Valid Names Results

Howardia biclavis (Comstock, 1883) (Diaspididae: Howardia)

Nomenclatural History

Chionaspis? biclavis Comstock 1883: 98-100. Type data: United States: Washington D.C., Department of Agriculture conservatory, on Diospyrus ebenum, Ficus laurifolia, Tamarindus sp., Ochras sapota and Etaecarpus cyanus. Syntypes, female, Type depository: Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA; accepted valid name Illustr.
Aspidiotus theae; Green 1890: 13. misidentification (discovered by Ferris1941e, 49).
Chionaspis biclavis detecta Maskell 1895b: 49. Type data: SANDWICH ISLANDS: Hawaii, Kona. Unknown type status, unknown, Type depository: Auckland: New Zealand Arthropod Collection, Landcare Research, New Zealand; London: The Natural History Museum, England, UK; Washington: United States National Entomological Collection, U.S. National Museum of Natural History, District of Columbia, USA; junior synonym (discovered by Borchs1966, 156).
Howardia biclavis (Comstock, 1883); Berlese & Leonardi 1896: 348. change of combination
Chionaspis (Howardia) biclavis (Comstock, 1883); Maxwell-Lefroy 1902: 250. change of combination
Howardia biclavis detecta (Maskell, 1895); Fernald 1903b: 227. change of combination
Megalodiaspis biclavis (Comstock, 1883); Paoli 1915: 256. change of combination
Howardia (Chionaspis) biclavis (Comstock, 1883); Ramakrishna Ayyar 1921a: 353. change of combination

Common Names

Mining scale Comsto1883
Burrowing scale Borchs1966
cochinilla gris de la jagua WatsonKo2022a
kawa-muguri-kaigara-mushi WatsonKo2022a

Ecological Associates

Hosts:

Families: 76 | Genera: 221

Acanthaceae

Graptophyllum pictum | Ballou1926
Justicia | Borchs1966
Justicia spicigera | Ballou1926 | (= Jacobinia mohintli)
Strobilanthes persicifolia | MacGow1982 | (= Strobilanthes isophyllus)
Thunbergia grandiflora | MestreHaEv2011

Achariaceae

Hydnocarpus | Borchs1966
Hydnocarpus wightianus | Mamet1943a

Amaranthaceae

Alternanthera | MillerDa2005
Iresine | HodgsoHi1990

Anacardiaceae

Cotinus | HodgsoHi1990
Mangifera | MillerDa2005
Mangifera indica | Ballou1926
Rhus copallina | Ballou1926
Rhus leucantha | Ballou1926
Schinus | HodgsoHi1990
Toxicodendron | HodgsoHi1990

Annonaceae

Annona | MillerDa2005
Annona muricata | Morgan1892
Annona reticulata | Mamet1949
Annona squamosa | Marlat1921a
Cananga | MillerDa2005
Monodora | MillerDa2005
Polyalthia longifolia | Sankar1984

Apiaceae

Centella | MillerDa2005

Apocynaceae

Allamanda | MillerDa2005
Allamanda cathartica | Ballou1926 | (= Allamanda hendersonii)
Allamanda schottii | Cohic1958 | (= Allamanda cathartica schottia)
Anodendron | MillerDa2005
Dischidia | MillerDa2005
Echites | Archan1929
Landolphia heudelotii | Archan1929
Melodinus orientalis | MilleHeCa2016 | (= Tabernaemontana orientalis)
Nerium oleander | ColonFMe1998
Plumeria | MillerDa2005
Plumeria rubra | Cohic1958 | (= Plumeria acuminata)
Saba comorensis | Archan1929 Archan1929 | (= Landolphia florida)
Tabernaemontana | MillerDa2005
Tabernaemontana citrifolia | Ballou1926
Tabernaemontana divaricata | Maxwel1902 | (= Tabernaemontana coronaria)
Trachelospermum jasminoides | Kuwana1909a

Aquifoliaceae

Ilex nitida | ColonFMe1998 | var. repanda (= Ilex riedlaei)

Araceae

Lasia spinosa | BugajNJuKa2021
Monstera | MillerDa2005

Arecaceae

Dypsis lutescens | ColonFMe1998 | (= Chrysalidocarpus lutescens)

Asteraceae

Artemisia abrotanum | Ballou1926
Centaurea | MillerDa2005
Cosmos caudatus | ColonFMe1998
Eupatorium | HodgsoHi1990
Microglossa zeylanica | Ali1969a
Tithonia | Borchs1966
Tithonia diversifolia | Mamet1943a

Begoniaceae

Begonia | Borchs1966

Bignoniaceae

Bignonia | Brain1919
Delostoma | MillerDa2005 | MillerDa2005
Spathodea | MillerDa2005
Tabebuia | MillerDa2005
Tecomaria capensis | Ballou1926 | (= Tecoma capensis)

Bixaceae

Bixa | MillerDa2005
Bixa orellana | Ballou1926

Boraginaceae

Cordia | MillerDa2005
Cordia curassavica | Mamet1943a Mamet1949 | (= Cordia interrupta) (= Cordia macrostachya)
Varronia polycephala | ColonFMe1998 | (= Cordia polycephala)

Burseraceae

Bursera simaruba | Ballou1926 | (= Bursera gummifera)

Cactaceae

Opuntia ficus-indica | BugajNJuKa2021

Calophyllaceae

Calophyllum | Hinckl1963
Calophyllum brasiliense | ColonFMe1998
Calophyllum calaba | MestreHaEv2011
Mammea | MillerDa2005
Mammea americana | Beatty1944 DonesEv2011

Calycanthaceae

Calycanthus | MillerDa2005

Cannabaceae

Celtis sinensis | Kuwana1909a
Trema micrantha | ColonFMe1998

Capparaceae

Steriphoma | MillerDa2005

Caprifoliaceae

Lonicera | Borchs1966 MillerDa2005

Caricaceae

Carica | MillerDa2005
Carica papaya | Mamet1943a

Caryocaraceae

Caryocar | MillerDa2005

Casuarinaceae

Casuarina | Borchs1966 MillerDa2005

Celastraceae

Euonymus | HodgsoHi1990

Chrysobalanaceae

Atuna racemosa | Hinckl1963 | (= Parinari laurinum)
Licania | MillerDa2005
Parinari | Hinckl1963

Clusiaceae

Clusia | Hua2000 MillerDa2005
Clusia flava | Balach1954e
Clusia rosea | ColonFMe1998

Combretaceae

Combretum fruticosum | Archan1929 | (= Combretum micropetalum)
Combretum indicum | Ballou1926 | (= Quisqualis indica)
Terminalia | MillerDa2005
Terminalia catappa | Kuwana1909a
Terminalia elliptica | Balach1954e | (= Terminalia tomentosa)

Convolvulaceae

Ipomoea tiliacea | ColonFMe1998

Ebenaceae

Diospyros ebenum | Comsto1883
Royena | Borchs1966

Elaeocarpaceae

Elaeocarpus | Borchs1966 MillerDa2005

Erythroxylaceae

Erythroxylum | MillerDa2005

Euphorbiaceae

Acalypha | Maxwel1902
Acalypha grandis | Mamet1943a
Aleurites | WatsonKo2022a
Codiaeum variegatum | ColonFMe1998
Croton | LincanHoCa2010 MillerDa2005
Croton discolor | Maxwel1902
Croton lucidus | Ballou1926
Euphorbia | Borchs1966 MillerDa2005
Euphorbia pulcherrima | Brain1919 | (= Poinsettia)
Hippomane | Mosque1976
Hippomane mancinella | Ballou1926
Jatropha integerrima | Ballou1926 | (= Jatropha hastata)
Mabea | Kuwana1909a
Macaranga | Hinckl1963
Sapium | Borchs1966 MillerDa2005

Fabaceae

Acacia macracantha | LincanHoCa2010
Acacia mollissima | Hinckl1963
Acacia oshanesii | MillerDa2005
Acacia tenuifolia | MestreHoHa2015
Albizia | WatsonKo2022a
Ammodendron | MillerDa2005
Andira inermis | ColonFMe1998
Bauhinia | MillerDa2005
Bauhinia monandra | ColonFMe1998
Bauhinia purpurea | DeLott1967a
Bauhinia tomentosa | BugajNJuKa2021
Bauhinia variegata | Ballou1926
Caesalpinia | MillerDa2005
Caesalpinia insolita | MillerDa2005
Cajanus cajan | Brown1965
Cassia | HodgsoHi1990
Derris | MillerDa2005
Desmodium incanum | ColonFMe1998
Erythrina | MillerDa2005
Erythrina droogmansiana | MillerDa2005
Hymenaea | MillerDa2005
Indigofera | Watson2002a
Inga | MillerDa2005
Inga laurina | ColonFMe1998
Leucaena | Borchs1966 MillerDa2005
Leucaena leucocephala | MestreHaEv2011
Mundulea sericea | Cohic1958 | (= Mundulea suberosa)
Pithecellobium dulce | ColonFMe1998
Pongamia | MillerDa2005
Pterocarpus indicus | ColonFMe1998
Tamarindus | Comsto1883
Wisteria | Borchs1966 MillerDa2005

Fagaceae

Castanea | Borchs1966

Gesneriaceae

Gesneria pedunculosa | ColonFMe1998

Hypericaceae

Hypericaceae | HodgsoHi1990 | (= Ascyrum)

Juglandaceae

Juglans | Borchs1966

Lamiaceae

Coleus amboinicus | ColonFMe1998 | (= Plectranthus amboinicus)

Lauraceae

Cinnamomum | Borchs1966 MillerDa2005
Ocotea insularis | PetersHaMo2020s1 | (= Ocotea ira)
Ocotea leucoxylon | ColonFMe1998
Persea | MillerDa2005
Persea americana | ColonFMe1998
Phoebe | MillerDa2005

Loranthaceae

Loranthus | MillerDa2005

Lythraceae

Cuphea | HodgsoHi1990
Lagerstroemia indica | Kuwana1909a
Punica | MillerDa2005
Punica granatum | Kuwana1909a

Malpighiaceae

Malpighia | HodgsoHi1990

Malvaceae

Althaea | HodgsoHi1990
Corchorus siliquosus | MestreHaEv2011
Hibiscus | MillerDa2005 Simmon1957
Hibiscus mutabilis | Ballou1926
Hibiscus rosa-sinensis | Mamet1943a
Hibiscus syriacus | Ballou1926
Hibiscus tiliaceus | Kuwana1909a
Matisia | MillerDa2005
Matisia cordata | Mosque1976
Pachira | MillerDa2005
Sida | Watson2002a
Theobroma | MillerDa2005
Thespesia | MillerDa2005 | (= Montezuma)
Triumfetta semitriloba | ColonFMe1998
Urena | Watson2002a
Waltheria | MillerDa2005
Waltheria ovata | LincanHoCa2010

Melastomataceae

Miconia laevigata | ColonFMe1998

Meliaceae

Cedrela | MillerDa2005
Guarea guidonia | ColonFMe1998
Khaya | Watson2002a
Melia azedarach | Simmon1957
Swietenia | Watson2002a
Trichilia | Brain1919 MillerDa2005

Moraceae

Artocarpus integer | Kuwana1909a | (= Artocarpus integrifolius)
Ficus | MillerDa2005
Ficus aspera | Ballou1926 | (= Ficus parcellii )
Ficus benghalensis | BugajNJuKa2021
Ficus benjamina | Morgan1892
Ficus carica | Ballou1926
Ficus laurifolia | Comsto1883
Ficus ulmifolia | BugajNJuKa2021
Morus nigra | Ballou1926

Moringaceae

Moringa oleifera | Ballou1926 Mamet1943a | (= Moringa pterygosperma)

Myricaceae

Morella cerifera | BesheaTiHo1973 | (= Myrica cerifera)
Myrica | MillerDa2005

Myristicaceae

Myristica macrantha | Hinckl1963

Myrtaceae

Metrosideros | Brown1965
Psidium | MillerDa2005
Psidium guajava | ColonFMe1998
Syzygium cleyerifolium | Kuwana1909a

Nyctaginaceae

Bougainvillea | MillerDa2005
Neea | Kuwana1909a

Ochnaceae

Ochna | Comsto1883

Oleaceae

Jasminum | Ballou1926 MillerDa2005
Jasminum humile | Ballou1926
Jasminum sambac | Cohic1958 Wilson1921 | (= Nyctanthes sambac)
Jasminum simplicifolium | Ballou1926
Ligustrum | Borchs1966 MillerDa2005
Olea europaea | Kuwana1909a

Onagraceae

Oenothera | HodgsoHi1990 | (= Raimannia)

Passifloraceae

Passiflora | MillerDa2005
Passiflora laurifolia | ColonFMe1998

Phytolaccaceae

Petiveria alliacea | BugajNJuKa2021
Phytolacca rivinoides | ColonFMe1998

Piperaceae

Piper | MillerDa2005
Piper aduncum | ColonFMe1998
Piper majusculum | WilliaBu1987
Piper nigrum | Tao1999
Piper rivinoides | Archan1929

Pittosporaceae

Pittosporum | HodgsoHi1990

Polygonaceae

Antigonon | MillerDa2005

Proteaceae

Grevillea | MillerDa2005
Grevillea robusta | Mamet1949
Macadamia | MillerDa2005 Willia1973
Macadamia ternifolia | Willia1973

Putranjivaceae

Drypetes alba | MestreHoHa2015

Rosaceae

Cydonia oblonga | Ballou1926
Malus | Borchs1966 MillerDa2005
Malus pumila | Hua2000
Osteomeles anthyllidifolia | Kuwana1909a
Photinia | MillerDa2005
Photinia wrightiana | Kuwana1909a
Prunus | MillerDa2005 Watson2002a
Pyracantha | MillerDa2005
Pyrus | Borchs1966 MillerDa2005
Rhaphiolepis | Brain1919

Rubiaceae

Catesbaea spinosa | Ballou1926
Chiococca | HodgsoHi1990
Cinchona | MillerDa2005
Cinchona pubescens | DEDAC1923 | (= Cinchona succirubra)
Coffea | MillerDa2005
Coffea arabica | Ballou1926
Gardenia | Hinckl1963 MillerDa2005
Gardenia jasminoides | Ballou1926 ColonFMe1998 | (= Gardenia augusta) (= Gardenia florida)
Genipa | MillerDa2005
Genipa americana | Ballou1926 | (= Genipa caruto)
Guettarda scabra | FDACSB1983
Hamelia patens | MestreHoHa2015
Ixora | BesheaTiHo1973 MillerDa2005
Ixora coccinea | Ballou1926 ColonFMe1998 | (= Ixora bandhuca)
Morinda | Borchs1966
Mussaenda | MillerDa2005
Neonauclea | WatsonKo2022a
Notopleura uliginosa | ColonFMe1998 | (= Psychotria uliginosa)
Randia | MillerDa2005
Randia aculeata | ColonFMe1998

Rutaceae

Citrus | MillerDa2005
Citrus aurantiifolia | ColonFMe1998
Citrus australasica | Ballou1926 | (= Microcitrus australasica)
Citrus limon | ColonFMe1998
Citrus maxima | JansenAl2023
Citrus medica | ColonFMe1998
Citrus sinensis | LincanHoCa2010
Glycosmis pentaphylla | Ballou1926
Harrisonia | MillerDa2005 | (= Feroniella)
Murraya paniculata | Ballou1926 | (= Murraya exotica)
Ruta chalepensis | Ballou1926

Salicaceae

Casearia | MillerDa2005
Casearia sylvestris | ColonFMe1998
Dovyalis | MillerDa2005
Flacourtia | Borchs1966

Santalaceae

Phoradendron | Mosque1976

Sapindaceae

Blighia sapida | Ballou1926
Cupania | Borchs1966 MillerDa2005
Dimocarpus longan | HowellBeTi1986 | (= Nephelium longan)
Dodonaea | HodgsoHi1990
Litchi | MillerDa2005 Watson2002a
Melicoccus bijugatus | ColonFMe1998
Nephelium | MillerDa2005
Thouinia striata | ColonFMe1998

Sapotaceae

Chrysophyllum | MillerDa2005
Chrysophyllum cainito | Ballou1926
Manilkara | MillerDa2005 | (= Achras) (= Lucuma)
Manilkara bidentata | ColonFMe1998
Manilkara zapota | Ballou1926 Cohic1958 ColonFMe1998 Marlat1921a | (= Achras zapota) (= Achras sapota) (= Lucuma mammosa)
Micropholis guyanensis | ColonFMe1998 | (= Micropholis chrysophylloides)
Mimusops | MillerDa2005
Planchonella obovata | Kuwana1909a | (= Sideroxylon ferrugineum)
Pouteria | MillerDa2005
Pouteria campechiana | Ballou1926 | (= Lucuma nervosa)
Pouteria dominigensis | Ballou1926 | (= Lucuma serpentaria)

Scrophulariaceae

Buddleja davidii | Ballou1926

Solanaceae

Brugmansia suaveolens | ColonFMe1998 | (= Datura suaveolens)
Brunfelsia nitida | Ballou1926
Brunsfelsia | MillerDa2005
Cestrum diurnum | Ballou1926
Solanum | ColonFMe1998 Watson2002a | (= Lycopersicon)
Solanum melongena | Ballou1926
Solanum seaforthianum | Ballou1926

Strelitziaceae

Strelitzia | BugajNJuKa2021

Theaceae

Camellia | Borchs1966 MillerDa2005 | (= Thea)
Camellia sinensis | Tao1999

Verbenaceae

Citharexylum | HodgsoHi1990
Duranta | MillerDa2005
Duranta erecta | Ballou1926 Mamet1943a | (= Duranta plumieri) (= Duranta repens)
Lantana | MillerDa2005
Lantana camara | ColonFMe1998
Petrea | MillerDa2005
Phyla stoechadifolia | ColonFMe1998 | (= Lippia stoechadifolia)
Stachytarpheta jamaicensis | ColonFMe1998

Vitaceae

Cissus verticillata | ColonFMe1998 | (= Cissus sicyoides)
Parthenocissus | HodgsoHi1990

Foes:

Families: 5 | Genera: 8

Aphelinidae

Aphytis diaspidis | Balach1954e | (= Aphytis fuscipennis)
Aphytis holoxanthus | LazaroGoRo2023
Aphytis proclia | Balach1954e
Aphytis theae | Morley1909 | (= Aphelinus theae)
Marietta busckii | LazaroGoRo2023
Pteroptrix albifemur | Fulmek1943 | (= Pteroptrix imitatrix)

Cecidomyiidae

Dyodiplosis generosi | Fulmek1943

Encyrtidae

Plagiomerus | Simmon1957
Zaomma | LazaroGoRo2023

Phlaeothripidae

Karnyothrips merrilli | Fulmek1943 | (= Haplothrips merilli)

Signiphoridae

Signiphora dozieri | LazaroGoRo2023
Signiphora perpauca | LazaroGoRo2023

Geographic Distribution

Countries: 84

Antigua and Barbuda

Antigua | Maxwel1902
Antigua | WalkerWaEv1974

Argentina

Buenos Aires | Watson2002a

Australia

Queensland | Watson2002a

Bahamas | Marlat1921a

Barbados | Maxwel1902

Bermuda | Brown1965

Bonin Islands (=Ogasawara-Gunto) | Kuwana1909a

Brazil | Bondar1914

Bahia | SilvadGoGa1968
Espirito Santo | MartinWoCu2022
Minas Gerais | SilvadGoGa1968
Para | MartinWoCu2022
Pernambuco | MartinWoCu2022
Rio Grande do Sul | MartinWoCu2022
Rio de Janeiro | SilvadGoGa1968
Santa Catarina | MartinWoCu2022
Sao Paulo | SilvadGoGa1968

Canary Islands | WatsonKo2022a

China

Guangdong (=Kwangtung) | Hua2000
Guangxi (=Kwangsi) | Hua2000
Yunnan | Tao1999

Colombia | Figuer1952 Mosque1976 NormarOkMo2019

Cook Islands | Watson2002a WilliaWa1988 | WatsonKo2022a

Costa Rica | WatsonKo2022a

Croatia | WatsonKo2022a

Cuba | Ballou1926 MestreHaEv2011

Czech Republic | Zahrad1953

Dominica | Mamet1943a

Dominican Republic | GomezM1941

Ecuador | Nakaha1982

Egypt | WatsonKo2022a

Federated States of Micronesia

Truk Islands | Takaha1942d

Fiji | Hinckl1963

France | DanzigPe1998 FoldiGe2018

French Guiana | WatsonKo2022a

French Polynesia

Tahiti | Green1907

Galapagos Islands | LincanHoCa2010 Willia1977ML

Germany | Lindin1909b

Ghana | Hall1946a

Grenada | WatsonKo2022a

Guadeloupe | Vayssi1955

Guatemala | JansenAl2023

Guiana | Nakaha1982

Guyana (=British Guiana) | Bodkin1914

Haiti | BerlesLe1898a

Hawaiian Islands | Maxwel1902 MillerDa2005

Maui | Nishid2002
Niihau | Nishid2002
Oahu | Nishid2002

India | Nakaha1982

Karnataka | Singh1964
Tamil Nadu | Fletch1919

Indonesia | Nakaha1982

Java | Clause1933

Ireland | Green1934d

Italy | Leonar1918

Sicily | RussoMa1997

Jamaica | Brown1965

Japan | Tao1999

Kenya | DeLott1967a

Madagascar | Mamet1959a

Madeira Islands | FrancoRuMa2011

Malawi | WatsonKo2022a

Malaysia | Suh2016

Martinique | Vayssi1955

Mauritius | Mamet1943a WilliaWi1988

Mexico | Cocker1899n MillerDa2005

Colima | LazaroGoRo2023
Michoacan | LazaroGoRo2023
Nayarit | LazaroGoRo2023

Montserrat | Maxwel1902

New Caledonia | Cohic1956

Niue | WilliaWa1988

Palau | Beards1966

Panama | Watson2002a

Papua New Guinea | WilliaWa1988

Peru | Nakaha1982

Philippines | Nakaha1982

Poland | Koteja1974b

Portugal | FrancoRuMa2011

Puerto Rico & Vieques Island

Puerto Rico | DonesEv2011 Maxwel1902

Rodriques Island | Mamet1956b WilliaWi1988

Russia

St. Petersburg (=Leningrad) Oblast | Archan1929

Saint Croix | MiskimBo1970

Saint Kitts and Nevis Islands

Saint Kitts | Maxwel1902

Saint Lucia | Malump2014a

Sao Tome and Principe

Sao Tome | Hall1946a

Singapore | Nakaha1982

South Africa | Brain1919

Spain | DanzigPe1998

Sri Lanka | Maxwel1902

Sweden | WatsonKo2022a

Switzerland | WatsonKo2022a

Taiwan | Nakaha1982

Tanzania | WatsonKo2022a

Tonga | Lindin1911

Trinidad and Tobago

Trinidad | Maxwel1902

United Kingdom

England | Maxwel1902

United States

California | MillerDa2005 Watson2002a
District of Columbia | Comsto1883
Florida | MerrilCh1923 MillerDa2005
Georgia | WatsonKo2022a
Kansas | Lawson1917
Louisiana | Miller2005
Maryland | MillerDa2005 Nakaha1982
Missouri | MillerDa2005 Nakaha1982
New York | MillerDa2005 Nakaha1982
Ohio | MillerDa2005 WebsteBu1902
Pennsylvania | MillerDa2005 Trimbl1928

Vanuatu (=New Hebrides) | WilliaBu1987

Venezuela | Ballou1945 | Watson2002a

Wallis and Futuna Islands

Wallis Island | WatsonKo2022a

Western Samoa | WilliaWa1988

Zanzibar | Mamet1943a

Zimbabwe | Hall1935

Keys

LazaroGoRo2023: pp.247-248 ( Adult (F) ) [Armored scale species on avocado in central Mexico]
SuhJi2009: pp.1041-1043 ( Adult (F) ) [Key to species of armored scales intercepted on imported plants (slide mounted females)]
MillerDa2005: pp.29-34 ( Adult (F) ) [Field Key to Economic Armored Scales]
MillerDa2005: pp.20 ( Adult (F) ) [Armored Scales]
WolffCo1993a: pp.153 ( Adult (F) ) [Diaspidid species on mango in Brazil]
WilliaWa1988: pp.137 ( ) [Key to species of Howardia]
Fullaw1932: pp.96 ( Adult (F) ) [Key to species of Hawaiian Diaspinae]
Comsto1916: pp.559 ( Adult (F) ) [Key to species of Chionaspis] Key as: Chionaspis biclavis
Green1899a: pp.108 ( Adult (F) ) [Synopsis of Chionaspis species] Key as: Chionaspis biclavis
Comsto1881a: pp.98 ( ) [Key to species of Chionaspis] Key as: Chionaspis biclavis

Remarks

Structure: Female scale white, but obscured by tissue of host, nearly circular. Exuviae marginal and project beyond the edge of scale (Comstock, 1883). Adult female scale cover circular and convex, 2.5‒3.0 mm in diameter, yellowish white to brown, with fibrous appearance, exuviae subcentral. Body of live adult female membranous, slightly brown, or gray, 2.4‒2. 9 mm long, occupying almost the entire diameter of the waxy cover. (Lazaro et al., 2023). Crawler pale purple (Watson & Kondo 2022).
Biology: According to Brown (1965) Howardia biclavis reproduces parthenogenetically without males. Kuwana (1911) and Murakami (1970) indicate that it has 1 generation each year in Japan. Of particular interest is the manner in which this pest becomes covered by a thin layer of the host's epidermis. Many comment in the literature about how the scale mines or burrows into the host, but these statements almost certainly are inaccurate since the crawlers have no unusual adaptations for burrowing. It seems more likely that crawlers find cracks in the bark and settle in the bark tissue. As the scale cover enlarges, it slowly forces its way under the epidermal layer. This hypothesis is substantiated by Fennah (1947) who observed that crawlers settle in minute crevices of the bark. To our knowledge, a detailed study of the biology of this species has not been under taken. According to Brown (1965) mining scale reproduces parthenogenetically without males. Kuwana (1911) and Murakami (1970) indicate that it has 1 generation each year in Japan. Of particular interest is the manner in which this pest becomes covered by a thin layer of the host's epidermis. There are many comments in the literature about how the scale mines or burrows into the host, but these statements almost certainly are inaccurate since the crawlers have no unusual adaptations for burrowing. It seems more likely that crawlers find cracks in the bark and settle in the bark tissue. As the scale cover enlarges, it slowly forces its way under the epidermal layer. This hypothesis is substantiated by Fennah (1947) who observed that crawlers settle in minute crevices of the bark. (Miller & Davidson, 2005).
Economic Importance: Miller & Davidson (1990) list this insect as a serious and widespread pest. Dekle (1977) reported the mining scale as an economic pest on many woody ornamentals in Florida despite the fact that it often is parasitized. Azevedo Marques (1923) considered it to be an important economic pest of coffee in Brazil, and Plank and Winters (1949) described it as causing heavy damage to Cinchona ledgeriana in Puerto Rico. It also has been reported to cause damage to tea in Sri Lanka (Ramakrishna Ayyar 1930), India, Japan, and Malawi (Nagarkatti and Sankaran 1990), to citrus in Cuba (Grillo et al. 1983), to coffee in Cuba (Köhler et al. 1980), and to macadamia in India (Ebeling 1959). Beardsley and González (1975) consider this scale to be one of 43 serious armored scale pests, and Miller and Davidson (1990) consider it to be a serious world pest. (Miller & Davidson, 2005).
General Remarks: Descriptions and illustrations by Comstock (1883) and Balachowsky (1954e). Color photograph in Watson & Kondo (2022a).

Illustrations

Citations

AAEE1931: taxonomy, 1284
Ali1969a: distribution, host, 51
AlmeidPeMa2018: distribution, host, 355
AndersWuGr2010: phylogeny, taxonomy, 997-1003
Archan1929: distribution, host, 197
Arnett1985: economic importance, 241
Balach1954e: biological control, description, distribution, host, illustration, taxonomy, 252-254
Ballou1926: catalog, distribution, host, 25-26
Ballou1945: distribution, host, 93
Beards1966: distribution, host, 533
Beatty1944: distribution, host, 126
BerlesLe1896: taxonomy, 348
BerlesLe1898a: description, distribution, host, illustration, taxonomy, 127-129
BesheaTiHo1973: distribution, host, 11
Bodkin1914: distribution, host, 114
BoedigSt1930: host, 51-52
Bondar1914: distribution, host, 1103
Borchs1937: taxonomy, 112
Borchs1937a: taxonomy, 115
Borchs1950b: distribution, host, taxonomy, 210
Borchs1966: catalog, distribution, host, taxonomy, 156-157
Boyce1950: taxonomy, 745
Brain1919: description, distribution, host, illustration, taxonomy, 219-220
Brown1965: chemistry, 132-137
BrunerScOt1945: catalog, distribution, host, 76
BugajNJuKa2021: distribution, host, list, 299
Butche1959: distribution, host, 364
Chou1982: description, distribution, host, taxonomy, 124-125
Chou1986: illustration, 525
Clause1933: distribution, 16
Cocker1896a: taxonomy, 256
Cocker1899n: distribution, host, 30
Cocker1899r: taxonomy, 900
Cocker1902p: distribution, 257
Cohic1956: distribution, host, 15, 16, 20, 58
Cohic1958: distribution, host, 16, 25, 30, 34
ColonFMe1998: description, distribution, host, illustration, taxonomy, 100-101
Comsto1883: description, distribution, host, illustration, taxonomy, 98-100
Comsto1916: description, distribution, host, illustration, taxonomy, 559-561
CoronaRuMo1997: distribution, economic importance, host, 40
CostaL1928: distribution, host, 119
CostaL1942: distribution, host, 272
Craw1896: taxonomy, 35, 40
DEDAC1923: distribution, host, 15
Danzig1993: description, distribution, host, illustration, taxonomy, 287
DanzigPe1998: catalog, distribution, host, taxonomy, 275-276
Dash1916: distribution, host, 42
DeLott1967a: distribution, host, 115
DeSilv1961: biological control, distribution, 119
DeitzTo1980: distribution, taxonomy, 35
Dekle1954: distribution, host, 227
Dekle1965c: distribution, host, taxonomy, 11, 74
DixLuMoKu2019: distribution, host,
DodgeRi1943: distribution, taxonomy, 346
DonesEv2011: distribution, host, 2
Draper1907: distribution, 10
FDACSB1983: distribution, host, 7
Felt1915: taxonomy, 183
Fennah1947: distribution, 65
Fernal1903b: distribution, host, taxonomy, 227
Ferris1920a: taxonomy, 63
Ferris1936a: illustration, taxonomy, 21, 22, 28, 60
Ferris1937: description, distribution, host, illustration, taxonomy, SI-64-65
Ferris1941e: taxonomy, 49
Figuer1952: distribution, 210
Fletch1914: distribution, 519
Fletch1917a: distribution, host, 34, 37
Fletch1919: distribution, host, 297
Foldi2001: distribution, 306
FoldiGe2018: distribution, 10
Fonsec1964: distribution, host, 515
FrancoRuMa2011: distribution, 12,24
Fullaw1932: distribution, host, 96
Fulmek1943: biological control, catalog, distribution, 36
Gavril2018: reproduction, 240
GermaiDeLa2016: host, list of species, 3
Gill1997: distribution, illustration, taxonomy, 163, 164
GomezM1941: distribution, host, 137
GrandpCh1899: distribution, host, 30
Green1890: distribution, host, 13
Green1896: distribution, host, 2
Green1899a: distribution, host, 108, 152
Green1907: distribution, host, 201
Green1908a: distribution, host, 36
Green1922: distribution, host, 460
Green1934d: distribution, 114
Green1937: catalog, distribution, 324
GruwelWoKo2016: phylogeny, 173, 175, 176
Hall1935: distribution, host, 84
Hall1946a: distribution, 548
Hart1896a: distribution, vi
HertinSi1972: biological control, 181
Hinckl1963: distribution, host, 45
HodgsoHi1990: distribution, host, 2, 3, 5-7, 9, 12-19,
HodgsoLa2011: distribution, host, 24
Houser1918: distribution, host, 161
HowellBeTi1986: description, distribution, host, illustration, 1-3
Hua2000: distribution, host, taxonomy, 153
JansenAl2023: dispersal, host, 26,34
Kawai1980: distribution, taxonomy, 260
KawaiMaUm1971: distribution, host, 22
Kirkal1902: distribution, 112
Kirkal1904b: distribution, host, 157
KondoMu2016: distribution, host, illustration, 6
KondoWa2022a: distribution, host, list, 17
Koteja1974b: structure, 84
KozarWa1985: catalog, distribution, 84
Kuwana1909a: distribution, host, 161
Kuwana1917a: distribution, host, 16
Kuwana1926: distribution, host, taxonomy, 4
LagowsGo2020: distribution, 69
Lamb1974: distribution, host, 42
Lawson1917: distribution, host, 206
LazaroGoRo2023: diagnosis, distribution, host, key, natural enemies, taxonomy, 248, 259-260
Leonar1918: description, distribution, taxonomy, 194
Leonar1920: description, distribution, host, illustration, taxonomy, 213-215
Lepage1938: catalog, distribution, host, 262, 408
LincanHoCa2010: distribution, host, 5
Lindin1909b: taxonomy, 223
Lindin1911: taxonomy, 177
Lindin1935: taxonomy, 148
Lobdel1937: structure, 78
LongoMaPe1995: distribution, 127
LongoMaPe1999a: distribution, 149
Lupo1938a: description, distribution, host, taxonomy, 256, 261
MacGil1921: distribution, host, taxonomy, 363
MacGow1982: distribution, host, 10
Malump2012b: distribution, 213
Malump2014a: distribution, host, 75
Mamet1943a: distribution, host, 161
Mamet1948: distribution, host, 12, 13
Mamet1949: catalog, distribution, host, taxonomy, 38
Mamet1956b: distribution, host, 305
Mamet1959a: distribution, host, 380
Marlat1921a: distribution, 11
MartinWoCu2022: distribution, host, 5
Martor1976: distribution, host, 43, 56, 84, 168, 253
Maskel1895b: distribution, host, 49
Maxwel1902: distribution, host, 250-251, 299
Maxwel1903: description, distribution, host, illustration, 48
McDani1924: distribution, host, 41-42
McDani1931: distribution, host, 61
McKenz1956: description, distribution, host, illustration, taxonomy, 32, 115
Medler1980: distribution, 89
Merril1953: distribution, host, 50
MerrilCh1923: distribution, host, 237
MestreHaEv2011: distribution, host, 12
MestreHoHa2015: distribution, host, 11-12
Meurge2011: distribution, 81
MilleHeCa2016: distribution, economic importance, host, 140
Miller2005: distribution, 487
MillerDa1990: economic importance, taxonomy, 302
MillerDa2005: description, distribution, economic importance, host, 234
MiskimBo1970: distribution, host, 30
Morgan1889a: taxonomy, 349
Morgan1892: description, distribution, host, taxonomy, 15, 16
Morley1909: biological control, 256
MorseNo2006: phylogeny, taxonomy, 340
Mosque1976: distribution, host, 41, 89
MoutiaMa1947: distribution, host, 10
MunroFo1936: catalog, distribution, host, 87
Muraka1970: distribution, host, 80
Nakaha1981a: distribution, 399
Nakaha1982: distribution, host, 44
NakahaMi1981: distribution, 34
Newste1901b: distribution, host, 180, 190
Newste1907a: distribution, 10
Newste1914: distribution, host, 311
Nishid2002: catalog, 141
NormarJo2010: ecology, host, 3
NormarOkMo2019: distribution, host, phylogeny, taxonomy, 23, 55, S6
OConnoGeMa2013: distribution, host, 39
Pace1939: host, 665
Paoli1915: distribution, host, 256
Parkin1906: distribution, host, 33, 56, 66, 69
PellizGe2010a: distribution, host, 502
PerezG2008: distribution, 215
PetersHaMo2020s1: DNA, distribution, host,
PooleGe1997: distribution, 349
PruthiMa1945: distribution, host, 8
Ramakr1919a: distribution, host, 10
Ramakr1921a: catalog, distribution, host, 353
Ramakr1930: distribution, host, 15
RaoCh1950: taxonomy, 23, 27
Ronna1933: distribution, host, 51-53
Ruhl1930: biological control, taxonomy, 20
Russo1927: distribution, taxonomy, 48, 203, 205
Ryan1946: distribution, economic importance, 124
Ryan1946: distribution, 124
Sander1904a: distribution, host, 51
Sankar1984: biological control, distribution, host, 27
Sassce1920: taxonomy, 182
Schmut1952: taxonomy, 580
Schmut1959: distribution, taxonomy, 204
ShiLi1991: host, 164
SilvadGoGa1968: catalog, distribution, host, 174
Simmon1957: biological control, catalog, distribution, host, 4
Singh1964: distribution, host, 219
South1910: distribution, 20
Suh2016: distribution, host, key, 316, 327
SuhJi2009: distribution, illustration, taxonomy, 1039-1054
Takagi1970: description, distribution, host, taxonomy, 32
Takagi1992b: description, distribution, host, illustration, taxonomy, 44-45, 55, 67, 72
Takaha1942d: distribution, host, 355
Tao1999: distribution, host, 91-92
Timber1924: biological control, distribution, 439
Trimbl1928: distribution, host, 45
Varshn2002: distribution, host, 47
Vayssi1955: distribution, 257-267
WalkerWaEv1974: distribution, host, 23
Wang1980: distribution, host, 175
Wang1982c: distribution, host, 74-76
Watson2001: list, 177
Watson2002: taxonomy, 117
Watson2002a: biological control, description, distribution, economic importance, host, illustration, taxonomy,
Watson2021: distribution, 523
WatsonBe1937: host, 15
WatsonKo2022a: biology, control, diagnosis, dispersal, distribution, economic importance, host, taxonomy, 489-491
WattMa1903: distribution, host, 309
WebsteBu1902: distribution, host, 113
Willia1960c: taxonomy, 393
Willia1973: distribution, host, 91
Willia1973: distribution, host, 91
Willia1977ML: distribution, host, 94
Willia2017a: catalog, list of species, 219
WilliaBu1987: distribution, host, 95
WilliaWa1988: description, distribution, host, illustration, taxonomy, 137-138, 140
WilliaWi1988: distribution, host, taxonomy, 67
Wilson1917: distribution, host, 40
Wilson1921: taxonomy, 24
Wolcot1936: catalog, distribution, taxonomy, 134
WongChCh1999: distribution, illustration, 26, 67
Yang1982: distribution, host, 264, 265
Zahrad1953: distribution, host, 143
Zahrad1990c: distribution, host, 16
ZarkanApTu2021: distribution, host, 165
Zimmer1948: distribution, host, 410