Formicidae

Workers of Leptanilla, photographed by Alex Wild.

Belongs within: Euaculeata.
Contains: Amblyoponinae, Proceratiinae, Ponerinae, Dorylinae, Cerapachys, Ectatommini, Myrmicinae, Formicinae, Myrmeciinae, Pseudomyrmecinae, Dolichoderinae.

The Formicidae are the ants, a diverse and familiar clade of social wasps with a caste division between wingless workers and winged reproductives, with the female reproductives (the 'queens') shedding their wings after mating. An elongate scape (first antennal segment) is present in living ants, but was absent in the Cretaceous Sphecomyrminae (Grimaldi & Engel 2005). The morphology of Sphecomyrma indicates that it was an epigean (above-ground) forager, but two of the most basally diverging living ant lineages, the Leptanillinae and Martialis heureka, are hypogean (below-ground) foragers with reduced pigmentation and eyes. Leptanillinae are found in the tropics of the Old World and are characterised by a two-segmented petiole and twelve-segmented antennae without frontal lobes concealing the base.

Other small subfamilies of ants include the Agroecomyrmecinae, little-known ants from Africa and South America with eyes placed far back on the head and antennae spaced widely apart. The shape of the gaster in Agroecomyrmecinae is also distinctive, with abdominal sternite 4 reduced while the corresponding tergite is much larger and vaulted.

Synapomorphies (from Grimaldi & Engel 2005): Eusocial habits, with wingless female caste(s) present; metapleural gland opening on propodeum; petiole present.

<==Formicidae [Formicariae, Formiciformes, Formicoidea, Paleosminthuridae, Poneridae]
    |--Sphecomyrminae [Sphecomyrmidae] BS06
    |    |--Cretomyrma WB03
    |    |--Baikuris mandibularis Dlussky 1987 RJ93
    |    |--Dlusskyidris zherichini WB03, GE05
    |    |--Paleomyrmex Dlussky 1975 H02
    |    |    `--P. zherichini H02
    |    `--Sphecomyrma Wilson & Brown 1967 RBV08, V10
    |         |--S. canadensis GE05
    |         `--S. freyi RBV08
    `--+--Martialis Rabeling & Verhaagh in Rabeling, Brown & Verhaagh 2008 [Martialinae] RBV08
       |    `--*M. heureka Rabeling & Verhaagh in Rabeling, Brown & Verhaagh 2008 RBV08
       `--+--Leptanillinae RBV08
          |    |--Protanilla RBV08
          |    `--Leptanilla Emery 1870 FT08
          |         |--*L. revelierii Emery 1870 TB85
          |         |--L. japonica Baroni Urbani 1977 I92
          |         |--L. kubotai Baroni Urbani 1977 I92
          |         |--L. morimotoi Yasumatsu 1960 I92
          |         |--L. oceanica Baroni Urbani 1977 I92
          |         |--L. swani Wheeler 1932 TB85
          |         `--L. tanakai Baroni Urbani 1977 I92
          `--+--+--Tatuidris [Agroecomyrmecinae] RBV08
             |  `--Amblyoponinae RBV08
             `--+--+--Proceratiinae RBV08
                |  `--Paraponera [Paraponerinae, Paraponerini] RBV08
                |       `--P. clavata RBV08
                `--+--Ponerinae RBV08
                   `--+--+--Dorylinae BS06
                      |  `--Cerapachyinae [Cerapachyini, Cerapachysii, Prodorylinae] WW90
                      |       |--Cerapachys BS06
                      |       `--+--Leptanilloides BS06
                      |          |    |--L. mckennae BS06
                      |          |    `--L. nomada BS06
                      |          `--+--Cylindromyrmex striatus BS06
                      |             `--Acanthostichus BS06
                      |                  |--A. kirbyi BS06
                      |                  `--A. texanus WW90
                      `--+--Dolichoformica helferi GE05
                         `--+--+--Ectatommini BS06
                            |  `--+--Myrmicinae BS06
                            |     `--Formicinae BS06
                            `--+--+--Myrmeciinae BS06
                               |  `--Pseudomyrmecinae BS06
                               `--+--Dolichoderinae BS06
                                  `--Aneuretinae BS06
                                       |--Aneuretellus GE05
                                       `--Aneuretus Emery 1893 B03
                                            `--*A. simoni Emery 1893 B03

Formicidae incertae sedis:
  Huberia striata (Smith 1876) H82
  Brownimecia clavata GE05
  Haidomyrmex cerberus GE05
  Strongylognathus E89a
    |--S. afer E89a
    |--S. caeciliae ES12
    |--S. christophi Emery 1889 E89a
    |--S. huberi E89a
    |--S. koreanus Pisarski 1965 I92
    `--S. testaceus A71
  Anergates atratulus A71
  Eoponera berryi GE05
  Gerontoformica cretacica GE05
  Canapone dentata GE05
  Protopone GE05
  Oxyopomyrmex E89a
    |--O. oculatus E89a
    `--O. saulcyi Emery 1889 E89a
  Trapeziopelta maligna E89b
  Lordomyrmex caledonica Andre 1889 E14
  Sphecomyrmodes occidentalis Perrichot, Nel et al. 2007 V10
  Labidus praedator B10
  Eusphinctus Emery 1893 TB85
  Chalepoxenus muellerianus ES12
  Cretacoformica explicata Jell & Duncan 1986 Z01
  Formicium giganteum S12
  Eomyrmex Hong 1974 H02
    `--E. guchengziensis Z01
  Aeromyrma sophiae Emery 1890 H02

*Type species of generic name indicated

REFERENCES

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

[B03] Bingham, C. T. 1903. The Fauna of British India, including Ceylon and Burma. Hymenoptera vol. 2. Ants and Cuckoo-Wasps. Taylor and Francis: London.

[BS06] Brady, S. G., T. R. Schultz, B. L. Fisher & P. S. Ward. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the USA 103 (48): 18172–18177.

[B10] Brown, B. V. 2010. Phoridae (hump-backed flies, scuttle flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 725–761. NRC Research Press: Ottawa.

[E89a] Emery, C. 1889a. Intorno ad alcune formiche della fauna palearctica. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 439–443.

[E89b] Emery, C. 1889b. Viaggio di Leonardo Fea in Birmania e regioni vicine. XX.—Formiche di Birmania e del Tenasserim raccolte da Leonardo Fea (1885–87). Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 485–520.

[E14] Emery, C. 1914. Les fourmis de la Nouvelle-Calédonie & des îles Loyalty. In: Sarasin, F., & J. Roux (eds) Nova Caledonia: Forschungen in Neu-Caledonian und auf den Loyalty-Inseln. A. Zoologie vol. 1 pt 4 pp. 393–436, pl. 12. C. W. Kreidels Verlag: Wiesbaden.

[ES12] Espadaler, X., & S. Santamaria. 2012. Ecto- and endoparasitic fungi on ants from the Holarctic region. Psyche 2012: 168478.

[FT08] Framenau, V. W., & M. L. Thomas. 2008. Ants (Hymenoptera: Formicidae) of Christmas Island (Indian Ocean): identification and distribution. Records of the Western Australian Museum 25 (1): 45–85.

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

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

[H02] Hong Y. 2002. Amber Insect of China. Beijing Scientific and Technological Publishing House.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[RBV08] Rabeling, C., J. M. Brown & M. Verhaagh. 2008. Newly discovered sister lineage sheds light on early ant evolution. Proceedings of the National Academy of Sciences of the USA 105 (39): 14913–14917.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[S12] Schaal, S. 2012. Messel Pit fossil site—the legacy of the environment and life of the Eocene. In: Talent, J. A. (ed.) Earth and Life: Global biodiversity, extinction intervals and biogeographic perturbations through time pp. 225–236. Springer.

[TB85] Taylor, R. W., & D. R. Brown. 1985. Formicoidea. In: Walton, D. W. (ed.) Zoological Catalogue of Australia vol. 2. Hymenoptera: Formicoidea, Vespoidea and Sphecoidea pp. 1–149. Australian Government Publishing Service: Canberra.

[V10] Vršanský, P. 2010. Cockroach as the earliest eusocial animal. Acta Geologica Sinica (English Edition) 84 (4): 793–808.

[WB03] Ward, P. S., & S. G. Brady. 2003. Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera: Formicidae). Invertebrate Systematics 17: 361–386.

[WW90] Wheeler, G. C., & J. Wheeler. 1990. Insecta: Hymenoptera Formicidae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1277–1294. John Wiley & Sones: New York.

[Z01] Zhou, S. 2001. Ants of Guangxi. Guangxi Normal University Press: Guilin (China).

Last updated: 14 June 2020.

Trichosphaeriaceae

Trichosphaeria notabilis on Clematis vitalba, from Mycologues Associes.

Belongs within: Sordariomycetes.

The Trichosphaeriaceae are a family of saprobic fungi most commonly found on wood and bark (Cannon & Kirk 2007).

Characters (from Cannon & Kirk 2007): Stromata absent, or reduced to a hyphal subiculum. Ascomata perithecial, superficial, often aggregated, ± globose, black, often thick-walled, usually setose, the ostiole papillate, periphysate; peridium composed of thick-walled angular cells, sometimes larger and more strongly pigmented towards the apex. Interascal tissue of narrow, persistent, thin-walled, true paraphyses formed from a basal meristem. Asci cylindrical, persistent, thin-walled, not fissitunicate, usually with a small J- apical ring. Ascospores variously shaped, hyaline to versicoloured, usually septate and rarely muriform, sometimes fragmenting at the septa, without germ pores, sometimes with a sheath. Anamorphs varied, hyphomycetous, with simple pigmented conidiophores and often with complex conidia.

Trichosphaeriaceae [Trichosphaeriales]
    |--Acanthosphaeria EB03
    |--Collematospora EB03
    |--Coniobrevicolla larsenii EB03, E99
    |--Crassochaeta EB03
    |--Cresporhaphis EB03
    |--Eriosphaeria EB03
    |--Fluviostroma EB03
    |--Kananascus EB03
    |--Miyoshiella EB03
    |--Rhynchomeliola EB03
    |--Rizalia EB03
    |--Schweinitziella EB03
    |--Setocampanula EB03
    |--Trichosphaeria EB03
    |--Umbrinosphaeria EB03
    `--Neorehmia von Höhnel 1902 EB03, RS99 [incl. Larseniella Munk 1942 RS99]
         |--*N. ceratophora von Höhnel 1902 (see below for synonymy) RS99
         `--N. arecae [=Oplothecium arecae, Trichosphaerella arecae] RS99

*Neorehmia ceratophora von Höhnel 1902 [=Trichosphaerella ceratophora (von Höhnel) Müller in Müller & von Arx 1962; incl. *Larseniella globulispora Munk 1942] RS99

*Type species of generic name indicated

REFERENCES

Cannon, P. F., & P. M. Kirk. 2007. Fungal Families of the World. CABI.

[E99] Eriksson, O. E. (ed.) 1999. Notes on ascomycete systematics. Nos 2440-2755. Myconet 2: 1-41.

[EB03] Eriksson, O. E., H. O. Barah, R. S. Currah, K. Hansen, C. P. Kurtzman, G. Rambold & T. Laessøe (eds.) 2003. Outline of Ascomycota—2003. Myconet 9: 1-89.

[RS99] Rossman, A. Y., G. J. Samuels, C. T. Rogerson & R. Lowen. 1999. Genera of Bionectriaceae, Hypocreaceae and Nectriaceae (Hypocreales, Ascomycetes). Studies in Mycology 42: 1-248.

Strumigeniti

Strumigenys dicomas, copyright April Nobile/antweb.org.

Belongs within: Myrmicinae.

The Strumigeniti are a group of often litter-dwelling ants characterised by ventrolateral eyes and greatly reduced palps. Many species, known as trapjaw ants, have distinctive elongate mandibles. However, others have more ordinary mandibles and the elongate mandible form may have arisen on more than one occasion. In some genera the mandibles are massive and downturned whereas others have the mandibles extended into long narrow blades (Bolton 1983).

Characters (from Bolton 1983): Eyes ventrolateral, placed within or beneath scrobes which are universally present though shallow and reduced in some. Antennae 4- or 6-segmented, apical segment much the longest of the funiculars. Palp formula is 1,1.

<==Strumigeniti B83
    |--Neostruma B83
    |--Asketogenys B83
    |--Chelystruma B83
    |--Codioxenus B83
    |--Dorisidris B83
    |--Dysedrognathus B83
    |--Gymnomyrmex B83
    |--Tingimyrmex B83
    |--*Codiomyrmex thaxteri B83
    |--Kyidris Brown 1949 B83, Z01 [incl. Polyhomoa B83]
    |    `--*K. mutica Brown 1949 Z01
    |--Cladarogenys Brown 1976 B83
    |    `--*C. lasia Brown 1976 B83
    |--Pentastruma Forel 1912 B83, Z01
    |    |--*P. sauteri Forel 1912 Z01
    |    `--P. canina Brown & Boisvert 1978 Z01
    |--Epitritus Emery 1869 B83
    |    |--*E. argiolus Emery 1869 B83
    |    |--E. hexamerus B83
    |    |--E. laticeps Brown 1962 B83
    |    |--E. minimus Bolton 1972 B83
    |    |--E. murphyi B83
    |    |--E. roomi Bolton 1972 B83
    |    `--E. tiglath Bolton 1983 B83
    |--Serrastruma Brown 1948 B83
    |    |--*S. simoni (Emery 1895) (see below for synonymy) B83
    |    |--S. concolor (Santschi 1914) [=Strumigenys (Trichoscapa) concolor] B83
    |    |--S. dotaja Bolton 1983 B83
    |    |--S. geoterra Bolton 1983 B83
    |    |--S. inquilina Bolton 1983 B83
    |    |--S. ludovici (Forel 1904) (see below for synonymy) B83
    |    |--S. lujae (Forel 1902) (see below for synonymy) B83
    |    |--S. maynei (Forel 1916) (see below for synonymy) B83
    |    |--S. miccata Bolton 1983 B83
    |    |--S. serrula (Santschi 1910) (see below for synonymy) B83
    |    `--S. sulumana Bolton 1983 B83
    |--Glamyromyrmex Wheeler 1915 [incl. Borgmeierita Brown 1953] B83
    |    |--*G. beebei Wheeler 1915 TB85
    |    |--G. africanus Bolton 1983 B83
    |    |--G. crypturus Bolton 1983 B83
    |    |--G. dagon Bolton 1983 B83
    |    |--‘Codiomyrmex’ excisus Weber 1934 [=*Borgmeierita excisa] B83
    |    |--G. flagellatus (Taylor 1962) [=Codiomyrmex flagellatus] TB85
    |    |--G. loveridgei (Brown 1953) [=Codiomyrmex loveridgei] B83
    |    |--G. ravidurus Bolton 1983 B83
    |    |--G. sahurus Bolton 1983 B83
    |    |--G. semicomptus (Brown 1959) [=Codiomyrmex semicomptus] TB85
    |    |--G. sistrurus Bolton 1983 B83
    |    |--G. tetragnathus (Taylor 1965) [=Codiomyrmex tetragnathus] B83
    |    |--G. thuvidus Bolton 1983 B83
    |    |--G. trymalus Bolton 1983 B83
    |    `--G. tukultus Bolton 1983 B83
    |--Smithistruma Brown 1948 (see below for synonymy) B83
    |    |--*S. pulchella (Emery 1895) [=Strumigenys pulchella] B83
    |    |--S. alberti B83
    |    |--S. anarta Bolton 1983 B83
    |    |--S. arahana Bolton 1983 B83
    |    |--S. behasyla Bolton 1983 B83
    |    |--S. cavinasis Brown 1950 B83
    |    |--S. chyatha Bolton 1983 B83
    |    |--S. datissa Bolton 1983 B83
    |    |--S. dendexa Bolton 1983 B83
    |    |--S. depressiceps (Weber 1934) [=Strumigenys (Cephaloxys) depressiceps, Sm. (*Platystruma) depressiceps] B83
    |    |--S. dietrichi B83
    |    |--S. dubia B83
    |    |--S. emarginata (Mayr 1901) [=Strumigenys emarginata] B83
    |    |--S. enkara Bolton 1983 B83
    |    |--S. fenkara Bolton 1983 B83
    |    |--S. fulda Bolton 1983 B83
    |    |--S. gatuda Bolton 1983 B83
    |    |--S. hensekta Bolton 1983 B83
    |    |--S. hyphata B83
    |    |--S. impidora Bolton 1983 B83
    |    |--S. kerasma Bolton 1983 B83
    |    |--S. leptothrix (Wheeler 1929) [=Strumigenys (Cephaloxys) leptothrix, Sm. (*Weberistruma) leptothrix] B83
    |    |--S. malaplax Bolton 1983 B83
    |    |--S. mandibularis (Szabo 1909) [=Epitritus mandibularis, *Miccostruma mandibularis] B83
    |    |--S. marginata (Santschi 1914) [=Epitritus marginatus, Miccostruma marginata] B83
    |    |--S. mekaha Bolton 1983 B83
    |    |--S. minkara Bolton 1983 B83
    |    |--S. ninda Bolton 1983 B83
    |    |--S. nykara Bolton 1983 B83
    |    |--S. ornata B83
    |    |--S. oxysma Bolton 1983 B83
    |    |--S. pergrandei (Emery 1895) [=Strumigenys pergrandei, Sm. (*Wessonistruma) pergrandei] B83
    |    |--S. placora Bolton 1983 B83
    |    |--S. rostrata B83
    |    |--S. rusta Bolton 1983 B83
    |    |--S. sharra Bolton 1983 B83
    |    |--S. synkara Bolton 1983 B83
    |    |--S. tacta Bolton 1983 B83
    |    |--S. terroni Bolton 1983 B83
    |    |--S. tigrilla (Brown 1973) [=Miccostruma tigrilla] B83
    |    |--S. tolomyia Bolton 1983 B83
    |    |--S. transversa (Santschi 1913) [=Strumigenys transversa] B83
    |    |--S. truncatidens Brown 1950 B83
    |    |--S. vodensa Bolton 1983 B83
    |    `--S. weberi Brown 1959 B83
    `--Strumigenys Smith 1860 FT08 (see below for synonymy)
         |  i. s.: *S. mandibularis Smith 1860 TB85
         |         S. adrasora Bolton 1983 B83
         |         S. arnoldi Forel 1913 B83
         |         S. bernardi Brown 1960 B83
         |         S. bitheria Bolton 1983 B83
         |         S. bryanti N-ZLI11
         |         S. cacaoensis Bolton 1971 B83
         |         ‘*Cephaloxys’ capitata Smith 1865 [=Smithistruma capitata] B83
         |         S. dextra Brown 1954 B83
         |         S. dromoshaula Bolton 1983 B83
         |         S. dyshaula Bolton 1983 B83
         |         S. eggersi WW90
         |         S. emmae (Emery 1890) FT08 (see below for synonymy)
         |         S. emdeni Forel 1915 TB85
         |         S. ettillax Bolton 1983 B83
         |         ‘Quadristruma’ eurycera [=Epitritus eurycerus] B83
         |         S. faurei Arnold 1948 B83
         |         S. feae Emery 1894 B03
         |         S. ferocior Brown 1973 TB85
         |         S. formosensis Forel 1912 [=S. faea var. formosensis] Z01
         |         S. friedae Forel 1915 TB85
         |         S. godeffroyi Mayr 1866 FT08 [incl. S. godeffroyi var. indica B03]
         |         S. gundlachi WW90
         |         S. guttulata Forel 1902 TB85
         |         S. hastyla Bolton 1983 B83
         |         S. havilandi Forel 1905 B83
         |         S. helytruga Bolton 1983 B83
         |         S. hispida Lin & Wu 1996 Z01
         |         S. irrorata Santschi 1913 B83
         |         S. katapelta Bolton 1983 B83
         |         S. koningsbergeri N-ZLI11
         |         S. korahyla Bolton 1983 B83
         |         S. labidogenys N-ZLI11
         |         S. lacunosa Z01
         |         S. leptorhina Bolton 2000 Z01
         |         S. lewisi Cam. 1887 B03
         |         S. londianensis (Patrizi 1946) [=*Proscopomyrmex londianensis] B83
         |         S. loriae [incl. *Eneria excisa Donisthorpe 1948] B83
         |         S. lyroessa (Roger 1862) [=*Labidogenys lyroessa] B83
         |         S. marleyi Arnold 1914 [=S. havilandi race marleyi] B83
         |         S. mayri Emery 1897 TB85
         |         S. mesahyla Bolton 1983 B83
         |         S. murshila Bolton 1983 B83
         |         S. nimbrata Bolton 1983 B83
         |         S. omalyx Bolton 1983 B83
         |         S. opaca Brown 1954 TB85
         |         S. pallestes Bolton 1971 B83
         |         S. paranax Bolton 1983 B83
         |         S. perplexa (Smith 1876) [=Orectognathus perplexus; incl. S. leae Forel 1913] TB85
         |         S. petiolata Bernard 1952 B83
         |         S. pilosa Zhou 2001 Z01
         |         S. pretoriae Arnold 1949 B83
         |         S. quinquedentata Crawley 1923 TB85
         |         S. relahyla Bolton 1983 B83
         |         S. rogeri Emery 1890 [incl. S. incisa Godfrey 1907, S. sulfurea Santschi 1915] B83
         |         S. rufobrunea Santschi 1914 B83
         |         S. rukha Bolton 1983 B83
         |         S. sarissa Bolton 1983 B83
         |         S. scotti Forel 1912 B83
         |         S. shaula Bolton 1983 B83
         |         S. signaea N-ZLI11
         |         S. silvestrii WW90
         |         S. smythiesi Forel 1902 B03
         |         S. spathoda Bolton 1983 B83
         |         S. stygia Santschi 1913 B83
         |         S. szalayi Emery 1897 [incl. S. szalayi australis Forel 1910] TB85
         |         S. tetraphanes Brown 1954 B83
         |         S. totyla Bolton 1983 B83
         |         S. traegaordhi Santschi 1913 B83
         |         S. vazerka Bolton 1983 B83
         |         S. xenohyla Bolton 1983 B83
         |         S. xenos Brown 1955 TB85
         |         S. zandala Bolton 1983 B83
         |--S. propiciens SB08
         `--+--S. dicomas SB08
            `--Pyramica Roger 1862 SB08, FT08 [incl. Trichoscapa Emery 1869 B83]
                 |--*P. gundlachi Roger 1862 B83
                 |--P. hoplites SB08
                 |--P. jacobsoni N-ZLI11
                 `--P. membranifera (Emery 1869) FT08 (see below for synonymy)

Pyramica membranifera (Emery 1869) FT08 [=Strumigenys (*Trichoscapa) membranifera B83, S. (Cephaloxys) membranifera B83; incl. S. (Cephaloxys) foochowensis Wheeler 1928 B83, S. (Cephaloxys) membranifera var. marioni Wheeler 1933 B83, S. membranifera var. santschii Forel 1904 B83, S. (Cephaloxys) silvestriana Wheeler 1928 B83, S. membranifera race simillima Emery 1890 B83, S. (Cephaloxys) vitiensis Mann 1921 B83, S. (Cephaloxys) membranifera var. williamsi B83]

Serrastruma ludovici (Forel 1904) [=Strumigenys ludovici; incl. St. alluaudi Santschi 1910, Serrastruma alluaudi, St. (Cephaloxys) escherichi lotti Weber 1943, Se. lotti, St. (Trichoscapa) alluaudi st. nigeriensis Santschi 1914, St. rothkirchi Wasmann 1918] B83

Serrastruma lujae (Forel 1902) [=Strumigenys lujae; incl. St. (Cephaloxys) aequalis Menozzi 1942, St. (Xephaloxys [l. c.]) bequaerti Santschi 1923, Serrastruma bequaerti, St. (Cephaloxys) calypso Santschi 1923, Se. calypso, St. (Cephaloxys) gerardi Santschi 1923, St. (Cephaloxys) glanduscula Santschi 1919, St. reticulata Stitz 1910] B83

Serrastruma maynei (Forel 1916) [=Strumigenys (Trichoscapa) maynei; incl. St. (T.) maynei var. latiuscula Forel 1916] B83

Serrastruma serrula (Santschi 1910) [=Strumigenys lujae var. serrula; incl. St. (Cephaloxys) uelensis Santschi 1923] B83

*Serrastruma simoni (Emery 1895) [=Strumigenys simoni, Smithistruma (*Serrastruma) simoni; incl. St. biconvexa Santschi 1913, St. cognata st. boerorum Santschi 1913, St. escherichi var. cliens Forel 1913, St. cognata Santschi 1910, St. escherichi Forel 1910, St. (Trichoscapa) escherichi st. cognata var. fusciventris Santschi 1915 (nom. inv.), St. escherichi race limbata Forel 1913, St. (T.) escherichi st. cognata var. obscuriventris Santschi 1914 (nom. inv.), St. (Cephaloxys) raymondi Donisthorpe 1945] B83

Smithistruma Brown 1948 [incl. Miccostruma Brown 1948, Platystruma Brown 1953, Weberistruma Brown 1948, Wessonistruma Brown 1948] B83

Strumigenys Smith 1860 FT08 [incl. Cephaloxys Smith 1864 non Signoret 1847 Z01, Eneria Donisthorpe 1948 B83, Labidogenys Roger 1862 B83, Proscopomyrmex Patrizi 1946 B83, Quadristruma Brown 1949 TB85]

Strumigenys emmae (Emery 1890) FT08 [=Epitritus emmae TB85, *Quadristruma emmae TB85, FT08; incl. E. clypeatus Szabo 1909 B83, E. clypeatus var. malesiana Forel 1913 B83, E. wheeleri Donisthorpe 1916 B83]

*Type species of generic name indicated

REFERENCES

[B03] Bingham, C. T. 1903. The Fauna of British India, including Ceylon and Burma. Hymenoptera vol. 2. Ants and Cuckoo-Wasps. Taylor and Francis: London.

[B83] Bolton, B. 1983. The Afrotropical dacetine ants (Formicidae). Bulletin of the British Museum (Natural History): Entomology series 46 (4): 267–416.

[FT08] Framenau, V. W., & M. L. Thomas. 2008. Ants (Hymenoptera: Formicidae) of Christmas Island (Indian Ocean): identification and distribution. Records of the Western Australian Museum 25 (1): 45–85.

[N-ZLI11] Nur-Zati A. M., G. T. Lim & A. B. Idris. 2011. Checklist of ants of selected hill dipterocarp forests of peninsular Malaysia. Serangga 16 (1): 91–105.

[SB08] Schultz, T. R. & S. G. Brady. 2008. Major evolutionary transitions in ant agriculture. Proceedings of the National Academy of Sciences of the USA 105 (14): 5435–5440.

[TB85] Taylor, R. W., & D. R. Brown. 1985. Formicoidea. In: Walton, D. W. (ed.) Zoological Catalogue of Australia vol. 2. Hymenoptera: Formicoidea, Vespoidea and Sphecoidea pp. 1–149. Australian Government Publishing Service: Canberra.

[WW90] Wheeler, G. C., & J. Wheeler. 1990. Insecta: Hymenoptera Formicidae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1277–1294. John Wiley & Sones: New York.

[Z01] Zhou, S. 2001. Ants of Guangxi. Guangxi Normal University Press: Guilin (China).

Last updated: 31 May 2020.

Attini

Worker of Trachymyrmex septentrionalis tending fungus, photographed by Alex Wild.

Belongs within: Myrmicinae.

The Attini, including the leafcutter ants and allied taxa, are a clade of ants characterised by the cultivation of fungi for food (Schultz & Brady 2008). Female attins, upon leaving the parental nest, carry a sample of their food fungus with them in order to start a culture in their new nest. The ancestral attins cultivated fungi of the Leucocoprineae, but one subgroup of the genus Apterostigma cultivates coral fungi of the Pterulaceae. Leafcutter ants, including the genera Atta and Acromyrmex, cultivate a single derived fungus species on pieces of leaves that they collect and carry back to their nest (Schultz & Brady 2008).

<==Attini
    |--Paleoattini SB08
    |    |--Apterostigma SB08
    |    |    |  i. s.: A. electropilosum SB08
    |    |    |         A. pilosum SB08
    |    |    |--A. auriculatum SB08
    |    |    `--+--A. collare SB08
    |    |       |--+--A. dentigerum SB08
    |    |       |  `--A. dorotheae SB08
    |    |       `--+--A. goniodes SB08
    |    |          `--A. manni SB08
    |    `--+--Mycocepurus SB08
    |       |    |--M. smithi SB08
    |       |    `--+--M. curvispinosus SB08
    |       |       `--M. tardus SB08
    |       `--Myrmicocrypta SB08
    |            |--M. infuscata SB08
    |            `--+--M. buenzlii SB08
    |               `--+--M. ednaella SB08
    |                  `--M. urichi SB08
    `--Neoattini SB08
         |--Mycetophylax emeryi SB08
         `--+--Mycetarotes SB08
            |    |--M. acutus SB08
            |    `--M. parallelus SB08
            `--Mycetosoritis SB08
                 |--M. hartmanni SB08
                 `--+--M. clorindae SB08
                    `--Cyphomyrmex SB08
                         |  i. s.: C. maya SB08
                         |         C. taino SB08
                         |--+--+--C. fanulus SB08
                         |  |  `--+--‘Mycetophylax’ conformis SB08
                         |  |     `--C. morschi SB08
                         |  `--+--C. rimosus SB08
                         |     `--+--C. cornutus SB08
                         |        `--C. minutus SB08
                         `--+--+--C. costatus SB08
                            |  `--+--C. longiscapus SB08
                            |     `--C. muelleri SB08
                            `--+--Mycetagroicus triangularis SB08
                               `--Trachymyrmex SB08
                                    |  i. s.: T. neomexicanus WW90
                                    |         T. nogalensis WW90
                                    |         T. primaevus SB08
                                    |         T. seminole H79
                                    |         T. urichi H79
                                    |--+--Sericomyrmex parvulus SB08
                                    |  `--+--T. papulatus SB08
                                    |     `--+--T. opulentus SB08
                                    |        `--T. zeteki SB08
                                    `--+--+--T. intermedius SB08
                                       |  `--+--T. diversus SB08
                                       |     `--+--T. cornetzi SB08
                                       |        `--+--T. bugnioni SB08
                                       |           `--T. irmgardae SB08
                                       `--+--+--T. arizonensis SB08
                                          |  `--+--T. septentrionalis SB08
                                          |     `--T. smithi SB08
                                          `--+--Atta SB08
                                             |    |  i. s.: A. dissimilis Jerdon 1851 (n. d.) B03
                                             |    |         A. domicola Jerdon 1851 (n. d.) B03
                                             |    |         A. sexdens H79
                                             |    |--+--A. cephalotes SB08
                                             |    |  `--A. laevigata SB08
                                             |    `--+--A. mexicana SB08
                                             |       `--A. texana SB08
                                             `--Acromyrmex BS06
                                                  |  i. s.: A. lobicornis CM07
                                                  |--A. versicolor SB08
                                                  `--+--A. octospinosus SB08
                                                     `--+--Pseudoatta SB08
                                                        |--+--A. balzani SB08
                                                        |  `--A. landolti SB08
                                                        `--+--A. heyeri SB08
                                                           `--A. lundi SB08

*Type species of generic name indicated

REFERENCES

[B03] Bingham, C. T. 1903. The Fauna of British India, including Ceylon and Burma. Hymenoptera. Vol. II. Ants and Cuckoo-Wasps. Taylor and Francis: London.

[BS06] Brady, S. G., T. R. Schultz, B. L. Fisher & P. S. Ward. 2006. Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences of the USA 103 (48): 18172–18177.

[CM07] Curtis, D. J., & G. Machado. 2007. Ecology. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 280–308. Harvard University Press: Cambridge (Massachusetts).

[H79] Howse, P. E. 1979. The uniqueness of insect societies: Aspects of defense and integration. In: Larwood, G. & B. R. Rosen (eds) Biology and Systematics of Colonial Organisms pp. 345–374. Academic Press: London.

[SB08] Schultz, T. R. & S. G. Brady. 2008. Major evolutionary transitions in ant agriculture. Proceedings of the National Academy of Sciences of the USA 105 (14): 5435–5440.

[WW90] Wheeler, G. C., & J. Wheeler. 1990. Insecta: Hymenoptera Formicidae. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1277–1294. John Wiley & Sones: New York.

Scolytinae

Female Phloeosinus dentatus, copyright N. Wright.

Belongs within: Curculionidae.
Contains: Xyloterini, Micracini, Bothrosternini, Scolytus, Diamerini, Scolytodes, Phloeotribus, Hylurgops, Tomicini, Polygraphini, Crypturgini, Pityophthorina, Corthylina, Cryphalus, Xyloctonini, Hypoborini, Ips, Hypothenemus, Scolytogenes, Cyrtogenius, Xyleborus.

The Scolytinae, bark beetles, are a group of small weevils that live and feed in the subcortical tissues of plants. They are characterised by a reduced rostrum, geniculate antennae with a distinct, compact club, and tibiae marginally armed by spines or socketed denticles (Wood 1986). The majority of species live in galleries in the phloem layer of their hosts but some feed on other tissues such as xylem or seeds. Some species feed on ambrosia fungi growing on host xylem. Many bark beetles are monogamous with a single mating pair living in a gallery; eggs are laid within the gallery and the parents guard the burrow entrance while the larvae develop. Other species exhibit various modes of polygyny.

Wood (1986) recognised the Scolytidae as a separate family from the Curculionidae, divided between two subfamilies Scolytinae and Hylesininae on the basis of features such as form of the pronotum and base of the elytra, but more recent studies have not supported monophyly of the latter two groups. Tribes of the Scolytinae include the Scolytini, members of which are most diverse in the Americas and lack armature on the lateral margins of the pro- and metatibiae except for a single, apical, spinelike process curving towards and extending beyond the process of the inner apical angle (Wood 1986).

Characters (from Wood 1986, as Scolytidae): Adult with head narrower than pronotum; eyes nearly flat against head, often emarginate to divided; pregular sutures present and reinforced internally by massive apodemal ridges; rostrum short to nonexistent; hypostomal area small, posterior mandibular condyle more anterior in position; axis of mandibular hinge oblique to transverse, cutting edge on mesal margin; pronotum often asperate, almost never with constriction near middle; when visible, scutellum usually flush with elytral surface, often flattened; metepisternum stout to elongate, anterior margin usually procurved, anteroventral angle displaced slightly caudad and distinctly extended ventrad; mesepimeron flat, oblique; mesepisternum smaller, declivous cephalad; tibiae armed on apical and/or lateral margins either by spines or socketed denticles, apical mucro formed by mesal element; tarsal segment 1 about equal in length to 2 or 3; male spiculum gastrale present. Larva with clypeus distinct, separate from frons; frontal sutures never reaching articulating membrane of mandibles.

<==Scolytinae [Hylesininae, Scolytarii] D09
    |--Scolytini [Camptoceridae] GC-P14
    |    |--Scolytus W86
    |    |--Scolytopsis Blandford 1896 W86
    |    |    `--*S. puncticollis W86
    |    |--Camptocerus Latreille 1829 W86
    |    |    |--*C. aeneipennis [=Hylesinus aeneipennis] W86
    |    |    `--C. auricomus W86
    |    `--Cnemonyx Eichhoff 1868 (see below for synonymy) W86
    |         |--*C. galeritus W86
    |         |--C. barbatus [=*Minulus barbatus] W86
    |         |--C. flavicornis [=*Loganius flavicornis] W86
    |         |--C. hylurgoides [=*Coptodryas hylurgoides, *Coptosomus hylurgoides] W86
    |         |--C. jucundus [=*Ceratolepis jucundus] W86
    |         `--C. panamensis W86
    `--+--+--+--Diamerini GC-P14
       |  |  `--+--Hexacolini [Ctenophoridae, Ctenophorini, Hexacolidae, Problechilidae] GC-P14
       |  |     |    |--Scolytodes W86
       |  |     |    |--Gymnochilus Eichhoff 1867 [=Problechilus Eichhoff 1878; incl. Meringopalpus Hagedorn 1905] W86
       |  |     |    |    `--*G. zonatus [=*Problechilus zonatus; incl. *Meringopalpus fallax] W86
       |  |     |    |--Pycnarthrum Eichhoff 1878 (see below for synonymy) W86
       |  |     |    |    `--P. hispidum (see below for synonymy) W86
       |  |     |    `--Microborus Blandford 1897 [incl. Pseudocrypturgus Eggers 1919] W86
       |  |     |         `--*M. boops [incl. *Pseudocrypturgus camerunus] W86
       |  |     `--Phloeotribini [Phloeotribidae, Phthorophloeides] GC-P14
       |  |          |--Phloeotribus W86
       |  |          `--Aricerus Blandford 1894 [incl. Hylesinosoma Lea 1910] W86
       |  |               |--*A. chapuisi W86
       |  |               `--A. eichhoffi [incl. Hylesinus fici, *Hylesinosoma fici] W86
       |  `--+--Hylesinini (see below for synonymy) GC-P14
       |     |    |--Hylurgops W86
       |     |    |--Hylastinus Bedel 1888 W86
       |     |    |    `--*H. obscurus [=Ips obscurus] W86
       |     |    |--Kissophagus Chapuis 1869 W86
       |     |    |    `--*K. hederae [=Hylesinus hederae] W86
       |     |    |--Pteleobius Bedel 1888 W86
       |     |    |    `--*P. vittatus [=Bostrichus vittatus] W86
       |     |    |--Cryptocurus Schedl 1957 [incl. Hyloperus Browne 1970] W86
       |     |    |    `--*C. spinipennis [incl. *Hyloperus bicornis]
       |     |    |--Neopteleobius Nobuchi 1971 W86
       |     |    |    `--*N. scutulatus [=Hylesinus scutulatus] W86
       |     |    |--Scierus LeConte 1876 W86
       |     |    |    `--*S. annectens W86
       |     |    |--Hylurgus Latreille 1807 M94
       |     |    |    `--*H. ligniperda (Fabricius 1775) [=Hylesinus ligniperda] M94
       |     |    |--Dendrosinus Chapuis 1869 W86
       |     |    |    `--*D. globosus [=Hyulesinus globosus] W86
       |     |    |--Hyleops Schedl 1938 W86
       |     |    |    `--*H. glabratus W86
       |     |    |--Pseudochramesus Blackman 1939 W86
       |     |    |    `--*P. acuteclavatus [=Chramesus acuteclavatus] W86
       |     |    |--Phloeosinopsoides Schedl 1964 [=Phloeosinopsis Schedl 1964 non Schedl 1936] W86
       |     |    |    `--*P. triseriatus [=*Phloeosinopsis triseriatus] W86
       |     |    |--Phloeocranus Schedl 1942 [incl. Diamerides Browne 1949] W86
       |     |    |    `--*P. bruchoides [incl. *Diamerides litseae] W86
       |     |    |--Phloeoditica Schedl 1962 W86
       |     |    |    `--*P. curta [=Kissophagus curtus] W86
       |     |    |--Carphotoreus Wood 1973 W86
       |     |    |    `--*C. alni [=Chaetophloeus alni] W86
       |     |    |--Catenophorus Nunberg 1956 W86
       |     |    |    `--*C. congonus W86
       |     |    |--Phloeosinites Hagedorn 1907 W86
       |     |    |    `--*P. rehi W86
       |     |    |--Phloeoborus Erichson 1836 [incl. Phloeotrupes Erichson 1836] W86
       |     |    |    |--*P. rudis W86
       |     |    |    `--P. grandis [=*Phloeotrupes grandis] W86
       |     |    |--Dactylipalpus Chapuis 1869 (see below for synonymy) W86
       |     |    |    |--D. transversus [=*Dactylopselaphus transversus] W86
       |     |    |    `--D. cicatricosus [=*Ethadopselaphus cicatricosus] W86
       |     |    |--Alniphagus Swaine 1918 [incl. Hylastinoides Spessivitzev 1919] W86
       |     |    |    |--*A. aspericollis [=Hylesinus aspericollis] W86
       |     |    |    `--A. alni [=Hylastes alni, *Hylastinoides alni] W86
       |     |    |--Rhopalopselion Hagedorn 1909 [incl. Hapalogenius Hagedorn 1912] W86
       |     |    |    |--*R. bituberculatum W86
       |     |    |    `--R. globosus [=*Hapalogenius globosus] W86
       |     |    |--Ficicis Lea 1910 [incl. Ficiphagus Murayama 1958] W86
       |     |    |    |--*F. varians W86
       |     |    |    `--‘Hylesinus’ porcatus Chap. 1869 W86, M87 [incl. *Ficiphagus goliathoides W86]
       |     |    |--Pachycotes Sharp 1877 M94
       |     |    |    |--P. peregrinus M94 [=Hylesinus peregrinus W86; incl. *P. ventralis Sharp 1877 M94]
       |     |    |    `--P. australis Schedl 1938 M94
       |     |    |--Hylesinus Fabricius 1801 [incl. Leperisinus Reitter 1913] W86
       |     |    |    |--*H. crenatus W86
       |     |    |    |--H. antipodius W86
       |     |    |    `--H. varius W86 (see below for synonymy)
       |     |    |--Cladoctonus Strohmeyer 1911 (see below for synonymy) W86
       |     |    |    |--*C. affinis W86
       |     |    |    |--C. banosus [=*Hoplites banosus, *Hoplitontus banosus] W86
       |     |    |    `--C. interruptus [=Hoplites interruptus; incl. *Hoplitophthorus sentosus] W86
       |     |    |--Olonthogaster Motschulsky 1866 (see below for synonymy) W86
       |     |    |    |--*O. nitidicollis [=*Holonthogaster nitidicollis] W86
       |     |    |    |--O. asper [=*Hyledius asper] W86
       |     |    |    |--O. spinifer [=Phloeosinus spinifer; incl. *Phloeosinopsis armatus] W86
       |     |    |    `--O. sumatranus [=*Hylurgulus sumatranus] W86
       |     |    |--Chramesus LeConte 1868 (see below for synonymy) W86
       |     |    |    |--*C. hickoriae W86
       |     |    |    |--C. annectans [=*Prochramesus annectans] W86
       |     |    |    |--C. incomptus W86
       |     |    |    |--C. quadridens W86
       |     |    |    |--C. rotundatus [incl. Dendrosinus bonnairei, *Thaumasinulus bonnairei] W86
       |     |    |    `--C. tuberculatus [=*Rhopalopleurus tuberculatus] W86
       |     |    |--Hylesinopsis Eggers 1920 (see below for synonymy) W86
       |     |    |    |--*H. dubius W86
       |     |    |    |--H. africanus [=*Pseudophloeotribus africanus] W86
       |     |    |    |--H. baphiae [=*Glochicopterus baphiae] W86
       |     |    |    |--H. brinckei [=Metahylesinus brinckei, *Hapalophloeus brinckei] W86
       |     |    |    |--H. endroedyi [=*Hemihylesinus endroedyi] W86
       |     |    |    |--H. joveri [=*Trypographus joveri] W86
       |     |    |    |--H. planicollis [=*Chilodendron planicolle] W86
       |     |    |    `--H. togonus [=*Pseudohylesinus togonus, *Metahylesinus togonus] W86
       |     |    `--Phloeosinus Chapuis 1873 M94
       |     |         |--*P. dentatus [=Hylurgus dentatus] M94
       |     |         |--P. bicolor W86
       |     |         |--P. canadensis M94
       |     |         |--P. cristatus W86
       |     |         |--P. cupressi Hopkins 1903 M94
       |     |         |--P. perlatus B14
       |     |         |--P. punctatus W86
       |     |         `--P. thujae [=Hylesinus thujae] W86
       |     `--+--Hylastes Erichson 1836 M94 [Hylastini GC-P14, Hylastinides]
       |        |    |--*H. ater (Paykull 1800) [=Bostrichus ater, Hylesinus ater] M94
       |        |    |--H. brunneus RW02
       |        |    |--H. cunicularius RW02
       |        |    |--H. glabratus C01
       |        |    `--H. nigrinus W86
       |        `--Tomicini GC-P14
       `--+--+--Polygraphini GC-P14
          |  `--Crypturgini GC-P14
          `--+--+--Corthylini [Amphicranidae, Araptidae, Corthyli, Pityophthoridae] GC-P14
             |  |    |--Pityophthorina W86
             |  |    `--Corthylina W86
             |  `--+--Cryphalus GC-P14
             |     `--Xyloctonini GC-P14
             `--+--Hypoborini GC-P14
                `--+--+--Premnobius Eichhoff 1878 W86 [incl. Premnophilus Browne 1962 W86; Premnobiini GC-P14]
                   |  |    |--*P. cavipennis W86
                   |  |    `--P. joveri [=Xyleborus joveri, *Premnophilus joveri] W86
                   |  `--Ipini [Eidophelinae, Ernoporinae, Trypophloeinae] GC-P14
                   |       |--Ips W86
                   |       |--Hypothenemus W86
                   |       |--Scolytogenes W86
                   |       |--Dendrochilus Schedl 1957 W86
                   |       |    `--*D. strombosiopsis W86
                   |       |--Cryphalities Cockerell 1917 W86
                   |       |    `--*C. rugosissimus Cockerell 1917 W86
                   |       |--Periocryphalus Wood 1971 W86
                   |       |    `--*P. pullus W86
                   |       |--Trischidias Hopkins 1915 W86
                   |       |    `--*T. georgiae W86
                   |       |--Neocryphus Nunberg 1956 W86
                   |       |    `--*N. argentinensis W86
                   |       |--Stegomerus Wood 1967 W86
                   |       |    `--*S. vulgaris W86
                   |       |--Allernoporus Kurentzov 1941 W86
                   |       |    `--*A. euonymi W86
                   |       |--Coriacephilus Schedl 1939 W86
                   |       |    `--*C. coriaceus [=Stephanoderes coriaceus] W86
                   |       |--Ernocladius Wood 1980 W86
                   |       |    `--*E. corpulentus [=Cryphalus corpulentus] W86
                   |       |--Cryphalogenes Wood 1980 W86
                   |       |    `--*C. euphorbiae W86
                   |       |--Hemicryphalus Schedl 1963 W86
                   |       |    `--*H. argutus [=Eidophelus argutus] W86
                   |       |--Margadillius Hopkins 1915 W86
                   |       |    `--*M. margadilaonis W86
                   |       |--Ptilopodius Hopkins 1915 W86
                   |       |    `--*P. stephegynis W86
                   |       |--Xyleborinus Reitter 1913 M94
                   |       |    `--*X. saxeseni (Ratzeburn 1837) [=Bostrichus saxeseni] M94
                   |       |--Pityokteines Fuchs 1911 [incl. Orthotomides Wood 1951] W86
                   |       |    |--*P. curvidens [=Ips curvidens] W86
                   |       |    `--P. lasiocarpi [=Orthotomicus lasiocarpi, *Orthotomides lasiocarpi] W86
                   |       |--Orthotomicus Ferrari 1867 [=Neotomicus Fuchs 1911, Onthotomicus] W86
                   |       |    |--*O. laricis [=Bostrichus laricis, *Neotomicus laricis] W86
                   |       |    `--O. erosus JKW03
                   |       |--Trypophloeus Fairmaire 1868 [=Glyptoderus Eichhoff 1878 non LaFerté Sénectère 1851] W86
                   |       |    |--*T. binodulus [=Bostrichus binodulus, *Glyptoderus binodulus] W86
                   |       |    `--T. populi [=Cryphalus populi] W86
                   |       |--Eidophelus Eichhoff 1875 [incl. Phellodendrophagus Krivolutskaya 1958] W86
                   |       |    |--*E. imitans W86
                   |       |    `--E. elegans [=*Phellodendrophagus elegans] W86
                   |       |--Procryphalus Hopkins 1915 W86
                   |       |    |--P. mucronatus [=Cryphalus mucronatus; incl. *P. populi] W86
                   |       |    `--P. utahensis W86
                   |       |--Cryptocarenus Eggers 1937 [incl. Tachyderes Blackman 1943] W86
                   |       |    |--*C. diadematus W86
                   |       |    `--C. seriatus [incl. *Tachyderes floridensis] W86
                   |       |--Acorthylus Brèthes 1922 [incl. Phacrylus Schedl 1938] W86
                   |       |    |--*A. asperatus W86
                   |       |    `--A. bosqui [=*Phacrylus bosqui] W86
                   |       |--Stephanopodius Schedl 1941 (see below for synonymy) W86
                   |       |    |--*S. dispar [=Stephanoderes dispar] W86
                   |       |    `--S. ghanaensis (see below for synonymy) W86
                   |       |--Pityogenes Bedel 1888 [incl. Eggersia Lebedev 1926, Pityoceragenes Balachowsky 1947] W86
                   |       |    |--*P. chalcographus [=Dermestes chalcographus] W86
                   |       |    |--P. bidentatus [=Bostrichus bidentatus, *Eggersia bidentata] W86
                   |       |    `--P. quadridens [=Bostrichus quadridens, *Pityoceragenes quadridens] W86
                   |       |--Ernoporus Thomson 1859 (see below for synonymy) W86
                   |       |    |--*E. tiliae [=Bostrichus tiliae; incl. Cryphalus lederi, *Cryphalops lederi] W86
                   |       |    |--E. concentralis [=*Euptilius concentralis] w86
                   |       |    `--E. melodori [=*Stephanorhopalus melodori] W86
                   |       |--Ernoporicus Berger 1917 (see below for synonymy) W86
                   |       |    |--*E. spessivtzevi W86
                   |       |    |--E. caucasicus [=Ernoporus caucasicus, *Ernopocerus caucasicus] W86
                   |       |    `--E. semenovi [=*Eocryphalus semenovi] W86
                   |       |--Cosmoderes Eichhoff 1878 (see below for synonymy) W86
                   |       |    |--*C. monilicollis [incl. *Vitaderes luffa (n. n.)] W86
                   |       |    |--C. granulicollis [=*Dendriops granulicollis] W86
                   |       |    `--C. setistriatus [=Cryphalus setistriatus, *Erioschidias setistriatus] W86
                   |       |--Hypocryphalus Hopkins 1915 [incl. Dacryphalus Hopkins 1915] W86
                   |       |    |--*H. rotundus W86
                   |       |    |--H. mangiferae W86
                   |       |    `--H. obesus [=*Dacryphalus obesus] W86
                   |       |--Acanthotomicus Blandford 1894 [incl. Isophthorus Schedl 1938, Mimips Eggers 1932] W86
                   |       |    |--*A. spinosus W86
                   |       |    |--A. mimicus W86
                   |       |    |--A. pilosus [=Ips pilosus, *Mimips pilosus] W86
                   |       |    `--A. quadrituberculatus [=*Isophthorus quadrituberculatus] W86
                   |       `--Coccotrypes Eichhoff 1879 M94 (see below for synonymy)
                   |            |--*C. dactyliperda [=Bostrichus dactyliperda] W86
                   |            |--C. advena [incl. *Thamnurgides persicae] W86
                   |            |--C. carpophagus (Hornung 1842) M94 (see below for synonymy)
                   |            |--C. rhizophorae [=*Spermatoplex rhizophorae, *Dendrurgus rhizophorae] W86
                   |            `--C. sannio [=*Poecilips sannio] W86
                   `--+--Dryocoetini [=Dryocoetoideae, Taphrorychini, Thamnurginae] GC-P14
                      |    |--Cyrtogenius W86
                      |    |--Toxophthorus Wood 1986 [=Toxophorus Eggers 1920 (preoc.)] W86
                      |    |    `--*T. africanus (Eggers 1920) [=*Toxophorus africanus] W86
                      |    |--Taphramites Schedl 1947 W86
                      |    |    `--*T. gnathotrichus Schedl 1947 W86
                      |    |--Thamnurgus Eichhoff 1864 W86
                      |    |    `--*T. euphorbiae [=Bostrichus euphorbiae] W86
                      |    |--Chiloxylon Schedl 1959 W86
                      |    |    `--*C. rufulus W86
                      |    |--Dactylotrypes Eggers 1927 W86
                      |    |    `--D. longicollis [=Xyloterus longicollis; incl. *D. uyttenboogaarti] W86
                      |    |--Dendrocranulus Schedl 1937 W86
                      |    |    `--*D. tardus W86
                      |    |--Lymantor Lovendal 1889 W86
                      |    |    `--‘Tomicus’ coryli [incl. *Lymantor sepicola] W86
                      |    |--Taphronurgus Reitter 1913 W86
                      |    |    `--*T. exul [=Thamnurgus exul] W86
                      |    |--Dolurgocleptes Schedl 1965 W86
                      |    |    `--*D. malgassicus W86
                      |    |--Dendrographus Schedl 1964 [incl. Protopityophthorus Schedl 1973] W86
                      |    |    `--*D. pygmaeus [=Pelicerus pygmaeus; incl. *Protopityophthorus durus] W86
                      |    |--Peridryocoetes Wood 1984 W86
                      |    |    `--*P. nitens [=Ozodendron nitens] W86
                      |    |--Ozopemon Hagedorn 1908 W86
                      |    |    `--*O. regius W86
                      |    |--Dryocoetiops Schedl 1957 W86
                      |    |    |--*D. laevis [=Ozopemon laevis] W86
                      |    |    `--D. coffeae W86
                      |    |--Triotemnus Wollaston 1864 [incl. Cladoctoproctus Schedl 1975] W86
                      |    |    |--*T. subretusus W86
                      |    |    `--T. scrafa [=*Cladoctoproctus scrafa] W86
                      |    |--Dryocoetes Eichhoff 1864 (see below for synonymy) W86
                      |    |    |--*D. autographus [=Bostrichus autographus, *Anodius autographus] W86
                      |    |    |--D. confusus W86
                      |    |    `--D. villosus [=Bostrichus villosus, *Dryocoetinus villosus] W86
                      |    |--Tiarophorus Schreiner 1882 (see below for synonymy) W86
                      |    |    |--*T. elongatus W86
                      |    |    |--T. camerunus [=*Hypaspistes camerunus, *Orthaspistes camerunus] W86
                      |    |    `--T. scrutator [=Thamnurgus scrutator, *Pseudothamnurgus scrutator] W86
                      |    |--Xylocleptes Ferrari 1867 [incl. Hylonius Nunberg 1973, Xestips Hagedorn 1912] W86
                      |    |    |--*X. bispinus [=Bostrichus bispinus] W86
                      |    |    |--X. brunneus [=*Hylonius brunneus] W86
                      |    |    `--X. marginatus [=*Xestips marginatus] W86
                      |    `--Taphrorychus Eichhoff 1878 (see below for synonymy) W86
                      |         |--*T. bicolor [=Bostrichus bicolor] W86
                      |         |--T. machnovskii [=Hypothenemus machnovskii, *Saliciphilus machnovskii] W86
                      |         |--T. mikuniyamensis [=*Pseudopoecilips mikuniyamensis] W86
                      |         `--T. primitus [=*Taphroterus primitus] W86
                      `--Xyleborini [Webbinae, Xylebori] GC-P14
                           |--Australoplatypus incompertus GE05
                           |--Xyleborus W86
                           |--Xyleborites Wickham 1913 W86
                           |    `--*X. longipennis Wickham 1913 W86
                           |--Sampsonius Eggers 1935 W86
                           |    `--*S. sexdentatus W86
                           |--Schedlia Browne 1950 W86
                           |    `--*S. sumatrana [=Xyleborus sumatranus] W86
                           |--Dryocoetoides Hopkins 1915 W86
                           |    `--D. capucinus [=Xyleborus capucinus; incl. *D. guatemalensis] W86
                           |--Leptoxyleborus Wood 1980 W86
                           |    `--*L. sordicauda [=Phloeotrogus sordicauda] W86
                           |--Theoborus Hopkins 1915 W86
                           |    `--*T. theobromae W86
                           |--Euwallacea Hopkins 1915 W86
                           |    `--*E. wallacei [=Xyleborus wallacei] W86
                           |--Taurodemus Wood 1980 W86
                           |    `--*T. sharpi [=Xyleborus sharpi] W86
                           |--Mesoscolytus Broun 1904 W86
                           |    `--*M. inurbanus W86
                           |--Hadrodemius Wood 1980 W86
                           |    `--*H. globus [=Xyleborus globus] W86
                           |--Taphrodasus Wood 1980 W86
                           |    `--*T. percorthylus [=Xyleborus percorthylus] W86
                           |--Cryptoxyleborus Schedl 1937 W86
                           |    `--*C. naevus W86
                           |--Cnestus Sampson 1911 [incl. Tosaxyleborus Murayama 1950] W86
                           |    |--*C. magnus W86
                           |    `--C. murayamai [incl. *Tosaxyleborus pallidipennis] W86
                           |--Coptoborus Hopkins 1915 [incl. Streptocranus Schedl 1939] W86
                           |    |--C. vespatorius [=Xyleborus vespatorius; incl. *C. emarginatus] W86
                           |    `--C. mirabilis [=*Streptocranus mirabilis] W86
                           |--Arixyleborus Hopkins 1915 [incl. Xyleboricus Eggers 1923] W86
                           |    |--*A. rugosipes W86
                           |    `--A. canaliculatus [=*Xyleboricus canaliculatus] W86
                           |--Terminalinus Hopkins 1915 [incl. Kelantanius Nunberg 1961] W86
                           |    |--*T. terminaliae W86
                           |    `--T. punctatopilosus [=Xyleborus punctatopilosus, *Kelantanius punctatopilosus] W86
                           |--Eccoptopterus Motschulsky 1863 (see below for synonymy) W86
                           |    |--E. spinosus [=Scolytus spinosus; incl. *Eccoptopterus sexspinosus] W86
                           |    `--E. gracilipes [=*Platydactylus gracilipes, *Eurydactylus gracilipes] W86
                           |--Ambrosiodmus Hopkins 1915 (see below for synonymy) W86
                           |    |--*A. tachygraphus [=Xyleborus tachygraphus] W86
                           |    |--A. obliquecaudata [=*Phloeotrogus oliquecaudata] W86
                           |    `--A. obliquus (see below for synonymy) W86
                           |--Amasa Lea 1894 [incl. Anaxyleborus Wood 1980, Pseudoxyleborus Eggers 1930] W86
                           |    |--*A. thoracicus W86
                           |    |--‘*Pseudoxyleborus’ beesoni W86
                           |    `--A. truncatus (Erichs. 1842) W86, M87 (see below for synonymy)
                           |--Coptodryas Hopkins 1915 (see below for synonymy) W86
                           |    |--*C. confusa W86
                           |    |--C. pullus [=Xyleborus pullus; incl. *Adryocoetes nitidus (n. n.)] W86
                           |    `--C. theae [=Xyleborus theae, *Microperus theae] W86
                           |--Webbia Hopkins 1915 (see below for synonymy) W86
                           |    |--*W. dipterocarpi W86
                           |    |--W. pabo W86
                           |    |--W. subuculae [=*Prowebbia subuculae] W86
                           |    `--W. trepanicauda [=Xyleborus trepanicauda, *Pseudowebbia trepanicauda] W86
                           |--Xylosandrus Reitter 1913 [incl. Apoxyleborus Wood 1980] W86
                           |    |--*X. morigerus [=Xyleborus morigerus] W86
                           |    |--X. compactus MHG04
                           |    |--X. mancus [=Xyleborus mancus, *Apoxyleborus mancus] W86
                           |    `--X. retusus W86
                           `--Anisandrus Ferrari 1867 GC-P14, W86
                                `--*A. dispar W86, GC-P14 [=Xyleborus (Apate) dispar W86]

Scolytinae incertae sedis:
  Bufonus Eggers 1919 W86
    `--*B. obscurus Eggers 1919 W86
  Cactopinus Schwarz 1899 [incl. Cactopinorus Bright 1967; Cactopinini] W86
    |--*C. hubbardi Schwarz 1899 B38
    |--C. cactophthorus [=*Cactopinorus cactophthorus] W86
    |--C. depressus W86
    |--C. desertus W86
    |--C. koebelei Blackman 1938 B38
    |--C. pini Blackman 1938 B38
    `--C. rhois Blackman 1938 B38
  Scolytoplatypus Schaufuss 1893 (see below for synonymy) W86
    |--*S. permirus W86
    |--S. congonus W86
    |--S. mikado [=*Strophionocerus mikado, *Taeniocerus mikado] W86
    |--S. papuanus W86
    `--S. tycon [=*Spongocerus tycon] W86
  Carphodicticini W86
    |--Dendrodicticus Schedl 1958 W86
    |    `--*D. argentiniae W86
    |--Carphodicticus Wood 1971 W86
    |    `--*C. cristatus W86
    `--Craniodicticus Blandford 1895 W86
         `--*C. mucronatus W86
  Xyloterini W86
  Micracini W86
  Phrixosoma Blandford 1897 (see below for synonymy) W86
    |--*P. rude W86
    |--P. magna W86
    |--P. psaltes [=*Bothryperus psaltes] W86
    |--P. quadrioculatum [=*Neohylesinus quadrioculatus] W86
    `--P. striatum [=*Sphaerosinus striatus] W86
  Hyorrhynchini [Hyorrhynchinae, Sueinae] W86
    |--Sueus Murayama 1951 [incl. Neohyorrhynchus Schedl 1962] W86
    |    `--S. niisimai (see below for synonymy) W86
    |--Hyorrhynchus Blandford 1894 W86
    |    `--*H. lewisi W86
    `--Pseudohyorrhynchus Murayama 1950 W86
         `--*P. wadai W86
  Bothrosternini W86

Amasa truncatus (Erichs. 1842) W86, M87 [=Tomicus truncatus W86, *Anaxyleborus truncatus W86]

Ambrosiodmus Hopkins 1915 [incl. Browneia Nunberg 1963, Phloeotrogus Motschulsky 1863 (nom. rej.)] W86

Ambrosiodmus obliquus [=Pityophthorus obliquus; incl. Xyleborus illepidus, *Browneia illepida] W86

Chramesus LeConte 1868 [incl. Prochramesus Wood 1956, Rhopalopleurus Chapuis 1869, Thaumasinulus Reitter 1913] W86

Cladoctonus Strohmeyer 1911 [incl. Hoplites Eggers 1923 nec Koch 1869 nec Neumayr 1875, Hoplitontus Wood 1961, Hoplitophthorus Wood 1961] W86

Cnemonyx Eichhoff 1868 [incl. Ceratolepis Chapuis 1869, Coptodryas Schedl 1948 non Hopkins 1915, Coptosomus Schedl 1952, Loganius Chapuis 1869, Minulus Eggers 1912] W86

Coccotrypes Eichhoff 1879 M94 [incl. Cryphaloides Formenek 1908 W86, Dendrurgus Eggers 1923 W86, Hyphaene Hagedorn 1913 (n. n.) W86, Poecilips Schaufuss 1897 W86, Spermatoplex Hopkins 1915 W86, Thamnurgides Hopkins 1915 W86]

Coccotrypes carpophagus (Hornung 1842) M94 [=Bostrichus carpophagus M94; incl. *Cryphaloides donisthorpei W86, *Hyphaene guineensis (n. n.) W86]

Coptodryas Hopkins 1915 [incl. Adryocoetes Eggers in Schedl 1952 (n. n.), Microperus Wood 1980] W86

Cosmoderes Eichhoff 1878 [incl. Dendriops Schedl 1953, Erioschidias Schedl 1938, Vitaderes Beeson 1941 (n. n.)] W86

Dactylipalpus Chapuis 1869 [=Dactylopselaphus Gemminger & Harold 1872; incl. Ethadopselaphus Blandford 1896] W86

Dryocoetes Eichhoff 1864 [=Anodius Motschulsky 1860 (nom. rej.); incl. Dryocoetinus Balachowsky 1949] W86

Eccoptopterus Motschulsky 1863 [incl. Eurydactylus Hagedorn 1909 non LaFerté Sénectère 1851, Platydactylus Eichhoff 1886] W86

Ernoporicus Berger 1917 [incl. Eocryphalus Kurentzov 1941, Ernopocerus Balachowsky 1949] W86

Ernoporus Thomson 1859 [incl. Cryphalops Reitter 1889, Euptilius Schedl 1940, Stephanorhopalus Hopkins 1915] W86

Hylesinini [Alniphagini, Dactylipalpi, Hylesinen, Phloeobori, Phloeosinides, Phloeosinini, Phloeotrupides] GC-P14

Hylesinopsis Eggers 1920 [incl. Chilodendron Schedl 1953, Glochicopterus Schedl 1954, Hapalophloeus Schedl 1966, Hemihylesinus Schedl 1967, Metahylesinus Eggers 1922, Pseudohylesinus Eggers 1919 non Swaine 1917, Pseudophloeotribus Eggers 1933, Trypographus Schedl 1950] W86

Hylesinus varius W86 [=Bostrichus varius W86; incl. B. fraxini W86, Hylesinus fraxini L86, *Leperisinus fraxini W86]

Olonthogaster Motschulsky 1866 [=Holonthogaster Gemminger & Harold 1872; incl. Hyledius Sampson 1921, Hylurgulus Eggers 1927, Phloeosinopsis Schedl 1936] W86

Phrixosoma Blandford 1897 [incl. Bothryperus Hagedorn 1909, Neohylesinus Eggers 1920, Sphaerosinus Eggers 1929; Phrixosomini] W86

Pycnarthrum Eichhoff 1878 [incl. Monebius Hopkins 1914, Nemobius Chapuis 1869 (preoc.), Nomebius Navas 1915] W86

Pycnarthrum hispidum [=Hypoborus hispidus; incl. *P. gracile, *Nemobius lambottei, *Monebius lambottei, *Nomebius lambottei] W86

Scolytoplatypus Schaufuss 1893 [incl. Spongocerus Blandford 1893, Strophionocerus Sampson 1921, Taeniocerus Blandford 1893; Scolytoplatypini, Scolytoplatypodini, Spongocerinae, Taeniocerini] W86

Stephanopodius Schedl 1941 [incl. Cryphalomimetes Browne 1963, Cryphalomimus Browne 1962 non Eggers 1927] W86

Stephanopodius ghanaensis [=Hypocryphalus ghanaensis, *Cryphalomimetes ghanaensis, *Cryphalomimus ghanaensis] W86

Sueus niisimai [=Hyorrhynchus niisimai, *Neohyorrhynchus niisimai; incl. *Sueus sphaerotrypoides] W86

Taphrorychus Eichhoff 1878 [incl. Pseudopoecilips Murayama 1957, Saliciphilus Sokanovskii 1954, Taphroterus Schedl 1965] W86

Tiarophorus Schreiner 1882 [incl. Hypaspistes Hagedorn 1908 (preoc.), Orthaspistes Hagedorn 1909, Pseudothamnurgus Eggers 1912] W86

Webbia Hopkins 1915 [incl. Prowebbia Browne 1962, Pseudowebbia Browne 1961, Xelyborus Schedl 1939 (n. n.)] W86

*Type species of generic name indicated

REFERENCES

[B38] Blackman, M. W. 1938. New species of Cactopinus Schwarz (Coleoptera: Scolytidae). Proceedings of the Entomological Society of Washington 40 (6): 151–157.

[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.

[GC-P14] Gillett, C. P. D. T., A. Crampton-Platt, M. J. T. N. Timmermans, B. H. Jordal, B. C. Emerson & A. P. Vogler. 2014. Bulk de novo mitogenome assembly from pooled total DNA elucidates the phylogeny of weevils (Coleoptera: Curculionoidea). Molecular Biology and Evolution 31 (8): 2223–2237.

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

[JKW03] Jacobs, K., T. Kirisits & M. J. Wingfield. 2003. Taxonomic re-evaluation of three related species of Graphium, based on morphology, ecology and phylogeny. Mycologia 95 (4): 714–727.

[L86] Lindquist, E. E. 1986. The world genera of Tarsonemidae (Acari: Heterostigmata): a morphological, phylogenetic, and systematic revision, with a reclassification of family-group taxa in the Heterostigmata. Memoirs of the Entomological Society of Canada 118 (S136): 1–517.

[M87] Masters, G. 1887. Catalogue of the described Coleoptera of Australia. Part VI. Proceedings of the Linnean Society of New South Wales, series 2, 1 (4): 979–1036.

[M94] May, B. M. 1994. An introduction to the immature stages of Australian Curculionoidea. In: Zimmerman, E. C. Australian Weevils (Coleoptera: Curculionoidea) vol. 2. Brentidae, Eurhynchidae, Apionidae and a chapter on immature stages by Brenda May pp. 365–728. CSIRO Australia.

[MHG04] Maynard, G. V., J. G. Hamilton & J. F. Grimshaw. 2004. Quarantine—phytosanitary, sanitary and incursion management: an Australian entomological perspective. Australian Journal of Entomology 43 (3): 318–328.

[RW02] Reay, S. D., & P. J. Walsh. 2002. Relative attractiveness of some volatiles to the introduced pine bark beetles, Hylastes ater and Hylurgus ligniperda (Curculionidae: Scolytinae). New Zealand Entomologist 25: 51–56.

[W86] Wood, S. L. 1986. A reclassification of the genera of Scolytidae (Coleoptera). Great Basin Naturalist Memoirs 10: 1–126.

Last updated: 19 September 2018.