10urn:lsid:arphahub.com:pub:6BEA2F36-F41A-5C4C-8F71-33F322F204B5Nature ConservationNC1314-69471314-3301Pensoft Publishers10.3897/natureconservation.27.2497724977Review ArticleBostrichidaeBostrichoideaColeopteraEcology & Environmental sciencesHabitats Ecosystems & Natural SpacesNeogeneEastern EuropeEuropePolandStephanopachyslinearis (Kugelann, 1792) (Coleoptera, Bostrichidae) in PolandBorowskiJerzyjerzy_borowski@sggw.pl1GutowskiJerzy M.2SławskiMarek1SućkoKrzysztof2ZubKarol3Department of Forest Protection and Ecology, Warsaw University of Life Sciences, ul. Nowoursynowska 159/34, 02-776 Warsaw, PolandWarsaw University of Life SciencesWarsawPolandDepartment of Natural Forests, Forest Research Institute, 17-230 Białowieża, PolandForest Research InstituteBiałowieżaPolandMammal Research Institute, Polish Academy of Science, 17-230 Białowieża, PolandMammal Research Institute, Polish Academy of ScienceBiałowieżaPoland
Corresponding author: Jerzy Borowski (jerzy_borowski@sggw.pl)
Academic editor: K. Henle
201803072018277584FFA62905-FF80-C816-5C05-FFD9FFA00F11861DAABD-8AC6-4811-BC3B-065A7BBDDB6613090471203201819062018Jerzy Borowski, Jerzy M. Gutowski, Marek Sławski, Krzysztof Sućko, Karol ZubThis is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.http://zoobank.org/861DAABD-8AC6-4811-BC3B-065A7BBDDB66
Stephanopachyslinearis (Kugelann, 1792) belongs to the family of horned powderpost beetles (Bostrichidae), represented in the fauna of Europe by 29 native species. It is a characteristic element of the northern, boreal zone of the Palaearctic and alpine areas of central and southern Europe. This species as a rare beetle important for the European Union, has been placed in Annex II of the Habitats Directive, on the IUCN Red List of Threatened Species and European Red List of Saproxylic Beetles. S.linearis was described from Poland in 1792 and, after 220 years, again encountered in this country. The zoogeographical distribution is shown and elements of its biology and ecology are discussed.
Borowski J, Gutowski JM, Sławski M, Sućko K, Zub K (2018) Stephanopachys linearis (Kugelann, 1792) (Coleoptera, Bostrichidae) in Poland. Nature Conservation 27: 75–84. https://doi.org/10.3897/natureconservation.27.24977
Introduction
Stephanopachyslinearis (Kugelann, 1792) is a representative of the forest fauna characteristic of boreal and alpine regions of the Palaearctic. It belongs to the family of horned powderpost beetles (Bostrichidae), of which 29 native species occur in Europe (Borowski 2007), 9 of them – including S.linearis – having been reported from Poland (Dominik 1958, Burakowski et al. 1986).
S.linearis was discovered and described in 1792 by the German entomologist J.G. Kugelann, working as an apothecary in Ostróda in the (then) Olsztyn regency of East Prussia (now NE Poland). Besides short Latin and German descriptions of Apatelinearis, Kugelann gave a diagnosis allowing its distinction it from the now common bark beetle, Anisandrusdispar (Fabricius, 1792) (Curculionidae, Scolytinae), with information that the new species had been found only once, on an old hoarding. Kugelann’s publication has become the basis of the inclusion – after World War II – of S.linearis in the Polish fauna (Dominik 1958, Burakowski et al. 1986, Borowski 2007), but during the 220 years since its description, it has never been reported from this country again and, for example, the renowned German faunist Adolf Horion (1961) considered its occurrence in Poland or Baltic countries as doubtful.
The current area of the distribution of S.linearis extends over the boreal zone of the Palaearctic, from Norway and Denmark, through Sweden, Finland and Siberia to the Far East (it was recently discovered in northern Manchuria – Zhang et al. 1995), reaching – through Estonia, Latvia and Lithuania – as far south as NE Poland, Belarus and Ukraine; relict localities are also spread over the mountainous regions of central and southern Europe (France, Switzerland, Germany, Austria, Italy, Slovenia and Bohemia – Borowski 2007), as well as the Caucasus (Horion 1961, Geis 2002) and on Corsica (Sainte-Claire Deville 1902). Its recent discovery in northern Iran (Liu et al. 2016, Nardi and Audisio 2016) seems highly doubtful: the locality, albeit close to the border of Azerbaijan, is practically devoid of vegetation, with the climate unsuitable for the development of this species, so the specimen seems either misidentified or mislabelled; one of us (JB) contacted the Iranian author of the respective publication for possible verification of its taxonomic identity, but the specimen proved unavailable not only for loan but even for taking a photograph.
In the years 2009–2017, in the course of the study of saproxylic insects on a burned site in Białowieża Primaeval Forest, the existence of this species has been discovered. Information concerning this discovery and the data supplementing the current knowledge on the ecology of the development of S.linearis are provided below.
Materials and methodsLocality and collection data
In 2009, the research programme was launched in Białowieża Primaeval Forest (Faliński 1986) to study the dynamics of the changes in species composition and abundance of (especially saproxylic) beetles after the disturbance caused by the forest brushwood fire on 28.04.2009. The studied area of the burned site (ca. 7 ha), in the compartment 105B of the Białowieża National Park, included natural stands with high proportion of old (120 years or more) trees in humid forest (60.5% of the disturbed area), boggy forest (24.5%) and humid mixed forest (13.2%). The site has been left without economic activity, with actions having been restricted to scientific research. The observations started a few days after the fire. Beetles were caught in 2009–2011 and 2015–2017 using 9 non-baited Moericke’s and 18 Netocia-type traps (Piętka and Borowski 2015) as well as 3 black funnel traps each baited with α-pinen, ethanol, fuscumol, ipsenol and ketol (https://www2.gov.bc.ca/assets/gov/farming-natural-resources-and-industry/forestry/forest-health/forest-health-docs/spruce-beetle-docs/spruce_beetle_funnel__traps.pdf). All traps were suspended 1-2 metres above ground.
In June 2017, we measured the diameter at breast height and extent of trunk scorching (from the ground level) of 140 living pine-trees in the burned area and evaluated (in a three-level scale: slight, medium, strong) the degree of insolation of each trunk. All trees had been carefully examined for the presence of S.linearis larval galleries.
Statistical methods
To assess the preferences of Stephanopachyslinearis for particular types of trees, we used a Generalised Linear Model (GLM), with binomial distribution for response variable (0 = tree not occupied by beetle, 1 = tree occupied by beetle). In the model, we used three explanatory variables: diameter of a tree degree of burning of the trunk (continuous, log-transformed variables) and exposure to the sun (categorical, three-level variable). All analyses were done using R (version 3.2.2) software.
Results
The first specimen of Stephanopachyslinearis (Fig. 1) was caught between 26.04.2015 and 11.05.2015 in the Netocia-type trap on living but scorched pine; it was identified in autumn 2016 and we decided to look for the species at the same place in 2017.
23499461-AFFC-57BD-AF42-D1F2EC77E7A7
Stephanopachyslinearis; the specimen collected in Białowieża Primaeval Forest in 2015 – dorsal view (photo by M. Sławski, 2017). Scale bar = 1 mm.
https://binary.pensoft.net/fig/214795
On 9.05.2017, we searched for the species on living insolated pines with visible scorching of the outer layer of bark (Fig. 2). The trees did not show any signs of weakness and, had well developed crowns with green needles. On the black, scorched surface of bark, we saw small (1.4–2.0 mm in diameter) rounded exit holes of S.linearis from the previous year (Fig. 3). Whittling the outer bark layer away exposed galleries in the inner layer or under the bark (Fig. 4). Tortuous, variously directed, frequently inter-crossing or overlapping galleries 1.0–2.2 mm in width, filled with fine brown sawdust, did not enter the xylem. Humidity in the feeding places was moderate with a tendency to low. In the galleries, one dead individual, remnants of five others and a living larva of S.linearis were found. Galleries occurred only in the scorched part of the trunk, at a height of 0.3 to 2 m, all around the tree.
9D707818-3077-5CA4-9D02-4279BC14E01F
Scots pines burned by fire – host trees of Stephanopachyslinearis (photo by J.M. Gutowski, 2017).
Larval galleries of Stephanopachyslinearis in the Scots pine bark (photo by J. Borowski, 2017).
https://binary.pensoft.net/fig/214798
Also in 2017, black, baited funnel traps were suspended near the place where S.linearis had been previously collected and, in one of them one more specimen (22.05.–5.06.2017) and remnants of another one (20.06.–3.07.2017) were found.
Tree preferences
Amongst the measured and analysed trees, 16, i.e. 11.5%, were infested. Logistic regression analysis showed that the beetles preferred trees significantly thinner (mean diameter at breast height: 28 cm) than the average (32 cm; GLM, z = -2.303, p = 0.021) and more strongly scorched (mean height of scorching 245 cm for infested trees vs. 195 cm for all trees, GLM, z = 3.050, p = 0.002), while insolation of trunks was irrelevant (GLM, z = 0.272, p = 0.786) (Fig. 5).
FFB6E6ED-805D-54DF-ADC2-56554824422B
Result of logistic regression in the study of significance between the trees colonised by Stephanopachyslinearis and the size of breast height and the height of the trunks’ blackened by fire.
https://binary.pensoft.net/fig/214799
Thus, it seems that Stephanopachyslinearis prefers thinner trees (it has not been found on the thickest ones), most frequently those of 15-34 cm diameter at breast height (14 trees) and only once it was found feeding on a pine of 38 cm diameter. On the basis of our observations, this species feeds only on living trees, on small subcortical surfaces, where fire had locally damaged the cambium and phloem; it heavily infests scorched pines (up to the height of 135–350 cm), avoiding those blackened only below 130 cm.
S.linearis was accompanied by the deadwatch beetle Ernobiusmollismollis (Linnaeus, 1758) (Ptinidae, Ernobiinae); their galleries were almost identical, but the anobiid prefers those places where the fire had killed the cambium, its larvae feed deeper, frequently damaging the cambium and outer layer of xylem. Its exit holes are somewhat larger and less regular.
DiscussionBionomy of Stephanopachyslinearis
Stephanopachyslinearis is bound biologically to conifers: in the northern zone mainly with Scotch pine (Pinussylvestris L.), in the alpine areas on various species of pines, larches, much less frequently spruces and firs (Brustel et al. 2013, Ranius et al. 2014, Nardi and Audisio 2016). As a pyrophilous species, it primarily infests living trees damaged by fire, being only rarely found on those showing mechanical wounds like bark flayings or local necrosis. Before intensive forest management, the species probably proliferated on areas of natural fires and, when fire did not occur, on lightning-damaged trees. Like the majority of bostrichids, swarming in Poland occurs in spring, from late April to June. As a representative of the subfamily Dinoderinae, females probably gnaw into the bark to lay eggs. Larval galleries, filled with brown cortical sawdust mixed with faeces, terminate with ovate pupal chambers, from which imagines gnaw their round exit holes through bark. The beetles may develop on the infested tree during at least several years (Wikars 2006).
The galleries of S.linearis run at the contact surface between bark and phloem, but larvae feed only in bark, not in living phloem, selecting the layer of dead dry phloem previously killed by fire and then overgrown with new, living tissue. The species prefers the unusual environment produced by, for example, fire. As thick bark protects phloem from being killed by fire, the beetles infest mainly thin-barked trees where the subcortical tissues could have been more easily “boiled”. Infestation occurs only after partial regeneration of phloem has occurred, usually 2–5 years after the fire (Wikars 2006).
Status of Stephanopachyslinearis in Europe
Stephanopachyslinearis is the 20-th – first of the family Bostrichidae (Gutowski and Przewoźny 2013) – of the beetle species listed in Annexes I and II of the Habitats Directive (Council Directive 92/43/EEC), which have been found in Poland. As a rare species, important for the European Union, it has been placed in Annex II of the Habitats Directive (code: 1926) (Gutowski and Przewoźny 2013; Council Directive 97/62/EC), as well as in the IUCN Red List and European red list of saproxylic beetles (Nieto and Alexander 2010) – in both cases with the category LC – and Red list of near-extinct and threatened animals in Poland (Pawłowski et al. 2002) as probably extinct (EX?). S.linearis could probably be considered at the Polish regional scale as “Critically Endangered (CR)” following the IUCN criteria B1ab (i,ii,iv) (IUCN 2014).
Apart from S.linearis, considered as an obligatory pyrophile, many other beetle species are associated with burned areas, which – as important causes of natural ecological disturbances – make a significant element of the strategy for preservation of biodiversity (Wikars 1992). In the past, ground fires were frequent in the Białowieża Primaeval Forest (Niklasson et al. 2010, Zin et al. 2015), as in Scandinavia (e.g. on the Swedish island Gotska Sandön – Niklasson 2015). The occurrence of many rare pyrophilous beetles (Gutowski and Jaroszewicz 2001, Gutowski et al. 2012, Niklasson 2015) in these areas is probably a consequence of that “igneous” past.
Concluding remarks
Unfortunately, despite its being listed in the EU Habitats Directive, S.linearis remains poorly known with regards to its biology of development, and many data from various reports placed from time to time in the internet are contradictory, making them difficult to interpret reliably.
The confirmation of the occurrence of S.linearis in Poland will probably stimulate more focused attention to the trees – especially pines – damaged by fire. The authors are convinced that the species can also be found elsewhere in the northeastern parts of Poland, perhaps on larger surfaces with more numerous infested trees (in Białowieża the relevant area was relatively small: 7 ha, 16 pines with galleries of S.linearis). This would allow more detailed studies hopefully resulting in the clarification of its bionomy, at the level of other, well recognised species listed on the EU Habitats Directive.
Acknowledgements
Our studies were partly funded by the Polish Ministry of Science and Higher Education, through the statute activity of the Forest Research Institute, in the frames of the project No. 24 06 07.
ReferencesBorowskiJ (2007) Bostrichidae. In: LöblISmetanaA (Eds) Catalogue of Palearctic Coleoptera., 320–328.BrustelHGouixNBouyonHRogéJ (2013) Les Stephanopachys de la faune ouest-paléarctique (ColeopteraBostrichidae): Distribution et reconnaissance des trois espèces françaises au service de l'application de la directive Habitats, Faune, Flore.69(1): 41–50.BurakowskiBMroczkowskiMStefańskaJ (1986) Katalog fauny Polski. Część XXIII, t. 11, Chrząszcze, Coleoptera. Dermestoidea, Bostrichoidea, Cleroidea i Lymexyloidea. PWN, Warszawa, 1–243.Council Directive (1992) 92/43/EEC of 21 May 1992 on the conservation of natural habitats and of wild fauna and flora. http://eur-lex.europa.eu/legal-content/EN/TXT/?uri=CELEX:31992L0043 [accessed 10 December 2017]Council Directive (1997) 97/62/EC of 27 October 1997 adapting to technical and scientific progress Directive 92/43/EEC on the conservation of natural habitats and of wild fauna and flora. https://publications.europa.eu/en/publication-detail/-/publication/4cd5bfb6-8e37-46f8-bcf3-78ba2052fe82 [accessed 10 December 2017]DominikJ (1958) PWN, Warszawa, 20 pp.FabriciusJCh (1792) Secundum classes, ordines, genera, species, adjectis synonimis, locis observationibus, descriptionibus. Tom 1. Pars II. Hafniae, 538 pp. https://doi.org/10.5962/bhl.title.125869FalińskiJB (1986) Junk Publishers, Dordrecht, 537 pp. https://doi.org/10.1007/978-94-009-4806-8GeisK-U (2002) Gebietsfremde Splintholz- und Bohrkäfer, nach Mitteleuropa mit Importholz und anderen Gütern eingeschleppt. Eine Bestandsaufnahme (Coleoptera: Lyctidae, Bostrichidae). Mitteilungen des Internationalen Entomologischen Vereins (Supplement 10): 1–106.GutowskiJMJaroszewiczB (Eds) (2001) Instytut Badawczy Leśnictwa, Warszawa, 403 pp.GutowskiJMKubiszDSućkoK (2012) Nacerdescarniolica (Gistel, 1834) (Coleoptera: Oedemeridae) – nowy chrząszcz dla polskiej fauny.31(4): 267–273.GutowskiJMPrzewoźnyM (2013) Program NATURA 2000 jako narzędzie ochrony chrząszczy (Coleoptera) w Polsce. Wiadomości Entomologiczne 32(Supl.): 5–40.HorionA (1961) Teil (Thorictidae bis Cisidae), Teredilia, Coccinellidae. Überlingen – Bodensee, 375 pp.IUCN (2014) Guidelines for using the IUCN Red List categories and criteria. Version 11. Prepared by the Standards and Petitions Subcommittee. http://www.iucnredlist.org/documents/redlistguidelines.pdf [accessed January 8, 2015]KugelannJG (1792) Verzeichniss der in einigen Gegenden Preussens bis jetzt entdeckten Käfer-Arten, nebst kurzen Nachrichten von denselben.1(4): 477–512.LinnaeusC (1758) Editio decima, reformata. Tomus 1. Holmiae, 823 pp.LiuL-YGhahariHBeaverRA (2016) An annotated synopsis of the powder post beetles of Iran (Coleoptera: Bostrichoidea: Bostrichidae).4(14): 1–22. https://doi.org/10.12976/jib/2016.4.14NardiGAudisioP (2016) Italian account for Stephanopachyslinearis (Kugelann, 1792), a species listed in Annex II of the Habitats Directive (Coleoptera: Bostrichidae).48(2): 131–136. https://doi.org/10.4081/fe.2016.185NietoAAlexanderKNA (Eds) (2010) European Red List of Saproxylic Beetles. Publications Office of the European Union, Luxembourg, VIII+46 pp.NiklassonM (2015) Skogshistoria och bränder på Gotska Sandön. Länsstyrelsen i Gotlands Lan. Visby, 58 pp.NiklassonMZinEZielonkaTFeijenMKorczykAFChurskiMSamojlikTJędrzejewskaBGutowskiJMBrzezieckiB (2010) 350-year tree-ring fire record from Białowieża Primeval Forest, Poland: Implications for Central European lowland fire history.98(6): 1319–1329. https://doi.org/10.1111/j.1365-2745.2010.01710.xPawłowskiJKubiszDMazurM (2002) Coleoptera Chrząszcze. In: GłowacińskiZ (Ed.) Czerwona lista zwierząt ginących i zagrożonych w Polsce., 88–110.PiętkaJBorowskiJ (2015) Substrat trocinowy z grzybnią w pułapkach do odłowu chrząszczy mycetobiontycznych. Studia i materiały CEPL w Rogowie 17, 44(3): 165–172.RaniusTBohmanPHedgrenOWikarsLOCarusoA (2014) Metapopulation dynamics of a beetle species confined to burned forest sites in a managed forest region.37(8): 797–804. https://doi.org/10.1111/ecog.00475Sainte-ClaireDeville J (1902) Sur l'existence en Corse d'espèces actuellement confinees dans la zone arctique (Col.). Bulletin de la Société Entomologique de France 1902: 332.WikarsL-O (1992) Skogsbränder och insekter.113(4): 1–11.WikarsL-O (2006) Naturvårdsverket, Stockholm, 77 pp. http://www.naturvardsverket.se/Documents/publikationer/620-5610-7.pdf [accessed 20 February 2018]ZhangQChuDYangY (1995) A new record genus of Bostrichidae (Coleoptera) from China.17(1): 75–76.ZinEDrobyshevIBernackiDNiklassonM (2015) Dendrochronological reconstruction reveals a mixed-intensity fire regime in Pinussylvestris – dominated stands of Białowieża Forest, Belarus and Poland.26(5): 934–945. https://doi.org/10.1111/jvs.12290