Hydaticus

Hydaticus seminiger, copyright Udo Schmidt.

Belongs within: Dytiscinae.

Hydaticus is a cosmopolitan genus of diving beetles characterised by straight anterolateral margins on the metasternum, a stridulatory device in males consisting of a region of reticulate cells on the dorsum of the second protarsal segment and a series of pegs along the dorsal margin of the protibia, and an irregularly rugulose surface to the pronotum and the humeral regions of the elytra in females (Miller 2003).

<==Hydaticus Leach 1817 [Hydaticini] LM87
    |--H. (Hydaticus) M03
    |    |  i. s.: *H. (H.) transversalis (Pontoppidan 1763) [=Dytiscus transversalis] LM87
    |    |--H. (H.) seminiger M03
    |    `--+--H. (H.) aruspex M03
    |       `--H. (H.) cinctipennis M03
    `--H. (Guignotites) M03
         |  i. s.: H. (G.) bivittatus M01
         |--+--H. (G.) subfasciatus M03
         |  `--H. (Hydaticinus) xanthomelas M03
         `--+--H. (G.) parallelus Clark 1864 M03, LM87 [incl. H. consimilis Régimbart 1887 LM87]
            `--+--+--H. (G.) bowringi M03
               |  `--H. (G.) major M03
               `--+--H. (G.) fabricii M03
                  `--H. (G.) grammicus M03

Hydaticus incertae sedis:
  H. consanguineus Aubé 1838 LM87
  H. conspersus Régimbart 1899 I92
  H. daemeli Sharp 1882 LM87
  H. fabricii (Macleay 1825) [=Colymbetes fabricii] LM87
  H. finus Watts 1978 LM87
  H. flavocinctus M86
  H. goryi Aubé 1836 [incl. H. bihamatus Aubé 1838] LM87
  H. jucundus M03
  H. microdaemeli Watts 1978 LM87
  H. pacificus I92
  H. pulcher (Clark 1863) [=Colymbetes pulcher] LM87
  H. thermonectoides Sharp 1884 I92
  H. variegatus Watts 1978 LM87
  H. vittatus (Fabricius 1775) [=Dytiscus vittatus] LM87
  H. wattsi Daussin 1980 [=H. modestus Watts 1978 non Sharp 1882] LM87

*Type species of generic name indicated

REFERENCES

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[M86] Macleay, W. 1886. The insects of the Fly River, New Guinea, "Coleoptera". Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 136–157.

[M01] Miller, K. B. 2001. On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32: 45–92.

[M03] Miller, K. B. 2003. The phylogeny of diving beetles (Coleoptera: Dytiscidae) and the evolution of sexual conflict. Biological Journal of the Linnaean Society 79: 359–388.

Sternopriscus

Sternopriscus hansardii, from here.

Belongs within: Hydroporini.

Sternopriscus is a genus of diving beetles found in Australia and New Guinea, characterised by a small, elongate, rugose-punctate form with a prominent mesosternum and distinctly visible fourth segment on the pro- and mesotarsi.

<==Sternopriscus Sharp 1882 LM87
    |--*S. browni Sharp 1882 LM87
    |--S. cervus Watts 1978 LM87
    |--S. hansardii (Clark 1862) [=Hydroporus hansardii; incl. S. clavatus Sharp 1882] LM87
    |--S. marginatus Watts 1978 LM87
    |--S. meadfootii (Clark 1862) [=Hydroporus meadfootii] LM87
    |--S. minimus Lea 1899 LM87
    |--S. montanus Watts 1978 LM87
    |--S. multimaculatus (Clark 1862) [=Hydroporus multimaculatus; incl. H. sinuatocollis Clark 1862] LM87
    |--S. mundanus Watts 1978 LM87
    |--S. obscurus Sharp 1882 LM87
    |--S. signatus Sharp 1882 LM87
    |--S. tarsalis Sharp 1882 LM87
    |    |--S. t. tarsalis LM87
    |    `--S. t. oscillator Sharp 1882 LM87
    |--S. tasmanicus Sharp 1882 LM87
    `--S. wehnckei Sharp 1882 LM87

*Type species of generic name indicated

REFERENCES

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

Dytiscoidea

Hygrobia hermanni, copyright Frank Koehler.

Belongs within: Adephaga.
Contains: Noteridae, Dytiscidae.

The Dytiscoidea is a clade of predatory beetles united by the possession of extensively fused mesal metacoxal walls (Ribera et al. 2002). Though all dytiscoids are found in damp habitats, advanced swimming abilities through simultaneous strokes of the hind legs are only found in species of Noteridae and Dytiscidae. Hygrobiidae swim with alternate movements of the hind legs, while Amphizoa and Aspidytes do not swim. Meru phyllisae is known from a whitewater cascade in Venezuela, where it crawls over submerged dead leaves lying on the exposed bedrock (Bouchard 2014). Simultaneous strokes also appear to have been the swimming mode used by the Coptoclavidae, known from the Jurassic and Early Cretaceous; these beetles were mostly excellent swimmers and several genera even became adapted to marine environments (Beutel et al. 2013).

<==Dytiscoidea
    |  i. s.: Necronectulus S02
    |         Colymbothethis [Colymbotethidae] BW13
    |           `--C. antecessor Ponomarenko 1993 BW13
    |         Protodytiscus johillaensis BW13
    |         Liadytidae BW13
    |           |--Liadytes longus Ponomarenko 1977 RB02, BW13
    |           `--Memptus S02
    |--+--Noteridae RB02
    |  `--Meru [Meruidae] BW13
    |       `--M. phyllisae Spangler & Steiner 2005 B14
    |--Coptoclavidae BW13
    |    |  i. s.: Coptoclavella minor BW13, S02
    |    |         Pseudohydrophilus avitus BW13
    |    |         Ovonectes BW13
    |    |--Charonoscapha [Charonoscaphinae] BW13
    |    |    `--C. grossa Ponomarenko 1977 BW13
    |    |--+--Exedia plana Ponomarenko 1977 BW13
    |    |  `--+--Daohugounectes primitivus Wang et al. 2009 BW13
    |    |     `--Hispanoclavina BW13
    |    `--+--Stygeonectes jurassicus Ponomarenko 1977 BW13
    |       `--Coptoclava [Coptoclavinae] BW13
    |            |--C. africana S02
    |            `--C. longipoda Ping 1928 BW13
    `--+--Dytiscidae MW15
       `--+--+--Aspidytes Ribera, Beutel et al. 2002 MW15, RB02 [Aspidytidae]
          |  |    |--*A. niobe Ribera, Beutel et al. 2002 RB02
          |  |    `--A. wrasei B14
          |  `--Amphizoa LeConte 1854 MW15, PX01 [incl. Dysmathes Mannerheim 1853 PX01; Amphizoidae, Amphizoinae]
          |       |--A. insolens LeConte 1853 B14
          |       |--A. leconti RB02
          |       `--A. striata PX01
          `--+--Parahygrobia [Parahygrobiidae] BW13
             |    `--P. natans Ponomarenko 1977 BW13
             `--Hygrobia Latreille 1804 MW15, LM87 (see below for synonymy)
                  |  i. s.: H. davidi B14
                  |         H. nigra (Clark 1862) [=Pelobius niger] LM87
                  |         H. tarda M01
                  |         H. wattsi B14
                  |--*H. hermanni (Fabricius 1775) LM87, RB02, B14 [=Dystiscus hermanni LM87, *Hydrachna hermanni LM87]
                  `--+--H. australasiae (Clark 1862) RB02, LM87 [=Pelobius australasiae LM87]
                     `--H. maculata Britton 1981 RB02, LM87

Hygrobia Latreille 1804 MW15, LM87 [=Hydrachna Fabricius 1801 non Müller 1776 LM87, Hygriobia LM87; Hygrobiidae RB02, Pelobiidae]

*Type species of generic name indicated

REFERENCES

[BW13] Beutel, R. G., B. Wang, J.-J. Tan, S.-Q. Ge, D. Ren & X.-K. Yang. 2013. On the phylogeny and evolution of Mesozoic and extant lineages of Adephaga (Coleoptera, Insecta). Cladistics 29 (2): 147–165.

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[MW15] McKenna, D. D., A. L. Wild, K. Kanda, C. L. Bellamy, R. G. Beutel, M. S. Caterino, C. W. Farnum, D. C. Hawks, M. A. Ivie, M. L. Jameson, R. A. B. Leschen, A. E. Marvaldi, J. V. McHugh, A. F. Newton, J. A. Robertson, M. K. Thayer, M. F. Whiting, J. F. Lawrence, A. Ślipiński, D. R. Maddison & B. D. Farrell. 2015. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology 40 (4): 835–880.

[M01] Miller, K. B. 2001. On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32: 45–92.

[PX01] Philips, T. K., & W. Xie. 2001. Amphizoidae LeConte, 1853. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 153–155. CRC Press: Boca Raton.

[RB02] Ribera, I., R. G. Beutel, M. Balke & A. P. Vogler. 2002. Discovery of Aspidytidae, a new family of aquatic Coleoptera. Proceedings of the Royal Society of London Series B—Biological Sciences 269: 2351–2356.

[S02] Sinitshenkova, N. D. 2002. Ecological history of the aquatic insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 388–426. Kluwer Academic Publishers: Dordrecht.

Last updated: 28 October 2018.

Dytiscidae

Laccophilus pictus, photographed by Alex Wild.

Belongs within: Dytiscoidea.
Contains: Agabinae, Dytiscinae, Colymbetinae, Copelatinae, Hydroporinae.

The Dytiscidae are the predacious diving beetles, aquatic beetles with enlarged hind legs that are moved simultaneously when swimming. They first appeared in the Early Cretaceous, where Liadytiscus cretaceus is probably the sister group of living dytiscids as indicated by its having the metacoxae rounded anteriorly but the metaventrite less narrowed than in modern dytiscids (Beutel et al. 2013). Miller (2001) identified the earliest-diverging clade among living dytiscids as the Matinae, which retain a plesiomorphic female genital morphology and lack apical subelytral setae. Other subfamilies include the Laccophilinae, characterised by hind legs lacking natatory setae along the tibia though such setae remain present on the tarsus (Miller 2001).

Synapomorphies (from Miller 2001): Anterolateral prothoracic glands present; metacoxae strongly expanded anteriorly with anterior margin strongly curved, metasternal wings narrowed laterally; large rounded posteromedial metacoxal lobes present; posteromedial metacoxal rim continuous medially; metacoxal cavities contiguous, separated only by a fine medial wall; medial portion of metacoxa not concave laterally.

<==Dytiscidae
    |  i. s.: Cretodytes S02
    |         Palaeogyrinus S02
    |         Schistomerus californicus S02
    |         Angaragabus jurassicus GE05, P02
    |         Allopachria [incl. Nipponhydrus Guignot 1954] NR97
    |--Liadytiscus [Liadytiscinae] BW13
    |    `--L. cretaceus Prokin & Ren 2010 BW13
    |--Matinae [Matini] M01
    |    |--Allomatus Mouchamps 1964 M01, LM87
    |    |    |--*A. wilsoni Mouchamps 1964 LM87
    |    |    `--A. nannup Watts 1978 LM87
    |    |--Matus Aubé 1836 M01
    |    |    |--M. bicarinatus M01
    |    |    `--M. ovatus M01
    |    `--Batrachomatus Clark 1863 RL01, LM87
    |         |--*B. wingi Clark 1863 LM87
    |         `--B. daemeli (Sharp 1882) [=Matus daemeli; incl. B. burnsi Mouchamps 1964] LM87
    `--+--+--Agabinae M01
       |  `--+--Dytiscinae M01
       |     `--+--Colymbetinae M01
       |        `--Lancetes Sharp 1882 [Lancetinae, Lancetini] M01
       |             |--*L. varius (Fabricius 1775) [=Dytiscus varius] LM87
       |             |--L. lanceolatus (Clark 1863) [=Colymbetes lanceolatus] LM87
       |             |--L. marginatus R89
       |             `--L. nigriceps M01
       `--+--+--Coptotomus Say 1834 [Coptotominae, Coptotomini] M01
          |  |    |--C. interrogatus M01
          |  |    |--C. lenticus M01
          |  |    |--C. longulus M01
          |  |    `--C. venustus M01
          |  `--+--Copelatinae M01
          |     `--+--Hydroporinae M01
          |        `--Hydrodytes Miller 2001 [Hydrodytinae, Hydrodytini] M01
          |             `--*H. opalinus (Zimmermann 1921) [=Agaporomorphus opalinus] M01
          `--Laccophilinae M01
               |  i. s.: Australphilus Watts 1978 LM87
               |           |--*A. saltus Watts 1978 LM87
               |           `--A. montanus Watts 1978 LM87
               |--Agabetes Crotch 1873 RL03 [Agabetinae, Agabetini M01]
               |    `--A. acuductus M01
               `--Laccophilini M01
                    |--Laccodytes Régimbart 1895 M01, RL01
                    |    `--L. pumilio RL01
                    `--+--+--Neptosternus M01
                       |  `--Philaccolilus belissimus M01
                       `--Laccophilus Leach 1817 M01
                            |--*L. minutus (Linnaeus 1758) [=Dytiscus minutus] LM87
                            |--L. angustus Régimbart 1889 R89
                            |--L. balzani Régimbart 1889 R89
                            |--L. biguttatus M01
                            |--L. cingulatus Sharp 1882 (see below for synonymy) LM87
                            |--L. clarki Sharp 1882 LM87
                            |--L. hyalinus C01
                            |--L. maculosus M01
                            |--L. nubilus Régimbart 1889 R89
                            |--L. obliquatus Régimbart 1889 R89
                            |--L. ovatus R89
                            |--L. pictus M01
                            |    |--L. p. pictus B14
                            |    |--L. p. coccinelloides Régimbart 1889 B14
                            |    `--L. p. insignis B14
                            |--L. quadrimaculatus Sharp 1882 LM87
                            |--L. religatus Sharp 1882 LM87
                            |--L. seminiger Fauvel 1883 LM87
                            |--L. sharpi Régimbart 1889 LM87
                            |--L. univittatus Régimbart 1891 LM87
                            |--L. variegatus C01
                            `--L. walkeri Balfour-Browne 1939 LM87

Laccophilus cingulatus Sharp 1882 [incl. L. basifasciatus Zimmermann 1922, L. unifasciatus Sharp 1882] LM87

*Type species of generic name indicated

REFERENCES

[BW13] Beutel, R. G., B. Wang, J.-J. Tan, S.-Q. Ge, D. Ren & X.-K. Yang. 2013. On the phylogeny and evolution of Mesozoic and extant lineages of Adephaga (Coleoptera, Insecta). Cladistics 29 (2): 147–165.

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[M01] Miller, K. B. 2001. On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32: 45–92.

[NR97] Nilsson, A. N., & R. E. Roughley. 1997. The genus- and family-group names of the Dytiscidae—additions and corrections. Beiträge zur Entomologie 47 (2): 359–364.

[P02] Ponomarenko, A. G. 2002. Superorder Scarabaeidea Laicharting, 1781. Order Coleoptera Linné, 1758. The beetles. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 164–176. Kluwer Academic Publishers: Dordrecht.

[R89] Régimbart, M. 1889. Enumération des Haliplidae, Dytiscidae et Gyrinidae recueillis par M.^r le Prof. L. Balzan dans l’Amérique méridionale et description de quelques autres espèces voisines. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 256–268.

[RL01] Roughley, R. E., & D. J. Larson. 2001. Dytiscidae Leach, 1815. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 156–186. CRC Press: Boca Raton.

[S02] Sinitshenkova, N. D. 2002. Ecological history of the aquatic insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 388–426. Kluwer Academic Publishers: Dordrecht.

Last updated: 27 October 2018.

Agabinae

Agabus unguicularis, photographed by N. Sloth.

Belongs within: Dytiscidae.
Contains: Ilybius.

The Agabinae is a group of diving beetles sharing the characters of an externally visible scutellum, and metatarsal claws that are equal in length. Members of the Agabini are also united by the presence of a linear series of setae at the anteroventral angle of the metafemur (secondarily lost in some taxa) (Miller 2001). Miller (2001) classified Hydronebrius in the Agabinae, despite the latter's lack of the aforementioned setal series, due to its similar genitalia to derived members of the Agabini; its relationship to the Agabini requires investigation. The genus Hydrotrupes has also been excluded from Agabinae on the basis of the presence of open mandibular grooves in its larva (in contrast to the closed grooves found in other Agabinae and most other Dytiscidae except Copelatinae) but this separation was not supported by Miller (2001). The type genus Agabus is a primarily Holarctic group of species with a continuous bead along the anterior clypeal margin.

<==Agabinae
    |--Hydronebrius [Hydronebriini] M01
    `--Agabini M01
         |  i. s.: Andonectes M01
         |--Hydrotrupes palpalis M01
         `--+--+--Agametrus M01
            |  `--Leuronectes M01
            |       |--L. curtulus M01
            |       `--L. gaudichaudii M01
            |--+--Ilybius M01
            |  `--+--‘Agabus’ ambiguus M01
            |     |--‘Agabus’ bifarius M01
            |     |--‘Agabus’ disintegratus M01
            |     `--‘Agabus’ punctulatus M01
            |--Platynectes Regimbart 1878 M01, LM87 (see below for synonymy)
            |    |--*P. decemnotatus (Aubé 1838) [=Agabus decemnotatus] LM87
            |    |--P. aenescens Sharp 1882 [incl. P. subaenescens Lea 1899] LM87
            |    |--P. australicus Guéorguiev 1972 LM87
            |    |--P. bakewelli (Clark 1863) [=Agabus bakewelli; incl. P. obscurus Sharp 1882] LM87
            |    |--P. beroni S86
            |    |--P. brownei Guéorguiev 1972 [=P. (*Australonectes) brownei] LM87
            |    |--P. chapmani S86
            |    |--P. darlingtoni Guéorguiev 1972 [incl. P. multimaculatus Watts 1978] LM87
            |    |--P. decempunctatus (Fabricius 1775) (see below for synonymy) LM87
            |    |--P. laurianus Watts 1978 LM87
            |    |--P. magellanicus (Babington 1841) [=Colymbetes magellanicus, P. (*Neoplatynectes) magellanicus] LM87
            |    |--P. monostigma (Hope 1842) [=Colymbetes monostigma; incl. Agabus latissimus Clark 1863] LM87
            |    |--P. octodecimmaculatus (Macleay 1825) [=Colymbetes octodecimmaculatus] LM87
            |    |--P. reticulosus (Clark 1863) [=Agabus reticulosus; incl. P. limbatus Sharp 1882] LM87
            |    `--P. tasmaniae (Clark 1863) (see below for synonymy) LM87
            `--+--+--Agabinus glabrellus M01
               |  `--Platambus M01
               |       |--P. (Platambus) maculatus M01
               |       `--P. (Anagabus) lineatus M01
               `--Agabus Leach 1817 M01
                    |  i. s.: A. heydeni RB02
                    |         A. paludosus C01
                    |         A. ragazzii G89
                    |--A. spinipes M01
                    `--+--+--+--A. cordatus M01
                       |  |  `--A. seriatus M01
                       |  `--+--A. punctatus M01
                       |     |--A. semivittatus M01
                       |     `--A. tristis M01
                       `--+--A. unguicularis M01
                          |--+--A. antennatus M01
                          |  `--A. crassipes [=Carrhydrus crassipes] M01
                          |--+--A. approximatus M01
                          |  |--A. griseipennis M01
                          |  `--A. obliteratus M01
                          `--+--A. japonicus M01
                             `--+--A. minnesotensis M01
                                `--A. regularis M01

Platynectes Regimbart 1878 M01, LM87 [incl. Australonectes Guéorguiev 1972 LM87, Neoplatynectes Guéorguiev 1972 LM87]

Platynectes decempunctatus (Fabricius 1775) [=Dytiscus decempunctatus; incl. Agabus lugubris Blanchard 1853, A. mastersii Macleay 1871, Lancetes ocularis Lea 1895, Platynectes polygrammus Régimbart 1899, Agabus spilopterus Germar 1848] LM87

Platynectes tasmaniae (Clark 1863) [=Agabus tasmaniae; incl. P. daemeli Sharp 1882, P. gagatinus Lea 1899] LM87

*Type species of generic name indicated

REFERENCES

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso «Esploratore» dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 5–72.

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[M01] Miller, K. B. 2001. On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32: 45–92.

[RB02] Ribera, I., R. G. Beutel, M. Balke & A. P. Vogler. 2002. Discovery of Aspidytidae, a new family of aquatic Coleoptera. Proceedings of the Royal Society of London Series B—Biological Sciences 269: 2351–2356.

[S86] Spangler, P. J. 1986. Insecta: Coleoptera. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 622–631. E. J. Brill/Dr W. Backhuys: Leiden.

Last updated: 5 February 2018.

Aciliini

Sunburst diving beetle Thermonectus marmoratus, from Insectarium de Montréal, René Limoges.

Belongs within: Dytiscinae.

The Aciliini are a clade of diving beetles, distinguished by the possession of bifid metatibial spurs, and extremely long gonocoxae with expanded and rounded apices and apicolateral margins concave (Miller 2003). Members of the Aciliini lay their eggs in cracks in wood, moss or other cryptic locations (Miller 2001). Within the Aciliini, members of the Holarctic genus Acilius are characterised by large, dense punctation over much of the body surface in contrast to the finer, sparser punctation found in other genera. The genus Thermonectus includes two species, T. marmoratus and T. zimmermanni, that are strikingly patterned with bright yellow markings (Bouchard 2014).

<==Aciliini [Thermonectini]
    |  i. s.: Tikoloshanes eretiformis M01
    |         Aethionectes M01
    |           |--A. apicalis M01
    |           `--A. optatus M01
    |--Thermonectus Dejean 1833 M03, RL01
    |    |  i. s.: T. succinctus R89
    |    |         T. zimmermani B14
    |    |--T. marmoratus Hope 1832 M03, B14
    |    `--+--T. basilaris M01
    |       |--T. circumscriptus M03
    |       |--T. margineguttatus M01
    |       |--T. nigrofasciatus M03
    |       |--T. nobilis M01
    |       `--T. ornaticollis M01
    `--+--Rhantaticus Sharp 1882 M03, LM87
       |    `--R. congestus (Klug 1833) (see below for synonymy) LM87
       `--+--Sandracottus Sharp 1882 M03, LM87
          |    |--*S. mixtus (Blanchard 1853) [=Acilius mixtus] LM87
          |    `--S. bakewelli (Clark 1864) [=Hydaticus bakewelli; incl. S. guttatus Sharp 1882] LM87
          `--+--Graphoderus Dejean 1833 M03, RL01 (see below for synonymy)
             |    |  i. s.: ‘Dytiscus’ cinereus Linnaeus 1758 [=*Prosciastes cinereus] NR97
             |    |         G. occidentalis M01
             |    |         G. perplexus M01
             |    |--G. liberus M03
             |    `--+--G. fascicollis M03
             |       `--G. zonatus M03
             `--Acilius Leach 1817 M03, RL01 [incl. Heteroelytrus Ádám in Mahunka 1993 NR97]
                  |  i. s.: A. athabascae RL01
                  |         A. japonicus I92
                  |         A. kishii Nakane 1963 I92
                  |         A. mediatus RL01
                  |--A. sulcatus (Linnaeus 1758) M03, NR97 [=Dytiscus sulcatus NR97, *Heteroelytrus sulcatus NR97]
                  `--A. (Acilius) M01
                       |  i. s.: A. (A.) abbreviatus M01
                       |--A. (A.) semisulcatus M01, M03
                       `--+--A. (A.) sylvanus M01, M03
                          `--A. (Homoeodytes Régimbart 1878) M01, LM87
                               |--‘*Homoeodytes’ scutellaris (Germar 1848) (see below for synonymy) LM87
                               |--‘Homoeodytes’ atratus (Fabricius 1801) (see below for synonymy) LM87
                               `--A. (H.) mediatus M01

Graphoderus Dejean 1833 M03, RL01 [incl. Derographus Portevin 1929 RL01, Graphothorax Motschulsky 1853 RL01, Prosciastes Gistel 1856 RL01]

‘Homoeodytes’ atratus (Fabricius 1801) [=Dytiscus atratus; incl. Cybister goryi Aubé 1838] LM87

‘*Homoeodytes’ scutellaris (Germar 1848) [=Cybister scutellaris; incl. C. incognitus Gschwendtner 1931] LM87

Rhantaticus congestus (Klug 1833) [=Hydaticus congestus; incl. H. signatipennis Castelnau 1834, *Rhantaticus signatipennis] LM87

*Type species of generic name indicated

REFERENCES

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[M01] Miller, K. B. 2001. On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32: 45–92.

[M03] Miller, K. B. 2003. The phylogeny of diving beetles (Coleoptera: Dytiscidae) and the evolution of sexual conflict. Biological Journal of the Linnaean Society 79: 359–388.

[NR97] Nilsson, A. N., & R. E. Roughley. 1997. The genus- and family-group names of the Dytiscidae—additions and corrections. Beiträge zur Entomologie 47 (2): 359–364.

[R89] Régimbart, M. 1889. Enumération des Haliplidae, Dytiscidae et Gyrinidae recueillis par M.^r le Prof. L. Balzan dans l’Amérique méridionale et description de quelques autres espèces voisines. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 256–268.

[RL01] Roughley, R. E., & D. J. Larson. 2001. Dytiscidae Leach, 1815. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 156–186. CRC Press: Boca Raton.

Last updated: 26 October 2018.

Dytiscinae

Megadytes latus, photographed by Wolfgang Walz Hillermann.

Belongs within: Dytiscidae.
Contains: Hydaticus, Aciliini.

The Dytiscinae are a large clade of diving beetles. With the exception of the basal Cybistrini, males of the Dytiscinae have the pro- and mesotarsomeres broadly expanded with ventral sucker-like setae, used to grip the female during mating. In a number of subsequent lineages, females have conversely evolved ornamented elytra that may reduce the effectiveness of such gripping setae (Miller 2003). Members of the tribe Cybistrini and Dytiscus have knife-like gonocoxae that they use to insert eggs into plant tissue. Other tribes have long, blunt gonocoxae and lay eggs in cracks in wood, moss or other cryptic positions (Miller 2001).

Cybistrini and Dystiscus are much larger than most other dytiscids; Cybistrini are more wedge-shaped than the elongate-oval Dytiscus (Roughley & Larson 2001). The Cybistrini are further united by a number of features including a small apicoventral region of very stiff setae on the elytra, a dorsal series of natatory setae on the metafemur, a relatively large cluster of setae at the apicodorsal angle of the metatibia, and an anterior metatibial spur that is acuminate and distinctly broader basally than the posterior spur (Miller 2001).

Chracters (from Miller 2001): Anterolateral margin of eye evenly rounded; female genitalia of dytiscine type, with bursa absent and spermatophore placed in membranous pouch ventrad of gonocoxae; gonocoxae strongly fused dorsally; median lobe bilaterally symmetrical; larvae with fringe of fine setae along last abdominal segments.

<==Dytiscinae
    |--Cybistrini M03
    |    |  i. s.: Regimbartina pruinosa M01
    |    |--+--Spencerhydrus Sharp 1882 M03, LM87
    |    |  |    |--*S. latecinctus Sharp 1882 [incl. S. semiflavus Sharp 1882] LM87
    |    |  |    `--S. pulchellus Sharp 1882 LM87
    |    |  `--+--Austrodytes Watts 1978 M01, LM87
    |    |     |    `--*A. insularis (Hope 1842) [=Cybister insularis] LM87
    |    |     `--Onychohydrus M03
    |    |          |--O. (Onychohydrus) M01
    |    |          |    |--O. (O.) hookeri M01
    |    |          |    `--O. (O.) scutellaris M01
    |    |          `--O. (Sternhydrus) atratus M01
    |    `--+--Megadytes Sharp 1882 M03, RL01
    |       |    |  i. s.: M. aeneus Ormancey 1843 R89
    |       |    |         M. fallax R89
    |       |    |         M. puncticollis R89
    |       |    |--M. (Bifurcitus) giganteus M01
    |       |    `--+--M. (Paramegadytes) glaucus M01
    |       |       `--M. (Megadytes) M01
    |       |            |--M. (M.) fraternus M01
    |       |            |--M. (M.) laevigatus M01
    |       |            |--M. (M.) latus M01
    |       |            `--M. (M.) marginithorax M01
    |       `--Cybister Curtis 1827 M03, LM87 (see below for synonymy)
    |            |  i. s.: *C. (Cybister) virens (Müller 1776) [=Dytiscus virens] LM87
    |            |         C. cephalotes G89
    |            |         C. godeffroyi (Wehncke 1876) [=Trogus godeffroyi; incl. C. simoni Régimbart 1877] LM87
    |            |         C. loxidiscus Wilke 1919 LM87
    |            |         C. yulensis Guignot 1956 LM87
    |            |         C. (Scaphinectes Ádám in Mahunka 1993) [=Trochalus Dejean 1833 non Laporte 1832] NR97
    |            |           `--‘Dytiscus’ laterimarginalis De Geer 1774 (see below for synonymy) NR97
    |            |--C. marginicollis M01
    |            |--+--C. ellipticus M01
    |            |  `--C. fimbriolatus M01
    |            `--+--C. puncticollis M01
    |               `--C. tripunctatus (Olivier 1795) M01, LM87 (see below for synonymy)
    |                    |--C. t. tripunctatus I92
    |                    `--C. t. orientalis Gschwendtner 1931 I92
    `--+--+--Hyderodes Hope 1838 M03, LM87 [Hyderodini]
       |  |    |--*H. shuckardi Hope 1838 LM87
       |  |    `--H. crassus Sharp 1882 [incl. H. collaris Sharp 1882] LM87
       |  `--+--Hydaticini RL01
       |     |    |--Hydaticus M03
       |     |    `--Prodaticus RL01
       |     `--+--+--Aciliini M03
       |        |  `--Notaticus [Aubehydrinae, Aubehydrini] M03
       |        |       `--N. fasciatus M03
       |        `--Eretes Laporte 1833 M03, RL01 [incl. Eunectes Erichson 1832 (preoc.) RL01; Eretini]
       |             |--*E. stictus (Linnaeus 1767) [=Dytiscus stictus] LM87
       |             |--E. australis (Erichson 1842) [=Eunectes australis; incl. Eu. punctipennis Macleay 1871] LM87
       |             |--E. explicitus B14
       |             |--E. griseus M03
       |             `--E. occidentalis RL01
       `--Dytiscus Linnaeus 1758 M01 [incl. Leionotus Kirby 1837 RL01, Macrodytes Thomson 1859 RL01; Dytiscini]
            |--D. alaskanus M01
            |--D. caraboides Linnaeus 1758 L58
            |--D. circumcinctus Ahrens 1811 BW04
            |--D. cordieri M01
            |--D. dauricus M01
            |--D. delictus Zaitsev 1906 I92
            |--D. erytrocephalus Linnaeus 1758 L58
            |--D. fasciventris M03
            |--D. flavocinctus B35
            |--D. fuscipes Linnaeus 1758 L58
            |--D. fuscus Linnaeus 1758 L58
            |--D. hermanni [=Hydrachne hermanni] L02
            |--D. latissimus Linnaeus 1758 L58
            |--D. maculatus Linnaeus 1758 L58
            |--D. marginalis Linnaeus 1758 B14 [incl. D. marginalis var. conformis C01]
            |--D. marginicollis M01
            |--D. persicus B14
            |--D. piceus Linnaeus 1758 L58
            |--D. scarabaeoides Linnaeus 1758 L58
            |--D. semistriatus Linnaeus 1758 L58
            |--D. sexpustulatus L02
            |--D. sharpi Wehncke 1875 I92
            |--D. striatus Linnaeus 1758 L58
            `--D. verticalis M03

Cybister Curtis 1827 M03, LM87 [incl. Alocomerus Brinck 1945 RL01, Cybisteter Bedel 1881 RL01, Gschwendtnerhydrus Brinck 1945 RL01, Megadytoides Brinck 1945 RL01, Meganectes Brinck 1945 RL01, Nealocomerus Brinck 1945 RL01, Trogus Leach 1817 RL01]

Cybister tripunctatus (Olivier 1795) M01, LM87 [=Dytiscus tripunctatus LM87; incl. C. gayndahensis Macleay 1871 LM87, C. granulatus Blackburn 1888 LM87]

‘Dytiscus’ laterimarginalis De Geer 1774 [incl. D. roeselii Füessly 1775, *Scaphinectes roeselii] NR97

*Type species of generic name indicated

REFERENCES

[BW04] Balke, M., C. H. S. Watts, S. J. B. Cooper, W. F. Humphreys & A. P. Vogler. 2004. A highly modified stygobiont diving beetle of the genus Copelatus (Coleoptera, Dytiscidae): taxonomy and cladistic analysis based on mitochondrial DNA sequences. Systematic Entomology 29: 59–67.

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso «Esploratore» dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 5–72.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[M01] Miller, K. B. 2001. On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32: 45–92.

[M03] Miller, K. B. 2003. The phylogeny of diving beetles (Coleoptera: Dytiscidae) and the evolution of sexual conflict. Biological Journal of the Linnaean Society 79: 359–388.

[NR97] Nilsson, A. N., & R. E. Roughley. 1997. The genus- and family-group names of the Dytiscidae—additions and corrections. Beiträge zur Entomologie 47 (2): 359–364.

[R89] Régimbart, M. 1889. Enumération des Haliplidae, Dytiscidae et Gyrinidae recueillis par M.^r le Prof. L. Balzan dans l’Amérique méridionale et description de quelques autres espèces voisines. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 256–268.

[RL01] Roughley, R. E., & D. J. Larson. 2001. Dytiscidae Leach, 1815. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 156–186. CRC Press: Boca Raton.

Last updated: 27 October 2018.

Copelatinae

Copelatus haemorrhoidalis, photographed by Gábor Keresztes.

Belongs within: Dytiscidae.

The Copelatinae is a speciose clade of diving beetles, most species of which are included in the genus Copelatus. Larvae, where known, are presumed solid feeders with serrated mandibles and a foregut including a crop.

Characters (from Miller 2001): Metacoxal lines closely approximated medially (metacoxal lines lost in Lacconectus); spermatheca strongly reduced; scutellum externally visible; metatarsal claws equal in length in both sexes.

<==Copelatinae [Copelatini]
    |  i. s.: Aglymbus M01
    |--Agaporomorphus M01
    |    |--A. knischi M01
    |    `--A. perairai M01
    `--+--Lacconectus M01
       |    |--L. formosanus M01
       |    `--L. peguensis RB02
       `--Copelatus Erichson 1832 M01
            |  i. s.: *C. posticatus (Fabricius 1801) [=Dystiscus posticatus] LM87
            |         C. ater Sharp 1882 LM87
            |         C. australiae Clark 1863 (see below for synonymy) LM87
            |         C. australis (Clark 1863) [=Celina australis; incl. Co. torcutus Guignot 1954] LM87
            |         C. bakewelli Balfour-Browne 1939 LM87
            |         C. boulevardi Watts 1978 LM87
            |         C. clarki Sharp 1882 LM87
            |         C. daemeli Sharp 1882 LM87
            |         C. distinctus M01
            |         C. divisus Balfour-Browne 1939 [=C. gentilis divisus] LM87
            |         C. elongatulus Macleay 1871 LM87
            |         C. ferrugineus Sharp 1882 [incl. C. erraticus Balfour-Browne 1939, C. lividus Sharp 1882] LM87
            |         C. gapa Watts 1978 LM87
            |         C. glyptus Guignot 1954 LM87
            |         C. gracilis Sharp 1882 LM87
            |         C. irregularis Macleay 1871 LM87
            |         C. longicornis R89
            |         C. marginatus Sharp 1882 LM87
            |         C. melanarius Sharp 1882 LM87
            |         C. nakamurai Guéorguiev 1970 I92
            |         C. nigrolineatus Sharp 1882 LM87
            |         C. nomax S86
            |         C. parvulus S62
            |         C. politus M86
            |         C. punctipennis Lea 1899 LM87
            |         C. ragazzii G89
            |         C. rasilis Lea 1899 LM87
            |         C. simplex Clark 1863 [incl. C. labratus Sharp 1882, C. victoriae Clark 1863] LM87
            |         C. strinatii Guignot 1958 S86
            |         C. subterraneus Gueorguiev 1978 S86
            |         C. tenebrosus Régimbart 1880 LM87
            |--C. haemorrhoidalis M01
            `--+--C. glyphicus M01
               `--C. kalaharii M01

Copelatus australiae Clark 1863 [incl. C. acuductus Clark 1863, C. extensus Sharp 1882, C. nigritulus Sharp 1882, C. tarsalis Balfour-Browne 1939] LM87

*Type species of generic name indicated

REFERENCES

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso «Esploratore» dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 5–72.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[M86] Macleay, W. 1886. The insects of the Fly River, New Guinea, "Coleoptera". Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 136–157.

[M01] Miller, K. B. 2001. On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32: 45–92.

[R89] Régimbart, M. 1889. Enumération des Haliplidae, Dytiscidae et Gyrinidae recueillis par M.^r le Prof. L. Balzan dans l’Amérique méridionale et description de quelques autres espèces voisines. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 256–268.

[RB02] Ribera, I., R. G. Beutel, M. Balke & A. P. Vogler. 2002. Discovery of Aspidytidae, a new family of aquatic Coleoptera. Proceedings of the Royal Society of London Series B—Biological Sciences 269: 2351–2356.

[S62] Spangler, P. J. 1962. Natural history of Plummers Island, Maryland. XIV. Biological notes and description of the larva and pupa of Copelatus glyphicus (Say) (Coleoptera: Dytiscidae). Proceedings of the Biological Society of Washington 75: 19–24.

[S86] Spangler, P. J. 1986. Insecta: Coleoptera. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 622–631. E. J. Brill/Dr W. Backhuys: Leiden.

Last updated: 30 January 2018.

Colymbetinae

Rhantus frontalis, photographed by Josef Dvořák.

Belongs within: Dytiscidae.

The Colymbetinae is a clade of diving beetles diagnosed by the presence of transverse rugae on the dorsal surface of the lateral margins of abdominal sternum two, and usually of unequal metatarsal claws in both sexes. The majority of species are included in a tribe Colymbetini, characterised by a female genital system with a separate bursa and vagina, with the bursa copulatrix long, slender and slightly twisted, and a large and broad spermathecal duct attaching near the base of the bursa. A single New Guinean species, Carabdytes upin, is included in a distinct tribe characterised by a lack of lobes on the metatarsomeres and a hydroporine-type genital system (Miller 2001).

<==Colymbetinae
    |--Anisomeria [Anisomeriini] M01
    |--Carabdytes Balke, Hendrich & Wewalka 1992 [Carabdytini] M01
    |    `--*C. upin Balke, Hendrich & Wewalka 1992 M01
    `--Colymbetini M01
         |--Meladema coriacea M01
         |--Batrachomatus Clark 1863 LM87
         |    |--*B. wingi Clark 1863 LM87
         |    `--B. daemeli (Sharp 1882) [=Matus daemeli; incl. B. burnsi Mouchamps 1964] LM87
         `--Rhantus Dejean 1833 [=Rantus] LM87
              |  i. s.: R. suturalis (Macleay 1825) LM87 (see below for synonymy)
              |         R. flavicollis G89
              |         R. impar Guignot 1956 LM87
              |         R. morneri Falkenström 1937 LM87
              |         R. simulans Régimbart 1908 LM87
              |--R. pseudopacificus M01
              `--+--R. calidus M01
                 `--+--+--R. atricolor M01
                    |  |--R. binotatus M03
                    |  |--R. frontalis M01
                    |  |--R. gutticollis Say 1834 M01, ZL74 [incl. R. hoppingi Wallis 1933 ZL74, R. mexicanus ZL74]
                    |  |--R. sericans M01
                    |  |--R. signatus M01
                    |  `--Hoperius planatus M01
                    `--+--+--R. (Nartus) M01
                       |  |    |--R. (N.) grapii M01
                       |  |    `--R. (N.) sinuatus M01, M03
                       |  `--Neoscutopterus M03
                       |       |--N. angustus M03
                       |       `--N. hornii M01
                       `--Colymbetes Clairville 1806 M03, M01
                            |--C. affinis B35
                            |--C. brunneus B35
                            |--C. decemstriatus B35
                            |--C. densus M01
                            |--C. dolabatus CS77
                            |--C. exaratus M03
                            |--C. niger B35
                            |--C. pacificus B35
                            |--C. parvulus B35
                            `--C. paykulli M03

Rhantus suturalis (Macleay 1825) LM87 [=Colymbetes suturalis LM87; incl. R. australis Aubé 1836 LM87, C. pulverosus Stephens 1828 LM87, *R. pulverosus LM87, R. suturalis var. virgulatus C01]

*Type species of generic name indicated

REFERENCES

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso «Esploratore» dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 5–72.

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[M01] Miller, K. B. 2001. On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32: 45–92.

[M03] Miller, K. B. 2003. The phylogeny of diving beetles (Coleoptera: Dytiscidae) and the evolution of sexual conflict. Biological Journal of the Linnaean Society 79: 359–388.

[ZL74] Zimmerman, J. R., & J. A. Ludwig. 1974. Multiple-discriminant analysis of geographical variation in the aquatic beetle, Rhantus gutticollis (Say) (Dytiscidae). Systematic Zoology 24 (1): 63–71.

Last updated: 30 January 2018.

Hydroporini

Necterosoma penicillatum, copyright Michael Munich.

Belongs within: Hydroporinae.
Contains: Paroster, Sternopriscus, Hydroporus.

The Hydroporini is an assemblage of mostly Holarctic diving beetles, with a group of species found in Australia. The monophyly of this group as generally recognised was not resolved by Miller (2001), who found it to be at least paraphyletic with regard to the Vatellini. The latter are characterised by a prosternal process that is completely excluded from the metasternum, metepisterna not reaching the mesocoxal cavities, and female genitalia with a heavily sclerotised, triangular process extending off the posterior end of the spermatheca (Miller 2001). Members of the Hydroporini include Necterosoma, a primarily Australian genus of diving beetles with outlying species in New Caledonia and Timor, with distinctly pentamerous protarsi, a small subapical spine on each elytron, and males with protibiae notched on the inner side.

<==Hydroporini
    |--+--+--‘Hydroporus’ paugus M01
    |  |  `--Sanfilippodytes Franciscolo 1979 M01, S86
    |  |       `--S. sbordonii Franciscolo 1979 S86
    |  `--Vatellini M01
    |       |--Macrovatellus M01
    |       `--Derovatellus M01
    |            |--D. lentus M01
    |            `--D. peruanus M01
    `--+--Nebrioporus macronychus M01
       |--Porhydrus lineatus M01
       |--Hydroporus M01
       |--Graptodytes M01
       |    |--G. flavipes M01
       |    `--G. granularis M01
       |--Heterosternuta M01
       |    |--H. pulcher M01
       |    `--H. wickhami M01
       |--Oreodytes M01
       |    |--O. congruus M01
       |    |--O. obesus M01
       |    `--O. scitulus M01
       |--Stictotarsus M01
       |    |--S. corpulentus M01
       |    |--S. griseostriatus M01
       |    `--S. striatellus M01
       |--Neoporus M01
       |    |--N. clypealis M01
       |    |--N. dimidiatus M01
       |    |--N. solitarius M01
       |    |--N. superioris M01
       |    `--N. undulatus M03
       `--Necterosoma Macleay 1871 M01, LM87
            |--N. penicillatum (Clark 1862) (see below for synonymy) LM87
            |--N. aphrodite Watts 1978 LM87
            |--N. darwini (Babington 1841) [=Hydroporus darwini] LM87
            |--N. dispar (Germar 1848) [=Hydroporus dispar] LM87
            |--N. regulare Sharp 1882 [incl. N. arcuatum Sharp 1882] LM87
            |--N. schmelzi Sharp 1882 LM87
            |--N. susanna Zwick 1979 LM87
            |--N. thoreyi (Clark 1862) [=Hydroporus thoreyi] LM87
            `--N. undecimlineatum (Babington 1841) (see below for synonymy) LM87

Hydroporini incertae sedis:
  Deronectes M01
  Haideoporus Young & Longley 1976 S86
    `--H. texanus Young & Longley 1976 S86
  Siettitia Abeille de Perrin 1904 S86
    |--S. avenionensis Guignot 1925 S86
    `--S. balsetensis Abeille de Perrin 1904 S86
  Morimotoa Ueno 1957 S86
    `--M. phreatica Ueno 1957 S86
         |--M. p. phreatica S86
         `--M. p. miurai Ueno 1957 S86
  Paroster LM87
  Chostonectes Sharp 1882 LM87
    |--*C. gigas (Boheman 1858) [=Hydroporus gigas; incl. Hyphydrus humeralis Clark 1862] LM87
    |--C. johnsonii (Clark 1862) [=Hydroporus johnsonii; incl. C. latus Sharp 1882] LM87
    |--C. nebulosus (Macleay 1871) [=Hydroporus nebulosus] LM87
    `--C. sharpi Sharp 1882 LM87
  Antiporus Sharp 1882 LM87
    |--*A. femoralis (Boheman 1858) [=Hydroporus femoralis] LM87
    |--A. bakewellii (Clark 1862) [=Hydroporus bakewellii; incl. A. curtulus Sharp 1882] LM87
    |--A. blakei (Clark 1862) [=Hydroporus blakeii] LM87
    |--A. gilbertii (Clark 1862) [=Hydroporus gilbertii] LM87
    |--A. interrogationis (Clark 1862) [=Hydroporus interrogationis] LM87
    |--A. simplex Watts 1978 LM87
    `--A. wilsoni Watts 1978 LM87
  Hypodes Watts 1978 [=Tiporus Watts 1985] LM87
    |--*H. undecimmaculatus (Clark 1862) [=Hydroporus undecimmaculatus, *Tiporus undecimmaculatus] LM87
    |--H. alastairi Watts 1978 [=Tiporus alastairi] LM87
    |--H. centralis Watts 1978 [=Tiporus centralis] LM87
    |--‘Hydroporus’ collaris Hope 1842 (see below for synonymy) LM87
    |--H. denticulatus Watts 1978 [=Tiporus denticulatus] LM87
    |--H. giuliani Watts 1978 [=Tiporus giuliani] LM87
    |--H. josepheni Watts 1978 [=Tiporus josepheni] LM87
    `--H. tambreyi Watts 1978 [=Tiporus tambreyi] LM87
  Sternopriscus LM87
  Barretthydrus Lea 1927 LM87
    |--*B. geminatus Lea 1927 LM87
    |--B. stepheni Watts 1978 LM87
    `--B. tibialis Lea 1927 LM87
  Megaporus Brinck 1943 [=Macroporus Sharp 1882 non Uhler 1875] LM87
    |--*M. hamatus (Clark 1862) (see below for synonymy) LM87
    |--M. fischeri Mouchamps 1964 LM87
    |--M. gardenerii (Clark 1862) [=Hydroporus gardenerii; incl. H. brunnipennis Macleay 1871] LM87
    |--M. howittii (Clark 1862) [=Hydroporus howittii; incl. H. foveiceps Macleay 1871] LM87
    |--M. nativigi Mouchamps 1964 LM87
    |--M. ruficeps (Sharp 1882) [=Macroporus ruficeps] LM87
    |--M. solidus (Sharp 1882) [=Macroporus solidus; incl. Ma. dilatatus Régimbart 1908] LM87
    `--M. wilsoni Mouchamps 1964 LM87

‘Hydroporus’ collaris Hope 1842 [=Tiporus collaris; incl. H. gravidus Clark 1862, H. thoracicus Schaum 1847] LM87

*Megaporus hamatus (Clark 1862) [=Hydroporus hamatus, *Macroporus hamatus; incl. Ma. lateralis Sharp 1882] LM87

Necterosoma penicillatum (Clark 1862) [=Hydroporus penicillatus; incl. N. costipenne Lea 1899, *N. vittipenne Macleay 1871] LM87

Necterosoma undecimlineatum (Babington 1841) [=Hydroporus undecimlineatus; incl. N. flavicolle Macleay 1871, H. wollastonii Clark 1862] LM87

*Type species of generic name indicated

REFERENCES

[LM87] Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[M01] Miller, K. B. 2001. On the phylogeny of the Dytiscidae (Insecta: Coleoptera) with emphasis on the morphology of the female reproductive system. Insect Systematics and Evolution 32: 45–92.

[M03] Miller, K. B. 2003. The phylogeny of diving beetles (Coleoptera: Dytiscidae) and the evolution of sexual conflict. Biological Journal of the Linnaean Society 79: 359–388.

[S86] Spangler, P. J. 1986. Insecta: Coleoptera. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 622–631. E. J. Brill/Dr W. Backhuys: Leiden.

Last updated: 30 january 2018.