Family Matsucoccidae


Eomatsucoccus Koteja

NOMENCLATURE:

Eomatsucoccus Koteja, 1988: 143. Type species: Eomatsucoccus sukachevae Koteja, by monotypy and original designation.

Eumatsucoccus; Foldi, 2001a: 201. Misspelling of genus name.

GENERAL REMARKS: Definition and characters by Koteja (1988).

SYSTEMATICS: All three species of this genus are fossil scale insects.

KEYS: Koteja 2000: 164 (male) [fossil related genera]; Koteja 1999: 864 (male) [Species of genus]; Koteja 1998: 143 (male) [Species of Eomatsucoccus.].

CITATIONS: BenDov2005a [taxonomy, catalogue: 112]; Foldi2001a [taxonomy: 201]; Foldi2005 [taxonomy: 148]; Koteja1988 [taxonomy, description: 143-147]; Koteja1999 [taxonomy: 863-866]; Koteja2000 [taxonomy: 163-167]; Koteja2000c [taxonomy: 204].



Eomatsucoccus andrewi Koteja

NOMENCLATURE:

Eomatsucoccus andrewi Koteja, 1999: 864. Type data: ENGLAND: West Sussex, Keymer Tileworks, wing impression of adult male, in Lower Weald Clay (Upper Hauterivian), Lower Cretaceous. Holotype male. Type depository: BMBM; type no. Coc0424. Described: male. Illust.

Eumatsucoccus andrewi; Foldi, 2001a: 201. Misspelling of genus name.

DISTRIBUTION: Palaearctic: United Kingdom (England [Koteja1999]).

BIOLOGY: A fossil insect found in England, West Sussex, Keymer Tileworks, wing impression of adult male, in Lower Weald Clay (Upper Hauterivian), Lower Cretaceous (Koteja, 1999)..

KEYS: Koteja 1999: 864 (male) [Species of genus].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 112]; Foldi2001a [taxonomy: 201]; Koteja1999 [taxonomy, description, illustration, distribution: 863-866]; Koteja2000c [taxonomy: 205].



Eomatsucoccus casei Koteja

NOMENCLATURE:

Eomatsucoccus casei Koteja, 2000: 165. Type data: USA: New Jersey, White Oaks Pits (Sayreville, Middlesex County), alate male inclusion in New Jersey amber; leg. K. Luzzi. Holotype male. Type depository: New York: American Museum of Natural History, Department of Entomology Collection, New York, USA; type no. Coc0465. Described: male. Illust.

Eumatsucoccus casei; Foldi, 2001a: 201. Misspelling of genus name.



HOST: Pinaceae [Koteja2000].

DISTRIBUTION: Nearctic: United States of America (New Jersey [Koteja2000]).

BIOLOGY: A fossil insect (Koteja, 2000).

KEYS: Koteja 2000: 164 (male) [Genus].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 112]; Foldi2001a [taxonomy: 201]; HodgsoHa2013 [phylogeny, taxonomy: 799]; Koteja2000 [taxonomy, description, illustration, distribution: 164-167, 206-209]; Koteja2000c [taxonomy: 205].



Eomatsucoccus popovi Koteja

NOMENCLATURE:

Eomatsucoccus popovi Koteja, 1988: 147. Type data: RUSSIA: Buryat, ASSR, Eravnenskii District, included in deposits (Zaza Formation), Neocomian (Lower Cretaceous). Holotype. Type depository: Moscow: Paleontological Institute, Russia; type no. Cocc168. Described: male. Illust.

Eumatsucoccus popovi; Foldi, 2001a: 201. Misspelling of genus name.

DISTRIBUTION: Palaearctic: Russia (Buryatia (=Buryat) Aut. Oblast [Koteja1988]).

BIOLOGY: A fossil species, found in deposits (Zaza Formation) that were formed in the early Neocomian (Lower Cretaceous), about 125-115 million years ago (Koteja, 1988).

KEYS: Koteja 2000: 164 (male) [Genus]; Koteja 1988: 143 (male) [Species of genus].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 113]; Foldi2001a [taxonomy: 201]; Koteja1988 [taxonomy, description, illustration, distribution: 147-153]; Koteja2000 [taxonomy: 164]; Koteja2000c [taxonomy: 205].



Eomatsucoccus sukachevae Koteja

NOMENCLATURE:

Eomatsucoccus sukachevae Koteja, 1988: 143. Type data: RUSSIA: Southern Siberia, Baia, Eravnenskii District, included in deposits (Zaza Formation) formed in the early Neocomian, Lower Cretaceous. Holotype male. Type depository: Moscow: Paleontological Institute, Russia; type no. Cocc169. Described: male. Illust.

DISTRIBUTION: Palaearctic: Russia (Aga Buryat Aut. Okrug [Koteja1988]).

BIOLOGY: A fossil species, found in deposits (Zaza Formation) that were formed in the early Neocomian (Lower Cretaceous), about 125-115 million years ago (Koteja, 1988).

KEYS: Koteja 2000: 164 (male) [Genus]; Koteja 1988: 143 (male) [Species of the genus].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 113]; Foldi2001a [taxonomy: 201]; Foldi2005 [taxonomy: 148]; Koteja1988 [taxonomy, description, illustration, distribution: 143-147]; Koteja2000 [taxonomy: 163-167]; Koteja2000c [taxonomy: 204].



Matsucoccus Cockerell

NOMENCLATURE:

Acreagris Koch & Berendt, 1854: 17. Type species: Acreagris crenata Koch & Berendt, by monotypy. Synonymy by Ben-Dov & Koteja, 2002: 380. Notes: Ben-Dov & Koteja (2002) designated this name as a nomen oblitum.

Matsucoccus Cockerell, 1909: 56. Type species: Xylococcus matsumurae Kuwana, by monotypy and original designation. Notes: Ben-Dov & Koteja (2002) designated this name as a nomen protectum.

Americoccus MacGillivray, 1921: 78. Type species: Matsucoccus fasciculensis Herbert, by monotypy and original designation. Synonymy by Morrison, 1928: 219.

Agreagris; Buchner, 1966: 280. Misspelling of genus name.

Sonsaucoccus Young, 1980: 186. Type species: Matsucoccus sinensis Chen, by monotypy and original designation. Synonymy by Tang & Hao, 1995: 95.

BIOLOGY: Kutscher & Koteja (2000) reported, described and illustrated a female of Matsucoccus species held between mandibles of an ant, Azteca sp. that both were included in Dominican amber.

GENERAL REMARKS: Definition and characters by Morrison (1928), Borchsenius (1950b), Danzig (1980b), Kosztarab & Kozar (1988F), Kosztarab (1996) and by Foldi (2004).

SYSTEMATICS: Ben-Dov & Koteja (2002) applied Article 29.3 of the Fourth edition of the International Code of Zoological Nomenclature, in order to stabilize the name Matsucoccus in scale insects nomenclature. They designated the junior synonym Matsucoccus Cockerell, 1909, as nomen protectum and gave it precedence over the senior synonym Acreagris Koch & Berendt, 1854, nomen oblitum. Young (1980, 1986a) clearly distinguished the genus Sonsaucoccus Young, 1980 from Matsucoccus. Tang & Hao (1995) regarded the former as a synonym of Matsucoccus. Main characteristics (adult female) of this genus are:

KEYS: Koteja 2008: 24 (male) [Genera of Baltic amber archeococcoids]; Foldi 2004: 164 (female) [Matsucoccus species in the Mediterranean]; Koteja 2000: 164 (male) [fossil related genera]; Tang, Wu & Li 1997: 21-22 (female) [Palaearctic region]; Kosztarab 1996: 44 (female) [Northeastern North America]; Tang & Hao 1995: 101-102 (female) [Species of Palearctic]; Gill 1993: 35, 42 (female) [USA, California]; Ray & Williams 1991: 190 (female) [North America]; Kosztarab & Kozar 1988F: 47 (female) [central Europe]; Ray & Williams 1984: 768-769 (female) [North America]; Yang 1982: 18-19 (female) [China]; Danzig 1980b: 86 (female) [Genera of Far East Russia]; Kosztarab & Kozar 1978: 16 (female) [Hungary]; Boratynski 1952a: 321-324 (female) [World]; Borchsenius 1950b: 32 (female) [Genera of USSR].

CITATIONS: BenDov2005a [taxonomy, catalogue: 113-114]; BenDovKo2002 [taxonomy: 379-380,434]; BodenhNe1955 [taxonomy, description: 7-11]; Boraty1952a [taxonomy, description: 285-326]; Borchs1950b [taxonomy: 32, 33]; Buchne1966 [taxonomy: 280]; Cocker1909 [taxonomy, description: 56]; Danzig1972a [taxonomy: 261]; Danzig1980b [taxonomy, description: 86, 94-95]; Ferris1941 [taxonomy, description: 6-10]; Ferris1950 [taxonomy: 11]; Foldi1997a [taxonomy: 192]; Foldi1998 [catalogue: 428]; Foldi2001a [taxonomy: 207]; Foldi2005 [taxonomy, description: 145-168]; Gill1993 [taxonomy, description: 35, 40-42]; Goidan1972a [taxonomy, host, life history: 290-294]; GullanCo2007 [taxonomy: 413-425]; GullanSj2001 [taxonomy: 257-278]; HodgsoFo2005 [taxonomy, phylogeny: 35-48]; HodgsoFo2006 [taxonomy: 50]; Kawai1980 [taxonomy: 90]; KochBe1854 [taxonomy, description: 17]; Koszta1996 [taxonomy, description: 43-44]; KosztaKo1978 [taxonomy, description: 16-17]; KosztaKo1988F [taxonomy, description: 47, 48]; Koteja1984d [taxonomy, description: 437-496]; Koteja1999 [taxonomy: 863-866]; Koteja2000 [taxonomy: 163-164]; Koteja2000c [taxonomy: 205]; KutschKo2000 [life history, host, distribution: 179-185]; Lindin1937 [taxonomy: 178, 189]; MacGil1921 [taxonomy, description: 78]; MendelCaPo1991 [host, distribution, biological control: 502-507]; Miller1971b [taxonomy: 66]; MillerPa1987 [taxonomy: 49-50, 56]; MkrtchSa2001 [structure, taxonomy: 167-168]; Morris1928 [taxonomy, description: 6-21, 27-53, 219]; MorrisMo1966 [taxonomy: 3,7,117]; Ray1982 [taxonomy: 1-295]; RayWi1984 [taxonomy, description: 765, 768-769]; RayWi1991 [taxonomy, description: 186, 190, 192]; Rieux1975 [taxonomy, host, distribution: 157-168]; TangHa1995 [taxonomy, description: 95-102]; TangWuLi1997 [taxonomy: 17-23]; Tao1999 [taxonomy: 4]; Wang1986 [taxonomy, ecology: 225-228]; Xie1998 [taxonomy, description: 16]; Yang1982 [taxonomy: 18-21]; Young1980 [taxonomy, description, distribution: 183-189]; Young1987a [taxonomy, description: 45-48]; Zahrad1959a [taxonomy: 354].



Matsucoccus acalyptus Herbert

NOMENCLATURE:

Matsucoccus acalyptus Herbert, 1921: 20. Type data: USA: Idaho, southern Idaho, on single-leaf pinon, Pinus monophylla, collected R.W. Doane. Holotype female and first instar. Type depository: Davis: The Bohart Museum of Entomology, University of California, California, USA. Described: female. Illust.

COMMON NAME: pinyon needle scale [Gill1993].



FOE: HEMIPTERA Anthocoridae: Elatophilus pullus Kelton & Anderson [MendelCaPo1991].

HOSTS: Pinaceae [Gill1993], Pinus monophylla [Herber1921FBB].

DISTRIBUTION: Nearctic: United States of America (Arizona [Gill1993], California [Gill1993], Colorado [Gill1993], Idaho [Herber1921FBB, Gill1993], New Mexico [Gill1993], Utah [Gill1993]).

BIOLOGY: Develops one annual generation in USA, California (McCambridge & Pierce, 1964), Furniss & Carolin (1977). In the southwestern United States, the pinyon needle scale, Matsucoccus acalyptus Herbert, can cause defoliation and mortality of pinyon pine (Pinus edulis Engelmann) after repeated feeding. (Mech, et al., 2013)

GENERAL REMARKS: Description and illustration of adult female given by Herbert (1921) and by Gill (1993).

STRUCTURE: Colour photograph of adult female given by Gill (1993).

KEYS: Gill 1993: 42 (female) [USA, California]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World]; Morrison 1928: 53 (female, larva) [Species of Matsucoccus.].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue, economic importance: 115]; BodenhNe1955 [taxonomy: 10]; BoothGu2006 [taxonomy, molecular biology: 749-760]; Boraty1952 [taxonomy: 322]; Buchne1966 [life history, structure: 282-283]; ClasseHaWh2005 [host, distribution: 2049-2057]; Foldi2001a [taxonomy: 207]; Foldi2005 [taxonomy: 165]; Gill1993 [taxonomy, description, illustration, host, distribution, economic importance: 42-43]; GullanCo2007 [taxonomy: 413-425]; Herber1921FBB [taxonomy, description, illustration, host, distribution: 20-22]; McCamb1974 [host, distribution, life history, economic importance, control: 1-4]; McCambPi1964 [host, distribution, life history, economic importance: 197-200]; MechAsCr2013 [distribution, economic importance, host: 2391]; MendelCaPo1991 [host, distribution, biological control: 502-507]; Morris1928 [taxonomy, distribution: 52-53,77,222]; Rieux1975 [taxonomy, host, distribution: 157-168]; SchmutKlLu1957 [host, distribution, economic importance: 419]; SchustCoWh2005 [host, distribution, life history, ecology: 62-72]; UnruhLu1987 [host, distribution, life history, ecology: 439-449]; Young1980 [taxonomy: 183].



Matsucoccus alabamae Morrison

NOMENCLATURE:

Matsucoccus alabamae Morrison, 1939: 2. Type data: USA: Alabama, Calhoun, on Pinus sp., March 16 & 31, April 21, 1902; collected A.M. Troyer. Syntypes, female and first instar. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female and first instar.

Matsucoccus alabama; Mendel et al., 1991: 503. Misspelling of species name.



FOE: HEMIPTERA Anthocoridae: Elatophilus pinophilus Blatchley [MendelCaPo1991].

HOST: Pinaceae: Pinus [Morris1939].

DISTRIBUTION: Nearctic: United States of America (Alabama [Morris1939]).

KEYS: Ray & Williams 1991: 190 (female) [North America]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 115-116]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 324]; Foldi2001a [taxonomy: 207]; Foldi2005 [taxonomy: 165]; MechAsCr2013 [distribution, host: 2391-2392]; MendelCaPo1991 [host, distribution, biological control: 502-507]; Morris1939 [taxonomy, description, host, distribution: 2-3]; Rieux1975 [taxonomy, host, distribution: 157-168]; Tang1978 [taxonomy: 165]; TangWuLi1997 [taxonomy: 17-23]; Young1980 [taxonomy: 183].



Matsucoccus apachecae Ray & Williams

NOMENCLATURE:

Matsucoccus apachecae Ray & Williams, 1984: 765. Type data: USA: Arizona, Chiricahua Mts., Cave Creek, on Pinus engelmanni; collected G.F. Ferris, 11 July 1940. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female. Illust.



HOSTS: Pinaceae: Pinus engelmanni [RayWi1984], Pinus ponderosa [RayWi1984].

DISTRIBUTION: Nearctic: United States of America (Arizona [RayWi1984]).

GENERAL REMARKS: Description and illustration of adult female by Ray & Williams (1984).

KEYS: Ray & Williams 1984: 768-769 (female) [North America].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 116]; Foldi2001a [taxonomy: 207]; Foldi2005 [taxonomy: 165]; RayWi1984 [taxonomy, description, illustration, host, distribution: 765-766, 769].



Matsucoccus apterus Koteja

NOMENCLATURE:

Matsucoccus apterus Koteja, 1984d: 486. Type data: BALTIC AMBER: Apterous adult male, Baltic amber inclusion. Holotype male. Type depository: Copenhagen: Zoological Museum, University of Copenhagen, Department of Entomology, Denmark; type no. Cocc57. Described: male. Illust.

GENERAL REMARKS: Description and illustration of adult male and adult female by Koteja (1984d).

SYSTEMATICS: A fossil species, described from inclusions of male and female in Baltic amber (Koteja, 1984d).

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 116]; Foldi2001a [taxonomy: 207]; Koteja1984d [taxonomy, description, illustration, distribution: 486-488]; Koteja1999 [taxonomy: 864]; Koteja2000c [taxonomy: 205].



Matsucoccus banksianae Ray & Williams

NOMENCLATURE:

Matsucoccus banksianae Ray & Williams, 1991: 186. Type data: USA: Minnesota, Ely, on Pinus banksiana, June 1962, collected J.C. Bean. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female. Illust.



FOE: HEMIPTERA Anthocoridae: Elatophilus minutus Kelton [MendelCaPo1991].

HOST: Pinaceae: Pinus banksiana [RayWi1991].

DISTRIBUTION: Nearctic: United States of America (Minnesota [RayWi1991]).

GENERAL REMARKS: Description and illustration of adult female by Ray & Williams (1991).

KEYS: Ray & Williams 1991: 190 (female) [North America].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 116-117]; Foldi2001a [taxonomy: 207]; Foldi2005 [taxonomy: 165]; MechAsCr2013 [distribution, host: 2391-2392]; MendelCaPo1991 [host, distribution, biological control: 502-507]; RayWi1991 [taxonomy, description, illustration, host, distribution: 186-188].



Matsucoccus bisetosus Morrison

NOMENCLATURE:

Matsucoccus bisetosus Morrison, 1939: 3. Type data: USA: California, Lassen National Forest, near Willow Spring, on Pinus ponderosa, August 8, 1936. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female.

COMMON NAME: ponderosa pine twig scale [Gill1993].



FOE: HEMIPTERA Anthocoridae: Elatophilus pullus Kelton & Anderson [MendelCaPo1991].

HOSTS: Pinaceae: Pinus contorta [McKenz1942a], Pinus jeffreyi [Morris1939, McKenz1942a], Pinus ponderosa [Morris1939, McKenz1942a], Pinus sabiniana [McKenz1942a], Pinus scopulorum [McKenz1942a].

DISTRIBUTION: Nearctic: United States of America (Arizona [McKenz1942a], California [McKenz1942a], Colorado [Morris1939, McKenz1942a]).

GENERAL REMARKS: Description and illustration of adult female by Morrison (1939) and by Gill (1993). Description and illustration of adult male by Beardsley (1968).

ECONOMIC IMPORTANCE AND CONTROL: Has been found in USA, California to be associated with branch decadence in both young and mature pines of the species Pinus ponderosa and P. jeffreyi (McKenzie, 1942). McKenzie (1942a) suggested that the principal natural control factor is the pressure exerted by the growing tree forcing the bark plates against the insects or by the growing scales forcing themselves out of shape between the narrow bark plates, or both.

KEYS: Hodgson & Foldi 2006: 57 (male); Gill 1993: 42 (female) [USA, California]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: Beards1968 [taxonomy, description, illustration, host, distribution: 1449-1459]; BenDov2005a [taxonomy, host, distribution, catalogue: 117]; BodenhNe1955 [taxonomy: 10]; BoothGu2006 [taxonomy, molecular biology: 749-760]; Boraty1952 [taxonomy: 323]; Foldi2001a [taxonomy: 207]; Foldi2005 [taxonomy: 165]; HodgsoFo2006 [taxonomy: 57]; HodgsoHa2013 [phylogeny, taxonomy: 796]; McKenz1941a [host, distribution, economic importance: 783-785]; McKenz1942a [life history, host, distribution, biological control: 19-24]; MendelCaPo1991 [host, distribution, biological control: 502-507]; Morris1939 [taxonomy, description, host, distribution: 3-5]; Rieux1975 [taxonomy, host, distribution: 157-168]; SchmutKlLu1957 [host, distribution, economic importance: 419]; Tang1978 [taxonomy: 164]; Young1980 [taxonomy: 183].



Matsucoccus boratynskii Bodenheimer & Neumark

NOMENCLATURE:

Matsucoccus pini; Borchsenius, 1950b: 33. Misidentification; discovered by Boratynski, 1952a: 286.

Matsucoccus Russian specimens Boratynski, 1952: 507. Nomen nudum; discovered by Bodenheimer & Neumark, 1955: 10.

Matsucoccus new species Boratynski, 1952a: 286. Nomen nudum; discovered by Bodenheimer & Neumark, 1955: 10.

Matsucoccus boratynskii Bodenheimer & Neumark, 1955: 10. Type data: RUSSIA: St. Petersburg Oblast, Park of Forestry Academy and Park Pavlov, on Pinus sylvestris. Syntypes, female. Type depository: St. Petersburg: Zoological Museum, Academy of Science, Russia. Described: female.

Matsucoccus insignis Borchsenius, 1955c: 222. Type data: RUSSIA: St. Petersburg Oblast, Park of Forestry Academy and Park Pavlov, on Pinus sylvestris. Syntypes, female. Type depository: St. Petersburg: Zoological Museum, Academy of Science, Russia. Described: female. Illust. Synonymy by Miller & Park, 1987: 56.



HOST: Pinaceae: Pinus sylvestris [Borchs1955].

DISTRIBUTION: Palaearctic: Russia (St. Petersburg (=Leningrad) Oblast [Borchs1955c]).

GENERAL REMARKS: Description and illustration of adult female by Borchsenius (1955c).

SYSTEMATICS: The first indications to this species were by Boratynski (1952:286, as "... Matsucoccus ... probably a new one ...") and by Boratynski (1952:507, as "... Matsucoccus... Russian specimens.."). Boratynski (1952: 286) indicated that the record of Matsucoccus pini (Green), from Leningrad oblast USSR, by Borchsenius (1950b: 33) represented a species different from M. pini. Bodenheimer & Neumark (1955:10) clearly referred to Borchsenius' (1950b: 33) description and illustrations of M. pini from Russia, St. Petersburg, and by naming the latter Matsucoccus boratynskii validated the latter. Borchsenius' (1955c) description of Matsucoccus insignis was based on material that Borchsenius (1950b) misidentified as M. pini. The book by Bodenheimer & Neumark (1955) was published before May, 1955, whereas Borchsenius (1955c) was published after 2 November, 1955 (K. Boratynski, personal communication to Yair Ben-Dov; Yair Ben-Dov, unpublished data; Miller & Park, 1987). Therefore, by the law of priority, M. boratynskii is the valid name, and M. insignis is its junior synonym.

KEYS: Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 118]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 507-508]; Boraty1952a [taxonomy: 286]; Borchs1950b [taxonomy, description, host, distribution: 33]; Borchs1955c [taxonomy, description, illustration, host, distribution: 222-224]; Foldi1998 [catalogue: 428]; Foldi2001a [taxonomy: 207]; Foldi2005 [taxonomy: 165]; HuHu1981 [taxonomy: 178-180]; MillerPa1987 [taxonomy: 56]; Rieux1975 [taxonomy, host, distribution: 157-168]; Rieux1976 [taxonomy: 242]; Siewni1976 [taxonomy: 339]; Tang1978 [taxonomy: 168]; TangHa1995 [taxonomy, description: 102-103]; Young1979 [taxonomy: 109]; Young1980 [taxonomy: 183].



Matsucoccus californicus Morrison

NOMENCLATURE:

Matsucoccus californicus Morrison, 1939: 5. Type data: USA: California, Lassen national Forest, Harvey Valley, Burgess Spring, on Pinus ponderosa; collected K.A. Salman, August 2, 1936. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA; type no. 36-10. Described: female.

COMMON NAME: California matsucoccus [Gill1993].



HOSTS: Pinaceae: Pinus jeffreyi [Gill1993], Pinus ponderosa [Morris1939, Gill1993].

DISTRIBUTION: Nearctic: United States of America (California [Morris1939, Gill1993]).

BIOLOGY: Develops one annual generation in USA, California (Gill, 1993).

GENERAL REMARKS: Description and illustration of adult female by Morrison (1939) and by Gill (1993).

ECONOMIC IMPORTANCE AND CONTROL: McKenzie (1942a) suggested that the principal natural control factor is the pressure exerted by the growing tree forcing the bark plates against the insects or by the growing scales forcing themselves out of shape between the narrow bark plates, or both.

KEYS: Gill 1993: 42 (female) [USA, California]; Ray & Williams 1991: 190 (female) [North America]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 119]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 324]; Foldi2001a [taxonomy: 207]; Foldi2005 [taxonomy: 165]; Gill1993 [taxonomy, description, illustration, host, distribution: 42,44,58]; McKenz1942a [life history: 22]; Morris1939 [taxonomy, description, host, distribution: 5-6]; Rieux1975 [taxonomy, host, distribution: 157-168]; SchmutKlLu1957 [host, distribution, economic importance: 420]; Young1980 [taxonomy: 183].



Matsucoccus crenatus (Koch & Berendt)

NOMENCLATURE:

Acreagris crenata Koch & Berendt, 1854: 123. Type data: Inclusion in Baltic amber. Holotype female. Described: female. Illust. Notes: The type was deposited in Koenigsberg Collection. It disappeared during the Second World War (Ben-Dov & Koteja, 2002).

Matsucoccus crenata; Foldi, 2001a: 207. Change of combination.

Matsucoccus crenatus; Pellizzari & Williams, 2013: 410. Change of combination requiring emendation of specific epithet for agreement in gender.

SYSTEMATICS: Koteja (2000c) summarized the status of this fossil as follows: " According to Menge (in Berendt 1854: 123, footnote; 1856: 13) it is a female member of Monophlebus sensu Germar & Berendt, and according to Ferris (1941:6), Acreagris is a senior synonym of Matsucoccus Cockerell; according to Koteja (1984: 451), a nomen dubium in the Orthezioidea. Ben-Dov & Koteja (2002) provided evidence that the type material of Acreagris crenata Koch & Berendt, 1854 was lost.

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 119]; BenDovKo2002 [taxonomy: 379-380,434]; Ferris1941 [taxonomy, description, illustration, distribution: 6-10]; Foldi2001a [taxonomy: 207]; Kaweck1965a [taxonomy, structure: 205-211]; KochBe1854 [taxonomy, description, illustration, distribution: 123]; Koteja1984d [taxonomy: 451-454]; Koteja1987c [taxonomy: 29]; Koteja1999 [taxonomy: 863]; Koteja2000c [taxonomy: 205].



Matsucoccus dahuriensis Hu & Hu

NOMENCLATURE:

Matsucoccus dahuriensis Hu & Hu, 1981: 178. Type data: CHINA: Heilongjiang, Tahe virgin Forest, on Pinus sylvestris var. Mongolica; collected June 1980. Holotype female. Type depository: Beijing: Institute of Entomology, Academy of Sciences, China. Described: female. Illust.



HOST: Pinaceae: Pinus sylvestris mongolicus [HuHu1981, TangHa1995].

DISTRIBUTION: Palaearctic: China (Heilongjiang (=HeilungKiang) [HuHu1981, TangHa1995]).

GENERAL REMARKS: Description and illustration of adult female by Hu & Hu (1981).

KEYS: Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 120]; Foldi2001a [taxonomy: 207]; Foldi2005 [taxonomy: 165]; HuHu1981 [taxonomy, description, illustration, host, distribution: 178-180]; HuSuZh1984 [life history, ecology: 14-20]; MillerPa1987 [taxonomy: 56]; TangHa1995 [taxonomy, description, host, distribution: 103-104].



Matsucoccus degeneratus Morrison

NOMENCLATURE:

Matsucoccus degeneratus Morrison, 1939: 6. Type data: USA: Arizona, Tonto National Forest, Colcord Mountain, on Pinus ponderosa, December 3, 1935, collected R.W. Davidson. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female.



HOSTS: Pinaceae: Pinus oocarpa [Foldi1995a], Pinus ponderosa [Morris1939].

DISTRIBUTION: Nearctic: Mexico [Morris1939, Foldi1995a]; United States of America (Arizona [Morris1939]).

KEYS: Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 120]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 322]; Foldi1995a [host, distribution: 177]; Foldi2001a [taxonomy: 207]; Foldi2005 [taxonomy: 165]; Morris1939 [taxonomy, description, host, distribution: 6-7]; Rieux1975 [taxonomy, host, distribution: 157-168]; Young1980 [taxonomy: 183].



Matsucoccus eduli Morrison

NOMENCLATURE:

Matsucoccus eduli Morrison, 1939: 7. Type data: USA: Arizona, along the Jerome-Flagstaff road about 10 miles east of Clemenceau, on Pinus edulis, April 27, 1936, collected H. Morrison. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female.

Matsucoccus edulis; Young, 1980: 183. Misspelling of species name.



HOST: Pinaceae: Pinus edulis [Morris1939].

DISTRIBUTION: Nearctic: United States of America (Arizona [Morris1939]).

KEYS: Miller & Park 1987: 54-55 (female) [World]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 120]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 324]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 165]; Morris1939 [taxonomy, description, host, distribution: 7-9]; Rieux1975 [taxonomy, host, distribution: 157-168]; Young1980 [taxonomy: 183].



Matsucoccus electrinus Koteja

NOMENCLATURE:

Matsucoccus electrinus Koteja, 1984d: 481. Type data: BALTIC AMBER: Holotype, alate male, Baltic amber inclusion. Holotype male. Type depository: Copenhagen: Zoological Museum, University of Copenhagen, Department of Entomology, Denmark; type no. Cocc72. Described: male. Illust.

GENERAL REMARKS: Description and illustration of adult male and adult female by Koteja (1984d).

SYSTEMATICS: A fossil species, described from inclusions of male and female in Baltic amber (Koteja, 1984d).

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 121]; Foldi2001a [taxonomy: 208]; Koteja1984d [taxonomy, description, illustration, distribution: 481-486, 493]; Koteja1996a [taxonomy, illustration, description: 75]; Koteja1999 [taxonomy: 864]; Koteja2000c [taxonomy: 205].



Matsucoccus fasciculensis Herbert

NOMENCLATURE:

Matsucoccus fasciculensis Herbert, 1919: 157. Type data: USA: California, Mt. St. Helena, on Pinus sabiniana; paratypes from P. ponderosa. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA; type no. 15406. Described: female. Illust.

Americoccus fasciculensis; MacGillivray, 1921: 78. Change of combination.

Matsucoccus facciculensis; Kozarzhevskaya, 1992: 145. Misspelling of species name.

COMMON NAME: needle fascicle scale [Gill1993].



HOSTS: Pinaceae: Pinus ponderosa [Herber1919, Gill1993], Pinus sabiniana [Herber1919, Gill1993].

DISTRIBUTION: Nearctic: United States of America (California [Herber1919, Gill1993], Idaho [Gill1993], Oregon [Gill1993]).

BIOLOGY: Develops one annual generation in USA, California (Gill, 1993).

GENERAL REMARKS: Description and illustration of adult female By Herbert (1919, 1921) and by Gill (1993).

STRUCTURE: Colour photograph of adult female given by Gill (1993).

KEYS: Gill 1993: 42 (female) [USA, California]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World]; Morrison 1928: 53 (female, larva) [Species of Matsucoccus.].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 121]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 322]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 165]; Gill1993 [taxonomy, description, illustration, host, distribution: 42,44,59]; GullanCo2007 [taxonomy: 413-425]; Herber1919 [taxonomy, description, illustration, host, distribution: 157-160]; Herber1921FBB [taxonomy, description: 20]; Kozarz1992 [taxonomy: 145]; MacGil1921 [taxonomy, description: 78]; Morris1928 [taxonomy: 48,52,53,223]; MorrisMo1966 [taxonomy: 7]; Rieux1975 [taxonomy, host, distribution: 157-168]; Young1980 [taxonomy: 183].



Matsucoccus feytaudi Ducasse

NOMENCLATURE:

Matsucoccus matsumurae; Ducasse, 1938: 1. Misidentification; discovered by Ducasse, 1941: 217.

Matsucoccus feytaudi Ducasse, 1941: 217. Type data: FRANCE: Les Landes, at the vicinity of Dax, Leon and Bordeaux, on Pinus maritima; collected G. Ducasse, 29.ix.1938. Lectotype female and first instar, by subsequent designation Foldi, 2004: 148. Type depository: Paris: Museum National d'Histoire naturelle, France; type no. 5760-1. Described: female and first instar.

Matsucoccus feutaudi; Kozarzhevskaya, 1992: 141. Misspelling of species name.

COMMON NAMES: La cochenille du Pin maritime [Foldi2005]; Maritime pine bast scale [Foldi2005].



FOES: COLEOPTERA Coccinellidae: Oenopia conglobata [CovassBiTo1991], Oenopia lyncea [CovassBiTo1991], Rhyzobius chrysomeloides (Herbst) [CovassBiTo1991, CovassToBi1993], Scymnus interruptus [CovassBiTo1991], Scymnus suturalis [CovassBiTo1991]. HEMIPTERA Anthocoridae: Dufuriellus ater [CovassBiTo1991], Elatophilus crassicornis [CovassBiTo1991], Elatophilus nigricornis (Zetterstedt) [BiliotRi1967, Schves1974, FabreDeRi1982, CovassBiTo1991, MendelCaPo1991, CovassToBi1993], Elatophilus pini (Baerensprung) [CovassPo1986, CovassBiTo1991, MendelCaPo1991, CovassToBi1993]. NEUROPTERA Hemerobiidae: Hemerobius simulans [CovassBiTo1991], Hemerobius stigma [CovassBiTo1991]. Inocelliidae: Inocellia bicolor [CovassBiTo1991].

HOSTS: Pinaceae: Pinus maritima [Ducass1941, Schves1974, Foldi2005], Pinus pinaster [ArzoneVi1981, Trembl1988a, CovassBi1992, Foldi2000, Foldi2005].

DISTRIBUTION: Palaearctic: Algeria [Foldi2005]; Bulgaria [Battis2005]; Corsica [Foldi2003, JactelMe2005]; France [Ducass1938, Ducass1941, Foldi1998, Foldi2000, Foldi2005]; Italy [ArzoneVi1981, CovassPo1986, CovassBiTo1991, CovassBi1992, Foldi1998, Foldi2005]; Morocco [Foldi1998, Foldi2005]; Portugal [Foldi1998, Foldi2005]; Spain [CadahiMo1967, Foldi1998, Foldi2005]; Tunisia [Foldi2005].

GENERAL REMARKS: Description and illustration of adult female by Ducasse (1941). Foldi (2005) described and illustrated all instars, adult female and adult male.

STRUCTURE: Colour photograph of adult female and male given by Covassi & Binazzi (1992).

ECONOMIC IMPORTANCE AND CONTROL: Schvester (1971) considered this species as the primary agent causing decline of Pinus maritimus in France. Early detection and risk assessment are considered a fundamental aspect of a management plan. Monitoring of adult males with pheromone traps has proved to be very effective in locating M. feytaudi infestations before the onset of tree decay and in implementing timely measures for prevention and control. Current silvicultural practices such as selective cutting and removal of infested or sensitive trees can help in preventing the establishment of M. feytaudi invasion, while other control methods include mass-trapping of emerging males and use of kairomones to attract natural enemies. (Roversi, et al., 2013)

KEYS: Hodgson & Foldi 2006: 57 (male); Foldi 2004: 164 (female) [Matsucoccus species in the Mediterranean]; Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic]; Miller & Park 1987: 54-55 (female) [World]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: ArzoneVi1981 [host, distribution, economic importance: 3-10]; Aubert2005 [host, distribution, life history, ecology: 47-62]; BarbagBiBo1995 [distribution: 39]; Battis2005 [host, distribution, economic importance: 15-22]; BenDov2005a [taxonomy, host, distribution, catalogue, economic importance: 122-123]; BiliotRi1967 [biological control, life history: 1103-1108]; BinazzCo1989 [host, distribution, economic importance, life history, biological control: 197-222]; BinazzFrPe2004 [chemical ecology: 901-905]; BinazzFrPe2004a [chemical ecology: 897-900]; BinazzPeFr2002 [host, distribution, control, chemical ecology: 155-171]; Blanck1966 [economic importance, chemical control, biological control: 15-26]; BodenhNe1955 [taxonomy: 10]; BonneaTiSc1975 [life history, ecology, biological control: 181-188]; Boraty1952 [taxonomy: 323]; BrancoFrCa2002 [ecology, biological control, chemical ecology: 397-407]; BrancoFrDu2006 [chemical ecology, biological control, life history, chemical ecology: 179-185]; BrancoFrOf2005 [life history, chemical ecology, biological control: 234-235]; BrancoJaSi2004 [biological control, chemical ecology: 233-239]; BrancoLeFr2006 [chemical ecology, biological control: 1577-1586]; BrancoSoBe2008 [life history, biological control: 273-274]; BurbanPeCa1999 [distribution, chemistry, structure: 1593-1602]; CABI2001 [host, distribution: 1-2]; CadahiMo1967 [host, distribution, life history, economic importance: 95-102]; Carle1968 [life history, ecology: 57-68]; CarleCaBo1970 [life history, ecology, economic importance: 89-104]; CarleDe1986 [economic importance, chemical control: 41-45]; CarleRiSc1970 [host, distribution, biological control: 648-650]; CarleSc1975 [economic importance, life history, host, distribution, biological control: 339-350]; CovassBi1992 [host, distribution, economic importance: 453-466]; CovassBiTo1991 [host, distribution, biological control: 575-598]; CovassPo1986 [host, distribution, biological control: 1-10]; CovassToBi1993 [host, distribution, biological control: 101-108]; CywinKa1991 [chemistry, chemical ecology: 1664-1667]; Delage1978 [host, distribution, life history, economic importance: 19-23]; Ducass1938 [taxonomy, life history, economic importance: 129-133]; Ducass1941 [taxonomy, description, illustration, host, distribution, life history: 217-225]; Dunkel1999 [chemistry, chemical ecology: 251-276]; DunkelMeGr1996 [life history, chemistry, biological control: 489-494]; DunkelMoMe2001 [life history, chemistry, biological control: 391-396]; EinhorMeMa1990 [chemistry, chemical ecology: 6633-6636]; Fabre1980 [host, distribution, economic importance: 40-42]; FabreDeRi1982 [life history, biological control, economic importance: 31-42]; FabreMeFo2000 [economic importance, host, distribution, biological control, life history: 777-792]; Foldi1998 [catalogue: 428]; Foldi1999 [structure, taxonomy: 326]; Foldi2000 [host, distribution: 77]; Foldi2001 [distribution: 303]; Foldi2001a [taxonomy: 208]; Foldi2003 [host, distribution: 148]; Foldi2005 [taxonomy, description, illustration, host, distribution: 148-158]; FrancoRuMa2011 [distribution: 17,25]; Germai2011 [distribution, economic importance: 31-34]; Germai2011a [distribution, economic importance: 8]; Grison1984a [host, distribution, economic importance: 827-834]; Grizon1976 [life history, host, distribution, economic importance, control: 68-78]; GullanCo2007 [taxonomy: 413-425]; HodgsoFo2006 [taxonomy: 57]; HodgsoHa2013 [phylogeny, taxonomy: 796]; JactelMe2005 [host, distribution, economic importance: 37-46]; JactelMeBu1996 [ecology, life history, chemistry, chemical ecology: 145-152]; JactelMeCe1998 [life history, ecology, chemical ecology: 33-45]; JactelMeLe1994 [chemistry, chemical ecology: 2159-2170]; JactelMeVe2006 [host, distribution, ecology: 314-323]; JactelPeMe1996 [chemical ecology, life history, ecology: 1143-1156]; Kozarz1992 [taxonomy: 141]; KreiteAuGe2006 [distribution, economic importance, host: 143]; KurosaTaDu2000 [chemistry: 55-66]; Landma2000 [host, distribution, economic importance: 497-506]; Mahieu1986 [host, distribution, economic importance: 43-44]; MAPF1995 [host, distribution: 1-107]; MAPF1996 [host, distribution: 1-80]; Marche2003 [host, distribution: 1-89]; Masutt1973 [economic importance, host, distribution: 41-42]; Masutt1983 [host, distribution: 357]; MendelAsDu2004 [biological control, chemical ecology: 134-140]; MendelCaPo1991 [host, distribution, biological control: 502-507]; MendelDuBr2003 [life history, chemistry, chemical ecology, biological control: 313-317]; MillerPa1987 [taxonomy: 56]; MoriHa1991 [chemistry, chemical ecology: 5995-5998]; MoriHa1993 [chemistry, chemical ecology: 391-401]; Rieux1975 [taxonomy, host, distribution: 157-168]; Riom1977 [life history, ecology: 709-732]; Riom1979 [host, distribution, life history, ecology: 397-456]; Riom1984 [host, distribution, ecology, control: 857-870]; Riom1994 [host, distribution, economic importance: 437-445]; RiomFa1977 [host, distribution, life history, ecology, economic importance: 181-209]; RiomFa1979 [life history, ecology, economic importance: 89-92]; RiomGe1977 [host, distribution, life history, ecology, phenology: 11-50]; RiomGeBo1971 [host, distribution, life history, ecology, biological control: 153-176]; RiomGoFa1979 [life history, economic importance, ecology, host, distribution: 299-320]; RoversPeTo1997 [host, distribution, biological control: 87-97]; RoversScMa2013 [distribution, ecology, host: 9-16]; Schves1967 [life history, economic importance: 375-377]; Schves1971 [life history, economic importance, host, distribution: 139-151]; Schves1974 [life history, economic importance, biological control: 241-256]; SmithMcSc1997 [economic importance]; SoriaMoVi2000 [host, distribution: 335-348]; Trembl1988a [taxonomy, life history, host, distribution: 204-205]; Trembl1999 [economic importance: 19-28]; vanLen2003b [biological control: 167-179]; VennetRiCh2004 [host, distribution, economic importance: 23-36]; Young1979 [taxonomy: 109]; Young1980 [taxonomy: 183].



Matsucoccus gallicolus Morrison

NOMENCLATURE:

Matsucoccus matsumurae; Herbert, 1921: 15. Misidentification; discovered by Morrison, 1939: 9-13.

Matsucoccus gallicolus Morrison, 1939: 9. Type data: USA: Pennsylvania, Stroudsburg, farm of Mrs. E. Bethel, on Pinus rigida, April, 1936, collected E.C. Pyle. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female.

COMMON NAME: pine twig gall scale [Koszta1996].



FOES: HEMIPTERA Anthocoridae: Elatophilus inimicus Drake & Harris [MendelCaPo1991], Elatophilus pinophilus Blatchley [MendelCaPo1991].

HOSTS: Pinaceae: Pinus echinata [Morris1939, Koszta1996], Pinus elliottii [Koszta1996], Pinus glabra [Morris1939, Koszta1996], Pinus ponderosa [Koszta1996], Pinus pungens [Koszta1996], Pinus resinosa [Koszta1996], Pinus rigida [Morris1939, Koszta1996], Pinus serotina [Koszta1996], Pinus taeda [Morris1939, Koszta1996], Pinus virginiana [Morris1939, Koszta1996].

DISTRIBUTION: Nearctic: United States of America (Connecticut [Morris1939, Koszta1996], District of Columbia [Morris1939, Koszta1996], Florida [Morris1939], Georgia [Morris1939], Kentucky [Koszta1996], Maine [Koszta1996], Maryland [Morris1939, Koszta1996], Massachusetts [Morris1939, Koszta1996], Missouri [Morris1939], New Hampshire [Morris1939, Koszta1996], New Jersey [Morris1939, Koszta1996], New York [Morris1939, Koszta1996], North Carolina [Morris1939], Ohio [Morris1939, Koszta1996], Pennsylvania [Morris1939, Koszta1996], Rhode Island [Morris1939, Koszta1996], Tennessee [Morris1939], Virginia [Morris1939, Koszta1996]).

BIOLOGY: Develops one annual generation; egg overwinter (Kosztarab, 1996).

GENERAL REMARKS: Description and illustration of adult female by Morrison (1939) and by Kosztarab (1996).

STRUCTURE: Occurring during the growing stage in small pustule galls on the twigs of the host pines; adult females usually emerging and migrating to oviposit (Morrison, 1939).

KEYS: Kosztarab 1996: 44 (female) [Northeastern North America]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 123-124]; BodenhNe1955 [taxonomy: 10]; BoothGu2006 [taxonomy, molecular biology: 749-760]; Boraty1952 [taxonomy: 323]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; GullanMiCo2005 [taxonomy, structure: 168-169,210=218]; Heptin1971 [host, distribution, economic importance: 1-5]; Koszta1996 [taxonomy, description, illustration, host, distribution: 44-46]; MechAsCr2013 [distribution, host: 2391-2392]; MendelCaPo1991 [host, distribution, biological control: 502-507]; Morris1939 [taxonomy, description, host, distribution, life history: 9-13]; Parr1939 [taxonomy, host, distribution: 624-630]; Rieux1975 [taxonomy, host, distribution: 157-168]; SchmutKlLu1957 [host, distribution, economic importance: 420]; Tang1978 [taxonomy: 165]; Wheele2004 [biological control: 85-89]; Young1980 [taxonomy: 183].



Matsucoccus josephi Bodenheimer & Harpaz {in}: Bodenheimer & Neumark

NOMENCLATURE:

Matsucoccus iosephi; Bodenheimer & Neumark, 1955: 12. Misspelling of species name.

Matsucoccus josephi Bodenheimer & Harpaz {in}: Bodenheimer & Neumark, 1955: 12. Type data: ISRAEL: Zikhron Ya'aqov, on Pinus halepensis, 6.vii.1949. Lectotype female (examined), by subsequent designation Ben-Dov, 1981: 44. Type depository: Bet Dagan: Department of Entomology, The Volcani Center, Israel. Described: female. Illust.

Matsucoccus josefi; Tremblay, 1988a: 205. Misspelling of species name.



FOES: ACARI Smarididae: Smaris pinus Zhang, Z.Q. [Zhang1995]. COLEOPTERA Coccinellidae: Coccinella decempunctata L. [BodenhNe1955], Exochomus flavipes Thnbg. [BodenhNe1955], Synharmonia conglobata L. [BodenhNe1955]. HEMIPTERA Anthocoridae: Elatophilus hebraicus Pericart [MendelCaPo1991, MendelAsSa1994]. NEUROPTERA Chrysopidae: Chrysopa carnea St. [BodenhNe1955]. Sympherobiidae: Sympherobius pygmaeus Rbr. [BodenhNe1955].

HOSTS: Pinaceae: Pinus brutia brutia [BodenhNe1955, BenDov1981, LiphscMe1989a, Zhang1995, Mendel1998], Pinus brutia eldarica [BenDov1981, LiphscMe1989a], Pinus canariensis [BenDov2012], Pinus eldarica [BenDov2012], Pinus halepensis [BenDov1981, Mendel1998], Pinus halepensis [BenDov2012].

DISTRIBUTION: Palaearctic: Crete [Mendel1998, PellizPoSe2011]; Cyprus [Mendel1992, Mendel1998]; Israel [BodenhNe1955, BenDov1981]; Jordan [MendelAsSa1994, Mendel1998]; Lebanon [MendelAsSa1994, Mendel1998]; Turkey [Mendel1992, Mendel1998, KaydanUlEr2007].

BIOLOGY: The synthetic female sex pheromone of this species attracts outdoors the males as well as the adult males and females of the anthocorid Elatophilus hebraicus Pericart (Mendel et al., 1995). The males of scale insects are short lived. Those of the pine bast scale Matsucoccus josephi Bodenheimer and Harpaz (Coccoidea; Matsucoccidae) emerge in the early morning and are dead at midday (Mendel et al., 2012).

GENERAL REMARKS: Description and illustration of adult female and nymphs By Bodenheimer & Harpaz (1955) and by Ben-Dov (1981). Description and illustration of adult female by Foldi (2004). Description and illustration of adult male by Hodgson & Foldi (2006).

STRUCTURE: Ben-Dov (1981) reported on seasonal differences in numbers and measurements of certain morphological characters of the adult female.

ECONOMIC IMPORTANCE AND CONTROL: A destructive pest to Aleppo pine, Pinus halepensis in Israel (Bodenheimer & Neumark, 1955; Ben-Dov, 1981; Golan et al., 1983).

KEYS: Hodgson & Foldi 2006: 57 (male); Foldi 2004: 164 (female) [Matsucoccus species in the Mediterranean]; Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov1981 [taxonomy, description, illustration, host, distribution: 35-51]; BenDov1981a [taxonomy, description, illustration, host, distribution: 18-23, 45]; BenDov2005a [taxonomy, host, distribution, catalogue, economic importance: 124-126]; BenDov2012 [catalogue, distribution, host: 34, 42]; BenDovHa1986 [taxonomy: 32]; BlumbeIsGo1994 [chemical control, biological control: 434-440]; BodenhNe1955 [taxonomy, description, illustration, host, distribution, life history, economic importance, biological control: 1-122]; BrancoFrDu2006 [chemical ecology, biological control, life history, chemical ecology: 179-185]; BrancoFrOf2005 [life history, chemical ecology, biological control: 234-235]; BrancoLeFr2006 [chemical ecology, biological control: 1577-1586]; Dunkel1999 [chemistry, chemical ecology: 251-276]; DunkelGrGr1995 [chemistry, chemical ecology: 849-858]; DunkelHaAs2000 [chemistry, life history, biological control: 1649-1657]; DunkelMeAs1993 [chemistry,: 2805-2808]; DunkelMeGr1996 [life history, chemistry, biological control: 489-494]; DunkelMoMe2001 [life history, chemistry, biological control: 391-396]; Foldi1998 [catalogue: 428]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy, description, illustration, host, distribution: 158-160]; GolanMaMe1983 [economic importance, host, distribution: 357-360]; GullanCo2007 [taxonomy: 413-425]; HalevyMeAs1995 [host, distribution, biological control: 265]; Halper1988 [chemical control, host, distribution: 957-958]; HodgsoFo2006 [taxonomy, description, illustration, host, distribution: 50-57]; HodgsoGo2006 [taxonomy: 30]; HodgsoHa2013 [phylogeny, taxonomy: 796]; KaydanUlEr2007 [host, distribution: 98]; KurosaTaDu2000 [chemistry: 55-66]; LiphscMe1987 [life history, structure, anatomy, economic importance, host, distribution: 369-376]; LiphscMe1989 [life history, structure, anatomy, economic importance, host, distribution: 2692-2703]; LiphscMe1989a [life history, structure, anatomy, economic importance, host, distribution: 135-142]; MadarSoKi2005 [host, distribution, economic importance, ecology: 28-32]; MadmonScMo2003 [host, life history, ecology: 213-222]; Mendel1984 [host, distribution, life history, economic importance: 259-266]; Mendel1988 [economic importance: 329-336]; Mendel1992 [host, distribution, economic importance: 299-306]; Mendel1998 [host, distribution: 323-330]; Mendel2000 [host, distribution, life history, ecology, economic importance: 223-224]; MendelAdNe1997 [life history, chemistry, biological control: 231-240]; MendelAsDu2004 [biological control, chemical ecology: 134-140]; MendelAsSa1994 [host, distribution, biological control: 9-18]; MendelAsSa1995 [economic importance, life history, ecology, host, distribution: 169-177]; MendelAsSa1997 [economic importance, life history, ecology, host, distribution: 327-333]; MendelBlIs1994 [chemical control, biological control: 199-209]; MendelCaPo1991 [host, distribution, biological control: 502-507]; MendelCaPo1995 [life history, biological control: 856-861]; MendelDuBr2003 [life history, chemistry, chemical ecology, biological control: 313-317]; MendelDuRo1990 [life history, ecology, chemistry, physiology, chemical ecology: 2341-2352]; MendelFaGo1983 [host, ecology, biological control: 33-36, 48]; MendelLi1988 [life history, economic importance, host: 951-959]; MendelNeGa1994 [chemistry, distribution: 165-169]; MendelPrJa2012 [behaviour: 390]; MendelRo1988 [chemical control: 1143-1147]; MendelSaRo1990 [host, distribution, life history, ecology: 532-537]; MendelSc1993 [host, distribution: 383-388]; MendelZeHa1995 [chemistry, life history, biological control, ecology: 331-341]; MendelZeTa1990 [life history, ecology: 603-608]; MillerPa1987 [taxonomy: 49, 56]; MoriAm1994 [chemistry, chemical ecology: 2727-2733]; NestelCoSa1995 [life history, ecology: 506-512]; PellizPoSe2011 [distribution: 293]; Rieux1975 [taxonomy, host, distribution: 157-168]; SchillMe1995 [host, distribution: 159-163]; SchmutKlLu1957 [host, distribution, economic importance: 420]; TangHa1995 [taxonomy, description: 104-106]; Trembl1988a [taxonomy, host, distribution: 205]; Young1979 [taxonomy: 109]; Young1980 [taxonomy: 183]; ZegelmHaMe1993 [life history, chemistry, ecology: 5641-5644]; Zhang1995 [biological control, host, distribution: 69-74].



Matsucoccus koraiensis Young & Hu, in: Young, Hu & Ren

NOMENCLATURE:

Matsucoccus matsumurae; Danzig, 1972: 261. Misidentification; discovered by Danzig, 1980b: 95.

Matsucoccus koraiensis Young & Hu, in: Young, Hu & Ren, 1976: 203. Type data: CHINA: on transported logs of Pinus koraiensis delivered from Heilungkiang Province to Nanking Province. Syntypes, female. Type depository: Beijing: Institute of Entomology, Academy of Sciences, China. Described: female. Illust.

Matsucoccus koraiensis; Kozarzhevskaya, 1992: 144. Notes: Incorrect citation of "Young, Hu & Ren" as authors.



HOST: Pinaceae: Pinus koraiensis [YoungHuRe1976, Danzig1980b, TangHa1995].

DISTRIBUTION: Palaearctic: China (Heilongjiang (=HeilungKiang) [YoungHuRe1976, TangHa1995], Jilin (=Kirin) [TangHa1995]); Russia (Primor'ye Kray [Danzig1980b]).

GENERAL REMARKS: Description and illustration of adult female by Young et al. (1976), Danzig (1980b) and by Tang & Hao (1995).

KEYS: Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 126]; Danzig1980b [taxonomy, description, illustration, host, distribution: 95-96]; Foldi1998 [catalogue: 428]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 165]; HuDaDu1983 [life history, biological control: 30-38]; Jashen1999 [taxonomy, host, distribution: 44]; Kozarz1992 [taxonomy, host, distribution: 144-145]; TangHa1995 [taxonomy, description, host, distribution: 106-108]; Yang1982 [host, distribution: 21]; Young1980 [taxonomy: 183]; YoungHuRe1976 [taxonomy, description, illustration, host, distribution: 203-204].



Matsucoccus larssoni Koteja

NOMENCLATURE:

Matsucoccus larssoni Koteja, 1984d: 470. Type data: BALTIC REGION: in Baltic amber inclusion. Holotype alate male and 4 paratype adult females. Holotype male. Type depository: Copenhagen: Zoological Museum, University of Copenhagen, Department of Entomology, Denmark; type no. Cocc15a. Described: male. Illust.

GENERAL REMARKS: Description and illustration of adult male and adult female by Koteja (1984d).

SYSTEMATICS: A fossil species, described from inclusions of male and female in Baltic amber (Koteja, 1984d).

KEYS: Koteja 1984d: 469 (male) [Fossil males of Matsucoccus spp.].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 126-127]; Foldi2001a [taxonomy: 208]; HodgsoHa2013 [phylogeny, taxonomy: 799]; Koteja1984d [taxonomy, description, illustration, distribution: 469-475, 489]; Koteja1987 [taxonomy, structure: 42]; Koteja1996a [taxonomy, illustration, description: 75]; Koteja1999 [taxonomy: 864]; Koteja2000c [taxonomy: 205].



Matsucoccus leiophyllae Ray & Williams

NOMENCLATURE:

Matsucoccus leiophyllae Ray & Williams, 1984: 766. Type data: USA: Arizona, Chiricahua Mts., Cave Creek, on Pinus leiophylla, July 11, 1940; collected G.F. Ferris. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female. Illust.



HOSTS: Pinaceae: Pinus [RayWi1984], Pinus leiophylla [RayWi1984].

DISTRIBUTION: Nearctic: Mexico (Distrito Federal [RayWi1984]); United States of America (Arizona [RayWi1984]).

GENERAL REMARKS: Description and illustration of adult female by Ray & Williams (1984).

KEYS: Ray & Williams 1984: 768-769 (female) [North America].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 127]; Foldi1995a [host, distribution: 177]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; RayWi1984 [taxonomy, description, illustration, host, distribution: 766-768].



Matsucoccus liaoningensis Tang

NOMENCLATURE:

Matsucoccus liaoningensis Tang, 1978: 164. Type data: CHINA: Liaoning Province, on pine. Syntypes, female, male and first instar. Type depository: Shanxi: Entomological Institute, Shanxi Agricultural University, Taigu, Shanxi, China. Described: female, male and first instar. Illust.

Matsucoccus liaoningensis; Young, 1979: 114. Incorrect synonymy; discovered by Tang & Hao, 1995: 558. Notes: M. liaoningensis Tang, 1978 is a distinct species, not a synonym of M. matsumurae.



HOST: Pinaceae: Pinus tabulaeformis [TangHa1995].

DISTRIBUTION: Palaearctic: China (Liaoning [Tang1978, TangHa1995]).

GENERAL REMARKS: Description and illustration of adult female by Tang (1978) and by Tang & Hao (1995).

KEYS: Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 127]; Foldi1998 [catalogue: 428]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 165]; Tang1978 [taxonomy, description, illustration, host, distribution: 166-170]; TangHa1995 [taxonomy, description, illustration, host, distribution: 108-109,558-559]; TangWuLi1997 [taxonomy: 17-23]; Yang1982 [host, distribution: 21]; Young1979 [taxonomy: 108-114].



Matsucoccus macrocicatrices Richards

NOMENCLATURE:

Matsucoccus macrocicatrices Richards, 1960: 179. Type data: CANADA: Ontario, Lynedoch, on Pinus strobus, June 3, 1958, collected N.W.B. Watson. Holotype female. Type depository: Ottawa: Canadian National Collection of Insects, Centre for Land and Biological Resources Research, Ontario, Canada; type no. 7070. Described: female, male and first instar. Illust.

COMMON NAMES: Canadian pine scale [Koszta1996]; canadian pine scale [Koszta1996].



ASSOCIATE: FUNGI Septobasidiaceae: Septobasidium pinicola Snell [MechAsCr2013].

FOE: HEMIPTERA Anthocoridae: Elatophilus brimleyi Kelton [MendelCaPo1991].

HOST: Pinaceae: Pinus strobus [Richar1960WR, Koszta1996].

DISTRIBUTION: Nearctic: Canada (New Brunswick [Koszta1996], Nova Scotia [Koszta1996], Ontario [Richar1960WR, Koszta1996], Quebec [Martin1964]); United States of America (Georgia [MechAsCr2013], Massachusetts [MechAsCr2013], New Hampshire [Koszta1996], North Carolina [MechAsCr2013], South Carolina [MechAsCr2013], Tennessee, Vermont [MechAsCr2013], Virginia, West Virginia [MechAsCr2013]).

BIOLOGY: Develops one generation every two years; overwinters at cyst stage in fungal mats of Septobasidium pinicola Snell, apparently in a mutualistic relationship. Eggs are deposited in lichens and bark crevices in spring; crawlers migrate to fungal mats, feed and molt to cyst stage, where they remain for two years, then mature during the third spring. Males are rare (Miller, 1985). The lack of overlapping generations and synchronous adult emergences in early 2012 and again in early 2013 suggests that M. macrocicatrices may have a 1-yr life cycle in Georgia. (Mech, et al., 2013)

GENERAL REMARKS: Description and illustration of adult female by Richards (1960) and by Kosztarab (1996).

ECONOMIC IMPORTANCE AND CONTROL: Unlike some of the other pine bast scales, M. macrocicatrices has not historically been associated with tree dieback or mortality, The variability of fungi found with cankers and tissue associated with feeding individuals of M. macrocicatrices reviewed in Mech, et al. (2013) indicates that the insect is most likely not associated with a single pathogen. Because scales are found either deeply embedded in the cankers or present on top of the bark with clear necrotic tissue under their feeding area, they hypothesizee that they may be creating wounds that are then infested by opportunistic fungi such as C. pinea.

KEYS: Kosztarab 1996: 44 (female) [Northeastern North America]; Ray & Williams 1984: 768-769 (female) [North America].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 127-128]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; Koszta1996 [taxonomy, description, illustration, host, distribution: 46-47]; Martin1964 [host, distribution: 42-46]; MechAsCr2013 [behaviour, description, distribution, ecology, host, illustration, molecular data, taxonomy: 2391-2398]; MendelCaPo1991 [host, distribution, biological control: 502-507]; Richar1960WR [taxonomy, description, illustration, host, distribution: 179-181]; Rieux1975 [taxonomy, host, distribution: 157-168]; WatsonUnRe1960 [life history, ecology: 662-667]; Young1980 [taxonomy: 183]; YoungLuZh1986 [taxonomy: 198].



Matsucoccus massonianae Young & Hu, in: Young, Hu & Ren

NOMENCLATURE:

Matsucoccus massonianae Young & Hu, in: Young, Hu & Ren, 1976: 202. Type data: CHINA: Chekiang Province, on Pinus massoniana. Syntypes, female. Type depository: Beijing: Institute of Entomology, Academy of Sciences, China. Described: female. Illust.

Matsucoccus massoniana; Foldi, 1998: 428. Misspelling of species name.

COMMON NAME: Massonian pine bast scale [YoungWu1983].



FOES: COLEOPTERA Coccinellidae: Harmonia axyridis (Pallas) [Wang1982]. HEMIPTERA Anthocoridae: Elatophilus nipponensis Hiura [MendelCaPo1991]. NEUROPTERA Sympherobiidae: Sympherobius weisong Yang [Wang1984].

HOSTS: Pinaceae: Pinus massoniana [YoungHuRe1976, TangHa1995], Pinus thunbergiana [TangHa1995], Pseudolarix kaempferi [TangHa1995].

DISTRIBUTION: Oriental: China (Zhejiang (=Chekiang) [YoungHuRe1976, TangHa1995]).

BIOLOGY: Develops one annual generation in Chekiang, China (Young et al., 1976). Young & Qi (1983) demonstrated that extracts of the female sex pheromone attracted adult males of this species, and also at a lower rate the adult males of M. matsumurae.

GENERAL REMARKS: Description and illustration of adult female given by Young et al. (1976) and by Tang & Hao (1995).

STRUCTURE: Young & Wu (1983) described the ultrastructure of sperm bundle.

ECONOMIC IMPORTANCE AND CONTROL: Regarded a pest of Pinus massoniana in China (Young et al. 1976).

KEYS: Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue, economic importance: 128-129]; Foldi1998 [catalogue: 428]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; HuWa1976 [life history, host, biological control, chemical control: 383-392]; LiSuYu1985 [host, distribution, life history, biological control : 21-23]; MendelCaPo1991 [host, distribution, biological control: 502-507]; MillerPa1987 [taxonomy: 56]; TangHa1995 [taxonomy, description, host, distribution: 109-110,559]; Wang1982 [life history, biological control: 37-39]; Wang1984 [life history, host, distribution, biological control: 95-96]; Yang1982 [host, distribution: 21]; Young1980 [taxonomy: 183]; YoungHuRe1976 [taxonomy, description, illustration, host, distribution: 199-204]; YoungQi1983 [taxonomy, life history, ecology, chemistry: 273-278]; YoungWu1983 [structure: 263-265].



Matsucoccus matsumurae (Kuwana)

NOMENCLATURE:

Xylococcus matsumurae Kuwana, 1905: 91. Type data: JAPAN: Tokyo, at Sugamo, on bark of the trunk of pine-tree; collected May 20, 1903. Syntypes, female. Described: female. Illust. Notes: The type-series was destroyed in an earthquake in 1923 (Kuwana, 1925; Booth & Gullan, 2006).

Xylococcus matsumurae; Kuwana, 1907: 209. Notes: Described again as "n. sp.".

Matsucoccus matsumurae; Cockerell, 1909: 56. Change of combination.

Matsucoccus resinosae Bean & Godwin, 1955: 166. Type data: USA: Connecticut, Easton, on Pinus resinosa, June 2, 1948; collected George H. Plumb. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female, male and first instar. Illust. Synonymy by Booth & Gullan, 2006: 757.

Matsucoccus thunbergianae Miller & Park, 1987: 50. Type data: SOUTH KOREA: Collanam, on Pinus thunbergiana, 1984, collected S.C. Park. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female, male and first instar. Illust. Synonymy by Booth & Gullan, 2006: 757.

Matsucoccus matsunurae; Zhao & Li, 1989: 13. Misspelling of species name.

Matsococcus matsumurae; Gertsson, 2001: 125. Misspelling of genus name.

COMMON NAMES: Japanese pine bast scale [YoungWu1983]; red pine scale [BeanGo1955, Koszta1996].



FOES: ACARINA Anystidae: Anystis baccarum [UlgentSzUy2013]. COLEOPTERA Coccinellidae: Harmonia axyridis (Pallas) [McClur1986, ChoiLeKi1995]. HEMIPTERA Anthocoridae: Dufouriella ater (Dufour) [Gao1987], Elatophilus inimicus Drake & Harris [MendelCaPo1991, NelsonLaGr2002], Elatophilus nipponensis Hiura [ChengMiGe1983, MendelCaPo1991]. HYMENOPTERA Formicidae: Formica lemani [HuoLiZh1983].

HOSTS: Pinaceae: Pinus [Kuwana1905], Pinus densiflora [BeanGo1971], Pinus resinosa [BeanGo1955], Pinus tabulaeformis [BeanGo1971, LiZh1997], Pinus thunbergiana [MillerPa1987], Pinus thunbergii [BeanGo1971].

DISTRIBUTION: Nearctic: United States of America (Connecticut [BeanGo1955, Koszta1996], Massachusetts [MechAsCr2013], New Hampshire [MechAsCr2013], New Jersey [BeanGo1971, Koszta1996], New York [BeanGo1971, Koszta1996], Pennsylvania [MechAsCr2013], Rhode Island [MechAsCr2013]). Palaearctic: China (Beijing (=Peking) [YoungMiMc1984, YoungQi1983]); Japan [Kuwana1917a, Kawai1980]; South Korea [MillerPa1987]; Sweden [Gertss2001].

BIOLOGY: Young & Qi (1983) demonstrated that extracts of the female sex pheromone attracted adult males of this species, and also at a lower rate the adult males of M. massonianae. Young et al. (1984) found that adult males of M. resinosae Bean & Godwin, and adult males of Japanese M. matsumurae are strongly attracted to the sex pheromone of adult females of Chinese populations of M. matsumurae. The results of these bioassays provided evidence that M. resinosae may be a junior synonym of M. matsumurae (Young et al., 1984).Develops two annual generations in USA, Connecticut (Bean & Godwin, 1955, 1971). Young et al. (1984) found that adult males of M. resinosae Bean & Godwin, and adult males of Japanese M. matsumurae are strongly attracted to the sex pheromone of adult females of Chinese populations of M. matsumurae. The results of these bioassays provided evidence that M. resinosae may be a junior synonym of M. matsumurae (Young et al., 1984). Park et al. (1986) found in laboratory bioassays, that males of Matsucoccus resinosae from Pinus resinosa in New York, and males of Matsucoccus sp. from Pinus thunbergiana in Korea, were strongly attracted to crude hexane extract of M. resinosae females. Males of M. resinosae responded strongly to extracts of Matsucoccus sp. females. Males of both species responded similarly to gas chromatographic fractions and subfractions of a pentate extract of M. resinosae females. Sex pheromones of these two species appear to be the same or very similar (Park et al., 1986). This species is regarded a pest of pine trees in eastern USA.Park et al. (1986) found in laboratory bioassays, that males of Matsucoccus resinosae from Pinus resinosa in New York, and males of Matsucoccus sp. from Pinus thunbergiana in Korea, were strongly attracted to crude hexane extract of M. resinosae females. Males of M. resinosae responded strongly to extracts of Matsucoccus sp. females. Males of both species responded similarly to gas chromatographic fractions and subfractions of a pentate extract of M. resinosae females. Sex pheromones of these two species appear to be the same or very similar (Park et al., 1986).

GENERAL REMARKS: Description and illustration of adult female by Kuwana (1905, 1907), Morrison (1928), Bean & Godwin (1955) (as M. resinosae), Young et al. (1976), Miller & Park (1987) (as M. thunbergianae), Tang & Hao (1995), Kosztarab (1996) and by Foldi (2004). The redescription by Herbert (1921), under the name Matsucoccus matsumurae is a misidentification of M. gallicolus; see discussion by Booth & Gullan (2006).

STRUCTURE: Young & Wu (1983) described the ultrastructure of sperm bundle. Young & Yao (1986) studied and described the ultrastructure and histology of the dorsal cicatrices in the adult females from Chinese population of this species. They demonstrated that the gland cells undergo distinct changes after mating, and hypothesized that the dorsal cicatrices function in the secretion of the sex pheromone. Miller & Park (1987) reported on seasonal differences in numbers and measurements of certain morphological characters of the adult female.

SYSTEMATICS: Danzig (1972a: 261-263) recorded under (M. matsumurae material from Russia (Leningrad region and Primorye Territory) and from Kazakhstan. Later, Danzig (1980b) indicated that the material recorded from Russia (Leningrad region and Primorye Territory) belongs to M. matsumurae, while the record from Russia, Primorye Territory belongs to M. koraiensis. Young et al. (1976) recorded and described M. koraiensis from the Liaoning, Shantung, Kiangsu and Chekiang provinces in China. However, Tang & Hao (1995) interpreted that this species does not occur in China. McClure (1983a) suggested that M. resinosae and M. matsumurae are synonyms, but did not formally introduce the synonymy.McClure (1983a) suggested that M. resinosae and M. matsumurae are synonyms, but did not formally introduce the synonymy.

ECONOMIC IMPORTANCE AND CONTROL: Lee et al. (1991) reported on chemical control of this pest in South Korea. M. matsumurae was responsible for foliage discoloration and mortality of plantation-grown P. resinosa in Connecticut. (McClure, 1963)

KEYS: Foldi 2004: 164 (female) [Matsucoccus species in the Mediterranean]; Kosztarab 1996: 44 (female) [Northeastern North America]; Tang & Hao 1995: 101-102 (female) [Palearctic]; Miller & Park 1987: 54-55 (female) [World]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World]; Morrison 1928: 53 (female, larva) [Species of Matsucoccus.].

CITATIONS: Abraha1977 [life history, host, distribution, economic importance: 1-2]; Aldric1996 [chemical ecology: 199-233]; AndersFoKe1976 [life history, host, distribution: 1-5]; Bean1956 [taxonomy, host, distribution, life history, economic importance: 1-4]; BeanGo1955 [taxonomy, description, illustration, host, distribution, life history, economic importance: 164-176]; BeanGo1971 [taxonomy, host, distribution, life history, economic importance: 1-6]; BenDov2005a [taxonomy, host, distribution, catalogue, economic importance: 129-131, 134-138]; BodenhNe1955 [taxonomy: 10]; BoothGu2006 [taxonomy, molecular biology: 749-760]; Boraty1952 [taxonomy: 323]; BrancoFrOf2005 [life history, chemical ecology, biological control: 234-235]; BrancoLeFr2006 [chemical ecology, biological control: 1577-1586]; Buchne1966 [life history, structure: 280-281]; ChengMiGe1983 [economic importance, biological control: 11-30]; ChoiLeKi1995 [host, distribution, biological control: 115-118]; ChoiLeSh1997 [host, distribution, chemical control: 82-86]; ChoiLeSh1998 [host, distribution, life history, ecology: 146-150]; ChungBaPa2000 [host, distribution: 85-89, 306-309]; Cocker1909 [taxonomy: 55-56]; CollinCo1958 [host, distribution, biological control, economic importance: 335-340]; CywinWeKa1991 [chemistry, chemical ecology: 2953-2955]; Danzig1972a [taxonomy, host, distribution: 261-263]; Doane1965 [host, distribution, economic importance: 1-7]; Dunkel1999 [chemistry, , chemical ecology: 251-276]; DunkelMeGr1996 [life history, chemistry, biological control: 489-494]; DunkelMoMe2001 [life history, chemistry, physiology, biological control: 391-396]; Floren1917 [taxonomy: 148]; Foldi1998 [catalogue: 429]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy, description, illustration, host, distribution: 160-162]; Gao1987 [biological control: 271-276]; Gertss2001 [distribution: 123-130]; GrimblPa1976 [life history, ecology: 1-3]; GuoYaFa2005 [host, distribution, life history: 17-19]; Herber1921FBB [taxonomy, description, host, distribution: 15-19]; HeZh2001 [host, distribution, chemical control: 24-26]; HibbarLaPa1991 [chemistry, chemical ecology: 89-102]; HuoLi1990 [host, distribution, economic importance: 21-22]; HuoLiZh1983 [host, distribution, life history, biological control: 243-245]; Jashen1999 [taxonomy, host, distribution: 44]; Kawai1972 [host, distribution: 3]; Kawai1980 [taxonomy, host, distribution: 90]; Kaweck1965a [taxonomy, structure: 205-211]; KishiAs2003 [host, distribution, life history: 131-135]; Koszta1996 [taxonomy, description, illustration, host, distribution: 46-50]; Kozarz1992 [taxonomy, host, distribution: 143-144]; KurosaTaDu2000 [chemistry: 55-66]; Kuwana1905 [taxonomy, description, illustration, host, distribution: 91-95]; Kuwana1907 [taxonomy, description, illustration, host, distribution: 209-212]; Kuwana1917a [taxonomy, distribution: 165]; LanierQiWe1989 [taxonomy, chemistry: 1645-1659]; LeePaPa1991 [host, distribution, chemical control, economic importance: 140-145]; LeeWiPa2013 [distribution, physiology: 349-356]; LiHaXu2005 [host, distribution, ecology, economic importance: 95-100]; LinXu1993 [chemistry, chemical ecology: 5931-5934]; LinXu1996 [chemistry, chemical ecology: 375-380]; LiZh1997 [host, distribution, life history, biological control: 307-320]; LiZhHa1980 [host, distribution, life history, economic importance, biological control: 1-27]; Mao1986 [host, distribution, life history: 201-204]; McClur1976 [host, distribution, life history, ecology: 943-947]; McClur1977b [host, distribution, life history, ecology: 789-795]; McClur1981a [host, distribution, life history, ecology, economic importance: 4-6]; McClur1982 [host, distribution: 150-157]; McClur1983 [host, distribution, life history, ecology: 710-718]; McClur1983a [host, distribution, life history, ecology: 761-765]; McClur1985a [host, distribution, life history, ecology: 535-538]; McClur1986 [host, distribution, life history, ecology, biological control: 976-983]; McClur1990 [host, distribution, life history, ecology: 672-676]; McClur1991 [host, distribution, life history, ecology: 256-270]; McClurDaDe1983 [host, distribution, life history, ecology, biological control, chemical control: 440,475-477]; McInty1958 [host, distribution, economic importance: 369-371]; MechAsCr2013 [distribution, host: 2391-2392]; MendelAsDu2004 [biological control, chemical ecology: 134-140]; MendelCaPo1991 [host, distribution, biological control: 502-507]; MillerPa1987 [taxonomy, description, illustration, host, distribution: 49-62]; MingGeZh1983 [biological control: 19-29]; MoriHa1993a [chemistry, chemical ecology: 993-1001]; Morris1928 [taxonomy, description, illustration, host, distribution: 13,48-53,225]; NelsonLaGr2002 [biological control, host, distribution: 284-285]; Park1988 [life history, chemical ecology: 1-179]; Park1996 [life history, chemistry, chemical ecology, ecology, host, distribution]; ParkAb1991a [chemistry, chemical ecology, physiology: 94-99]; ParkWeAb1986 [taxonomy, life history, chemistry: 609-617]; ParkWiKi1994 [chemistry, physiology, control, chemical ecology: 2185-2196]; ParkWiKi2000 [chemistry, chemical ecology, physiology: 135-140]; ParkWiMo1994 [chemistru, chemical ecology: 250-256]; QiFuXu1984 [life history, ecology, chemistry: 49-52]; QiFuXu1985 [life history, chemistry: 13-17]; QiYaZh1983 [life history, chemistry: 21-28]; Rieux1975 [taxonomy, host, distribution: 157-168]; SchmutKlLu1957 [host, distribution, economic importance: 420]; ShenLoJi2003 [host, distribution, chemical control: 27-28]; ShinLeLe2003 [host, distribution, economic importance: 1-10]; Smetni1991 [chemistry: 92-129]; Stephe1977 [life history, ecology: 4-5]; StepheAy1978 [life history, ecology: 556-563]; Taketa1972 [host, distribution, life history, ecology: 1-9]; Tang1978 [taxonomy, description: 164-170]; TangHa1995 [taxonomy, description: 111,551-558]; TangWuLi1997 [taxonomy: 17-23]; WatanaWaKi1997 [chemistry, chemical ecology: 127-130]; Wei2002 [economic importance: 66-70]; WiPa2001 [life history, behaviour, ecology, chemistry: 139-145]; Xie1998 [taxonomy, description, illustration, host, distribution: 16-19]; XuYaZh1985 [host, distribution, life history: 389-395]; Yang1982 [host, distribution: 21]; Young1979 [taxonomy, life history, host, distribution: 108-114, 183]; Young1980 [taxonomy: 183]; YoungHuRe1976 [taxonomy, description, illustration, host, distribution: 199-204]; YoungHuRe1980 [taxonomy, description: 42-46]; YoungMiMc1984 [life history, taxonomy, chemistry: 1-20]; YoungQi1983 [taxonomy, life history, ecology, chemistry: 273-278]; YoungWu1983 [structure: 263-265]; YoungYa1986 [life history, physiology, structure: 253-260]; YoungYa1987 [structure, life history: 79-82]; Zhao1987 [biological control: 126-127]; Zhao1988 [host, distribution, biological control: 1-109]; ZhaoLi1989 [taxonomy, host, distribution, biological control: 7-13].



Matsucoccus monophyllae McKenzie

NOMENCLATURE:

Matsucoccus monophyllae McKenzie, 1941: 2. Type data: USA: California, Los Padres National Forest, Chuchupate Ranger Station, on Pinus monophylla, November 5, 1939; collected J.E. Patterson & S.T. Carlson. Holotype female. Type depository: Davis: The Bohart Museum of Entomology, University of California, California, USA. Described: female and first instar. Illust.



HOSTS: Pinaceae: Pinus edulis [FurnisCa1977, Gill1993], Pinus monophylla [McKenz1941, Gill1993].

DISTRIBUTION: Nearctic: United States of America (California [McKenz1941]).

BIOLOGY: Develops one annual generation in USA, California (Gill, 1993).

GENERAL REMARKS: Description and illustration of adult female by McKenzie (1941) and by Gill (1993).

STRUCTURE: Colour photograph of adult female, male and cysts given by Gill (1993).

ECONOMIC IMPORTANCE AND CONTROL: Heavily infested trees exhibit branch dieback and "flagging" (McKenzie, 1941).

KEYS: Gill 1993: 42 (female) [USA, California]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue, economic importance: 131]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 322]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; Gill1993 [taxonomy, description, illustration, host, distribution: 42,45,60]; McKenz1941 [taxonomy, description, illustration, host, distribution: 2-5]; Rieux1975 [taxonomy, host, distribution: 157-168]; SchmutKlLu1957 [host, distribution, economic importance: 420]; Young1980 [taxonomy: 183].



Matsucoccus mugo Siewniak

NOMENCLATURE:

Matsucoccus mugo Siewniak, 1970: 168. Type data: GERMANY: Altenberg, Georgenfelder Hochmoor, on Pinus mugo, 3.vi.1969. Holotype female; type no. II-1. Described: female. Illust. Notes: Depository of type material unknown.



HOST: Pinaceae: Pinus mugo [Siewni1970].

DISTRIBUTION: Palaearctic: Germany [Siewni1970].

GENERAL REMARKS: Description and illustration of adult female by Siewniak (1970).

KEYS: Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 131]; Foldi1998 [catalogue: 429]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; MillerPa1987 [taxonomy: 56]; Rieux1975 [taxonomy, host, distribution: 157-168]; Siewni1970 [taxonomy, description, illustration, host, distribution: 168-172]; TangHa1995 [taxonomy, description: 111-112]; Young1979 [taxonomy: 109]; Young1980 [taxonomy: 183].



Matsucoccus oocarpae Ray & Williams

NOMENCLATURE:

Matsucoccus oocarpae Ray & Williams, 1991: 188. Type data: GUATEMALA: Ostuncalco, on Pinus oocarpa, July 10, 1944; collected E.J. Hambleton. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female. Illust.



FOE: HEMIPTERA Anthocoridae: Elatophilus antennatus Kelton [MendelCaPo1991].

HOST: Pinaceae: Pinus oocarpa [RayWi1991].

DISTRIBUTION: Neotropical: Guatemala [RayWi1991].

GENERAL REMARKS: Description and illustration of adult female by Ray & Williams (1991).

KEYS: Ray & Williams 1991: 190 (female) [North America].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 132]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; MendelCaPo1991 [host, distribution, biological control: 502-507]; RayWi1991 [taxonomy, description, illustration, host, distribution: 188-190].



Matsucoccus paucicicatrices Morrison

NOMENCLATURE:

Matsucoccus paucicicatrices Morrison, 1939: 45. Type data: USA: California, Yosemite National Park, Eight Mile, on Pinus lambertiana, June 27, 1936 & May 10, 1937, collected Hopkins. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA; type no. 31905. Described: female.

COMMON NAME: sugar pine matsucoccus [Gill1993].



HOSTS: Pinaceae: Pinus [Morris1939], Pinus flexilis [Morris1939], Pinus lambertiana [Morris1939], Pinus monticola [Morris1939].

DISTRIBUTION: Nearctic: United States of America (California [Morris1939, Gill1993], Montana [Morris1939], Oregon [Morris1939], Wyoming [Morris1939]).

BIOLOGY: Develops one annual generation in USA, California (Gill, 1993).

GENERAL REMARKS: Description and illustration of adult female by Morrison (1939) and by Gill (1993).

KEYS: Gill 1993: 42 (female) [USA, California]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 132]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 322]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; Gill1993 [taxonomy, description, illustration, host, distribution: 42, 45, 61]; Morris1939 [taxonomy, description, host, distribution: 13-14]; Rieux1975 [taxonomy, host, distribution: 157-168]; SchmutKlLu1957 [host, distribution, economic importance: 420]; Young1980 [taxonomy: 183]; YoungLuZh1986 [taxonomy: 198].



Matsucoccus pini Green

NOMENCLATURE:

Matsucoccus pini Green, 1925: 34. Type data: ENGLAND: Surrey, Oxshott, on Pinus sylvestris, collected F.C. Withycombe. Syntypes, female. Type depository: London: The Natural History Museum, England, UK. Described: female. Illust.

Matsucoccus pine; Hu & Hu, 1981: 180. Misspelling of species name.



FOES: COLEOPTERA Coccinellidae: Adalia bipunctata (L.) [Siewni1976], Coccinella quinquepunctata (L.) [Siewni1976], Harmonia quadripunctata (Pont.) [Siewni1976], Myrrha octodecimguttata (L.) [Siewni1976, UlgentSzUy2013]. Nitidulidae: Scymnus suturalis Thunb. [Siewni1976]. HEMIPTERA Anthocoridae: Elatophilus crassicornis Reuter [MendelCaPo1991], Elatophilus nigricornis Zetterstedt [MendelCaPo1991], Elatophilus pachycnemis Horvath [MendelCaPo1991, UlgentSzUy2013], Elatophilus pini Baerensprung [MendelCaPo1991], Elatophilus roubali Stys [MendelCaPo1991], Elatophilus stigmatellus Zetterstedt [MendelCaPo1991].

HOSTS: Pinaceae: Pinus nigra [Pelliz1987, Trembl1988a], Pinus nigra italica [Pelliz1987], Pinus sylvestris [Green1925, Boraty1952a, Siewni1969, Ossian1972, Rieux1976, Loytty1978, Pelliz1987, Trembl1988a].

DISTRIBUTION: Palaearctic: Austria [Foldi2005]; Bulgaria [Foldi2005]; Czech Republic [Siewni1972]; Finland [Loytty1978]; France [Rieux1976, Foldi2005]; Germany [Siewni1971, Siewni1972]; Hungary [Foldi2005]; Italy [Pelliz1987, Trembl1988a, BarbagBiBo1995, Foldi2005]; Morocco [Foldi2005]; Netherlands [Foldi2005]; Poland [Siewni1971, Siewni1972, LagowsKo1996]; Romania [Foldi2005]; Russia (St. Petersburg (=Leningrad) Oblast [Borchs1950b]); Spain [Cadahi1971, Foldi2005]; Sweden [Ossian1972, Gertss2011]; Turkey [Foldi2005, KaydanUlEr2007]; United Kingdom (England [Green1925]).

BIOLOGY: A parthenogenetic species in Sussex, England (Boratynski, 1951a). Develops two annual generations in Sussex, England (Boratynski, 1951a) and in Katowice, Poland (Siewniak, 1976).

GENERAL REMARKS: Description and illustration of adult female and nymphs by Green (1925), Boratynski (1952a) and by Borchsenius (1950b). Description and illustration of adult female, male and nymphs by Siewniak (1976) and by Rieux (1976). Description and illustration of adult female by Foldi (2004).

STRUCTURE: Boratynski (1952a) reported on seasonal differences in numbers and measurements of certain morphological characters of the adult female.

SYSTEMATICS: Rieux (1976) analysed the intraspecific variation of taxonomic characters in populations from seven localities in France, and concluded that all belong to M. pini.

KEYS: Foldi 2004: 164 [Matsucoccus species in the Mediterranean]; Tang & Hao 1995: 101-102 (female) [Palearctic]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BarbagBiBo1995 [distribution: 39]; BenDov2005a [taxonomy, host, distribution, catalogue: 132-134]; BodenhNe1955 [taxonomy: 10]; Boraty1951a [host, distribution, life history: 28-29]; Boraty1952a [taxonomy, description, illustration, host, distribution, life history: 285-326]; Boraty1960a [life history, host, distribution: 240]; Borchs1950b [taxonomy, description, host, distribution: 33]; BrancoFrOf2005 [life history, chemical ecology, biological control: 234-235]; Buchne1966 [life history, structure: 281-283]; BurzynRo1976 [host, distribution: 1-10]; Cadahi1971 [life history, host, distribution, biological control: 81-91]; CovassPo1986 [host, distribution, biological control: 1-10]; Foldi1998 [catalogue: 429]; Foldi2001 [distribution: 303]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy, description, illustration, host, distribution: 162-164]; Germai2011 [distribution, economic importance: 31-34]; Germai2011a [distribution, economic importance: 8]; Green1925 [taxonomy, description, illustration, host, distribution: 34-35]; HuHu1981 [taxonomy: 178-180]; KaydanUlEr2007 [host, distribution: 98]; Kohler2009a [host, distribution: 24]; KohlerEi2005 [host, distribution: 161]; KohlerEi2006 [host, distribution: 14]; KotejaPyVo2003 [taxonomy, structure: 253]; KozarKoFe2013 [distribution, taxonomy: 56]; KreiteAuGe2006 [distribution, economic importance, host: 143]; LagowsKo1996 [host, distribution: 30]; Lindin1936 [taxonomy: 158]; Loytty1978 [host, distribution: 115-116]; MendelCaPo1991 [host, distribution, biological control: 502-507]; MillerPa1987 [taxonomy: 49, 56]; Morris1928 [taxonomy: 53,226]; Ossian1972 [host, distribution: 97]; Pelliz1987 [taxonomy, host, distribution: 118]; Rieux1975 [taxonomy, host, distribution: 157-168]; Rieux1976 [taxonomy, description, illustration, host, distribution: 231-263]; Siewni1969 [life history, economic importance, host, distribution: 1043-1047]; Siewni1971 [economic importance, life history, host, distribution: 35-41]; Siewni1972a [life history, host, distribution: 29-37]; Siewni1976 [taxonomy, description, illustration, host, distribution, life history: 337-362]; Siewni1983 [economic importance, life history, host, distribution: 49-77]; Siewni1983a [economic importance, life history, host, distribution: 79-93]; SimonKa2011 [distribution: 234]; Tang1978 [taxonomy: 164-170]; TangHa1995 [taxonomy, description: 112]; Trembl1988a [taxonomy, life history, host, distribution: 203-204]; Young1979 [taxonomy: 109]; Young1980 [taxonomy: 183]; Zahrad1959a [taxonomy: 534].



Matsucoccus pinnatus (Germar & Berendt)

NOMENCLATURE:

Monophlebus pinnatus Germar & Berendt, 1856: 3. Type data: Inclusion in Baltic amber. Holotype male. Described: male. Illust. Notes: The type was deposited in Koenigsberg Collection. It disappeared during the Second World War; Jan Koteja, personal information to Yair Ben-Dov; February 2001.

Palaeococcus pinnatus; Cockerell, 1902q: 233. Change of combination.

Acreagris crenata; Ferris, 1941: 6. Misidentification; discovered by Koteja, 1984d: 475.

Monophlebus pinnatus; Ferris, 1941: 7. Incorrect synonymy; discovered by Koteja, 1984d: 453. Notes: Incorrect synonymy with Acreagris crenata.

Acreagris crenata; Beardsley, 1968: 1457. Misidentification; discovered by Koteja, 1984d: 475.

Matsucoccus pinnatus; Koteja, 1984d: 475. Change of combination.

Monophlebus pinatus; Koteja, 1987c: 29. Misspelling of species name.

GENERAL REMARKS: Description and illustration of adult male, adult female and nymph by Koteja (1984d).

SYSTEMATICS: A fossil species, described from inclusions of male and female in Baltic amber (Koteja, 1984d). Koteja (2000c: 2055) summarized the status of this species: "According to Ferris (1941: 6), a junior synonym of Matsucoccus crenata (Koch & Berendt). Redescribed by Ferris (1941) and by Koteja (1984d)."

CITATIONS: Beards1968 [taxonomy: 1457]; BenDov2005a [taxonomy, host, distribution, catalogue: 134]; Cocker1902q [taxonomy: 233]; Ferris1941 [taxonomy: 6-10]; Koteja1984d [taxonomy, description, illustration, distribution: 475-481, 490-492]; Koteja1987c [taxonomy: 29]; Koteja1999 [taxonomy: 863]; Koteja2000c [taxonomy: 205].



Matsucoccus saxonicus Koteja

NOMENCLATURE:

Matsucoccus saxonicus Koteja, 1986: 56. Type data: GERMANY: inclusion in Saxonian amber (Bitterfeld). Holotype female and first instar. Type depository: Berlin: Museum fur Naturkunde der Humboldt Universitat zu Berlin, Germany; type no. MB.I411. Described: male. Illust.



HOST: Pinaceae: Cupressospermum saxonicum [Koteja1986].

DISTRIBUTION: Palaearctic: Germany [Koteja1986].

BIOLOGY: A fossil scale insects, described from two male inclusions (holotype and paratype) in Saxonian amber, dated from the Miocene, about 22 million years (Koteja, 1986).

GENERAL REMARKS: Description and illustration of adult male by Koteja (1986).

CITATIONS: BenDov2005a [pp. 135-136]; Foldi2001a [taxonomy: 208]; Koteja1986 [taxonomy, description, illustration, host, distribution: 55-63]; Koteja1999 [taxonomy: 864]; Koteja2000c [taxonomy: 205].



Matsucoccus secretus Morrison

NOMENCLATURE:

Matsucoccus secretus Morrison, 1939: 14. Type data: U.S.A.: Colorado, Monument, on Pinus ponderosa; collected Carl Hartley, summer 1935. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female.

COMMON NAME: secretive pine scale [Gill1993].



HOST: Pinaceae: Pinus ponderosa [Morris1939, Gill1993].

DISTRIBUTION: Nearctic: Mexico (Chihuahua [Morris1939]). Nearctic: Mexico (Sonora [Morris1939]); United States of America (Arizona [Morris1939], California [Morris1939], Colorado [Morris1939], Nevada [Morris1939], New Mexico [Morris1939]).

GENERAL REMARKS: Description and illustration of adult female by Morrison (1939) and by Gill (1993).

KEYS: Gill 1993: 42 (female) [USA, California]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 136]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 322]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; Gill1993 [taxonomy, description, illustration, host, distribution: 42-46, 62]; Morris1939 [taxonomy, description, host, distribution: 14-17]; Rieux1975 [taxonomy, host, distribution: 157-168]; Young1980 [taxonomy: 183].



Matsucoccus shennongjiaensis Young & Lu in: Young, Lu & Zhan

NOMENCLATURE:

Matsucoccus shennongjiaensis Young & Lu in: Young, Lu & Zhan, 1986: 195. Type data: CHINA: Hubei Province, Shennongjia, on Pinus armandi, July 1986. Syntypes, female. Type depository: Beijing: Institute of Entomology, Academy of Sciences, China. Described: female. Illust.



HOST: Pinaceae: Pinus armandi [YoungLuZh1986, TangHa1995].

DISTRIBUTION: Oriental: China (Hubei (=Hupei) [YoungLuZh1986, TangHa1995]).

GENERAL REMARKS: Description and illustration of adult female by Young et al. (1986).

KEYS: Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 136]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; LuZh1989 [life history, economic importance: 577-582]; TangHa1995 [taxonomy, description, host, distribution: 112-113]; YoungLuZh1986 [taxonomy, description, illustration, host, distribution: 195-198].



Matsucoccus sinensis Chen, F.G.

NOMENCLATURE:

Matsucoccus sinensis Chen, F.G., 1937: 382. Type data: CHINA: Chekiang, Hwengyen (Chi-Lon-Shan), on needles of Pinus sp.; collected January 30 and May 14, 1936. Syntypes, female and first instar. Described: female and first instar. Illust. Notes: The author stated that the types were in his collection.

Sonsaucoccus sinensis; Young, 1980: 186. Change of combination.

Matsucoccus sinensis; Tang & Hao, 1995: 113. Revived combination.

COMMON NAME: Chinese pine needle scale [YoungHuRe1980].



HOSTS: Pinaceae: Pinus [Chen1937], Pinus massoniana [TangHa1995], Pinus tabulaeformis [TangHa1995].

DISTRIBUTION: Oriental: China (Jiangsu (=Kiangsu) [TangHa1995], Shanghai [TangHa1995], Yunnan [Ferris1950], Zhejiang (=Chekiang) [Chen1937, TangHa1995]). Palaearctic: China (Shaanxi (=Shensi) [TangHa1995]).

GENERAL REMARKS: Description and illustration of adult female by Chen (1937), Ferris (1950), Young et al. (1980) and by Tang & Hao (1995). Description and illustration of nymphs by Ferris (1950).

KEYS: Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 136-137]; BodenhNe1955 [taxonomy: 10]; Chen1937 [taxonomy, description, illustration, host, distribution: 382-386]; DaiSuSh2006 [host, distribution, life history, ecology, control: 33-35]; Ferris1950 [taxonomy, description, illustration, host, distribution: 11-12,30-31]; Foldi1998 [catalogue: 429]; Foldi2001a [taxonomy: 208]; Foldi2005 [taxonomy: 166]; Rieux1975 [taxonomy, host, distribution: 157-168]; Tang1978 [taxonomy: 168]; TangHa1995 [taxonomy, description, illustration, host, distribution: 113-114,560,680]; TangWuLi1997 [taxonomy: 17-23]; Yang1982 [host, distribution: 21]; Young1980 [taxonomy: 183,186,188]; YoungHuRe1980 [taxonomy, description, illustration, host, distribution: 42-46].



Matsucoccus subdegeneratus Morrison

NOMENCLATURE:

Matsucoccus subdegeneratus Morrison, 1939: 17. Type data: DOMINICAN REPUBLIC: from herbarium sample (Arnold Arboretum Herbarium) of Pinus occidentalis, collected at Moncion, Monte Cristi Province, October 1929, E.J. Valeur. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female.



HOST: Pinaceae: Pinus occidentalis [Morris1939].

DISTRIBUTION: Neotropical: Dominican Republic [Morris1939, PerezG2008].

KEYS: Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 137]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 322]; Foldi2001a [taxonomy: 208]; Morris1939 [taxonomy, description, host, distribution: 17-18]; PerezG2008 [distribution: 217]; Rieux1975 [taxonomy, host, distribution: 157-168]; Young1980 [taxonomy: 183].



Matsucoccus vexillorum Morrison

NOMENCLATURE:

Matsucoccus vexillorum Morrison, 1939: 18. Type data: USA: Arizona, Prescott National Forest, Copper Basin Area, near Prescott, on Pinus ponderosa, March 11, 1936. Holotype female. Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA. Described: female.

COMMON NAME: prescott scale [McKenzGiEl1948].



HOST: Pinaceae: Pinus ponderosa [Morris1939, McKenzGiEl1948].

DISTRIBUTION: Nearctic: United States of America (Arizona [Morris1939, McKenzGiEl1948], New Mexico [Morris1939, McKenzGiEl1948]).

GENERAL REMARKS: Description and illustration of adult female by Morrison (1939) and by McKenzie (1943a, 1943b).

ECONOMIC IMPORTANCE AND CONTROL: In Arizona, USA, this species was reported to cause twig blight and 'flagging' of ponderosa pine, Pinus ponderosa (McKenzie et al., 1948)

KEYS: Ray & Williams 1991: 190 (female) [North America]; Miller & Park 1987: 54-55 (female) [World]; Ray & Williams 1984: 768-769 (female) [North America]; Boratynski 1952a: 321-324 (female) [World].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue, economic importance: 138-139]; BodenhNe1955 [taxonomy: 10]; Boraty1952 [taxonomy: 323]; Foldi2001a [taxonomy: 208]; McKenz1943a [taxonomy, host, distribution, life history, structure: 42-52]; McKenz1943b [taxonomy, description, illustration, host, distribution: 53-57]; McKenzGiEl1948 [economic importance, life history, host, distribution: 33-51]; Morris1939 [taxonomy, description, host, distribution: 18-20]; Rieux1975 [taxonomy, host, distribution: 157-168]; SchmutKlLu1957 [host, distribution, economic importance: 420]; Young1980 [taxonomy: 183].



Matsucoccus yunnanensis Ferris

NOMENCLATURE:

Matsucoccus yunnanensis Ferris, 1950: 12. Type data: CHINA: Yunnan Province, An-lin-wen-chian, near Kunming, on Pinus yunnanensis, April 29, 1949,. Syntypes, female. Type depository: Davis: The Bohart Museum of Entomology, University of California, California, USA. Described: female. Illust.



HOST: Pinaceae: Pinus yunnanensis [Ferris1950, YoungHuRe1976, TangHa1995].

DISTRIBUTION: Oriental: China (Yunnan [Ferris1950, YoungHuRe1976, TangHa1995]).

BIOLOGY: Occurring in and beneath the bark of the host, the apodous stage flattened between the layers of bark (Ferris, 1950).

GENERAL REMARKS: Description and illustration of adult female by Ferris (1950), Young et al. (1976) and by Tang & Hao (1995).

KEYS: Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 139]; BodenhNe1955 [taxonomy: 10]; Ferris1950 [taxonomy, description, illustration, host, distribution: 12, 31]; Foldi1998 [catalogue: 429]; Foldi2001a [taxonomy: 208]; MillerPa1987 [taxonomy: 56]; QiWa1981 [economic importance, host, distribution: 20-25]; Rieux1975 [taxonomy, host, distribution: 157-168]; TangHa1995 [taxonomy, description, host, distribution: 114-115,560]; Yang1982 [host, distribution: 21]; Young1980 [taxonomy: 183].



Matsucoccus yunnansonsaus Young & Hu, in: Young, Hu & Ren

NOMENCLATURE:

Matsucoccus yunnansonsaus Young & Hu, in: Young, Hu & Ren, 1980: 46. Type data: CHINA: Yunnan Province, Kunming, on Pinus yunnanensis. Syntypes, female. Type depository: Beijing: Institute of Entomology, Academy of Sciences, China. Described: female. Illust.

COMMON NAMES: Yunnan pine needle scale [YoungHuRe1980].



HOST: Pinaceae: Pinus yunnanensis [YoungHuRe1980, TangHa1995].

DISTRIBUTION: Oriental: China (Yunnan [YoungHuRe1980, TangHa1995]).

GENERAL REMARKS: Description and illustration of adult female by Young et al. (1980) and by Tang & Hao (1995).

KEYS: Tang & Hao 1995: 101-102, 608-609 (female) [Palearctic].

CITATIONS: BenDov2005a [taxonomy, host, distribution, catalogue: 139]; Foldi2001a [taxonomy: 208]; TangHa1995 [taxonomy, description, host, distribution: 115-116,560]; Yang1982 [host, distribution: 21]; Young1980 [taxonomy: 183]; YoungHuRe1980 [taxonomy, description, illustration, host, distribution: 42-46].