Published January 23, 2026 | Version v1
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A taxonomic review of Boreus (Mecoptera, Boreidae) with descriptions of two new Alaskan species

  • 1. University of Alaska Museum, Fairbanks, United States of America|University of Alaska Fairbanks, Fairbanks, United States of America
  • 2. University of Alaska Museum, Fairbanks, United States of America

Description

Boreus (Mecoptera, Boreidae) species are reviewed. Two new Alaskan species are described (Boreus tananaensis Kane, sp. nov., Boreus timaryi Kane, sp. nov.), a previously synonymized species is resurrected (Boreus gracilis Carpenter, 1935, stat. res.), and morphological descriptions are provided for all five Alaskan species. A key to male and female Alaskan Boreus species is provided. An estimate of the mitochondrial gene tree, based on COI DNA barcodes, is used to infer relationships, and morphological species are tested using five molecular species delimitation methods. What is known about the subgeneric classification of Boreus, and how Alaskan species are classified, is discussed. A checklist of all 33 currently valid species of the family Boreidae is provided.

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References

  • Banks N (1923) Mecoptera. In: McAtee WL, Hebard M, Ferris GF, Patch EM, Banks M, Forbes WT, Wickham HF, Alexander CP, Malloch JR, Walton WR, Viereck HL, Chamberlin RV (Eds) Insects, Arachnids, and Chilopods of the Pribilof Islands, Alaska. U.S.D.A Bureau of Biological Survey, North American Fauna 46: 158.
  • Beutel RG, Russell LK, Friedrich F, Hünefeld F, Simiczyjew B, Suzuki N, Dallai R, Jandausch K, Pohl H (2019) Neomecoptera, Boreidae, Boreinae, Boreus and Hesperoboreus. Handbook of Zoology. Nannomecoptera and Neomecoptera. De Gruyter, Berlin, Boston, 113–162. https://doi.org/10.1515/9783110272543-003
  • Blades DC (2002) A new species of Boreus (Mecoptera: Boreidae) from Vancouver Island, British Columbia. Journal of the Entomological Society of British Columbia 99: 133–140.
  • Bouckaert R, Vaughan TG, Barido-Sottani J, Duchêne S, Fourment M, Gavryushkina A, Drummond AJ (2019) BEAST 2.5: An advanced software platform for Bayesian evolutionary analysis. PLoS Computational Biology 15(4): e1006650. https://doi.org/10.1371/journal.pcbi.1006650
  • Brauer F (1876) The Neuroptera of Europe and especially of Austria with regard to their geographical distribution. Festschrift der zoologisch-botanischen Gesellschaft in Wien, 263–300. [In German]
  • Burrows M (2011) Jumping mechanisms and performance of snow fleas (Mecoptera, Boreidae). The Journal of Experimental Biology 214(14): 2362–2374. https://doi.org/10.1242/jeb.056689
  • Byers GW (1955) A new species of Boreus (Mecoptera: Boreidae) from Colorado. Occasional Papers of the Museum of Zoology University of Michigan 562: 1–4.
  • Byers GW (1961) An Unusual New Species of Boreus (Mecoptera: Boreidae) from Oregon. Journal of the Kansas Entomological Society 34(2): 74.
  • Byers GW, Shaw SR (1999) A new species of Boreus (Mecoptera: Boreidae) from Wyoming. Journal of the Kansas Entomological Society 72(3): 322–326.
  • Caron DM (1967) Habitat and distribution of Mecoptera in East Tennessee. Journal. Tennessee Academy of Science 42(3): 73–77.
  • Carpenter FM (1931) Revision of the Nearctic Mecoptera. Bulletin of the Museum of Comparative Biology 72: 205–277.
  • Carpenter FM (1933) A new Boreus from British Columbia (Mecoptera). Canadian Entomologist 65(4): 94–95. https://doi.org/10.4039/Ent6594-4
  • Carpenter FM (1935) New Nearctic Mecoptera, with notes on other species. Psyche (Cambridge, Massachusetts) 42(2): 105–122. https://doi.org/10.1155/1935/29562
  • Carpenter FM (1936) Descriptions and records of Nearctic Mecoptera. Psyche (Cambridge, Massachusetts) 43(2–3): 56–64. https://doi.org/10.1155/1936/26817
  • Cockle JW (1917) Notes on the hybernation of some larvae and the movement of Boreus in the snow. Journal of the Entomological Society of British Columbia 10: 14–15.
  • Cooper KW (1951) Compound sex chromosomes with anaphasic precocity in the male mecopteran, Boreus brumalis Fitch. Journal of Morphology 89(1): 37–57. https://doi.org/10.1002/jmor.1050890104
  • Cooper KW (1972) A southern Californian Boreus, B. notoperates n. sp. I. Comparative morphology and systematics (Mecoptera: Boreidae). Psyche (Cambridge, Massachusetts) 79(4): 269–283. https://doi.org/10.1155/1972/56294
  • Cooper KW (1974) Sexual biology, chromosomes, development, life histories and parasites of Boreus, especially of B. notoperates. A Southern California Boreus. II. (Mecoptera: Boreidae). Psyche (Cambridge, Massachusetts) 81(1): 84–120. https://doi.org/10.1155/1974/48245
  • Dellicour S, Flot JF (2018) The hitchhiker's guide to single‐locus species delimitation. Molecular Ecology Resources 18(6): 1234–1246. https://doi.org/10.1111/1755-0998.12908
  • Esben-Petersen P (1915) A synonymic list of the Order Mecoptera together with descriptions of new species. Entomologiske Meddelelser 10: 216–242.
  • Ezard T, Fujisawa T, Barraclough TG (2009) splits: SPecies' LImits by Threshold Statistics. R package version 1.0-14/r31.
  • Fitch A (1847) Winter insects of Eastern New York. American Journal of Agriculture and Science 5: 274–284.
  • Fujisawa T, Barraclough TG (2013) Delimiting species using single-locus data and the Generalized Mixed Yule Coalescent approach: A revised method and evaluation on simulated data sets. Systematic Biology 62(5): 707–724. https://doi.org/10.1093/sysbio/syt033
  • Gabdullina AU, Nikolajev GV (2015) A new species of the scorpion-fly of the genus Boreus Latreille (Mecoptera, Boreidae) from Altai mountains, Kazakhstan. Eurasian Entomological Journal 14(1): 32–34. [In Russian]
  • Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: Assessing the performance of PhyML 3.0. Systematic Biology 59(3): 307–321. https://doi.org/10.1093/sysbio/syq010
  • Hagen HA (1866) Synopsis of the genus Boreus. Entomologist's Monthly Magazine 3: 132.
  • Hågvar S, Østbye E (2011) Distribution of Boreus westwoodi Hagen, 1866 and Boreus hyemalis (L., 1767) (Mecoptera) in Norway. Norwegian Journal of Entomology 58: 73–80.
  • Hendricks P (2025) Snow-surface activity of California Snow Scorpionfly, Boreus californicus (Mecoptera: Boreidae), in western Montana, USA. Canadian Field Naturalist 138(3): 251–256. https://doi.org/10.22621/cfn.v138i3.3147
  • Hine JS (1901) A review of the Panorpidae of America north of Mexico. Bulletin of the Scientific Laboratories of Denison University 11: 241–164.
  • Hori S, Morimoto K (1996) Discovery of the family Boreidae (Mecoptera) from Japan, with description of a new species. Japanese Journal of Entomology 64(1): 75–81.
  • iNaturalist (2025) iNaturalist Research-grade Observations. iNaturalist.org. Occurrence dataset. [https://doi.org/10.15468/ab3s5x accessed via GBIF.org on 26 Nov 2025] https://www.gbif.org/occurrence/4018287737
  • Kalyaanamoorthy S, Minh BQ, Wong TK, Von Haeseler A, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14(6): 587–589. https://doi.org/10.1038/nmeth.4285
  • Kapli P, Lutteropp S, Zhang J, Kobert K, Pavlidis P, Stamatakis A, Flouri T (2017) Multi-rate Poisson tree processes for single-locus species delimitation under maximum likelihood and Markov chain Monte Carlo. Bioinformatics (Oxford, England) 33(11): 1630–1638. https://doi.org/10.1093/bioinformatics/btx025
  • Kari J (1985) Athabaskan Place Names in the Fairbanks Area. New River Times, 2–3.
  • Kaspřák D (2012) Morphological and molecular-taxonomic revision of the genus Boreus (Mecoptera: Boreidae) in the territory of the Czech and Slovak Republics. Master's thesis, University of Ostrava. [In Czech]
  • Klapálek F (1901) On new and little-known species of Palearctic Neuropteroids. Discourses of the Czech Academy of Emperor Franz Joseph for Sciences, Literature and the Arts. Class II. Mathematical and Natural Sciences 21: 1–19. [In Czech]
  • Kreithner A (2001) Boreus – species from the Alps: taxonomic characters and geographical distribution (Insecta, Mecoptera, Boreidae). Berichte des naturwissenschaftlichen-medizinischen Verein Innsbruck 88: 213–236. [In German]
  • Latreille PA (1805) Histoire Naturelle générale et particulière des Crustacés et des Insectes 13: 17–21. [In French]
  • Latreille PA (1816) Nouveau dictionnaire d'histoire naturelle 4: 152–153. [In French]
  • Lattin JD (1956) The first California records of Boreus californicus fuscus Carpenter (Mecoptera: Boreidae). The Pan-Pacific Entomologist 32(1): 81–82.
  • Lestage JA (1940) Pour l'historie des Boreus (Stegopteres - Mecopteres) I. Annales de la Société royale zoologique de Belgique 71: 5–22. [In French]
  • Lestage JA (1941) Pour l'historie des Boreus (Stegopteres - Mecopteres) II & III. Annales de la Société royale zoologique de Belgique 72: 5–29, 105–125. [In French]
  • Letunic I, Bork P (2021) Interactive Tree Of Life (iTOL) v5: An online tool for phylogenetic tree display and annotation. Nucleic Acids Research 49(W1): W293–W296. https://doi.org/10.1093/nar/gkab301
  • Lindsay BA, Lindsay JA (2010) Pribilof Islands, Alaska: The people: A historical account told through illustrated biographies. U.S. Department of Commerce, National Oceanic and Atmospheric Administration.
  • Linnaeus C (1767) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus differentiis, synonymis, locis (12th ed.). Laurentii Salvii, Holmiae 1(2): 915. https://doi.org/10.5962/bhl.title.156772
  • Lloyd LC (1934) Two species of Mecoptera from Alaska. The Pan-Pacific Entomologist 10(3): 119–120.
  • Looney C, Freeman MM, Asche M, Zack RS (2019) Snow Scorpionflies (Mecoptera: Boreidae) of the Hanford Reach National Monument. The Pan-Pacific Entomologist 95(1): 1–12. https://doi.org/10.3956/2019-95.1.1
  • Luo A, Ling C, Ho SY, Zhu CD (2018) Comparison of methods for molecular species delimitation across a range of speciation scenarios. Systematic Biology 67(5): 830–846. https://doi.org/10.1093/sysbio/syy011
  • Martynova OM (1954) Scorpionflies (Mecoptera) of the USSR fauna. Proceedings of the Zoological Institute of the USSR Academy of Sciences 15: 54–66. [In Russian]
  • Mayer K (1938) The genus Boreus (Panorpata Boreidae) in Czechoslovakia. Entomologické listy (Folia Entomologica) 2: 129–140. [In Czech]
  • McAtee WL, Hebard M, Ferris GF, Patch EM, Banks M, Forbes WTM, Wickham HF, Alexander CP, Malloch JR, Walton WR, Viereck HL, Chamberlin RV (1923) Insects, Arachnids, and Chilopods of the Pribilof Islands, Alaska. In: Biological Survey of the Pribilof Islands, Alaska. North American Fauna; no. 46: 129–244. https://doi.org/10.3996/nafa.46.0002
  • McLachlan R (1869) Note on Boreus hyemalis and B. Westwoodii. Transactions of the Royal Entomological Society of London 17(5): 399–401. https://doi.org/10.1111/j.1365-2311.1869.tb01115.x
  • Meinander M (1962) The Neuroptera and Mecoptera of Eastern Fennoscandia. Fauna Fennica 13: 1–87.
  • Mickoleit G, Mickoleit E (1976) Functional importance of the Tergal Apophysis of Boreus westwoodi (Hagen) (Insecta, Mecoptera). Zoomorphologie 85(2): 157–164. https://doi.org/10.1007/BF00995409 [In German]
  • Minh BQ, Nguyen MAT, Von Haeseler A (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution 30(5): 1188–1195. https://doi.org/10.1093/molbev/mst024
  • Minh BQ, Schmidt HA, Chernomor O, Schrempf D, Woodhams MD, Von Haeseler A, Lanfear R (2020) IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution 37(5): 1530–1534. https://doi.org/10.1093/molbev/msaa015
  • Navás SJ (1911) Two new European Boreus (Neuroptera). Revue Russe d'Entomologie 11: 277–278. [In Latin]
  • Navás SJ (1925) Entomological results of the Swedish Kamchatka Expedition 1920–1922 Part 6, Neuroptera and Mecoptera. Arkiv för Zoologi 18B(2): 1–6. [In German and Latin]
  • Nikolajev GV (1998) New and little-known species of insects for Central Asia (Insecta: Mantoptera, Raphidioptera, Neuroptera, Coleoptera, Mecoptera). Vestnik KazGU. Seriya Biologicheskaya 5: 38–46. [In Russian]
  • Nikolajev GV (2003) Scorpionflies of the family Boreidae (Mecoptera) of Asia. Tethys Entomological Research 8: 239–248. [In Russian]
  • Packard AS (1870) New or rare American Neuroptera, Thysanura, and Myriapoda. Proceedings of the Boston Society of Natural History 13: 405–411.
  • Panzer GWF (1796) Favnae insectorvm Germanicae initia, oder, Deutschlands Insecten. Nürnberg, In den Felseckerschen Buchhandlung. [In Latin] https://doi.org/10.5962/bhl.title.15007
  • Penny ND (1977) A systematic study of the family Boreidae (Mecoptera). The University of Kansas Science Bulletin 51: 141–217.
  • Pliginsky V (1914) New species of the genus Boreus Latr. from Crimea (Neuroptera, Panorpidae). Revue Russe d'Entomologie 14: 363–367. [In Russian]
  • Pliginsky V (1930) A new species of the genus Boreus Latr. (Neuroptera, Panorpidae). Revue Russe d'Entomologie 24: 15–16. [In Russian]
  • Plutenko AV (1984) A new species of Boreus (Mecoptera, Boreidae) from the Soviet Far East. Zoologicheskii Jurnal 63: 778–781. [In Russian]
  • Plutenko AV (1985) New and little-known species of Mecoptera from the Soviet Far East. Entomological Review 64(2): 113–118.
  • Puillandre N, Lambert A, Brouillet S, Achaz GJME (2012) ABGD, Automatic Barcode Gap Discovery for primary species delimitation. Molecular Ecology 21(8): 1864–1877. https://doi.org/10.1111/j.1365-294X.2011.05239.x
  • Puillandre N, Brouillet S, Achaz G (2021) ASAP: Assemble species by automatic partitioning. Molecular Ecology Resources 21(2): 609–620. https://doi.org/10.1111/1755-0998.13281
  • Pyszko P, Drgová M, Plášek V, Drozd P (2024) Non-random distribution of Boreus hyemalis among Bryophyte hosts: Evidence from field and laboratory tests. Insects 15(11): 878. https://doi.org/10.3390/insects15110878
  • Ratnasingham S, Hebert PDN (2013) A DNA-based registry for all animal species: The Barcode Index Number (BIN) system. PLOS ONE 8(7): e66213. https://doi.org/10.1371/journal.pone.0066213
  • Russell LK (1979) A new genus and a new species of Boreidae from Oregon (Mecoptera). Proceedings of the Entomological Society of Washington 81: 22–31.
  • Russell LK (2017) Collecting techniques for Caurinus (Mecoptera: Boreidae). Bulletin of the Oregon Entomological Society 2016(4): 1–3.
  • Schneider CA, Rasband WS, Eliceiri KW (2012) NIH Image to ImageJ: 25 years of image analysis. Nature Methods 9(7): 671–675. https://doi.org/10.1038/nmeth.2089
  • Shorthouse JD (1979) Observations on the snow scorpionfly Boreus brumalis Fitch (Boreidae: Mecoptera) in Sudbury, Ontario. Quaestiones Entomologicae 15: 341–344.
  • Sikes DS, Stockbridge J (2013) Description of Caurinus tlagu, new species, from Prince of Wales Island, Alaska (Mecoptera, Boreidae, Caurininae). ZooKeys 316: 35–53. https://doi.org/10.3897/zookeys.316.5400
  • Sikes DS, Bowser M, Morton JM, Bickford C, Meierotto S, Hildebrandt K (2017) Building a DNA barcode library of Alaska's non-marine arthropods. Genome 60(3): 248–259. https://doi.org/10.1139/gen-2015-0203
  • Stolbov VA, Galich DE, Nizovtsev DS (2018) New data on distribution of winter insects (Mecoptera: Boreidae; Diptera: Limoniidae) in Western Siberia. Far Eastern Entomologist = Dal'nevostochnyi Entomolog 359: 12–15. https://doi.org/10.25221/fee.359.3 [In Russian]
  • Strohmeier T (2023) Cryptic diversity in snow scorpionflies (Boreus spp.) in relation to cuticular chemistry and geography. Master's thesis. University of Graz. https://unipub.uni-graz.at/obvugrhs/content/titleinfo/9276561
  • Svensson SA (1972) Boreus Latreille, 1825 (Mecoptera). A synopsis of described species. Insect Systematics & Evolution 3(1): 26–32. https://doi.org/10.1163/187631272X00049
  • Tarbinsky SP (1962) Finding scorpion-flies, Boreus sp. n. (Mecoptera, Boreidae) in the foothills of the Kirghiz Ala-Too. Sbornik Entomologicheskikh Rabot. Akademiia Nauk Kirgiszk 1: 131–136. [In Russian]
  • Vertyankin AV, Zaycev AV (2021) New data on distribution of winter insects (Mecoptera, Boreidae; Diptera: Limoniidae) on the Sakhalin Island. Bulletin of the Sakhalin Museum 4: 101–106. [In Russian]
  • Wang JS, Jia LJ (2024) First discovery of snow scorpionflies (Mecoptera: Boreidae) from China. The Indochina Entomologist 1(11): 75–93. https://doi.org/10.70590/ice.2024.01.11
  • Willman R (1978) Redescription of Boreus gigas Brauer (Boreidae, Mecoptera), also a contribution to the variability of B. hyemalis (L.). Annalen des Naturhistorischen Museums in Wien 81: 525–532. [In German]
  • Withycombe CL (1922) On the Life‐history of Boreus hyemalis L. Transactions of the Royal Entomological Society of London 69(3–4): 312–318. https://doi.org/10.1111/j.1365-2311.1922.tb02813.x
  • Zangl L, Glatzhofer E, Schmid R, Randolf S, Koblmüller S (2021) DNA barcoding of Austrian snow scorpionflies (Mecoptera, Boreidae) reveals potential cryptic diversity in Boreus westwoodi. PeerJ 9: e11424. https://doi.org/10.7717/peerj.11424