- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Pathway Causes
- Pathway Vectors
- Plant Trade
- Impact Summary
- Economic Impact
- Environmental Impact
- Risk and Impact Factors
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Gaps in Knowledge/Research Needs
- Links to Websites
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Amylostereum areolatum (Chaillet ex Fr.) Boidin 1958
Other Scientific Names
- Lloydellopsis areolata (Chaillet ex Fr.) Pouzar 1959
- Stereum areolatum (Chaillet ex Fr.) Fr. 1838
- Thelephora areolata Chaillet ex Fr. 1828
- Xylobolus areolatus (Chaillet ex Fr.) P. Karst 1881
Summary of InvasivenessTop of page
A. areolatum is a basidiomycete that causes a white rot of a broad range of conifers. Its invasiveness arises from a symbiotic association with woodwasps of the genus Sirex. The species Sirexnoctilio is listed as “highly invasive” on the ISSG/IUCN website (ISSG, 2008) and is a Regulated Pest for the USA (APHIS, 2009a). The wasp and the fungus are native to Europe, North Africa and western Asia where their damage is considered secondary (Spradbery and Kirk, 1978). Introduced to areas of the Southern Hemisphere where exotic pine species are grown in plantations, these organisms have caused major losses. The insect invaded New Zealand by at least 1900, but did not cause serious concern until the 1940s (Talbot, 1977). It later spread to Tasmania and the southern parts of Australia and the wasp/fungus association was introduced into southern South America, beginning in Uruguay in 1980 (Ciesla, 2003). Invasion of South Africa occurred in 1994 (Tribe, 1995). Woodwasps are repeatedly detected in material imported to the USA, but were successfully excluded until 2004 (Wilson et al., 2009). The wasp and fungus were later found in nearby Canada (Ontario), although apparently due to a separate introduction (Bergeron et al., 2008; Wilson et al., 2009). Recent surveys found the wasp in four states of the USA (Evans-Goldner and Bunce, 2009) and 25 counties of Ontario in Canada (Shields, 2009).
Schiff (2008) summarizes differences in complexity of the ecological situations in the Southern Hemisphere countries and North America that could affect spread and impact of the fungus and wasp.
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Fungi
- Phylum: Basidiomycota
- Subphylum: Agaricomycotina
- Class: Agaricomycetes
- Subclass: Agaricomycetidae
- Order: Russulales
- Family: Amylostereaceae
- Genus: Amylostereum
- Species: Amylostereum areolatum
Notes on Taxonomy and NomenclatureTop of page
The genus Amylostereum, including A. areolatum, was separated by Boidin (1958) from Stereum on the basis of the encrusted cystidia in the hymenium. Lloydellopsis is a redundant later genus, based on the same type species. Xylobolus is another group of Stereum-like fungi having smooth amyloid basidiospores that is distinguished from Stereum and Amylostereum principally by the presence of acanthophyses in the hymenium (Boidin, 1958).
DescriptionTop of page
Basidiomata perennial on conifers, corky or leathery, partly resupinate, flattened on bark or wood with hymenium exposed, partly reflexed, with shield-shaped or broad areas projected 1-3 cm upwards/outwards. Upper/outer surface rust-brown to black, matted-hairy, undulating, with furrows or grooves. Hymenial surface generally smooth, sometimes cracked or uneven, grey to brown with a violet or lilac tint, margin thin, white. Basidia in hymenium narrowly clavate, 20-30 x 4-5 µm; basidiospores smooth, colourless, ellipsoidal to cylindrical, amyloid (staining blue in iodine). Context dimitic; cystidia encrusted, 40-60 x 6-9 mm. Thomsen (1998) found the hymenium of a limited sample of basidiocarps to be brown-violet, with a white margin; basidiospores were 3.5-5.5 x 2.0-3.0 mm.
DistributionTop of page
Due to the mutualistic relationship between the fungus and the wasp Sirex noctilio, an identical distribution is presumed. The wasp is considered native to Europe, North Africa and western Asia (Spradbery and Kirk, 1978), but records of the fungus only represent Europe. A. areolatum is more commonly seen in central Europe than in the northern countries (Eriksson et al., 1978; Jahn, 1979). On the other hand, the fungus has been identified as a pathogen of native tree species in Japan where it has a mutualistic relationship with at least one other wasp, Sirex nitobei (Fukuda and Hijii, 1996). Sirex juvencus is also a vector of A. areolatum in the Northern Hemisphere (Spradbery and Kirk, 1978).
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.Last updated: 17 Feb 2021
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|South Africa||Present||Introduced||Invasive||On Pinus radiata|
|Japan||Present||Present based on regional distribution.|
|Russia||Present||Present based on regional distribution.|
|-Eastern Siberia||Present||Wasp distribution|
|-Russian Far East||Present|
|-Western Siberia||Present||Wasp distribution|
|Serbia and Montenegro||Present||Native||Wasp distribution|
|United Kingdom||Present||Native||Wasp distribution|
|-Ontario||Present, Localized||Introduced||Invasive||On Pinus sylvestris|
|United States||Present||Present based on regional distribution.|
|-Michigan||Present, Localized||Introduced||Invasive||Wasp survey result|
|-New York||Present, Localized||Introduced||Invasive|
|-Pennsylvania||Present, Localized||Introduced||Invasive||Wasp survey result|
|-Vermont||Present, Localized||Introduced||Invasive||Wasp survey result|
|Australia||Present||Present based on regional distribution.|
|-New South Wales||Present||Introduced||Invasive||Wasp distribution|
|-Victoria||Present, Widespread||Introduced||Invasive||Wasp distribution|
|New Zealand||Present, Widespread||Introduced||Invasive|
|Brazil||Present||Present based on regional distribution.|
|-Rio Grande do Sul||Present||Introduced||Invasive||Wasp introduced|
|-Santa Catarina||Present||Introduced||Invasive||Wasp introuced|
|Chile||Present, Localized||Introduced||2001||Invasive||Wasp introduced 2001|
|Uruguay||Present||Introduced||1980||Invasive||Wasp introduced 1980|
History of Introduction and SpreadTop of page
The fungus and wasp larvae were introduced into New Zealand before 1900, apparently in shipments of logs from Europe (Gilmour, 1965). This was not of great concern until damage to exotic tree plantations appeared in the drought years of the late 1940s (Rawlings, 1955). In 1950-51, the wasp/fungus appeared in Tasmania (Gilbert and Miller, 1952). By 1961, they had arrived in Victoria eventually reaching New South Wales and Southern Australia (Carnegie et al., 2005). Despite the awareness of its threat, the wasp/fungus was introduced into South America, beginning in Uruguay in 1980, spreading to Argentina, Brazil, and Chile (Ciesla, 2003). Invasion of South Africa occurred in 1994 (Tribe, 1995). Woodwasps are repeatedly detected in material imported to the USA, but Sirex noctilio was successfully excluded until 2004 (Ciesla, 2003; Wilson et al., 2009). The wasp and fungus were later found in nearby Canada (Ontario) although apparently due to a separate introduction (Bergeron et al., 2008; Wilson et al., 2009). Recent surveys found the wasp in four states of the USA (Evans-Goldner and Bunce, 2009) and 25 counties of Ontario in Canada (Shields, 2009).
IntroductionsTop of page
|Introduced to||Introduced from||Year||Reason||Introduced by||Established in wild through||References||Notes|
|Natural reproduction||Continuous restocking|
|Argentina||1985-1986||Self-propelled (pathway cause)||Yes||Echeverria (1991); Echeverría (1991)||Thought to be introduced from Uruguay|
|Australia||New Zealand||1950-1961||Yes||Talbot (1977)||Tasmania - 1950, Victoria by 1961|
|Brazil||1988||Self-propelled (pathway cause)||Yes||Iede et al. (1988)||Thought to be introduced from Uruguay|
|Canada||2006||Bergeron et al. (2008)|
|Chile||2001||No||Ciesla (2003)||Limited areas|
|New Zealand||Europe||<1900||Timber trade (pathway cause)||Yes||Talbot (1977)|
|South Africa||Europe||1994||Yes||Tribe (1995)|
|USA||2004||Yes||Wilson et al. (2009)|
Risk of IntroductionTop of page
The most recent introductions into the USA and Canada indicate the level of risk for this pest/pathogen threat. Despite a high level of awareness and frequent interceptions of woodwasps by plant protection personnel (Ciesla, 2003), the wasp and fungus have arrived in North America. The likely routes and means were not identified (Bergeron et al., 2008; Wilson et al, 2009). Ciesla (2003) cites a number of consumer goods that incorporate logs or wood products as potential carriers. Bark-bearing and debarked logs, as well as untreated lumber, might carry the fungus, but the wasp appears necessary for its inoculation into trees. Once introduced, the wasp, a strong flier seeking out suitable tree hosts (Ciesla, 2003), can transport the fungus over land and possibly across narrow seas. Northwestern South America, eastern Africa, and Western Australia are areas for which accidental introduction by humans would be required (Carnegie et al., 2006). Likewise, afforestation in China with introduced Pinus species suggests that Sirex/Amylostereum is of quarantine concern for China (Bi et al., 2008).
Habitat ListTop of page
|Terrestrial||Managed||Managed forests, plantations and orchards||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Natural forests||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Natural forests||Principal habitat||Natural|
Hosts/Species AffectedTop of page
In Europe, the fungus is reported on Abies, Picea (Hansen and Knudsen, 1997) and Cryptomeria species (Cardosa et al., 1992). Gadgil (2005) considers the introduced species of Abies,Larix, and Picea to be relatively resistant in New Zealand, but the numerous Pinus species, also introduced, are generally susceptible. Abies holophylla is a susceptible native species in the Russian Far East (Boidin and Lanquetin, 1984). In Japan, Pinus densiflora and Pinus thunbergii are hosts (Kobayashi, 2007); the fungusis reported from China on a Picea sp. (Zhang, 2005). Pseudotsugamenziesii (Gilmour, 1965) and Cedrus atlantica are also hosts in New Zealand (Burnip et al., 2008). The tree species known to be attacked by Sirex noctilio, thus are also hosts for this fungus, are listed by CABI (2009).
Host Plants and Other Plants AffectedTop of page
Growth StagesTop of page
SymptomsTop of page
Symptoms on infected trees are a combination of those caused by the fungus and wasp vector. Resin droplets appear at sites of wasp oviposition (King, 1966; Kobayashi et al., 1978). On Pinus radiata, yellowing of needles occurs on older parts of the tree within several weeks, followed by yellowing and loss of needles on younger parts, or death of the tree with foliage remaining green or turning red-brown. The yellowing of younger needles is sometimes accompanied by wilting of the youngest ones at branch tips, the whole fascicle drooping, particularly in periods of active growth and adequate moisture (Coutts, 1969). However, these symptoms are due to a mucus material produced by the wasp and not the fungus (Coutts, 1969; Talbot, 1977).
List of Symptoms/SignsTop of page
|Growing point / dieback|
|Leaves / wilting|
|Leaves / yellowed or dead|
|Stems / dieback|
|Stems / discoloration|
|Stems / gummosis or resinosis|
|Stems / internal feeding|
|Stems / mycelium present|
|Stems / rot|
|Whole plant / plant dead; dieback|
|Whole plant / wilt|
Biology and EcologyTop of page
A. areolatum has a tetrapolar outcrossing mating system (Vasiliauskas and Stenlid, 1999). The species also exhibits vegetative compatibility groups in culture (Vasiliauskas and Stenlid, 1999), which appear to represent genetic clones (Vasiliauskas et al., 1998).
The fungus is not known to fruit in Australia or New Zealand (King, 1966; Hood, 1992), although isolates from infected trees were able to fruit on wood blocks in culture (Gaut, 1969). Arthrospores carried by adult female wasps are injected into trees during oviposition. Neither arthrospores nor basidiospores were able to grow beyond germination in pure culture, but Gaut (1969) was able to obtain cultures from arthrospores by starting them on a tissue culture of Pinus radiata. The sexual state is observed in nature in central Europe (Breitenbach and Kranzin, 1988), although it is rare in northern Europe (Eriksson et al., 1978). Based on the low number and wide distribution of vegetative compatibility groups around the Baltic Sea, Vasiliauskas and Stenlid (1999) and Thomsen and Koch (1999) considered distribution of the pathogen in that region more likely to have occurred through the insect vector than by aerial dispersal of the basidiospores.
Based on cultural tests, King (1966) suggested that A. areolatum is a poor competitor for growth in Sirex-infested trees; poor competition with saprobes in dead trees may explain the low frequency of fruiting. A. areolatum grows slowly in the tree and only in dry wood (70% moisture content) (Gaut, 1969; Webber and Gibbs, 1989) The fungus colonies are limited to areas around the oviposition site and larval tunnels while the tree is alive (King, 1966) and do not grow much beyond the inoculation site in resistant trees (Coutts, 1969). Nevertheless, Klepzig et al. (2009) found that in culture, A. areolatum can exclude the Ophiostoma species associated with the southern pine beetle, indicating that it might be successful in early competition in infected trees. Strains of the fungus may differ. Williams and Mastro (2009) report a significant growth rate difference between a North American isolate and one used in Australian biological control work. The Australian isolate is different from the single clone, which was introduced with the wasp into South Africa and South America (Slippers et al., 2001). Associations The fungus has a mutualistic association with Sirex spp.: Sirex noctilio, Sirex juvencus, Sirexnotobei (Gilbertson, 1984). It is carried in specific structures (mycangia) on the female woodwasp body at the base of the ovipositor, and is introduced into the holes drilled by the wasp when it is laying eggs, although, in some cases, only the spores will be deposited (Gaut, 1969; Fukuda and Hijii, 1996). The fungus grows in a limited area around the hole, rotting the sapwood, and providing a dry, resin-free habitat for the wasp larvae, which tunnel in the wood and feed on the mycelium (Madden, 1981; Webber and Gibbs, 1989). When fungal growth results from wasp deposits of spores alone in relatively healthy trees, this may weaken the trees sufficiently to provide additional breeding sites for the wasp (Gilbertson, 1984; Fukuda and Hijii, 1996). Wasp larvae carry hyphae encased in a wax-like material inside hypopleural organs on the exoskeleton, but these are absent in the pupae (Gilmour, 1965). The adult female then must acquire quantities of hyphae as it is emerging from pupation and out of the tree (Webber and Gibbs, 1989). Male larvae and adults do not carry the fungus.
Strains of the fungus may differ. Williams and Mastro (2009) report a significant growth rate difference between a North American isolate and one used in Australian biological control work. The Australian isolate is different from the single clone, which was introduced with the wasp into South Africa and South America (Slippers et al., 2001).
The fungus has a mutualistic association with Sirex spp.: Sirex noctilio, Sirex juvencus, Sirexnotobei (Gilbertson, 1984). It is carried in specific structures (mycangia) on the female woodwasp body at the base of the ovipositor, and is introduced into the holes drilled by the wasp when it is laying eggs, although, in some cases, only the spores will be deposited (Gaut, 1969; Fukuda and Hijii, 1996).
The fungus grows in a limited area around the hole, rotting the sapwood, and providing a dry, resin-free habitat for the wasp larvae, which tunnel in the wood and feed on the mycelium (Madden, 1981; Webber and Gibbs, 1989). When fungal growth results from wasp deposits of spores alone in relatively healthy trees, this may weaken the trees sufficiently to provide additional breeding sites for the wasp (Gilbertson, 1984; Fukuda and Hijii, 1996).
Wasp larvae carry hyphae encased in a wax-like material inside hypopleural organs on the exoskeleton, but these are absent in the pupae (Gilmour, 1965). The adult female then must acquire quantities of hyphae as it is emerging from pupation and out of the tree (Webber and Gibbs, 1989). Male larvae and adults do not carry the fungus.
ClimateTop of page
|Cf - Warm temperate climate, wet all year||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year|
|Cs - Warm temperate climate with dry summer||Preferred||Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers|
|Df - Continental climate, wet all year||Preferred||Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)|
|Dw - Continental climate with dry winter||Preferred||Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)|
Notes on Natural EnemiesTop of page
CABI (2009) presents information on natural enemies used for the biological control of Sirex noctilio.
Means of Movement and DispersalTop of page
Basidiospores are distributed by the wind from the fruiting bodies produced on the outside surface of erect or fallen trees (Breitenbach and Kranzlin, 1988). Basidiocarps are rare in northern Europe (Eriksson et al., 1978). Vasiliauskas and Stenlid (1999) and Thomsen and Koch (1999) suggest that this means of dispersal is unlikely to account for trans-Baltic occurrence of clones of this species. Instead, the fungus is likely to be wasp-transmitted across significant distances, but probably carried across international boundaries in imported wood. Basidiocarps have not been observed in overseas areas of introduction (King, 1966; Hood, 1992), thus basidiospores are not a significant means of spread there.
Pathway CausesTop of page
Pathway VectorsTop of page
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Bark||hyphae||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Stems (above ground)/Shoots/Trunks/Branches||hyphae||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
|Wood||hyphae||Yes||Pest or symptoms not visible to the naked eye but usually visible under light microscope|
Impact SummaryTop of page
ImpactTop of page
Talbot (1977) notes that the impact of the Sirex/Amylostereum association can vary depending on the silvicultural situation. The fungus is relatively rare in Europe, where the several Amylostereum species/siricid wasp associations are presumably in balance in the natural woodland. However, in plantations of exotic species this balance may be disrupted. Vasiliauskas et al. (1996) report A. areolatum as one of the primary parasites on wounded trees in Swedish plantations. The large outbreak of Sirex noctilio in 1946-1949 in New Zealand, despite the significant number of trees killed, may have had a beneficial effect by reducing the density of the long-unthinned plantations of Pinus radiata (Rawlings, 1955). Eliminating the more susceptible smaller and weaker trees through thinning is a tactic to reduce the severity and impact of the fungus/wasp association on plantations in Australia (Neumann and Marks, 1990). In the USA and Canada, a number of native species, some of which are attacked as exotics in overseas plantations, may be threatened (Wilson et al., 2009), particularly where they occur in denser stands or are grown in plantations in southern states (Dodds, 2009). The existing complex ecological situation that is more similar to that of the fungus native range may reduce or complicate the impact (Schiff, 2008).
Economic ImpactTop of page
Economic effects of the introduction of the Sirex noctilio/A. areolatum complex in countries of the Southern Hemisphere are summarized by CABI (2009). Haugen (2006) reports some projections of possible impacts on forests of the USA, and Yemshanov et al. (2009) have presented models of possible impacts on the forest industry in eastern Canada. The total harvest losses over 28 years in Canada could equal as much as $2.1 billion depending on model assumptions.
Environmental ImpactTop of page
Attention on the effects in the southern hemisphere has focused primarily on introduced tree species. No data are available on the impacts on the native species in forests in Japan (Kobayashi et al., 1978; Fukuda and Hijii, 1996). Haugen (2006) identified possible impacts in the USA through changes in forest composition, reduction in native siricid wasp populations, and increases in other insect pests and fungal pathogens that attack weakened or dying trees.
Risk and Impact FactorsTop of page
- Proved invasive outside its native range
- Has a broad native range
- Has high reproductive potential
- Reproduces asexually
- Host damage
- Increases vulnerability to invasions
- Modification of fire regime
- Negatively impacts forestry
- Threat to/ loss of native species
- Difficult to identify/detect as a commodity contaminant
- Difficult to identify/detect in the field
- Difficult/costly to control
DiagnosisTop of page
The production of arthrospores in culture is necessary to identify A. areolatum (Gaut, 1969). These appear in cultures grown on standard fungal media, including 2% malt extract agar and cornmeal agar (King, 1966; Stalpers, 1978). Gaut (1969) utilized matings of homokaryotic cultures, observation of hyphal anastomosis, and electrophoretic protein profiles to identify the pathogen in Australia. Bergeron et al. (2008) compared the ITS sequences of rDNA extracted from cultured isolates with those of standard strains of A. areolatum. The Canadian isolates matched European and Asian isolates at the 99% level; they matched Amylostereum chailletii at about 97%. Slippers et al. (2002) found that RFLP analysis of DNA easily distinguished the two species.
Detection and InspectionTop of page
In Europe, the fungus may eventually produce fruiting bodies on infected trees. Elsewhere, observation of holes made by emerging siricid wasps are a clue to the presence of the fungus associated with larval tunnels (Spradbery and Kirk, 1978; Schiff, 2008). Due to the fact that there may be more than one species of woodwasp in a given area, examination of culture characteristics or DNA sequence data is required to establish the identity of the fungus.
Similarities to Other Species/ConditionsTop of page
Amylostereum chailletii, also a woodwasp-vectored fungus in Northern Hemisphere forests, is reported to be distinguished by larger basidiospores, 6.2-8 x 3-4 mm, according to Boidin and Lanquetin (1984), although Thomsen (1998) found a range of 4.8-8 x 2.4-4 mm and Chamuris (1988) reports 5-8 x 3-5 mm. Thomsen (1998) describes the hymenium colour of fruiting bodies as varying in pale to darker brown shades (cream, orange-grey, reddish-brown, greyish-brown or brick-red) without the purple or lilac tint of A. areolatum; the paler colours are most common. Cultures on 5% MEA or 4% PDA are paler than those of A. areolatum (pale-yellow or yellowish-white vs. yellow-brown to rust-brown) and do not produce arthrospores (Thomsen, 1998). King (1966) reported that many basidiospores of A. chailletii germinated and produced mycelium on agar, whereas those of A. areolatum seldom germinated and, if so, did not produce mycelium.
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.
Gaps in Knowledge/Research NeedsTop of page
ReferencesTop of page
Bedding RA, 1995. Biological control of Sirex noctilio using the nematode Deladenus siricidicola. In: Nematodes and Biological Control of Insect Pests [ed. by Bedding, R. A. \Akhurst, R. J. \Kaya, H.]. Melbourne, Australia: CSIRO, 11-20.
Bergeron MJ; Hamelin RC; Leal I; Davis C; Groot Pde, 2008. First report of Amylostereum areolatum, the fungal symbiont of Sirex noctilio, on Pinus spp. in Canada. Plant Disease, 92(7):1138. HTTP://www.apsnet.org
Bi HuiQuan; Simpson J; Eldridge R; Sullivan S; Li RongWei; Xiao YuGui; Zhou JianHua; Wu ZhongXing; Yan Hong; Huang Quan; Liu QianLi, 2008. Survey of damaging pests and preliminary assessment of forest health risks to the long term success of Pinus radiata introduction in Sichuan, southwest China. Journal of Forestry Research, 19(2):85-100. http://jfr.nefu.edu.cn
Boidin J; Lanquetin P, 1984. [English title not available]. (Le genre Amylostereum (Basidiomycetes). Intercompatibilites partielles entre especes allopatriques.) Bulletin Trimestriel de la Societe Mycologique de France, 100:211-236.
Boroczky K; Crook DJ; Francese JA; Mastro VC; Tumlinson JH, 2009. Chemical ecology of Sirex noctilio. Proceedings of the 19th Interagency Research Forum on Invasive Species 2008 [ed. by U. S. D. A.]. Pennsylvania, USA: USDA Forest Service, 8-9. www.nrs.fs.fed.us/pubs/gtr/gtr_nrs-p-36.pdf
BPI (US National Fungus Collections), 2009. Fungal Databases - Specimens. Beltsville, USA: Systematic Mycology and Microbiology Laboratory, Agricultural Research Service, USDA. www.nt.ars-grin.gov/fungaldatabases/specimens/specimens.cfm
Carnegie AJ; Matsuki M; Haugen DA; Hurley BP; Ahumada R; Klasmer P; Sun JiangHua; Iede ET, 2006. Predicting the potential distribution of Sirex noctilio (Hymenoptera: Siricidae), a significant exotic pest of Pinus plantations. Annals of Forest Science, 63(2):119-128.
Coutts MP, 1969. The mechanism of pathogenicity of Sirex noctilio on Pinus radiata. I. Effects of the symbiotic fungus Amylostereum sp. (Thelephoraceae). Australian Journal of Biological Sciences, 22(4):915-24.
Croft P, 2007. Monitoring the introduction, development and damage of Sirex infestations in KwaZulu-Natal. The Sirex Woodwasp: Expanding Frontiers. Programme, International Sirex Symposium, Pretoria and Pietermaritzburg, South Africa, 9-16 May 2007., South Africa: Forestry and Agricultural Biotechnology Institute, 37. http://www.fabinet.up.ac.za/sirex/sirexsymposium
Dodds KJ, 2009. Sirex noctilio impacts on native and exotic pine stands in the northeastern United States. Proceedings of the 19th Interagency Research Forum on Invasive Species 2008 [ed. by U. S. D. A.]. Pennsylvania, USA: USDA Forest Service, 17-18. www.nrs.fs.fed.us/pubs/gtr/gtr_nrs-p-36.pdf
Evans-Goldner L; Bunce LK, 2009. United States Sirex woodwasp program update. Proceedings of the 19th Interagency Research Forum on Invasive Species 2008 [ed. by U. S. D. A.]. Pennsylvania, USA: USDA Forest Service. www.nrs.fs.fed.us/pubs/gtr/gtr_nrs-p-36.pdf
Gilbert JM; Miller LW, 1952. An outbreak of Sirex noctilio (F.) in Tasmania. Australian Forestry, 16:63-69.
Gilbertson RL, 1984. Relationships between insects and wood-rotting Basidiomycetes. In: Fungus-Insect Relationships. Perspectives in Ecology and Evolution [ed. by Wheeler, Q. \Blackwell, M.]. New York, USA: Columbia University Press, 514 pp.
Haugen DA, 2006. Sirex noctilio. Exotic Forest Pest Information System for North America. Pennsylvania, USA: USDA Forest Service. http://spfnic.fs.fed.us/exfor/data/pestreports.cfm?pestidval=33&langdisplay=English
Iede ET; Penteado S do RC; Bisol JC, 1988. First record of Sirex noctilio attacking Pinus tpda in Brazil. Circular Tecnica - Centro Nacional de Pesquisa de Florestas Curitiba, Parana, Brazil, No. 20:ii + 12 pp.
Klepzig KD; Slippers B; Wingfield MJ, 2009. Competition between fungi associated with Sirex woodwasp and Southern Pine Beetle. Proceedings of the 19th Interagency Research Forum on Invasive Species 2008 [ed. by U. S. D. A.]. Pennsylvania, USA: USDA Forest Service. www.nrs.fs.fed.us/pubs/gtr/gtr_nrs-p-36.pdf
Kobayashi T; Sasaki K; Enda N, 1978. Correlation between Sirex nitobei and Amylostereum areolatum, associated with the death of Japanese pine trees during winter season. Journal of the Japanese Forestry Society, 60(11):405-411.
Martínez AS; Fernández-Arhex V; Corley JC, 2006. Chemical information from the fungus Amylostereum areolatum and host-foraging behaviour in the parasitoid Ibalia leucospoides. Physiological Entomology, 31(4):336-340. http://www.blackwell-synergy.com/doi/pdf/10.1111/j.1365-3032.2006.00523.x
Shields L, 2009. Update on Sirex noctilio in Canada. Proceedings of the 19th Interagency Research Forum on Invasive Species 2008 [ed. by U. S. D. A.]. Pennsylvania, USA: USDA Forest Service, 69. www.nrs.fs.fed.us/pubs/gtr/gtr_nrs-p-36.pdf
Slippers B; Wingfield BD; Coutinho TA; Wingfield MJ, 2002. DNA sequence and RFLP data reflect geographical spread and relationships of Amylostereum areolatum and its insect vectors. Molecular Ecology, 11(9):1845-1854.
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