Abstract 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.
Table 1.
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 |
Checklists
Checklist of West African Coccinellidae in IFAN and IITAB museums (see Table 1 for more details)
Serangium kunowi
Weise, 1892
6C08AE98-8273-5ED6-90CC-F87BBC52F548
Distribution
Zambia
Brumoides foudrasii
(Mulsant, 1850)
F145D8A6-249B-5FE8-B59E-2F1B70C4CE19
Distribution
Benin, Gambia, Guinea, Nigeria, Senegal
Chilocorus distigma
(Klug, 1835)
987BB720-000F-53CD-8C9D-D5F81C0C0A0E
Distribution
Mozambique
Chilocorus dorhni
Mulsant, 1850
611CE9A1-C8DB-5C3A-A1A9-AEFDC24B20DA
Distribution
Senegal
Chilocorus schioedtei
Mulsant, 1850
5C66671C-194C-51EE-8692-7949F89AEC69
Distribution
Benin, Cameroon
Chilocorus simoni
Sicard, 1907
E283D897-7362-5CE2-8821-FC62B886D75D
Distribution
South Africa
Exochomus flavipes
(Thunberg, 1781)
2FBF8BEF-EB91-561E-A288-6E2885FA8404
Distribution
Gabon
Exochomus laeviusculus
Weise, 1909
7A6E4183-D702-56C1-A936-89A0CD234694
Distribution
Benin, Côte d’Ivoire, Guinea, Mali, Mauritania, Senegal, Togo
Exochomus nigrifrons
Gerstäcker, 1871
A0B866A5-E30C-5C9D-94E9-7BB968407E2E
Distribution
Mali, Senegal
Parexochomus nigripennis
(Erichson, 1843)
82867BBC-4337-5664-A99D-98112C1AFBC4
Distribution
Mali, Senegal
Exochomus pulchellus
Gerstäcker, 1871
79E7BA08-3F1C-5E17-8503-DB5E015151E6
Distribution
Gambia, Niger, Rwanda, Senegal
Exochomus troberti
Mulsant, 1850
7D78E719-1F19-5868-A5EA-F2E3B07E46CF
Distribution
Burkina Faso
Aulis annexa
Mulsant, 1850
9D2EAB15-7F2A-50C7-B256-3E49AC7395DA
Distribution
Senegal
Clitostethus flavotestaceus
Mader, 1955
DBD81906-B263-5595-8373-5D2DFA36624F
Distribution
Senegal
Nephus flavomaculatus
Fürsch, 1966
5B760B86-9AF4-5EDD-BF4D-B9BBB9A99371
Distribution
Benin, Nigeria
Nephus vetustus
Weise, 1915
357A9314-78D6-52AD-81D3-032870015377
Distribution
Gabon
Nephus phenacoccophagus
Fürsch, 1987
A72D8A23-7357-59FE-9037-0E22ACE44349
Distribution
Nigeria
Nephus kamburovi
Fürsch, 1992
4BA9A0A8-ECD4-5017-B0DB-602EF59323B0
Distribution
Malawi
Nephus oblongosignatus
Mulsant, 1850
D324C8AD-A466-5D0D-9AF9-71FDDE06243D
Distribution
Tanzania
Nephus ornatulus
Korschefsky, 1931
7A6DD1B1-6B22-5268-B455-C5D3AE0F3FB8
Distribution
DRC
Nephus sudanicus
Weise, 1925
3BFEBA83-41A1-55C9-9D2F-B724A8F97EEF
Distribution
Mauritania
Scymnobius bilucernarius
(Mulsant, 1850)
F4DF6361-FEBD-5CE8-B642-8189CA31C54B
Distribution
Mexico
Scymnus canariensis
Wollaston, 1864
D8670A86-01D1-5BA1-AAC1-C2B8EC3FE969
Distribution
São Tome and Principe, Senegal
Scymnus casstroemi
Mulsant, 1850
1B350FE1-0669-5324-A7EB-930E943913AA
Distribution
Guinea, Senegal
Scymnus gnavus
Weise, 1895
E054571A-7356-56D5-BE4E-6B94C2B09653
Distribution
Guinea
Scymnus kibonotensis
Weise, 1910
44A344EB-1700-553A-BA65-7BFC9EC9A047
Distribution
Côte d’Ivoire, Guinea, Nigeria
Scymnus levaillanti
Mulsant, 1850
81A41B21-DE38-5E9E-A631-FB762FFF45DF
Distribution
Nigeria, Malawi
Scymnus pruinosus
Weise, 1895
E43D7300-A485-50A7-A955-4845D1EA219A
Distribution
Zambia
Scymnus monroviae
Casey, 1899
2FCF8766-B290-5EE8-8720-162B04FC155C
Distribution
Benin, Côte d’Ivoire, Guinea, Niger, Senegal, Togo
Scymnus nigrosellatus
Mader, 1950
63025A4B-B79C-5181-82B7-CF554921EDEC
Distribution
Zambia
Scymnus quadrivittatus
Mulsant, 1850
9738E438-BFE1-51B5-A1C7-67A1F8E4DF58
Distribution
Nigeria
Scymnus rubiginosus
Mader, 1950
7D91E966-C223-5568-8A88-898FEA4CC113
Distribution
Côte d’Ivoire, Guinea, Senegal
Scymnus scapuliferus
Mulsant, 1850
D1BBC229-5A5A-5C02-9D47-745AC3692421
Distribution
Benin, Côte d’Ivoire, Guinea, Nigeria, Madagascar, Senegal, Togo
Scymnus schoutedeni
Mader, 1950
99E145E3-EC00-5746-AAA7-74C9F1D031B2
Distribution
Senegal
Scymnus senegalensis
Mader, 1955
E492343F-636F-54F2-A535-55D45B8E03BD
Distribution
Côte d’Ivoire, Gambia, Guinea, Mali, Mauritania, São Tome and Principe, Senegal
Scymnus villiersi
Mader, 1955
3CA780B0-326B-5355-BA7E-44D6A693023D
Distribution
Niger, Senegal
Stethorus aethiops
Weise, 1899
ACB57C94-B1ED-5044-9C74-1D78DA667457
Distribution
Benin, Ghana, Guinea-Bissau, Mozambique
Stethorus endroedyi
Fürsch, 1970
4F6EC29C-B255-5DB5-9C5A-B38A91AEF55B
Distribution
Malawi
Stethorus jejunus
Casey, 1899
0EA19E48-8314-5F7B-8E8B-E797B0C48599
Distribution
Ghana, Nigeria, Mozambique
Adalia bipunctata
(Linnaeus, 1758)
6AB2552E-C104-5828-998E-01858BDCCFEC
Distribution
Cameroon
Anisolemnia decempustulata
Weise, 1888
8740EBD4-5BD2-5CFC-9201-EF9F5539BC4A
Distribution
Togo
Bulaea anceps
(Mulsant, 1850)
F830F859-02F9-5B71-91D5-A402E4D1217A
Distribution
Mozambique
Caria welwitschii
Crotch, 1874
54F76728-8648-5E70-8053-6340CDCA98A7
Distribution
Guinea
Cheilomenes aurora
Gerstäcker, 1871
5C30D7D2-C9DD-54D7-BCB8-807AF27F1E69
Distribution
Tanzania
Cheilomenes lunata
(Fabricius, 1775)
039ACC42-54E5-5D9A-A850-C7A70EBCE416
Distribution
Benin, Burkina Faso
Cheilomenes sulphurea
(Olivier, 1791)
38306156-309E-5BA4-9296-14E9F6B99C49
Distribution
Angola, Cameroon
Cheilomenes propinqua
(Mulsant, 1850)
43D175AD-0B9F-500C-8B15-92160DB6C9B3
Distribution
Côte d’Ivoire, Gabon
Cheilomenes quadrilineata
(Mulsant, 1850)
B08456B4-BCB2-5E03-9998-A16257FCD339
Distribution
Senegal
Coccinella intermedia
(Crotch, 1874)
B8500504-FED1-5405-8BAE-650EFE34F58D
Distribution
São Tome and Principe
Coccinella septempunctata
Linnaeus, 1758
11792B51-197D-5D4E-88C3-8A1CD7677D44
Distribution
Cape Verde
Declivitata hamata
(Thunberg, 1808)
8104F430-F761-5685-B0DA-BE91FF3D2B94
Distribution
Senegal
Declivitata uncifera
Fürsch, 1967
9AB6D0F0-4055-587B-89A5-333EE7FE78C7
Distribution
Cameroon
Harmonia vigintiduomaculata
(Fabricius, 1792)
A22708E4-F70C-584C-BAF7-AB0EB9AECB94
Distribution
Benin, Liberia, Nigeria, Togo
Hippodamia variegata
(Goeze, 1777)
335DC6A9-8C00-5153-B368-131F70AFD80C
Distribution
Colombia, Nigeria, Senegal, Tanzania
Lemnia machadoi
Mader, 1952
D8C4A968-F1B7-5CBC-8E09-3D5840269877
Distribution
Cameroon
Megalocaria dilatata
(Fabricius, 1775)
917472C2-D77D-5B71-821A-698C574109B4
Distribution
Benin
Micraspis lineola
(Fabricius, 1775)
C64F6216-086F-5E31-B90F-90F84ADFCC7A
Distribution
Togo
Micraspis striata
(Fabricius, 1792)
B32DF781-9B4D-5549-8D88-C504FB949886
Distribution
Côte d’Ivoire, Gabon
Psyllobora bisoctonotata
(Mulstant, 1850)
CC91E27C-F108-51AC-B82F-7D266005964E
Distribution
Senegal
Psyllobora lutescens
(Crotch, 1874)
9E9682D5-3020-5A68-B710-5F8F9DE7F939
Distribution
Guatemala
Psyllobora variegata
(Fabricius, 1781)
036CE85C-635D-56D7-B4A9-2A0DEF6586DA
Distribution
South Africa
Xanthadalia effusa
(Erichson, 1843)
921AEBB7-F849-57B5-A109-FB7E9CF46487
Distribution
Benin, DRC
Xanthadalia rufescens
Mulsant, 1850
E376E7EB-580B-562B-B482-66AB41987CAD
Distribution
Benin, Mali, Mauritania, Senegal
Diomus hennesseyi
Fürsch, 1987
A7DE0649-BBBC-596D-BCAB-018BAAD32D50
Distribution
Nigeria
Chnootriba elaterii
(Rossi, 1794)
491F19F1-D594-5926-9780-01BB715C3F53
Distribution
Benin, Côte d’Ivoire, Gambia, Guinea, Liberia, Mali, Mauritania, Nigeria, São Tome and Principe, Senegal
Chnootriba hirta
(Thunberg, 1781)
2AA87447-7A03-5FB5-9A30-2FCC5A71CD1A
Distribution
Guinea, Tanzania
Chnootriba similis
(Thunberg, 1781)
C9DE0E27-4C4D-5753-9B16-5BC11A21F09D
Distribution
Benin, Burkina Faso
Cleta punctipennis
(Mulsant, 1850)
7EEC920E-FDA5-502A-9255-AACD0D177DCE
Distribution
Togo
Cleta sahlbergi
(Mulsant, 1850)
053E7477-493C-50EA-A6F3-AB75BF474A4F
Distribution
Côte d’Ivoire, Kenya
Epilachna bissexguttata
Weise, 1895
A8FC50D7-2455-5275-A2E5-EC0EB5BA9B3A
Distribution
Côte d’Ivoire, DRC
Epilachna bomparti
Mulsant, 1850
7DBB54EC-77C6-515A-AC5C-8D676E6E294D
Distribution
Liberia, Senegal
Epilachna colorata
Mulsant, 1850
462C8061-1250-5D39-A245-B2A973EC8EF8
Distribution
Cameroon
Epilachna iocosa
(Mader, 1941)
169A3C8E-B305-528B-AB91-C49F2E6EAA5C
Distribution
South Africa
Epilachna nigritarsis
Mulsant, 1850
5B67661B-0EF8-594A-ABCC-FC704A942111
Distribution
Cameroon
Epilachna vigintipunctata
Mulsant, 1850
A68D6FAC-0F18-58E7-94F6-75C3167514D3
Distribution
Liberia, Tanzania
Henosepilachna atropos
(Sicard, 1912)
846C306C-466E-5593-BFFA-5AEB9B66C8C7
Distribution
Equatorial Guinea
Henosepilachna bisseptemnotata
(Mulsant, 1853)
7498AF38-2BFF-51C4-B1F2-579CC58DC36B
Distribution
Tanzania
Henosepilachna clavareaui
(Weise, 1901)
3E22E012-F4C3-54E2-A68A-67148C5BA9B9
Distribution
Benin
Henosepilachna ertli
(Weise, 1906)
5311A71F-2F85-54DF-8655-261A24D2A557
Distribution
Côte d’Ivoire, Liberia
Henosepilachna fulvosignata
(Reiche, 1847)
BAEFAE7F-C8D1-555A-A0FE-FBEC61AC0C5C
Distribution
Côte d’Ivoire
Henosepilachna moseri
(Weise, 1903)
CB309483-53C2-5A92-8435-760FC9FC9233
Distribution
Equatorial Guinea
Henosepilachna reticulata
(Olivier, 1791)
D62E1FD2-D258-5916-9EF7-CF6392EA5AE5
Distribution
Benin, Mali, Niger, Nigeria, Senegal
Henosepilachna simplex
(Weise, 1895)
62A58A88-ABD5-538B-B541-73DA01945900
Distribution
Liberia
Solanophila canina
(Fabricius, 1781)
8B2CA986-4056-5CD1-99E1-C95064859686
Distribution
Guinea
Solanophila dregei
(Mulsant, 1850)
7F669D7A-815E-59FA-8089-C8E3F350AFE2
Distribution
Côte d’Ivoire
Solanophila scalaris
(Gerstäcker, 1871)
6DE54F2B-3C49-53B2-AA36-5A177259740A
Distribution
Tanzania
Hyperaspis aestimabilis
Mader, 1955
80C4E12E-B249-57BD-AA30-EE2211E4D6B5
Distribution
Angola, DRC
Hyperaspis centralis
Mulsant, 1850
475B8890-4115-511A-AB08-2665D8EFF7F5
Distribution
Mexico
Hyperaspis delicatula
(Mulsant, 1850)
E8896E7D-4093-5429-9D43-577243C41EE8
Distribution
Benin, Gambia, Ghana, Guinea-Bissau, Nigeria, Malawi
Hyperaspis lugubris
(Randall, 1838)
C5DD9997-6D2F-52C6-AFE0-5BF69639E608
Distribution
Ghana, Nigeria
Hyperaspis maindroni
(Sicard, 1929)
FFB7EE79-1179-56DC-939C-03827DAA19F6
Distribution
Mauritania, Niger, Senegal
Hyperaspis merckii
(Mulsant, 1850)
6D4A602C-4D50-5731-A1B2-A04B4B382279
Distribution
Mauritania, Senegal
Hyperaspis pumila
Mulsant, 1850
EEE926FA-E163-5A5E-B24A-8E187F04874F
Distribution
Gambia, Guinea, Guinea-Bissau, Niger, Nigeria, Senegal, Togo
Hyperaspis senegalensis
(Mulsant, 1850)
C2267008-C788-552B-8DA3-52EB950FE9DA
Distribution
Gambia, Ghana, Nigeria, Senegal, Sierra Leone, Malawi
Hyperaspis sericea
Fürsch, 1972
4007B2E6-E527-5021-B877-DA72F665FABC
Distribution
Malawi
Hyperaspis vinciguerra
Capra, 1929
FA4C8459-72D9-5A29-8E62-ED272E16E384
Distribution
Gambia, Senegal, Malawi
Tenuisvalvae notata
(Mulsant, 1850)
A8D79B2C-99B8-5003-87EE-440F982C3128
Distribution
Benin, Bolivia
Ortalia ovulum
Weise, 1898
75D4E5F3-20DA-557A-A8B5-CCE07B685A8C
Distribution
Liberia, Mali, Togo
Rodolia cardinalis
(Mulsant, 1850)
53162E98-068F-5245-8F30-FFBB6C292FA8
Distribution
Kenya
Rodolia iceryae
Janson in Ormerod, 1887
8B18A899-5B4C-5D0C-9388-9CC83F7AE0EE
Distribution
Senegal
Rodolia occidentalis
Weise, 1898
0436B09A-DA2A-591D-9C96-114C54EFBD50
Distribution
Benin, Ghana, Nigeria, Senegal
Rodolia senegalensis
Weise, 1913
837B3D0F-00B4-5CBE-970A-C9F92DD91DA8
Distribution
Senegal
Platynaspis capicola
Crotch, 1874
01612FAC-08A3-59B6-AAFD-488C1968F5F0
Distribution
DRC
Platynaspis ferruginea
Weise, 1895
59D78062-FFAA-545D-A636-942A48B43856
Distribution
Benin, Togo
Platynaspis kollari
Mulsant, 1850
5D3B29FC-524D-5FDF-AFB5-A8BF1385A688
Distribution
Liberia
Platynaspis obscura
Gorham, 1901
7A2101B4-7A6D-5847-952D-56F0B403E16A
Distribution
Côte d’Ivoire, Liberia
Platynaspis pilosa
Sicard, 1930
2B97EBBA-2DFC-5408-805D-1C8A61FE1EB5
Distribution
South Africa
Platynaspis rufipennis
Gerstäcker, 1871
F05691A6-54E1-5457-B8F3-FE32316291F4
Distribution
Côte d’Ivoire, Liberia, Niger
Platynaspis vittigera
Weise, 1895
027F9CEB-94FF-5737-A613-16DA63A24284
Distribution
DRC
Pharoscymnus sexguttatus
(Gyllenhall, 1808)
3357EFFA-9D97-513E-A7ED-3E536B63A400
Distribution
Ghana
“Leis” “maculata”
F9412911-A954-57F8-9927-21B56604D202
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.
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).
Figure 4.
Geographic source of coccinellid specimens in the IFAN Collection.
Figure 5.
Geographic source of coccinellid specimens in the IITAB Collection.
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.
Footnotes
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”.
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.
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.
This is probably a nomen nudum.
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