Invasive Species Compendium

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Datasheet

Dendroctonus micans
(great spruce bark beetle)

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Datasheet

Dendroctonus micans (great spruce bark beetle)

Summary

  • Last modified
  • 28 March 2018
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Preferred Scientific Name
  • Dendroctonus micans
  • Preferred Common Name
  • great spruce bark beetle
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta

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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
Adults exposed under bark.
TitleAdults
CaptionAdults exposed under bark.
CopyrightAsko Lehtijarvi
Adults exposed under bark.
AdultsAdults exposed under bark.Asko Lehtijarvi

Identity

Top of page

Preferred Scientific Name

  • Dendroctonus micans (Kugelann, 1794)

Preferred Common Name

  • great spruce bark beetle

Other Scientific Names

  • Bostrichus micans Kugelann, 1794
  • Hylesinus ligniperda Gyllenhal, 1813

International Common Names

  • English: beetle, European spruce
  • French: hylesine geant; le dendroctonus geant de l'epicea; le hylesine geant de l'epicea

Local Common Names

  • Denmark: kjemperbarkbille
  • Finland: ukkoniluri
  • Germany: Riesenbastkaefer; Riesenbastkäfer
  • Netherlands: Sparrebastkever
  • Norway: kjemperbarkbille
  • Sweden: jättebastborre

EPPO code

  • DENCMI (Dendroctonus micans)

Taxonomic Tree

Top of page
  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Coleoptera
  •                         Family: Scolytidae
  •                             Genus: Dendroctonus
  •                                 Species: Dendroctonus micans

Notes on Taxonomy and Nomenclature

Top of page D. micans was originally described as Bostrichus micans by Kugelann in 1794 and as Hylesinus ligniperda by Gyllenhal in 1813 (Grüne, 1979). This insect was designated as the type species of the genus Dendroctonus, when Erichson described the genus in 1836 (Wood, 1982).

Description

Top of page Eggs

Scolytidae eggs are smooth, ovoid, white and translucent. They are 1.2 mm long and deposited in clusters of 100-150 in the egg gallery.

Larvae

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 abdominal segment has two to three tergal folds and the pleuron is not longitudinally divided. The larvae do not change as they grow. Spruce beetle larvae have four larval instars and are 4-6 mm long when mature (Holsten et al., 1989).

Pupae

Scolytid pupae are white and mummy-like. They are exarate, with legs and wings free from the body. Some species have paired abdominal urogomphi. The elytra are either rugose or smooth, sometimes with a prominent head and thoracic tubercles.

Adults

The adults are 6-9 mm long, dark-brown and cylindrical. The legs and antennae are yellow-brown. The head is visible when viewed dorsally and the elytral declivity is smooth and rounded.

Distribution

Top of page

D. micans is believed to have originated in the conifer forests of Asia. When determining the geographic distribution of this insect it is difficult to establish whether it is native or introduced because of its unique history.

It has steadily spread westward over the past 100 years, undoubtedly aided by the increased trade in unprocessed logs. At present, it is found throughout Eurasia and has adapted to a wide range of forest conditions. This insect is now established across most of western Europe from European Russia, west to Belgium and France, south to Turkey, and north to Finland and Sweden. It was discovered in the UK in 1982.

Distribution Table

Top 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.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ChinaPresentEPPO, 2014
-HeilongjiangPresentEPPO, 2014
-LiaoningPresentEPPO, 2014
-QinghaiPresentEPPO, 2014
-SichuanPresentEPPO, 2014
Georgia (Republic of)PresentIntroduced Invasive Kobakhidze, 1965; EPPO, 2014
JapanRestricted distributionEPPO, 2014
-HokkaidoPresentEPPO, 2014
TurkeyRestricted distributionIntroduced Invasive DKOA, 2001; EPPO, 2014

Europe

AustriaPresentWood and Bright, 1992; EPPO, 2014
BelgiumPresentIntroduced Invasive Wood and Bright, 1992; EPPO, 2014
Bosnia-HercegovinaPresentEPPO, 2014
BulgariaWidespread****Wood and Bright, 1992; EPPO, 2014
CroatiaRestricted distributionEPPO, 2014
Czech RepublicWidespread****Wood and Bright, 1992; EPPO, 2014
DenmarkRestricted distributionWood and Bright, 1992; EPPO, 2014
EstoniaRestricted distributionBright and Skidmore, 2002; EPPO, 2014
FinlandRestricted distributionBright and Skidmore, 2002; EPPO, 2014
FranceRestricted distributionIntroduced Invasive Grégoire et al., 1989; Grégoire, 1988; EPPO, 2014
GermanyPresent, few occurrences193*Wood and Bright, 1992; EPPO, 2014
GreeceAbsent, confirmed by surveyWood and Bright, 1992; EPPO, 2014
HungaryRestricted distribution****Wood and Bright, 1992; EPPO, 2014
IrelandAbsent, confirmed by surveyEPPO, 2014
ItalyRestricted distribution****Battisti, 1984; Wood and Bright, 1992; EPPO, 2014
LatviaPresentBright and Skidmore, 2002
LithuaniaPresent, few occurrencesEPPO, 2014
LuxembourgPresentWood and Bright, 1992; EPPO, 2014
NetherlandsRestricted distributionNPPO of the Netherlands, 2013; Wood and Bright, 1992; EPPO, 2014
NorwayWidespread****Wood and Bright, 1992; EPPO, 2014
PolandRestricted distribution****Wood and Bright, 1992; EPPO, 2014
PortugalAbsent, confirmed by surveyEPPO, 2014
RomaniaRestricted distributionWood and Bright, 1992; EPPO, 2014
Russian FederationRestricted distributionNativeWood and Bright, 1992; EPPO, 2014
-Central RussiaPresentNativeWood and Bright, 1992; EPPO, 2014
-Eastern SiberiaPresentNativeWood and Bright, 1992; EPPO, 2014
-Northern RussiaPresentNativeWood and Bright, 1992; EPPO, 2014
-Russian Far EastPresentNativeWood and Bright, 1992; EPPO, 2014
-Southern RussiaPresentNativeWood and Bright, 1992; EPPO, 2014
-Western SiberiaPresentNativeWood and Bright, 1992; EPPO, 2014
SerbiaPresentEPPO, 2014
SlovakiaRestricted distributionEPPO, 2014; Vakula et al., 2016
SloveniaPresentJurc, 2006
SpainAbsent, confirmed by surveyWood and Bright, 1992; EPPO, 2014
SwedenWidespread****Wood and Bright, 1992; EPPO, 2014
SwitzerlandRestricted distribution****Wood and Bright, 1992; EPPO, 2014
UKRestricted distributionEPPO, 2014
-Channel IslandsAbsent, confirmed by surveyEPPO, 2014
-England and WalesRestricted distributionIntroduced Invasive Bevan and King, 1983; Evans and King, 1989; EPPO, 2014
-Northern IrelandAbsent, confirmed by surveyEPPO, 2014
-ScotlandAbsent, confirmed by surveyEPPO, 2014
UkraineRestricted distributionEPPO, 2014
Yugoslavia (Serbia and Montenegro)PresentWood and Bright, 1992

Risk of Introduction

Top of page The adults can fly at least 2 to 3 km in search of new hosts but prefer to attack either the same trees in which they developed or immediately adjacent trees. The adults can disperse on air currents. Other life stages are confined to the cambium layer and inner bark, and do not naturally disperse.

Pathways for human-assisted dispersal include the transport of unprocessed pine logs or lumber, crates, pallets and dunnage, containing bark strips. It is conceivable that larvae, pupae and overwintering adults could survive an ocean voyage and be introduced into a new location. Should this new location contain forests with a component of spruce, D. micans could become established and cause severe damage. The related North American red turpentine beetle, Dendroctonus valens, has recently been introduced to and established in, China, and has killed more than 6 million pines in recent years (Sun et al., 2003).

Most Eurasian conifer forests now have populations of D. micans. However, this insect is not present in North America. Therefore, North American conifer forests are at risk from the introduction and establishment of this insect.

Habitat

Top of page D. micans normally occurs in mature spruce forests. Within its natural range, this insect typically causes low levels of tree mortality and is not considered to be a major pest.

Hosts/Species Affected

Top of page D. micans primarily breeds in spruce (Picea spp.), especially Picea abies (Norway spruce), Picea sitchensis (Sitka spruce) and Picea orientalis (Oriental spruce) (Grégoire, 1988). It also breeds in other Eurasian spruce species such as Picea asperata (dragon spruce), Picea crassifolia (Qinghai spruce), Picea ajanensis [Picea jezoensis] (ezo spruce), Picea obovata (Siberian spruce) and Picea omorika (Serbian spruce), as well as in several North American spruce species that have been introduced into its geographic range. These include Picea breweriana (Brewer spruce), Picea engelmannii (Engelman spruce), Picea glauca (white spruce), Picea mariana (black spruce) and Picea pungens (blue spruce) (Grégoire, 1988; Evans and King, 1989). Pines (Pinus spp.), especially Pinus sylvestris (Scotch pine), have also been attacked, especially in Estonia, Poland, Scandinavia and Siberia (Kolomiets and Isaev, 1981; Markov, 1985; Voolma, 1993).

D. micans is also known to attack several other pines, including Pinus contorta (lodgepole pine), Pinus nigra (black pine), Pinus sylvestris var. hamata, Pinus strobus (white pine) and Pinus uncinata (mountain pine). It also attacks fir trees; Abies alba (silver fir), Abies nordmanniana (Nordmann fir), Abies sibirica (Siberian fir) and Pseudotsuga menziesii (Douglas fir); and larch, Larix decidua (European larch) (Grégoire, 1988; Smith et al., 1992; Wood and Bright, 1992; Wainhouse and Beech-Garwood, 1994).

Growth Stages

Top of page Vegetative growing stage

Symptoms

Top of page Heavily infested trees have large numbers of conspicuous purplish-brown pitch tubes on the bark surface of the lower boles. Lightly infested trees usually survive beetle attack. Therefore, several generations of beetles could emerge from a tree that still has green foliage. The bark often easily peels away in areas where the larvae have consumed the inner bark. Removal of the bark from infested portions of trees will reveal characteristic galleries, larvae, pupae and adults (Bevan and King, 1983; Grégoire, 1988).

List of Symptoms/Signs

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SignLife StagesType
Leaves / yellowed or dead
Stems / gummosis or resinosis
Stems / internal feeding
Stems / necrosis
Whole plant / frass visible
Whole plant / internal feeding
Whole plant / plant dead; dieback

Biology and Ecology

Top of page Life History and Habits

The genus Dendroctonus consists of 19 species, worldwide. Most occur on conifers in North and Central America. Dendroctonus armandi (native to China) and D. micans are found in Palearctic conifer forests (Wood, 1982). Several species are important forest pests, capable of reaching epidemic levels and killing thousands of trees. The genus Dendroctonus contains some of the most destructive forest insects in North and Central America, including the southern pine beetle (Dendroctonus frontalis), mountain pine beetle (Dendroctonus ponderosae), Douglas-fir beetle (Dendroctonus pseudotsugae) and the spruce beetle (Dendroctonus rufipennis).

D. micans exhibits a number of life cycle characteristics that make it unique among Dendroctonus species. Mating takes place under the bark prior to emergence and before the adult beetles are fully chitinized. Sibling males normally mate the females (incestuous mating). The ratio of males to females is low. Typically the sex ratio is one male per 10 females but can be as low as one male per 45 females. The phenomenon of pre-emergence mating precludes the need for females to attract males. Therefore, there is no adult aggregation pheromone.

The adult beetles can remain underneath the bark of the trees in which they developed for long periods, if the conditions for emergence are not suitable. They often mine in large groups, among their original larval galleries, chewing the larval frass and sometimes forming 'nose to tail' columns within the brood excavations. When emergence does occur, the adults cut round emergence holes through the thin bark that covers the brood system. The emergence holes can be constructed well ahead of the actual emergence and large quantities of powdery frass are ejected. Emergence can occur over a protracted period, with many beetles using the same emergence hole.

The mated females emerge to attack either new trees or unattacked portions of the host tree from which they emerged. Adult flight and, more commonly, walking, play an important part in adult dispersal. This typically leads to small groups of attacked trees. Sometimes no adult emergence occurs and new brood areas are established in the same tree, along the margins of existing galleries.

D. micans is different from the more aggressive North American Dendroctonus species in that it usually attacks its hosts in low numbers, killing the bark in patches. Successive attacks, over a period of 5 to 8 years, may be necessary to kill a tree, except during outbreaks.

The temperature threshold for adult flight is reported to be 21-23°C. However, in Britain, initial flight at 20°C with sustained flight at 14°C has been observed.

Beetle attacks often occur around areas of damage on a tree, which may have been caused by lightning or logging. Attacks are often associated with decreased resin pressure and commonly occur in forked or multi-stemmed trees, just below the branch nodes. In some countries, there appears to be an association between beetle attack and the occurrence of root disease caused by fungi such as Heterobasidium annosum or Armellaria sp.. However, apparently healthy trees are also commonly attacked.

The female bores through the bark and establishes a brood chamber. She clears the resin that accumulates during the attack process by mixing it with frass and expelling it through the entrance hole. The expelled resin mixed with frass is purplish-brown and gives rise to resin tubes, which are characteristic of D. micans. These can be seen on the bark surface of infested trees. When the female reaches the cambium, she bores upwards for approximately 2 cm, constructs an egg chamber and deposits a cluster of between 100 and 150 eggs. These are covered with frass and wood dust. Then she may produce additional egg chambers, leading to a mix of several larval instars in the same family group, or she may attack other portions of the tree or adjacent trees.

Newly hatched larvae feed gregariously, side by side, in a brood gallery that becomes larger as the larvae feed. The size of the brood gallery varies according to the number of larvae present. A large brood of larvae can construct a gallery that is 30-60 cm long and 10-20 cm wide. When several females oviposit close to each other, the individual galleries coalesce. This can cause extensive injury to the tree.

The larval colony feeds upwards and outwards from its origin. The frass and dead bodies of siblings are tightly packed into the area behind the feeding front. The larvae produce an aggregation pheromone; a mixture of trans- and cis-verbenol, verbenone and myrtenol, which sustains larval aggregation. There are five larval instars. When feeding is completed, they move back into the islands of tightly packed frass and construct single pupal chambers.

The time required for D. micans to complete one generation in the field ranges from 1 to 3 years, depending on local temperatures. In Great Britain, the time to complete a generation ranges from 10 to 18 months (Fielding and Evans, 1997). In Turkey and the Former Soviet Republic of Georgia, 12 to 15 months are required to complete a generation and in the Nordic countries, 2 to 3 years may be required (Grégoire, 1988).

D. micans does not appear to be associated with any major pathogenic fungi (Serez, 1979; Kolomiets and Isaev, 1981; Grégoire, 1988; King and Fielding, 1989).

Natural enemies

Top of page
Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Bacillus thuringiensis thuringiensis Pathogen
Beauveria bassiana Pathogen
Bracon hylobii Parasite Estonia Picea abies; Pinus sylvestris
Dolichomitus tenebrans Parasite
Dolichomitus terebrans Parasite Pupae Estonia Picea abies; Pinus sylvestris
Dromius quadrimaculatus Predator Estonia Picea abies; Pinus sylvestris
Lonchaea collini Parasite Estonia Picea abies; Pinus sylvestris
Melanotus villosus Predator Estonia Picea abies; Pinus sylvestris
Nudobius lentus Predator Estonia Picea abies; Pinus sylvestris
Placusa depressa Predator Estonia Picea abies; Pinus sylvestris
Raphidia ophiopsis Predator Estonia Picea abies; Pinus sylvestris
Raphidia xanthostigma Predator Estonia Picea abies; Pinus sylvestris
Rhizophagus grandis Predator Larvae Estonia; Italy; Republic of Georgia; UK Picea; Picea abies; Pinus sylvestris
Scoloposcelis pulchella Predator Estonia Picea abies; Pinus sylvestris
Thanasimus dubius Predator USSR
Thanasimus femoralis Predator Estonia Picea abies; Pinus sylvestris
Thanasimus formicarius Predator Estonia Picea abies; Pinus sylvestris

Notes on Natural Enemies

Top of page D. micans has few natural enemies. This may be due to its unique biology, which seems to protect it from competitors and generalist natural enemies (Everaerts et al., 1988). One specific predator, Rhizophagus grandis, is abundant in areas where D. micans has been present for long periods of time. This beetle is believed to be responsible for maintaining a low and stable D. micans population in these areas (Kobakhidzi, 1965; Moeck and Safranyik, 1983). Woodpeckers (Picoides major) prey on the larvae and pupae.

Plant Trade

Top of page
Plant parts liable to carry the pest in trade/transportPest stagesBorne internallyBorne externallyVisibility of pest or symptoms
Bark adults; eggs; larvae; nymphs; pupae 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
Plant parts not known to carry the pest in trade/transport
Bulbs/Tubers/Corms/Rhizomes
Flowers/Inflorescences/Cones/Calyx
Fruits (inc. pods)
Growing medium accompanying plants
Leaves
Roots
Seedlings/Micropropagated plants
True seeds (inc. grain)
Wood

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 Picea spp., Pinus spp. Yes
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

Top of page Within most of its natural range, D. micans normally occurs at low levels and causes little tree mortality. However, occasionally outbreaks do occur and result in widespread tree mortality. Most outbreaks occur along the leading edge of the geographic range of D. micans and not within the interior portion of the range. Trees are killed as a result of the girdling action of larval feeding. This can take place over a period of several years. As D. micans extended its range westward into Europe (France and the UK) and southwestern Asia (Georgia and Turkey) during the late 1900s, outbreaks occurred on more than 200,000 ha of spruce forests (Grégoire, 1988; Vouland and Schvester, 1994; Konca, 1995). In some cases, older trees were preferentially attacked (Carle et al., 1979) while in other instances, all age classes of spruce trees were attacked (Battisti, 1984; Benz, 1984; Evans et al., 1984). Similarly, when D. micans attacked Pinus sylvestris in Estonia (Voolma, 1978) and Siberia (Kolomiets and Bogdanova, 1976), both young and old trees were infested. Normally, D. micans only colonizes green standing trees, but it will attack trees that are stressed as a result of logging damage, frost, snow, wind, lightning, poor soil nutrition and drought (Chararas 1960; Bejer-Petersen, 1976; Novak, 1976; Voolma, 1978; Battisti, 1984; Shavliashvili and Zharkov, 1985; Gabeev and Gnat, 1986; Grégoire, 1988).

Environmental Impact

Top of page Sustained attack on individual trees can result in tree mortality. Widespread outbreaks are common, especially along the leading edge of the D. micans range. Outbreaks have occurred in spruce forests and, to a lesser extent, in Scotch pine forests. In some cases more than 50% tree mortality has occurred. Outbreaks are often more common in forests that are stressed by drought, poor soil nutrition and logging damage, for example.

Detection and Inspection

Top of page The bark surface should be inspected for pitch tubes and/or boring dust. The cambium and inner bark of unprocessed logs, or dunnage, crates or pallets, containing bark strips, should be inspected for the presence of galleries and insect life stages.

Similarities to Other Species/Conditions

Top of page Morphologically, D. micans is difficult to distinguish from the North American species, Dendroctonus punctatus and the two species were thought to be conspecific. However, it has now been established that they are distinct species (Furniss, 1996; Kegley et al., 1997). The communal larval galleries of D. micans resemble the galleries of the North American black turpentine beetle, Dendroctonus terebrans, the red turpentine beetle, Dendroctonus valens and the Mexican species, Dendroctonus rhizophagus. However, all of the latter species confine their attacks to Pinus spp.. To ensure positive identification, a taxonomist, with expertise in the family Scolytidae, should examine bark beetles believed to be a new introduction.

Prevention and Control

Top of page Cultural Control

The felling, removal and rapid processing of infested trees to destroy broods is a widely used control method for D. micans. The thinning of overstocked forests will help to reduce their susceptibility to attack.

Biological Control

Classical biological control, involving the mass rearing and release of the predaceous beetle, Rhizophagus grandis, has been used in France (Grégoire et al., 1989, 1992; van Averbeke and Grégoire, 1995), the Former Soviet Republic of Georgia (Tvaradze, 1984; Evans, 1987), Turkey (Alkan and Aksu, 1990) and the UK (King and Evans, 1984; Fielding et al., 1991; Evans and Fielding, 1996; Fielding and Evans, 1997).

Chemical Control

Insecticide application to infested portions of the lower boles of trees has been undertaken but with questionable success. For example, extensive chemical control operations were undertaken in Turkey between 1967 and 1985. However, treatments were not successful, except for a slight decrease in the populations of the pest (DKOA, 2001).

Pheromonal Control

Since this insect does not produce an aggregating pheromone, pheromonal control is not a viable control method.

Field Monitoring

In the UK, surveys designed for the early detection of new infestations or for general forest health monitoring have failed to detect infestations until the beetle has been present for approximately 3 years and has become well-established. However, ground surveys, especially near areas of old infestations, can be conducted to detect infested trees that should be removed from stands.

D. micans does not produce an aggregating pheromone, therefore the use of pheromones for the early detection of infestations is not feasible.

Integrated Pest Management

Integrated pest management (IPM) of this insect consists of the timely detection of infestations, the rapid removal and processing of infested trees, thinning of overstocked stands to reduce their susceptibility to attack and the release of the predator, Rhizophagus grandis, into areas where this insect has recently spread.

References

Top of page

Alkan S; Aksu Y, 1990. Research on rearing techniques for Rhizophagus grandis Gyll. (Coleoptera, Rhizophagidae). Proceedings of the Second Turkish National Congress of Biological Control Izmir, Turkey; Ege Universitesi, 173-179

Averbeke A van; Grégoire JC, 1995. Establishment and spread of Rhizophagus grandis Gyll. (Coleoptera: Rhizophagidae) 6 years after release in the Foret domaniale du Mezenc (France). Annales des Sciences Forestieres, 52(3):243-250

Battisti A, 1984. Dendroctonus micans (Kugelann) in Italy (Coleoptera Scolytidae). Frustula Entomologica, 7-8:631-637

Bejer-Petersen B, 1976. Dendroctonus micans Kug. in Denmark: the situation 25 years after a 'catastrophe'. Zeitschrift fur Pflanzenkrankheiten und Pflanzenschutz, 83(1/3):16-21

Benz G, 1984. Dendroctonus micans in Turkey: the situation today. In: Proceedings of the EEC Seminar on the Biological Control of Bark Beetles (Dendroctonus micans), Brussels, Belgium, 43-47.

Bevan D; King CJ, 1983. Dendroctonus micans Kug. - a new pest of spruce in U.K. Commonwealth Forestry Review, 62(1):41-51

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

CABI/EPPO, 1998. Distribution maps of quarantine pests for Europe (edited by Smith IM, Charles LMF). Wallingford, UK: CAB International, xviii + 768 pp.

Carle P; Granet AM; Perrot JP, 1979. Contribution to the study of the dispersal and aggressivity of Dendroctonus micans Kug. (Col. Scolytidae) in France. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 52(2/3):185-196

Chararas C, 1960. Variations de la pression osmotique de Picea excelsa a la suite des attaques de Dendroctonus micans Kug. (Coleoptera, Scolytidae). Comptes Rendus Hebdomadaires des Siances de I'Acadgmie des Sciences, 251(18):1917-1919.

DKOA, 2001. Bark beetles. Eastern Black Sea Research Institute, Trabzon, Turkey, http://www.angelfire.com/fl4/yuksel/yukselenglish.htm.

EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm

Evans H, 1987. Biological control of Dendroctonus micans in the USSR. Entopath News, No. 89:5-7

Evans HF; Fielding NJ, 1996. Restoring the natural balance: biological control of Dendroctonus micans in Great Britain. Biological control introductions - opportunities for improved crop production. In: Proceedings of an International Symposium Brighton, UK 18 November 1996., 45-57; 23 ref.

Evans HF; King CJ, 1989. Biological control of Dendroctonus micans (Coleoptera: Scolytidae): British experience of rearing and release of Rhyzophagus grandis (Coleoptera: Rhizophagidae). In: Kulhavy DL, Miller MC , eds. Potential for Biological Control of Dendroctonus and Ips beetles. Texas, USA: Center for Applied Studies, School of Forestry, Stephen F. Austin State University, 109-128.

Evans HF; King CJ; Wainhouse D, 1984. Dendroctonus micans in the United Kingdom. The result of two years experience in survey and control. In: Proceedings of the EEC Seminar on the Biological Control of Bark Beetles (Dendroctonus micans), Brussels, Belgium, 20-34.

Everaerts C; Grégoire JC; Merlin J, 1988. The toxicity of spruce monoterpenes to bark beetles and their associates. In: Mattson WJ, et al. eds. Mechanisms of Woody Plant Resistance to Insects and Pathogens. New York, USA: Springer-Verlag.

Fielding NJ; Evans HF, 1997. Biological control of Dendroctonus micans (Scolytidae) in Great Britain. Biocontrol News and Information, 18(2):51N-60N; 35 ref.

Fielding NJ; Evans HF; Williams JM; Evans B, 1991. Distribution and spread of the great European spruce bark beetle, Dendroctonus micans, in Britain - 1982 to 1989. Forestry (Oxford), 64(4):345-358

Furniss MM, 1996. Taxonomic status of Dendroctonus punctatus and D. micans (Coleoptera: Scolytidae). Annals of the Entomological Society of America, 89(3):328-333; 21 ref.

Gabeev VN; Gnat EV, 1986. Silvicultural and physiological characteristics of trees damaged by Dendroctonus. Izvestiya Sibirskogo Otdeleniya Akademii Nauk SSSR, Biologicheskikh Nauk, No. 3:24-28

Grégoire JC, 1988. The greater European spruce beetle. In: Berryman AA, ed. Dynamics of Forest Insect Populations. New York, USA: Plenum Publishing Corporation, 455-478.

Grégoire JC; Balsier M; Merlin J, 1989. Interactions between Rhizophagus grandis (Coleoptera: Rhizophagidae) and Dendroctonus micans (Coleoptera: Scolytidae) in the field and laboratory: Their application for the biological control of D. micans in France. In: Kulhavy DL, Miller MC, eds. Potential for Biological Control of Dendroctonus and Ips beetles. Texas, USA: Center for Applied Studies, School of Forestry, Stephen F. Austin State University, 95-108.

Grégoire JC; Couillien D; Drumont A; Meyer H; Francke W, 1992. Semiochemicals and the management of Rhizophagus grandis Gyll. (Col., Rhizophagidae) for the biocontrol of Dendroctonus micans Kug. (Col., Scolytidae). Journal of Applied Entomology, 114:110-112.

Grüne S, 1979. Handbuch zur bestimmung der Europäischen Borkenkäfer (Brief illustrated key to European bark beetles). Hannover, Germany: Verlag M & H Schapfer.

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