Triketrion flabelliforme, overgrown by zoanthid Palythoa, from the Queensland Museum.

Belongs within: Poecilosclerida.

The Raspailiidae are a group of often branching sponges, with a distinct outer skeleton composed of spicule brushes grouped around extra-axial spicules.

Characters (from Hooper 1991): Typically branching, less often massive, flabelliform or encrusting. Surface typically hispid, even, but forms with microconules or entirely smooth surfaces also known. Ectosomal skeleton with ectosomal spicule brushes (fine styles, oxeas or anisoxeas) grouped around one or more long, central, extra-axial spicules (styles, oxeas or anisoxeas). Skeletal construction usually well developed, consisting of at least two differentiated structures: axially or basally condensed choanosomal skeleton, and extra-axial subectosomal skeleton, which may be radial, plumose or plumo-reticulate. Taxa with predominantly reticulate skeletal choanosomal architecture preserve at least some rudiments of extra-axial skeleton, in form of single (or groups of) subectosomal spicules. Spongin fibres usually echinated by acanthostyles which may be secondarily lost. Acanthostyle geometry extremely diverse. Structural megascleres varying from true styles to true oxeas, with intermediate forms. Microscleres occurring rarely, when present consisting of raphides (or microxeas).

    |--Cyamon aruense Hentschel 1912 BJ06
    |--*Axechina raspailiodes Hentschel 1912 BJ06
    |--Aulospongus MG-H11
    |--Ceratopsion dichotoma (Whitelegge 1907) MG-H11
    |--Echinodictyum cancellatum (Lamarck 1814) MG-H11
    |--Lithoplocamia MG-H11
    |--Thrinacophora cervicornis Ridley & Dendy 1886 MG-H11
    |--Trikentrion flabelliforme Carter 1882 MG-H11
    |--Raspailia MG-H11
    |    |  i. s.: R. fruticosa BJ06
    |    |           |--R. f. fruticosa BJ06
    |    |           `--R. f. aruensis Hentschel 1912 BJ06
    |    |--R. (Raspailia) vestigifera Dendy 1896 MG-H11
    |    |--R. (Clathriodendron) desmoxyiformis Hooper 1991 MG-H11
    |    `--R. (Syringella) rhaphidophora Hentschel 1912 BJ06
    `--Hymeraphia BJ06
         |--H. aruensis Hentzschel 1912 BJ06
         |--H. calochela Hentschel 1912 BJ06
         |--H. distincta Thiele 1903 BJ06
         |--H. lendenfeldi Hentschel 1912 BJ06
         |--H. longitoxa Hentschel 1912 BJ06
         |--H. rhopalophora Hentschel 1912 BJ06
         |--H. similis Thiele 1903 BJ06
         |--H. tetrastyla Hentschel 1912 BJ06
         |--H. thielei Hentschel 1912 BJ06
         `--H. toxifera Hentschel 1912 BJ06

*Type species of generic name indicated


[BJ06] Barnich, R., & D. Janussen. 2006. Die Typen und Typoide des Naturmuseums Senckenberg, Nr. 86. Type catalogue of the Porifera in the collections of the Senckenberg Museum in Frankfurt am Main, Germany. Senckenbergiana Biologica 86 (2): 127–144.

Hooper, J. N. A. 1991. Revision of the family Raspailiidae (Porifera : Demospongiae), with description of Australian species. Invertebrate Taxonomy 5: 1179–1418.

[MG-H11] McEnnulty, F. R., K. L. Gowlett-Holmes, A. Williams, F. Althaus, J. Fromont, G. C. B. Poore, T. D. O'Hara, L. Marsh, P. Kott, S. Slack-Smith, P. Alderslade & M. V. Kitahara. 2011. The deepwater megabenthic invertebrates on the western continental margin of Australia (100–1100 m depths): composition, distribution and novelty. Records of the Western Australian Museum Supplement 80: 1–191.


Sumatran rhinoceroses Dicerorhinus sumatrensis at Cincinnati Zoo, copyright Charles W. Hardin.

Belongs within: Ceratomorpha.

The Rhinocerotidae, rhinoceroses, are a group of large, heavy-bodied ungulates with living species in Africa and tropical Asia, though fossil representatives have a broader distribution in Eurasia and North America. Members of this group have more or less well-developed keratinous horns; Diceros, Ceratotherium and Dicerorhinus have one horn on the snout and one above the eyes whereas species of Rhinoceros possess the nose horn only.

Characters (from Animal Diversity Web): Body massive; skin thick, scantily-haired and wrinkled, furrowed or pleated, producing appearance of riveted armor plates in some species. Head large with one or two dermal horns composed of compressed, fibrous keratin; eyes small; ears fairly short but prominent and erect. Skull elongate, elevated posteriorly. Braincase small; nasal bones projecting forward freely, may extend beyond and above premaxillae; surface of nasals roughened beneath horns; occipital crest strongly developed. 24-34 teeth present, dental formula 1-2/0-1, 0/1-1, 3-4/3-4, 3/3; canines and incisors vestigial except for sometimes lower incisors which may be developed into powerful slashing tusks; cheek teeth with prominent transverse lophs of enamel. Chest broad; legs short, stumpy legs. Radius/ulna and tibia/fibula only slightly moveable but well-developed and separate. Fore- and hind feet mesaxonic with 3 digits each; each digit with a small hoof. Tail with stiff bristles.

<==Rhinocerotidae [Rhinocerotoidea]
    |--Diceros Gray 1821 FS15, H78
    |    |--D. bicornis H78
    |    |    |--D. b. bicornis R86
    |    |    |--D. b. brucii R86
    |    |    `--D. b. michaeli R86
    |    |--D. douariensis H78
    |    `--D. pachygnathus H78
    `--+--Ceratotherium Gray 1867 FS15, H78
       |    |--C. praecox Hooijer & Patterson 1972 H78
       |    `--C. simum H78
       |         |--C. s. simum H78
       |         |--C. s. cottoni BP87
       |         `--C. s. germanoafricanum H78
       `--+--Rhinoceros Linnaeus 1758 FS15, L58
          |    |--R. bicornis Linnaeus 1758 L58
          |    |--R. indicus C77
          |    |--R. sondaicus SM06
          |    |    |--R. s. sondaicus SM06
          |    |    `--R. s. annamiticus Heude 1892 SM06
          |    `--R. unicornis Linnaeus 1758 L58
          `--+--Dicerorhinus Gloger 1841 FS15, H78
             |    |--D. africanus Arambourg 1970 H78
             |    |--D. leakeyi Hooijer 1966 H78
             |    |--D. merckii FP64
             |    |--D. primaevus Arambourg 1959 H78
             |    |--D. ringstromi DW04
             |    |--D. sansaniensis H78
             |    |--D. schleiermacheri H78
             |    |--D. sumatrensis H78
             |    `--D. tagicus H78
             `--+--Coelodonta Blumenbach 1807 FS15, D07
                |    |--C. antiquitatis FS15
                |    `--C. nihowanensis DW04
                `--Stephanorhinus FS15
                     |--S. etruscus [=Rhinoceros etruscus, Dicerorhinus etruscus] ME05
                     |    |--S. e. etruscus ME05
                     |    `--‘Dicerorhinus’ e. brachycephalus ME05
                     |--S. hemitoechus FS15
                     |--S. hundsheimensis ME05
                     `--S. kirchbergensis FS15

Rhinocerotidae incertae sedis:
  Allacerops DW04
  Chilotherium H78
    |--C. anderssoni DW04
    |--C. palaeosinense CS04
    `--C. wimani DW04
  Tichorhinus antiquitates B61
  Trigonias Lucas 1900 D07
    |--T. osborni D07
    |--T. wellsi D07
    `--T. yoderensis D07
  Chilotheridium Hooijer 1971 H78
    `--*C. pattersoni Hooijer 1971 H78
  Aceratherium Kaup 1832 H78
    |--A. acutirostratum (Deraniyagala 1951) [=Turkanatherium acutirostratus] H78
    |--A. incisivum H78
    |--A. lemanense H78
    `--A. occidentale C77
  Brachypotherium Roger 1904 H78
    |--B. aurelianense (Nouel 1866) [=Rhinoceros aurelianensis, Teleoceras aurelianense] H78
    |--B. brachypus H78
    |--B. goldfussi H78
    |--B. heinzelini Hooijer 1963 H78
    |--B. lewisi Hooijer & Patterson 1972 H78
    |--B. perimense CS04
    |--B. snowi (Fourtau 1920) [=Teleoceras snowi] H78
    `--B. stehlini Viret 1961 H78
  0--Aphelops Cope 1873 H78, C77
  |    |--A. crassus [=Rhinocerus crassus] C77
  |    |--A. jemezanus Cope 1877 C77
  |    |--A. megalodus C77
  |    `--A. meridianus [=Rhinocerus meridianus] C77
  `--Teleoceras Hatcher 1894 H78
       |--T. fossiger D07
       |--T. guymonense D07
       |--T. hicksi D07
       |--T. major D07
       |--T. medicornutum H78
       |--T. meridianum D07
       `--T. proterum D07
  Paradiceros Hooijer 1968 H78
    `--P. mukirii Hooijer 1968 H78
  Hyrachyus Leidy 1870 HUG17, D07 [Hyrachyidae]
    |--H. asiaticus D07
    |--H. eximius D07
    |--H. modestus D07
    `--H. singularis Cope 1875 C77

Inorganic: Rhinoceros unicornis protominilorientalis Okamura 1987 O87

*Type species of generic name indicated


[B61] Banfield, A. W. F. 1961. A revision of the reindeer and caribou, genus Rangifer. National Museum of Canada Bulletin 177: i–vi, 1–137.

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[CS04] Chaimanee, Y., V. Suteethorn, P. Jintasakul, C. Vidthayanon, B. Murandat & J.-J. Jaeger. 2004. A new orang-utan relative from the Late Miocene of Thailand. Nature 427: 439–441.

[C77] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4 (2): i–iv, 1–370.

[DW04] Deng T., Wang X., Ni X. & Liu L. 2004. Sequence of the Cenozoic mammalian faunas of the Linxia Basin in Gansu, China. Acta Geologica Sinica (English Edition) 78 (1): 8–14.

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[FS15] Faurby, S., & J.-C. Svenning. 2015. A species-level phylogeny of all extant and late Quaternary extinct mammals using a novel heuristic-hierarchical Bayesian approach. Molecular Phylogenetics and Evolution 84: 14–26.

[FP64] Fisher, J., & R. T. Peterson. 1964. The World of Birds: A comprehensive guide to general ornithology. Macdonald: London.

[HUG17] Halliday, T. J. D., P. Upchurch & A. Goswami. 2017. Resolving the relationships of Paleocene placental mammals. Biological Reviews 92 (1): 521–550.

[H78] Hooijer, D. A. 1978. Rhinocerotidae. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 371–378. Harvard University Press: Cambridge (Massachusetts).

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[ME05] Moullé, P.-E., A. Echassoux, F. Lacombat, E. Desclaux & S. Bailon. 2005. L’environnement animal des premiers habitants de l’Europe méditerranéenne: les grands mammifères contemporains de l’homme du Vallonnet, données taxonomiques et biostratigraphiques pour la deuxième moitie du Pléistocène inférieur. BAR International Series 1364: 105–113.

[O87] Okamura, C. 1987. New facts: Homo and all Vertebrata were born simultaneously in the former Paleozoic in Japan. Original Report of the Okamura Fossil Laboratory 15: 347–573.

[R86] Ryder, O. A. 1986. Species conservation and systematics: the dilemma of subspecies. Trends in Ecology and Evolution 1 (1): 9–10.

[SM06] Schwendinger, P., & J. Martens. 2006. A taxonomic revision of the family Oncopodidae V. Gnomulus from Vietnam and China, with the description of five new species (Opiliones, Laniatores). Revue Suisse de Zoologie 113 (3): 595–615.

Last updated: 31 July 2018.


Tumulitermes soldiers, copyright Melinda Hillery.

Belongs within: Termitinae.
Contains: Eutermes, Nasutitermes, Bulbitermes.

The Nasutitermitinae, nasute termites, are a distinct group of termites in which the head capsule of the soldier caste has an enlarged frontal gland and the fontanelle placed on a protruding rostrum, allowing the ejection of defensive chemicals at threats to the colony. The development of the rostrum is correlated with the reduction of the mandibles which are usually non-functional in the soldiers. The mandibles retain sharp points in the basal genera Angularitermes and Mimeutermes (Ahmad 1950). Members of the African genera Mimeutermes and Postsubulitermes feed on humus, presumably digesting the decomposed vegetable matter it contains.

<==Nasutitermitinae [Nasutitermitidae]
    |--+--Angularitermes nasutissimus IVE07, A50
    |  |--Mimeutermes giffardii A50
    |  `--Postsubulitermes parviconstrictus IVE07
    `--+--+--Cyranotermes caete IVE07
       |  |--Atlantitermes oculatissimus IVE07
       |  |--Obtusitermes panamae IVE07, A50
       |  |--Coatitermes kartaboensis IVE07
       |  `--+--Velocitermes IVE07
       |     |    |--V. beebei S57
       |     |    `--V. heteropterus [=Eutermes heteropterus] S57
       |     `--+--Convexitermes manni A50
       |        `--Subulitermes IVE07
       |             |--*S. microsoma A50 [=Nasutitermes microsoma S57]
       |             |--S. baileyi IVE07
       |             |--S. oculatissimus A50
       |             `--S. undecimus RD77
       `--+--+--Fulleritermes coatoni IVE07
          |  `--+--Leptomyxotermes doriae IVE07
          |     `--Rhadinotermes coarctus IVE07
          `--+--+--Oriensubulitermes inanis IVE07
             |  `--+--‘Nasutitermes’ longinasus IVE07
             |     `--Havilanditermes IVE07
             |          |--H. atripennis A50
             |          `--H. proatripennis IVE07
             `--+--Nasutitermes IVE07
                |--Leucopitermes leucops IVE07
                |--+--Aciculitermes maymyoensis IVE07
                |  `--Malaysiotermes spinocephalus IVE07
                |--+--Australitermes IVE07
                |  |    |--A. dilucidus WG91
                |  |    |--A. insignitus [=Termes insignitus] WG91
                |  |    `--A. perlevis [=Macrosubulitermes perlevis] WG91
                |  `--+--+--Eutermellus convergens A50
                |     |  `--Occasitermes IVE07
                |     |       |--O. occasus (Silvestri 1909) [=Eutermes (Occasitermes) occasus] H42
                |     |       `--O. watsoni WG91
                |     `--Tumulitermes IVE07
                |          |  i. s.: T. peracutus (Hill 1925) J13
                |          |         T. recalvus WG91
                |          |--T. westraliensis IVE07
                |          `--+--T. apiocephalus (Silvestri 1909) IVE07, H42 (see below for synonymy)
                |             `--+--T. hastilis (Froggatt 1898) IVE07, J13
                |                `--‘Nasutitermes’ eucalypti (Mjöberg 1920) IVE07, J13
                `--+--Bulbitermes IVE07
                   `--+--Constrictotermes S57
                      |    |--C. cavifrons S57
                      |    `--C. cyphergaster S57
                      `--+--Lacessititermes IVE07
                         |    |--L. batavus A50
                         |    `--L. laborator S57
                         `--+--Longipeditermes longipes IVE07
                            `--Hospitalitermes IVE07
                                 |--H. asahinai SVT04
                                 |--H. ataramensis SVT04
                                 |--H. bicolor SVT04
                                 |--H. birmanicus SVT04
                                 |--H. butteli [=Eutermes butteli] S57
                                 |--H. hospitalis A50
                                 |--H. jepsoni SVT04
                                 |--H. medioflavus SVT04
                                 `--H. monoceros IVE07

Nasutitermitinae incertae sedis:
  Eutermes H42
  Macrosubulitermes greavesi WG91
  Occultitermes WG91
    |--O. aridus WG91
    `--O. occultus WG91
  Diversitermes diversimilis [=Nasutitermes diversimiles] S57
  Triacitermes triacifer A50
  Rhynchotermes nasutissimus A50
  Hirtitermes spinocephalus A50
  Grallatotermes A50
    |--G. africanus S57
    `--G. splendidus A50
  Rotunditermes rotundiceps A50
  Parvitermes A50
    |--P. discolor A50
    `--P. pallidiceps A50

Tumulitermes apiocephalus (Silvestri 1909) IVE07, H42 [=Eutermes apiocephalus H42; incl. E. (Tumulitermes) aagaardi Mjöberg 1920 H42]

*Type species of generic name indicated


[A50] Ahmad, M. 1950. The phylogeny of termite genera based on imago-worker mandibles. Bulletin of the American Museum of Natural History 95 (2): 37–86.

[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.

[J13] Jones, D. T. 2013. The termites of Barrow Island, Western Australia. Records of the Western Australian Museum Supplement 83: 241–244.

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

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

[SVT04] Sornnuwat, Y., C. Vongkaluang & Y. Takematsu. 2004. A systematic key to termites of Thailand. Kasetsart Journal of Natural Science 38 (3): 349–368.

[WG91] Watson, J. A. L., & F. J. Gay. 1991. Isoptera (termites). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 330–347. Melbourne University Press: Carlton (Victoria).

Last updated: 21 May 2022.


Colony of Nasutitermes luzonicus, copyright Choon Huat Quah.

Belongs within: Nasutitermitinae.

Nasutitermes is a pantropical genus of nasute termites in which soldiers are generally monomorphic and lack a constriction on the head capsule behind the antennal sockets. The genus is poorly defined relative to other nasute genera and may not be monophyletic.

<==Nasutitermes Banks 1920 H42
    |--N. neoparvus IVE07
    `--+--+--N. latifrons IVE07
       |  `--+--N. infuscatus IVE07
       |     `--Trinervitermes IVE07
       |          |--T. biformis [=Eutermes biformis] S57
       |          |--T. geminatus IVE07
       |          |--T. heimi [=Eutermes heimi] S57
       |          |--T. rapulum [=Eutermes rapulum] S57
       |          |--T. rhodesiensis N55
       |          |--T. trinerviformis [=Eutermes trinerviformis] S57
       |          |--T. trinervius S57
       |          `--T. trinervoides HD-S02
       `--+--+--+--N. ceylonicus IVE07 [=Eutermes ceylonicus S57]
          |  |  `--N. lacustris IVE07
          |  `--+--+--N. canaliculatus IVE07
          |     |  `--N. lujae IVE07 [incl. Eutermes lujae S57]
          |     `--+--Malagasitermes milloti IVE07
          |        `--Coarctotermes IVE07
          |             |  i. s.: C. brunneus Noirot 1955 N55
          |             |--C. beharensis IVE07
          |             `--+--C. clepsydra IVE07
          |                `--C. pauliani IVE07
          `--+--+--N. arborum IVE07
             |  `--N. fulleri IVE07
             `--+--N. glabritergus IVE07
                `--+--+--N. similis IVE07
                   |  `--+--N. corniger IVE07
                   |     `--+--N. matangensis IVE07 [=Eutermes matangensis S57]
                   |        `--N. voeltzkowi IVE07
                   `--+--N. graveolus IVE07
                      |--N. triodea (Froggatt 1898) IVE07, J13
                      |--+--N. dasyopsis IVE07
                      |  `--N. nigriceps IVE07 [incl. N. acajutlae S57, N. pilifrons S57]
                      `--+--N. longipennis (Hill 1915) IVE07, J13
                         `--‘Tumulitermes’ pastinator IVE07

Nasutitermes incertae sedis:
  N. amboinensis RD77
  N. aquilinus S57
  N. arenarius S57
  N. bivalens S57
  N. brachynasutus SVT04
  N. brevirostris S57
  N. carnarvonensis WG91
  N. chaquimayensis S57
  N. columbicus S57
  N. corporaali S57
  N. costalis S57
  N. coxipoensis S57
  N. dendrophilus S57
  N. dimorphus SVT04
  N. dixoni WG91
  N. ehrhardti S57
  N. electrodominicus GE05
  N. ephratae S57
  N. exitiosus WG91
  N. fulviceps S57
  N. fumipennis S57
  N. fuscipennis SVT04
  N. gaigei S57
  N. globiceps GE05
  N. guayanae S57
  N. havilandi SVT04
  N. incurvus S57
  N. intermedius S57
  N. itapocuensis S57
  N. jaraguae S57
  N. johoricus SVT04
  N. luzonicus [incl. Eutermes laspinasensis, N. laspinasensis] S57
  N. maculiventris S57
  N. magnus (Froggatt 1898) J13
  N. matagensiformis SVT04
  N. meinerti [=Eutermes meinerti] S57
  N. minimus S57
  N. mojosensis S57
  N. novarumhebridarum S57
  N. octopilis S57
  N. olidus S57
  N. peruanus S57
  N. pilifrons S57
  N. pilosus S57
  N. pluriarticulatus S57
  N. preparvus SVT04
  N. profuscipennis SVT04
  N. proximus S57
  N. rufirostris S57
  N. sanctaeanae S57
  N. surinamensis S57
  N. torquatus S57
  N. tungsalangensis SVT04
  N. usambarensis S57
  N. walkeri WG91

*Type species of generic name indicated


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

[HD-S02] Haddad, C. R., & A. S. Dippenaar-Schoeman. 2002. The influence of mound structure on the diversity of spiders (Araneae) inhabiting the abandoned mounds of the snouted harvester termite Trinervitermes trinervoides. Journal of Arachnology 30 (2): 403–408.

[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.

[J13] Jones, D. T. 2013. The termites of Barrow Island, Western Australia. Records of the Western Australian Museum Supplement 83: 241–244.

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

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

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

[SVT04] Sornnuwat, Y., C. Vongkaluang & Y. Takematsu. 2004. A systematic key to termites of Thailand. Kasetsart Journal of Natural Science 38 (3): 349–368.

[WG91] Watson, J. A. L., & F. J. Gay. 1991. Isoptera (termites). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 330–347. Melbourne University Press: Carlton (Victoria).


Mango Mangifera indica, copyright Fpalli.

Belongs within: Anacardiaceae.

Mangifera is a genus of trees found primarily in lowland rainforest in tropical Asia, including the mango Mangifera indica and related species.

Characters (from Flora of China): Evergreen trees, andro-monoecious with functionally staminate and bisexual flowers in same inflorescence. Leaves petiolate, simple; leaf blade margin entire. Inflorescence a terminal pleiothyrsoid. Flowers small, 4- or 5-merous, with imbricate aestivation. Sepals sometimes united at base. Petals with 1-5 prominent veins adaxially. Stamens 5, free or united with disk basally, usually 1(or 2) noticeably larger. Disk of 5 separate extrastaminal glands. Ovary glabrous, 1-locular, 1-ovulate; style 1, terminal. Fruit drupaceous; mesocarp fleshy or fibrous; endocarp thick and bony, compressed. Embryo 1 to several by polyembryony.

<==Mangifera [=M. subg. Mangifera] YH02
    |  i. s.: M. altissima YH02
    |         M. caesia YH02
    |         M. kemanga YH02
    |         M. pajang YH02
    |         M. zeylanica YH02
    |--M. macrocarpa [=M. (subg. Limus sect. Perennis) macrocarpa] YH02
    |--M. sect. Rawa YH02
    |    |--M. griffithii YH02
    |    `--+--M. gracilipes YH02
    |       `--M. sect. Euantherae YH02
    |            |--M. caloneura YH02
    |            `--M. pentandra YH02
    `--+--M. cochinchinensis [=M. (sect. Euantherae) cochinchinensis] YH02
       |--M. subg. Limus sect. Perennis YH02
       |    |--M. foetida YH02
       |    `--M. odorata YH02 [M. foetida × M. indica KTG03]
       `--M. sect. Mangifera YH02
            |--+--M. flava YH02
            |  `--M. (sect. Marchandora) gedebe YH02
            `--+--M. oblongifolia YH02
               `--+--M. indica YH02
                  |--M. laurina YH02
                  `--M. sylvatica YH02

*Type species of generic name indicated


[KTG03] Kiew, R., L. L. Teo & Y. Y. Gan. 2003. Assessment of the hybrid status of some Malesian plants using Amplified Fragment Length Polymorphism. Telopea 10 (1): 225–233.

[YH02] Yonemori, K., C. Honsho, S. Kanzaki, W. Eiadthong & A. Sugiura. 2002. Phylogenetic relationships of Mangifera species revealed by ITS sequences of nuclear ribosomal DNA and a possibility of their hybrid origin. Plant Systematics and Evolution 231: 59–75.


Paederus riparius, from here.

Belongs within: Paederini.

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

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

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

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

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

*Type species of generic name indicated


[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

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

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

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso “Esploratore” dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 5–72.

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

[M86] Macleay, W. 1886. The insects of the Fly River, New Guinea, "Coleoptera". Proceedings of the Linnean Society of New South Wales, series 2, 1 (1): 136–157.

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

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

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

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

Last updated: 10 April 2022.


Abroteles beaumonti, from Seevers (1957).

Belongs within: Corotocini.

The Abrotelina are a group of distinctively frail-bodied physogastric staphylinid beetles found in association with nasutitermitine termites in the Neotropical region (Seevers 1957).

Characters (from Seevers 1957): Frail-bodied, frequently with very thin sclerites; head and thorax slender, abdomen ovoidal, weakly to strongly physogastric. Entire clypeus very weakly sclerotised or not sclerotised; very little sclerotisation anterior to tentorial pit level. Pronotum small, its surface not impressed. Tergites relatively narrow, about three-fifths as broad as abdomen; inner paratergites large, about one-third as broad as tergites. Elytra generalized, contiguous. Wings normal. Antennae 11-segmented. Tarsi 4, 4, 4-segmented or 5, 5, 5-segmented. Abdominal segments 7-9 distinctive.

<==Abrotelina S57
    |--Oecidiophilus Silvestri 1946 S57
    |    `--*O. mimellus Silvestri 1946 [=O. oglobinii ms] S57
    |--Parvidolum Silvestri 1946 S57
    |    `--*P. microsomatis Silvestri 1946 [=P. microsamatis] S57
    |--Mormellus Silvestri 1946 S57
    |    `--*M. bicolor Silvestri 1946 S57
    |--Termitosius Silvestri 1901 S57
    |    `--*T. pauciseta Silvestri 1901 S57
    |--Termitoiceus Silvestri 1901 S57
    |    |--*T. anastrephoproctus Silvestri 1901 [=T. anastrephoroctus (l. c.)] S57
    |    `--T. simulans Silvestri 1946 [=T. anastrephoproctus simulans] S57
    |--Perlinctus Silvestri 1946 S57
    |    |--*P. quaesitus Silvestri 1946 S57
    |    `--P. foveicollis Seevers 1957 S57
    |--Eburniogaster Seevers 1938 S57
    |    |--*E. termitocola Seevers 1938 S57
    |    |--E. anahuaci Seevers 1957 S57
    |    `--E. texana (Brues 1902) [=Termitogaster texana] S57
    |--Fonsechellus Silvestri 1946 [incl. Trianellus Silvestri 1946] S57
    |    |--*F. diversicolor Silvestri 1946 S57
    |    |--F. bicolor Silvestri 1946 [=F. (*Trianellus) bicolor] S57
    |    `--F. fragilis Seevers 1957 S57
    |--Termitonidia Seevers 1938 S57
    |    |--*T. lunata Seevers 1938 S57
    |    |--T. michoacani Seevers 1957 S57
    |    |--T. jaliscensis Seevers 1957 S57
    |    `--T. tarascani Seevers 1957 S57
    `--Abroteles Casey 1890 [incl. Termitothymus Silvestri 1901] S57
         |--*A. beaumonti Casey 1890 S57
         |--A. badia Seevers 1957 S57
         |--A. bisetosus Seevers 1937 S57
         |--A. compacticornis Borgmeier 1950 S57
         |--A. lobata Seevers 1957 S57
         |--A. philetaerus (Silvestri 1901) (n. d.) [=*Termitothymus philetaerus] S57
         |--A. pubicollis Seevers 1957 S57
         `--A. rurrenabaquensis Seevers 1946 S57

*Type species of generic name indicated


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


Thyreoxenus brevitibialis, copyright Taro Eldredge.

Belongs within: Staphylinidae.
Contains: Abrotelina, Termitogastrina.

The Corotocini are a group of physogastric staphylinid beetles found mostly in association with nasutitermitine termites, with a very few species found with the termitine genus Anoplotermes (Seevers 1957).

See also: The Corotocini in their gut-swollen glory.

Characters (from Seevers 1957): Mentum fused to submentum; mesocoxal acetabula not margined; metepimera abbreviated; hind coxae triangular; physogastric.

<==Corotocini S57
    |  i. s.: Termitochara Wasmann 1893 [Termitocharina] S57
    |           `--*T. kraatzi Wasmann 1893 S57
    |--Timeparthenina S57
    |    |--Termituncula Borgmeier 1950 S57
    |    |    `--*T. gracilipes Borgmeier 1950 S57
    |    `--+--Termitozophilus Silvestri 1901 [incl. Corymbogaster Mann 1923] S57
    |       |    |--*T. laetus Silvestri 1901 S57
    |       |    `--T. mirandus (Mann 1923) [=*Corymbogaster miranda] S57
    |       `--+--Autuoria Silvestri 1946 S57
    |          |    `--*A. elegantulum Silvestri 1946 [incl. A. orthocephali ms] S57
    |          `--+--Ptocholellus Silvestri 1946 S57
    |             |    `--*P. mimus Silvestri 1946 S57
    |             `--Timeparthenus Silvestri 1901 S57
    |                  |--*T. regius Silvestri 1901 S57
    |                  `--T. oglobini Silvestri 1946 [=Timeparthemus (l. c.) oglobini] S57
    `--+--+--Abrotelina S57
       |  `--Termitogastrina S57
       `--Corotocina S57
            |  i. s.: Eutermitoptochus Silvestri 1921 S57
            |           `--*E. novaehollandiae Silvestri 1921 S57
            |--Corotoca Schiødte 1853 S57
            |    |--*C. melantho Schiødte 1854 S57
            |    |--C. araujoi Seevers 1957 S57
            |    |--C. guyanae Mann 1923 S57
            |    `--C. phylo Schiødte 1854 S57
            `--+--+--Termitomimus Trägärdh 1907 S57
               |  |    |--*T. entendveniensis Trägärdh 1907 S57
               |  |    |--T. emersoni Seevers 1957 S57
               |  |    `--T. latipes Seevers 1957 S57
               |  `--+--Termitopullus Reichensperger 1922 [incl. Termitoscapha Bernhauer 1938] S57
               |     |    `--*T. sociusculus Reichensperger 1922 [incl. *Termitoscapha gestroi Bernhauer 1938] S57
               |     |--Termitoptocinus Silvestri 1921 S57
               |     |    `--*T. australiensis Silvestri 1921 S57
               |     `--Eburniola Mann 1923 S57
               |          |--*E. leucogaster Mann 1923 S57
               |          |--E. gastrovittata Seevers 1937 S57
               |          `--E. lujae Seevers 1957 S57
               `--+--Spirachtha Schiødte 1853 S57
                  |    |--*S. eurymedusa Schiødte 1854 S57
                  |    |--S. mirabilis Mann 1923 S57
                  |    `--S. schioedtei Mann 1923 S57
                  `--+--Oideprosoma Silvestri 1920 S57
                     |    `--*O. mirandum Silvestri 1920 S57
                     `--Thyreoxenus Mann 1923 S57
                          |--*T. parviceps Mann 1923 S57
                          |--T. albidus Seevers 1946 S57
                          |--T. autuorii Silvestri 1946 S57
                          |--T. boliviae Seevers 1946 S57
                          |--T. brevitibialis Seevers 1946 S57
                          |--T. convexinotus Seevers 1946 S57
                          |--T. cucullatus Seevers 1946 S57
                          |--T. major S57
                          |--T. pulchellus Mann 1923 S57
                          `--T. solomonensis Seevers 1937 S57

*Type species of generic name indicated


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


Termitonannus vagans, from Seevers (1957).

Belongs within: Staphylinidae.

The Termitonannini are a group of mostly limuloid staphylinid beetles found in association with nasutitermitine termites (Seevers 1957).

Characters (from Seevers 1957): Body usually non-physogastric and limuloid, very rarely physogastric. Head either prognathous or hypognathous; antennae 9-11-segmented, terminal antennal segments bearing one or more pairs of polytrichous coeloconic sensilla; mentum free except in certain derived genera. Mesocoxal cavities margined; hind coxae and metepimera generalised except in certain derived genera. Tarsi 4, 4, 4-segmented or 4, 4, 5-segmented.

<==Termitonannini S57
    |--Termitonannina S57
    |    |  i. s.: Termitocomes Seevers 1941 S57
    |    |           |--*T. wasmanni Seevers 1941 S57
    |    |           `--T. trinidadensis Seevers 1957 S57
    |    |--+--Eunannodes Silvestri 1946 S57
    |    |  |    `--*E. reconditi Silvestri 1946 S57
    |    |  `--Nannellus Silvestri 1946 S57
    |    |       `--*N. anoplotermitis Silvestri 1946 S57
    |    `--+--Termitonilla Borgmeier 1950 S57
    |       |    `--*T. luteola Borgmeier 1950 S57
    |       `--+--Macrotrichuris Silvestri 1946 S57
    |          |    |--*M. brasiliensis Silvestri 1946 S57
    |          |    `--M. notabilis Silvestri 1946 S57
    |          `--Termitonannus Wasmann 1902 [incl. Tetraphilus Silvestri 1946] S57
    |               |--*T. schmalzi Wasmann 1902 S57
    |               |--T. brachycerus Silvestri 1946 [=T. (*Tetraphilus) brachycerus] S57
    |               |--T. domunculi Silvestri 1946 S57
    |               |--T. echinoides Seevers 1946 S57
    |               |--T. gatuni Seevers 1941 S57
    |               |--T. major Wasmann 1902 S57
    |               |--T. microsomatis Silvestri 1946 S57
    |               |--T. parvulus Silvestri 1946 S57
    |               |--T. proximatus Silvestri 1946 S57
    |               |--T. setosus Seevers 1941 S57
    |               |--T. silvestrii Wasmann 1902 S57
    |               `--T. vagans Silvestri 1946 S57
    `--Perinthina [Perinthini] S57
         |--Termitopelta Borgmeier 1950 S57
         |    `--*T. fulgens Borgmeier 1950 S57
         `--+--Termitonicus Mann 1926 S57
            |    |--*T. mahout Mann 1926 S57
            |    `--T. uroclaviger Silvestri 1946 S57
            `--+--Macrognathellus Silvestri 1946 S57
               |    `--M. crassicornis (Wasmann 1902) (see below for synonymy) S57
               `--+--Poduroides Mann 1926 S57
                  |    `--*P. bovingi Mann 1926 S57
                  `--+--Perinthus Casey 1890 S57
                     |    |--*P. dudleyanus Casey 1890 S57
                     |    |    |--P. d. dudleyanus S57
                     |    |    |--P. d. major Seevers 1957 S57
                     |    |    `--P. d. wasmanni Mann 1923 S57
                     |    |--P. bolivari Seevers 1946 S57
                     |    |--P. guatemalae Seevers 1946 S57
                     |    |--P. hageni Seevers 1937 S57
                     |    |--P. mayae S57
                     |    |--P. tarsatus Mann 1923 S57
                     |    `--P. xenocostalis Seevers 1937 S57
                     `--+--Termitocola Seevers 1937 S57
                        |    |--*T. cylindricornis Seevers 1937 S57
                        |    `--T. silvestrii (Wasmann 1902) [=Perinthus silvestrii] S57
                        `--+--Lauella Mann 1921 S57
                           |    |--*L. vitiensis Mann 1921 [incl. Hetairotermes leai Cameron 1927] S57
                           |    |--L. javana Seevers 1957 S57
                           |    `--L. palauensis Seevers 1957 S57
                           `--+--Eutermitophila Cameron 1939 S57
                              |    `--*E. fletcheri Cameron 1939 S57
                              `--+--Perinthodes Seevers 1957 S57
                                 |    `--*P. africanus Seevers 1957 S57
                                 `--Paraperinthus Seevers 1957 S57
                                      `--*P. pauciseta Seevers 1957 S57

Macrognathellus crassicornis (Wasmann 1902) [=Perinthus crassicornis; incl. *M. paraguayensis Silvestri 1946] S57

*Type species of generic name indicated


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


Termitogaster insolens, from Seevers (1957).

Belongs within: Corotocini.

The Termitogastrina are a lineage of physogastric staphylinids found in association with nasutitermitine termites of the genera Nasutitermes, Trinervitermes and Grallatotermes. Though difficult to characterise as a whole due to the derived morphology of some genera, but all carry the variously inflated abdomen recurved over the foreparts. In less derived species, the tarsi are 5, 5, 5-segmented, albeit with the fifth segment partially fused and not movable relative to the fourth segment; in some derived species the tarsi have become 4, 4, 4-segmented. These species have the sixth and seventh tergites characteristically enlarged with the seventh tergite shield-shaped (Seevers 1957).

Termitogastrina S57
    |--Termitopithus Seevers 1957 S57
    |    `--*T. crassiusculus Seevers 1957 S57
    |--Termitophya Wasmann 1902 S57
    |    |--*T. heyeri Wasmann 1902 S57
    |    |--T. amica Mann 1923 S57
    |    |--T. araujoi Borgmeier 1950 S57
    |    |--T. ecuadoriensis Seevers 1937 S57
    |    |--T. emersoni Seevers 1937 S57
    |    |--T. flaviventris Mann 1923 S57
    |    |--T. holmgreni Wasmann in Wasmann & Holmgren 1911 [incl. T. wasmanni Holmgren in Wasmann & Holmgren 1911] S57
    |    |--T. inornata Seevers 1937 S57
    |    |--T. piliventris Seevers 1937 S57
    |    `--T. punctata Mann 1923 S57
    `--+--+--Termitomorpha Wasmann 1894 [incl. Termitosomus Seevers 1939, Thaxteria Fenyes 1921] S57
       |  |    |--*T. meinerti Wasmann 1894 (see below for synonymy) S57
       |  |    |--T. costaricensis (Seevers 1946) [=Termitosomus costaricensis] S57
       |  |    |--T. fissipennis (Casey 1890) (see below for synonymy) S57
       |  |    |--T. huachii Seevers 1946 S57
       |  |    `--T. manni Seevers 1946 S57
       |  `--+--Trachopeplus Mann 1923 S57
       |     |    |--*T. setosus Mann 1923 S57
       |     |    `--T. disjunctipennis Seevers 1939 S57
       |     `--+--Termitoides Seevers 1939 S57
       |        |    `--*T. marginatus Seevers 1939 S57
       |        `--+--Xenopelta Mann 1923 S57
       |           |    |--*X. cornuta Mann 1923 S57
       |           |    `--X. tricornis (Mann 1923) [=Ceratoxenus tricornis] S57
       |           `--Xenogaster Wasmann 1891 S57
       |                |--*X. inflata Wasmann 1891 S57
       |                |--X. fossulata Mann 1923 S57
       |                |--X. glabriventris Seevers 1957 S57
       |                |--X. inquilina Borgmeier 1950 S57
       |                |--X. nana Seevers 1957 S57
       |                |--X. pilosula Seevers 1957 S57
       |                |--X. reichenspergeri Seevers 1957 S57
       |                `--X. subnuda Seevers 1957 S57
       `--+--+--Termitosynodes Seevers 1957 S57
          |  |    `--*T. williamsi Seevers 1957 S57
          |  `--Termitosyne Seevers 1957 S57
          |       `--*T. platygastra Seevers 1957 S57
          |--+--Neotermitogaster Seevers 1939 S57
          |  |    `--*N. colonus Seevers 1939 S57
          |  `--Termitogaster Casey 1889 S57
          |       |--*T. insolens Casey 1889 S57
          |       |--T. beniensis Seevers 1946 S57
          |       |--T. bicolor Seevers 1937 S57
          |       |--T. brevis Mann 1923 S57
          |       |--T. chavantinae Seevers 1957 S57
          |       |--T. diversicollis Seevers 1957 S57
          |       |--T. emarginatus Seevers 1946 S57
          |       |--T. emersoni Mann 1923 S57
          |       |--T. impressicollis Seevers 1937 S57
          |       |--T. magdalenae Seevers 1946 S57
          |       |--T. nigricollis (Silvestri 1901) [=Xenogaster nigricollis] S57
          |       |--T. puncticeps Seevers 1946 S57
          |       |--T. testaceus Seevers 1946 S57
          |       |--T. wasmanni (Holmgren in Wasmann & Holmgren 1911) [=Xenogaster wasmanni] S57
          |       `--T. wenzeli Seevers 1957 S57
          `--+--Termitotima Wasmann 1916 S57
             |    `--*T. assmuthi Wasmann 1916 S57
             `--+--Idiogaster Wasmann 1912 S57
                |    `--*I. escherichi Wasmann 1912 S57
                `--+--+--Affinoptochus Kemner 1925 S57
                   |  |    `--*A. exclusus Kemner 1925 S57
                   |  `--+--Termitella Wasmann 1911 (see below for synonymy) S57
                   |     |    |--*T. lujae Wasmann 1911 S57
                   |     |    |--T. foveolata (Reichensperger 1922) [=*Termitissa foveolata] S57
                   |     |    |--T. gridelli (Bernhauer 1938) [=*Termitoctesis gridelli] S57
                   |     |    `--T. rubricollis Seevers 1957 S57
                   |     `--+--Termitellodes Seevers 1957 S57
                   |        |    `--*T. lativentris Seevers 1957 S57
                   |        `--Idioptochus Seevers 1957 S57
                   |             `--*I. lehmensicki Seevers 1957 S57
                   `--+--Paracorotoca Warren 1920 S57
                      |    `--*P. akermani (Warren 1914) [=Corotoca akermani] S57
                      |--Millotoca Paulian 1948 S57
                      |    |--*M. mirotermitidis Paulian 1948 S57
                      |    `--M. tanganyuikae Seevers 1957 S57
                      `--Termitoptochus Silvestri 1910 S57
                           |--*T. indicus Silvestri 1910 S57
                           |--T. ceylonicus Silvestri 1921 S57
                           |--T. luzonicus Silvestri 1921 S57
                           |--T. peninsularis Silvestri 1921 S57
                           |--T. philippinus Silvestri 1921 S57
                           `--T. sumatranus Silvestri 1921 S57

Termitella Wasmann 1911 [incl. Termitissa Reichensperger 1912, Termitoctesis Bernhauer 1938] S57

Termitomorpha fissipennis (Casey 1890) [=Termitogaster fissipennis, *Termitosomus fissipennis; incl. Termitogaster simopelta Mann 1923] S57

*Termitomorpha meinerti Wasmann 1894 [incl. *Thaxteria insularis Fenyes 1921, Termitogaster simulans Mann 1923, Termitosomus simulans] S57

*Type species of generic name indicated


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