A new subspecies of Papilio saharae Oberthür, 1879 (Lepidoptera: Papilionidae) from Lampedusa, Italy.

An investigation into the only representative of the genus Papilio on the island of Lampedusa, the largest island of the Pelagian group, was conducted as part of a broader Mediterranean-wide study on the machaon complex. Over a three-year period, adults, larvae and ova were collected for further research during field visits, while an in-house breeding programme involving wild-collected gravid females was initiated with a view to examine an adequately sized series of specimens. A total of 38 adults, including 23 male specimens, >150 ova, 233 larvae, and 220 pupae were examined. In addition to a thorough morphometric assessment, statistical tests were performed using one-way ANOVA and multivariate analysis. Results demonstrate morphological characters of P. saharae and, to a lesser degree, of P. machaon, suggesting that the taxon is plausibly the outcome of a hybrid swarm whose occurrence on the island was facilitated by Pleistocene lowstands (when the island was physically connected with the north African continental landmass, the taxon's centre of origin). Based on results, involving all four stages of metamorphosis, it is proposed to 'anchor' the taxon to Papilio saharae Oberthür, 1879 and assign it subspecific status, aferpilaggi ssp. nov.

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

The swallowtail genus Papilio (Lepidoptera: Papilionidae) is species rich, distributed worldwide, and has broad morphological habits and ecological niches. Because of its elevated species richness, it has been historically difficult to reconstruct a densely sampled phylogeny for this clade. Here we provide a taxonomic working list for the genus, resulting in 235 Papilio species, and assemble a molecular dataset of seven gene fragments representing ca. 80% of the currently described diversity. Phylogenetic analyses reconstructed a robust tree with highly supported relationships within subgenera, although a few nodes in the early history of the Old World Papilio remain unresolved. Contrasting with previous results, we found that Papilio alexanor is sister to all Old World Papilio and that the subgenus Eleppone is no longer monotypic. The latter includes the recently described Fijian Papilio natewa with the Australian Papilio anactus and is sister to subgenus Araminta (formerly included in subgenus Menelaides) occurring in Southeast Asia. Our phylogeny also includes rarely studied (P. antimachus, P. benguetana) or endangered species (P. buddha, P. chikae). Taxonomic changes resulting from this study are elucidated. Molecular dating and biogeographic analyses indicate that Papilio originated ca. 30 million years ago (Oligocene), in a northern region centered on Beringia. A rapid early Miocene radiation in the Paleotropics is revealed within Old World Papilio, potentially explaining their low early branch support. Most subgenera originated in the early to middle Miocene followed by synchronous southward biogeographic dispersals and repeated local extirpations in northern latitudes. This study provides a comprehensive phylogenetic framework for Papilio with clarification of subgeneric systematics and species taxonomic changes enumerated, which will facilitate further studies to address questions on their ecology and evolutionary biology using this model clade.

Species Richness of Papilionidae Butterflies (Lepidoptera: Papilionoidea) in the Hengduan Mountains and Its Future Shifts under Climate Change.

The family of Papilionidae (Lepidoptera: Papilionoidea) is a group of butterflies with high ecological and conservation value. The Hengduan Mountains (HMDs) in Southwest China is an important diversity centre for these butterflies. However, the spatial distribution pattern and the climate vulnerability of Papilionidae butterflies in the HDMs remain unknown to date. The lack of such knowledge has already become an obstacle in formulating effective butterfly conservation strategies. The present research compiled a 59-species dataset with 1938 occurrence points. The Maxent model was applied to analyse the spatial pattern of species richness in subfamilies Parnassiinae and Papilioninae, as well as to predict the response under the influence of climate change. The spatial pattern of both subfamilies in the HDMs has obvious elevation prevalence, with Parnassiinae concentrated in the subalpine to alpine areas (2500-5500 m) in western Sichuan, northwestern Yunnan and eastern Tibet, while Papilioninae is concentrated in the low- to medium-elevation areas (1500-3500 m) in the river valleys of western Yunnan and western Sichuan. Under the influence of climate change, both subfamilies would exhibit northward and upward range shifts. The majority of Parnassiinae species would experience drastic habitat contraction, resulting in lower species richness across the HDMs. In contrast, most Papilioninae species would experience habitat expansion, and the species richness would also increase significantly. The findings of this research should provide new insights and a clue for butterfly diversity and climatic vulnerability in southwestern China. Future conservation efforts should be focused on species with habitat contraction, narrow-ranged distribution and endemicity with both in situ and ex situ measures, especially in protected areas. Commercialised collecting targeting these species must also be regulated by future legislation.

Checklist of Yunnan Papilionidae (Lepidoptera: Papilionoidea) with nomenclatural notes and descriptions of new subspecies.

A checklist of the Papilionidae of Yunnan is presented, with nomenclatural and taxonomic changes made. In the nomenclatural section, the junior homonym Papilio bootes nigricans Rothschild, 1895 is replaced by Papilio bootes nigricauda Lamas & Cotton nom. nov., Chilasa (Cadugoides) epycides muhabbet Koak, 2005 is synonymised with Papilio epycides camilla Rousseau-Decelle, 1947 syn. nov., Graphium cloanthus nyghmat Koak & Kemal, 2000 is placed as a junior objective synonym syn. nov. of Graphium cloanthus clymenus (Leech, 1893), and Papilio astorion Westwood, 1842 is shown to have priority over Papilio varuna White, 1842, thus the valid species name is Atrophaneura astorion (Westwood, 1842) comb. nov. In the main checklist, five new subspecies are described: Parnassius cephalus haba Hu & Cotton ssp. nov., Lamproptera curius hsinningae Hu, Zhang & Cotton ssp. nov., Lamproptera curius yangtzeanus Hu & Cotton ssp. nov., Graphium macareus vadimi Cotton & Hu ssp. nov., and Papilio krishna benyongi Hu & Cotton ssp. nov. The First Reviser Principle under the ICZN Code is invoked to solve four taxonomic problems, and 18 names are synonymised with explanations, notably Papilio machaon venchuanus Moonen, 1984 syn. nov., which is synonymised with Papilio machaon schantungensis Eller, 1936. Byasa genestieri (Oberthr, 1918) stat. nov. is separated from Byasa latreillei (Donovan, 1826), and Papilio everesti Riley, 1927 stat. nov. and P. verityi Fruhstorfer, 1907 stat. nov. are separated from Papilio machaon Linnaeus, 1758 as species. Taxa that require further confirmation of their presence in Yunnan and those that do not occur in Yunnan are enumerated.

Graphium septentrionicolus Page amp; Treadaway, 2013 (Lepidoptera: Papilionidae) is a distinct species.

After molecular and morphological analyses, the taxon septentrionicolus Page Treadaway, 2013 is shown to be a distinct species, and Graphium adonarensis (Rothschild, 1896) is placed as conspecific with Graphium sarpedon (Linnaeus, 1758). Graphium huangshanensis Wu Ma, 2016 syn. nov. is synonymised with G. septentrionicolus.

Genome-wide macroevolutionary signatures of key innovations in butterflies colonizing new host plants.

The mega-diversity of herbivorous insects is attributed to their co-evolutionary associations with plants. Despite abundant studies on insect-plant interactions, we do not know whether host-plant shifts have impacted both genomic adaptation and species diversification over geological times. We show that the antagonistic insect-plant interaction between swallowtail butterflies and the highly toxic birthworts began 55 million years ago in Beringia, followed by several major ancient host-plant shifts. This evolutionary framework provides a valuable opportunity for repeated tests of genomic signatures of macroevolutionary changes and estimation of diversification rates across their phylogeny. We find that host-plant shifts in butterflies are associated with both genome-wide adaptive molecular evolution (more genes under positive selection) and repeated bursts of speciation rates, contributing to an increase in global diversification through time. Our study links ecological changes, genome-wide adaptations and macroevolutionary consequences, lending support to the importance of ecological interactions as evolutionary drivers over long time periods.

Revision of Pazala Moore, 1888: The Graphium (Pazala) alebion and G. (P.) tamerlanus Groups, with Notes on Taxonomic and Distribution Confusions (Lepidoptera: Papilionidae).

Three Graphium species belonging to two species groups of the subgenus Pazala, the alebion and tamerlanus groups, were examined in molecular and morphological studies, and their female genitalia are reported for the first time. Their relationship with other species groups within the subgenus is assessed and their divergence times are estimated. We find that G. (P.) alebion is the first lineage to diverge within Pazala in the early Miocene (20 Ma) and that G. (P.) tamerlanus and G. (P.) parus are sister species and diverged from each other in the late Miocene (7 Ma). A revision of the four recognised taxa belonging to three species is presented, and historical misidentification of these taxa and their geographic ranges are explained.

Are the Yellow and Red Marked Club-Tail Losaria coon the Same Species?

Losaria coon (Fabricius, 1793) is currently comprised of ten subspecies, which were originally described under two names, Papilio coon and P. doubledayi before 1909, when they were combined as one species. The main difference between them is the colour of abdomen and hindwing subterminal spots-yellow in coon and red in doubledayi. Wing morphology, male and female genitalia, and molecular evidence (DNA barcodes) were analysed for multiple subspecies of L. coon and three other Losaria species-rhodifer, neptunus, and palu. Our molecular data support the separation of L. coon and L. doubledayi stat. rev. as two distinct species, with L. rhodifer positioned between them in phylogenetic analyses. Wing morphology and genitalic structures also confirm the molecular conclusions. Our findings divide L. coon into two species occupying different geographic ranges: with L. coon restricted to southern Sumatra, Java, and Bawean Island, while L. doubledayi occurs widely in regions from North India to northern Sumatra, including Hainan and Nicobar Islands. Hence, future conservation efforts must reassess the status and threat factors of the two species to form updated strategies.

A New Species of the Graphium (Pazala) mandarinus Group from Central Vietnam (Lepidoptera: Papilionidae).

The Graphium (Pazala) mandarinus group was recently defined and the status of taxa as well as the number of species was revised. We report here the discovery of a new species from Kon Tum plateau of the Truong Son (Annamite) Range of Central Vietnam, which we describe based on morphological and molecular evidence. Molecular phylogeny shows that the new taxon, G. (P.) wenlingae Hu, Cotton Monastyrskii sp. nov., is sister to G. (P.) daiyuanae Hu, Zhang Cotton, 2018 plus G. (P.) confucius Hu, Duan Cotton, 2018. Molecular dating analysis further suggests that this new species diverged from its sister clade in the Pliocene (~3.5 million years ago). The new taxon constitutes the eighth and southernmost species of the mandarinus group.

Revision of Pazala Moore, 1888: The Graphium (Pazala) mandarinus (Oberthür, 1879) Group, with Treatments of Known Taxa and Descriptions of New Species and New Subspecies (Lepidoptera: Papilionidae).

The previously recognised closely related species Graphium (Pazala) mandarinus (Oberthür, 1879) and G. (P.) sichuanica (Koiwaya, 1993) are shown to comprise seven species as a result of both molecular and morphological analysis. Molecular dating analysis is also performed on the mandarinus group in order to investigate the divergence time of the taxa. Two taxa, G. (P.) garhwalica (Katayama, 1988) stat. nov. and G. (P.) paphus (de Nicéville, 1886) stat. nov., are raised from subspecific to specific status; G. (P.) hoeneanus Cotton Hu nom. nov., stat. rev. is separated from sichuanica at species level; and two previously unrecognised new species, G. (P.) daiyuanae Hu, Zhang Cotton sp. nov. and G. (P.) confucius Hu, Duan Cotton sp. nov. are described from Vietnam and China respectively, the latter being sympatric with nominate G. (P.) mandarinus. The identity of the lectotype of G. (P.) mandarinus is confirmed and a lectotype is designated for the taxon Papilio Glycerion Gray, 1831. A new subspecies of G. (P.) mandarinus is described from western Yunnan and northern Myanmar, G. (P.) mandarinus stilwelli Cotton Hu ssp. nov.

Role of Caribbean Islands in the diversification and biogeography of Neotropical Heraclides swallowtails.

Numerous hypotheses on the evolution of Neotropical biodiversity have stimulated research to provide a better understanding of diversity dynamics and distribution patterns of the region. However, few studies integrate molecular and morphological data with complete sampling of a Neotropical group, and so there has been little synthesis of the multiple processes governing biodiversity through space and time. Here, a total-evidence phylogenetic approach is used to reconstruct the evolutionary history of the butterfly subgenus Heraclides. We used DNA sequences for two mitochondrial genes and one nuclear gene and coded 133 morphological characters of larvae and adults. A robust and well-resolved phylogeny was obtained using several analytical approaches, while molecular dating and biogeographical analyses indicated an early Miocene origin (22 Mya) in the Caribbean Islands. We inferred six independent dispersal events from the Caribbean to the mainland, and three from the mainland to the Caribbean, and we suggest that cooling climates with decreasing sea levels may have contributed to these events. The time-calibrated tree is best explained by a museum model of diversity in which both speciation and extinction rates remained constant through time. By assessing both continental and fine-scale biodiversity patterns, this study provides new findings, for instance that islands may act as source of diversity rather than as a sink, to explain spatio-temporal macroevolutionary processes within the Neotropical region.

Discovery of a third species of Lamproptera Gray, 1832 (Lepidoptera: Papilionidae).

A newly discovered, third species of the genus Lamproptera (Lepidoptera: Papilionidae) is described, 183 years after the second currently recognised species was first named. Lamproptera paracurius Hu, Zhang & Cotton sp. n., from N.E. Yunnan, China, is based on marked differences in external morphology and male genital structure. The species is confirmed as a member of the genus, and detailed comparisons are made with other taxa included in the genus. Keys to Lamproptera species based on external characters and male genitalia are included.

Fine-scale biogeographical and temporal diversification processes of peacock swallowtails (Papilio subgenus Achillides) in the Indo-Australian Archipelago.

© The Willi Hennig Society 2012. ABSTRACT: Explanations for the high species diversity of the Indo-Australian Archipelago are often challenged by the region's complex climatic and geological histories. Here, we investigated the evolutionary history of swallowtail butterflies of the Papilio subgenus Achillides, comprising up to 25 recognized species and about 100 subspecies distributed across the Indo-Australian Archipelago. To estimate the relative contributions of factors influencing their biodiversity, we used DNA sequences to infer the phylogeny and species limits of 22 species including most of their subspecies. We recovered a highly resolved and well-supported phylogeny for the subgenus, and clarified some taxonomic ambiguities at the species level. The corresponding DNA-based species phylogeny was then employed to reconstruct their historical biogeography using relaxed-clock and parametric-based analyses. Molecular dating and biogeographical analyses showed that Achillides originated around 19 Ma in Sunda + Wallacea. Biogeographical reconstructions indicated that geological vicariance shaped the early evolutionary history of Achillides whereas dispersal influenced late diversification. Birth-death likelihood analyses allowed exploration of their tempo and mode of diversification. We detected several shifts in diversification rates that are attributable to past climate-induced biogeographical events. By assessing both regional and fine-scale biodiversity patterns, this study brings new findings to a biogeographical understanding of the Indo-Australian Archipelago.

Phylogenetics and divergence times of Papilioninae (Lepidoptera) with special reference to the enigmatic genera Teinopalpus and Meandrusa.

© The Willi Hennig Society 2010. ABSTRACT: Phylogenetic relationships of 18 genera of the swallowtail subfamily Papilioninae, four genera of Parnassiinae, and the monobasic Baroniinae are inferred based on 94 morphological characters and 5616 bp DNA from seven genes (16S, COI, COII, ND1, ND5, EF-1 alpha and wingless). Bayesian likelihood analyses show that Baroniinae are the sister of a clade comprising Parnassiinae and Papilioninae. Four Papilioninae tribes are recognized, Leptocircini, Teinopalpini, Papilionini and Troidini, with Leptocircini being the sister of the remaining tribes. Meandrusa and Teinopalpus are sister taxa and comprise the tribe Teinopalpini, which is the sister of a clade comprising Papilionini and Troidini. The tribe Troidini (pipevine swallowtails) comprises two subtribes: Battina (including only Battus) and Troidina. The endemic Madagascan genus Pharmacophagus is consistently placed as the sister to the remaining Troidina. The non-Pharmacophagus Troidina are tentatively divided into a Neotropical lineage and an Australasian lineage. Dispersal-vicariance analyses indicate that past dispersal events are most important for explaining current distribution patterns of Papilionidae. However, the division of the non-Pharmacophagus Troidina into a Neotropical lineage and an Australasian lineage is possibly due to the final break-up of southern Gondwana. A fossil-calibrated relaxed Bayesian molecular clock analysis confirms that the ages of the lineages fit this scenario. The basal lineages leading to the current subfamily-level diversity of Papilionidae probably arose around the K/T boundary. Analyses of larval host-plant relationships within Papilionidae show very little phylogenetic pattern. However, Aristolochiaceae-feeding apparently evolved independently in non-Parnassiini parnassiines and Troidini.