Orthotomicus erosus (Mediterranean pine beetle)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Plant Trade
- Wood Packaging
- Environmental Impact
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Orthotomicus erosus (Wollaston, 1857)
Preferred Common Name
- Mediterranean pine beetle
Other Scientific Names
- Ips erosus (Wollaston, 1857)
- Ips erosus var. robustus Knotek
- Tomicus erosus Wollaston, 1857
- Tomicus rectangulus Ferrari, 1867
Local Common Names
- Germany: Borkenkaefer, Suedeuropaeischer Kiefern-; Sudeuropäischer Kiefernborkenkäfer,
- Italy: Bostrico corroso
- IPSXER (Orthotomicus erosus)
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Coleoptera
- Family: Scolytidae
- Genus: Orthotomicus
- Species: Orthotomicus erosus
Notes on Taxonomy and NomenclatureTop of page
DescriptionTop of page
Scolytidae eggs are smooth, ovoid, white and translucent. The eggs of O. erosus are approximately 1 mm long and laid separately in niches along the egg gallery.
All scolytidae larvae are similar in appearance and difficult to separate. They are white, 'C'-shaped and legless. The head capsule is lightly sclerotized and amber with dark, well-developed mouthparts. Each of the abdominal segments has two to three tergal folds and the pleuron is not longitudinally divided. The larvae do not change as they grow.
Scolytidae pupae are white and mummy-like. They are exarate, with the legs and wings free from the body. Some species have paired abdominal urogomphi. The elytra are either rugose or smooth and they sometimes have a prominent head and thoracic tubercles.
The adults are, on average, 3 to 3.8 mm long and reddish-brown. They are typical bark beetles of the subfamily Ipinae, in the family Scolytidae. The head is covered by a thoracic shield and is not visible when viewed dorsally and the declivity is concave Each side is armed with four spines and the second from the top is more conspicuous (Grüne, 1979).
DistributionTop of page
O. erosus is widely distributed across the Mediterranean and southern Europe, Asia and North Africa. It has been introduced into Fiji, South Africa, Swaziland (Wood and Bright, 1992) and the USA (Haack, 2004).
A record of O. erosus in Norway (ISSG, 2009) published in previous versions of the Compendium is invalid. A Norwegian population of Orthotomicus which was thought to be O. erosus (Kirkendall, 1989) is now known to be Orthotomicus proximus (Seybold et al., 2016). There are no current records of native or introduced populations of O. erosus in Norway (Seybold et al., 2016)
A record of O. erosus in Chile (Ciesla, 1988; Ciesla and Parra Sanhueza, 1988; Wood and Bright, 1992) published in previous versions of the Compendium is invalid. The invasive pine bark beetle commonly referred to in Chilean research as O. erosus is a misidentification of Orthotomicus laricis (Kirkendall, 2018).
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|
|Cyprus||Present, Widespread||Native||Invasive||Original citation: Ciesla (2003)|
|Finland||Absent, Intercepted only|
|Norway||Absent, Invalid presence record(s)|
|United States||Present||Present based on regional distribution.|
|-Florida||Absent, Intercepted only||Original citation: Haack (2001)|
|-Maryland||Absent, Intercepted only||Original citation: Haack (2001)|
|-New Jersey||Absent, Intercepted only||Original citation: Haack (2001)|
|-South Carolina||Absent, Intercepted only||Original citation: Haack (2001)|
|-Texas||Absent, Intercepted only||Original citation: Haack (2001)|
|Chile||Absent, Invalid presence record(s)||Original citation: Kirkendall et al. (2018)|
Risk of IntroductionTop of page
O. erosus has a broad host range. Moreover, its successful introduction and establishment into five countries demonstrates its ability to adapt to new pine hosts. Chararas (1973) reported that in Turkey, O. erosus was successfully reared on a variety of hosts, including several North American pines. In this study, O. erosus was not significantly affected by differences in host terpenes, provided that they did not differ drastically from their indigenous hosts, or did not contain repellents such as heptane, which is found in the North American Pinus jeffreyi.
Although obviously adaptable, the ability of O. erosus to adapt to varying climatic conditions and hosts or compete with other insects, may be questionable. In Chile, for example, O. erosus was introduced around 1986, but was either not successful in competing with another introduced bark beetle, Hylurgus ligniperda, or unable to adapt to the host, Pinus radiata, or the local climatic conditions were unsuitable for its development. It is now difficult to find O. erosus in Chile. In contrast, Karnavar (1984) reported that O. erosus first appeared in Swaziland in the early 1980s and soon became a major pest.
HabitatTop of page
Habitat ListTop of page
Hosts/Species AffectedTop of page
Near Eastern hosts include two varieties of Calabrian pine; Pinus brutia var. eldarica and Pinus brutia var. pityusa [P. brutia]. Known Asian pine hosts are Pinus massoniana (Masson pine), Pinus armandii (Armand pine), Pinus kesiya (Khasi pine), Pinus yunnanensis (Yunnan pine) and Pinus taiwanensis (Chinese pine).
Several North American pines that have been planted in areas where this insect is native or has become established, include Pinus coulteri (Coulter pine), Pinus caribaea (Caribbean pine), Pinus echinata (shortleaf pine), Pinus radiata (Monterey pine), Pinus patula (Mexican weeping pine) and Pinus strobus (Eastern white pine).
Occasionally, maturing beetles feed in Pseudotsuga menziesii (Douglas-fir), Picea spp. (spruce), Abies spp. (fir) and Cedrus spp. (cedar). However, this insect does not breed in trees other than pines (Eglitis, 2000).
Host Plants and Other Plants AffectedTop of page
Growth StagesTop of page
SymptomsTop of page
Breeding attacks are characterized by the presence of reddish-brown boring dust on the bark surface of trees, freshly cut logs or wind-thrown trees. If relatively vigorous trees are attacked, pitch tubes may be found in the bark crevasses. The gallery pattern in the cambial region of infested trees consists of a nuptial chamber and one to five longitudinal egg galleries, but this could vary depending on the host and location. A blue stain in the woody tissue accompanies the breeding attacks.
Round exit holes are visible on the bark surface of trees where this insect has completed its life cycle and the adults have emerged.
List of Symptoms/SignsTop of page
|Leaves / yellowed or dead|
|Stems / gummosis or resinosis|
|Stems / visible frass|
|Whole plant / frass visible|
|Whole plant / plant dead; dieback|
Biology and EcologyTop of page
Orthotomicus is a small genus of bark beetles found in Asia, Europe and North America, and is closely related to the Ips genus. Twelve species are known worldwide. Eleven species have Palearctic distributions and one species, Orthotomicus caelatus, is found in North America (Wood, 1982).
The adults of O. erosus are polygamous. The males bore through the bark to the cambium layer where they construct a nuptial chamber. They are joined by one to three females, each of which mates with the male and then constructs an individual egg gallery from the nuptial chamber, parallel to the grain of the wood. Typically, a female oviposits between 26 and 75 eggs in niches along the sides of the galleries (Mendel and Halperin, 1982). The females will make ventilation holes in their egg galleries and they sometimes abandon the gallery and finish their egg laying in another tree. The larvae mine at right angles to the parent gallery and have three instars.
The adults must feed before reaching sexual maturation. This occurs beneath the bark of the tree in which the brood developed, providing that the bark is still moist. If the inner bark is too dry, maturation feeding can take place in a different host tree, sometimes of a different species. Occasionally, maturation feeding will take place during normal egg gallery construction and oviposition (Mendel and Halperin, 1982).
O. erosus completes two to seven generations per year, depending on the temperature. Two generations per year are common in Turkey, France and Morocco. In Tunisia, Chararas and M'Sadda (1973) found that O. erosus completed three and sometimes four generations in a year, with the development time being a function of the nutritive quality of the wood, as well as the temperature. In Israel, where the beetle can complete three to five generations in a year, the time required for the development of a brood varies from 25 days in the summer to 76 days in the winter (Mendel, 1983). Based on these development times, Mendel (1983) concluded that coastal Israel could have as many as seven generations per year. Tribe (1990) estimated that four generations per year occur in South Africa, based on an average development time of 35 days for one generation.
The winter is spent in the adult stage. From mid-October to December, the adults aggregate beneath the bark of the host in which they developed, or in a new one. They enter the new host through a single hole and then concentrate as many as several hundred individuals into the phloem/cambium region (Mendel, 1983). Beetle flight can occur through a broad temperature range of 14 to 38°C (Chararas and M'Sadda, 1973). In Israel, the threshold for flight is even lower during winter (12°C) (Mendel, 1983).
Breeding generally takes place in the rough-barked sections of the bole and in branches larger than 5 cm in diameter. Smooth-barked areas are primarily used for maturation feeding. The lower trunk of relatively old pines is not suitable for attack because the bark is too thick. In Israel, trees that are younger than 5-years-old are not usually attacked (Mendel and Halperin, 1982).
O. erosus commonly occurs in association with other bark beetles. Common associates on pines in Israel included Pityogenes calcaratus, Tomicus destruens [Hylurgus destruens] and Carphoborus minimus (Mendel and Halperin, 1982). In South Africa, Tribe (1990) found Hylastes angustatus, Hylurgus ligniperda and Pissodes nemorensis associated with O. erosus. Ciesla (2004) reported the occurrence of Tomicus minor in the upper boles of trees infested by O. erosus in Cyprus.
A number of wood-staining fungi are associated with O. erosus in a symbiotic relationship. These also play a role in the death of the host tree and cause the loss of lumber quality. A number of these fungi have been identified and include Ophiostoma ips and Verticicladiella alacris (Wingfield and Marasas, 1980).
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Aulonium ruficorne||Predator||Eggs; Arthropods|Larvae|
|Corticeus pini||Predator||Eggs; Arthropods|Larvae|
|Eurytoma morio||Parasite||Arthropods|Larvae; Arthropods|Pupae|
|Platysoma oblongum||Predator||Arthropods|Larvae; Arthropods|Pupae|
|Roptrocerus xylophagorum||Parasite||Arthropods|Larvae; Arthropods|Pupae|
Notes on Natural EnemiesTop of page
Tribe (2003) noted that Dendrosoter caenopachoides is dispersing slowly, which he attributes to the fact that it can only parasitize hosts in thin-barked trees but in these it can achieve very high rates of parasitism.
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||adults||Yes||Yes||Pest or symptoms usually visible to the naked eye|
|Stems (above ground)/Shoots/Trunks/Branches||adults; eggs; larvae; nymphs; pupae||Yes||Pest or symptoms usually visible to the naked eye|
Wood PackagingTop of page
|Wood Packaging liable to carry the pest in trade/transport||Timber type||Used as packing|
|Solid wood packing material with bark||Pine: crating, dunnage, pallets||Yes|
|Wood Packaging not known to carry the pest in trade/transport|
|Loose wood packing material|
|Processed or treated wood|
|Solid wood packing material without bark|
ImpactTop of page
Occasionally other insects or diseases are the stressing agents that lead to attacks by O. erosus. In France, Carle (1971) noted that O. erosus was one of the beetles involved in the decline of P. pinaster following the weakening of the host by the scale insect, Matsucoccus feytaudi. Zwolinski et al. (1995), working in South Africa, described the occurrence of O. erosus on P. radiata infected with the canker-causing fungus, Diplodia pinea [Sphaeropsis sapinea]. O. erosus was one of two beetles attacking these trees and was confined to the zone of discoloration produced by the fungus.
Throughout its geographic range, there have been reports of population build-ups of O. erosus in weakened or stressed host material, followed by attacks on healthy trees. In Israel, Halperin et al. (1982) related the increase of bark beetle outbreaks in P. halepensis to the maturation of plantations and increased thinning. Also in Israel, Mendel and Halperin (1982) reported that the agents predisposing stands to attack were thinning operations, followed by dry winters or fires in neighbouring pine stands. Ferreira and Ferreira (1986) indicated that O. erosus periodically reaches epidemic levels and has caused the death of many P. pinaster trees in Portugal.
Environmental ImpactTop of page
Detection and InspectionTop of page
Similarities to Other Species/ConditionsTop of page
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.
The most effective means of reducing the losses caused by O. erosus are cultural methods designed to maintain healthy, vigorous pine forests that are capable of resisting attack. Available tactics include: avoiding planting pines on unsuitable sites; periodic thinning to reduce stocking levels and competition for available moisture; and the rapid removal of wind-thrown trees or broken tops caused by ice and snow, which can serve as breeding sites.
Within its natural range, the population levels of O. erosus are undoubtedly held in check by a complex of indigenous natural enemies unless forest conditions favour a population increase. However, in South Africa, where O. erosus has been introduced, a classical biological control programme, involving the rearing and release of the parasitoid, Dendrosoter caenopachoides, was conducted. The programme resulted in the establishment of this parasitoid (Tribe and Kfir, 2001).
In trials in Israel, pine trap-logs containing the larvae and pupae of O. erosus were sprayed with carbaryl, chloropyrifos, or cypermethrin at concentrations of 0.1, 0.5 or 1.25% a.i.. At 1.25%, the effectiveness of the treatment (based on the rate of prevention of beetle emergence) was 100% with chloropyrifos (chlorpyrifos) and cypermethrin, and 94% with carbaryl. At 0.5%, chloropyrifos was 98% effective, but in general the lower concentrations did not provide an effective control (Mendel et al., 1983).
The chemical control of O. erosus has been attempted in Turkey but, as is the case with the chemical control of bark beetles in general, this has proven to be costly and of marginal effectiveness. Therefore the chemical control of this insect is not recommended (Mercikoglu, 2001).
The response of O. erosus to pheromones was studied in south-western France. O. erosus adults were caught mostly in traps baited with a combination of the insect-produced compounds 2-methyl-3-buten-2-ol and ipsdienol. When used together with 2-methyl-3-buten-2-ol, an increase in the concentration of up to 1000-fold of racemic ipsdienol resulted in a continual increase in the numbers of O. erosus and Ips sexdentatus caught. This was accompanied by a steady increase in the percentage of females collected. It is assumed that 2-methyl-3-buten-2-ol influences the landing behaviour of O. erosus, whereas ipsdienol acts as a long-distance signal (Klimetzek and Vité, 1986). The use of pheromones for the mass-trapping of adult beetles has reportedly been met with some degree of success in Turkey and was found to be less costly than mechanical or chemical control methods (Mercikoglu, 2001).
Aerial and ground surveys can be conducted over pine forests to detect tree mortality caused by O. erosus and other bark beetles. The population levels of adults can be monitored with funnel traps baited with commercial Ips lures. The occurrence of O. erosus outside its geographic range could also be monitored with pheromone traps.
Integrated Pest Management
The management of O. erosus can be accomplished by a combination of monitoring designed to detect low levels of bark beetle-caused tree mortality, the rapid removal and processing of infested trees, and the management of pine forests to keep them in a vigorous condition.
ReferencesTop of page
Arias, M., Robertson, L., Garcia-Alvarez, A., Arcos, S. C., Escuer, M., Sanz, R., Mansilla, J. P., 2005. Bursaphelenchus fungivorus (Nematoda: Aphelenchida) associated with Orthotomicus erosus (Coleoptera: Scolitydae) in Spain. Forest Pathology, 35(5), 375-383. doi: 10.1111/j.1439-0329.2005.00422.x
Bright DE, Skidmore RE, 2002. A catalogue of Scolytidae and Platypodidae (Coleoptera), Supplement 2 (1995-1999). Ottawa, Canada: NRC Research Press, 523 pp.
Chararas C, 1973. Adaptability of Orthotomicus erosus to conifers other than its usual host species. Comptes Rendus Hebomadaires de Seances de l'Acadamie de Sciences, 275(4):555-558.
Chararas C, M'Sadda K, 1973. Study of the biology, behaviour and the action of cobalt60 ionising radiations in Orthotomicus erosus Woll., (Coleoptera Scolytidae), a specific pest of conifers. Archives de l'Institut Pasteur de Tunis, 50(3):243-265
Ciesla WM, 2004. Forests and forest protection in Cyprus. Forestry Chronicle, 79(6): in press.
Ciesla WM, Parra Sanhueza P, 1988. Orthotomicus erosus Corporación Nacional Forestal, Protección Fitosanitaria Forestal (Chile). Folleto de Divulgación, Ano 8 N 16.
Eglitis A, 2000. Exotic Pest Information System for North America Prototype Application (EXFOR): Orthotomicus erosus. http://spfnic.fs.fed.us/exfor/data/pestreports.cfm?pestidval=9&langdisplay=english.
Gómez D, Martínez G, 2013. Bark beetles in pine tree plantations in Uruguay: first record of Orthotomicus erosus Wollaston (Coleoptera: Curculionidae: Scolytinae). Coleopterists Bulletin, 67(4):470-472. http://www.bioone.org/loi/cole
Grüne S, 1979. Handbuch zur bestimmung der Europäischen Borkenkäfer (Brief illustrated key to European bark beetles). Hannover, Germany: Verlag M & H Schapfer.
Haack RA, 2001. Intercepted scolytidae (Coleoptera) at U.S. ports of entry: 1985-2000. Bark Beetles and Reforestation Pests: Facing a Crisis. Proceedings of a meeting of IUFRO Working Parties S7.03.03 (Integrated control of scolytid beetles) and S7.03.03 (Insects affecting reforestation), Velaine-en-Haye, France, 5-7 September, 2001. Integrated Pest Management Reviews, 6(3/4)253-282.
Haack RA, 2001. Intercepted Scolytidae (Coleoptera) at US ports of entry: 1985-200. Integrated Pest Management Reviews 6: 253-282.
Karnavar GK, 1984. Preliminary studies on the use of 2-methyl-3-buten-2-ol as an attractant for the pine bark beetle, Orthotomicus erosus. Journal of the Royal Swaziland Society of Science and Technology, 5(2):2-4
Kirkendall, L. R., 2018. Invasive Bark Beetles (Coleoptera, Curculionidae, Scolytinae) in Chile and Argentina, Including Two Species New for South America, and the Correct Identity of the Orthotomicus Species in Chile and Argentina, Diversity, 10(40), doi: 10.3390/d10020040
Klimetzek D, Vite JP, 1986. The effect of insect-produced attractants on the aggregation behaviour of the Mediterranean pine engraver beetle Orthotomicus erosus. Journal of Applied Entomology, 101(3):239-243
Lee, J. C., Flint, M. L., Seybold, S. J., 2008. Suitability of pines and other conifers as hosts for the invasive Mediterranean pine engraver (Coleoptera: Scolytidae) in North America. Journal of Economic Entomology, 101(3), 829-837. doi: 10.1603/0022-0493(2008)101[829:SOPAOC]2.0.CO;2
Mercikoglu AM, 2001. Biological struggles against bark beetles in Izmir Forest Conservancy (Abstract). Proceedings XI World Forestry Congress, Antalya, Turkey 13-22 October 1997, Summaries of Voluntary Papers 1(5).
Questienne P, 1979. Notes sur quelques insectes nuisibles aux pins au Maroc. Annales de Gembloux, 85(2):113-130.
Sarikaya O, Yildirim S, 2011. Scolytinae (Coleoptera: Curculionidae) species of the coniferous forests in Isparta-Aksu province. (Isparta-Aksu yöresi igne yaprakli ormanlariniin scolytinae (Coleoptera: Curculionidae) türleri.) Bartin Orman Fakültesi Dergisi, 13(20):38-50. http://bof.bartin.edu.tr/journal/1302-0943/2011/Cilt13/Sayi20/2011-05.pdf
Serrao-Nogueira CD, 1976. O problem fitossanitario dos pinhais do Estaril. Anais do Instituto Superior de Agronomia, 36:203-240.
Seybold SJ, Downing M, 2009. What risks do invasive bark beetles and woodborers pose to forests of the western United States? A case study of the Mediterranean pine engraver, Orthotomicus erosus. General Technical Report - Pacific Northwest Research Station, USDA Forest Service. The Western Bark Beetle Research Group: a unique collaboration with Forest Health Protection. Proceedings of a symposium at the 2007 Society of American Foresters conference, Portland, Oregon, 23-28 October 2007. Portland, USA: Pacific Northwest Research Station, USDA Forest Service, 111-134.
Seybold, S. J., Penrose, R. L., Graves, A. D., 2016. Invasive Bark and Ambrosia Beetles in California Mediterranean Forest Ecosystems. In: Insects and Diseases of Mediterranean Forest Systems, [ed. by Paine, T. D., Lieutier, F.]. Heidelberg, New York, Dordrecht, London, Springer, 583-662.
Takov, D., Doychev, D., Linde, A., Draganova, S., Pilarska, D., 2011. Pathogens of bark beetles (Coleoptera: Curculionidae) in Bulgarian forests. Phytoparasitica, 39(4), 343-352. doi: 10.1007/s12600-011-0167-3
Tribe GD, 1990. Phenology of Pinus radiata log colonization and reproduction by the European bark beetle Orthotomicus erosus (Wollaston) (Coleoptera: Scolytidae) in the south-western Cape Province. Journal of the Entomological Society of Southern Africa, 53(2):117-126
Tribe GD, 2003. Biological control of defoliating and phloem- or wood-feeding insects in commercial forestry in Southern Africa. In: Neuenschwander P, Borgemeister C, Langewald J, eds. Biological control in IPM systems in Africa. Wallingford, UK: CAB International, 113-129.
Tribe GD, Kfir R, 2001. The establishment of Dendrosoter caenopachoides (Hymenoptera: Braconidae) introduced into South Africa for the biological control of Orthotomicus erosus (Coleoptera: Scolytidae), with additional notes on D. sp. nr. labdacus. African Entomology, 9(2):195-198; 23 ref.
Wood SL, Bright DE, 1992. A catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxonomic index. Great Basin Naturalist Memoirs, 13: 1-1553.
Arias M, Robertson L, Garcia-Alvarez A, Arcos S C, Escuer M, Sanz R, Mansilla J P, 2005. Bursaphelenchus fungivorus (Nematoda: Aphelenchida) associated with Orthotomicus erosus (Coleoptera: Scolitydae) in Spain. Forest Pathology. 35 (5), 375-383. DOI:10.1111/j.1439-0329.2005.00422.x
Bright DE, Skidmore RE, 2002. A catalogue of Scolytidae and Platypodidae (Coleoptera), Supplement 2 (1995-1999., Ottawa, Canada: NRC Research Press. 523 pp.
CABI, Undated. Compendium record. Wallingford, UK: CABI
CABI, Undated a. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI
CABI, Undated b. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI
Gómez D, Martínez G, 2013. Bark beetles in pine tree plantations in Uruguay: first record of Orthotomicus erosus Wollaston (Coleoptera: Curculionidae: Scolytinae). Coleopterists Bulletin. 67 (4), 470-472. http://www.bioone.org/loi/cole DOI:10.1649/0010-065X-67.4.470
Lee J C, Flint M L, Seybold S J, 2008. Suitability of pines and other conifers as hosts for the invasive Mediterranean pine engraver (Coleoptera: Scolytidae) in North America. Journal of Economic Entomology. 101 (3), 829-837. http://www.bioone.org/doi/full/10.1603/0022-0493%282008%29101%5B829%3ASOPAOC%5D2.0.CO%3B2 DOI:10.1603/0022-0493(2008)101[829:SOPAOC]2.0.CO;2
Mercikoglu AM, 2001. Biological struggles against bark beetles in Izmir Forest Conservancy (Abstract). [Proceedings XI World Forestry Congress, , Antalya, Turkey 13-22 October 1997, Summaries of Voluntary Papers], 1 (5)
Mouton M, Wingfield M J, Wyk P S van, Wyk P W J van, 1994. Graphium pseudormiticum sp. nov.: a new hyphomycete with unusual conidiogenesis. Mycological Research. 98 (11), 1272-1276. DOI:10.1016/S0953-7562(09)80298-0
Seybold SJ, Penrose RL, Graves AD, 2016. Invasive Bark and Ambrosia Beetles in California Mediterranean Forest Ecosystems. In: Insects and Diseases of Mediterranean Forest Systems, Heidelberg, New York, Dordrecht, London, Springer. 583-662.
Siitonen J, 2000. Beetles (Coleoptera) imported to Finland on Russian roundwood. In: Aktuelt fra Skogforskningen. ?s, Norway: Norsk Institutt for Skogforskning (Norwegian Forest Research Institute). 11-17.
Sousa E, Naves P, Bonifácio L, Bravo M A, Penas A C, Pires J, Serrão M, 2002. Preliminary survey for insects associated with Bursaphelenchus xylophilus in Portugal. Bulletin OEPP. 32 (3), 499-502. DOI:10.1046/j.1365-2338.2002.00597.x
Tribe G D, 1990. Phenology of Pinus radiata log colonization and reproduction by the European bark beetle Orthotomicus erosus (Wollaston) (Coleoptera: Scolytidae) in the south-western Cape Province. Journal of the Entomological Society of Southern Africa. 53 (2), 117-126.
Wood SL, Bright DE, 1992. A catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxonomic index. In: Great Basin Naturalist Memoirs, 13 1-1553.
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