Family-Group Names in the Scale Insects (Hemiptera: Sternorrhyncha: Coccoidea)—A Supplement

Zootaxa 3616 (4): 325–344 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3616.4.2 http://zoobank.org/urn:lsid:zoobank.org:pub:351A064C-8CEC-4A85-A466-0E2F139856BB Family-group names in the scale insects (Hemiptera: Sternorrhyncha: Coccoidea)—a supplement

D.J. WILLIAMS Department of Entomology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK. E-mail [email protected]

Abstract

Williams (1969) published a list of the family-group names in the Coccoidea (scale insects) recognised at that time. The present paper supplements this earlier list and includes all nominal genera that have had family-group names based on them, including those in the earlier paper, in case it is not readily available to some workers. Nominal genera and their family-group names are listed alphabetically in catalogue form. There are now 49 families generally recognised in the scale insects, of which 16 are only known as fossils. Furthermore, 180 nominal genera have now had family-group names based on them. As stated in the 1969 list, all categories in the family group are deemed to be of co-ordinate status in nomenclature.

Key words: archaeococcoids, neococcoids, ranks, genera

Introduction

The following list of family-group names in the superfamily Coccoidea or scale insects is a supplement to the earlier list in Williams (1969). When the first list was compiled, most scale insect workers recognised fewer than 20 extant families. This figure has now risen to 33 with the newest addition Rhizoecidae as shown in the database of scale insects (Ben-Dov et al. 2012). Furthermore, the total excludes 16 families only known from fossils that have been described in recent years, so the total of all families is now 49. Moreover, a little over 100 of the nominal genera listed in Williams (1969) had family-group names based on them; there are now 180 in the present list. There are probably far too many families for a group the size of the scale insects. In recent years, there have been many developments in the study of scale insects. It was shown by Koteja (1974, a, b) from studies based on the mouthparts, that many of the subfamilies and tribes established in the family Margarodidae (sensu Morrison, 1928), should be raised to distinct family level. Some later phylogenetic studies based on morphological and molecular data (Hodgson, 2002; Cook et al. 2002; Gullan & Cook, 2007; Hodgson & Foldi, 2005, 2006) have shown that Koteja’s action was justified. A new extinct family Naibiidae Shcherbakov and the subfamily Naibiinae based on the nominal genus Naibia Shcherbakov, described by Shcherbakov (2007) and purporting to be a scale insect stem group, is so far excluded because the position and relationships of the family to scale insects is uncertain (Gullan & Cook, 2007). At present, the scale insects are divided into two major informal groups, the archaeococcoids and the neococcoids. These names are not formally recognised using the “The International Code of Zoological Nomenclature” but they were derived from the names Archaeococcoidea and Neococcoidea of Bodenheimer (1953). Borchsenius (1949) preferred the name Paleococcoidea for the archaeococcoids. Unfortunately, these names were not based on nominal genera. The archaeococcoids comprise the Ortheziidae, all the families elevated from the lower ranks in Morrison (1928), and some other families, mostly with a plesiomorphic sex chromosome system XX-XO (Hughes-Schrader, 1948; Yokogawa & Yahara, 2009). Based on adult male morphology, the archaeococcoid group includes many of the families only known as fossils. The neococcoids contain all the other families, mostly with a sex chromosome system based on paternal genome elimination (PGE) as discussed by

Accepted by C.J. Hodgson: 23 Jan. 2013; published: 21 Feb. 2013 325 Normark (2003), Yokogawa and Yahara (2009) and Ross et al. (2010), plus a few extinct families based on morphological characters. Some workers, mostly in eastern Europe (e.g. Koteja, 1974a; Danzig, 1980) favour a classification accepting the scale insects as belonging to a suborder, the Coccinea, with the two superfamilies Orthezioidea and Coccoidea, names based on nominal genera and replacing the names archaeococcoids and neococcoids. It is not clear if workers in the other hemipteran groups, currently accepted as the superfamilies Psylloidea, Aphididoidea and Aleyrodoidea in the present suborder Sternorrhyncha, would agree with this elevation in rank and accept their study groups as suborders. Family-group ranks below those of family and subfamily are not always popular or necessary. Concepts of generic groupings are changing rapidly with new phylogenetic studies. For example, based on morphological and molecular studies by Hardy et al. (2008), it was shown that there are two subfamilies only in the family Pseudococcidae. All family-group names that had been erected in the two subfamilies of Pseudococcidae were listed by Williams & Gullan (2010). More recently, Ben-Dov (2011) has listed all the genera and species in most of the extant families of the archaeococcoids belonging to the former family Margarodidae of Morrison (1928). Recent printed catalogues of the family Diaspididae by Ben-Dov (2003) and Miller & Gimpel (2009) have listed genera and species belonging to the subfamilies Aspidiotinae, Comstockiellinae, Diaspidinae, Leucaspidinae, Odonaspidinae and Ullucoccinae without using lower ranks. Published catalogues on all the other families have used the family rank only. A recent work on the family Diaspididae (Miller & Davidson, 2005) has dealt with genera and species in strict alphabetical order with keys but without recourse to any ranks below that of family. Other taxonomic works on families in some zoogeographical regions have been published in recent years with the same approach without using ranks below that of family, and all appear to be readily usable. Some of the family- group names have already been treated as synonyms and, detailed work based on morphological and molecular studies, may show new relationships (e.g. Morse & Normark (2006) and Andersen et al. (2010)) for the family Diaspididae) so that some of the present ranks may become obsolete. The following account lists all the nominal genera that have family-group names based on them including names listed in Williams (1969). For the latter, the page reference in Williams (1969) is provided. Names of new ranks, published after the first list, are preceded by an asterisk [*]. The family name, presently accepted in the latest published catalogues and in ScaleNet (Ben-Dov et al. 2012), is added after each nominal genus together with the main groups (either archaeococcoids or neococcoids) following the phylogeny given in Gullan & Cook (2007). I have searched most of the pertinent literature published after 1969 to the end of 2012 but there may be some omissions and there is always the possibility that the list may not be complete. Some major works, published mainly in Chinese, have more precise dates of publication than is generally understood by non-Chinese workers. For instance, the work by Tang (1991) was published in April 1991, that of Tang (1992) on 31 May 1992, and the work by Tang & Hao (1995) in June 1995 (San-an Wu, personal communication).

List of nominal genera and family-group names

Acanthococcus Signoret, 1875. Eriococcidae (neococcoids). Williams, 1969: 318. After a proposal by Miller & Williams (1976), the family-group name Eriococcini Cockerell, 1889, was given precedence over Acanthococcidae Signoret, 1875, by the International Commission on Zoological Nomenclature (1982) (Opinion, 1203). The family-group name based on the nominal genus Acanthococcus is still available.

Aclerda, Signoret, 1874. Aclerdidae (neococcoids). Williams, 1969: 318.

Albicoccus Koteja, 2004. Albicoccidae (archaeococcoids). *Albicoccidae Koteja, 2004: 111. This family was described from a fossil adult male in Burmese amber.

Ancepaspis Ferris, 1920. Diaspididae (neococcoids). Williams, 1969: 319.

Andaspis MacGillivray 1921. Diaspididae (neococcoids). Williams, 1969: 319.

Antakaspis Mamet, 1959. Diaspididae (neococcoids). Williams, 1969: 319.

Anomalococcus Green, 1902. Lecanodiaspididae (neococcoids). *Anomalococcina Tang & Hao, 1995: 211, 244, 245, 623. Tang & Hao placed this subtribe in the tribe Lecanodiaspidini.

Antonina Signoret, 1875. Pseudococcidae (neococcoids). Williams 1969: 319. *Antoninina Borchenius, Tang, 1992: 18. Tang placed this subtribe in the subfamily Sphaerococcinae, now Pseudococcinae.

Antoninoides Ferris, 1953. Pseudococcidae (neococcoids). *Antonoidina Han, Hao, Tang & Tang, 1992: 31 (nomen nudum and misspelling). This subtribal name was placed in the tribe Sphaerococcini, subfamily Sphaerococcinae. Although the authors stated that the subtribe would be included in a key, no such key has been found so the name is without a description (San-an Wu, personal communication).

Aonidia Targioni Tozzetti, 1868. Diaspididae (neococcoids). Williams, 1969: 319.

Apiomorpha Rübsaamen, 1894. Eriococcidae (neococcoids). Williams, 1969: 319.

Araucaricoccus Brimblecombe 1960: 183. Steingeliidae (archaeococcoids). *Araucaricoccini Tang & Hao, 1995: 74, 606. This was erected as a tribe of the subfamily Kuwaniinae. The genus was listed in the subfamily Monophlebinae by Ben-Dov (2005: 140) and transferred later to the family Steingeliidae by Gullan & Cook (2007: 417). The tribal rank is still available in the family Steingeliidae.

Arctorthezia Cockerell, 1900. Ortheziidae (archaeococcoids). *Arctortheziini Kozár 2004: 257. This tribe was placed in the subfamily Ortheziinae.

Arnoldus Koteja, 2008. Arnoldidae (archaeococcoids). *Arnoldidae Koteja, 2008: 27. This family was described from Baltic amber fossils.

Aspidiella Leonardi, 1898. Diaspididae (neococcoids). Williams, 1969: 320.

Aspidoproctus Newstead, 1901. Monophlebidae (archaeococcoids). *Aspidoproctina Tang & Hao, 1995: 121, 173, 616. The authors placed this subtribe in the subfamily Monophlebinae.

Asterococcus Borchsenius, 1960. Cerococcidae (neococcoids). *Asterococcina Tang & Hao, 1995: 211, 213, 619, 620. The authors placed this subtribe in the subfamily Cerococcinae.

Asterodiaspis Signoret, 1877. Asterolecaniidae (neococcoids). *Asterodiaspidina Tang & Hao, 1995: 211, 288. The authors placed this subtribe in the tribe Cerococcini of the family Asterolecaniidae but that tribe is now a member of the family Cerococcidae, whereas the subtribe is assigned to the Asterolecaniidae.

Asterolecanium Targioni Tozzetti, 1868. Asterolecaniidae (neococcoids). Williams, 1969: 320. *Asterolecaniina Cockerell, Tang & Hao, 1995: 211, 288, 334, 663.

Augulaspis MacGillivray, 1921. Diaspididae (neococcoids). Williams, 1969: 320.

Austrotachardia Chamberlin, 1923. Kerriidae (neococcoids). Williams, 1969: 320. The spelling of the subtribe Austrotachardina in Williams (1969) should have been Austrotachardiina. This rank was elevated to the tribe Austrotachardiini in Varshney (1990: 73).

Beesonia Green, 1926. Beesoniidae (neococcoids). Williams, 1969: 320. *Beesoniini Ferris, Foldi, 1995: 287.

Brachyscelis Schrader, 1863. Eriococcidae (neococcoids). Williams, 1969: 321.

Burmacoccus Koteja, 2004. Burmacoccidae (archaeococcoids). *Burmacoccidae Koteja: 2004: 110. This family was described from an amber fossil adult male from Lower Cretaceous (Upper Albian).

Callipappus Guérin-Méneville, 1841. Callipappidae (archaeococcoids). Williams, 1969: 321. *Callipappina MacGillivray, Tang & Hao, 1995: 10, 607. *Callipappidae MacGillivray, Koteja, 1996: 68. This group is now accepted as a family.

Calycicoccus Brain, 1918. Eriococcidae (neococcoids). Williams, 1969: 321. *Calycicoccidae Brain, Koteja, 1974a: 50. *Calycicoccina Brain, Tang & Hao, 1995: 425, 435, 642. The subtribe was erected in the subfamily Eriococcinae. Its present status is in the family Eriococcidae and the group name is available.

Canceraspis Hempel, 1934. Beesoniidae (neococcoids). Williams, 1969: 321. The genus Canceraspis has been treated as a synonym of Limacoccus, a genus placed in the family Beesoniidae (Foldi, 1995). The family-group names based on the nominal genus Canceraspis listed in Williams (1969) are now presumably treated as synonyms of Limacoccini Foldi, 1995, although this action may not be necessary according to Article 40.1 of the International Code of Zoological Nomenclature (1999).

Cardiococcus Cockerell, 1903. Coccidae (neococcoids). *Cardiococcina Tang, Hao, Xie & Tang, 1990: 76, 77. *Cardiococcinae Tang, Hao, Xie & Tang, Hodgson, 1994: 42. *Cardiococcini Tang, Hao, Xie & Tang, Hodgson, 1994: 43.

Caryonema Richard, 1986. Caryonemidae (archaeococcoids). *Caryonemidae Richard, 1986: 268. * Caryoneminae Richard, Kozár, in Kozár & Konczné Benedicty, 2000: 72. *Caryonemini Richard, Kozár, in Kozár & Konczné Benedicty, 2000: 72. The family rank is generally recognised at present.

Cerococcus Comstock, 1882. Cerococcidae (neococcoids). Williams, 1969: 321. *Cerococcidae Balachowsky, Koteja, 1974a: 49. *Cerococcina Balachowsky, Tang & Hao, 1995: 211, 213, 224, 619, 621. The family rank is now widely accepted.

Ceronema Masskell, 1895. Coccidae (neococcoids). *Ceronemina Tang, Hao, Xie & Tang, 1990: 75, 76. The authors placed this subtribe in the tribe Filippiini.

Ceroplastes Gray, 1828. Coccidae (neococcoids). Williams, 1969: 321. *Ceroplastini Maskell, Tang, Hao, Xie & Tang, 1990: 76, 77.

Ceroplastodes Cockerell, 1893. Coccidae (neococcoids). *Ceroplastina Tang, Hao, Xie & Tang, 1990: 75, 76. See note below. *Ceroplastodina Tang, 1991: 11, 29. According to the keys to the subfamily Filippiinae in Tang, 1991: 11, 29, it seems that the name Ceroplastina in Tang, Hao, Xie & Tang, 1990: 75, 76 was a misspelling of Ceroplastodina.

Chionaspis Signoret, 1869. Diaspididae (neococcoids). Williams, 1969: 322.

Cissococcus Cockerell, 1902. Coccidae (neococcoids). Williams, 1969: 322. *Cissococcidae Brain, Koteja, 1974b: 277, 307. *Cissococcini Brain, Ben-Dov, 1993: xxi.

Coccomytylus Leonardi, 1898. Diaspididae (neococcoids). Williams: 1969: 322.

Coccus Linnaeus, 1758. Coccidae (neococcoids). Williams, 1969: 322. Coccus is the type genus for all the scale insects and the family-group name based on it was the first to be established for the scale insects. The type species of Coccus was discussed in detail by Morrison & Morrison (1966) and a proposal by Danzig & Kerzhner (1981) to establish the name Coccus hesperidum Linnaeus as type species was agreed by the Internatonal Commission on Zoological Nomenclature (1985). The generic name Coccus was placed on the Official List of Generic Names in Zoology (Name Number 2244) (Opinion 1303). The family rank Coccidae is widely accepted.

Coelostomidia Cockerell, 1900. Coelostomidiidae (archaeococcoids). Williams, 1969: 323. *Coelostomidiidae Morrison, Koteja, 1974a: 38. This family status is now generally accepted.

Comstockiella Cockerell, 1896. Diaspididae (neococcoids). Williams, 1969: 323. *Comstockiellinae Borchsenius, Ben-Dov, 2003: 8, 816.

Conchaspis Cockerell, 1893. Conchaspididae (neococcoids). Williams, 1969: 323. The family status is generally accepted.

Conifericoccus Brimblecombe, 1960. Steingeliidae (archaeococcoids). *Conifericoccina Tang & Hao, 1995: 9, 608. The authors placed this subtribe in the family Steingeliidae.

Cryptococcus Douglas, 1890. Eriococcidae (neococcoids). Williams, 1969: 323. *Cryptococcinae Kosztarab, Tang & Hao, 1995: 425, 426, 641. This subfamily is presently in the family Eriococcidae but some workers (e.g. Koteja, 1974a) have given it family status—Cryptococcidae.

Cryptokermes Hempel, 1900. Coelostomidiidae (archaeococcoids). *Cryptokermini Tang & Hao, 1995: 121. The authors placed this tribe in the subfamily Coelostomidiinae, presently the family Coelostomidiidae. The tribal name is here corrected to Cryptokermesini.

Ctenochiton Maskell, 1879. Coccidae (neococcoids). Williams, 1969: 323. Ctenochitini Cockerell, Koteja, 1974b: 305. Misspelling. *Ctenochitoninae Cockerell, Koteja & Żak-Ogaza, 1983: 479. *Ctenochitonina Cockerell, Tang, Hao, Xie & Tang, 1990: 76, 77.

Cylindrococcus Maskell, 1892. Eriococcidae (neococcoids). Williams, 1969: 324.

Cyphococcus Laing, 1925. Coccidae (neococcoids). *Cyphococcinae Hodgson, 1994: 51. *Cyphococcini Hodgson, Hodgson, 1994: 214.

Desmococcus McKenzie, 1942. Pityococcidae (archaeococcoids). *Desmococcina Tang & Hao, 1995: 10. The authors placed this subtribe in the tribe Pityococcini, presently in the family Pityococcidae.

Diaspis O.G. Costa, 1828. Diaspididae (neococcoids). Williams, 1969: 324.

Dimargarodes Sivestri, 1938. Margarodidae (archaeococcoids). Williams, 1969: 325.

Drosicha Walker, 1838. Monophlebidae (archaeococcoids). Williams, 1969: 325.

Dysmicoccus Ferris, 1950. Pseudococcidae (neococcoids). *Dysmicoccina Tang, 1992: 187.

Electrococcus Beardsley, 1969. Electrococcidae (archaeococcoids). *Electrococcidae Koteja, 2000: 171. This family was described from a single amber adult male fossil.

Eremococcus Ferris, 1919. Asterolecaniidae (neococcoids). Williams, 1969: 325.

Eriochiton Maskell, 1887. Eriococcidae (neococcoids). *Eriochitonini Hodgson, 1994a: 172. This tribe was placed in the subfamily Eriococcinae of the family Eriococcidae.

Eriococcus Targioni Tozzetti, 1868. Eriococcidae (neococcoids). Williams, 1969: 325. *Eriococcina Cockerell, Tang & Hao, 1995: 425, 444, 643. This subtribe was erected in the subfamily Eriococcinae of the Eriococcidae. The family-group name Eriococcini was given precedence over the family-group name Acanthococcidae by the International Commission on Zoological Nomenclature (1982), after a proposal by Miller & Williams (1972).

Eriopeltis Signoret, 1872. Coccidae (neococcoids). Williams, 1969: 325. *Eriopeltina Šulc, Tang, Hao, Xie & Tang, 1990: 75, 76.

Eulecanium Cockerell, 1893. Coccidae (neococcoids). *Eulecaniinae Koteja in Kosztarab & Kozár, 1988: 173, 174. *Eulecaniina Koteja, Tang, Hao, Xie & Tang, 1990: 75, 76. *Eulecaniini Koteja, Hodgson, 1994: 221.

Eumargarodes Jakubski, 1950. Margarodidae (archaeococcoids). Williams, 1969: 325.

Eurhizococcus Silvestri, 1936. Margarodidae (archaeococcoids). Williams, 1969: 326.

SCALE INSECT FAMILY-GROUP NAMES Zootaxa 3616 (4) © 2013 Magnolia Press · 331 Filippia Targioni Tozzetti, 1868. Coccidae (neococcoids). Williams, 1969: 326. *Filippiina Bodenheimer, Tang, Hao, Xie & Tang, 1990: 75, 76.

Fiorinia Targioni Tozzetti, 1868. Diaspididae (neococcoids). Williams, 1969: 326. *Fioriniina Leonardi, Takagi, 1989: 124, 141; 1989a: 152.

Foldicoccus Kozár in Kozár & Konczné Benedicty, 2000. Caryonemidae (archaeococcoids). *Foldicoccinae Kozár in Kozár & Konczné Benedicty, 2000: 76. *Foldicoccini Kozár in Kozár & Konczné Benedicty, 2000: 76.

Furcaspis Lindinger, 1908. Diaspididae (neococcoids). Williams, 1969: 326.

Gallinococcus Lambdin & Kosztarab, 1976. Lecanodiaspididae (neococcoids). *Gallinococcina Tang & Hao, 1995: 221, 623. The authors placed this subtribe in the subfamily Lecanodiaspidinae.

Geococcus Green, 1902. Rhizoecidae (neococcoids). Geococcina Kozár in Kozár & Konczné Benedicty, 2007: 36. This subtribe was placed originally in the subfamily Rhizoecinae.

Grimaldiella Koteja, 2000. Grimaldiellidae (archaeococcoids). *Grimaldiellidae Koteja, 2000: 158. This family was described from adult male amber fossils.

Grohnus Koteja, 2008. Grohnidae (archaeococcoids). *Grohnidae Koteja, 2008: 46. This family was described from Baltic amber fossils.

Gymnaspis Newstead, 1898. Diaspididae (neococcoids). Williams, 1969: 326.

Halimococcus Cockerell, 1902. Halimococcidae (neococcoids). Williams, 1969: 326. The Family status is widely recognised.

Hammanococcus Koteja & Azar, 2008: 139. Hammanococcidae (archaeococcoids). *Hammanococcidae Koteja & Azar, 2008: 139. This family was described from a Lebanese amber fossil.

Heterococcus Ferris, 1918. Pseudococcidae (neococcoids). *Heterococcina Tang, 1992: 415, 569, 570. This subtribe was placed in the subfamily Phenacoccinae.

Howardia Berlese & Leonardi, 1896. Diaspididae (neococcoids). Williams, 1969: 326.

Humococcus Ferris, 1953. Pseudococcidae (neococcoids). *Humococcina Tang, 1992: 91, 92. This subtribe was placed in the tribe Mirococcopsidini.

Inka Koteja, 1989. Inkaidae (neococcoids). *Inkaidae Koteja, 1989: 78. This family was described from an Upper Cretaceous Taymyrian amber fossil.

Jersicoccus Koteja, 2000. Jersicoccidae (archaeococcoids). *Jersicoccidae Koteja, 2000: 155. This family was described from a single adult male New Jersey amber fossil.

Kermes Boitard, 1828. Kermesidae (neococcoids). Williams, 1969: 327.

Kermococcus Silvestri, 1911. Kermesidae (neococcoids). Williams, 1969: 327.

Kerria Targioni Tozzetti, 1884. Kerriidae (neococcoids). Williams, 1969: 327. The family-group name Kerriidae is now widely used in place of Tachardiidae after the proposal to stabilise the name Kerriidae for all the lac insects (Ben-Dov & Lit, 1998). Varshney (1966), however, quoting the International Code of Zoological Nomenclature, then in use, proposed that the name Tachardiidae should have priority over Kerriidae because the family-group name Tachardiinae of Green in 1906 has priority over Tachardiidae of Lindinger in 1937. Varshney’s action still holds under Article 40.1 of the present Code of Zoological Nomenclature (1999) in the absence of any formal proposal to the International Commission on Zoological Nomenclature to give Kerriidae priority over Tachardiidae.

Kukaspis Koteja & Poinar, 2001. Kukaspididae (archaeococcoids). *Kukaspididae Koteja & Poinar, 2001: 357. This family was described from a single adult male Alaskan amber fossil.

Kuwanaspis MacGillivray, 1921. Diaspididae (neococcoids). Williams, 1969: 328. *Kuwanaspidini Borchsenius, Varshney, 1985: 29.

Kuwania Fernald, 1903. Kuwaniidae (archaeococcoids). Williams, 1969: 328. *Kuwaniina MacGillivray, Danzig 1980: 58. *Kuwaniidae MacGillivray, Koteja, 1974a: 40. The family rank is widely recognised now.

Labiococcus Koteja, 2000. Labiococcidae (neococcoids). Labiococcidae Koteja, 2000: 178. This family was described from a first-instar nymph from New Jersey amber.

Laccifer Oken, 1815. Kerriidae (neococcoids). Williams, 1969: 328.

Lebanococcus Koteja & Azar, 2008. Lebanococcidae (archaeococcoids). *Lebanococcidae Koteja & Azar, 2008: 142. This is an extinct family described from a Lebanese amber fossil.

Lecanodiaspis Targioni Tozzetti, 1869. Lecanodiaspididae (neococcoids). Williams, 1969: 329. *Lecanodiaspidina Tang & Hao, 1995: 211, 224, 253, 623. The family rank for this group is now widely accepted.

Lecanopsis Targioni Tozzetti, 1868. Coccidae (neococcoids). Lecanopsiini Tang, Hao, Xie & Tang, 1990: 75, 76. Lecanopsidini. Corrected spelling (Hodgson, 1994: 39).

Lepidosaphes Shimer, 1868. Diaspididae (neococcoids). Williams, 1969: 329.

Leucaspis Targioni Tozzetti, 1868. Diaspididae (neococcoids). Williams, 1969: 329.

Leucodiaspis Signoret, 1869. Diaspididae (neococcoids). Williams, 1969: 330.

Limacoccus Bondar, 1929. Beesoniidae (neococcoids). *Limacoccini Foldi, 1995: 288. See note under Canceraspis.

Lithuanicoccus Koteja, 2008. Lithuanicoccidae (archaeococcoids). *Lithuanicoccidae Koteja: 2008: 33. This is an extinct family described from Baltic amber fossils.

Llaveia Signoret, 1876. Monophlebidae (archaeococcoids). Williams, 1969: 330.

Mahunkacoccus Kozár in Kozár & Konczné Benedicty, 2000. Caryonemidae (archaeococcoids). *Mahunkacoccinae Kozár in Kozár & Konczné Benedicty, 2000: 74. *Mahunkacoccini Kozár in Kozár & Koncné Benedicty, 2000: 72.

Marchalina Vayssière, 1923. Marchalinidae (archaeococcoids). Williams, 1969: 330. *Marchalinidae Morrison, Koteja, 1996: 68. The family rank of this group is now widely accepted.

Margarodes Guilding, 1829, Margarodidae (archaeococcoids) Williams, 1969: 330.

Matileortheziola Kozár & Foldi, 2000. Ortheziidae (archaeococcoids). Matileortheziolini Kozár, 2004: 432, 434. This tribe was placed originally in the subfamily Ortheziolinae but the tribal name has been treated as a synonym of the tribe Ortheziolini (Vea & Grimaldi, 2012: 780).

Matsucoccus Cockerell, 1909. Matsucoccidae (archaeococcoids). Williams, 1969: 331. *Matsucoccinae Morrison, Beardsley, 1968: 1458.

Melanaspis Cockerell, 1897. Diaspididae (neococcoids). Williams, 1969: 331.

Mempelaspis Takagi, 2000. Diaspididae (neococcoids). *Mempelaspidina Takagi, 2000: 53. The author placed this subtribe in the tribe Lepidosaphidini.

Micrococcus Leonardi, 1907. Micrococcidae (neococcoids). Williams, 1969: 331. *Micrococcidae Silvestri, Koteja, 1974a: 56. The family rank of this group is now widely accepted.

Mirococcopsis Borchesenius, 1948. Pseudococcidae (neococcoids). *Mirococcopsini Tang, 1992: 91. *Mirococcopsina Tang, 1992: 91. Spelling here corrected to Mirococcopsidina. *Mirococcopsidini Tang, Williams & Gullan, 2010: 67. This is a corrected spelling.

Mixorthezia Morrison, 1926. Ortheziidae (archaeococcoids). *Mixortheziini Kozár, 2004: 172. This tribe was placed in the subfamily Nipponortheziinae.

Monophlebulus Cockerell, 1902. Monophlebidae (archaeococcoids). Williams, 1969: 331.

Monophlebus Guérin-Méneville, 1827. Monophlebidae (archaeococcoids). Williams, 1969: 331. The family rank is now widely accepted.

Mytilaspis Targioni Tozzetti, 1868. Diaspididae (neococcoids). Williams, 1969: 331.

Mytilococcus Amerling, 1838. Diaspididae (neococcoids). Williams, 1969: 332.

Myzolecanium Beccari, 1877. Coccidae (neococcoids). *Myzolecaniinae Hodgson, 1994: 61. *Myzolecaniini Hodgson, Hodgson, 1994: 99.

Neomargarodes Green, 1914. Margarodidae (archaeococcoids). Williams 1969: 332.

Neosteingelia Morrison, 1927: 101. Kuwaniidae (archaeococcoids). *Neosteingeliini Tang & Hao, 1995: 74. This tribe was placed originally in the subfamily Kuwaniinae.

Newsteadia Green, 1902. Ortheziidae (archaeococcoids). *Newsteadiinae Kozár, 2004: 29, 31. *Newsteadiini Kozár, 2004: 35.

SCALE INSECT FAMILY-GROUP NAMES Zootaxa 3616 (4) © 2013 Magnolia Press · 335 Nipponorthezia Kuwana, 1916. Ortheziidae (archaeococcoids). Nipponortheziinae Kozár, 2004: 171. Nipponortheziini Kozár, 2004: 223.

Odonaspis Leonardi, 1897. Diaspididae (neococcoids). Williams, 1969: 332. *Odonaspidina Ferris, Tang, 1984: 187.

Orthezia Bosc d’Antic, 1784. Ortheziidae (archaeococcoids). Williams, 1969: 332. The family rank is generally accepted.

Ortheziola Šulc, 1895. Ortheziidae (archaeococcoids). *Ortheziolinae Kozár, 2004: 429. *Ortheziolini Kozár, 2004: 432, 495.

Ortheziolacoccus Kozár, 2004. Ortheziidae (archaeococcoids). *Ortheziolacoccini Kozár, 2004: 432, 438. This tribe was placed originally in the subfamily Ortheziolinae but the tribal name has now been treated as a synonym of Ortheziolini (Vea & Grimaldi, 2012: 780).

Ortheziolamameti Kozár, 2004. Ortheziidae (archaeococcoids). *Ortheziolamametiini Kozár, 2004, 432, 482. This tribe was placed originally in the subfamily Ortheziolinae but the tribal name has now been treated as a synonym of Ortheziolini (Vea & Grimaldi, 2012: 780).

Ovaticoccus Kloet, 1944. Eriococcidae (neococcoids). *Ovaticoccini Kozár & Konczné Benedicty, 2008: 223. The authors Kozár & Konczné Benedicty credited this family-group name to Miller & McKenzie (1967) and referred to Koteja (1974a), but Kozár & Konczné Benedicty were the first to use the name.

Palaeococcus Cockerell, 1894. Monophlebidae (archaeococcoids). Williams, 1969: 332.

Paralecanium Cockerell, 1894. Coccidae (neococcoids). Williams, 1969: 333.

Paraputo Laing, 1929. Pseudococcidae (neococcoids). *Paraputoina Tang 1992: 275, 276. This tribe is now placed in the subfamily Pseudococcinae (Williams & Gullan, 2010).

Parlatoreopsis Lindinger, 1912. Diaspididae (neococcoids). Williams, 1969: 333.

Parlatoria Targioni Tozzetti, 1868. Diaspididae (neococcoids). Williams, 1969: 333.

Pennygullania Koteja & Azar, 2008: 158. Pennygullaniidae (neococcoids). Pennygullaniidae Koteja & Azar, 2008: 158. This family was described from a Lebanon amber fossil. Gavrilov-Zimin & Danzig (2012) regard the genus Pennygullania to be one of the ancient genera of the family Pseudococcidae.

Phenacoccus Cockerell, 1893. Pseudococcidae (neococcoids). Williams, 1969: 333. *Phenacoccina Tang, 1992: 415.

Phenacoleachia Cockerell, 1899. Phenacoleachiidae (archaeococcoids). Williams, 1969: 333. The family rank of this group is generally accepted.

Phoenicococcus Cockerell, 1899. Phoenicococcidae (neococcoids). Williams, 1969: 334. The family rank is now widely accepted.

Physokermes Targioni Tozzetti, 1868. Coccidae (neococcoids). Williams, 1969: 334.

Pityococcus McKenzie, 1942. Pityococcidae (archaeococcoids). Williams, 1969: 334. *Pityococcinae McKenzie, Beardsley, 1969: 278. *Pityococcina McKenzie, Danzig, 1980: 94. *Pityococcidae McKenzie, Koteja, 1985: 492; 2000a: 196, 206. The family rank is now widely accepted.

Planococcus Ferris, 1950. Pseudococcidae (neococcoids). Williams, 1969: 334.

Platycoelostoma Morrison, 1923. Callipappidae (archaeococcoids). Williams, 1969: 324.

Pollinia Targioni Tozzetti, 1868. Asterolecaniidae (neococcoids). Williams, 1969: 324.

Porphyrophora Brandt, 1833. Margarodidae (archaeococcoids). Williams, 1969: 325. *Porphyrophorina Signoret, Tang & Hao, 1995: 10, 26, 602. The authors placed this subtribe in the tribe Margarodini in the family Margarodidae.

Protodiaspis Cockerell, 1898. Diaspididae (neococcoids). *Protodiaspidinae Varshney, 1985: 29 (nomen nudum). *Protodiaspidina Takagi, 1993: 21. *Protodiaspidinae Takagi, Varshney, 1995: 154. Varshney (1985) did not give a description of Protodiaspidinae and therefore Takagi becomes the author of this higher taxon because he was the first to provide a valid description (Takagi, 1993).

Pseudaonidia Cockerell, 1897. Diaspididae (neococcoids). Williams, 1969: 325.

Pseudococcus Westwood, 1840. Pseudococcidae (neococcoids). Williams, 1969: 325. *Pseudococcina Tang, 1992: 187, 218.

SCALE INSECT FAMILY-GROUP NAMES Zootaxa 3616 (4) © 2013 Magnolia Press · 337 The family rank is generally accepted. A proposal to give precedence of the family-group name Pseudococcini Cockerell, 1905, over Sphaerococcini Cockerell, 1899 (Hardy & Gullan, 2008), was agreed by the International Commission on Zoological Nomenclature (2009) (Opinion 2236).

Pseudopulvinaria Atkinson, 1889. Coccidae (neococcoids). Pseudopulvinariinae Tang, Hao, Xie & Tang, 1990: 75, 76. *Pseudopulvinariini Tang, Hao, Xie & Tang, Hodgson, 1994: 496.

Pulvinaria Targioni Tozzerti, 1866. Coccidae (neococcoids). Williams, 1969: 325. *Pulvinariina Targioni Tozzetti, Tang, Hao, Xie & Tang, 1990: 75, 76.

Puto Signoret, 1876. Putoidae (archaeococcoids). *Putoidae Beardsley, 1969: 278 (nomen nudum). *Putoini Tang, 1992: 382, 383. Putoidae Tang, Gill, 1993: 3. Although the family rank is widely accepted, there is some disagreement by Gavrilov-Zimin & Danzig (2012) about the placement of the genus Puto. These authors consider Puto to be a genus within the family Pseudococcidae.

Rhizoecus Künckel d’Herculais. Rhizoecidae (neococcoids). Williams, 1969: 325. *Rhizoecinae Williams, Koteja 1974a: 45. *Rhizoecina Williams, Tang, 1982: 44. *Rhizoecidae Williams, Hodgson, 2012: 8. The subtribe designation by Tang in 1982 preceded that of Kozár in Kozár & Konczné Benedicty (2007: 82). The family rank is recent.

Rhodesaclerda McConnell, 1954. Aclerdidae (neococcoids). *Rhodesaclerdinae Hodgson & Millar, 2002: 513.

Ripersiella Tinsley, 1899. Rhizoecidae (neococcoids). Ripersiellina Kozár in Kozár & Konczné Benedicty, 2007: 360. This subtribe was placed in the subfamily Rhizoecinae.

Ritsemia Lichtenstein, 1879. Pseudococcidae (neococcoids). *Ritsemimi Tang, 1992: 382, 591. This name is corrected here to Ritsemiini. This subtribe was placed in the subfamily Phenacoccinae.

Rugaspidiotus MacGillivray, 1921. Diaspididae (neococcoids). Williams, 1969: 336. *Rugaspidiotini, Balachowsky, Takagi, 1969: 7.

Saissetia Déplanche, 1867. Coccidae (neococcoids). *Saissetiini Hodgson, 1994: 50. *Saissetiinae Hodgson, Hodgson 2009: 57.

Sclopetaspis MacGillivray, 1921. Diaspididae (neococcoids). Williams, 1969: 336.

Serafinus Koteja, 2008. Serafinidae (archaeococcoids). *Serafinidae Koteja, 2008: 47. This family was described from Baltic amber fossils.

Serrolecanium Shinji, 1935. Pseudococcidae (neococcoids). Williams, 1969: 336. *Serrolecaniinae Shinji, Ferris, 1950: 72, corrected spelling instead of Serrolecaniniinae Shinji, 1935. *Serrolecaniina Shinji, Tang, 1992: 32. *Serrolecaniini Shinji, Hendricks & Kosztarab, 1999: ix, 25. Tang and Hendricks & Kosztarab placed these taxa in the subfamily Sphaerococcinae, now Pseudococcinae (Williams & Gullan, 2010: 11).

Signoretia Targioni Tozzetti, 1868. Coccidae (neococcoids). Williams, 1969: 336.

Smilacicola Takagi, 1969. Diaspididae (neococcoids). Smilacicolini Takagi, 2002: 59. The tribe was placed in the subfamily Aspidiotinae. Miller & Gimpel (2009: 853) list it in the subfamily Diaspidinae.

Sphaerococcus Maskell, 1892. Pseudococcidae (neococcoids). Williams, 1969: 337. *Sphaerococcinae Cockerell, Koteja 1974a: 44. *Sphaerococcina Cockerell, Han, Hao, Tang & Tang, 1992: 31. The family-group name Pseudococcini Cockerell, 1905, was given precedence over Sphaerococcini Cockerell, 1899, by the International Commission on Zoological Nomenclature (2009) (Opinion 2236), after a proposal by Hardy & Gullan (2008). The family-group name based on the genus Sphaerococcus is still available.

Steingelia Nassonov, 1908. Steingeliidae (archaeococcoids). Williams, 1969: 337. *Steingeliidae Morrison, Koteja & Żak-Ogaza, 1983: 470. *Steingeliina Morrison, Tang & Hao, 1995: 85, 707. The family rank is now widely accepted.

Stictococcus Cockerell, 1905. Stictococcidae (neococcoids). Williams, 1969: 337. The family rank for this group is generally accepted.

Stigmacoccus Hempel, 1900. Stigmacoccidae (archaeococcoids). Williams, 1969: 337. Stigmacoccidae Morrison, Hodgson & Foldi, 2006: 107. The family rank has now wide acceptance.

Tachardia Blanchard, 1886. Kerriidae (neococcoids). Williams, 1969: 337. *Tachardii Green, Varshney, 1966: 118. This subtribe was omitted in Williams (1969). It was erected in the family Tachardiidae presently accepted as the family Kerriidae but see note under Kerria.

Tachardina Cockerell, 1901. Kerriidae (neococcoids). Williams, 1969: 338.

Takahashia Cockerell, 1896. Coccidae (neococcoids). *Takahashina Tang, Hao, Xie & Tang, 1990: 75, corrected to Takahashiina on p. 76. This subtribe was placed in the tribe Pulvinariini.

Targionia Signoret, 1869. Diaspididae (neococcoids). Williams, 1969: 338.

Termitococcus Silvestri 1901. Margarodidae (archaeococcoids). Williams, 1969: 338.

Thysanaspis Ferris, 1955. Diaspididae (neococcoids). *Thysanaspidini Takagi, 2002: 58. Takagi placed this tribe in the subfamily Aspidiotinae. Miller & Gimpel (2009) list it in the subfamily Diaspidinae.

Trabutina Marchal, 1908. Pseudococcidae (neococcoids). Williams, 1969: 338. *Trabutinini Silvestri, Tang, 1992: 274 (corrected spelling). *Trabutinina Silvestri, Tang, 1992: 275, 319. These groups are now placed in the family Pseudococcinae (Williams & Gullan, 2010).

Trionymus Berg, 1899. Pseudococcidae (neococcoids). *Trionymini Tang, 1992: 120.

Ulucoccus Takagi, Tho & Khoo, 1990. Diaspididae (neococcoids). *Ulucoccinae Takagi, 2002: 59. This subfamily was erected for a single genus. It is one of three subfamilies listed in Miller & Gimpel (2009).

Walkeriana Signoret, 1876. Monophlebidae (archaeococcoids). *Walkerianina Tang & Hao, 1995, 121, 182, 616, 617. The authors placed this subtribe in the tribe Monophlebini.

Weitschatus Koteja, 2008. Weitschatidae (archaeococcoids). *Weitschatidae Koteja, 2008: 39. This family was described from Baltic amber fossils.

Xanthophthalma Cockerell & Parrott, 1899. Diaspididae (neococcoids). Williams, 1969: 338.

Xenococcus Silvestri, 1924. Rhizoecidae (neococcoids). *Xenococcini Tang, 1982: 43, 73. *Xenococcinae Tang, Hodgson, 2012: 51. Hodgson placed the subfamilies Xenococcinae and Rhizoecinae in the family Rhizoecidae.

Xerococcus Ferris, 1921. Eriococcidae (neococcoids). *Xerococini Tang & Hao, 1995: 425, 433. The authors erected this subfamily in the family Eriococcidae.

Xylococculus Morrison, 1927: 101. Xylococcidae (archaeococcoids). *Xylococculina Tang & Hao, 1995: 90, 608. The authors placed this subtribe in the tribe Xylococcini, presently in the family Xylococcidae.

Xylococcus Löw, 1883. Xylococcidae (archaeococcoids). Williams, 1969: 339. *Xylococcina Tang & Hao, 1995: 90. The authors included this subtribe in the tribe Xylococcini, presently in the family Xylococcidae. The family rank is now widely accepted.

Acknowledgements

San-an Wu, Beijing Forestry University, Beijing, China, has kindly checked the substantial works by Wang, T., Wang, T.C. and Xie, Y.P. on Chinese scale insects, works that have not been readily available to me. I am much indebted to Dr Wu for this kind gesture and for some of the literature not accessible to me. I am most grateful to Penny Gullan, Australian National University, Canberra, Australia, Danièle Matile-Ferrero, Muséum national d’Histoire naturelle, Paris, and Douglass R. Miller and Barbara Denno, Systematic Entomology Laboratory, U.S.D.A., Beltsville, Maryland, U.S.A., for their kind help in supplying or checking some of the literature. Penny Gullan and Douglass R. Miller have also read the manuscript to this work and made important comments and suggestions for which I extend my sincere thanks. I acknowledge with deep gratitude the help given by Chris Hodgson for checking the details throughout the manuscript and for general improvements to the text. My thanks are due also to Takumasa (Demian) Kondo and Giuseppina Pellizzari for reviewing the manuscript and for further recommendations, most of which I have implemented.

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