Skip to main content Skip to main navigation menu Skip to site footer
Type: Monograph
Published: 2018-09-12
Page range: 1–104
Abstract views: 283
PDF downloaded: 19

Phylogeny and taxonomic revision of Deronectina Galewski, 1994 (Coleoptera: Dytiscidae: Hydroporinae: Hydroporini)

Räuschstraße 73, 13509 Berlin, Germany.
Institute of Evolutionary Biology (CSIC-Universitat Pompeu Fabra), Passeig Maritim de la Barceloneta 37, 08003 Barcelona, Spain.
Coleoptera Dytiscidae Hydroporinae Deronectina phylogeny new classification new genera new combinations


The subtribe Deronectina Galewski, 1994 (Dytiscidae, Hydroporinae, Hydroporini) is distributed in the Nearctic, in the north of the Neotropical region, and in the Palaearctic and Afrotropical regions. It is currently composed of 194 species and 13 subspecies in eight genera: Amurodytes Fery & Petrov, 2013, Boreonectes Angus, 2010, Deronectes Sharp, 1882, Nebrioporus Régimbart, 1906, Oreodytes Seidlitz, 1887, Scarodytes Gozis, 1914, Stictotarsus Zimmermann, 1919, and Trichonectes Guignot, 1941. We present a morphological and a molecular phylogeny of the species of the subtribe, and a revision of their taxonomy to accommodate our phylogenetic results. The morphological phylogeny is based on the study of 54 characters of the adults of 189 species and 2 subspecies, of which 114 species and the 2 subspecies were coded in the morphological matrix. For the molecular phylogeny we investigated 115 species and 11 subspecies, using a combination of fragments of four mitochondrial (COI, 16S rRNA, tRNA-Leu and NAD1) and two nuclear genes (18S rRNA and H3), analysed with maximum likelihood and Bayesian methods. For both datasets we included the type species of all genus-group taxa. The morphological, molecular and combined phylogenies mostly agree with the current classification of the group, but in some cases our results are in contradiction with established genera. Most remarkable are the polyphyly of Stictotarsus and Nebrioporus, the low support for the monophyly and internal phylogeny of Oreodytes, and the low support for the monophyly of Deronectina with molecular data. Thus, we introduce some taxonomic changes in the current classification to accommodate the generic concepts to our phylogenetic results. Nine new genera are established: Clarkhydrus n. gen. (type species Hydroporus roffii Clark, 1862), Hornectes n. gen. (type species Hydroporus quadrimaculatus Horn, 1883), Iberonectes n. gen. (type species Deronectes bertrandi Legros, 1956), Larsonectes n. gen. (type species Potamonectes minipi Larson, 1991), Leconectes n. gen. (type species Hydroporus striatellus LeConte, 1852), Mystonectes n. gen. (type species Deronectes neomexicanus Zimmerman & Smith, 1975), Nectoboreus n. gen. (type species Hydroporus aequinoctialis Clark, 1862), Nectomimus n. gen. (type species Oreodytes okulovi Lafer, 1988), and Zaitzevhydrus n. gen. (type species Hydroporus formaster Zaitzev, 1908). Three genera are reinstated as valid: Deuteronectes Guignot, 1945 (stat. rest.) (type species Hydroporus picturatus Horn, 1883), Nectoporus Guignot, 1950 (stat. rest.) (type species Hydroporus abbreviatus Fall, 1923), and Neonectes J. Balfour-Browne, 1940 (stat. rest.) (type species Hydroporus natrix Sharp, 1884). Thirty-six new combinations for species and subspecies thus far treated in the genera Boreonectes, Nebrioporus, Oreodytes and Stictotarsus result from the new classification: Clarkhydrus corvinus (Sharp, 1887) n. comb., C. decemsignatus (Clark, 1862) n. comb., C. deceptus (Fall, 1932) n. comb., C. eximius (Motschulsky, 1859) n. comb., C. falli (Nilsson, 2001) n. comb., C. interjectus (Sharp, 1882) n. comb., C. minax (Zimmerman, 1982) n. comb., C. opaculus (Sharp, 1882) n. comb., C. roffii (Clark, 1862) n. comb., C. spectabilis (Zimmerman, 1982) n. comb., Deuteronectes angustior (Hatch, 1928) n. comb., Hornectes quadrimaculatus (Horn, 1883) n. comb., Iberonectes bertrandi (Legros, 1956) n. comb., Larsonectes minipi (Larson, 1991) n. comb., Leconectes striatellus (LeConte, 1852) n. comb., Mystonectes coelamboides (Fall, 1923) n. comb., M. grammicus (Sharp, 1887) n. comb., M. neomexicanus (Zimmerman & Smith, 1975) n. comb., M. panaminti (Fall, 1923) n. comb., M. titulus (Leech, 1945) n. comb., Nectoboreus aequinoctialis (Clark, 1862) n. comb., N. dolerosus (Leech, 1945) n. comb., N. funereus (Crotch, 1873) n. comb., Nectomimus okulovi (Lafer, 1988) n. comb., Nectoporus angelinii (Fery, 2015) n. comb., N. congruus (LeConte, 1878) n. comb., N. crassulus (Fall, 1923) n. comb., N. obesus obesus (LeConte, 1866) n. comb., N. obesus cordillerensis (Larson, 1990) n. comb., N. rhyacophilus (Zimmerman, 1985) n. comb., N. sanmarkii sanmarkii (C.R. Sahlberg, 1826) n. comb., N. sanmarkii alienus (Sharp, 1873) n. comb., N. sierrae (Zimmerman, 1985) n. comb., N. subrotundus (Fall, 1923) n. comb., Zaitzevhydrus formaster formaster (Zaitzev, 1908) n. comb., and Z. formaster ulanulana (C.-K. Yang, 1996) n. comb.



  1. Abellán, P., Sánchez-Fernández, D., Picazo, F., Millán, A., Lobo, J.M. & Ribera, I. (2013) Preserving the evolutionary history of freshwater biota in Iberian National Parks. Biological Conservation, 62, 116–126.

    Alarie, Y. (1993) A systematic review of the North American species of the Oreodytes alaskanus clade (Coleoptera: Dytiscidae: Hydroporinae). The Canadian Entomologist, 125, 847–867.

    Alarie, Y. (1997) Taxonomic revision and phylogenetic analysis of the genus Oreodytes Seidlitz (Coleoptera: Dytiscidae: Hydroporinae) based on larval morphology. The Canadian Entomologist, 129, 399–503.

    Alarie, Y. & Nilsson, A.N. (1996) The larvae of Neonectes J. Balfour-Browne, (Coleoptera: Dytiscidae: Hydroporinae) with a description of N. natrix (Sharp) and a discussion of its phylogenetic relationships with members of the tribe Hydroporini. The Coleopterists Bulletin, 50, 107–121.

    Ali, H.A. (1978) A list of some aquatic beetles of Iraq (Coleoptera: Dytiscidae). Bulletin of the Natural History Research Centre, 7 (2), 11–14.

    Ali, H.A. & Abdul-Karim, R.M. (1989) A key to the Dytiscidae (Insecta, Coleoptera) of Iraq. Iraq National History Museum, 35, 1–21.

    Angus, R.B. (2010) Boreonectes gen. n., a new genus for the Stictotarsus griseostriatus (De Geer) group of sibling species (Coleoptera: Dytiscidae), with additional karyosystematic data on the group. Comparative Cytogenetics, 4 (2), 123–131.

    Angus, R.B., Ribera, I. & Jia, F. (2017) Further studies on Boreonectes Angus, 2010, with a molecular phylogeny of the Palaearctic species of the genus. Comparative Cytogenetics, 11(2), 189–201.

    Angus, R.B. & Tatton, A.G. (2011) A karyosystematic analysis of some water beetles related to Deronectes Sharp (Coleoptera, Dytiscidae). Comparative Cytogenetics, 5 (3), 173–190. v5i3.1185

    Aubé, C. (1838) Hydrocanthares et gyriniens. In: Dejean, P.F.M.A., Species général des coléoptères de la collection de M. le Comte Dejean. Vol. 6. Méquignon Père et Fils, Paris, xvi + 804 pp. [published on 29 September 1838]

    Aykut, M. & Fery, H. (2017) Scarodytes costatus nov.sp. from the Bingöl Province in Turkey, the first species of the genus with costate elytra (Insecta: Coleoptera: Dytiscidae: Deronectina). Linzer Biologische Beiträge, 49 (1), 395–414.

    Aykut, M., Taşar, G.E. & Fery, H. (2018) Deronectes taron sp. n. from the eastern Anatolian region of Turkey (Coleoptera, Dytiscidae, Hydroporinae). Zootaxa, 4422, 403–410.

    Balfour-Browne, F. (1934a) Systematic notes upon British aquatic Coleoptera. Part III. (Deronectes). The Entomologist's Monthly Magazine, 70, 175–180.

    Balfour-Browne, F. (1934b) Systematic notes upon British aquatic Coleoptera. Part V(1). Hydroporus (Suphrodytes, Scarodytes and Hydroporus s. str.) and Laccornis. The Entomologist's Monthly Magazine, 70, 224–230.

    Balfour-Browne, F. (1934c) The proventriculus in the Dytiscidae (Col.) as a taxonomic character. Stylops, 3 (11), 241–244.

    Balfour-Browne, F. (1936) Systematic notes upon British aquatic Coleoptera. Part XI. Haliplidae. (Peltodytes, Haliplus and Brychius). The Entomologist's Monthly Magazine, 72, 68–77.

    Balfour-Browne, F. (1940) British water beetles. Vol. I. Ray Society, London, xix + 375 pp., 5 pls.

    Balfour-Browne, F. (1944) The proventriculus of the Coleoptera (Adephaga) and other insects–a study in evolution. Journal of the Royal Microscopical Society, 64, 68–117.

    Balfour-Browne, J. (1944) Remarks on the Deronectes-complex (Col., Dytiscidae). The Entomologist, 77, 186–189.

    Bergsten, J., Brilmyer, G., Crampton-Platt, A. & Nilsson, A.N. (2012) Sympatry and colour variation disguised well-differentiated sister species: Suphrodytes revised with integrative taxonomy including 5 kbp of housekeeping genes (Coleoptera: Dytiscidae). DNA Barcodes, 1, 1–18.

    Bergsten, J., Nilsson, A.N. & Ronquist, F. (2013) Bayesian tests of topology hypotheses with an example from diving beetles. Systematic Biology, 62 (5), 660–673.

    Bertrand, H. & Legros, C. (1971) Notes complémentaires sur Deronectes bertrandi Legros (Col. Dytiscidae). Bulletin de la Société Entomologique de France, 76, 193–196.

    Bilton, D.T. (2012) Stictonectes rebeccae sp. n. from the Iberian Peninsula, with notes on its phylogenetic position (Coleoptera, Dytiscidae). Zootaxa, 3188 (2011), 42–54.

    Colgan, D.J., McLauchlan, A., Wilson, G.D.F., Livingston, S.P., Edgecombe, G.D., Macaranas, J., Cassis, G. & Gray, M.R. (1998) Histone H3 and U2 snRNA DNA sequences and arthropod molecular evolution. Australian Journal of Zoology, 46, 419–437.

    Clark, H. (1862) On the Mexican species of Hydropori. The Annals and Magazine of Natural History, 10, 173–184.

    De Geer, C. (1774) Mémoires pour servir à l'histoire des insectes. Vol. IV. Pierre Hesselberg, Stockholm, xii + 456 pp.

    Désamoré, A., Laenen B., Miller, K.B. & Bergsten, J. (2018) Early burst in body size evolution is uncoupled from species diversification in diving beetles (Dytiscidae). Molecular Ecology, 27, 979–993.

    Drummond, A.J., Suchard, M.A., Xie, D. & Rambaut, A. (2012) Bayesian phylogenetics with BEAUti and the BEAST 1.7. Molecular Biology and Evolution, 29, 1969–1973.

    Dutton, L.A. & Angus, R.B. (2007) A karyosystematic investigation of a group of sibling species related to Stictotarsus griseostriatus (De Geer) (Coleoptera: Dytiscidae). Comparative Cytogenetics, 1 (1), 3 –16.

    Fabricius, J.C. (1777) Genera Insectorum eorumque characteres naturales secundum numerum, figuram, situm et proportionem omnium partium oris adiecta mantissa speciorum nuper detectarum. M.F. Bartschii, Chilonii, xvi + 310 pp.

    Fabricius, J.C. (1787) Mantissa Insectorum sistens species nuper detectas adiectis characteribus genericis, differentiis specificis, emendationibus, observationibus. 2 Vols. C.G. Proft, Hafniae, xx + 348 + 382 pp.

    Fabricius, J.C. (1792–1794) Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species adjectis synonimis, locis, observationibus, descriptionibus. 4 vols. C.G. Proft, Hafniae, xx + 330 pp. + 538 pp., viii + 519 pp., vi + 487 pp. + 349 pp., vi + 472 pp. [Vol. 1. Part 1: xx + 330 pp. in 1792, Part 2: 538 pp. in 1793, Vol. 2: viii + 519 pp. in 1793, Vol. 3. Part 1: vi + 487 pp. in 1793, Part 2: 349 pp. in 1794, Vol. 4: vi + 472 pp. in 1794]

    Falkenström, G.A. (1939) Beitrag zur Revision einiger Dytisciden-Gattungen, vor allem Deronectes Sharp und Oreodytes Seidlitz. Entomologisk Tidskrift, 60, 69–101.

    Fall, H.C. (1923) A revision of the North American species of Hydroporus and Agaporus. Privately printed, 129 pp. [city location not identified]

    Fall, H.C. (1926) A list of the Coleoptera taken in Alaska and adjacent parts of the Yukon Territory in the summer of 1924. The Pan-Pacific Entomologist, 2 (3), 127–154.

    Fall, H.C. (1932) Random notes and descriptions (Coleoptera). Bulletin of the Brooklyn Entomological Society, 27, 145–148.

    Fery, H. (1992) Revision der saginatus-Gruppe der Gattung Coelambus Thomson (Coleoptera: Dytiscidae). Linzer Biologische Beiträge, 24 (1), 339–358.

    Fery, H. (1995) Ergänzungen zur saginatus-Gruppe sowie Bemerkungen zu weiteren Arten der Gattung Coelambus Thomson (Coleoptera, Dytiscidae). Linzer Biologische Beiträge, 27 (2), 1045–1061.

    Fery, H. (1999) Revision of a part of the memnonius-group of Hydroporus Clairville, 1806 (Insecta: Coleoptera: Dytiscidae) with the description of nine new taxa, and notes on other species of the genus. Annalen des Naturhistorischen Museums in Wien, 101B, 217–269.

    Fery, H. (2003) Dytiscidae: V. Taxonomic and distributional notes on Hygrotus Stephens, with emphasis on the Chinese fauna and a key to the Palearctic species (Coleoptera). In: Jäch, M.A. & Ji, L. (Eds), Water Beetles of China. Vol. 3. Zoologisch-Botanische Gesellschaft in Österreich and Wiener Coleopterologenverein, Wien, pp. 133–193.

    Fery, H. (2011) Scarodytes roberti nov. sp. from Greece (Coleoptera, Dytiscidae). Linzer Biologische Beiträge, 43 (1), 381–398.

    Fery, H. (2013) Rhithrodytes—are we all blind? Latissimus, 34, 13–15.

    Fery, H. & Brancucci, M. (1997) A taxonomic revision of Deronectes Sharp, 1882 (Insecta: Coleoptera: Dytiscidae) (part I). Annalen des Naturhistorischen Museums in Wien, 99B, 217–302.

    Fery, H. & Erman, Ö.K. (2009) Five new species of the longulus-group of Hydroporus Clairville, 1806 from north-eastern Turkey (Coleoptera: Dytiscidae). Zootaxa, 2033, 1–12.

    Fery, H., Fresneda, J. & Millán, A. (1996) Bemerkungen zur Nebrioporus ceresyi-Gruppe sowie Beschreibung von Nebrioporus schoedli n. sp. (Coleoptera: Dytiscidae). Entomologische Zeitschrift, 106 (8), 306–328.

    Fery, H., Hasanshahi, G. & Abbasipour, H. (2014) Deronectes tashk nov.sp. from the Fars Province (Iran) (Insecta: Coleoptera: Dytiscidae). Linzer Biologische Beiträge, 46 (2), 1255–1265.

    Fery, H. & Hosseinie, S.O. (1998) A taxonomic revision of Deronectes Sharp, 1882 (Insecta: Coleoptera: Dytiscidae) (part II). Annalen des Naturhistorischen Museums in Wien, 100 B, 219–290.

    Fery, H. & Petrov, P.N. (2013) Amurodytes belovi nov.gen. et nov.sp. from eastern Russia (Coleoptera, Dytiscidae, Hydroporinae). Linzer Biologische Beiträge, 45 (2), 1821–1838.

    Fery, H. & Petrov, P.N. (2014) Notes on the identity of Oreodytes okulovi Lafer 1988 (Coleoptera: Dytiscidae). Linzer Biologische Beiträge, 46 (2), 1241–1254.

    Fery, H., Petrov, P.N. & Alarie, Y. (2007) Notes on the identity of Oreodytes dauricus (Motschulsky, 1860) and other members of the O. alaskanus-clade (Coleoptera: Dytiscidae). Linzer Biologische Beiträge, 39 (1), 65–78.

    Fery, H. & Štastný, J. (2007) Notes on the Scarodytes savinensis-complex with the description of two new taxa (Coleoptera: Dytiscidae). Linzer Biologische Beiträge, 39 (2), 877–899.

    Galewski, K. (1994) The description of third-stage larvae of Hydroporus tartaricus Lec. and H. marginatus (Duft.) (Coleoptera, Dytiscidae) with a key to third-stage larvae of Central European species of Hydroporini. Bulletin of the Polish Academy of Sciences, Biological Sciences, 42, 95–113.

    García-Vázquez, D., Bilton, D.T., Alonso, R., Benetti, C.J., Garrido, J., Valladares, L.F. & Ribera, I. (2016) Reconstructing ancient Mediterranean crossroads in Deronectes diving beetles. Journal of Biogeography, 43, 1533–1545.

    Gozis, M. des (1914) Tableaux de détermination des dytiscides, notérides, hyphydrides, hygrobiides et haliplides de la Faune Franco-Rhénane (part). Miscellania Entomologica, 22 (3–4), 129–160.

    Guignot, F. (1931–1933) Les hydrocanthares de France. Toulouse: Les Frères Douladoure, xv + 1057 pp. [pp. 1–188 in 1931; pp. 189–786 in 1932; pp. 787–1057 in 1933]

    Guignot, F. (1941) Description d'un Potamonectes nouveau du Maroc et considérations sur la systématique du genre. Bulletin de la Société des Sciences Naturelles du Maroc, 21, 57–60.

    Guignot, F. (1945) Dix-neuvième note sur les hydrocanthares. Bulletin de la Société d'Étude des Sciences Naturelles de Vaucluse, 14 (3–4), 5–9. [1943]

    Guignot, F. (1947) Coléoptères hydrocanthares. Faune de France, 48, 1–287.

    Guignot, F. (1950) Trente-et-unième note sur les hydrocanthares. Bulletin de la Société Linnéenne de Lyon, 19 (1), 25–28.

    Gyllenhal, L. (1826) Hydroporus borealis. In: Sahlberg, C.R., Insecta fennica, dissertationibus academicis, A. 1817–1834 Editis. Vol. 1 A. Part 12. Frenckel, Helsingforsiae, pp. 174

    Hájek, J. (2017) Dytiscidae. In: Löbl, I. & Löbl, D. (Eds), Catalogue of Palaearctic Coleoptera. Vol. 1. Revised and updated edition. Archostemata-Myxophaga-Adephaga. Brill, Leiden and Boston, pp. 846–914

    Hájek, J., Šťastný, J., Boukal, M. & Fery, H. (2011) Updating the eastern Mediterranean Deronectes (Coleoptera: Dytiscidae) with the description of two new species from Turkey. Acta Entomologica Musei Nationalis Pragae, 51 (2), 463–476.

    Hebert, P.D.N., Cywinska, A., Ball, S.L. & Waard, J.R. de (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Entomological Society of London (B), 270, 313–321.

    Hebert, P.D.N., Penton, E. H., Burns, J. M., Janzen, D. H., & Hallwachs, W. (2004). Ten species in one: DNA barcoding reveals cryptic species in the Neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences of the United States of America, 101 (41), 14812–14817.

    Hernando, C., Aguilera, P., Castro, A. & Ribera, I. (2012) A new interstitial species of the Hydroporus ferrugineus group from north-western Turkey, with a molecular phylogeny of the H. memnonius and related groups (Coleoptera: Dytiscidae: Hydroporinae). Zootaxa, 3173, 37–53.

    Horn, G.H. (1883) Miscellaneous notes and short studies of North American Coleoptera. Transactions of the American Entomological Society, 10, 269–312.

    ICZN (1999) International code of zoological nomenclature. 4th Edition. The International Trust for Zoological Nomenclature, London, 306 pp.

    Katoh, K. & Toh, H. (2008) Recent developments in the MAFFT multiple sequence alignment program. Briefings in Bioinformatics, 9, 286–298.

    Kholin, S.K. & Nilsson, A.N. (2000) Biogeography of the Dytiscidae (Coleoptera) of Kamchatka: Preliminary note. Natural History Research, Special Issue, Chiba, 7, 93–99.

    Lafer, G.S. (1988) New species of diving beetles (Coleoptera, Dytiscidae) from the Far East of the USSR. In: Fauna, systematics and biology of freshwater invertebrates. Akademija Nauk SSSR, Vladivostok, pp. 52–59 [in Russian]

    Larson, D.J. (1975) The predaceous water beetles (Coleoptera: Dytiscidae) of Alberta: systematics, natural history and distribution. Quaestiones Entomologicae, 11, 245–498.

    Larson, D.J. (1991) A new species of Potamonectes Zimmermann (Coleoptera: Dytiscidae) from Labrador. The Coleopterists Bulletin, 45, 280–284.

    Larson, D.J., Alarie, Y. & Roughley, R.E. (2000) Predaceous diving beetles (Coleoptera: Dytiscidae) of the Nearctic Region, with emphasis on the fauna of Canada and Alaska. NRC Research Press, Ottawa, 982 pp.

    Lawrence, J.F., Hastings, A.M., Dallwitz, M.J., Paine, T.A. & Zurcher, E.J. (1999) Beetles of the world: A key and information system for families and subfamilies. Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne. [CD-ROM]

    LeConte, J.L. (1851–1852) Descriptions of new species of Coleoptera, from California. Annals of the Lyceum of Natural History of New York, 5, 125–216. [pp. 125–184 in 1851; pp. 185–216 in 1852]

    Legros, C. (1956) Un Deronectes nouveau d'Espagne (Col. Dytiscidae). Bulletin de la Société Entomologique de France, 61, 134–136.

    Lewis, P.O. (2001) A likelihood approach to estimating phylogeny from discrete morphological character data. Systematic Biology, 50, 913–925.

    Maddison, W.P. & Maddison, D.R. (2015) Mesquite: A modular system for evolutionary analysis. Version 3.02 Available from: 29 November 2017)

    Madsen, B.L. (2011) Pore respiration in some small diving beetles. Latissimus, 29, 7–8.

    Madsen, B.L. (2012) Submersion respiration in small diving beetles (Dytiscidae). Aquatic Insects, 34 (Supplement 1), 57–76.

    Mazzoldi, P. & Toledo, M. (1998) A new Stictotarsus (Insecta: Coleoptera: Dytiscidae) from the Sahara and observations on Stictotarsus sensu Nilsson & Angus, 1992. Annalen des Naturhistorischen Museums in Wien, 100 (B), 203–218.

    Michat, M., Alarie, Y. & Miller, K.B. (2017) Higher-level phylogeny of diving beetles (Coleoptera: Dytiscidae) based on larval characters. Systematic Entomology, 42, 734–767.

    Millán, A., Picazo, F., Fery, H., Moreno, J.M. & Sánchez-Fernández, D. (2013) Stictonectes abellani sp. n. (Coleoptera: Dytiscidae: Hydroporinae) from the Iberian Peninsula, with notes on the phylogeny, ecology and distribution of the Iberian species of the genus. Zootaxa, 3745 (5), 533–550.

    Miller, K.B. (2001) On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics & Evolution, 32, 45–92.

    Miller, K.B. & Bergsten, J. (2014) The phylogeny and classification of predaceous diving beetles. In: Yee, D.A. (Ed.), Ecology, systematics and the natural history of predaceous diving beetles (Coleoptera: Dytiscidae). Springer, Berlin, pp. 49–172.

    Miller, K.B. & Bergsten, J. (2016) Diving beetles of the world: Systematics and biology of the Dytiscidae. Totness: Johns Hopkins University Press, ix + 305 pp.

    Miller, K.B., Wolfe, G.W. & Biström, O. (2006) The phylogeny of the Hydroporinae and classification of the genus Peschetius Guignot, 1942 (Coleoptera: Dytiscidae). Insect Systematics & Evolution, 37, 257–279.

    Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: Proceedings of the Gateway Computing Environments Workshop (GCE). New Orleans, 2010, pp. 1–8.

    Nascimento, F.F., Reis, M. dos & Yang, Z. (2017) A biologist's guide to Bayesian phylogenetic analysis. Nature Ecology & Evolution, 1, 1446–1454.

    Nilsson, A.N. (1992) A revision of the East African Nebrioporus abyssinicus group (Coleoptera, Dytiscidae). Entomologica Fennica, 3, 81–93.

    Nilsson, A.N. (2001) World catalogue of insects. Vol. 3. Dytiscidae Coleoptera. Stenstrup: Apollo Books, 395 pp.

    Nilsson, A.N. & Angus, R.B. (1992) A reclassification of the Deronectes-group of genera (Coleoptera: Dytiscidae) based on a phylogenetic study. Entomologica Scandinavica, 23, 275–288.

    Nilsson, A.N. & Hájek, J. (2018a) A world catalogue of the family Dytiscidae (Coleoptera, Adephaga), 304 pp. Internet version 31.I.2018. Available from: (accessed 7 March 2018)

    Nilsson, A.N. & Hájek, J. (2018b) Catalogue of Palaearctic Dytiscidae (Coleoptera), 88 pp. Internet Version 31.I.2018. Available from: (accessed 7 March 7 2018)

    Nilsson, A.N. & Holmen, M. (1995) The aquatic Adephaga (Coleoptera) of Fennoscandia and Denmark. II. Dytiscidae. Fauna Entomologica Scandinavica, 32, 1–192.

    Nilsson, A.N., Roughley, R.E. & Brancucci, M. (1989) A review of the genus- and family-group names of the family Dytiscidae Leach (Coleoptera). Entomologica Scandinavica, 20, 287–316.

    Paykull, G. von (1798) Fauna Suecica. Insecta. Vol. 1. Edman, Uppsaliae, x + 358 pp.

    Pederzani, F. (1995) Keys to the identification of the genera and subgenera of adult Dytiscidae (sensu lato) of the world (Coleoptera Dytiscidae). Atti dell'Accademia Roveretana degli Agiati, 244 VII (IV, B), 5–83. [1994]

    Pederzani, F. & Schizzerotto, A. (2000) Sulla presenza di angoli preapicali nelle elitre di Potamonectes martini sardus (Gemminger & Harold, 1868) (Coleoptera Dytiscidae). Atti dell'Accademia Roveretana degli Agiati, 250, VII (IX, B), 139–148.

    Rambaut, A., Suchard, M.A., Xie, D. & Drummond, A.J. (2014) Tracer v1. 6. 2014. Available from: (accessed 10 November 2017)

    Régimbart, M. (1906) Voyage de M. Ch. Alluaud dans l'Afrique orientale. Dytiscidae, Gyrinidae, Hydrophilidae. Annales de la Société Entomologique de France, 75, 235–278.

    Ribera, I. (2003) Are Iberian endemics Iberian? A case-study using water beetles of family Dytiscidae (Coleoptera). Graellsia, 59 (2–3), 475–502.

    Ribera, I. & Faille, A. (2010) A new microphthalmic stygobitic Graptodytes Seidlitz from Morocco, with a molecular phylogeny of the genus (Coleoptera, Dytiscidae). Zootaxa, 2641, 1–14.

    Ribera, I., Fresneda, J., Bucur, R., Izquierdo, A., Vogler, A.P., Salgado, J.M. & Cieslak, A. (2010) Ancient origin of a Western Mediterranean radiation of subterranean beetles. BMC Evolutionary Biology, 10, 29.

    Ribera, I., Nilsson, A.N. & Vogler, A.P. (2004) Phylogeny and historical biogeography of Agabinae diving beetles (Coleoptera) inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 30, 545–562.

    Ribera, I., Vogler, A.P. & Balke, M. (2008) Phylogeny and diversification of diving beetles (Coleoptera: Dytiscidae). Cladistics, 24, 563–590.

    Ronquist, F. & Huelsenbeck, J.P. (2003) MRBAYES 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574.

    Seidlitz, G. (1887) Bestimmungs-Tabelle der Dytiscidae und Gyrinidae des europäischen Faunengebietes. Verhandlungen des Naturforschenden Vereines in Brünn, 25 (1886), 3–136.

    Shaverdo, H.V. & Fery, H. (2006) Oreodytes shorti sp.n. from Mongolia (Coleoptera: Dytiscidae). Koleopterologische Rundschau, 76, 35–42.

    Shaverdo, H.V., Short, A.E.Z. & Davaadorj, E. (2008) Diving beetles of Mongolia (Coleoptera: Dytiscidae). Koleopterologische Rundschau, 78, 43–53.

    Sharp, D. (1882) On aquatic carnivorous Coleoptera or Dytiscidae. The Scientific Transactions of the Royal Dublin Society, 2, 179–1003, pls. 7–18.

    Sharp, D. (1884) The water beetles of Japan. Transactions of the Entomological Society of London, 4 (2), 439–464.

    Shull, V.L., Vogler, A.P., Baker, M.D., Maddison, D.R. & Hammond, P.M. (2001) Sequence alignment of 18S ribosomal RNA and the basal relationships of Adephagan beetles: evidence for monophyly of aquatic families and the placement of Trachypachidae. Systematic Biology, 50, 945–969.

    Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H., & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene sequences and a compilation of conserved polymerase chain reaction primers. Annals of the Entomological Society of America, 87, 651–701.

    Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web-servers. Systematic Biology, 57, 758–771.

    Šťastný, J. & Hájek, J. (2016) Agabus abaensis sp. nov., a distinctive species of the confinis-group from central China (Coleoptera: Dytiscidae: Agabinae). Zootaxa, 4121 (3), 311–318.

    Stephens, J.F. (1828–1829) Illustrations of British entomology. Mandibulata. Vol. II. London: Baldwin & Cradock, 200 pp. [pp. 1–112 in 1828; pp. 113–200 in 1829].

    Swofford, D.L. (2002) PAUP*. Phylogenetic Analysis Using Parsimony (* and other methods), Version 4.0b10. Sinauer Associates, Sunderland, MA.

    Toledo, M. (1998) Dytiscidae: II. The genus Nebrioporus Régimbart, 1906 in China (Coleoptera). In: Jäch, M.A. & Ji, L. (Eds), Water beetles of China. Vol. 2. Zoologisch-Botanische Gesellschaft in Österreich and Wiener Coleopterologverein, Wien, pp. 69–91.

    Toledo, M. (2009) Revision in part of the genus Nebrioporus Régimbart, 1906, with emphasis on the N. laeviventris-group (Coleoptera: Dytiscidae). Zootaxa, 2040, 1–111.

    Villastrigo, A., Fery, H., Manuel, M., Millán, A. & Ribera, I. (2018) Evolution of salinity tolerance in the diving beetle tribe Hygrotini (Coleoptera, Dytiscidae). Zoologica Scripta, 47, 63–71.

    Villastrigo, A., Ribera, I., Manuel, M., Millán, A. & Fery, H. (2017) A new classification of the tribe Hygrotini Portevin, 1929 (Coleoptera: Dytiscidae: Hydroporinae). Zootaxa, 4317 (3), 499–529.

    Watts, C.H.S. (1978) A revision of the Australian Dytiscidae (Coleoptera). Australian Journal of Zoology, Supplementary Series, 57, 1–166.

    Wewalka, G. (1977) Die Arten der Gattung Scarodytes aus Griechenland und eine neue Art dieser Gattung aus Israel (Dytiscidae, Col.). Koleopterologische Rundschau, 53, 137–144.

    Wolfe, G.W. & Zimmerman, J.R. (1984) Sensilla, punctation, reticulation, and body shape in the Hydroporinae (Coleoptera: Dytiscidae). International Journal of Insect Morphology and Embryology, 13 (5–6), 373–387.

    Yang, C. (1996) Dytiscidae. In: Wu, S. & Feng, Z. (Eds), The biology and human physiology in the Hoh Xil region. The comprehensive scientific expedition to the Hoh Xil region. Beijing: Science Press, pp. 142–145. [in Chinese with English abstract]

    Zaitzev, F.A. (1908) Zwei neue sibirische Dytisciden (Coleoptera). Russkoe Entomologicheskoe Obozrenie, 7 (4), 268–269. [1907]

    Zaitzev, F.A. (1953) Fauna SSSR, 58. Nasekomyye zhestkokrylyye, 4. Plavuntsovyye i vertyachki. Izdatelstvo Akademii Nauk SSSR, Moscow, Leningrad, 377 pp. [in Russian]

    Zaitzev, F.A. (1972) Fauna of the U.S.S.R. Coleoptera. Families: Amphizoidae, Hygrobiidae, Haliplidae, Dytiscidae, Gyrinidae. Israel Program for Scientific Translations, Jerusalem, 401 pp.

    Zimmerman, J.R. (1982) The Deronectes (Coleoptera: Dytiscidae) of the southwestern U.S.A., Mexico and Guatemala. The Coleopterists Bulletin, 36 (2), 412–438.

    Zimmerman, J.R. (1985) A revision of the genus Oreodytes in North America (Coleoptera: Dytiscidae). Proceedings of the Academy of Natural Sciences of Philadelphia, 137 (1), 99–127.

    Zimmerman, J.R. & Smith, A.H. (1975) A survey of the Deronectes (Coleoptera: Dytiscidae) of Canada, the United States and northern Mexico. Transactions of the American Entomological Society, 101, 651–722.

    Zimmermann, A. (1919) Die Schwimmkäfer des Deutschen Entomologischen Museums in Berlin-Dahlem. Archiv für Naturgeschichte, 83A (12), 68–249. [1917]

    Zimmermann, A. (1933) Monographie der paläarktischen Dytiscidae, IV. Hydroporinae (4. Teil). Koleopterologische Rundschau, 19 (5), 153–193.