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 2a, 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 2a, 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 2nd 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 9th ed. Brooks/Cole Cengage Learning.
Last updated: 19 December 2020.
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