Pityogenes chalcographus (sixtoothed spruce bark beetls)

Pictures

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Picture

Title

Caption

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Male

P. chalcographus; museum set specimen of adult male.

Milos Knizek

Female

P. chalcographus; museum set specimen of adult female.

Milos Knizek

Adult male

P. chalcographus; elytral declivity of adult male (museum set specimen).

Milos Knizek

Adult female

P. chalcographus; elytral declivity of adult female (museum set specimen).

Milos Knizek

Entrance hole

P. chalcographus attacking timber; entrance hole with frass.

Milos Knizek

Gallery system

Initial gallery system of P. chalcographus; maternal galleries with females laying eggs. The mating chamber is hidden within the phloem.

Milos Knizek

Gallery system

Gallery system of P. chalcographus; maternal and larval galleries, and hatching eggs.

Milos Knizek

Mature gallery system

Fully developed gallery system of P. chalcographus.

Milos Knizek

Entrance and exit holes

Entrance and exit holes of P. chalcographus on spruce (Picea abies).

Milos Knizek

Damage symptoms

Picea abies attacked by P. chalcographus.

Milos Knizek

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

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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

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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

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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

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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: 21 Jul 2022

Continent/Country/Region	Distribution	Origin	Invasive	Reference	Notes
Africa
Algeria	Present	Introduced		CABI (Undated)	Original citation: Gaubil (1849)
Asia
China	Present	Native		Fang et al. (1988) Zhou (2012) CABI (Undated)
-Beijing	Present			Zhou (2012)
-Heilongjiang	Present	Native		CABI (Undated) Fang et al. (1988)	Original citation: Wood and Bright (1992)
-Sichuan	Present	Native		CABI (Undated)	Original citation: Wood and Bright (1992)
Israel	Present			CABI (Undated)	Original citation: Schendl, 1978
Japan	Present, Widespread	Native		CABI (Undated)	Original citation: JPPA, 1980
Mongolia	Present	Native		CABI (Undated)	Original citation: Yanovskii and Tegshzhargal (1984)
North Korea	Present, Widespread	Native		CABI (Undated)
South Korea	Present, Widespread	Native		CABI (Undated)
Turkey	Present, Widespread	Native		CABI (Undated) Ünal (2010)	Original citation: Íymen, 1992
Europe
Austria	Present, Widespread	Native		CABI (Undated) Rosner and Führer (2002) Avtzis et al. (2008)	Original citation: Amman and Knabl (1913)
Belgium	Present, Widespread	Native		CABI (Undated)
Bosnia and Herzegovina	Present, Widespread	Native		CABI (Undated) Avtzis et al. (2008) Bojić et al. (2020)	Original citation: Wood and Bright (1992)
Bulgaria	Present, Widespread	Native		Takov et al. (2011) CABI (Undated)
Croatia	Present, Localized	Native		Avtzis et al. (2008) CABI (Undated)
Czechia	Present, Widespread	Native		CABI (Undated) Avtzis et al. (2008) Foit (2012) Čermák et al. (2013) Kula et al. (2013) Fiala et al. (2022)	Original citation: Jaminicky, 1960
Czechoslovakia	Present, Widespread	Native		CABI (Undated)	Original citation: Pfeffer (1931)
Federal Republic of Yugoslavia	Present, Widespread	Native		Androic (1951) CABI (Undated)
Denmark	Present, Widespread	Native		CABI (Undated) Harding et al. (1986)	Original citation: Byers (1993)
Estonia	Present, Widespread	Native		CABI (Undated) Voolma et al. (2004) Avtzis et al. (2008)	Original citation: Voolma and et al. (1996)
Finland	Present, Widespread	Native		Martikainen et al. (2001) Voolma et al. (2004) Avtzis et al. (2008) CABI (Undated)
France	Present, Widespread	Native		Avtzis et al. (2008) Bertheau et al. (2009) Bertheau et al. (2009a) CABI (Undated)
-Corsica	Present			CABI (Undated)	Original citation: Barthe (1896)
Germany	Present, Widespread	Native		CABI (Undated) Niemeyer (1992) Avtzis et al. (2008)	Original citation: Andersch (1851)
Greece	Present			Avtzis et al. (2008)
Hungary	Present, Widespread	Native		CABI (Undated)
Ireland	Present, Few occurrences	Introduced	Invasive	CABI (Undated)	Original citation: O’Conner and et al. (1991)
Italy	Present, Widespread	Native		CABI (Undated) Avtzis et al. (2008)	Original citation: Beffa (1949)
Latvia	Present, Widespread	Native		CABI (Undated)	Original citation: Telnov et al., 1997
Lithuania	Present, Widespread	Native		CABI (Undated) Gedminas et al. (2007) Avtzis et al. (2008)	Original citation: Silfverberg (1992)
Netherlands	Present, Widespread	Native		CABI (Undated)
North Macedonia	Present, Widespread	Native		CABI (Undated)	Original citation: Karaman (1971)
Norway	Present, Widespread	Native		Krokene and Solheim (1996) Avtzis et al. (2008) CABI (Undated)
Poland	Present, Widespread	Native		Grodzki (1997) Avtzis et al. (2008) CABI (Undated)
Romania	Present, Widespread	Native		Avtzis et al. (2008) Fora et al. (2012) CABI (Undated)
Russia	Present, Widespread	Native		Voolma et al. (2004) CABI (Undated)
-Central Russia	Present, Widespread	Native		CABI (Undated)	Original citation: Stark (1952)
-Eastern Siberia	Present, Widespread	Native		CABI (Undated)	Original citation: Stark (1952)
-Northern Russia	Present, Widespread	Native		CABI (Undated)	Original citation: Belousov (1916)
-Russian Far East	Present, Widespread	Native		CABI (Undated)	Original citation: Krivolutskaya (1996)
-Southern Russia	Present, Widespread	Native		CABI (Undated)	Original citation: Yanovskii (1999)
-Western Siberia	Present, Widespread	Native		CABI (Undated)	Original citation: Stark (1952)
Serbia	Present, Widespread	Native		Androic (1951) Markovic and Stojanovic (2010)
Slovakia	Present, Widespread	Native		Turčáni and Hlásny (2007) Avtzis et al. (2008) CABI (Undated)
Slovenia	Present, Widespread	Native		CABI (Undated) Jurc et al. (2006) Repe et al. (2013)	Original citation: Jurc (2003)
Spain	Present, Few occurrences	Introduced	Invasive	CABI (Undated)	Original citation: Riba (1996)
Sweden	Present, Widespread	Native		CABI (Undated) Byers (1993) Göthlin et al. (2000) Avtzis et al. (2008)	Original citation: Klefbeck and Sjoberg (1960)
Switzerland	Present, Widespread	Native		Avtzis et al. (2008) CABI (Undated)
Ukraine	Present, Widespread	Native		CABI (Undated)	Original citation: Rudnev (1965)
United Kingdom	Absent, Intercepted only			CABI (Undated)
North America
Canada
-British Columbia	Absent, Intercepted only			CABI (Undated)	Original citation: Humble et al., 1994
Jamaica	Absent, Intercepted only			CABI (Undated)	Original citation: Wood and Bright (1992)
Puerto Rico	Absent, Intercepted only			CABI (Undated)	Original citation: USDA, 1976
United States
-Georgia	Absent, Intercepted only			CABI (Undated)	Original citation: Haack (2001)
-Kentucky	Absent, Intercepted only			CABI (Undated)	Original citation: Haack (2001)
-Louisiana	Absent, Intercepted only			CABI (Undated)	Original citation: Haack (2001)
-Michigan	Absent, Intercepted only			CABI (Undated)	Original citation: Haack (2001)
-Minnesota	Absent, Intercepted only			CABI (Undated)	Original citation: MDA, 2003
-New York	Absent, Intercepted only			CABI (Undated)	Original citation: Haack (2001)
-Texas	Absent, Intercepted only			CABI (Undated)	Original citation: Haack (2001)
Oceania
New Zealand	Absent, Intercepted only			CABI (Undated)	Original citation: Bain (1974)

History of Introduction and Spread

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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

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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

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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	Managed	Cultivated / agricultural land	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Protected agriculture (e.g. glasshouse production)	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Managed forests, plantations and orchards	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Managed grasslands (grazing systems)	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Disturbed areas	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	Rail / roadsides	Present, no further details	Harmful (pest or invasive)
Terrestrial	Managed	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)
Terrestrial	Natural / Semi-natural	Natural grasslands	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	Riverbanks	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	Wetlands	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	Cold lands / tundra	Present, no further details	Harmful (pest or invasive)
Terrestrial	Natural / Semi-natural	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

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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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Plant name	Family	Context	References
Abies alba (silver fir)	Pinaceae	Other	Bertheau et al. (2009)
Abies grandis (grand fir)	Pinaceae	Unknown	Bertheau et al. (2009)
Juniperus communis (common juniper)	Cupressaceae	Other
Larix decidua (common larch)	Pinaceae	Other
Larix gmelinii (Dahurian larch)	Pinaceae	Other
Larix sibirica (Siberian larch)	Pinaceae	Other
Picea (spruces)	Pinaceae	Unknown	Turčáni and Hlásny (2007)
Picea abies (common spruce)	Pinaceae	Main	Bertheau et al. (2009); Bertheau et al. (2009); G÷thlin et al. (2000); Kula et al. (2013)
Picea glehnii (Sakhalin spruce)	Pinaceae	Main
Picea jezoensis (Yeddo spruce)	Pinaceae	Main
Picea obovata (Siberian spruce)	Pinaceae	Main
Picea omorika (Pancic spruce)	Pinaceae	Main
Picea orientalis (oriental spruce)	Pinaceae	Main	Ünal (2010)
Picea pungens (blue spruce)	Pinaceae	Unknown	Kula et al. (2013)
Picea sitchensis (Sitka spruce)	Pinaceae	Main	Bertheau et al. (2009)
Pinus banksiana (jack pine)	Pinaceae	Other
Pinus cembra (arolla pine)	Pinaceae	Other
Pinus densiflora (Japanese umbrella pine)	Pinaceae	Other
Pinus koraiensis (fruit pine)	Pinaceae	Other	Fang et al. (1988)
Pinus mugo (mountain pine)	Pinaceae	Other
Pinus parviflora var. pentaphylla	Pinaceae	Other
Pinus peuce (macedonian pine)	Pinaceae	Other
Pinus pumila (Dwarf Siberian pine)	Pinaceae	Other
Pinus sibirica (Siberian stone pine)	Pinaceae	Other
Pinus strobus (eastern white pine)	Pinaceae	Other	Bertheau et al. (2009)
Pinus sylvestris (Scots pine)	Pinaceae	Other	Bertheau et al. (2009); Foit (2012)
Pinus uncinata (mountain pine)	Pinaceae	Other
Pseudotsuga menziesii (Douglas-fir)	Pinaceae	Other	Bertheau et al. (2009)
Thuja plicata (western redcedar)	Cupressaceae	Unknown	Bertheau et al. (2009)

Growth Stages

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Post-harvest, Vegetative growing stage

Symptoms

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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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Sign	Life Stages	Type
Leaves / yellowed or dead
Leaves / yellowed or dead
Stems / gummosis or resinosis
Stems / gummosis or resinosis
Stems / internal feeding
Stems / internal feeding
Stems / visible frass
Stems / visible frass
Whole plant / discoloration
Whole plant / discoloration
Whole plant / frass visible
Whole plant / frass visible
Whole plant / internal feeding
Whole plant / internal feeding
Whole plant / plant dead; dieback
Whole plant / plant dead; dieback

Biology and Ecology

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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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Natural enemy	Type	Life stages
Beauveria bassiana	Pathogen	Adults; Arthropods\|Larvae
Beauveria caledonica	Pathogen	Adults; Arthropods\|Larvae
Caenopachys hartigii	Parasite	Arthropods\|Larvae
Chytridiopsis typographi	Pathogen	Adults; Arthropods\|Larvae
Coeloides bostrichorum	Parasite	Arthropods\|Larvae
Corticeus linearis	Predator	Eggs; Arthropods\|Larvae
Cosmophorus cembrae	Parasite	Adults
Dendrosoter middendorffii	Parasite	Arthropods\|Larvae
Dinotiscus eupterus	Parasite	Arthropods\|Larvae
Ecphylus hylesini	Parasite	Arthropods\|Larvae
Ecphylus silesiacus	Parasite	Arthropods\|Larvae
Epuraea marseuli	Predator	Eggs
Eurytoma arctica	Parasite	Arthropods\|Larvae
Eurytoma morio	Parasite	Arthropods\|Larvae
Gregarina typographi	Pathogen	Adults; Arthropods\|Larvae
Macromesus amphiretus	Parasite	Arthropods\|Larvae
Malamoeba scolyti	Pathogen	Adults; Arthropods\|Larvae
Mattesia	Pathogen	Adults; Arthropods\|Larvae
Medetera adjaniae	Predator	Adults; Arthropods\|Larvae
Medetera dendrobaena	Predator	Adults; Arthropods\|Larvae
Medetera dichrocera	Predator	Adults; Arthropods\|Larvae
Medetera setiventris	Predator	Adults; Arthropods\|Larvae
Menzbieria chalcographi	Pathogen	Adults; Arthropods\|Larvae
Metacolus azureus	Parasite	Arthropods\|Larvae
Metarhizium anisopliae	Pathogen	Adults; Arthropods\|Larvae
Nemosoma elongatum	Predator	Adults; Arthropods\|Larvae
Nemozoma elongatum	Predator
Nudobius lentus	Predator	Adults; Arthropods\|Larvae
Paromalus parallelepipedus	Predator
Phaonia	Predator	Arthropods\|Larvae
Placusa depressa	Predator	Adults; Eggs; Arthropods\|Larvae
Platysoma angustatum	Predator	Arthropods\|Larvae
Platysoma lineare	Predator	Arthropods\|Larvae
Pteromalus abieticola	Parasite	Arthropods\|Larvae
Pyemotes dryas	Predator	Arthropods\|Larvae
Rhizophagus depressus	Predator	Eggs; Arthropods\|Larvae
Rhopalicus quadratus	Parasite	Arthropods\|Larvae
Rhopalicus tutela	Parasite	Arthropods\|Larvae
Roptrocerus brevicornis	Parasite	Arthropods\|Larvae
Roptrocerus mirus	Parasite	Arthropods\|Larvae
Roptrocerus xylophagorum	Parasite	Arthropods\|Larvae
Scoloposcelis pulchella	Predator	Arthropods\|Larvae
Thanasimus femoralis	Predator	Adults; Arthropods\|Larvae
Thanasimus formicarius	Predator	Adults; Arthropods\|Larvae
Tolypocladium cylindrosporum	Pathogen	Adults; Arthropods\|Larvae
Tomicobia pityophthori	Parasite	Adults
Troxochrus nasutus	Predator	Adults
Unikaryon	Pathogen	Adults; Arthropods\|Larvae

Notes on Natural Enemies

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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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Vector	Notes	Long Distance	Local	References
Soil, sand and gravel	Soil	Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transport	Pest stages	Borne internally	Borne externally	Visibility of pest or symptoms
Bark	arthropods/adults; arthropods/eggs; arthropods/larvae; arthropods/nymphs; arthropods/pupae	Yes		Pest or symptoms usually visible to the naked eye
Stems (above ground)/Shoots/Trunks/Branches	arthropods/adults; arthropods/eggs; arthropods/larvae; arthropods/nymphs; arthropods/pupae	Yes		Pest or symptoms not visible to the naked eye but usually visible under light microscope

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/transport	Timber type	Used as packing
Solid wood packing material with bark	Conifer	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 Summary

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Category	Impact
Animal/plant collections	Negative
Animal/plant collections	Negative
Animal/plant products	None
Animal/plant products	None
Biodiversity (generally)	Positive
Biodiversity (generally)	Positive
Crop production	None
Crop production	None
Environment (generally)	Positive
Environment (generally)	Positive
Fisheries / aquaculture	None
Fisheries / aquaculture	None
Forestry production	Negative
Forestry production	Negative
Human health	None
Human health	None
Livestock production	None
Livestock production	None
Native fauna	None
Native fauna	None
Native flora	None
Native flora	None
Rare/protected species	None
Rare/protected species	None
Tourism	None
Tourism	None
Trade/international relations	Negative
Trade/international relations	Negative
Transport/travel	None
Transport/travel	None

Impact

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

Economic Impact

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

Detection and Inspection

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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

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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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GISD/IASPMR: Invasive Alien Species Pathway Management Resource and DAISIE European Invasive Alien Species Gateway	https://doi.org/10.5061/dryad.m93f6	Data source for updated system data added to species habitat list.

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Pityogenes chalcographus (sixtoothed spruce bark beetls)

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