Biodiversity Data Journal : Taxonomic Paper
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Taxonomic Paper
Documenting museum records of West African Coccinellidae (Coleoptera) in Benin and Senegal
expand article infoKwevitoukoui Hounkpati‡,§, Joseph V. McHugh, Abdoul Aziz Niang|, Georg Goergen
‡ Department of Entomology, University of Georgia, Athens, United States of America
§ Grain de Sel Togo, Inc., Athens, United States of America
| Laboratoire de Zoologie des Invertébrés Terrestres, Institut Fondamental d’Afrique Noire Cheikh Anta Diop, Dakar, Senegal
¶ International Institute of Tropical Agriculture, Cotonou, Benin
Open Access

Abstract

Background

This work provides a preliminary inventory of West African Coccinellidae.

This was based on the West African Coccinellidae (WAC) specimens in the holdings of insect collections at the Laboratoire de Zoologie des Invertébrés Terrestres at the Institut Fondamental d’Afrique Noire Cheikh Anta Diop (IFAN), Senegal and the Biodiversity Center at the International Institute of Tropical Agriculture (IITAB), Benin.

New information

A total of 129 species representing 11 tribes and 40 genera is reported, including one species of the subfamily Microweiseinae and 128 species of the subfamily Coccinellinae. The geographic distribution of collection localities is presented for these species. Cheilomenes lunata (Fabricius, 1775), Cheilomenes propinqua (Mulsant, 1850), Cheilomenes sulphurea (Olivier, 1791), Chnootriba elaterii (Rossi, 1794), Chnootriba similis (Thunberg, 1781), Exochomus laeviusculus Weise, 1909, Hyperaspis delicatula (Mulsant, 1850) and Hyperaspis pumila Mulsant, 1850 are the best represented species in these collections.

Keywords

Africa, Coccinelloidea, Coccinellinae, distribution, diversity, insect collection, inventory, Microweiseinae, taxonomy, museum, systematics, lady beetles, ladybugs.

Introduction

Natural history museums play a critical role in science and education (Gropp 2018, Holmes et al. 2016). They contribute fundamental data, necessary for understanding the biodiversity of Earth across temporal and geographic ranges. Biological information derived from specimen data provides essential information supporting a wide range of basic and applied biological endeavours. Museum specimens often play an important role in the recognition of threatened taxa, allowing the scientific community to propose appropriate action (e.g. Thomson et al. 2018, Mikula et al. 2018). Other activities that benefit from specimen-based biological information include the monitoring of environmental change, protecting public health and safety and enhancing agriculture, to name a few (Suarez and Tsutsui 2004).

In agriculture, biological collections can provide vitally important information about pest species, including phenology, food preferences, behaviour, ecological associations etc. (Escalona et al. 2017). For invasive species, museum data can help to identify the point of entry, the date of introduction, the rate of expansion and the native distributional range, which is where natural enemies (potential biocontrol agents) might be found.

The full benefits of natural history museums can only be realised if these collections are known and accessible to the public and scientific communities worldwide, thereby enabling cooperation between local and distant scientists to explore and advance our knowledge of global biodiversity. Unfortunately, access to biological collections is limited for researchers in some regions, like Africa. Much of the available museum material of African origin is housed in natural history museums or private collections in Europe (Klopper et al. 2002, Miller and Rogo 2001). There are some major African national or regional natural history museums. The majority of their holdings comprise subsets of material that was collected by missionaries, explorers and scientists from European institutions (Medler 1980, Scholtz and Mansell 2017). There are some natural history museums in West Africa, but these have remained overlooked by the research community.

Coccinellidae Latreille, 1807, commonly known as ladybugs, are small beetles ranging from 0.8 to 18 mm (Seago et al. 2011). Although some coccinellids are phytophagous or fungivorous (Iperti 1999), nearly 70% of species are considered predaceous, preying on aphids, mealybugs, scale insects, thrips, leaf hoppers, mites and other soft bodied insects (Giorgi et al. 2009, Riddick 2017, Shah and Khan 2014, Ślipiński and Tomaszewska 2010, Szawaryn et al. 2015). One of the earliest and most successful examples of biocontrol was the management of cottony-cushion scale, Icerya purchasi Maskell, 1879, on citrus crops using the vedalia lady beetle, Rodolia cardinalis (Mulsant, 1850), in California during the 1890s.

Due to their economic importance, major regional taxonomic works have been published for the coccinellid faunas of North America, Europe, Palearctic Region, Russian Far East, other parts of the former USSR, Oriental region, Central Asia, Japan, Vietnam, China and Australia (e.g. Giorgi et al. 2009, Tomaszewska and Szawaryn 2013, Tomaszewska and Ślipiński 2013, Ślipiński et al. 2012). Unfortunately, nothing comparable exists for Africa. The identification of African lady beetles has been done almost exclusively by European entomologists and explorers (Medler 1980). As a result, valuable reference specimens and their associated collection data are often inaccessible to researchers and agencies in Africa (Scholtz and Mansell 2017).

Coccinellidae has been the focus of several recent phylogenetic studies as researchers attempt to understand the evolution of the group and to improve its classification (e.g. Escalona and Ślipiński 2012, Kobayashi et al. 2011, Nedved and Kovář 2012, Robertson et al. 2015, Seago et al. 2011, Tomaszewska and Szawaryn 2013, Tomaszewska and Ślipiński 2013, Ślipiński et al. 2012). Unfortunately, endemic African taxa are very poorly represented in these studies, probably because appropriate material was unavailable and because the state of taxonomy for those groups was not mature.

Although Africa is well known for its rich and charismatic vertebrate diversity, there is far less appreciation for the great diversity of other taxa there. It is estimated that 100,000 species of insects are currently known from the continent and conservative estimates put the total number of insect species there at about 600,000, yet few research collections of insects exist on the continent (Miller and Rogo 2001). While entomological research collections in some African countries (e.g. Algeria, Egypt, Ghana, Kenya, Libya, Malawi, Morocco, Mozambique, South Africa, Tanzania, Uganda and Zambia) have appeared for some time in registries of museums (e.g. Arnett et al. 1993, Evenhuis 2019), others remain virtually unknown to the outside world, especially those in French West African countries (Afrique Occidentale Française). This is true for the two largest reference collections of arthropods in West Africa, the Biodiversity Center at the International Institute of Tropical Agriculture (IITAB) in Benin and the Laboratoire de Zoologie des Invertébrés Terrestres at the Institut Fondamental d’Afrique Noire Cheikh Anta Diop (IFAN) in Senegal. These two important resources were absent from compilations of insect and spider collections of the world until just recently when they were added to a web-based listing of biological collections by Evenhuis (2019) following an enquiry by the senior author.

The Laboratoire de Zoologie des Invertébrés Terrestres (formerly Section Entomologie) was created in 1945 by André Villiers. It is housed in the Institut Fondamental d’Afrique Noire Cheikh Anta Diop (IFAN), Université Cheikh Anta Diop, Dakar in Senegal (Fig. 1a–b). The IFAN insect collection was established to serve as a centre for entomological collection-based research in West Africa. With over 400,000 specimens, the IFAN insect museum is the largest insect collection in West Africa. Dr. Abdoul Aziz Niang, a specialist of Phlebotomine sandflies (Diptera: Psychodidae), is the current Curator and Director of the IFAN insect collection.

Figure 1.  

Photos of the two largest West African insect museums: a. IFAN, outside view of the museum; b. IFAN, inside view of the museum showing insect boxes; c. IITAB, outside view of the museum; d. IITAB, inside view of the museum showing insect cabinets and drawers.

The International Institute of Tropical Agriculture (IITA) is a non-profit international research organisation founded in 1967. Headquartered in Ibadan, Nigeria, IITA is a member of the Consultative Group for International Agricultural Research. IITA has stations and hubs in Central, Eastern, Southern and West Africa (www.iita.org). The IITAB, IITA Biodiversity Center (formerly IITA insect centre or museum) is housed at the Benin Station of IITA in Calavi, Cotonou, Benin (Fig. 1c–d). With over 365,000 specimens, the IITAB collection is the second largest insect collection in West Africa. Dr. Georg Goergen, Entomologist and Biosystematist, is the current Curator and Director of the IITAB.

The taxonomic impediment, which affects biologists around the world, impacts West African researchers especially hard. Most African scientists must rely on distant experts for the identification of insect specimens. Africa produces far fewer trained insect taxonomists than any other continent in the world (de Carvalho et al. 2005). This gap in taxonomic knowledge has also limited the assessment of biodiversity in Africa (Coleman 2015). Currently, the Biodiversity Center of IITA, Benin (IITAB) is one of the major insect identification hubs in Africa.

The IFAN and IITA insect collections are the two largest in West Africa, a region situated between the Tropic of Cancer and Equator, covering 6,140,000 km2, approximately one fifth of Africa. More than 75% of the land consists of plains lying below an elevation of 300 m. The region includes 16 countries: Benin, Burkina Faso, Cape Verde, Côte d’Ivoire, Gambia, Ghana, Guinea, Guinea-Bissau, Liberia, Mali, Mauritania, Niger, Nigeria, Senegal, Sierra Leone and Togo. Additionally, the United Nations recognises Saint Helena (a United Kingdom Overseas Territory), Ascension and Tristan da Cunha as part of West Africa (Comité Inter-états de Lutte contre la Sécheresse dans le Sahel 2016, Kabo-bah and Diji 2018) (Fig. 2).

Figure 2.  

Map of West Africa (United Nations 2014).

Given the importance of coccinellids and the patchy knowledge of their diversity in West Africa, we summarised data about the taxonomic holdings of this family in IFAN and IITAB insect collections with the goals of raising the visibility of those institutions and providing a preliminary inventory of West African Coccinellidae (WAC). This work represents part of ongoing research on the WAC by the senior author, who is currently developing a formal taxonomic catalogue of West African Coccinellidae.

Materials and methods

Museum specimens were studied at the IFAN and IITAB insect collections in West Africa. We photographed specimens and recorded label data (e.g. taxonomic determination, collection locality, collection date, associated plants, collector, determiner etc.).

The taxonomic names that were recorded on museum specimens were put in a database and updated to currently valid names using the systematics literature. Aberrations, variations and subspecies were not included in the database except when museum specimens were identified as such. References are provided, when appropriate, to clarify the current classification for species. In some cases, specimen determinations were made (by KH) using the literature along with examination of reference collections of authoritatively identified material at the Musée Royal de l'Afrique Centrale (MRAC), Museum für Naturkunde der Humboldt-Universität (ZMHB), Museum of Comparative Zoology (MCZ), National Museum of Natural History (NMNH) and University of Georgia Collection of Arthropods (UGCA). Annotations are given when an invalid determination on a specimen label has been updated to the currently valid taxonomic name.

Many museums and institutions kindly assisted this study by hosting visits, providing data or initiating loans of specimens for this and related studies. These collections and institutions include the following:

CERAAS, Centre d’Etude Régional pour l’Amélioration de l’Adaptation à la Sécheresse, Université Cheikh Anta Diop de Dakar, Senegal.

IFAN, Institut Fondamental d’Afrique Noire Cheikh Anta Diop, Dakar, Senegal.

IITAB, International Institute of Tropical Agriculture, Benin Station, Cotonou, Benin.

MCZ, Museum of Comparative Zoology, Harvard University, Cambridge, MA, U.S.A.

MRAC, Musée Royal de l'Afrique Centrale, Tervuren, Belgium.

NMNH, National Museum of Natural History (formerly USNM: United States National Museum), Washington D.C., U.S.A.

UCAD, Université Cheikh Anta Diop de Dakar, Dakar, Senegal.

UEM/IPD, Unité d'Entomologie Médicale, Institut Pasteur de Dakar, Dakar, Senegal.

UGA, University of Georgia, Athens GA, U.S.A.

UGCA, University of Georgia Collection of Arthropods, Georgia Museum of Natural History, Athens, GA, U.S.A.

ZMHB, Museum für Naturkunde der Humboldt-Universität, Berlin, Germany.

Other abbreviations used in the text:

DRC, Democratic Republic of the Congo.

The updated specimen database was used to produce the first list of West African coccinellid diversity (Table 1) and to explore general characteristics about the museum holdings. Valid taxonomic names and tribal placement follow Seago et al. (2011). Generic and species names are arranged alphabetically. The lists of synonyms provided below valid names are not exhaustive, but instead include only junior synonyms that have been used in literature regarding West African taxa or ones that appear on specimen determination labels in the focal collections.

Updated list of coccinellid holdings of the IFAN and IITAB collections, where: a*: present at IFAN, but not from West African locality, b*: present at IITAB, but not from West Africa locality, a: present at IFAN, West African source, b: present at IITAB, West African source and ab: in both museums. The current list is limited to specific rank only. Subspecies and aberrations are mentioned only if they were found on specimen determination labels.

Taxon Collection localities References
Family Coccinellidae Latreille, 1807
Subfamily Microweiseinae Leng, 1920
Tribe Serangiini Pope, 1962
Serangium kunowi Weise, 1892*1 Zambiab* Escalona and Ślipiński 2012
Subfamily Coccinellinae Latreille, 1807
Tribe Chilocorini Mulsant, 1846
Brumoides foudrasii (Mulsant, 1850) Beninb, Gambiab, Guineab, Nigeriab, Senegala Mader 1954
= Exochomus foudrasii Mulsant, 1850
Chilocorus distigma (Klug, 1835) Mozambiqueb*, Nigeriab, Senegala
Chilocorus dorhni Mulsant, 1850 Senegala
Chilocorus schioedtei Mulsant, 1850 Beninab, Cameroona*, Ghanab*, Guineaa, Nigeriab, Senegala Chazeau and Couturier 1985, Samways et al. 1999
= Chilocorus discoideus Crotch, 1874
Chilocorus simoni Sicard, 1907 South Africaab*
Exochomus flavipes (Thunberg, 1781) Gabonb, Madagascarb*, Nigeriab, Senegala
Exochomus laeviusculus Weise, 1909 Benina, Côte d’Ivoirea, Guineaa, Malia, Mauritaniaa, Senegala, Togoa
Exochomus nigrifrons Gerstäcker, 1871*2 Malia, Senegalab Biranvand et al. 2017a, Łączyński and Tomaszewska 2012
= Brumus nigrifrons Gerstäcker, 1871
= Brumus fulviventris Fairmaire, 1884
= Brumus trivittatus Weise, 1891
= Brumus nigeriana Korschefsky, 1938
= Brumus nigrifrons nigerianus Korschefsky, 1938
Parexochomus nigripennis (Erichson, 1843) Malia, Senegala Biranvand et al. 2017b, Li et al. 2016, Kovář 2007
= Exochomus nigripennis (Erichson, 1843)
= Exochomus nigromaculatus nigripennis Crotch, 1874
Exochomus pulchellus Gerstäcker, 1871 Gambiab, Nigera, Rwandab*, Senegala
Exochomus troberti Mulsant, 1850 Burkina Fasoa, Nigeriab, Senegala Hodek et al. 2012, Mohamed et al. 2018, Kovář 2007
= Exochomus flavipes troberti Mulsant, 1850
Tribe Coccidulini Mulsant, 1846
= Scymnini Mulsant, 1846
= Exoplectrini Crotch, 1874
Aulis annexa Mulsant, 1850 Senegala
Clitostethus flavotestaceus Mader, 1955 Senegala
Nephus flavomaculatus Fürsch, 1966 Beninb, Nigeriab
Nephus vetustus Weise, 1915 Gabonb, Nigeriab
Nephus phenacoccophagus Fürsch, 1987 Nigeriab
Nephus kamburovi Fürsch, 1992 Malawib*
Nephus oblongosignatus Mulsant, 1850 Tanzaniab*
Nephus ornatulus Korschefsky, 1931 DRCb*, Ghanab, Nigeriab, Rwanda*, Senegalb, Sierra Leoneb, Togob Fürsch 1966
= Scymnus ornatulus Korschefsky, 1931
Nephus sudanicus Weise, 1925 Mauritaniaa
Scymnobius bilucernarius (Mulsant, 1850) Mexicob* Gordon and González 2002
= Nephus bilucernarius (Mulsant, 1850)
Scymnus canariensis Wollaston, 1864 São Tome and Principea*, Senegala
Scymnus casstroemi Mulsant, 1850 Guineaa, Senegala
Scymnus gnavus Weise, 1895 Guineaa
Scymnus kibonotensis Weise, 1910 Côte d’Ivoirea, Guineaa, Nigeriab
Scymnus levaillanti Mulsant, 1850 Nigeriab, Malawib*
Scymnus pruinosus Weise, 1895 Zambiab*
Scymnus monroviae Casey, 1899 Benina, Côte d’Ivoirea, Guineaa, Nigera, Senegala, Togoa
Scymnus nigrosellatus Mader, 1950 Zambiab*
Scymnus quadrivittatus Mulsant, 1850 Nigeriab
Scymnus rubiginosus Mader, 1950 Côte d’Ivoirea, Guineaa, Senegala
Scymnus scapuliferus Mulsant, 1850 Benina, Côte d’Ivoirea, Guineab, Nigeriab, Madagascarb*, Senegala, Togoa
Scymnus schoutedeni Mader, 1950 Senegala
Scymnus senegalensis Mader, 1955 Côte d’Ivoirea, Gambiaa, Guineaa, Malia, Mauritaniaa, São Tome and Principea, Senegala
Scymnus villiersi Mader, 1955 Nigera, Senegala
Stethorus aethiops Weise, 1899 Beninb, Ghanab, Guinea-Bissaub, Mozambiqueb*, Nigeriab, Sierra Leoneb, Tanzaniab*
Stethorus endroedyi Fürsch, 1970 Malawib*
Stethorus jejunus Casey, 1899 Ghanab, Nigeriab, Mozambiqueb*, Tanzaniab*
Tribe Coccinellini Latreille, 1807
Adalia bipunctata (Linnaeus, 1758) Cameroona*
Anisolemnia decempustulata Weise, 1888 Togoa Chazeau and Couturier 1985, Mader 1954
= Anisolemnia 10-pustulata Weise, 1888
Bulaea anceps (Mulsant, 1850) Mozambiqueb* Fürsch 2005
= Isora circularis Mader, 1941
Caria welwitschii Crotch, 1874 Guineaa
Cheilomenes aurora Gerstäcker 1871 Tanzaniaa* Fürsch 1975
= Cydonia aurora Gerstäcker, 1871
Cheilomenes lunata (Fabricius, 1775) Benina, Burkina Fasoa, Cameroona*, Côte d’Ivoirea, Gabona*, Gambiaa, Guineaa, Guinea-Bissaua, Liberiaa, Malia, Senegala, West Africaa, South Africaa*, Tanzaniaa* Fürsch 1991a
= Cydonia lunata (Fabricius, 1775)
= Cydonia lunata vulpina (Fabricius, 1798)
= Cydonia lunata vulpiphursa Olivier, 1791
Cheilomenes sulphurea (Olivier, 1791) Angolab*, Cameroona*, Côte d’Ivoirea, Democratic Republic of Congo (DRC)b*, Gabonb*, Ghanab, Nigeriab, Malawib*, Rwandab*, Senegala Eizaguirre 2007
= Cheilomenes orbicularis Casey, 1899
= Cheilomenes sulphurea sulphurea (Olivier, 1791)
Cheilomenes propinqua (Mulsant, 1850) Côte d’Ivoire, Gabonb* , Guinea-Bissauab, Malia, Mauritaniaa, Nigera, Nigeriab, Senegalab Eizaguirre 2007, Fürsch 1979
= Cheilomenes vicina (Mulsant, 1850)
= Cydonia vicina Mulsant, 1850
= Cheilomenes vicina vicina Mulsant, 1850
=Cheilomenes propinqua vicina Mulsant, 1850
Cheilomenes quadrilineata (Mulsant, 1850) Senegala
= Cydonia 4-lineata Mulsant, 1850
Coccinella intermedia (Crotch, 1874) São Tome and Principe a* Gordon 1987a, Nematollahi 2016
= Lioadalia intermedia Crotch, 1874*3
= "Cydonia intermedia" Cramer
Coccinella septempunctata Linnaeus, 1758 Cape Verdea
Declivitata hamata (Thunberg, 1808) Senegala Fürsch 1987
= Alesia hamata (Mulsant, 1850)
= Micraspis striata (Crotch, 1874)
= Alesia striata (Gemminger & Harold, 1876)
= Alesia striata hamata (Weise, 1898)
= Declivitata hamata Fürsch, 1964
Declivitata uncifera Fürsch, 1967 Cameroona*, DRCb*, Guinea-Bissaub
Harmonia vigintiduomaculata (Fabricius, 1792) Beninb, Liberiaa, Nigeriab, Togob Coutanceau 2008
= Stictoleis vigintiduomaculata (Fabricius, 1792)
= Stictoleis 22-maculata (Fabricius, 1792)
Hippodamia variegata (Goeze, 1777) Colombiab*, Nigeriab, Senegalb, Tanzaniaa* GBIF Secretariat 2019, Gordon 1987b
= Adonia variegata (Goeze, 1777)
Lemnia machadoi Mader, 1952 Cameroona Fürsch 2002
= Dysis sicardi Mader, 1954
Megalocaria dilatata (Fabricius, 1775) Beninb Hodek et al. 2012, Poorani 2002
= Anisolemnia dilatata (Fabricius, 1775)
Micraspis lineola (Fabricius, 1775) Togob Nattier et al. 2015, Ślipiński 2007
= Alesia lineola (Fabricius, 1775)
Micraspis striata (Fabricius, 1792) Côte d’Ivoirea, Gabonb*, Guineaa, São Tome and Principea, Senegala* Kovář 2007, Ślipiński 2007
= Alesia striata (Fabricius, 1792)
Psyllobora bisoctonotata (Mulstant, 1850) Senegala Ali et al. 2018
Psyllobora lutescens (Crotch, 1874) Guatemalab*
Psyllobora variegata (Fabricius, 1781) South Africaa* Nicolas 2013
= Thea variegata (Fabricius, 1781)
Xanthadalia effusa (Erichson, 1843) Beninb, DRCa
Xanthadalia rufescens Mulsant, 1850 Beninb, Malia, Mauritaniaa, Senegala Fürsch 1987
Tribe Diomini Gordon, 1999
Diomus hennesseyi Fürsch, 1987 Nigeriab
Tribe Epilachnini Mulsant, 1846
Chnootriba elaterii (Rossi, 1794) Benina, Côte d’Ivoirea, Gambiaa, Guineaa, Liberiaa, Malia, Mauritaniaa, Nigeriab, São Tome and Principea*, Senegala Hodek et al. 2012, Jadwiszczak and Węgrzynowicz 2003, Tomaszewska and Szawaryn 2016
= Henosepilachna elaterii (Rossi, 1794)
= Epilachna chrysomelina (Fabricius, 1775)
= Epilachna chrysomelina manca Mader, 1929
= Henosepilachna elaterii voltaensis senegalensis Fürsch, 1964
Chnootriba hirta (Thunberg, 1781) Guineaa, Tanzaniaa* Tomaszewska and Szawaryn 2016
= Henosepilachna hirta (Thunberg, 1781)
= Epilachna hirta (Thunberg, 1781)
Chnootriba similis (Thunberg, 1781) Benina, Burkina Fasoa, Côte d’Ivoirea, DRCa*, Guineaa, Liberiaa, Nigeriab, Senegala Chazeau and Couturier 1985, Jadwiszczak and Węgrzynowicz 2003, Tomaszewska and Szawaryn 2016
= Chnootriba assimilis Mulsant, 1850
= Chnootriba similis ab. repanda Sicard, 1930
Cleta punctipennis (Mulsant, 1850) Togoab Tomaszewska and Szawaryn 2016
= Epilachna punctipennis Mulsant, 1850
Cleta sahlbergi (Mulsant, 1850) Côte d’Ivoirea, Kenyaa* Szawaryn et al. 2015, Tomaszewska and Szawaryn 2016
= Solanophila sahlbergi Mulsant, 1850
Epilachna bissexguttata Weise, 1895 Côte d’Ivoirea, DRCa*, Malia, Nigera, Senegala Jadwiszczak and Węgrzynowicz 2003
= Epilachna monticola Weise, 1899
= Solanophila monticola Weise, 1898
Epilachna bomparti Mulsant, 1850 Liberiaa, Senegala
Epilachna colorata Mulsant, 1850 Cameroona*, Liberiaa Jadwiszczak and Węgrzynowicz 2003
= Epilachna subsignata Mulsant, 1895
= Solanophila subsignata Mulsant, 1895
= Solanophila elliptica Weise, 1912
Epilachna iocosa (Mader, 1941) South Africaa* Jadwiszczak and Węgrzynowicz 2003
= Solanophila 20-punctata Mader, 1941
Epilachna nigritarsis Mulsant, 1850 Cameroona*, Liberiaa Jadwiszczak and Węgrzynowicz 2003
= Epilachna impatiens Fürsch, 1960
Epilachna vigintipunctata Mulsant, 1850 Liberiaa, Tanzaniaa*, Togoa Jadwiszczak and Węgrzynowicz 2003
= Epilachna punctipennis multinotata Gerstäcker, 1873
Henosepilachna atropos (Sicard, 1912) Equatorial Guineaa*, Senegala Jadwiszczak and Węgrzynowicz 2003
= Epilachna atropos Sicard, 1912
Henosepilachna bisseptemnotata (Mulsant, 1853) Tanzaniaa* Jadwiszczak and Węgrzynowicz 2003
= Epilachna bisseptemnotata Mulsant, 1853
Henosepilachna clavareaui (Weise, 1901) Benina Fürsch 1991b
= Epilachna clavareaui Weise, 1901
Henosepilachna ertli (Weise, 1906) Côte d’Ivoirea, Liberiaa Jadwiszczak and Węgrzynowicz 2003
= Epilachna ertli Weise, 1906
Henosepilachna fulvosignata (Reiche, 1847) Côte d’Ivoirea Fürsch 1991b
Henosepilachna moseri (Weise, 1903) Equatorial Guineaa* Jadwiszczak and Węgrzynowicz 2003
= Epilachna moseri Weise, 1903
Henosepilachna reticulata (Olivier 1791) Beninb, Malia, Nigerb, Nigeriab, Senegala Jadwiszczak and Węgrzynowicz 2003, Heinrichs and Barrion 2004
= Epilachna reticulata (Olivier 1791)
Henosepilachna simplex (Weise, 1895) Liberiaa Jadwiszczak and Węgrzynowicz 2003
= Epilachna simplex Weise, 1895
Solanophila canina (Fabricius, 1781) Guineaa
Solanophila dregei (Mulsant, 1850) Côte d’Ivoirea Tomaszewska and Szawaryn 2016
= Epilachna dregei Mulsant, 1850
Solanophila scalaris (Gerstäcker, 1871) Tanzaniaa* Mader 1941
= Epilachna scalaris (Gerstäcker, 1871)
Tribe Hyperaspini Mulsant, 1846
Hyperaspis aestimabilis Mader, 1955 Angolab*, DRCb*, Malawib*, Zambiab*
Hyperaspis centralis Mulsant, 1850 Mexico
Hyperaspis delicatula (Mulsant, 1850) Beninb, Gambiab, Ghanab, Guinea-Bissaub, Nigeriab, Malawib*, Sénégalb, Sierra Leoneb, Togob
Hyperaspis lugubris (Randall, 1838) Ghanab, Nigeriab Gordon 1985
= Hyperaspis jucunda LeConte, 1852
Hyperaspis maindroni (Sicard, 1929) Mauritaniaa, Nigera, Senegala Biranvand et al. 2017b, Ali et al. 2018
Hyperaspis merckii (Mulsant, 1850) Mauritaniaa, Senegala
Hyperaspis pumila Mulsant, 1850 Gambiab, Guineab, Guinea-Bissaub, Nigera, Nigeriab, Senegalb, Togob Biranvand et al. 2017b
Hyperaspis senegalensis (Mulsant, 1850) Gambiab, Ghanab, Nigeriab, Senegalb, Sierra Leoneb, Malawib*
Hyperaspis sericea Fürsch, 1972 Malawib*
Hyperaspis vinciguerra Capra, 1929 Gambiab, Senegalb, Malawib*
Tenuisvalvae notata (Mulsant, 1850) Beninb, Boliviab*, Nigeriab Gordon and Canepari 2008
= Hyperaspis notata Crotch, 1874)
Tribe Ortaliini Mulsant, 1850
Ortalia ovulum Weise, 1898 Liberiaa, Malia, Togob
Tribe Noviini Mulsant, 1846
Rodolia cardinalis (Mulsant, 1850) Kenyab*
Rodolia iceryae Janson in Ormerod, 1887 Senegala Hounkpati et al. 2019
= Rodolia iceryae Janson, 1887
= Rodolia obscura Weise, 1898
Rodolia occidentalis Weise, 1898 Benina, Ghanaa, Nigeriaa, Senegalab
Rodolia senegalensis Weise, 1913 Senegala
Tribe Platynaspini Mulsant, 1846
Platynaspis capicola Crotch, 1874 DRCb*, Malawib*
Platynaspis ferruginea Weise, 1895 Beninb, Togob
Platynaspis kollari Mulsant, 1850 Liberiaa
Platynaspis obscura Gorham, 1901 Côte d’Ivoirea, Liberiaa
Platynaspis pilosa Sicard, 1930 South Africaa*
Platynaspis rufipennis Gerstäcker, 1871 Côte d’Ivoirea, Liberiaa, Nigera
Platynaspis vittigera Weise, 1895 DRCb*
Tribe Sticholotidini Weise, 1901
Pharoscymnus sexguttatus (Gyllenhall, 1808) Ghanab
Nomen nudum
Leis maculata*4 Côte d’Ivoirea

Checklist of West African Coccinellidae in IFAN and IITAB museums (see Table 1 for more details)

Serangium kunowi Weise, 1892

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Distribution: 
Zambia

Brumoides foudrasii (Mulsant, 1850)

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Distribution: 
Benin, Gambia, Guinea, Nigeria, Senegal

Chilocorus distigma (Klug, 1835)

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Distribution: 
Mozambique

Chilocorus dorhni Mulsant, 1850

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Distribution: 
Senegal

Chilocorus schioedtei Mulsant, 1850

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Distribution: 

Benin, Cameroon

Chilocorus simoni Sicard, 1907

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Distribution: 
South Africa

Exochomus flavipes (Thunberg, 1781)

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Distribution: 

Gabon

Exochomus laeviusculus Weise, 1909

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Distribution: 

Benin, Côte d’Ivoire, Guinea, Mali, Mauritania, Senegal, Togo

Exochomus nigrifrons Gerstäcker, 1871

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Distribution: 
Mali, Senegal

Parexochomus nigripennis (Erichson, 1843)

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Distribution: 
Mali, Senegal

Exochomus pulchellus Gerstäcker, 1871

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Distribution: 
Gambia, Niger, Rwanda, Senegal

Exochomus troberti Mulsant, 1850

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Distribution: 
Burkina Faso

Aulis annexa Mulsant, 1850

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Distribution: 
Senegal

Clitostethus flavotestaceus Mader, 1955

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Distribution: 
Senegal

Nephus flavomaculatus Fürsch, 1966

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Distribution: 
Benin, Nigeria

Nephus vetustus Weise, 1915

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Distribution: 
Gabon

Nephus phenacoccophagus Fürsch, 1987

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Distribution: 
Nigeria

Nephus kamburovi Fürsch, 1992

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Distribution: 
Malawi

Nephus oblongosignatus Mulsant, 1850

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Distribution: 
Tanzania

Nephus ornatulus Korschefsky, 1931

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Distribution: 
DRC

Nephus sudanicus Weise, 1925

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Distribution: 
Mauritania

Scymnobius bilucernarius (Mulsant, 1850)

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Distribution: 
Mexico

Scymnus canariensis Wollaston, 1864

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Distribution: 

São Tome and Principe, Senegal

Scymnus casstroemi Mulsant, 1850

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Distribution: 
Guinea, Senegal

Scymnus gnavus Weise, 1895

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Distribution: 
Guinea

Scymnus kibonotensis Weise, 1910

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Distribution: 

Côte d’Ivoire, Guinea, Nigeria

Scymnus levaillanti Mulsant, 1850

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Distribution: 
Nigeria, Malawi

Scymnus pruinosus Weise, 1895

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Distribution: 
Zambia

Scymnus monroviae Casey, 1899

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Distribution: 
Benin, Côte d’Ivoire, Guinea, Niger, Senegal, Togo

Scymnus nigrosellatus Mader, 1950

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Distribution: 
Zambia

Scymnus quadrivittatus Mulsant, 1850

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Distribution: 
Nigeria

Scymnus rubiginosus Mader, 1950

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Distribution: 

Côte d’Ivoire, Guinea, Senegal

Scymnus scapuliferus Mulsant, 1850

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Distribution: 
Benin, Côte d’Ivoire, Guinea, Nigeria, Madagascar, Senegal, Togo

Scymnus schoutedeni Mader, 1950

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Distribution: 
Senegal

Scymnus senegalensis Mader, 1955

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Distribution: 

Côte d’Ivoire, Gambia, Guinea, Mali, Mauritania, São Tome and Principe, Senegal

Scymnus villiersi Mader, 1955

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Distribution: 
Niger, Senegal

Stethorus aethiops Weise, 1899

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Distribution: 
Benin, Ghana, Guinea-Bissau, Mozambique

Stethorus endroedyi Fürsch, 1970

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Distribution: 
Malawi

Stethorus jejunus Casey, 1899

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Distribution: 
Ghana, Nigeria, Mozambique

Adalia bipunctata (Linnaeus, 1758)

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Distribution: 

Cameroon

Anisolemnia decempustulata Weise, 1888

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Distribution: 
Togo

Bulaea anceps (Mulsant, 1850)

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Distribution: 
Mozambique

Caria welwitschii Crotch, 1874

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Distribution: 
Guinea

Cheilomenes aurora Gerstäcker, 1871

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Distribution: 
Tanzania

Cheilomenes lunata (Fabricius, 1775)

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Distribution: 
Benin, Burkina Faso

Cheilomenes sulphurea (Olivier, 1791)

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Distribution: 

Angola, Cameroon

Cheilomenes propinqua (Mulsant, 1850)

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Distribution: 
Côte d’Ivoire, Gabon

Cheilomenes quadrilineata (Mulsant, 1850)

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Distribution: 
Senegal

Coccinella intermedia (Crotch, 1874)

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Distribution: 

São Tome and Principe

Coccinella septempunctata Linnaeus, 1758

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Distribution: 
Cape Verde

Declivitata hamata (Thunberg, 1808)

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Distribution: 
Senegal

Declivitata uncifera Fürsch, 1967

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Distribution: 

Cameroon

Harmonia vigintiduomaculata (Fabricius, 1792)

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Distribution: 
Benin, Liberia, Nigeria, Togo

Hippodamia variegata (Goeze, 1777)

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Distribution: 
Colombia, Nigeria, Senegal, Tanzania

Lemnia machadoi Mader, 1952

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Distribution: 

Cameroon

Megalocaria dilatata (Fabricius, 1775)

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Distribution: 
Benin

Micraspis lineola (Fabricius, 1775)

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Distribution: 
Togo

Micraspis striata (Fabricius, 1792)

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Distribution: 
Côte d’Ivoire, Gabon

Psyllobora bisoctonotata (Mulstant, 1850)

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Distribution: 
Senegal

Psyllobora lutescens (Crotch, 1874)

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Distribution: 
Guatemala

Psyllobora variegata (Fabricius, 1781)

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Distribution: 
South Africa

Xanthadalia effusa (Erichson, 1843)

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Distribution: 
Benin, DRC

Xanthadalia rufescens Mulsant, 1850

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Distribution: 
Benin, Mali, Mauritania, Senegal

Diomus hennesseyi Fürsch, 1987

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Distribution: 
Nigeria

Chnootriba elaterii (Rossi, 1794)

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Distribution: 

Benin, Côte d’Ivoire, Gambia, Guinea, Liberia, Mali, Mauritania, Nigeria, São Tome and Principe, Senegal

Chnootriba hirta (Thunberg, 1781)

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Distribution: 
Guinea, Tanzania

Chnootriba similis (Thunberg, 1781)

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Distribution: 

Benin, Burkina Faso

Cleta punctipennis (Mulsant, 1850)

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Distribution: 
Togo

Cleta sahlbergi (Mulsant, 1850)

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Distribution: 
Côte d’Ivoire, Kenya

Epilachna bissexguttata Weise, 1895

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Distribution: 
Côte d’Ivoire, DRC

Epilachna bomparti Mulsant, 1850

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Distribution: 
Liberia, Senegal

Epilachna colorata Mulsant, 1850

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Distribution: 

Cameroon

Epilachna iocosa (Mader, 1941)

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Distribution: 
South Africa

Epilachna nigritarsis Mulsant, 1850

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Distribution: 

Cameroon

Epilachna vigintipunctata Mulsant, 1850

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Distribution: 
Liberia, Tanzania

Henosepilachna atropos (Sicard, 1912)

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Distribution: 
Equatorial Guinea

Henosepilachna bisseptemnotata (Mulsant, 1853)

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Distribution: 
Tanzania

Henosepilachna clavareaui (Weise, 1901)

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Distribution: 
Benin

Henosepilachna ertli (Weise, 1906)

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Distribution: 
Côte d’Ivoire, Liberia

Henosepilachna fulvosignata (Reiche, 1847)

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Distribution: 

Côte d’Ivoire

Henosepilachna moseri (Weise, 1903)

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Distribution: 
Equatorial Guinea

Henosepilachna reticulata (Olivier, 1791)

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Distribution: 
Benin, Mali, Niger, Nigeria, Senegal

Henosepilachna simplex (Weise, 1895)

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Distribution: 
Liberia

Solanophila canina (Fabricius, 1781)

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Distribution: 
Guinea

Solanophila dregei (Mulsant, 1850)

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Distribution: 
Côte d’Ivoire

Solanophila scalaris (Gerstäcker, 1871)

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Distribution: 
Tanzania

Hyperaspis aestimabilis Mader, 1955

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Distribution: 
Angola, DRC

Hyperaspis centralis Mulsant, 1850

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Distribution: 
Mexico

Hyperaspis delicatula (Mulsant, 1850)

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Distribution: 
Benin, Gambia, Ghana, Guinea-Bissau, Nigeria, Malawi

Hyperaspis lugubris (Randall, 1838)

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Distribution: 
Ghana, Nigeria

Hyperaspis maindroni (Sicard, 1929)

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Distribution: 
Mauritania, Niger, Senegal

Hyperaspis merckii (Mulsant, 1850)

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Distribution: 
Mauritania, Senegal

Hyperaspis pumila Mulsant, 1850

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Distribution: 
Gambia, Guinea, Guinea-Bissau, Niger, Nigeria, Senegal, Togo

Hyperaspis senegalensis (Mulsant, 1850)

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Distribution: 

Gambia, Ghana, Nigeria, Senegal, Sierra Leone, Malawi

Hyperaspis sericea Fürsch, 1972

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Distribution: 
Malawi

Hyperaspis vinciguerra Capra, 1929

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Distribution: 
Gambia, Senegal, Malawi

Tenuisvalvae notata (Mulsant, 1850)

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Distribution: 
Benin, Bolivia

Ortalia ovulum Weise, 1898

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Distribution: 
Liberia, Mali, Togo

Rodolia cardinalis (Mulsant, 1850)

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Distribution: 
Kenya

Rodolia iceryae Janson in Ormerod, 1887

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Distribution: 
Senegal

Rodolia occidentalis Weise, 1898

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Distribution: 
Benin, Ghana, Nigeria, Senegal

Rodolia senegalensis Weise, 1913

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Distribution: 
Senegal

Platynaspis capicola Crotch, 1874

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Distribution: 
DRC

Platynaspis ferruginea Weise, 1895

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Distribution: 
Benin, Togo

Platynaspis kollari Mulsant, 1850

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Distribution: 
Liberia

Platynaspis obscura Gorham, 1901

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Distribution: 
Côte d’Ivoire, Liberia

Platynaspis pilosa Sicard, 1930

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Distribution: 
South Africa

Platynaspis rufipennis Gerstäcker, 1871

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Distribution: 
Côte d’Ivoire, Liberia, Niger

Platynaspis vittigera Weise, 1895

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Distribution: 
DRC

Pharoscymnus sexguttatus (Gyllenhall, 1808)

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Distribution: 

Ghana

“Leis” “maculata”

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Distribution: 
Côte d’Ivoire

Analysis

The taxonomically updated list of coccinellid species, present in the IFAN and IITAB collections, includes 129 species, representing 40 genera assigned to 11 tribes and two subfamilies following the classification of Seago et al. (2011). A total of 751 West African coccinellid specimens was recorded for the two collections. Of those, 385 specimens (68 spp., 30 genera) are deposited at IITA, while 366 specimens (84 spp., 31 genera) are at IFAN Table 1.

Most specimens (62%) were curated under currently valid names; however, 38% of specimens were labelled using junior synonyms. At IITA, 83% of the specimens were labelled using currently valid species names, while at IFAN, 39% of specimens were labelled using valid names.

Five genera comprise 57% of the specimens: Exochomus (6%), Chnootriba (9%), Scymnus (13%), Cheilomenes (14%) and Hyperaspis (15%) (Fig. 3). Twenty-five (of 40) genera each represent less than 1% of the total specimens. The remaining genera (Chilocorus, Declivitata, Epilachna, Henosepilachna, Nephus, Platynaspis, Rodolia, Stethorus and Xanthadalia) each account for between 2 and 5% of the overall specimen total.

Figure 3.  

Generic-level specimen representation in the IFAN and IITAB Collections.

Figure 4.  

Geographic source of coccinellid specimens in the IFAN Collection.

Figure 5.  

Geographic source of coccinellid specimens in the IITAB Collection.

Cheilomenes lunata (Fabricius, 1775), Cheilomenes propinqua (Mulsant, 1850), Cheilomenes sulphurea (Olivier, 1791), Chnootriba elaterii (Rossi, 1794), Chnootriba similis (Thunberg, 1781), Exochomus laeviusculus Weise, 1909, Hyperaspis delicatula (Mulsant, 1850) and Hyperaspis pumila Mulsant, 1850, are the most abundant species in the collections (Fig. 6). Specimens identified as Scymnus sp. make up the third most numerous group appearing in these collections.

Figure 6.  

Specific-level specimen representation of coccinellid holdings of the IFAN and IITAB Collections.

Geographic distribution

The coccinellid holdings in these two collections originated in 35 countries with 85% of specimens coming from West African countries, 14% coming from other African countries (DRC, Gabon, Madagascar, Malawi, Mozambique, Rwanda, Tanzania and Zambia) and 1% from non-African countries. More than half (66%) of West African material, housed in these two collections, came from just five countries: Senegal (27%), Nigeria (26%), Benin (5%), Liberia (4%) and Côte d'Ivoire (4%).

West African specimens housed in the IFAN museum were collected from 22 African countries. Most of these specimens (77%) were from five countries: Senegal (49%), Liberia (8%), Côte d’Ivoire (8%), Guinea (6%) and Mali (6%) (Fig. 4). The coccinellid specimens in the IITA originated in 25 countries, including 10 non-West African countries and four non-African countries (Bolivia, Columbia, Guatemala and Mexico). Most lady beetle specimens in the IITA museum (72%) were collected from five West African countries: Nigeria (50%), Benin (6%), Gambia (5%), Ghana (5%) and Senegal (5%). Two non-West African countries were represented: DRC (5%) and Malawi (5%) (Fig. 5).

Temporal distribution

Coccinellid material in the IFAN and IITA insect collections differ in temporal coverage (IFAN: 1900–1994; IITAB: 1950–2009) (Fig. 7). In the IFAN collection, the oldest specimens were all collected in 1900 and represent the following species: Adalia bipunctata (Linnaeus, 1758), Anisolemnia decempustulata Weise, 1888, Aulis annexa Mulsant, 1850, Brumoides foudrasii (Mulsant, 1850), Caria welwitschii Crotch, 1874, Cheilomenes aurora Gerstäcker 1871, Cheilomenes lunata (Fabricius, 1775) and Chnootriba elaterii (Rossi, 1794). A single specimen of Stethorus aethiops Weise, 1899, collected in 1950, represents the oldest coccinellid record in the IITAB collection.

Figure 7.  

Temporal distribution of IFAN and IITAB collection of coccinellids.

Both collections show a spike in growth of coccinellid holdings during one decade, but not the same one (Fig. 7). While 17% of the IFAN WAC specimens were collected between 1900 and 1944, the great majority (73%) were collected between 1945 and 1954. Only 5% of the specimens were added between 1955 and 1994. No new coccinellid material has been added since 1994.

The IITAB WAC records indicate that 6% of specimens were collected between 1950 and 1979, 80% from 1980 to 1989 and 14% between 1990 and 2009. No new coccinellid material was added after 2009.

Discussion

Data records compiled from collection labels in the IFAN and IITAB insect collections show that both collections combined provide an historical record of West African coccinellid diversity spanning over a century. It is clear that much coccinellid diversity in this region remains unrecorded though.

Very little published information is available about African coccinellids. Fürsch (1992) reports 70 species from Western Uganda. In Algeria, 75 species were recorded (Lakhal et al. 2018). The West African region, with its surface area of 6,140,000 km2, is nearly 26 times the size of Uganda (236,040 km2) and more than twice the size of Algeria.

West Africa, with its diverse ecosystems, landscapes, bioclimatic regions and vegetation (desert, rain forest, savannah), should support one of the highest diversities of coccinellids in all of Africa. The current total of 129 known coccinellid species from West Africa is surprisingly low for such a heterogeneous region.

The two focal collections of this study, the largest biodiversity centres in West Africa, differ in their taxonomic coverage. The IFAN holds more West African coccinellid diversity (31 gen., 84 spp.) than IITA (30 gen., 68 spp.). One possible explanation for the higher taxonomic diversity at IFAN is that their coccinellid records span nearly a century (1900–1994) while records at IITA only range from 1950 to 2009. In addition, the holdings at IFAN were enhanced by many expeditions to other West African countries, especially Côte d’Ivoire, Liberia, Mali, Mauritania, Senegal and Togo (Paulian et al. 1983).

A species name can become invalid due to the discovery of an older valid name or due to subsequent reclassification of the species in a different genus. Even though there was more taxonomic diversity represented at IFAN, 61% of species names used in the collection have not been updated to the valid names used in the current classification. At IITAB, however, most coccinellid species names (83%) were current and valid. It should also be noted that the various researchers, who have served as curators of IFAN insect collection, were taxonomists. Even though their expeditions and fieldwork focused on insect biodiversity in general, their efforts were concentrated on their respective specialities. These researchers each left Africa after some time and were no longer involved in the curation of these collections (e.g. André Villiers: 1945–1956, Michel Condamin: 1950–1973 and 1978–1988; Roger Roy: 1958–1992, Bernadette Soltani: 1988, Aïssatou Dramé: 1988–1991, Sun Heat Han: 1992–1996) (A. Niang pers. comm.). These are some of the potential reasons why the taxonomy of the coccinellid holdings at these museums was not current.

The IITA arthropod collection plays a crucial taxonomic role by providing essential, authoritative insect identifications amongst other services (e.g. biodiversity monitoring, pest management control etc.). IITA research has contributed to the description of more than 120 arthropod species (Ortiz 2017). This position of the IITA helps to explain why the identifications of its WAC specimens are more current: the IITAB WAC collection is newer than the one at IFAN.

Considering the numbers of specimens, Exochomus (6%), Chnootriba (9%), Scymnus (13%), Cheilomenes (14%) and Hyperaspis (15%) are the most strongly represented West African genera in the two collections. Cheilomenes lunata, Cheilomenes propinqua, Cheilomenes sulphurea, Chnootriba elaterii and Chnootriba similis are the most commonly collected species. Whereas Cheilomenes lunata, C. propinqua and C. sulphurea are widespread aphid predators, Chnootriba elaterii and Ch. similis are serious herbivorous pests of major staple crops. All these species may have been collected more often because they are relatively large, more colourful than many other coccinellids in the region and are regularly occurring on many cultivated and wild plants. In contrast, the collection, preparation and identification of tiny, brown coccinellids, like Scymnus species, are more difficult and time consuming. Drab, minute coccinellids could have been abundantly collected in field samples, but might never have been prepared, identified and curated. As a result, these less conspicuous coccinellids could be greatly under-represented in museum holdings even though they might be very common and important in various agroecosystems.

Many predaceous species are represented in the holdings, such as Exochomus flavipes (Thunberg, 1781), Exochomus laeviusculus Weise, 1909, Stethorus jejunus Casey, 1899, Hyperaspis delicatula, H. pumila, Rodolia cardinalis (Mulsant, 1850) and Scymnus senegalensis Weise, 1913. Some of these species are poorly represented in these collections, but this is likely due to collection and preparation biases, rather than actual rarity in the region. Although these relative abundance numbers of specimens in the collections are not the result of systematic and long-term sampling efforts, the simple spatial and temporal records of occurrence for these species in the region provide important information that could facilitate entomological research and pest management programmes in the sub-region.

Records of material in both IFAN and IITAB show that more than 60% of WAC specimens were collected from five countries (Senegal, Nigeria, Benin, Liberia and Côte d'Ivoire). These countries might have experienced more collecting effort because they either house the museums (Benin and Senegal) or because they are neighbouring countries where museum expeditions could be easily conducted (Côte d'Ivoire, Liberia, Nigeria). Benin, Côte d’Ivoire, Nigeria and Senegal have been agricultural research and trade centres in West Africa since the colonial period (Paulian et al. 1983). IFAN (Senegal), IITA (Benin, Nigeria) and ORSTOM (Côte d’Ivoire) were originally established to promote scientific research before the 1950s. Studies conducted through those organisations have continued to be published in more recent years (Chazeau and Couturier 1985, Fürsch 1991a, Sæthre et al. 2011), generating specimens for the museums.

The collecting efforts that built these museum holdings were haphazard, not the result of long-term, systematic monitoring efforts in the region. More than 23% of IITAB WAC specimens were collected from the IITA Station in Ibadan, Nigeria, while 17% of IFAN WAC were collected in Dakar, Senegal. Despite the high historical value of both collections, the geographic record is uneven. Some countries were far more heavily sampled over the years than others.

The coccinellid material housed in both collections is diverse, but there is a surprising lack of overlap in taxa between the two collections, even though they are in neighbouring countries that have similar ecological habitats. At IFAN, there are 56 WAC species that are not present in IITAB. There are 35 WAC species represented in the IITAB holdings that are not found in the IFAN collection. The lack of overlap could be due to collecting biases in the projects or expeditions that occurred at each institution. A large percentage of taxa represented in these museums was collected by only a few individuals (Fig. 3) which is consistent with that scenario. In fact, the combined WAC diversity at these two collections (40 genera, 129 species), represents only 75% of genera and 50% of the species already known to occur in the region as reported in the literature (KH). This lack of overlap between collections may be due in part from being at an early stage of discovery and collection development for WAC diversity. It is clear that there is a need for much more thorough study and sampling in order to accurately assess the diversity of this important group across West Africa.

It is noteworthy that both collections include African coccinellids from outside West Africa. The IITAB also has material from Central and South America (Bolivia, Columbia, Guatemala and Mexico). These non-West African specimens were probably received as exchanges between international collaborators who were conducting general systematics research or were collaborating with the various Insect Pest Management programmes carried out by IITA. These collaborations with researchers from around the world might help to explain why the identifications of material at IITAB were taxonomically more current than at IFAN. In fact, most of the IITAB coccinellids were identified by a German researcher, Helmut Fürsch, a taxonomic authority of Afrotropical Coccinellidae.

The IFAN and IITA WAC collections each show a history that is marked by three distinct periods of development. For IFAN, these significant periods occurred in 1900-1944, 1945-1954 and 1955-1994. IFAN holds some material that predates the establishment of the insect collection in the 1940s. This older material (collected 1900-1944) came from many European collections and collectors (e.g. P. Daget, E. Fleutiaux, M. Griaule, T. Jackson, Delattre, H. Junod, A. Vuillet etc.). In 1945, André Villiers established the IFAN entomology section and began to organise many expeditions to African locations, including Cameroon (1939, 1951), Senegal (1945–1956), Casamance, Senegal (1946), Mali (1946, 1947), Guinea (1946, 1954), Côte-d'Ivoire (1946, 1955), Guinea Bissau (1947), Aïr Mountains, Niger (1947), Senegalese Ferlo (1948, 1950), Mauritania (1948–1953), southern Nigeria (1949), Benin and Togo (1950), southern Togo (1950), Sudan region (border of Senegal with South of Sahara), Fernando-Poo Island, Equatorial Guinea (1951) and Niokolo-Koba National Park, Senegal (1955, 1956). These field trips and Villiers’ collaborations with a network of foreign entomologists resulted in a decade (1945–1954) during which most of IFAN WAC specimens (73% of the total WAC holdings) were collected. The period of rapid growth of WAC holdings at IFAN ceased after Villiers returned to the Muséum National d'Histoire Naturelle (Paris) in 1956. Although he organised many subsequent trips to Africa between 1961 and 1977, that was a period of great change in the region. In the 1960s, most West African countries became independent and experienced major transitions and restructuring of administration.

The noteworthy periods of development for the IITAB WAC collection were 1950–1979, 1980–1989 and 1990–2009. Not surprisingly, IFAN’s slower growth in the 1960s coincided with the lowest rate of growth for IITA’s new collection. In the late 1970s, however, there were major pest outbreaks (e.g. Maize streak virus, cassava mealybug, cassava green mite, mango mealybug, fruit tree mealybug etc.) that led IITA scientists to establish collaborative integrated pest management programmes with Central and South American researchers at the International Center for Tropical Agriculture (CIAT) (Agounké et al. 1988, Ortiz 2017). These biological control programmes were very successful in importing, rearing and releasing natural enemies to manage these pests (Agounké et al. 1988, Ortiz 2017). During research trials for those projects, insects were sampled from agricultural lands. In fact, more than 40% of WAC specimens in the IITA museum were collected between 1979 and 1989 on cassava alone. It is likely that many coccinellids in the IITA collection were collected because the 1970s pest outbreaks pushed the institution to establish partnerships with stakeholders, international researchers and other African research institutions. From 1990 to the present, records from the IITA museum show a very significant decrease in the number of coccinellids collected. This decreased rate of growth could be explained by the spectacular success of the biological pest control programme carried out by IITA on cassava, mango trees and other crops, thereby reducing the need for field sampling. In addition to the identified material referred to in the present paper, there is still a huge backlog of unidentified coccinellid specimens at IITAB. The non-WAC specimens obtained at that same time are another benefit from those same events and resulting collaborations.

Despite the success of IPM programmes and taxonomic expeditions led by IFAN and IITA in many African countries, it is clear that some groups of West African coccinellid genera with high known diversity were poorly sampled (e.g. species of Adalia, Anisolemnia, Clitostethus, Coccinella, Diomus, Megalocaria, Micraspis, Nephus, Psyllobora, Rodolia, Scymnus etc.). For example, 22 species of Nephus have been reported to occur in West Africa (KH), yet only 6 are represented in these collections.

This gap in taxonomic knowledge about lady beetles mirrors the situation seen in many other insect taxa in West Africa. The assessment of biodiversity in the region has been hampered historically by a lack of local taxonomic expertise, inaccessibility of scientific literature, rarity of reliable arthropod reference collections, limited scientific infrastructure and a lack of financial resources. Recent advances in systematics, especially in “cybertaxonomy,” now provide web-based taxonomic tools, diverse publication outlets and easy access to a wealth of digitised scientific resources including technical literature, high quality photographs, specimen data etc., thereby reducing the taxonomic impediment for researchers in places like West Africa. If coupled with strategic development of international, institutional collaborations to conduct biodiversity surveys and inventory projects, great progress could be made towards filling large taxonomic and geographical gaps in our knowledge of West African insects.

Acknowledgements

This work was made possible through the assistance of numerous volunteers and staff at various institutions. We are grateful to the following people for their kind assistance: Kyky Komla Ganyo (CERAAS/UCAD), team members of Grain de Sel Togo, Inc., Mawufe Agbodzavu and Mawuko Sokame (ICIPE), Mouhamadou M. Ndiaye (IFAN), Ghislain Tepa-Yotto and Francis Tchibozo (IITA), Philip Perkins (MCZ), Muriel Van Nuffel and Eliane De Coninck (MRAC), Floyd W. Shockley (NMNH), Ibrahima Dia (UEM/IPD), Demba Sarr (UGA), Edward R. Hoebeke (UGCA) and Johannes Frisch and Bernd Jaeger (ZMHB). We also thank Wioletta Tomaszewska and anonymous reviewers for their helpful comments and criticism on an earlier version of this manuscript.

This study was done in partial fulfillment of the senior author’s Ph.D. dissertation requirements at the University of Georgia. The senior author wishes to thank his advisory committee members K.G. Ross, M.D. Toews and J.V. McHugh.

This project was supported by the following grants: Borlaug LEAP grant #016258-97 (to KH), H.H. Ross Fund grant (to KH), Fulbright Foreign Student Program Scholarship from Institute of International Education (to KH), Global Programs Graduate International Travel Award from the University of Georgia (to KH), West African Research Center (WARC) Travel Grant (to KH) and the Department of Entomology at the University of Georgia.

References

Endnotes
*1

This record is based on two specimens collected by R. Borowka on 12 AUG 1992 Citrus mussel scale in Chipata, Zambia. They were originally identified as “Chilocorus serangium Kunowi”.

*2

This was treated as subspecies of Exochomus nigrifrons Gerstäcker, 1871 by Mader (1954: 88) (Brumus nigrifrons nigerianus Korschefsky). Brumus nigrifrons nigerianus was misspelled. Species of the genus Brumus Mulsant, 1850 were transferred to Exochomus Redtenbacher, 1843.

*3

This record is based on a single specimen determined as “Cydonia intermedia Cramer” by Mader. The holotype for this species is from Abyssinia, now Ethiopia.

*4

This is probably a nomen nudum.