Grape Phylloxera
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Grape phylloxera is an insect pest of
grapevine ''Vitis'' (grapevine) is a genus of 81 accepted species of vining plants in the flowering plant family Vitaceae. The genus consists of species predominantly from the Northern Hemisphere. It is economically important as the source of grapes, bot ...
s worldwide, originally native to eastern
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
. Grape phylloxera (''Daktulosphaira vitifoliae'' (Fitch 1855) belongs to the family
Phylloxeridae Phylloxeridae is a small family of plant-parasitic hemipterans closely related to aphids with only 75 described species. This group comprises two subfamilies (Phylloxerininae and Phylloxerinae) and 11 genera with one that is fossil. The genus ...
, within the order
Hemiptera Hemiptera (; ) is an order of insects, commonly called true bugs, comprising more than 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from ...
, bugs); originally described in France as ''Phylloxera vastatrix''; equated to the previously described ''Daktulosphaera vitifoliae'', ''Phylloxera vitifoliae''. The insect is commonly just called phylloxera (; from , leaf, and , dry). These almost microscopic, pale yellow sap-sucking
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s, related to
aphid Aphids are small sap-sucking insects in the Taxonomic rank, family Aphididae. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white Eriosomatinae, woolly ...
s, feed on the roots and leaves of grapevines (depending on the phylloxera genetic strain). On ''
Vitis vinifera ''Vitis vinifera'', the common grape vine, is a species of flowering plant, native to the Mediterranean Basin, Mediterranean region, Central Europe, and southwestern Asia, from Morocco and Portugal north to southern Germany and east to northern ...
'', the resulting deformations on roots ("nodosities" and "tuberosities") and secondary fungal infections can
girdle A belt without a buckle, especially if a cord or rope, is called a girdle in various contexts, especially historical ones, where girdles were a very common part of everyday clothing from antiquity until perhaps the 15th century, especially for w ...
roots, gradually cutting off the flow of nutrients and water to the vine.Wine & Spirits Education Trust ''"Wine and Spirits: Understanding Wine Quality"'' pgs 2–5, Second Revised Edition (2012), London,
Nymphs A nymph (; ; sometimes spelled nymphe) is a minor female nature deity in ancient Greek folklore. Distinct from other Greek goddesses, nymphs are generally regarded as personifications of nature; they are typically tied to a specific place, land ...
also form protective
gall Galls (from the Latin , 'oak-apple') or ''cecidia'' (from the Greek , anything gushing out) are a kind of swelling growth on the external tissues of plants. Plant galls are abnormal outgrowths of plant tissues, similar to benign tumors or war ...
s on the undersides of grapevine leaves of some ''Vitis'' species and overwinter under the bark or on the vine roots; these leaf galls are typically only found on the leaves of American vines. American vine species (such as ''
Vitis labrusca ''Vitis labrusca'', the fox grape, is a species of grapevines belonging to the ''Vitis'' genus in the flowering plant family Vitaceae. The vines are native to eastern North America and are the source of many grape cultivars, including Catawba, ...
'') have evolved to have several natural defenses against phylloxera. The roots of the American vines exude a sticky sap that repels the nymph form by clogging its mouth when it tries to feed from the vine. If the nymph is successful in creating a feeding wound on the root, American vines respond by forming a protective layer of tissue to cover the wound and protect it from secondary bacterial or fungal infections. Currently there is no cure for phylloxera and unlike other
grape diseases This is a list of diseases of grapes (''Vitis'' spp.). Bacterial diseases Fungal diseases Miscellaneous diseases and disorders Nematodes, parasitic Phytoplasma, virus and viruslike diseases See also *'' Ampeloglypter ater'' *'' Am ...
such as powdery or
downy mildew Downy mildew refers to any of several types of oomycete microbes that are obligate parasites of plants. Downy mildews exclusively belong to the Peronosporaceae family. In commercial agriculture, they are a particular problem for growers of c ...
, there is no chemical control or response. The only successful means of controlling phylloxera has been the
grafting Grafting or graftage is a horticulture, horticultural technique whereby tissues of plants are joined so as to continue their growth together. The upper part of the combined plant is called the scion () while the lower part is called the roots ...
of phylloxera-resistant American rootstock (usually
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two diff ...
varieties created from the ''
Vitis berlandieri ''Vitis berlandieri'' is a species of grape native to the southern North America, primarily Texas, New Mexico and Arkansas. It is primarily known for good tolerance against soils with a high content of lime, which can cause chlorosis in many vi ...
'', ''
Vitis riparia ''Vitis riparia'' Michx, with common names riverbank grape or frost grape, is a vine indigenous to North America. As a climbing or trailing vine, it is widely distributed across central and eastern Canada and the central and northeastern parts ...
'' and ''
Vitis rupestris ''Vitis rupestris'' is a species of grape native to the United States that is known by many common names including July, Coon, sand, sugar, beach, bush, currant, ingar, rock, and mountain grape. It is used for breeding several French-American hy ...
'' species) to more susceptible European ''vinifera'' vines.


Biology

Phylloxera has a complex life-cycle of up to 18 stages, that can be divided into four principal forms: sexual form, leaf form, root form, and winged form. The ''sexual form'' begins with male and female eggs laid on the underside of young grape leaves. The male and female at this stage lack a digestive system, and once hatched, they mate and then die. Before the female dies, she lays one winter egg in the bark of the vine's trunk. This egg develops into the ''leaf form''. This nymph, the fundatrix (stem mother), climbs onto a leaf and lays eggs
parthenogenetic Parthenogenesis (; from the Greek + ) is a natural form of asexual reproduction in which the embryo develops directly from an egg without need for fertilization. In animals, parthenogenesis means the development of an embryo from an unfertiliz ...
ally in a leaf gall that she creates by injecting saliva into the leaf. The nymphs that hatch from these eggs may move to other leaves, or move to the roots where they begin new infections in the ''root form''. In this form they perforate the root to find nourishment, infecting the root with a poisonous secretion that stops it from healing. This poison eventually kills the vine. This nymph reproduces by laying eggs for up to seven more generations (which also can reproduce parthenogenetically) each summer. These offspring spread to other roots of the vine, or to the roots of other vines through cracks in the soil. The generation of nymphs that hatch in the autumn hibernate in the roots and emerge next spring when the sap begins to rise. In humid areas, the nymphs develop into the ''winged form'', else they perform the same role without wings. These nymphs start the cycle again by either staying on the vine to lay male and female eggs on the bottom side of young grape leaves, or flying to an uninfected vine to do the same. Many attempts have been made to interrupt this life cycle to eradicate phylloxera, but it has proven to be extremely adaptable, as no one stage of the life cycle is solely dependent upon another for the propagation of the species.


Fighting the "phylloxera plague"

In the late 19th century the phylloxera epidemic destroyed most of the vineyards for
wine Wine is an alcoholic drink made from Fermentation in winemaking, fermented fruit. Yeast in winemaking, Yeast consumes the sugar in the fruit and converts it to ethanol and carbon dioxide, releasing heat in the process. Wine is most often made f ...
grape A grape is a fruit, botanically a berry, of the deciduous woody vines of the flowering plant genus ''Vitis''. Grapes are a non- climacteric type of fruit, generally occurring in clusters. The cultivation of grapes began approximately 8,0 ...
s in Europe, most notably in France. Phylloxera was introduced to
Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
when avid botanists in Victorian England collected specimens of American vines in the 1850s. Because phylloxera is native to North America, the native grape species are at least partially resistant. By contrast, the European wine grape ''
Vitis vinifera ''Vitis vinifera'', the common grape vine, is a species of flowering plant, native to the Mediterranean Basin, Mediterranean region, Central Europe, and southwestern Asia, from Morocco and Portugal north to southern Germany and east to northern ...
'' is very susceptible to the insect. The epidemic devastated vineyards in Britain and then moved to the European mainland, destroying most of the European grape growing industry. In 1863, the first vines began to deteriorate inexplicably in the southern Rhône region of France. The problem spread rapidly across the continent. In France alone, total wine production fell from 84.5 million
hectolitre The litre ( Commonwealth spelling) or liter (American spelling) (SI symbols L and l, other symbol used: ℓ) is a metric unit of volume. It is equal to 1 cubic decimetre (dm3), 1000 cubic centimetres (cm3) or 0.001 cubic metres (m3). A cub ...
s in 1875 to only 23.4 million hectolitres in 1889.winepros.com.au Some estimates hold that between two-thirds and nine-tenths of all European vineyards were destroyed. In France, one of the desperate measures of grape growers was to bury a live
toad Toad (also known as a hoptoad) is a common name for certain frogs, especially of the family Bufonidae, that are characterized by dry, leathery skin, short legs, and large bumps covering the parotoid glands. In popular culture (folk taxonomy ...
under each vine to draw out the "poison". Areas with
soil Soil, also commonly referred to as earth, is a mixture of organic matter, minerals, gases, water, and organisms that together support the life of plants and soil organisms. Some scientific definitions distinguish dirt from ''soil'' by re ...
s composed principally of sand or
schist Schist ( ) is a medium-grained metamorphic rock generally derived from fine-grained sedimentary rock, like shale. It shows pronounced ''schistosity'' (named for the rock). This means that the rock is composed of mineral grains easily seen with a l ...
were spared, and the spread was slowed in dry climates, but gradually phylloxera spread across the continent. A significant amount of research was devoted to finding a solution to the phylloxera problem, and two major solutions gradually emerged: grafting cuttings onto resistant rootstocks and hybridization.


Response

By the end of the 19th century, hybridization became a popular avenue of research for stopping phylloxera. Hybridization is the breeding of ''
Vitis vinifera ''Vitis vinifera'', the common grape vine, is a species of flowering plant, native to the Mediterranean Basin, Mediterranean region, Central Europe, and southwestern Asia, from Morocco and Portugal north to southern Germany and east to northern ...
'' with resistant species. Most native American grapes are naturally phylloxera resistant (''
Vitis aestivalis ''Vitis aestivalis'', the summer grape, or pigeon grape is a species of grape native to eastern North America from southern Ontario east to Maine, west to Oklahoma, and south to Florida and Texas. It is a vigorous vine, growing to 10 m or more hi ...
'', ''rupestris'', and ''riparia'' are particularly so, while ''
Vitis labrusca ''Vitis labrusca'', the fox grape, is a species of grapevines belonging to the ''Vitis'' genus in the flowering plant family Vitaceae. The vines are native to eastern North America and are the source of many grape cultivars, including Catawba, ...
'' has a somewhat weak resistance to it) but have
aroma An odor (American English) or odour (Commonwealth English; see spelling differences) is a smell or a scent caused by one or more volatilized chemical compounds generally found in low concentrations that humans and many animals can perceive v ...
s that are off-putting to palates accustomed to European grapes. The intent of the cross was to generate a hybrid vine that was resistant to phylloxera but produced wine that did not taste like the American grape. The hybrids tend not to be especially resistant to phylloxera, although they are much hardier with respect to climate and other vine diseases. The new hybrid varieties have never gained the popularity of the traditional ones. In the EU they are generally banned or at least strongly discouraged from use in quality wine, although they are still in widespread use in much of North America, such as Missouri, Ontario, and upstate New York.


Grafting with resistant rootstock

Use of a resistant, or tolerant, rootstock, developed by
Charles Valentine Riley Charles Valentine Riley (18 September 1843 – 14 September 1895) was a British-born American entomologist and artist. He was one of the first individuals to use biological pest control and authored over 2,400 publications. He convinced Congress ...
in collaboration with J. E. Planchon and promoted by T. V. Munson, involved grafting a ''
Vitis vinifera ''Vitis vinifera'', the common grape vine, is a species of flowering plant, native to the Mediterranean Basin, Mediterranean region, Central Europe, and southwestern Asia, from Morocco and Portugal north to southern Germany and east to northern ...
''
scion Scion may refer to: Horticulture *Scion (grafting), in horticulture, the upper part of a combined plant Arts, entertainment, and media Characters *Atlantean Scion, a device in the ''Tomb Raider'' video game series *Scion, avatar of the warrior en ...
onto the roots of a resistant ''
Vitis aestivalis ''Vitis aestivalis'', the summer grape, or pigeon grape is a species of grape native to eastern North America from southern Ontario east to Maine, west to Oklahoma, and south to Florida and Texas. It is a vigorous vine, growing to 10 m or more hi ...
'' or other American native species. This is the preferred method today, because the rootstock does not interfere with the development of the wine grapes (more technically, the
genes In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
responsible for the grapes are not in the rootstock but in the scion), and it furthermore allows the customization of the rootstock to soil and weather conditions, as well as desired vigor. Not all rootstocks are equally resistant. Between the 1960s and the 1980s in
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
, many growers used a rootstock called AxR1. Even though it had already failed in many parts of the world by the early twentieth century, it was thought to be resistant by growers in California. Although phylloxera initially did not feed heavily on AxR1 roots, within twenty years, mutation and selective pressures within the phylloxera population began to overcome this rootstock, resulting in the eventual failure of most vineyards planted on AxR1. The replanting of afflicted vineyards continues today. Many have suggested that this failure was predictable, as one parent of AxR1 is in fact a susceptible ''V. vinifera'' cultivar. But the transmission of phylloxera tolerance is more complex, as is demonstrated by the continued success of 41B, an
F1 hybrid F1 hybrid (also known as filial 1 hybrid) is the first filial generation of offspring of distinctly different parental types. F1 hybrids are used in genetics, and in selective breeding, where the term F1 crossbreed may be used. The term is somet ...
of ''
Vitis berlandieri ''Vitis berlandieri'' is a species of grape native to the southern North America, primarily Texas, New Mexico and Arkansas. It is primarily known for good tolerance against soils with a high content of lime, which can cause chlorosis in many vi ...
'' and ''
Vitis vinifera ''Vitis vinifera'', the common grape vine, is a species of flowering plant, native to the Mediterranean Basin, Mediterranean region, Central Europe, and southwestern Asia, from Morocco and Portugal north to southern Germany and east to northern ...
''. Modern phylloxera infestation also occurs when wineries are in need of fruit immediately, and choose to plant ungrafted vines rather than wait for grafted vines to be available. The use of resistant American rootstock to guard against phylloxera also brought about a debate that remains unsettled to this day: whether self-rooted vines produce better wine than those that are grafted. Of course, the argument is essentially irrelevant wherever phylloxera exists. Had American rootstock not been available and used, there would be no ''V. vinifera'' wine industry in Europe or most places other than Chile, Washington State, and most of Australia.
Cyprus Cyprus (), officially the Republic of Cyprus, is an island country in the eastern Mediterranean Sea. Situated in West Asia, its cultural identity and geopolitical orientation are overwhelmingly Southeast European. Cyprus is the List of isl ...
was spared by the phylloxera plague, and thus its wine stock has not been grafted for phylloxera resistant purposes.


Aftermath

The only European grapes that are natively resistant to phylloxera are the
Assyrtiko Assyrtiko or Asyrtiko is a white Greek wine grape indigenous to the island of Santorini. Assyrtiko is widely planted in the arid volcanic-ash-rich soil of Santorini and other Aegean islands, such as Paros. It is also found on other scattered r ...
grape which grows on the volcanic island of
Santorini Santorini (, ), officially Thira (, ) or Thera, is a Greek island in the southern Aegean Sea, about southeast from the mainland. It is the largest island of a small, circular archipelago formed by the Santorini caldera. It is the southern ...
, Greece, although it is not clear whether the resistance is due to the rootstock itself or the volcanic ash on which it grows; and the Juan Garcia grape variety, autochthonous to the medieval village of
Fermoselle Fermoselle is a small medieval village located in the province of Zamora, western Spain, and is part of the region of Castile and León in the south-west region of the province. It has a population of fewer than 1500. The village of Fermoselle d ...
in
Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...
. The Juan Garcia variety remained—untouched by phylloxera—sheltered on the vineyards planted on the man-made land terraces along the mountainous skirts on the gigantic and steep Duero Arribes / Douro River Canyon, where the microclimatic conditions discourage the growth of phylloxera. To escape the threat of phylloxera, wines have been produced since 1979 on the sandy beaches of Provence's
Bouches-du-Rhône Bouches-du-Rhône ( ; , ; ; "the Mouths of the Rhône") is a Departments of France, department in southern France. It borders Vaucluse to the north, Gard to the west and Var (department), Var to the east. The Mediterranean Sea lies to the sout ...
, which extends from the coastline of the
Gard Gard () is a department in Southern France, located in the region of Occitanie. It had a population of 748,437 as of 2019;Saintes Maries de la Mer. The sand, sun and wind in this area has been a major deterrent to phylloxera. The wine produced here is called "Vins des Sables" or "wines of the sands". In the same department, where the canal irrigation system built by the Romans still partly persists to this day, winter flooding is also practiced where possible, for instance south of the city of
Tarascon Tarascon (; ), sometimes referred to as Tarascon-sur-Rhône, is a commune situated at the extreme west of the Bouches-du-Rhône department of France in the Provence-Alpes-Côte d'Azur region. Inhabitants are referred to as Tarasconnais or Tara ...
. Flooding the vineyards for 50 days kills all the nymphs that overwinter in the roots or the bark at the bottom of the plant. Some regions were so blighted by phylloxera that they never recovered, and instead the producers switched crops entirely. The island of
Mallorca Mallorca, or Majorca, is the largest of the Balearic Islands, which are part of Spain, and the List of islands in the Mediterranean#By area, seventh largest island in the Mediterranean Sea. The capital of the island, Palma, Majorca, Palma, i ...
is one example, where almonds now substitute for vines.


Vines that survived phylloxera

According to wine critic and author
Kerin O'Keefe Kerin O'Keefe is a wine critic specialized in Italian wine and author of four books. She reviews wines and writes articles on the growing areas, wines and producers for kerinokeefe.com. O'Keefe was Italian Editor for Wine Enthusiast from May 2 ...
, thanks to tiny parcels of vineyards throughout Europe which were inexplicably unscathed, some vineyards still exist as they were before the phylloxera devastation. So far, most
Chilean wine Chile has a long history in the production of wine, with roots dating back to the 16th century when the Spanish conquistadors introduced ''Vitis vinifera'' vines to the region. In the mid-19th century, French wine variety (botany), varieties s ...
has remained phylloxera free. It is isolated from the rest of the world by the
Atacama Desert The Atacama Desert () is a desert plateau located on the Pacific Ocean, Pacific coast of South America, in the north of Chile. Stretching over a strip of land west of the Andes Mountains, it covers an area of , which increases to if the barre ...
to the north, the Pacific Ocean to the west, and the
Andes Mountains The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long and wide (widest between 18°S ...
to the east. Phylloxera has also never been found in several wine-growing regions of Australia, including Tasmania, Western Australia and South Australia. Australia has strong internal biosecurity controls to minimise the risk of phylloxera spreading beyond 'Phylloxera Infested Zones' in parts of Victoria and New South Wales. The
Riesling Riesling ( , ) is a white grape variety that originated in the Rhine region. Riesling is an aromatic grape variety displaying flowery, almost perfumed, aromas as well as high acidity. It is used to make dry, semi-sweet, sweet, and sparkling ...
of the Mosel region has also remained untouched by phylloxera; the parasite is unable to survive in the slate soil. Until 2005, three tiny parcels of ungrafted
Pinot noir Pinot noir (), also known as Pinot nero, is a red-wine grape variety of the species ''Vitis vinifera''. The name also refers to wines created predominantly from Pinot noir grapes. The name is derived from the French language, French words fo ...
that escaped phylloxera were used to produce
Bollinger Bollinger () is a French Champagne house, a producer of luxury sparkling wines from the Champagne (wine region), Champagne region. They produce several labels of Champagne (wine), Champagne under the Bollinger name, including the vintage ''Vieil ...
Vieilles Vignes Françaises, one of the rarest and most expensive Champagnes available. In 2004, one of the parcels, Croix Rouge in
Bouzy Bouzy () is a commune of the Marne department in northeastern France, the Montagne de Reims subregion of Champagne. Population Champagne The village's vineyards are located in the Montagne de Reims subregion of Champagne, and are classified ...
, died from phylloxera and was replanted with grafted rootstock. A rare vintage
port A port is a maritime facility comprising one or more wharves or loading areas, where ships load and discharge cargo and passengers. Although usually situated on a sea coast or estuary, ports can also be found far inland, such as Hamburg, Manch ...
is made from ungrafted vines grown on a small parcel, called Nacional, in the heart of the Quinta do Noval. It is unknown why this plot survived. Another vineyard unaffected by the phylloxera is the Lisini estate in
Montalcino Montalcino is a hill town and ''comune'' in the province of Siena, Tuscany, central Italy. The town is located to the west of Pienza, close to the Crete Senesi in Val d'Orcia. It is from Siena, from Florence and from Pisa. Monte Amiata is l ...
in Italy, a half-hectare vineyard of Sangiovese with vines dating back to the mid-1800s. Since 1985, the winery has produced a few bottles of Prefillossero (Italian for "before the phylloxera"). The wine has a following, including Italian wine critic Luigi Veronelli who inscribed on a bottle of the 1987 at the winery that drinking Prefillossero was like listening to 'the earth singing to the sky'. Jumilla (DO), Jumilla in southeastern Spain is an important area of ungrafted vineyards, mainly from Monastrell grapes. Those vineyards, however, are not immune to the insect, which is slowly advancing and destroying the Pie Franco vineyard of the Casa Castillo estate, planted in 1942, i.e., when phylloxera had already been in the region for five decades. Large swaths of vineyards on the slopes of Sicily's volcano Mount Etna also remain free of phylloxera. Some vines are more than one-hundred-fifty years old, predating the phylloxera infestation in Sicily (1879–1880). Part of the reason for this is the high concentration of silica sand and very low (less than 3%) concentration of clay in the volcanic soils. In this environment (> 400m AMSL), the surface water from heavy bouts of rain seals the soil so perfectly that it drowns phylloxera before they are able to thrive.Campbell, Christy. "Phylloxera: How Wine Was Saved For the World." Harper Perennial, 2004, p. 129–130 Bien Nacido Vineyards, Bien Nacido Vineyard in Santa Maria Valley AVA of Santa Barbara, CA is a phylloxera-free vineyard. Despite being planted on its own roots, with UC Davis virus-free clones, the vineyard has never been affected by phylloxera. The high percentage of sand in the soil creates a mostly uninhabitable substrate for phylloxera. While Bien Nacido has not been affected, there is a potential, as all of the vines are true ''Vitis vinifera'' without scions or grafting. Many of the old vines were planted in 1973 and fall within the blocks G, N, Q and W. The wines of Bien Nacido Estate have a high percentage of these ungrafted and phylloxera-free vines within the cuvée. Colares DOC, Colares vineyards, native to the Portugal, Portuguese region of Sintra are grown on 3–4 metres of sand, and are therefore unaffected by phylloxera.


References


Further reading

* Boubals, Denis, "Sur les attaques de Phylloxera des racines dans le monde", Progres Agricole et Viticole, Montpellier, 110:416-421, 1993. * Campbell, Christy, "The Botanist and the Vintner: How Wine Was Saved for the World", Algonquin Books, 2005. * Ordish, George, "The Great Wine Blight", Pan Macmillan, 1987. * Powell, Kevin, "Grape phylloxera: An Overview". In Root feeders An Ecosystem perspective (Eds S.N. Johnson & P.J. Murray) CAB International 2008. * Benheim, Devin et al., "Grape phylloxera (Daktulosphaira vitifoliae) – a review of potential detection and alternative management options", Annals of Applied Biology, Volume 161, Issue 2, pages 91–115, September 2012


External links

* *
Web page of the Phylloxera and Grape Industry Board of South Australia

The Vine's Enemy
A profile of phylloxera drawn from the 2nd edition of ''The Oxford Companion to Wine'' {{Authority control Phylloxeridae Grape pest insects Gall-inducing insects Insects in culture Monotypic Hemiptera genera