Terpnacarus gibbosus, photographed by D. E. Walter.

Belongs within: Arachnida.
Contains: Parhypochthonioidea, Enarthronota, Holonota, Palaeacaridae, Ctenacaroidea, Archeonothridae, Alycidae.

The Sarcoptiformes are a major clade of mites including the armoured oribatids, the dust mites and related taxa in the Astigmata, and a small number of plesiomorphic taxa generally grouped as the Endeostigmata (a potentially paraphyletic grouping). Many endeostigmates retain plesiomorphic features of the clade such as one or two pairs of trichobothria on the prodorsum, one or two pairs of simple ocelli, and well-developed rutella on either side of the two-segmented chelicerae (Walter & Proctor 1999). These characters may be modified or lost in derived subgroups. Members of the families Terpnacaridae and Bimichaeliidae also possess transverse folds in the cuticle of the opisthosoma reminiscent of segmentation, a feature that has been mostly lost in mites. The Nematalycoidea, which lack both trichobothria and ocelli, are soil-burrowing mites with an elongate body shape that is most exaggerated in the worm-like Nematalycidae.

The Oribatida, beetle mites, are a major group of mites characterised by a more or less sclerotised body. A number of basal forms retain a segmented notogaster, but in various subgroups the notogaster has become fused into a single sclerite. In the Palaeosomata, the weakly sclerotised hysterosoma bears small, separate sclerites; these mites also have six segments in the legs as opposed to the five-segmented legs of other oribatids (Balogh & Balogh 1992).

    |  i. s.: Protacarus crani Hirst 1923 WP99, S61
    |--Oribatida (see below for synonymy) DW10
    |    |  i. s.: Jureremus foveolatus W91
    |    |--Parhypochthonioidea DW10
    |    `--+--Enarthronota DW10
    |       `--+--Holonota DW10
    |          `--Palaeosomata [Bifemorata, Bifemoratina, Palaeacariformes, Palaecari, Palaeosomatides] DW10
    |               |--Palaeacaridae S04
    |               |--Ctenacaroidea NB-P09
    |               `--Acaronychoidea [Archeonothroidea] NB-P09
    |                    |--Archeonothridae NB-P09
    |                    `--Acaronychidae S04
    |                         |--Amuracarus Lange 1975 S04
    |                         |    `--*A. voskresenskii Lange 1975 S04
    |                         `--Acaronychus Grandjean 1932 S04
    |                              |--*A. traegardhi Grandjean 1932 S04
    |                              |--A. grandjeani Lange 1975 S04
    |                              `--A. proximus Schubart 1968 S04
    `--Endeostigmata DW10
         |  i. s.: Nematalycoidea [Nematolycina] LKW09
         |           |--Proteonematalycus K90 [Proteonematalycidae W01]
         |           |    `--P. wagneri W09
         |           |--Micropsammus Coineau & Théron 1983 CT83 [Micropsammidae W01]
         |           |    `--*M. littoralis Coineau & Théron 1983 CT83
         |           `--Nematalycidae W01
         |                |--Psammolychus delamarei W09
         |                |--Gordialycus tuzetae CT83
         |                `--Nematalycus WL09
         |         Proterorhagia [Proterorhagiidae] W09
         |           `--P. oztotloica W09
         |         Nanorchestidae W09
         |           |--Neonanorchestes McDaniel & Bolen 1981 V96
         |           |--Speleorchestes Trägårdh 1909 V96
         |           |    `--S. poduroides WP99
         |           |--Caenonychus Oudemans 1902 G39
         |           |    `--*C. fallax G39
         |           `--Nanorchestes Grandjean 1937 V96 [incl. Monalichus Berlese 1904 G39]
         |                |--N. africanus Theron & Ryke 1969 V96
         |                |--N. amphibius Tops. & Trouess. 1890 G42
         |                |--N. antarcticus Strandtmann 1963 K91
         |                |--N. arboriger (Berl. 1904) G42
         |                |--N. collinus Hirst. 1918 G42
         |                |--N. hoffmannae Vázquez 1996 V96
         |                |--N. memelensis WP99
         |                |--‘Gainia’ nivalis W01
         |                |--N. pollicaris Theron & Ryke 1969 V96
         |                `--N. pulvinar Grandjean 1942 G42
         |         Grandjeanicus Theron 1974 W01 [Grandjeanicidae W09]
         |           |--*G. uncus Theron 1974 W01
         |           |--G. gabonensis W01
         |           `--G. theroni Walter 2001 W01
         |         Coccorchestes Jacot 1938 G45
         |           `--*C. humicolus Jacot 1938 G45
         |--+--Alycidae DW10
         |  `--Alicorhagiidae [Alicorhagiida, Alichorhagioidea] DW10
         |       |--Archaeacarus dubinini W01, WP99
         |       |--Pseudoprotacarus W01
         |       |--Stigmalychus Theron, Meyer & Ryke 1970 W01
         |       |    `--*S. veretrum Theron, Meyer & Ryke 1970 [=Stigmalichus veretrum] W01
         |       `--Alicorhagia Berlese 1910 W01 (see below for synonymy)
         |            |--*A. fragilis Berlese 1910 W01 (see below for synonymy)
         |            `--A. usitata Theorn, Meyer & Ryke 1970 W01
         `--Terpnacarina [Terpnacarida] LKW09
              |--Oehserchestes Jacot 1939 W01 [Oehserchestidae, Oehserchestoidea LKW09]
              |    |--*O. humicolus Jacot 1939 W01
              |    `--O. arboriger (Theron 1974) [=Hybalicus arboriger] W01
              `--Terpnacaridae [Terpnacaroidea] DW10
                   |--Alycosmesis Grandjean 1939 W01
                   |    |--*A. palmata (Oudemans 1904) [=Sebaia palmata] W01
                   |    `--A. corallium WP99
                   `--Terpnacarus Grandjean 1939 W01
                        |--*T. bouvieri Grandjean 1939 W01
                        |--T. carolinaensis Theron 1976 W01
                        `--T. gibbosus (Womersley 1944) (see below for synonymy) W01

Alicorhagia Berlese 1910 W01 [incl. Epistomalycus Thor 1931 W01, G39, Willania Oudemans 1931 W01, G39]

*Alicorhagia fragilis Berlese 1910 W01 [incl. Willania mira Oudemans 1931 G39, Epistomalycus plumipilus Thor 1931 G39]

Oribatida [Archoribatida, Cryptostigmata, Dichostomata, Macropylina, Monofemorata, Novoribatida, Oribatei, Oribatidae, Oribatini] DW10

Terpnacarus gibbosus (Womersley 1944) [=Hybalicus gibbosus; incl. T. glebulentus Theron 1976, T. variolus Shiba 1968] W01

*Type species of generic name indicated


[CT83] Coineau, Y., & P. Théron. 1983. Les Micropsammidae, n. fam. d'acariens Endeostigmata des sables fins. Acarologia 24 (3): 275–280.

[DW10] Dabert, M., W. Witalinski, A. Kazmierski, Z. Olszanowski & J. Dabert. 2010. Molecular phylogeny of acariform mites (Acari, Arachnida): strong conflict between phylogenetic signal and long-branch attraction artifacts. Molecular Phylogenetics and Evolution 56 (1): 222–241.

[G39] Grandjean, F. 1939. Quelques genres d'acariens appartenant au groupe des Endeostigmata. Annales des Sciences Naturelles, Séries Botanique et Zoologie, 11e Série, 2: 1–122.

[G42] Grandjean, F. 1942. Observations sur les acariens (7e série). Bulletin du Muséum National d’Histoire Naturelle, 2e Série 14 (4): 264–267.

[G45] Grandjean, F. 1945. Observations sur les acariens (8e série). Bulletin du Muséum National d’Histoire Naturelle, 2e Série 17 (5): 399–406.

[K90] Kethley, J. 1990. Acarina: Prostigmata (Actinedida). In: Dindal, D. L. (ed.) Soil Biology Guide pp. 667–756. John Wiley & Sones: New York.

[K91] Kethley, J. 1991. Calyptostatic nymphs of Neonanorchestes (Nanorchestidae): a third example of alternating calyptostasy. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 279–282. SPB Academic Publishing: The Hague.

[LKW09] Lindquist, E. E., G. W. Krantz & D. E. Walter. 2009. Classification. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 97–103. Texas Tech University Press.

[NB-P09] Norton, R. A., & V. M. Behan-Pelletier. 2009. Suborder Oribatida. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 430–564. Texas Tech University Press.

[S61] Southcott, R. V. 1961. Studies on the systematics and biology of the Erythraeoidea (Acarina), with a critical revision of the genera and subfamilies. Australian Journal of Zoology 9: 367–610.

[S04] Subías, L. S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758–2002). Graellsia 60 (número extraordinario): 3–305.

[V96] Vázquez, I. M. 1996. Especie nueva de Nanorchestes (Endeostigmata, Nanorchestidae) de arena de playa en La Mancha, Veracruz, México. Anales Inst. Biol. Univ. Nac. Autón. México, Ser. Zool. 67 (2): 205–211.

[W01] Walter, D. E. 2001. Endemism and cryptogenesis in 'segmented' mites: a review of Australian Alicorhagiidae, Terpnacaridae, Oehserchestidae and Grandjeanicidae (Acari: Sarcoptiformes). Australian Journal of Entomology 40: 207–218.

[W09] Walter, D. E. 2009. Suborder Endeostigmata. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 421–429. Texas Tech University Press.

[WP99] Walter, D. E., & H. C. Proctor. 1999. Mites: Ecology, Evolution and Behaviour. CABI Publishing: Wallingford (UK).

[W91] Woas, S. 1991. Distribution of characters and phylogenetic age—systematic problems in the higher taxa of the Oribatida. In: Schuster, R., & P. W. Murphy (eds) The Acari: Reproduction, development and life-history strategies pp. 329–333. Chapman & Hall: London.

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