Molecular phylogeny of Hesperiidae (Lepidoptera) with an emphasis on Asian and African genera.

Despite considerable research efforts in recent years, the deeper phylogenetic relationships among skipper butterflies (Hesperiidae) remain unresolved. This is primarily because of limited sampling, especially within Asian and African lineages. In this study, we consolidated previous data and extensively sampled Asian and African taxa to elucidate the phylogenetic relationships within Hesperiidae. The molecular dataset comprised sequences from two mitochondrial and two nuclear gene regions from 563 species that represented 353 genera. Our analyses revealed seven subfamilies within Hesperiidae: Coeliadinae, Euschemoninae, Eudaminae, Pyrginae, Heteropterinae, Trapezitinae, and Hesperiinae. The systematics of most tribes and genera aligned with those of prior studies. However, notable differences were observed in several tribes and genera. Overall, the position of taxa assigned to incertae sedis in Hesperiinae is largely clarified in this study. Our results strongly support the monophyly of the tribe Tagiadini (Pyrginae), and the systematics of some genera are clarified with comprehensive discussion. We recognize 15 tribes within the subfamily Hesperiinae. Of these, nine tribes are discussed in detail: Aeromachini, Astictopterini, Erionotini, Unkanini (new status), Ancistroidini, Ismini (confirmed status), Plastingini (new status), Gretnini (confirmed status), and Eetionini (confirmed status). We propose four subtribes within Astictopterini: Hypoleucina subtrib.n., Aclerosina, Cupithina, and Astictopterina. Furthermore, we describe a new genus (Hyarotoidesgen.n.) and reinstate two genera (Zeareinst.stat. and Separeinst.stat.) as valid. Additionally, we propose several new combinations: Zea mythecacomb.n.,Sepa bononiacomb.n. & reinst.stat., and Sepa umbrosacomb.n. This study, with extensive sampling of Asian and African taxa, greatly enhances the understanding of the knowledge of the skipper tree of life.

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 favorable-warmer and wetter. Our results suggest changing Neogene climate, uplift of eastern African orogens, and biotic interactions have had different effects on the various subclades of Bicyclus, producing one of the most spectacular butterfly radiations in Africa. [Afrotropics; biodiversity; biome; biotic interactions; Court Jester; extinction; grasslands; paleoclimates; Red Queen; refugia forests; dependent-diversification; speciation.].

The Only Blue Mimeresia (Lepidoptera: Lycaenidae: Lipteninae) Uses a Color-Generating Mechanism Widely Applied by Butterflies.

The butterfly Mimeresia neavei (Joicey & Talbot, 1921) is the only species in the exclusively African subtribal clade Mimacraeina (Lipteninae: Lycaenidae: Lepidoptera) having sexual dimorphism expressed by structurally blue-colored male and pigmentary colored orange-red female phenotypes. We investigated the optical mechanism generating the male blue color by various microscopic and experimental methods. It was found that the blue color is produced by the lower lamina of the scale acting as a thin film. This kind of color production is not rare in day-flying Lepidoptera, or in other insect orders. The biological role of the blue color of M. neavei is not yet well understood, as all the other species in the clade lack structural coloration, and have less pronounced sexual dimorphism, and are involved in mimicry-rings.

Magnifying the hotspot: descriptions of nine new species of many-plumed moths (Lepidoptera, Alucitidae), with an identification key to all species known from Cameroon.

This study confirms Mount Cameroon as an unprecedented hotspot for the diversity of many-plumed moths, with the discovery and description of nine new species: Alucitafako Ustjuzhanin & Kovtunovich, sp. nov., Alucitapyrczi Ustjuzhanin & Kovtunovich, sp. nov., Alucitasroczki Ustjuzhanin & Kovtunovich, sp. nov., Alucitapotockyi Ustjuzhanin & Kovtunovich, sp. nov., Alucitasedlaceki Ustjuzhanin & Kovtunovich, sp. nov., Alucitatonda Ustjuzhanin & Kovtunovich, sp. nov., Alucitaerzayi Ustjuzhanin & Kovtunovich, sp. nov., Alucitasokolovi Ustjuzhanin & Kovtunovich, sp. nov., and Alucitahirsuta Ustjuzhanin & Kovtunovich, sp. nov. Additionally, four additional species are reported from the Mount Cameroon area as new for the country: Alucitaagassizi, Alucitadohertyi, Alucitaplumigera, and Alucitarhaptica. Of the 89 Alucitidae known from the Afrotropics, the studied area hosts 36 species, most of which are endemic to the area. This unprecedented level of diversity and endemism within this lepidopteran family highlights Mount Cameroon's significance as a stronghold for specialised insect taxa. Efficient conservation efforts are necessary to protect these ecosystems and their associated unique microlepidopteran diversity.

Comprehensive phylogeny of Pieridae butterflies reveals strong correlation between diversification and temperature.

Temperature is thought to be a key factor influencing global species richness patterns. We investigate the link between temperature and diversification in the butterfly family Pieridae by combining next generation DNA sequences and published molecular data with fine-grained distribution data. We sampled nearly 600 pierid butterfly species to infer the most comprehensive molecular phylogeny of the family and curated a distribution dataset of more than 800,000 occurrences. We found strong evidence that species in environments with more stable daily temperatures or cooler maximum temperatures in the warm seasons have higher speciation rates. Furthermore, speciation and extinction rates decreased in tandem with global temperatures through geological time, resulting in a constant net diversification.

Two new Aphnaeus (Lepidoptera: Lycaenidae: Theclinae) species from Liberia.

Two new butterfly species belonging to the genus Aphnaeus (Lycaenidae: Theclinae), A. mirabilis sp. n., A. nimbaensis sp. n. were found in the Nimba Mountains, Western Range, Liberia in February 2012. Both are illustrated and described in comparison with closely related species: A. flavescens Stempffer, 1954 and A. liberti Bouyer, 1996, respectively. Genitalia of the male holotype of A. nimbaensis are also illustrated.

A new species in the genus Precis Hübner, [1819] (Lepidoptera: Nymphalidae: Nymphalinae), another surprising discovery in the Nimba Mountains, Guinea and the broader Guinea Highlands in West Africa.

A new species in the genus Precis (Papilionoidea, Nymphalidae, Nymphalinae), described in this paper as P. koivoguii sp. n., was discovered in the Nimba Mountains, Guinea, West Africa, and specimens were also identified from the neighbouring mountainous area in Ivory Coast in the broader Guinea Highlands. In appearance the species is similar to some wet season forms of the widely distributed P. pelarga and the Madagascan P. andremiaja, but in size and wingshape, it also resembles P. ceryne, and could have been a natural hybrid between this latter species and P. pelarga. The specific distinction from these taxa was confirmed by marked differences in male genitalia and constant phenotypic characteristics. Based on the type series, it was thought that the new species might be endemic to mid- and high-altitude habitats, particularly forest-scrub-grassland edges or wetlands in the broader Guinea Highlands but a new photographic record from western Ghana suggests a much wider distribution in suitable habitats of the Upper Guinean forest zone.

Measuring and Modelling Structural Colours of Euphaedra neophron (Lepidoptera: Nymphalidae) Finely Tuned by Wing Scale Lower Lamina in Various Subspecies.

The nymphalid butterfly Euphaedra neophron (Hopffer, 1855) is the only structurally coloured species representing the genus along the Indian Ocean coast in East Africa and Southern Africa, with a distribution from southern Somalia to the Kwa-Zulu-Natal region of South Africa. The range of E. neophron is subdivided to several, geographically distinct populations, currently recognised as subspecies by taxonomists on the basis of violet, blue, and green-coloured morphs. We investigated the optical mechanism of all these morphs by various materials science techniques. We found that the structural colour is generated by the lower lamina of the cover scales and the different colours are tuned according to their thickness, which was also proved by modelling. The colour tuning of the different subspecies does not reflect any clinal pattern, be it geographical or altitudinal.

Rapid radiation of ant parasitic butterflies during the Miocene aridification of Africa.

Africa has undergone a progressive aridification during the last 20 My that presumably impacted organisms and fostered the evolution of life history adaptations. We test the hypothesis that shift to living in ant nests and feeding on ant brood by larvae of phyto-predaceous Lepidochrysops butterflies was an adaptive response to the aridification of Africa that facilitated the subsequent radiation of butterflies in this genus. Using anchored hybrid enrichment we constructed a time-calibrated phylogeny for Lepidochrysops and its closest, non-parasitic relatives in the Euchrysops section (Poloyommatini). We estimated ancestral areas across the phylogeny with process-based biogeographical models and diversification rates relying on time-variable and clade-heterogeneous birth-death models. The Euchrysops section originated with the emerging Miombo woodlands about 22 million years ago (Mya) and spread to drier biomes as they became available in the late Miocene. The diversification of the non-parasitic lineages decreased as aridification intensified around 10 Mya, culminating in diversity decline. In contrast, the diversification of the phyto-predaceous Lepidochrysops lineage proceeded rapidly from about 6.5 Mya when this unusual life history likely first evolved. The Miombo woodlands were the cradle for diversification of the Euchrysops section, and our findings are consistent with the hypothesis that aridification during the Miocene selected for a phyto-predaceous life history in species of Lepidochrysops, with ant nests likely providing caterpillars a safe refuge from fire and a source of food when vegetation was scarce.

Revisional notes on Iolaus aequatorialis Stempffer Bennett, 1958 and related species in the subgenus Philiolaus Stempffer Bennett, 1958 (Lepidoptera, Lycaenidae, Theclinae), with description of two new species.

Similarities in male and female genitalic structure isolate a small group of taxa in the genus Iolaus Hübner, [1819], subgenus Philiolaus Stempffer & Bennett, 1958, related to Iolaus aequatorialis Stempffer & Bennett, 1958. They are assigned to the newly proposed I. aequatorialis species group with two new species recognised and described. This action also clarifies previous taxonomic misunderstandings involving a few other species with similar facies because of a lack of information on male and female genital morphology.

Description of five new species of Epitola sensu lato (Lepidoptera: Lycaenidae: Poritiinae) from West and Central Africa.

Five new species of Epitola s.l. (Lycaenidae: Poritiinae) are described in the genera Stempfferia Jackson, 1962 and Cephetola Libert, 1999. Stempfferia salonga sp. n. was found in the 'Cuvette Centrale' of the Democratic Republic of Congo and a single specimen was identified also from southern Cameroon. The similar S. buea sp. n. is known only from the unique holotype collected at the foothills of Mount Cameroon, while S. michelliberti sp. n. is distributed in lowland rainforest between southeast Ivory Coast and Guinea. The new species of Cephetola were found only in Liberia. Cephetola praecox sp. n. is known from a single upland forest locality in the Putu Range, whereas C. wologizi sp. n. was collected also in upland forest in the Wologizi Mountains.

Alpha-taxonomy and phylogeny of African Junoniini butterflies based on morphological data, with an emphasis on genitalia, and COI barcode (Lepidoptera: Nymphalidae).

The tribe Junoniini is a predominantly Paleotropical group of the cosmopolitan butterfly subfamily Nymphalinae (Nymphalidae), with highest diversity in the Afrotropical region. Its systematics and relationships are not entirely resolved. Question marks remain concerning the validity of some genera; and the apparently close relationship between the Indo-Australian genus Yoma and the Afrotropical Protogoniomorpha, as evidenced by molecular phylogenies, remains a puzzle. Here, we present a cladistic analysis, based on 42 characters of the male and female genitalia of 41 species of Junoniini belonging to six genera, nearly all of them continental Afrotropical, and 3 species of two Indo-Australian genera Yoma and Rhinopalpa. A ML COI-based tree is produced for 36 species of Afrotropical Junoniini and Yoma. The molecular data are consistent with previous studies. However, morphological analysis does not confirm a close relationship between Protogoniomorpha and Yoma. Despite the evolution of a number of modifications, the male genitalia within all genera and species of the Junoniini share a cohesive build plan, in particular a transformed sacculus, from which Yoma is highly divergent. The position of the genus Kamilla, previously synonymized with Junonia, is discussed. Three East African coast taxa, Junonia elgiva stat. reinst., Protogoniomorpha nebulosa stat. reinst. and Salamis amaniensis stat. reinst., and one from central Africa, Precis silvicola stat. reinst. are raised to species level, based on comparative analysis of their male genitalia.

Elevational and seasonal patterns of butterflies and hawkmoths in plant-pollinator networks in tropical rainforests of Mount Cameroon.

Butterflies and moths are conspicuous flower visitors but their role in plant-pollinator interactions has rarely been quantified, especially in tropical rainforests. Moreover, we have virtually no knowledge of environmental factors affecting the role of lepidopterans in pollination networks. We videorecorded flower-visiting butterflies and hawkmoths on 212 plant species (> 26,000 recorded hrs) along the complete elevational gradient of rainforests on Mount Cameroon in dry and wet seasons. Altogether, we recorded 734 flower visits by 80 butterfly and 27 hawkmoth species, representing only ~ 4% of all flower visits. Although lepidopterans visited flowers of only a third of the plant species, they appeared to be key visitors for several plants. Lepidopterans visited flowers most frequently at mid-elevations and dry season, mirroring their local elevational patterns of diversity. Characteristics of interaction networks showed no apparent elevational or seasonal patterns, probably because of the high specialisation of all networks. Significant non-linear changes of proboscis and forewing lengths were found along elevation. A positive relationship between the lengths of proboscis of hesperiid butterflies and tube of visited flowers was detected. Differences in floral preferences were found between sphingids and butterflies, revealing the importance of nectar production, floral size and shape for sphingids, and floral colour for butterflies. The revealed trait-matching and floral preferences confirmed their potential to drive floral evolution in tropical ecosystems.

Description of the previously unknown sexes in four West African Lycaenidae (Lepidoptera) with new data on their distribution.

Several new butterfly species in the family Lycaenidae were found during recent field surveys in Liberia. Amongst these, quite a few were collected as singletons or in small series, represented by only one of the sexes. This paper describes and illustrates the previously unknown sexes of four of those species which were subsequently found, adding new locality records to their known distribution.

Even hotter hotspot: description of seven new species of many-plumed moths (Lepidoptera, Alucitidae) from Mount Cameroon.

Mount Cameroon, SW Cameroon, has already been described as a unique hotspot of the many-plumed moth (Lepidoptera, Alucitidae), with their local diversity unrivalled in the entire Afrotropics. We confirm its importance with description of seven new species: Alucita bakweri Ustjuzhanin & Kovtunovich, sp. nov., Alucita jana Ustjuzhanin & Kovtunovich, sp. nov., Alucita bakingili Ustjuzhanin & Kovtunovich, sp. nov., Alucita tatjana Ustjuzhanin & Kovtunovich, sp. nov., Alucita zuza Ustjuzhanin & Kovtunovich, sp. nov., Alucita deja Ustjuzhanin & Kovtunovich, sp. nov., and Alucita bokwango Ustjuzhanin & Kovtunovich, sp. nov. These descriptions have raised the known local diversity of many-plumed moth species on Mount Cameroon to 22, i.e., over a quarter of the known Afrotropical biodiversity of this group. This study also emphasises the great conservation importance of the area.

First records of 31 species of butterflies and moths (Lepidoptera) in Cameroon, with remarks on their elevational ranges.

BACKGROUND: The biodiversity of West and Central Africa is understudied, including butterflies and moths (Lepidoptera). Cameroon, through its position in between few biogeographic regions and diversity of habitats, is an important hotspot of lepidopteran diversity. However, the country also ranks low when it comes to local biodiversity knowledge. During our long-term ecological projects in the Cameroonian part of the Gulf of Guinea Highlands, we collected rich material of butterflies and moths, including a number of interesting faunistic records. NEW INFORMATION: In this study, we report 31 species of butterflies and moths which have not yet been recorded in Cameroon. These species comprised eight new genera records for the country. In many cases, our records represented an important extension of the species' known distribution, including ten species whose distribution ranges extended into the Guinean biogeographic region. We also comment on the species' elevational distribution ranges on Mount Cameroon where most of our records originated. Additionally, we confirm the presence of a butterfly Telchinia encedena, after more than a century since its first and so far its only record in Cameroon.

Further information on Lolibaia Gorbunov Gurko, 2017 (Lepidoptera: Sesiidae: Synanthedonini) with description of three new species from tropical Africa.

The knowledge of the clearwing moth genus Lolibaia Gorbunov Gurko, 2017, recently described from South Sudan and until now known only from a single, damaged specimen, is extended by three new species from different parts of Africa including the as yet unknown female of the genus. L. jurateae sp. nov. is based on a single male from Kakamega Forest, Kenya; L. triangularis sp. nov. is based on six males from different localities in north-western Zambia, and L. polythyris sp. nov. is described from two females from the Eastern Cape and KwaZulu-Natal Provinces, South Africa.

Two new species in the genus Andronymus Holland, 1896 (Lepidoptera, Hesperiidae).

During extensive field work in West Africa (Cameroon, Ivory Coast, Liberia, Sierra Leone), the authors collected two skipper species in the genus Andronymus (Lepidoptera, Hesperiidae), which would not fit the descriptions of any existing taxa. Both are described as new, A. magma sp. nov. is known only from Cameroon, while A. fenestra sp. nov. was found in a few localities in the Liberian sub-region of West Africa.

Amerila (Lepidoptera: Erebidae: Arctiinae) of Cameroon with morphological remarks on male and female genitalia.

Amerila is one of the most studied Afrotropical genera of Arctiinae. However, based on a regionally constrained sample of specimens from Mount Cameroon, we show how superficial our knowledge on these tiger moths is. Among six collected Amerila species, A. femina's female is described here for the first time, and A. mulleri and A. roseomarginata had never been recorded before in the country. Moreover, novel biological data are presented, including individual species' elevational ranges. Finally, female reproductive organs of the genus are illustrated here for the first time. The value of such regional studies is highlighted, with some remarks on necessary requirements of such small-scaled field sampling.

A new species of Conopsia Strand, 1913 (Lepidoptera: Sesiidae: Tinthiinae) from West Africa.

A new species of Sesiidae, discovered during a Lepidoptera survey in the Nimba Mountains, Guinea (West Africa) is described as Conopsia puehringeri sp. nov. The new species is compared to its potential sister species C. bicolor (Le Cerf, 1917), in sense of external morphology and genitalia structure. Male specimens and genitalia of both studied species are depicted.