Grape Phylloxera

Daktulosphaira vitifoliae FITCH

Family: Phylloxeridae

Order: Hemiptera

Other Names:

Phylloxera vastatrix Planchon, Phylloxera vitifoliae, Viteus vitifolii Shimer, Viteus vitifoliae Fitch, Dactylosphaera vitifolii Shimer, Peritymbia (Phylloxera) vitifolii pervastatrix C. B. vastatrix Planchon and Peritymbia pervastatrix Borner
Grapevine phylloxera
Filoxera
Vine louse
Reblaus
Vine fretter

Summary:

Grape phylloxera (Daktulosphaira vitifoliae Fitch) is a small, invasive, sap-sucking insect of grape vines that causes substantial physical and economic effects on commercial grapevines. It is a monophagous and sedentary gall forming phylloxerid that establishes an intimate association with the roots of grape vines (Powell et al., 2013; Forneck and Huber, 2009; Granett et al., 2001). The insect forms organoid galls on fibrous root tips (nodosities) and callus tissue on mature roots (tuberosities).
Nodosities serve as the insect's nutritional basis and represent the exclusive feeding site required for growth and development (Griesser et al., 2015a; Du et al., 2008; Kellow et al., 2004).

Description:

Almost microscopic, related to the aphids.
Five distinct forms of adults.
The amphigonic individuals of both sexes do not feed and have no rostrum.
The virginoparous forms have a rostrum but their hind gut is functionless and the anal orifice is closed
Adult -
Colour - Pale yellow.
Body - The root infesting form (“radicicoles”) is 0.8-1.2 mm long x 0.5-0.9 mm wide, sub oval to pear shaped and with many hair-bearing dorsal tubercles. It is yellowish green to yellow.
The leaf-infesting form (“gallicoles”) is 1-1.25 mm long x 1 mm wide, has a similar colour but without dorsal tubercles. It is greenish yellow.
The alate form (“sexupara”) is orange-yellow with a darker mesothorax, body length about 1.2 mm. It has hyaline wings more or less tinted.
Wings - Hyaline, nearly tinted, horizontal in repose on alate forms. Other forms are wingless.
Mouthparts - Stylet sucking
Antennae - 3-4 joints on winged forms.
Legs - 6
Head -
Thorax -
Abdomen -
Habits -
Caterpillar -
Has nymphs and crawlers.

Key Characters:

Forms galls on leaves and roots of grape vines.

Biology:

Reproduces by viviparous parthenogenesis. Completing a generation on susceptible cultivars requires 16 days at 24ºC and longer on resistant cultivars of grape vine.
The threshold for development was calculated to be at 9.3ºC. Generation time could require one month, and each female produces about 50-70 eggs.

Life Cycle:

Its life cycle has sexual and asexual portions with forms that feed from leaf and root galls. Not all forms occur throughout the insect's range. Root forms predominate on Vitis vinifera cultivars; leaf forms predominate on other Vitis species characteristic of the American native range. Damage to grapevines is by secondary soil borne pathogens attacking the feeding site and by physiological interaction of the insect with the grapevine (Granett et al, 2001).

Figure 1 The life cycle of grape phylloxera. Asexuals occur on Vitis leaves and roots. Individuals forming galls on leaves are called gallicoles, and on roots, radicicoles. The root galls are termed tuberosities if they occur on mature, suberised roots; they are called nodosities when occurring near root tips. The eggs hatch into first instars, crawlers, that are mobile and can move between
roots and leaves to establish new feeding sites. After the first instar, gallicoles and radicicoles tend to feed in one place. There are four nymphal instars prior to the wingless adult stage. Alates are generated on roots (R), have four nymphal instars, and emerge from the ground prior to adult eclosion. After eclosion these winged forms do not feed but disperse and then lay male and female eggs asexually. The newly hatched sexuals moult four times into wingless adults without feeding; they mate, and each female lays a single overwintering egg. The fundatrix for the succeeding year's gallicoles hatches from the overwintering egg. An alternative route to the sexual forms occurs
in the south western and, to a limited extent, in the south eastern United States, with leaf feeding adults laying male or female eggs in leaf galls (L) without the intermediary alate. Overwintering of radicicoles occurs as first instar hibernants.
In order for phylloxera to successfully feed on a Vitis species, the plant must produce a leaf gall or a homologous root gall. Feeding activity, the making of the feeding wound and injection of saliva or gut contents, induces formation of the gall. Gallicoles (Figure 1) induce formation of galls on newly expanding leave, shoots, and tendrils; galls do not form on mature leaves. Galls
extend below the leaf surface and fully enclose the feeding gallicoles. The gall
opens to the upper leaf surface allowing crawlers to exit. Guard hairs at the opening restrict the entryway, presumably reducing predation and maintaining humidity. Since galls persist, canes record phylloxera activity seasonally. Some crawlers may remain within the mother's gall to produce a second generation, but most disperse. Outside they may attack newly forming leaves on the same or close-by canes, disperse in the wind with the chance of landing on another suitable host, or fall to the ground and attack available roots. The proportion of crawlers that survive dispersal is unknown but is probably small. In some areas non-winged adults in leaf galls may produce sexual offspring. (Granett et al, 2001).
Radicicoles live on root swellings that are homologous to leaf galls.
Tuberosities appear as localized enlargements and are the result of proliferation and expansion of phloem parenchyma cells. Nodosities are enlarged, fleshy growths that cause rootlets to bend at the point of feeding. The radicicole crawlers are able to move from root to root searching for a suitable feeding place but have difficulty infesting leaves. Crawlers disperse over the soil surface and onto vine foliage. The downwind pattern in the spread of phylloxera-induced damage within and between vineyards suggests that this dispersal results in successful establishment on roots. Distance of this movement and survival must be low but has not been quantified. Floodwaters may also move phylloxera between vineyards. Radicicole adults and nymphs, except for the crawlers, tend not to move. (Granett et al, 2001).
Radicicoles can give rise to wing pad-bearing nymphs (in the literature these
individuals are sometimes called larvae). Crowding or degraded root quality may stimulate development of nymphal stages of the alates, as may high soil moisture. Alate adults are not strong flyers and, without feeding, asexually oviposit a small number of eggs on a woody portion of the grape vine. These eggs are of two sizes and hatch into sexual males or females. The sexuals do not have complete mouthparts and develop without feeding. Sexual nymphs are immobile pupiform individuals. Upon becoming adults, the wingless sexuals mate and the female lays the overwintering egg. The fundatrix hatching from the overwintering egg in the spring is capable of founding new colonies by inducing formation of a leaf or root gall. Survival rates for overwintering eggs are not known. (Granett et al, 2001).
Leaf galling of Vitis vinifera is commonly absent but does occur in Australia (Froggat, 1922).
Australian strains of phylloxera in Rutherglen, Victoria, are able to induce leaf galling and are genetically distinct from neighbouring phylloxera strains that lack this capacity (Corrie et al, 1998).
Optimal temperatures for insect survival and gall formation are between 22 and 260C. In this range, the egg stage lasts about 6 days, and if eggs are maintained in a moist condition, more than 90% hatch (Granett and Timper, 1987).
On mature, suberised roots in vitro, crawlers may take up to 2 weeks to induce a tuberosity. They are successful 20 to 80% of the time. If moisture is optimized within the rearing chamber, where the roots and phylloxera are held, survival increases. Once a tuberosity is established, the insect progresses through its instars rapidly, reaching adulthood in less than 1 week. Under optimal conditions, adults on excised roots survive 1-2 months with oviposition of between 2 and 10 eggs per female per day, more at the beginning of adult life and fewer at the end. Radicicoles tend to develop
more rapidly on nodosities than on tuberosities; however, duration of the
adult stage on nodosities is short lived owing to rapid rootlet degradation. The
insect's lower temperature threshold for gall formation is about 180C, although
development can occur at temperatures as low as 50C if galls are present. The upper threshold for radicicoles is about 280C, whereas the upper threshold for gallicoles may be higher.
Grape phylloxera are unable to initiate feeding sites or develop beyond hatching at soil temperatures lower than 15-18?C (Turley et al., 1996).
Gall formation is critical to radicicole survival and in general, resistant vines suppress tuberosity formation in the field.
Radicicole population size in the field varies over the growing season and has
myriad direct and indirect influences. Populations overwinter in small numbers as first instar hibernants that are arrested in their development by low temperatures. As temperatures become favourable in the spring, the population increases exponentially, initially on new roots and then spreading to mature roots. In California, the population peaks in mid summer and then declines; a second population peak occurs after harvest (Omer et al, 1997).
Survival rates of the overwintering forms, hibernants, and the overwintering
egg have not been studied, but numbers decrease dramatically during the winter. Alates determine the site for overwintering eggs (Granett et al, 2001).
Radicicoles cause more vine damage than gallicoles. Whereas radicicoles can cause severe vine decline or death, gallicole damage is limited to decreases in cane growth and quality. Damage is first seen as decreased cane growth followed by faded-leaf and potassium-deficiency symptoms in midsummer when vines are heat stressed. Over time, the root system collapses. Death of vines in a newly infested vineyard block is initially seen in epicentres of as few as ten vines (151). From epicentres, vine decline and death expand radially, the number of vines affected increasing at a two to
tenfold rate per year. As epicentres enlarge, newly damaged vines not contiguous with the original focus develop downwind. Population levels are highest at the periphery of the epicentres, the vines in the centre having much reduced and necrotic root systems with few remaining phylloxera. In heavy clay soils with moderate temperatures, vine decline tends to be very rapid; it is slower in very hot or cold climates and in sandy soils. Populations spread at
a faster rate than vines decline. A vineyard block with a relatively small number of symptomatic vines may have infestations on the majority of vines. The economic threshold is low but unknown. (Granett et al, 2001)
Damage may be caused by:
(a) Removal of photosynthates may cause loss of vine vigour but is unlikely as the biomass of grape phylloxera on root systems is small, with damaging populations averaging fewer than 100 individuals per gram of dry root weight;
(b) Root mortality caused by secondary pathogens entering feeding wounds is most likely and can cause water and nutrient stresses leading to eventual vine death. Fusarium oxysporum Schlecht was found to be the pathogen most often involved;
(c) Physiological disruption other than through direct removal of photosynthates or water stress can also occur (Granett et al, 2001)
Unlike aphids, phylloxera are clean feeders, producing no honeydew or obvious, externally excreted waste products (Granett et al, 2001).
Damage by gallicoles is not simply the result of ingestion of photosynthate from infested leaves. They may consume about 2% of the photosynthate produced but more importantly reduce photosynthesis of the leaf by up to 50% and the leaf does not recover, even after the gallicoles have died. Some affected leaves may drop. Early infestations (bloom to two weeks post bloom) have a more severe effect on fruit quality and yield than do later infestations but economic thresholds have not been determined (Granett et al, 2001).
Breeders focused on phylloxera-resistant rootstocks upon which Vitis vinifera cultivars could be grafted because production of hybrids has not been very successful. Cultivars bred and selected prior to 1930 constitute a large
percentage of the rootstocks used today and their use where phylloxera is a threat is almost universal (Granett et al, 2001).
Resistant rootstocks rarely eradicate a phylloxera infestation so the site remains a potential source of infection for clean vineyards.
Phylloxera can live on grape roots several meters deep in the soil (quoted in Granett et al, 2001).
Phylloxera has potential for rapid population growth. Generation time may be less than a month, giving vineyards three to ten generations per year and gross reproductive rate of 50 eggs per female has been noted. Population
regeneration by survivors of insecticide treatments may be rapid.
Grape phylloxera, due to their compartmentalised gut structure and limited ability to excrete waste (Andrews et al., 2012), can survive without food under suitable ambient conditions and even when immersed in water for several days (Korosi et al., 2009).

Habitats:

Vines grown in sandy soils do not support damaging phylloxera populations. This has been attributed to the inability of phylloxera crawlers to disperse from infested to uninfested roots because of sand texture. Phylloxera populations in clay soils tends to be large (Granett et al, 2001).

Soil

Characteristics such as low aluminium exchange capacity and acidic pH are associated with high grape phylloxera abundance above and below ground in commercial vineyards in Victoria, Australia (Powell et al., 2003).
They are much worse on cracking soils as the radicicole form can move through the cracks to infest new roots. It is less of a problem on sandy soils.

Origin and History:

Described in 1855 from native American Vitis (Granett et al, 2001).
Imported into Europe before the 1860s where it devastated the European grape vineyards initially in France from 1868 destroying over 1 million ha of ungrafted vineyards by the turn of the century and then spread across the world (Benheim et al, 2012; Granett et al, 2001).
Native to North America and possibly down to Venezuela.
Discovered in Australia in 1877 (Buchanan, 1990).

Distribution:

It is a worldwide pest of grapevines.
Only 2%of all Australian vineyards are known to be infested with grape phylloxera (Nicol et al., 1999)

Courtesy https://www.gbif.org/species/2049512

Significance:

Historically, one of the most economically significant insect pests of vineyards has been grape phylloxera.

Beneficial:


Detrimental:

The California failure of the AXR#1 rootstock resulted in an estimated billion-dollar loss to growers. Phylloxera has still not infested a number of important viticultural regions around the world. In these regions phylloxera-resistant rootstocks are not used and when phylloxera arrives, the billion-dollar loss experienced in California with AXR#1's failure will seem modest. Alternative and supplemental control methods are needed to back up rootstock use and prevent such losses in the future (Granett et al, 2001).
An estimated cost of replanting and lost production in Australia is
AUS$20000/ha (DAFWA, 2006)

Legislation:

Australian state legislation requires the declaration of a new grape phylloxera infested zone (PIZ) with a minimum 5 km boundary from the initial detection point.
The disinfestation of viticultural machinery, such as grape harvesters, requires low humidity heating at 450C for a minimum of 75 minutes or 400C for 2 hours (Korosi et al., 2009, 2012; NVHSC, 2009)
Heat treatment of soil samples for diagnostic purposes is also recommended where samples are dispatched from a PIZ to a processing laboratory for general diagnostic testing (NVHSC, 2009).

Management and Control:

Tolerant grape vine (Vitis spp.) rootstocks form organoid root galls on fibrous root tips to protect the plant (Eitle et al, 2019).
Resistant rootstocks derived from native American Vitis species are the primary control tool. Resistant rootstocks have controlled grape phylloxera for more than 120 years.
Flooding of vineyards for 40-50 days during the winter months has been shown to limit phylloxera populations (Riley, 1874). Grape phylloxera first instars and eggs have been shown to survive for up to 8 days when submersed in water and the treatment is more effective at low temperatures (Korosi et al., 2009).
Dipping footwear and shovels in sodium hypochlorite solution to kill phylloxera is not completely effective; eggs survived 7 minute dips in undiluted commercial bleach (Grzegorczyk and Walker, 1997).
Grape phylloxera is unable to survive 4 days of composting of green waste and winery waste when conducted using commercial standards with temperature being the predominant factor influencing mortality (Keen et al, 2002).
Fumigation of table grapes with sulphur dioxide (Buchanan, 1990) has also achieved 100% grape phylloxera mortality.
Handheld viticultural equipment and footwear can be disinfested with a 2% NaOCl solution for a minimum of 30 seconds.
Gamma irradiation of grapevine plant material as a disinfestation procedure is reportedly an effective but relatively slow process, with treated organisms taking weeks to reach 100% mortality (Witt & Van de Vrie, 1985).
Use of insecticides is limited in effect, and other control methods are not proven (Granett et al, 2001).
In the past it has been ameliorated to a very limited extent by the use of chemicals such as carbon bisulfide (Pouget R., 1990).
Insecticide treatments (Stevenson, 1962) have been approved for treatment of plant material before transport, as have hot water dips at 52-540C for 3-5 min (Flaherty et al, 1992).
Grape vine cuttings will tolerate 450C for 24 hours, 500C for 2.5 hours and 550C for 10 minutes (Goheen et al, 1973; Gramaje et al, 2009)
Insecticides have been effective for controlling gallicoles on the leaves but have generally not been effective on radicicoles on the roots. Recently a downwardly mobile insecticides (thiamethoxam and imidacloprid) has become available for field-testing against phylloxera and shows efficacy against gallicoles and radicicoles. It is available in Australia but is no registered for use in grape vines.
Spirotetramat has been registered for use against grape phylloxera in Canada.
Imidacloprid, acetamiprid, fenopropathrin and spirotetramat are currently registered against the foliar form of grape phylloxera in the USA.
Systemic insecticides, such as imidacloprid and spirotetramat, have shown suppression of grape phylloxera in laboratory, glasshouse and field based trials (Herbert et al., 2008a).
Fenamiphos and disulfoton have some action but are under review to limit continued registration.
Aldicarb provided good control (Loubser et al, 1992) but is no longer available in Australia.
The US EPA has registered soil application (dinotefuran and imidacloprid) or a foliage application (acetamiprid, endosulfan, fenpropathrin, spirotetramat, thiamethoxam and a mixture of thiamethoxam and chlorantraniliprole).
Chemicals can stimulate induced host plant resistance. Foliar applications of
jasmonic acid reduce phylloxera populations on roots of greenhouse vines (Omer et al 2000).
Quarantine regulations have been coupled with eradication procedures in Australia (Buchanan GA and Amos, 1984).
Grape phylloxera management in Australia has therefore evolved into an integrative approach consisting of: (a) early detection and surveillance;
(b) extensive quarantine regulations which encompass disinfestation procedures for plant material, machinery, hand-held equipment and footwear aimed at preventing the spread of grape phylloxera outside of designated
grape phylloxera infested zones (PIZ's); and
(c) the use of grape phylloxera resistant rootstocks.
No biological control programs have been implemented.

Thresholds:

The detection of leaf-galling grape phylloxera strains is evident by visual inspection during the spring and summer.
Because of its predominantly subterranean habitat and relatively high economic damage once an incursion occurs, several different approaches for
improved detection of root-galling grape phylloxera strains have been explored and are currently under development (Bruce et al., 2011a).
Typically it is isolated to only a few vines, principally expressed as a
gradual decline in canopy vigour followed by premature yellowing of foliage and an incremental reduction in grape yield. These symptoms alone are not strong indications of grape phylloxera infestation as some grapevine phytoplasma diseases such as flavescence doree and bois noir, and field conditions such as dehydration and sustained high temperatures, cause similar symptoms. The infestation eventually leads to reduced functional root mass, canopy decline, reduced crop yield, vine death and occurrence of satellite spots throughout the infested vineyard as a result of spread by machinery, wind or human traffic.
Conventional detection of grape phylloxera infestation can use manual ground surveys either alone or in combination with some form of remote aerial imaging to assess canopy decline and rate of spread.
Conventional detection of grape phylloxera involves manual excavation and visual inspection of the grapevine root system for the presence of grape phylloxera and associated galls.
Emergence, trunk and pitfall traps are also used to detect infestations.
Hyperspectral, multispectral and infra-red (NDVI) have been used to detect infestations with varying success.
Metabolomic methods are being increasingly applied.
Molecular methods for grape phylloxera detection have been explored (Herbert et al., 2008b) resulting in the development of a commercially available grape phylloxera-specific DNA soil probe.
Sleezer et al 2010 has a model for foliar grape phylloxera and table grape cultivars (Vitis labrusca L.) are not attacked by foliar phylloxera.

Related Species:

It is a single species in this genera.

Similar Species:

References:

CSIRO. The Insects of Australia. Melbourne University Press. (1991)

Andrews K.B., Kemper D., Powell K.S., Cooper P.D. (2012) Spatial trade-offs in the digestive and reproductive systems of grape phylloxera. Australian Journal of Zoology, 59, 392-399.

Avidov, Z. and Harpaz, I. (1969) Plant Pest of Israel. Israel University Press. P140-142.

Bruce R.J., Hoffmann A.A., Runting J., Lamb D., Powell K.S. (2011) Towards improved early detection of grapevine phylloxera (Daktulosphaira vitifoliae Fitch) using a risk based assessment. Acta Horticulturae, 904, 123-132.

Buchanan G.A. (1990) The distribution, biology and control of grape phylloxera, Daktulosphaira vitifolii (Fitch), in Victoria. PhD Thesis, La Trobe University, Melbourne, 177 pp.

Buchanan GA, Amos TG, eds. 1984. The biology, quarantine and control of grape phylloxera in Australia and New Zealand. Proc. Standing Committee Agriculture Workshop, Rutherglen Res. Inst. Rutherglen, Vic., Dep. Agric., 78 pp.

Corrie AM, Buchanan GA, Van Heeswijck R. 1998. DNA typing of populations of phylloxera [Daktulosphaira vitifoliae (Fitch)] from Australian vineyards. Austr. J. Grape Wine Res. 3:50-56.

DAFWA (2006) Fact Sheet: Grape phylloxera Daktulosphaira vitifoliae - Exotic threat to Western Australia [WWW Document]. Department of Agriculture and Food, Western Australia. URL http://www.agric.wa.gov.au/objtwr/imported_assets/content/pw/ins/pp/hort/fs00200.pdf

Flaherty DL, Christensen LP, Lanini WT, Marois JJ, Phillips PA, et al, eds. (1992). Grape Pest Management. Oakland, CA: Div. Agric. Natl. Res., Univ. Calif. 400 pp. 2nd ed.

Froggat WW. (1922). Leaf galls of phylloxera at Howlong. Agric. Gaz. NSW xxxiii:360.

Goheen A.C., Nyland G., Lowe S.K. (1973). Association of a Rickettsialike organism with Pierce's disease of grapevines and alfalfa dwarf and heat therapy of the disease in grapevines. Phytopathology 63, 341-345.

Gramaje D., Armengol J., Salazar D., Lo´ pez-Corte´ L., Garcýa-Jimenez J. (2009). Effect of hot-water treatments above 500C on grapevine viability and survival of Petri disease pathogens. Crop Protection 28 (2009) 280-285.

Granett J., Walker M., Kocsis L. and Omer, A.D. (2001).Biology and Management of Grape Phylloxera. Annu. Rev. Entomol. 2001. 46:387-412.

Granett J, Timper P. 1987. Demography of grape phylloxera (Daktulosphaira vitifoliae) (Homoptera: Phylloxeridae). J. Econ. Entomol. 80:327-29

Grzegorczyk W, Walker MA. 1997. Surface sterilization of grape phylloxera eggs in preparation for in vitro culture with Vitis species. Am. J. Enol. Vitic. 48:157-59.

Herbert K.S., Hoffmann A.A., Powell K.S. (2008a) Assaying the potential benefits of thiamethoxam and imidacloprid for phylloxera suppression and improvements to grapevine vigour. Crop Protection, 27, 1229- 1236.

Herbert K.S., Powell K.S., McKay A., Hartley D., Herdina, Ophel-Keller K., Schiffer M., Hoffmann A.A. (2008b) Developing and testing a diagnostic probe for grape phylloxera applicable to soil samples. Journal of Economic Entomology, 101, 1934-1943.

Keen B.P., Bishop A.L., Gibson T.S., Spohr L.J., Wong P.T.W. (2002) Phylloxera mortality and temperature profiles in compost. Australian Journal of Grape and Wine Research, 8, 56-61.

Korosi G.A., Trethowan C.J., Powell K.S. (2009) Reducing the risk of phylloxera transfer on viticultural waste and machinery. Acta Horticulturae, 816, 53-62.

Korosi G.A., Mee P.T., Powell K.S. (2012) Influence of temperature and humidity on mortality of grapevine phylloxera Daktulosphaira vitifoliae clonal lineages - a scientific validation of a disinfestation procedure for viticultural machinery. Australian Journal of Grape and Wine Research, 18, 43-47.

Loubser J.T., van Aarde I.M.F. and Hoppner G.F.J. (1992) Assessing the Control Potential of Aldicarb against Grapevine Phylloxera. Nietvoorbij Institute for Viticulture and Oenology (Nietvoorbij), Private Bag X5026, 7599 Stellenbosch, Republic of South Africa.

Nicol J.M., Stirling G.R., Rose B.J., May P., van Heeswijck R. (1999) Impact of nematodes on grapevine growth and productivity: current knowledge and future directions, with special reference to Australian viticulture. Australian Journal of Grape and Wine Research, 5, 109-127.

NVHSC (2009) National Phylloxera Management Protocol [WWW Document]. National Vine Health Steering Committee, Australia. URL http://www.gwrdc.com.au/webdata/resources/files/NationalProtocolCombinedv 2009_4_endorsed_forweb.pdf

Omer AD, Granett J, Downie DA,Walker MA. 1997. Population dynamics of grape phylloxera in California vineyards. Vitis 36:199-205

Omer AD, Thaler JS, Granett J, Karban R. (2000). Jasmonic acid induced resistance in grapes to a root and leaf feeder. J. Econ. Entomol. 93:840-45

Pouget R. 1990. Histoire de la Lutte Contre le Phyllox´era de la Vigne en France (1868-1895). Paris: Inst. Natl. Rech. Agron. 157 pp.

Powell K.S., Slattery W.J., Deretic J., Herbert K., Hetherington S. (2003) Influence of soil type and climate on the population dynamics of grapevine phylloxera in Australia. Acta Horticulturae, 617, 33-41.

Riley CV. 1874. The grape phylloxera Phylloxera vastatrix Planchon. In 6th Annu. Rep. Noxious, Beneficial, and Other Insects of the State of Missouri, pp. 30-65. 9th Annu. Rep. State Board Agric. State of Missouri. Jefferson City, MO: Regan & Carter.

Sleezer S., Johnson D.T., Lewis B. and Goggin F. (2010). Foliar Grape Phylloxera, Daktulosphaira vitifoliae (Fitch), Seasonal Biology, Predictive Model and Management in the Ozarks Region of the United States. Proc. 5th International Phylloxera Symposium Eds.: M. Griesser and A. Forneck. Acta Hort. 904, ISHS 2011

Stevenson AB. 1962. Insecticide dips to control grape phylloxera on nursery stock. J. Econ. Entomol. 55:804-5

Turley M., Granett J., Omer A.D., De Benedictis J. (1996) Grape phylloxera (Homoptera: Phylloxeridae) temperature threshold for establishment of feeding sites and degree-day calculations. Environmental Entomology, 25, 842-847.

Witt A.K.H., Van de Vrie M. (1985) Gamma radiation for post-harvest control of insects and mites in cut flowers. Mededelingen van de Faculteit Landbouwwetenschappen, Rijksuniversiteit Gent, 50, 697-704.

Acknowledgments:

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Appendix



Table 1 Summary of a range of insecticides used in phylloxera control trials
Compound ClassActive Ingredient(Trade name) Trial Location Trial Type Phylloxera Type Selected Sources
Carbon disulphide France Field Radicicolae Ordish, 1972
Sulphocarbonates France Field Radicicolae Ordish, 1972; Campbell, 2004
Enzone USA Field Radicicolae R. Loveless (as cited by Herbert, 2005); Weber
et al., 1996
Carbamates and
Organophosphates  Carbofuran USA, Australia Field and Laboratory Radicicolae Rammer, 1980; Granett & Timper, 1987;Buchanan, 1990
Fenamiphos Germany, USA and AustraliaField Radicicolae Homeyer & Wagner, 1981; Buchanan, 1990; de Klerk, 1979
Phosphorothioic acid Canada Field Radicicolae Stevenson, 1968
Baygono-isopropoxyphenyl methylcarbamateCanada Field Radicicolae Stevenson, 1968
Disulfoton South Africa and Canada Field Radicicolae Stevenson, 1968; de Klerk, 1979
Oxamyl Australia Field Radicicolae Buchanan & Godden, 1989; Nazer et al ., 2006
Aldicarb Australia Field Radicicolae Buchanan & Godden, 1989; Loubser et al.,1992
Organochlorines   Hexachlorobutadiene South Africa Field Radicicolae de Klerk, 1979
Hexachlorocyclopentadiene USA Field and Laboratory Radicicolae Cox et al., 1960
Endosulfan USA and Canada Field Gallicolae Stevenson, 1970; Williams, 1979
Neonicotinoids   Thiamethoxam Australia and USA Laboratory Radicicolae Granett et al., 2001; Nazer et al., 2006; Herbert et al., 2008a,b
Imidacloprid South Africa, Jordan, USA and Australia Field and Laboratory Radicicolae and Gallicolae C. Coetzee & R. Loveless (as cited by Herbert,2005); Herbert et al., 2008; Nazer et al.,2006; Al-Antary et al., 2008
Spirotetramat USA Field Gallicolae Nauen et al., 2008; van Steenwyk et al., 2009;Johnson et al. 2010