Broscinae

Apotomus rufus, copyright elleelle.

Belongs within: Conjunctae.
Contains: Broscini.

The Broscinae are a group of ground beetles characterised by a pedunculate connection between the pro- and mesothorax, and a single supraorbital seta. Members of the genus Apotomus, which have elongate palps and an unmargined prothorax, are found on mudflats adjacent to rivers (Bouchard 2014).

<==Broscinae [Broschidae, Broscitae]
    |--+--Broscini R-JC01
    |  `--Melaenitae [Melaeninae] R-JC01
    |       |--Melaenus [Melaenini] R-JC01
    |       |    `--M. piger R-JC01
    |       `--Cymbionotum B14 [Cymbionotini R-JC01]
    |            `--C. (Procoscinia) B14
    |                 |--C. (P.) fernandezi Ball & Shpeley 2005 B14
    |                 `--C. (P.) negrei B14
    `--Apotomus Illiger 1807 R-JC01, LM87 [Apotominae, Apotomini, Apotomitae]
         |--*A. rufus (Rossi 1790) [=Scarites rufus] LM87
         |--A. australis Castelnau 1867 LM87
         |--A. fuscus B89
         |--A. hirsutulus R-JC01
         |--A. mastersii Macleay 1871 LM87
         |--A. novaehollandiae Castelnau 1867 LM87
         |--A. reichardti Erwin 1980 B14
         `--A. xanthotelus Bates 1874 B89

Broscinae incertae sedis:
  Brithysternum Macleay 1873 LM87
    |--*B. calcaratum Macleay 1873 LM87
    |--B. macleayi Sloane 1910 LM87
    `--B. nodosum Sloane 1910 LM87
  Adotela Castelnau 1867 LM87
    |--A. carenoides Putzeys 1873 LM87
    |--A. concolor Castelnau 1867 LM87
    |--A. esmeralda Castelnau 1867 LM87
    |--A. frenchi Sloane 1890 LM87
    |--A. nigerrima Macleay 1873 LM87
    |--A. striolata Putzeys 1873 LM87
    `--A. viridis (Macleay 1871) [=Promecoderus viridis] LM87
  Parroa Castelnau 1867 LM87
    |--P. apicalis Sloane 1893 LM87
    |--P. atronitens (Sloane 1890) [=Adotela atronitens] LM87
    |--P. australis (Sloane 1890) [=Adotela australis] LM87
    |--P. bicolor Castelnau 1867 LM87
    |--P. carbonaria Castelnau 1867 LM87
    |--P. grandis Castelnau 1867 LM87
    |--P. howittii Castelnau 1867 LM87
    |--P. laevigata Sloane 1893 LM87
    |--P. noctis Sloane 1893 LM87
    `--P. violacea Castelnau 1867 LM87
  Cerotalis Castelnau 1867 LM87
    |--C. amabilis Sloane 1890 LM87
    |--C. brachypleura Sloane 1898 LM87
    |--C. longipes Sloane 1898 LM87
    |--C. majuscula (Putzeys 1868) [=Promecoderus majusculus] LM87
    |--C. semiviolacea Castelnau 1867 LM87
    |--C. substriata Castelnau 1867 LM87
    `--C. versicolor Castelnau 1867 LM87
  Gnathoxys Westwood 1842 LM87
    |--G. barbatus Macleay 1864 LM87
    |--G. cicatricosus Reiche 1842 LM87
    |--G. crassipes Sloane 1898 LM87
    |--G. foveatus Macleay 1866 LM87
    |--G. granularis Westwood 1842 [incl. G. blissii Macleay 1866] LM87
    |--G. humeralis M80
    |--G. insignitus Macleay 1864 LM87
    |--G. irregularis Westwood 1842 LM87
    |--G. macleayi Putzeys 1868 LM87
    |--G. murrumbidgensis Macleay 1865 LM87
    |--G. obscurus Reiche 1842 LM87
    |--G. punctipennis Macleay 1873 LM87
    |--G. submetallicus Macleay 1864 LM87
    |--G. sulcicollis Sloane 1910 LM87
    |--G. tesselatus Macleay 1864 LM87
    `--G. westwoodi Putzeys 1868 LM87
  Chylnus Sloane 1920 [=Lychnus Putzeys 1868 non Matheson 1832] LM87
    `--*C. ater (Putzeys 1868) (see below for synonymy) LM87
  Percolestus Sloane 1892 LM87
    `--*P. blackburni Sloane 1892 LM87
  Percosoma Schaum 1858 LM87
    |--*P. carenoides (White 1846) (see below for synonymy) LM87
    |--P. concolor Sloane 1892 LM87
    |--P. montanum (Castelnau 1867) [=Mecodema montanum] LM87
    |--P. substriatum Moore 1960 LM87
    `--P. sulcipenne Bates 1878 LM87

*Chylnus ater (Putzeys 1868) [=*Lychnus ater; incl. L. strangulatus Bates 1878, L. striatulus Bates 1878] LM87

*Percosoma carenoides (White 1846) [=Broscus carenoides; incl. B. hopei Sturm 1843 (n. n.), Mecodema percoides Castelnau 1867] LM87

*Type species of generic name indicated

REFERENCES

[B89] Bates, H. W. 1889. Viaggio di Leonardo Fea in Birmani e regioni vicine. XVI.—On some Carabidae from Burma collected by Mr. L. Fea. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 100–111.

[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.

[M80] Matthews, E. G. 1980. A Guide to the Genera of Beetles of South Australia. Part 1. Archostemata and Adephaga. South Australian Museum: Australia.

[R-JC01] Roig-Juñent, S., & A. C. Cicchino. 2001. Chaltenia patagonica, new genus and species belonging to Chalteniina, a new subtribe of Zolini (Coleoptera: Carabidae). Canadian Entomologist 133: 651–670.

Last updated: 3 November 2018.

Broscini

Broscus cephalotes, copyright Udo Schmidt.

Belongs within: Broscinae.
Contains: Mecodema, Promecoderus.

The Broscini are a group of large ground beetles with a pedunculate thorax, most diverse in the Australasian region (Britton 1949). The genus Broscus, species of which are relatively large and have two pairs of setae on the pronotum and eight or nine setae laterally on the elytra, is found in the Palaearctic region, though the species B. cephalotes has also been introduced to eastern Canada (Ball & Bousquet 2001).

Characters (from Britton 1949): Size large; colour dark brown or black, occasionally slightly aeneous; mandibles usually with scrobal seta; labrum with angles rounded, with six setae on front margin and row of small setae arising just beneath anterolateral margins; prothorax connected to mesothorax by peduncle formed by mesothoracic projection bearing scutellum; pronotum without setiferous puncture at posterior angle; border base of elytra not complete, elytra fused along suture, hind wings vestigial; anterior coxal cavities with single perforation; mesepimera not reaching border of middle coxal cavities; anterior tarsi not widened in male, without modified setae ventrally; male genitalia with parameres prolonged into long styles at apices, right-hand style bearing fringe of long setae, base of aedeagus closed on dorsal side so orifice is circular.

<==Broscini [Cnemacanthides] B49
    |--Axonya [Axonyina] R-JC01
    |    `--A. championi R-JC01
    `--+--Nothobroscus [Nothobroscina] R-JC01
       |    `--N. chilensis R-JC01
       `--Broscina R-JC01
            |--Zacotus LeConte 1869 BB01
            |    `--Z. matthewsii LeConte 1869 BB01
            |--Miscodera Eschscholtz 1830 BB01
            |    `--M. arctica (Paykull 1800) BB01
            |--Broscodera Lindroth 1961 BB01
            |    |--B. (Broscodera) insignis (Mannerheim 1852) BB01
            |    `--B. (Sinobrosculus Deuve 1990) BB01
            `--Broscus Panzer 1813 B49
                 |--B. cephalotes (Linnaeus 1758) B14
                 |--B. insularis B14
                 |--B. laevigatus G89
                 |--B. nobilis B14
                 |--B. politus B14
                 `--B. punctatus G89

Broscini incertae sedis:
  Rawlinsius R-JC01
  Broscodes R-JC01
  Diglymma Sharp 1886 [incl. Snofru Broun 1908] B49
    |--*D. clivinoides (Castelnau 1867) (see below for synonymy) B49
    |--D. castigatum Broun 1909 B49
    |--D. marginale Broun 1914 B49
    `--D. obtusum (Broun 1893) (see below for synonymy) B49
  Oregus Putzeys 1868 B49
    |--*O. aereus (White 1846) [=Promecoderus aereus] B49
    `--O. inaequalis (Castelnau 1867) [=Mecodema inaequale] B49
  Mecodema B49
  Metaglymma Bates 1867 [incl. Maoria Castelnau 1867] B49
    |--*M. monilifer Bates 1867 [incl. M. minor Broun 1905, M. rugipenne Broun 1893] B49
    |--M. aberrans Putzeys 1868 (see below for synonymy) B49
    `--M. tibiale (Castelnau 1867) (see below for synonymy) B49
  Brullea Castelnau 1867 B49
    `--*B. antarctica Castelnau 1867 B49
  Promecoderus B49
  Eurylychnus Bates 1891 B49, LM87
    |--E. dyschirioides (Castelnau 1867) LM87 [=Maoria dyschirioides B49; incl. *E. olliffi Bates 1891 LM87]
    |--E. blagravei (Castelnau 1867) [=Mecodema blagravei; incl. Percosoma mastersi Rainbow 1899] LM87
    |--E. cylindricus Sloane 1916 LM87
    |--E. femoralis Sloane 1915 LM87
    |--E. kershawi Sloane 1915 LM87
    |--E. ovipennis Sloane 1915 LM87
    |--E. regularis Sloane 1911 LM87
    `--E. victoriae Sloane 1892 LM87

*Diglymma clivinoides (Castelnau 1867) [=Maoria clivinoides; incl. D. basale Broun 1917, D. dubium Sharp 1886, D. ovipenne Sharp 1886] B49

Diglymma obtusum (Broun 1893) [=Metaglymma obtusum; incl. Snofru aemulator Broun 1908, Diglymma nigripes Broun 1893, D. punctipenne Broun 1893, D. tarsale Broun 1908, D. thoracicum Broun 1917] B49

Metaglymma aberrans Putzeys 1868 [incl. M. rugiceps Broun 1903, M. tersatum Broun 1893, M. thoracicum Broun 1893] B49

Metaglymma tibiale (Castelnau 1867) [=*Maoria tibialis; incl. Mecodema asperum Broun 1893, Mec. calcaratum Broun 1903, Mec. junctum Broun 1893, Mec. rufipes Broun 1886] B49

*Type species of generic name indicated

REFERENCES

[BB01] Ball, G. E., & Y. Bousquet. 2001. Carabidae Latreille, 1810. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 32–132. CRC Press: Boca Raton.

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

[B49] Britton, E. B. 1949. The Carabidae (Coleoptera) of New Zealand. Part III. A revision of the tribe Broscini. Transactions and Proceedings of the Royal Society of New Zealand 77 (4): 533–581.

[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.

[R-JC01] Roig-Juñent, S., & A. C. Cicchino. 2001. Chaltenia patagonica, new genus and species belonging to Chalteniina, a new subtribe of Zolini (Coleoptera: Carabidae). Canadian Entomologist 133: 651–670.

Last updated: 3 November 2018.

Brachininae

Pheropsophus verticalis, copyright Peter Halasz.

Belongs within: Conjunctae.

The Brachininae, bombadier beetles, are a group of ground beetles capable of explosively emitting a defensive spray from the abdominal apex. They are most diverse in the tropics, particularly in Africa. Larvae of at least some species are ectoparasitoids of other insects with Brachinus attacking pupae of water beetles and Pheropsophus feeding on eggs of mole crickets (Ball & Bousquet 2001).

Characters (from Ball & Bousquet 2001): Size small to large (total length ca. 2–30 mm); head capsule ventrally without antennal grooves; antennal insertions laterad, each between base of mandible and anterior margin of eye; antenna filiform, number of antennomeres 11; antennomeres 2–4 not nodose, 2–6 without unusually long setae; labrum with six to eight setae; mandible with scrobe unisetose or plurisetose, occlusal margin with one terebral tooth, retinacular teeth about size of terebral tooth, not enlarged and molariform; terminal palpomeres about cylindrical not securiform; pterothorax with mesonotal scutellum dorsally visible, not concealed beneath elytra; front coxal cavities closed, with projection of proepimeron inserted each side into cavity of prosternum; middle coxal cavities conjunct-confluent, and hind coxal cavities lobate (disjunct)-confluent; elytra striate (intervals and striae broad) to smooth dorsally, extended to apex of abdominal tergum VI, and apical margins truncate; without flange of Coanda; metathoracic wings large to variously reduced; front tibia anisochaete, antenna cleaner emarginate, type C; abdomen with seven (II–VIII) or eight (II–IX) sterna normally exposed, sterna III and IV separated by suture; male genitalia with parameres glabrous, reduced, right minute, left larger, and in adults of most species baldric-like in position, across basal part of median lobe; ovipositor with gonocoxae rotated 90° in repose, without ramus, gonocoxa with two gonocoxites, gonocoxite 2 without accessory lobe; reproductive tract with spermatheca 1 only; secretion of pygidial glands quinones, delivered hot; discharge by crepitation.

<==Brachininae [Brachinides, Brachinini] B12
    |  i. s.: Mastax B55
    |           |--M. burgeoni B55
    |           `--M. ornatellus B55
    |         Brachynillus varendorffi Reitter 1904 BB01
    |--+--Aptinus OM08
    |  |    |--A. alpinus OM08
    |  |    `--A. displosor OM08
    |  `--+--Styphlodromus OM08
    |     `--Styphlomerus OM08
    |          |--S. ludicrus [incl. S. ludicrus var. grandis] B55
    |          `--S. vittaticollis OM08
    `--+--Crepidogaster [Crepidigastrini] OM08
       |    |--C. (Tyronia) ambreana OM08
       |    |--C. atrata OM08
       |    `--C. picipennis B55
       `--+--Brachinus Weber 1801 OM08, BB01
          |    |  i. s.: B. bimaculatus B35
          |    |         B. caligatus Bates 1889 B89
          |    |         B. frontalis B89
          |    |         B. oblongus G89
          |    |--B. (Brachinus) crepitans G20
          |    |--B. (Metabrachinus) armiger OM08 [incl. Metabrachinus armiger var. natalicus B55]
          |    `--B. (Neobrachinus Erwin 1970) BB01
          |         `--B. (N.) hirsutus OM08
          `--Pheropsophus Solier 1833 [Pherapsophina] B12
               |  i. s.: P. aequinoctialis (Linnaeus 1763) B14
               |         P. africanus [=Brachinus africanus] G89
               |         P. alexandrae Giachino 2003 B12
               |         P. arcanus B55
               |         P. darwini Giachino 2003 B12
               |         P. dimidiatus B55
               |         P. gregoryi Giachino 2003 B12
               |         P. jessoensis LB91
               |         P. livingstonei B55
               |         P. macleayi Sloane 1895 B12
               |         P. minor B55
               |         P. straneoi Giachino 2003 B12
               |         P. windjanae Baehr 2012 B12
               |--P. (Pheropsophus) B12
               |    |--*P. (P.) madagascariensis (Dejean 1831) [=Brachinus madagascariensis] B12
               |    `--P. (P.) complanatus (Fabricius 1775) LM87 [=Carabus complanatus LM87, Nebria complanata L02]
               `--P. (Stenaptinus Maindron 1906) [incl. Parapheropsophus Hubenthal 1914] LM87
                    |--P. (*S.) krichna (Maindron 1914) [=*Stenaptinus krichna] LM87
                    |--‘Stenaptinus’ insignis SBM11
                    `--P. (S.) verticalis (Dejean 1825) LM87 (see below for synonymy) LM87

Pheropsophus (Stenaptinus) verticalis (Dejean 1825) LM87 [=Brachinus verticalis LM87, P. (*Parapheropsophus) verticalis LM87; incl. P. australis Castelnau 1867 LM87, P. baliothorax Heller 1910 LM87, P. hypoxanthus Chaudoir 1876 LM87, P. (Parapheropsophus) intermedius Hubenthal 1914 LM87, Aptinus jamissoni B35, P. (Pa.) niger Hubenthal 1914 LM87, P. papuensis Macleay 1877 LM87]

*Type species of generic name indicated

REFERENCES

[B12] Baehr, M. 2012. A new species of the genus Pheropsophus (Coleoptera: Carabidae: Brachininae) from northern Australia. Records of the Western Australian Museum 27 (1): 62–67.

[BB01] Ball, G. E., & Y. Bousquet. 2001. Carabidae Latreille, 1810. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 32–132. CRC Press: Boca Raton.

[B55] Basilewsky, P. 1955. Carabidae (Coleoptera) de l’Angola (première partie). Diamang Publicações Culturais 27: 91–137.

[B89] Bates, H. W. 1889. Viaggio di Leonardo Fea in Birmani e regioni vicine. XVI.—On some Carabidae from Burma collected by Mr. L. Fea. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 100–111.

[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).

[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.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

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

[LB91] Lawrence, J. F., & E. B. Britton. 1991. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 543–683. Melbourne University Press: Carlton (Victoria).

[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.

[OM08] Ober, K. A., & D. R. Maddison. 2008. Phylogenetic relationships of tribes within Harpalinae (Coleoptera: Carabidae) as inferred from 28S ribosomal DNA and the wingless gene. Journal of Insect Science 8: 63.

[SBM11] Solomon, E. P., L. R. Berg & D. W. Martin (eds) 2011. Biology 9^th ed. Brooks/Cole Cengage Learning.

Last updated: 19 December 2020.

Craspedophorus

Craspedophorus sublaevis, copyright Peter Geissler.

Belongs within: Panagaeitae.

Craspedophorus is a genus of ground beetles widespread in the Palaeotropics, species of which have a distinctive colour pattern of four bright spots on the elytra and are often flightless. This coloration reflects the beetle's glandular production of defensive chemicals that are held in large reservoirs in the abdomen.

Craspedophorus Hope 1838 [incl. Epicosmus Chaudoir 1846, Eudema Castelnau 1840, Isotarsus LaFerté-Sénectère 1851] LM87
    |--*C. reflexus (Fabricius 1781) [=Carabus reflexus] LM87
    |--C. alternans (Castelnau 1867) [=Eudema alternans] LM87
    |--C. angulatus (Fabricius 1781) B14
    |--C. angusticeps Sloane 1923 LM87
    |--C. australasiae (Chaudoir 1850) [=Epicosmus australasiae; incl. E. froggatti Sloane 1903 LM87
    |--C. australis (Dejean 1831) [=Panagaeus australis; incl. Epicosmus mastersi Sloane 1903] LM87
    |--C. banksi Sloane 1923 LM87
    |--C. clasispila B55
    |--C. comptus (LaFerté-Sénectère 1851) [=Isotarsus comptus; incl. Epicosmus corpulentus Chaudoir 1879] LM87
    |--C. elongatus (Castelnau 1867) [=Eudema elongata] LM87
    |--‘Epicosmus’ feae Bates 1889 B89
    |--C. hexagonus B14
    |--C. impictus B55
    |--C. insignis (Schaum 1853) [=Isotarsus insignis; incl. C. convexus Macleay 1864] LM87
    |--C. longicollis (Chaudoir 1879) [=Epicosmus longicollis] LM87
    |--C. macleayi (Sloane 1903) [=Epicosmus macleayi, E. nobile Macleay 1888 non Dejean 1826] LM87
    |--C. ornatus B55
    |--C. parvulus (Macleay 1888) [=Eudema parvulum] LM87
    |--C. pubiger B14
    |--‘Panageus’ regalis Gory 1833 [=*Eudema regale, *Isotarsus regalis] LM87
    |--C. rikatlensis B55
    |--C. rockhamptonensis (Castelnau 1867) [=Eudema rockhamptonensis; incl. Epicosmus obesulus Sloane 1903] LM87
    |--C. rufipalpis OM08
    |--C. sublaevis B14
    |--‘Panageus’ tormentosus Vigors 1825 [=*Epicosmus tormentosus] LM87
    `--‘Epicosmus’ transversalis B89

*Type species of generic name indicated

REFERENCES

[B55] Basilewsky, P. 1955. Carabidae (Coleoptera) de l’Angola (première partie). Diamang Publicações Culturais 27: 91–137.

[B89] Bates, H. W. 1889. Viaggio di Leonardo Fea in Birmani e regioni vicine. XVI.—On some Carabidae from Burma collected by Mr. L. Fea. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 100–111.

[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.

[OM08] Ober, K. A., & D. R. Maddison. 2008. Phylogenetic relationships of tribes within Harpalinae (Coleoptera: Carabidae) as inferred from 28S ribosomal DNA and the wingless gene. Journal of Insect Science 8: 63.

Last updated: 3 November 2018.

Chlaenius

Chlaenius festivus, copyright Siga.

Belongs within: Oodini.

Chlaenius is a cosmopolitan genus of often metallic ground beetles with pubescent elytra that have the epipleura interrupted by a twisted fold near the apex. Members of the genus have been divided between over fifty subgenera and monophyly of the genus perhaps warrants investigation.

<==Chlaenius Bonelli 1810 [Chlaeniina] B12
    |--C. (Chlaenius) (see below for synonymy) BB01
    |    |--*C. (C.) festivus (Fabricius 1810) [=Carabus festivus] LM87
    |    |--C. (C.) aestivus Say 1823 BB01
    |    |--C. (C.) augustus Newman 1838 BB01
    |    |--C. (C.) cordicollis Kirby 1837 BB1
    |    |--C. (C.) cursor BB01
    |    |--C. (C.) laticollis BB01
    |    |--C. (C.) platyderus Chaudoir 1856 BB01
    |    |--C. (C.) ruficauda BB01
    |    |--C. (C.) sericeus (Forster 1771) BB01
    |    |--C. (C.) solitarius BB01
    |    `--C. (C.) viduus Horn 1871 BB01
    |--C. (Agostenus Motschulsky 1850) (see below for synonymy) BB01
    |    |--‘Agostenus’ glaucus BB01
    |    |--C. (A.) impunctifrons Say 1823 BB01
    |    |--‘Agostenus’ lithophilus BB01
    |    |--‘Agostenus’ nemoralis BB01
    |    |--‘Agostenus’ pennsylvanicus BB01
    |    |--C. (A.) purpuricollis Randall 1838 BB01
    |    |--C. (A.) tricolor Dejean 1826 BB01
    |    |--‘Agostenus’ vafer BB01
    |    `--‘Agostenus’ variabilipes BB01
    |--C. (Anomoglossus Chaudoir 1856) BB01
    |--C. (Diapheropsophus) B66
    |    |--C. (D.) mellyi B66
    |    `--C. (D.) swinhoei Bates 1866 B66
    |--C. (Epomis) B14
    |    |--C. (E.) circumscriptus (Duftschmid 1812) B14
    |    `--C. (E.) dejeanii B14
    |--C. (Eurydactylus LaFerté Sénectère 1851) [incl. Glyptoderus LaFerté Sénectère 1851] BB01
    |    |--C. (E.) pimalicus Casey 1914 BB01
    |    `--C. (E.) tomentosus (Say 1823) BB01
    |--C. (Lissauchenius) rufifemoralis OM08
    |    |--C. r. rufifemoralis OM08
    |    `--C. r. bimaculatus Dejean 1826 OM08, LM87
    |--C. (Pelasmomimus Grundmann 1955) LM87
    |    `--C. (*P.) greyanus White 1841 [=C. grayianus] LM87
    `--C. (Pseudanomoglossus Bell 1960) BB01
         `--C. (P.) maxillosus Horn 1876 BB01

Chlaenius incertae sedis:
  C. australis Dejean 1831 LM87
  C. darlingensis Castelnau 1867 LM87
  C. flaviguttatus Macleay 1825 (see below for synonymy) LM87
  C. hamifer Chaudoir 1856 [incl. C. bihamatus Chaudoir 1856, C. queenslandicus Sloane 1910] LM87
  C. maculiger Castelnau 1867 LM87
  C. nigricornis [incl. C. nigricornis var. melanocornis] C01
  C. nigripes Macleay 1886 M86
  C. ophonoides Fairmaire 1843 (see below for synonymy) LM87
  C. planicornis B66
  C. rudicollis Chaudoir 1876 [incl. C. bimaculatus Macleay 1864 non Dejean 1826] LM87
  C. spoliatus C01
  C. subcostatus Macleay 1864 LM87
  C. tenellus G89
  C. virgulifer C01

Chlaenius flaviguttatus Macleay 1825 [incl. Lissauchenius biguttatus Montrouzier 1860, C. binotatus Dejean 1826, C. guttatus Eschscholtz 1833, C. flaviguttatus immaculipennis Jedlicka 1951, C. maculifer Castelnau 1867, C. punctatus Chaudoir 1856, C. puncticeps Gemminger & Harold 1868] LM87

Chlaenius ophonoides Fairmaire 1843 [incl. C. peregrinus Chaudoir 1856, C. viridescens Gemminger & Harold 1868, C. viridis Montrouzier 1860 non Ménétries 1832] LM87

Chlaenius (Agostenus Motschulsky 1850) [inc. Brachylobus Chaudoir 1876, Chlaeniellus Reitter 1908, Pelasmus Motschulsky 1850] BB01

Chlaenius (Chlaenius) [incl. Callistometus Grundmann 1956, Hemichlaenius Lutschnik 1933 (preoc.), Lithochlaenius Kryzhanovskij 1976] BB01

*Type species of generic name indicated

REFERENCES

[B12] Baehr, M. 2012. A new species of the genus Pheropsophus (Coleoptera: Carabidae: Brachininae) from northern Australia. Records of the Western Australian Museum 27 (1): 62–67.

[BB01] Ball, G. E., & Y. Bousquet. 2001. Carabidae Latreille, 1810. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 32–132. CRC Press: Boca Raton.

[B66] Bates, H. W. 1866. On a collection of Coleoptera from Formosa sent home by R. Swinhoe, Esq., H. B. M. Consul, Formosa. Proceedings of the Zoological Society of London 1866: 339–355.

[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.

[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.

[OM08] Ober, K. A., & D. R. Maddison. 2008. Phylogenetic relationships of tribes within Harpalinae (Coleoptera: Carabidae) as inferred from 28S ribosomal DNA and the wingless gene. Journal of Insect Science 8: 63.

Last updated: 3 November 2018.

Zuphiitae

Zuphium olens, copyright Amy.

Belongs within: Harpalinae.
Contains: Galeritina, Gigadema, Planetina, Helluomorphoides, Dryptini.

The Zuphiitae are a group of carabid ground beetles whose members possess spination on the first stylomere of the ovipositor (Ober & Maddison 2008). They have been divided between several tribes; however, Ober & Maddison's (2008) phylogenetic analysis did not support monophyly of the tribes Zuphiini, Galeritini and Helluonini. Members of the Anthiini and Helluonina are large beetles with an enlarged labrum covering most of the mandibles in dorsal view. Species of the genus Anthia are capable of ejecting a spray of defensive chemicals from glands in the abdomen up to a distance of a metre. Members of the Zuphiini are characterised by truncate elytra and uniformly setose antennae with the first antennomere longer than the following two together. The genus Zuphium, with the head markedly constricted behind the eyes to form a 'neck', has a cosmopolitan distribution concentrated in the tropics.

See also: Ground beetles for today.

<==Zuphiitae [Anthiitae, Dryptitae] OM08
    |--Colasidia [Leleupidiina, Leleupidiini] B86
    |    `--C. monteithi Baehr 1988 Ba14b
    |--Anthiini OM08
    |    |--Cypholoba [Cypholobina] OM08
    |    `--Anthia [Anthiina] OM08
    |         |--A. cinctipennis B55
    |         |    |--A. c. cinctipennis B55
    |         |    `--A. c. actaeon B55
    |         |--A. circumscripta B55
    |         |    |--A. c. circumscripta B55
    |         |    `--A. c. omostigma [incl. A. niveicincta] B55
    |         |--A. maxillosa Bo14
    |         |--A. sexguttata L02
    |         `--A. thoracica (Thunberg 1784) Bo14
    `--Zuphiini (see below for synonymy) OM08
         |  i. s.: Progalerita B86
         |         Eunostus B86
         |         Meladroma lugubre B55
         |         Triaenogenius B55
         |           |--T. anthioides (Chaud. 1872) [=Acanthogenius anthioides] G89
         |           |--T. arabicus Gestro 1889 G89
         |           |--T. carinulatus (Fairm. 1887) [=Acanthogenius carinulatus] G89
         |           |--T. corpulentus Chaud. 1877 G89
         |           |--T. ferox (Erichs. 1843) [=Helluo ferox] G89
         |           |--T. helluonoides (Ancey 1881) [=Anthia helluonoides] G89
         |           `--T. sculpturatus (Gerst. 1866) [=Acanthogenius sculpturatus] G89
         |--+--Galeritina OM08
         |  `--+--+--Omphra OM08
         |     |  `--Macrocheilus Kirby 1838 OM08, BR05
         |     |       |--M. bimaculatus B55
         |     |       `--M. biplagiatus B55
         |     `--+--Acrogenys Macleay 1864 OM08, Ba14b [Dicrodontina]
         |        |    |  i. s.: A. centralis GWW10
         |        |    |--A. (Acrogenys) LM87
         |        |    |    |--*A. (A.) hirsuta Macleay 1864 LM87
         |        |    |    |--A. (A.) demarzi Baehr 1984 LM87
         |        |    |    `--A. (A.) laticollis Baehr 1984 LM87
         |        |    `--A. (Paracrogenys Baehr 1984) LM87
         |        |         `--A. (*P.) longicollis Gestro 1875 LM87
         |        `--Helluonina OM08
         |             |  i. s.: Helluo Bonelli 1813 LM87
         |             |           |--*H. costatus Bonelli 1813 [incl. H. carinatus Chaudoir 1848] LM87
         |             |           `--H. insignis Sloane 1890 LM87
         |             |--Dicranoglossus Chaudoir 1872 OM08, LM87
         |             |    `--*D. resplendens (Castelnau 1867) (see below for synonymy) LM87
         |             `--+--Gigadema OM08
         |                `--Aenigma Newman 1836 OM08, LM87
         |                     `--*A. iris Newman 1836 (see below for synonymy) LM87
         `--+--+--Thalpius LeConte 1851 OM08, LM87 (see below for synonymy)
            |  |    |--*T. pygmaeus (Dejean 1826) [=Helluo pygmaeus, Pseudaptinus (*Thalpius) pygmaeus] LM87
            |  |    |--T. australis (Blackburn 1889) [=Acrogenys australis, Pseudaptinus (Thalpius) australis] LM87
            |  |    `--T. fulvus (Castelnau 1867) [=*Zuphiosoma fulva, Pseudaptinus (Thalpius) fulvus] LM87
            |  `--+--Planetina OM08
            |     `--+--Helluomorphoides OM08
            |        `--+--‘Thalpius’ rufulus OM08
            |           `--Pseudaptinus Castelnau 1834 OM08, LM87 (see below for synonymy)
            |                |--*P. albicornis Castelnau 1834 LM87
            |                |--P. brittoni Baehr 1985 B86
            |                |--P. hirsutulus Baehr 1985 B86
            |                |--P. iridescens Baehr 1985 B86
            |                |--P. lecontei (Dejean 1831) BB01
            |                |--P. monteithi Baehr 1985 B86
            |                `--P. tenuicollis (LeConte 1851) BB01
            `--+--+--Dryptini OM08
               |  `--Pogonoglossus Chaudoir 1862 OM08, LM87 (see below for synonymy)
               |       |--*P. validicornis Chaudoir 1862 LM87
               |       |--P. danielsi Baehr 2008 Ba14a
               |       |--P. frater Baehr 2014 Ba14a
               |       |--P. inarmatus Baehr 1988 Ba14a
               |       |--P. inflaticeps (Sloane 1904) [=Carpaulum inflaticeps] LM87
               |       |--P. parvus Darlington 1968 Ba14a
               |       |--P. porosus (Sloane 1904) [=Carpaulum porosum] LM87
               |       |--P. rufopiceus Baehr 1993 Ba14a
               |       `--P. soror Baehr 2014 Ba14a
               `--+--Catapiesini BS02
                  |    |--Catapiesis brasiliensis OM08
                  |    `--Homalomorpha BS02
                  `--+--Mischocephalus [Mischocephalina] OM08
                     `--Zuphium Latreille 1806 OM08, LM87 [=Zophium Gistel 1839 BB01, Zoyphium Agassiz 1847 BM76]
                          |--*Z. olens (Rossi 1790) [=Carabus olens] LM87
                          |--Z. australe Chaudoir 1862 LM87
                          |--Z. castelnaui Gestro 1875 LM87
                          |--Z. fitzroyense Macleay 1888 LM87
                          |--Z. flavum GWW10
                          |--Z. numidicum Lucas 1846 E12
                          |--Z. pindan Macleay 1888 LM87
                          |--Z. rockhamptonense Castelnau 18967 [=Z. rockhamptonensis] LM87
                          `--Z. thouzeti Castelnau 1867 B86

Zuphiitae incertae sedis:
  Helluarchus Sloane 1914 M80, LM87
    |--*H. robustus Sloane 1914 LM87
    `--H. whitei Lea 1914 LM87
  Speozuphium Moore 1995 Ba14b
  Speothalpius Moore 1995 Ba14b
    `--S. grayi Ba14b
  Typhlozuphium Baehr 2014 Ba14b
    |--*T. humicolum Baehr 2014 Ba14b
    `--T. longipenne Baehr 2014 Ba14b
  Parazuphium Jeannel 1942 B86, Ba14b
    |--*P. (Parazuphium) chevrolati (Castelnau 1833) [=Zuphium chevrolati] Ba14b
    `--P. (Austrozuphium Baehr 1985) Ba14b
         |--P. (*A.) mastersii (Castelnau 1867) [=Zuphium mastersii] Ba14b
         |--P. (A.) flavescens Baehr 1985 Ba14b
         `--P. (A.) pilbarae Baehr 1985 Ba14b

*Aenigma iris Newman 1836 [incl. A. newmanni Castelnau 1867, A. splendens Castelnau 1867] LM87

*Dicranoglossus resplendens (Castelnau 1867) [=Helluosoma resplendens; incl. Aenigma parvulum Macleay 1874] LM87

Pogonoglossus Chaudoir 1862 OM08, LM87 [incl. Carpaulum Sloane 1904 LM87; Physocrotaphinae, Physocrotaphini LB91]

Pseudaptinus Castelnau 1834 OM08, LM87 [incl. Diaphorus Dejean 1831 non Meigen 1824 BB01, Tiphys Gistel 1848 (preoc.) BB01]

Thalpius LeConte 1851 OM08, LM87 [incl. Enaphorus LeConte 1851 BB01, Zuphiosoma Laporte de Castelnau 1867 BB01]

Zuphiini [Galeritini, Galeritinini, Galeritulini, Helluomorphina, Helluonides, Helluoninae, Helluonini, Omphrina, Patriziina, Zuphiina, Zuphiinae] OM08

*Type species of generic name indicated

REFERENCES

[B86] Baehr, M. 1986. Revision of the Australian Zuphiinae. 6. The genus Planetes Macleay. Supplement to the other genera (Insecta, Coleoptera, Carabidae). Spixiana 9 (2): 151–168.

[Ba14a] Baehr, M. 2014a. Two new species of the beetle genus Pogonoglossus from the Pilbara region of north-western Australia (Coleoptera: Carabidae: Physocrotaphini). Records of the Western Australian Museum 29 (2): 89–94.

[Ba14b] Baehr, M. 2014b. Three new species and a new genus of subterranean Zuphiini from the Pilbara region of north-western Australia (Coleoptera: Carabidae: Harpalinae). Records of the Western Australian Museum 95–104.

[BB01] Ball, G. E., & Y. Bousquet. 2001. Carabidae Latreille, 1810. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 32–132. CRC Press: Boca Raton.

[BS02] Ball, G. E., & D. Shpeley. 2002. Ginemini, Ginema thomasi, new tribe, new genus and new species, from Amazonian Bolivia (Coleoptera: Carabidae: Harpalinae). Transactions of the American Entomological Society 128 (1): 75–98.

[B55] Basilewsky, P. 1955. Carabidae (Coleoptera) de l’Angola (première partie). Diamang Publicações Culturais 27: 91–137.

[BM76] Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

[Bo14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[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.

[GWW10] Guthrie, N. A., T. Weir & K. Will. 2010. Localised and regional patterns in ground-dwelling beetle assemblages in a semi-tropical arid zone environment. Records of the Western Australian Museum Supplement 78 (1): 169–184.

[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.

[M80] Matthews, E. G. 1980. A Guide to the Genera of Beetles of South Australia. Part 1. Archostemata and Adephaga. South Australian Museum: Australia.

[OM08] Ober, K. A., & D. R. Maddison. 2008. Phylogenetic relationships of tribes within Harpalinae (Coleoptera: Carabidae) as inferred from 28S ribosomal DNA and the wingless gene. Journal of Insect Science 8: 63.

Last updated: 3 November 2018.

Cicindelinae

Mating pair of green tiger beetles Cicindela campestris, copyright Ian Alexander.

Belongs within: Conjunctae.
Contains: Distipsidera, Megacephala (Pseudotetracha), Cicindela (Myriochile), Cicindela (Rivacindela).

The Cicindelinae include the tiger beetles, fast-moving and often brightly coloured predatory beetles. They have commonly been treated in the past as a distinct family, but phylogenetic studies have agreed that they are nested within the ground beetles of the Carabidae. The greater number of species are included in the genus Cicindela, characterised by an anterior pronotal sulcus continuous with the anterior prosternal sulcus, and anterior pronotal angles that are not more prominent than the prosternal margin. Members of the tribes Omini and Megacephalini have the anterior pronotal and prosternal sulci separated, and the anterior pronotal angles exceed the prosternal margin (Ball & Bousquet 2001).

Characters (from Ball & Bousquet 2001): Head capsule ventrally without antennal grooves; antennal insertions mediad, above and closer together than bases of mandibles; antenna filiform, antennomeres 2–4 not nodose, 2–6 without unusually long setae, with 11 antennomeres; labrum with six to twelve setae; mandible with scrobe multisetose or asetose, occlusal margin with two to four terebral teeth, retinacular tooth very large, molariform; terminal palpomeres about cylindrical; pterothorax with mesonotal scutellum dorsally visible, not concealed beneath elytra; front coxal cavities open, middle and hind coxal cavities disjunct-confluent; elytra smooth, not striate dorsally, extended to apex of abdominal tergum VIII, apical margins subsinuate; metathoracic wings large to variously reduced; front tibia anisochaete, antenna cleaner sulcate, type A; abdomen with six (II–VII) pregenital sterna normally exposed, sterna III and IV separated by suture; male genitalia with parameres but similar to one another in form (slender) and in length, asetose, joined medio-dorsally to narrow jugum; ovipositor with gonocoxae rotated in retracted position, with or without ramus, gonocoxa with two gonocoxites, and gonocoxite 2 with accessory lobe, not articulated or articulated; reproductive tract with spermatheca 2 only; secretions of pygidial glands aromatic aldehydes, discharged by oozing. Larva with large prothorax with pronotum semicircular; abdominal tergum V with two or three pairs of sclerotized, hook-like projections; abdominal segment IX without urogomphi.

<==Cicindelinae [Collyridae, Collyridinae, Ctenostomidae, Mantichoridae, Megacephalinae]
    |--Manticora B14 [Manticorini BB01]
    |    |--M. latipennis Waterhouse 1837 B14
    |    `--M. maxillosa L02
    |--Omini BB01
    |    |--Omus Eschscholtz 1829 [incl. Leptomus Casey 1914, Megomus Casey 1914] BB01
    |    `--Amblycheila Say 1830 BB01
    |         |--A. cylindriformis (Say 1823) B14
    |         `--A. hoversoni B14
    |--Megacephalini B14
    |    |--Picnochile Motschulsky 1856 B14, BB01
    |    |    `--P. fallaciosa (Chevrolat 1854) B14
    |    |--Tetracha Hope 1838 LM87
    |    |    `--*T. carolina (Linnaeus 1767) LM87, B14 [=Cicindela carolina LM87, Megacephala carolina L02]
    |    `--Megacephala Latreille 1802 LM87
    |         |  i. s.: M. affinis [incl. M. affinis f. angustata Chevrolat 1841] F54
    |         |         M. australasiae Hope 1842 (see below for synonymy) LM87
    |         |         M. howittii Castelnau 1867 LM87
    |         |         M. kimberleyensis Mjöberg 1916 LM87
    |         |         M. mediopilosa Horn 1934 LM87
    |         |         M. murchisona (Fleutiaux 1896) [=Tetracha murchisona] LM87
    |         |           |--M. m. murchisona LM87 (see below for synonymy)
    |         |           `--M. m. elongata Horn 1934 LM87
    |         |         M. scapularis Macleay 1863 [=M. (Tetracha) scapularis] LM87
    |         |         M. sobrina [incl. M. sobrina f. ignea Bates 1878] F54
    |         |         M. spenceri Sloane 1897 LM87
    |         |         M. virginica BB68
    |         |--*M. (Megacephala) megacephala (Olivier 1790) [=Cicindela megacephala] LM87
    |         |--M. (Australicapitona Sumlin 1992) G07
    |         |    |--M. (A.) basalis (Macleay 1866) G07 (see below for synonymy)
    |         |    |--M. (A.) bostockii (Castelnau 1867) G07, LM87 (see below for synonymy)
    |         |    |--M. (A.) crucigera Macleay 1863 G07 [=M. (Tetracha) crucigera LM87]
    |         |    |--M. (A.) hopei (Castelnau 1867) G07 [=Tetracha hopei LM87]
    |         |    |--M. (A.) intermedia Sloane 1906 G07 [incl. M. sloanei Horn 1920 LM87]
    |         |    `--M. (A.) karratha Sumlin 1992 G07
    |         `--M. (Pseudotetracha) G07
    `--Cicindelini BB01
         |--Oxycheila BB01
         |--Cheiloxya BB01
         `--Cicindela Linnaeus 1758 LM87
              |  i. s.: C. aeneicollis Bates 1881 F54
              |         C. anchoralis Chevrolat 1845 I92
              |         C. andersonii Gestro 1889 G89b
              |         C. aquatica Linnaeus 1758 L58
              |         C. argentata B66
              |         C. aulica G89a
              |         C. auricollis G89b
              |         C. aurulenta [incl. C. flavomaculata Moore 1878] G89b
              |         C. bonina Nakane & Kurosawa 1959 I92
              |         C. burmeisteri C01
              |         C. catena G89b
              |         C. chloris Hope 1831 [incl. C. himalayica Moore 1831] G89b
              |         C. chlorochila Chaud. 1852 G89b
              |         C. craspedota B66
              |         C. cuprascens VD94
              |         C. cyaniventris [incl. C. cyaniventris f. papillosa Chaudoir 1854] F54
              |         C. curvata VD94
              |         C. davisonii Gestro 1889 G89b
              |         C. decemguttata B35
              |         C. dorsalis VD94
              |           |--C. d. dorsalis VD94
              |           `--C. d. media VD94
              |         C. durvillei B35
              |         C. elisae I92
              |         C. eugenia Chaud. 1865 G89b
              |         C. exornata G89b
              |         C. feae Gestro 1889 G89b
              |         C. flavopunctata [incl. C. flavopunctata f. rectilatera Chaudoir 1843] F54
              |         C. foveolata Schaum. 1863 G89b
              |         C. fuliginosa G89b
              |         C. fulvipes B35
              |         C. funerata Boisduval 1835 M86
              |         C. gracilis [incl. C. gracilis var. angustata] C01
              |         C. guttula B35
              |         C. hamata VD94
              |         C. helmsi H15
              |         C. heros B35
              |         C. hybrida Linnaeus 1758 L58 (see below for synonymy)
              |         C. immanis Bates 1874 G89a
              |         C. inspecularis Horn 1904 I92
              |         C. interruptofasciata G89b
              |         C. kaleela Bates 1866 B66
              |         C. latecincta F02
              |         C. latreillii [=C. latreilliana Boisduval 1835] B35
              |         C. lewisi Bates 1873 I92
              |         C. limbata C90
              |           |--C. l. limbata C90
              |           `--C. l. albissima C90
              |         C. limosa G89b
              |         C. lineifrons Chaud. 1865 [=C. lineicollis (l. c.)] G89b
              |         C. longicornis F54
              |         C. macra VD94
              |         C. macrocnema Chaudoir 1852 F54
              |         C. marginata VD94
              |         C. maura Linnaeus 1758 L58
              |         C. minuta [incl. C. pumila] G89b
              |         C. niveicincta B66
              |         C. octonotata G89b
              |         C. patruela B14
              |         C. peletieri Lucas 1846 E12
              |         C. psilica Bates 1866 B66
              |         C. punctatissima B66
              |         C. punctulata BB68
              |         C. puritana VD94
              |         C. quadripunctata [=Euryoda quadripunctata] G89b
              |         C. riparia Linnaeus 1758 L58
              |         C. roseiventris Chevrolat 1834 F54
              |         C. scalaris B35
              |         C. scutellaris B14
              |         C. sexguttata Fabricius 1775 B14
              |         C. sexpunctata G89b
              |         C. spinolae Gestro 1889 G89b
              |         C. striolata Illig. 1800 [incl. C. semivittata] G89b
              |         C. sumatrensis Herbst 1806 G89b
              |           |--C. s. sumatrensis I92
              |           `--C. s. niponensis Bates 1883 I92
              |         C. sylvatica Linnaeus 1758 TKM93 [incl. C. silvatica var. fennica C01]
              |         C. transbaikalica C01
              |         C. tricolor C01
              |         C. trifasciata I92
              |         C. tritoma G89b
              |         C. tuberculata B35
              |         C. vasseleti Chevrolat 1834 F54
              |         C. viridilabris G89b
              |         C. yuasai I92
              |--*C. (Cicindela) campestris Linnaeus 1758 LM87 [incl. C. campestris ab. desertorum C01]
              |--C. (Antennaria Dokhtouroff 1883) LM87
              |    |--C. (A.) ioscelis Hope 1841 (see below for synonymy) LM87
              |    |--C. (A.) crassicornis Macleay 1888 LM87
              |    |--C. (A.) doddi Sloane 1905 [incl. C. doddi semiviridis Sloane 1906] LM87
              |    `--C. (A.) sparsimpilosa Horn 1913 LM87
              |--C. (Archidela Rivalier 1963) LM87
              |    |--C. (*A.) nigrina Macleay 1863 (see below for synonymy) LM87
              |    `--C. (A.) darwini Sloane 1909 LM87
              |--C. (Cylindera Westwood 1831) LM87
              |    |--C. (*C.) germanica Linnaeus 1758 LM87
              |    `--C. (C.) discreta Schaum 1863 (see below for synonymy) LM87
              |--C. (Euzona Rivalier 1963) LM87
              |    |--C. (*E.) tetragramma Boisduval 1835 (see below for synonymy) LM87
              |    |--C. (E.) aeneodorsis Sloane 1917 LM87
              |    |--C. (E.) albolineata Macleay 1888 [incl. C. queenslandica Sloane 1909] LM87
              |    |--C. (E.) aurita Sloane 1904 LM87
              |    |--C. (E.) gilesi Sloane 1914 LM87
              |    |--C. (E.) levitetragramma Freitag 1979 LM87
              |    `--C. (E.) trivittata Macleay 1888 LM87
              |--C. (Grandopronotalia Horn 1936) LM87
              |    `--C. (G.) carnarvona Freitag 1979 [=Prothyma (*G.) browni Horn 1936 non C. browni Sloane 1913] LM87
              |--C. (Hypaetha LeConte 1860) LM87
              |    |--C. (*H.) quadrilineata Fabricius 1781 LM87
              |    |--C. (H.) antiqua Lea 1917 LM87
              |    |--C. (H.) frenchi Sloane 1904 LM87
              |    |--C. (H.) rafflesia Chaudoir 1852 LM87
              |    |    |--C. r. rafflesia [incl. C. montraveli Blanchard 1853] LM87
              |    |    |--‘Hypaetha’ r. expandosa (Sumlin 1992) G07
              |    |    `--C. r. pseudorafflesia Horn 1925 LM87
              |    `--C. (H.) upsilon Dejean 1825 LM87
              |         |--C. u. upsilon LM87
              |         `--C. u. albicans Chaudoir 1854 LM87
              |--‘Cylindera’ (Ifasina Jeannel 1946) G07
              |    `--C. (I.) centropunctata (Dejean 1831) G07
              |--C. (Macfarlandia Sumlin 1981) LM87
              |    `--C. (*M.) arachnoides Sumlin 1981 LM87
              |--C. (Micromentignatha Sumlin 1981) LM87
              |    |--C. (*M.) leai Sloane 1905 LM87
              |    |    |--C. l. leai [inc. C. l. nigella Sloane 1906] LM87
              |    |    `--C. l. demarzi Mandl 1960 [=C. nigella demarzi] LM87
              |    `--C. (M.) oblongicollis Macleay 1888 LM87 (see below for synonymy)
              |--C. (Myriochile) LM87
              `--C. (Rivacindela) LM87

Cicindelinae incertae sedis:
  Oxychila M02
  Therates G89b
    |--T. basalis M86
    |--T. concinnus G89b
    |--T. fasciata B35
    |--T. festiva Dupont in Boisduval 1835 B35
    |--T. flavilabris B35
    `--T. labiata [=Eurychile labiata] B35
  Lophyra reliqua [incl. L. obliquograciliaenea] B55
  Hipparidium interruptum B55
  Bennigsenium nubiferum [incl. B. flavipes (preoc.)] B55
  Cratohaerea confusa Basilewsky 1954 B55
  Prothyma concinna [incl. P. concinna var. anosignata] B55
  Calochroa Hope 1838 G07
  Odontochila Laporte 1834 F54
    `--O. salvini Bates 1874 F54
  Habrodera Motschulsky 1862 G07
    `--H. capensis (Linnaeus 1764) G07
  Tricondyla Latreille 1822 LM87
    `--*T. aptera (Olivier 1790) LM87 (see below for synonymy)
  Rhysopleura Sloane 1906 LM87
    `--*R. orbicollis (Sloane 1904) [=Distypsidera orbicollis] LM87
  Nickerlea Horn 1899 LM87
    |--*N. distypsideroides Horn 1899 LM87
    `--N. sloanei (Lea 1898) [=Cicindela sloanei] LM87
  Neocicindela H82
    |--N. latecincta (White 1846) E02
    `--N. tuberculata (Fabricius 1775) H82
  Lophyridia FG03
    |--L. caucasica (Adams 1817) [=Cicindela caucasica] FG03
    `--L. sturmi (Ménétriés 1832) [=Cicindela caucasica var. sturmi] FG03
  Rhopaloteres B55
    |--R. marshalli B55
    |--R. mechowi Quedenfeldt 1883 B55
    |    |--R. m. mechowi B55
    |    |--R. m. pseudomechowi Horn 1913 B55
    |    `--R. m. transversefasciatus Horn 1913 B55
    |--R. nysus B55
    `--R. regina B55
  Ophryodera B55
    |--O. foliicornis B55
    |    |--O. f. foliicornis B55
    |    |--O. f. divergentemaculata B55
    |    `--O. f. trimaculata B55
    `--O. rufomarginata B55
         |--O. r. rufomarginata (see below for synonymy) B55
         `--O. r. poggei [incl. O. maculatovittata, O. richteri] B55
  Distipsidera LM87
  Ctenostoma (Neoprocepalus) B14
    |--C. (N.) davidsoni B14
    |--C. (N.) guatemalensis B14
    |--C. (N.) laeticolor B14
    `--C. (N.) maculicorne (Chaudoir 1860) B14
  Habroscelimorpha dorsalis (Say 1817) B14
  Phaeoxantha B14
    |--P. aequinoctialis Dejean 1825 B14
    |    |--P. a. aequinoctialis B14
    |    `--P. a. bifasciata B14
    `--P. cruciata B14
  Aniara B14
  Collyris B66
    |--C. filiformis Chaud. 1864 B66
    |--C. formosana Bates 1866 B66
    `--C. loochooensis Kano 1929 TYM08

Cicindela hybrida Linnaeus 1758 L58 [incl. C. hybrida var. restricta C01, C. hybrida var. sahlbergi C01]

Cicindela (Antennaria) ioscelis Hope 1841 [incl. C. platycera Gestro 1879, *Antennaria platycera, C. ioscelis seminuda Mandl 1960, C. ioscelis setosoabdominalis Horn 1913] LM87

Cicindela (*Archidela) nigrina Macleay 1863 [incl. C. nigrina rugosicollis Horn 1913, C. scitula Sloane 1909] LM87

Cicindela (Cylindera) discreta Schaum 1863 [incl. C. froggatti Macleay 1887, C. discreta subfasciata Horn 1892] LM87

Cicindela (*Euzona) tetragramma Boisduval 1835 [incl. C. lineifera Sloane 1913, C. macleayi Castelnau 1867] LM87

Cicindela (Micromentignatha) oblongicollis Macleay 1888 LM87 [incl. C. clarki Sloane 1921 LM87, C. minutissima Mandl 1967 LM87, Micromentignatha minutissima G07, C. tenuicollis Macleay 1888 LM87]

Megacephala australasiae Hope 1842 [incl. M. (Tetracha) humeralis Macleay 1863, Tetracha australasiae nickerli Srnka 1895] LM87

Megacephala murchisona murchisona LM87 [incl. M. murchisona corpulenta Horn 1907 LM87, M. (Pseudotetracha) corpulenta G07]

Megacephala (Australicapitona) basalis (Macleay 1866) G07 [=Tetracha basalis LM87; incl. T. basalis phaeoxantha Horn 1894 LM87]

Megacephala (Australicapitona) bostockii (Castelnau 1867) G07, LM87 [=Tetracha bostockii LM87; incl. T. excisilatera Sloane 1897 LM87]

Ophryodera rufomarginata rufomarginata [incl. O. bohemani, O. bradshawi, O. circumcinctoides, O. distanti, O. oberthueri] B55

*Tricondyla aptera (Olivier 1790) LM87 [=Cicindela aptera LM87, Collyris aptera B35; incl. Tricondyla connata M86]

*Type species of generic name indicated

REFERENCES

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[BB68] Basford, N. L., J. E. Butler, C. A. Leone & F. J. Rohlf. 1968. Immunologic comparisons of selected Coleoptera with analyses of relationships using numerical taxonomic methods. Systematic Zoology 17 (4): 388–406.

[B55] Basilewsky, P. 1955. Carabidae (Coleoptera) de l’Angola (première partie). Diamang Publicações Culturais 27: 91–137.

[B66] Bates, H. W. 1866. On a collection of Coleoptera from Formosa sent home by R. Swinhoe, Esq., H. B. M. Consul, Formosa. Proceedings of the Zoological Society of London 1866: 339–355.

[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).

[C90] Crawford, C. S. 1990. Scorpiones, Solifugae, and associated desert taxa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 421–475. John Wiley & Sones: New York.

[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.

[E02] Emberson, R. M. 2002. The beetle (Coleoptera) fauna of the Chatham Islands: Additions and corrections. New Zealand Entomologist 25: 69–77.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[F02] Foord, M. 2002. Entomological Society conference field trip 2002. Weta 24: 37–38.

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[FG03] Franzen, M. & J. Gebert. 2003. Distribution of the tiger beetle Lophyridia caucasica (Adams, 1817) in Iran (Coleoptera: Cicindelidae). Zoology in the Middle East 29: 79–82.

[G89a] Gestro, R. 1889a. 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.

[G89b] Gestro, R. 1889b. Viaggio di Leonardo Fea in Birmania e regioni vicine. XV.—Primo studio delle cicindele. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 77–91.

[G07] Golding, M. R. 2007. A Pictorial Field Guide to the Beetles of Australia. Part 2: Cicindelidae. Ocean Publishing.

[H15] Hedley, C. 1915. Presidential address. Journal and Proceedings of the Royal Society of New South Wales 49 (1): 1–77, pls 1–7.

[H82] Helmore, D. W. 1982. Drawings of New Zealand insects. Bulletin of the Entomological Society of New Zealand 8: 1–52.

[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.

[M02] Machado, G. 2002. Maternal care, defensive behavior, and sociality in Neotropical Goniosoma harvestmen (Arachnida, Opiliones). Insectes Sociaux 49: 388–393.

[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.

[TYM08] Tennent, W. J., M. Yasuda & K. Morimoto. 2008. Lansania Journal of arachnology and zoology—a rare and obscure Japanese natural history journal. Archives of Natural History 35 (2): 252–280.

[TKM93] Thaler, K., B. Knoflach & E. Meyer. 1993. Fragmenta faunistica Tirolensia—X. Berichte des Naturwissenschaftlich-Medizinischen Vereins in Innsbruck 80: 311–325.

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Last updated: 10 November 2018.