Family Cryptococcidae


Cryptococcus Douglas

NOMENCLATURE:

Cryptococcus Douglas, 1890a: 155. Type species: Coccus fagi Baerensprung (= Cryptococcus fagisuga Lindinger), by monotypy.

BIOLOGY: Adult females in bark crevices of deciduous trees, covered with fine, white, waxy secretion or felt-like ovisac. Apparently with one yearly generation. (Kozár, et al., 2013)

GENERAL REMARKS: The type species Coccus fagi Baerensprung, 1849 is a junior primary homonym of Coccus fagi Sulzer, 1776. This genus was placed in a separate family (Cryptococcidae) by Kosztarab (1968), but the characters given as diagnostic of the family are typical of other legless adult female eriococcids.

STRUCTURE: Adult female rotund (Miller & Miller, 1993). Adult females covered with fine, white, waxy secretion. Female oval, 0.5-1.0 mm; antenna normally 1 or 2, rarely 4 or 5 segmented; labium 3 segmented, basal segment without setae; legs absent, rarely reduced to unsegmented stubs; spiracles with sclerotized frame and a few associated quinquelocular pores; with a cluster pore plate below each hind spiracle; anal ring with 4 or 5 setae, rarely with pores; anal ring forming a broad plate surround by setae; macrotubular ducts present, except in one species; microtubular with simple pores, without cruciform pores; tubular ducts, normally two types on both surfaces. (Kozár, et al., 2013)

SYSTEMATICS: Slide-mounted adult female with: hind pair of legs represented by small sclerotized protrusion; other legs reduced or absent; antennae reduced; anal ring forming broad plate with setae surrounding plate; macrotubular ducts present, except in one species; microtubular ducts (Miller & Miller, 1993). In Kozár, et al., 2013 Cryptococcus was placed in the family Cryptococcidae Kosztarab, 1967.

KEYS: Henderson 2007a: 3-4 (female, adult) [Key to genera of Eriococcidae in New Zealand (adult females) Modified from Henderson (2006)]; Miller 2005: 491 (female) [Genera of Eriococcidae of the Eastern U.S.]; Kosztarab 1996: 262 (adult female) [Cryptococcus species of Northeastern North America]; Tang & Hao 1995: 427, 641 (adult female) [Cryptococcus species]; Miller, D. R. and Miller, G. L 1993: 68 (adult female) [All species of U.S. Cryptococcus]; Danzig 1971d: 821 (female) [Key to genera of Eriococcidae]; Kosztarab, M. and Hale, D. L. 1968: 11 (adult female) [All described species of Cryptococcus].

CITATIONS: AndersWuGr2010 [phylogeny, taxonomy: 996]; Arnett1985 [distribution, taxonomy: 239]; Balach1942 [taxonomy: 42]; Balach1948b [taxonomy: 254]; Beards1974 [taxonomy: 325]; BoratyWi1964 [taxonomy: 91]; Borchs1937 [taxonomy: 40]; Borchs1949 [description, taxonomy: 31, 42, 45, 370-371]; Borchs1958b [taxonomy: 772]; Cocker1894v [distribution, taxonomy: 1050]; Cocker1896b [taxonomy: 324]; Cocker1899j [taxonomy: 263]; Cocker1899m [taxonomy: 279]; Cocker1902g [taxonomy: 114]; Danzig1964 [distribution, taxonomy: 632, 634]; Danzig1971d [taxonomy: 821]; Danzig1986a [description, taxonomy: 271-272]; Danzig1988 [taxonomy: 707, 709]; Dougla1890a [taxonomy: 155]; Ferris1921b [taxonomy: 60]; Ferris1922b [taxonomy: 247]; Ferris1937 [taxonomy: 5]; Ferris1955a [description, distribution, taxonomy: 83]; Ferris1957c [taxonomy: 85]; Gill1993 [taxonomy: 153]; Green1915a [distribution, taxonomy: 180]; Green1922b [description, taxonomy: 21]; Green1928 [description, taxonomy: 9]; Hoy1962 [distribution, host, taxonomy: 6, 12, 13, 21, 25]; Hoy1963 [catalogue, taxonomy: 53-54]; Koszta1968 [taxonomy: 12]; Koszta1996 [description, distribution, taxonomy: 225, 261, 262]; KosztaHa1968 [taxonomy: 7]; KosztaKo1988F [catalogue, taxonomy: 269]; Koteja1974 [structure, taxonomy: 269, 275, 295, 300]; Koteja1974a [taxonomy: 249]; Koteja1974b [taxonomy: 78]; Kozar2009 [distribution, host, taxonomy: 114]; KozarDr1998e [catalogue, distribution, taxonomy: 167]; KozarKaKo2013 [description, distribution, host, illustration, structure, taxonomy: 594-607]; KozarKo2008a [taxonomy: 247-248]; KozarWa1985 [taxonomy: 75]; Lindin1923 [taxonomy: 140, 142, 146]; Lindin1932f [taxonomy: 188]; Lindin1937 [taxonomy: 183, 187]; MacGil1921 [distribution, host, taxonomy: 132, 147]; Miller1985b [taxonomy: 101]; Miller2005 [distribution,, taxonomy: 491]; MillerGi2000 [catalogue, taxonomy: 83-84]; MillerMi1993 [description, taxonomy: 68]; MorrisMo1966 [taxonomy: 50]; Newste1903 [description, taxonomy: 214]; PooleGe1997 [distribution: 354]; Schmut1952 [taxonomy: 405, 406, 417]; Sulc1895a [description, taxonomy: 13, 22]; TangHa1995 [description, distribution, taxonomy: 427]; Terezn1981 [distribution, taxonomy: 13, 46]; Terezn1982 [description, taxonomy: 35-36]; Willia1985h [distribution, host, taxonomy: 352]; Wise1977 [distribution, taxonomy: 96]; Xie1998 [taxonomy: 101]; Zahrad1959a [taxonomy: 538].



Cryptococcus aceris Borchsenius

NOMENCLATURE:

Cryptococcus aceris Borchsenius, 1937: 62. Type data: AZERBAIJAN: Abkhazia, Gagra District, Bzyb Gorge, on Acer sp., 1931. Lectotype female, by subsequent designation Danzig, 1996b: 521. Type depository: St. Petersburg: Zoological Museum, Academy of Science, Russia. Described: female. Illust. Notes: There are 5 female paralectotypes on the same slide as the lectotype.



HOSTS: Aceraceae: Acer pseudoplatanus [KosztaKo1988F], Acer sp. [Hoy1963]. Rosaceae: Pyrus sp. [KosztaKo1988F]. Tiliaceae: Tilia sp. [KosztaKo1988F]

DISTRIBUTION: Palaearctic: Azerbaijan [Hoy1963]; China [TangHa1995]; Germany [KosztaKo1988F]; Hungary [KosztaKo1988F, KozarKoFe2013]; Russia (Stavrapol Oblast [Danzig1985]); Ukraine (Zakarpat'ye (=Transcarpathia) Oblast [KosztaKo1988F]).

GENERAL REMARKS: Detailed description and illustration by Borchsenius (1949). Kosztarab & Kozár (1988) also provide a description.

STRUCTURE: Adult female hemispherical and yellow, completely enclosed within white sac (Borchsenius, 1949). Kosztarab & Kozár (1988) state that the female is covered by a white waxy secretion.

SYSTEMATICS: Slide-mounted adult female with: antennae 4-segmented; anal ring with 6 setae; 10 setae surrounding anal ring (Kosztarab & Kozár, 1988).

KEYS: Kozár et al. 2013: 596 (female) [Key to species of Cryptococcus]; Tang & Hao 1995: 427, 641 (adult female) [Cryptococcus species]; Kosztarab & Kozár 1988: 269 (adult female) [Cryptococcus species of Central Europe]; Kosztarab & Hale 1968: 11 (adult female) [Adult females of Cryptococcus]; Borchsenius 1949: 371 (adult female) [USSR Species of Cryptococcus].

CITATIONS: Borchs1937 [distribution, illustration: 62]; Borchs1939a [description, distribution, host, illustration: 43, 45, 50]; Borchs1949 [description, distribution, host, taxonomy: 48-49, 371-372]; Borchs1963a [distribution, host, taxonomy: 87, 156, 162, 164]; Borchs1973 [distribution, host, taxonomy: 87, 156, 164]; CookGu2004 [taxonomy: 444]; Danzig1971a [host, taxonomy: 1415]; Danzig1972 [distribution, taxonomy: 198]; Danzig1985 [taxonomy: 111]; Danzig1985 [distribution, taxonomy: 147]; Danzig1996a [distribution, host, taxonomy: 574]; Danzig1996b [distribution, host, taxonomy: 521]; Ferris1955a [taxonomy: 83]; Hadzib1950 [distribution, host, taxonomy: 256, 257, 263]; Hadzib1983 [distribution, host, taxonomy: 270]; Hoy1962 [distribution, host, taxonomy: 25]; Hoy1963 [catalogue, distribution, host, taxonomy: 54]; KosztaHa1968 [taxonomy: 11]; KosztaKo1988F [description, distribution, host, life history, taxonomy: 269-270]; Kozar2009 [distribution, taxonomy: 96]; KozarDr1998e [catalogue, distribution, host, taxonomy: 167]; KozarKaKo2013 [description, distribution, host, illustration, physiology, taxonomy: 597-599]; KozarKoFe2013 [distribution, taxonomy: 53]; KozarWa1985 [distribution: 75]; Kunkel1967 [taxonomy: 47]; Lagows2002 [distribution: 243]; LongoMaPe1999a [distribution: 145]; MillerGi2000 [catalogue, description, distribution, host, taxonomy: 84-85]; Schmut1955 [host, taxonomy: 160]; Schmut1980 [distribution, host, taxonomy: 50]; StoetzMi1979 [taxonomy: 4]; TangHa1995 [description, distribution, host, taxonomy: 427, 641]; Terezn1959b [p. 448]; TerGri1962 [distribution, host, taxonomy: 132, 156]; TerGri1969a [distribution, host, taxonomy: 79]; TerGri1983 [taxonomy: 882]; Willia1985h [distribution, host, taxonomy: 352]; Zahrad1972 [distribution, host: 405].



Cryptococcus fagisuga Lindinger

NOMENCLATURE:

Coccus fagi Baerensprung, 1849: 174. Type data: GERMANY: Berlin. Syntypes, female. Described: female. Homonym of Coccus fagi Sulzer 1776; discovered by Lindinger, 1936b: 444. Notes: Type material has probably been lost (Williams, 1985h).

Coccus fagi Walker, 1852: 1086. Type data: ENGLAND. Lectotype female, by subsequent designation Williams, 1985h: 353-356. Described: female. Synonymy by Lindinger, 1936: 444. Homonym. Notes: Coccus fagi Walker is a junior homonym of Coccus fagi Sulzer 1776 and Coccus fagi Baerensprung 1849.

Pseudococcus fagi; Douglas, 1886c: 152. Described: female. Change of combination.

Cryptococcus fagi; Douglas, 1890a: 155. Change of combination.

Eriococcus fagi; Perrier, 1926: 122. Described: female. Change of combination.

Cryptococcus fagisuga Lindinger, 1936b: 444. Replacement name for Coccus fagi Baerensprung 1849.

Kermes fagi; Lindinger, 1957: 549. Change of combination. Notes: Lindinger (1957) meant to reiterate the placement of fagi in Cryptococcus and he cites Guérin (1818) as the creator of the Kermes fagi combination. The date of this is probably wrong, and no publication by Guérin mentioning the combination can be found. Therefore, at least for the time being, this combination will be listed as by Lindinger (1957) even though it is not what he intended.

COMMON NAMES: beech bark scale [MalumpBa2012]; beech coccus [KozarDr1998e]; beech eriococcin [MillerMi1993]; beech scale [Westco1973]; felted beech coccus [Collin1908].



ASSOCIATES: HYPOCREALES Nectriaceae: Nectia galligena Bres. [GwiazdVaDe2006], Nectria coccinea faginata Lohm. & Watson [GwiazdVaDe2006].

FOES: Arachnida: Histiogaster entomophagus [KosztaKo1988F]. COLEOPTERA : Enicmus minutus [KosztaKo1988F]. Coccinellidae: Chilocorus bipustulatus [KosztaKo1988F], Chilocorus renipustulatus [KosztaKo1988F], Chilocorus stigma [KosztaKo1988F], Exochomus quadripustulatus [KosztaKo1988F]. HETEROPTERA Microphysidae: Microphysa pselaphiformis [KosztaKo1988F]. LEPIDOPTERA Psychidae: Psyche casta [KosztaKo1988F]. NEUROPTERA Chrysopidae: Chrysopa ciliata [KosztaKo1988F], Chrysopa ventralis [KosztaKo1988F].

HOSTS: Fagaceae: Fagus grandifolia [Hoy1963], Fagus orientalis [Hoy1963, Moghad2013a], Fagus sylvatica [Hoy1963]. Pinaceae: Pinus sylvestris [KosztaKo1988F].

DISTRIBUTION: Nearctic: Canada [Hoy1963] (New Brunswick [Ehrlic1932], Nova Scotia [Koszta1996], Ontario [MawFoHa2000], Prince Edward Island [Ehrlic1932]); United States of America (Connecticut [Hoy1963, MillerMi1993], Maine [Hoy1963, MillerMi1993], Massachusetts [Hoy1963, MillerMi1993], Michigan [WieferMcHa2011], New Hampshire [Hoy1963, MillerMi1993], New Jersey [Hoy1963, MillerMi1993], New York [Hoy1963, MillerMi1993], Ohio? [SinclaLyJo1987] (Although Sinclair et al. (1987) state that Cryptococcus fagisuga was found in Ohio, they give no reference and the species has not been recorded in Ohio by any other author known to us. We suspect that the Ohio record is erroneous.), Pennsylvania [Hoy1963, MillerMi1993], Rhode Island [Hoy1963, MillerMi1993], Tennessee [Koszta1996], Vermont [Hoy1963, MillerMi1993], Virginia [Koszta1996], West Virginia [Stimme1993]). Palaearctic: Armenia [Hoy1963]; Austria [Hoy1963]; Belgium [Hoy1963]; Bulgaria [KosztaKo1988F]; Croatia [MastenSi2008]; Czechoslovakia [Hoy1963]; Denmark [Hoy1963, Gertss2001]; France [Hoy1963, Foldi2001]; Georgia [Hoy1963]; Germany [Hoy1963]; Hungary [KosztaKo1988F]; Iran [KozarFoZa1996]; Italy [KosztaKo1988F]; Netherlands [Hoy1963, Jansen2001]; Poland [Hoy1963, SimonKa2011]; Romania [Hoy1963]; Russia [Hoy1963] (Stavrapol Oblast [Danzig1985]); Sicily [LongoMaPe1995]; Slovenia [KozarKaKo2013]; Sweden [Hoy1963, Gertss2001]; Switzerland [Hoy1963]; Turkey [Hoy1963]; Ukraine (Krym (=Crimea) Oblast [Mateso1958], Zakarpat'ye (=Transcarpathia) Oblast [Terezn1959]); United Kingdom [Hoy1963, MalumpBa2012] (Channel Islands [Hoy1963], England [Parker1974]); Yugoslavia [KosztaKo1988F].

BIOLOGY: The general pattern of life cycle is as follows. Yellow eggs are laid in the spring or early summer within the ovisac. Eggs hatch in late summer and early fall. Crawlers are quite mobile and may move some distance before settling on the bark. The first instar is usually the overwintering stage. During the winter, yellow first instar nymphs change in body form from oval to rotund. Molting occurs during the spring and the short lived apodous second instar is produced. Adult females occur throughout the spring and summer. Males have never been collected (Miller & Miller, 1993). In adult females of C. fagisuga having restricted access of oxygen to apiracles the spiracles are located in deep cavities of scleotised walls which probably contain airr reserves. (Podsiadlo, 2006)

GENERAL REMARKS: Detailed description and illustration by Miller & Miller (1993). Description of first and second instars and adult female by Williams (1985h). Discussion of population dynamics relating to physiological defense reactions of attacked trees by Gora et al. (1996).

STRUCTURE: Adult female body is nearly hemispherical, bright yellow and covered with a filamentous white sac (Miller & Miller, 1993).

SYSTEMATICS: Slide-mounted adult female with: antennae 1- to 3-segmented; anal ring with 4 setae; 6 or 8 setae surrounding anal ring (Kosztarab & Kozár, 1988; Miller & Miller, 1993); anal ring with small pores; multiloculars absent from dorsum; macrotubular ducts of 2 sizes (Miller & Miller, 1993). This species was first described as Coccus fagi Baerensprung (1849). The next description was as Coccus fagi Walker (1852). C. fagi Baerensprung is the junior primary homonym of C. fagi Sulzer (1776). C. fagi Walker is a synonym of C. fagi Baerensprung and a junior primary homonym of C. fagi Sulzer. The next three names Pseudococcus fagi Douglas (1886c), Cryptococcus fagi Douglas (1890) and Eriococcus fagi Perrier (1926) are change of combinations. The last and final change is to Cryptococcus fagisuga Lindinger (1936) who realized the homonymy of the species (Miller & Miller, 1993). C. fagisuga is part of a clade that includes one other Palearctic species (P. fraxini), one Nearctic species (C. williamsi), and two species endemic to New Zealand (C. nudatus and M. totarae). Since this clade is nested within a larger "Gondwanan" clade consisting of species from Australia and New Zealand, it appears to have originated in New Zealand and to have somehow spread to the North Temperate Zone at an ancient date. (Gwiazdowski et al., 2006)

ECONOMIC IMPORTANCE AND CONTROL: This scale, together with a fungus, can cause extensive damage to the native beech trees in the northeastern United States (Miller & Miller, 1993). Van Driesche & Japoshvili (2012) found that scale densities were 45.4-fold higher per unit area of bark in Massachusetts on F. grandifolia than in their native range in the country of Georgia on F. orientalis. Also, F. orientalis trees at sample sites in Georgia were 2.9-fold larger in DBH and much healthier that were F. grandifolia trees in Massachusetts.

KEYS: Kozár et al. 2013: 596 (female) [Key to species of Cryptococcus]; Kosztarab 1996: 262 (adult female) [Cryptococcus species of Northeastern North America]; Tang & Hao 1995: 427, 641 (adult female) [Cryptococcus species]; Miller & Miller 1993: 68 (adult female) [Adult female of Cryptococcus]; Kosztarab & Kozár 1988: 269 (adult female) [Cryptococcus species of central Europe]; Tereznikova 1982: 36 (adult female) [as Cryptococcus fagi; Cryptococcus species]; Danzig 1971d: 824 (female) [Key to species of family Eriococcidae]; Kosztarab & Hale 1968: 11 (adult female) [Adult females of Cryptococcus]; Danzig 1964: 635 (adult female) [Cryptococcus species of the USSR]; Borchsenius 1949: 371 (adult female) [USSR species of Cryptococcus].

CITATIONS: AAEE1930 [taxonomy: 639]; Apstei1915 [taxonomy: 158]; Arnett1985 [distribution, taxonomy: 239]; Baeren1849 [description: 174]; Baker1972 [distribution, host, life history, illustration: 97-98]; Balach1937c [distribution, taxonomy: 6]; BarbagBiBo1995 [distribution: 43]; BarethVa1976 [description, distribution, host, illustration, structure: 209-221]; Blicke1965 [taxonomy: 289, 306]; Bodenh1935 [taxonomy: 271]; Bodenh1953a [distribution, host: 134, 153]; BoratyWi1964 [taxonomy: 91]; Borchs1934 [distribution, host: 12]; Borchs1937 [distribution, illustration: 62]; Borchs1939a [taxonomy: 43, 45]; Borchs1949 [description, distribution, host, taxonomy: 371]; Britti1915a [description, taxonomy: 157]; CCNI1989 [taxonomy: 158]; CebeciKu2005 [distribution, host: 97-102]; Cocker1896b [taxonomy: 324]; Collin1908 [distribution: 23]; CookGuSt2000 [taxonomy: 889]; Danzig1964 [taxonomy: 635]; Danzig1971a [host, taxonomy: 1415]; Danzig1971d [taxonomy: 824]; Danzig1985 [taxonomy: 111]; Davids1974 [chemical control, distribution, host: 3]; Dougla1886c [description, distribution, host, taxonomy: 152]; Dougla1890a [taxonomy: 155]; Ehrlic1932 [chemical control, distribution, host, life history, taxonomy: 75-80]; Ehrlic1934 [biological control, chemical control, description, distribution, economic importance, host, life history, taxonomy: 593-692]; Fernal1903b [catalogue, distribution, host, taxonomy: 120]; Ferris1955a [description, distribution, host, taxonomy: 83]; Ferris1957c [taxonomy: 85]; Foldi2001 [distribution, economic importance: 305, 307]; ForresMcMi2003 [economic importance: 70]; Germai2011 [distribution, economic importance: 31-34]; Germai2011a [distribution, economic importance: 8]; Gertss1997 [distribution, host, illustration: 113, 114]; Gertss2000 [distribution, host: 151]; Gertss2001 [distribution: 126]; Gertss2011 [distribution: 40]; GoraKoLu1996 [ecology, host: 112-120]; GordonHi1990 [biological control: 287]; Gottsc1995 [host: 222]; Goux1931 [distribution, host: 332]; Green1922b [description, distribution, host, taxonomy: 21]; Green1925b [distribution, host: 517]; Green1927a [distribution: 29]; Green1928 [description, host, taxonomy: 9]; GullanKo1997 [host: 42]; GullanMa2003 [economic importance: 1089]; GwiazdVaDe2006 [distribution, economic importance, host, molecular data, phylogenetics, taxonomy: 9-18]; Hadzib1983 [distribution, host, taxonomy: 270]; Harris1916a [distribution: 94]; Hendry1962 [taxonomy: 127]; Henrik1921 [distribution, host, taxonomy: 313]; Herric1935 [description, distribution, host, taxonomy: 45]; Hopkin1904 [distribution, host: 51, 52]; Housto1994 [distribution, ecology, host: 75-87]; Hoy1962 [distribution, taxonomy: 25]; Hoy1963 [catalogue, distribution, host, taxonomy: 54-55]; Jaap1914 [taxonomy: 136]; Jansen2001 [distribution: 200]; Johnso1982 [chemical control, description, economic importance, illustration,: 114, 116, 122]; JohnsoLy1976 [description, distribution, host, illustration, life history: 286-287]; Jonsso1998 [ecology, distribution, host: 297-305]; Kaweck1935 [host: 74]; Kaweck1985 [distribution, taxonomy: 33-34]; KaydanUlEr2007 [distribution, host: 90-106]; Kiritc1931 [description, distribution, host, taxonomy: 312]; Kiritc1935 [distribution, host: 1]; Koszta1959 [distribution, host: 402]; Koszta1968 [taxonomy: 12]; Koszta1996 [biological control, description, distribution, economic importance, host, illustration, life history, taxonomy: 262-264]; KosztaHa1968 [taxonomy: 11]; KosztaKo1988F [biological control, description, distribution, host, illustration, life history, taxonomy,: 270-272]; KosztaRh1999 [distribution, economic importance, host: 121]; Koteja1971a [distribution, host: 322]; Koteja1974b [structure: 272]; Koteja1974b [taxonomy: 78]; Koteja1983a [distribution, host: 676]; Koteja2000a [distribution: 172]; Koteja2000d [distribution: 242]; KotejaZa1966 [distribution, host: 310, 320]; KotejaZa1969 [distribution, host, taxonomy: 362]; KotejaZa1983 [distribution, host, taxonomy: 478]; Kozar1980 [distribution, host, taxonomy: 66, 67]; Kozar1983a [distribution, host, taxonomy: 146]; Kozar1991 [taxonomy: 81]; Kozar1999a [distribution, host: 139]; Kozar2009 [distribution, taxonomy: 96]; KozarDr1998e [catalogue, distribution, host, taxonomy: 167-168]; KozarFoZa1996 [distribution: 65]; KozarGuBa1994 [distribution, host, taxonomy: 154]; KozarKaKo2013 [description, distribution, host, illustration, structure, taxonomy: 600-603]; KozarKiSa2004 [distribution: 60]; KozarKo1982 [distribution, economic importance, host, taxonomy: 204, 210]; KozarKo2002b [distribution: 375]; KozarKoFe2013 [distribution, taxonomy: 53]; KozarWa1985 [distribution: 75]; Kozarz1986 [distribution, taxonomy: 306, 307]; KozarzRe1975 [distribution, economic importance, host, taxonomy: 7, 16]; Lagows2002 [distribution: 243]; LagowsKo1996 [distribution: 31]; Lindin1907d [taxonomy: 158]; Lindin1912b [host, taxonomy: 152]; Lindin1932f [taxonomy: 204]; Lindin1935 [taxonomy: 135]; Lindin1936a [taxonomy: 444]; Lindin1938 [distribution, taxonomy: 4-5]; Lindin1957 [taxonomy: 549]; Lindin1958 [taxonomy: 367]; LongoMaPe1995 [distribution: 121]; Lunder1998 [biological control, distribution, host: 319-322]; MalumpBa2012 [distribution: 26]; Marcha1908 [description, distribution, host, illustration, taxonomy: 263]; MastenSi2008 [catalogue, distribution, host: 105-119]; MawFoHa2000 [distribution: 43]; Miller1985b [distribution, host, life history: 101-102]; Miller1991b [economic importance: 101]; Miller2005 [distribution: 491]; MillerGi2000 [biological control, catalogue, description, distribution, economic importance, host, taxonomy: 85-89]; MillerMi1993 [description, distribution, host, illustration, taxonomy: 68-70]; MillerWi1995aDR [taxonomy: 200, 242]; Moghad2013a [distribution, host: 55]; MoraalJa2011 [ecology: 53, 57]; NanDeWu2013 [phylogenetics, taxonomy: 171, 173-174]; NastChKl1990 [distribution, taxonomy: 121]; Newste1900a [description, distribution, host, illustration, taxonomy: 249]; Newste1903 [description, distribution, host, illustration, taxonomy: 215-221]; Nikols1936 [taxonomy: 155]; Nitsch1895 [description, distribution, host, illustration, taxonomy: 1250-1252]; Ossian1951 [distribution, taxonomy: 4]; OuvrarKo2009 [host, phylogeny, taxonomy: 101-115]; Parker1974 [chemical control, distribution, host, taxonomy: 3-15]; PellizKo2011 [distribution: 66]; Perrie1926 [description, distribution, taxonomy: 122]; Perrin1980 [biological control, distribution, economic importance, host: 319-331]; Pierce1917 [distribution, economic importance, host: 39]; Piltz1952 [distribution, taxonomy: 304]; Podsia2006a [illustration, physiology: 277-281]; PooleGe1997 [distribution: 354]; Reh1903 [description, distribution, host, taxonomy: 351-353]; Reh1904 [taxonomy: 34]; Reh1926 [taxonomy: 322]; Reyne1961 [taxonomy: 138]; RossHaOk2012 [phylogeny, taxonomy: 199]; Schmut1952 [description, distribution, host, illustration, taxonomy: 378, 406, 417-418]; Schmut1955 [host, taxonomy: 159, 160]; Schmut1974 [host, taxonomy: 49]; Signor1877 [description, host, taxonomy: 615]; SimonKa2011 [distribution: 237]; Step1929 [distribution, host: 183]; Stimme1993 [chemical control, description, distribution, economic importance, host, illustration, life history, taxonomy: 17-18]; Stocki1998 [distribution, host: 89, 91, 97]; StoetzMi1979 [taxonomy: 15]; Storka1925 [distribution, taxonomy: 70]; Sulc1895a [description, distribution, taxonomy: 13, 22]; Sulc1912 [distribution, taxonomy: 34]; Suvak1999 [description, distribution, economic importance, host, taxonomy: 73-83]; SwaineHu1926 [description, distribution, host: 47-48]; Szklar1998 [host, illustration, physiology: 168, 169, 171]; TangHa1995 [description, distribution, host, taxonomy: 416, 427, 641]; TealeLeMa2009 [ecology: 1235-1240]; Terezn1959 [distribution: 683-685]; Terezn1959b [distribution: 447]; Terezn1959d [taxonomy: 92]; Terezn1966 [distribution, host: 27]; Terezn1967a [distribution: 474, 475]; Terezn1975 [taxonomy: 29]; Terezn1981 [taxonomy: 46]; Terezn1982 [distribution, illustration, taxonomy: 36]; TerGri1983 [distribution, taxonomy: 882]; TranfaMa1988 [taxonomy: 609]; Tsalev1968 [distribution, host: 207]; VanDriJa2012 [distribution, ecology, host: 422-427]; Walczu1932 [taxonomy: 625]; Walker1852 [description, distribution: 1086]; Westco1973 [chemical control, description, distribution, host, life history,: 388-389]; WieferMcHa2011 [description, distribution, host, life history: 173-179]; Willia1985h [description, distribution, host, illustration, taxonomy: 353-356]; WilliaBe2009 [catalogue: 21]; Wunn1925 [distribution, host: 122]; Wunn1925b [description, distribution, host, taxonomy: 281, 284]; Wunn1925c [distribution, host: 438]; Zahrad1959a [host, taxonomy: 538]; Zahrad1972 [biological control, description, distribution, host, illustration, taxonomy: 403-404]; Zahrad1977 [taxonomy: 121]; ZakOgaKo1964 [distribution, host: 418].



Cryptococcus integricornis Danzig

NOMENCLATURE:

Cryptococcus integricornis Danzig, 1971a: 1415. Type data: RUSSIA: Primorsky Kray, Suchan (Partizansk) District, Tigrovoy, on Tilia amurensis, 29/06/1963, E. Danzig. Holotype female, by original designation. Type depository: St. Petersburg: Zoological Museum, Academy of Science, Russia; type no. 181-64. Described: female. Illust. Notes: 1 female paratype on same slide as holotype. 2 female paratypes on 1 slide with same data. 2 paratypes on 1 slide RUSSIA: Primorsky Kray, Suputinsky (Ussuriysky) Reserve, on Tilia amurensis, 12/07/1969, E. Danzig; all in ZMAS (Danzig, personal communication, 1996)



HOST: Tiliaceae: Tilia amurensis [Danzig1986a].

DISTRIBUTION: Palaearctic: China [TangHa1995]; Russia (Primor'ye Kray [Danzig1986a]).

GENERAL REMARKS: Detailed description and illustration by Danzig (1986a).

STRUCTURE: Adult female pink, body hemispherical. Nymphs and females grow in small white silky wax sacks. Colonies observed in late June and early July consisted of nymphs and adult females (Danzig, 1986a).

SYSTEMATICS: Slide-mounted adult female with: with 3 pairs of legs; anal ring with pores and 6 setae; macrotubular and microtubular ducts; antennae 4-segmented (Danzig, 1986a).

KEYS: Kozár et al. 2013: 596 (female) [Key to species of Cryptococcus]; Tang & Hao 1995: 427, 642 (adult female) [Cryptococcus species].

CITATIONS: BarethVa1976 [taxonomy: 210]; Danzig1971a [description, distribution, host, illustration, taxonomy: 1415]; Danzig1977b [distribution, host, taxonomy: 44, 51]; Danzig1980b [distribution, host, taxonomy: 235]; Danzig1986a [description, distribution, host, illustration, taxonomy: 272-273]; Danzig1988 [taxonomy: 709]; Kozar2009 [distribution, taxonomy: 96]; KozarDr1998e [catalogue, distribution, host, taxonomy: 168]; KozarKaKo2013 [description, distribution, host, illustration, structure, taxonomy: 604-606]; KozarWa1985 [distribution: 75]; MillerGi2000 [catalogue, description, distribution, host, taxonomy: 89-90]; TangHa1995 [description, distribution, host, taxonomy: 427, 428, 642]; Willia1985h [distribution, host, taxonomy: 352].



Cryptococcus nudatus Brittin

NOMENCLATURE:

Cryptococcus nudata Brittin, 1915a: 160. Type data: NEW ZEALAND: South Island, Cashmere Hill, Christchurch, on Hoheria sp. Syntypes, female. Type depository: Auckland: New Zealand Arthropod Collection, Landcare Research, New Zealand. Described: female. Illust.

Kuwanina parva; Green, 1916d: 52. Misidentification; discovered by Hoy, 1962: 26.

Chaetococcus parvus; Thomson, 1922: 338. Misidentification; discovered by Hoy, 1962: 25-26.



HOSTS: Malvaceae: Hoheria populnea? [Hoy1962], Hoheria sexstylosa [Hoy1962], Hoheria sp. [Britti1915a]

DISTRIBUTION: Australasian: New Zealand (South Island [Hoy1963]).

GENERAL REMARKS: Detailed description and illustration by Hoy (1962).

STRUCTURE: According to Hoy (1962), "...females in crevices in bark on main trunk of tree. Some associated loose white wax but female not forming a sac."

SYSTEMATICS: Slide-mounted adult female with: front 2 pairs of legs absent; no macrotubular ducts; anal ring with pores and 6 setae; quinquelocular pores on dorsum and venter (Hoy, 1962).

KEYS: Kosztarab & Hale 1968: 11 (adult female) [Adult female of Cryptococcus].

CITATIONS: Britti1915a [taxonomy: 157, 160]; Danzig1971a [distribution, host, taxonomy: 1415]; Fernal1903b [catalogue, taxonomy]; Green1916d [taxonomy: 52]; GwiazdNo2008 [phylogenetics: 10-17]; Hoy1962 [description, distribution, host, illustration, taxonomy: 25, 26-27]; Hoy1963 [catalogue, distribution, host, taxonomy: 55]; KosztaHa1968 [taxonomy: 11]; Koteja1974b [taxonomy: 78]; Koteja1976 [structure: 272]; Kozar2009 [distribution, taxonomy: 96]; Lindin1932f [taxonomy: 188]; MillerGi2000 [catalogue, description, distribution, host, taxonomy: 90]; NanDeWu2013 [phylogenetics, taxonomy: 171, 173]; Thomso1922 [distribution, host, taxonomy: 338]; Willia1985h [taxonomy: 353]; Wise1977 [distribution, taxonomy: 96].



Cryptococcus williamsi Kosztarab & Hale

NOMENCLATURE:

Cryptococcus williamsi Kosztarab & Hale, 1968: 7. Type data: UNITED STATES: Vermont, N. of Perry, on Acer sp., 01/11/1966, by R.L. Murray. Holotype female (examined), by original designation. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female. Illust. Notes: Type material also at CDAE.

COMMON NAME: maple bark scale [Koszta1996].



FOE: HYMENOPTERA Encyrtidae: Coccophagiodes sp. [Koszta1996].

HOSTS: Aceraceae: Acer rubrum [Koszta1996], Acer saccharum [Koszta1996], Acer sp. [MillerMi1993]

DISTRIBUTION: Nearctic: Canada (Ontario [Koszta1996], Quebec [Koszta1996]); United States of America (Iowa [MillerMi1993], Maine [MillerMi1993], New Hampshire [MillerMi1993], New York [MillerMi1993], Vermont [MillerMi1993], Virginia [MillerMi1993]).

BIOLOGY: In spring and early summer, crawlers are either born as nymphs or hatch from eggs. Adult female is apparently the overwintering stage and there is 1 generation per year. 20-30 eggs are laid per female (Kosztarab, 1996). The life cycle is carried out in bark crevices of the maple host (Kosztarab & Hale, 1968).

GENERAL REMARKS: Original description and illustration by Kosztarab & Hale (1968). Detailed description and illustration also by Miller & Miller (1993).

STRUCTURE: Adult female is salmon orange, enclosed in a filamentous ovisac. A filamentous secretion encloses all stages, but may be worn off on some individuals (Kosztarab, 1996).

SYSTEMATICS: Slide-mounted adult female with: anal ring without pores; multiloculars present on dorsum; macrotubular ducts of 1 size (Miller & Miller, 1993).

ECONOMIC IMPORTANCE AND CONTROL: This species can be a pest on sugar maple trees (Kosztarab & Hale, 1968).

KEYS: Kosztarab 1996: 262 (adult female) [Cryptococcus species of Northeastern North America]; Miller & Miller 1993: 68 (adult female) [Cryptococcus species of the Eastern United States].

CITATIONS: Arnett1985 [distribution, taxonomy: 239]; BarethVa1976 [taxonomy: 210, 219]; Baylac1980 [biological control, distribution, host, taxonomy: 207]; Danzig1971a [host, taxonomy: 1415]; Dziedz1977 [taxonomy: 59]; GwiazdVaDe2006 [biological control, phylogenetics: 10-17]; HowellWi1976 [distribution, host, taxonomy: 187]; Koszta1996 [biological control, description, distribution, economic importance, host, illustration, life history, taxonomy: 262, 264-266]; KosztaHa1968 [description, distribution, host, illustration, taxonomy: 7-11]; KosztaRh1999 [distribution, economic importance, host: 122]; Koteja1974b [taxonomy: 78]; Kozar2009 [distribution, taxonomy: 96]; KozarKaKo2013 [description, distribution, host, illustration, structure, taxonomy: 613-615]; Mason1977 [biological control, distribution, host, taxonomy: 490-191]; MawFoHa2000 [distribution: 43]; Miller1985b [distribution, host: 102-103]; Miller2005 [distribution: 491]; MillerGi2000 [catalogue, description, distribution, economic importance, host, life history, taxonomy: 91-92]; MillerMi1993 [description, distribution, host, illustration, taxonomy: 70-72]; NanDeWu2013 [phylogenetics, taxonomy: 171, 173]; PooleGe1997 [distribution: 354]; Weidha1968 [taxonomy: 255]; Willia1985h [distribution, host, taxonomy: 352].



Pseudochermes Nitsche

NOMENCLATURE:

Pseudochermes Nitsche, 1895: 1247-1249. Type species: Chermes fraxini Kaltenbach, by original designation.

Apterococcus Newstead, 1898: 97. Type species: Ripersia fraxini Newstead, by monotypy and original designation. Synonymy by Lindinger, 1937: 179, 185.

STRUCTURE: antennae 6 segmented. Frontal tubercle present. Labium 3 segmented. Stylet loop much longer than body. Quinquelocular pores present on both surfaces in small number. Legs well developed.

SYSTEMATICS: Generic characteristics that distinguish this genus from all other eriococcids are: anal ring crescent shaped; frontal tubercles present; dorsal setae slightly enlarged posteriorly; multilocular pores present on dorsum (Williams, 1985h). The nearest genus to Pseudochermes seems the Rhopalotococcus Williams, 2007 from New Caledonia by having only one pair of setae on basal segment of labium or basal segment not developed.In Palaearctic Region the Borchseniococcus Kayden and Kozár, 2008 and Hispaniococcus are similar to Pseudochermes by sclerotized and reduced anal ring, not well developed anal lobes, and by the presence of frontal tubercles. Both genera differ by two pairs of setae on the basal segment of the labium. In Kozár, et al., 2013 Pseudochermes was placed in the family Cryptococcidae Kosztarab, 1967.

KEYS: Danzig 1988: 709 (adult female) [Pseudochermes species of far East USSR]; Williams 1985h: 352 (female) [Key to genera of British Eriococcidae].

CITATIONS: BoratyWi1964 [taxonomy: 92]; Borchs1937 [taxonomy: 40, 58]; Borchs1949 [taxonomy: 43, 322, 365]; Cocker1899a [taxonomy: 392]; Cocker1899m [taxonomy: 278]; Danzig1964 [taxonomy: 632, 634]; Danzig1980b [taxonomy: 231]; Danzig1986 [description, taxonomy: 269]; Danzig1988 [taxonomy: 707, 709]; Fernal1903b [catalogue, taxonomy: 114]; Ferris1921b [taxonomy: 60]; Ferris1922b [taxonomy: 246]; Ferris1937 [taxonomy: 5]; Ferris1955a [taxonomy: 178]; Ferris1957c [distribution, taxonomy: 88]; GomezM1937 [taxonomy: 322, 360]; GomezM1948 [taxonomy: 97]; Green1922 [distribution: 345]; Green1922b [description, taxonomy: 20-21]; Hoy1962 [taxonomy: 13, 201]; Hoy1963 [catalogue, taxonomy: 187-188]; JudeicNi1895 [taxonomy: 1247-1249]; Kiritc1940 [taxonomy: 123]; Kohler1998 [catalogue, distribution, taxonomy: 394-395]; KosztaKo1978 [taxonomy: 67]; KosztaKo1988F [catalogue, distribution, taxonomy: 272]; Koteja1974 [taxonomy: 295]; Koteja1974b [taxonomy: 78]; KotejaZa1981 [taxonomy: 502]; Kozar2009 [distribution, host: 111]; KozarKaKo2013 [description, distribution, host, taxonomy: 607-609]; KozarKaKo2013 [description, distribution, host, illustration, structure, taxonomy: 608-615]; KozarKo2008a [description, taxonomy: 247-250]; KozarWa1985 [distribution: 75]; Lindin1908 [taxonomy: 94]; Lindin1933 [description, taxonomy: 31]; Lindin1937 [taxonomy: 179, 185, 194]; MacGil1921 [taxonomy: 129]; MillerGi2000 [catalogue, taxonomy: 443]; MorrisMo1966 [taxonomy: 164]; Newste1898 [taxonomy: 97]; Newste1903 [description, taxonomy: 209-210]; Nitsch1895 [description, taxonomy: 1247-1249]; Schmut1952 [taxonomy: 418]; Sulc1912 [taxonomy: 33]; Terezn1981 [taxonomy: 13, 58]; Willia1985h [taxonomy: 385-386]; Willia2007a [structure: 1357]; Zahrad1959a [taxonomy: 539].



Pseudochermes betula (Wu & Liu)

NOMENCLATURE:

Kuwanina betula Wu & Liu, 2009: 221-223. Type data: CHINA: Henan Province, Songxian County, Mt. Baiyun, (34.08şN, 112.05şE) on Betula ablo-sinensis Burkill, 8/15/2008, by S. Wu & J. Liu. Holotype female (examined). Type depository: Beijing: Forestry University, Beijing, China. Described: female. Illust.

Kuwanina betulae; Kozár et al., 2013: 642. Misspelling of species name.

Pseudochermes betula; Wu & Nan, 2015: 576–584. Change of combination.



HOST: Betulaceae: Betula ablo-sinensis Burkill [WuLi2009].

DISTRIBUTION: Palaearctic: China (Henan (=Honan) [WuLi2009]).

BIOLOGY: It lives in the crevices of bank on the main trunk of Betula ablo-sinensis. (Wu & Liu, 2009)

GENERAL REMARKS: Detailed description and illustration in Wu & Liu, 2009. The immature stages are described and illustrated in Wu & Nan, 2015.

STRUCTURE: In life, adult female hemispherical or spherical, pink, covered with white cottony secretion of fine waxy threads. Mounted specimen is circular. Anal lobe absent, antenna stub-like, 3 segmented; dypeolabral shield nearly rectangular. Fore and mid-legs absent, each hind leg reduced to a transverse oval pore plate; spiracles surrounded by a sclerotized plate. Vulva distinct, circular. (Wu & Liu, 2009)

SYSTEMATICS: Nan et al. (2013) studied the relationships among eriococcids and related scale insects, including K. betula, based on molecular data. In two phylogenetic trees, based on 18S and 28S rDNA sequences, K. betula always fell within the "Gondwanan" clade of Cook and Gullan (2004), along with Cryptococcus spp. (including the type species C. fagisuga Lindinger), Pseudochermes fraxini (Kaltenbach) and members of the Acanthococcidae, showing a strong sister relationships with P. fraxini with 100% Bayesian posterior probability support. The first-instar nymphs of K. betula are most similar to those of P. fraxini in the presence of frontal tubercles, and the adult females are more similar to those of C. fagisuga. Based on these similarities and molecular relationships of K. betula, plus the presence of 6-locular pores around the inner orifice of outer part of the macrotubular ducts in Pseudochermes williamsi Kozár & Konczné Benedicty (Kozár & Konczné Benedicty, 2008), which is similar to that found in adult female K. betula, Wu and Nan transferred K. betula to Pseudochermes.(Wu & Nan, 2015)

KEYS: Wu & Nan 2015: 579 (female, other) [Key to female instars of Pseudochermes betula.]; Kozár et al. 2013: 641 (female) [as Kuwanina betulae; Key to species of Kuwanina]; Wu & Liu 2009: 221 (female) [as Kuwanina betula; Key to Kuwanina species].

CITATIONS: KozarKaKo2013 [description, distribution, host, illustration, structure, taxonomy: 642-643]; NanDeWu2013 [phylogenetics, taxonomy: 171, 173-174]; WuLi2009 [description, distribution, host, illustration, taxonomy: 221-223]; WuNa2015 [description, distribution, illustration, molecular data, structure, taxonomy: 576–584].



Pseudochermes fraxini (Kaltenbach)

NOMENCLATURE:

Chermes fraxini Kaltenbach, 1860: 259. Type data: GERMANY. Syntypes, female. Notes: Williams (1985h) states that the type material, which was held in Germany, is probably lost.

Eriococcus fraxini Newstead, 1891: 165-166. Type data: ENGLAND: Cheshire, Ince, on Fraxinus excelsior, ?/08/1890, by R. Newstead. Lectotype female, by subsequent designation Williams, 1985h: 386-389. Type depository: London: The Natural History Museum, England, UK. Synonymy by Judeich & Nitsche, 1895: 1249. Notes: In addition to the lectotype, there are nine paralectotypes on the same slide in the BMNH (Williams, personal communication, June 19, 1996).

Ripersia fraxini; Newstead, 1892: 147. Change of combination.

Coccus fraxini; Judeich & Nitsche, 1895: 1247. Illust. Change of combination.

Pseudochermes fraxini; Nitsche, 1895: 1247-1249. Change of combination.

Fonscolombia fraxini; Cockerell, 1899j: 264. Change of combination.

Apterococcus fraxini; Newstead, 1903: 210-213. Described: both sexes. Change of combination.

Fonscolombea fraxini; Lindinger, 1909: 223. Misspelling of genus name.

Pseudichermes fraxini; Gertsson, 2011: 41. Misspelling of genus name.

COMMON NAMES: ash bark scale [Gertss1997]; ash coccus [Kohler1998]; ash scale [Kohler1998]; felted ash coccus [Collin1908]; felted ash scale [Kohler1998].



FOES: Arachnida: Platybunus pinetorum [KosztaKo1988F]. COLEOPTERA Coccinellidae: Chilocorus bipustulatus [KosztaKo1988F], Exochomus quadripustulatus [KosztaKo1988F]. HETEROPTERA Anthocoridae: Anthocoris minki [KosztaKo1988F].

HOSTS: Oleaceae: Fraxinus angustifolia [KozarDr1991], Fraxinus caroliniana [Hoy1963], Fraxinus excelsior [Bodenh1944b], Fraxinus mandtschurica [Hoy1963], Fraxinus ornus [Kozar1999a], Fraxinus sp. [Hoy1962], Ligustrum sp. [Kohler1998], Syringa sp. [Hoy1963]. Rosaceae: Sorbus aucuparia [Hoy1963]. Salicaceae: Populus tremula [Willia1985h].

DISTRIBUTION: Palaearctic: Austria [Kohler1998]; Belgium [Willia1985h]; Bulgaria [Kozar1985]; China [TangHa1995]; Croatia [MastenSi2008]; Czech Republic [Kohler1998]; Denmark [Kohler1998]; Finland [KozarzRe1975, Gertss2001]; France [Hoy1963, Foldi2001, Germai2011]; Germany [Hoy1963]; Greece [Kohler1998]; Hungary [Willia1985h, KozarKoFe2013]; Iran [Hoy1963, KozarFoZa1996]; Italy [Marott1993]; Lithuania [MalumpOsPy2009]; Luxembourg [Kohler1998]; Netherlands [Hoy1963, Jansen2001]; Norway [Kohler1998]; Poland [Szulcz1926, ZakOgaKo1964, SimonKa2011]; Portugal [KozarFr1995]; Romania [Kozar1985]; Russia [Hoy1963] (Caucasus [Danzig1986a], Primor'ye Kray [Danzig1986a], Stavrapol Oblast [Danzig1985]); Slovenia [KozarKaKo2013]; Spain [Hoy1963]; Sweden [KozarzRe1975, Gertss2001]; Switzerland [KozarGuBa1994]; Turkey [Hoy1963]; Ukraine (Krym (=Crimea) Oblast [Danzig1986a]); United Kingdom (England [Willia1985h, MalumpBa2012], Wales [Willia1985h]); Yugoslavia [Kozar1983a].

BIOLOGY: Females had completed egg-laying by the 21st of May, and on the 10th of June the first instars were swarming. Pupation takes place in September and the males appear early in October, and continue to emerge throughout the month and during the early part of November. Males are very active (Newstead, 1903). According to Danzig (1986a), females are the overwintering stage and eggs are laid from May to June. P. fraxini feeds in bark crevices of trees. The access of air to spiracles is limited since it remains pressed with the ventral side into a tight space when feeding. Thereofre, spiracles are situated in deep cavities of sclerotized walls. In addition, the clypeo-labral shild is very large which is the adaptation to feeding on woody parts of plants, and to living in unfavorable oxic conditions. (Podsiadlo, 2006)

GENERAL REMARKS: Detailed description of most stages, including adults, by Newstead (1903). Detailed descriptions and illustrations of adult and immature by Williams (1985h). Afifi (1968) provides an illustration and description of adult male. Photograph of adult fremal in Malumphy, et al., 2009)

STRUCTURE: Adult female bright red, or orange-red, more or less globular. Females are ovate during fecundation but at the time of parturition they become very tumid and almost globose. Sac of adult female is more or less spherical, white and closely felted. A single sac contains 2 or more individuals. Second stage female pale red or yellowish-red, form short ovate, partially covered with a thin cottony secretion. Male apterous bright orange-red or yellowish-red, last 6 segments of the abdomen greenish-yellow, eyes black. Pupae usually the same color as the male, more or less ovate and narrowed posteriorly. First instars elongate-ovate, pale crimson-yellow, mottled with bright crimson, the latter color being also the color of the eyes. Ova pink when first laid, but become mottled with reddish-crimson (Newstead, 1903).

SYSTEMATICS: Slide-mounted adult female with: enlarged setae absent with exception of dorsal setae on posterior 2 or 3 abdominal segments which are slightly enlarged; protruding anal lobes absent; dorsal quinquelocular pores present; legs with translucent pores on hind coxa; anal ring primarily sclerotized posteriorly, forming "u"; microtubular ducts short, with 1 sclerotized area (Williams, 1985h).

KEYS: Kozár et al. 2013: 609 (female) [Key to species of Pseudochermes]; Danzig 1988: 709 (adult female) [Pseudochermes species of the far eastern USSR]; Danzig 1971d: 823 (female) [as Pseudochermes fraxini; Key to species of family Eriococcidae]; Danzig 1964: 634 (adult female) [Pseudochermes species of the far east USSR].

CITATIONS: Afifi1968 [taxonomy: 8]; AndersWuGr2010 [phylogeny, taxonomy: 996]; Balach1935b [distribution, host: 265]; Balach1937c [distribution, host: 6]; BalasSa1982 [distribution, host, life history: 399]; BarbagBiBo1995 [distribution: 43]; Bodenh1935 [taxonomy: 244, 271]; Bodenh1944b [distribution, host: 93, 98]; Bodenh1953a [distribution, host, life history: 127-128, 159]; BognarVi1979 [distribution, host, illustration: 16, 19]; BoratyWi1964 [taxonomy: 92]; Borchs1936 [distribution: 104]; Borchs1937 [illustration: 58]; Borchs1949 [description, distribution, host, illustration, taxonomy: 365]; Borchs1950b [distribution, host: 126]; Borchs1963a [distribution, host: 201, 202]; Borchs1973 [distribution, host: 202]; CebeciKu2005 [distribution, host: 97-102]; Cocker1899j [taxonomy: 264]; Collin1908 [distribution, host: 23]; Collin1937 [chemical control, distribution, host: 101, 107]; Danzig1964 [taxonomy: 634]; Danzig1971d [taxonomy: 823]; Danzig1977b [distribution, host: 39, 57]; Danzig1980b [distribution, host: 233]; Danzig1985 [taxonomy: 111]; Danzig1985 [distribution, host: 147]; Danzig1986a [description, distribution, host, life history: 271]; Danzig1988 [taxonomy: 709]; Dingle1924 [distribution, taxonomy: 378]; Dziedz1970 [distribution, host: 25]; Elliot1933 [distribution: 142]; Ellis1924 [chemical control, distribution, host: 143-144]; Eysell1979 [host: 122]; Ferris1957c [distribution, taxonomy: 88]; FetykoKoDa2010 [distribution: 296]; Fjeldd1996 [distribution: 14]; Foldi2001 [distribution, economic importance: 305, 307]; FrancoRuMa2011 [distribution: 2,16,25]; Germai2011 [distribution, economic importance: 31-34]; Germai2011a [distribution, economic importance: 8]; Gertss1997 [distribution, host, illustration: 113, 114]; Gertss2000 [distribution, host: 151]; Gertss2001 [distribution: 126]; Gertss2011 [distribution: 41]; Gillan1908 [description, distribution, illustration: 238-239]; GomezM1937 [description, distribution, host, taxonomy: 361]; GomezM1958a [distribution, host: 8]; Goot1912 [distribution, host: 289]; Goux1931 [distribution, host: 331]; Green1915a [distribution, host: 181]; Green1922b [description, distribution, host, taxonomy: 21]; Green1928 [description, host, taxonomy: 9]; GwiazdVaDe2006 [phylogenetics: 16-17]; Hadzib1950 [distribution, host: 257, 262]; Hadzib1983 [distribution, host, taxonomy: 269]; Harris1916 [distribution, host: 173]; Harris1916a [distribution, host: 94]; Harris1944 [distribution, host: 110]; Hendry1962 [taxonomy: 127]; HertinSi1972 [biological control: 134]; HodekHo2009 [biological control: 235]; Hopkin1904 [host, distribution: 51, 52]; Howell1979 [taxonomy: 558]; Hoy1963 [catalogue, distribution, host, taxonomy: 188-190]; Jaap1914 [taxonomy: 137]; Jansen2001 [distribution: 200]; JudeicNi1895 [taxonomy: 1249]; Kalten1860 [description, taxonomy: 259]; Kalten1874 [distribution, taxonomy: 433]; Kaweck1935 [distribution, host: 75]; Kaweck1985 [distribution, taxonomy: 34]; KaydanUlEr2007 [distribution, host: 90-106]; Kiritc1931 [distribution, taxonomy: 312]; Kiritc1940 [taxonomy: 123, 124]; Komosi1987 [distribution, host,: 96]; Komosi1987a [distribution, host: 106]; Koszta1959 [distribution, host: 402]; KosztaKo1978 [taxonomy: 65]; KosztaKo1988F [biological control, description, distribution, host, illustration, taxonomy: 272-274]; Koteja1971a [distribution, host: 322]; Koteja1974a [taxonomy: 249]; Koteja1974b [taxonomy: 78]; Koteja1976 [taxonomy: 274]; Koteja1980 [distribution, host, illustration: 79]; Koteja1983a [distribution, host: 676]; Koteja2000a [distribution: 172]; KotejaLi1976 [structure: 667]; KotejaZa1966 [distribution, host: 310, 320]; KotejaZa1969 [distribution, host, taxonomy: 362]; KotejaZa1983 [distribution, host, taxonomy: 478]; Kozar1980 [distribution, host, taxonomy: 65, 67]; Kozar1983a [host: 144]; Kozar1985 [distribution: 202]; Kozar1991 [taxonomy: 81]; Kozar1999a [distribution, host: 139]; Kozar2009 [distribution, taxonomy: 105]; KozarDr1991 [distribution, host: 363]; KozarFoZa1996 [distribution: 64]; KozarFr1995 [distribution, host: 70]; KozarKaKo2013 [description, distribution, host, illustration, structure, taxonomy, phylogeny: 31,35, 609-612]; KozarKiSa2004 [distribution]; KozarKo1982 [distribution, economic importance, host, taxonomy: 204, 210]; KozarKo2002b [distribution: 375]; KozarKo2008a [taxonomy: 148]; KozarKoFe2013 [distribution, taxonomy: 53]; KozarKoSa2002 [catalogue, distribution: 38]; KozarOrKo1977 [distribution, host: 71]; KozarTzVi1979 [distribution, host, taxonomy: 130]; KozarWa1985 [distribution: 75]; Kozarz1986 [distribution, taxonomy: 307]; KozarzRe1976 [distribution, host: 1, 2]; KozarzVl1981 [distribution, host: 16, 20, 23]; KozarzVl1982 [distribution, host: 189]; Krasuc1922 [distribution, host: 54]; Kunkel1967 [distribution, host: 47]; Kuwana1923b [description, taxonomy: 57]; Lagows1998a [ecology: 65]; Lagows2002 [distribution: 243]; LagowsKo1996 [distribution: 31]; Lindin1909b [distribution, host, taxonomy: 223]; Lindin1910 [taxonomy: 192]; Lindin1911 [taxonomy: 358]; Lindin1912b [host, taxonomy: 159]; Lindin1921 [distribution, host: 433]; Lindin1923 [taxonomy: 142, 146]; Lindin1928 [distribution, taxonomy: 103]; Lindin1931 [distribution, host: 121]; Lindin1933 [taxonomy: 31]; Lindin1934b [taxonomy: 175]; Lindin1935 [taxonomy: 135]; Lindin1938 [distribution, taxonomy: 5]; Lindin1941a [distribution, host: 75]; Lindin1957 [taxonomy: 545]; Lindin1958 [taxonomy: 369, 372]; LongoMaPe1995 [distribution: 122]; MacGil1921 [distribution, host: 141]; MalumpBa2012 [distribution: 27]; MalumpOsPy2009 [description, distribution, host: 120-147]; Marcha1908 [description, distribution, host, illustration, taxonomy: 262-263]; Marott1993 [description, distribution, host, illustration, taxonomy: 160, 166-167]; MartinFo1982 [host: 221]; MastenSi2008 [catalogue, distribution, host: 105-119]; MillerGi2000 [biological control, catalogue, description, distribution, host, life history, taxonomy: 443-447]; MilonaKoKo2008a [distribution: 143-147]; Moghad2013a [distribution, host: 58]; MorrisMo1966 [taxonomy: 164]; NanDeWu2013 [phylogenetics, taxonomy: 171, 173-174]; NastChKl1990 [distribution, taxonomy: 121]; Newste1891 [description, distribution, host, taxonomy: 165-166]; Newste1892 [taxonomy: 147]; Newste1896 [description, distribution, host, illustration, taxonomy: 57]; Newste1898 [taxonomy: 97]; Newste1900a [description, distribution, host, illustration, taxonomy: 250-251]; Newste1903 [description, distribution, host, illustration, taxonomy: 210-213]; Nitsch1895 [description, distribution, host, illustration, taxonomy: 1247-1249]; Ossian1951 [distribution, taxonomy: 4]; Ossian1959 [distribution, host: 195]; OuvrarKo2009 [host, phylogeny, taxonomy: 101-115]; Pelliz1991a [distribution, host, taxonomy: 18]; PellizKo2011 [distribution: 66]; Perrie1926 [description, distribution, taxonomy: 122]; Pierce1917 [economic importance: 26]; Podsia2006a [illustration, physiology: 277-281]; Rasina1955 [distribution, host: 69]; Reh1903 [description, distribution, host, taxonomy: 353-354]; Reh1904 [taxonomy: 34]; RipkaReKo1996 [distribution, taxonomy: 13]; RossemBuBu1965 [distribution, host: 148-150]; Schmut1952 [description, distribution, host, taxonomy: 370, 378, 418]; Schmut1955 [host, taxonomy: 159]; Schmut1974 [host, taxonomy: 49]; Schmut1980 [host: 50, 53]; SimonKa2011 [distribution: 237]; Storka1925 [distribution, taxonomy: 71-72]; Sulc1912 [distribution, taxonomy: 33]; Szklar1998 [host, illustration, physiology: 168, 170, 171]; Szulcz1926 [distribution, host, taxonomy: 138]; TangHa1995 [description, distribution, host, taxonomy: 430]; Terezn1966 [distribution: 27]; Terezn1968b [distribution, host: 48]; Terezn1968c [distribution, host: 49]; Terezn1975 [taxonomy: 29]; Terezn1981 [taxonomy: 59]; Thiem1931 [taxonomy: 121]; Tsalev1968 [taxonomy: 207]; Tschor1939 [taxonomy: 89]; Willia1985h [description, distribution, host, illustration, taxonomy: 386-389]; WilliaBe2009 [catalogue: 23]; Wunn1925 [distribution, host: 123]; Wunn1925b [description, distribution, host, taxonomy: 281, 285-286]; Wunn1926 [distribution, host: 41, 45, 48]; Zahrad1959a [taxonomy: 539]; Zahrad1972 [biological control, distribution, host: 403]; Zahrad1977 [taxonomy: 121]; ZahradRo1995 [distribution, host: 204]; ZakOgaKo1964 [distribution, host: 426]; Zielke1942 [taxonomy: 293].



Pseudochermes williamsi Kozár & Konczné Benedicty

NOMENCLATURE:

Pseudochermes williamsi Kozár & Konczné Benedicty, 2008a: 250-252. Type data: CANARY ISLAND:Tenerife, Orotaro Botanical Garden, on Coffea arabica, 4/13/1976, by D.J. Williams. Holotype female (examined), by original designation. Type depositories: London: The Natural History Museum, England, UK, and Budapest: Hungarian Natural History Museum, Zoological Department, Hungary. Described: female. Illust. Notes: Holotype and 3 paratypes deposited in BMNH and one paratype deposited in HNHM.



HOST: Rubiaceae: Coffea arabica [KozarKo2008a].

DISTRIBUTION: Palaearctic: Canary Islands [KozarKo2008a].

GENERAL REMARKS: Detailed description and illustration in Kozár & Konczné Benedicty (2008a).

STRUCTURE: Body of slide-mounted female, elongate-oval. Antennae 6 segmented. Frontal tubercles present. Eyes present on ventral margin. Labium 3 segmented, basal segment not well developed, with a pair of setae. Legs well developedHindcoxae with large translucent pores on dorsal surface. Anal ring on dorsum, well developed with single row of pores and 6 strong setae. Anal lobes not well developed, not sclerotized with 2 spine-like setae. (Kozár & Konczné Benedicty, 2008a)

SYSTEMATICS: P. williamsi is similar to P. fraxini by having sclerotized reduced anal ring with a row of pores and strong setae; by the presence of quinquelocular pores on venter and dorsum, by the small body size, antennae and legs; by one pair of setae on the basal segment of the labium; and by the presence of frontal tubercle. It differs by sthe smaller number of large tubular ducts and quinquelocular pores on the dorsum; by longer ductules than tube ending with a flower-like terminal gland; by unique structure of smaller tubular ducts having 6-locular ends; and by the number, shape and size of the translucent pores on the posterior coxae. (Kozár & Konczné Benedicty, 2008a) Spines of the first instar nymph of P. williamsi are conical, sharply pointed, not truncate as in P. fraxini. (Kozár, et al., 2013)

KEYS: Kozár et al. 2013: 609 (female) [Key to species of Pseudochermes].

CITATIONS: KozarKaKo2013 [distribution, host, illustration, structure, taxonomy: 613-615]; KozarKo2008a [description, distribution, structure, taxonomy: 250-252]; OuvrarKo2009 [host, phylogeny, structure, taxonomy: 105].