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California Red Scale Aonidiella
aurantii Maskell --
Hemiptera: Diaspididae |
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History
California red scale infests all parts of a citrus tree except the
roots. When it is moderately abundant
or dense on the fruit, the fruit is reduced in grade or culled in the packinghouse. When the red scale is dense on leaves and
wood, it defoliates portions of the tree and either kills all or parts of
branches within the tree. The scale occurs in all of the citrus
growing regions of California and Arizona, including coastal groves, inland
southern California, the San Joaquin Valley, and the low desert. Events leading to the biological
control of California red scale have been variously presented in previous
sections, as it marks one of the most thorough and scientific approaches in
biological control. It is also the
longest campaign in the history of biological control. Kennett et al. (1999) summarized the history of these studies,
and Ebeling (1959) regarded red scale as the most important citrus pest
worldwide. Although it is not as
widely distributed as purple scale, it causes greater damage and is more
difficult to control. California red
scale is believed to be of oriental origin (Quayle 1938). Natural
Enemies During 1889-1947
a variety of exotic natural enemies were introduced into California for
control, the earliest efforts emphasizing introductions of coccinellid
predators, of which more than 40 species were imported during 1889-1892
(Compere 1961). Most failed to
established and attention was then given to parasitoids. During 1900-1908 several species,
including Aspidiotiphagus
citrinus (Craw), Comperiella bifasciata Howard and Pteroptrix (= Casca) chinensis (Howard) were imported from southern China,
but none established (Compere 1961).
One parasitoid that became associated with the scale in southern
California during this time was Aphytis
chrysomphali (Mercet),
although its origin remained obscure.
This parasitoid was propagated during 1902-1904 and distributed to
citrus orchards on request, but dramatic results were not obtained (Compere
1961). Following this poor
performance of A. chrysomphali, additional parasitoids
in the genus Aphytis
were discovered in foreign areas, but they were erroneously believed to be
the same species. During 1916-1924 additional introductions of Comperiella bifasciata were made from Japan
(Compere 1961). It did not rear on
red scale in the laboratory so a factitious host, Chrysomphalus bifasciculatus Ferris was used. Releases in the field did not result in
parasitism of red scale, but rather the yellow scale, Aonidiella citrina (Coquillett) (Smith
1942). Later the original host scale
in Japan was identified as Aonidiella
taxus Leonardi Chrysomphalus bifasciculatus and not A. aurantii and C.
aonidium (L.) as
originally believed. Several coccinellid predators were imported from
South Africa, Australia, South America and China during 1918-1934 (Compere
1961). An Aphytis sp. (probably A. lingnanensis DeBach) was imported from southern China
in 1924 under the erroneous name of A.
chrysomphali, but did
not become established. The
endoparasitoid Habrolepis
rouxi Compere was
imported from South Africa in 1937, and was widely colonized. Establishment was obtained in a very small
area, however (Flanders 1944a). Comperiella bifasciata was tried once more in
1940, but this time the collections were made on A. auranti in southern China (Smith
1942), and were then successfully propagated on that host in California
(Flanders 1943a). The parasitoid
eventually spread throughout all of the southern California area except near
the coast (DeBach et al. 1955).
Flanders (1944b) and Teran & DeBach (1963) distinguished two
distinct biological races of this parasitoid, one from A. auranti in China and the other from A. taxus and C.
bifasciculatus in Japan,
which solved the earlier puzzle. The period 1947-1949 marked many shipments of
parasitized red scales being sent to California from southern China and
Taiwan. Propagation was successful
with many species, but only Aphytis
lingnanensis from
southern China and Encarsia
(= Prospaltella) perniciosi (Tower) from Taiwan
became established (DeBach et al. 1950, DeBach 1953, Rosen & DeBach
1978). Encarsia perniciosi
readily established in the coastal areas of southern California. However, A. lingnanensis
became dominant by 1958, displacing the long established A. chrysomphali everywhere but in a few coastal areas
(DeBach & Sundby 1963). Although
producing good biological control in coastal areas, A. lingnanensis was relatively ineffective in the warmer,
drier interior areas of southern California.
Expanded Exploration Explorations were expanded in Asia during 1956-57, resulting
in the importation of some new parasitoids, including two Aphytis spp. (DeBach 1959). Only Aphytis
melinus DeBach <PHOTO> from northern India
and Pakistan was established in California, however. Aphytis
fisheri DeBach, Aphytis proclia (Walker), Coccobius (= Physcus) debachi (Compere & Annecke) and Aspidiotiphagus citrinus were liberated but did not
become established. Importations
during 1960-1964 of Aphytis
coheni DeBach and Aphytis holoxanthus DeBach from Israel and Aphytis africanus Quednau from South Africa,
all failed to establish (Rosen & DeBach 1978). Aphytis melinus spread rapidly following
liberation in 1958, and soon displaced A.
lingnanensis throughout
the area. By 1964 A. melinus was the dominant parasitoid on A. aurantii in southern California except along coastal
areas, where A. lingnanensis remained common or
dominant (DeBach 1966, 1969). There
was a gradual decline in red scale abundance in southern California after 1962
which was attributed to these two parasitoids. Presently A.
melinus is complemented
by C. bifasciata in the interior and
intermediate areas while A. lingnanensis is complemented by E. perniciosi in coastal areas (DeBach 1965a, 1969). Control varies from partial to complete
depending on the climatic zone in which red scale occurs. Extreme temperatures cause abnormal sex
ratios and progeny production in A.
melinus and A. lingnanensis which helps to explain some of the poor
performance witnessed with these parasitoids in certain climatic zones (Kfir
& Luck 1979). The discovery of Aphytis melinus was the result of increased
taxonomic knowledge and of previous ecological field studies with A. chrysomphali beginning in 1946 and with A. lingnanensis starting in 1948 (DeBach 1974). Studies with these parasitoids showed the
importance of the genus Aphytis,
and led to the search for additional species from climatic zones similar to
the interior areas of southern California.
Biological control of red scale was investigated in
Australia on the same pattern as that for California. Around 1902 several species of natural
enemies were imported from different countries, but only one species, Aphytis chrysomphali established in Western
Australia (Wilson 1960). Comperiella bifasciata became established in the
1940's, and the subsequent importations of Aphytis melinus
in 1961 and Encarsia perniciosi in 1970 established these
parasitoids. Attempts to establish in
Australia Aphytis lingnanensis, A. coheni, A.
riyahdi DeBach and Habrolepis sp. after 1977 were not
successful (Furness et al. 1983). Aphytis chrysomphali was displaced by A. melinus after 1972 in the Lower Murray Valley in the
states of Victoria and South Australia (Furness et al. 1983). A decline in A. aurantii
abundance following colonization of Aphytis
melinus was documented
(Campbell 1976). In Queensland
effective biological control was caused by Aphytis lingnanensis
and Comperiella bifasciata (Smith 1978a). Importations of Aonidiella
aurantii parasitoids
were also made in South Africa, France, Sicily, Cyprus, Greece, Morocco and
Israel, with the same pattern of displacement by A. melinus of A.
chrysomphali witnessed
(DeBach & Argyriou 1967, Rosen 1967, Orphanides 1984). On the other hand, in Israel A. melinus displaced A.
chrysomphali, but it did
not displace A choeni (Rosen 1967); and in South
Africa A. africanus is still dominant despite
the establishment of A. melinus, A. lingnanensis and A.
coheni (Annecke &
Moran 1982). Partial to complete biological
control was reported in other countries, such as Greece (DeBach &
Argyriou 1967), France (Benassy & Bianchi 1974), Argentina (Crouzel et
al. 1974), Cyprus (Orphanides 1984), and Chile (Gonzalez 1969). For some unknown reason, Comperiella bifasciata alone was responsible for
the successful biological control witnessed in France (Kennett et al. 1999)
(also see Coquillett 1893, Coquillett & Koebele 1893, Jones 1936,
Sokoloff & Klotz 1942, Campbell 1943, Jenkins 1945, Gressitt &
Flanders 1949, Bodenheimer 1951, Flanders 1953, Flanders & Gressitt 1958,
Landi & DeBach 1960, Harpaz 1961, Quednau 1964, 1965; Quednau &
Hübsch 1964, Rosen 1965, Gonzalez & Rojas 1966, Benassy & Euverte
1967). Management The strategy of managing red scale in California by 2004
had developed into a periodic parasitoid release program in those orchards
where climate posed restrictions on the natural annual increase of the
parasitoids or where insecticidal drift from neighboring areas upset natural
balances. The boost of adding
additional parasitoid individuals in such areas provides satisfactory red
scale control in many orchards.
Nevertheless, additional natural enemies might be sought in Pakistan
and India. REFERENCES: FURTHER RELATED REFERENCES Annecke, D. P. 1958.
Ants and citrus. Citrus Grower
196: 1–3. Annecke, D. P. & V.
C. Moran. 1982. Insects and Mites of cultivated plants in
South Africa. Butterworth & Co.,
South Africa. 383 p. Argyriou, L. C.
1990. Olive". In:
D. Rosen (ed.), Armoured scale insects, their biology, natural enemies and control.
Vol. 4B. World Crop Pests. Elsevier, Amsterdam, the Netherlands: 579–583. Bedford, E. C. G. 1998. Red scale Aonidiella
auranii (Maskell). In: E. C. G. Bedford,
M. A. Van den Berg & E. A. De Villiers (eds.), Citrus pests in the
Republic of South Africa. Dynamic Ad., Nelspruit, South Africa: 132–134. Benassy, C. &
G. Euverte. 1967. Perspectives nouvelles dans la lutte
contre Aonidiella auranti au Maroc (Hom.
Diaspididae). Entomophaga 12: 449-59. Benassy, C. &
H. Bianchi. 1974. Observations sur Aonidiella aurantii Mask. et son parasite
indigene Comperiella bifasciata How. (Hymenoptera,
Encyrtidae). Bull. SROP 3:
39-50. Borchsenius, N. S. 1966. A catalogue of the armored scale insects
(Diaspidoidea) of the world. (In Russian.) Nauka, Moscow, Leningrad, Russia. 449 pp. Bodenheimer, F. S. 1951. Citrus Entomology in the Middle East. Junk Publ., The Hague. 663 p. Campbell, M. M.
1976. Colonization of Aphytis melinus DeBach [Hymenoptera:
Aphelinidae] in Aonidiella
aurantii (Mask.) [Hemiptera:
Coccidae] on citrus in South Australia.
Bull. Ent. Res. 65: 659-68. Campbell, T. F.
1943. Introduction of scale
parasites from California. Austral.
Council Sci. & Indus. Res. Journal 16: 41-2. Chua, T. H.
& B. J. Wood, 1990.
Other tropical fruit trees and shrubs". In: D. Rosen (ed.),
Armoured scale insects, their biology, natural enemies and control. Vol. 4B. World Crop Pests. Elsevier,
Amsterdam, the Netherlands: 543–552. Compere, H.
1961. The red scale and its
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31: 173-278. Coquillet, D. W.
1893. Report on the
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Botto. 1973. Lucha biológica contra la "cochinilla
roja australiana" Aonidiella
aurantii (Maskell)
(Hom.: Diaspididae) en cítricos.
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perniciosi Tower on the
California red scale. J. Econ.
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1959. New species and strains
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Eulophidae) parasitic on the California red scale, Aonidiella aurantii (Mask.) in the Orient. Ann. Ent. Soc. Amer.
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