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.

Identifying Immigrating Sogatella furcifera (Hemiptera: Delphacidae) Using Field Cages: A Case Study in the Yuanjiang (Red River) Valley of Yunnan, China.

The white-backed planthopper, Sogatella furcifera (Horváth), is a devastating migratory rice pest in South China; lack of effective methods to identify immigrating populations is the main cause of difficulties in outbreak forecasting, active prevention, and control. The current study set up field cages (2 × 2 × 3 m each, US-80 standard nylon mesh) in both early- and mid-season paddies in Yuanjiang (Red River) Valley in Yunnan, China, in 2012 and 2014. The immigrating population was successfully separated from the local population of S. furcifera and identified using statistical comparisons. The findings showed that densities of macropterous adults outside the cages were all significantly higher than those inside the cages on both early- and mid-season rice in both years, whereas the densities of young nymphs and old nymphs showed no significant differences. This indicated that immigrations were occurring, the earliest of which occurred on early-season rice in early May and reached its peak in mid-late May before a rapid collapse in both years. In contrast, the immigration on mid-season rice showed a continuous decline or fluctuation throughout the entire period. Analyses demonstrated that the migration process of S. furcifera in the Yuanjiang Valley features continuous immigration from the adjacent southern parts of Yunnan, which may represent most migration events in Yunnan during the outbreak period of a year. The findings of this case study could benefit our understanding of planthopper migration and outbreaks in other parts of China, especially where the outbreak pattern is very different from Yunnan.

Density, Demography, and Influential Environmental Factors on Overwintering Populations of Sogatella furcifera (Hemiptera: Delphacidae) in Southern Yunnan, China.

Sogatella furcifera (Horváth) (Hemiptera: Delphacidae) is the most serious pest on rice in southwestern China. Yunnan province is within this region and is a major overwintering area for S. furcifera in China. This field study was carried out over 4 yr (2010-2013) and focused on S. furcifera distribution, population density, and demography, as well as the relationship between various environmental factors and the distribution and density of overwintering S. furcifera in Yunnan. Our study demonstrated that overwintering populations of S. furcifera mainly occurred in valleys and lowlands below 25.02°N and 1,608 m above sea level (a.s.l.), where ratooning rice was present [corrected]. The overwintering range of S. furcifera has expanded in Yunnan compared with 20 yr ago, and regional climate change is believed to be the main contributing cause for this expansion. Environmental factor analysis showed that the mean air temperature of the coldest quarter and precipitation of the coldest quarter were two key factors that were strongly linked to the overwintering distribution and density of S. furcifera in Yunnan. Wintertime temperature was the principal influencing factor to determine the distribution and density of S. furcifera, while the effect of precipitation was indirect in that it influenced the insect's distribution via its host. This study documented the major overwintering areas of S. furcifera in Yunnan, which can be used to predict outbreak potential in the following spring. Hence, key climatic factors, overwintering distribution, and density of S. furcifera should be used when forecasting outbreaks in spring.

Oviposition preference and offspring performance of Mechoris ursulus (Coleoptera: Attelabidae) in Cyclobalanopsis glaucoides (Fagales: Fagaceae) and Quercus franchetii (Fagales: Fagaceae) in Central Yunnan, China.

Mechoris ursulus (Roelofs) (Coleoptera: Attelabidae) is a pest weevil of Fagaceae oak trees in eastern Asia. The female has a distinct branch-cutting behavior in conjunction with oviposition in the acorns of its host plant. This study analyzed the factors influencing oviposition preference by carrying out continuous field surveys over the course of 2009-2010 and through laboratory rearing. The field survey showed that for both of the hosts, Cyclobalanopsis glaucoides Schottky (Fagales: Fagaceae) and Quercus franchetii Skan, M. ursulus preferred branches with fewer acorns and larger acorns on the same branch for oviposition. Laboratory rearing experiments showed offspring performance (i.e., survival rate and fresh weight of larvae) was significantly and positively correlated with acorn size. Preference for larger acorns could maximize the fitness of offspring by providing sufficient food source and space.

Projecting distribution of the overwintering population of Sogatella furcifera (Hemiptera: Delphacidae), in Yunnan, China with analysis on key influencing climatic factors.

Sogatella furcifera (Horváth) is the most threatening migratory rice pest in Yunnan, China. S. furcifera overwinters in low- altitude basins and valleys in southern Yunnan and migrates northward in spring and summer of the following year, causing serious damage during migration. The overwintering distribution, areas, and spatial pattern of S. furcifera are relevant to the migration and outbreak of this pest. Based on a 4-yr field survey (2010-2013), this study projected areas suitable for S. furcifera to overwinter using a species distribution model, and analyzed the key influencing climatic factors using principal component analysis (PCA) and ecological niche factor analysis (ENFA). Our field survey showed that the northern latitudinal- and upper elevation limits of overwintering S. furcifera was 25.4° N and 1,608 m in western Yunnan and 24.2° N and 1,563 m in eastern Yunnan. The species distribution model produced a fragmented distribution pattern, with most of which in western Yunnan and only a few in eastern Yunnan. The PCA and ENFA analyses showed that the mean temperature of the driest quarter and the precipitation of the coldest quarter significantly influenced the distribution of S. furcifera in winter. The results suggested that the complex topography, spatial differences in winter temperatures, and host availability altogether determined the distribution of overwintering S. furcifera. Compared with previous surveys, the northern latitudinal- and upper elevation limits of overwintering S. furcifera were higher, while the population became rarer in some suitable areas due to change of farmland utilization in winter and possibly climate change.

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.

First description of the females of Qinorapalaqinlingana Chou & Wang, 1995 (Lepidoptera, Lycaenidae) from Shaanxi and Sichuan Provinces, western China.

Background: The family Lycaenidae is a widely distributed and species-rich group with approximately 5300 described species. The rare genus Qinorapala Chou & Wang, with Q.qinlingana Chou & Wang as its type species was established as monotypic. In the original description, Q.qinlingana was described from a male holotype; the female remained unknown. To date, the genus is only recorded from the Qinling Mountains (Shaanxi and Gansu Provinces). In this study, two female specimens, from Shaanxi Province and western Sichuan Province (bordering Yunnan Province) are described and illustrated for the first time. New information: Female specimens of Q.qinlingana from Shaanxi and Sichuan are described for the first time. The species' distribution is updated and a distribution map is provided.

Revision of the Genus Rhagastis Rothschild & Jordan, 1903 (Lepidoptera: Sphingidae) from China, Based on Morphological and Phylogenetic Analyses.

Here, the taxonomy of the genus Rhagastis Rothschild & Jordan, 1903 (Lepidoptera, Sphingidae, Macroglossinae, Macroglossini) from China is revised based on differences in wing morphology, male and female genitalia, and the phylogenetic relationship of the DNA barcodes. Subspecies of Rhagastis albomarginatus (Rothschild, 1894) and R. castor (Walker, 1856) are treated as "good" species, namely Rhagastis dichroae Mell, 1922 stat. nov.; R. everetti Rothschild & Jordan, 1903 stat. nov.; R. aurifera (Butler, 1875) stat. rev.; R. chinensis Mell, 1922 stat. nov.; R. formosana Clark, 1925 stat. nov.; and R. jordani Oberthür, 1904 stat. rev. The distribution maps, biological notes, and ecological records of the genus Rhagastis Rothschild & Jordan, 1903 from China are given, and a species inventory of genus Rhagastis in the world is also included.

Complete mitochondrial genome of the little-known regional endemic Aporia hastata (Oberthür, 1892) (Lepidoptera: Pieridae).

The present study reports the mitochondrial genome of A. hastata (Oberthür, 1892), a little-known Aporia species endemic to the southern margin of the Hengduan Mountains in Yunnan Province. This genome is circular, 15,148 bp in length, and consists of 13 PCGs, 22 tRNAs, and two rRNAs. The Bayesian phylogenetic tree clusters A. hastata with other Aporia taxa inside tribe Pierini Duponchel, [1835]. The findings of this study add valuable new information to the genus Aporia and are beneficial to a better understanding of phylogeography of these butterflies.

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.

The life history of Hayesiana triopus (Westwood, 1847), with taxonomic notes on both present and former species of the genus Hayesiana Fletcher, 1982 (Lepidoptera: Sphingidae).

The two species previously included in the genus Hayesiana Fletcher, 1982 were studied. The life history of the sole currently included species, Hayesiana triopus (Westwood, 1847) is illustrated in colour for the first time. Field records of Hayesiana triopus and Dahira farintaenia (Zhu & Wang, 1997) (previously Hayesiana farintaenia) are given, with the first description of the female genitalia of the latter. The diagnostic features and DNA barcoding data of Hayesiana triopus and Dahira farintaenia are also discussed.

Review of the Narrow-Banded Hawkmoth, Neogurelca montana (Rothschild & Jordan, 1915) (Lepidoptera: Sphingidae) in China, with Morphological and Phylogenetic Analysis.

Neogurelca montana (Rothschild & Jordan, 1915) is a species of the genus Neogurelca Hogenes & Treadaway, 1993, that was previously known from Sichuan, Yunnan, and Tibet, China. Recently, however, this species was also found in Beijing and Hebei. These populations differ from those in southwest China in body colour and the shape of the yellow patches of the hindwing-a paler body colour and triangular patches in the former and darker body colour and fan-like patches in the latter. Wing morphology, male and female genitalia, and molecular evidence (DNA barcodes) were analysed for the different localities of this species and three other Neogurelca species-N. hyas, N. himachala, and N. masuriensis. Our molecular data support the Beijing population of montana as a valid subspecies, which we describe as N. montana taihangensisssp. nov. Wing and genital morphology confirm the molecular conclusions. We also collected larvae of the new subspecies in the Beijing suburbs and describe its life history and larval hosts and compare them with those of N. himachala.

Cut to Disarm Plant Defence: A Unique Oviposition Behaviour in Rhynchites foveipennis (Coleoptera: Attelabidae).

Female weevils of the family Attelabidae (Coleoptera: Curculionoidea) possess a unique behaviour of partially cutting the branches connecting egg-bearing organs of their host plants during oviposition. However, the consequence of such behaviour remains unclear. Using Rhynchites foveipennis and its host pear (Pyrus pyrifolia), the present study tested the hypothesis that the oviposition behaviour could disarm the host plants' defence. We compared the survival rates, growth rates, and performance of eggs and larvae under two conditions: (1) the fruit stems were naturally damaged by the females before and after oviposition, and (2) the fruit stems were artificially protected from the females. When fruit stems were protected from female damage, the survival rates of eggs and larvae were only 21.3-32.6%, respectively; and the larval weight was 3.2-4.1 mg 30 days after laying eggs. When the fruit stems were damaged, the survival rates of eggs and larvae reached 86.1-94.0%, respectively; and the larval weight reached 73.0-74.9 mg 30 days after laying eggs. The contents of tannin and flavonoids in the pears did not change significantly along with the oviposition and larval feeding, but weevil eggs were crushed and killed by the callus in the pears. Once the stunted larvae in branch-growing pears were moved into the picked-off ones, the growth and development recovered. The findings indicate that the oviposition behaviour can significantly increase the survival of the offspring. Our study suggested that the oviposition behaviour of attelabid weevils is a strategy to overcome plant defence.

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.

Diversity of planthoppers associated with the winter rice agroecosystems in southern Yunnan, China.

A field survey of the overwintering planthoppers (Hemiptera: Delphacidae) associated with the rice agroecosystems in southern Yunnan was carried out during January-February in 2010 and 2011. 22 species of planthoppers were collected and identified, with one species representing the subfamily Stenocraninae and the other 21 species in Delphacinae. Nycheuma cognatum (Muir), Peregrinus maidis (Ashmead), and Pseudosogata vatrenus (Fennah) were new provincial records for Yunnan. The pest species, Sogatella furcifera (Horváth), Nilaparvata lugens (Stål), and Laodelphax striatellus (Fallén) were able to overwinter in part of the survey range. 13 species were listed to be of economic importance. Abandoned rice paddies with dense Poaceae grasses (Poaceae) were the most favorable overwintering habitat. The survey range was divided into four regions and five areas based on natural geographical characteristics. The study demonstrated that winter temperature differentiation, terrains, and habitat differences were three factors affecting planthopper diversity. Planthopper species diversity showed a reductive trend from south to north and reflected a gradient of more severe winter temperatures. In addition, planthopper diversity was influenced by smaller-scale differences in terrain and habitat, as evidenced by greater diversity in the valleys and low-altitude areas as compared to mid-mountain and Karst plain areas.

Complete mitochondrial genome of the recently discovered multivoltine Graphium (Pazala) confucius Hu, Duan & Cotton, 2018 (Lepidoptera: Papilionidae).

Graphium (Pazala) confucius Hu, Duan & Cotton, 2018 is a recently discovered, wide ranging, multivoltine swordtail butterfly in China and Vietnam. The present study reports the complete mitochondrial genome of this butterfly, which is the fifth mitochondrial genome record for subgenus Pazala Moore, 1888. The mitochondrial genome of G. (P.) confucius is circular and 15,212 bp in length, and consists of 37 genes, including 13 PCGs, 22 tRNAs, and two rRNAs. The Bayesian phylogenetic tree containing the focal species and 33 other Papilioninae members clusters G. (P.) confucius with other Pazala taxa inside tribe Leptocircini, which agrees with its taxonomic position. The findings of this study added data to the complex subgenus Pazala and are beneficial to future understanding and conservation planning of butterfly diversity.

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.

A new species of Rhodambulyx Mell, 1939 (Lepidoptera: Sphingidae) from Southwest Chongqing, China.

A new species of the genus Rhodambulyx Mell, 1939, Rhodambulyx xinyuae sp. nov., is described from Simianshan Nature Reserve in Southwest Chongqing, China. This species is similar to R. davidi Mell, 1939 and R. kitchingi Brechlin, 2015 in habitus, but can be distinguished by a different wing pattern, male genitalia structure and DNA barcode sequence. In addition, Rhodambulyx namvui Eitschberger Nguyen, 2017 is removed from synonymy with R. kitchingi and synonymized instead with R. davidi, although whether it would be better treated as a subspecies of R. davidi requires further investigation.

Complete mitochondrial genomes of two insular races of Pazala swordtails from Taiwan, China (Lepidoptera: Papilionidae: Graphium).

The mitogenomes of two insular subspecies of Pazala, G. (P.) eurous asakurae and G. (P.) mullah chungianus from Taiwan Island, are reported. Both mitogenomes are circular, 15,228 bp and 15,240 bp in length respectively, and consist of 37 genes, including 13 PCGs, 22 tRNAs, and two rRNAs. The Bayesian phylogenetic tree containing the focal taxa and 31 other Papilioninae members clustered them with G. (P.) mullah (Alphéraky, 1897) and then G. (P.) parus (Nicéville, 1900) inside tribe Leptocircini, which agrees with their taxonomic positions. The findings of this study would benefit future understanding of phylogeography and conservation of subgenus Pazala.

A New Species of Aporia/ (Lepidoptera: Pieridae) from Northwest Yunnan, China with Taxonomic Notes on Its Similar Sympatric Taxa.

The northwest corner of Yunnan Province, China is the world-class diversity centre of the genus Aporia Hübner, 1819 (Lepidoptera: Pieridae). During our expeditions to this area in 2019 and 2020, a new species of Aporia was discovered from the upper Yangtze River valley in northwest Yunnan, China. The new taxon, A. chunhaoi sp. nov., is similar to A. lhamo (Oberthür, 1893) but can be easily identified by larger size, much paler (creamy) male hindwing, as well as different genitalic structures. Our field surveys and comparative studies also confirmed a new range of A. lhamo in the upper Yangtze River valley, and the morphological variation of this species is discussed herein.