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Datasheet

Pityogenes chalcographus
(sixtoothed spruce bark beetls)

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Datasheet

Pityogenes chalcographus (sixtoothed spruce bark beetls)

Summary

  • Last modified
  • 11 December 2020
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Preferred Scientific Name
  • Pityogenes chalcographus
  • Preferred Common Name
  • sixtoothed spruce bark beetls
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta
  • Summary of Invasiveness
  • 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 sui...

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Pictures

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PictureTitleCaptionCopyright
P. chalcographus; museum set specimen of adult male.
TitleMale
CaptionP. chalcographus; museum set specimen of adult male.
CopyrightMilos Knizek
P. chalcographus; museum set specimen of adult male.
Male P. chalcographus; museum set specimen of adult male.Milos Knizek
P. chalcographus; museum set specimen of adult female.
TitleFemale
CaptionP. chalcographus; museum set specimen of adult female.
CopyrightMilos Knizek
P. chalcographus; museum set specimen of adult female.
FemaleP. chalcographus; museum set specimen of adult female. Milos Knizek
P. chalcographus; elytral declivity of adult male (museum set specimen).
TitleAdult male
CaptionP. chalcographus; elytral declivity of adult male (museum set specimen).
CopyrightMilos Knizek
P. chalcographus; elytral declivity of adult male (museum set specimen).
Adult maleP. chalcographus; elytral declivity of adult male (museum set specimen).Milos Knizek
P. chalcographus; elytral declivity of adult female (museum set specimen).
TitleAdult female
CaptionP. chalcographus; elytral declivity of adult female (museum set specimen).
CopyrightMilos Knizek
P. chalcographus; elytral declivity of adult female (museum set specimen).
Adult femaleP. chalcographus; elytral declivity of adult female (museum set specimen).Milos Knizek
P. chalcographus attacking timber; entrance hole with frass.
TitleEntrance hole
CaptionP. chalcographus attacking timber; entrance hole with frass.
CopyrightMilos Knizek
P. chalcographus attacking timber; entrance hole with frass.
Entrance holeP. chalcographus attacking timber; entrance hole with frass.Milos Knizek
Initial gallery system of P. chalcographus; maternal galleries with females laying eggs. The mating chamber is hidden within the phloem.
TitleGallery system
CaptionInitial gallery system of P. chalcographus; maternal galleries with females laying eggs. The mating chamber is hidden within the phloem.
CopyrightMilos Knizek
Initial gallery system of P. chalcographus; maternal galleries with females laying eggs. The mating chamber is hidden within the phloem.
Gallery systemInitial gallery system of P. chalcographus; maternal galleries with females laying eggs. The mating chamber is hidden within the phloem.Milos Knizek
Gallery system of P. chalcographus; maternal and larval galleries, and hatching eggs.
TitleGallery system
CaptionGallery system of P. chalcographus; maternal and larval galleries, and hatching eggs.
CopyrightMilos Knizek
Gallery system of P. chalcographus; maternal and larval galleries, and hatching eggs.
Gallery systemGallery system of P. chalcographus; maternal and larval galleries, and hatching eggs. Milos Knizek
Fully developed gallery system of P. chalcographus.
TitleMature gallery system
CaptionFully developed gallery system of P. chalcographus.
CopyrightMilos Knizek
Fully developed gallery system of P. chalcographus.
Mature gallery systemFully developed gallery system of P. chalcographus.Milos Knizek
Entrance and exit holes of P. chalcographus on spruce (Picea abies).
TitleEntrance and exit holes
CaptionEntrance and exit holes of P. chalcographus on spruce (Picea abies).
CopyrightMilos Knizek
Entrance and exit holes of P. chalcographus on spruce (Picea abies).
Entrance and exit holesEntrance and exit holes of P. chalcographus on spruce (Picea abies).Milos Knizek
Picea abies attacked by P. chalcographus.
TitleDamage symptoms
CaptionPicea abies attacked by P. chalcographus.
CopyrightMilos Knizek
Picea abies attacked by P. chalcographus.
Damage symptomsPicea 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: 17 Feb 2021
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

AlgeriaPresentIntroducedOriginal citation: Gaubil (1849)

Asia

ChinaPresentNative
-BeijingPresent
-HeilongjiangPresentNativeOriginal citation: Wood and Bright (1992)
-SichuanPresentNativeOriginal citation: Wood and Bright (1992)
IsraelPresentOriginal citation: Schendl, 1978
JapanPresent, WidespreadNativeOriginal citation: JPPA, 1980
MongoliaPresentNativeOriginal citation: Yanovskii and Tegshzhargal (1984)
North KoreaPresent, WidespreadNative
South KoreaPresent, WidespreadNative
TurkeyPresent, WidespreadNativeOriginal citation: Íymen, 1992

Europe

AustriaPresent, WidespreadNativeOriginal citation: Amman and Knabl (1913)
BelgiumPresent, WidespreadNative
Bosnia and HerzegovinaPresent, WidespreadNativeOriginal citation: Wood and Bright (1992)
BulgariaPresent, WidespreadNative
CroatiaPresent, LocalizedNative
CzechiaPresent, WidespreadNativeOriginal citation: Jaminicky, 1960
CzechoslovakiaPresent, WidespreadNativeOriginal citation: Pfeffer (1931)
Federal Republic of YugoslaviaPresent, WidespreadNative
DenmarkPresent, WidespreadNativeOriginal citation: Byers (1993)
EstoniaPresent, WidespreadNativeOriginal citation: Voolma and et al. (1996)
FinlandPresent, WidespreadNative
FrancePresent, WidespreadNative
-CorsicaPresentOriginal citation: Barthe (1896)
GermanyPresent, WidespreadNativeOriginal citation: Andersch (1851)
GreecePresent
HungaryPresent, WidespreadNative
IrelandPresent, Few occurrencesIntroducedInvasiveOriginal citation: O’Conner and et al. (1991)
ItalyPresent, WidespreadNativeOriginal citation: Beffa (1949)
LatviaPresent, WidespreadNativeOriginal citation: Telnov et al., 1997
LithuaniaPresent, WidespreadNativeOriginal citation: Silfverberg (1992)
NetherlandsPresent, WidespreadNative
North MacedoniaPresent, WidespreadNativeOriginal citation: Karaman (1971)
NorwayPresent, WidespreadNative
PolandPresent, WidespreadNative
RomaniaPresent, WidespreadNative
RussiaPresent, WidespreadNative
-Central RussiaPresent, WidespreadNativeOriginal citation: Stark (1952)
-Eastern SiberiaPresent, WidespreadNativeOriginal citation: Stark (1952)
-Northern RussiaPresent, WidespreadNativeOriginal citation: Belousov (1916)
-Russian Far EastPresent, WidespreadNativeOriginal citation: Krivolutskaya (1996)
-Southern RussiaPresent, WidespreadNativeOriginal citation: Yanovskii (1999)
-Western SiberiaPresent, WidespreadNativeOriginal citation: Stark (1952)
SerbiaPresent, WidespreadNative
SlovakiaPresent, WidespreadNative
SloveniaPresent, WidespreadNativeOriginal citation: Jurc (2003)
SpainPresent, Few occurrencesIntroducedInvasiveOriginal citation: Riba (1996)
SwedenPresent, WidespreadNativeOriginal citation: Klefbeck and Sjoberg (1960)
SwitzerlandPresent, WidespreadNative
UkrainePresent, WidespreadNativeOriginal citation: Rudnev (1965)
United KingdomAbsent, Intercepted only

North America

Canada
-British ColumbiaAbsent, Intercepted onlyOriginal citation: Humble et al., 1994
JamaicaAbsent, Intercepted onlyOriginal citation: Wood and Bright (1992)
Puerto RicoAbsent, Intercepted onlyOriginal citation: USDA, 1976
United States
-GeorgiaAbsent, Intercepted onlyOriginal citation: Haack (2001)
-KentuckyAbsent, Intercepted onlyOriginal citation: Haack (2001)
-LouisianaAbsent, Intercepted onlyOriginal citation: Haack (2001)
-MichiganAbsent, Intercepted onlyOriginal citation: Haack (2001)
-MinnesotaAbsent, Intercepted onlyOriginal citation: MDA, 2003
-New YorkAbsent, Intercepted onlyOriginal citation: Haack (2001)
-TexasAbsent, Intercepted onlyOriginal citation: Haack (2001)

Oceania

New ZealandAbsent, Intercepted onlyOriginal 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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CategorySub-CategoryHabitatPresenceStatus
Terrestrial ManagedCultivated / agricultural land Present, no further details Harmful (pest or invasive)
Terrestrial ManagedProtected agriculture (e.g. glasshouse production) Present, no further details Harmful (pest or invasive)
Terrestrial ManagedManaged forests, plantations and orchards Present, no further details Harmful (pest or invasive)
Terrestrial ManagedManaged grasslands (grazing systems) Present, no further details Harmful (pest or invasive)
Terrestrial ManagedDisturbed areas Present, no further details Harmful (pest or invasive)
Terrestrial ManagedRail / roadsides Present, no further details Harmful (pest or invasive)
Terrestrial ManagedUrban / peri-urban areas Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalNatural forests Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalNatural grasslands Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalRiverbanks Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalWetlands Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalCold lands / tundra Present, no further details Harmful (pest or invasive)
Terrestrial Natural / Semi-naturalDeserts Present, no further details Harmful (pest or invasive)
LittoralCoastal 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 nameFamilyContextReferences
Abies alba (silver fir)PinaceaeOther
  • Bertheau et al. (2009)
Abies grandis (grand fir)PinaceaeUnknown
  • Bertheau et al. (2009)
Juniperus communis (common juniper)CupressaceaeOther
    Larix decidua (common larch)PinaceaeOther
      Larix gmelinii (Dahurian larch)PinaceaeOther
        Larix sibirica (Siberian larch)PinaceaeOther
          Picea abies (common spruce)PinaceaeMain
          • Bertheau et al. (2009);
          • Bertheau et al. (2009);
          • G÷thlin et al. (2000);
          • Kula et al. (2013)
          Picea glehnii (Sakhalin spruce)PinaceaeMain
            Picea jezoensis (Yeddo spruce)PinaceaeMain
              Picea obovata (Siberian spruce)PinaceaeMain
                Picea omorika (Pancic spruce)PinaceaeMain
                  Picea orientalis (oriental spruce)PinaceaeMain
                  • Ünal (2010)
                  Picea pungens (blue spruce)PinaceaeUnknown
                  Picea sitchensis (Sitka spruce)PinaceaeMain
                  • Bertheau et al. (2009)
                  Pinus banksiana (jack pine)PinaceaeOther
                    Pinus cembra (arolla pine)PinaceaeOther
                      Pinus densiflora (Japanese umbrella pine)PinaceaeOther
                        Pinus koraiensis (fruit pine)PinaceaeOther
                        • Fang et al. (1988)
                        Pinus mugo (mountain pine)PinaceaeOther
                          Pinus parviflora var. pentaphyllaPinaceaeOther
                            Pinus peuce (macedonian pine)PinaceaeOther
                              Pinus pumila (Dwarf Siberian pine)PinaceaeOther
                                Pinus sibirica (Siberian stone pine)PinaceaeOther
                                  Pinus strobus (eastern white pine)PinaceaeOther
                                  • Bertheau et al. (2009)
                                  Pinus sylvestris (Scots pine)PinaceaeOther
                                  Pinus uncinata (mountain pine)PinaceaeOther
                                    Pseudotsuga menziesii (Douglas-fir)PinaceaeOther
                                    • Bertheau et al. (2009)
                                    Thuja plicata (western redcedar)CupressaceaeUnknown
                                    • 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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                                    SignLife StagesType
                                    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 enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
                                    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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                                    VectorNotesLong DistanceLocalReferences
                                    Soil, sand and gravelSoil Yes

                                    Plant Trade

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                                    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 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/transportTimber typeUsed 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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                                    CategoryImpact
                                    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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                                    Virkki N, 1960. Cytology of male meiosis in certain European forest beetles of the families Scolytidae, Cleridae, and Anobiidae. Annales Academiae Scientiarum Fennicae, Helsinki (Ser. A. IV.) No. 49, 1960. pp. 16 + 13 photomicrographs. 7 refs.

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                                    Vité JP; Pitmann GP, 1969. Aggregation behaviour of Dendroctonus brevicomis in response to synthetic pheromones. J. Ins. Phys. 15: 1617-1622.

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                                    Weiser J, 1955. Beitrag zur Kenntnis der Parasiten des Borkenkäfers Ips typographus. II. Vestnik Ceskoslovenske Zoologicke Spolecnosti (Acta Societatis Zoologicae Bohemoslovenicae; Transactions of the Czechoslovak Zoological Society) 19, 374-380.

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                                    Wulf A, 1983. Studies on the entomopathogenic fungus Beauveria bassiana (Bals.) Vuill. as a parasite of the bark-beetle Pityogenes chalcographus L. (Col., Scolytidae). Zeitschrift fur Angewandte Entomologie, 95(1):34-46

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