Invasive Species Compendium

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Hylastes ater
(black pine bark beetle)

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Datasheet

Hylastes ater (black pine bark beetle)

Pictures

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PictureTitleCaptionCopyright
Adult beetle, museum set specimen.
TitleAdult
CaptionAdult beetle, museum set specimen.
Copyright©Rune Axelsson
Adult beetle, museum set specimen.
AdultAdult beetle, museum set specimen.©Rune Axelsson

Identity

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Preferred Scientific Name

  • Hylastes ater (Paykull, 1800)

Preferred Common Name

  • black pine bark beetle

Other Scientific Names

  • Bostrichus ater Paykull, 1800
  • Hylesinus chloropus Duftschmidt, 1825
  • Tomicus pinicola Bedel, 1888

International Common Names

  • English: bark beetle, black pine
  • French: hylesine noir du pin

Local Common Names

  • Denmark: rodbille, fyrrens
  • Finland: ruskoniluri
  • Germany: Bastkaefer, Schwarzer Kiefern-; Schwarzer Kiefernbastkär
  • Italy: ilaste nero dei pini
  • Japan: matsuno-kuro-kikuimushi
  • Netherlands: Dennebastkever, zwarte; zwarte dennebastkever
  • Norway: fururotbille
  • Sweden: tallbastborre, Svart

EPPO code

  • HYASAR (Hylastes ater)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Coleoptera
  •                         Family: Curculionidae
  •                             Genus: Hylastes
  •                                 Species: Hylastes ater

Notes on Taxonomy and Nomenclature

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There has been some disagreement as to whether H. ater and Hylastes brunneus are distinct species. Blair (1949), Duffy (1953), Hansen (1955) and Lekander (1965) stated that they are distinct but Schedl (1968) considered that H. brunneus was only a form of H. ater. Beaver (1970) presented evidence of both adult and larval characters that demonstrate they are distinct species.

Description

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The eggs are ovoid, pearly-white, 0.9 mm long and 0.5 mm wide (Clark, 1932).

The larvae are of a typical scolytine form. For a detailed description see Lekander (1968) and Beaver (1970).

The pupa is soft, yellowish-white and exarate, with two conspicuous caudal processes on the ninth abdominal segment. Spines are conspicuous on the head, pronotum and abdomen (Clark, 1932).

There are about 30 species of Hylastes that are native to the Holarctic region and they are all superficially quite similar. See Wood (1982) for a generic description of the adults. Hylastes is closely related to Hylurgops but can be distinguished by having narrower, emarginated third tarsal segments. In Hylurgops these segments are broader and bilobed, and the pronotum is usually anteriorly constricted.

H. ater adults are 3.5-4.5 mm long and black (light-brown when teneral). The pronotum is strongly elongate and parallel-sided for the basal half with a median, impunctate, conspicuous ridge. The elytral interstices are dull and microscopically reticulate between the punctures.

Distribution

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H. ater is indigenous to virtually the whole of the Palearctic region and is present as an introduced species in Australia, Chile and New Zealand.

Distribution Table

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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: 10 Feb 2022
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Asia

ChinaPresentPresent based on regional distribution.
-FujianPresentNative
-HeilongjiangPresentNative
JapanPresentNative
North KoreaPresent
South KoreaPresent
TurkeyPresentNative

Europe

AustriaPresentNative
BelgiumPresentNative
BulgariaPresentNative
CyprusPresentNative
CzechoslovakiaPresentNative
Federal Republic of YugoslaviaPresentNative
DenmarkPresentNative
EstoniaPresentNative
FinlandPresentNative
FrancePresentNative
-CorsicaPresent
GermanyPresentNative
GreecePresentNative
ItalyPresentNative
LatviaPresentNative
LithuaniaPresentNative
NetherlandsPresentNative
NorwayPresentNative
PolandPresentNative
PortugalPresentNative
-AzoresPresentNative
RussiaPresentNative
SpainPresentNative
SwedenPresentNative
SwitzerlandPresentNative
United KingdomPresentNative

North America

GreenlandPresent

Oceania

AustraliaPresentPresent based on regional distribution.
-New South WalesPresentIntroduced
-QueenslandPresentIntroduced
-South AustraliaPresentIntroduced
-TasmaniaPresentIntroduced
-VictoriaPresentIntroduced
New ZealandPresentIntroduced1929

South America

ChilePresentIntroduced1981

Risk of Introduction

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H. ater is readily transported around the world in logs with bark on, in dunnage and casewood that has bark strips. However, a thorough inspection should reveal its presence in such material.

Hosts/Species Affected

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The main hosts of H. ater are Pinus spp. but other conifers are utilized to a lesser extent.

Growth Stages

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Post-harvest, Seedling stage

List of Symptoms/Signs

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SignLife StagesType
Growing point / dieback
Growing point / discoloration
Roots / external feeding
Stems / external feeding
Stems / gummosis or resinosis
Whole plant / external feeding
Whole plant / plant dead; dieback

Biology and Ecology

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H. ater primarily breeds in the inner bark and cambium of pine roots and fresh stumps, the bases of dead and dying trees, and in recently felled logs, particularly those in contact with the ground (Milligan, 1978; Pasek, 1998). The adults also feed on the bark around the root collars, and the roots of pines and other conifers. Broods are mainly reared in pines (Clark, 1932; Milligan, 1978).

The brood galleries may be initiated in any month of the year. Each gallery that is initiated by the female consists of a short entrance tunnel leading to an oblique nuptial chamber from which issues a single egg gallery. This is 80-130 mm long and usually parallel with the grain of the wood. These brood galleries reach, but do not engrave, the surface of the sapwood. The male is usually in the nuptial chamber, close to the entrance and the female is further in towards the egg gallery, but in some chambers the male may be absent. Approximately 100 eggs are laid in individual notches that the female cuts in the lateral walls of the egg gallery. The females may sometimes leave the first egg gallery and start a second one elsewhere. The males may assist with the gallery construction, at least by pushing out frass from the entrance and sometimes making short feeding tunnels that extend from the nuptial chamber. The success of brood galleries in the absence of the males suggests that their part in establishing the galleries is only minor. The larvae initially make feeding tunnels at right angles to the maternal gallery, but these are later randomly directed and eventually obliterate both the early larval tunnels and those made by the parent adults. There are four larval instars and the rate of development of the feeding larvae appears to depend on seasonal temperatures. Emergence does not immediately follow eclosion of the adults but the length of the delay has not been investigated. Some adults feed in the material in which they were reared but the majority emerge and feed in fresh material where groups of approximately 40 feeding adults are found in communal galleries (Milligan, 1978).

In Canterbury, New Zealand the development from egg to adult can take from 60 to 300 days. The duration of development is least when the eggs are laid at the end of January and the beginning of February, and greatest when they are laid in approximately the second week of March. These later broods overwinter as mature larvae and appear to be subject to diapause so that they do not pupate until late December. Variations in the rate of development of both feeding and non-feeding larvae cause the adults from both the fast-developing and slow-developing broods to appear in January (Milligan, 1978).

There are two or three overlapping broods of H. ater per year in New Zealand (Clark, 1932) and Munro (1917) reported two in Scotland. Milligan (1978) noted that in September to October and in April to May swarming flights of H. ater occur in New Zealand. These do not coincide with the times when the greatest numbers reach the adult stage and probably involve many beetles that have been feeding for some time. What initiates this swarming, or what purpose it serves, is not understood. It has been suggested that it is associated with the local depletion of host material but this would not seem to apply in those forests where felling continues throughout the year (Milligan, 1978).

Reay and Walsh (2001) discuss the results of a trapping programme, which indicate that H. ater is now univoltine in New Zealand. They hypothesize that this has happened since Hylurgus ligniperda (another northern hemisphere scolytine) established in New Zealand in 1974. The trapping programme used raw turpentine in Lindgren funnel traps as the attractant and this must make the results of the programme open to question because Reay and Walsh (2002a) reported that raw turpentine performed no better as an attractant than a control.

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Parasitylenchus hylastis Parasite Adults; Arthropods|Larvae; Arthropods|Pupae
Rhizophagus bipustulatus Predator New Zealand Pinus
Rhizophagus dispar Predator New Zealand Pinus
Rhizophagus ferrugineus Predator New Zealand Pinus
Rhopalicus tutele Parasite New Zealand Pinus
Thanasimus formicarius Predator Adults; Arthropods|Larvae New Zealand Pinus

Notes on Natural Enemies

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Herting (1973) lists 11 species (in the families Staphylinidae, Histeridae, Nitidulidae, Cleridae, Raphididae and Asilidae) as predators but it is not clear whether these have been specifically recorded from H. ater or from other species of bark beetles in similar niches.

In 1976, the adults and larvae of Thanasimus formicarius were imported into New Zealand from Austria. The adults were successfully reared, and between 1977 and 1987 over 12,000 adults were released in numerous pine plantations. The first recoveries were made in 1984 (Faulds, 1989).

Rhopalicus tutele and Dinotiscus eupterus may be parasitic on H. ater in Europe. In 1975, numbers of both of these species were shipped to New Zealand from Austria as potential biological control agents. They were reared from logs from which H. ater constituted less than 1.7% of emerging bark beetles compared with over 90% for Hylurgops palliatus (Faulds, 1989).

Ten species of nematodes (Allantonema morosa, Parasitylenchus kleini, Contortylenchus cunicularii, Neoditylenchus panurgus, Cryptaphelenchus koerneri, Ektaphelenchus hylastophilus, Parasitaphelenchus uncinatus, Parasitylenchus hylastis, Micoletzkya thalenhorsti and Parasitorhabditis ateri) are associated with H. ater in Europe. Except for P. hylastis, the nature of the relationships with H. ater is unknown and it may be just phoretic. Three of these species: P. hylastis, M. thalenhorsti and P. ateri have also been found in New Zealand as has Bursaphelenchus eggersi which is not associated with H. ater in Europe but is found with Hylurgops palliatus. Anguilluloides zondagi has also been found associated with H. ater in New Zealand (Dale, 1967).

Means of Movement and Dispersal

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H. ater is a strong flier that disperses readily and responds to host volatiles.

Plant Trade

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Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Roots arthropods/adults; arthropods/eggs; arthropods/larvae; arthropods/pupae Yes Pest or symptoms usually visible to the naked eye
Seedlings/Micropropagated plants arthropods/adults Yes Pest or symptoms usually visible to the naked eye
Stems (above ground)/Shoots/Trunks/Branches arthropods/adults; arthropods/eggs; arthropods/larvae; arthropods/pupae Yes Pest or symptoms usually visible to the naked eye

Wood Packaging

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Wood Packaging liable to carry the pest in trade/transportTimber typeUsed as packing
Solid wood packing material with bark Conifers, mainly Pinus spp. No
Wood Packaging not known to carry the pest in trade/transport
Loose wood packing material
Non-wood
Processed or treated wood
Solid wood packing material without bark

Impact

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Bevan (1987) stated that the damage caused by H. ater in Britain is severe and noted that over the years it has played a major role in causing planting losses, particularly of pine. Bevan also noted that it has proved to be rather more aggressive under British conditions than in many other parts of Europe. Munro (1917) recorded 45% mortality of newly planted Pinus sylvestris in a plantation in Britain.

Neumann (1979) stated that the damage is sometimes severe on nursery stock and young plantings in Australia but Elliott et al. (1998) noted that it is a relatively minor pest of Pinus spp., particularly Pinus radiata. However, Elliott also noted that in the early 1990s, 770 ha of second rotation P. radiata in Tasmania sustained numerous deaths of young seedlings.

Ciesla (1988) reported that up to 70% of pine seedlings in some areas of Chile have died as a result of H. ater attack.

Zondag (1965) reported that H. ater was the most troublesome insect in P. radiata regeneration in New Zealand and recorded over 50% mortality but with the qualification that it was only significant in poorly stocked areas. This implies that he was referring to natural regeneration and not planted areas. Clark (1932) reported that it caused severe loss in a young pine stand that had been re-established after logging operations.

Reay et al. (2001) discussed seedling mortality surveys that had been conducted within 1 year following planting in the central North Island of New Zealand. Seedling mortality in most forest compartments was less than 5% but was as high as 30% in a few compartments. The destructive sampling of seedlings showed that there was a high level of sub-lethal attack on seedlings, it was greater than 50% in two-thirds of the compartments sampled. Similar results were reported from a trial in the South Island of New Zealand much earlier, where 82% of seedlings had superficial resin-encrusted wounds on the stem base and larger roots (New Zealand Forest Service, 1970). Obviously the sub-lethal or abortive attack on seedlings is very common and it would seem that the presence of feeding damage or the beetles themselves on dead seedlings is not conclusive evidence of cause and effect. Milligan (1978) suggested that in general healthy seedlings do not succumb to adult feeding and that those seedlings that do die are in some way debilitated or weakened by environmental factors such as extreme climatic conditions and poorly aerated soils. Several writers have observed that the quality of nursery stock and its treatment prior to and during planting, affect its susceptibility to damage by H. ater (e.g. Munro, 1917; Clark, 1932).

Reay et al. (2001) suggested that there might be ramifications from the adult feeding on seedlings on future wood quality because it is known that the beetles can carry sapstain fungi to seedlings. The following species of sapstain fungi have been isolated from H. ater in New Zealand: Ophiostoma ips, Ophiostoma setosum, Ophiostoma querci, Ophiostoma huntii, Ophiostoma galeiformis, Ophiostoma pluriannulatum, Leptographium truncatum and Leptographium procerum. The following fungi have been isolated from surface-sterilized seedlings following attack by H. ater: O. galeiformis, O. huntii, O. setosum, O. querci, Ophiostoma floccosum, Ophiostoma piceae, L. procerum and L. truncatum (MacKenzie, 1992; Reay et al., 2001). It would be of interest to know what species of fungi could be isolated from seedlings that have not been attacked by H. ater. Clearly more work is warranted.

Pasek (1998) suggested that there is the potential for H. ater to vector root diseases associated with intensive management. In England, several species of sapstain fungi can be consistently isolated from brood galleries of H. ater. These include Ophiostoma coerulescens, Ophiostoma penicillata (Dowding, 1973), Leptographium serpens and Leptographium sp. (Wingfield and Gibbs, 1991).

In New Zealand, infested logs are either refused for export or must be treated (usually by fumigation) immediately before shipping. Green sawn timber for export may also have to be treated (fumigation or kiln sterilization) if the adults that have been attracted to the freshly sawn wood are present. Sapstain fungi are transmitted by the beetles to the outer sapwood of logs but rarely significantly penetrate unless there is abnormal delay between felling and sawing (Milligan, 1978).

Environmental Impact

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Although the possible economic damage caused by this insect would not cause environmental problems, control measures using pesticides, if warranted, would raise environmental concerns.

Detection and Inspection

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Adult feeding on seedlings can be detected by a close examination of the root collar. This is greatly facilitated by removing the seedlings from the ground.

Removal of the bark from stumps, larger roots, logs and sawn timber will reveal the presence of insects under the bark.

Reay and Walsh (2002a) have shown that H. ater is attracted to raw turpentine and beta-pinene with the addition of ethanol but was not attracted to alpha-pinene, either in combination with ethanol or alone. Eight-funnel traps (Lindgren, 1983) were used.

Similarities to Other Species/Conditions

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H. ater is most closely related to Hylastes brunneus and they are difficult to distinguish. In H. brunneus the pronotum is less elongate and broadest behind the middle. See Beaver (1970) for further details.

Prevention and Control

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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.

Infestations in logs may be minimized by:

- The rapid turnover of stockpiles in the forest.

- A selection of unshaded skid sites.

- The stacking of logs on skids rather than on the ground (Milligan, 1978).

Logs for export can be mechanically debarked and treated with an insecticide (USDA, 1992).

Green sawn timber that is intended for export can either be kiln sterilized just prior to shipping or treated with an insecticidal dip (Milligan, 1978).

Seedlings can be protected from adult feeding damage by the use of controlled release granular insecticides (Reay and Walsh, 2002b). However, this measure is not usually necessary.

Biological control agents were imported into New Zealand, but only Thanasimus formicarius established. Because the larvae of H. ater are principally below ground, they are protected from natural enemies and T. formicarius is ineffective (Faulds, 1989).

References

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Beaver RA, 1970. On Hylastes ater Payk. and H. brunneus Er. (Coleoptera: Scolytidae), with a description of the larva of H. ater. The Entomologist, 103(1287):198-206.

Bevan D, 1987. Forest insects: a guide to insects feeding on trees in Britain. Forestry Commission Handbook, UK, No. 1:153 pp.

Blair KG, 1949. Hylastes brunneus Er. (Col., Scolytidae) in Britain. Entomologist's Monthly Magazine, 85:89.

Bright DE; Skidmore RE, 1997. A catalog of Scolytidae and Platypodidae (Coleoptera), Supplement 1 (1990-1994). Ottawa, Canada: NRC Research Press, 368 pp.

Bright DE; Skidmore RE, 2002. A catalogue of Scolytidae and Platypodidae (Coleoptera), Supplement 2 (1995-1999). Ottawa, Canada: NRC Research Press, 523 pp.

Browne FG, 1968. Pests and diseases of forest plantation trees: an annotated list of the principal species occurring in the British Commonwealth. Oxford, UK: Clarendon Press.

Ciesla WM, 1988. Pine bark beetles: a new pest management challenge for Chilean foresters. Journal of Forestry, 86(12):27-31

Clark AF, 1932. The pine-bark beetle, Hylastes ater, in New Zealand. New Zealand Journal of Science and Technology, 14:1-20.

Dale PS, 1967. Nematodes associated with the pine-bark beetle, Hylastes ater, in New Zealand. New Zealand Journal of Science, 10:222-234.

Dowding P, 1973. Effects of felling time and insecticidal treatment on the interrelationships of fungi and arthropods in pine logs. Oikos, 24:422-429.

Duffy EAJ, 1953. Handbooks for the Identification of British Insects. Coleoptera: Scolytidae and Platypodidae. London, UK: Royal Entomological Society of London, 5(15).

Elliott HJ; Ohmart CP; Wylie FR, 1998. Insect Pests of Australian Forests: Ecology and Management. Melbourne, Australia: Inkata Press.

Hansen V, 1955. Notes on some species of Hylastes Er. and Trypophloeus Fairm. (Coleopt. Scolytidae). Entomologiske Meddeleiser, 27:169-185.

Herting B, 1973. A catalogue of parasites and predators of terrestrial arthropods, Section A, Volume III, Coleoptera to Strepsiptera. Wallingford, UK: CAB International.

Jakaitis B; Valenta V, 1976. Faunistic complexes of invertebrates living under the bark of pine stumps in the forests of the Lithuanian SSR. Acta Entomologica Lituanica, 3:11-26

Lekander B, 1965. On Hylastes ater Payk. and Hylastes brunneus Er. (Col. Scolytidae). Entomologisk Tidskrift, 86:184-185.

Lekander B, 1968. Scandinavian bark beetle larvae. Royal College of Forestry, Sweden, Research Notes, 4:1-186.

Lindgren BS, 1983. A multiple funnel trap for scolytid beetles (Coleoptera). Canadian Entomologist, 115(3):299-302

MacKenzie M, 1992. Potential forest exports and pests from New Zealand. In Log imports and introduced forest pests into the Pacific Northwest. Symposium Proceedings. Oregon State University, Corvallis, USA, 21-23 April 1992.

Milligan RH, 1970. Overseas wood- and bark-boring insects intercepted at New Zealand ports [in 1948-65]. Technical Paper, Forest Research Institute, New Zealand Forest Service, No. 57:80 pp.

Milligan RH, 1978. Hylastes ater (Paykull) (Coleoptera: Scolytidae): Black pine bark beetle. Forest and timber insects in New Zealand. No. 29. Rotorua, New Zealand: New Zealand Forest Service, Forest Research Institute.

Munro JW, 1917. The genus Hylastes Er., and its importance in forestry: a study in scolytid structure and biology. Proceedings of the Royal Physical Society of Edinburgh, 20:123-158.

Neumann FG, 1979. Insect pest management in Australian radiata pine plantations. Australian Forestry, 42(1):30-38

New Zealand Forest Service, 1970. Report of Forest Research Institute for 1 January to 31 December 1969. Wellington, New Zealand: New Zealand Forest Service.

Ojeda Gomez P, 1985. Hylastes ater (Paykull). Programma de control de plagas y enfermadades forestales. Santiago, Chile: Corporacion Nacional Forestal.

Pasek JE, 1998. Hylastes ater (Paykull). USDA-APHIS, North Carolina, USA. http://www.exoticforestpests.org/english/Detail.CFM?tblEntry__PestID=21.

Reay SD; Walsh PJ, 2001. Observations on the flight activity of Hylastes ater and Hylurgus ligniperda (Curculionidae: Scolytinae) in Pinus radiata forests in the central North Island, New Zealand. New Zealand Entomologist, 24:79-85.

Reay SD; Walsh PJ, 2002. A carbosulfan insecticide to protect pine seedlings from Hylastes ater (Coleoptera: Scolytidae) damage. New Zealand Plant Protection, 55:80-84.

Reay SD; Walsh PJ, 2002. Relative attractiveness of some volatiles to the introduced pine bark beetles, Hylastes ater and Hylurgus ligniperda (Curculionidae: Scolytinae). New Zealand Entomologist, 25:51-56.

Reay SD; Walsh PJ; Thwaites JM; Farrell; RL, 2001. The black pine bark beetle Hylastes ater in Zealand. New Zealand Tree Grower, November 2001, 32-33.

Reay, S. D., Thwaites, J. M., Farrell, R. L., 2005. A survey of Ophiostoma species vectored by Hylastes ater to pine seedlings in New Zealand. Forest Pathology, 35(2), 105-113. doi: 10.1111/j.1439-0329.2004.00393.x

Schedl KE, 1968. Die Gattung Hylastes Er. 256 Beitrag. Anzeiger für Schadlingskunde, 41:155-158.

USDA, 1992. Pest risk assessment of the importation of Pinus radiata and Douglas-fir logs from New Zealand. USDA Forest Service Miscellaneous Publication 1508.

Voolma K; Ounap H; Süda I, 1996. Bark beetles (Coleoptera, Scolytidae) in the insect collections of Estonia. Metsanduslikud Uurimused, 27:125-132; 31 ref.

Wingfield MJ; Gibbs JN, 1991. Leptographium and Graphium species associated with pine-infesting bark beetles in England. Mycological Research, 95(11):1257-1260

Wood SL, 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. Great Basin Naturalist Memoirs, No. 6:1359 pp.

Wood SL; Bright DE, 1992. A catalog of Scolytidae and Platypodidae (Coleoptera), Part 2: Taxonomic index. Great Basin Naturalist Memoirs, 13: 1-1553.

Zondag R, 1965. Insect pests of forest nurseries and younf plantations in New Zealand. Proceedings of the 12th International Congress of Entomology, London, UK, 674-675.

Distribution References

Bright DE, Skidmore RE, 1997. A catalog of Scolytidae and Platypodidae (Coleoptera), (1990-1994)., Ottawa, Canada: NRC Research Press. 368 pp.

Bright DE, Skidmore RE, 2002. A catalogue of Scolytidae and Platypodidae (Coleoptera), Supplement 2 (1995-1999)., Ottawa, Canada: NRC Research Press. 523 pp.

Browne FG, 1968. Pests and diseases of forest plantation trees: an annotated list of the principal species occurring in the British Commonwealth., Oxford, UK: Clarendon Press.

CABI, Undated. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

Clark A F, 1932. The Pine-bark Beetle, Hylastes ater, in New Zealand. New Zealand Journal of Science and Technology. 14 (1), 1-20 pp.

Elliott HJ, Ohmart CP, Wylie FR, 1998. Insect Pests of Australian Forests: Ecology and Management., Melbourne, Australia: Inkata Press.

Jakaitis B, Valenta V, 1976. Faunistic complexes of invertebrates living under the bark of pine stumps in the forests of the Lithuanian SSR. Acta Entomologica Lituanica. 11-26.

Ojeda Gomez P, 1985. Hylastes ater (Paykull).,

Reay S D, Thwaites J M, Farrell R L, 2005. A survey of Ophiostoma species vectored by Hylastes ater to pine seedlings in New Zealand. Forest Pathology. 35 (2), 105-113. DOI:10.1111/j.1439-0329.2004.00393.x

Seebens H, Blackburn T M, Dyer E E, Genovesi P, Hulme P E, Jeschke J M, Pagad S, Pyšek P, Winter M, Arianoutsou M, Bacher S, Blasius B, Brundu G, Capinha C, Celesti-Grapow L, Dawson W, Dullinger S, Fuentes N, Jäger H, Kartesz J, Kenis M, Kreft H, Kühn I, Lenzner B, Liebhold A, Mosena A (et al), 2017. No saturation in the accumulation of alien species worldwide. Nature Communications. 8 (2), 14435. http://www.nature.com/articles/ncomms14435

Voolma K, Õunap H, Süda I, 1996. Bark beetles (Coleoptera, Scolytidae) in the insect collections of Estonia. (Ürasklased (Coleoptera, Scolytidae) eesti entomoloogilistes kollektsioonides.). Metsanduslikud Uurimused. 125-132.

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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