Diaspidiotus perniciosus (San José scale)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Hosts/Species Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Diaspidiotus perniciosus (Comstock) Cockerell, 1899; Danzig, 1993
Preferred Common Name
- San José scale
Other Scientific Names
- Aonidia fusca Maskell, 1895
- Aonidiella fusca (Maskell) Berlese & Leonardi, 1898
- Aonidiella perniciosa (Comstock) Balachowsky & Mesnil, 1935
- Aspidiotus (Comstockaspis) perniciosus (Comstock) Borschenius, 1935
- Aspidiotus (Diaspidiotus) andromelas Cockerell, 1897
- Aspidiotus (Diaspidiotus) perniciosus (Comstock) Brain, 1918
- Aspidiotus (Hemiberlesiana) perniciosus (Comstock) Thiem & Gerneck, 1934
- Aspidiotus (Quadraspidiotus) perniciosus (Comstock) Merril, 1953
- Aspidiotus albopunctatus Cockerell, 1896
- Aspidiotus fuscus (Maskell) Ferris, 1941
- Aspidiotus perniciosus Comstock, 1881
- Comstockaspis perniciosa (Comstock) MacGillivray, 1921
- Hemiberlesia perniciosa (Comstock) Lindinger, 1957
- Quadraspidiotus (Aspidiotus) perniciosus (Comstock) Rahman & Ansari, 1941
- Quadraspidiotus perniciosus (Comstock) Ferris, 1938
International Common Names
- English: California scale (Russia); Chinese scale; perniciosus scale
- Russian: Kaliforniyskaya schitova
- Portuguese: piolho de San José
Local Common Names
- Denmark: san José skjoldlus
- Finland: hirmukilpikirva
- France: cochenille de San José; pou de San José
- Germany: Kalifornische Schildlaus; San José Schildlaus
- Hungary: Kaliforniai pajzstetü
- Italy: cocciniglia di San José; pidochio di San José
- Japan: Sanhoze kaigaramusi
- Netherlands: San José schildluis
- Norway: san-José skjoldlus
- Spain: cochinilla de San José; escama de San José; piojo de San José
- Sweden: san José-sköldlus
- Turkey: san Jose kabuklu biti
- QUADPE (Quadraspidiotus perniciosus)
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Hemiptera
- Suborder: Sternorrhyncha
- Unknown: Coccoidea
- Family: Diaspididae
- Genus: Diaspidiotus
- Species: Diaspidiotus perniciosus
Notes on Taxonomy and NomenclatureTop of page Taxonomists disagree as to the generic placement of D. perniciosus. Danzig (1993) put it in Diaspidiotus, whereas Kosztarab (1996) left it in the widely used Quadraspidiotus genus. For further information see Bazarov and Shmelev (1971), Borchsenius (1966) and Kosztarab and Kozar (1988).
DescriptionTop of page The testa of the female is almost circular, slightly convex, light to dark grey, diameter ca 1.5-2.2 mm. Exuviae central or subcentral, the first-instar exuviae has a craterlike appearance. Young adult female pear-shaped, yellow; egg-laying females almost circular. Pygidium with two pairs of well-developed lobes, notched on outer margin. Plates between lobes partly pointed or fimbriated. Perivulvar pores absent (Kosztarab and Kozar, 1988).
The adult male was described by Bustshik (1958) and Gauri (1962). Morphometric analyses and a key for Quadraspidiotus (=Diaspidiotus) males are given by Kozar et al. (1996). The male is similar to the male of D. ostreaeformis, but the third segment of the antennae is shorter and elliptical in shape. For a long time there was no possibility of identifying the male and female in the first-instar stages. Later, Stoetzel and Davidson (1974) found some microscopic characters for separating the different sexes, and supposed these to be general for all members of the Aspidiotini. However, Kozar et al. (1996) found that first-instar female larva of D. perniciosus differs from other species in at least one character (an extra pair of dorsal setae on the second segment).
DistributionTop of page D. perniciosus originated from Eastern Asia and was accidentally introduced to the USA, and subsequently to other parts of the world including Argentina, Australia and New Zealand. It is widely distributed in the Palearctic and Nearctic regions (Kozar, 1990a). There are many publications about its detailed distribution and importance in different parts of the world (Konstantinova, 1976; Baker, 1977; Chowdhuri, 1977; Kozar and Konstantinova, 1981; Anon., 1986; APPPC, 1987; Kozar and Drozdjak, 1988; Davidson and Miller, 1990; Kozar et al., 1994; Mani et al., 1995).
The distribution map includes records based on specimens of D. perniciosus from the collection in the Natural History Museum (London, UK): dates of collection are noted in the List of Countries (NHM, various dates).
See also CABI/EPPO (1998, No. 131).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Afghanistan||Present||Mohammad, 1988; EPPO, 2014|
|Azerbaijan||Present||Danzig, 1993; EPPO, 2014|
|Brunei Darussalam||Present||Kosztarab and Kozár, 1988|
|-Hong Kong||Present, few occurrences||EPPO, 2014|
|-Nei Menggu||Present||EPPO, 2014|
|Georgia (Republic of)||Present||Borchsenius, 1966; EPPO, 2014|
|-Andhra Pradesh||Present||EPPO, 2014|
|-Himachal Pradesh||Present||EPPO, 2014|
|-Indian Punjab||Present||NHM, 1983; Sharma, 2011; EPPO, 2014|
|-Jammu and Kashmir||Present||EPPO, 2014|
|-Tamil Nadu||Present||EPPO, 2014|
|-Uttar Pradesh||Present||EPPO, 2014|
|-Uttarakhand||Present||Singh et al., 2001|
|-West Bengal||Present||EPPO, 2014|
|Iran||Restricted distribution||Kosztarab and Kozár, 1988; EPPO, 2014|
|Iraq||Present||Kosztarab and Kozár, 1988; EPPO, 2014|
|Jordan||Absent, unreliable record||EPPO, 2014|
|Kazakhstan||Absent, unreliable record||Danzig, 1993; EPPO, 2014|
|Korea, DPR||Present||Danzig, 1993; EPPO, 2014|
|Korea, Republic of||Present||Danzig, 1993; EPPO, 2014|
|Nepal||Present||Nakahara, 1982; EPPO, 2014|
|Pakistan||Present||Danzig, 1993; EPPO, 2014|
|Taiwan||Absent, formerly present||EPPO, 2014|
|Tajikistan||Restricted distribution||EPPO, 2014|
|Turkey||Restricted distribution||****||Kosztarab and Kozár, 1988; EPPO, 2014|
|Uzbekistan||Restricted distribution||Danzig, 1993; EPPO, 2014|
|Algeria||Widespread||Kosztarab and Kozár, 1988; EPPO, 2014|
|Congo Democratic Republic||Present||EPPO, 2014|
|Morocco||Restricted distribution||EPPO, 2014|
|South Africa||Present||Danzig, 1993; EPPO, 2014|
|-Canary Islands||Restricted distribution||Kosztarab and Kozár, 1988; EPPO, 2014|
|Zimbabwe||Restricted distribution||EPPO, 2014|
|Canada||Restricted distribution||Borchsenius, 1966; EPPO, 2014|
|-British Columbia||Restricted distribution||Agr. Canada; EPPO, 2014|
|-Nova Scotia||Eradicated||Agr. Canada; EPPO, 2014|
|-Ontario||Restricted distribution||Agr. Canada; Kosztarab, 1996; EPPO, 2014|
|-Quebec||Restricted distribution||Agr. Canada; EPPO, 2014|
|Mexico||Present||Borchsenius, 1966; EPPO, 2014|
|USA||Widespread||Borchsenius, 1966; EPPO, 2014|
|-New Jersey||Present||EPPO, 2014|
|-New York||Present||EPPO, 2014|
|-Washington||Present||Knight et al., 2001; EPPO, 2014|
|-Washington||Present||Knight et al., 2001; EPPO, 2014|
|-West Virginia||Present||Kozar et al., 1994|
Central America and Caribbean
|Cuba||Present||Nakahara, 1982; EPPO, 2014|
|Argentina||Widespread||Borchsenius, 1966; EPPO, 2014|
|Bolivia||Widespread||Nakahara, 1982; EPPO, 2014|
|-Minas Gerais||Present||EPPO, 2014|
|-Rio de Janeiro||Present||EPPO, 2014|
|-Rio Grande do Sul||Present||EPPO, 2014|
|-Santa Catarina||Present||EPPO, 2014|
|-Sao Paulo||Present||EPPO, 2014|
|Chile||Widespread||Borchsenius, 1966; EPPO, 2014|
|Ecuador||Restricted distribution||EPPO, 2014|
|Paraguay||Restricted distribution||Nakahara, 1982; EPPO, 2014|
|Peru||Widespread||Nakahara, 1982; EPPO, 2014|
|Uruguay||Widespread||Nakahara, 1982; EPPO, 2014|
|Venezuela||Widespread||Nakahara, 1982; EPPO, 2014|
|Austria||Widespread||Kosztarab and Kozár, 1988; EPPO, 2014|
|Belgium||Absent, formerly present||EPPO, 2014|
|Bulgaria||Restricted distribution||196*||Kosztarab and Kozár, 1988; EPPO, 2014|
|Croatia||Widespread||Kosztarab and Kozár, 1988; EPPO, 2014|
|Cyprus||Absent, confirmed by survey||EPPO, 2014|
|Czech Republic||Restricted distribution||EPPO, 2014|
|Denmark||Absent, formerly present||EPPO, 2014|
|Estonia||Absent, no pest record||EPPO, 2014|
|Finland||Absent, no pest record||EPPO, 2014|
|France||Widespread||Kosztarab and Kozár, 1988; EPPO, 2014|
|Germany||Widespread||1946||Kosztarab and Kozár, 1988; EPPO, 2014; Schönfeld, 2015|
|Greece||Restricted distribution||Kosztarab and Kozár, 1988; EPPO, 2014|
|Guernsey||Absent, no pest record||EPPO, 2014|
|Hungary||Widespread||1928||Kosztarab and Kozár, 1988; EPPO, 2014|
|Italy||Widespread||Kosztarab and Kozár, 1988; EPPO, 2014|
|Latvia||Absent, no pest record||EPPO, 2014|
|Lithuania||Absent, intercepted only||EPPO, 2014|
|Malta||Absent, no pest record||EPPO, 2014|
|Moldova||Restricted distribution||EPPO, 2014|
|Netherlands||Restricted distribution||EPPO, 2014|
|Norway||Absent, intercepted only||EPPO, 2014|
|Poland||Absent, formerly present||Kosztarab and Kozár, 1988; EPPO, 2014|
|Portugal||Widespread||Kosztarab and Kozár, 1988; EPPO, 2014|
|-Madeira||Present||Kosztarab and Kozár, 1988; EPPO, 2014|
|Romania||Widespread||Kosztarab and Kozár, 1988; EPPO, 2014|
|Russian Federation||Restricted distribution||EPPO, 2014|
|-Central Russia||Present||Danzig, 1993|
|-Eastern Siberia||Absent, formerly present||Danzig, 1993; EPPO, 2014|
|-Russian Far East||Present||Danzig, 1993; EPPO, 2014|
|-Southern Russia||Present||Danzig, 1993; EPPO, 2014|
|Slovakia||Present||Kosztarab and Kozár, 1988; EPPO, 2014|
|Slovenia||Restricted distribution||Kosztarab and Kozár, 1988; EPPO, 2014|
|Spain||Restricted distribution||Kosztarab and Kozár, 1988; EPPO, 2014|
|Sweden||Absent, intercepted only||EPPO, 2014|
|Switzerland||Present||****||Kosztarab and Kozár, 1988; EPPO, 2014|
|UK||Absent, intercepted only||Kosztarab and Kozár, 1988; EPPO, 2014|
|Ukraine||Restricted distribution||1952||Kosztarab and Kozár, 1988; EPPO, 2014|
|Yugoslavia (Serbia and Montenegro)||Widespread||Kosztarab and Kozár, 1988|
|Australia||Restricted distribution||****||Kosztarab and Kozár, 1988; EPPO, 2014|
|-New South Wales||Present||EPPO, 2014|
|-South Australia||Present||EPPO, 2014|
|-Victoria||Restricted distribution||EPPO, 2014|
|-Western Australia||Present||EPPO, 2014|
|New Zealand||Widespread||1908||Borchsenius, 1966; New Zealand MAF Biosecurity, 2011; EPPO, 2014|
Risk of IntroductionTop of page This species is considered an object of quarantine regulations on almost all continents. In the past few years, however, the European Union has deleted it from its quarantine list, because of its presence in most of the member countries.
Hosts/Species AffectedTop of page D. perniciosus is a polyphagous species. The host plant range could probably be much wider. However, on some of the recorded host plants this insect cannot finish development, which means that these species are not true host plants. The importance of the host plant is different in different parts of the world, so for each region only the local literature sources can give a true picture. In general, this species is very dangerous on deciduous fruit trees including apple, pear, peach, plum, currants and some woody ornamental plants (Konstantinova, 1976; APPPC, 1987; Davidson and Miller, 1990; Kozar, 1990a; Kosztarab, 1996). Different varieties of fruit species show different susceptibility in different parts of the world, and many fruit species have been studied in this respect (Bichina and Gatina, 1976; Shalamberidze, 1978; Verma and Sriravasta, 1990).
Growth StagesTop of page Fruiting stage, Post-harvest, Vegetative growing stage
SymptomsTop of page D. perniciosus infests mostly the bark on the stem and branches of the tree. Sometimes it can be found on the fruits, where it causes red spot. In cases of heavy infestation, the branches can die.
The red spots on the fruits and leaves do not cause great damage; however, it can cause problems in quarantine inspection, because the species is a quarantine insect in many countries.
List of Symptoms/SignsTop of page
|Fruit / lesions: black or brown|
|Leaves / abnormal colours|
|Leaves / abnormal leaf fall|
|Leaves / necrotic areas|
|Stems / dieback|
|Stems / discoloration of bark|
|Whole plant / dwarfing|
|Whole plant / early senescence|
|Whole plant / plant dead; dieback|
Biology and EcologyTop of page D. perniciosus has 1-5 generations in different parts of the world. Its development differs in these regions and detailed information can be found only in local publications. It overwinters in colder countries as first-instar larvae. The female has three stages (first- and second-instar larvae and the adult) while the male has five (first- and second-instar larvae, the pronymph, nymph and adult). In central Europe, the adults appear at the end of April, and in northern Europe 1 or 2 months later. The crawlers continue to appear for 1-2 months: the females have about 100 crawlers. The development of one generation requires about 45-80 days (Kosztarab and Kozar, 1988).
The first-instar larva is the dispersal phase. It can be distributed by wind, birds or flying insects. The most important means of transport, however, is infested nursery material. Using this method, this pest was accidentally introduced throughout the world.
The development was studied in detail in different countries. A comprehensive biological monograph has not yet been published. There are some smaller, but important, works available in different countries: For example, see Benassy (1969). In general, development starts at 7°C and this pest is tolerant of cold winter temperatures.
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Ablerus kashmirensis||Parasite||Adults/Nymphs||India; Jammu and Kashmir||apples|
|Aphytis mytilaspidis||Parasite||Adults/Nymphs||India||fruit trees|
|Aphytis proclia||Parasite||Adults/Nymphs||India; Jammu and Kashmir; Republic of Georgia||apples; fruit trees|
|Aphytis sp. nr. diaspidis||Parasite||Adults/Nymphs||India||fruit trees|
|Chilocorus bijugus||Predator||Adults/Nymphs||Himachal Pradesh; India; USSR|
|Chilocorus circumdatus||Predator||Adults/Nymphs||Australia||apricots; peaches; plums|
|Chilocorus geminus||Predator||Adults/Nymphs||China; Xinjiang; USSR||Ziziphus|
|Chilocorus infernalis||Predator||Adults/Nymphs||Pakistan; USSR|
|Chilocorus kuwanae||Predator||Adults/Nymphs||India; USA||fruit trees|
|Chilocorus renipustulatus||Predator||Adults/Nymphs||Europe||fruit trees|
|Chilocorus similis||Predator||Adults/Nymphs||Czechoslovakia; USA||fruit trees|
|Chilocorus stigma||Predator||Adults/Nymphs||Australia||apricots; peaches; plums|
|Comperiella bifasciata||Parasite||New Zealand||pears|
|Cybocephalus gibbulus||Predator||Adults/Nymphs||India||fruit trees|
|Encarsia citrina||Parasite||Adults/Nymphs||New Zealand||pears|
|Encarsia fasciata||Parasite||Adults/Nymphs||France; Germany||fruit trees|
|Encarsia perniciosi||Parasite||Adults/Nymphs||Austria; Bulgaria; Chile; Czechoslovakia; France; Germany; Greece; India; India; Jammu and Kashmir; Italy; New South Wales; New Zealand; Pakistan; Republic of Georgia; South Africa; Spain; Switzerland; USSR; Yugoslavia||apples; fruit trees; pears|
|Habrolepis aspidioti||Parasite||California||fruit trees|
|Marietta carnesi||Parasite||Adults/Nymphs||India; Jammu and Kashmir||apples|
|Orius insidiosus||Predator||Adults/Nymphs||USA; Virginia||apples|
|Rhyzobius lindi||Predator||Adults/Nymphs||California||fruit trees|
|Rhyzobius lophanthae||Predator||Adults/Nymphs||Czechoslovakia; Europe; India||fruit trees|
|Thomsonisca shutovae||Parasite||USSR||fruit trees|
Notes on Natural EnemiesTop of page D. perniciosus has a high number of parasitoids and predators, which have been studied in detail, especially in the Palearctic region (Trjapitzin, 1978; Kosztarab and Kozar, 1988). There are many papers on natural enemies of this pest in different countries (Ahmad and Ghani, 1971; Rosen and De Bach, 1976; Folkina, 1978; Myartzeva, 1978; Trjapitzin, 1978; Popova, 1979; Darling and Johnson, 1984; Katsoyanos, 1984; Yasnosh, 1985; Bhagat et al., 1988; Titayavan and Davis, 1988; Thakur et al., 1989; Sharma et al., 1990; Bull et al., 1993).
These natural enemies are efficient regulators of D. perniciosus, and they keep down the population density in natural habitats. Where chemical control is inefficient against D. perniciosus, but kills the natural enemies, local outbreaks can occur in orchards. The efficiency of natural enemies is also reduced in urban regions by pollution, where subsequently the pest can cause damage on ornamental plants.
There have been many successful special biological control projects against D. perniciosus in different parts of the world, especially in the USA, Europe and Russia. Usually the main specialized parasitoid Encarsia perniciosi was reared, released and established, according to Rosen (1991). Concerning the list of parasitoids it is important to notice that some parasitoids are only parasitoids, but that others may be both parasitoids and hyperparasitoids, and in many published cases this is not clarified.
All data cited in the table are based on original publications. Therefore there could be misidentifications of the parasitoids or even the host. The generic names also could be not updated. Very often the origin, and the real distribution of the natural enemies cannot be established from the original source and needs to be surveyed. The time of the introduction into a particular country also needs clarification: for example there was not a special release of Encarsia perniciosi in Hungary, but the parasitoid appeared very quickly because of the mass release in the neighbouring countries.
ImpactTop of page Crop loss caused by D. perniciosus on different trees is difficult to assess. The trees lose their vigour and have a shorter lifespan. Shortly after introduction to a new country, this pest can infest and kill whole trees and plantations. This species is a quarantine pest in different parts of the world. The import and export of infested fruits or plants could be refused.
Local outbreaks have been observed in different parts of the world on fruit trees and ornamental plants, as in Hungary (Kozar and Drozdjak, 1988), Switzerland (Kozar et al, 1994; Mani et al, 1995), the European part of Russia (Kozar and Konstantinova, 1981), Australia (Baker, 1977) and Canada (Ker and Sears, 1986).
Detection and InspectionTop of page In cases of heavy infestation, greyish scales can be found on the bark of the trees. Lighter infestations can be found by laboratory examination with stereomicroscope analyses of the branches, following the survey system described in detail by Kozar (1990b, c). The pheromone of the species is also known. The different compounds of the D. perniciosus pheromone attract the males and parasitoids. Colour and sticky traps have been developed to follow the flight of the males and parasitoids (Kozar, 1990c). The red spots on the fruits can also indicate infestation.
Similarities to Other Species/ConditionsTop of page D. perniciosus can be identified only by its characteristics on microscope slides. A method of slide mounting is described by Kosztarab and Kozar (1988). This species is often found together with other species of Diaspidiotus. According to microscopic characters, it is easy to distinguish, because D. perniciosus has no groups of perivulvar pores, which are present in other Diaspidiotus species infesting fruit trees. D. perniciosus overwinters as first-instar larvae, whereas other species overwinter as second instars or females. D. perniciosus is ovoviviparous, whereas the other species lay eggs.
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
Regulatory Control (Plant Quarantine and Certification)
Many different countries have special regulations against this pest, the most detailed elaborated by EPPO (1996).
Cultural Control and Sanitary Methods
Scale-free nursery material is very important, because young plants can die very quickly once infested. Cutting the heavily infested parts of the trees and cleaning the bark from infestation can help to increase the efficiency of chemical treatments. There is discussion in the literature about the role of the surrounding infested vegetation: it is partly a source for the re-infestation of the plantations, but it could also be a refuge for the natural enemies.
There are no resistant varieties against this pest. There are several tolerant varieties of different fruit species; however, these varieties are not usually very marketable.
Biological control against D. perniciosus is well studied and is one of the more successful projects of this nature in the world, especially using Encarsia perniciosi. Such programmes can give substantially good results only after a relatively long time, and only in the absence of toxic insecticides. The additional release of different predators, and conserving the local natural enemies can increase the efficiency of control.
For a long time fumigation of seedlings was the most important means of preventing early infestation. For this purpose different chemicals were used, such as hydrogen cyanide, fostoxin or methylbromide (Obretenchev and Stanev, 1992; Anon., 1993; Mordkovich and Chernei, 1993). There are also quarantine recommendations, or requirements for treatment of the fruits to prevent infestation by this pest (although the infestation of plants from fruits has never been proved). For this purpose different methods were proposed as irradiation, fumigation, cold treatment or warm treatment (Angerilli and Fitzgibbon, 1990 and others).
For the treatment of plantations, dormant oil spraying was proposed and is very effective (Ker and Sears, 1986; Chander and Kakar, 1994; Hippe et al., 1995; Isufi and Myrtal, 1996). Almost all the toxic insecticides used (phosphororganics, carbamates, pyrethroids) are effective against crawlers of this pest. The use of these chemicals in summer against other pests such as codling moth, leafrollers, leafminers or mites can often keep down the density of this pest.
Early Warning Systems, Field Monitoring/Economic Threshold Levels
Monitoring and forecasting systems were analysed in detail by Kozar (1990b, c). It was established that there are several methods for monitoring and detection, like white sticky traps (for the males), suction traps (for the males and parasitoids). Pheromone traps collecting the males and parasitoids are the most widely used. Because of the quarantine requirements an economic threshold was not developed. However, in some countries (where there is no strict prohibition, like in Hungary or Russia) there is a special quarantine threshold, only 5% of the fruits can be infested with this scale.
Special integrated control projects have been developed against orchard pests including scale insects (Kozar and Varjas, 1976; Ripa and Caltagirone, 1994; Paloukis and Navzoridis, 1995; Jenser et al., 1997). These programmes propose the use of new, selective, non-toxic products (different types of juvenoids and chitin inhibitors, for example) against pests, which have a good effect against different groups of pests, including scale insects. In these orchards the introduced and local natural enemies could reach a high density and reduce the density of pests. If such programmes are implemented, it is possible to have fruits with very low numbers of D. perniciosus.
ReferencesTop of page
1993. Quarantine procedure. Hydrogen cyanide fumigation of woody plants to control Quadraspidiotus perniciosus. Bulletin OEPP, 23(2):204
Angerilli NPD, Fitzgibbon F, 1990. Effects of cobalt gamma radiation on San Jose scale (Homoptera: Diaspididae) survival on apples in cold and controlled-atmosphere storage. Journal of Economic Entomology, 83(3):892-895.
APPPC, 1987. Insect pests of economic significance affecting major crops of the countries in Asia and the Pacific region. Technical Document No. 135. Bangkok, Thailand: Regional Office for Asia and the Pacific region (RAPA).
Bazarov BB, Shmelev GP, 1971, Akad. Nauk Tadzh. SSR Inst. Zool. Parazitol. 11:220.
Benassy C, 1969. Contribution a l'etude du peuplement d'un vegetal par Quadraspidiotus perniciosus Comst. (Homoptera Coccidae) et a l'influence de ce dennier sur son hote. These, Faculte des Sciences de L'Universite de Lyon, No 319.
Bichina TI, Gatina Esh, 1976. Varietal susceptibility of apple trees to San Jose scale and its integrated control. Ustoichivost' Vinograda i Plodov. Kul'tur k Zabolevaniyam i Vreditelyam. 124-137. Stiinca, Kishinev: Moldavian SSR.
Borchsenius NS, 1966. A Catalogue of the Armoured Scale Insects (Diaspidoidea) of the World. (In Russian). Leningrad, Russia: Akademii Nauk SSR Zoologicheskogo Instituta, 449 pp.
Bull BC, Raupp MJ, Hardin MR, Sadof CS, 1993. Suitability of five horticulturally important armored scale insects as hosts for an exotic predaceous lady beetle. Journal of Environmental Horticulture, 11(1):28-30
Bustshik TN, 1958. A contribution to the comparative morphology of the males of the scale insects (Homoptera, Coccoidea, Diaspididae). Trudy Akad. Nauk. SSSR. Vsesoiuzn. Entomol. Obsh., 46:162-269.
Chowdhuri AN, 1977. San Jose scale. Quadraspidiotus perniciosus (Comstock). Science and Culture, 43:512.
Danzig EM, 1993. Fauna of Russia. Families Phoenicoccidae and Diaspididae. St. Petersburg, Russia: Nauka, 450 pp.
Davidson JA, Miller DR, 1990. Ornamental plants. In: Rosen D, ed. Armoured Scale Insects, their Biology, Natural Enemies and Control. Vol. 4B. Amsterdam, Netherlands: Elsevier, 603-632.
EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm
Ghauri M, 1962. The Morphology and Taxonomy of Male Scale Insects (Homoptera, Coccoidea). British Museum (Natural History). Dorking, UK: Adlard and Son, 221 pp.
Jenser G, Balßzs K, ErdTlyi C, Haltrich A, Kozßr F, Mark= V, Rßcz V, Samu F, 1997. The effect of an integrated pest management program on the arthropod populations in a Hungarian apple orchard. Zahradnictvi^acute~, 24(2):63-76; 16 ref.
Ker KW, Sears MK, 1986. Effectiveness of superior oil applied to apple for control of the San Jose scale, Quadraspidiotus perniciosus, and the European fruit scale, Quadraspidiotus ostrepformis (Homoptera: Diaspididae). Proceedings of the Entomological Society of Ontario, 117:45-48
Kosztarab M, 1996. Scale Insects of North Eastern North America: identification, biology, and distribution. Martinsville, USA: Virginia Museum of Natural History, 650 pp.
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