Skip to main content Skip to main navigation menu Skip to site footer
Article
Published: 2021-10-14

Towards the identification of the scale insects (Hemiptera: Coccomorpha) of continental Africa: 1. Identification of the families

Department of Life Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.
ANSES, Laboratoire de la Santé des Végétaux, Unité entomologie et plantes invasives, 755 avenue du campus Agropolis, CS 30016, FR-34988 Montferrier-sur-Lez Cedex, France.
Hemiptera identification key to families Sternorrhyncha

Abstract

Scale insects (Hemiptera: Sternorrhyncha: Coccomorpha) are obligate plant parasites feeding on plant sap; some are damaging pests in agriculture, horticulture and forestry. Despite their economic importance, the scale insects found in continental Africa have not been extensively studied and the keys for identifying them are incomplete and scattered through the literature in several languages. The aim of this study is to improve our understanding of the African scale insect fauna. As a first step towards their identification, we provide a key to the 23 families currently known from continental Africa, based on slide-mounted adult females, covering Aclerdidae, Asterolecaniidae, Cerococcidae, Coccidae, Conchaspididae, Dactylopiidae, Diaspididae, Eriococcidae, Halimococcidae, Kermesidae, Kerriidae, Kuwaniidae, Lecanodiaspididae, Margarodidae, Matsucoccidae, Micrococcidae, Monophlebidae, Ortheziidae, Phoenicococcidae, Pseudococcidae, Putoidae, Rhizoecidae and Stictococcidae.

 

References

  1. Balachowsky, A.S. (1956) Les cochenilles du continent africain noir. Vol. 1. Aspidiotini (1ère Partie). Annales du Musée royal du Congo belge, New Series, 3, 1–142.
    Balachowsky, A.S. (1958) Les cochenilles du continent africain noir. Vol. 2. Aspidiotini (2me Partie), Odonaspidini et Parlatorini. Annales du Musée royal du Congo belge, New Series, 4, 145–356.
    Ben-Dov, Y. (1977) New species of Aclerda Signoret (Homoptera: Aclerdidae) from southern Africa. Journal of Natural History, 11, 371–376. https://doi.org/10.1080/00222937700770281
    Ben-Dov, Y. (1988) A taxonomic analysis of the armored scale tribe Odonaspidini of the world (Homoptera: Coccoidea: Diaspididae). United States Department of Agriculture Technical Bulletin, 1723, 1–142.
    Bullington, S.W. & Kosztarab, M.P. (1985) Studies on the morphology and systematics of scale insects. No. 12.I. Revision of the family Kermesidae (Homoptera) in the Nearctic region based on adult and third instar females. Bulletin of the Virginia Polytechnic Institute and State University Agricultural Experiment Station, 85 (11), 1–118.
    Cave, R.D., Chao, J.-T., Kumashiro, B., Marler, T., Miles, J., Moore, A., Muniappan, R. & Watson, G.W. (2013) Status and biological control of Cycad Aulacaspis Scale. Biocontrol News and Information, 34 (1), 1N–8N.
    Cox, J.M. (1987) Pseudococcidae (Insecta: Hemiptera). In: Duval, C.T. (Series Ed.), Fauna of New Zealand. Vol. 11. DSIR Science Information Publishing Centre, Wellington, pp. 1–229. Available from: https://www.landcareresearch.co.nz/publications/books/fauna-of-nz-series/extracts/fnz11 (accessed 22 October 2020)
    Danzig, E.M. (1993) [Fauna of Russia and neighbouring countries. Rhynchota, Volume X: suborder scale insects (Coccinea): families Phoenicococcidae and Diaspididae.] Nauka Publishing House, St. Petersburg, 452 pp.
    Danzig, E.M. & Gavrilov-Zimin, I.A. (2014) Palaearctic mealybugs (Homoptera: Coccinea: Pseudococcidae), Part 1: Subfamily Phenacoccinae. Russian Academy of Sciences, Zoological Institute, St. Petersburg, 678 pp. 
    De Lotto, G. (1954a) Three apparently new coccids (Homopt.: Coccidae) from Eritrea. Journal of the Entomological Society of Southern Africa, 17, 213–218.
    De Lotto, G. (1954b) Three apparently new mealy bugs from Kenya. Proceedings of the Royal Entomological Society of London, 23, 110–114. https://doi.org/10.1111/j.1365-3113.1954.tb00111.x
    De Lotto, G. (1955) Three new coccids (Hemipt.: Coccoidea) attacking coffee in East Africa. Bulletin of Entomological Research, 46, 267–273. https://doi.org/10.1017/S000748530003090X
    De Lotto, G. (1956a) The identity of some East African species of Saissetia (Homoptera, Coccidae). Bulletin of Entomological Research, 47, 239–249.  https://doi.org/10.1017/S0007485300046691
    De Lotto, G. (1956b) A new Ceroplastodes (Hom.: Coccoidea) from Kenya. Journal of the Entomological Society of Southern Africa, 19, 310–312.
    De Lotto, G. (1957a) Notes on some African species of Saissetia (Homoptera: Coccoidea: Coccidae). Journal of the Entomological Society of Southern Africa, 20, 170–182.
    De Lotto, G. (1957b) On some Ethiopian species of the genus Coccus (Homoptera: Coccoidea: Coccidae). Journal of the Entomological Society of Southern Africa, 20, 295–314. 
    De Lotto, G. (1957c) The Pseudococcidae (Hom.: Coccoidea) described by H.C. James from East Africa. Bulletin of the British Museum (Natural History) Entomology, Supplement 5, 185–232.
    De Lotto, G. (1958a) A new species of Saissetia from South Africa (Homoptera: Coccoidea: Coccidae). Journal of the Entomological Society of Southern Africa, 21, 66–68.
    De Lotto, G. (1958b) New soft scales (Homoptera: Coccoidea: Coccidae) from Africa. Proceedings of the Royal Entomological Society of London (B), 27, 165–172. https://doi.org/10.1111/j.1365-3113.1958.tb00402.x
    De Lotto, G. (1958c) The Pseudoccoidae (Hom.: Coccoidea) described by C.K. Brain from South Africa. Bulletin of the British Museum (Natural History) Entomology, Supplement 7, 79–120.
    De Lotto, G. (1959a) Further notes on Ethiopian species of the genus Coccus (Homoptera: Coccoidea: Coccidae). Journal of the Entomological Society of Southern Africa, 22, 150–173.
    De Lotto, G. (1959b) Two new Margarodidae (Homopt.: Coccoidea) from Kenya. Journal of the Entomological Society of Southern Africa, 22, 385–389.
    De Lotto, G. (1960) The green scales of coffee in Africa south of the Sahara (Homoptera, Coccidae). Bulletin of Entomological Research, 51, 389–403. https://doi.org/10.1017/S0007485300058041
    De Lotto, G. (1961a) Two new Ceroplastes species from Africa (Homopt.: Coccidae). Journal of the Entomological Society of Southern Africa, 24, 318–321.
    De Lotto, G. (1961b) New Pseudococcidae (Homoptera: Coccoidea) from Africa. Bulletin of the British Museum (Natural History) Entomology, 10, 211–238. https://doi.org/10.5962/bhl.part.16262
    De Lotto, G. (1962) A new Coccus from South Africa (Homoptera: Coccidae). Journal of the Entomological Society of Southern Africa, 25, 263–265.
    De Lotto, G. (1963) New species and a new genus of hard scales from East Africa (Homoptera: Coccidae). Proceedings of the Royal Entomological Society of London (B), 32, 191–200. https://doi.org/10.1111/j.1365-3113.1963.tb01590.x
    De Lotto, G. (1964a) A new species of Pulvinaria (Homopt.: Coccidae) attacking sugar cane in South Africa. South African Journal of Agricultural Science, 7, 863–866.
    De Lotto, G. (1964b) Observations on African mealy bugs (Hemiptera: Coccoidea). Bulletin of the British Museum (Natural History) Entomology, 14, 343–397. https://doi.org/10.5962/bhl.part.8787
    De Lotto, G. (1964c) A new species of Mombasinia (Hom.: Pseudococcidae) from South Africa. Proceedings of the Linnean Society of London, 175, 135–136. https://doi.org/10.1111/j.1095-8312.1964.tb00926.x
    De Lotto, G. (1965) On some Coccidae (Homoptera), chiefly from Africa. Bulletin of the British Museum (Natural History) Entomology, 16, 175–239. https://doi.org/10.5962/bhl.part.21865
    De Lotto, G. (1966a) Descriptions of three new species of Coccus (Homoptera: Coccidae) from South Africa. Proceedings of the Royal Entomological Society of London (B), 35, 41–46. https://doi.org/10.1111/j.1365-3113.1966.tb00465.x
    De Lotto, G. (1966b) Another new species of Pulvinaria (Hom.: Coccidae) from sugar cane. South African Journal of Agricultural Science, 9, 467–472.
    De Lotto, G. (1966c) A new genus and four new species of Coccidae (Homoptera) from South Africa. Proceedings of the Linnean Society of London, 177, 143–149. https://doi.org/10.1111/j.1095-8312.1966.tb00956.x
    De Lotto, G. (1967a) The soft scales (Homoptera: Coccidae) of South Africa. I. South African Journal of Agricultural Sciences, 10, 781–810.
    De Lotto, G. (1967b) The mealybugs of South Africa (Homoptera: Pseudococcidae), I. Entomology Memoirs, Department of Agricultural Technical Services, Republic of South Africa. Pretoria, 12, 1–28.
    De Lotto, G. (1969a) On a few old and new soft scales and mealybugs (Homoptera: Coccoidea). Journal of the Entomological Society of Southern Africa, 32, 413–422. 
    De Lotto, G. (1969b) The mealybugs of South Africa (Homoptera: Pseudococcidae), II. Entomology Memoirs, Department of Agricultural Technical Services, Republic of South Africa. Pretoria, 20, 1–30.
    De Lotto, G. (1970) The soft scales (Homoptera: Coccidae) of South Africa, II. Journal of the Entomological Society of Southern Africa, 33, 143–156. 
    De Lotto, G. (1971) On some genera and species of wax scales (Homoptera: Coccidae). Journal of Natural History, 5, 133–153. https://doi.org/10.1080/00222937100770091
    De Lotto, G. (1974a) New species of Filippia Targioni-Tozzetti, 1868 (Homoptera: Coccoidea: Coccidae) from South Africa. Journal of the Entomological Society of Southern Africa, 37, 207–214. 
    De Lotto, G. (1974b) On the status and identity of the cochineal insects (Homoptera: Coccoidea: Dactylopiidae). Journal of the Entomological Society of Southern Africa, 37, 167–193.
    De Lotto, G. (1974c) On two genera of mealybugs (Homoptera: Coccoidea: Pseudococcidae). Journal of the Entomological Society of Southern Africa, 37, 109–115.
    De Lotto, G. (1975a) New species of Paracoccus Ezzat & McConnell, 1956. (Homoptera: Coccoidea: Pseudococcidae) from southern Africa. Journal of the Entomological Society of Southern Africa, 38, 65–73.
    De Lotto, G. (1975b) Two new genera of soft scales from Africa (Homoptera: Coccoidea: Coccidae). Journal of the Entomological Society of Southern Africa, 38, 61–63.
    De Lotto, G. (1978) The soft scales (Homoptera: Coccidae) of South Africa, III. Journal of the Entomological Society of Southern Africa, 41, 135–147.
    De Lotto, G. (1979) The soft scales (Homoptera: Coccidae) of South Africa, IV. Journal of the Entomological Society of Southern Africa, 42, 245–256.
    Ferris, G.F. (1937) s.n. In: Atlas of the scale insects of North America. Series 1. Stanford University Press, Palo Alto, California, S1–S136.
    Ferris, G.F. (1950) Atlas of the Scale Insects of North America. Seris 5. Vol. 5. The Pseudococcidae. Part I. Stanford University Press, Palo Alto, California, 278 pp.
    Ferris, G.F. (1953) Atlas of the Scale Insects of North America. Vol. 6. The Pseudococcidae (Part II). Stanford University Press, Palo Alto, California, 506 pp.
    Foldi, I. (2005a) Ground pearls: a generic revision of the Margarodidae sensu stricto. (Hemiptera: Sternorrhyncha: Coccoidea). Annales de la Société entomologique de France, 41, 81–125. https://doi.org/10.1080/00379271.2005.10697442
    Foldi, I. (2005b) The Matsucoccidae in the Mediterranean basin with a world list of species (Hemiptera: Sternorrhyncha: Coccoidea). Annales de la Société entomologique de France, 40 (2), 145–168. https://doi.org/10.1080/00379271.2004.10697412
    Foldi, I. (2010) Trois espèces nouvelles du genre Crypticerya Cockerell de la région méditerranéenne (Hemiptera, Coccoidea, Monophlebidae). Bulletin de la Société entomologique de France, 115 (3), 289–304.
    García Morales, M., Denno, B., Miller, D.R., Miller, G.L., Ben-Dov, Y. & Hardy, N.B. (2016) ScaleNet: a literature-based model of scale insect biology and systematics. Database. Available from: http://scalenet.info (accessed 22 October 2020)
    Giliomee, J.H. & Munting, J. (1968) A new species of Asterolecanium Targ. (Homoptera: Coccoidea: Asterolecaniidae) from South Africa. Journal of the Entomological Society of Southern Africa, 31, 221–229.
    Giliomee, J.H. & Kozár, F. (2008) Review of the family Asterolecaniidae (Hemiptera: Coccoidea) in South Africa, with the description of a new species. African Entomology, 16 (2), 276–286. Available from: http://www.bioone.org/doi/full/10.4001/1021-3589-16.2.276 (accessed 22 October 2020) https://doi.org/10.4001/1021-3589-16.2.276
    Gill, R.J. (1988). The Scale Insects of California. Part 1. The Soft Scales (Homoptera: Coccoidea: Coccidae). California Department of Food & Agriculture, Sacramento, California, 132 pp.
    Gill, R.J. (1993) The Scale Insects of California. Part 2. The Minor Families (Homoptera: Coccoidea). California Department of Food & Agriculture, Sacramento, California, 241 pp.
    Gill, R.J. (1997). The Scale Insects of California. Part 3. The Armored Scales (Homoptera: Diaspididae). California Department of Food & Agriculture, Sacramento, California, 307 pp.
    Githae, M.M., Ong’amo, G.O., Nderitu, J., Watson, G.W. & Kinuthia, W. (2021) Diversity of scale insects (Hemiptera: Coccomorpha) attacking citrus trees in Machakos, Makueni, Kilifi and Kwale Counties, Kenya. Journal of Agricultural Science and Practice, 6 (3), 79‒85.
    Gullan, P.J., Giliomee, J.H., Hodgson, C.J. & Cook, L.G. (2006) The systematics and biology of the South African gall-inducing scale insect, Calycicoccus merwei Brain (Hemiptera: Coccoidea: Eriococcidae). African Entomology, 14 (1), 13–33.
    Hall, W.J. (1946) On the Ethiopian Diaspidini (Coccoidea). Transactions of the Royal Entomological Society of London, 97, 497–592. https://doi.org/10.1111/j.1365-2311.1946.tb00275.x
    Hambleton, E.J. (1976) A revision of the new world mealybugs of the genus Rhizoecus (Homoptera: Pseudococcidae). United States Department of Agriculture Technical Bulletin, 1522, 1–88.
    Hodgson, C.J. (1967a) Notes on Rhodesian Coccidae (Homoptera: Coccoidea): Part 1: The genera Coccus, Parasaissetia, Saissetia and a new genus Mashona. Arnoldia, Rhodesia, 3, 1–22.
    Hodgson, C.J. (1967b) Some Pulvinaria species (Homoptera: Coccidae) of the Ethiopian region. Journal of the Entomological Society of Southern Africa, 30, 198–211.
    Hodgson, C.J. (1967c) A species of Conchaspis (Homoptera: Conchaspididae) from central Africa. Arnoldia, Rhodesia, 3, 1–3.
    Hodgson, C.J. (1967d) A revision of the species of the genus Ceronema Maskell (Homoptera: Coccoidea) recorded from the Ethiopian Region. Arnoldia, Rhodesia, 3, 1–8.
    Hodgson, C.J. (1967e) A revision of the species of Inglisia Maskell (Homoptera: Coccoidea) recorded from the Ethiopian region. Arnoldia, Rhodesia, 3, 1–11.
    Hodgson, C.J. (1968a) Further notes on the genus Pulvinaria Targ. (Homoptera: Coccoidea) from the Ethiopian region. Journal of the Entomological Society of Southern Africa, 31, 141–174.
    Hodgson, C.J. (1968b) Four new species and a new genus of Coccidae (Homoptera: Coccoidea) from Africa. Proceedings of the Royal Entomological Society of London, Series B: Taxonomy, 37, 114–120. https://doi.org/10.1111/j.1365-3113.1968.tb00208.x
    Hodgson, C.J. (1969a) Notes on Rhodesian Coccidae (Homoptera: Coccoidea): Part II: The genera Ceroplastes and Gascardia. Arnoldia, Rhodesia, 4, 1–43.
    Hodgson, C.J. (1969b) Notes on Rhodesian Coccidae (Homoptera: Coccoidea): Part III. Arnoldia, Rhodesia, 4, 1–42.
    Hodgson, C.J. (1969c) The status of Hemilecanium imbricans (Green) (Homoptera: Coccoidea) in Africa south of the Sahara. Journal of Natural History, 3, 321–327. https://doi.org/10.1080/00222936900770281
    Hodgson, C.J. (1969d) Pests of citrus and their control in citrus in Rhodesia. Rhodesia Agricultural Journal Technical Bulletin, No. 7, 18–29.
    Hodgson, C.J. (1970a) Status of Pulvinaria vayssierei Castel-Branco (Homoptera: Coccoidea). Journal of the Entomological Society of Southern Africa, 33, 279–283.
    Hodgson, C.J. (1970b) A new species of Coccus (Homoptera: Coccoidea) from Malawi. Entomologist’s Monthly Magazine, 106, 35–37.
    Hodgson, C.J. (1971) The species assigned to the genus Ceroplastodes (Homoptera: Coccoidea) in the Ethiopian region. Journal of Entomology (B) Taxonomy, 40, 49–61. https://doi.org/10.1111/j.1365-3113.1971.tb00104.x
    Hodgson, C.J. (1973a) A new name to replace Mashona Hodgson, 1967 (Hem. Coccoidea). Entomologist’s Monthly Magazine, 109, 63.
    Hodgson, C.J. (1973b) A revision of the Lecanodiaspis Targioni-Tozzetti (Homoptera: Coccoidea) of the Ethiopian Region. Bulletin of the British Museum (Natural History). Entomology, 27, 413–452.
    Hodgson, C.J. (1990) The scale insect genus Houardia Marchal (Homoptera: Coccidae). Systematic Entomology, 15, 219–226. https://doi.org/10.1111/j.1365-3113.1990.tb00313.x
    Hodgson, C.J. (1991) A revision of the scale insect genera Etiennea and Platysaissetia (Homoptera: Coccidae) with particular reference to Africa. Systematic Entomology, 16, 173–221. https://doi.org/10.1111/j.1365-3113.1991.tb00684.x
    Hodgson, C.J. (1993) The immature instars and adult male of Etiennea (Homoptera: Coccidae) with a discussion of its affinities. Journal of African Zoology, 107 (3), 193–215.
    Hodgson, C.J. (1994) The scale insect family Coccidae: an identification manual to genera. CAB International, Wallingford, 639 pp.
    Hodgson, C.J. (2008) A new genus and two new species of soft scale insect (Sternorrhyncha, Coccoidea, Coccidae) from Africa. ZooKeys, 3, 57–76. https://doi.org/10.3897/zookeys.3.45
    Hodgson, C.J. & Foldi, I. (2006) A review of the Margarodidae sensu Morrison (Hemiptera: Coccoidea) and some related taxa based on the morphology of adult males. Zootaxa, 1263 (1), 1–250. https://doi.org/10.11646/zootaxa.1263.1.1
    Hodgson, C.J. & Gullan, P.J. (2005) Unusual abdominal segmentation in a South African gall-forming Eriococcid (Hemiptera: Coccoidea) (Abstract only). Proceedings of the X International Symposium on Scale Insect Studies, held at Plant Protection Research Institute, Adana/ Turkey, 19–23 April 2004. Adana Zirai Muscadele Arastirma Enstitusu, Adana, 408 pp.
    Hodgson, C.J. & Henderson, R.C. (2000) Coccidae (Insecta: Hemiptera: Coccoidea). Fauna of New Zealand, 41, 1–259.
    Hodgson, C.J. & Millar, I.M. (2002) A new subfamily, two new genera and three new species of Aclerdidae (Hemiptera: Coccoidea) from southern Africa, with a phylogenetic analysis of relationships. Systematic Entomology, 27 (4), 469–517. https://doi.org/10.1046/j.1365-3113.2002.00189.x
    Hodgson, C.J. & Peronti, A.L.B.G. (2012) A revision of the wax scale insects (Hemiptera: Sternorrhyncha: Coccoidea: Ceroplastinae) of the Afrotropical Region. Zootaxa, 3372 (1), 1–265. https://doi.org/10.11646/zootaxa.3372.1.1
    Hodgson, C.J. & Williams, D.J. (2016) A revision of the family Cerococcidae Balachowsky (Hemiptera: Sternorrhyncha, Coccomorpha) with particular reference to species from the Afrotropical, western Palaearctic and western Oriental Regions, with the revival of Antecerococcus Green and description of a new genus and fifteen new species, and with ten new synonymies. Zootaxa, 4091 (1), 1–175. https://doi.org/10.11646/zootaxa.4091.1.1
    Hodgson, C.J., Abbas, G., Arif, M.J., Saeed, S. & Karar, H. (2008) Phenacoccus solenopsis Tinsley (Sternorrhyncha: Coccoidea: Pseudococcidae), an invasive mealybug damaging cotton in Pakistan and India, with a discussion on seasonal morphological variation. Zootaxa, 1913 (1), 1–35. https://doi.org/10.11646/zootaxa.1913.1.1
    Hodgson, C.J., Millar, I.M. & Gullan, P.J. (2011) Cissococcus Cockerell (Hemiptera: Coccoidea: Coccidae), a unique gall-inducing soft scale genus on Vitaceae from South Africa, with description of a new species. Zootaxa, 2996 (1), 1–32. https://doi.org/10.11646/zootaxa.2996.1.1
    Hodgson, C.J., Williams, D.J. & Giliomee, J.H. (2009) The identity of the wax scale insect, Ceroplastes myricae (Linnaeus) (Sternorrhyncha: Coccoidea: Coccidae), and a key to the wax scales recorded from South Africa. African Entomology, 17 (1), 95–105. https://doi.org/10.4001/003.017.0111
    Howell, J.O. & Williams, M.L. (1976) An annotated key to the families of scale insects (Homoptera: Coccoidea) of America, North of Mexico, based on characteristics of the adult female. Annals of the Entomological Society of America, 69 (2), 181–189. https://doi.org/10.1093/aesa/69.2.181
    Johnson, C., Agosti, D., Delabie, J.J., Dumpert, K., Williams, D.J., Tschirnhaus, M. von & Maschwitz, U. (2001) Acropyga and Azteca ants (Hymenoptera: Formicidae) with scale insects (Sternorrhyncha: Coccoidea): 20 million years of intimate symbiosis. American Museum Novitates, 3335, 1–18. https://doi.org/10.1206/0003-0082(2001)335%3C0001:AAAAHF%3E2.0.CO;2
    Klerk, C.A. de, Ben-Dov, Y. & Giliomee, J.H. (1982a) Redescriptions of four vine infesting species of Margarodes Guilding (Homoptera: Coccoidea: Margarodidae) from South Africa. Phytophylactica, 14, 61‒76.
    Klerk, C.A. de, Ben-Dov, Y. & Giliomee, J.H. (1982b) Redescriptions of three Margarodes Guilding species (Homoptera: Coccoidea: Margarodidae) found on grasses in South Africa. Phytophylactica, 14, 77‒83.
    Klerk, C.A. de, Ben-Dov, Y. & Giliomee, J.H. (1983) Adult female morphology of South African species of Margarodes (Homoptera, Coccoidea, Margarodidae) with keys to nymphs and adult females. Phytophylactica, 15, 133‒144.
    Kondo, T., Gullan, P.J. & Pemberton, R.W. (2011) A new species of Paratachardina Balachowsky (Hemiptera: Coccoidea: Kerriidae) related to the lobate lac scale, P. pseudolobata Kondo & Gullan. Journal of Asia-Pacific Entomology, 14, 141–146. https://doi.org/10.1016/j.aspen.2010.09.003
    Kosztarab, M. (1996) Scale insects of Northeastern North America. Identification, biology, and distribution. Virginia Museum of Natural History, Martinsburg, Virginia, 650 pp.
    Kosztarab, M. & Kozár, F. (1988) Scale Insects of Central Europe. Akademiai Kiado, Budapest, 456 pp. https://doi.org/10.1007/978-94-009-4045-1
    Kozár, F. (2004) Ortheziidae of the World. Plant Protection Institute, Hungarian Academy of Sciences, Budapest, 525 pp. 
    Kozár, F. & Konczné Benedicty, Z. (2007) Rhizoecinae of the world. Plant Protection Institute, Hungarian Academy of Sciences, Budapest, 617 pp.
    Kozár, F., Kaydan, M.B., Konczné Benedicty, Z. & Szita, É. (2013) Acanthococcidae and Related Families of the Palaearctic Region. Hungarian Academy of Sciences, Budapest, 680 pp.
    Łagowska, B. & Hodgson, C.J. (2019a) On some new and old soft scale insects (Hemiptera: Coccomorpha: Coccidae) from Africa, with description of a new Coccus species and introduction of a new genus. Zootaxa, 4612 (3), 373–386. https://doi.org/10.11646/zootaxa.4612.3.4
    Łagowska, B. & Hodgson, C.J. (2019b) Addendum to Zootaxa 4612 (3): On some new and old soft scale insects (Hemiptera: Coccomorpha: Coccidae) from Africa, with description of a new Coccus species and introduction of a new genus by B. Łagowska and Chris J. Hodgson. Zootaxa, 4629 (4), 600. https://doi.org/10.11646/zootaxa.4629.4.8
    Lambdin, P.L. & Kosztarab, M. (1977) Morphology and systematics of the adult females of the genus Cerococcus (Homoptera: Coccoidea: Cerococcidae). Bulletin of the Virginia Polytechnic Institute and State University Research Division, Blacksburg, 128, 1–252.
    Macharia, I., Kimani, E., Koome, F., Kosiom, T., Heya, H., Otipa, M. & Oronje, M.L. (2018) First report and distribution of the papaya mealybug, Paracoccus marginatus, in Kenya. Journal of Agricultural and Urban Entomology, 33 (1), 142–150. https://doi.org/10.3954/JAUE17-02.1
    Macharia, I., Kibwage, P., Heya, H.M., Koome, F., Olobayo, D., Kinuthia, W., Ouvrard, D. & Watson, G.W. (2021) Recent new records of scale insects and mealybugs in Kenya. EPPO Bulletin. [in press]
    Mamet, R.J. (1954) A monograph of the Conchaspididae Green (Hemiptera: Coccoidea). Transactions of the Royal Entomological Society of London, 105, 189–239. https://doi.org/10.1111/j.1365-2311.1954.tb00764.x
    Mamet, R.J. (1958) The Selenaspidus complex (Homoptera, Coccoidea). Annales du Musée Royal du Congo Belge. New Series, 4, 359–429.
    Matile-Ferrero, D. (1977) Une cochenille nouvelle nuisible au manioc en Afrique Équatoriale, Phenacoccus manihoti n. sp. (Hom., Coccoidea, Pseudococcidae). Annales de la Société entomologique de France, 13, 145–152.
    McKenzie, H.L. (1967) Mealybugs of California with taxonomy, biology, and control of North American species (Homoptera: Coccoidea: Pseudococcidae). University of California Press, Berkeley, California, 526 pp. https://doi.org/10.1525/9780520338227
    Mekuria, T.A., Smith, T.J., Beers, E., Watson, G.W. & Eastwell, K.C. (2013) First report of transmission of Little Cherry Virus 2 to sweet cherry by Pseudococcus maritimus (Ehrhorn) (Hemiptera: Pseudococcidae). Disease Note. Plant Disease, 97 (6), 851. https://doi.org/10.1094/PDIS-12-12-1115-PDN
    Miller, D.R. & Davidson, J.A. (2005) Armored Scale Insect Pests of Trees and Shrubs. Cornell University Press, Ithaca, New York, 442 pp.
    Miller, D.R. & Williams, D.J. (1995) Systematic revision of the family Micrococcidae (Homoptera: Coccoidea), with a discussion of its relationships, and a description of a gynandromorph. Bollettino del Laboratorio di Entomologia Agraria ‘Filippo Silvestri’. Portici, 50 (1993), 199–247.
    Miller, D., Rung, A., Parikh, G., Venable, G., Redford, A.J., Evans, G.A. & Gill, R.J. (2014) Scale Insects, Edition 2. USDA APHIS Identification Technology Program (ITP), Fort Collins, Colorado, U.S.A. Available from: http://idtools.org/id/scales/key.php?families (accessed 2 April 2020)
    Morrison, H. (1925) Classification of scale insects of the subfamily Ortheziinae. Journal of Agricultural Research, 30, 97–154.
    Morrison, H. (1928) A classification of the higher groups and genera of the coccid family Margarodidae. United States Department of Agriculture Technical Bulletin, 52, 1–239.
    Morrison, H. (1952) Classification of the Ortheziidae. Supplement to “Classification of scale insects of the subfamily Ortheziinae”. United States Department of Agriculture Technical Bulletin, 1052, 1–80.
    Munting, J. (1973) Further descriptions of African lac insects (Homoptera: Lacciferidae). Phytophylactica, 5, 59–64.
    N’Guessan, W.P., Watson, G.W., Brown, J. & N’Guessan, K.F. (2014) First record of Pseudococcus jackbeardsleyi (Hemiptera: Pseudococcidae) from Africa, Côte d’Ivoire. Florida Entomologist, 97 (4), 1690–1693. https://doi.org/10.1653/024.097.0443
    Normark, B.B., Okusu, A., Morse, G.E., Peterson, D.A., Itioka, T. & Schneider, S.A. (2019) Phylogeny and classification of armored scale insects (Hemiptera: Coccomorpha: Diaspididae) Zootaxa, 4616 (1), 1–98. https://doi.org/10.11646/zootaxa.4616.1.1
    Parsa, S., Kondo, T. & Winotai, A. (2012) The cassava mealybug (Phenacoccus manihoti) in Asia: first records, potential distribution, and an identification key. PLoS ONE, 7 (10), e47675, 1–12. https://doi.org/10.1371/journal.pone.0047675
    Peters, T. & Watson, G.W. (1999) The biological control of hibiscus mealybug in Grenada. In: Bell, K. (Ed.), Paths to prosperity—science & technology in the Commonwealth 1999/2000. Kensington Publications Ltd for the Commonwealth Secretariat, London, pp. 130–132.
    Porcelli, F. & Pellizzari, G. (2019) New data on the distribution of scale insects (Hemiptera, Coccomorpha). Bulletin de la Société entomologique de France, 124 (2), 183–188. https://doi.org/10.32475/bsef_2077
    Richard, C. (1976) Révision du groupe des Stictococcus, et création de taxa nouveaux (Homoptera, Coccoidea). Annales de la Société entomologique de France, 12, 653–669.
    Russell, L.M. (1941) A classification of the scale insect genus Asterolecanium. United States Department of Agriculture, Miscellaneous Publications, 424, 1–319. https://doi.org/10.5962/bhl.title.65621
    Schneider, S.A., Fizdale, M. & Normark, B.B. (2019) Key to commonly intercepted armored scales: Aspidiotini. An interactive key to species of Diaspididae: tribe Aspidiotini. Available from: https://idtools.org/id/scales/aspidiotini/about_index.php (accessed 22 October 2020)
    Spodek, M. & Ben-Dov, Y. (2014) A taxonomic revision of the Kermesidae (Hemiptera: Coccoidea) in Israel, with a description of a new species. Zootaxa, 3781 (1), 1‒99.
    Stickney, F.S. (1934) The external anatomy of the red date scale Phoenicococcus marlatti Cockerell, and its allies. United States Department of Agriculture Technical Bulletin, 404, 1–162.
    Stumpf, C.F. & Lambdin, P.L. (2006) Pit scales (Sternorrhyncha: Coccoidea) of North and South America. Agricultural Experiment Station, Institute of Agriculture, University of Tennessee, Knoxville, Tennessee, xix + 231 pp.
    Takagi, S. (1977) Appendix: A new species of Aulacaspis associated with a cycad in Thailand (Homoptera: Coccoidea). Insecta Matsumurana, New Series, 11, 68–72.
    Tinsley, J.D. (1898) An ants’-nest coccid from New Mexico. Canadian Entomologist, 30, 47–48. https://doi.org/10.4039/Ent3047-2
    Unruh, C.M. & Gullan, P.J. (2008) Identification guide to species in the scale insect tribe Iceryini (Coccoidea: Monophlebidae). Zootaxa, 1803 (1), 1–106. https://doi.org/10.11646/zootaxa.1803.1.1
    Vahedi, H.A. & Hodgson, C.J. (2007) Some species of the hypogeal scale insect Porphyrophora Brandt (Hemiptera: Sternorrhyncha: Coccoidea: Margarodidae) from Europe, the Middle East and North Africa. Systematics and Biodiversity, 5 (1), 23–122. https://doi.org/10.1017/S1477200006002039
    Varshney, R.K. (1984) A review of the family Tachardiidae (Kerriidae) in the Orient (Homoptera: Coccoidea). Oriental Insects, New Delhi, 18, 361–384. https://doi.org/10.1080/00305316.1984.10432212
    Varshney, R.K. (1990) A synoptic updated catalogue of lac insects (Homoptera: Coccoidea: Tachardiidae). In: Koteja, J. (Ed.), Proceedings of the Sixth International Symposium of Scale Insect Studies. Part II. Cracow, Poland. 6–12 August 1990. Agricultural University Press, Cracow, pp. 71–74.
    Watson, G.W. (1997) The role of taxonomy in biological control. In: Syarif Hidayat, Sadeli Natasasmita, Entun Santosa, Sumeno, Delon Suwantaputra, Tohidin, Sudarjat, Bey Permadi, M. Suhunan Sianipar, Nenet Susniahti, Elly Rosmaria (Eds.), Pros. Kongres Perhimpunan Entomologi Indonesia V dan Simposium Entomologi: Pengelolaan Serangga Secara Berkelanjutan, Bandung, 24–26 Juni 1997. Perhimpunan Entomologi Indonesia in collaboration with Universitas Padjadjaran, Bandung, West Java, pp. 1–6.
    Watson, G.W. (2002) Arthropods of economic importance: Diaspididae of the world. An illustrated identification guide and information source. Expert Center for Taxonomic Identification (ETI), University of Amsterdam, CD-Rom. Available from: https://diaspididae.linnaeus.naturalis.nl/linnaeus_ng/app/views/introduction/topic.php?id=3377&epi=155 (accessed 22 October 2020)
    Watson, G.W. & Chandler, L.R. (2000) Identification of mealybugs important in the Caribbean region. 2nd Edition. Revised. CABI Bioscience, Egham, Surrey, 40 pp.
    Watson, G.W., Ouvrard, D., Kasina, M., Achieng, J.C., Githae, M.M., Mulwa, J., Kinuthia, W., Macharia, I., Heya, H.M. & Polaszek, A. (2021) New scale insect country records for Kenya (Hemiptera: Coccomorpha) from old samples in insect collections. African Phytosanitary Journal, 2 (1), 72‒104.
    Williams, D.J. (1986) Rastrococcus invadens sp. n. (Hemiptera: Pseudococcidae) introduced from the Oriental Region to West Africa and causing damage to mango, citrus and other trees. Bulletin of Entomological Research, 76, 695–699. https://doi.org/10.1017/S0007485300015182
    Williams, D.J. (1989) The mealybug genus Rastrococcus Ferris (Hemiptera: Pseudococcidae). Systematic Entomology, 14 (4), 433–486. https://doi.org/10.1111/j.1365-3113.1989.tb00298.x
    Williams, D.J. (2004) Mealybugs of southern Asia. The Natural History Museum, London and Southdene SDN. BHD., Kuala Lumpur, 896 pp.
    Williams, D.J. & Granara de Willink, M.C. (1992) Mealybugs of Central and South America. CAB International, Wallingford, 635 pp.
    Williams, D.J. & Watson, G.W. (1988a) The Scale Insects of the Tropical South Pacific Region. Pt. 1. The Armoured Scales (Diaspididae). CAB International, Wallingford, 290 pp.
    Williams, D.J. & Watson, G.W. (1988b) The Scale Insects of the Tropical South Pacific Region. Pt. 1. The Mealybugs (Pseudococcidae). CAB International, Wallingford, 260 pp.
    Williams, D.J. & Watson, G.W. (1990) The Scale Insects of the Tropical South Pacific Region. Pt. 3. The Soft Scales (Coccidae) and Other Families. CAB International, Wallingford, 267 pp.
    Williams, D.J., Matile-Ferrero, D. & Miller, D.R. (2010) A study of some species of the genus Stictococcus Cockerell (Hemiptera: Sternorrhyncha: Coccoidea: Stictococcidae), and a discussion on Stictococcus vayssierei Richard, a species injurious to cassava in Equatorial Africa with a description of a new species from Nigeria. Zootaxa, 2527 (1), 1–27. https://doi.org/10.11646/zootaxa.2527.1.1
    Zeddies, J., Schaab, R.P., Neuenschwander, P. & Herren, H.R. (2001) Economics of biological control of cassava mealybug in Africa. Agricultural Economics, Amsterdam, 24 (2), 209–219. https://doi.org/10.1111/j.1574-0862.2001.tb00024.x
    Zimmerman, E.C. (1948) Homoptera: Sternorrhyncha. Insects of Hawaii, 5, 1‒464.

How to Cite

WATSON, G. W. ., & OUVRARD, D. . (2021). Towards the identification of the scale insects (Hemiptera: Coccomorpha) of continental Africa: 1. Identification of the families. Zootaxa, 5052(2), 1–40. https://doi.org/10.11646/zootaxa.5052.2.1