Pselaphinae

Fustigerodes majusculus, from Péringuey (1897).

Belongs within: Staphylinidae.
Contains: Pselaphitae, Goniaceritae, Batrisitae, Faronini, Bythinoplectitae, Euplectini, Trogastrini, Jubini, Raffrayia, Claviger, Articerus, Fustiger.

The Pselaphinae are a cosmopolitan group of small, often grotesque-looking staphylinid beetles with a robust, relatively inflexible body (Newton et al. 2001). Members include the Clavigeritae, specialised beetles that are all associates of ants. Clavigeritae have antennae with one to six segments and greatly reduced maxillary palpi. An extensive hollow at the base of the abdomen surrounded by tufts of golden-yellow setae secretes a substance attractive to the beetle's ant hosts.

Characters (from Newton et al. 2001): Body robust, relatively inflexible, usually reddish-brown, with elytra exposing some abdominal segments; deep conical foveae present on vertex and/or other parts of the body; antennae inserted anterior to a line drawn between anterior margins of eyes; maxillary palps with 1-4 palpomeres plus a lightly sclerotized or unsclerotized apical pseudosegment, therefore often appearing to have five palpomeres; tarsal formula 3-3-3, rarely apparently 2-2-2; abdominal sterna III and IV usually with very dense setal fringe covering a basal impression on sternum IV.

<==Pselaphinae [Clavigeridae, Pselaphi, Pselaphidae, Pselaphii] NG05
    |  i. s.: Pselaphaulax MW01
    |         Seracamaurops PN02
    |         Troglamaurops PN02
    |         Cordiamaurops PN02
    |         Imtemps NG05
    |         Machadoites termitophilus N55
    |         Tinaroo ZS10
    |         Malleecola myrmecophila ZS10
    |--+--Pselaphitae R03
    |  `--Brachyscelidae R03
    |       |--Goniaceritae R03
    |       |--Batrisitae R03
    |       |--Faronini R03
    |       |--Bythinoplectitae R03
    |       `--Euplectitae [Euplectinae] R03
    |            |--Euplectini R03
    |            |--Trogastrini NT01
    |            |--Jubini NT01
    |            |--Raffrayia NT01
    |            `--Pteracmes Raffray 1890 R03 [Pteracmini NT01]
    |                 `--P. schaufussi Raffray 1890 R03
    `--Clavigeritae [Clavigerinae] R03
         |--Tiracerus LB91 [Tiracerini NG05]
         |    `--T. foveicollis LB91
         |--Colilodion Besuchet 1991 [Colilodionini] NG05
         |    |--C. incredibilis Besuchet 1991 NG05
         |    |--C. inopinatus Besuchet 1991 NG05
         |    |--C. mirus Besuchet 1991 NG05
         |    `--C. wuesti Löbl 1994 NG05
         `--Clavigerini [Adranini, Clavigerodini, Fustigerini] NG05
              |--Claviger R04
              |--Cerylambus reticulatus (Raffray 1895) NG05
              |--Articerus R04
              |--Fustiger NT01
              |--Pseudacerus Raffray 1895 R04
              |    `--P. furcatus Raffray 1895 NG05
              |--Anaclasiger Raffray 1890 R04
              |    `--A. sinuaticollis Raffray 1890 NG05
              |--Elasmatus Raffray 1903 R04
              |    `--E. lamellipes Raffray 1903 R04
              |--Theocerus Raffray 1896 R04
              |    `--T. crenulatus Raffray 1896 R04
              |--Amblycerus Raffray 1895 R04
              |    `--A. reticulatus Raffray 1895 R04
              |--Neocerus Wasmann 1893 R04
              |    `--N. compressicornis Wasmann 1893 R04
              |--Commatocerus Raffray 1882 R04
              |    `--C. elegantulus Raffray 1882 R04
              |--Apoderiger Wasmann 1897 R04
              |    `--A. cervinus Wasmann 1897 R04
              |--Trymalius Fairmaire 1898 R04
              |    `--T. foveicollis Fairmaire 1898 R04
              |--Hadrophorus Fairmaire 1898 R04
              |    `--H. humerosus Fairmaire 1898 R04
              |--Novofustiger Wasmann 1893 R04
              |    `--N. raffrayi Wasmann 1893 R04
              |--Pseudofustiger Reitter 1884 R04
              |    `--P. stricticornis (Reitter 1883) [=Articerus stricticornis] R04
              |--Commatoceropsis Raffray 1890 R04
              |    `--C. madagascariensis (Raffray 1882) [=Commatocerus madagascariensis] R04
              |--Rhynchoclaviger Wasmann 1891 R04
              |    `--R. crematogastris Wasmann 1891 R04
              |--Articeropsis Wasmann 1893 R04
              |    `--A. sikorae Wasmann 1893 R04
              |--Articeronomus Raffray 1897 R04
              |    `--A. nitidus Raffray 1897 R04
              |--Fusifer Raffray 1899 R04
              |    `--F. perrieri (Fairmaire 1898) [=Commatoceropsis perrieri] R04
              |--Paussiger Wasmann 1893 R04
              |    `--P. limicornis Wasmann 1893 R04
              |--Commatocerodes Péringuey 1888 R04
              |    `--C. raffrayi Péringuey 1888 R04
              |--Syrraphesina Raffray 1903 R04
              |    `--S. pliciventris Raffray 1903 R04
              |--Clavigerodes Raffray 1877 R04
              |    `--C. abyssinicus Raffray 1877 R04
              |--Bironia Raffray 1903 R04
              |    `--B. cavernosa Raffray 1903 R04
              |--Braunsiella Raffray 1901 R04
              |    `--B. pubiventris Raffray 1901 R04
              |--Clavigeropsis Raffray 1882 R04
              |    `--C. formicaria Raffray 1882 R04
              |--Semiclaviger Wasmann 1893 R04
              |    `--S. sikorae Wasmann 1893 R04
              |--Mastiger Motschulsky 1851 R04
              |    |--M. abruptus Motschulsky 1851 R04
              |    `--M. brevicornis Raffray 1890 NG05
              |--Disarthricerus Raffray 1895 R04
              |    |--D. integer Raffray 1895 NG05
              |    `--D. moultoni Bryant 1915 NG05
              |--Diartiger Sharp 1883 R04
              |    |--D. fossulatus Sharp 1883 R04
              |    `--D. spinipes Sharp 1883 R04
              |--Fustigeropsis Raffray 1890 R04
              |    |--F. peringueyi (Raffray 1887) [=Commatocerus peringueyi] R04
              |    `--F. simplex Raffray 1904 R04
              |--Thysdarius Fairmaire 1904 [incl. Thysdrus Fairmaire 1898 (preoc.)] R04
              |    |--T. gracilis Raffray 1899 R04
              |    `--T. perrieri Fairmaire 1898 R04
              |--Radamellus Raffray 1904 R04
              |    |--R. minutus (Raffray 1899) [=Radamides minutus] R04
              |    `--R. spinipennis (Raffray 1883) [=Radama spinipennis] R04
              |--Imerina Raffray 1896 R04
              |    |--I. breviceps Raffray 1896 R04
              |    `--I. wasmanni Raffray 1896 R04
              |--Miroclaviger Wasmann 1893 R04
              |    |--M. alluaudi Raffray 1897 R04
              |    `--M. cervicornis Wasmann 1893 R04
              |--Radamides Wasmann 1897 R04
              |    |--R. sulcicollis Fairmaire 1899 R04
              |    `--R. trifoveolatus Wasmann 1897 R04
              |--Articerodes Raffray 1890 R04
              |    |--A. borneensis Bryant 1915 NG05
              |    |--A. quadriscopulatus Schaufuss 1883 R04
              |    `--A. syriacus Saulcy 1865 [incl. A. bucharicus Reitter 1900, A. ponticus Sharp 1878] R04
              |--Radama Raffray 1883 R04
              |    |--R. fimbriatus Wasmann 1893 R04
              |    |--R. inflatus Raffray 1883 R04
              |    `--R. sulcatus Wasmann 1897 R04
              |--Adranes LeConte 1849 NT01
              |    |--A. coecus Leconte 1850 R04
              |    |--A. lecontei Brendel 1865 R04
              |    |--A. pacificus Wickham 1901 R04
              |    `--A. taylori Wickham 1901 R04
              `--Fustigerodes Reitter 1884 [incl. Novoclaviger Wasmann 1894] R04
                   |--F. articulatus Wasmann 1898 R04
                   |--F. braunsi Wasmann 1897 R04
                   |--F. capensis Péringuey 1888 R04
                   |--F. majusculus Péringuey 1888 R04
                   `--F. wroughtoni [=Novoclaviger wroughtoni] R04

*Type species of generic name indicated

REFERENCES

[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).

[MW01] Molleman, F., & D. E. Walter. 2001. Niche segregation and can-openers: scydmaenid beetles as predators of armoured mites in Australia. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 283–288 Publishing: Melbourne.

[NT01] Newton, A. F., M. K. Thayer, J. S. Ashe & D. S. Chandler. 2001. Staphylinidae Latreille, 1802. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 272–418. CRC Press: Boca Raton.

[N55] Noirot, C. 1955. Termites du centre et du sud-ouest de l’Angola récoltés par A. de Barros Machado. Diamang Publicações Culturais 27: 139–150.

[NG05] Nomura, S., & I. Abd Ghani. 2005. Faunistic notes on the pselaphine species of the supertribes Goniaceritae, Pselaphitae and Clavigeritae from Malaysia and Singapore (Coleoptera: Staphylinidae: Pselaphinae). Serangga 10 (1–2): 1–36.

[PN02] Poggi, R., G. Nonveiller, A. Colla, D. Pavićević & T. Rađa. 2001–2002. Thaumastocephalini, a new tribe of Pselaphinae for Thaumastocephalus folliculipalpus n. gen., n. sp., an interesting new troglobious species from central Dalmatia (Croatia) (Coleoptera, Staphylinidae, Pselaphinae). Annali del Museo Civico di Storia Naturale “Giacomo Doria” 94: 1–20.

[R03] Raffray, A. 1903. Genera et catalogue des psélaphides. Annales de la Société Entomologique de France 72: 484–604.

[R04] Raffray, A. 1904. Genera et catalogue des psélaphides [part III]. Annales de la Société Entomologique de France 73: 401-476, pls 1-3.

[ZS10] Zborowski, P., & R. Storey. 2010. A Field Guide to Insects in Australia 3^rd ed. Reed New Holland: Sydney.

Scydmaeninae

Microscydmus, copyright Edenis.

Belongs within: Staphylinidae.
Contains: Euconnini.

The Scydmaeninae, ant-like stone beetles, are a group of minute beetles found in habitats such as leaf litter or moss, or in association with ants or termites (O'Keefe 2001). Though long treated as their own family, more recent phylogenetic studies have established that they are derived within the Staphylinidae and they are now treated as a subfamily within that larger grouping.

Characters (from O'Keefe 2001, as Scydmaenidae): Body elongate, slender to ovoid; slightly to very convex, 0.6–2.7 mm long; often distinctly constricted between head and pronotum and between pronotum and elytra, usually light to dark brown, sometimes black in color; often densely pubescent on head, pronotum, and elytra, pubescence usually long. Head short and broad to ovoid; deflexed; usually distinctly constricted between vertex and occiput; occiput formed into a “neck”; clypeofrontal region moderate in length, slightly narrowed anteriorly. Eyes anterior, median, or posterior; moderate in size or absent. Antennae inserted anterior to eyes, antennal insertions adjacent to widely separated; antennae composed of scape, pedicel and nine flagellomeres; antennal club composed of distal three to five antennomeres, from indistinct to distinct in form. Mandibles usually planar, subtriangular with large basal area, narrowed to acute incisor, subapical teeth usually absent. Maxillary palpus composed of four palpomeres; palpomere III large, clavate; palpomere IV variable, distinctly smaller than III; apices of lacinea and galea densely covered with long setae. Labial palpus composed of three palpomeres; mentum large. Thorax with pronotum distinctly wider than head, usually with fovea or transverse furrow along posterior margin. Scutellum small, visible, occasionally hidden. Elytra ovoid, convex to rectangular, flat; humeri varied in form with 0-2 basal foveae in broad basal impression; elytral apices usually entire, occasionally truncate or fused. Hind wings, if present, well-developed. Legs relatively short to moderate in length; procoxae projecting beyond prosternum, procoxal cavities open; mesocoxal cavities closed; trochanters relatively large and sublinear or rounded triangular; femora strongly clavate in distal half; protibiae and mesotibiae usually with dense patches of setae at distal end. Prosternum varied from subquadrate to very narrow before conical, contiguous procoxae, pubescence varied. Mesosternum usually raised, sometimes densely setose, bearing mid-ventral carina separating otherwise contiguous mesocoxae. Metasternum long, broad, occasionally carinate or medially excavate; metacoxae round to transverse, conical, widely separate to contiguous. Abdomen with six visible sternites. Male genitalia with median lobe bulbous, large, to elongate, to reduced, lightly sclerotised, distinctly curved, parameres long, slender, articulated on dorsal surface of aedeagus, occasionally absent. Ovipositor composed of paired, fused or unfused dorsolateral paraprocts, triangular dorsal proctiger, paired elongate ventral valvifers, paired elongate gonocoxae; gonostyli absent; spermatheca present, variable in shape from elongate, slender to spherical, accessory gland sometimes present.

<==Scydmaeninae [Anisosphaeridae, Scydmaenidae, Scydmaenides] B14
    |  i. s.: Adrastia MW15
    |         Phagonophana B70
    |         Heterognathus myrmecophilus B70
    |         Syndicus [Syndicini] LB91
    |         Palaeostigus Newton 1982 GN09
    |           `--P. bifoveolatus (Boheman 1851) GN09
    |         Palaeoleptochromus GN09
    |--Hapsomela [Hapsomelinae, Hapsomelitae] GN09
    |--Mastigitae [Mastiginae] GN09
    |    |--Leptomastacini GN09
    |    |--Mastigini GN09
    |    `--Clidicini GN09
    |         |--Clidicus AY04
    |         `--Papusus Casey 1897 O01
    |              `--P. macer O01
    `--Scydmaeinitae GN09
         |  i. s.: Cephenniini GN09
         |           |--Coatesia LB91
         |           |--Neseuthia LB91
         |           |--Chelonoidum Strand 1935 [=Chelonoides Croissandeau 1894] O01
         |           |    `--C. corporosum O01
         |           `--Cephennium Müller & Kunze 1822 O01
         |                |--C. anophthalmicum O01
         |                `--C. thoracicum (Müller & Kunze 1822) B14
         |         Plaumanniolini GN09
         |         Chevrolatia Jacquelin du Val 1850 O01 [Chevrolatiini GN09]
         |           |--C. amoena LeConte 1866 B14
         |           `--C. occidentalis B14
         |         Leptoscydmus Casey 1897 O01 [Leptoscydmini GN09]
         |           `--L. caseyi O01
         |--Scydmaenini LB91
         |    |--Palaeoscydmaenus LB91
         |    |--Ceramphis Casey 1897 O01
         |    |    `--C. deformata O01
         |    `--Scydmaenus Latreille 1802 GN09 (see below for synonymy)
         |         |--S. angustatus Lucas 1846 E12
         |         |--S. hellwigii [=Pselaphus hellwigii] L02
         |         |--S. motschulskii O01
         |         |--S. myrmecophilus LB91
         |         |--S. nanus Lucas 1846 E12
         |         `--S. schaumii Lucas 1846 E12
         `--+--+--Euconnini GN09
            |  `--+--Cephennodes Reitter 1884 GN09
            |     `--Eutheiini O01
            |          |--Eutheia Stephens 1830 [=Euthia Agassiz 1847; incl. Euthiopsis Müller 1925] O01
            |          |--Euthiconus Reitter 1881 [incl. Ascydmus Casey 1897, Euthiodes Brendel 1893] O01
            |          |    `--E. lata O01
            |          `--Veraphis Casey 1987 GN09
            |               `--V. cristata O01
            `--Cyrtoscydmini MW01
                 |--Neuraphus MW01
                 |--Parascydmus Casey 1897 O01
                 |--Brachycepsis Brendel 1889 O01
                 |    `--B. subpunctatus O01
                 |--Catalinus Casey 1897 O01
                 |    `--C. angustus O01
                 |--Lophioderus Casey 1897 O01
                 |    `--L. arcifer O01
                 |--Taphroscydmus Casey 1897 O01
                 |    `--T. californicus O01
                 |--Stenichnus Thomson 1859 GN09 [incl. Cyrtoscydmus Motschulsky 1870 O01]
                 |    |--S. perforatus O01
                 |    `--S. turbatus Casey 1897 GN09
                 `--Microscydmus Saulcy & Croissandeau 1893 O01
                      |--M. (Microscydmus) [incl. Opresus Casey 1897] O01
                      |    `--M. misellus O01
                      |--M. (Delius Casey 1897) O01
                      |    `--M. (D.) robustulus O01
                      `--M. (Neladius Casey 1897) O01
                           `--M. (N.) tenuis O01

Scydmaenus Latreille 1802 GN09 [incl. Acholerops Casey 1897 O01, Eumicrus LaPorte 1840 O01, Microstemma Motschulsky 1858 O01]

*Type species of generic name indicated

REFERENCES

[AY04] Austin, A. D., D. K. Yeates, G. Cassis, M. J. Fletcher, J. La Salle, J. F. Lawrence, P. B. McQuillan, L. A. Mound, D. J. Bickel, P. J. Gullan, D. F. Hales & G. S. Taylor. 2004. Insects ‘Down Under’—diversity, endemism and evolution of the Australian insect fauna: examples from select orders. Australian Journal of Entomology 43 (3): 216–234.

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

[B70] Britton, E. B. 1970. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 495–621. Melbourne University Press.

[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.

[GN09] Grebennikov, V. V., & A. F. Newton. 2009. Good-bye Scydmaenidae, or why the ant-like stone beetles should become megadiverse Staphylinidae sensu latissimo (Coleoptera). European Journal of Entomology 106: 275–301.

[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).

[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.

[MW01] Molleman, F., & D. E. Walter. 2001. Niche segregation and can-openers: scydmaenid beetles as predators of armoured mites in Australia. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 283–288 Publishing: Melbourne.

[O01] O’Keefe, S. T. 2001. Scydmaenidae Leach, 1915. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 259–267. CRC Press: Boca Raton.

Last updated: 27 January 2019.

Lomechusini

Drusilla canaliculata, copyright Udo Schmidt.

Belongs within: Aleocharinae.
Contains: Zyras.

The Lomechusini are a diverse but poorly defined group of staphylinid beetles that have been recognised in the past (under the name Myrmedoniini) as including almost all Aleocharinae with 4,5,5-segmented tarsi. Many species possess mouthparts with extensive maxillary sinuses that extend caudally beyond the cardo, and an elongated galea and lacinia (Seevers 1957). Many, but not all, species are associated with ants and members of the subtribe Lomechusina possess prominent tufts of golden-yellow trichomes on the abdomen (Newton et al. 2001).

Characters (from Newton et al. 2001): Tarsi 4-5-5; metasternal process distinctly longer than very short mesosternal process; galea and lacinia of maxilla usually moderately to greatly elongated, length of galea equal to or greater than distance to base of galea from cardo; mesocoxae moderately to very broadly separated by broad meso- and metasternal processes.

Lomechusini [Myrmedoniini, Zyrasini] NT01
    |--Schistogeniae S57
    |--Termitonda Seevers 1957 [Termitondina] S57
    |    |--*T. liberiae Seevers 1957 S57
    |    `--T. tachyporoides Seevers 1957 S57
    |--Lomechusina NT01
    |    |--Lomechusa strumosa A71
    |    `--Xenodusa Wasmann 1894 [incl. Pseudolomechusa Mann 1914] NT01
    |         |--X. cava B14
    |         `--X. reflexa (Walker 1866) B14
    |--Feldina S57
    |    |--Felda Blackwelder 1952 [=Asticta Wasmann 1916 (preoc.)] S57
    |    |    `--*F. butteli (Wasmann 1916) [=*Asticta butteli] S57
    |    |--Zunia Blackwelder 1952 [=Disticta Wasmann 1916 (preoc.)] S57
    |    |    `--*Z. capritermitis (Wasmann 1916) [=*Disticta capritermitis] S57
    |    `--Termitobiella Wasmann 1916 S57
    |         `--*T. setipes Wasmann 1916 S57
    |--Termitozyrina S57
    |    |--Termitophagus Silvestri 1945 S57
    |    |    `--*T. synterminus Silvestri 1945 S57
    |    |--Termitonusa Borgmeier 1950 S57
    |    |    `--*T. sequax Borgmeier 1950 S57
    |    |--Iheringocantharus Bernhauer 1912 S57
    |    |    `--*I. ypiranganus Bernhauer 1912 S57
    |    |--Termitosymbia Seevers 1957 S57
    |    |    `--*T. nitida Seevers 1957 S57
    |    `--Termitozyras Seevers 1957 S57
    |         |--*T. adamsoni Seevers 1957 S57
    |         |--T. boliviae Seevers 1957 S57
    |         `--T. guianae Seevers 1957 S57
    `--Myrmedoniina [Myrmedoniae, Zyrasina] NT01
         |--Atemeles S57
         |--Zyras NT01
         |--Termozyras Cameron 1930 S57
         |    `--*T. politus Cameron 1930 S57
         |--Urodonia Silvestri 1946 S57
         |    `--*U. notabilis Silvestri 1946 S57
         |--Apalonia Casey 1906 NT01
         |    `--A. seticornis NT01
         |--Meronera Sharp 1887 [incl. Merona Sharp 1883 non Norman 1865] NT01
         |    `--M. venustula NT01
         |--Tetradonia Wasmann 1894 [incl. Chlorotusa Casey 1906] NT01
         |    `--T. megalops (Casey 1906) NT01
         |--Trachyota Casey 1906 NT01
         |    `--T. lativentris NT01
         `--Drusilla Leach 1819 [incl. Astilbus Dillwyn 1829, Myrmedonia Erichson 1837] NT01
              |--*D. canaliculata (Fabricius 1787) S57, NT01 [=Astilbus canaliculatus C01]
              |--D. cavicollis Casey 1906 NT01
              |--‘Myrmedonia’ clavigera Fauvel 1878 O86
              |--‘Myrmedonia’ humeralis C01
              |--D. impressicollis [incl. D. sculpticollis] S57
              |--D. incola S57
              |--‘Myrmedonia’ insignicornis Fauvel 1878 O86
              |--D. planaticollis S57
              `--‘Myrmedonia’ tristis Lucas 1846 E12

Lomechusini incertae sedis:
  Dinocoryna Casey 1893 [incl. Ecitonusa Wasmann 1897] NT01
    `--D. carolinensis NT01
  Ecitocala Frank 1981 NT01
    `--E. rugosa Frank 1981 NT01
  Ecitonidia Wasmann 1900 NT01
    `--E. wheeleri Wasmann 1900 NT01
  Ecitoxenidia Wasmann 1909 NT01
    `--E. brevicornis NT01
  Microdonia Casey 1893 NT01
    `--M. nitidiventris NT01
  Xesturida Casey 1906 NT01
    `--X. laevis Casey 1906 NT01

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[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.

[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.

[NT01] Newton, A. F., M. K. Thayer, J. S. Ashe & D. S. Chandler. 2001. Staphylinidae Latreille, 1802. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 272–418. CRC Press: Boca Raton.

[O86] Olliff, A. S. 1886. A revision of the Staphylinidae of Australia. Part I. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 403–473, pl. 7.

[S57] Seevers, C. H. 1957. A monograph on the termitophilous Staphylinidae (Coleoptera). Fieldiana Zoology 40: 1–334.

Staphylinini

Acylophorus pratensis, copyright Richard Migneault.

Belongs within: Staphylininae.
Contains: Amblyopinina, Cyrtoquediina, Indoquedius, Quedius, Philonthina, Staphylinina.

The Staphylinini are a cosmopolitan group of relatively large staphylinid beetles, including the rove beetles that are the most familiar members of the family. Representatives of the Staphylinini have antennae inserted closer to the eyes than to each other, a well developed apical maxillary palpomere, a relatively wide neck (if any), and no separate sclerotised plate anterior to the prosternum (Newton et al. 2001). A major clade within the group has been referred to as the 'Staphylinini propria', members of which are often distinguished by a dorsal basal ridge on the head capsule, a complete fusion of the pronotum and prosternum inside the procoxal cavity, and a transverse ridge on the hind coxa (Brunke et al. 2016).

Staphylinini [Quediini]
    |  i. s.: Valdiviodes BC16
    |         Alesiella BC16
    |         Quediomacrus BC16
    |         Lonia BC16
    |         Quelaestrygon BC16
    |         Strouhalium gracilicorne BC16
    |--Cretoquedius BC16
    `--+--Atanygnathus Jacobson 1909 NT01 (see below for synonymy)
       |    `--A. acuminatus BC16
       `--+--Antimerus punctipennis BC16
          |--+--Afroquedius sexpunctatus BC16
          |  `--Amblyopinina BC16
          `--+--Cyrtoquediina BC16
             `--+--+--Erichsonius Fauvel 1874 NT01 (see below for synonymy)
                |  |    |--E. nanus BC16
                |  |    `--E. patella N90
                |  `--Acylophorina BC16
                |       |  i. s.: Australotarsius grandis BC16
                |       |         Acylohsellus BC16
                |       |         Stevensia longipennis BC16
                |       |--Hemiquedius Casey 1915 BC16, NT01
                |       |    `--H. ferox (LeConte 1878) NT01
                |       `--+--Anaquedius Casey 1915 BC16, NT01
                |          |    `--A. vernix (LeConte 1878) NT01
                |          `--+--+--Anchocerus BC16
                |             |  `--Paratolmerus siamensis BC16
                |             `--Acylophorus Nordmann 1837 BC16, NT01 (see below for synonymy)
                |                  |--A. capensis BC16
                |                  |--A. pratensis BC16
                |                  `--A. wagenschieberi BC16
                `--+--Indoquedius BC16
                   `--+--Quediina BC16
                      |    |--Quedius BC16
                      |    |--Pseudorientis BC16
                      |    |--Velleiopsis BC16
                      |    `--Beeria Hatch 1957 NT01
                      |         `--B. nematocera (Casey 1915) [incl. B. punctata Hatch 1957] NT01
                      `--Staphylinini propria BC16
                           |  i. s.: Eucibdelus BC16
                           |         Ioma BC16
                           |         Weiserianum BC16
                           |         Haematodes BC16
                           |         Euristus BC16
                           |--+--Philothalpus BC16
                           |  |    |--P. bilobus BC16
                           |  |    `--P. falini BC16
                           |  `--Xanthopygina BC16
                           |       |  i. s.: Gastrisus Sharp 1876 NT01
                           |       |         Trigonopselaphus Gemminger & Harold 1868 (see below for synonymy) NT01
                           |       |--+--Isanopus sallaei BC16
                           |       |  `--+--Nordus fungicola BC16
                           |       |     `--Plociopterus BC16
                           |       `--+--Glenus flohri BC16
                           |          `--Xanthopygus Kraatz 1857 BC16, GN09 (see below for synonymy)
                           |               |--X. chapareanus BC16
                           |               `--X. xanthopygus Nordmann 1837 [incl. X. cacti Horn 1868] NT01
                           `--+--+--Philonthina BC16
                              |  `--Algon BC16
                              |       |--A. hollowayae BC16
                              |       `--A. oculatus BC16
                              `--+--Staphylinina BC16
                                 `--Anisolinina BC16
                                      |--+--Misantlius gebieni BC16
                                      |  `--Hesperosoma pederseni BC16
                                      `--+--Tolmerinus BC16
                                         `--+--Tympanophorus Nordmann 1837 BC16, NT01
                                            |    |--T. concolor Sharp 1884 NT01
                                            |    `--T. puncticollis Erichson 1840 (see below for synonymy) NT01
                                            `--+--Anisolinus BC16
                                               `--Pammegus ruficollis BC16

Acylophorus Nordmann 1837 BC16, NT01 [incl. Amacylophorus Smetana 1971 NT01, Palpacylophorus Smetana 1971 NT01]

Atanygnathus Jacobson 1909 NT01 [incl. Tanygnathinus Reitter 1909 NT01, Tanygnathus Erichson 1839 non Wagler 1832 NT01; Tanygnathinina BC16]

Erichsonius Fauvel 1874 NT01 [incl. Actobius Fauvel 1876 NT01, Sectophilonthus Tottenham 1949 NT01; Erichsoniina BC16]

Trigonopselaphus Gemminger & Harold 1868 [incl. Trigonophorus Nordmann 1837 nec Stephens 1829 nec Solier 1851] NT01

Tympanophorus puncticollis Erichson 1840 [incl. Xanthopygus borealis Hatch 1957, T. borealis] NT01

Xanthopygus Kraatz 1857 BC16, GN09 [incl. Heteropygus Bernhauer 1906 NT01, Lampropygus Sharp 1884 NT01]

*Type species of generic name indicated

REFERENCES

[BC16] Brunke, A. J., S. Chatzimanolis, H. Schillhammer & A. Solodovnikov. 2016. Early evolution of the hyperdiverse rove beetle tribe Staphylinini (Coleoptera: Staphylinidae: Staphylininae) and a revision of its higher classification. Cladistics 32 (4): 427–451.

[GN09] Grebennikov, V. V., & A. F. Newton. 2009. Good-bye Scydmaenidae, or why the ant-like stone beetles should become megadiverse Staphylinidae sensu latissimo (Coleoptera). European Journal of Entomology 106: 275–301.

[N90] Newton, A. F., Jr. 1990. Insecta: Coleoptera Staphylinidae adults and larvae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1137–1174. John Wiley & Sones: New York.

[NT01] Newton, A. F., M. K. Thayer, J. S. Ashe & D. S. Chandler. 2001. Staphylinidae Latreille, 1802. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 272–418. CRC Press: Boca Raton.

Last updated: 16 December 2020.

Aleochara

Aleochara curtula, copyright U. Schmidt.

Belongs within: Aleocharini.

Aleochara is a genus of aleocharine staphylinids characterised by a partially concealed head and five-segmented tarsi with the first segment much longer than the second. They are commonly associated with decaying matter (Olliff 1886).

Characters (from Olliff 1886): Mentum very short, narrowed and slightly emarginate in front. Ligula very short, broad, bifid. Maxillary palpi 5-segmented, the second and third segment subequal, the fifth very small. Labial palpi 4-segmented, the first segment longer than the others, the fourth very small. Mandibles unarmed. Labrum transverse, truncate in front. Head ovate, retracted, not narrowed behind. Antennae short, thickened towards the extremity, 1st, 2nd and 3rd segments elongate, the following, with the exception of the 11th, transverse. Prothorax broadly transverse, narrowed in front, rounded at the sides. Elytra slightly rounded behind. Abdomen either parallel-sided or narrowed behind. Legs rather short; intermediate coxae somewhat remote; tarsi 5:5:5, first segment of posterior pair distinctly longer than 2nd.

Aleochara Gravenhorst 1802 O86
    |--A. actae Olliff 1886 O86
    |--A. analis (Macleay 1871) [=Oxypoda analis] O86
    |--A. asiatica O86
    |--A. baliola Olliff 1886 O86
    |--A. bilineata P91
    |--A. bipunctata C01
    |--A. bisulcata (Redt. 1867) [=Oxypoda bisulcata] O86
    |--A. brachialis Jekel 1873 O86
    |--A. crassiuscula G89
    |--A. croceipennis Motschulsky 1858 [incl. A. maculipennis Kraatz 1859, A. sanguinipennis Kraatz 1859] O86
    |--A. curtula (Goeze 1777) M04
    |--A. fuscipes O86
    |--A. haemorrhoidalis O86
    |--A. insuavis Olliff 1886 O86
    |--A. lata Gravenhorst 1802 M04
    |--A. marginata Fauvel 1877 O86
    |--A. mastersi Macleay 1871 O86
    |--A. moesta C01
    |--A. nitida C01
    |--A. (Isochara) perturbans Fenyes 1921 F21
    |--A. puberula Klug 1883 [incl. A. armitagei Wollaston 1854, A. decorata Aubé 1850, A. dubia Fauvel 1863] O86
    |--A. punctum Fauvel 1878 O86
    |--A. scutellaris Lucas 1846 E12
    |--A. semirubra Fauvel 1877 O86
    |--A. speculifera Erichson 1842 O86
    |--A. subaenea ED04
    |--A. tristis C01
    `--A. vicina Olliff 1886 O86

*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] Band II. 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.

[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.

[ED04] Ewers, R. M., & R. K. Didham. 2004. A first assessment of New Zealand beetles restricted to large forest areas. Weta 27: 33–40.

[F21] Fenyes, A. 1921. New genera and species of Aleocharinae with a polytomic synopsis of the tribes. Bulletin of the Museum of Comparative Zoology 65 (2): 15–36.

[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.

[M04] Maruyama, M. 2004. Redescription of the genus Creochara (Coleoptera: Staphylinidae: Aleocharinae: Aleocharini) and its systematic position. Canadian Entomologist 136: 621–637.

[O86] Olliff, A. S. 1886. A revision of the Staphylinidae of Australia. Part I. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 403–473, pl. 7.

[P91] Petrushov, A. Z. 1991. Selection and mechanisms of ambush resistance in Metaseiulus occidentalis N. (Acarina: Phytoseiidae). In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 741–748. SPB Academic Publishing: The Hague.

Paederus

Paederus riparius, from here.

Belongs within: Paederini.

Paederus is a diverse genus of staphylinid beetles, generally coloured in contrasting patterns of red and black. Many species are found in damp habitats alongside rivers or in marshy fields. A number of species produce pederin and other haemolymph compounds that can cause severe dermatitis upon contacting human skin (Newton et al. 2001).

Characters (from Fagel 1961): Pronotum and prosternum not separate; weak prosternal extension; abdomen thick; pleurites well marked; sternite 1 with strong longitudinal carina; median lobe of aedeagus with ventral and dorsal lamella, median opening, parameres generally symmetrical, fused to median lobe over large part of their length and carrying numerous rows of bristles which do not reach the apex.

Paederus Fabricius 1775 NT01 (see below for synonymy)
    |--P. adelaidae Blackb. 1887 M96
    |--P. angusticeps F61
    |--P. australis B35
    |--P. caffer Boheman 1848 F61
    |--P. cruenticollis [incl. P. cingulatus] M96
    |--P. fastuosus F61
    |--P. fuscipes Curtis 1823 (see below for synonymy) F61
    |--P. gestroi M86
    |--P. impressipennis F61
    |--P. littoralis LC20
    |--P. littorarius LC20
    |--P. longipennis G89
    |--P. meyricki Blackb. 1891 M96
    |--P. nigripes Bernhauer 1932 F61
    |--P. nigrolineatus F61
    |--P. obliteratus N90
    |--P. riparius (Linnaeus 1758) B14
    |--P. rudebecki Fagel 1961 F61
    |--P. simsoni Blackb. 1894 M96
    `--P. termitophilus Wasmann 1911 S57

Paederus Fabricius 1775 NT01 [incl. Leucopaederus Casey 1905 NT01, Neopaederus Blackwelder 1939 F61, Paederillus Casey 1905 NT01, Paederomorphus Gautier des Cottes 1862 F61; Paederina]

Paederus fuscipes Curtis 1823 [incl. P. abyssinicus Cam. 1950, P. aestuans Erichson 1839-1840, P. alfierii Koch 1934, P. angolensis Er. 1843, P. erichsoni Wollaston 1867, P. mayumbeanus Cameron 1939] F61

*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.

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

[F61] Fagel, G. 1961. Coleoptera (Staphylinidae): Paederinae. In: Hanström, B., P. Brinck & G. Rudebeck (eds) South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 259–295. Almqvist & Wiksell: Uppsala.

[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.

[LC20] Lü, L., C.-Y. Cai, X. Zhang, A. F. Newton, M. K. Thayer & H.-Z. Zhou. 2020. Linking evolutionary mode to palaeoclimate change reveals rapid radiations of staphylinoid beetles in low-energy conditions. Current Zoology 66 (4): 435–444.

[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.

[M96] Masters, G. 1896. Catalogue of the described Coleoptera of Australia. Supplement, part II. Dytiscidae, Gyrinidae, Hydrophyllidae, Staphylinidae, Pselaphidae, Paussidae, Silphidae, Scaphididae, Histeridae, Phalacridae, Nitidulidae, Trogositidae, Colydiidae, Cucujidae, Cryptophagidae, Lathridiidae, Mycetophagidae, Dermestidae, Byrrhidae, Parnidae, Heteroceridae. Proceedings of the Linnean Society of New South Wales 21 (Suppl.): 695–754.

[N90] Newton, A. F., Jr. 1990. Insecta: Coleoptera Staphylinidae adults and larvae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1137–1174. John Wiley & Sones: New York.

[NT01] Newton, A. F., M. K. Thayer, J. S. Ashe & D. S. Chandler. 2001. Staphylinidae Latreille, 1802. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 272–418. CRC Press: Boca Raton.

[S57] Seevers, C. H. 1957. A monograph on the termitophilous Staphylinidae (Coleoptera). Fieldiana Zoology 40: 1–334.

Last updated: 10 April 2022.

Staphylinidae

Oxyporus maxillosus, copyright Nikolai Vladimirov.

Belongs within: Staphylinoidea.
Contains: Pseudopsinae, Scaphidiinae, Osoriinae, Piestinae, Oxytelinae, Proteininae, Omaliinae, Paederinae, Staphylininae, Mycetoporini, Phloeocharinae, Silphidae, Tachyporinae, Scydmaeninae, Aleocharinae, Steninae, Euaesthetinae, Leptotyphlinae, Pselaphinae, Thoracophorini.

The Staphylinidae is an extremely large family of usually small, gracile beetles with a flexible abdomen. Many staphylinids have reduced elytra that expose the greater part of a long, flexible abdomen, but there are notable exceptions. Some subfamilies, such as Scydmaeninae and many Omaliinae, have most or all of the abdomen covered by the elytra. Molecular phylogenetic analyses also indicate that some other groups with complete elytra classified to date in separate families, such as the Silphidae, may also be derived from within the Staphylinidae (McKenna et al. 2015). Staphylinids with complete elytra can usually be distinguished from other beetles by the presence of six visible abdominal ventrites and the lack of lobes on the fourth tarsomeres (Newton et al. 2001). Ocelli are commonly absent but are present in some subfamilies such as Omaliinae.

The majority of staphylinids are predatory, but a number of species feed on carrion, algae or fungi (Grimaldi & Engel 2005). Some, such as a number of members of the Aleocharinae and Pselaphinae, are specialised inquilines of social insects. They may have a flattened, limuloid body form that possibly provides physical protection from their hosts, or they may have a physogastric abdomen and/or exudatory trichomes that gives them a physical or chemical resemblance to the host insects (Seevers 1957). Some staphylinids are subsocial with adults caring for larvae in nests or burrows (Newton et al. 2001).

Subgroups of the Staphylinidae include the Apateticinae which have elytra covering most of the abdomen, lobed tarsomeres and gradually clavate antennae. The Habrocerinae have a very compact, shining body with few large setae and extremely slender, verticillate antennae. The genus Trichophya has similar antennae to the Habrocerinae but the body is more densely punctate and setose (Newton et al. 2001).

Staphylinidae [Oxyporidae, Oxytelidae, Piestides, Staphilinii, Staphyliniae]
    |--+--+--Apateticinae LC20
    |  |  |    |--Apatetica princeps LC20
    |  |  |    `--Nodynus Waterhous 1876 GN09
    |  |  |         `--N. leucofasciatus (Lewis 1879 GN09
    |  |  `--+--Habrocerinae LC20
    |  |     |    |--Nomimocerus Coiffait & Sáiz 1965 NT01
    |  |     |    `--Habrocerus Erichson 1839 NT01
    |  |     |         |--H. capillaricornis (Gravenhorst 1806) NT01
    |  |     |         |--H. magnus LeConte 1878 NT01
    |  |     |         `--H. schwarzi Horn 1877 NT01
    |  |     `--+--Trichophya Mannerheim 1830 NT01 [incl. Eumitocerus Casey 1886 NT01; Trichophyinae LC20]
    |  |        |    |--T. pilicornis (Gyllenhal 1810) NT01
    |  |        |    `--T. tarsalis LC20
    |  |        `--Trigonurinae LC20
    |  |             |--Trigonurus Mulsant 1847 GN09
    |  |             |    |--T. caelatus LeConte 1874 GN09
    |  |             |    `--T. crotchii LC20
    |  |             `--Abolescus glabratus LC20
    |  `--+--+--Pseudopsinae LC20
    |     |  `--Scaphidiinae LC20
    |     `--+--Osoriinae LC20
    |        `--+--Piestinae LC20
    |           `--Oxytelinae LC20
    `--+--+--Micropeplinae [Micropeplidae] LC20
       |  |    |--Peplomicrus mexicanus Campbell 1978 B14
       |  |    |--Micropeplus Latreille 1809 NT01
       |  |    |    |--M. dokuchaevi GE05
       |  |    |    |--M. fulvus LC20
       |  |    |    `--M. punctatus LC20
       |  |    |--Kalissus LeConte 1874 NT01
       |  |    |    `--K. nitidus LeConte 1874 NT01
       |  |    `--Protopeplus cretaceus LC20
       |  `--+--Proteininae LC20
       |     `--Omaliinae LC20
       `--+--+--+--Paederinae LC20
          |  |  `--Staphylininae LC20
          |  `--Olisthaerus Dejean 1833 LC20, NT01 [Olisthaerinae]
          |       |--O. megacephalus (Zetterstedt 1828) LC20, NT01
          |       `--+--O. substriatus (Gyllenhal 1827) LC20, NT01
          |          `--Mycetoporini LC20
          `--+--+--Phloeocharinae LC20
             |  `--+--Silphidae LC20
             |     `--Tachyporinae LC20
             `--+--+--Scydmaeninae LC20
                |  `--Solieriinae LC20
                |       |--Solierius Bernhauer 1921 GN09
                |       |    `--S. obscurus (Solier 1849) B14
                |       `--Prosolierius B14
                |            |--P. crassicornis B14
                |            |--P. mixticornis B14
                |            `--P. tenuicornis B14
                `--+--Aleocharinae LC20
                   `--+--+--+--Steninae LC20
                      |  |  `--Euaesthetinae LC20
                      |  `--+--+--Megalopinus Eichelbaum 1915 GN09 (see below for synonymy)
                      |     |    |--M. caelatus (Gravenhorst 1802) NT01
                      |     |    |--M. cruciger (Sharp 1886) B14
                      |     |    |--M. rufipes (LeConte 1863) NT01
                      |     |    `--M. sanguinitriguttatus (Scheerpeltz 1972) GN09
                      |     `--+--Leptotyphlinae LC20
                      |        `--Oxyporus Fabriciaus 1775 GN09 (see below for synonymy)
                      |             |--O. femoralis BC16
                      |             |--O. lateralis Gravenhorst 1802 GN09
                      |             |--O. mannerheimi C01
                      |             |--O. maxillosus B14
                      |             |--O. occipitalis Fauvel 1864 GN09
                      |             |--O. rufipennis LeConte 1963 GN09
                      |             |--O. rufus (Linnaeus 1758) B14 [=Staphylinus rufus G20]
                      |             |--O. vittatus NT01
                      |             `--O. yixianus LC20
                      `--+--+--Neophonus [Neophoninae] LC20
                         |  |    `--N. bruchi Fauvel 1905 B14
                         |  `--+--Dasycerinae [Dasyceridae] LC20
                         |     |    |--Dasycerus Brongniart 1800 B00
                         |     |    |    |--*D. sulcatus Brongniart 1800 B00
                         |     |    |    |--D. angulicollis LC20
                         |     |    |    |--D. bicolor B14
                         |     |    |    `--D. carolinensis Horn 1882 B14
                         |     |    `--Protodasycerus aenigmaticus LC20
                         |     `--Pselaphinae LC20
                         `--+--Charhyphus Sharp 1887 LC20, NT01 (see below for synonymy)
                            |    |--C. arizonensis Herman 1972 NT01
                            |    `--C. picipennis (LeConte 1863) NT01
                            `--+--+--Thoracochirus LC20
                               |  `--Plastus LC20
                               |       |--P. magnificus LC20
                               |       `--+--P. miles LC20
                               |          `--P. unicolor LC20
                               `--+--Neolispinus LC20
                                  `--+--Eleusinini NT01
                                     |    |--Renardia Motschulsky 1865 [incl. Eumalus Sharp 1887] NT01
                                     |    |    |--R. canadensis (Horn 1871) NT01
                                     |    |    `--R. nigrella (LeConte 1863) [incl. R. jubilaea Motschulsky 1865] NT01
                                     |    `--Eleusis Laporte 1835 LC20, NT01 [incl. Isomalus Erichson 1839 NT01]
                                     |         |--E. humilis (Erichson 1840) [incl. E. fasciata (LeConte 1863)] NT01
                                     |         |--E. mutica LC20
                                     |         |--E. pallida (LeConte 1863) NT01
                                     |         `--E. parva Blackb. 1892 M96
                                     `--Thoracophorini LC20

Staphylinidae incertae sedis:
  Paratorchus microphthalmus (Fauvel 1900) E02
  Stenomalium E02
  Anicula inferna P02
  Caririderma S02
  Prorhinopsenius Zhe02
  Brevicula gradus Whalley 1985 Zha02
  Achenium G89
    |--A. cribriceps Fauv. 1886 G89
    |--A. distinctum Lucas 1846 E12
    `--A. haemorrhoidale Lucas 1846 E12
  Hesperomimus DK08
  Canariobolitobius Schülke 2004 F05
    `--*C. filicornis (Wollaston 1864) [=Bolitobius filicornis] F05
  Paratyphlus maestratensis Hernando & Fancello 2004 F05
  ‘Heterosoma’ Bernhauer 1903 non Schaum 1845 LT61
  Protopselaphus Newton & Thayer 1995 [Protopselaphinae] NG05
  Ischnoglossa prolixa O86
  Coptolimulus B74
  Coptophilus B74
  Eulissus chalcopterus B70
  Myrmelibia kingi NT01
  Pareiobledius pruinosus NT01
  Lamprinus RD77
  Bembicidiodes P92
  Planeustomus P92
  Palaeopsenius Seevers 1971 P92
    `--P. mexicanus Seevers 1971 P92
  Paracyptus Seevers 1971 P92
    `--P. minutissima Seevers 1971 P92
  Paracraspedomerus Moore 1960 [=Xystridiogaster Scheerpeltz 1966] M72
    `--*P. speculifrons [=Cafius speculifrons, *Xystridiogaster speculifrons] M72

Charhyphus Sharp 1887 LC20, NT01 [incl. Pseudeleusis Bernhauer 1923 NT01, Triga Fauvel 1878 non Gray 1867 NT01, Trigites Handlirsch 1907 NT01]

Megalopinus Eichelbaum 1915 GN09 [incl. Megalops Erichson 1839 non Lacepède 1803 NT01, Megalopsidia Leng 1918 NT01; Megalopsidiinae LC20]

Oxyporus Fabriciaus 1775 GN09 [incl. Pseudoxyporus Nakane & Sawada 1956 NT01; Oxyporinae LC20]

*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).

[B70] Britton, E. B. 1970. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 495–621. Melbourne University Press.

[B74] Britton, E. B. 1974. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 62–89. Melbourne University Press.

[B00] Brongniart, A. 1800. Sur un nouveau genre d'insecte des environs de Paris. Bulletin des Sciences, par la Societé Philomathique de Paris 2 (39): 115–116.

[BC16] Brunke, A. J., S. Chatzimanolis, H. Schillhammer & A. Solodovnikov. 2016. Early evolution of the hyperdiverse rove beetle tribe Staphylinini (Coleoptera: Staphylinidae: Staphylininae) and a revision of its higher classification. Cladistics 32 (4): 427–451.

[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.

[DK08] Dohm, P., D. Kovač, A. Freidberg & R. Bin Hashim. 2008. Biology of the Oriental bamboo-inhabiting fly Felderimyia gombakensis and observations on mating trophallaxis in Felderimyia (Insecta, Diptera, Tephritidae, Phytalmiinae, Acanthonevrini). Senckenbergiana Biologica 88 (2): 311–318.

[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.

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261–282.

[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.

[GN09] Grebennikov, V. V., & A. F. Newton. 2009. Good-bye Scydmaenidae, or why the ant-like stone beetles should become megadiverse Staphylinidae sensu latissimo (Coleoptera). European Journal of Entomology 106: 275–301.

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

[LT61] Loeblich, A. R., Jr & H. Tappan. 1961. Remarks on the systematics of the Sarkodina (Protozoa), renamed homonyms and new and validated genera. Proceedings of the Biological Society of Washington 74: 213–234.

[LC20] Lü, L., C.-Y. Cai, X. Zhang, A. F. Newton, M. K. Thayer & H.-Z. Zhou. 2020. Linking evolutionary mode to palaeoclimate change reveals rapid radiations of staphylinoid beetles in low-energy conditions. Current Zoology 66 (4): 435–444.

[M96] Masters, G. 1896. Catalogue of the described Coleoptera of Australia. Supplement, part II. Dytiscidae, Gyrinidae, Hydrophyllidae, Staphylinidae, Pselaphidae, Paussidae, Silphidae, Scaphididae, Histeridae, Phalacridae, Nitidulidae, Trogositidae, Colydiidae, Cucujidae, Cryptophagidae, Lathridiidae, Mycetophagidae, Dermestidae, Byrrhidae, Parnidae, Heteroceridae. Proceedings of the Linnean Society of New South Wales 21 (Suppl.): 695–754.

[M72] Moore, I. 1972. Xystridiogaster Scheerpeltz, a synonym of Paracraspedomerus Moore (Coleoptera: Staphylinidae). Pan-Pacific Entomologist 48 (3): 218.

[NT01] Newton, A. F., M. K. Thayer, J. S. Ashe & D. S. Chandler. 2001. Staphylinidae Latreille, 1802. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 272–418. CRC Press: Boca Raton.

[NG05] Nomura, S., & I. Abd Ghani. 2005. Faunistic notes on the pselaphine species of the supertribes Goniaceritae, Pselaphitae and Clavigeritae from Malaysia and Singapore (Coleoptera: Staphylinidae: Pselaphinae). Serangga 10 (1–2): 1–36.

[O86] Olliff, A. S. 1886. A revision of the Staphylinidae of Australia. Part I. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 403–473, pl. 7.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[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.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[S02] Shcherbakov, D. E. 2002. Order Forficulida Latreille, 1810. The earwigs and protelytropterans (=Dermaptera DeGeer, 1773 +Protelytroptera Tillyard, 1931). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 288–291. Kluwer Academic Publishers: Dordrecht.

[Zha02] Zhang, J.-F. 2002. The most primitive earwigs (Archidermaptera, Dermaptera, Insecta) from the Upper Jurassic of Nei Monggol Autonomous Region, northeastern China. Acta Micropalaeontologica Sinica 19 (4): 348–362.

[Zhe02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 331–388. Kluwer Academic Publishers: Dordrecht.

Last updated: 10 April 2022.