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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Coleopterida
Ordo: Coleoptera
Subordo: Adephaga

Familia: Carabidae
Subfamilia: Paussinae
Tribus (4): Metriini - Ozaenini - Paussini - Protopaussini
Overview of genera (51)

AnentmetusArthropterusCarabidomemnusCerapteritesCerapterusCeratoderusCrepidozaenaDhanyaEntomoantyxEohomopterusEopaussusEuplatyrhopalusEustraFilicerozaenaGibbozaenaGranulopaussusHeteropaussusHexaplatarthrusHomopterusHylopaussusHylotorusInflatozaenaItamusLebioderusLeleupaussusMegalopaussusMelanospilusMesarthropterusMetriusMicrozaenaMimozaenaMystropomusOzaenaPachytelesPaussomorphusPaussopsisPaussusPentaplatarthrusPhyseaPhyseomorphaPlatycerozaenaPlatyrhopalopsisPlatyrhopalusProozaenaProtopaussusPseudozaenaPterorhopalusSerratozaenaSinometriusSphaerostylusTachypeles
Name

Paussinae Latreille, 1807
Original spelling(s): Paussili, Paussiles
Original rank(s): familia
Type genus: Paussus Linnaeus, 1775
Stem: Pauss-

References

Carvalho de, E.L., 2001: A new species of paussine ground-beetle from Guinea-Bissau (Coleoptera: Carabidae: Paussinae). Boletin Sociedad Entomológica Aragonesa 29: 45–46. Full article: [1].
Linnaeus 1775: Bigae Insect., 6.
Latreille, P.A. 1807. Genera crustaceorum et insectorum secundem ordinem naturalem in familias disposita, iconibus exemplisque plurimus explicata. Tomus tertius. Parisiis et Argentorati: A. Koenig, 258 pp. BHL Reference page.
Nagel, P. 2009: Flanged bombardier beetles from Laos (Carabidae: Paussinae). Entomologica Basiliensia et Collectionis Frey, 31: 101–113. PDF

Links

Carabidae of the World
Ground beetles of the Subfamily Paussinae Latreille, 1807

Vernacular names
Deutsch: Fühlerkäfer
English: Paussid beetle
日本語: ヒゲブトオサムシ亜科
русский: Пауссиды

Ant nest beetles (subfamily Paussinae) or paussines, some members of which are known also as flanged bombardier beetles, are a large subfamily within the ground beetles (Carabidae).The tribes Metriini, Ozaenini, Paussini and Protopaussini are included in the subfamily.

Rarely seen in the open, except at lights, most Paussinae are obligate or facultative myrmecophiles, living within the nests of ants, predatory on ant larvae and workers. Many have elaborate antennal structures and body parts flattened. Paussines are moderate sized (6–20 mm), characterised by glandular hairs that produce secretions attractive to ants and by the odd antennal structures of many species. Their pygidial glands can produce explosive secretions, with a spray of quinones that are directed forward by flanges at the posterior end of the elytra, giving them the other name of flanged bombardier beetles although they are not particularly close relatives of the typical bombardier beetles (Brachininae).

Biology

Very little is known about the immature stages of ant nest beetles. Most appear to live in ant nests in their early stages of life. Although many are facultative or obligate myrmecophiles, most do not appear like ants (i.e. myrmecomorphic) and unlike in the case of myrmecophilous larval Lycaenidae, there appears to be no benefit gained by the ants in this association.[1] Many species follow the trails of worker ants of specific species. Glandular secretions on their antennae and body allow them to interact with ant workers.[2] It is believed that the beetles use an acoustic mechanism to successfully imitate the sounds of an ant queen thus permitting them access to the nest without alarming the ants.[3] Worker ants groom the beetles. Adult Paussus favieri have been observed to antennate and interact by stridulation with the queen of Pheidole pallidula in the brood chamber.[4] The defensive bombardier behaviour is never used against ants. Unlike bombardier beetles in the family Brachininae, which have two glands opening close together on the abdomen, the glands are located close to the lateral margins and lie under a flange formed by the tip of the elytra. The flange is curved and the explosive hot spray of quinones is directed forward by Coandă effect.[5] The flange is not present in members of the tribe Metriini but is replaced by similar internal structures.[6] Males of some species are attracted to lights during some parts of the season and are thought to disperse from one ant nest to another. Males are thought to be short-lived. These beetles feed on ant eggs, larvae and adults by piercing their mandibles into the abdomen or other soft part and sucking the contents. Apart from chemical mimicry and communication with their hosts, they also make use of vibrations. Several stridulatory structures are found in these beetles including alary-elytral, abdomen-femur and thorax-femur combinations of surfaces.[7] Larvae of only about 10 species have ever been examined. The first instar larvae of Paussus favieri has a terminal disc which is raised at times and bent back over the head. They also exhibit some behaviours and mouth part adaptations which may be involved in eliciting trophallaxis by the ant hosts but this has not been confirmed. They opportunistically feed on the haemolymph of ant larvae.[8]
Systematics

The subfamily Paussinae contains 49 genera, with around 800 species divided into the following tribes and subtribes:[9]

While some tribes like Metriini and Ozaenini appear quite similar to typical carabids, others have modified antennae and body shapes. The Protopaussini and Paussini are slender or compact in body shape with enlarged antennae in the Paussini.
Arthropterus cerapteroides (Paussini: Cerapterina)
Paussina species from South Africa
Heteropaussus hastatus (Paussini: Heteropaussina)
Protopaussus sp.
Tribe Metriini

Metrius Eschscholtz, 1829
Sinometrius Wrase & Schmidt, 2006
†Kryzhanovskiana Kataev et al, 2019[10] Burmese amber, Myanmar, Cenomanian

Tribe Ozaenini

Anentmetus Andrewes, 1924
Crepidozaena Deuve, 2001
Dhanya Andrewes, 1919
Entomoantyx Ball & McCleve, 1990
Eustra Schmidt-Goebel, 1846
Filicerozaena Deuve, 2001
Gibbozaena Deuve, 2001
Goniotropis Gray, 1831
Inflatozaena Deuve, 2001
Itamus Schmidt-Goebel, 1846
Microzaena Fairmaire, 1901
Mimozaena Deuve, 2001
Mystropomus Chaudoir, 1848
Ozaena Olivier, 1812
Pachyteles Perty, 1830
Physea Brulle, 1834
Physeomorpha Ogueta, 1963
Platycerozaena Banninger, 1927
Proozaena Deuve, 2001
Pseudozaena Laporte de Castelnau, 1834
Serratozaena Deuve, 2001
Sphaerostylus Chaudoir, 1848
Tachypeles Deuve, 2001

Tribe Paussini

Subtribe Carabidomemnina Wasmann, 1928

Carabidomemnus H. Kolbe, 1924
Eohomopterus Wasmann, 1919

Subtribe Cerapterina Billberg, 1820

Arthropterus MacLeay, 1838
Cerapterus Swederus, 1788
Megalopaussus Lea, 1906
Mesarthropterus Wasmann, 1926

Subtribe Heteropaussina Janssens, 1953

Heteropaussus Thomson, 1860

Subtribe Homopterina Janssens, 1953

Homopterus Westwood, 1838

Subtribe Paussina Latreille, 1807

Ceratoderus Westwood 1841
Eopaussus Wasmann, 1926
Euplatyrhopalus Desneux, 1905
Granulopaussus H. Kolbe, 1938
Hylopaussus Luna de Carvalho, 1989
Hylotorus Dalman, 1823
Lebioderus Westwood, 1838
Leleupaussus Luna de Carvalho, 1962
Melanospilus Westwood, 1845
Paussomorphus Raffray, 1885
Paussus Linnaeus, 1775
Platyrhopalopsis Desneux, 1905
Platyrhopalus Westwood, 1838
Pterorhopalus Maruyama, 2011

Subtribe Pentaplatarthrina Jeannel, 1946

Hexaplatarthus Jeannel, 1955
Pentaplatarthus Westwood, 1833

Tribe Protopaussini

Protopaussus Gestro, 1892

References

Moore, W; Xiao-bin Song; Andrea Di Giulio (2011). "The larva of Eustra (Coleoptera, Paussinae, Ozaenini):a facultative associate of ants". ZooKeys (90): 63–82. doi:10.3897/zookeys.90.1136. PMC 3084492. PMID 21594107.
Cammaerts, R; C Detrain; M-C Cammaerts. "Host trail following by the myrmecophilous beetle Edaphopaussus favieri (Fairmaire) (Carabidae, Paussinae)" (PDF).
Andrea Di Giulio; Emanuela Maurizi; Francesca Barbero; Marco Sala; Simone Fattorini; Emilio Balletto; Simona Bonelli (2015). "The Pied Piper: A Parasitic Beetle's Melodies Modulate Ant Behaviours". PLOS ONE. 10 (7): e0130541. doi:10.1371/journal.pone.0130541. PMC 4496082. PMID 26154266.
Robertson, James A.; Moore, Wendy (2017). "Phylogeny of P aussus L . ( C arabidae: P aussinae): unravelling morphological convergence associated with myrmecophilous life histories". Systematic Entomology. 42 (1): 134–170. doi:10.1111/syen.12205. ISSN 0307-6970.
Eisner, Thomas; Aneshansley, Daniel J. (1982). "Spray Aiming in Bombardier Beetles: Jet Deflection by the Coanda Effect". Science. 215 (4528): 83–85. doi:10.1126/science.215.4528.83. ISSN 0036-8075.
Muzzi, Maurizio; Moore, Wendy; Di Giulio, Andrea (2019). "Morpho-functional analysis of the explosive defensive system of basal bombardier beetles (Carabidae: Paussinae: Metriini)". Micron. 119: 24–38. doi:10.1016/j.micron.2019.01.003.
Geiselhardt, Stefanie F.; Klaus Peschke; Peter Nagel (2007). "A review of myrmecophily in ant nest beetles (Coleoptera: Carabidae: Paussinae): linking early observations with recent findings". Naturwissenschaften. 94 (11): 871–894. doi:10.1007/s00114-007-0271-x. PMID 17563864. S2CID 945199.
Di Giulio, Andrea; Maurizi, Emanuela; Hlaváč, Peter; Moore, Wendy (2011). "The long-awaited first instar larva of Paussus favieri (Coleoptera: Carabidae: Paussini)". European Journal of Entomology. 108 (1): 127–138. doi:10.14411/eje.2011.016.
"Paussinae Latreille, 1807". Carabidae of the World. 2011. Retrieved 21 Jul 2011.
Kataev, Boris M.; Kirejtshuk, Alexander G.; Manukyan, Andranik R.; Anokhin, Boris A. (November 2019). "Kryzhanovskiana olegi gen. et sp. nov., a remarkable eyeless representative of the tribe Metriini (Coleoptera: Carabidae: Paussinae) from Upper Cretaceous amber of northern Myanmar". Cretaceous Research. 103: 104168. doi:10.1016/j.cretres.2019.06.014.

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