Hostname: page-component-586b7cd67f-2plfb Total loading time: 0 Render date: 2024-11-25T09:47:25.992Z Has data issue: false hasContentIssue false

GEOGRAPHICAL VARIATION, CLASSIFICATION, RECONSTRUCTED PHYLOGENY, AND GEOGRAPHICAL HISTORY OF THE CICINDELA SEXGUTTATA GROUP (COLEOPTERA: CICINDELIDAE)1

Published online by Cambridge University Press:  31 May 2012

Michael M. Kaulbars
Affiliation:
Department of Biology, Lakehead University, Thunder Bay, Ontario, Canada P7B 5El
Richard Freitag
Affiliation:
Department of Biology, Lakehead University, Thunder Bay, Ontario, Canada P7B 5El

Abstract

A systematic review was conducted for the Cicindela sexguttata group taxa. Comparative methods of examination were applied to adult structural and ecological characters for purposes of taxa diagnoses, and to establish a basis for the derivation of the group’s history.Based on 13 adult exoskeletal characters and 25 tests on populations throughout the group’s geographical range, a discriminant analysis identifies the most significant characters as being elytral maculation, body size, pilosity of the stipes, and number of sensory setae on the antennal scape. In addition, selected characters of the male and female genitalia are shown to identify all taxa within the group. Among biological comparisons the different number of mature eggs found in adult females of C. sexguttata Fabricius and C. denikei Brown indicates that their fecundity differs; and the larval burrow of C. denikei opens directly beneath rocks and stones, a habit unique in Cicindela. Seasonality profiles of taxa appear to be dictated by geographical location, and by phylogeny to a smaller degree. Species–soils associations indicate that C. sexguttata has a strong affinity to warm, moist and loamy soils, but C. denikei is correlated with sandy, silty till. For all species of the group, habitats occupied and limits of distribution to eastern Canada and the United States appear to be governed by soil and forest types.Three species of the group are recognized: C. sexguttata consisting of geographical populations varying considerably in adult characteristics and the problematic form C. harrisii Leng which may be considered a cryptic species; C. denikei; and C. patruela Dejean consisting of two subspecies C. p. patruela and C. p. consentanea Dejean.A reconstructed phylogeny of the C. sexguttata group based on methods of Hennig (1966) allies C. sexguttata and C. denikei as sister species, and C. patruela as an earlier lineage. Recognized as a stem group of the C. purpurea complex, the C. sexguttata group is postulated to have had its origins in forested eastern North America during the Late Miocene. Speciations of C. patruela and lineage C. sexguttataC. denikei are perceived to have occurred in the Pliocene, followed by speciations of C. sexguttata and C. denikei in the Late Pleistocene effected by continental ice mass advances and recessions.

Résumé

On trouvera ici la révision systématique des taxons du groupe Cicindela sexguttata. Des examens comparatifs des structures adultes et des caractéristiques écologiques ont abouti à la diagnose des taxons et ont permis d’établir les bases de l’histoire évolutive du groupe.Une analyse discriminante basée sur 13 caractères de l’exosquelette adulte et sur 25 tests démographiques dans toute la répartition géographique du groupe a permis de reconnaître que les principaux caractères diagnostiques sont la maculation des élytres, la taille du corps, la pilosité des stipes et le nombre de soies sensorielles sur le scape antennaire. De plus, des caractères particuliers reliés aux structures génitales mâles et femelles permettent d’identifier tous les taxons au sein du groupe. D’après les comparaisons biologiques, le nombre d’oeufs à maturité est différent chez les femelles de C. sexguttata Fabricius et chez les femelles de C. denikei Brown, ce qui indique que la fécondité diffère chez ces deux espèces; en outre, l’embouchure du terrier larvaire de C. denikei se trouve directement sous une pierre ou un caillou, ce qui ne s’est jamais vu chez une autre espèce de Cicindela. Les habitudes saisonnières des taxons semblent régies par la position géographique et, à un degré moindre, par la phylogénie. Les associations espèce – type de sol indiquent que C. sexguttata a une forte affinité pour les terreaux chauds, humides et riches, alors que C. denikei préfère les sols d’argile sablonneux et limoneux. Chez toutes les espèces du groupe, le choix des habitats et la répartition limitée à l’est du Canada et aux États-Unis semblent fonction des types de sol et de forêt.Trois espèces sont reconnues au sein du groupe : C. sexguttata, qui comprend d’une part des populations géographiques chez lesquelles les adultes ont des caractéristiques très variables, et d’autre part la forme problématique C. harrisii Leng (qui constitue peut-être une espèce cryptique), C. denikei et C. patruela Dejean, celle-là représentée par deux sous-espèces, C. p. patruela et C. p. consentanea Dejean.Une reconstitution de la phylogénie du groupe C. sexguttata selon les méthodes proposées par Hennig (1966) reconnaît C. sexguttata et C. denikei comme des espèces-soeurs et C. patruela comme une lignée plus ancienne. Reconnu comme une branche du complexe C. purpurea, le groupe C. sexguttata a probablement prix ses origines dans les forêts de l’est nord-américain vers la fin du Miocène. Chez C. patruela et chez la lignée C. sexguttata/C. denikei, la spéciation remonte probablement au Pliocène, suivie des spéciations respectives de C. sexguttata et de C. denikei vers la fin du Pleistocene à la suite des mouvements d’avancée et de retrait de la masse glaciaire continentale.

[Traduit par la rédaction]

Type
Articles
Copyright
Copyright © Entomological Society of Canada 1993

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

REFERENCES

Agriculture Handbook 296. 1978. Map. Land resource regions and major land resource areas of the United States. 1:7,500,000. Albers equal area projection, 42 × 68 cm, coloured. USDA Soil Conservation Service, Washington, DC.Google Scholar
Blackwelder, R.E. 1939. Fourth Supplement 1933 to 1938 (incl.) to the Leng Catalogue of Coleoptera of America, North of Mexico. J.D. Sherman, Jr., Mount Vernon, NY. 146 pp.Google Scholar
Borror, D.J., DeLong, D.M., and Triplehorn, C.A.. 1981. An Introduction to the Study of Insects. Saunders College Publishing, New York, NY. 827 pp.Google Scholar
Boyd, H.P. 1973. Collecting tiger beetles in the Pine Barrens of New Jersey. Cicindela 5: 112.Google Scholar
Boyd, H.P. 1978. The tiger beetles (Coleoptera: Cicindelidea) of New Jersey with special reference to their ecological relationships. Transactions of the American Entomological Society 104: 191242.Google Scholar
Boyd, H.P., and Associates. 1982. Checklist of Cicindelidae: The Tiger Beetles. Plexus Publishing, Inc., Marlton, NJ. 31 pp.Google Scholar
Brown, E.L. 1950. Deciduous Forests of Eastern North America. Blakiston Co., Philadelphia, PA. 596 pp.Google Scholar
Brown, W.J. 1934. New species of Coleoptera V. The Canadian Entomologist 66: 2224.CrossRefGoogle Scholar
Butzer, K.W. 1964. Environment and Archeology: Introduction to Pleistocene Geography. Aldine Publishers, Chicago, IL. 524 pp.Google Scholar
Casey, T.L. 1909. Studies in the Caraboidea and Lamellicornia. The Canadian Entomologist 41: 253284.CrossRefGoogle Scholar
Chapman, R.F. 1979. The Insects: Structure and Function. Elsevier Publishers, New York, Oxford. 819 pp.Google Scholar
Charpentier, T. de. 1825. Horae Entomologicae, Adjectis Tabulis Novem Coloratis, Apud A. Gosohorsky, Bibliopolam, Wratislaviae. 255 pp.CrossRefGoogle Scholar
Clayton, J.S., Ehrlich, W.A., Cann, D.B., Day, J.H., and Marshall, I.B.. 1977. Soils of Canada. Department of Agriculture, Supply and Services, Ottawa, Ont. 3 volumes.Google Scholar
Criddle, N. 1910. Habits of some Manitoba tiger beetles (Cicindela). The Canadian Entomologist 42: 916.CrossRefGoogle Scholar
Davis, C.A. 1903. The Cicindelidae of Rhode Island. Entomological News, Philadelphia 14: 270273.Google Scholar
Davis, W.T. 1910. Miscellaneous notes on collecting in Georgia. Journal of the New York Entomological Society 18: 8285.Google Scholar
Davis, W.T. 1912. Notes on the distribution of several species of tiger beetles. Journal of the New York Entomological Society 20: 1720.Google Scholar
Dawson, R.W. 1928. Biology of the tiger beetles with a key to the species of Cicindela found in Minnesota. University of Minnesota Agricultural Experiment Station, Technical Bulletin 56: 28.Google Scholar
Dejean, P.F.M.A. 1821. Catalogue des Coléoptères de la collection d'Auguste Dejean, Paris. pages 12.Google Scholar
Dejean, P.F.M.A. 1825. Species général des Coléoptères de la collection d'Auguste Dejean. Mequignon Marvais, Paris 1: 1463.Google Scholar
Dillon, L.S. 1956. Wisconsin climate and life zones in North America. Science 123: 167176.CrossRefGoogle ScholarPubMed
Dunn, G.A. 1978. Tiger Beetles of New Hampshire (Coleoptera: Cicindelidae). M.Sc. thesis, University of New Hampshire, Durham, NH. 161 pp.Google Scholar
Easton, N.S. 1909. A list of Coleoptera collected within 10 miles of Fall River, Massachusetts. Psyche 16: 3542.CrossRefGoogle Scholar
Eckhoff, D.E. 1939. Cicindelidae of Iowa (Coleoptera). Iowa State College Journal of Science 13: 201230.Google Scholar
Emmons, E. 1854. The Natural History of New York Agriculture. IV. Coleoptera. Gould, Kendall and Lincoln, Albany, NY. 272 pp.Google Scholar
Fabricius, J.C. 1775. Systema Entomologiae. Flensburg and Lipsiae, Korte. 832 pp.Google Scholar
Fabricius, J.C. 1801. Systema Eleutheratorum Secundum Ordines, Genera, Species 1: 232243.Google Scholar
Fowells, H.A. 1965. Silvics of forest trees of the United States. U.S. Department of Agriculture, Washington, DC. 762 pp.Google Scholar
Fox, H. 1910. Observations on Cicindelidae in northern Cape May Co., New Jersey during the summers of 1908–09. Entomological News 21: 7582.Google Scholar
Freitag, R. 1965. A revision of the North American species of the Cicindela maritima group with a study of hybridization between Cicindela duodecimguttata and oregona. Quaestiones Entomologicae 1: 87170.Google Scholar
Freitag, R. 1966. The female genitalia of four species of tiger beetles. The Canadian Entomologist 98: 942952.CrossRefGoogle Scholar
Freitag, R. 1972. Female genitalia of the North American species of the Cicindela maritima group (Coleoptera: Cicindelidae). The Canadian Entomologist 104: 12771306.CrossRefGoogle Scholar
Freitag, R. 1979. Reclassification, phylogeny, and zoogeography of the Australian species of Cicindela (Coleoptera: Cicindelidae). Australian Journal of Zoology, Supplementary Series 66: 199.CrossRefGoogle Scholar
Freitag, R., and Tropea, R.. 1969. Twenty-one cicindelid species in thirty-eight days. Cicindela 1: 1423.Google Scholar
Gaumer, G.C. 1977. The Variation and Taxonomy of Cicindela formosa Say (Coleoptera: Cicindelidae). Ph.D. thesis, Texas A & M University, College Station, TX. 253 pp.Google Scholar
Gaumer, G.C., Kurczewski, E.J., and Kurczewski, F.E.. 1970. The tiger beetles of Presque Isle State Park, Pennsylvania. Cicindela 2: 47.Google Scholar
Glaser, J.D. 1976. Cicindelids of Chesapeake Bay revisited. Cicindela 8: 1720.Google Scholar
Goldsmith, W.M. 1916. Field notes on the distribution and life habits of the tiger beetles (Cicindelidae) of Indiana. Proceedings of the Indiana Academy of Science 26: 447455.Google Scholar
Gould, A.A. 1834. On the Cicindelidae of Massachusetts. Boston Journal of Natural History 1: 4152.Google Scholar
Graves, R.C. 1963. The Cicindelidae of Michigan (Coleoptera). American Midland Naturalist 69: 492507.CrossRefGoogle Scholar
Graves, R.C. 1965. The distribution of tiger beetles in Ontario (Coleoptera: Cicindelidae). Proceedings of the Entomological Society of Ontario 95: 6370.Google Scholar
Graves, R.C., and Pearson, D.L.. 1973. The tiger beetles of Arkansas, Louisiana and Mississippi (Coleoptera: Cicindelidae). Transactions of the American Entomological Society 99: 157203.Google Scholar
Hamilton, C.C. 1925. Studies on the morphology, taxonomy, and ecology of holarctic tiger beetles (family Cicindelidae). Proceedings of the Unites States National Museum 65: 187.CrossRefGoogle Scholar
Harris, E.D. 1902. Notes on Cicindelidae in North Carolina. The Canadian Entomologist 34: 217218.CrossRefGoogle Scholar
Harris, T.W. 1852. On some American species of Cicindela. Family Visitor 2(39): 1.Google Scholar
Harris, T.W. 1869. Entomological Correspondence of Thaddeus William Harris, M.D. In Scudder, S.H. (Ed.), Occasional Papers of the Boston Society of Natural History 1: 1375.Google Scholar
Hennig, W. 1966. Phylogenetic Systematics. University of Illinois Press, Urbana, IL. 263 pp.Google Scholar
Hicks, S.D. 1965. The northern limits of several species of Coleoptera with special reference to their occurrence in the Ottawa district, Ontario. The Coleopterists' Bulletin 19: 3742.Google Scholar
Horn, W. 1915. Coleoptera. Adephaga. Family Carabidae. Subfamily Cicindelinae. Wytsman Genera Insectorum, fasc. 82A (1908), 82B (1910), 82C (1915). 486 pp., 23 pls. Brussels.Google Scholar
Horn, W. 1926. Carabidae: Cicindelinae. Coleopterum Catalogus Pars 86: 1345. W. Junk, Berlin.Google Scholar
Horn, W. 1928. Notes and records on the tiger beetles of Minnesota. University of Minnesota Agricultural Experiment Station, Technical Bulletin 56: 913.Google Scholar
Horn, W. 1930. Notes on the races of Omus californicus and a list of the Cicindelidae of America north of Mexico (Coleoptera). Transactions of the American Entomological Society 56: 7386.Google Scholar
Howden, H.F. 1969. Effects of the Pleistocene on North American insects. Annual Review of Entomology 14: 3956.CrossRefGoogle Scholar
Huber, R.L. 1969. Homonymy in the nearctic Cicindela. Cicindela 1: 1922.Google Scholar
Johnson, W.M. 1989. A new subspecies of Cicindela patruela from west-central Wisconsin. Cicindela 21: 2732.Google Scholar
Kaulbars, M.M. 1982. Systematics and Ecology of the sexguttata Species Group, Genus Cicindela (Coleoptera: Cicindelidae). M.Sc. thesis, Lakehead University, Thunder Bay, Ont. 237 pp.Google Scholar
King, P.B. 1959. The Evolution of North America. Princeton University Press, Princeton, NJ. 189 pp.Google Scholar
Knaus, W. 1900. The Cicindelidae of Kansas. The Canadian Entomologist 32: 109116.CrossRefGoogle Scholar
Knaus, W. 1929. A new name for Cicindela violacea Fab. Journal of the Kansas Entomological Society 2: 2324.Google Scholar
Knisley, C.B. 1978. Collecting Cicindela willistoni estancia beneath alkali encrustations. Cicindela 10: 3132.Google Scholar
Knisley, C.B. 1979. Distribution, abundance, and seasonality of tiger beetles (Cicindelidae) in Indiana Dunes region. Proceedings of the Indiana Academy of Science 88: 209217.Google Scholar
Knisley, C.B., and Pearson, D.L.. 1981. The function of turret building behaviour in the larval tiger beetle Cicindela willistoni (Coleoptera: Cicindelidae). Ecological Entomology 6: 401410.CrossRefGoogle Scholar
Larochelle, A. 1972. The Cicindelidae of Quebec. Cicindela 4: 4966.Google Scholar
Larochelle, A. 1978. Techniques for catching tiger beetles. Cicindela 10: 2326.Google Scholar
Larochelle, A. 1980. Cicindelidae of the maritime provinces of Canada. Cicindela 12: 3539.Google Scholar
Lawton, J.K. 1970. A new colour variant of Cicindela patruela. Cicindela 2: 13.Google Scholar
Lawton, J.K. 1974. Unusual activity of Cicindela sexguttata. Cicindela 6: 1718.Google Scholar
Leconte, J.L. 1856. Revision of the Cicindelidae of the United States. Transactions of the American Philosophical Society 11: 2764.Google Scholar
Leconte, J.L. 1861. New species of Coleoptera inhabiting the Pacific district of the United States. Proceedings of the Academy of Natural Sciences, Philadelphia 13: 338389.Google Scholar
Leffler, S.R. 1979. Tiger Beetles of the Pacific Northwest (Coleoptera: Cicindelidae). Ph.D. thesis, University of Washington, College of Forest Resources, Seattle, WA. 790 pp.Google Scholar
Lehmkuhl, D.M. 1980. Temporal and spatial changes in the Canadian insect fauna: Patterns and explanation, the prairies. The Canadian Entomologist 112: 11451160.CrossRefGoogle Scholar
Leng, C.W. 1902. Revision of the Cicindelidae of boreal America. Transactions of the American Entomological Society 28: 93186.Google Scholar
Leng, C.W. 1912. The geographical distribution of Cicindelidae in eastern North America. Journal of the New York Entomological Society 20: 117.Google Scholar
Leng, C.W. 1918. A new race of Cicindela with notes on other races and species. Journal of the New York Entomological Society 26: 138141.Google Scholar
Leng, C.W. 1920. Catalogue of the Coleoptera of America, North of Mexico. J.D.Sherman, Jr. Mount Vernon, NY. 470 pp.Google Scholar
Leng, C.W., and Mutchler, A.J.. 1927. First Supplement (1919–1924) to the Leng Catalogue of Coleoptera of America, North of Mexico, J.D. Sherman, Jr., Mount Vernon, NY. 78 pp.Google Scholar
Leng, C.W., and Mutchler, A.J.. 1933. Second and Third Supplements to the Leng Catalogue of Coleoptera of America, North of Mexico, J.D. Sherman, Jr., Mount Vernon, NY. 112 pp.Google Scholar
Ljungh, S.I. 1799. Nya insector uturegen Sammling beskritne. Kungliga Svenska Vetenskajes Academiens Nya Handlingar 20: 145149.Google Scholar
Macnamara, C. 1922. Tiger beetle larvae. The Canadian Entomologist 54: 241246.CrossRefGoogle Scholar
Mares, A. 1921. A new species and a new variety of Cicindela (Coleoptera). Entomological News, Philadelphia 32: 310.Google Scholar
Matthews, J.V. 1979. Tertiary and Quaternary environments: Historical background for an analysis of the Canadian insect fauna. pp. 31–86 in Danks, H.V. (Ed.), Canada and its Insect Fauna. Memoirs of the Entomological Society of Canada 108: 573 pp.Google Scholar
Morgan, A.V., and Morgan, A.. 1980. Faunal assemblages and distributional shifts of Coleoptera during late pleistocene in Canada and northern United States. The Canadian Entomologist 112: 11051128.CrossRefGoogle Scholar
Nie, H.N., Hull, C.H., Jenkins, J.G., Steinbrenner, K., and Bent, D.H.. 1975. Statistical Package for the Social Sciences, 2nd ed. McGraw-Hill, New York, Toronto. 675 pp.Google Scholar
Palmer, M.K. 1978. Growth rates and survivorship of tiger beetle larvae. Cicindela 10: 4966.Google Scholar
Palmer, M.K. 1979. Rearing tiger beetles in the laboratory. Cicindela 11: 111.Google Scholar
Papp, H. 1952. Morphologische und phylogenetische Untersuchungen an Cicindela-Arten: unter besonderer Berücksichtigung der Ableitung der nearktischen Formen. Oesterreichische Zoologische Zeitschrift 3: 494533.Google Scholar
Rivalier, E. 1954. Démembrement du genre Cicindela Linné II. Faune Américaine. Revue française d'entomologie 21: 249268.Google Scholar
Ross, H.H. 1953. On the origin and composition of the nearctic insect fauna. Evolution 7: 145158.CrossRefGoogle Scholar
Ross, H.H. 1965. Pleistocene events and insects. pp. 583–594 in Wright, H.E., and Frey, D.G. (Eds.), The Quaternary of the United States. Princeton University Press, Princeton, NJ. 922 pp.Google Scholar
Ross, H.H. 1974. Biological Systematics. Addison-Wesley Publishing Co., Inc., Reading, MA; Menlo Park, CA; London; Don Mills, Ont. 345 pp.Google Scholar
Say, T. 1817. Descriptions of several new species of North American insects. Journal of the Academy of Natural Science, Philadelphia 1: 1923.Google Scholar
Say, T. 1818. Monograph of the North American Cicindela. Transactions of the American Philosophical Society 1: 401426.CrossRefGoogle Scholar
Schaupp, F.G. 1884. Synoptic tables of Cicindelidae. Bulletin of the Brooklyn Entomological Society 6: 121124.Google Scholar
Schincariol, L.A., and Freitag, R.. 1991. Biological character analysis, classification, and history of the North American Cicindela splendida Hentz group taxa (Coleoptera: Cicindelidae). The Canadian Entomologist 123: 13271353.CrossRefGoogle Scholar
Scudder, G.G.E. 1979. Present patterns in the fauna and flora of Canada. pp. 87–179 in Danks, H.V. (Ed.), Canada and its Insect Fauna. Memoirs of the Entomological Society of Canada 108: 573 pp.Google Scholar
Shelford, V.E. 1907. Preliminary notes on the distribution of tiger beetles and its relation to plant succession. Biological Bulletin 14: 914.CrossRefGoogle Scholar
Shelford, V.E. 1908. Life histories and larval habits of the tiger beetles (Cicindelidae). Journal of the Linnean Society (Zoology) 30: 157184.Google Scholar
Shelford, V.E. 1911. Physiological animal geography. Journal of Morphology (and Physiology) 22: 551618.CrossRefGoogle Scholar
Shelford, V.E. 1917. Color and color-pattern mechanism of tiger beetles. Illinois Biological Monographs 3: 395532.Google Scholar
Smyth, E.G. 1905. Notes on collecting Cicindelidae. Kansas Academy of Science 19: 425432.CrossRefGoogle Scholar
Soil Survey Staff. 1960. Soil Classification: A Comprehensive System, 7th approximation. U.S. Department of Agriculture, Washington, DC. 265 pp.Google Scholar
Soil Survey Staff. 1967. Supplement to Soil Classification System. U.S. Department of Agriculture, Washington, DC. 207 pp.Google Scholar
Tanner, V.M. 1927. A preliminary study of the genitalia of female Coleoptera. Transactions of the Entomological Society of America 53: 550.Google Scholar
Townsend, C.H.T. 1884. The proportion of the sexes in Cicindela vulgaris Say and other notes on the species. The Canadian Entomologist 16: 227231.CrossRefGoogle Scholar
Wallis, J.B. 1961. The Cicindelidae of Canada. University of Toronto Press, Toronto, Ont. 74 pp.CrossRefGoogle Scholar
Ward, R.D. 1971. On spring collecting in Louisiana, Texas, and Arkansas with notes on Cicindela sexguttata tridens Casey. Cicindela 3: 6977.Google Scholar
Wickham, H.F. 1902. The habits of American Cicindelidae. Proceedings of the Davenport Academy of Natural Science 6: 206228.Google Scholar
Wickham, H.F. 1911. A list of the Coleoptera of Iowa. Bulletin from the Laboratories of Natural History of the State University of Iowa 6: 115.Google Scholar
Wiedemann, C.R.W. 1821. Neue exotische Käfer beschrieben von C.R.W. Wiedemann u. E.F. Germar. Magazin d'Entomologique 4: 107183.Google Scholar
Wiedemann, C.R.W. 1823. Zeihundert neue Käfer von Java, Bengalen und dem Vorgebirge der guten Hoffnung. Zoological Magazine 2(1): 1135.Google Scholar
Wiley, E.O. 1981. Phylogenetics: The Theory and Practice of Phylogenetic Systematics. John Wiley and Sons, New York, NY. 439 pp.Google Scholar
Willis, H.L. 1967. Bionomics and zoogeography of tiger beetles of saline habitats in the central United States. University of Kansas Science Bulletin 47: 145313.Google Scholar
Willis, H.L. 1968. Artificial key to the species of Cicindela of North America north of Mexico. Journal of the Kansas Entomological Society 41: 303317.Google Scholar
Willis, H.L. 1970. The Cicindelidae of Kansas. Cicindela 2: 127.Google Scholar
Zikan, J.J. 1929. Zur biologie der cicindeliden Brasiliens. Zoologischer Anzieger 82: 269414.Google Scholar
Zoltai, S.C. 1961. Glacial history of part of northwestern Ontario. Proceedings of the Geological Association of Canada 13: 6183.Google Scholar
Zoltai, S.C. 1968. Glacial history of the upper Great Lakes. Proceedings of the Entomological Society of Ontario 99: 1520.Google Scholar