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''Pseudococcus viburni'' (formerly ''Pseudococcus affinis'' ( Maskell), and commonly known as the obscure mealybug and tuber mealybug) is a close relative of the grape mealybug (''P. maritimus'') and a
pest Pest or The Pest may refer to: Science and medicine * Pest (organism), an animal or plant deemed to be detrimental to humans or human concerns ** Weed, a plant considered undesirable * Infectious disease, an illness resulting from an infection ** ...
of the
vineyard A vineyard (; also ) is a plantation of grape-bearing vines, grown mainly for winemaking, but also raisins, table grapes and non-alcoholic grape juice. The science, practice and study of vineyard production is known as viticulture. Vine ...
s around the world.


Distribution

''P. viburni'' is a pest of vineyards in
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 List of islands of New Zealand, smaller islands. It is the ...
, Hamlet (2005), p. 1. the Central Coast of
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the ...
, and the tea gardens of northern
Iran Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkm ...
. Abbasipour ''et al.'' (2007), p. A120. Unlike the grape mealybug, the obscure mealybug is not native to California, having most likely been introduced to the region from either Australia or
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the souther ...
in the latter part of the 19th century. Varela ''et al.'' (2006), p. 2. It is known to infest apple and pear in South Africa.


Taxonomy

The obscure mealybug is thought to have evolved in Australia or South America. Its history in North America is not clear; from 1900 (when ''P. maritimus'', the grape mealybug, was first described) to 1960, the obscure mealybug was variably misidentified as or synonymized with ''P. maritimus'', ''P. longispinus'', ''P. obscurus'' Essig, ''P. capensis'' Brian, ''P. malacearum'' Ferris, and ''P. affinis'' (Maskell). This
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. A ...
confusion has hampered attempts to trace the exact origin of the species, but its presence in both Australia and South America strongly suggests that it is of
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final st ...
n origin.


Morphology

Obscure mealybugs exhibit a high degree of
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
; females are flightless, larger, and longer-lived than the winged males, who cannot feed and die immediately after
mating In biology, mating is the pairing of either opposite- sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. '' Copulation'' is the union of the sex organs of two sexually rep ...
.


Females

The bodies of nymphal and adult female obscure mealybugs are rectangular, with rounded
anterior Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
and
posterior Posterior may refer to: * Posterior (anatomy), the end of an organism opposite to its head ** Buttocks, as a euphemism * Posterior horn (disambiguation) * Posterior probability, the conditional probability that is assigned when the relevant evidenc ...
ends. Adult females range from 1-5mm in length. New England Greenhouse Floriculture Guide (2003–2004), p. B-14. Like all
mealybug Mealybugs are insects in the family Pseudococcidae, unarmored scale insects found in moist, warm habitats. Many species are considered pests as they feed on plant juices of greenhouse plants, house plants and subtropical trees and also act as a ...
s, the obscure mealybug covers its body in a white, waxy secretion which accumulates in clumps along thin
filaments The word filament, which is descended from Latin ''filum'' meaning "thread", is used in English for a variety of thread-like structures, including: Astronomy * Galaxy filament, the largest known cosmic structures in the universe * Solar filament, ...
protruding from its
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton ( endoskeleton) in for example, a human. In usage, some of the ...
. This clumpy secretion lends the mealybug its characteristic mealy appearance.


Males

Male obscure mealybugs are tiny, fragile insects with long antennae and a single pair of wings. They bear a strong resemblance to common
house flies The housefly (''Musca domestica'') is a fly of the suborder Cyclorrhapha. It is believed to have evolved in the Cenozoic Era, possibly in the Middle East, and has spread all over the world as a commensal of humans. It is the most common fly ...
, but have two white wax tail filaments. Hamlet (2005), p. 5.


Identification

It is important for vintners to distinguish between the obscure mealybug and its close relatives, the grape and
vine mealybug ''Planococcus ficus'', commonly known as the vine mealybug, is a species of mealybug, belonging to the family Pseudococcidae, native to tropical and subtropical regions. The vine mealybug is found in Europe, Northern Africa, Southern Africa, the ...
. The obscure mealybug has longer, thinner, and more crooked filaments than does the vine mealybug, making the obscure mealybug look comparatively untidy. The most distinctive feature of the obscure mealybug is the set of two to four exceptionally long caudal filaments growing from the posterior of large nymphs and adult females. The
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs ...
's waxy secretion accumulates heavily on these filaments, making it appear to have several long "tails". The obscure mealybug more closely resembles the grape mealybug than it does the vine mealybug, but the two can be distinguished by the color of the defensive fluid they secrete when disturbed; grape mealybugs secrete reddish-orange fluid, while obscure mealybugs secrete clear fluid. When crushed, the obscure mealybug's body contents (
guts The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans an ...
) are pinkish-grey.


Life cycle


Females

Being in the order Hemiptera (so-called "true bugs"), obscure mealybugs undergo
incomplete metamorphosis Hemimetabolism or hemimetaboly, also called incomplete metamorphosis and paurometabolism,McGavin, George C. ''Essential Entomology: An Order-by-Order Introduction''. Oxford: Oxford University Press, 2001. pp. 20. is the mode of development of cert ...
; nymphal young closely resemble adults in body shape, take six to nine weeks to mature, and retain the use of all six legs throughout their entire lives. Depending on temperature, obscure mealybugs may complete 2-3 generations per year; females will lay clutches of several hundred orange
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
in cottony sacs, from which nymphs will hatch and emerge after about 5–10 days. If environmental conditions are too cold, young nymphs will remain in the sac until temperatures rise. Obscure mealybugs lay eggs all year long, and during winter, under the bark of trees and vines (though there is no true
dormancy Dormancy is a period in an organism's life cycle when growth, development, and (in animals) physical activity are temporarily stopped. This minimizes metabolic activity and therefore helps an organism to conserve energy. Dormancy tends to be c ...
). These overwintering populations include individual mealybugs from all stages of development, but are dominated by eggs and first instars; overwinter mortality for young nymphs is high, but a few individuals (normally the ones quickest to hatch) will survive and feed on the first spring leaves. Mortality in non-overwintering generations is greatly decreased. Hamlet (2005), pp. 5–6.


Males and mating

Male obscure mealybugs do not feed, and have very short lifespans (2–3 days); males will spin
cocoon Cocoon may refer to: *Cocoon (silk), a pupal casing made by moth caterpillars and other insect larvae *Apache Cocoon, web development software * ''Cocoon'' (film), a 1985 science fiction-fantasy film **'' Cocoon: The Return'', 1988 sequel to ''Coco ...
s shortly after hatching, where they develop wings. Upon emerging from their cocoons, male obscure mealybugs will fly toward the scent of the female's
sex pheromone Sex pheromones are pheromones released by an organism to attract an individual of the same species, encourage them to mate with them, or perform some other function closely related with sexual reproduction. Sex pheromones specifically focus on ind ...
, mate as many times as possible, then die. Given the male's short lifespan, the timing of the female's emission of sex pheromone is crucial; females will emit the pheromone day and night around the time of male emergence (especially in the spring), then cease immediately after fertilization. Obscure mealybugs generally mate at dusk and at dawn. The female obscure mealybug sex pheromone has the unfortunate property of sometimes attracting parasitic wasps (such as ''Tetracnemoidea peregrina''), and is therefore a
kairomone A kairomone (a coinage using the Greek καιρός ''opportune moment'', paralleling pheromone"kairomone, n.". OED Online. September 2012. Oxford University Press. http://www.oed.com/view/Entry/241005?redirectedFrom=kairomone (accessed 3 Octobe ...
.


Ecology


Diet

Obscure mealybugs feed on the
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
of vines and woody-stemmed plants, especially pear and apple trees and grape vines. Some individuals are vectors for infectious
pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a ger ...
s and can transmit them from plant to plant while feeding; mealybug-spread ''grapevine leafroll associated virus type III'' (GRLaV-3), in particular, has wreaked havoc among the grapes of New Zealand, reducing the crop yield of infected vineyards by up to 60%.


Symbiosis

Female mealybugs excrete honeydew, a thick, sugary fluid created as a byproduct of
digestion Digestion is the breakdown of large insoluble food molecules into small water-soluble food molecules so that they can be absorbed into the watery blood plasma. In certain organisms, these smaller substances are absorbed through the small intest ...
(large colonies of mealybugs can produce enough honeydew to seep through bark and leaves, leaving shiny, sticky patches on the exterior of the plant). Some
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22,0 ...
s have developed a mutualistic
symbiotic relationship Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasi ...
with obscure mealybugs, tending and protecting the insects from natural enemies to increase the production of honeydew, on which the ants feed. This relationship is similar to the one that some ants have with
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A ...
s.


Pestilence

This species is a
pest Pest or The Pest may refer to: Science and medicine * Pest (organism), an animal or plant deemed to be detrimental to humans or human concerns ** Weed, a plant considered undesirable * Infectious disease, an illness resulting from an infection ** ...
in North America, South America, Australia, New Zealand, and Iran, especially in vineyards and fruit orchards. The obscure mealybug has caused particularly substantial damage to vineyards in the Central Coast of California, where it is an
introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived the ...
and has no natural
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
s.


Control

Many attempts have been made to control obscure mealybug populations in commercial
greenhouse A greenhouse (also called a glasshouse, or, if with sufficient heating, a hothouse) is a structure with walls and roof made chiefly of transparent material, such as glass, in which plants requiring regulated climatic conditions are grown.These ...
s and vineyards, using both synthetic pesticides and introduced mealybug
parasites Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
. Since the obscure mealybug's waxy coating and habit of seeking sheltered feeding sites protect it from water-based pesticides, oil-based
organophosphate In organic chemistry, organophosphates (also known as phosphate esters, or OPEs) are a class of organophosphorus compounds with the general structure , a central phosphate molecule with alkyl or aromatic substituents. They can be considered ...
pesticides (among the more toxic of pesticides) are most effective in reducing population density, though such pesticides can harm plants if applied after the first budding. Even so, some New Zealand obscure mealybug populations have developed a resistance to organophosphate pesticides. Because of the high toxicity and strict regulation of organophosphate pesticides, some Californian vintners have imported natural parasites of the obscure mealybug from
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the eas ...
, particularly '' Pseudaphycus flavidulus'' and '' Leptomastix epona''. Such efforts have so far produced indeterminate results; the obscure mealybug is capable of encapsulating and killing the eggs of ''L. epona'' and ''L. dactylopii'', for example, rendering those parasites ineffective. On the other hand, significant success in reducing obscure mealybug populations has resulted from isolating them from their
symbiont Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or paras ...
ants. Daane ''et al.'' (2008), pp. 45–46. Late-season insecticide has proven ineffective in apple and pear in South Africa. Nel 1983 and Kaiser et al 2001 find ''P. viburni'' to have taken residence in these pomes by that time in the season, at the
calyx Calyx or calyce (plural "calyces"), from the Latin ''calix'' which itself comes from the Ancient Greek ''κάλυξ'' (''kálux'') meaning "husk" or "pod", may refer to: Biology * Calyx (anatomy), collective name for several cup-like structures ...
or
distal Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
end, and unreachable to sprays.


References


Bibliography

* * * * * * {{Taxonbar, from=Q14621706 Pseudococcidae Grape pest insects Insect vectors of plant pathogens Insects described in 1875