(Linnaeus, 1758)
Taxonomic note
Lepidosaphes ulmi was originally described in the genus Coccus. It has since been described as different species on different hosts, resulting in numerous synonyms. Biparental and parthenogenetic strains of L. ulmi occur; in addition, there are populations with different host preferences (Kozár, 1990b), which suggests that this may be a species complex. Thiem, 1933, separated the two forms into Lepidosaphes ulmi ulmi Linnaeus and L. ulmi bisexualis Thiem (Zahradník, 1990). These forms have been recognized as valid by many workers. However, there is no morphological evidence that there is more than one species present (Gill, 1997) and they are all treated together here. Further study is required.
Diagnosis
In life, scale cover of adult female 1.0-3.5 mm long, convex, mussel-shaped (curved or straight, depending on the texture of the host and population density of the scales), strongly tapered towards the exuvial end, silvery grey or light brown initially but becoming purplish to coppery brown with age LEULL3.jpg , with yellow-brown exuviae at the narrow end. Occasionally scale covers appear banded (Johnson and Lyon, 1976) LEULL2.jpg . Ventral scale thick, white. Female scales may adhere to branches for several years and become quite bleached (Gill, 1997). Scale cover of male, if present, light brown, smaller, slenderer and more parallel-sided than that of female, with yellow terminal exuviae. Body of female in life white to yellowish, with yellowish-brown pygidium (Zahradník, 1990); lateral lobes distinct. Adult male winged (Ghauri, 1962).
Body of slide-mounted adult female elongate, more than 1.8x as long as wide, and membranous; head rounded; eye not developed into a spur; sharply pointed lateral marginal spurs present on margins of prepygidial segments II, III and IV, each spur moderately sclerotized, not containing any duct openings LEPULS.jpg . Pygidium with median lobes with a pair of gland spines between them, not yoked, and without club-shaped basal scleroses; perivulvar pores present. Six large, marginal macroducts present on each side of pygidium; small dorsal pygidial macroducts numerous, all similar in size; abdominal segments V and VI each with a small dorsal, sclerotized submarginal boss on either side LEPULP.jpg .
The first instar nymph was described and illustrated by Howell and Tippins, 1977.
Host range
Lepidosaphes ulmi is a highly polyphagous, temperate species. It is known to infest over 130 host plants in the USA (Kosztarab, 1996), representing 85 genera in 33 families (Davidson and Miller, 1990). Rosaceae and Leguminosae are favoured hosts, although in California it prefers willows (Gill, 1997). In northern Europe it is important on apple, whereas around 49°-50° latitude it is important on plum; further south again, L. ulmi is very polyphagous and may represent a different race or even species (Kozár, 1990b). The parthenogenetic and bisexual forms have different host preferences with the former found on hosts of the Rosaceae, and the latter found on a variety of hosts including species of Acer, Betula, Buxus, Fagus, Fraxinus, Liriodendron, Malus, Populus, Prunus, Pyrus, Salix, Syringa and Ulmus. Recorded hosts of L. ulmi sensu lato include species of: Abies, Acer, Aesculus, Ailanthus, Amelanchier, Andromeda, Angelonia, Arbutus, Arctostaphylos, Berberis, Betula, Buxus, Caesalpinia, Cajanus, Calluna, Carpinus, Castanea, Ceanothus, Ceratonia, Chimaphila, Clematis, Cornus, Corylus, Cotinus, Cotoneaster, Crataegus, Cyanotis, Cydonia, Cytisus, Elaeagnus, Erica, Euonymus, Euphorbia, Fagus, Fraxinus, Ficus carica, Genista, Geranium, Ginkgo, Hippophae, Hypericum, Juglans spp., Laburnum, Ligustrum, Lonicera, Lycium, Maackia, Malus spp., Melica, Mespilus, Myrica, Olea europaea, Pachysandra, Philadelphus, Platanus, Populus spp., Potentilla, Prunus spp., Ptelea, Pyrus communis, Quercus, Rhamnus, Rhododendron, Ribes, Ricinus, Robinia, Rosa, Rubus, Salix, Sapium, Sassafras, Sesbania, Sorbaria, Sorbus, Spartium, Spiraea, Staphylea, Stillingia, Symphoricarpos, Syringa, Tilia, Ulex, Ulmus, Vaccinium myrtillus, Viburnum, Vincetoxicum, Viscum, Vitis vinifera and Ziziphus.
Affected plant stages: vegetative growing, flowering, fruiting and post-harvest stages
Affected plant parts: on aerial parts, especially on the bark of trunk and branches LEULL1.jpg , and on fruit (less often on leaves)
Biology and ecology
Depending on the strain of L. ulmi, development can be either univoltine or bivoltine; it is the parthenogenetic form that is found on apples. The parthenogenetic "apple strain" is oviparous and potentially multivoltine. The bisexual "poplar strain" is viviparous (Gharib, 1978).
Bisexual L. ulmi overwinters in the egg stage and has one or two slightly overlapping generations annually, depending upon climate (Carillo et al., 1995; Gill, 1997). Those populations in more northerly regions have one generation per year, while two generations occur in more southern areas (Johnson and Lyon, 1976; Kozár, 1990b). Egg production varies according to host plant, with each female producing 10-120 eggs (Bazarov and Smelev, 1971). The larvae require an effective temperature sum exceeding 130.8°C to hatch (Savkovskij, 1969).
Lepidosaphes ulmi is parthenogenetic in California, although a biparental strain occurs on the East coast of North America. In Europe, around the Mediterranean and in Chile, both strains occur (Gharib, 1978; Carillo et al., 1995; Kozár, 1990b). Wang et al., 2000, recorded parthenogenetic reproduction of L. ulmi in China (Zhejiang), where there are two generations per year. Each female lays about 40 white eggs (Kozár, 1990b; Zahradník, 1990). Mussey and Potter, 1997, found that overwintering eggs began to hatch at 761 (687-834) degree days in Kentucky, USA. The eggs laid on apple were found to contain primitive embryos that develop when conditions become favourable; the eggs laid by bisexual females on poplar contain embryos in a more advanced state (Gharib and Bénassy, 1983). Development from first-instar nymph to adult ranges from 51 to 58 days.
Lepidosaphes ulmi is most often found on the trunk, limbs and fruit (Merrill, 1953; Murakami, 1970), and occasionally on the leaves of the host plant. The females may develop in a straight pattern or become twisted or curved to follow the surface structure of the bark. Males tend to be more numerous on the leaves. The first instar crawlers are the dispersal stage, and are mainly spread by wind and animal agencies and, over longer distances, by human transport of infested plant material.
Symptoms
Infestation by L. ulmi causes leaf discolouration and premature leaf fall; encrusted branches may die back; infested fruit become unsightly, spotted and discoloured and may drop prematurely. This scale insect can kill twigs, branches and young trees, and if left unchecked will seriously weaken and stunt mature plants (Gill, 1997). Shoots and leaf petioles may be deformed as a result of infestation by L. ulmi (Kosztarab, 1990).
Hegde et al., (1977) reported that a mycoplasma-like organism [phytoplasma] was transmitted by L. ulmi. This organism was found to produce little-leaf symptoms on Angelonia grandiflora grown in India.
Economic impact
Lepidosaphes ulmi is a very destructive species, causing dieback and death of ornamental trees (Kosztarab, 1996). Infestations may cause leaf yellowing, fruit deformity, leaf drop and dieback of branches. If left uncontrolled, heavy infestations can weaken or stunt plants. In addition to reducing plant vigour (Groot, 1967), heavy infestations reduce plant growth and lower frost resistance, endangering trees and possibly leading to death in 2-3 years. Helsen et al., 1996, noted that even minor infestations of fruit may cause major economic losses as a result of the zero tolerance policies for export produce.
Lepidosaphes ulmi was listed as a pest of deciduous fruit trees of world importance by Kozár, 1990b. It is often found infesting apples, and heavy populations may seriously damage fruit. It was one of the most frequently encountered scale insect pests infesting fruit trees in European orchards (Kozár et al., 1979; Kozár and Konstantinova, 1981) and occasionally becomes an important pest of forest trees and ornamental trees, especially under urban conditions, and in neglected orchards (Kosztarab and Kozár, 1988). It was regarded as one of the most important pests of apple in the northern USSR (Konstantinova, 1976). Yanin, 1975, listed it as one of the more injurious species in the Caucasus. The species is also an occasional pest of fruits and nuts, e.g. on walnuts in California (Ebeling, 1959). The greatest damage is caused on apples in northern Europe and plums further south (Kozár, 1990b), and on poplars and willows (Zahradník, 1990). Lepidosaphes ulmi was reported to be the most important pest of apples in Turkey (Kozár et al., 1979) and northern Russia (Konstantinova, 1976; Korchagin, 1987) and on bilberry (Vaccinium myrtillus) in Poland (Koteja, 1983). It has also been a problem on olives in Greece where it, along with Parlatoria oleae, causes serious injury to the tree and fruit (Argyriou and Kourmadas, 1981); it has also caused serious damage to olives in Spain (Argyriou, 1990). In the USA, this species has caused considerable losses of ornamental trees in the East and Midwest, but it is not usually a serious pest in California (Gill, 1997). Charles and Henderson, submitted, recorded it as a pest on economically important plants in New Zealand. In Chile it is abundant on abandoned fruit trees (Claps et al., 2001a) so it presumably has the potential to reach high populations in cultivated orchards. Danzig and Pellizzari, 1998, described L. ulmi as a dangerous pest in the Palaearctic, and Foldi, 2001, listed it as an occasional pest in France. In China (Zhejiang) it is an important pest of Arbutus, attacking two-year-old branches (Wang et al., 2000).
In California, L. ulmi is being considered for use as a biological control agent against brush species (e.g. Ceanothus) that compete with young conifers for light and nutrients on reafforestation programmes; this might reduce the need for herbicides (Gill, 1997).
Detection and inspection methods
Lepidosaphes ulmi may be found on all parts of the plant with the exception of the roots. Examine potential host plants for grey to brown mussel-shaped scale covers on trunks, branches, leaves, or fruit.
Phytosanitary protection
Lepidosaphes ulmi is mentioned on quarantine lists (Burger and Ulenberg, 1990).
Natural enemies
Some 41 species of natural enemies have been recorded to be associated with L. ulmi. The species listed below are from Kosztarab and Kozár, 1988, and CABI, 2000.
Parasitoids:
- Adelencyrtus mayrai, attacking: nymphs, adults, in Canada
- Anabrolepis zetterstedtii
- Anaphes gracilis
- Aphytis abnormis
- Aphytis aonidiae, attacking: nymphs, adults, in Bulgaria
- Aphytis caucasicus
- Aphytis diaspidis, in USA (Ohio)
- Aphytis moldavicus
- Aphytis mytilaspidis, attacking: nymphs, adults, in Europe, Bulgaria, Canada, USA (Ohio)
- Aphytis proclia, attacking: nymphs, adults, in Bulgaria, Canada
- Apteroptrix longiclava
- Coccobius testaceus
- Coccophagus niger
- Coccophagus parvipennis
- Coccophagus similis
- Encarsia citrina, in USA (Ohio)
- Epitetracnemus zetterstedtii, attacking: nymphs, adults, in Russia
- Eusemion cornigerum
- Gyranusa matritensis
- Habrolepis dalmanni
- Microterys macharus
- Pteroptrix lauri
- Pteroptrix maritimus
- Pteroptrix longicornis
- Pteroptrix opaca
- Zaomma lambinus, in USA (Ohio)
Predators:
- Chilocorus bipustulatus, attacking: nymphs, adults, in Europe, Bulgaria, Russia
- Chilocorus renipustulatus, attacking: nymphs, adults, in Europe, Bulgaria, Russia
- Chilocorus stigma, attacking: nymphs, adults, in Georgia, Russia, USA (Ohio)
- Dometorina sp., in USA (Ohio)
- Haplothrips kurdjumovi, in Europe, North America, New Zealand
- Hemisarcoptes malus, in Russia, Bulgaria, Canada
- Karnyothrips flavipes
- Octobdellodes sp., in USA (Ohio)
- Oripoda elongata, in USA (Ohio)
- Scapheremaeus marginalis, in USA (Ohio)
- Thyreophagus entomophagus, in USA (Ohio)
Distribution
See Lepidosaphes ulmi distribution.
Microscopic examination of slide-mounted adult females is required for authoritative identification to species.
Lepidosaphes camelliae Hoke (camellia scale) LECAMS.jpg could be misidentified as L. ulmi, but differs in having no sclerotized spurs on the margins of abdominal segments II-IV; fewer than 10 submarginal and submedian ducts on each side of segment VI; and the marginal and submarginal ducts are almost the same size LACAMP.jpg. In contrast, L. ulmi LEPULS.jpg possesses sclerotized spurs on the margins of abdominal segments II-IV; more than 15 submarginal and submedian ducts on each side of segment VI; and the submarginal ducts are much smaller than the marginal ducts LEPULP.jpg. Lepidosaphes camelliae is known from USA (Alabama, Arkansas, Connecticut, Delaware, Florida, Georgia, Louisiana, Massachusetts, Maryland, Missouri, Mississippi, North Carolina, New York, Oklahoma, Oregon, South Carolina, Texas and Virginia), Mexico, China (Guangdong, Hunnan) and Japan (Tao, 1999; Nakahara, 1982; Kawai, 1980) on leaf undersides of species of Camellia, Cleyera, Cupressus, Ilex, Ligustrum, Magnolia, Rhaphiolepis and Ternstroemia (Tao, 1999; Danzig and Pellizzari, 1998; Kosztarab, 1996; Miller, 1996; Nakahara, 1982). Lepidosaphes camelliae is occasionally a serious pest on camellia and holly in Florida (Dekle, 1976). Heavy infestations cause yellow and orange leaf discolouration and premature drop (Johnson and Lyon, 1991). This species has 4-5 generations per year in California, where it is considered to be a 'B'-rated pest on camellias, holly, cuttings and small plants (Gill, 1997). Scale cover of adult female mussel-shaped, slightly convex, light or dark brown, with yellow terminal exuviae; scale cover of male similar to that of female but smaller and narrower, with green-gold terminal exuviae (Davidson and Miller, 1990). Female scale cover LECAML1.jpg ; male and female scale covers LECAML2.jpg ; female scaleLECAML3.jpg ; habit sketch LECAML.jpg
Lepidosaphes conchiformis (Gmelin) (red oystershell scale) LECONS.jpg is a species not featured in the picture key, which has some similarities with L. ulmi. It lacks sclerotized marginal spurs between the lateral lobes of the free abdominal segments that are present in L. ulmi LEPULS.jpg. Lepidosaphes conchiformis is a polyphagous species known from Belgium, Cyprus, former Czechoslovakia, France (including Corsica), Germany, Greece, Hungary, Italy, Sardinia, Sicily, Poland, Romania, Spain (mainland and Balearic Is), Switzerland, Turkey, former USSR (South European Territory, Ukraine, Transcaucasus, Middle Asia), Algeria, Egypt, Iran, Iraq, Israel, India (Assam, Tamil Nadu), Pakistan, China, Japan, North America (including California) and South America, on leaves and branches of woody hosts including species of Acer, Betula, Carpinus, Celtis, Corylus, Crataegus, Diospyros, Fagus, Ficus, Fraxinus, Juglans, Malus, Olea, Pistacia, Prunus, Pyrus, Syringa, Tilia, Ulmus and Zelkova (Amparo Blay Golcoechea, 1993; Kozár et al., 1994; Longo et al., 1995; Gill, 1997; Danzig and Pellizzari, 1998; Foldi, 2001; Abivardi, 2001). Fig is the favoured host, and heavy infestations reduce the quality of the fruit due to discolouration and distortion (Gill, 1997). Lepidosaphes conchiformis is commonest on the periphery of forests, in towns (often in alleys and parks) (Zahradník, 1990a). Scale covers mussel-shaped; those formed on the leaves tend to be pale and sigmoid, whereas those formed on bark are brown and straight. Reproduction is sexual and there is one generation per year; each female produces 5-8 eggs in Poland (Komosinska, 1975), but an average of 26 each in Germany (Schmutterer, 1959). Habit sketch LECONL.jpg ; appearance in life LECONCL1.jpg , LECONCL2.jpg , LECONL0.jpg ; adult female scale cover LECONL2.jpg ; adult female scale covers LECONL3.jpg
Foldi, 2001, lists this species as an economically important pest in France. Abivardi, 2001, discusses the morphology, biology and natural enemies of L. conchaspiformis. This species has morphologically distinct leaf- and twig forms; the twig form has a dark brown female scale cover, whereas the in leaf form the scale cover is white; the exuviae are terminal. In both forms the male scale cover is elongate, light tan or white with terminal exuviae. In California there are 2-3 generations per year, and the insect overwinters as the twig form adult female (Gill, 1997). In Europe there are 1 or 2 generations per year, and the overwintering female (whose body is red-violet) lays about 25 eggs in spring (Zahradník, 1990a).
Lepidosaphes conchiformis LECONP.jpg can be distinguished from L. camelliae (above) because it lacks small macroducts anterior to the second lobe that are present in L. camelliae LACAMP.jpg.
Comments
Lepidosaphes ulmi is probably native to temperate Eurasia, but is now a cosmopolitan species (Gill, 1997). It is generally found distributed throughout the temperate regions, but has not been recorded from Africa south of the Sahara, or from the tropical Pacific islands.
Europe
Austria: present, no further details (Danzig and Pellizzari, 1998)
Belgium: restricted distribution (Busschère et al., 1970)
Bulgaria: widespread (Dirimanov et al., 1984; Danzig and Pellizzari, 1998)
Croatia: restricted distribution (Kozarzevskaja and Vlainic, 1981)
Cyprus: present, no further details (Nakahara, 1982)
Denmark: present, no further details (Gertsson, 2001)
Finland: present, no further details (Gertsson, 2001)
Former Czechoslovakia: present, no further details (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Former USSR
Eastern Siberia: present, no further details (Danzig and Pellizzari, 1998)
Armenia: present, no further details (Babayan and Oganesyan, 1979)
Caucasus: present, no further details (Yanin, 1975)
Central European Territory: present, no further details (Danzig and Pellizzari, 1998)
Central Russia: present, no further details (Kulikova, 1987)
Crimea: present, no further details (CIE, 1958)
Far East: present, no further details (Danzig and Pellizzari, 1998)
Georgia, Republic of: widespread (Aleksidze, 1995; Yasnosh, 1995)
Latvia: present, no further details (CIE, 1958)
Lithuania: present, no further details (CIE, 1958)
Middle Asia: present, no further details (Danzig and Pellizzari, 1998)
Moldavia: present, no further details (CIE, 1958)
Kazakhstan: present, no further details (Matesova, 1980; Danzig and Pellizzari, 1998)
Russian Federation: widespread (Borisoglebskaya, 1979; Konashevich, 1980)
Southern European Territory: present, no further details (Danzig and Pellizzari, 1998)
Transcaucasus: present, no further details (Danzig and Pellizzari, 1998)
Turkmenistan: present, no further details (CIE, 1958)
Ukraine: present, no further details (CIE, 1958)
Uzbekistan: present, no further details (CIE, 1958)
Former Yugoslavia: restricted distribution (Kozarzevskaja and Vlainic, 1982; Velimirovic, 1990)
France: present, no further details (Gharib and Bénassy, 1983; Danzig and Pellizzari, 1998; Foldi, 2001)
Germany: restricted distribution (Witte and Lein, 1992; Danzig and Pellizzari, 1998)
Greece: restricted distribution (Kozár and Konstantinova, 1981; Kosztarab and Kozár, 1988)
Hungary: restricted distribution (Kozár and Drozdjak, 1988; Danzig and Pellizzari, 1998)
Isle of Man: The Natural History Museum collection, London, UK
Italy: present, no further details (Longo et al., 1995; Danzig and Pellizzari, 1998)
Sardinia: present, no further details (Longo et al., 1995)
Sicily: present, no further details (Longo et al., 1995)
Luxemburg: The Natural History Museum collection, London, UK
Macedonia: present, no further details (Merrill, 1953; Soulis, 1960)
Malta: present, no further details (CIE, 1958)
Netherlands: restricted distribution (Dijke and Frankenhuyzen, 1985; Helsen et al., 1996)
Norway: present, no further details (Kozár and Konstantinova, 1981; Gertsson, 2001)
Poland: restricted distribution (Komosinska, 1986; Danzig and Pellizzari, 1998)
Portugal
Madeira: present, no further details (Nakahara, 1982)
Romania: present, no further details (Kosztarab and Kozár, 1988; Danzig and Pellizzari, 1998)
Spain: widespread (Amparo Blay Golcoechea, 1993)
Balearic Is: present (Amparo Blay Golcoechea, 1993)
Canary Is: restricted distribution (Torres et al., 1992)
Sweden: quite widespread (Gertsson, 2001)
Switzerland: present, no further details (Kozár et al., 1994; Kozár and Hippe, 1996)
United Kingdom: widespread (Easterbrook et al., 1985)
England: present, no further details (Tao, 1999)
Channel Is (Guernsey): The Natural History Museum collection, London, UK
Northern Ireland: The Natural History Museum collection, London, UK
Asia
Afghanistan: restricted distribution (Fowjhan and Kozár, 1994)
China: restricted distribution (Tang, 1984)
Shanxi: present, no further details (Xie, 1982)
Yunnan: present, no further details (CIE, 1958)
Zhejiang: present, no further details (Wang et al., 2000)
India: present, no further details (Hegde et al., 1977)
Iran: present, no further details (Seghatoleslami, 1977; Babayan and Oganesyan, 1979)
Iraq: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Israel: restricted distribution (Mendel et al., 1992; Danzig and Pellizzari, 1998)
Japan: present (Murakami, 1970; Kawai, 1980; Tao, 1999)
Hokkaido: present, no further details (CIE, 1958)
Honshu: present, no further details (CIE, 1958)
Korea: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Saudi Arabia: present, no further details (Nakahara, 1982; Danzig and Pellizzari, 1998)
Syria: present, no further details (Nakahara, 1982)
Taiwan: present, no further details (Nakahara, 1982)
Turkey: restricted distribution (Acatay, 1970)
Africa
Algeria: present, no further details (Merrill, 1953; CIE, 1958)
Egypt: restricted distribution (El-Minshawy et al., 1974; Danzig and Pellizzari, 1998)
Morocco: present, no further details (CIE, 1958)
Tunisia: present, no further details (CIE, 1958)
Western Hemisphere
Argentina: uncommon (Claps et al., 2001a)
Rio Negro: present, no further details (Claps et al., 2001a)
Brazil: present, no further details (Nakahara, 1982)
Rio de Janeiro: present, no further details (Claps et al., 2001a)
Rio Grande do Sul: present, no further details (Claps et al., 2001a)
Sao Paulo: present, no further details (Claps et al., 2001a)
Canada: restricted distribution (Karsemeijer, 1973; Nakahara, 1982)
Alberta: present, no further details (CIE, 1958)
British Columbia: present, no further details (Kosztarab, 1996)
Manitoba: present, no further details (CIE, 1958)
New Brunswick: present, no further details (Kosztarab, 1996)
Nova Scotia: restricted distribution (Macphee and Maclellan, 1971; Kosztarab, 1996)
Ontario: present, no further details (Kosztarab, 1996)
Prince Edward I.: present, no further details (Kosztarab, 1996)
Quebec: restricted distribution (Parent, 1970; Parent, 1973)
Chile: abundant on abandoned fruit trees (Claps et al., 2001a)
Antofagasta: present, no further details (Claps et al., 2001a)
Atacama: present, no further details (Claps et al., 2001a)
Biobío: present, no further details (Claps et al., 2001a)
Coquimbo: present, no further details (Claps et al., 2001a)
La Araucanía: present, no further details (Claps et al., 2001a)
Los Lagos: present, no further details (Claps et al., 2001a)
Maule: present, no further details (Claps et al., 2001a)
O'Higgins: present, no further details (Claps et al., 2001a)
Santiago: present, no further details (Claps et al., 2001a)
Tarapacá: present, no further details (Claps et al., 2001a)
Valparaiso: present, no further details (Claps et al., 2001a)
Colombia: present, no further details (Nakahara, 1982)
Mexico: present, no further details (Nakahara, 1982; Miller, 1996)
Peru: present, no further details (Nakahara, 1982)
Uruguay: present, no further details (Nakahara, 1982)
USA: most states in the continental USA (Nakahara, 1982; Gill, 1997)
California: widespread and common (Gill, 1997)
Colorado: present, no further details (CIE, 1958)
Connecticut: present, no further details (CIE, 1958)
Delaware: present, no further details (CIE, 1958)
Florida: present, no further details (Merrill, 1953)
Idaho: present, no further details (CIE, 1958)
Illinios: present, no further details (CIE, 1958)
Indiana: present, no further details (Dietz and Morrison, 1916; CIE, 1958)
Iowa: present, no further details (CIE, 1958)
Kentucky: present, no further details (Mussey and Potter, 1997)
Maine: present, no further details (CIE, 1958)
Maryland: restricted distribution (Garrett, 1973)
Massachusetts: present, no further details (King, 1901; CIE, 1958)
Michigan: present, no further details (McDaniel, 1939; CIE, 1958)
Minnesota: present, no further details (CIE, 1958)
Mississippi: present, no further details (Herrick, 1911)
Missouri: present, no further details (Hollinger, 1923; CIE, 1958)
Montana: present, no further details (CIE, 1958)
Nebraska: present, no further details (CIE, 1958)
New Hampshire: present, no further details (CIE, 1958)
New Jersey: present, no further details (CIE, 1958)
New York: restricted distribution (Groot, 1967)
Ohio: present, no further details (CIE, 1958)
Oregon: present, no further details (CIE, 1958)
North Carolina: present, no further details (CIE, 1958)
North Dakota: present, no further details (CIE, 1958)
Pennsylvania: restricted distribution (Hart and Byrnes, 1960)
Rhode I.: present, no further details (CIE, 1958)
South Dakota: present, no further details (CIE, 1958)
Tennessee: present, no further details (Lambdin and Watson, 1980)
Utah: present, no further details (CIE, 1958)
Vermont: present, no further details (CIE, 1958)
Virginia: present, no further details (Kosztarab, 1996)
Washington: present, no further details (CIE, 1958)
West Virginia: present, no further details (CIE, 1958)
Wisconsin: present, no further details (CIE, 1958)
Wyoming: present, no further details (CIE, 1958)
Oceania
Australia
New South Wales: present, no further details (CIE, 1958; CSIRO, 2001)
South Australia: present, no further details (CIE, 1958; CSIRO, 2001)
Tasmania: present, no further details (Hudson, 1967; CSIRO, 2001)
Victoria: present, no further details (CIE, 1958; CSIRO, 2001)
Western Australia: present, no further details (CIE, 1958; CSIRO, 2001)
New Zealand: present, no further details (Nakahara, 1982; Charles and Henderson, submitted)