3.1. Material sources

For the period between 1858 and 2023, we collected data from entomological collections and literature. Most of the material consists of the authors’ unpublished data from field sampling of beetles.

3.1.1. Entomological collections

We examined specimens of bark beetles and pinhole borers in 10 entomological collections which are currently held at the Natural History Museum of Slovenia – NHMS, https://www.pms-lj.si/; the Jovan Hadži Institute of Biology ZRC SAZU, the Barje Research Station; the Department of Forestry and Renewable Forest Resources at the Biotechnical Faculty, as well as three collections held privately by collectors.

The largest part of the material is deposited in five collections kept by the Natural History Museum of Slovenia. (1) The Josef Stussiner Collection (cJSs), also known as the Stussiner Entomological Collection, was reviewed. It was created at the end of the 19th and the beginning of the 20th century. The specimens are mainly from Slovenia and Greece. Its author is Josef Stussiner, 1850-1917. It consists of 167 entomological boxes, about 40,844 specimens, including 6,500 species of beetles (Coleoptera). The beetles in the collection are identified and systematically arranged. (2) The Historical collection of Josef Staudacher (1876-1945), the Staudacher Entomological Collection (cJSD), created between both world wars in Slovene territory (former Drava Banovina) and Dalmatia. The collection is systematically arranged, the beetles are identified. It comprises 40 entomological boxes (4,346 specimens) of butterflies (Lepidoptera) and 139 entomological boxes (about 100,000 specimens) of beetles (Coleoptera). (3) The largest collection is the Central Slovene Collection of Beetles (Coleoptera) (cCCS), authored by Savo Brelih (1927-2012) (Figure 3). The collection includes the collections of Anton Karl Vincenc Bianchi (1858-1933) (cABi), dr. Evgen Jaeger (1892-1959) (cEJe), Josef Peyer (??-1940/45?) (cJPe) and Savo Brelih (1927-2012) (cSBr). The collection comprises 72,313 specimens in 623 entomological boxes. (4). The Gspan Collection of Beetles (Coleoptera) (cAGs) was created in the first half of the 20th century in Slovenia, the Balkan Peninsula and the Palearctic. Its author is Alfonz Gspan, 1878-1963. The collection consists of 293 entomological boxes and about 64,537 specimens. Well-arranged and administered, it represents the largest collection of beetles from this period in Slovenia. (5). The aforementioned Furlan Entomological Collection (Vincenc Furlan, 1934-2012), comprises 151 entomological boxes containing true bugs (Hemiptera), beetles (Coleoptera) and butterflies (Lepidoptera).

We also analysed the collections of Dr. Božo Drovenik (cBDr). The Coleoptera Collection of the Jovan Hadži Biological Institute ZRC SAZU, has ben created between 1920 and 2020. In 2017 the museum took it over from the ZRC SAZU, now it is stored in the ZRC SAZU’s Jovan Hadži Institute of Biology, Barje Research Station). It contains 363 entomological boxes of beetles and 35 entomological boxes of butterflies.

The contents of the collection of bark beetles (one box) donated by Prof. J. Titovšek from the BFG to the Faculty of Forestry in Sarajevo (cŠFS) were also reviewed.

A number of beetle specimens are also held in private collections. Materials from the collections of Andrej Kapla (cAKa), Alja Pirnat (cAPi) and Vladimir Kodrič (cVKo) have been reviewed, confirmed or redefined for the contribution. These collections represent an important source of data for the subfamilies analysed here.

3.1.2. Literature data

The first works from the 19th and early 20th centuries on the bark beetles and pinhole borers in the territory of present-day Slovenia were published by three authors: Moritz Siegel (1866), Karl Brancsik (1871) and Gilbert Fuchs (1905).

Siegel (1866), in Versuch einer Käfer-Fauna Krains, lists  bark beetles, which are given in the list below. The names according to Siegel are written in the first place, and the now valid names are in the second place.

Brancsik (1871), in Die Käfer der Steiermark, gives data for both the Austrian and the Slovene part of Styria. For some, especially the more common species of bark beetles, he does not give the exact sites, nor the names of the finders. We have not included these in our list, as it is not certain that the sites were in the territory of present-day Slovenia. However, we summarise the data where the finder is listed as J. N. Spitzy, who collected material in the wider area of Lenart in the Slovenske Gorice, and of course those where the exact site is mentioned. Where it was not possible to determine with certainty from Brancsik’s text whether the site was in North Styria (Austria) or Lenart in Slovenske Gorice, we used a question mark (?). Brancsik obtained material from the Slovene part of Styria from the aforementioned Spitzy and from Prof. Dr. Leitgeb from Celje. In the list, the names according to Brancsik are given first, the currently valid scientific names, followed by the sites and notes as given by the author.

In Die Borkenkäfer Kärntens und der angrenzenden Gebirge, Fuchs (1905) lists the presence of 70 species of bark beetles and one species of pinhole borer for Carinthia and neighbouring mountain ranges. For these species, more detailed site information is often lacking, and the dates of collection are not given. For only three species of bark beetles the book lists sites which are undoubtedly in present-day Slovenia.

Only the above information is included in the main part of Scopolia. For all the other species mentioned in the author’s work, it cannot be stated with certainty that their sites are in the Slovene part of Carinthia. For example, for the species Ips amitinus Eich. = Ips amitinus (Eichhoff, 1872), he states that it occurs “on spruce, pine, and less frequently on larch, both south and north of the Drava River”. For 30 species of bark beetle and the species Platypus cylindrus Fabr. = Platypus cylindrus (Fabricius, 1792), the Karavanke Mountains are listed as the site of occurrence, and for a further 15 species Fuchs states that they are present “throughout the whole country”. Theodor Prossen, in his 1913 article I. Nachtrag zum Verzeichnisse der bisher in Kärnten beobachtete Käfer, lists 48 species of bark beetle for Carinthia in the Ipidae chapter, but mostly limits himself to quoting Gilbert Fuchs and does not mention any additional sites in present-day Slovenia.

Previous posts are followed by short technical papers that mainly show the economic importance of bark beetles. In 1876, the Imperial Royal Forest Supervisor Ivan Salzer published a paper entitled A short inventory of the spruce bark beetle, with an emphasis on its killing (Teacher’s Library of the Folk School in Stari Trg pri Ložu) (Salzer 1876). This was followed by contributions by professors of forestry Šlander (1948, 1951, 1953, 1958) and Titovšek (1969, 1973, 1974, 1977, 1983, 1988, 1994).

The results of research work on bark beetles in Slovenia have been published: Babuder & Pohleven 1993, 1995; Babuder et al. 1996; Benkovič 1951; De Groot 2018; Demšar & Jurc 2007; Geister 2004; Hauptman 2018, 2019a, 2019b, 2020, 2022a, 2022b, 2022c; Hauptman et al. 2018, 2019, 2019a, 2019b; Hočevar & Jurc 1982; Jug 1967; Jerina et al. 2008; Jurc 1993,1996, 2000, 2000a, 2001, 2001a, 2001b, 2002,  2002a, 2002b, 2004a, 2004b, 2006a, 2006b, 2006c, 2006d,  2006e, 2006f, 2007, 2008, 2008a, 2010, 2012, 2013, 2016, 2019, 2020, 2021a, 2021b, Jurc 2021c; Jurc & Bojović 2004c; Jurc & Brelih 2001; Jurc & Miljašević 2006g; Jurc & Nève Repe 2009; Jurc & Reščič, 2013; Jurc & Škulj 2018; Jurc et al. 2003, 2006, 2009a, 2010, 2010a, 2011, 2012, 2016, 2017, 2021, 2022; Jurc D. 1988; Jurc D. et al. 2003, 2004, 2005; Kavčič 2017; Kavčič et al. 2017, 2023; Kovačević 1953; Meterc et al. 2010; Nève Repe et al. 2013, 2015, 2018; Ogris & Jurc 2010; Ogris et al. 2019, 2020; Pavlin 1991a, 1991b, 1991c, 1992a, 1993, 1993a, 1993b, 1993c, 1993d, 1993e, 1993f, 1994, 1994a, 1994c, 1994d, 1995a, 1995b, 1996, 1997, 2001, 2003, 2004, 2005, 2008, 2010a, 2011a, 2012, 2013a, 2013b, 2014a, 2014b, 2014c, 2015a, 2015b, 2016a, 2016b, 2016c, 2016e, 2017a, 2017b, 2017c, 2017d, 2018, 2018a, 2018, 2018c, 2018d, 2018e, 2019a, 2019b, 2019c, 2019d, 2019e, 2020a, 2020b, 2020c, 2020d, 2020e, 2021a, 2021b, 2022a, 2022b, 2023a, 2023b; Pavlin et al. 1994b, 2010b, 2011b, 2012; Pavlin & Trdan 2007; Perko, 1969; Podlesnik & Jurc 2012, 2018; Podlesnik et al. 2014; Rener 2002; Rener & Jurc 2001; Ropret et al. 2007; Smolnikar et al. 2019; Stauffer et al. 2001; Urleb 1957; Vrezec et al. 2012.

Results of the work on bark beetles are also published in bachelor/master and PhD theses: Bjelić 2014; Brudar 2020; Demšar 2006; Groznik 2019; Habjan 2009; Jagodic 1997; Jene 2005; Kalič 2006; Komjanc 2005; Koželj 2010; Krašna 1998; Kresevič 2005; Križnar 2012; Lesar 2004; Lukman 2010; Pajek 2019; Perko 2001; Nève Repe 2014; Petelin 2017; Podlesnik 2016; Rener 2000; Ribič 2007; Ropret 2006; Rus 2007; Rutar 2014; Sever 2012; Škulj [Jurc], 1988;  Šimon 2011; Vrhovnik 2008; Vučko 2016; Župnek 2020.

The use of synthetic population (aggregation) pheromones in the management of bark beetles has been introduced in the world and in Slovenia. Foreign publications in this field include Bakke 1970, 1975; Bakke & Hugges 1974; Bakke et al. 1977; Bauer & Vité 1975; Blight et al. 1977; Francke  1973; Francke & Heemann 1974; Francke et al. 1974, 1977;  Gerken 1978; Gerken et al. 1978; Grüne 1979; Harring & Candace 1978; Harring et al. 1975; Harrtig et al. 1975; Lanier et al. 1977; Leuze 1977; Peacock et al. 1975; Pearce et al. 1975; Stoakley et al. 1978; Vité 1978. Slovenian authors who have published papers on bark beetle pheromones were Titovšek 1988; Pavlin 1991b, 1992a, 1997, 2001; and Jurc et al. 2012, 2017. In recent years, pest distribution surveys (MKGP, UVHVVR) have been carried out by trapping bark beetles in different types of traps (Econex black cross trap, Witasek black cross trap, ground trap, Theysohn® black flat slot trap) using different attractants (ethanol, conifer twigs, Galloprotect Pack®, Linoprax®, Pheroprax®, Chalcoprax®, α-pinene, Chalcowit®, PC-Ecolure®, Curviwit®) (Jurc et al. 2016, Pavlin et al. 2016c, Hauptman et al. 2018, Kavčič 2018).

We consulted older catalogues and taxonomic keys such as Siegel 1866, Barncsik 1871, Grüne 1979, Freude et al. 1981, Titovšek 1988, Pfeffer & Knížek 1993, Lucht 1994; Pfeffer 1995 and aligned the taxonomy of the covered bark beetles and pinhole borers with the current catalogue Löbl & Smetana 2011.

3.1.3. Baza podatkov FloVegSi

From the FloVegSi database (= Fauna, flora and vegetation of Slovenia, BIJH ZRC SAZU (https://bijh.zrc-sazu.si/sl/zbirka/podatkovna-zbirka-flovegsi-favna-flora-in-vegetacija-slovenije-bijh-zrc-sazu) we used data for 10 species of bark beetles which were the result of a faunal inventory.

3.1.4. Sampling beetles in the field

More than half of the data collected is the result of the authors’ fieldwork. Field collection of bark beetles is carried out in two basic ways:

1. by collecting specimens on host plants, other woody material or in the soil;
2. (and) with trapping.

3.1.4.1 Collecting beetles on host plants

The easiest way to collect bark beetles is on infested (host) plants. On trees showing signs of attack (change in canopy colour, entrance or exit holes on the trunk, presence of droppings and, in conifers, resin oozing), the bark is usually removed in phloemophagous species and the imagoes are collected directly from the tunnel systems. Even if the young beetles have already fledged, the remains of dead imago are often found in the tunnel systems. Often, even parts of damaged beetles are sufficient for reliable species identification. If the bark beetles are in the ‘white’ developmental stages (egg, larva, unpigmented pupa), it is advisable to take the pieces of infested wood material to the laboratory, place them in insectaries and wait for the end of the development. This way, we have adult specimens (imagoes) that are easier to identify, and species that we have previously overlooked are often present as well. Other ways of collecting bark beetles on woody plants include collecting beetles from hollow of young shoots, from overwintering sites under the bark or directly on infested trees before or during boring into trunks or branches. The latter method is also suitable for collecting xylomycetophagous species of bark beetles, where the only way to access the imagoes is usually by splitting the wood and collecting them from the tunnel systems. Both phloemophagous and xylomycetophagous species of bark beetles can also be collected from logs, stumps, logging residue and construction timber. For phloemophagous species, any reasonably fresh piece of wood that has not been (or has only been partially) debarked is suitable for colonisation. We can also obtain samples of bark beetles through traps used in the control of economically important bark beetles: control (or trapping) trees, trunks or piles of branches. Some species of bark beetles overwinter (also) in the ground, near infested trees. Digging bark beetles from the ground was first used in Slovenia by ŠLANDER (1958).

3.1.4.2. Catching bark beetles in traps

In the mid-1980s, foresters in Slovenia started using traps based on synthetic bark beetle pheromones. Initially, Pheroprax® (for Ips typographus), Chalcoprax® (for Pityogenes chalcographus) and Linoprax® (for Trypodendron lineatum) were available. The original tube traps were soon replaced by the more effective Theysohn® flat slot traps. The first peer-reviewed papers on traps focused on their effectiveness or on the capture of target species (CIMPERŠEK 1986, 1988). Later, several studies were carried out to analyse the additional capture of non-target species, in particular other species of bark beetle and their predators (PAVLIN 1991a, 1991b). A similar study was carried out in timber yards (BABUDER ET AL. 1996). The development of new pheromones also enabled the capture or inventorying of other species of bark beetle. A systematic zoogeographical inventory of Ips cembrae in the northern part of Slovenia was carried out by PAVLIN (1997, 2001c) using the Cemprax® trap and pheromone. An important source of data on the distribution of bark beetles in Slovenia was provided by a detailed analysis of additional catches from cross-traps designed to monitor sawyers (Cerambycidae: Monochamus), which are potential vectors of the pinewood nematode (Bursaphelenchus xylophilus) (JURC ET AL. 2016). The study was carried out from 2007 to 2015 in coniferous stands using different pheromones for Monochamus galloprovincialis, which also contained “kairomone components” in the form of a mixture of different bark beetle pheromones (PAVLIN 2010b, PAVLIN ET AL. 2016c). Traps equipped with attractants in the form of alpha-pinene and ethanol are also suitable for catching xylomycetophagous bark beetles (HAUPTMAN ET AL. 2019a ). The use of traps with different attractants has provided a completely new perspective on the zoogeography of individual bark beetle species. Among the species found for the first time in Slovenia after 2000, there are many (especially xylomycetophagous) species that were found only in traps but never on host plants.

3.2. Display of data

3.2.1. Collection and processing of data in the table

Data from various sources were entered into an Excel spreadsheet which made it possible to review and unify the data. The table facilitated transparency, in particular the standardization of data. In addition to the Latin name of the species and the author, the table contained the region, place name, longitude (X), latitude (Y), UTM quadrant, altitude, host plant / trap type, plant part attacked / type of pheromone, author name, collection / literature, and date. All available data were entered. Particularly for older records, some information may be missing or entered with a lower level of accuracy. The contents of the table are presented in the form of text for each species in Chapter 4. RESULTS.

Landscape. The territorial division of Slovenia into regions were taken from older volumes of Scopolia in which the division is based on the Map of the Slovene Land and its Regions (Kozler & Knorr 1853) and the map by Gabrovec, M. & Rajšp, V. (1998),  published in the book Slovenia – Regions and People (p. 18-19), which is based on the 1914 borders of Austrian provinces. Bela Krajina region is specifically discussed, as well as Ljubljana and its surroundings, the borders of which are not historical.

Place name: The first place name refers to the narrower name of the site, the second (if provided) to the broader name of the site. The first name is taken from the Atlas of Slovenia, Third Edition (Kos 1996). It may contain different geographical terms. In exceptional cases, microtoponyms or names of landowners are added. Individual sites which are located close to the same place and are at least 30 m apart are also indicated with additional numbers. Place names are spelled according to the 8th version of the Slovene Orthography. The Slovene preposition pri or a hyphen is used only if it is part of the official place name. Place names are sometimes accompanied by a footnote referring to a more specific location (e.g., a tree nursery).

Geographical coordinates: Geographical coordinates are based on the WGS84 reference ellipsoid projection. For most of the older sites, the coordinates are taken from the Atlas of Slovenia (Kos 1996), rounded to the nearest minute. More recent locations are based on the online Atlas of the Environment and are rounded to the nearest second. In the case of older locations where the geographical concept extends tens of kilometres (Pohorje, Ljubljana, etc.), the central coordinates are given in brackets.

UTM quadrant: UTM quadrants are defined based on longitude and latitude via the online Coordinate System Converter. For approximate coordinates, UTM quadrants are provided in brackets.

Altitude: For older data, altitude is determined with maps or barometric altimeters. Altitudes marked with the symbol ~ are only approximate, with a tolerance of ± 50 m. In cases where altitude may change rapidly (mountains, etc.) and could not be determined reliably, it is replaced by the * symbol. In more recent data, altitude is usually determined via the digital Environmental Atlas and has been read at the same time as the geographical coordinates.

Host plant / Trap type: The Latin name of the plant on which the specimen was collected is listed. With rare exceptions, this is the name of the host plant. The names have been taken from various editions of the botanical key to the Small Flora of Slovenia (Martinčič et al. 2007). In older data, the host plant may be entered only as a genus, and even more often this information is missing altogether. If the bark beetle was not collected on the host plant or a part of it, it was trapped. There are different types of traps, which are abbreviated in the text. The abbreviation is usually based on the abbreviated name of the trap manufacturer.

Plant part attacked / Pheromone type: For most of the more recent entries, we also recorded the part of the plant attacked and its dimensions. If the plant part was irregularly shaped (e.g., a stump), the number in cm represents the average of the measurements of the largest and smallest dimensions. The abbreviation rs means that the species was determined solely based on observations of the tunnel system. In the genus Crypturgus, the abbreviation nrs is used to indicate that the species was found in tunnel systems derived from the tunnel system of another species which is referred to below by its Latin name.

Authors’ information: Abbreviations are used for legators (l), observers (o), determiners (d), renamers (r) and validators (v). One or more persons may be named and their names are abbreviated to the first letter of the first name and the first two letters of the surname (e.g. MJu for Maja Jurc).

Collection / Literature: The data must be obtained from a collection (museum or private) or from a source / literature. Both are abbreviated. Collections begin with the letter c and are followed by the abbreviation of the founder of the collection or institution. Only the source from which the information was actually taken is provided. Sources begin with the abbreviation lit., followed by the abbreviated name of the author and the year of publication (e.g. lit.JTi1983).

Date: The date is indicated at the end of each entry. This is due to the periodic collection of material at specific locations (especially in traps), so the shortest and easiest way to read the entry is with the dates listed at the end. In older data, often only the month is entered instead of the day, e.g., 8.1966. In rarer cases, only the year is available. If the year is also missing, the data are classified into three periods: <1951 (up to and including 31.12.1950), 1951-2000 (1.1.1951 – 31.12.2000) or >2000 (from 1.1.2001). The periods have been defined on the basis of the period of activity of the legators and the method of preparation.

Shortening entries: In addition to the dates entered at the end, we have also shortened the text by omitting identical information in similar entries. When ibidem is used, it means that the entry is identical to the previous one, only the part of the entry following the word ibidem is different. After the first change, the entry is then continued to the end, even in cases where the subsequent data match the previous entry.

3.2.3. Distribution in the Palearctic region, maps, photographs, mentions in catalogues

Distribution in the Palearctic region: The Palearctic covers Europe, North Africa, Western Asia, Siberia, Northern China and Japan. The distribution of the considered species in the Palearctic is presented according to the LÖBL & SMETANA (2011) catalog, with country abbreviations (see chapter 3.4.1). In cases where the presence of the species in Slovenia was not indicated in the catalog, the labels for Slovenia (SL*) was added subsequently. Maps: The basis for creating maps of the distribution of bark beetles was an edited excel table with historical and recent data. Based on the geographic coordinates for each entry, a graphical representation was prepared using the ArcGIS program. Distribution maps are drawn for all represented species, on which the labels are divided into three periods: <1951 (up to and including 31.12.1950), 1951-2000 (1.1.1951 – 31.12.2000) or >2000 (from 1.1.2001). Older, incomplete data which do not provide a place name are not shown in the maps.

Photographs: Most species are presented with a dorsal (top) and lateral (side) photograph of the specimen, for some species photographs of females and males are available (OLYMPUS, EFI, PMS, Maja Jurc, 17.10.2004).