Abstract
All members of Sophiodela Nakane 1955, hitherto a subgenus of Cicindela Linnaeus, 1758, are reviewed based on the structure of the everted internal sac of the male genitalia. In addition, many species of other genera and subgenera such as the genera Cosmodela Rivalier, 1961, Calochroa Hope, 1838, and Cicindela and its subgenus Pancallia Rivalier, 1961, are also reviewed regarding their reproductive structures. Their relationships are discussed using a comparison between morphological and molecular analyses. Through this study, the taxonomic rank of Sophiodela is raised from a subgenus to a genus as it is demonstrated to be a distinct monophyletic group by both the morphological and molecular analyses. Amongst the species of Sophiodela, S. cyanea Fabricius, 1787 is changed to the subgenus Pancallia. We conclude that the everted internal sac is a useful tool for grouping the genera and subgenera and can be used as a taxonomic character.
References
Acciavatti, R.E. & Pearson, D.L. (1989) The tiger beetle Genus Cicindela (Coleoptera, Insecta) from the indian subcontinent.Annals of Carnegie Museum, 58, 77–353.
Ádám, L. & Merkl, O. (1986) Adephaga of the Kiskunság National Park, I.: Carabidae (Coleoptera). In: Mahunka, S. (Ed.), The Fauna of the Kiskunság National Park, I. Akadémiai Kiadó, Budapest, pp. 119–142.
Bates, H.W. (1873) On the geodephagous Coleoptera of Japan. Transactions of the Entomological Society of London, 1873, 219–322.
https://doi.org/10.1111/j.1365-2311.1873.tb00643.x
Bates, H.W. (1883) Supplement to the geodephagous Coleoptera of Japan, chiefly from the collection of Mr. George LEWIS, made during his second visit, from February, 1880, to September, 1881. Transactions of the Entomological Society of London, 1883, 205–290.
https://doi.org/10.1111/j.1365-2311.1883.tb02947.x
Beuthin, H. (1905) Neue Varietäten paläarktischer Cicindelen. Societas Entomologica, 19 (24), 185–186.
Cassola, F. & Brouerius van Nidek, C.M.C (1984) Checklist of Cicindela (s. auct.) of the Palaearctic region (Coleoptera: Cicindelidae). Cicindela, 16, 7–17.
Cassola, F. (1999) Studies on tiger beetles. CVII. The cicindelid fauna of anatolia: faunistics and biogeography (Coleoptera, Cicindelidae). Biogeographia, Biogeografia de’ll Anatolia, 20, 229–276.
https://doi.org/10.21426/B6110089
Chaudoir, M. de (1863) Enumération des Cicindélètes et des Carabiques recueillis dans la Russe méridionale, dans la Finlande septentrionale et dans la Sibérie orientale par MM. Alexandre et Artur de Nordmann. Bulletin de la Société Imperial des Naturalistes de Moscou, 36 (1), 201–232.
Chaudoir, M. de (1865) Catalogue de la collection de Cicindélètes de M. le Baron de Chaudoir. Nys, Bruxelles, 64 pp.
https://doi.org/10.5962/bhl.title.134840
DeGeer, C. (1774) Mémoires pour servir à l’histoire des insectes and quadrième by quatrième. Tome quadrième. Hesselberg, Stockholm, xii + 457 pp.
Dejean P.F.M.A. (1826) Species général des Coléoptères de la collection de M. le Comte Dejean. Tome Second. Crevot, Paris, 195–276 pp.
https://doi.org/10.5962/bhl.title.2086
Fabricius, J.C. (1775) Systema Entomologiae, sistens insectorum classes, ordines, genera, species, adiectis synonymis, locis, descriptionibus, observationibus. Kortii, Flensburgi et Lipsiae, xxxii + 832 pp.
https://doi.org/10.5962/bhl.title.36510
Fabricius, J.C. (1781) Species Insectorum exhibentes eorum differentias specificas, synonyma, auctorum, loca natalia, metamorphosin adiectis observationibus, descriptionibus. Tomus I. Hamburgi et Kilonii: Carol. Ernest. Bohnii, 552 pp.
https://doi.org/10.5962/bhl.title.36509
Fabricius, J.C. (1787) Mantissa insectorum, sistens eorum species nuper detectas adiectis characteribus genericis, differentiis specificis, emendationibus, observationibus. Tom. I. C.G. Proft, Hafniae, 185–187.
https://doi.org/10.5962/bhl.title.36471
Fabricius J.C. (1801) Systema Eleutheratorum secundum ordines, genera, species; adiectis synonymis, locis, observationibus, descriptionibus. Tomus I & II. Bibliopolii Academici Novi, Kiliae, 226 pp.
Faldermann, F. (1835) Coleopterorum ab illustrissimo Bungio in China boreali, Mongolia, et montibus Altaicis collectorum, nec non ab ill. Turczaninoffio et Stchukino e provincia Irkutzk missorum illustrationes. Mémoires présentés à l’Académie Impériale des Sciences de Saint-Pétersbourg, 2, 337–464., 5 pls.
Fleutiaux, E. (1892) Catalogue systématique des Cicindelidae. Imprimerie H. Vaillant-Carmanne, Liège, 186 pp.
https://doi.org/10.5962/bhl.title.48634
Fleutiaux, E. (1893) Remarques sur quelques Cicindelidae et descriptions d’espèces nouvelles. Annales de la Société Entomologique de France, 62, 483–502.
Fleutiaux, E. (1895) Description d’une nouvelle espèce de Cicindela du groupe des Calochroa vrais et une note sur deux espèces du genre Peridexia. Bulletin de la Société Entomologique de France, 1894, cclxxxi–cclxxxii.
Fleutiaux, E. (1898) Description d’une espèce et d’une variété nouvelles de Cicindela (Col.). Bulletin de la Société Entomologique de France, 1898, 146–147.
Fleutiaux, E. (1919) Sur quelques Cicindelidae du Laos (Col.). Bulletin de la Société Entomologique de France, 1919, 126–128.
Fowler, W.W. (1912) The fauna of British India including Ceylon and Burma. Coleoptera. General introduction and Cicindelidae and Paussidae, Taylor and Francis, London, 529 pp.
https://doi.org/10.5962/bhl.title.83566
Fukuda, Y., Yamasako, J., Ogawa, R. & Sakai, M. (2015) Review of the Japanese species of the genus Cicindela (Sophiodela) (Coleoptera, Carabidae, Cicindelinae) based on the characters of internal sac. Elytra, New Series, 5 (2), 269–280.
Gotô, M. & Iga, M. (1950) On a new variety of Cicindela chinensis DeGeer from Japan (Coleoptera, Cicindelidae). Entomological Review of Japan, 5, 14.
Guérin-Méneville F.E. (1847) Décade entomologique. Revue Zoologique, 10, 2–11.
Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Vinh, L.S. (2018) UFBoot2: improving the ultrafast bootstrap approximation. Molecular Biology and Evolution, 35, 518–522.
https://doi.org/10.1093/molbev/msx281
Hope, F.W. (1831) Synopsis of the new species of Nepal insects in the collection of Major General Hardwicke. Zoological Miscellany, 1, 21–32.
Hope, F.W. (1838) The coleopterist’s manual, part the second, containing predaceous land and water beetles of Linnaeus and Fabricius. Bohn, London, 168 pp.
Horn, W. & Roeschke, H. (1891) Monographie der paläarktischen Cicindelen. Analytisch bearbeitet mit besonderer Berücksichtigung der Variationsfähigkeit und geographischen Verbreitung. Berlin, 197 pp.
https://doi.org/10.5962/bhl.title.9205
Horn, W. (1894) Beitrag zur Cicindeliden-Fauna von Vorder-Indien. Deutsche Entomologische Zeitschrift, 2, 169–175.
https://doi.org/10.1002/mmnd.48018940507
Horn, W. (1900) De novis Cicindelidarum speciebus. Deutsche entomologische Zeitschrift, 1, 193–212.
https://doi.org/10.1002/mmnd.48019000126
Horn, W. (1908) Coleoptera Adephaga (Family Carabidae, Subfamily Cicindelidae). in: Wytsman, P. (Ed.), Genera insectorum. 82 A, pp. 1–484.
Horn, W. (1910) Coleoptera Adephaga, Fam. Carabidae, Subfam. Cicindelinae. In: Wytsman, P. (Ed.), Genera Insectorum, 82, pp. 105–208, pls. 1–15.
Horn, W. (1913) Matériaux pour servir à l’étude de la faune entomologique de l’Indo-Chine. Annales de la Société entomologique de Belgique, 57, 362–366.
https://doi.org/10.5962/bhl.part.4597
Horn, W. (1915) Coleoptera Adephaga, Fam. Carabidae, Subfam. Cicindelinae. In: Wytsman, P. (Ed.), Genera Insectorum, 82, pp. 209–487, pls. 16–23.
Horn, W. (1920) Cicindelinen-Studien aus dem schwedischen Reichsmuseum nebst einigen Bemerkungen neuer Arten etc. Arkiv för Zoologi, 13 (11), 1-21.
Horn, W. (1926) Carabidae, Cicindelinae (Pars 86). In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus. Junk, Berlin, pp. 1–345.
Iga, M. (1955) [new taxon] P. 15. In: Nakane, T. (Ed.), Coloured Illustrations of the Insects of Japan. Coleoptera. Osaka, Hoikusha, 274 pp., 67 pl.
Lewis, G. (1891) Note on a new Cicindela from north Japan. The Entomologist’s Monthly Magazine, 27, 20.
Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Tomus I. Laurentii Salvii, Holmiae, 828 pp.
https://doi.org/10.5962/bhl.title.542
López-López, A., Abdul, A. & Galián, J. (2015) Molecular phylogeny and divergence time estimation of Cosmodela (Coleoptera: Carabidae: Cicindelinae) tiger beetle species from Southeast Asia. Zoologica Scripta, 44 (4), 437–445.
https://doi.org/10.1111/zsc.12113
Mandl, K. (1975) Ergebnisse der Bhutan-Expedition 1972 des Naturhistorischen Museums in Basel. Coleoptera: Fam. Cicindelidae. Entomologica Basiliensia, 1, 135–143.
Matalin, A.V. (1998) The tiger-beetles of “hybrida”-species group (Coleoptera, Carabidae, Cicindelinae). III. A taxonomic review of the Iberian Cicindela lagunensis Gautier, 1872 complex. Graellsia, 54, 75–96.
https://doi.org/10.3989/graellsia.1998.v54.i0.346
Matalin, A.V. (1999) The tiger-beetles of the “hybrida” species-group. II. A taxonomic review of subspecies in Cicindela sahlbergii Fischer von Waldheim, 1824 (Coleoptera, Carabidae, Cicindelinae). In: Zamotajlov, A. & Sciaky, R. (Eds.), Advances in Carabidology. MUISO publishers, Krasnodar, pp. 13–55.
Matalin, A.V. & Chikatunov, V.I. (2016) The tiger beetles (Coleoptera, Carabidae, Cicindelinae) of Israel and adjacent lands. ZooKeys, 578, 115–160.
https://doi.org/10.3897/zookeys.578.7383
Miwa, Y. (1935) Coleoptera from Amami-Islands in Loo-Choo Archipelago. Transactions of the Kansai Entomological Society of Osaka, 6, 11–30.
Moravec, J. (2010) Tiger beetles of the Madagascan Region (Madagascar, Seychelles, Comoros, Mascarenes, and other islands. Taxonomic revision of the 17 genera occurring in the region (Coleoptera: Cicindelidae). Biosférická rezervace Dolní Morava, o.p.s., Lednice na Moravě, 429 pp.
Morawitz, A. (1862) Vorlaufige Diagnosen neuer Carabiciden aus Hakodate. Bulletin de l’Académie imperiale des Sciences de St.-Petersbourg, 4, 237–247.
Motschulsky, V. de (1857) VI. Nouveautés. Etudes Entomologiques, 6, 108-112.
Motschulsky, V. de (1860) Coléoptères de la Sibérie orientale et notamment en particulier des rives de l’Amour. In: Schrenck, L. von (Ed.), Reisen und Forschungen im Amur-Lande in den Jahren 1854-1856 / im Auftrage der Kaiserl. Akademie der Wissenschaften zu St. Petersburg ausgeführt und in Verbindung mit mehreren Gelehrten herausgegeben von Leopold v. Schrenck. Band II. Zweite lieferung. Coleopteren. St. Petersburg: Eggers & Comp., pp. 77–257, pls. VI–XI, 1 map.
Motschulsky, V. de (1862) Entomologie spéciale. Remarques sur la collection d’insectes de V. Motschulsky. Coléoptères. Etudes Entomologiques, 11, 15–55.
Nakane, T. (1955) New or little-known Coleoptera from Japan and its adjacent regions, XII. The Scientific Report of the Saikyo University, 2 (1), 24–40.
Nakane, T. (1957) New or little-known Coleoptera from Japan and its adjacent regions, XIV. The Scientific Report of the Saikyo University, 2 (4), 41–43.
Nakane, T. (1963) Cicindelidae. In: Nakane, T., Ohbayashi, K., Nomura, S. & Kurosawa, Y. (Eds.), Iconographia Insectorum Japonicorum Colore naturali edita, Volumen II. Hokuryûkan, Tokyo, pp. 2–4. [In Japanese]
Nakane, T. (1976) Check-list of Coleoptera of Japan, Cicindelidae. The Coleopterists’ Association of Japan, 3, 1–7.
Naviaux, R. (1986) Coleoptera, Cicindelidae. Premier complément aux Cicindèles du Sri Lanka. Revue Scientifique Du Bourbonnais, 58–69.
Naviaux, R. (1991) Les Cicindèles de Thaïlande, étude faunistique (Coleoptera Cicindelidae). Bulletin Mensuel de la Société Linnéenne de Lyon, 60 (7), 209–288.
https://doi.org/10.3406/linly.1991.10944
Nguyen, L.T., Schmidt, H.A. & Minh, B.Q. (2015) IQ-TREE: a fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular biology and evolution, 32 (1), 268–274.
https://doi.org/10.1093/molbev/msu300
Nylander, J.A.A. (2004) MrModeltest v2. Program distributed by the author. Evolutionary Biology Centre,Uppsala University.
Oike, K. (1939) Cicindela gemmata shikokuana subsp. nov. Transactions of the Kansai Entomological Society, 8, 15–16.
Osozawa, S., Fukuda, H., Kwon, H.-Y. & Wakabayashi, J. (2016) Quaternary vicariance of tiger beetle, Cicindela chinensis, in Ryukyu, Japan, Taiwan and Korea–China. Entomological Research, 46 (2), 122–127.
Parry, F.J.S. (1844) A decade of new Coleoptera from Assam. The Annals and Magazine of Natural History, 14, 454.
Parry, F.J.S. (1848) Description and notes upon some new and rare Coleoptera. The Transactions of the Entomological Society of London, 4, 80–84.
Pons, J., Barraclough, T., Theodorides, K., Cardoso, A. & Vogler, A. (2004) Using exon and intron sequences of the gene Mp20 to resolve basal relationships in Cicindela (Coleoptera:Cicindelidae). Systematic Biology, 53 (4), 554–570.
https://doi.org/10.1080/10635150490472940
Rivalier, E. (1950) Démembrement du genre Cicindela Linné (Travail préliminaire limité a la faune paléarctique). Revue française d’Entomologie, 17, 249–268.
Rivalier, E. (1958) Démembrement du genre Cicindela Linné, III. Faune III. Africano-Malgache. Revue française d’Entomologie, 24, 312–341.
Rivalier, E. (1961) Démembrement du genre Cicindela Linné, IV. Faune indomalaise. Revue française d’Entomologie, 28, 121 –149.
Sasakawa, K. (2008) Geographical variation of the flightless tiger beetle Cylindera ovipennis (Bates, 1883) (Coleoptera, Carabidae, Cicindelinae): an approach using male genital morphology. Biogeography, 10, 103–105.
Sawada, H. & Wiesner, J. (2002) Further new records of tiger beetles collected in Laos (Coleoptera: Cicindelidae). Entomological Review of Japan, 57, 65–99.
Schaum, H.R., (1863) Contributions to the knowledge of the Cicindelidae of tropical Asia, containing descriptions of new species, a list of those hitherto described, and synonymical notes. The Journal of Entomology. Descriptive and Geographical, 2, 57–73.
Schilder, F.A., (1953) Studien zur Evolution von Cicindela. Wissenschaftliche Zeitschrift der Martin-Luther-Universität Halle-Wittenberg, 3, 539–576.
Silvestro, D. & Michalak, I. (2012) raxmlGUI: a graphical front-end for RAxML. Organisms Diversity & Evolution, 12 (4), 335–337.
https://doi.org/10.1007/s13127-011-0056-0
Sota, T., Liang, H., Enokido, Y. & Hori, M. (2011) Phylogeny and divergence time of island tiger beetles of the genus Cylindera (Coleoptera: Cicindelidae) in East Asia. Biological Journal of the Linnean Society, 102, 715–727.
https://doi.org/10.1111/j.1095-8312.2011.01617.x
Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts.
Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: molecular evolutionary genetics analysis version 6.0. Molecular biology and evolution, 30 (12), 2725–2729.
https://doi.org/10.1093/molbev/mst197
Thomson, J. (1857) Voyage entomologique à Londres. Archives Entomologiques, 1, 321–325.
Thunberg, C.P. (1781) Dissertatio entomologica novas insectorum species sistens. Edman, Upsaliae, 28 pp.
Trifinopoulos, J., Nguyen, L.T., von Haeseler, A. & Minh, B.Q. (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic acids research, 44 (W1), W232–W235.
https://doi.org/10.1093/nar/gkw256
Tsuji, K, Hori, M., Phyu, M.H., Liang, H. & Sota, T. (2016) Colorful patterns indicate common ancestry in diverged tiger beetle taxa: Molecular phylogeny, biogeography, and evolution of elytral coloration of the genus Cicindela subgenus Sophiodela and its allies. Molecular Phylogenetics and Evolution, 95, 1–10.
https://doi.org/10.1016/j.ympev.2015.11.006
Vigors, M. (1825) Mr. Vigors on some new subjects of zoology. The Zoological Journal, 1, 410–415.
Werner, K. (1992) Cicindelidae Regionis Palaearcticae. Cicindelini 2: Cosmodela – Platydela – Lophyra – Habrodera – Chaetodera – Neolaphyra – Cephalota – Cassolai – Homodela – Cylindera – Eugrapha – Myriochile – Salpingophora – Hypaeth – Abroscelis – Callytron. Die Käfer der Welt, Vol.15, Sciences Nat, Venette: 94 pp., pls. 31–57.
Wiedemann, C.R.W. (1819) Neue Kafer aus Bengalen und Java. Zoologische Magazin, 1 (3), 157–183.
Wiesner, J. (1992) Checklist of the Tiger Beetles of the World. Verlag Erna Bauer, Keltern, 364 pp.
Wiesner, J. (2004) Three new tiger beetle species from Myanmar and notes on further species (Coleoptera: Cicindelidae). 86. contribution towards the knowledge of Cicindelidae. Entomologische Zeitschrift, 114 (4), 175–181.
Wood, J.G. (1874) Insects abroad, being a popular account of foreign insects, their structure, habits, and transformations. Longmans, Green, and Co., London, xiii+648 pp.