The Butterflies and Skippers (Lepidoptera: Papilionoidea) of Angola: An Updated Checklist

Open Access


Presently, 792 species/subspecies of butterflies and skippers (Lepidoptera: Papilionoidea) are known from Angola, a country with a rich diversity of habitats, but where extensive areas remain unsurveyed and where systematic collecting programmes have not been undertaken. Only three species were known from Angola in 1820. From the beginning of the twenty-first century, many new species have been described and more than 220 faunistic novelties have been assigned. As a whole, of the 792 taxa now listed for Angola, 57 species/subspecies are endemic and almost the same number are known to be near-endemics, shared by Angola and by one or another neighbouring country. The Nymphalidae are the most diverse family. The Lycaenidae and Papilionidae have the highest levels of endemism. A revised checklist with taxonomic and ecological notes is presented and the development of knowledge of the superfamily over time in Angola is analysed.


Africa Conservation Ecology Endemism Taxonomy 


Angola is a large country of 1,246,700 km2, notable for its great diversity of physiography, climates, habitats and resultant biodiversity). The country includes seven biomes and 15 ecoregions, ranging from equatorial rainforests of the northwest (Cabinda) and along the northern border with the Democratic Republic of Congo, through the moist miombo woodlands and savannas of the central plateaus, to the dry forests and woodlands of the southeast, and to the arid shrublands and Namib Desert of the southwest. Isolated forests with Congolian affinities are found along the Angolan Escarpment, and similar remnant patches of Afromontane forests are found on some of the highest mountains such as Mount Moco and Mount Namba.

Despite the fact that at the beginning of the nineteenth century only a few species of butterflies and skippers (Insecta: Lepidoptera: Papilionoidea) were recorded from Angola, today a large number of taxa (at least 792 species and subspecies: Fig. 10.1, Table 10.1 and Appendix) are known to occur in the country. However, extensive areas are still poorly surveyed for butterflies, or have not been surveyed at all (Fig. 10.2). This applies in particular to the southern provinces of Namibe, Cunene and Cuando Cubango and the northwestern province of Zaire as well as most of southern Moxico. Furthermore, the Baixa de Cassanje (Malanje), separated from surrounding areas by steep escarpments, appears to have distinctive vegetation and may produce some interesting butterflies. Although most of the localities where butterflies and skippers have been collected in Angola have been determined (Mendes et al. 2013b), some localities previously reported for a few species remain untraced despite searches by us, using the detailed maps of the Junta de Investigações do Ultramar (JIU 1948–1963).
Fig. 10.1

Cumulative number of species/subspecies of Papilionoidea reported from Angola from 1801–1819 (first records) to the recent decade – 2011–2017 – according to Appendix. For practical reasons, species which first reference to the country was untraceable (marked in Appendix with a ▲) were included in the decade 2001–2010; species that are now assigned as faunistic novelties to Angola (marked in Appendix with a ◘) are included in the last decade (2011–2017)

Table 10.1

Number of species of Papilionoidea families and subfamilies known to occur in the Afrotropical Region and Angola (with % of Afrotropical species present in the country), and number of species endemic to Angola (with % of endemism shown)






N° | %

N | %



134 | 22

5 | 3.7



7 | 33




48 | 22




5 | 19




74 | 22




33 | 33

3 | 9.1



67 | 34

5 | 7.5



2 | 40




8 | 57




57 | 32




210 | 11

18 | 8.6



11 | 9




53 | 8




42 | 14




21 | 8




83 | 17




4 | 27

0 | 0



344 | 21

26 | 7.6



2 | 40




9 | 35




50 | 14




56 | 30




1 | 33




35 | 48




1 | 100




16 | 52




97 | 14




77 | 30




792 | 18

Fig. 10.2

Map of Angola showing, marked in orange, the known areas surveyed for the Papilionoidea from the beginning of their study, in the nineteenth century to the present day – each square ca. 33 × 33 km. The collecting pressure varies across the country, from ‘squares’ where samples were obtained only once, in passing, to others where the collectors were based for months

The accumulation of knowledge in regard to Angolan butterflies has been constrained by several factors. The two largest Angolan entomological collections, deposited in the Museu do Dundo (Lunda-Norte) and in the Instituto de Investigação Agronómica (Huambo), have never been studied in detail. In addition little fieldwork was carried out in Angola during the post-independence period because of the protracted civil war. Finally, the vastness of the country and the difficulty in accessing many remote regions has impeded progress.

Until recently the Hesperiidae (skippers) were placed in the superfamily Hesperioidea, separate from the rest of the butterflies, which were placed in the superfamily Papilionoidea. However, today the skippers and butterflies are all placed in the Papilionoidea (e.g. Heikkilä et al. 2012). The classification used for butterflies in this chapter is based on Williams (2018), Espeland et al. (2018) and Dhungel and Wahlberg (2018). Six families of butterflies are represented in Angola, namely Papilionidae, Hesperiidae, Pieridae, Riodinidae, Lycaenidae and Nymphalidae.

History of Research on the Papilionoidea of Angola

The first known reference to butterflies obtained in Angola is by Latreille and Godart (1819), who reported the presence of Colotis euippe (Linnaeus, 1758) and described Acraea parrhasia servona. In the decade between 1871 and 1880, Druce (1875) reported about 90 species from Angola for the first time, a number of these being descriptions of species new to science. By the end of the nineteenth century a total of 214 butterfly taxa were known from Angola.

The first contributions to our knowledge of Angolan butterflies by Portuguese researchers were only made in the middle nineteen hundreds. These were the result of the activities in Angola of the Centro de Zoologia (CZ) of the Junta de Investigações do Ultramar, coordinated by its first director Fernando Frade. In 2014 this research institution was renamed the Instituto de Investigação Científica Tropical (IICT). Working from these large zoological collections, as well as from further specimens obtained in Angola by Amélia Bacelar (1948, 1956, 1958a, b, 1961) and Miguel Ladeiro (1956), considerably expanded the list of Angolan butterflies. Most of this material, obtained during colonial times, was stored in Lisbon, with corrections to the published identifications only being made recently. All of this material has now been integrated into the collections of the Museu Nacional de História Natural e da Ciência (MUHNAC). Albert Monard (1956) of the La Chaux-de-Fonds Swiss Museum also studied other material obtained by the CZ missions. Significant contributions in the twentieth century were also made by Weymer (1901) on the southern Angolan species, and by Evans (1937) on the Hesperiidae. All of the then known Angolan butterflies were listed by Aurivillius (in Seitz) in 1928. All of the Angolan Charaxinae were dealt with by Henning in his 1988 book on the African taxa of this family. The 339 taxa added to the faunal list during the twentieth century brought the total to 553 known butterfly taxa for Angola.

During the first 18 years of the twenty-first century, 239 further taxa were added to the total. In the first decade of the present century, most of the new information was due to several contributions by Libert (1999, 2000, 2004) on the Lycaenidae, and by Gardiner (2004). The latter author listed taxa from the southeastern Cuando Cubango province, which borders the Caprivi Strip of Namibia. Cuando Cubango and the easternmost province of Moxico are the only provinces in Angola with Zambezian fauna. To these taxa we add our own contributions (Bivar-de-Sousa and Mendes 2006, 2007, 2009a, b; Mendes and Bivar-de-Sousa 2006a, b, 2007a, b, 2009a, b, c, d). Over the last 8 years 33 species were described as new or recorded for the first time from Angola by Mendes and Bivar-de-Sousa (2012, 2017) Mendes et al. (2013a, 2017, 2018), Bivar-de-Sousa and Mendes (2014) and Bivar-de-Sousa et al. (2017), Turlin and Vingerhoedt (2013) and Pierre and Bernaud (2013). Finally, 66 further taxa are now recorded as faunistic novelties for the country (Appendix). The current total number of butterfly taxa for Angola now stands at 792.

Sources Consulted for the Checklist

In preparing this revised checklist of the Papilionoidea, the following collections of Angolan butterflies held by institutions in Portugal were examined: Museu Nacional de História Natural e da Ciência (MUHNAC) in Lisbon, Museu de História Natural da Universidade do Porto (MHNC-UP), Liceu Nun’Álvares in the Caldas da Saúde and the Singeverga Order of St Benedict Abbey in Areias. Major contributions to these collections were made by A Bivar-de-Sousa (Luanda district and Cuanza-Norte, Cuanza-Sul and Moxico Provinces), António Figueira, (northwestern Angola), Mário Macedo (northern Angola), Passos de Carvalho (Huambo and Cuanza-Norte Provinces), Carneiro Mendes and Pessoa Guerreiro. The Angolan insect collection of Nozolino de Azevedo (mainly Huambo Province), maintained and made available by his widow, was also studied.

The collections in the MUHNAC in Lisbon were destroyed by a fire in March 1978. However, prior to the fire BS had studied some of the material and published his findings. In 1995 LM studied the collections, mainly of Barros Machado and Luna de Carvalho, in the Dundo Museum in Angola but there was insufficient time to do a detailed analysis. We did not inspect the entomological collections in the former Instituto de Investigação Agronómica de Angola, collected mainly by Passos de Carvalho, but they are apparently in good condition. No entomological collections were found by LM, in 1995 and 2013, at the Museu de História Natural de Luanda. Material collected from 2010 to 2014 by Ruben Capela and Carmen Van-Dúnen Santos of Agostinho Neto University, Luanda and Artur Serrano of the Faculty of Sciences, Lisbon University, was examined by us.

In addition, images of live specimens published by Lautenschläger and Neinhhuis (2014) were examined, as were several images presented by Jorge Palmeirim of Lisbon University and Pedro Vaz Pinto of the Kissama Foundation.

Taxa Excluded from the Checklist

A number of taxa have erroneously been reported to occur in Angola. This was due mainly to probable misidentifications or mislabelled specimens. Some older records are omitted because the known range of the taxon is unlikely to include Angola. A list of the omitted taxa is given below.

  • Hesperiidae: Eretis djaelaelae (Wallengren, 1857), Metisella metis (Linnaeus, 1764), Kedestes chaca (Trimen, 1873), Platylesches chamaeleon (Mabille, 1891).

  • Papilionidae: Papilio menestheus Drury, 1773, Graphium taboranus (Oberthür, 1886), Graphium (Arisbe) junodi (Trimen, 1893).

  • Pieridae: Eurema brigitta (Stoll, 1780), Colotis chrysonome (Klug, 1829), Colotis ephyia (Klug, 1829), Belenois theora (Doubleday, 1846), Mylothris rubricosta (Mabille, 1890), Mylothris similis Lathy, 1906.

  • Lycaenidae: Telipna acraea (Westwood, 1851), Cooksonia abri Collins & Larsen, 2008, Mimacraea darwinia Butler, 1872, Liptena bassae Bethune-Baker, 1926, Aethiopana honorius honorius (Fabricius, 1793), Stempfferia uniformis (Kirby, 1887), Stempfferia dorothea (Bethune-Baker, 1904), Oxylides faunus (Drury, 1773), Dapidodigma hymen (Fabricius, 1775), Aloeides molomo (Trimen, 1870), Leptomyrina lara (Linnaeus, 1764), Deudorix livia (Klug, 1834), Neurellipes onias (Hulstaert, 1924), Zintha hintza (Trimen, 1864).

  • Riodinidae: Afriodinia caeca semicaeca (Riley, 1932), Afriodinia gerontes (Fabricius, 1781).

  • Nymphalidae: Bicyclus milyas (Hewitson, 1864), Ypthima congoana Overlaet, 1955, Charaxes jahlusa argynnides Westwood, 1864, Junonia touhilimasa Vuillot, 1892, Neptis continuata Holland, 1892, Neptis strigata Aurivillius, 1894, Evena oberthueri (Karsch, 1894), Euriphene atrovirens (Mabille, 1878), Bebearia mardania (Fabricius, 1793), Euphaedra morini Hecq, 1983, Euphaedra xypete (Hewitson, 1865), Euphaedra campaspe (Felder & Felder, 1867), Euphaedra inanum (Butler, 1873), Euphaedra eupalus (Fabricius, 1781).

A Revised Checklist of the Papilionoidea of Angola

A revised and annotated checklist of the Papilionoidea of Angola (Appendix) confirms the presence of at least 792 taxa in the country. Their presence is based mainly on verification by the authors of this chapter. Some taxa, recorded by other authors, are accepted because they were, with rare exceptions, reported by more than one author, are based on reliable literature records, or because Angola falls within their putative geographical range. In the Checklist, the first reference to their occurrence in Angola is given, followed by the sources of validation of the record and their preferred habitat(s). Occasionally more than one subspecies of a particular species occurs in the country. This is due to both the size and ecological diversity of Angola. A number of forests, especially gallery-forests, are independent of each other as are the fragmented forests of the Angolan Escarpment. In addition, the southeastern parts of Moxico and the Cuando Cubango provinces are part of the Zambesi Basin; consequently their fauna has affinities with that of eastern Africa.

As far as habitats are concerned the great majority of the Angolan Papilionoidea, as might be expected, occur in forest, both wet and dry (Appendix). However, the Hesperiidae and Pieridae appear to be almost as diverse in moist woodland (miombo) and dry woodland as they are in forest. The number of Pieridae in dry woodland and miombo is similar, while the number of species in dry woodland, arid shrubland and grassland surpasses that of wet forest. The subfamily Nymphalinae is more diverse in miombo than forest and equally diverse in savanna. The Heliconiinae (Nymphalidae) in savanna are almost as diverse as they are in wet forest.

Composition, Diversity and Endemism

All six families and all of the subfamilies (except the Lycaeninae, Leach, 1815) of Afrotropical butterflies are represented in Angola (Table 10.1).

One genus and 56 species/subspecies of Papilionoidea are endemic to Angola, many of which were described over the last few decades. The endemic genus Mashunoides, Mendes and Bivar-de-Sousa 2009a, b, c, d (Nymphalidae: Satyrinae) is confined to Cuando Cubango Province, in the ecotone between miombo and savanna/dry woodland mosaic. Endemism rates for Angolan butterfly families are highest for the Papilionidae and Lycaenidae and lowest for the Hesperiidae and Riodinidae (Tables 10.1 and 10.2). Examples of endemic species are illustrated in Fig. 10.3.
Table 10.2

Endemic butterfly species and subspecies in Angola


Endemic species

Endemic subspecies


Eagris multiplagata

Abantis bergeri

Calleagris jamesoni ansorgei

Eretis herewardi rotundimacula

Spialia colotes colotes


Papilio bacelarae

Papilio chitondensis

Papilio macinnoni benguellae


Mylothris carvalhoi

Appias epaphia angolensis

Appias phaola uigensis

Appias sylvia ribeiroi

Mylothris spica gabela


Alaena rosei

Cooksonia nozolinoi

Falcuna lacteata

Deloneura barca

Aloeides angolensis

Zeritis krystyna

Cupidesthes vidua

Uranothauma nozolinoi Lepidochrysops ansorgei

Lepidochrysops flavisquamosa

Lepidochrysops fulvescens

Lepidochrysops hawker

Lepidochrysops nacrescens

Lepidochrysops reichenowi

Liptena homeyeri straminea

Falcuna libyssa angolensis

Cigarits modestus modestus

Leptomyrina henningi angolensis


Brakefieldia angolensis

Brakefieldia ochracea

Neita bikuarica

Mashunoides carneiromendesi

Charaxes figuerai

Charaxes ehmckei

Precis larseni

Bebearia hassoni

Euphaedra divoides

Euphaedra uigensis

Acraea bellona

Acraea lapidorum,

Acraea onerata

Amauris crawshayi angola,

Amauris dannfelti dannfelti

Charaxes fulvescens rubenarturi,

Charaxes macclouni carvalhoi,

Charaxes lucretius saldanhai

Charaxes jahlusa angolensis

Charaxes minor karinae

Charaxes trajanus bambi

Palla ussheri hassoni

Sevenia occidentalium penricei

Euphaedra harpalyce commineura

Acraea violarum anchietai

Fig. 10.3

The holotype specimens of endemic Angolan papilionoidea: Left to right, top to bottom (V Ventral, D Dorsal): 1. Abantis bergeri male D (Mendes and Bivar-de-Sousa 2009a, b, c, d), 2. Eagris multiplagata male V (Bivar-de-Sousa and Mendes 2007), 3. Cooksonia nozolinoi female D (Mendes and Bivar-de-Sousa 2007), 4. Papilio bacelarae male D (Bivar-de-Sousa and Mendes 2009a, b), 5. Mashunoides carneiromendesi male V (Mendes and Bivar-de-Sousa 2009a, b, c, d), 6. Charaxes jahlusa angolensis male D (Mendes et al. 2017), 7. Euxanthe trajanus bambi male D (Bivar-de-Sousa and Mendes 2006), 8. Euphaedra (Euphaedrana) divoides male V (Bivar- de-Sousa and Mendes 2018)


Because butterflies are sensitive to changing environmental conditions and are taxonomically well known, they are valuable as indicators of ecological dynamics. They are also key drivers of ecological processes. In particular, adult butterflies are active pollinators of many plants and the imagos and larvae are an important source of nutrition for a diverse range of vertebrate and invertebrate predators and insect parasitoids. Their conservation importance is also due to their positive and occasionally negative economic impacts. Although humans utilise mainly moth caterpillars as a food source, the larvae of the skipper Coeliades libeon is much appreciated. A limited number of butterfly species are agricultural pests, including Papilio demodocus (young citrus orchards), Lampides boeticus (cultivated Leguminosae) and Acraea acerata (sweet-potatoes). A few species, such as Pyrrhochalcia iphis and Zophopetes dysmephila, may cause damage in coconut and oil-palm plantations.

In terms of species of conservation concern, information on the status of Angolan butterflies is very limited. Many species of Angolan butterflies are obviously abundant and widespread, both within and outside the country. Those taxa that appear to be rare and/or more localised may be genuinely rare or local but this may simply reflect a paucity of information. This makes it difficult or impossible to propose rational conservation measures at present. The urgent need for more fieldwork, particularly in regard to the endemic taxa, is thus highlighted. In the meantime habitat conservation, especially with respect to isolated forest patches, can be considered as part of a wider effort to conserve both the fauna and flora of the country.

Potential Future Discoveries and Research

Considering the number of taxa new to science described in the last few decades there are almost certainly further undiscovered butterfly taxa in Angola. Vast areas of the country remain unexplored, mainly because of inaccessibility and post-independence political instability. Not only will new taxa be found but also known taxa from bordering countries will be added to the list of Angolan butterflies during future fieldwork. This work will also improve our knowledge in regard to the distribution of the taxa in the country. Finally, almost nothing is known about the habitats, behaviours, early stages and larval host plants of Angolan butterflies, making these fertile areas for future research on the fauna. More information concerning all the endemic taxa is urgently needed in order to determine conservation priorities.


  1. Ackery PR, Smith CR, Vane Wright RI (1995) Carcasson’s African Butterflies. An annotated catalogue of the Papilionoidea and Hesperioidea of the Afrotropical Region. CSIRO, Victoria, i–xii + 803 ppGoogle Scholar
  2. Aduse-Poku K, Vingerhoedt E, Wahlberg N (2009) Out-of-Africa again: A phylogenetic hypothesis of the genus Charaxes (Lepidoptera: Nymphalidae) based on five genes region. Mol Phylogenet Evol 53:463–478CrossRefGoogle Scholar
  3. Aurivillius C In Seitz A (1928) Les Macrolepidopteres du Globe. Les Macrolepidoptères de la Faune Ethiopienne 13(4):615 pp + 80 pl. E. Le Moult, ParisGoogle Scholar
  4. Bacelar A (1948) Lepidópteros de África principalmente das colónias portuguesas (colecção do Museu Bocage). Arquivos do Museu Bocage 19:165–207Google Scholar
  5. Bacelar A (1956) Lepidópteros (Rhopalocera) de Buco Zau, enclave de Cabinda, Angola. Anais da Junta de Investigações do Ultramar 11(3):175–197Google Scholar
  6. Bacelar A (1958a) Alguns Lepidópteros (Rhopalocera) do enclave de Cabinda. Revista portuguesa de Zoologia e Biologia Geral 1(2/3):197–217Google Scholar
  7. Bacelar A (1958b) Alguns Lepidópteros (Rhopalocera) da África Ocidental portuguesa. Revista portuguesa de Zoologia e Biologia Geral 1(4):311–330Google Scholar
  8. Bacelar A (1961) Lepidópteros do Bié (Rhopalocera) da colecção do Colégio de São Bento, em Luso (Angola). Memórias da Junta de Investigações do Ultramar (2)23:61–81Google Scholar
  9. Berger LA (1979) Espèces peu connues et descriptions de nouvelles sous-espèces de Mylothris (Lepidoptera Pieridae). Revue de Zoologie Africaine 93(1):1–9Google Scholar
  10. Bethune-Baker GT (1914) Notes on the taxonomic value of genital armature in Lepidoptera. Trans Entomol Soc London 1914:314–337Google Scholar
  11. Bivar-de-Sousa A (1983) Contribuição para o conhecimento dos lepidópteros de Angola (3ª nota). Dados sobre a ocorrência do género Charaxes (Lep. Nymphalidae) em Angola (1ª parte). Actas do I Congresso Ibérico de Entomologia 1:107–119Google Scholar
  12. Bivar-de-Sousa A, Fernandes JA (1964) Contribuição para o conhecimento dos Lepidópteros de Angola. Boletim da Sociedade Portuguesa de Ciências Naturais (2) 10(25):104–115Google Scholar
  13. Bivar-de-Sousa A, Mendes LF (2006) On the genus Euxanthe Hübner, 1819 in Angola, with description of a new subspecies (Lepidoptera, Nymphalidae, Charaxinae). Nouvelle Revue d’Entomologie 23(4):369–376Google Scholar
  14. Bivar-de-Sousa A, Mendes LF (2007) New data on the Uranothauma from Angola, with description of a new species (Lepidoptera: Lycaenidae: Polyommatinae). Boletin Sociedad Entomológica Aragonesa 41:73–76Google Scholar
  15. Bivar-de-Sousa A, Mendes LF (2009a) On a new species of the genus Princeps Hübner, (1807) from Cabinda (Angola) (Lepidoptera: Papilionidae). SHILAP, Revista de Lepidopterologia 37(147):313–318Google Scholar
  16. Bivar-de-Sousa A, Mendes LF (2009b) New data on the Angolan Charaxes of the “etheocles group” with description of a new species (Lepidoptera, Nymphalidae, Charaxinae). Boletim da Sociedade portuguesa de Entomologia 8(15)(229):293–309Google Scholar
  17. Bivar-de-Sousa A, Mendes LF (2014) New data on the Angolan Charaxesof the “etheocles group” with description of a new species (Lepidoptera, Nymphalidae, Charaxinae). Boletim da Sociedade portuguesa de Entomologia 229(8–15):293–309Google Scholar
  18. Bivar-de-Sousa A, Mendes LF, Vasconcelos S (2017) Description of one new species and one new subspecies of Nymphalidae from Angola (Lepidoptera: Papilionoidea). SHILAP, Revista de Lepidopterologia 45(178):227–236Google Scholar
  19. Bivar-de-Sousa A, Mendes LF (2018, in press) The “themis group” of Euphaedra (Euphaedrana) in Angola. Revision and description of one new species (Lepidoptera: Nymphalidae: Limenetidinae). Boletim da Sociedade portuguesa de EntomologiaGoogle Scholar
  20. Bouyer T (1999) Note sur les Charaxes du «groupe eupale» avec description d’une nouvelle sous-espèce (Lepidoptera Nymphalidae). Entomologia Africana 4(1):37–40Google Scholar
  21. Butler AG (1872) On a small collection of butterflies from Angola. Proc Zool Soc London 1871:721–725Google Scholar
  22. Carvalho EL (1962) Alguns Papilionídeos da Lunda (Lepidoptera). Publicações Culturais da Companhia dos Diamantes de Angola 60:163–170Google Scholar
  23. Condamin M (1966) Mise au point sur les Neptis au facies d’ «14avann» (Lepidoptera: Nymphalidae). Bulletin de l’Institut Fondamental de l’Afrique Noire 28(A)(3):1008–10029Google Scholar
  24. D’Abrera B (1980) Butterflies of the afrotropical region. Lansdowe Ed., Melbourne, pp i–xx + 1–593Google Scholar
  25. Dhungel B, Wahlberg N (2018) Molecular systematics of the subfamily Limenitidinae (Lepidoptera: Nymphalidae). PeerJ 6:e4311CrossRefGoogle Scholar
  26. Druce H (1875) A list of the collections of diurnal Lepidoptera made by J. J. Monteiro in Angola with description of some new species. Proc Zool Soc London 27:406–417Google Scholar
  27. Eltringham H (1912) A monograph of the African species of the genus Acraea Fab., with a supplement of those of the Oriental Region. Trans Entomol Soc London 1912(1):1–374Google Scholar
  28. Espeland M, Breinholt J, Willmott KR et al (2018) A comprehensive and dated phylogenomic analysis of butterflies. Curr Biol 28:1–9CrossRefGoogle Scholar
  29. Evans WH (1937) Catalog of African Hesperiidae (Indicating the classification and nomenclature adopted in the British Museum). British Museum, London, 212pp + 30 plGoogle Scholar
  30. Gardiner A (2004) Chapter 10. Butterflies of the four corners area. In: Timberlake JR, Childers SL (eds) Biodiversity of the Four Corners Area: Technical Review, vol 15. Occasional Publications on Biodiversity, Bulawayo, pp 381–397Google Scholar
  31. Hecq J (1997) Euphaedra. Lambillionea & Hecq, Tervuren & Mont-Sur-Marchienne, 121 pp + 48 plGoogle Scholar
  32. Heikkilä M, Kaila L, Mutanen M et al (2012) Cretaceous origin and repeated Tertiary diversification of the redefined butterflies. Proc R Soc Lond B 279:1093–1099CrossRefGoogle Scholar
  33. Henning SF (1988) The Charaxes Butterflies of Africa. Aloe Books, Johannesburg, 457 ppGoogle Scholar
  34. Holmes CWN (2001) A reaprisal of the Bebearia mardania complex (Lepidoptera, Nymphalidae). Trop Zool 14:31–62CrossRefGoogle Scholar
  35. Homeyer A, Dewitz H (1882) Drei neue westafrikanische Charaxes. Berliner Entomologisches Zeitschrift 26(II):381–383Google Scholar
  36. JIU (1948–1963) Cartas do levantamento aerofotogramétrico de Angola (escala 1: 250,000). Junta de Investigações do Ultramar: folhas:8–471Google Scholar
  37. Kielland J (1982) Revision of the genus Ypthima in the Ethiopian Region excluding Madagascar (Lepidoptera, Satyridae). Tijdschrift voor Entomologie 125(5):99–154Google Scholar
  38. Kielland J (1994) A revision of the genus Henotesia (excluding Madagascar and other Indian Ocean islands) (Lepidoptera Satyridae). Lambillionea 94(2):235–273Google Scholar
  39. Ladeiro JM (1956) Lepidópteros de Angola (estudo de uma colecção oferecida ao Museu Zoológico de Coimbra). Anais da Junta de Investigações do Ultramar 11(3):151–172Google Scholar
  40. Larsen TB (2005) Butterflies of West Africa. Text volume: 1–595. Apollo Books, StenstrupGoogle Scholar
  41. Latreille PA, Godart JB (1819) Encyclopédie Méthodologique. Histoire Naturelle (Zoologie), 9, Entomologie, Paris, i–iv + 1–328Google Scholar
  42. Lautenschläger T, Neinhuis C (eds) (2014) Riquezas Naturais do Uíge – Uma breve Introdução sobre o estado atual. A utilização, a Ameaça e a Preservação da Biodiversidade. Technische Universität Dresden, Dresden, 125 ppGoogle Scholar
  43. Libert M (1999) Revision des Epitola (ls). Révision des genres Epitola Westwood, Hypophytala Clench et Stempfferia Jackson et description de trois nouveaux genres (Lepidoptera Lycaenidae). ABRI & Lambillionea, Nairobi/Tervuren, pp 1–219, pl. I–XVIGoogle Scholar
  44. Libert M (2000) Révision du genre Mimacraea Butler, avec description de quatre nouvelles espèces et deux nouvelles sous-espèces (Lepidoptera, Lycaenidae). Lambillionea & ABRI, Tervuren/Nairobi, pp 1–73Google Scholar
  45. Libert M (2004) Revision du genre Oxylides Hübner (Lepidoptera, Lycaenidae). Lambillionea 104(2):143–158Google Scholar
  46. Mendes LF, Bivar-de-Sousa A (2006a) A new species of Neita van Son (Nymphalidae, Satyrinae) from southern Angola. Boletin Sociedad Entomológica Aragonesa 39:95–96Google Scholar
  47. Mendes LF, Bivar-de-Sousa A (2006b) Notes and descriptions of Afrotropical Appias butterflies (Lepidoptera: Pieridae). Boletin Sociedad Entomológica Aragonesa 39:151–160Google Scholar
  48. Mendes LF, Bivar-de-Sousa A (2007a) New species of Cooksonia Druce, 1905 from Angola (Lepidoptera: Lycaenidae, Lipteninae). SHILAP, Revista de Lepidopterologia 35(138):265–268Google Scholar
  49. Mendes LF, Bivar-de-Sousa A (2007b) On the genus Eagris Guenée, 1863 in Angola (Lepidoptera: Hesperiidae). SHILAP, Revista de Lepidopterologia 35(139):311–316Google Scholar
  50. Mendes LF, Bivar-de-Sousa A (2009a) Description of a new species of Mylothris from northern Angola (Lepidoptera Pieridae). Bolletino della Societá Entomológica Italiana 141(1):55–58Google Scholar
  51. Mendes LF, Bivar-de-Sousa A (2009b) New account on the butterflies of Angola. The genus Leptomyrina (Lepidoptera Lycaenidae). Bolletino della Societá Entomológica Italiana 141(2):109–112Google Scholar
  52. Mendes LF, Bivar-de-Sousa A (2009c) On a new south-eastern Angolan Satyrine butterfly belonging to a new genus (Lepidoptera, Nymphalidae). Entomologia Africana 14(2):5–8Google Scholar
  53. Mendes LF, Bivar-de-Sousa A (2009d) The genus Abantis Hopffer, 1855 in Angola and description of a new species (Lepidoptera: Hesperiidae, Pyrginae). SHILAP, Revista de Lepidopterologia 37(147):313–318Google Scholar
  54. Mendes LF, Bivar-de-Sousa A (2012) Notes on the species of Hypolycaena (Lepidoptera, Lycaenidae, Theclinae) known to occur in Angola. Boletin Sociedad Entomológica Aragonesa 50:193–197Google Scholar
  55. Mendes LF, Bivar-de-Sousa A, Figueira R (2013a) Butterflies of Angola / Borboletas diurnas de Angola. Lepidoptera. Papilionoidea, I. Hesperiidae, Papilionidae. IICT and CIBIO, Lisboa and Porto, 288 ppGoogle Scholar
  56. Mendes LF, Bivar-de-Sousa A, Figueira R et al (2013b) Gazetteer of the Angolan localities known for beetles (Coleoptera) and butterflies (Lepidoptera: Papilionoidea). Boletim da Sociedade Portuguesa de Entomologia 8(14/228):257–290Google Scholar
  57. Mendes LF, Bivar-de-Sousa A, Vasconcelos S et al (2017) Description of two new subspecies and notes on Charaxes Ochenheimer, 1816 of Angola (Lepidoptera: Nymphalidae). SHILAP, Revista de Lepidopterologia 45(178):299–315Google Scholar
  58. Mendes LF, Bivar-de-Sousa A, Vasconcelos S (2018, in press) On the butterflies of genus Precis Hübner, 1819 from Angola and description of a new species (Lepidoptera: Nymphalidae: Nymphalinae). SHILAP Revista de LepidopterologiaGoogle Scholar
  59. Monard A (1956) Compendium Entomologicum Angolae – 1. Insecta. VI – Ord. Lepidoptera. Anais da Junta de Investigações do Ultramar 11(3):119–128Google Scholar
  60. NHM – Natural History Museum (2004). Wallowtails. Available at:
  61. Pierre J (1988) Les Acraea du super-group «egina»: Révision et phylogénie (Lepidoptera : Nymphalidae). Annales de la Société Entomologique de France 24(3):263–287Google Scholar
  62. Pierre J, Bernaud D (2013) Butterflies of the world. NymphalidaeXXIII. Acraea subgenus Acraea. Goecke & Evers, Keltern, 39:1–8, pl. 1–28Google Scholar
  63. Rothschild W, Jordan K (1903) Lepidoptera collected by Oscar Neumann in north-east Africa. Novitates Zoologicae 10(3):491–542Google Scholar
  64. Smith CR, Vane-Wright RI (2001) A review of the Afrotropical species of the genus Graphium (Lepidoptera: Rhopalocera: Papilionidae). Bull Nat His Mus London (Entomol) 70(2):503–719Google Scholar
  65. Stempffer H (1957) Les Lépidoptères de l’Afrique noire française. Lycaenidés. Initiations Africaines 14(3):1–228Google Scholar
  66. Stempffer H, Bennett NH (1963) A new genus of Lipteninae (Lepidoptera: Lycaenidae). Bull Brit Mus (Nat His) (Entomol) 13:171–194Google Scholar
  67. Talbot G (1944) A preliminary revision of the genus Mylothris Hübn. (1819) (Lep. Rhop. Pieridae). T Roy Ent Soc London 94(2):155–185CrossRefGoogle Scholar
  68. Tite GE (1959) New species and notes on the genus Lepidochrysops (Lepidoptera, Lycaenidae). The Entomologist 92:158–163Google Scholar
  69. Tite GE (1961) New species of the genus Lepidochrysops (Lepidoptera, Lycaenidae). The Entomologist 94:21–25Google Scholar
  70. Tite GE, Dickson CGC (1973) The genus Aloeides and allied genera. Bull Brit Mus (Nat His) (Entomol) 29:227–280Google Scholar
  71. Trimen R (1891) On butterflies collected in tropical south-western Africa by Mr. A. W. Eriksson. Proc Zool Soc 1891:59–107Google Scholar
  72. Turlin B, Vingerhoedt E (2013) Butterflies of the World, supl. 23. Les Charaxinae de la faune Afrotropicale. Les genres Palla et Euxanthe. Nymphalidae: Charaxinae: Pallini et Euxanthini. Goecke & Evers, KelternGoogle Scholar
  73. Weymer G (1901) Beitrag zur Lepidopterofauna von Angola. Entomologischen Zeitschriften, Stuttgart 15(17):61–64, 65–67, 69–70Google Scholar
  74. Weymer G (1908) Einige neuer Lepidopteren des Deutschen Entom. National-Museums, gesammelt von Dr. F. Cr. Wellman in Benguella. Deutsche Entomologische Zeitschrift 1908:507–413Google Scholar
  75. Willis CK (2009) Amist the butterflies of southwestern Angola. Metamorphosis 20(3):74–88Google Scholar
  76. Williams MC (2018) Afrotropical butterflies. Available at:

Copyright information

© The Author(s) 2019

Open Access This chapter is licensed under the terms of the Creative Commons Attribution 4.0 International License (, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license and indicate if changes were made.

The images or other third party material in this chapter are included in the chapter's Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the chapter's Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder.

Authors and Affiliations

  1. 1.Museu Nacional de História Natural e da CiênciaUniversidade de LisboaLisboaPortugal
  2. 2.CIBIO, Centro de Investigação em Biodiversidade e Recursos GenéticosVairãoPortugal
  3. 3.Sociedade Portuguesa de EntomologiaLisboaPortugal
  4. 4.Pretoria UniversityPretoriaSouth Africa

Personalised recommendations