Abstract
For almost a century, the scale insect genus Puto Signoret (Hemiptera: Sternorrhyncha: Coccoidea) was considered to belong to the family Pseudococcidae (the mealybugs), but recent consensus accords Puto its own family, the Putoidae. This paper reviews the taxonomic history of Puto and family Putoidae, compares the morphology of Puto to that of Ceroputo Šulc and Phenacoccus Cockerell, and reassesses the status of all species that have been placed in Puto to determine whether they belong to the Putoidae or to the Pseudococcidae. For 49 of 57 species that have been placed in Puto, as listed in the online database ScaleNet, we score and tabulate features that are diagnostic for Putoidae and then list all species in their correct family placement. For comparison, we include a few species of Pseudococcidae, namely five species of Phenacoccus, including the type species Phenacoccus aceris (Signoret), and the type species of Ceroputo, C. pilosellae Šulc. We provide revised synonymy lists for Puto and Ceroputo, a brief diagnosis of each genus, synonymy lists and notes for several species for which we suggest recombinations or additional synonyms, or for which we have additional data on morphology. We provide a brief diagnosis of Phenacoccus for comparison with Ceroputo and Puto. As a result of our study, we recognise 47 extant and two fossil species of Puto, and six species of Ceroputo. The New World species Puto mimicus McKenzie and Puto nulliporus McKenzie are transferred to the mealybug genus Ceroputo as Ceroputo mimicus (McKenzie) comb. nov. and Ceroputo nulliporus (McKenzie) comb. nov., respectively, and the Old World species Puto pini Danzig and Puto vaccinii Danzig are recognised as Ceroputo pini (Danzig) comb. rev. and Ceroputo vaccinii (Danzig) comb. rev., respectively, in agreement with Tang (1992). The Old World species Puto graminis Danzig is transferred to Ceroputo as Ceroputo graminis (Danzig) comb. nov. Based only on a study of the literature, the following two names are treated here as junior subjective synonyms of Ceroputo pilosellae: Phenacoccus asteri Takahashi syn. nov. and Puto jarudensis Tang syn. nov. We agree with Tang (1992) that Leococcus erigeroneus Kanda should be treated as a junior subjective synonym of C. pilosellae and thus the genus name Leococcus Kanda, erected for L. erigeroneus and formerly treated as a junior synonym of Puto, is a junior synonym of Ceroputo.References
Afifi, S.A. (1968) Morphology and taxonomy of the adult males of Pseudococcidae and Eriococcidae (Homoptera : Coccoidea). Bulletin of the British Museum (Natural History) Entomology, Supplement, 13, 1–210.
Balachowsky, A.S. (1953a) Sur deux Pseudococcini Hom. (Coccoidea) nouveaux appartenant à la faune alpine d'altitude. Mitteilungen der Sweizerischen Entomologischen Gesellschaft, 26, 295–300.
Balachowsky, A.S. (1953b) Sur le statut des genres Puto Signoret, Ceroputo Sulc. et Macrocerococcus Leonardi. Mitteilungen der Sweizerischen Entomologischen Gesellschaft, 26, 301–304.
Beardsley, J.W. (1960) A preliminary study of the males of some Hawaiian mealybugs (Homoptera: Pseudococcidae). Proceedings of the Hawaiian Entomological Society, (1959), 17, 199–243.
Beardsley, J.W. (1962) Descriptions and notes on male mealybugs (Homoptera: Pseudococcidae). Proceedings of the Hawaiian Entomological Society, 28, 81–98.
Beardsley, J.W. (1962) Descriptions and notes on male mealybugs (Homoptera: Coccoidea). Proceedings of the Hawaiian Entomological Society, 18, 81–98.
Beardsley, J.W. (1969) A new fossil scale insect (Homoptera: Coccoidea) from Canadian amber. Psyche, 76, 270–279.
Ben-Dov, Y. (1994) A Systematic Catalogue of the Mealybugs of the World (Insecta: Homoptera: Coccoidea: Pseudococcidae and Putoidae) with Data on Geographical Distribution, Host Plants, Biology and Economic Importance. Intercept Ltd, Andover, U.K., 686 pp.
Ben-Dov, Y. & Kaydan, M.B. (2008) Puto pilosellae (Šulc), new synonymy and distribution records (Hem. Coccoidea, Putoidae). Bulletin de la Société entomologiue de France, 113, 285–286.
Ben-Dov, Y. & Matile-Ferrero, D. (1995) The identity of the mealybug taxa described by V.A. Signoret (Homoptera, Coccoidea, Pseudococcidae). Bulletin de la Société entomologique de France, 100, 241–256.
Ben-Dov, Y., Miller, D.R. & Gibson, G.A.P. (2010) ScaleNet. http://www.sel.barc.usda/scalenet/scalenet.htm (Last accessed 12 December 2010)
Bodenheimer, F.S. (1943) A first survey of the Coccoidea of Iraq. Government of Iraq, Ministry of Economics, Directorate General of Agriculture, Bulletin, 28, 1–33.
Borchsenius, N.S. (1948) [Review of Palaearctic soft scales of the genus Macrocerococcus Leon. (Coccoidea, Pseudococcidae).] (in Russian) Entomologicheskoe Obozrenye, 30, 31–39.
Borchsenius, N.S. (1949) [Insects Homoptera. suborders mealybugs and scales (Coccoidea). Family mealybugs (Pseudococcidae). Vol. VII.] (in Russian) Fauna SSSR. Zoologicheskii Institut Akademii Nauk SSSR. N.S., 38, 1–382.
Borchsenius, N.S. (1958) [On the evolution and phylogenetic interrelations of Coccoidea (Insecta, Homoptera).] (in Russian with English summary) Zoologicheskiĭ Zhurnal, 37, 765–780.
Cook, L.G., Gullan, P.J. & Trueman, H.E. (2002) A preliminary phylogeny of the scale insects (Hemiptera: Sternorrhyncha: Coccoidea) based on nuclear small-subunit ribosomal DNA. Molecular Phylogenetics and Evolution, 25, 43–52.
Cox, J.M. (1984) Relationships of the Phenacoleachiidae (Homoptera: Coccoidea). Proceedings of the 10th International Symposium of Central European Entomofaunistics, Budapest, 15–20 August 1983, 339–341.
Danzig, E.M. (1972a) [New and little-known species of the scale insects (Homoptera, Coccoidea) from Siberia and the far east of the USSR.] (in Russian) Trudy Zoologicheskogo Instituta, Leningrad, 52, 261–276.
Danzig, E.M. (1972b) [Contribution to the fauna of the white flies and scale insects (Homoptera: Aleyrodoidea, Coccoidea) of Mongolia.] (in Russian) Insects of Mongolia, 1, 325–348.
Danzig, E.M. (1978) [New species of mealybugs (Homoptera, Coccoidea) from Siberia and the Far East. In: New Species of Animals.] (in Russian) Trudy Akademii Nauk SSR Zoologicheskogo Instituta. 61, 124–132.
Danzig, E.M. (1980) Scale insects of the Far East of the USSR (Homoptera, Coccinea) SSSR [Keys to the fauna of the USSR]. (in Russian) Nauka, Leningrad, 368 pp.
Danzig, E.M. (1999) [Mealybugs of the genus Puto Signoret (Homoptera, Pseudococcidae) from Russia and adjacent countries.] (in Russian) Entomologicheskoye Obozrenie, 78, 79–91.
De Lotto, G. (1964) Observations on African mealy bugs (Hemiptera: Coccoidea). Bulletin of the British Museum (Natural History) Entomology, 14, 343–397.
De Lotto, G. (1979) On two podal features in the mealybugs (Homoptera: Pseudococcidae). Journal of the Entomological Society of Southern Africa, 42, 11–15.
Ferris, G.F. (1918) The California species of mealy bugs. Leland Stanford Junior University Publications, University Series, Palo Alto, 78 pp.
Ferris, G.F. (1950) Atlas of the Scale Insects of North America. Series V. The Pseudococcidae (Part 1). Palo Alto, California, Stanford University Press, 278 pp.
Foldi, I. (1983) Structure et fonctions des glandes tégumentaires de cochenilles Pseudococcines et de leurs secretions. Annales de la Société entomologique de France, 19, 155–156.
Foldi, I. & Kozár, F. (2006) New species of Cataenococcus and Puto from Brazil and Venezuela, with data on others species (Hemiptera, Coccoidea). Nouvelle Revue d'Entomologie, 22, 305–312.
Giliomee, J.H. (1961) Morphological and taxonomic studies on the males of three species of Pseudococcus (Hemiptera: Coccoidea). Annale Universittett van Stellenbosch (Serie A), 36, 239–296.
Giliomee, J.H. (1967) Morphology and taxonomy of adult males of the family Coccidae (Homoptera: Coccoidea). Bulletin of the British Museum (Natural History) Entomology, Supplement, 7, 1–168.
Gómez-Menor Ortega, J. (1948) Adiciones a los "Cóccidos de España" (2a nota). EOS - Revista Espanola de Entomologia, 24, 73–121.
Green, E.E. (1930) A new species of Phenacoccus (Coccidae) from the eastern Alps. Annals and Magazine of Natural History, 5, 320–322.
Gullan, P.J. & Cook, L.G. (2002) Phenacoleachia Cockerell (Hemiptera: Coccoidea: Phenacoleachiidae). Bollettino di Zoologia agraria e Bachicoltura, 33, 163–173.
Gullan, P.J. & Cook, L.G. (2007) Phylogeny and higher classification of the scale insects (Hemiptera: Sternorrhyncha: Coccoidea). Zootaxa, 1668, 413–425.
Hadzibejli, Z.K. (1956) A new species of the genus Puto Sign. (Homoptera, Coccoidea) from the mountain zone of Georgia. Soobshchenya Akademii Nauk Gruzinskoy SSR, 17, 515–518.
Hardy, N.B., Gullan, P.J. & Hodgson, C.J. (2008) A classification of mealybugs (Hemiptera: Coccoidea) based on integrated molecular and morphological data. Systematic Entomology, 33, 51–71.
Hodgson, C.J. (2002) Preliminary phylogeny of some non-margarodid Coccoidea (Hemiptera) based on adult male characters. Bollettino di Zoologia agraria e Bachicolura, Serie II, 33, 129–137.
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, 1–35.
Hodgson, C.J. & Foldi, I. (2005) Preliminary phylogenetic analysis of the Margarodidae sensu Morrison and related taxa (Hemiptera: Coccoidea) based on adult male morphology. Proceedings of the X International Symposium of Scale Insect Studies held at Adana, Turkey, 19–23 April, 2004, pp. 35–47.
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, 1–250.
Hodgson, C.J. & Henderson, R.C. (2004) Coccidae (Insecta: Hemiptera: Coccoidea): adult males, pupae and prepupae of indigenous species. Fauna of New Zealand, 51, 1–229.
Hughes-Schrader, S. (1944) A primitive coccid chromosome cycle in Puto sp. Biological Bulletin, 87, 167–176.
Hughes-Schrader, S. (1948) Cytology of coccids (Coccoidea-Homoptera). Advances in Genetics, 2, 127–203.
Kanda, S. (1959) Two new genera and two new species of the Pseudococcidae (Homoptera). Kontyu, 27, 239–243.
Kawai, S. (1980) Scale Insects of Japan in Colors. Tokyo, Japan, National Agricultural Education Association, 455 pp.
Kiritshenko, A.N. (1936) [Some new Pseudococcinae of the fauna of USSR (Hemiptera, Coccoidea).] (in Russian) Entomologicheskoye Obozrenie (1935) 26, 130–59.
Kosztarab, M. & Kozár, F. (1988) Scale Insects of Central Europe. Akademiai Kiado, Budapest. 456 pp.
Koteja, J. (1974a) Comparative studies on the labium in the Coccinea (Homoptera). Zeszyty Naukowe Akademii Rolniczej w Krakowie, 27, 1–162.
Koteja, J. (1974b) On the phylogeny and classification of the scale insects (Homoptera, Coccinea) (discussion based on the morphology of the mouthparts). Acta Zoologica Cracoviensia, 19, 267–326.
Koteja, J. (1974c) The occurrence of a campaniform sensillum on the tarsus in the Coccinea (Homoptera). Polskie Pismo Entomologiczne, 44, 243–252.
Koteja, J. (1980) Campaniform, basiconic, and intersegmental sensilla on the antennae in the Coccinea (Homoptera). Acta Biologica Cracoviensia, Series Zoologica, 22, 73–88.
Koteja, J. (1996) Scale Insects (Homoptera: Coccinea) a Day After. In: Schaefer, C.W. (Ed.), Studies on Hemipteran Phylogeny. Thomas Say Publications in Entomology, Proceedings. Entomological Society of America, Lanham, Maryland, pp. 65–88.
Koteja, J. (2000) Scale insects (Homoptera, Coccinea) from Upper Cretaceous New Jersey amber, In: Grimaldi, D. (Ed.), Studies on Fossils in Amber with particular reference to Cretaceous of New Jersey, Backhys Publishers, Leiden, The Netherlands, viii, pp. 147–229.
Koteja, J. & Azar, D. (2008) Scale insects from Lower Cretaceous amber. Alavesia, 2, 133–167.
Koteja, J. & Poinar, G.O. (2001) A new family, genus, and species of scale insect (Hemiptera: Coccoidea: Kukaspididae, new family) from Cretaceous Alaskan amber. Proceedings of the Entomological Society of Washington, 103, 356–363.
Koteja, J. & Zak-Ogaza, B. (1972) Morphology and taxonomy of the male Kermes quercus (L.) (Homoptera, Coccoidea). Acta Zoologica Cracoviensia, 17, 193–215.
Kozár, F. & Walter, J. (1985) Check-list of the Palaearctic Coccoidea (Homoptera). Folia Entomologica Hungarica, 46, 63–110.
Łagowska, B. (2000) Puto superbus (Leonardi, 1907) (Homoptera: Pseudococcidae) new to the Polish fauna. Polskie Pismo Entomologiczne, 69, 3–6.
Leonardi, G. (1907) Contribuzione alla conoscenza delle cocciniglie Italiane. Bollettino del Laboratorio del Zoologia generale e agraria, 1, 135–169.
Leonardi, G. (1909) Chermotheca Italica Continens Exsiccata, In Situ, Coccidarum Plantis Precipue Cultis, In Italia Occurrentibus Obnoxiarum. Cocciniglie Raccolte in Italia. Vesuviano, Portici, Fascicolo V, (106) 125 pp.
Lindinger, L. (1912) Die Schildläuse (Coccidae) Europas, Nordafrikas und Vorder-Asiens, einschliesslich der Azoren, der Kanaren und Madeiras. Ulmer, Stuttgart. 388 pp.
Lindinger, L. (1935) Neue Beiträge zur Kenntnis der Schildläuse (Coccidae). Entomologische Zeitschrift, 49, 121–123.
Malumphy, C. (2010) Barber giant mealybug Puto barberi (Cockerell) (Hemiptera: Pseudococcidae), a Neotropical pest of ornamental plants established in Gran Canaria, Spain. Entomologist’s Monthly Magazine, 146, 21–25.
Marotta, S. (1992) Osservazioni bio-etologiche su Puto superbus (Leonardi) (Homoptera Coccoidea Pseudoccidae) su Arrhenatherum elatius Mert. E. K. in Aruzzo. Redia, 75, 485–499.
Marotta, S. & Tranfaglia, A. (1985) Ridescrizione di Macrocerococcus superbus Leonardi e considerazioni sulla sua posizione generica, con note su Puto Signoret. Bollettino del Laboratorio di Entomologia Agraria 'Filippo Silvestri', Portici, 42, 212–219.
Marotta, S. & Tranfaglia, A. (1993) Le specie del genere Puto Signoret (Homoptera Coccoidea Pseudococcidae) del’Europa centrale e del bacino del Mediterraneo. Bollettino del Laboratorio di Entomologia Agraria ‘Filippo Silvestri’, Portici, 48, 171–205.
Marotta, S. & Tranfaglia, A. (1995) Variability of morphological characters and its use in the systematics of mealybugs (Homoptera: Pseudococcidae). Israel Journal of Entomology, 29, 67–73.
Matesova, G.I. (1957) [New species of coccids (Homoptera, Coccoidea) from southeastern Kazakhstan.] (in Russian) Entomologicheskoe Obozrenye, 36, 163–174.
McKenzie, H.L. (1960) Taxonomic study of California mealybugs with descriptions of new species (Homoptera: Coccoidea: Pseudococcidae). Hilgardia, 29, 681–770.
McKenzie, H.L. (1967) Mealybugs of California with taxonomy, biology, and control of North American species (Homoptera: Coccoidea: Pseudococcidae). University of California Press, Berkeley, 526 pp.
Miller, D.R. (1975) A revision of the genus Heterococcus Ferris with a diagnosis of Brevennia Goux (Homoptera: Coccoidea: Pseudococcidae). United States Department of Agriculture Technical Bulletin, 1497, 1-61.
Miller, D.R. (1984) Phylogeny and classification of the Margarodidae and related groups (Homoptera: Coccoidea). Proceedings of the 10th International Symposium of Central European Entomofaunistics, Budapest, 15–20 August 1983, 321–324.
Miller, D.R. (1991) The scales, scale insects or coccoids, In: Stehr, F.W. (Ed.), Immature Insects, Volume 2. Kendall/Hunt Publishing Company, Iowa, pp. 90–117.
Miller, D.R. & Appleby, J.E. (1971) A redescription of Phenacoccus dearnessi (Homoptera: Coccoidea: Pseudococcidae). Annals of the Entomological Society of America, 64, 1342–1357.
Miller, D.R. & Kosztarab, M. (1979) Recent advances in the study of scale insects. Annual Review of Entomology, 24, 1–27.
Miller, D.R. & Miller, G.L. (1993a) Description of a new genus of scale insect with a discussion of relationships among families related to the Kermesidae (Homoptera: Coccoidea). Systematic Entomology, 18, 237–251.
Miller, D.R. & Miller, G.L. (1993b) A new species of Puto and preliminary analysis of the phylogenetic position of the Puto group within the Coccoidea (Homoptera: Coccoidea). Jeffersoniana: Contributions from the Virginia Museum of Natural History, 4, 1–35.
Miller, D.R., Miller, G.L. & Watson, G.W. (2002) Invasive species of mealybugs (Hemiptera: Pseudococcidae) and their threat to U.S. agriculture. Proceedings of the Entomological Society of Washington, 104, 825–835.
Miller, G.L. & Williams, M.L. (2002) Systematics of the adult male soft scales from America north of Mexico (Hemiptera: Coccidae). Contributions on Entomology, International, 5, 53–126.
Morrison, H. (1927) Descriptions of the genera and species belonging to the coccid family Margarodidae. Proceedings of the Biological Society of Washington, 40, 99–109.
Morrison, H. (1928) A classification of the higher groups and genera of the coccid family Margarodidae. Technical Bulletin United States Department of Agriculture, 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.
Nasonov, N.V. (1908) Steingelia gorodetskia, nov. gen. et nov. sp. nouveau genre et espèce des coccides du groupe Xylococcini. Annuaire du Musée Zoologique de l'Académie Impériale des Sciences de St. Pétersbourg, 13, 345–352.
Normark, B.B. (2003) The evolution of alternative genetic systems in insects. Annual Review of Entomology, 48, 397–423.
Nur, U. (1980) Evolution of unusual chromosome systems in scale insects (Coccoidea: Homoptera). In: Blackman, R.L., Hewitt, G.M. and Ashburner, M. (Eds). Insect Cytogenetics, Symposia of the Royal Entomological Society of London, 10, 97–117.
Reyne, A. (1954) A redescription of Puto antennatus Sign. (Homoptera: Coccoidea) with notes on Ceroputo pilosellae Šulc and Macrocerococcus superbus Leon. Zoologische Mededelingen, 32, 291–324.
Ross, L., Pen, I. & Shuker, D.M. (2010) Genomic conflict in scale insects: the causes and consequences of bizarre genetic systems. Biological Reviews, 85, 807–828.
Schumacher, F. (1917) Eine neue Schildlaus aus Brandenburg. Zeitschrift für Deutsche Entomologen, 4, 429–430.
Signoret, V. (1875) Essai sur les cochenilles ou gallinsectes (Homoptères - Coccides), 15e partie. Annales de la Société entomologique de France (série 5), 5, 305–352.
Signoret, V. (1876) Essai sur les cochenilles ou gallinsectes (Homoptères – Coccides) 17e partie. Annales de la Société entomologique de France (série 5.), 5, 374–394.
Sternlicht, M. (1969) Kermes bytinskii n. spec. (Coccoidea, Kermesidae) in Israel and observations on its life history. Israel Journal of Entomology, 4, 251–270.
Šulc, K. (1898) Studie o coccidech. -II. Sitzungsberichte der K. Bohmisch Gesellschaft der Wissenschaften (1897) 66, 1–19.
Takahashi, R. (1932) Records and descriptions of the Coccidae from Formosa. Part 2. Journal of the Society of Tropical Agriculture Formosa, 4, 41–48.
Tang, F.D. (1992) The Pseudococcidae of China (Homoptera: Coccinea of Insects). Beijing, Chinese Agricultural Science Technology Press, 767 pp.
Tereznikova, E.M. (1975a) Coccids, Issue 18, Scale insects, gigantic scales and mealybugs, Ortheziidae, Margarodidae, Pseudococcidae. Fauna Ukraini, 20, 295 pp.
Tereznikova, E.M. (1975b) [About the synonymy of two coccoid species.] (In Russian) Vestnik Zoologii. Institut Zoologii Akademiya Nauk Ukrainskoi SSR. Kiev, No. 4, 32.
Theron, J.G. (1958) Comparative studies on the morphology of male scale insects. (Hemiptera: Coccoidea). Annals of the University of Stellenbosch, 34, 1–71.
Vayssière, P. (1923) Un nouveau coccide (Hem.) de la faune Africaine. Bulletin de la Société Entomologique de France, 11/12, 152–55.
Vayssière, P. (1927) Sur quelques Coccidae (Hem.) de l'Afrique du Nord. Bulletin de la Société Entomologique de France, 6, 107–111.
Washburn, R.I. (1965) Description and bionomics of a new species of Puto from Utah (Homoptera: Coccoidea: Pseudococcidae). Annals of the Entomological Society of America, 58, 203–297.
Williams, D.J. (2004) Mealybugs of Southern Asia. The Natural History Museum, London, U.K. and Southdene SDN, BHD, Kuala Lumpur, Malaysia, 896 pp.
Williams, D.J. & Granara de Willink, C.M. (1992) Mealybugs of Central and South America. Wallingford, U.K., CAB International, 635 pp.
Yokogawa, T. & Yahara, T. (2009) Mitochondrial phylogeny certified PGL (paternal genome loss) is of single origin and haplodiploidy sensu stricto (arrhenotoky) did evolve from PGL in the scale insects (Hemiptera: Coccoidea). Genes and Genetic Systems, 84, 57–66.
Zacharuk, R.Y. & Shields, V.D. (1991) Sensilla of immature insects. Annual Review of Entomology, 36, 331–354.