Pityogenes chalcographus
(sixtoothed spruce bark beetls)

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Identity

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Preferred Scientific Name

• Pityogenes chalcographus (Linnaeus, 1761)

Preferred Common Name

• sixtoothed spruce bark beetls

Other Scientific Names

• Bostrichus bicolor Chevrolat, 1838
• Bostrichus xylographus Sahlberg, 1836
• Dermestes chalcographus Linnaeus, 1761
• Ips chalcographus Linnaeus
• Ips spinosus DeGeer, 1775
• Scolytus sexdentatus Olivier, 1795
• Tomicus chalcographus Linnaeus

International Common Names

• English: bark beetle, six-dentated; engraver, spruce wood
• Spanish: barenillo pequeno de los abetos; barrenillo pequeño de los abetos
• French: bostryche chalcographe; petit rongeur de l'épicéa; petit rongeur du sapin
• Russian: obyknovennyj graver
• Chinese: xue xing keng xiao chong

Local Common Names

• Croatia: bestozubu smrekov potkornjak
• Czech Republic: lykozrout leskly
• Denmark: chalcograf
• Finland: kuusentähtikirjaaja
• Germany: Borkenkaefer, Sechszaehniger; Borkenkaefer, Sechszaehniger Fichten-; Kupferstecher; Sechszähniger Fichtenborkenkäfer
• Hungary: rèzmetszöszù
• Italy: bostrico calcografo
• Netherlands: koperetser
• Norway: liten barkbille
• Poland: rytownnik pospolity
• Serbia: mali trozubi smrcin potkornjak; sesterozubi smrcin potkornja
• Slovakia: lykozrut leskly
• Slovenia: mali smrekov lubadar; sesterozobi smrekov lubadar
• Sweden: sextandad barkborre
• Yugoslavia (former): mal jelkin potkornik

EPPO code

• PITYCH (Pityogenes chalcographus)

Summary of Invasiveness

Top of page P. chalcographus is considered in its native region a serious secondary insect pest, occasionally outbreaking and attacking trees. It can easily establish itself in new areas, mainly in temperate and sub-tropical zones, where the species can find suitable conditions for its development due to its wide polyphagy.

Taxonomic Tree

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• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Arthropoda
•             Subphylum: Uniramia
•                 Class: Insecta
•                     Order: Coleoptera
•                         Family: Scolytidae
•                             Genus: Pityogenes
•                                 Species: Pityogenes chalcographus

Notes on Taxonomy and Nomenclature

Top of page The species was described under the genus Dermestes by Linnaues in 1761 under the species name spelled "calcographus". In the 18th Century different authors spelled its name both ways, "calcographus" and "chalcographus". For more than the last 200 years the spelling commonly used is "chalcographus", which seems to be correctly expressing the original meaning, presumably referring to the gallery system below the bark resembling a copper (chalkos) engraving (grapho). It was transfered by Bedel (1888) into the genus Pityogenes.

It is rather unique within the Scolytids that such a widespread species has so few synonyms. P. chalcographus is a rather distinctive species but can be commonly mixed up with other morphologically related species, such as P. trepanatus, P. bidentatus or other species in museum and private collections.

Description

Top of page Eggs
Round, shiny, whitish.

Larvae
Whitish, curved, apode, 2.7-3.0 mm long in its last instar, there are three larval instars. Lekander (1968) described its morphology. The appearance of the larvae supports the division of this genus into two groups; species in which the females have a pit in their frons (e.g. P. chalcographus and P. trepanatus), and species where the females do not have this pit (e.g. P. bidentatus and P. quadridens).

Adults

Males
The beetle is about 1.6-3.0 mm in length, has a black head and thorax, the elytra has a characteristic red-brownish shine. Frons is simply convex and scarce punctured. Elytral declivity longitudinally impressed in the middle and armed with three strong lateral spines on each side. (Schwerdtfeger, 1957; Postner, 1974; Freude et al., 1981).

Females
Females have the same appearance as males, but frons has a deep transversal impression above epistomal margin and elytral spines are markedly smaller than the males, while the declivity is not so impressed (Schwerdtfeger, 1957; Postner, 1974; Freude et al., 1981).

Distribution

Top of page P. chalcographus is a common bark beetle species throughout the entire natural range of Picea abies in Europe and other Picea species over its distribution in Asia. It also occurs in plantations in western Europe, outside the natural range of the host. Generally, it has trans-Palaearctic distribution.

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 Jan 2020

History of Introduction and Spread

Top of page The species is native to the Palaearctic region but is frequently introduced into new areas, probably through commerce e.g. New Zealand (Bain, 1974); North America (Wood and Bright, 1992; Humble et al., 1994); Africa (Gaubil, 1849).

Risk of Introduction

Top of page The risk of introduction for P. chalcographus must be considered as high. Because of world wide commerce it can be introduced into new areas; once established its eradication is nearly impossible. The main pathways of introduction could be the trade of seedlings, wood with bark and dunnage with bark.

Habitat

Top of page P. chalcographus occurs in both natural and managed forests. Under optimal conditions it can become a serious pest with significant economic importance.

Habitat List

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Category	Sub-Category	Habitat	Presence	Status
Terrestrial
	Terrestrial – Managed	Cultivated / agricultural land	Present, no further details	Harmful (pest or invasive)
		Protected agriculture (e.g. glasshouse production)	Present, no further details	Harmful (pest or invasive)
		Managed forests, plantations and orchards	Present, no further details	Harmful (pest or invasive)
		Managed grasslands (grazing systems)	Present, no further details	Harmful (pest or invasive)
		Disturbed areas	Present, no further details	Harmful (pest or invasive)
		Rail / roadsides	Present, no further details	Harmful (pest or invasive)
		Urban / peri-urban areas	Present, no further details	Harmful (pest or invasive)
	Terrestrial ‑ Natural / Semi-natural	Natural forests	Present, no further details	Harmful (pest or invasive)
		Natural grasslands	Present, no further details	Harmful (pest or invasive)
		Riverbanks	Present, no further details	Harmful (pest or invasive)
		Wetlands	Present, no further details	Harmful (pest or invasive)
		Cold lands / tundra	Present, no further details	Harmful (pest or invasive)
		Deserts	Present, no further details	Harmful (pest or invasive)
Littoral
		Coastal areas	Present, no further details	Harmful (pest or invasive)
Freshwater			Present, no further details	Harmful (pest or invasive)
Marine			Present, no further details	Harmful (pest or invasive)

Hosts/Species Affected

Top of page P. chalcographus occurs generally on all coniferous tree species in its main area of distribution; Central and Eastern Europe, and from northern Asia to the Far East.

Host Plants and Other Plants Affected

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Growth Stages

Top of page Post-harvest, Vegetative growing stage

Symptoms

Top of page Trees infested by P. chalcographus can be easily identified by the change in colour of their needles, which start by yellowing and then become brown later on. In the initial stages of attack, the entrance holes can be seen together with frass that the beetles push out during the construction of the gallery system. The colour changes are easily visible by terrestrial investigation as well as by aerial survey.

List of Symptoms/Signs

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Biology and Ecology

Top of page A comprehensive review on the literature of P. chalcographus is given by Bright and Skidmore (1997; 2002) and Wood and Bright (1987; 1992).

The ecology and biology of P. chalcographus has been described extensively by Ratzeburg (1839), Schwerdtfeger (1929), Stark (1930), Karpinski (1933), Galoux (1948), Klausert (1954), Pfeffer (1955), Schwerdtfeger (1957), Postner (1974), Novak (1976) and Zumr and Soldán (1981).

P. chalcographus is a polygamous species. The males dig a nuptial chamber in the bark. Inside the nuptial chamber, every male copulates with three to six females. After copulation, females begin to bore the mother galleries in a star-like arrangement, depositing about 40 egg niches on both sides of the mother gallery. After hatching, larvae generate larval galleries horizontally to the mother galleries, ending in a pupal chamber where development is completed. Here, after metamorphosis to immature adults, maturation feeding takes place, enabling the imago to undergo sexual development. The nuptial chamber does not touch the wood if the bark is thick. However, in thin parts of the phloem of spruce and pine trees the nuptial chamber can carve into the wood making the frass similar to that of P. bidentatus.

P. chalcographus has one to two generations per year, while under especially favourable conditions a third generation may also be established. The swarming period of the first generation begins in the last ten days of April, dependant on the temperature. Vité (1965) states that the adults appear only as soon as the temperature reaches 16°C. The second generation swarms during July and August (Schwerdtfeger, 1929; Postner, 1974). P. chalcographus hibernates either in the larval, pupal or in the young adult stage inside the gallery. Adult beetles can also hibernate in the soil (Renner, 1974; Postner, 1974). Eggs can survive hibernation, however, under extreme temperatures they are more likely to die than the other ontogenetic stages. Embryonic development is heavily dependant on photoperiod and temperature.

If females are disturbed or if the tree is too heavily infested by other P. chalcographus, females can emerge and start another mother gallery without mating, on another tree.

P. chalcographus is attracted to the host tree by a mixture of monoterpenes such as (±)-alpha-pinene,(-)-beta-pinene and camphene (Chararas, 1962; Kangas, 1968; Vite and Pitmann, 1969; Byers et al., 1988). The detoxification of these substances in the gut, results in their transformation into the aggregation pheromones that compose of two chemical substances, dioxaspiro[4.4]nonane, 2-ethyl-1,6 or chalcogran (Francke, 1977) and methyl-E,Z-2,4-decadienoate, which are found to act synergistically (Byers et al., 1988; Byers et al., 1990). The aggregation pheromone attracts males as well as females, resulting often in mass infestation.

Intra-specific investigations showed that P. chalcographus did not colonize its current distribution area in a uniform way but instead had formed races within Europe. Crossing experiments among Northeast and Central European populations revealed incompatibilities (Führer, 1976) and differences in fertility and reproductive incompatibility were recognised (Führer, 1977). Analysis of morphological parameters - proportion of the antennae and spine formation on the elytra revealed significant differences between Northeast and Central European populations (Führer, 1978). Allopatric females were partly rejected by males in certain strain combinations, in which sympatric and allopatric females were simultaneously offered as pairing partners (Sturies and Führer, 1979). Crossings of Scandinavian, Polish and Alpine strains resulted in different degrees of inter-population heterosis (Führer and Klipstein, 1980). Hybrids tend to have a superior epidemiological potential when compared to the parental strains (Führer, 1984). Inter-popular hybrids showed polyploid sperm. Increased polyploid rates are indicative of inter-popular hybridisation (Führer, 1980; Führer and Krehan, 1985). Allozyme electrophoreses revealed two groups, however, races were not clustered according to previous findings (Ritzengruber, 1990). The bacterial endosymbiont Wolbachia, often responsible for race formation due to induction of cytoplasmic incompatibility, was not found in P. chalcographus (Riegler, 1999).

P. chalcographus was influenced by the postglacial history of its host, Picea abies. After the temperature amelioration about 10 000 years ago, vegetation and fauna began to re-invade the previously frozen and incompatible northern zones. According to pollen analysis, Picea abies had three refugial areas: Dinaric Alps, Carpatic Alps and area north of Moscow, Russia (Schmidt-Vogt, 1977; Lagercrantz and Ryman, 1990). Further, the Apennines were important for the re-colonization of the Southern Alps (Giannini et al., 1991). It is suggested that Picea abies re-migrated from the Dinaric and the Apennine back to the Central Alps; from the Carpathic back to the northern Alps and from the area north of Moscow (Kostroma), Russia, back to western Europe and over Finland to Scandinavia (Schmidt-Vogt, 1977; Lagercrantz and Ryman, 1990). It is likely that P. chalcographus had the same refugial areas and the same remigration routes. However, according to allozyme findings in P. chalcographus (Ritzengruber, 1990), this scolytid species might have had a geographic barrier preventing migration from Kostroma, Russia, to Scandinavia. Further there might be several other barrier zones (e.g., the Danube river) which might be responsible for the race formation within Central Europe. The genetic background of the formation of the races in P. chalcographus is not yet known although the understanding of their evolution would be interesting from the phylogeographic standpoint but also could be exploited in forest protection. It may open new ways for integrated pest management such as the use of new semiochemicals.

Virkki (1960) studied the cytology of male meiosis in P. chalcographus. The main features appeared to be the same as in Pityogenes quadridens (Hartig). The chromosome number was 20 in spermatogonia, and 10 bivalents in the first meiotic metaphase (9+Xyp). Abnormal spermatogenesis in P. chalcographus results in oversized spermatozoa in all the populations investigated. They can be identified by light microscopy and classified as 2n up to 16n polyploid. The percentage of polyploid sperm increases when allopatric parents are crossed: Parental populations with less than 1% polyploid, result in male F-1 with more than 20% polyploid. Populations from allochthonous sites for the host tree are distinguished by higher rates of sperm polyploidy than those from autochthonous areas. Thus, it is assumed that polyploid sperm indicates populations originating from the mixing of partially incompatible beetles (Führer, 2004).

The preference of P. chalcographus in the upper, thin barked parts of the trunk enables its association with Ips typographus, whose presence is located in the lower, thick barked parts. (Ratzeburg 1839, Schwerdtfeger 1957, Postner 1974, Zumr and Soldán 1981, Benz & Zuber 1993). The proportion of windthrown trees attacked by I. typographus increases with stem diameter, whereas the opposite is true for P. chalcographus. There is a positive interspecific association between the species on the lower, middle and upper third parts of the trees (Gothlin et al, 2000). Byers (1993) concluded that the avoidance of interspecific competition between these two scolytid species is achieved by the pheromone components, drawing a barrier to their distribution on the trunk.

P. chalcographus, is associated with blue-stain fungi, in a rather intimate manner, since a large proportion of individuals carry spores of ophiostomatoid fungi (Kirisits 2004). The spectrum of the mycobiota associated with P. chalcographus comprises of Graphium fimbriisporum (Kirisits 1996; Kirisits et al., 2000), many Ophiostoma species, Ophiostoma ainoae (Kirisits 1996; Kirisits et al., 2000) Ophiostoma bicolour (Krokene and Sollheim 1996; Kirschner 1998, 2001) Ophiostoma piceae (Kirschner 1998; 2001) Ophiostoma piceaperdum (Kirisits 1996; Kirisits et al., 2000; Kirschner 1998, 2001), as well as some representatives of Ceratocystiopsis (Ceratocystiopsis minuta (Kirisits 1996; Kirisits et al., 2000; Kirschner 1998, 2001) and Pesotum species (Mathiesen 1950; Kirisits 1996; Kirisits et al., 2000).

Natural enemies

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Notes on Natural Enemies

Top of page All parasitoids and predators mentioned in this datasheet are from Kenis et al. (2004). Parasitoid records considered by these authors as dubious are not included. For the predators, only those species clearly associated with P. chalcographus or its galleries are mentioned. The parasitoids and predators attacking the larvae usually also attack the pupae.

Pathway Vectors

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Plant Trade

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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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Impact Summary

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Impact

Top of page The EU funded COST action BAWBILT started to list the economic impact in European countries.

Economic Impact

Top of page The EU funded COST action BAWBILT started to list the economic impact in European countries.

Detection and Inspection

Top of page P. chalcographus can be easily detected by using pheromone traps bated with commercially available pheromone dispensers. Infested logs are easily detected by the presence of entrance holes, which are easily identifiable in the initial stage of infestation by the frass pushed out during the construction of the gallery system. Galleries of P. chalcographus are typical in shape and are easily visible on the lower surface of bark after debarking.

Similarities to Other Species/Conditions

Top of page This species can be easily distinguished from other Pityogenes species due to the characteristic brown-reddish tinge of the elytra, the shape of spines on the elytral declivity as well as the morphology of the female frons. Morphologically, the closest related species are P. trepanatus, P. fossifrons and other "three simple-spined" Pityogenes spp. with impressions on female the frons. If this species is found on pine species, the frass can be easily confused with P. bidentatus.

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.

The most effective measure is to remove infested trees from the forest before the new generation of adult beetles emerge. After logging, residue and other infested material should be burnt or chipped (Knízek and Zahradník, 2004). Thicker parts of trunks can be treated chemically by many commercially available pesticides (Zahradník, 2004). Forest management is recommended in order to increase the stability and vitality of forest stands (Thalenhorst, 1958; Christiansen and Bakke, 1988; Eidmann, 1992). Mass trapping with pheromone-baited traps or trap trees has also been successfully used to suppress beetle populations and prevent outbreak conditions (Bakke et al., 1977; Zumr, 1983; Bakke, 1985; Furuta et al., 1985; Weslien et al., 1989; Raty et al., 1995).

The pheromone traps with commercially available pheromone dispensers are used in central Europe mainly for monitoring but sometimes for mass trapping as well (Knízek and Zahradník, 2004).

Debarking of logs before export is the best and may be the only efficient way to prevent P. chalcographus from being introduced into isolated new areas. The EPPO Specific Quarantine Requirements (OEPP/EPPO, 1990) offer countries the choice of prohibiting import of bark of conifers from countries where the species occurs or of demanding an appropriate treatment. Wood of conifers should be debarked, kiln-dried or subjected to another appropriate treatment.

No biological control method of P. chalcographus is applied at the present. The percentage of mortality caused by natural enemies is not effective enough to stop the beetles in the outbreak stage.

References

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÷ymen T, 1992. The forest Scolytidae of Turkey. I^dot over~stanbul U^umlaut~niversitesi Orman Faku^umlaut~ltesi Dergisi. Seri A, 42(1):77-91; 76 ref.

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Bakke A, 1968. Ecological studies on bark beetles (Coleoptera: Scolytidae) associated with Scots Pine (Pinus sylvestris L.) in Norway with particular reference to the influence of temperature. Medd. Norske Skogforsoksv. 21 (6), (No. 83), (441-602). [9 pp. of refs.].

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Byers JA; Birgesson G; Löfqvist J; Appelgren M; Bergström G, 1990. Isolation of pheromone synergists of a bark beetle, Pityogenes chalcographus, from complex insect-plant odours by fractionation and subtractive-combination bioassay J. Chemical Ecology 16: 861-876.

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Fergusson A, 1920. The Clyde record of Pityogenes chalcographus L. Scottish Naturalist 1920:199-200.

Francke W, 1977. 2-Ethyl-1,6-dioxaspirol[4,4]nonane, Principal aggregation pheromone of Pityogenes chalcographus L. Naturwissenschafte 64: 590-591.

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Fuhrer E, 1976. Reproductive incompatibility in Pityogenes chalcographus - a new means of bark beetle control? Forstarchiv, 47(6):114-117; 10 ref.

Fuhrer E, 1977. Studies on intraspecific incompatibility in Pityogenes chalcographus L. (Col., Scolytidae). Zeitschrift fur Angewandte Entomologie, 83(3):286-297

Fuhrer E, 1980. Sperm polyploidy by inter-population hybridisation in Pityogenes chalcographus. Naturwissenschaften, 67(8):410-411

Fuhrer E, 1984. Race differentiation in Pityogenes chalcographus. Part 4: viability of interpopulation hybrids. [Rassendifferenzierung bei Pityogenes chalcographus L. (Col., Scolytidae). IV. Lebenserwartung interpopularer Bastarde.] Centralblatt fur das Gesamte Forstwesen, 101(1):24-33; 12 ref.

Fuhrer E; Klipstein EL, 1980. Race differentiation in Pityogenes chalcographus. Fertility of intraspecific F1 hybrids. [Rassendifferenzierung bei Pityogenes chalcographus L. (Col., Scolytidae). Fertilitat intraspezifischer F1-Bastarde.] Forstwissenschaftliches Centralblatt, 99(2):85-90; 7 ref.

Führer E, 2004. Polyploid spermatozoa in Pityogenes chalcographus and Ips typographus (Coleoptera: Scolytidae). European Journal of Entomology 101: 21-27.

Führer E; Krehan H, 1985. Zum Auftreten der Spermapolyploidie in Populationen des Fichtenborkenkäfers Pityogenes chalcographus. Zentralblatt für das gesamte Forstwesen 102, 134-149.

G÷thlin E; Schroeder LM; Lindel÷w +, 2000. Attacks by Ips typographus and Pityogenes chalcographus on windthrown spruces (Picea abies) during the two years following a storm felling. Scandinavian Journal of Forest Research, 15(5):542-549; 39 ref.

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Distribution References

Androic M, 1951. (Los mas importantes problemas de entomologia forestal en Yugoslavia)., 9 43-53.

CABI, Undated. Compendium record. Wallingford, UK: CABI

CABI, Undated a. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

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