. orsippus from P. ulysses in subgenus Achillides, and similar separation is found in subgenus Menelaides below.In the helenus group of subgenus Menelaides, bothMunroe (1961 -nephelus, nubilus and chaon) and Hancock (1985 -noblei and antonio) included species which have been found not to be monophyletic. Papilio noblei Nicéville, [1889] and P. antonio Hewitson, 1875 are shown here to belong to subgenus Araminta, which is also supported by morphology and wing pattern, particularly on the underside. Papilio nubilus has been shown to be a natural hybrid In the Lesser Sunda Islands, two more taxa in the helenus group are also separate species, Papilio mangarinusRothschild, 1908 (stat. nov.) and Papilio biseriatus Rothschild, 1895 (stat. rev.). P. biseriatus from Timor was previously treated as a separate species by[START_REF] Hancock | Classification of the Papilionidae (Lepidoptera): a phylogenetic approach[END_REF] and differs from the other species in the postdiscal white hindwing patch extending across four hindwing cells as in P. hystaspes from the Philippines, whereas in P. helenus, palawanicus and mangarinus the hindwing patch only covers three cells. As a result of this analysis the range of P. helenus is restricted to mainland Asia and the Malay Peninsula, , not polytes; presumably this similarity is plesiomorphic. Five more species are recognised within the polytes group, two of which are newly separated in this work. The two easternmost species, Papilio ambrax Boisduval, 1832 and Papilio phestus Guérin-Méneville, 1830, were treated as distinct species until[START_REF] Fujioka | Japanese Butterflies and Their Relatives in the World[END_REF] combined them based on morphology and allopatry. However, molecular phylogenies[START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF] and population genetic analyses[START_REF] Zhang | Tracing the origin and evolution of supergene mimicry in butterflies[END_REF][START_REF] Deshmukh | Reproductive barriers and genomic hotspots of adaptation during allopatric species divergence[END_REF] show P. ambrax and P. phestus to be specifically distinct (stat. rev.). Papilio alphenor Cramer, 1776 was separated from P. polytes based on genitalia by[START_REF] Hiura | Studies on the Philippine butterflies chiefly collected by the co-operative survey by the Osaka Museum of Natural History and the National Museum of the Philippines, 1969. Part 1: Papilionidae[END_REF], but subsequently sunk byPage and Treadaway (2003a) without explanation. Molecular phylogenies suggest that P. alphenor is sister to phestus + ambrax, not polytes[START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF], and thus must be treated as a separate species (stat. rev.). The remaining taxa placed in P. polytes also belong to two separate species: the mainland Asian populations are specifically distinct from the Sundaic island taxa, the oldest name for which isPapilio javanus C. Felder, 1862 (stat. nov.

	Papilio saharae Oberthür, 1879 is treated as a subspecies of P. machaon as per Dupuis and
	Sperling (2020), subject to further investigation.
	Yeats, 1776, namely Papilio meriones C. Felder & R. Felder, 1864 (stat. rev.) from
	Madagascar and Papilio humbloti Oberthür, 1888 (stat. rev.) from Comoros Islands.
	may be conspecific; but a recent genomic study clarified that mitochondrial gene exchange occurs between these two species in western China without significant exchange of nuclear In subgenus Pterourus, rev.). Taiwan and Japan.

genes

[START_REF] Xiong | Admixture of evolutionary rates across a butterfly hybrid zone[END_REF]

. As a result, we retain P. maackii and P. syfanius as separate species.

Subgenus Euchenor has previously been regarded as monobasic, but DNA analysis shows that the subgenus consists of two distinct species

[START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF]

): between P. nephelus and P. polytes rather than a valid species

(Tsukada and Nishiyama 1980: 307)

. We confirmed that Papilio hystaspes C. Felder & R. Felder, 1862, separated from helenus by

[START_REF] Hiura | Studies on the Philippine butterflies chiefly collected by the co-operative survey by the Osaka Museum of Natural History and the National Museum of the Philippines, 1969. Part 1: Papilionidae[END_REF]

based on male genitalia and treated as separate by

[START_REF] Hancock | Classification of the Papilionidae (Lepidoptera): a phylogenetic approach[END_REF]

but placed within P. helenus by

Page and Treadaway (2003b)

, is a distinct species (stat. rev.) which branches off before the traditional helenus taxa and sataspes. However, we also found that several taxa always considered to belong to P. helenus are separate species. The first species is Papilio daksha (stat. rev.) from Southern India. Joshi and Kunte (2022) analyzed specimens of ssp. enganius Doherty, 1891 from Sumatra, Java and Borneo and found that they are sister to P. sataspes from Sulawesi and associated islands, not P. helenus from mainland SE Asia and the Malay Peninsula. The oldest name for the taxon in the Sundaic islands of Sumatra, Java, Borneo and Palawan, is Papilio palawanicus Staudinger, 1888 (stat. nov.).

Within the polytes group, Papilio protenor Cramer, 1775 is sister to all other species. This does not correspond to its expected position, as the early stages are very similar to P. memnon). Papilio polytes, alphenor and javanus also show prezygotic and postzygotic barriers to hybridization including assortative mating and low fitness of hybrid progeny, thus being reproductively isolated, with genome-wide and population genetic signatures of being highly diverged distinct species

  Originally these were considered separate species, but they were treated as conspecific by[START_REF] Evans | The identification of Indian butterflies[END_REF], followed by[START_REF] Talbot | The Fauna of British India including Ceylon and Burma[END_REF] and subsequent authors. Previous molecular phylogenies show that P. bootes and P. janaka are significantly genetically distinct[START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF], and are treated as different species although they are sympatric in NE Myanmar. The other branch of the first clade contains Papilio acheron Grose-Smith, 1887,

	forbesi Grose-Smith, 1883 and lampsacus Boisduval, 1836 (this last taxon not sequenced due
	to rarity) with Papilio alcmenor C. Felder & R. Felder, 1865 and thaiwanus Rothschild, 1898

memnon and close relatives. There are two branches within the first clade, one branch containing just two species, Papilio bootes Westwood, 1842 and Papilio janaka Moore, 1857 (stat. rev.).

sisters within this group. Previously P. acheron, forbesi and lampsacus were considered as very close to P. memnon due to their similar appearance, but this relationship was not confirmed by our analysis.

Marginal likelihood (SS) Bayes factor

  Monophyly of the subgenus Druryia (i.e. Druryia + Nireopapilio) -97,269.70 190.31 Monophyly of the subgenus Princeps (i.e. Princeps + dardanus sp. gr.)

	Unconstrained MrBayes phylogenetic analysis	-97,079.39	-
	Monophyly of the subgenus Pterourus (i.e. excluding Agehana)	-97,127.68	48.29
		-97,277.63	198.24
	Monophyly of the subgenus Menelaides (i.e. Menelaides + Araminta)	-97,145.29	65.9
	Monophyly of the New World Papilio	-97,094.08	14.69
	P. alexanor sister to Agehana + Chilasa + Pterourus	-97,141.38	61.99

Table 3 .

 3 Phylogenetic, dating, and biogeographic results for the main nodes of Papilio. Clades 1 to 3 are shown in Fig.3. 1460

			Node supports		Median node ages, Ma (95% CI)
	Clades							Ancestral area estimates Biogeographic interpretation
		PPCAT	PPPART	BSPART	UFBSPART	Fossil calibrations	Secondary calibrations
	Papilio sensu lato	1	1	100	100	30.06 (20.66 -47.42)	29.95 (23.97 -37.34)	EP + WN + CA + WA	Origin centred on Beringia
	Clade 1: Heraclides	1	0.96	75	100	21.22 (13.95 -33.86)	21.2 (16.07 -27.48)	CA	Central America
	Clade 2: Papilio sensu lato excl. Heraclides	0.75	0.98	55	90	28.57 (19.98 -45.45)	28.5 (22.89 -35.55)	EP + WN + CA + WA	Origin centred on Beringia
	Chilasa + Pterourus + 'Agehana'	0.99	1	91	100	22.73 (15.36 -35.9)	22.64 (17.33 -28.74)	EP + WN + CA + WA	Origin centred on Beringia
	Chilasa	1	1	99	100	16.68 (12.49 -29.71)	18.63 (14.02 -24.44)	WA	Indonesia
	Pterourus + 'Agehana'	1	1	79	99	20.38 (13.8 -32.28)	20.3 (15.3 -25.92)	EP + WN + CA	Origin centred on Beringia
	'Agehana'	1	1	100	100	0.28 (0.09 -0.63)	0.27 (0.09 -0.53)	EP + WA	Asia and Indonesia
	Clade 3: Old World Papilio	1	0.99	58	85	26.93 (18.62 -42.62)	26.86 (21.52 -33.51)	EP	East Palearctic
	Old World Papilio excl. Alexanoria	1	1	86	100	23.74 (16.45 -37.49)	23.67 (18.96 -29.58)	WP + EP + AF + WA	Paleotropics
	Clade 3a: Druryia + Euchenor	0.82	0.99	74	99	22.32 (15.3 -35.23)	22.23 (17.46 -27.95)	WP + EP + AF + WA	Paleotropics
	Euchenor	1	1	100	100	5.49 (2.74 -9.71)	5.33 (2.84 -8.42)	AU	Australasia
	Druryia	1	1	99	100	17.95 (12.07 -28.62)	17.88 (13.61 -22.95)	AF	Afrotropics
	Clade 3b: Eleppone + Araminta	-	0.72	50	93	21.09 (14.82 -33.9)	20.99 (16.4 -26.47)	WA	Indonesia
	Eleppone	1	1	97	100	9.72 (4.41 -17.0)	9.54 (4.77 -15.03)	AU	Australasia
	Araminta	1	1	100	100	14.07 (8.87 -22.59)	13.9 (9.89 -18.61)	WA	Indonesia
	Clade 3c + Clade 3d + Clade 3e	0.99	1	72	99	21.35 (15.08 -33.97)	21.25 (16.97 -26.59)	WP + EP + AF + WA	Paleotropics
	Clade 3c: Sinoprinceps + Papilio sensu stricto	1	1	97	100	17.39 (11.36 -27.85)	17.27 (13.1 -22.34)	EP	East Palearctic
	Sinoprinceps	1	1	100	100	1.5 (0.63 -2.82)	1.44 (0.67 -2.45)	EP + WA	Asia and Indonesia
	Papilio sensu stricto	1	1	100	100	10.3 (6.26 -16.69)	10.2 (7.01 -13.85)	EP + WN	Origin centred on Beringia
	Clade 3d + Clade 3e	0.99	1	60	99	20.17 (14.01 -31.92)	20.09 (16.09 -25.23)	WP + EP + AF + WA	Paleotropics
	Clade 3d: Nireopapilio	1	1	76	98	16.1 (10.21 -25.51)	15.98 (11.78 -20.65)	AF	Afrotropics
	Clade 3e: Achillides + Princeps + Menelaides	-	0.91	54	96	19.52 (13.5 -30.98)	19.41 (15.57 -24.39)	WP + EP + AF + WA	Paleotropics
	Achillides	1	1	85	99	17.1 (11.45 -26.88)	17.0 (13.31 -21.62)	WA + AU	Indonesia and Australasia
	Princeps + Menelaides	0.93	1	70	99	18.21 (12.6 -29.05)	18.15 (14.37 -22.86)	WP + EP + AF + WA	Paleotropics
	Princeps	1	1	100	100	11.29 (6.93 -18.01)	11.21 (8.0 -14.97)	WP + EP + AF + MD	Paleotropics
	Menelaides	1	1	99	100	13.94 (9.4 -21.98)	13.94 (10.9 -17.65)	WA	Indonesia

Acknowledgements

This project has received funding from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (project GAIA, agreement no. 851188), and an 'Investissement d'Avenir' grant from the Agence Nationale de la Recherche (project CASMA, CEBA, ref. ANR-10-LABX-25-01) to F.L.C.; a Ramanujan Fellowship from the Dept. of Science and Technology, Govt. of India, to K.K., and support of the Dept. of Atomic Energy, Govt. of India, under project nos. 12-RandD-TFR-5.04-0800 and 12-RandD-TFR-5.04-0900 to TIFR/NCBS; a Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery Grant to F.A.H.S. Sequencing of most Indian Papilio was done at NCBS under research and voucher specimen collection permits issued to K.K. by the state forest departments in Kerala (permit no. WL 10-3781/2012 dated 18/12/2012, and GO (RT) No. 376/2012/FandWLD dated 26/07/2012) and Nagaland (permit no.

CWL/GEN/240/522-39 dated 14 August 2012). Alberto Fracasso (Italy) and Laurie Wills (New Zealand) are thanked for providing samples of some hard to obtain taxa. 1421 1422

Molecular phylogenies confirmed that the Philippine taxon rumanzovia Eschscholtz, 1821 is genetically conspecific with Moluccan P. deiphobus [START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF], as stated on morphological grounds by Page and Treadaway (2003a,b). There is a wide genetic divergence between the mainland Asian and the Sundaland taxa previously united under P. memnon, which clearly represent two separate species, Papilio agenor Linnaeus, 1758 (stat.

rev.) and P. memnon [START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF]. Two traditionally well-regarded species, Papilio polymnestor Cramer, 1775 from peninsular India and Sri Lanka, and Papilio mayo Atkinson, [1874] from the Andaman Islands, do not merit species status. Papilio polymnestor was nested within P. agenor [START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF] and should therefore be treated as a subspecies, Papilio agenor polymnestor Cramer, 1775 (stat. rev.). Likewise, P. mayo was nested within and therefore confirmed as conspecific with P. memnon, and should be treated as its subspecies, Papilio memnon mayo Atkinson, [1874] (stat. rev.).

The nephelus group comprises four species. Papilio castor Westwood, 1842 and P. dravidarum Wood-Mason, 1880 are sister species within this group, and the taxon Papilio mahadeva Moore, [1879] is confirmed to be conspecific with P. castor as stated on morphological grounds by [START_REF] Cotton | A preliminary annotated checklist of the Papilionidae of Laos with notes on taxonomy, phenology, distribution and variation (Lepidoptera, Papilionoidea)[END_REF]. Papilio chaon Westwood, 1844 and P. nephelus Boisduval, 1836, from mainland SE Asia and Sundaland respectively, were originally described as separate species but were treated as conspecific by [START_REF] Igarashi | Papilionidae and their early stages[END_REF] based on early stages. Molecular phylogenies showed that genetically nephelus and chaon are highly divergent [START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF], and thus must be treated as separate species, P. nephelus and P. chaon (stat. rev.).

Within the aegeus clade, P. godeffroyi was found to be the sister taxon to Papilio oritas Godman & Salvin, 1879 (stat. rev.) from New Britain, New Ireland and New Hannover, which was placed within P. aegeus by Jordan (1909). In molecular phylogenies, P. oritas diverged before the remaining species in the aegeus group [START_REF] Joshi | Polytypy and systematics: diversification of Papilio swallowtail butterflies in the biogeographically complex Indo-Australian Region[END_REF], and thus is not conspecific with P. aegeus. This is another example of speciation within the Bismarck Islands.

Papilio inopinatus Butler, 1883 from Timor was found to be nested within P. aegeus (syn. nov.), thus it becomes Papilio aegeus inopinatus (comb. nov.).

The most complex clade within subgenus Menelaides is the fuscus group. The species Papilio fuscus Goeze, 1779 as recognised by [START_REF] Hancock | The Princeps fuscus complex (Lepidoptera: Papilionidae)[END_REF] in the most recent revision of this species is not a monophyletic entity.

Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Table S1. Geographic distribution of Papilio species. The tables include the geographic data for (1) all species listed in Table 1, and (2) the species that have been sampled in the current phylogenetic tree. We coded the presence [START_REF]Changes to subspecies nomenclature resulting from literature review and this study. 1462 1463 Species New subspecies combinations Papilio (Achillides) orsippus Godman & Salvin[END_REF] or absence (0) in each of the 11 defined biogeographic regions. Histogram plots show the number of species per region.