Coptotermes

Castes of Coptotermes acinaciformis, from CSIRO.

Belongs within: Rhinotermitidae.

Coptotermes is a genus of drywood termites in which soldiers bear a large fontanelle opening near the front of the head.

Characters (from Maiti 2006): Imago with head dark to dark brown, head and pronotum fairly densely and abdomen densely hairy. Head subround. Eyes of medium size. Ocelli either subround or elongately oval, either almost touching or slightly separated from eyes. Fontanelle minute, placed almost at middle of head. Mandibles each with an apical and three marginal teeth. Left with apical only slightly longer than first marginal and almost equal to or slightly shorter than or subequal to second and third marginal. Right with apical thinner and slightly shorter than second, first marginal rudimentary placed at anterior base of second, third smaller with long posterior margin, generally longer than molar plate. Postclypeus flat or weakly swollen with or without median line. Apical tibial spur formula 3:2:2. Wings generally moderate to densely hairy, all veins arising independently from scale. In fore wing, costa and subcosta merged, strongly chitinised, running along anterior margin. Radius thick and running parallel and close to costa and subcosta. Medial thin and with one to three branches distally, cubitus also weak and with six to thirteen branches to posterior margin. Hind wing as in fore wing except medial arising from radius just outside scale. Soldier with head light yellow to deep brown in colour, moderately to densely hair, pyriform to broadly oval. Eyes generally absent, eye spots at times visible. Ocelli absent. Fontanelle large with round to oval large opening placed far anteriorly. Mandibles sabre-shaped, gradually narrowing anteriorly and strongly to weakly hooked at apices, left with a few to maximum of four crenulations in basal half and with a large knob-like projection. Right with a few basal crenulations also. Antennae with 13–17 segments, 3–4 variable in shape and size. Postmentum long and club-shaped with distinct widest and narrowest width, sometimes with hairs anteriorly. Pronotum narrower than head, flat, sometimes a little raised on anterior margin, anterior margin with median notch and posterior margin with weak median emargination.

<==Coptotermes Wasmann 1896 M06 (see below for synonymy)
    |--C. sjostedti Holmgren 1911 IVE07, M06 [incl. C. sjostedti var. modica M06]
    `--+--C. acinaciformis (Froggatt 1898) IVE07, M06 (see below for synonymy)
       |    |--C. a. acinaciformis M06
       |    `--C. a. raffrayi Wasmann 1900 [=Termes (Coptotermes) raffrayi] M06
       `--+--C. niger Snyder 1922 IVE07, M06
          `--C. testaceus (Linnaeus 1758) IVE07, M06 (see below for synonymy)

Coptotermes incertae sedis:
  *C. gestroi (Wasmann 1896) [=Termes (*Coptotermes) gestroi] M06
  C. amanii (Sjöstedt 1911) (see below for synonymy) M06
  C. amboinensis Kemner 1931 M06
  C. bannaensis Xia & He 1986 [=[=C. (Oligocrinitermes) bannaensis] XH86
  C. beckeri Mathur & Chhotani 1969 M06
  C. bentongensis Krishna 1956 M06 [=C. (Polycrinitermes) bentongensis XH86]
  C. boetonensis Kemner 1934 M06 [=C. (Polycrinitermes) boetonensis XH86]
  C. bornensis Oshima 1914 M06
  C. brevis DAF07
  C. brunneus Gay 1955 M06
  C. ceylonicus Holmgren 1911 XH86, M06 [=C. (Oligocrinitermes) ceylonicus] XH86
  C. changtaiensis Xia & He 1986 [=C. (Polycrinitermes) changtaiensis] XH86
  C. chaoxianensis Huang & Li 1985 [=C. varicapitatus chaoxianensis, [=C. (Polycrinitermes) chaoxianensis] XH86
  C. cochlearus Xia & He 1986 [=C. (Oligocrinitermes) cochlearus] XH86
  C. communis Xia & He 1986 [=C. (Polycrinitermes) communis] XH86
  C. crassus Snyder 1922 M06
  C. ‘crassus’ Ping 1985 non Snyder 1922 M06
  C. curvignathus Holmgren 1913 M06 (see below for synonymy)
  C. cyclocoryphus Zhu et al. 1984 [=C. (Polycrinitermes) cyclocoryphus] XH86
  C. dimorphus Xia & He 1986 [=C. (Oligocrinitermes) dimorphus] XH86
  C. dobonicus Oshima 1914 M06 [=C. (Polycrinitermes) dobonicus XH86]
  C. dreghorni Hill 1942 M06
  C. elisae (Desneux 1905) M06 [=Termes (Coptotermes) elisae M06; incl. C. hyaloapex Holmgren 1911 M06, H42]
  C. emersoni Ahmad 1953 M06
  C. eucalyptus Ping 1984 [=C. (Polycrinitermes) eucalyptus] XH86
  C. formosanus Shiraki 1909 XH86 (see below for synonymy)
  C. gaurii Roonwal & Krishna 1956 [incl. C. exignus Jepson 1930, C. exiguus] M06
  C. grandiceps Snyder 1925 [incl. C. froggatti Light & Davis 1929, C. solomonensis Snyder 1925] M06
  C. grandis Li & Huang 1985 [=C. (Polycrinitermes) grandis] XH86
  C. guangdongensis Ping 1985 M06
  C. guanzhouensis Ping 1985 M06
  C. hainanensis Li & Tsai 1985 [=C. (Polycrinitermes) hainanensis] XH86
  C. harrisi M06
  C. havilandi Holmgren 1911 [incl. C. javanicus Kemner 1934, C. pacificus Light 1932] M06
  C. heimi (Wasmann 1902) [=*Arrhinotermes heimi; incl. C. parvulus Holmgren 1913] M06
  C. hekouensis Xia & He 1986 [=C. (Polycrinitermes) hekouensis] XH86
  C. heteromorphus Ping 1985 M06
  C. intermedius Silvestri 1912 [incl. C. truncatus var. reducta Sjostedt 1926, C. reductus] M06
  C. jiaxingensis Xia & He 1986 [=C. (Polycrinitermes) jiaxingensis] XH86
  C. kalshoveni Kemner 1934 M06
  C. kishori Roonwal & Chhotani 1962 M06
  C. labiosus Hill 1926 H42 [incl. C. flavus Hill 1926 non Bugnion & Popeff 1910 M06, C. frenchi Hill 1932 M06]
  C. lacteus (Froggatt 1898) [=Termes lacteus; incl. C. sedulus Hill 1923] M06
  C. longignathus Xia & He 1986 [=C. (Oligocrinitermes) longignathus] XH86
  C. longistriatus Li & Huang 1985 [=C. (Polycrinitermes) longistriatus] XH86
  C. menadoensis Oshima 1914 M06
  C. menglunensis Tsai & Huang 1985 (see below for synonymy) XH86
  C. michaelseni Silvestri 1909 M06
  C. minutissimus Kemner 1934 M06
  C. minutus Li & Huang 1985 [=C. varicapitatus minutus, C. (Polycrinitermes) minutus] XH86
  C. monosetosus Tsai & Li 1985 [=C. (Oligocrinitermes) monosetosus] XH86
  C. obiratus Hill 1927 M06
  C. obliquus Xia & He 1986 [=[=C. (Oligocrinitermes) obliquus] XH86
  C. ochraceus Ping & Xu 1986 M06
  C. pamuae Snyder 1925 M06
  C. parradoxus M06
  C. perigrenator Kemner 1943 M06
  C. premrasmii Ahmad 1965 XH86, M06 [=C. (Oligocrinitermes) premrasmii XH86]
  C. priscus Emerson 1971 P92
  C. rectangularis Ping & Xu 1986 M06
  C. remotus Hill 1927 M06
  C. reticulatus M06
  C. sepangensis Krishna 1956 M06
  C. shanghaiensis Xia & He 1986 [=C. (Polycrinitermes) shanghaiensis] XH86
  C. sinabangensis Oshima 1923 M06 [=C. (Polycrinitermes) sinabangensis XH86]
  C. sucineus Emerson 1971 P92
  C. suzhouensis Xia & He 1986 [=C. (Polycrinitermes) suzhouensis] XH86
  C. travians (Haviland 1898) [=Termes (Coptotermes) travians] M06
  C. truncatus (Wasmann 1897) [=Termes (Coptotermes) truncatus, Eutermes truncatus] M06
  C. vastator Light 1929 M06
  C. xiaoliangensis Ping 1984 [=C. (Polycrinitermes) xiaoliangensis] XH86

Coptotermes Wasmann 1896 M06 [incl. Arrhinotermes Wasmann 1902 M06, Oligocrinitermes Xia & He 1986 (n. n.) XH86, Polycrinitermes Xia & He 1986 (n. n.) XH86, Vastitermes Sjöstedt 1926 M06; Coptotermitinae]

Coptotermes acinaciformis (Froggatt 1898) IVE07, M06 [=Termes acinaciformis M06, T. (Eutermes) acinaciformis M06]

Coptotermes amanii (Sjöstedt 1911) [=Eutermes (Coptotermes) amanii; incl. C. sjostedti var. subintactae Silvestri 1914-1915] M06

Coptotermes curvignathus Holmgren 1913 M06 [=C. (Polycrinitermes) curvignathus XH86; incl. C. robustus Holmgren 1913 M06]

Coptotermes formosanus Shiraki 1909 XH86 [=[=C. (Polycrinitermes) formosanus XH86; incl. C. formosae Holmgren 1911 M06, C. hongkonensis Oshima 1914 M06, C. intrudens Oshima 1920 M06]

Coptotermes menglunensis Tsai & Huang 1985 [=C. monosetosus menglunensis, [=C. (Oligocrinitermes) menglunensis] XH86

Coptotermes testaceus (Linnaeus 1758) IVE07, M06 [=Hemerobius testaceus M06, Termes (Eutermes) testaceus M06; incl. Perla fusca De Geer 1778 M06, T. (E.) marabitanus Hagen 1858 M06, Coptotermes marabitanus M06, T. morio Fabricius 1793 M06]

*Type species of generic name indicated

REFERENCES

[DAF07] Department of Agriculture and Food. 2007. Common Seasonal Pests. State of Western Australia.

[H42] Hill, G. F. 1942. Termites (Isoptera) from the Australian Region (including Australia, New Guinea and islands south of the Equator between 140°E. longitude and 170°W. longitude). Commonwealth of Australia Council for Scientific and Industrial Research: Melbourne.

[IVE07] Inward, D. J. G., A. P. Vogler & P. Eggleton. 2007. A comprehensive phylogenetic analysis of termites (Isoptera) illuminates key aspects of their evolutionary biology. Molecular Phylogenetics and Evolution 44: 953–967.

[M06] Maiti, P. K. 2006. A taxonomic monograph on the world species of termites of the family Rhinotermitidae (Isoptera: Insecta). Memoirs of the Zoological Survey of India 20 (4): i–viii, 1–272.

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

[XH86] Xia K.-L. & He X.-S. 1986. Study on the genus Coptotermes from China (Isoptera: Rhinotermitidae). Contr. Shanghai Inst. Entomol. 6: 157–182.

Neanuridae

Friesea sp., copyright Centre for Biodiversity Genomics.

Belongs within: Collembola.
Contains: Neanura, Ceratrimeria, Anurida, Pseudachorutes.

The Neanuridae are a group of springtails characterised by the presence of microsensilla on the third thoracic segment, a cardo between the stipes and the fulcrum, near-cylindrical antennae, and pointed buccal cones pointed (Greenslade 1991).

<==Neanuridae [Achorutinae, Neoarthropleona]
    |--Uchidanurinae G91
    |    |--Megalanura tasmaniae G91
    |    |--Womersleymeria G91
    |    `--Acanthanura Börner 1906 G91, W39
    |--Neanurinae [Achorutini] G91
    |    |--Protanura W39
    |    |--Australonura G91
    |    |--Neanura W39
    |    |--Oudemansia Schött 1893 nec Miq. 1854 (ICBN) nec Speg. 1880 (ICBN) CB80
    |    |    |--*O. coerulea Schött 1893 CB80 [=Polyacanthella coerulea W39]
    |    |    |--O. esakii CB80
    |    |    |--O. georgia Christiansen & Bellinger 1980 CB80
    |    |    `--O. hirashimai CB80
    |    `--Morulina Börner 1906 CB80
    |         |--*M. gigantea (Tullberg 1876) (see below for synonymy) CB80
    |         |--M. alia Christiansen & Bellinger 1980 CB80
    |         |--M. crassa Christiansen & Bellinger 1980 CB80
    |         |--M. mackenziana Hammer 1953 CB80
    |         |--M. multatuberculata (Coleman 1941) [=Neanura multatuberculata] CB80
    |         |--M. nuda Cassagnau 1955 CB80
    |         `--M. thulensis Hammer 1953 CB80
    |--Friesea Dalla Torre 1895 SS06 (see below for synonymy)
    |    |--*F. mirabilis (Tullberg 1871) [=Triaena mirabilis] CB80
    |    |--F. cera Christiansen & Bellinger 1980 CB80
    |    |--F. claviseta Axelson 1900 [incl. F. caldaria Guthrie 1903] CB80
    |    |--F. emucronata Stach 1922 CB80
    |    |--F. fara Christiansen & Bellinger 1980 CB80
    |    |--F. grandis Mills 1934 CB80
    |    |--F. grisea (Schäffer 1891) (see below for synonymy) W67
    |    |--F. hammeni H83
    |    |--F. kardosia (Wray 1952) [=*Prospinanura kardosia] CB80
    |    |--F. landwehri Christiansen & Bellinger 1980 SS06
    |    |--F. magnicornis Denis 1931 CB80
    |    |--F. mauriesi Cassagnau 1958 CB80
    |    |--F. millsi Christiansen & Bellinger 1974 CB80
    |    |--F. multispinosa Denis 1947 G86
    |    |--F. nauroisi Cassagnau 1958 CB80
    |    |--F. oteruelensis Simón-Benito 2005 F05
    |    |--‘Prospinanura’ oxfordi Wray 1953 (n. d.) [=Wrayella oxfordi] CB80
    |    |--F. pentacantha Mills 1934 CB80
    |    |--F. peruensis Smolis & Skarżyński 2006 SS06
    |    |--F. polla Christiansen & Bellinger 1980 CB80
    |    |--F. quinquespinosa CB80 [=Polyacanthella quinquispinosa W39]
    |    |--F. quinta Christiansen & Bellinger 1980 CB80
    |    |--F. sublimis Macnamara 1921 CB80
    |    |--‘Neoprospinanura’ tecala Wray 1963 (n. d.) CB80
    |    |--F. tilbrooki Wise 1970 [incl. F. viennei Deharveng 1981] G86
    |    |--F. wilkeyi Christiansen & Bellinger 1974 CB80
    |    `--F. woyciechowskii Weiner 1980 SS06
    `--Pseudachorutinae [Pseudachorutini] G91
         |--Echinanura W39
         |--Sericanura W39
         |--Stachia W39
         |--Platanurida W39
         |--Meganurida W39
         |--Ceratrimeria W39
         |--Anurida W39
         |--Pseudachorutes W39
         |--Tasmanura Womersley 1937 W39
         |    `--T. evansi Womersley 1937 W39
         |--Anuritelsa Womersley 1939 W39
         |    `--*A. maritima Womersley 1939 W39
         |--Anuridella Willem 1906 CB80
         |    |--*A. marina Willem 1906 [=Anurida (*Anuridella) marina] CB80
         |    `--A. calcarata Denis 1925 TM86 [=Anurida (Anuridella) calcarata CB80]
         |--Pseudanurida Schött 1901 [incl. Neachorutes Womersley 1933] W39
         |    |--*P. billitonensis Schött 1901 W39
         |    `--P. glauerti (Womersley 1933) G91, W39 [=Neachorutes glauerti W39]
         |--Pseudachorutoides Becker 1905 W39
         |    |--*P. bogoyawlensky W39
         |    |--P. clysmae Jackson 1927 W39
         |    `--P. dollfusi W39
         |--Polyacanthella Schaeffer 1897 [incl. Conotelsa Denis 1925] W39
         |    |--*P. michaelseni Schaeffer 1897 [=Friesea michaelseni] W39
         |    |--P. accuminata [=Conotelsa accuminata, Friesea accuminata] W39
         |    |--P. barnardi Womersley 1930 W39
         |    |--P. brevicaudata [=Friesea brevicaudata] W39
         |    `--P. parva W39
         `--Paranura Axelson 1902 W39
              |--*P. sexpunctata Axelson 1902 CB80
              |--P. anops Christiansen & Bellinger 1980 CB80
              |--P. australasiae Womersley 1935 W39
              |--P. caeca Folsom 1916 CB80
              |--P. colorata Mills 1934 [=P. sexpunctata var. colorata] CB80
              |--P. millsi Christiansen & Bellinger 1980 CB80
              |--P. quadrilobata Hammer 1953 CB80
              `--P. s-uenoi CB80

Neanuridae incertae sedis:
  Pseudachorutella SL06
    |--P. asigillata (Börner 1901) SL06
    `--P. stachi [=Pseudachorutes (Pseudachorutella) stachi] H83
  Pseudoxenylla Christiansen & Pike 2002 CP02
    `--*P. fovealis Christiansen & Pike 2002 CP02
  Kenyura MC13
  Cassagnaudina Massoud 1967 CB80
    `--*C. coiffaiti Cassagnau 1955 CB80
  Stachorutes demattiesi Dallai 1973 CB80
  Neanurodes neomexicanus Scott 1960 (n. d.) CB81

Friesea Dalla Torre 1895 SS06 [incl. Macgillivraya Grote 1894 (nom. rej.) CB80, Prospinanura Wray 1952 CB80; Frieseinae G91]

Friesea grisea (Schäffer 1891) [=Tullbergia grisea, Frisia (l. c.) grisea, Pseudotullbergia grisea; incl. Achorutoides antarcticus Willem 1901] W67

*Morulina gigantea (Tullberg 1876) [=Anura gigantea; incl. M. callowayia Wray 1953, M. kotzebuensis Bödvarsson 1960] CB80

*Type species of generic name indicated

REFERENCES

[CB80] Christiansen, K., & P. Bellinger. 1980. The Collembola of North America north of the Rio Grande: A taxonomic analysis vol. 1. Introduction; general; families Poduridae and Hypogastruridae. Grinnell College: Grinnell (Iowa).

[CB81] Christiansen, K., & P. Bellinger. 1981. The Collembola of North America north of the Rio Grande: A taxonomic analysis vol. 4. Families Neelidae and Sminthuridae; glossary; bibliography; index. Grinnell College: Grinnell (Iowa).

[CP02] Christiansen, K., & E. Pike. 2002. Cretaceous Collembola (Arthropoda, Hexapoda) from the Upper Cretaceous of Canada. Cretaceous Research 23: 165–188.

[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.

[G86] Greenslade, P. 1986. Additions to the collembolan fauna of Heard Island. Records of the South Australian Museum 19 (7): 91–96.

[G91] Greenslade, P. J. 1991. Collembola (springtails). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 252–264. Melbourne University Press: Carlton (Victoria).

[H83] Hammen, L. van der. 1983. Contribution to the knowledge of the soil-fauna of New Guinea. Zoologische Verhandelingen 206: 1–36.

[MC13] Majer, J. D., S. K. Callan, K. Edwards, N. R. Gunawardene & C. K. Taylor. 2013. Baseline survey of the terrestrial invertebrate fauna of Barrow Island. Records of the Western Australian Museum Supplement 83: 13–112.

[SL06] Schulz, H.-J., & P. Lymberakis. 2006. First contribution to the knowledge of the Collembola fauna of the White Mountains (Lefká Óri) in west Crete (Insecta, Collembola, Isotomidae). Senckenbergiana Biologica 86 (2): 229–234.

[SS06] Smolis, A., & D. Skarżyński. 2006. A new species of Friesea Dalla Torre 1895 from Peru (Insecta, Collembola, Neanuridae). Senckenbergiana Biologica 86 (1): 39–45.

[TM86] Thibaud, J.-M., & Z. Massoud. 1986. Insecta: Collembola. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 616–617. E. J. Brill/Dr W. Backhuys: Leiden.

[W67] Wise, K. A. J. 1967. Collembola (springtails). Antarctic Research Series 10: 123–148.

[W39] Womersley, H. 1939. Primitive Insects of South Australia: Silverfish, springtails, and their allies. South Australian Branch of the British Science Guild: Adelaide.

Hypogastruridae

Ceratophysella denticulata, copyright Scott Justis.

Belongs within: Collembola.
Contains: Hypogastrura (Hypogastrura).

The Hypogastruridae are a group of elongate springtails with a granular, ridged molar plate on the mandible but lacking pseudocelli. The body is usually pigmented and ocelli are usually present, as is the furca; the antenna 3 organ is simple, with only two or three rod-shaped sensilla and no papillae or sensorial bodies; and the postantennal organ, if present, is usually round or oval with a maximum of six vesicles (Greenslade 1991).

<==Hypogastruridae [Hypogastrurinae]
    |--Gomphiocephalus Carpenter 1908 W67
    |    `--G. hodgsoni Carpenter 1908 [=Gomphocephalus (l. c.) hodgsoni] W67
    |--Beckerella W39
    |--Proxenyllodes W39
    |--Xenyllogastrura W39
    |--Paraxenylla G91
    |--Cosberella conatoa Wray 1963 (n. d.) CB81
    |--Knowltonella Wray 1958 CB80a
    |    `--*K. idahoensis Wray 1958 CB80a
    |--Stenogastrura Christiansen & Bellinger 1980 CB80a
    |    `--*S. hiemalis Christiansen & Bellinger 1980 CB80a
    |--Bonetogastrura Thibaud 1975 CB80a
    |    `--*B. balazuci (Delamare 1951) [=Typhlogastrura balazuci] CB80a
    |--Tafallia Bonet 1947 [incl. Hoffia Scott 1961] CB80a
    |    |--*T. insularis Bonet 1947 CB80a
    |    `--T. robusta (Scott 1961) [=*Hoffia robusta] CB80a
    |--Microgastrura Stach 1922 CB80a
    |    |--*M. duodecimoculata Stach 1922 CB80a
    |    `--M. minutissima (Mills 1934) [=Achorutes (Schoettella) minutissima] CB80a
    |--Mesachorutes Absolon 1900 CB80a
    |    |--*M. (Mesachorutes) quadriocellata Absolon 1900 CB80a
    |    `--M. (Denigastrura Stach 1949) CB80a
    |         |--M. (*D.) tetrophthalma (Denis 1928) [=Hypogastrura tetrophthalma] CB80a
    |         `--M. (D.) thomomys (Chamberlain 1943) [=Achorutes thomomys, Hypogastrura thomomys] CB80a
    |--Acherontiella Absolon 1913 CB80a
    |    |  i. s.: A. prominentia MC13
    |    |--A. (Acherontiella) [incl. Acherontiellina Salmon 1964] CB80a
    |    |    |--*A. (A.) onychiuriformis Absolon 1913 CB80a
    |    |    |--A. (A.) epigea CB80a
    |    |    `--A. (A.) sabina Bonet 1945 [=*Acherontiellina sabina] CB80a
    |    `--A. (Acherontides) CB80a
    |         |--A. (A.) atoyacensis CB80a
    |         `--A. (A.) potosinus CB80a
    |--Willemia Börner 1901 CB80a
    |    |--*W. anophthalma Börner 1901 CB80a
    |    |--W. biseta Christiansen & Bellinger 1980 CB80a
    |    |--W. bulbosa CB80a
    |    |--W. denisi Mills 1932 CB80a
    |    |--W. dubia Christiansen & Bellinger 1980 CB80a
    |    |--W. intermedia Mills 1934 CB80a
    |    |--W. persimilis CB80a
    |    |--W. similis Mills 1934 CB80a
    |    `--W. vashtia Wray 1950 CB80a
    |--Triacanthella Schäffer 1897 CB80a
    |    |--*T. michaelseni Schäffer 1897 CB80a
    |    |--T. alba Carpenter 1909 W39
    |    |--T. biroi Stach 1924 W39
    |    |--T. clavata [=Triacanthura clavata] W39
    |    |--T. copelandi (Wray 1963) [=*Clingmantha copelandi] CB80a
    |    |--T. inopinata Denis 1924 W39
    |    |--T. perfecta Denis 1927 W39
    |    |--T. rosea Wahlgren 1906 W39
    |    `--T. violacea Womersley 1939 W39
    |--Schaefferia W39
    |    |  i. s.: S. coeca (Cassagnau 1959) TM86
    |    |         S. decemoculata (Stach 1939) DB21
    |    |         S. subterranea V73
    |    |--S. (Schaefferia) C90
    |    |    |--*S. (S.) emucronata Absolon 1900 CB80a
    |    |    |--S. (S.) cheoha Wray 1963 CB80a
    |    |    |--S. (S.) duodecimocellata CB80a
    |    |    `--S. (S.) guerrerense CB80a
    |    `--S. (Typhlogastrura Bonet 1930) CB80a
    |         |--S. (*T.) mendizabali (Bonet 1930) [=Hypogastrura (*T.) mendizabali] CB80a
    |         |--S. (T.) alabamensis (Thibaud 1975) [=Typhlogastrura alabamensis] CB80a
    |         |--S. (T.) christianseni (Thibaud 1975) CB80a
    |         `--S. (T.) variabilis Christiansen 1952 CB80a
    |--Hypogastrura Bourlet 1839 W39
    |    |  i. s.: H. antarctica Salmon 1962 W67
    |    |         H. papillata Gisin 1949 SL06
    |    |         H. pseudopurpurascens Womersley 1928 W39
    |    |         H. purpurascens H44
    |    |         H. rufescens W39
    |    |         H. sahlbergi RD77
    |    |--H. (Hypogastrura) CB80a
    |    |--H. (Mitchellania Wray 1953) [incl. Cyclograna Yosii 1961, Spinifacies Scott 1963] CB80a
    |    |    |--H. (*M.) hermosa (Wray 1953) [=*Mitchellania hermosa] CB80a
    |    |    |--H. (M.) californica (Bacon 1914) [=Achorutes californica] CB80a
    |    |    |--H. (M.) gibbomucronata (Hammer 1953) CB80a
    |    |    |--H. (M.) hiawatha (Yosii 1962) [=Cyclograna hiawatha] CB80a
    |    |    |--H. (M.) horrida Yosii 1960 CB80a
    |    |    |--H. (M.) loricata Yosii 1960 CB80a
    |    |    |--‘*Spinifacies’ oregonensis Scott 1963 CB80a
    |    |    |--H. (M.) virga Christiansen & Bellinger 1980 CB80a
    |    |    `--H. (M.) vulgaris Yosii 1960 [=H. (*Cyclograna) vulgaris] CB80a
    |    `--H. (Schoettella Schäffer 1896) CB80a
    |         |--H. (*S.) ununguiculata (Tullberg 1869) [=Achorutes ununguiculatus] CB80a
    |         |--H. (S.) albamaculata Scott 1960 CB80a
    |         |--H. (S.) banksi Maynard 1951 CB80a [incl. Tullbergia neomexicana Scott 1961 CB80b]
    |         |--H. (S.) glasgowi (Folsom 1916) [=Achorutes glasgowi; incl. A. magnoliana Wray 1952] CB80a
    |         |--‘Schoettella’ poppei S00
    |         `--‘Schoettella’ subcrassa S00
    |--Xenylla Tullberg 1869 CB80a
    |    |--*X. maritima Tullberg 1869 CB80a
    |    |--X. acauda Gisin 1947 CB80a
    |    |--X. californica Gama 1976 CB80a
    |    |--X. canadensis Hammer 1953 CB80a
    |    |--X. carolinensis Wray 1946 CB80a
    |    |--X. christianseni Gama 1974 CB80a
    |    |--X. collis Bacon 1914 (n. d.) CB80a
    |    |--X. duchesnea Wray 1958 CB80a
    |    |--X. grisea Axelson 1900 [incl. X. gracilis Guthrie 1903] CB80a
    |    |--X. humicola (Fabricius 1780) [=Podura humicola; incl. X. baconae Folsom 1916] CB80a
    |    |--X. littoralis Womersley 1934 W39
    |    |--X. louisiana Gama 1976 CB80a
    |    |--X. mucronata Axelson 1903 [incl. Achorutes speciosa Rainbow 1907] W39
    |    |--X. neomexicana Scott 1960 (n. d.) CB80a
    |    |--X. occidentalis Womersley 1934 W39
    |    |--X. pallescens (Scott 1960) [=Xenyllodes pallescens; incl. Xenyllodes alpina Scott 1960] CB80a
    |    |--X. paludis Bacon 1914 (n. d.) CB80a
    |    |--X. proxima Denis 1931 CB80a
    |    |--X. pseudomaritima James 1933 CB80a
    |    |--X. simberloffi Gama 1974 CB80a
    |    |--X. similata H83
    |    |--X. thibaudi H83
    |    |--X. welchi Folsom 1916 CB80a
    |    |--X. wilsoni Gama 1974 CB80a
    |    `--X. yucatana MC13
    |--Biscoia Salmon 1962 W67
    |    `--B. sudpolaris Salmon 1962 W67
    `--Ceratophysella Börner 1932 S07 [incl. Spinachorutes Scott 1962 CB80a]
         |  i. s.: C. alachuensis Skarżyński 2007 S07
         |         C. annae Babenko 1994 S07
         |         C. caucasica Martynova 1964 S07
         |         ‘Achorutes’ citri Bacon 1914 (n. d.) CB80a
         |         ‘Hypogastrura’ gravesi Wray 1971 CB80a
         |         C. kennethi Skarżyński 2007 S07
         |         C. kutyrevae Babenko 1994 S07
         |         C. longispina (Tullberg 1876) [=Achorutes longispina, Hypogastrura (Ceratophysella) longispina] CB80a
         |         ‘Hypogastrura’ macrospinata Maynard 1951 (n. d.) CB80a
         |         ‘Achorutes’ marmorata Packard 1873 (n. d.) CB80a
         |         C. maya [=Hypogastrura (Ceratophysella) maya] CB80a
         |         C. orizabae [=Hypogastrura (Ceratophysella) orizabae] CB80a
         |         C. robustiseta Skarżyński & Smolis 2006 S07
         |         ‘Achorutes’ scotica Carpenter & Evans 1899 CB80a
         |         ‘Achorutes’ sigillata Uzel 1891 CB80a
         |         C. stercoraria Stach 1963 S07
         |         ‘Achorutes’ texensis Packard 1873 (n. d.) CB80a
         |         C. tolteca Yosii 1962 S07 [=Hypogastrura (Ceratophysella) tolteca CB80a]
         |--C. armata group CB80a
         |    |--*C. armata (Nicolet 1841) CB80a, SL06 (see below for synonymy)
         |    |--C. bengtssoni (Ågren 1904) [=Achorutes bengtssoni, Hypogastrura (Ceratophysella) bengtssoni] CB80a
         |    |--C. brevisensillata Yosii 1961 [=Hypogastrura (Ceratophysella) brevisensillata] CB80a
         |    |--C. glancei (Hammer 1953) [=Hypogastrura (Ceratophysella) glancei] CB80a
         |    |--C. jondavi (Wray 1946) [=Hypogastrura (Ceratophysella) jondavi] CB80a
         |    `--C. pseudarmata (Folsom 1916) [=Hypogastrura (Ceratophysella) pseudarmata] CB80a
         |--C. boletivora group CB80a
         |    |--C. biloba (Christiansen & Bellinger 1980) [=Hypogastrura (Ceratophysella) biloba] CB80a
         |    |--C. boletivora (Packard 1873) [=Achorutes boletivorus, Hypogastrura (Ceratophysella) boletivora] CB80a
         |    |--C. communis (Folsom 1897) [=Achorutes communis, Hypogastrura (Ceratophysella) communis] CB80a
         |    `--C. pratorum (Packard 1873) [=Achorutes pratorum] CB80a
         |--C. densornata (Maynard 1951) [=Hypogastrura (Ceratophysella) densornata] CB80a
         |--C. denticulata group CB80a
         |    |--C. brevis (Christiansen & Bellinger 1980) [=Hypogastrura (Ceratophysella) brevis] CB80a
         |    |--C. denticulata (Bagnall 1941) (see below for synonymy) CB80a
         |    |--C. gibbosa (Bagnall 1940) CB80a, S07 (see below for synonymy)
         |    |--C. granulifera Yosii 1962 [=Hypogastrura (Ceratophysella) granulifera] CB80a
         |    |--C. lucifuga (Packard 1888) [=Achorutes lucifuga, Hypogastrura (Ceratophysella) lucifuga] CB80a
         |    |--C. pecki (Christiansen & Bellinger 1980) [=Hypogastrura (Ceratophysella) pecki] CB80a
         |    |--C. scotti Yosii 1962 [=Hypogastrura (Ceratophysella) scotti] CB80a
         |    `--C. succinea (Gisin 1949) [=Hypogastrura (Ceratophysella) succinea] CB80a
         `--C. maheuxi group CB80a
              |--C. guthriei (Folsom 1916) [=Achorutes guthriei, Hypogastrura (Ceratophysella) guthriei] CB80a
              |--C. krafti (Scott 1962) [=*Spinachorutes krafti, Hypogastrura (Ceratophysella) krafti] CB80a
              `--C. maheuxi (Butler 1966) [=Hypogastrura (Ceratophysella) maheuxi] CB80a

*Ceratophysella armata (Nicolet 1841) CB80a, SL06 [=Achorutes armatus W39, Hypogastrura (*Ceratophysella) armata CB80a; incl. H. armata var. inermis W39, Achorutes armatus var. pallens S00]

Ceratophysella denticulata (Bagnall 1941) [=Achorutes denticulata, Hypogastrura (Ceratophysella) denticulata; incl. H. engadinensis Gisin 1949, H. exilis Yosii 1956] CB80a

Ceratophysella gibbosa (Bagnall 1940) CB80a, S07 [=Achorutes gibbosa CB80a, Hypogastrura (Ceratophysella) gibbosa CB80a; incl. H. azteca Yosii 1962 CB80a]

*Type species of generic name indicated

REFERENCES

[C90] Christiansen, K. A. 1990. Insecta: Collembola. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 965–995. John Wiley & Sones: New York.

[CB80a] Christiansen, K., & P. Bellinger. 1980a. The Collembola of North America north of the Rio Grande: A taxonomic analysis vol. 1. Introduction; general; families Poduridae and Hypogastruridae. Grinnell College: Grinnell (Iowa).

[CB80b] Christiansen, K., & P. Bellinger. 1980b. The Collembola of North America north of the Rio Grande: A taxonomic analysis vol. 2. Families Onychiuridae and Isotomidae. Grinnell College: Grinnell (Iowa).

[CB81] Christiansen, K., & P. Bellinger. 1981. The Collembola of North America north of the Rio Grande: A taxonomic analysis vol. 4. Families Neelidae and Sminthuridae; glossary; bibliography; index. Grinnell College: Grinnell (Iowa).

[DB21] Derkarabetian, S., C. M. Baker, M. Hedin, C. E. Prieto & G. Giribet. 2021. Phylogenomic re-evaluation of Triaenonychoidea (Opiliones: Laniatores), and systematics of Triaenonychidae, including new families, genera and species. Invertebrate Systematics 35: 133–157.

[G91] Greenslade, P. J. 1991. Collembola (springtails). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 252–264. Melbourne University Press: Carlton (Victoria).

[H83] Hammen, L. van der. 1983. Contribution to the knowledge of the soil-fauna of New Guinea. Zoologische Verhandelingen 206: 1–36.

[H44] Hammer, M. 1944. Studies on the oribatids and collemboles of Greenland. Meddelelser om Grønland 141 (3): 1–210.

[MC13] Majer, J. D., S. K. Callan, K. Edwards, N. R. Gunawardene & C. K. Taylor. 2013. Baseline survey of the terrestrial invertebrate fauna of Barrow Island. Records of the Western Australian Museum Supplement 83: 13–112.

[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.

[S00] Schäffer, C. 1900. Ueber württembergische Collembola. Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg 56: 245–280, pl. 2.

[SL06] Schulz, H.-J., & P. Lymberakis. 2006. First contribution to the knowledge of the Collembola fauna of the White Mountains (Lefká Óri) in west Crete (Insecta, Collembola, Isotomidae). Senckenbergiana Biologica 86 (2): 229–234.

[S07] Skarżyński, D. 2007. Two new species and a new record of the genus Ceratophysella Börner 1932 from North America (Insecta, Collembola, Hypogastruridae). Senckenbergiana Biologica 87 (2): 137–142.

[TM86] Thibaud, J.-M., & Z. Massoud. 1986. Insecta: Collembola. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 616–617. E. J. Brill/Dr W. Backhuys: Leiden.

[V73] Vuilleumier, F. 1973. Insular biogeography in continental regions. II. Cave faunas from Tessin, southern Switzerland. Systematic Zoology 22 (1): 64–76.

[W67] Wise, K. A. J. 1967. Collembola (springtails). Antarctic Research Series 10: 123–148.

[W39] Womersley, H. 1939. Primitive Insects of South Australia: Silverfish, springtails, and their allies. South Australian Branch of the British Science Guild: Adelaide.

Brachystomellidae

Juvenile Brachystomella parvula, copyright Andy Murray.

Belongs within: Collembola.

The Brachystomellidae are a group of elongate-bodied springtails that lack mandibles, with a toothed head present on the maxilla and flattened, blunt buccal cones (Greenslade 1991).

<==Brachystomellidae
    |--Salvarella wallacei G91
    |--Setanodosa G91
    |--Subclavontella G91
    |--Rapoportella G91
    |--Bellingeria Christiansen & Pike 2002 CP02
    |    `--*B. cornua Christiansen & Pike 2002 CP02
    `--Brachystomella Ågren 1903 [incl. Guacharia] W39
         |--*B. maritima Ågren 1903 CB80
         |--B. acantha Womersley 1933 W39
         |--B. afurcata Womersley 1933 W39
         |--B. anomala Womersley 1933 W39
         |--B. arida Christiansen & Bellinger 1980 CB80
         |--B. capitata Womersley 1930 W39
         |--B. christianseni H83
         |--B. contorta CB80
         |--B. curvula CB80
         |--B. dianae G91
         |--B. fungicola Womersley 1933 W39
         |--B. geniculata Womersley 1934 W39
         |--B. globulosa Cassagnau & Rapoport 1962 CB80
         |--B. granulata Womersley 1935 W39
         |--B. hiemalis Yosii 1956 CB80
         |--B. parvula (Schäffer 1896) [=Schoettella parvula] CB80
         |--B. platensis G91
         |--B. stachi Mills 1934 CB80
         |--B. terricola Womersley 1930 W39
         `--B. villalobosi Cassagnau & Rapoport 1962 CB80

*Type species of generic name indicated

REFERENCES

[CB80] Christiansen, K., & P. Bellinger. 1980. The Collembola of North America north of the Rio Grande: A taxonomic analysis vol. 1. Introduction; general; families Poduridae and Hypogastruridae. Grinnell College: Grinnell (Iowa).

[CP02] Christiansen, K., & E. Pike. 2002. Cretaceous Collembola (Arthropoda, Hexapoda) from the Upper Cretaceous of Canada. Cretaceous Research 23: 165–188.

[G91] Greenslade, P. J. 1991. Collembola (springtails). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 252–264. Melbourne University Press: Carlton (Victoria).

[H83] Hammen, L. van der. 1983. Contribution to the knowledge of the soil-fauna of New Guinea. Zoologische Verhandelingen 206: 1–36.

[W39] Womersley, H. 1939. Primitive Insects of South Australia: Silverfish, springtails, and their allies. South Australian Branch of the British Science Guild: Adelaide.

Tomoceridae

Tomocerus minor, copyright Francisco Welter-Schultes.

Belongs within: Collembola.

The Tomoceridae are a group of elongate springtails in which the third abdominal segment is longer than the fourth (Womersley 1939).

Characters (from Womersley 1939): Postantennal organ absent. Abdominal segment 3 longer than 4, all tergites free. With or without scales, if present then these with longitudinal ribs. Ciliated sensory setae present on abdominal segments or not. Furca present and well developed.

<==Tomoceridae
    |  i. s.: Lasiofinus CP02
    |         Entomocerus Christiansen & Pike 2002 CP02
    |           `--*E. mirus Christiansen & Pike 2002 CP02
    |--Lepidophorellinae W39
    |    |--Neophorella Womersley 1934 [Neophorellini] S-AB08
    |    |--Lepidophorella Schäffer 1897 (see below for synonymy) S-AB08
    |    |    |--L. australis Carpenter 1925 W39
    |    |    `--L. brachycephala (Moniez 1894) [=Drepanura brachycephala] W39
    |    |--Antennacyrtus Salmon 1941 S-AB08
    |    |--Lasofinius Ireson & Greenslade 1990 S-AB08
    |    `--Novacerus Salmon 1942 S-AB08
    `--Tomocerinae S-AB08
         |--Aphaenomurus Yosii 1956 S-AB08
         |--Monodontocerus Yosii 1955 S-AB08
         |--Tritomurus Frauenfeld 1854 S-AB08
         `--Tomocerus Nicolet 1842 CB80
              |  i. s.: T. bicolor (Say 1821) (n. d.) [=Podura bicolor] CB80
              |         T. iricolor Say 1821 (n. d.) CB80
              |         T. longicornis (Müller 1776) [=Podura longicornis] CB80
              |         T. lubbocki Schäffer 1900 S00
              |         T. niger Bourlet 1839 (n. d.) CB80
              |         T. nigritus Maynard 1951 (n. d.) CB80
              |         T. tasmanicus Womersley 1935 W39
              |--T. (Tomocerus) CB80
              |    |--*T. (T.) minor (Lubbock 1862) CB80 (see below for synonymy)
              |    `--T. (T.) vulgaris (Tullberg 1871) CB80, W39
              |--T. (Lethemurus Yosii 1970) CB80
              |    `--T. (*L.) missus Mills 1949 [=T. (Tritomurus) missus] CB80
              |--T. (Plutomurus Yosii 1956) CB80
              |    |--‘Tritomurus’ riugadoensis Yosii 1939 [=*Plutomurus riugadoensis] CB80
              |    |--T. (P.) brevimucronatus Denis 1929 CB80
              |    |--T. (P.) californicus Folsom 1913 [=Tritomuruc californicus; incl. Tr. oregonensis Denis 1929] CB80
              |    |--T. (P.) grahami Christiansen 1980 CB80
              |    `--T. (P.) wilkeyi Christiansen 1965 CB80
              |--T. (Pogonognathellus Paclt 1944) (see below for synonymy) CB80
              |    |--T. (*P.) plumbeus (Linnaeus 1758) [=Podura plumbea, *Pogonognathus plumbeus] CB80
              |    |--T. (P.) bidentatus Folsom 1913 (see below for synonymy) CB80
              |    |--T. (P.) celsus Christiansen 1965 CB80
              |    |--T. (P.) dubius Christiansen 1965 CB80
              |    |--T. (P.) elongatus Maynard 1951 [=*Maynardia elongata] CB80
              |    `--T. (P.) flavescens (Tullberg 1871) (see below for synonymy) CB80
              |--T. (Tomocerina Yosii 1955) CB80
              |    |--T. (*T.) minutus Tullberg 1876 CB80
              |    |--T. (T.) curtus Christiansen 1965 CB80
              |    |--T. (T.) lamelliferus Mills 1934 CB80
              |    `--T. (T.) teres Christiansen 1965 CB80
              `--T. (Tomolonus Mills 1949) CB80
                   `--T. (*T.) reductus (Mills 1949) [=*Tomolonus reductus] CB80

Lepidophorella Schäffer 1897 [incl. Millsia Womersley 1942, Pseudolepidophorella Salmon 1941; Lepidophorellini] S-AB08

Tomocerus (Pogonognathellus Paclt 1944) [=Pogonognathus Börner 1908 non Agassiz 1846; incl. Architomocerura Denis 1931, Maynardia Yosii 1956] CB80

Tomocerus (Pogonognathellus) bidentatus Folsom 1913 [incl. Tritomurus henroti Delamare 1949, To. jeanneli Bonet 1934] CB80

Tomocerus (Pogonognathellus) flavescens (Tullberg 1871) [=Macrotoma flavescens; incl. T. albus Packard 1877, T. americanus Schött 1896, T. arcticus Schött 1893, *Architomocerura crassicauda Denis 1931, T. pallidus Packard 1888, T. separatus Folsom 1913] CB80

*Tomocerus (Tomocerus) minor (Lubbock 1862) CB80 [=Macrotoma minor CB80; incl. M. tridentifera Tullberg 1872 CB80, Tomocerus tridentifer S00]

*Type species of generic name indicated

REFERENCES

[CB80] Christiansen, K., & P. Bellinger. 1980. The Collembola of North America north of the Rio Grande: A taxonomic analysis vol. 3. Family Entomobryidae. Grinnell College: Grinnell (Iowa).

[CP02] Christiansen, K., & E. Pike. 2002. Cretaceous Collembola (Arthropoda, Hexapoda) from the Upper Cretaceous of Canada. Cretaceous Research 23: 165–188.

[S00] Schäffer, C. 1900. Ueber württembergische Collembola. Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg 56: 245–280, pl. 2.

[S-AB08] Soto-Adames, F. N., J.-A. Barra, K. Christiansen & R. Jordana. 2008. Suprageneric classification of Collembola Entomobryomorpha. Annals of the Entomological Society of America 101 (3): 501–513.

[W39] Womersley, H. 1939. Primitive Insects of South Australia: Silverfish, springtails, and their allies. South Australian Branch of the British Science Guild: Adelaide.

Campodeoidea

Campodea staphylinus, copyright Michel Vuijlsteke.

Belongs within: Diplura.

The Campodeoidea are a group of diplurans with filiform cerci and lacking abdominal spiracles (Ferguson 1990).

Characters (from Ferguson 1990): Trichobothria present, beginning on antennal segment 3; mandible with lacinia mobilis (prostheca); maxilla without palpus, lacinia without pectens; urosternite 1 without styli, with one pair of subcoxal appendages; styli present on urosternites 2–7, with several distinctly developed setae; abdominal spiracles absent; cerci multisegmented, filiform, long or short, without inner canal, closed at apex; anal valvules well developed.

<==Campodeoidea [Campodeida, Campodeomorpha] F90
    |--Procampodea [Procampodeidae] F90
    |    `--P. macswaini Condé & Pagés 1956 F90
    `--Campodeidae F05
         |  i. s.: Plusiocampa F05
         |           |--P. baetica Sendra 2004 F05
         |           `--P. bonneti F05
         |                |--P. b. bonneti F05
         |                `--P. b. condei Sendra & Escolà 2004 F05
         |--Lepidocampinae F90
         |    |--Syncampa Silvestri 1931 F90
         |    `--Lepidocampa Oudemans 1890 F90
         |         `--L. weberi Oudemans 1890 F90
         |--Hemicampinae F90
         |    |--Tritocampa Silvestri 1933 F90
         |    `--Hemicampa Silvestri 1912 F90
         |         `--H. boudreauxi Condé & Geeraert 1962 F90
         `--Campodeinae CP91
              |--Eutrichocampa hispanica Silvestri 1932 F90
              |--Haplocampa Silvestri 1912 F90
              |--Podocampa Silvestri 1932 [incl. Parallocampa Silvestri 1933] F90
              |--Meiocampa Silvestri 1933 F90
              |--Eumesocampa Silvestri 1933 F90
              |    |--E. danielsi Silvestri 1933 F90
              |    |--E. frigillis (Hilton 1936) F90
              |    `--E. lutzi Silvestri 1933 F90
              |--Campodella Silvestri 1913 CP91, W39
              |    |--*C. clavigera CP91
              |    |--C. milloti CP91
              |    `--C. tiegsi Womersley 1939 W39
              |--Notocampa Silvestri 1930 CP91, W39
              |    |--*N. leae (Silvestri 1931) [=Metriocampa (*Notocampa) leae] W39
              |    |--N. philpotti (Tillyard 1924) [=Campodea philpotti, Metriocampa (Notocampa) philpotti] W39
              |    `--N. westraliense (Wom. 1937) [=Metriocampa (Notocampa) westraliense] W39
              |--Litocampa Silvestri 1933 F90
              |    |  i. s.: L. cookei F90
              |    `--L. (Cocytocampa Paclt 1957) F90
              |         |--L. (C.) perkinsi (Silvestri 1934) F90
              |         `--L. (C.) solomonis Bareth & Condé 1972 F90
              |--Metriocampa Silvestri 1911 F90
              |    |--M. (Austrocampa Wom. 1937) W39
              |    |    `--M. (A.) spinigera Wom. 1937 W39
              |    |         |--M. s. spinigera CP91
              |    |         `--M. s. victoriense Wom. 1937 CP91, W39
              |    `--M. (Tricampa Silvestri 1933) F90
              |         `--M. (T.) paradoxa CP91
              `--Campodea Westwood 1842 F90
                   |  i. s.: C. darwinii RJ93
                   |         C. lubbocki WS10
                   |         C. remyi F90
                   |         C. staphylinus P92
                   |         C. waterhousei Wom. 1937 W39
                   |--C. (Campodea) F05
                   |    |--C. (C.) cossetana Sendra & Escolà 2004 F05
                   |    |--C. (C.) fragilis Meinert 1865 CP91, W39
                   |    |--C. (C.) maestrazgoensis Sendra & Escolà 2004 F05
                   |    |--C. (C.) majorica F05
                   |    |    |--C. m. majorica F05
                   |    |    `--C. m. valentina Sendra & Moreno 2004 F05
                   |    `--C. (C.) navasi Sendra & Moreno 2004 F05
                   |--C. (Dicampa Silvestri 1932) F90
                   |    `--C. (D.) escalerai Silvestri 1932 F90
                   |--C. (Hypercampa Silvestri 1933) F90
                   |    `--C. (H.) essigi Silvestri 1933 F90
                   |--C. (Indocampa) CP91
                   |    |--C. (I.) froggattii Silvestri 1921 CP91, W39
                   |    |--C. (I.) novaecaledoniae CP91
                   |    `--C. (I.) tillyardii Silvestri 1931 CP91, W39
                   `--C. (Monocampa Silvestri 1932) F90
                        |--C. (M.) morgani Silvestri 1911 F90
                        `--C. (M.) tonnoiri Wom. 1937 CP91, W39

*Type species of generic name indicated

REFERENCES

[CP91] Condé, B., & J. Pagés. 1991. Diplura. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 269–271. Melbourne University Press: Carlton (Victoria).

[F90] Ferguson, L. M. 1990. Insecta: Diplura. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 951–963. John Wiley & Sones: New York.

[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.

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

[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.

[WS10] Wang, M., S. Sun, C. Li & X. Shen. 2010. Distinctive mitochondrial genome of the calanoid copepod Calanus sinicus, a useful molecular marker for phylogenetic and population studies. In: China-Russia Bilateral Symposium: Proceedings of the China-Russia Bilateral Symposium of "Comparison on Marine Biodiversity in the Northwest Pacific Ocean", 10–11 October 2010, Qingdao (China) pp. 187–192. Institute of Oceanology, Chinese Academy of Sciences; A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Russian Academy of Sciences.

[W39] Womersley, H. 1939. Primitive Insects of South Australia: Silverfish, springtails, and their allies. South Australian Branch of the British Science Guild: Adelaide.

Diplura

Heterojapyx novaezelandae, copyright Bugman-NZ.

Belongs within: Panhexapoda.
Contains: Campodeoidea, Japygidae.

The Diplura are a cosmopolitan group of slender-bodied, wingless, blind hexapods that are mostly found in damp soil under logs and stones (Condé & Pagés 1991).

Characters (from Condé & Pagés 1991): Small to large, narrow-bodied hexapods, mostly unpigmented; eyes absent; antennae moniliform, with intrinsic muscles in the flagellar segments; thorax usually weakly developed; legs 5-segmented; abdomen 10-segmented, with styles and exsertile vesicles on at least some of the segments, gonopore between 8 and 9; cerci variously developed. Larval development epimorphic.

<==Diplura
    |  i. s.: Onychojapyx schmidti Pierce 1950 K-P87
    |         Testajapyx Kukalová-Peck 1987 [Testajapygidae] K-P87
    |           `--*T. thomasi Kukalová-Peck 1987 K-P87
    |         Ocellia [Ocelliidae] RJ93
    |           `--O. articulicornis RJ93
    |--Rhabdura F90
    |    |--Campodeoidea F90
    |    `--Projapygoidea F90
    |         |--Anajapyx Silvestri 1903 [Anajapygidae] F90
    |         |    |--*A. vesiculosus W39
    |         |    |--A. hermosus Smith 1960 F90
    |         |    `--A. menkei F90
    |         |--Projapygidae CP91
    |         |    |--Projapyx Cook 1899 F90
    |         |    |    `--*P. stylifer W39
    |         |    `--Symphylurinus Silvestri 1909 F90
    |         |         |--*S. grassi W39
    |         |         `--S. swani CP91
    |         `--Octostigma [Octostigmatidae] P01
    |              |--*O. herbivorum [=O. herbivora] P01
    |              |--O. sinense [=O. sinensis] P01
    |              `--O. spiniferum Pagés 2001 P01
    `--Japygoidea [Dicellurata, Japygomorpha] F90
         |--Dinjapyx W39 [Dinjapygidae F90, Dinjapyginae]
         |--Japygidae CP91
         |--Evalljapyx Silvestri 1911 F90 [Evalljapygidae CP91, Evalljapyginae]
         |--Heterojapyx CP91 [Heterojapygidae, Heterojapyginae F90]
         |    |--H. evansi Wom. 1934 W39
         |    |--H. gallardi Till. 1924 W39
         |    |--H. novaezelandae W39
         |    |--H. soulei RD77
         |    |--H. tambourinensis Wom. 1934 W39
         |    |--H. tillyardi K-P83
         |    `--H. victoriae Silvestri 1910 W39
         `--Parajapygidae [Parajapyginae] F90
              |--Ectasjapyx W39
              `--Parajapyx CP91
                   |  i. s.: P. emeryanus MG06
                   |         P. isabellae S62
                   |         P. swani Wom. 1934 W39
                   `--P. (Parajapyx) B86
                        |--P. (P.) botosaneanui Pagés 1975 B86
                        `--P. (P.) gerlachi Pagés 1967 B86

*Type species of generic name indicated

REFERENCES

[B86] Botosaneanu, L. 1986. Insecta: Diplura (avec une note sur les Acari: Nematalycidae). In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) p. 618. E. J. Brill/Dr W. Backhuys: Leiden.

[CP91] Condé, B., & J. Pagés. 1991. Diplura. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 269–271. Melbourne University Press: Carlton (Victoria).

[F90] Ferguson, L. M. 1990. Insecta: Diplura. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 951–963. John Wiley & Sones: New York.

[K-P83] Kukalová-Peck, J. 1983. Origin of the insect wing and wing articulation from the arthropodan leg. Canadian Journal of Zoology 61: 1618–1669.

[K-P87] Kukalová-Peck, J. 1987. New Carboniferous Diplura, Monura, and Thysanura, the hexapod ground plan, and the role of thoracic side lobes in the origin of wings (Insecta). Canadian Journal of Zoology 65: 2327–2345.

[MG06] Mallatt, J., & G. Giribet. 2006. Further use of nearly complete 28S and 18S rRNA genes to classify Ecdysozoa: 37 more arthropods and a kinorhynch. Molecular Phylogenetics and Evolution 40: 772–794.

[P01] Pagés, J. 2001. Notes sur des diploures rabdoures (insectes, aptérygotes) n^o 2: Octostigma spiniferum sp. n. (Projapygoidea, Octostigmatidae) de Java (Indonésie)—Diplura Genavensia XXV. Revue Suisse de Zoologie 108 (1): 31–39.

[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.

[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.

[S62] Smith, L. M. 1962. Japygidae of South America. 3: Japygidae of Chile. Proceedings of the Biological Society of Washington 75: 273–292.

[W39] Womersley, H. 1939. Primitive Insects of South Australia: Silverfish, springtails, and their allies. South Australian Branch of the British Science Guild: Adelaide.

Japygidae

Metajapyx suterraneus, copyright Christopher C. Wirth.

Belongs within: Diplura.

The Japygidae are a group of diplurans with forcipate cerci and trichobothria on segments 4–6 of the antennae (Condé & Pagés 1991).

Characters (from Condé & Pagés 1991): Trichobothria present on segments 4–6 of antenna; mandible without prostheca; labial palps present; four pairs of thoracic spiracles; pretarsus without empodium on ventral side of the two lateral claws; S1–7 with minute membranous vesicles; cerci one-segmented, forcipate.

<==Japygidae
    |--Provalljapyx Silvestri 1949 [Provalljapyginae] F90
    |--Japygellinae W39
    |    |--Allurjapyx W39
    |    |--Epijapyx W39
    |    |--Japyginus W39
    |    `--Japygellus W39
    |--Indiajapyginae W39
    |    |--Burmjapyx glauerti W39, WM70
    |    |--Parindjapyx W39
    |    |--Neojapyx W39
    |    `--Indjapyx W39
    `--Japyginae S62
         |--Opisthjapyx W39
         |--Catajapyx W39
         |--Mixojapyx W39
         |--Megajapyx W39
         |--Hecajapyx S62
         |--Occasjapyx japonicus S62, SC17
         |--Austrjapyx S62
         |--Japygianus wheeleri WM70
         |--Japyx Halliday 1863 W39
         |    |--J. solifugus WS10
         |    `--J. westraliense Wom. 1934 W39
         |--Chiljapyx Smith 1962 S62
         |    `--*C. caltagironei Smith 1962 S62
         |--Penjapyx Smith 1962 S62
         |    `--*P. altus Smith 1962 S62
         |--Hapljapyx Silvestri 1948-1949 S62
         |    `--*H. lopesi Silvestri 1948-1949 S62
         |--Nelsjapyx Smith 1962 S62
         |    |--*N. hichinsi Smith 1962 S62
         |    `--N. soldadi Smith 1962 S62
         |--Rossjapyx Smith 1962 S62
         |    |--*R. australis Smith 1962 S62
         |    `--R. anodus (Silvestri 1949) [=Hapljapyx anodus; incl. H. subanodus Silvestri 1949] S62
         |--Valpjapyx Smith 1962 S62
         |    |--*V. botani Smith 1962 S62
         |    `--V. talcae Smith 1962 S62
         |--Teljapyx Silvestri 1949 S62
         |    |--*T. riestrae Silvestri 1949 S62
         |    |--T. megalocerus (Silv. 1901) S62
         |    `--T. profundus Smith 1962 S62
         |--Merojapyx Silvestri 1949 S62
         |    |--*M. speggazzinii Silvestri 1949 S62
         |    |--M. bidentatus [=M. bidentaus (l. c.)] S62
         |    |--M. porteri Silvestri 1949 S62
         |    `--M. riverosi Silvestri 1949 S62
         |--Notojapyx WM70
         |    |--N. mjobergi CP91
         |    `--N. tillyardi CP91
         |         |--N. t. tillyardi CP91
         |         `--N. t. pagesi CP91
         `--Metajapyx W39
              |--M. froggatti (Silvestri 1930) [=Japyx (Metajapyx) froggatti; incl. J. augens Silv. 1930] W39
              |--M. glauerti (Wom. 1934) [=Japyx (Metajapyx) glauerti] W39
              |--M. leae (Silvestri 1930) [=Japyx (Metajapyx) leae] W39
              |--M. longiseta (Silvestri 1908) [=Japyx (Metajapyx) longiseta] W39
              |--M. michaelseni (Silvestri 1930) [=Japyx (Metajapyx) michaelseni] W39
              |--M. mjoebergi (Silvestri 1928) [=Japyx (Metajapyx) mjoebergi] W39
              |--M. nichollsi (Wom. 1934) [=Japyx (Metajapyx) nichollsi] W39
              |--M. onkaparinga (Wom. 1935) [=Japyx (Metajapyx) onkaparinga] W39
              |--M. suterraneus RS10
              `--M. tillyardii (Silvestri 1930) [=Japyx (Metajapyx) tillyardii] W39

*Type species of generic name indicated

REFERENCES

[CP91] Condé, B., & J. Pagés. 1991. Diplura. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 269–271. Melbourne University Press: Carlton (Victoria).

[F90] Ferguson, L. M. 1990. Insecta: Diplura. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 951–963. John Wiley & Sones: New York.

[RS10] Regier, J. C., J. W. Shultz, A. Zwick, A. Hussey, B. Ball, R. Wetzer, J. W. Martin & C. W. Cunningham. 2010. Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences. Nature 463: 1079–1083.

[SC17] Schwentner, M., D. J. Combosch, J. P. Nelson & G. Giribet. 2017. A phylogenomic solution to the origin of insects by resolving crustacean-hexapod relationships. Current Biology 27: 1818–1824.

[S62] Smith, L. M. 1962. Japygidae of South America. 3: Japygidae of Chile. Proceedings of the Biological Society of Washington 75: 273–292.

[WM70] Wallace, M. M. H., & I. M. Mackerras. 1970. The entognathous hexapods. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 205–216. Melbourne University Press.

[WS10] Wang, M., S. Sun, C. Li & X. Shen. 2010. Distinctive mitochondrial genome of the calanoid copepod Calanus sinicus, a useful molecular marker for phylogenetic and population studies. In: China-Russia Bilateral Symposium: Proceedings of the China-Russia Bilateral Symposium of "Comparison on Marine Biodiversity in the Northwest Pacific Ocean", 10–11 October 2010, Qingdao (China) pp. 187–192. Institute of Oceanology, Chinese Academy of Sciences; A. V. Zhirmunsky Institute of Marine Biology, Far East Branch of the Russian Academy of Sciences.

[W39] Womersley, H. 1939. Primitive Insects of South Australia: Silverfish, springtails, and their allies. South Australian Branch of the British Science Guild: Adelaide.