Systematics and historical biogeography of the old world butterfly subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrinae).

BACKGROUND: Butterflies of the subtribe Mycalesina have radiated successfully in almost all habitat types in Africa, Madagascar, the Indian subcontinent, Indo-China and Australasia. Studies aimed at understanding the reasons behind the evolutionary success of this spectacular Old World butterfly radiation have been hampered by the lack of a stable phylogeny for the group. Here, we have reconstructed a robust phylogenetic framework for the subtribe using 10 genes from 195 exemplar taxa. RESULTS: We recovered seven well supported clades within the subtribe corresponding to the five traditional genera (Lohora, Heteropsis, Hallelesis, Bicyclus, Mycalesis), one as recently revised (Mydosama) and one newly revised genus (Culapa). The phylogenetic relationships of these mycalesine genera have been robustly established for the first time. Within the proposed phylogenetic framework, we estimated the crown age of the subtribe to be 40 Million years ago (Mya) and inferred its ultimate origin to be in Asia. Our results reveal both vicariance and dispersal as factors responsible for the current widespread distribution of the group in the Old World tropics. We inferred that the African continent has been colonized at least twice by Asian mycalesines within the last 26 and 23 Mya. In one possible scenario, an Asian ancestor gave rise to Heteropsis on continental Africa, which later dispersed into Madagascar and most likely back colonised Asia. The second colonization of Africa by Asian ancestors resulted in Hallelesis and Bicyclus on continental Africa, the descendants of which did not colonise other regions but rather diversified only in continental Africa. The genera Lohora and Mydosama are derivatives of ancestors from continental Asia. CONCLUSION: Our proposed time-calibrated phylogeny now provides a solid framework within which we can implement mechanistic studies aimed at unravelling the ecological and evolutionary processes that culminated in the spectacular radiation of mycalesines in the Old World tropics

Revision of the Bicyclus ignobilis species-group (Lepidoptera: Nymphalidae: Satyrinae) with descriptions of two new species

The ignobilis-group of the genus Bicyclus Kirby 1871 is revised. The species-group contains six species with a distinct wing pattern, but limited intraspecific variation, distributed across tropical African rainforest. We investigate a set of more than 1000 specimens from a range of museum collections, including some type material, and thoroughly update the biogeographical knowledge for the group. We also describe two new species as members of the group. The included species are: Bicyclus ignobilis (Butler 1870) stat. rev., B. rileyi Condamin 1961, B. maesseni Condamin 1971, B. brakefieldi Brattström 2012, B. ottossoni sp. nov. and B. vandeweghei sp. nov. Due to observing a gradual morphological cline within B. ignobilis without any sharp transitions we suppress the previously identified subspecies B. ignobilis eurini Condamin & Fox 1963 syn. nov. and B. ignobilis acutus Condamin 1965 syn. nov

Revision of the Bicyclus sciathis species group (Lepidoptera: Nymphalidae) with descriptions of four new species and corrected distributional records

In this paper we present a thorough revision of the sciathis species group of the butterfly genus Bicyclus (Kirby). Type materials are discussed and in several cases lectotypes are assigned to specimens from original type series. Four new, and morphologically distinct, species are described (B. elishiae Brattström sp.n., B. heathi Brattström sp.n., B. sigiussidorum Brattström sp.n. and B. subtilisurae Brattström sp.n.), along with a comprehensive molecular phylogeny that includes exemplar taxa of all currently recognized species. We also investigate the types of all previously synonymized taxa and in the process invalidate the name B. ewondo Libert. This was done after finding the previously missing holotype of B. makomensis (Strand), which clearly belongs to the same species and thereby gives the older name priority. The phylogeny showed that some distinctly different species were surprisingly closely related, suggesting a high rate of morphological evolution in parts of the sciathis group. The distributional records for the group are updated after investigating over 1700 specimens kept in a range of museum collections. Many species previously thought to be broadly sympatric were found to have much more restricted ranges, with the previous overestimations probably based on misidentified specimens. The higher level of allopatry now established will make identification of many morphologically similar species easier. The fact that species often have smaller ranges than previously known, meaning that the level of endemism for African butterflies is likely to be higher than current estimates, has important implications for conservation management. An identification key for males of all 13 currently recognized species in the species group is included

Geographic origin and migration phenology of European red admirals (Vanessa atalanta) as revealed by stable isotopes.

BACKGROUND: Long-distance migration has evolved multiple times in different animal taxa. For insect migrants, the complete annual migration cycle covering several thousand kilometres, may be performed by several generations, each migrating part of the distance and reproducing. Different life-cycle stages and preferred orientation may thus, be found along the migration route. For migrating red admirals (Vanessa atalanta) it has been questioned if they reproduce in the most northern part of the range. Here we present migration phenology data from a two-year time series of migrating red admirals captured at Rybachy, Kaliningrad, in the northern part of Europe investigating time for migration, life-history stage (migration, reproduction) as well as site of origin in individual butterflies. METHODS: Red admirals were captured daily at a coastal site during spring, summer and autumn in 2004 and 2005. For the sampled individuals, reproductive status and fuel content were estimated by visual inspection, and hydrogen isotopes (δ 2H) were analysed in wing samples. δ 2H values was compared with samples from two nearby reference sites in Estonia and Poland. RESULTS: Analysis of hydrogen isotopes (δ 2H) in red admiral wings showed that the spring cohort were of a southerly origin, while those caught in August or later in the autumn were from the local region or areas further to the north. All females caught during spring had developing eggs in their abdomen, but no eggs were found in late summer/autumn. There was a male-biased sex ratio during autumn and a difference in lipid content between years. When comparing the isotopic data with inland nearby locations, it was clear that the range of δ 2H values (- 181 to - 78) was wider at Rybachy as compared to the two reference sites in Estonia and Poland (- 174 to - 100). CONCLUSIONS: During spring, migratory female red admirals arrived from the south and were ready to reproduce, while the autumn passage mainly engaged local and more northern individuals carrying large fuel deposits in preparation for long-distance migration. The phenology data suggest that individuals select to migrate in favourable weather conditions and that numbers may differ between years. Future studies should focus on individual sampling at a wide range of sites to reveal differential migration strategies and timing of migration between sexes and populations of migrating butterflies

Miocene climate and habitat change drove diversification in Bicyclus, Africa’s largest radiation of satyrine butterflies

Compared to other regions, the drivers of diversification in Africa are poorly understood. We studied a radiation of insects with over 100 species occurring in a wide range of habitats across the Afrotropics to investigate the fundamental evolutionary processes and geological events that generate and maintain patterns of species richness on the continent. By investigating the evolutionary history of Bicyclus butterflies within a phylogenetic framework, we inferred the group’s origin at the Oligo-Miocene boundary from ancestors in the Congolian rainforests of central Africa. Abrupt climatic fluctuations during the Miocene (ca. 19-17 Ma) likely fragmented ancestral populations, resulting in at least eight early-divergent lineages. Only one of these lineages appears to have diversified during the drastic climate and biome changes of the early Miocene, radiating into the largest group of extant species. The other seven lineages diversified in forest ecosystems during the late Miocene and Pleistocene when climatic conditions were more favourable–warmer and wetter. Our results suggest changing Neogene climate, uplift of eastern African orogens, and biotic interactions might have had different effects on the various subclades of Bicyclus, producing one of the most spectacular butterfly radiations in Africa

Whole-chromosome hitchhiking driven by a male-killing endosymbiont

Data Availability Statement: Raw genomic data and assemblies are available from GenBank (project accession numbers PRJNA448181 and PRJEB35880, and individual sample accessions are provided in S9 Table). All processed data files underlying all figures are available from the Dryad digital repository: https://doi.org/10.5061/dryad. 9kd51c5d0. Scripts used for data analysis are available from https://github.com/simonhmartin/ genomics_general.Neo-sex chromosomes are found in many taxa, but the forces driving their emergence and spread are poorly understood. The female-specific neo-W chromosome of the African monarch (or queen) butterfly Danaus chrysippus presents an intriguing case study because it is restricted to a single ‘contact zone’ population, involves a putative colour patterning supergene, and co-occurs with infection by the the male-killing endosymbiont Sprioplasma. We investigated the origin and evolution of this system using whole genome sequencing. We first identify the ‘BC supergene’, a large region of suppressed recombination that links two colour patterning loci. Association analysis suggests that the genes yellow and arrow control the forewing colour pattern differences between D. chrysippus subspecies. We then show that the same chromosome has recently formed a neo-W that has spread through the contact zone within ~2200 years. We also assembled the genome of the male-killing Spiroplasma, and find that it shows perfect genealogical congruence with the neo-W, suggesting that the neo-W has hitchhiked to high frequency as the male killer has spread through the population. The complete absence of female crossing-over in the Lepidoptera causes whole-chromosome hitchhiking of a single neo-W haplotype, carrying a single allele of the BC supergene, and dragging multiple non-synonymous mutations to high frequency. This has created a population of infected females that all carry the same recessive colour patterning allele, making the phenotypes of each successive generation highly dependent on uninfected male immigrants. Our findings show how hitchhiking can occur between the unlinked genomes of host and endosymbiont, with dramatic consequences.Biotechnology & Biological Sciences Research Council (BBSRC