Abstract
Morphological and genetic data indicate that Xyela fusca Blank, Kramp & Shinohara spec. nov. from Shikoku, Japan, plus an undescribed species from China are closely related to the Nearctic X. bakeri Konow, 1898. These species form the X. bakeri group, which is considered to be the sister clade of the Eurasian X. julii group. In addition to the X. alpigena group, the X. bakeri group is the second species group of Xyela with representatives occurring in both the Old and the New World. While X. bakeri is associated with Pinus (Pinus) species as the larval hosts, collection data of imagines suggest P. (Strobus) parviflora for X. fusca. This appears to be the second case, in which closely related Xyela species have switched hosts between species belonging to different subgenera of Pinus. Based on distribution data, migration of the common ancestor across Beringia appears likely, which would also have involved the change of the host subgenus. The six species of Xyela of Japan are keyed and new distribution data are presented. New, most northwesterly records of imagines from Yukon Territory (Canada) and field observations in Colorado and California (USA) suggest P. contorta as an additional larval host of X. bakeri.
References
Ashmead, W.H. (1898) Table of genera of Xyelidae. In: Dyar, H. G. Description of an unusual saw-fly larva belonging to the Xyelinae. Psyche, 8 (265), 212–214.
https://doi.org/10.1155/1898/68964
Benson, R.B. (1935) Swarming of Xyela julii Bréb. Entomologist's Monthly Magazine, Series 3, 71, 245.
Benson, R.B. (1938) European sawflies of the genus Xyela Dalman (sens. lat.) (Hymenoptera, Symphyta). Proceedings of the Royal Entomological Society of London, Series B: Taxonomy, 7 (2), 32–36.
https://doi.org/10.1111/j.1365-3113.1938.tb01240.x
Benson, R.B. (1960) Two new European species of Xyela Dalman (Hymenoptera: Xyelidae). Proceedings of the Royal Entomological Society of London, Series B: Taxonomy, 29 (7–8), 110–112.
https://doi.org/10.1111/j.1365-3113.1960.tb01153.x
Benson, R.B. (1961) The sawflies (Hymenoptera Symphyta) of the Swiss National Park and surrounding Area. Ergebnisse der wissenschaftlichen Untersuchungen im Schweizerischen Nationalpark, 7, 161–195.
Benson, R.B. (1962) Holarctic sawflies (Hymenoptera: Symphyta). Bulletin of the British Museum (Natural History), Entomology Series, 12, 379–409.
https://doi.org/10.5962/bhl.part.5879
Blank, S.M. (2002) The Western Palaearctic Xyelidae (Hymenoptera). In: Viitasaari, M. (Ed.), Sawflies (Hymenoptera, Symphyta). I. A review of the suborder, the Western Palaearctic taxa of Xyeloidea and Pamphilioidea. Tremex, Helsinki, pp. 197–233.
Blank, S.M. & Kramp, K. (2017) Xyela davidsmithi (Hymenoptera, Xyelidae), a new pine catkin sawfly with an unusual host association from the Sierra Nevada. Proceedings of the Washington Entomological Society. [submitted]
Blank, S.M., Kramp, K., Smith, D.R., Sundukov, Yu. N., Wei, M. & Shinohara, A. (2017) Big and beautiful: The Megaxyela species (Hymenoptera, Xyelidae) of East Asia and North America. European Journal of Taxonomy. [accepted]
Blank, S.M., Shinohara, A. & Byun, B.-K. (2005) The East Asian Xyela species (Hymenoptera: Xyelidae) associated with Japanese Red Pine (Pinus densiflora; Pinaceae) and their distribution history. Insect Systematics & Evolution, 36, 259–278.
https://doi.org/10.1163/187631205788838393
Blank, S.M., Shinohara, A. & Altenhofer, E. (2013) The Eurasian species of Xyela (Hymenoptera, Xyelidae): taxonomy, host plants and distribution. Zootaxa, 3629 (1), 1–106.
https://doi.org/10.11646/zootaxa.3629.1.1
Brébisson, J.B.G. de (1818) In: Blainville, H. de. Sur une nouveau genre d'insectes, de l'ordre des Hyménoptères (Pinicole). Bulletin des sciences, par la Société Philomatique, 1818 ([8]), 116–117.
Brikiatis, L. (2014) The De Geer, Thulean and Beringia routes: key concepts for understanding early Cenozoic biogeography. Journal of Biogeography, 41, 1036–1054.
https://doi.org/10.1111/jbi.12310
Burdick, D.J. (1961) A taxonomic and biological study of the genus Xyela Dalman in North America. University of California Publications in Entomology, 17, 285–355.
Critchfield, W.B. (1985) The late Quaternary history of lodgepole and jack pines. Canadian Journal of Forest Research, 15, 749–772.
https://doi.org/10.1139/x85-126
Critchfield, W.B. & Little, E.L. Jr. (1966) Geographic Distribution of the Pines of the World. U.S. Department of Agriculture, Forest Service, Miscellaneous Publication, 991, 1–97.
https://doi.org/10.5962/bhl.title.66393
Dalman, J.W. (1819) Några nya insect-genera. Kungliga Svenska Vetenskapsakademiens handlingar, 1819, 117–127.
Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9 (8), 772.
https://doi.org/10.1038/nmeth.2109
Debreczy, Z. & Rácz, I. (2011) Conifers Around the World. Conifers of the Temperate Zones and Adjacent Regions. Vols. 1–2. DendroPress Ltd., Budapest, 1089 pp.
Ebel, B.H. (1966) Rearing and occurrence of xyelid sawflies on slash and longleaf pines in North Florida (Hymenoptera: Xyelidae). Annals of the Entomological Society of America, 59, 227–229.
https://doi.org/10.1093/aesa/59.1.227
Eckenwalder, J.E. (2009) Conifers of the World. The Complete Reference. Timber Press, Portland & London, 720 pp.
Farjon, A. (2010) A Handbook of the World's Conifers. Vols. 1–2. Brill, Leiden & Boston, 1111 pp.
https://doi.org/10.1163/9789047430629Goulet, H. (1992) The Genera and Subgenera of the Sawflies of Canada and Alaska: Hymenoptera: Symphyta. In: The Insects and Arachnids of Canada. Part 20. Agriculture Canada, Publication 1876, pp. 1–235.
Guo, Q. & Ricklefs, R.E. (2000) Species richness in plant genera disjunct between temperate eastern Asia and North America. Botanical Journal of the Linnean Society, 134, 401–423.
https://doi.org/10.1006/bojl.2000.0345
Gussakovskij, V.V. (1935) Insectes Hyménoptères, Chalastogastra 1. In: Fauna SSSR. 2 (1). Moskva, Leningrad, pp. 1–453.
Hadley, A. (2010) CombineZP, Image Stacking Software. Available from: http://www.hadleyweb.pwp.blueyonder.co.uk/ (accessed 18 March 2011)
Hall, T.A. (1999) BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98.
Harris, R.A. (1979) A glossary of surface sculpturing. Occasional Papers in Entomology, 28, 1–31.
Hartig, T. (1837) Die Aderflügler Deutschlands mit besonderer Berücksichtigung ihres Larvenzustandes und ihres Wirkens in Wäldern und Gärten für Entomologen, Wald- und Gartenbesitzer. Die Familien der Blattwespen und Holzwespen nebst einer allgemeinen Einleitung zur Naturgeschichte der Hymenopteren. Vol. 1. Haude und Spener, Berlin, xiv + 416 pp, VIII tabs.
Huber, J.T. & Sharkey, M.J. (1993) Structure. In: Goulet, H. & Huber, J.T. (Eds.), Hymenoptera of the World: An Identification Guide to Families. Publication 1894/E. Agriculture Canada, Ottawa, pp. 13–59.
Available from: www.esc-sec.ca/aafcmonographs/hymenoptera_of_the_world.pdf (accessed 17 May 2017)
Jinno, T. & Yamamoto, S. (1960) [6. Catalog of plants.] In: Ishihara, T., Ito, T., Jinno, T., Soma, M., Nagai, K., Noguchi, M., Murakami, S. & Yamanaka, M. (Eds.), Scientific Researches of the Ishizuchi-Range. Ehime Shimbun Sha, Matsuyama, pp. 179–230. [in Japanese]
Kavanaugh, D.H. (1986) A systematic review of amphizoid beetles (Amphizoidae: Coleoptera) and their phylogenetic relationships to other Adephaga. Proceedings of the California Academy of Sciences, 44 (6), 67–109.
Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Meintjes, P. & Drummond, A.J. (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649.
https://doi.org/10.1093/bioinformatics/bts199
Kondo, T. & Miyake, M. (1974) [Observations on the biology of Xyela obscura (II). From the fall of the mature larvae to the ground to the emergence of the adults.] Heredity, 28, 100–102. [in Japanese]
Konow, F.W. (1897) Ueber die Xyelini (Tenthredinidarum tribus.). Entomologische Nachrichten, 23, 55–58.
Konow, F.W. (1898) Ueber einige neue Chalastogastra Arten. Entomologische Nachrichten, 24, 327–330.
Kremenetski, C.V., Liu, K. & MacDonald, G.M. (1998) The late Quaternary dynamics of pines in northern Asia. In: Richardson, D.M. (Ed.), Ecology and Biogeography of Pinus. Cambridge University Press, Cambridge, New York, Melbourne & Madrid, pp. 95–106 [paperback edition published 2000]
Kumar, S., Stecher, G. & Tamura, K. (2016) MEGA7: Molecular Evolutionary Genetics Analysis version 7.0 for bigger datasets. Molecular Biology and Evolution, msw054.
https://doi.org/10.1093/molbev/msw054
Marincovich, L. Jr. & Gladenkov, A. Yu. (1999) Evidence for an early opening of the Bering Strait. Nature, 397, 149–151.
Meiri, M., Lister, A.M., Collins, M.J., Tuross, N., Goebel, T., Blockley, S., Zazula, G.D., van Doorn, N., Guthrie, R.D., Boeskorov, G.G., Baryshnikov, G.F., Sher, A. & Barnes, I. (2014) Faunal record identifies Bering isthmus conditions as constraint to end-Pleistocene migration to the New World. Proceedings of the Royal Society B, 281, 20132167.
https://doi.org/10.1098/rspb.2013.2167
Millar, C.I. (1993) Impact of the Eocene on the evolution of Pinus L. Annals of the Missouri Botanical Garden, 80, 471–498.
https://doi.org/10.2307/2399795
Millar, C.I. (1998) Early evolution of pines. In: Richardson, D.M. (Ed.), Ecology and Biogeography of Pinus. Cambridge University Press, Cambridge, New York, Melbourne & Madrid, pp. 69–91 [paperback edition published 2000]
Mirov, N.T. (1967) The Genus Pinus. Ronald Press Company, New York, viii + 602 pp.
Miyake, M. & Kondo, T. (1974) [Discovery of the larvae of Xyela obscura and observations on their biology.] Heredity, 28, 92–95. [in Japanese]
Naito, T., Yoshida, H., Nakamine, H., Morita, T., Ikeda, T., Suzuki, H. & Nakanishi, A. (2004) [Species diversity of sawflies in Hyogo Prefecture, central Japan.] Museum of Nature and Human Activities, Hyogo, Monograph of Natural History and Environmental Science, 1, 1–2 + pl. 1–10 + 1–85. [in Japanese]
Nei M. & Kumar S. (2000) Molecular Evolution and Phylogenetics. Oxford University Press, New York, 352 pp.
Norton, E. (1869) Catalogue of the described Tenthredinidae and Uroceridae of North America. Transactions of the American Entomological Society, 2 [1868] (4), 321–368.
https://doi.org/10.2307/25076170
Nyman, T., Zinovjev, A.G., Vikberg, V. & Farrel, B.D. (2006) Molecular phylogeny of the sawfly subfamily Nematinae (Hymenoptera: Tenthredinidae). Systematic Entomology, 31, 569–583.
https://doi.org/10.1111/j.1365-3113.2006.00336.x
Okutani, T. (1977) Xyelidae. In: Ito, S., Okutani, T. & Hiura, I. (Eds.), Coloured Illustrations of the Insects of Japan. Vol. II. Hoikusha, Osaka, pp. 285. [in Japanese]
Okutani, T. (1982) [Symphyta of Japan IV.] Nature and Insects, 17, 19–21. [in Japanese]
Prous, M., Heidemaa, M. & Soon, V. (2011) Empria longicornis species group: taxonomic revision with notes on phylogeny and ecology (Hymenoptera, Tenthredinidae). Zootaxa, 2756, 1–39.
Prous, M., Vikberg V., Liston. A.D. & Kramp K. (2016) North-Western Palaearctic species of the Pristiphora ruficornis group (Hymenoptera, Tenthredinidae). Journal of Hymenoptera Research, 51, 1–54.
https://doi.org/10.3897/jhr.51.9162
Rasnitsyn, A.P. (1965) К познанию биологии, системы и филогенеза пилильщиков подсемейства Xyelinae (Hymenoptera, Xyelidae). [Notes on the biology, systematics and phylogeny of Xyelinae (Hymenoptera, Xyelidae).] Polskie Pismo Entomologiczne, 35, 483–519. [in Russian, English abstract]
Rasnitsyn, A.P. (1971) Эволюция перепончатокрылых насекомых семейства Xyelidae. [Evolution of the Hymenoptera of the Family Xyelidae.] Trudy Paleontologicheskogo Instituta, 130, 187–196. [in Russian]
Rasnitsyn, A.P. (1997) Xyela (Pinicolites) lata Smith (Vespida: Xyelidae), a living fossil sawfly from Western North America. Pan-Pacific Entomologist, 73, 231–235.
Richardson, D.M. & Rundel, P.W. (1998) Ecology and biogeography of Pinus: an introduction. In: Richardson, D.M. (Ed.), Ecology and Biogeography of Pinus. Cambridge University Press, Cambridge, New York, Melbourne & Madrid, pp. 3–46 [paperback edition published 2000]
Rohwer, S.A. (1910) Japanese sawflies in the collection of the United States National Museum. Proceedings of the United States National Museum, 39, 99–120.
https://doi.org/10.5479/si.00963801.1777.99
Rohwer, S.A. (1913) A synopsis of the Nearctic species of sawflies of the genus Xyela, with descriptions of other new species of sawflies. Proceedings of the United States National Museum, 45, 265–281.
https://doi.org/10.5479/si.00963801.45-1981.265
Sasaki, N. (2003) A 700-year landscape history of dwarf bamboo (Sasa)–Nikko fir community in the sub-alpine zone of Mt. Kamegamori, Shikoku Island, Japan. Japanese Journal of Ecology, 53, 219–232. [in Japanese, English abstract]
Say, T. (1824) [Insecta]. In: Keating, W.H. (Ed.), Narrative of an expedition to the source of the St. Peter's River, Lake Winnepeek (Winnepeg), Lake of the Woods 1823 under the command of Major Long. Vol. 2. H.C. Carey & I. Lea, Philadelphia, Appendix, pp. 268–378.
Schmidt, S., Taeger, A., Morinière, J., Liston, A., Blank, S.M., Kramp, K., Kraus, M., Schmidt, O., Heibo, E., Prous, M., Nyman, T., Malm, T. & Stahlhut, J. (2016) Identification of sawflies and horntails (Hymenoptera, ‘Symphyta’) through DNA barcodes: successes and caveats. Molecular Ecology Resource. [published online]
https://doi.org/10.1111/1755-0998.12614
Shinohara, A. (1983) Discovery of the families Xyelidae, Pamphiliidae, Blasticotomidae, and Orussidae from Taiwan, with description of four new species (Hymenoptera: Symphyta). Proceedings of the Entomological Society of Washington, 85, 309–320.
Shinohara, A. (1992) The sawfly genus Megaxyela (Hymenoptera, Xyelidae) in East Asia. Japanese Journal of Entomology, 60, 783–796.
Shinohara, A. (1995) The sawfly genus Pleroneura (Hymenoptera, Xyelidae) in East Asia. Japanese Journal of Entomology, 63, 825–840.
Shinohara, A. (1998) Collection records of two rare xyelid sawflies (Hymenoptera, Symphyta) in Japan. Japanese Journal of Systematic Entomology, 4, 389–390.
Shinohara, A. (2000) Sawflies and woodwasps of the Imperial Palace, Tokyo. Memoirs of the National Science Museum, 36, 295–305. [in Japanese, English summary]
Shinohara, A. (2005) Sawflies and woodwasps of the Akasaka Imperial Gardens and the Tokiwamatsu Imperial Villa, Tokyo. Memoirs of the National Science Museum, 37, 225–238. [in Japanese, English summary]
Shinohara, A. (2013) Sawflies of the shrine forest of Meiji Jingu, Tokyo, Japan. In: Committee of the Second General Survey on the Shrine Precincts of Meiji Jingu on the Occasion of 100th Anniversary after the Enshrinement (Ed.), Report of the Second General Survey on the Shrine Precincts of Meiji Jingu on the Occasion of 100th Anniversary after the Enshrinement. Shrine Office of Meiji Jingu, Tokyo, pp. 331–334. [in Japanese, English summary]
Shinohara, A. (2014) Sawflies and woodwasps of the Imperial Palace, Tokyo. Memoirs of the National Science Museum, 50, 461–475. [in Japanese, English summary]
Shinohara, A. (2016) The sawfly genus Pleroneura (Hymenoptera, Xyelidae) of Japan: P. itoi n. sp. and a key to species. Zootaxa, 4121 (4), 495–500.
https://doi.org/10.11646/zootaxa.4121.4.9
Shinohara, A., Hara, H., Kramp, K., Blank, S.M. & Kameda, Y. (2017) Bird droppings on chestnut leaves or sawfly larvae: DNA barcodes verify the occurrence of the archaic Megaxyela togashii (Hymenoptera, Xyelidae) in Hokkaido, Japan. Zootaxa, 4221 (2), 220–232.
https://doi.org/10.11646/zootaxa.4221.2.6
Smith, D.R. (1964) Description of the male of Xyelecia nearctica, with comments on the genus (Hymenoptera: Xyelidae). Pan-Pacific Entomologist, 40 (1), 54–56.
Smith, D.R. (1970) A new Nearctic Xyela causing galls on Pinus spp. (Hymenoptera: Xyelidae). Journal of the Georgia Entomological Society, 5, 69–72.
Smith, D.R. (1990) A new Xyela (Hymenoptera: Xyelidae) from western United States. Entomological News, 101, 9–12.
Stein, J.P.E.F. (1876) Einige neue dalmatinische, griechische und kleinasiatische Tenthredinoiden. Entomologische Zeitung, 37, 53–61.
Strobl, G. (1895) Beiträge zur geographischen Verbreitung der Tenthrediniden. V. Theil. Wiener entomologische Zeitung, 14, 277–279.
Taeger, A., Blank, S.M. & Liston, A.D. (2010) World catalog of Symphyta (Hymenoptera). Zootaxa, 2580, 1–1064.
Takeuchi, K. (1938) A systematic study on the suborder Symphyta (Hymenoptera) of the Japanese Empire (I). Tenthredo, 2, 173–229.
Takeuchi, K. (1955) [Coloured Illustrations of the Insects of Japan. Vol. II.] Hoikusha, Osaka, 190 pp., 68 pls. [in Japanese]
Togashi, I. (1954) [On the occurrence of Xyela kamtshatica in Honshu.] Kontyû, 22, 12. [in Japanese]
Togashi, I. (1961) Sawflies (Hym., Symphyta) of Mt. Hakusan. Life Study, 5, 27–42. [in Japanese, with English abstract]
Togashi, I. (1964) On the species of the genus Xyela (Hym., Xyelidae) of Japan. Life Study, 8, 51–52. [in English and Japanese]
Togashi, I. (1965) Tenthredinidae. In: Asahina, S., Ishihara, T. & Yasumatsu, K. (Eds.), Iconographia Insectorum Japonicorum Colore naturali edita, Volumen III. Hokuryûkan, Tokyo, pp. 245–254, pls. 123–126. [in Japanese]
Togashi, I. (1970) The comparative morphology of the internal reproductive organs of the Symphyta (Hymenoptera). Mushi, 43 (Supplement), 1–114.
Togashi, I. (1972) Discovery of the genus Xyelecia Ross (Hymenoptera, Xyelidae) from Japan. Kontyû, 40, 87–89.
Togashi, I. (1972) Sawflies of Mt. Hiko, Kyushu (Hym., Symphyta). Mushi, 46, 53–64.
Togashi, I. (1974) Symphyta of Shikoku, Japan (Hymenoptera). Transactions of the Shikoku Entomological Society, 12, 13–22.
Varga, O., Reshchikov, A. & Broad, G.R. (2014) First record of the genus Aplomerus Provancher, 1886 (Hymenoptera: Ichneumonidae: Xoridinae) from the Oriental region, with descriptions of two new species. Zootaxa, 3815 (4), 591–599.
https://doi.org/10.11646/zootaxa.3815.4.9Viitasaari, M. (2002) The Suborder Symphyta of the Hymenoptera. In: Viitasaari, M. (Ed.), Sawflies (Hymenoptera, Symphyta). I. A review of the suborder, the Western Palaearctic taxa of Xyeloidea and Pamphilioidea. Tremex, Helsinki, pp. 11–174.
Yates, H.O. III & Smith, D.R. (2009) History, distribution, damage, and life cycle of a pine shoot gall sawfly, Xyela gallicaulis (Hymenoptera: Xyelidae). Journal of Entomological Science, 44, 276–283.
Yoshida, H. (2006) [Symphyta (Hymenoptera) of Osaka Prefecture, Japan.] West Japan Hymenopterists' Club, Kakogawa, 4, 24 plates, 1–127. [in Japanese]
Zirngiebl, L. (1937) Neue oder wenig bekannte Tenthredinoiden (Hym.) aus dem Naturhistorischen Museum in Wien. Festschrift zum 60. Geburtstage von Professor Dr. Embrik Strand, 3, 335–350.