Vladimir Gusarov

• Kreiling, Agnes-Katharina; Aldará, Jón; Haraldsstovu, Kolbrún í; Andersen, Hans Jørgen; Bechberger, Oliver & Fahldieck, Moritz [Show all 15 contributors for this article] (2025). A species inventory on the island of Koltur: Setting the baseline for long-term monitoring. Fróðskaparrit - Faroese Scientific Journal. 71, p. 116–167. doi: 10.18602/ydr0xk51. Full text in Research Archive Show summary

  A species inventory was conducted on the island of Koltur (Faroe Islands) between 2019 and 2024, serving as a baseline for future monitoring of the terrestrial ecosystem of the island. The resulting species checklist is presented here. The inventory focussed on terrestrial species of birds, plants and invertebrates and resulted in a total of 519 species recorded from the island since 2019. Among those, 56 were birds, 146 plants, and 316 invertebrates, mainly insects. A high number of invertebrates were new records for Koltur (242) as well as for the Faroe Islands in general (49)

• Haugen, Marianne N.; Gusarov, Vladimir; Sikes, Derek S. & Struck, Torsten H. (2025). Insight Into the Phylogeny of the Intertidal Beetle Genus Aegialites (Coleoptera, Salpingidae). Zoologica Scripta. ISSN 0300-3256. doi: 10.1111/zsc.70035. Full text in Research Archive Show summary

  Aegialites Mannerheim, 1853 (Coleoptera: Salpingidae) is an understudied genus. These beetles inhabit rock crevices in the supralittoral to intertidal zones. They have widely separated coxae, large claws and are flightless. Despite their limited vagility they are broadly distributed along the northern Pacific coast, from California to northern Japan. Phylogenetic relationships and species richness within Aegialites remain unclear, with historical morphological assessments yielding 4–30 species. Recent molecular studies have suggested potential cryptic species may exist. We present the first molecular phylogenetic analysis of Aegialites based on one mitochondrial gene (cytochrome oxidase I) and one nuclear gene (ribosomal internal transcribed spacer 2) from over 160 specimens across 96 localities, as well as the complete mitochondrial genome and the 18S nuclear gene for several specimens. Our phylogenetic analyses revealed three major clades, one corresponding to Alaska, Canada and Northern California; a sister clade from Russia and Japan; and a separate clade for southern California. Haplotype networks, along with calculations of FST and P‐distance, supported our findings. Based on these analyses, we have evidence for seven species of Aegialites . However, fixing the taxonomy of Aegialites will require integration of morphology and further molecular analyses.

• Haugen, Marianne Nilsen; Gusarov, Vladimir & Struck, Torsten H (2024). The complete mitochondrial genome of Aegialites californicus (Motchoulsky, 1845) (insecta: coleoptera: salpingidae). Mitochondrial DNA Part B: Resources. 9(1), p. 214–218. doi: 10.1080/23802359.2024.2309255. Full text in Research Archive Show summary

  The flightless intertidal beetle genus Aegialites (family Salpingidae) is distributed along the Northern Pacific coasts, from California to Alaska and from Northern Japan to Kamchatka. Systematics of Aegialites and its phylogenetic relationships to other members of Salpingidae are unclear, and little genetic information is available. We here present the first complete mitochondrial genome of this genus, represented by Aegialites californicus (Motchoulsky, 1845) from Sonoma County, California, U.S.A. The complete mitochondrial genome of A. californicus is 15,899 bp long and comprises 13 protein-coding (PCG), two ribosomal RNA (rRNA) and 22 transfer RNA (tRNA) genes. The phylogenetic analysis places A.californicus as sister to other members of family Salpingidae. The mitochondrial genome sequence of A. californicus will contribute to future phylogenetic and taxonomic studies of genus Aegialites, family Salpingidae and superfamily Tenebrionoidea.

• Konstantinov, Alexander S.; Baselga, Andrés; Anderson, Robert S.; Carlton, Christopher; Gusarov, Vladimir & Ivie, Michael A. [Show all 9 contributors for this article] (2023). Moss-inhabiting beetles of the West Indies (Insecta: Coleoptera). Journal of Insect Biodiversity (JIB). ISSN 2538-1318. 38(2), p. 48–72. doi: 10.12976/jib/2023.38.2.3. Full text in Research Archive
• Kozlov, Mikhail V.; Zverev, Vitali; Gusarov, Vladimir; Korobushkin, Daniil I.; Krivosheina, Nina P. & Mattila, Jaakko [Show all 16 contributors for this article] (2022). Changes in Biomass and Diversity of Soil Macrofauna along a Climatic Gradient in European Boreal Forests. Insects. 13(1), p. 1–13. doi: 10.3390/insects13010094. Full text in Research Archive Show summary

  Latitudinal gradients allow insights into the factors that shape ecosystem structure and delimit ecosystem processes, particularly climate. We asked whether the biomass and diversity of soil macrofauna in boreal forests change systematically along a latitudinal gradient spanning from 60° N to 69° N. Invertebrates (3697 individuals) were extracted from 400 soil samples (20 × 20 cm, 30 cm depth) collected at ten sites in 2015–2016 and then weighed and identified. We discovered 265 species living in soil and on the soil surface; their average density was 0.486 g d·w·m−2. The species-level diversity decreased from low to high latitudes. The biomass of soil macrofauna showed no latitudinal changes in early summer but decreased towards the north in late summer. This variation among study sites was associated with the decrease in mean annual temperature by ca 5 °C and with variation in fine root biomass. The biomass of herbivores and fungivores decreased towards the north, whereas the biomass of detritivores and predators showed no significant latitudinal changes. This variation in latitudinal biomass patterns among the soil macrofauna feeding guilds suggests that these guilds may respond differently to climate change, with poorly understood consequences for ecosystem structure and functions.

• Kozlov, Mikhail V.; Oudendijk, Zowi; Forsman, Anders; Lanta, Vojtěch; Barclay, Maxwell V. L. & Gustafsson, Bert [Show all 17 contributors for this article] (2021). Climate shapes the spatiotemporal variation in color morph diversity and composition across the distribution range of Chrysomela lapponica leaf beetle. Insect Science. ISSN 1672-9609. doi: 10.1111/1744-7917.12966. Full text in Research Archive Show summary

  Color polymorphism offers rich opportunities for studying the eco-evolutionary mechanisms that drive the adaptations of local populations to heterogeneous and changing environments. We explored the color morph diversity and composition in a Chrysomela lapponica leaf beetle across its entire distribution range to test the hypothesis that environmental and climatic variables shape spatiotemporal variation in the phenotypic structure of a polymorphic species. We obtained information on 13 617 specimens of this beetle from museums, private collections, and websites. These specimens (collected from 1830–2020) originated from 959 localities spanning 33° latitude, 178° longitude, and 4200 m altitude. We classified the beetles into five color morphs and searched for environmental factors that could explain the variation in the level of polymorphism (quantified by the Shannon diversity index) and in the relative frequencies of individual color morphs. The highest level of polymorphism was found at high latitudes and altitudes. The color morphs differed in their climatic requirements; composition of colour morphs was independent of the geographic distance that separated populations but changed with collection year, longitude, mean July temperature and between-year temperature fluctuations. The proportion of melanic beetles, in line with the thermal melanism hypothesis, increased with increasing latitude and altitude and decreased with increasing climate seasonality. Melanic morph frequencies also declined during the past century, but only at high latitudes and altitudes where recent climate warming was especially strong. The observed patterns suggest that color polymorphism is especially advantageous for populations inhabiting unpredictable environments, presumably due to the different climatic requirements of coexisting color morphs.

• Anlaş, Si̇Nan & Gusarov, Vladimir (2020). Two new species of the genus leptobium casey from western anatolia, turkey (Coleoptera: Staphylinidae: Paederinae). Journal of Insect Biodiversity (JIB). ISSN 2538-1318. 14(2), p. 40–46. doi: 10.12976/jib/2020.14.2.2. Full text in Research Archive
• Gusarov, Vladimir (2018). Phylogeny of the Family Staphylinidae Based on Molecular Data: A Review. In Betz, Oliver; Irmler, Ulrich & Klimaszewski, Jan (Ed.), Biology of Rove Beetles (Staphylinidae). Life History, Evolution, Ecology and Distribution.. Springer. ISSN 9783319702551. p. 7–25. doi: 10.1007/978-3-319-70257-5_2. Full text in Research Archive
• Støa, Bente; Halvorsen, Rune; Mazzoni, Sabrina & Gusarov, Vladimir (2018). Sampling bias in presence-only data used for species distribution modelling: theory and methods for detecting sample bias and its effects on models. Sommerfeltia. ISSN 0800-6865. 38, p. 1–53. doi: 10.2478/som-2018-0001. Full text in Research Archive Show summary

  This paper provides a theoretical understanding of sampling bias in presence-only data in the context of species distribution modelling. This understanding forms the basis for two integrated frameworks, one for detecting sampling bias of different kinds in presence-only data (the bias assessment framework) and one for assessing potential effects of sampling bias on species distribution models (the bias effects framework). We exemplify the use of these frameworks to museum data for nine insect species in Norway, for which the distribution along the two main bioclimatic gradients (related to oceanicity and temperatures) are modelled using the MaxEnt method. Models of different complexity (achieved by use of two different model selection procedures that represent spatial prediction or ecological response modelling purposes, respectively) were generated with different types of background data (uninformed and background-target-group [BTG]). The bias assessment framework made use of comparisons between observed and theoretical frequency-of-presence (FoP) curves, obtained separately for each combination of species and bioclimatic predictor, to identify potential sampling bias. The bias effects framework made use of comparisons between modelled response curves (predicted relative FoP curves) and the corresponding observed FoP curves for each combination of species and predictor. The extent to which the observed FoP curves deviated from the expected, smooth and unimodal theoretical FoP curve, varied considerably among the nine insect species. Among-curve differences were, in most cases, interpreted as indications of sampling bias. Using BTG-type background data in many cases introduced strong sampling bias. The predicted relative FoP curves from MaxEnt were, in general, similar to the corresponding observed FoP curves. This indicates that the main structure of the data-sets were adequately summarised by the MaxEnt models (with the options and settings used), in turn suggesting that shortcomings of input data such as sampling bias or omission of important predictors may overshadow the effect of modelling method on the predictive performance of distribution models. The examples indicate that the two proposed frameworks are useful for identification of sampling bias in presence-only data and for choosing settings for distribution modelling options such as the method for extraction of background data points and determining the appropriate level of model complexity.

• Makarov, K.V.; Gusarov, Vladimir; Makarova, O.L.; Bizin, M.S. & Nekhaeva, A.A. (2018). The first data on beetles (Coleoptera) of the High Arctic Shokalsky Island (Kara Sea). Russian Entomological Journal. ISSN 0132-8069. 27(4), p. 387–398. doi: 10.15298/rusentj.27.4.06. Full text in Research Archive
• Struck, Torsten H; Feder, Jeffrey L.; Bendiksby, Mika; Birkeland, Siri; Oliveira, Josè Cerca De & Gusarov, Vladimir [Show all 13 contributors for this article] (2017). Finding Evolutionary Processes Hidden in Cryptic Species. Trends in Ecology & Evolution. ISSN 0169-5347. 33(3), p. 153–163. doi: 10.1016/j.tree.2017.11.007. Full text in Research Archive Show summary

  Cryptic species could represent a substantial fraction of biodiversity. However, inconsistent definitions and taxonomic treatment of cryptic species prevent informed estimates of their contribution to biodiversity and impede our understanding of their evolutionary and ecological significance. We propose a conceptual framework that recognizes cryptic species based on their low levels of phenotypic (morphological) disparity relative to their degree of genetic differentiation and divergence times as compared to non-cryptic species. We discuss how application of a more rigorous definition of cryptic species in taxonomic practice will lead to more accurate estimates of their prevalence in nature, better understanding of their distribution patterns on the tree of life, and increased abilities to resolve the processes underlying their evolution.

• Gusarov, Vladimir (2016). The importance of tracking introduced species: nine synonyms of Atheta (Dimetrotina) pasadenae Bernhauer, 1906 (Coleoptera, Staphylinidae, Aleocharinae). Zootaxa. ISSN 1175-5326. 4066(4), p. 390–398. doi: 10.11646/zootaxa.4066.4.2. Full text in Research Archive
• Osswald, Judith; Bachmann, Lutz & Gusarov, Vladimir (2013). Molecular phylogeny of the beetle tribe Oxypodini (Coleoptera: Staphylinidae: Aleocharinae). Systematic Entomology. ISSN 0307-6970. 38(3), p. 507–522. doi: 10.1111/syen.12011. Full text in Research Archive
• Elven, Hallvard; Bachmann, Lutz & Gusarov, Vladimir (2012). Molecular phylogeny of the Athetini-Lomechusini-Ecitocharini clade of aleocharine rove beetles (Insecta). Zoologica Scripta. ISSN 0300-3256. 41(6), p. 617–636. doi: 10.1111/j.1463-6409.2012.00553.x. Full text in Research Archive
• Epp, Laura; Boessenkool, Sanne; Bellemain, Eva; Haile, James; Esposito, Alfonso & Riaz, Tiayyba [Show all 19 contributors for this article] (2012). New environmental metabarcodes for analysing soil DNA: potential for studying past and present ecosystems. Molecular Ecology. ISSN 0962-1083. 21(8), p. 1821–1833. doi: 10.1111/j.1365-294X.2012.05537.x. Full text in Research Archive
• Gusarov, Vladimir (2011). ATHETINI Casey, 1910 and GEOSTIBINA Seevers, 1978 (Insecta, Coleoptera, STAPHYLINIDAE, ALEOCHARINAE): proposed conservation. Bulletin of Zoological Nomenclature. ISSN 0007-5167. 68(1), p. 54–60. doi: 10.21805/bzn.v68i1.a12. Full text in Research Archive
• Elven, Hallvard; Bachmann, Lutz & Gusarov, Vladimir (2010). Phylogeny of the tribe Athetini (Coleoptera: Staphylinidae) inferred from mitochondrial and nuclear sequence data. Molecular Phylogenetics and Evolution. ISSN 1055-7903. 57(1), p. 84–100. doi: 10.1016/j.ympev.2010.05.023. Full text in Research Archive
• Maruyama, Munetoshi; Klimaszewski, Jan & Gusarov, Vladimir (2008). Osakatheta yasukoae, a new intertidal genus and species of athetine rove beetles (Coleoptera, Staphylinidae, Aleocharinae) from Japan. Zootaxa. ISSN 1175-5326. p. 39–50. Full text in Research Archive
• Gusarov, Vladimir & Makranczy, Gyorgy (2004). Yosiityphlus Sawada, 1971, a new synonym of Thinobius Kiesenwetter, 1844 (Coleoptera: Staphylinidae). Zootaxa. ISSN 1175-5326. p. 1–4. Full text in Research Archive
• Gusarov, Vladimir (2004). A revision of the genus Lypoglossa Fenyes, 1918 (Coleoptera: Staphylinidae: Aleocharinae). Zootaxa. ISSN 1175-5326. p. 1–36. Full text in Research Archive
• Gusarov, Vladimir (2004). A revision of the Nearctic species of the genus Halobrecta Thomson, 1858 (Coleoptera: Staphylinidae: Aleocharinae) with notes on some Palaearctic species of the genus. Zootaxa. ISSN 1175-5326. p. 1–25. Full text in Research Archive
• Ashe, James S. & Gusarov, Vladimir (2004). Review of the genus Sableta Casey, 1910 (Coleoptera: Staphylinidae: Aleocharinae: Athetini). Zootaxa. ISSN 1175-5326. p. 1–15. Full text in Research Archive

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• Orlova-Bienkowskaja, Marina Yakovlevna; Bienkowski, Andrei; Volkovich, Mark & Gusarov, Vladimir (2019). Inventory on alien beetles of European Russia. Mukhametov G.V.. ISBN 9785904246808. 882 p. Full text in Research Archive

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• Gusarov, Vladimir & Struck, Torsten H (2025). Door 23: Who is walking on the snow? – Not just snow scorpionflies! doi: https:/blog.annelida.de/2025/12/23/door-23-who-is-walking-on-the-snow-not-just-snow-scorpionflies/. Full text in Research Archive
• Gusarov, Vladimir & Struck, Torsten H (2025). Door 9: Who is walking on the snow? – Snow scorpionflies, of course! doi: https:/blog.annelida.de/2025/12/09/door-9-who-is-walking-on-the-snow-snow-scorpionflies-of-course/. Full text in Research Archive
• Haugen, Marianne Nilsen; Gracia, Alberto Valero; Cerca, José; Fleming, James Frederick; Austin, Rita M & Gusarov, Vladimir [Show all 8 contributors for this article] (2020). International Happy Seasons – Blogs of the CEG research group. Full text in Research Archive
• Gusarov, Vladimir; Bachmann, Lutz & Struck, Torsten H (2020). Summer project at rocky shores – Blogs of the CEG research group. Full text in Research Archive
• Gusarov, Vladimir; Bachmann, Lutz & Struck, Torsten H (2020). Training in African Insect Biodiversity – Blogs of the CEG research group. Full text in Research Archive
• Gusarov, Vladimir; Bachmann, Lutz & Struck, Torsten H (2020). Beetles on the Falkland islands – Blogs of the CEG research group. Full text in Research Archive
• Struck, Torsten H; Feder, Jeffrey L.; Bendiksby, Mika; Birkeland, Siri; Oliveira, Josè Cerca De & Gusarov, Vladimir [Show all 13 contributors for this article] (2018). Cryptic Species – More Than Terminological Chaos: A Reply to Heethoff. Trends in Ecology & Evolution. ISSN 0169-5347. 33(5), p. 310–312. doi: 10.1016/j.tree.2018.02.008. Full text in Research Archive
• Gusarov, Vladimir; Makranczy, Gyorgy & Tronquet, Marc (2015). Staphylinidae: Oxytelinae. New synonym. In Loebl, Ivan & Loebl, D. (Ed.), Catalogue of Palaearctic Coleoptera. Hydrophyloidea - Staphylinoidea. Revised and updated edition. Volume 1. De Gruyter Brill. ISSN 9789004302761. p. 21–21. Full text in Research Archive
• Støa, Bente & Gusarov, Vladimir (2010). Den norske insekt-rødlista 2010 og 2015: bruk av stedfestede artsdata for objektiv evaluering av truethet. Full text in Research Archive
• Osswald, Judith; Bachmann, Lutz & Gusarov, Vladimir (2010). Towards a molecular phylogeny of the Aleocharine tribe Oxypodini (Coleoptera: Staphylinidae). Full text in Research Archive
• Osswald, Judith; Bachmann, Lutz & Gusarov, Vladimir (2010). First insight into phylogeny of the aleocharine tribe Oxypodini Thomson, 1859. Full text in Research Archive
• Elven, Hallvard; Bachmann, Lutz & Gusarov, Vladimir (2010). Disentangling the Athetini/Lomechusini knot. Molecular markers resolve relationships between the two largest aleocharine tribes (Coleoptera: Staphylinidae). Full text in Research Archive
• Elven, Hallvard & Gusarov, Vladimir (2009). Tribe Athetini Casey, 1910: The phylogeny of a very challenging beetle group. Full text in Research Archive
• Gusarov, Vladimir (2009). Naturmangfoldsmodellering, med insekteksempler. Full text in Research Archive
• Gusarov, Vladimir (2009). Taksonomiske utfordringer. Full text in Research Archive
• Gusarov, Vladimir (2009). Norwegian insect diversity: what we know and what we do not know. Introduction. Full text in Research Archive
• Gusarov, Vladimir (2009). Homologies in the internal structures of the aleocharine (Coleoptera: Staphylinidae) aedeagus: how widespread? Full text in Research Archive
• Epp, Laura; Boessenkool, Sanne; Bellemain, Eva; Esposito, Alfonso; Gusarov, Vladimir & Johnsen, Arild [Show all 13 contributors for this article] (2009). BarFrost – A new project for reconstructing past ecosystems by barcoding DNA from permafrost. Full text in Research Archive
• Elven, Hallvard & Gusarov, Vladimir (2009). Phylogeny of the tribe Athetini Casey, 1910: results and challenges. Full text in Research Archive
• Gusarov, Vladimir (2009). Staphylinids of the East African sky islands. Full text in Research Archive
• Elven, Hallvard & Gusarov, Vladimir (2008). Phylogeny of the tribe Athetini Casey, 1910 and its position in the subfamily Aleocharinae (Coleoptera: Staphylinidae). Full text in Research Archive
• Elven, Hallvard & Gusarov, Vladimir (2008). Phylogeny of the tribe Athetini Casey, 1910 and its position in the subfamily Aleocharinae (Coleoptera: Staphylinidae). Full text in Research Archive
• Gusarov, Vladimir (2008). Reproductive barriers on the beach: Speciation in the genus Aegialites Mannerheim (Coleoptera: Salpingidae). Full text in Research Archive
• Gusarov, Vladimir (2008). The Southern Appalachians as a biodiversity hotspot and a natural laboratory to study speciation (of staphylinid beetles, of course) (Coleoptera: Staphylinidae). Full text in Research Archive
• Gusarov, Vladimir (2008). Entomologisk kunnskapstatus i Norge - og hvordan øke denne? Full text in Research Archive
• Støa, Bente & Gusarov, Vladimir (2008). Økologisk nisjemodellering og utbredelsespredikering av norske rødlistede insektarter; bruk av stedfestede artsfunn for en objektiv evaluering av rødlistestatus. Full text in Research Archive
• Elven, Hallvard & Gusarov, Vladimir (2007). Phylogeny of the tribe Athetini Casey, 1910: a combined morphological/molecular approach. Full text in Research Archive
• Gusarov, Vladimir (2007). Local endemics of the southern Appalachian “sky islands”: Do all groups of staphylinid beetles (Coleoptera: Staphylinidae) fit the same distribution pattern? Full text in Research Archive
• Gusarov, Vladimir (2007). A review of Nearctic leptotyphlines (Coleoptera: Staphylinidae: Leptotyphlinae). Using ecological niche modelling to predict geographical distributions of leptotyphlines in America north of Mexico. Full text in Research Archive
• Gusarov, Vladimir (2006). Where are the Nearctic leptotyphlines (Coleoptera: Staphylinidae: Leptotyphlinae) and the obscure art of soil flotation? Full text in Research Archive
• Gusarov, Vladimir (2006). 1+1+1+1=4? or 5? or 6? Full text in Research Archive
• Gusarov, Vladimir (2006). Molecular phylogeny of Athetini: progress report and call for specimens suitable for DNA work. Full text in Research Archive
• Gusarov, Vladimir (2005). Fooling with Lee’s catalogue. Full text in Research Archive Show summary

  Despite the fact that the title is not very serious, the talk itself is (was). In my talk I analyzed the data from the World Catalogue of Staphylinidae (Herman 2001) to reveal trends in taxon description, revision and synonymy, produce species accumulation curves for different regions of the world, and make some predictions.

• Gusarov, Vladimir (2005). Phylogeny and classification of the staphylinid beetle tribe Athetini with particular reference to Holarctic taxa. Full text in Research Archive
• Gusarov, Vladimir (2005). Modeling the distribution of Lypoglossa (Insecta: Coleoptera: Staphylinidae: Aleocharinae). Full text in Research Archive
• Gusarov, Vladimir (2005). The amazing diversity of Staphylinid beetle. Full text in Research Archive
• Gusarov, Vladimir (2004). Revision of the tribe Athetini (Insecta: Coleoptera: Staphylinidae) of America north of Mexico: preliminary results. Full text in Research Archive
• Haugen, Marianne Nilsen; Struck, Torsten H; Gusarov, Vladimir & Sikes, Derek (2026). Species diversity, geographic ranges and genetic connectivity in organisms with limited dispersal capacity. A case study of the flightless intertidal beetle genus Aegialites Mannerheim, 1853. Universitetet i Oslo. Full text in Research Archive Show summary

  Flightless organisms are generally expected to have limited dispersal, yet some occupy remarkably large geographic ranges. This thesis investigates this apparent paradox in Aegialites, a genus of small, flightless beetles specialized to rocky intertidal habitats and distributed across much of the North Pacific. By combining molecular, genomic and population genomic approaches, the thesis establishes the first comprehensive evolutionary framework for Aegialites. The results support substantially fewer species than previously proposed based on morphology, while also revealing previously unrecognized diversity. The first mitochondrial and high-quality nuclear genomes for the genus were generated. The genomes were relatively compact, and comparisons with other beetles revealed no evidence that adaptation to the intertidal environment involved extensive changes in genome organization or gene content. Population genomic analyses revealed strong geographic structure, but striking differences in connectivity among regions. California populations were strongly genetically differentiated across relatively short distances, whereas populations from Russia and Japan retained evidence of connectivity across more than 1,500 kilometers. Together, these findings show that limited active dispersal can generate strong population structure locally without preventing broad geographic distributions over evolutionary timescales. Historical demographic processes and occasional long-distance dispersal may therefore both contribute to the distribution of these flightless beetles.

• Elven, Hallvard; Gusarov, Vladimir & Bachmann, Lutz (2013). Molecular phylogeny of the rove beetle tribes Athetini, Geostibini, and Lomechusini (Coleoptera: Staphylinidae: Aleocharinae). Akademika publishing. Full text in Research Archive

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