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. The African Brachynillus varendorffi is completely blind (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.: MastaxB55
    |           |--M. burgeoniB55
    |           |--M. ornatellusB55
    |           `--M. paecilus Schaum 1863B89b
    |         Brachynillus varendorffi Reitter 1904BB01
    |  |    |--A. alpinusOM08
    |  |    `--A. displosorOM08
    |  `--+--StyphlodromusOM08
    |     `--StyphlomerusOM08
    |          |--S. ludicrus [incl. S. ludicrus var. grandis]B55
    |          `--S. vittaticollisOM08
    `--+--Crepidogaster [Crepidigastrini]OM08
       |    |--C. (Tyronia) ambreanaOM08
       |    |--C. atrataOM08
       |    `--C. picipennisB55
       `--+--Brachinus Weber 1801OM08, BB01
          |    |  i. s.: B. barbarus Lucas 1846E12
          |    |         B. bimaculatusB35
          |    |         B. caligatus Bates 1889B89a
          |    |         B. causticusR26
          |    |         B. frontalisB89a
          |    |         B. explodens [incl. B. explodens var. glabratus]C01
          |    |         B. fimbriolatus Lucas 1846E12
          |    |         B. immaculicornisC01
          |    |         B. mexicanusE72b
          |    |         B. oblongusG89
          |    |         B. salleiE72a
          |    |         B. sclopetaC01
          |    |--B. (Brachinus) crepitansG20
          |    |--B. (Metabrachinus) armigerOM08 [incl. Metabrachinus armiger var. natalicusB55]
          |    `--B. (Neobrachinus Erwin 1970)BB01
          |         `--B. (N.) hirsutusOM08
          `--Pheropsophus Solier 1833 [Pherapsophina]B12
               |  i. s.: P. aequinoctialis (Linnaeus 1763)B14
               |         P. africanus [=Brachinus africanus]G89
               |         P. agnatusB89b
               |         P. alexandrae Giachino 2003B12
               |         P. arcanusB55
               |         P. darwini Giachino 2003B12
               |         P. dimidiatusB55
               |         P. gregoryi Giachino 2003B12
               |         P. hispanicusA90
               |         P. jessoensisLB91
               |         P. livingstoneiB55
               |         P. macleayi Sloane 1895B12
               |         P. marginalis [incl. P. nebulosus Chaud. 1876]B89b
               |         P. minorB55
               |         P. siamensisB89b
               |         P. straneoi Giachino 2003B12
               |         P. windjanae Baehr 2012B12
               |--P. (Pheropsophus)B12
               |    |--*P. (P.) madagascariensis (Dejean 1831) [=Brachinus madagascariensis]B12
               |    `--P. (P.) complanatus (Fabricius 1775)LM87 [=Carabus complanatusLM87, Nebria complanataL02]
               `--P. (Stenaptinus Maindron 1906) [incl. Parapheropsophus Hubenthal 1914]LM87
                    |--P. (*S.) krichna (Maindron 1914) [=*Stenaptinus krichna]LM87
                    |--‘Stenaptinus’ insignisSBM11
                    `--P. (S.) verticalis (Dejean 1825)LM87 (see below for synonymy)

Pheropsophus (Stenaptinus) verticalis (Dejean 1825)LM87 [=Brachinus verticalisLM87, P. (*Parapheropsophus) verticalisLM87; incl. P. australis Castelnau 1867LM87, P. baliothorax Heller 1910LM87, P. hypoxanthus Chaudoir 1876LM87, P. (Parapheropsophus) intermedius Hubenthal 1914LM87, Aptinus jamissoniB35, P. (Pa.) niger Hubenthal 1914LM87, P. papuensis Macleay 1877LM87]

*Type species of generic name indicated


[A90] Alluaud, C. 1889. Une lettre relative à son voyage entomologique. Annales de la Société Entomologique de France, 6e série 9: ccxxxi–ccxxxii.

[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.

[B89a] Bates, H. W. 1889a. 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.

[B89] Bates, H.-W. 1889b. Contributions a la faune Indo-Chinoise. 3e mémoire. Carabidae. Annales de la Société Entomologique de France, 6e série 9: 261–286.

[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.

[E72a] Erwin, T. L. 1972a. The lost type of Brachinus sallei Chaudoir (Coleoptera: Carabidae). Pan-Pacific Entomologist 48 (1): 63–64.

[E72b] Erwin, T. L. 1972b. Observations on the life history of bombadier beetles: mechanism of egg-hatching (Coleoptera: Carabidae). Pan-Pacific Entomologist 48 (1): 64.

[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 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.

[R26] Risso, A. 1826. Histoire naturelle des principales productions de l’Europe méridionale et particulièrement de celles des environs de Nice et des Alpes maritimes vol. 5. F.-G. Levrault: Paris.

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

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