Abstract
Scale insects (Hemiptera: Sternorrhyncha: Coccomorpha) feed on plant sap; some are damaging pests in agriculture and forestry. However, the scale insects found in continental Africa have not been extensively studied and the available means of identification are incomplete. This synoptic work, the second in this series, covers the six archaeococcoid scale insect families found in continental Africa: Kuwaniidae, Margarodidae, Matsucoccidae, Monophlebidae, Ortheziidae and Putoidae. The work provides identification keys to a total of 29 genera and 137 species, and a checklist for each family. Pseudaspidoproctus zimmermanni (Newstead) (Monophlebidae) is transferred to Aspidoproctus, as Aspidoproctus zimmermanni (Newstead), comb. n. Praelongorthezia praelonga (Douglas) (Ortheziidae) is recorded from Gambia for the first time.
References
Ahmed, M.Z., Ray, C.H., Moore, M.R. & Miller, D.R. (2020) The Matsucoccus Cockerell, 1909 of Florida (Hemiptera: Coccomorpha: Matsucoccidae): potential pests of Florida pines. Insecta Mundi, 0810, 1–31.
Balachowsky, A.S. (1932) Étude biologique des coccides du bassin occidental de la Méditerranée. Encyclopédie Entomologique. Vol. XV. P. Lechevalier & Fils, Paris, 214 + LXVII pp.
Bartlett, B.R. (1978) Ortheziidae. In: Clausen, C.P. (Ed.), Introduced Parasites and Predators of Arthropod Pests and Weeds: a World Review. Agricultural Research Service, United States Department of Agriculture, Washington, DC, pp. 136–137.
Borchsenius, N.S. (1950) [Mealybugs and Scale Insects of USSR (Coccoidea).] Akademii Nauk SSSR. Vol. 32. Zoological Institute, Moscow, 250 pp.
Borchsenius, N.S. (1960) [Fauna of USSR, Homoptera, Kermococcidae, Asterolecaniidae, Lecanidodiaspididae, Aclerdidae.] Akademiia Nauk SSSR. Series. Zoologicheskii Institut, Leningrad, 282 pp.
Branco, M., Brito, O., Jactel, H., Mendel, Z. & Franco, J.C. (2002) Seasonal history of the maritime pine bast scale Matsucoccus feytaudi Ducasse (Hemiptera: Matsucoccidae) in Portugal. Bollettino di Zoologia Agraria e di Bachicoltura, Milano, 33 (3), 319–329. [2001]
Chandra, A., Bhati, D.P.S. & Singh, K.M. (1987) Bionomics of mango mealy bug, Drosicha mangiferae Green (Margarodidae: Hemiptera). Bulletin of Entomology, 28 (2), 145–152.
Choi, J. & Lee, S. (2022) Higher classification of mealybugs (Hemiptera: Coccomorpha) inferred from molecular phylogeny and their endosymbionts. Systematic Entomology, 1–17. https://doi.org/10.1111/syen.12534
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. (1959) Two new Margarodidae (Homopt.: Coccoidea) from Kenya. Journal of the Entomological Society of Southern Africa, 22, 385–389.
Durán, M. (1971) [The investigations on ground pearls (Margarodes (Porphyrophora) tritici Bod.) a grain pest in central Anatolia]. Bitki Koruma Bulteni. Ankara (Plant Protection Bulletin), Supplement 1, 1–80.
Foldi, I. (1981) Margarodesia desmieri, n. gen. n. sp., cochenille Afro-tropicale nuisible au palmier à huile. Annales de la Société entomologique de France, 17, 331–335.
Foldi, I. (1999) Diversité morphologique chez les cochenilles mâles des Margarodidae: l’exemple de Neomargarodes erythrocephala Green and Stigmacoccus asper Hempel (Hemiptera: Coccoidea). Annales de la Société entomologique de France, 35 (3/4), 319–327.
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. [2004] 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. https://doi.org/10.3406/bsef.2010.2686
García Morales, M., Denno, B.D., 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 12 January 2022) https://doi.org/10.1093/database/bav118.
Gavrilov-Zimin, I.A. (2017) Contribution to the cytogenetics of Kuwaniini scale insects (Homoptera, Coccinea, Margarodidae s.l.). Comparative Cytogenetics, 11 (4), 659–663. https://doi.org/10.3897/CompCytogen.v11i4.20168
Gavrilov-Zimin, I.A. (2018) Ontogenesis, morphology and higher classification of archaeocococcids (Homoptera: Coccinea: Orthezoiidea). Zoosystematica Rossica, 2018 (2), 1–264. https://doi.org/10.31610/zsr/2018.supl.2.1
Gavrilov-Zimin, I.A. & Stekolshikov, A.V. (2018) A new species of the genus Steatococcus Ferris, 1921 (Homoptera, Coccinea: Margarodidae) with some additions to the fauna of the Republic of Mali. Entomological Review, 98 (7), 365‒367. https://doi.org/10.1134/S0013873818070060
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.
González, R.H. (1983) Manejo de plagas de la vid. Publicaciones en Ciencia Agricultura, Universidad de Chile, Facultad Ciencia Agrarias, Veterinarias, y Forestales, 13, 1–115.
Green, E.E. (1922) The Coccidae of Ceylon. Part V. Dulau & Co., London, pp. 345–472.
Gullan, P.J. & Cook, L.G. (2007) Phylogeny and higher classification of the scale insects (Hemiptera: Sternorrhyncha: Coccoidea). Zootaxa, 1668 (1), 413–425. https://doi.org/10.11646/zootaxa.1668.1.22
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. & Hardy, N.B. (2013) The phylogeny of the superfamily Coccoidea (Hemiptera: Sternorrhyncha) based on the morphology of extant and extinct macropterous males. Systematic Entomology, 38, 794–804. https://doi.org/10.1111/syen.12030
Hughes-Schrader, S. (1930) Contributions to the life history of the iceryine coccids, with special reference to parthenogenesis and hermaphroditism. Annals of the Entomological Society of America, 23, 359–380. https://doi.org/10.1093/aesa/23.2.359
Hughes-Schrader, S. (1963) Hermaphroditism in an African coccid, with notes on other Margarodids (Coccoidea-Homoptera). Journal of Morphology, 113, 173–184. https://doi.org/10.1002/jmor.1051130205
Karar, H., Arif, M.J., Sayyed, H.A., Saeed, S., Abbas, G. & Gogi, M.D. (2008) Integrated pest management of mango mealybug (Drosicha mangiferae) in mango orchards. International Journal of Agriculture & Biology, 11, 81–84.
Kawai, S. (1980) Scale insects of Japan in colors. National Agricultural Education Association, Tokyo, 455 pp.
Kinuthia, M.W. & Mwangi, R.W. (1988) Growth, development and life cycle of Icerya pattersoni (Newstead) (Homoptera: Margarodidae), a coffee pest in Kenya. Insect Science and its Application, 9 (1), 69–72. https://doi.org/10.1017/S1742758400010055
Klerk, C.A. de (2017) Identification, control and management of grapevine Margarodes. Viticulture research, Winetech Technical. Available from: https://www.wineland.co.za/identification-control-management-grapevine-margarodes/ (accessed 18 June 2021)
Klerk, C.A. de, Ben-Dov, Y. & Giliomee, J.H. (1982) 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. (1983) General morphology of South African species of Margarodes (Homoptera, Coccoidea, Margarodidae) with keys to nymphs and adult females. Phytophylactica, 15 (4), 133‒144.
Kondo, T. (2022a in press) 3. Archaeococcoids. In: Kondo, T. & Watson, G.W. (Eds.), Encyclopedia of Scale Insect Pests. CAB International, Wallingford, p. 38.
Kondo, T. (2022b in press) 4. Neococcoids. In: Kondo, T. & Watson, G.W. (Eds.), Encyclopedia of Scale Insect Pests. CAB International, Wallingford, p. 117.
Kondo, T. & Watson, G.W. (Eds.) (2022 in press) 3.7 Monophlebidae. In: Kondo, T. & Watson, G.W. (Eds.), Encyclopedia of Scale Insect Pests. CAB International, Wallingford, pp. 86–108.
Kondo, T., Peronti, A.L., Kozár, F. & Szita, É. (2013) Chapter 17. The citrus orthezia, Praelongorthezia praelonga (Douglas) (Hemiptera: Ortheziidae), a potential invasive species. In: Peña, J. (Ed.), Potential Invasive Pests of Agricultural Crops. CAB International, Wallingford, pp. 301–319. https://doi.org/10.1079/9781845938291.0301
Koteja, J. (1974) Comparative studies on the labium in the Coccinea (Homoptera). Zeszyty Naukowe Akademii Rolniczej w Warszawie, Rozprawy Naukowe, 89, 1–162.
Kosztarab, M.P. (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. & Foldi, I. (2000) Matileortheziola angolaensis n. gen., n. sp. [Hemiptera, Coccoidea, Ortheziidae]. Revue française d’Entomologie, New Series, 22 (4), 251–254.
Kozár, F. & Konczné Benedicty, Z. (2000) Revision of Newsteadia of the Australian and Pacific regions, with description of eleven new species (Homoptera: Coccoidea, Ortheziidae). Acta Zoologica Academiae Scientiarum Hungaricae, 46 (3), 197–229.
Kozár, F. & Konczné Benedicty, Z. (2001) Ortheziola of Asia with the descriptions of three new species, and world distribution of the genus (Homoptera: Coccoidea, Ortheziidae). Acta Zoologica Academiae Scientiarum Hungaricae, 47 (1), 15–25.
Kozár, F. & Miller, D.R. (2000) World revision of Ortheziola Šulc (Homoptera: Coccoidea: Ortheziidae) with descriptions of eleven new species. Systematic Entomology, 25, 15–45. https://doi.org/10.1046/j.1365-3113.2000.00090.x
Macharia, I., Kibwage, P., Heya, H.M., Koome, F. Olubayo, D., Kinuthia, W. & Ouvrard, D. & Watson, G.W. (2021) Recent new records of scale insects and mealybugs in Kenya. EPPO Bulletin, 51 (3), 639–647. https://doi.org/10.1111/epp.12786
McClure, M.S. (1977) Population dynamics of the red pine scale, Matsucoccus resinosae (Homoptera: Margarodidae): the influence of resinosis. Environmental Entomology, 6, 789–795. https://doi.org/10.1093/ee/6.6.789
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
Mendel, Z., Zehavi, A. & Zeidan, S. (1991) [Palaeococcus fuscipennis Burm. (Homoptera: Margarodidae), a new pest of pines in Israel.] Hassadeh, 71, 759–760. [In Hebrew]
Mendel, Z., Assael, F., Zeidan, S. & Zehavi, A. (1998) Classical biological control of Palaeococcus fuscipennis (Burmeister) (Homoptera: Margarodidae) in Israel. Biological Control, 12, 151–157. https://doi.org/10.1006/bcon.1998.0621
Miller, D.R. & Kozár, F. (2002) Systematic analysis of afrotropical Newsteadia (Hemiptera, Coccoidea: Ortheziidae) with descriptions of nine new species. Acta Phytopathologica et Entomologica Hungarica, 37 (103), 201–250. https://doi.org/10.1556/APhyt.37.2002.1-3.23
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. Available from: http://idtools.org/id/scales/factsheet.php?name=6726 (accessed 15 February 2022)
Mongue, A.J., Michaelides, S., Coombe, O., Tena, A., Kim, D-S., Normark, B.B., Hardner, A., Hoddle, M.S. & Ross, L. (2021) Sex, males, and hermaphrodites in the scale insect Icerya purchasi. Evolution. [published online] https://doi.org/10.1101/2020.09.04.281618
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.
Normark, B.B. (2003) The evolution of alternative genetic systems in insects. Annual Review of Entomology, 48, 397‒423. https://doi.org/10.1146/annurev.ento.48.091801.112703
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
Shivakumar, G. & Lakshmikantha, B.P. (2001) Biology of Orthezia insignis Browne (Ortheziidae: Homoptera) on coleus. Pest Management and Economic Zoology, 9 (2), 197–200.
Sirisena, U.G.A.I., Watson, G.W., Hemachandra, K.S. & Wijayagunasekara, H.N.P. (2013) A modified technique for the preparation of specimens of Sternorrhyncha for taxonomic studies. Tropical Agricultural Research Journal, 24 (2), 139–149.
Tang, F.T. (2000) [Neomargarodes (Coccoidea: Margarodidae) and a new species reported from China.] Wuyi Science Journal, 16, 1–5.
Tang, F.T. & Hao, J.J. (1995) [The Margarodidae and others of China.] Chinese Agricultural Science Technology Press, Beijing, 738 pp.
Vahedi, H.A. (2001) The biology of Porphyrophora tritici (Hemiptera: Coccoidea: Margarodidae) and the effect of some farming practices on its populations in Kermanshah, Iran. Entomologica, 33 (1999), 357–363.
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
Vayssière, P. (1926) Contribution à l’étude biologique et systématique des Coccidae. Annales des Epiphyties, 12, 187‒382.
Vayssière, P. (1932) Monophlebines et pseudococcines de l’Afrique du Nord. Hem. Coccidae. Bulletin de la Société entomologique de France, 37, 24–28. https://doi.org/10.3406/bsef.1932.14365
Vayssière, P. (1957) Monophlebines nuisible au cotonnier et a l’arachide en Afrique (Homoptera, Coccidae). Bulletin Institut Royal des Sciences Naturelles de Belgique, 33, 1–8.
Vayssière, P. (1968) Une cochenille nouvelle, Afrodrosicha nimbae n. g., n. sp., (Margarodidae) du Mont Nimba (Guinée). Bulletin de l’Institut Fondamental d’Afrique Noire (Ser. A) Science Naturelles, 30, 1493‒1500.
Vea, I.M. & Grimaldi, D.A. (2016) Putting scales into evolutionary time: the divergence of major scale insect lineages (Hemiptera) predates the radiation of modern angiosperm hosts. Scientific Reports, 6 (23487), 1–11. https://doi.org/10.1038/srep23487
Watson, G.W. (2002) Arthropods of economic importance: Diaspididae of the world. An illustrated identification guide and information source. CD-ROM. Expert Center for Taxonomic Identification (ETI), University of Amsterdam, The Netherlands. ISBN-13: 978-9231700033; ISBN-10: 9231700030. Available from: https://diaspididae.linnaeus.naturalis.nl/linnaeus_ng/app/views/introduction/topic.php?id=3377&epi=155 (accessed 12 February 2021)
Watson, G.W. (2021) A checklist of the scale insects (Hemiptera: Coccomorpha) of Kenya. Zootaxa, 5060 (4), 515–532. https://doi.org/10.11646/zootaxa.5060.4.3
Watson, G.W. & Chandler, L.R. (2000) Identification of mealybugs important in the Caribbean region. Second edition (revised). CABI Bioscience, Egham, 40 pp.
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), 151–190. https://doi.org/10.11646/zootaxa.5052.2.1
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 (2), 72–104.
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., Gullan, P.J., Miller, D.R., Matile-Ferrero, D. & Han, S.I. (2011) A study of the scale insect genera Puto Signoret (Hemiptera: Sternorrhyncha: Coccoidea: Putoidae) and Ceroputo Šulc (Pseudococcidae) with a comparison to Phenacoccus Cockerell (Pseudococcidae). Zootaxa, 2802 (1), 1–22. https://doi.org/10.11646/zootaxa.2802.1.1
Wu, S.A., Nan, N., Gullan, P. & Deng, J. (2013) The taxonomy of the Japanese oak red scale insect, Kuwania quercus (Kuwana) (Hemiptera: Coccoidea: Kuwaniidae), with a generic diagnosis, a key to species and description of a new species from California. Zootaxa, 3630 (2), 291–307. https://doi.org/10.11646/zootaxa.3630.2.5