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  • Darren A. Pollock


The classification of the nine world species of Pytho Latreille is reviewed by study of adult, larval, and pupal stages. Keys are provided for separation of species in these three life stages. Taxonomic changes (senior synonym in brackets) include synonymy of P. fallax Seidlitz 1916 [= P. niger Kirby 1837]; P. americanus Kirby 1837 [= P. planus (Olivier 1795)]; P. deplanatus Mannerheim 1843 is transferred from a junior subjective synonym of P. depressus (Linnaeus 1767) to a junior subjective synonym of P. planus (Olivier 1795). Lectotype designations are provided for the following: P. seidlitzi Blair 1925; P. nivalis Lewis 1888; P. niger Kirby 1837; P. fallax Seidlitz 1916; P. abieticola J. Sahlberg 1875; and P. americanus Kirby 1837. Eight larval stage, and 12 adult stage characters were selected for cladistic analysis. Lacking out-group material, pupal characters were not analysed. Character states were polarized using a generalized out-group composed of the three other genera of Pythinae (all monobasic). Phylogenetic analysis based on these 18 characters suggests four monophyletic species-groups: P. seidlitzi group (P. seidlitzi Blair — North America); P. kolwensis group (P. strictus LeConte – North America, P. kolwensis C. Sahlberg —Fennoscandia and the U.S.S.R., P. nivalis Lewis — Japan); P. niger group (P. niger Kirby — North America, P. abieticola J. Sahlberg — Europe, P. jezoensis Kôno — Japan); P. depressus group [P. planus (Olivier, 1795) — North America, P. depressus (Linnaeus, 1767) — Europe and the U.S.S.R.]. Larval stage synapomorphies are relatively more important in defining the species-groups than are those of the adult stage. The ancestor of Pythidae may have been associated with Coniferae as early as the Jurassic. The common ancestor of Northern Hemisphere Pythinae became isolated upon Laurasia once separation from Gondwanaland occurred near the end of the Jurassic. Two of the species-groups have similar disjunctions in North America, Europe, and Japan. The relatively eastern distributions of the North American member of each suggests that the ancestor of each species-group was Euramerican, and underwent vicariance with the opening of the North Atlantic in the Middle Cretaceous. The present distribution of both species-groups is thought to have been caused by the same vicariant event. The ancestor of the P. depressus group, which is presently circumboreal, was probably widespread and could have been Asiamerican in distribution. In the middle to late Tertiary, evidence suggests that Beringia was covered with coniferous forest, and the ancestor of the P. depressus group probably extended across this land bridge. Final separation between any North American and European/Asian species occurred in the Late Miocene or Pliocene, when a cooling climate made possible the evolution of treeless tundra in the north.



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Abdullah, M. 1964. A revision of the genus Pilipalpus (Coleoptera, Anthicidae: Pedilinae). Beitr. Ent. 14: 39.
Abdullah, M. 1966. The taxonomie position of the Australian Anaplopus tuberculatus, with a proposed new subfamily (Anaplopinae) of the Tenebrionidae, and including remarks on the family status of the Merycidae(Coleoptera). Ent. News 77: 143147.
Allen, R.T. 1983. Distribution patterns among arthropods of the north temperate deciduous forest biota. Ann. Mo. bot. Gdn 70: 616628.
Andersen, J., and Nilssen, A.C.. 1978. The food selection of Pytho depressus L. (Col., Pythidae). Norw. J. Ent. 25: 225226.
Andersen, J., and Nilssen, A.C.. 1983. Intrapopulation size variation of free-living and tree-living Coleoptera. Can. Ent. 115: 14531464.
Amett, R.H. Jr., 1968. The Beetles of the United States (A Manual for Identification). The American Entomological Institute, Ann Arbor, MI.
Amett, R.H. Jr., 1985. American Insects. A Handbook of the Insects of America North of Mexico, van Nostrand Rheinhold Co., New York, NY.
Askevold, I.S. 1988. The genus Neohaemonia Székessy in North America (Coleoptera: Chrysomelidae: Donaciinae): systematics, reconstructed phylogeny, and geographic history. Trans. Am. ent. Soc. 113: 360430.
Austin, E.P., and LeConte, J.L.. 1874. Catalogue of the Coleoptera of Mt. Washington, N.H., by E.P. Austin; with descriptions of new species, by J.L. LeConte, M.D. Proc. Bost. Soc. nat. Hist. 16: 265276.
Ball, G.E. 1963. The distribution of the species of the subgenus Cryobius (Coleoptera, Carabidae, Pterostichus) with special reference to the Bering land bridge and Pleistocene refugia, pp. 133151in Gressitt, J.L. (Ed.), Pacific Basin Biogeography. Bishop Museum Press, Honolulu, HI.
Bethune, C.J.S. 1872. Insects of the northern parts of British America. Compiled by the Editor from Kirby's fauna Boreali-Americana: Insecta. Can. Ent. 4: 5257.
Blackburn, T. 1890. Notes on Australian Coleoptera with descriptions of new species. Part vii. Proc. Linn. Soc. N.S.W. 5: 303366.
Blackwelder, R.E. 1946. Fabrician genotype designations. Bull. Brooklyn ent. Soc. 41: 7278.
Blackwelder, R.E. 1947. The genotypes (of Coleoptera) fixed by Fabricius. Bull. Brooklyn ent. Soc. 42: 5157.
Blair, K.G. 1925. Further notes on the Pythidae [Coleoptera]. Entomologist's mon. Mag. 61: 209219.
Blair, K.G. 1928. Pars 99: Pythidae. 56 pp. in Junk, W., and Schenkung, S. (Eds.), Coleopteroram Catalogus. Volume 17. W. Junk, Berlin and s'Gravenhage.
Blatchley, W.S. 1910. An Illustrated Descriptive Catalogue of the Coleoptera or Beetles (Exclusive of the Rhynchophora) Known to Occur in Indiana. Nature Publishing Co., Indianapolis, IN.
Böving, A.G., and Craighead, F.C.. 1931. An illustrated synopsis of the principal larval forms of the order Coleoptera. Entomologica am. (n.s.) 11: 1351.
Bowditch, F.C. 1896. List of Mt. Washington Coleoptera. Psyche, Camb 7, Supplement 2: 111.
Brimley, C.S. 1938. The Insects of North Carolina, Being a List of the Insects of North Carolina and their Close Relatives. North Carolina Department of Agriculture, Raleigh, NC.
Burakowski, B. 1962. Obserwacje biologiczno-morfologiczne nad Pytho kolwensis C. Sahib. (Coleoptera, Pythidae) w Polsce. Fragm. faun. 10: 173204.
Burakowski, B. 1976. Coleoptera: Pythidae-Lagriidae-Alleculidae. Zeszyt 88, Rozmiazgowate-Pythidae. pp. 318in Mikolajczyk, W. (Ed.), Klucze do oznaczania owadów Polski. Wroclawska Drukarnia Naukowa, Warsaw.
Crowson, R.A. 1955. The Natural Classification of the Families of Coleoptera. N. Lloyd, London.
Crowson, R.A. 1966. Observations on the constitution and subfamilies of the family Melandryidae. Eos, Madr. 41: 507513.
Crowson, R.A. 1975. The evolutionary history of Coleoptera, as documented by fossil and comparative evidence. Atti del X Congresso Nazionale Italiano di Entomologia: 4790.
Crowson, R.A. 1980. On amphipolar distribution patterns in some cool climate groups of Coleoptera. Ent. Gener. 6: 281292.
Crowson, R.A. 1981. The Biology of the Coleoptera. Academic Press, London.
Degeer, C. 1775. Mémoires pour servir à l'histoire des Insectes. Band 5. Pierre Hosselberg, Stockholm.
DeLeon, D. 1934. An annotated list of the parasites, predators, and other associated fauna of the mountain pine beetle in western white pine and lodgepole pine. Can. Ent. 66: 5161.
Donoghue, M.J. 1985. A critique of the biological species concept and recommendations for a phylogenetic alternative. Bryologist 88: 172181.
Emden, F.I. van. 1942. A key to the genera of larval Carabidae (Col.). Trans. R. ent. Soc. Lond. 92: 199.
Emmons, E. 1854. Agriculture of New York: Comprising an Account of the Classification, Composition, and Distribution of the Soils and Rocks, and of the Climate and Agricultural Productions of the State; Together with Descriptions of the More Common and Injurious Species of Insects. Volume 5. C. van Benthuysen, Albany, NY.
Fabricius, J.C. 1792. Entomologia Systematica emendata et aucta. Secundum classes, ordines, genera, species adjectis synonimis, locis, observationibus, descriptionibus. Tomus I, part 2, Proft., Hafniae.
Fabricius, J.C. 1801. Systema Eleutheratorum secundum ordines, genera, species, adjectis locis, observationibus, descriptionibus. Tomus I. Bibliopol. Acad., Kiliae.
Fauvel, A. 1889. Liste des Coléoptères communs a l'Europe et a l'Amérique du Nord, d'après le catalogue de M.J. Hamilton, avec remarques et additions. Revue Ent. 8: 92174.
Füessly, J.C. 1782. Archiv des Insectengeschichte. Heft 2. H. Steiner, Zürich and Winterthur.
Gistel, J.N. 1848. Naturgeschichte des Thierreichs für höhere Schulen. Hoffmann, Stuttgart.
Gistel, J.N. 1857. Achthundert und zwanzig neue oder unbeschreibene wirbellose Thiere. Vacuna 2: 513606.
Goulet, H. 1977. Technique for the study of immature Coleoptera in glycerin. Coleopts Bull. 31: 381382.
Grill, C. 1896. Förteckning öfver Skandinaviens, Danmarks och Finlands Coleoptera jämte deras synonymi och geografiska utbredning. Hospitals Boktryckeri, Stockholm.
Haack, R.A., and Slansky, F. Jr., 1987. Chapter 15. Nutritional ecology of wood-feeding Coleoptera, Lepidoptera, and Hymenoptera. pp. 449486in Slansky, F. Jr., and Rodriguez, J.G. (Eds.), Nutritional Ecology of Insects, Mites, and Spiders. J. Wiley & Sons, Inc., New York, NY.
Hallam, A. 1981. Relative importance of plate movements, eustasy, and climate in controlling major biogeographical changes since the early Mesozoic. pp. 303330in Nelson, G., and Rosen, D.E. (Eds.), Vicariance Biogeography, a Critique. Columbia-University Press, New York, NY.
Hamilton, J. 1889. Catalogue of the Coleoptera common to North America, northern Asia and Europe, with distribution and bibliography. Trans. Am. ent. Soc. 16: 88162.
Hamilton, J. 1890. Proposed corrections of specific names to harmonize Mr. Henshaw's catalogue of the North American Coleoptera, with the generally accepted European nomenclature, with relation to the species common to the two continents. Entomologica am. 6: 4144.
Hamilton, J. 1894 a. Catalogue of the Coleoptera of Alaska, with the synonymy and distribution. Trans. Am. ent. Soc. 21: 138.
Hamilton, J. 1894 b. Catalogue of the Coleoptera common to North America, northern Asia and Europe, with distribution and bibliography. Trans. Am. ent. Soc. 21: 345416.
Hamilton, W.D. 1978. Evolution and diversity under bark. pp. 154175in Mound, L.A., and Waloff, N. (Eds.), Diversity of Insect Faunas (Symposia of the Royal Entomological Society of London, 9). Blackwell Scientific Publications, New York, NY.
Hansen, V. 1945. Biller XII. Heteromerer. Larverne ved Sv. G. Larsson. Danm. Fauna 50: 1293.
Hatch, M.H. 1965. Family Pythidae. pp. 8488in The Beetles of the Pacific Northwest. Part IV: Macrodactyles, Palpicornes, and Heteromera. University of Washington Press, Seattle, WA.
Hayashi, N. 1969. On the larvae of some species of small families of Cucujoidea in Japan (Coleoptera). Insecta matsum. Suppl. 7: 19.
Hayashi, N., Fukuda, A., and Kurosa, K.. 1959. Coleoptera, pp. 392545in Kawada, et al. (Eds.), Illustrated Insect Larvae of Japan. [In Japanese.]
Hennig, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana, IL.
Heppner, J.B., and Lamas, G.. 1982. Acronyms for world museum collections of insects, with an emphasis on Neotropical Lepidoptera. Bull. ent. Soc. Am. 28: 305315.
Herbst, J.F.W. 1799. Natursystem aller bekannten in- und ausländischen Insekten. Der Käfer. VIII Theil. Pauli, Berlin.
Horn, G.H. 1872. Descriptions of some new North American Coleoptera. Trans. Am. ent. Soc. 4: 143152.
Horn, G.H. 1886. A review of the species described by Olivier in the “Entomologie”. Trans. Am. ent. Soc. 13: 135144.
Horn, G.H. 1888. Miscellaneous coleopterous studies. Trans. Am. ent. Soc. 15: 2648.
Horn, G.H., and LeConte, J.L.. 1883. Classification of the Coleoptera of North America. Smithson. misc. Collns 507.
Howden, A.T., and Howden, H.F.. 1981. The larva and adult biology of Rhinosimus viridiaeneus (Coleoptera: Salpingidae). Can. Ent. 113: 10551060.
Iablokoff-Khnzorian, S.M. 1985. Les Pythidae paléarctiques (Coleoptera). D. ent. Zt. (n.F.) 32: 193229.
International Commission on Zoological Nomenclature. 1985. International Code of Zoological Nomenclature, 3rd edition. University of California Press, Berkeley and Los Angeles, CA.
Kaszab, Z. 1969. 71. Fam. Pythidae. pp. 9299in Freude, H., Harde, K.W., and Lohse, G.A. (Eds.), Die Käfer Mitteleuropas. Band 8 Teredilia, Heteromera, Lamellicomia. Goecke & Evers, Krefeld.
Kavanaugh, D.H. 1972. Hennig's principles and methods of phylogenetic systematics. Biologist 54: 115127.
Kavanaugh, D.H. 1978. Hennigian phylogenetics in contemporary systematics: principles, methods, and uses. Chapter 8. pp. 139150in Romberger, J. et al. , (Eds.), Biosystematics in Agriculture. Beltsville Symposia in Agriculture Research [2]. Allanheld, Osmun & Co., Montclair, NJ.
Kiefer, H., and Moosbrugger, J.. 1942. Beitrag zur Coleopterenfauna des steirischen Ennstales und der angrenzenden Gebiete. Mitt. Münch, ent. Ges. 32: 486536.
Kirby, W. 1837. Part the fourth and last. The Insects. In Richardson, J. 1837. Fauna Boreali-Americana, or the Zoology of the Northern Parts of British America: Containing Descriptions of the Objects of Natural History Collected on the Late Northern Land Expeditions, under Command of Cpt. Sir John Franklin, R.N. J. Fletcher, Norwich and London.
Kôno, H. 1934. Die Heteromeren von Sachalin. Insecto matsum. 9: 2140.
Kôno, H. 1936. Beitrag zu den Pytho -arten Japans (Col.). Insecto matsum. 11: 3537.
Kurentsov, A.I. 1976. Significance of Beringian links in Holarctic insect zoogeography, pp. 529536in Kontrimavichus, V.L. (Ed.), Beringia in the Cenozoic Era. [Translation of Beringiia v kainozoe.] A.A. Balkema, Rotterdam.
Latreille, P.A. 1796. Precis des charactères générique des insectes, disposés dans un ordre naturel. Brive, Bourdeaux.
Latreille, P.A. 1810. Considerations générales sur l'ordre naturel des animaux composant les classes des Crustacés, des Arachnides et des Insectes avec un tableau méthodique de leurs genres disposés en familles. Schoell, Paris.
Lawrence, J.F. 1977. The family Pterogeniidae, with notes on the phylogeny of the Heteromera. Coleopts Bull. 31: 2556.
Lawrence, J.F. 1982. Coleoptera, pp. 482553in Parker, S.P. (Ed.), Synopsis and Classification of Living Organisms. Volume 2. McGraw-Hill Publ., New York, NY.
Lawrence, J.F. 1987. Notes on the classification of some Australian Tenebrionoidea (Coleoptera). J. Austr. ent. Soc. 26: 361362.
LeConte, J.L. 1850. Coleoptera, pp. 201242in Agassiz, J.L.R. (Ed.), Lake Superior, its Physical Character, Vegetation and Animals. Gould, Kendall and Lincoln, Boston, MA.
LeConte, J.L. 1866. New species of North American Coleoptera. Smithson. misc. Colins 6: 1177.
Lewis, G. 1888. On the capture of a new species of Pytho in Japan. Entomologist 21: 108109.
Lewis, G. 1895. On the Cistelidae and other heteromerous species of Japan. Ann. Nat. Hist. 15: 250278, 422–448.
Lindroth, C.H. 1953. Influence of Pleistocene climate changes on the insect fauna of northern Europe. Trans. IXth Int. Congr. Ent., Amsterdam 2: 145153.
Lindroth, C.H. 1957. The principal terms used for male and female genitalia in Coleoptera. Opusculo ent. 22: 241256.
Lindroth, C.H. 1969. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 1. Opuscula ent. Suppl. 25: I–XLVIII.
Lindroth, C.H. 1970. Survival of animals and plants on ice-free refugium during the Pleistocene glaciation. Endeavour 29: 129134.
Lindroth, C.H. 1979. The importance of Beringia as reflected in the present fauna, pp. 349354in Erwin, T.L., Ball, G.E., and Whitehead, D.R. (Eds.), Carabid Beetles: Their Evolution, Natural History and Classification. Dr W. Junk, The Hague.
Lindroth, C.H. 1960. Catalogus Coleopterorum Fennoscandiae et Daniae. Entomologiska Sällskapet i Lund.
Linnaeus, C. von. 1767. Systema Naturae, sive regna tria naturae systematice proposita per classes, ordines, genera et species… Lugduni Batavorum, Th. Haak. Edition XII, part 2, Laur. Savii., Hafniae.
Maca, J., Jelinek, J., and Svec, Z.. 1984. Faunistic records from Czechoslovakia: Coleoptera. Acta Entomologica Bohemoslovaca 81: 398.
Maddison, W.P., Donoghue, M.J., and Maddison, D.R.. 1984. Outgroup analysis and parsimony. Syst. Zool. 33: 83103.
Mannerheim, C.G. 1843. Beitrag zur Kaefer-Fauna der Aleutischen Inseln, de Insel Sitkha und neu-Californiens. Bull. Soc. Imp. Nat. Mose. 16: 175314.
Marshall, S.A. 1987. Systematics of Bitheca, a new genus of New World Sphaeroceridae (Diptera). Syst. Ent. 12: 355380.
Matsumura, S. 1931. 6000 Illustrated insects of Japan-empire. [Publishing data unknown, in Japanese.] Matthews J.V. Jr. 1979. Tertiary and Quaternary environments: historical background for an analysis of the Canadian insect fauna, pp. 310–86 in Danks H.V. (Ed.), Canada and its Insect Fauna. Mem. ent. Soc. Can. 108.
Matsumura, S. 1980. Tertiary land bridges and their climate: Backdrop for development of the present Canadian insect fauna. Can. Ent. 112: 10891103.
Matys, E.G., and Glushkova, L.A.. 1976. Orthoptera and Cassidinae: Zoogeographie peculiarities of northeastern U.S.S.R. and the role of Beringia in the formation of recent insect faunas, pp. 546554in Kontrimavichus, V.L. (Ed.), Beringia in the Cenozoic Era. [Translation of Beringiiavkainozoe.] A.A. Balkema, Rotterdam.
Morgan, A.V., and Morgan, A.. 1980. Faunal assemblages and distributional shifts of Coleoptera during the Lake Pleistocene in Canada and the northern United States. Can. Ent. 112: 11051128.
Nikitsky, N.B. 1986. The family Pilipalpidae Stat. N. (Coleoptera, Heteromera), its composition and taxonomie relationships. Zool. Zh. 65: 11781189. [In Russian.]
Noonan, G.R. 1986. Distribution of insects in the Northern Hemisphere: Continental drift and epicontinental seas. Bull. ent. Soc. Am. 32: 8084.
Nordal, I. 1987. Tabula rasa after all? Botanical evidence for ice-free refugia in Scandinavia reviewed. J. Biogeogr. 14: 377388.
Olivier, A.G. 1795. Entomologie, ou histoire naturelle des Insectes, avec leurs caractères génériques et spécifiques, leur description, leur synonymie et leur figure enluminée. Coléoptères. Tome 3, nos. 3565, Baudonin, Paris.
Paulus, H.F. 1971. Neue Pyrochroidae aus Nepal (Coleoptera, Heteromera), mit einer Diskussion der verwandtschaftlichen Verhältnisse der Familie. Z. ArbGem. öst. Ent. 23: 7585.
Payne, T.L. 1983. Nature of insect and host tree interactions. Z. angew. Ent. 96: 105109.
Perkins, P.D. 1980. Preventing distortion in preserved beetle pupae. Coleopts Bull. 34: 284.
Peterson, A. 1951. Larvae of Insects. Part II. Coleoptera, Diptera, Neuroptera, Siphonaptera, Mecoptera, Trichoptera. Edwards Bros. Inc., Ann Arbor, MI.
Pettersson, R.B. 1983. Pytho kolwensis C. Sahib. — en av skogsbruket hotad trädskalbagge. Natur i Norr 2: 2329.
Pettersson, R.B. 1988. Barkplattbaggarna (Col. Pythidae), en ny värdfamilj for bracksteklama CyanopterusflavatorÏabr. och Meteorus corax Marsh. Natur i Norr 7(1): 64.
Pic, M. 1912. Descriptions ou diagnoses et notes diverses. (Suite). Échange 28: 4951, 57–58.
Pic, M. 1929. Addenda et corrigenda du Coleopterorum Catalogus. Bull. Ann. Soc. ent. Belg. 69: 205208.
Pollock, D.A. 1988. A technique for rearing subcortical Coleoptera larvae. Coleopts Bull. 42(4): 311312.
Provancher, L'Abbe L. 1877. Petite faune entomologique du Canada précédée d'un traite elementaire d'entomologie. Volume I — les Coléoptères. C. Darveau, Québec.
Rafinesque, C.S. 1815. Analyse de la nature ou tableau de l'univers et des corps organisés. [Publ, unknown], Palermo.
Randall, J.W. 1838. Descriptions of new species of coleopterous insects inhabiting the state of Maine. Bost. J. nat. Hist. 2: 133.
Ring, R.A. 1981. The physiology and biochemistry of cold tolerance in Arctic insects. J. therm. Biol. 6: 219229.
Ring, R.A. 1982. Freezing-tolerant insects with low supercooling points. Comp. Blochem. Physiol. A 73: 605612.
Ring, R.A., and Tesar, D.. 1980. Cold-hardiness of the arctic beetle, Pytho americanus Kirby. Coleoptera, Pythidae (Salpingidae). J. Insect Physiol. 26: 763774.
Ring, R.A., and Tesar, D.. 1981. Adaptations to cold in Canadian Arctic insects. Cryobiology 18: 199211.
Ritchie, J.C., and MacDonald, G.M.. 1986. The patterns of post-glacial spread of white spruce. J. Biogeogr. 13: 527540.
Rozen, J.G. Jr., 1959. Systematic study of the pupae of the Oedemeridae (Coleoptera). Ann. ent. Soc. Am. 52: 299303.
Saalas, U. 1917. Die Fichtenkäfer Finnlands. Studien über die Entwicklungsstadien, Lebensweise, und geographische Verbreitung der an Picea excelsa Link, lebenden Coleopteren nebst einer Larvensbetimmungsta-belle. I. Allgemeiner Teil und Spezieller Teil 1. Suomalaisen Tiedeakatemian Justantama, Helsinki.
Sahlberg, C.R. 1833. Dissertatio entomologica Insecta Fennica enumerans. Band 1. P. 29. J.C. Frenckel, Aboae.
Sahlberg, J. 1875. Ueber die Finnischen arten der Käfer- Gattung Pytho und deren Larven. Dt. ent. Z. 19: 219224 [and Dt. ent. Z. 20: 380, figs. 18–20 (1876)].
Sahlberg, C.R. 1892. Meddelanden från Sällskapets sammanträden (den 7 Februari 1891). Meddn Soc. Fauna Flora fenn. 18: 143265.
Schoenfeldt, H. von. 1891. Catalog der Coleopteren von Japan. Jb. nassau. Ver. Naturk. 44: 237274.
Seidlitz, G. von. 1916, 1917. Die letzten Familien der Heteromeren. űr. ent. Z. 1916: 113128, 397–498; 1917: 65–116.
Sharp, D., and Muir, F.A.G.. 1912. The comparative anatomy of the male genital tube in Coleoptera. Trans. R. ent. Soc. Lond. part III 1912: 477642.
Shpeley, D. 1986. Genera of the subtribe Metallicina and classification, reconstructed phylogeny and geographical history of the species of Euproctinus Leng and Mutchler (Coleoptera: Carabidae: Lebiini). Quaest. Ent. 22: 261349.
Silfverberg, H. (Ed.). 1979. Enumeratio coleopterorum Fennoscandiae et Daniae. Helsingfors Entomologiska Bytesförening, Helsinki.
Smith, D.B., and Sears, M.K.. 1982. Mandibular structure and feeding habits of three morphologically similar coleopterous larvae: Cucujus clavipes (Cucujidae), Dendroides canadensis (Pyrochroidae), and Pytho depressus (Salpingidae). Can. Ent. 114: 173175.
Smith, J.B. 1910. Annual report of the New Jersey State Museum including a report of the insects of New Jersey. 1909. MacCrellish & Quigley, Trenton, NJ.
Spilman, T.J. 1952. The male genitalia of the Nearctic Salpingidae. Coleopts Bull. 6: 913.
Spilman, T.J. 1954. Generic names of the Salpingidae and their type species (Coleoptera). J. Wash. Acad. Sci. 44: 8594.
Stork, N.E. 1980. A scanning electron microscope study of tarsal adhesive setae in the Coleoptera. J. Linn. Soc. 68: 173306.
Tanner, V.M. 1927. A preliminary study of the genitalia of female Coleoptera. Trans. Am. ent. Soc. 53: 550.
Watrous, L.E., and Wheeler, Q.D.. 1981. The out-group comparison method of character analysis. Syst. Zool. 30: 111.
Watt, J.C. 1974. A revised subfamily classification of Tenebrionidae (Coleoptera). N.Z. J. Zool. 1: 381452.
Watt, J.C. 1987. The family and subfamily classification and New Zealand genera of Pythidae and Scraptiidae (Coleoptera). Syst. Ent. 12: 111136.
Westwood, J.O. 1839. An Introduction to the Modern Classification of Insects. Longman, Orme, Brown, Green, and Longmans, London.
Wheeler, Q.D. 1986. Revision of the genera of Lymexylidae (Coleoptera: Cucujiformia). Bull. Am. Mus. nat. Hist. 183: 113210.
Whitehead, D.R. 1972. Classification, phylogeny, and zoogeography of Schizogenius Putzeys (Coleoptera: Carabidae: Scantini). Quaest. Ent. 8: 131348.
Whitehead, D.R. 1973. Late-Wisconsin vegetational changes in unglaciated eastern North America. Quaternary Res. 3: 621631.
Wickham, H.F. 1899. The Coleoptera of Canada. XXXI. The Pythidae of Ontario and Quebec. Can. Ent. 31: 5761.
Wickham, H.F. 1913. Fossil Coleoptera from the Wilson ranch near Florissant, Colorado. Bull. Labs nat. Hist. St. Univ. Ia. 6: 329.
Wiley, E.O. 1981. Phylogenetics: The Theory and Practice of Phylogenetic Systematics. J. Wiley, New York, NY.
Yanovskiy, V.M.,andDmitrienko, V.K.. 1983. Forest beetles (Coleoptera) of the Sayanoshushenskoe state reserve. Ent. Rev. 62: 4351.
Young, D.K. 1975. A revision of the family Pyrochroidae (Coleoptera: Heteromera) for North America based on the larvae, pupae, and adults. Contr. Am. ent. Inst. 11: 139.
Young, D.K. 1976. The systematic position of Sphalma quadricollis Horn (Col: Salpingidae: Pythini) as clarified by discovery of its larva. Coleopts Bull. 30: 227232.
Young, D.K. 1985. The true larva of Lecontia discicollis and change in the systematic position of the genus (Coleoptera: Boridae). Great Lakes Ent. 18: 97101.
Zachariassen, K.E. 1977. Effects of glycerol in freeze-tolerant Pytho depressus L. (Col., Pythidae). Norw. J. Ent. 24: 2529.
Zachariassen, K.E. 1979. The mechanism of the cryoprotective effect of glycerol in beetles tolerant to freezing. J. Insect Physiol. 25: 2932.
Zachariassen, K.E. 1980. The role of polyols and nucleating agents in cold-hardy beetles. J. Comp. Physiol. B 140: 227234.

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