Sarcoptiformes

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).

<==Sarcoptiformes
    |  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

REFERENCES

[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.

Cymbaeremaeidae

Dorsal view of Ametroproctus oresbios, from Canadian Biodiversity Information Facility.


Belongs within: Circumdehiscentiae.

The Cymbaeremaeidae are a small but morphologically diverse family of oribatid mites. The genera Ametroproctus and Scapuleremaeus have been treated as a separate family Ametroproctidae due to their possession of well-developed prodorsal lamellae (absent in other cymbaeremaeid genera), but Behan-Pelletier (1988) united the two families on the basis of their similarly apheredermous, plicate nymphs with a characteristic arrangement of porose areas on the body. These porose areas are absent from the genus Scapheremaeus, commonly included in Cymbaeremaeidae but here excluded. The legs of Cymbaeremaeidae are usually tridactyle, but the Fijian Seteremaeus spinosus has monodactylous legs. In Cymbaeremaeus, the notogaster is divided into distinct marginal and central areas though both share the same ornamentation of rough, irregular wrinkles (Balogh & Balogh 1992).

Characters (from Behan-Pelletier 1988, excluding Scapheremaeus): Immatures with plicate integument. Apheredermous nymphs with unideficient setation. Paraproctal atrichosy in larva, protonymph and deutonymph. Seta d present on genua and tibiae of immatures, usually absent from adult (present in some Ametroproctus). Immatures with prodorsal and opisthosomal porose areas or sacculi. Tarsi of immatures with subunguinal bilobed pulvillus. Adult prodorsum with pedotecta I and II, without genal incision, without lamellae. Palpal eupathidium acm not attached to solenidion, inserted on large tubercle. Mentum with tectum. Notogaster with or without lenticulus and humeral projection; without tectal border posteriorly. Pteromorphs absent. Porose areas of octotaxic system absent. Ten or thirteen pairs of notogastral setae, c1 and c3 absent, d series present or absent. All leg femora, tibiae and tarsi generally with sacculi in adult (porose areas present in one species of Ametroproctus). Seta s on tarsus I non-eupathidial.

<==Cymbaeremaeidae [Ametroproctidae, Cymbaeremaeinae, Cymbaeremaeoidea] NB-P09
    |--Jujeremus foveolatus WP99
    |--Seteremaeus Hammer 1971 S04
    |    `--*S. spinosus Hammer 1971 S04
    |--Bulleremaeus Hammer 1966 S04
    |    |--*B. reticulatus Hammer 1966 S04
    |    `--B. tuberculatus Hammer 1966 S04
    |--Cymbaeremaeus Berlese 1896 S04
    |    |--*C. cymba (Nicolet 1855) [=Eremaeus cymba; incl. C. pulvillifer Willmann 1931] S04
    |    |--C. foliatus (Balogh & Csiszár 1963) [=Scutovertex foliatus] S04
    |    `--C. silva Fujikawa 2002 S04
    |--Scapuleremaeus Behan-Pelletier 1989 S04
    |    `--*S. kobauensis Behan-Pelletier 1989 S04
    `--Ametroproctus Higgins & Woolley 1968 NB-P09, S04
         |--A. (Ametroproctus) S04
         |    |--*A. (A.) oresbios Higgins & Woolley 1968 S04
         |    |--A. (A.) aridus Behan-Pelletier 1987 S04
         |    `--A. (A.) tuberculosus Behan-Pelletier 1987 S04
         `--A. (Coropoculia Aoki & Fujikawa 1972) S04
              |--A. (*C.) reticulata (Aoki & Fujikawa 1972) [=*Coropoculia reticulata] S04
              |--A. (C.) beringianus Behan-Pelletier 1987 S04
              |--A. (C.) canningsi Behan-Pelletier 1987 S04
              `--A. (C.) lamellatus (Schweizer 1956) [=Cymbaeremaeus lamellatus] S04

*Type species of generic name indicated

REFERENCES

Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

Behan-Pelletier, V. M. 1988. Systematic relationships of Ametroproctus, with modified definition of Cymbaeremaeidae (Acari: Oribatida). In: Channabasavanna, G. P., & C. A. Viraktamath (eds) Progress in Acarology vol. 1 pp. 301–308. E. J. Brill: Leiden.

[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.

[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.

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

Eremaeidae

Eueremaeus foveolatus, from here.


Belongs within: Circumdehiscentiae.

The Eremaeidae are a group of oribatid mites usually characterised by anal neotrichy, with at least three (usually more) pairs of anal setae. Members of the European genus Tricheremaeus also have a higher number of setae on the notogaster than other species (fifteen to twenty pairs versus ten or eleven), with the setae being particularly long. The genus Proteremaeus is exceptional in only having two pairs of anal setae, together with three pairs of adanal setae (Balogh & Balogh 1992a).

Characters (from Balogh & Balogh 1992a; Norton & Behan-Pelletier 2009): Length 400–800 µm. Prodorsum without true lamellae. Dorsosejugal suture usually arched. Ten to twenty pairs of well-developed notogastral setae; octotaxic system absent. Six pairs of genital setae; one pair of aggenital setae; two to nine pairs of anal setae; three to six pairs of adanal setae. Legs tridactyle; porose areas present distoventrally on tibiae I–IV and proximoventrally on tarsi I–IV.

<==Eremaeidae
    |--Asperemaeus Behan-Pelletier 1982 S04
    |    `--*A. longipilus Behan-Pelletier 1982 S04
    |--Carinabella Hammer 1977 S04
    |    |--*C. puchra Hammer 1977 S04
    |    `--C. tuberculata Bayoumi & Mahunka 1979 S04
    |--Tricheremaeus Berlese 1908 MW07
    |    |--*T. serratus (Michael 1885) [=Notaspis serrata] S04
    |    |--T. abnobensis Miko & Weigmann 2006 MW07
    |    |--T. conspicuus Berlese 1916 MW07
    |    |--T. grandjeani Bernini 1970 MW07
    |    |--T. nemossensis Grandjean 1963 MW07
    |    `--T. travei Miko 1993 MW07
    |--Proteremaeus Piffl 1965 NB-P09, S04 [incl. Sibiremaeus Rjabinin & Krivolutsky 1975 S04]
    |    |--*P. jonasi Piffl 1965 BB92a
    |    |--P. angarensis (Rjabinin & Krivolutsky 1975) [=Sibiremaeus angarensis] S04
    |    |--P. chadaevae Golosova 1983 S04
    |    |--P. elongatus (Rjabinin & Krivolutsky 1975) [=Sibiremaeus elongatus] S04
    |    |--P. lawariensis Hammer 1977 S04
    |    |--P. macleani Behan-Pelletier 1982 S04
    |    |--P. mongolicus (Golosova 1983) [=Sibiremaeus mongolicus] S04
    |    |--P. nebaikini Behan-Pelletier & Rjabinin 1991 S04
    |    `--P. punctulatus Bayartogtokh 2000 S04
    |--Eremaeus Koch 1835 S04
    |    |--*E. hepaticus Koch 1835 (see below for synonymy) S04
    |    |--E. appalachicus Bahen-Pelletier 1993 S04
    |    |--E. boreomontanus Behan-Pelletier 1993 S04
    |    |--E. brevitarsus (Ewing 1917) [=Damaeus brevitarsus] S04
    |    |--E. californiensis Behan-Pelletier 1993 S04
    |    |--E. gracilis Behan-Pelletier 1993 S04
    |    |--E. grandis Hammer 1952 S04
    |    |--E. insertus Grishina 1980 S04
    |    |--E. kananaskis Behan-Pelletier 1993 S04
    |    |--E. kevani Behan-Pelletier 1993 S04
    |    |--E. longiseta Djaparidze 1990 S04
    |    |--E. megistos Behan-Pelletier 1993 S04
    |    |--E. monticolus Behan-Pelletier 1993 S04
    |    |--E. neonominatus Subías 2004 [=E. borealis Wen 1988 non E. oblongus borealis Strenzke 1952] S04
    |    |--E. nortoni Behan-Pelletier 1993 S04
    |    |--E. occidentalis Behan-Pelletier 1993 S04
    |    |--E. oresbios Behan-Pelletier 1993 S04
    |    |--E. orientalis Golosova 1970 S04
    |    |--E. plumosus Woolley 1964 S04
    |    |--E. porosus Behan-Pelletier 1993 S04
    |    |--E. roissi Piffl 1996 S04
    |    |--E. salish Behan-Pelletier 1993 S04
    |    |--E. tenuisetiger Aoki 1970 S04
    |    |--E. translamellatus Hammer 1952 [incl. E. oblongus borealis Strenzke 1952] S04
    |    |--E. tuberosus Gordeeva 1970 S04
    |    `--E. walteri Behan-Pelletier 1993 S04
    `--Eueremaeus Mihelčič 1963 [incl. Kartoeremaeus Higgins 1979] S04
         |--*E. oblongus (Koch 1835) S04 (see below for synonymy)
         |--E. acostulatus Behan-Pelletier 1993 S04
         |--E. alvordensis Behan-Pelletier 1993 S04
         |--E. aridulus Behan-Pelletier 1993 S04
         |--E. aysineep Behan-Pelletier 1993 S04
         |--E. brevifer (Mahunka 1980) [=Eremaeus brevifer] S04
         |--E. carinatus Behan-Pelletier 1993 S04
         |--E. chiatous (Higgins 1979) [=Kartoeremaeus chiatous] S04
         |--E. columbianus (Berlese 1916) (see below for synonymy) S04
         |--E. cornutus (Lombardini 1943) (n. d.) [=Eremaeus cornutus] S04
         |--E. danos Behan-Pelletier 1993 S04
         |--E. elongatus (Fujikawa 1972) [=Eremaeus elongatus] S04
         |--E. foveolatus (Hammer 1952) [=Eremaeus foveolatus] S04
         |--E. higginsi Behan-Pelletier 1993 S04
         |--E. hokkaiensis Fujikawa 1991 S04
         |--E. lindquisti Behan-Pelletier 1993 S04
         |--E. magniporus (Wallwork 1972) [=Eremaeus magniporus] S04
         |--E. marshalli Behan-Pelletier 1993 S04
         |--E. masinasin Behan-Pelletier 1993 S04
         |--E. michaeli Behan-Pelletieri 1993 S04
         |--E. nahani Behan-Pelletier 1993 S04
         |--E. nemoralis Behan-Pelletier 1993 S04
         |--E. osoyoosensis Behan-Pelletier 1993 S04
         |--E. proximus (Berlese 1916) [=Eremaeus proximus] S04
         |--E. quadrilamellatus (Hammer 1952) (see below for synonymy) S04
         |--E. stiktos (Higgins 1962) [=Eremaeus stiktos] S04
         |--E. tetrosus (Higgins 1979) [=Kartoeremaeus tetrosus] S04
         |--E. travei Mihelčič 1963 [incl. Eremaeus oblongus major Mihelčič 1957 non E. major Mihelčič 1953] S04
         |--E. triglavensis (Tarman 1958) [=Eremaeus triglavensis] S04
         |--E. trionus (Higgins 1979) [=Kartoeremaeus trionus] S04
         |--E. woolleyi (Higgins 1979) [=Kartoeremaeus woolleyi] S04
         `--E. yukonensis Behan-Pelletieri 1993 S04

*Eremaeus hepaticus Koch 1835 [incl. E. hepaticus acruciata Mihelčič 1952, E. cordiformis Grandjean 1934, Notaspis dentilamellatus Storkán 1925, E. magnus Mihelčič 1957, E. ovalis Mihelčič 1955 non Koch 1841, E. setiger Mihelčič 1957] S04

Eueremaeus columbianus (Berlese 1916) [=Eremaeus columbianus; incl. Er. politus Banks 1947, Kartoeremaeus reevesi Higgins 1979] S04

*Eueremaeus oblongus (Koch 1835) S04 [=Eremaeus oblongus S04; incl. Er. areolatus Kunst 1959 S04, Er. figuratus Winkler 1956 S04, Er. granulatus Mihelčič 1955 S04, Eueremaeus granulatus BB92b, Er. intermedius Mihelčič 1955 S04, Eueremaeus kuehnelti Mihelčič 1963 S04, Er. major Mihelčič 1953 S04, Er. ornatus Mihelčič 1957 non Oudemans 1900 S04, Er. silvestris Forsslund 1956 S04, Er. valkanovi Kunst 1957 S04]

Eueremaeus quadrilamellatus (Hammer 1952) [=Eremaeus quadrilamellatus; incl. Er. fossulatus Kunst 1959, Eueremaeus valkanovi debilis Mihelčič 1963] S04

*Type species of generic name indicated

REFERENCES

[BB92a] Balogh, J., & P. Balogh. 1992a. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

[BB92b] Balogh, J., & P. Balogh. 1992b. The Oribatid Mites Genera of the World vol. 2. Plates. Hungarian Natural History Museum: Budapest.

[MW07] Miko, L., & G. Weigmann. 2007. Tricheremaeus abnobensis Miko & Weigmann 2006, a recently described oribatid mite from central Europe (Arachnida, Acarina, Oribatida, Eremaeidae). Senckenbergiana Biologica 87 (2): 131–134.

[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.

[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.

Hermanniellidae

Hermanniella sp., photographed by Pfliegler Walter.


Belongs within: Circumdehiscentiae.

The Hermanniellidae are a group of oribatid mites in which the tritonymphal scalp adheres very closely to the adult. In genera other than Hermanniella and Mahunkobates, the bothridia have become closely placed to each other and more or less removed from the prodorsal margin. The Venezuelan Mahunkobates singularis has the posteromarginal setae on the notogaster spiniform and ciliate, in contrast to the setiform and smooth setae on the remainder of the notogaster, and the more uniform notogastral setae in most species of Hermanniella (Balogh & Balogh 1992).

Characters (from Balogh & Balogh 1992): Prodorsum without true lamellae; lamellar setae well developed. Hysterosoma with lateral tube. Notogaster without concentrically arranged scalps, but tritonymphal scalp adheres closely to notogaster and bears the most conspicuous notogastral setae; 4-14 pairs of well developed setae, the remainder extremely small.

<==Hermanniellidae [Opsiopheredermata]
    |--Mahunkobates Calugar 1989 S04
    |    `--*M. singularis Calugar 1989 S04
    |--Ampullobates Grandjean 1962 S04
    |    `--*A. nigriclavatus Grandjean 1962 S04
    |--Akansilvanus Fujikawa 1993 S04
    |    `--*A. parvus Fujikawa 1993 S04
    |--Sacculobates Grandjean 1962 [incl. Bruneiella Mahunka 1997] S04
    |    |--*S. horologiorum Grandjean 1962 S04
    |    `--S. sultan (Mahunka 1997) [=Bruneiella sultan] S04
    |--Dicastribates Balogh & Balogh 1988 S04
    |    |--*D. heterotrichus (Mahunka 1984) [=Sacculobates heterotrichus] S04
    |    `--D. tenuisetosus Choi, Bayartogtokh & Aoki 2001 S04
    |--Baloghacarus Mahunka 1983 S04
    |    |--*B. hauseri Mahunka 1983 S04
    |    `--B. australis Balogh & Mahunka 1981 S04
    |--Issaniella Grandjean 1962 S04
    |    `--*I. mograbin Grandjean 1962 S04
    |         |--I. m. mograbin S04
    |         `--I. m. hauseri Mahunka 2001 S04
    |--Hermannobates Hammer 1961 S04
    |    |--*H. monstruosus Hammer 1961 S04
    |    |--H. cubanus Starý 1998 S04
    |    |--H. flagelliseta Balogh & Mahunka 1981 S04
    |    |--H. hammerae Starý 1998 S04
    |    |--H. horridus Pérez-Íñigo & Baggio 1988 S04
    |    |--H. intermedius Calugar 1990 S04
    |    |--H. ruseki Starý 1998 S04
    |    `--H. scoparius Pérez-Íñigo & Baggio 1993 S04
    `--Hermanniella Berlese 1908 S04
         |--*H. granulata (Nicolet 1855) [=Hermannia granulata] S04
         |--H. aristosa Aoki 1965 S04
         |--H. barbata Moraza 1990 S04
         |--H. canestrinii (Tietze 1899) [=Hypochthonius canestrinii] S04
         |--H. clavasetosa Hammer 1966 S04
         |--H. clavigera Berlese 1908 S04
         |--H. congoensis Balogh 1958 S04
         |--H. diversisetosa Hammer 1966 S04
         |--H. dolosa Grandjean 1931 S04
         |--H. dubinini Sitnikova 1973 S04
         |--H. gibber Kulijev 1979 S04
         |--H. grandis Sitnikova 1973 S04
         |--H. humilis Balogh 1962 S04
         |--H. incondita Pérez-Íñigo 1987 S04
         |--H. issanielloides Gil-Martín & Subías 1997 S04
         |--H. laurisilvae Pérez-Íñigo 1972 S04
         |--H. longisetosa Hammer 1966 S04
         |--H. madagascarensis Balogh 1962 S04
         |--H. mastys Mahunka 1983 S04
         |--H. microsetosa Hammer 1966 S04
         |--H. multipora Sitnikova 1973 S04
         |--H. oblitera Sitnikova 1973 S04
         |--H. occidentalis Ewing 1918 [=H. punctulata occidentalis] S04
         |--H. orbiculata Hammer 1979 S04
         |--H. picea (Koch 1839) [=Nothrus piceus; incl. Hermannia arrecta Nicolet 1855] S04
         |--H. punctulata Berlese 1908 SK10, S04
         |    |--H. p. punctulata S59
         |    `--H. p. columbiana Sellnick 1959 S59
         |--H. reticulata Sitnikova 1973 S04
         |--H. robusta Ewing 1918 [=H. punctulata robusta] S04
         |--H. septentrionalis Berlese 1910 [=H. punctulata septentrionalis] S04
         |--H. serrata Sitnikova 1973 S04
         |--H. similis Sitnikova 1973 S04
         |--H. subnigra (Ewing 1909) [=Hermannia subnigra] S04
         |    |--H. s. subnigra [incl. H. punctulata columbiana Berlese 1910] S04
         |    |--H. s. rigida Jacot 1938 S04
         |    `--H. s. setacea Jacot 1937 S04
         |--H. thani Mahunka 1987 S04
         `--H. yasumai Aoki 1973 S04

*Type species of generic name indicated

REFERENCES

Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

[SK10] Schäffer, S., S. Koblmüller, T. Pfingstl, C. Sturmbauer & G. Krisper. 2010. Ancestral state reconstruction reveals multiple independent evolution of diagnostic morphological characters in the "Higher Oribatida" (Acari), conflicting with current classification. BMC Evolutionary Biology 10: 246.

[S59] Sellnick, M. 1959. Acarina from southeastern Polynesia—II (Oribatidae). Occasional Papers of the Bernice P. Bishop Museum 22 (9): 109–152.

[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.

Eutegaeidae

Eutegaeus bostocki, photographed by Stephen Thorpe.


The Eutegaeidae are a family of oribatid bites characterised by the presence of long humeral processes on the notogaster, together with a thickened bothridium. In the genera Dudichella, Porrhotegaeus and Pterozetes, these processes are ventrally reflexed, and these genera have also lost the interlamellar setae, leading some authors to separate them as the Pterozetidae. Among other eutegaeids, the New Guinean Birotegaeus biroi is distinguished by the presence of nine pairs of setae on the notogaster (vs eight pairs otherwise) which also possesses a single small knob on the posterior edge (Balogh & Balogh 1992).

Characters (from Balogh & Balogh 1992): Prodorsum with true lamellae. Bothridium long, tubular. Notogaster with long horizontal humeral process, at least as long as half the length of prodorsum. Legs monodactyle.

Belongs within: Circumdehiscentiae.

<==Eutegaeidae [Pterozetidae]
    |--Atalotegaeus Luxton 1988 S04
    |    `--*A. mensarosi (Balogh & Balogh 1983) [=Eutegaeus mensarosi] S04
    |--Birotegaeus Luxton 1988 S04
    |    `--*B. biroi (Balogh 1970) [=Eutegaeus biroi] S04
    |--Dudichella Balogh 1970 S04
    |    `--*D. membranigera Balogh 1970 S04
    |--Pterozetes Hammer 1966 S04
    |    `--*P. novazealandicus Hammer 1966 S04
    |--Porrhotegaeus Balogh & Mahunka 1966 S04
    |    |--*P. ornatus Balogh & Mahunka 1966 S04
    |    `--P. herminae Balogh & Balogh 1983 S04
    |--Pareutegaeus Woolley 1965 S04
    |    |--*P. similis (Trägardh 1931) [=Eutegaeus similis] S04
    |    `--P. pulcher (Balogh & Csiszár 1963) [=Eutegaeus pulcher] S04
    |--Neoeutegaeus Aoki 1965 S04
    |    |--*N. silvicola (Hammer 1962) [=Eutegaeus silvicola] S04
    |    |--N. africanus Mahunka 1974 S04
    |    `--N. phyllophorus Balogh & Balogh 1983 S04
    |--Neseutegaeus Woolley 1965 S04
    |    |--*N. spinatus Woolley 1965 S04
    |    |--N. angustus Hammer 1966 S04
    |    |--N. consimilis Hammer 1966 S04
    |    |--N. denticulatus Balogh 1988 S04
    |    |--N. distentus Hammer 1966 S04
    |    |--N. latus Hammer 1966 S04
    |    `--N. monteithi Balogh & Balogh 1983 S04
    `--Eutegaeus Berlese 1916 S04
         |--*E. bostocki (Michael 1908) [=Oribata bostocki] S04
         |--E. biovatus Hammer 1972 S04
         |--E. curviseta Hammer 1966 DA07
         |--E. fueginus Arcidiacono 1993 S04
         |--E. lagrecai Arcidiacono 1993 S04
         |--E. membraniger Hammer 1966 S04
         |--E. papuensis Aoki 1965 S04
         |--E. pinnatus Hammer 1966 S04
         |--E. radiatus Hammer 1966 S04
         |--E. soror Balogh 1985 S04
         `--E. stylesi Hammer 1966 S04

*Type species of generic name indicated

REFERENCES

Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

[DA07] Domes, K., M. Althammer, R. A. Norton, S. Scheu & M. Maraun. 2007. The phylogenetic relationship between Astigmata and Oribatida (Acari) as indicated by molecular markers. Experimental and Applied Acarology 42 (3): 159–171.

[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.

Cepheidae

Cepheus corae, from here.


Belongs within: Circumdehiscentiae.

The Cepheidae are a family of often relatively large oribatid mites, usually with a sculptured integument. Species of Cepheus can be found in moss, litter and rotting wood (Norton & Behan-Pelletier 2009). The genera Ommatocepheus, Eupterotegaeus, Conoppia, Sphodrocepheus and Tritegeus have tridactylous legs; the remaining genera are monodactyle (Balogh & Balogh 1992).

Characters (from Norton & Behan-Pelletier 2009): Length 500–1200 µm. Integument usually sculptured. Subcapitulum diarthritic; chelicerae chelate-dentate; subcapitular mentum without tectum. Prodorsum with true lamellae. Scalps absent. Lenticulus absent. Notogaster without setae centrodorsally, all notogastral setae in marginal or posteromarginal position. Three pairs of adanal setae, positioned adanally and postanally. Tarsus I with two solenidia.

<==Cepheidae
    |--Protocepheus Jacot 1928 S04
    |    `--*P. hericius (Michael 1887) [=Tegeocranus hericius] S04
    |--Tikizetes Hammer 1967 S04
    |    `--*T. spinipes Hammer 1967 S04
    |--Pilocepheus Pérez-Íñigo 1992 S04
    |    `--*P. azoricus Pérez-Íñigo 1992 S04
    |--Conoppia Berlese 1908 [incl. Balzania Jacot 1929 non Spegazzini 1898 (ICBN), Phyllotegeus Berlese 1913] S04
    |    |--C. palmicincta (Michael 1884) S04 (see below for synonymy)
    |    `--C. setiformis Golosova & Karppinen 1985 S04
    |--Oribatodes Banks 1895 S04
    |    |--*O. mirabilis Banks 1895 [incl. Cepheus lamellatus Banks 1906, Tegeocranus longisetus Berlese 1910] S04
    |    `--O. crenulatus Csiszár 1962 S04
    |--Tereticepheus Bernini & Bernini 1990 S04
    |    |--*T. annamariae Bernini & Bernini 1990 S04
    |    `--T. undulatus (Willmann 1939) [=Cepheus undulatus] S04
    |--Hypocepheus Krivolutsky 1971 S04
    |    |--*H. mirabilis Krivolusky 1971 S04
    |    |--H. helveticus Mahunka & Mahunka-Papp 2002 S04
    |    `--H. krivolutskyi Calugar & Vasiliu 1976 S04
    |--Reticulocepheus Vasiliu & Calugar 1977 [incl. Cubacepheus Balogh & Mahunka 1979, Geocepheus Sitnikova 1979] S04
    |    |--*R. decoui Vasiliu & Calugar 1977 (see below for synonymy) S04
    |    |--R. areolatus (Sitnikova 1979) [=Geocepheus areolatus] S04
    |    |--R. grandis (Sitnikova 1979) [=Geocepheus grandis] S04
    |    `--R. reticulatus (Sitnikova 1979) [=Geocepheus reticulatus] S04
    |--Tritegeus Berlese 1913 S04
    |    |--T. bisulcatus Grandjean 1953 [incl. Cepheus bifidatus Michael 1880 non Nicolet 1855, *T. bifidatus] S04
    |    |--T. brevisetus Sitnikova 1975 S04
    |    |--T. major Golosova & Karppinen 1984 S04
    |    `--T. sculptus Bernini & Bernini 1990 S04
    |--Sphodrocepheus Woolley & Higgins 1963 S04
    |    |--*S. tridactylus Woolley & Higgins 1963 S04
    |    |--S. anthelionus Woolley & Higgins 1968 S04
    |    |--S. dentatus Fujikawa 1972 S04
    |    |--S. mitratus Aoki 1967 S04
    |    `--S. tuberculatus Mahunka 1988 S04
    |--Compactozetes Hammer 1966 [Compactozetidae] S04
    |    |--*C. rotoruensis Hammer 1966 S04
    |    |--C. hastatus Hammer 1973 S04
    |    |--C. niger Hammer 1966 S04
    |    |--C. serratus Balogh 1970 S04
    |    `--C. zeugus Luxton 1988 S04
    |--Ommatocepheus Berlese 1913 S04
    |    |--*O. ocellatus (Michael 1882) [=Cepheus ocellatus; incl. Scutovertex pulcherrimus Berlese 1913] S04
    |    |--O. clavatus Woolley & Higgins 1964 S04
    |    |    |--O. c. clavatus S04
    |    |    `--O. c. japonicus Aoki 1974 S04
    |    |--O. crassisetosus Pérez-Íñigo & Peña 1996 S04
    |    |--O. parvilamellatus Pérez-Íñigo & Pérez-Íñigo 1996 S04
    |    `--O. reticulatus Pérez-Íñigo & Peña 1997 S04
    |--Eupterotegaeus Berlese 1916 [incl. Diodontocepheus Mihelčič 1958] S04
    |    |--*E. ornatissimus (Berlese 1908) (see below for synonymy) S04
    |    |--E. armatus Aoki 1969 S04
    |    |--E. dentatus Sitnikova 1979 S04
    |    |--E. flavus (Ewing 1918) [=Cepheus flavus] S04
    |    |--E. nasalis Sitnikova 1979 S04
    |    |--E. pseudosculptus (Coggi 1900) [=Cepheus pseudosculptus] S04
    |    |--E. rhamphosus Higgins & Woolley 1968 S04
    |    |--E. rostratus Higgins & Woolley 1963 S04
    |    |--E. spinatus Higgins & Woolley 1963 S04
    |    `--E. xizangensis Wang & Solhoy 2001 S04
    |--Sadocepheus Aoki 1965 [incl. Hamotegeus Balogh & Mahunka 1969] S04
    |    |--*S. undulatus Aoki 1965 S04
    |    |    |--S. u. undulatus S04
    |    |    `--S. u. setiger Fujita & Fujikawa 1986 S04
    |    |--S. breviseta (Balogh 1986) [=Hamotegeus breviseta] S04
    |    |--S. dubius Hammer 1979 S04
    |    |--S. elevatus Mahunka 1987 S04
    |    |--S. foveolatus Luxton 1988 S04
    |    |--S. franzi (Balogh 1986) [=Hamotegeus franzi] S04
    |    |--S. granulatus (Balogh & Mahunka 1969) S04 [=*Hamotegeus granulatus BB92]
    |    |--S. longiseta (Balogh 1986) [=Hamotegeus longiseta] S04
    |    |--S. makarcevae Sitnikova 1975 S04
    |    `--S. subniger (Ewing 1917) [=Tegeocranus subniger] S04
    `--Cepheus Koch 1835 [incl. Tegeocranus Nicolet 1855] S04
         |--*C. latus Koch 1835 S04
         |--C. brachiatus Sitnikova 1975 S04
         |--C. caucasicus Sitnikova 1975 S04
         |--C. cepheiformis (Nicolet 1855) [=Tegeocranus cepheiformis; incl. C. michaeli Jacot 1928] S04
         |--C. corae Jacot 1928 S04
         |--C. dentatus (Michael 1888) [=Tegeocranus dentatus; incl. C. grandis Sitnikova 1975] S04
         |--C. feideri Suciu & Panu 1972 S04
         |--C. gracilipes (Canestrini 1897) (n. d.) [=Tegeocranus gracilipes] S04
         |--C. granulatus Mihelčič 1958 S04
         |--C. heterosetosus (Sitnikova 1975) [=Oribatodes heterosetosus] S04
         |--C. hokkaiensis Fujikawa 1992 S04
         |--C. incisus Mihelčič 1958 S04
         |--C. jindingensis Wen & Sun 1990 S04
         |--C. kurosawai Aoki 1986 S04
         |--‘Tegeocranus’ latus H99
         |--C. lobatus Mihelčič 1953 (n. d.) S04
         |--C. mirabiloides Jacot 1938 S04
         |--C. ovalis CF77
         |--C. pegazzanoae Bernini & Nannelli 1982 S04
         |--C. pertusus Haupt 1882 (n. d.) S04
         |--C. pustulatus (Pearce 1910) [=Tegeocranus pustulatus] S04
         |--C. takasago Aoki 1991 S04
         |--C. transversalis Balogh 1958 (n. d.) S04
         |--C. transylvanicus Cooreman 1951 S04
         |--C. tuberculosus Strenzke 1951 S04
         `--C. verrucosus Bernini 1971 S04

Conoppia palmicincta (Michael 1884) S04 [=Leiosoma palmicinctum S04, Phyllotegeus palmicinctus G46; incl. Oppia microptera Berlese 1885 S04, *Conoppia microptera S04, Oribata microptera Canestrini 1896 non Oppia microptera Berlese 1885 S04]

*Eupterotegaeus ornatissimus (Berlese 1908) [=Tegeocranus ornatissimus; incl. Diodontocepheus steinboecki Mihelčič 1958] S04

*Reticulocepheus decoui Vasiliu & Calugar 1977 [incl. Cepheus (Cubacepheus) lobatus Balogh & Mahunka 1979 non C. lobatus Mihelčič 1953] S04

*Type species of generic name indicated

REFERENCES

[BB92] Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

[CF77] Canestrini, G., & F. Fanzago. 1877. Intorno agli Acari Italiani. Atti del Reale Istituto Veneto di Scienze, Lettere ed Arti, Serie 5, 4: 69–208, pls 202–207.

[G46] Grandjean, F. 1946. Les poils et les organes sensitifs portés par les pattes et le palpe chez les oribates. Bulletin de la Société Zoologique de France 71: 10–29.

[H99] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

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.

[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.

Ameronothridae

Possible cluster of Ameronothrus, photographed by D. Sikes.


Belongs within: Circumdehiscentiae.

The Ameronothridae are a family of mostly semi-aquatic oribatid mites, inhabiting ephemeral rock pools or intertidal littoral zones. Some species are noteworthy for being found in extreme environments in the Arctic or Antarctic (Norton & Behan-Pelletier 2009). Members of the genera Antarcticola and Pseudantarcticola have fewer setae on the notogaster than other genera, with only ten or eleven pairs versus the twelve or more pairs of the latter (Balogh & Balogh 1992).

Characters (from Norton & Behan-Pelletier 2009): Length 550–800 µm. Gnathosoma normal, chelicerae with dentate chelae and two setae; rutella normally developed. Prodorsum without true lamellae. Bothridium developed, bothridial seta reduced. Scalps absent. Lyrifissure iad positioned anterior to ad3. Lateral plastron absent. Genital plates with five pairs of setae. Epimera III–IV distinctly delineated. Leg tibiae and tarsi completely separate. Tarsus II with two solenidia.

<==Ameronothridae
    |--Aquanothrus Engelbrecht 1975 [Aquanothridae] S04
    |    `--*A. montanus Engelbrecht 1975 S04
    |--Capillibates Hammer 1966 S04
    |    `--*C. stagaardi Hammer 1966 S04
    |--Chudalupia Wallwork 1981 S04
    |    `--*C. meridionalis Wallwork 1981 S04
    |--Pseudantarcticola Balogh 1970 S04
    |    `--*P. tropica Balogh 1970 S04
    |--Antarcticola Wallwork 1967 [incl. Petrozetes Sitnikova 1968] S04
    |    |--*A. meyeri Wallwork 1967 [incl. Petrozetes oblongus Sitnikova 1968] S04
    |    `--A. georgiae Wallwork 1970 S04
    |--Alaskozetes Hammer 1955 S04
    |    |--*A. coriaceus Hammer 1955 S04
    |    |--A. antarcticus (Michael 1903) S04 [=Notaspis antarctica S04, Halozetes antarctica CH98]
    |    |    |--A. a. antarcticus S04
    |    |    |--A. a. grandjeani (Dalenius 1958) S04 [=Halozetes antarctica grandjeani CH98]
    |    |    `--A. a. intermedius Wallork 1967 S04
    |    `--A. bouvetoyaensis Pletzen & Kok 1971 S04
    |--Ameronothrus Berlese 1896 [incl. Hygroribates Jacot 1934] S04
    |    |--*A. lineatus (Thorell 1871) (see below for synonymy) S04
    |    |--A. bilineatus (Michael 1888) [=Scutovertex bilineatus] S04
    |    |--A. dubinini Sitnikova 1975 S04
    |    |--A. harioti (Michael 1891) (n. d.) [=Scutovertex harioti] S04
    |    |--A. lapponicus Dalenius 1963 S04
    |    |--A. maculatus (Michael 1882) (see below for synonymy) S04
    |    |--A. marinus (Banks 1896) (see below for synonymy) S04
    |    |--A. nidicola Sitnikova 1975 S04
    |    |--A. nigrofemoratus (Koch 1879) [=Nothrus nigrofemoratus; incl. A. lineatus brevipes Willmann 1937] S04
    |    |--A. oblongus Sitnikova 1975 S04
    |    |--A. schneideri (Oudemans 1905) [=Scutovertex schneideri] S04
    |    |--A. schubarti Weigmann & Schulte 1975 S04
    |    `--A. schusteri Schubart 1970 S04
    `--Halozetes Berlese 1916 [=Anarea Dalenius 1958; incl. Pertorgunia Dalenius 1958] S04
         |--*H. marinus (Lohmann 1907) [=Notaspis marina, *Anarea marinus] CH98
         |    |--H. m. marinus [incl. Anarea macquariensis Dalenius 1958] S04
         |    |--H. m. devilliersi Engelbrecht 1974 S04
         |    `--H. m. minor Wallwork 1966 S04
         |--H. bathamae Luxton 1984 S04
         |--H. belgicae (Michael 1903) S04 [=Notaspis belgicae S04, Pertorgunia belgicae CH98]
         |    |--H. b. belgicae S04
         |    |--H. b. brevipilis Wallwork 1963 S04
         |    |--H. b. longisetae Wallwork 1967 S04
         |    `--H. b. mickii Coetzee 1999 S04
         |--H. crozetensis (Richters 1908) S04 (see below for synonymy)
         |--H. edwardensis Pletzen & Kok 1971 S04
         |--H. fulvus Engelbrecht 1975 S04
         |--H. impeditus Niedbala 1986 S04
         |--H. intermedius Wallwork 1963 S04
         |--H. littoralis Wallwork 1970 S04
         |--H. macquariensis (Dalenius 1958) [=Pertorgunia macquariensis] S04
         |--H. marionensis Engelbrecht 1974 S04
         |--H. necrophagus Wallwork 1967 S04
         |--H. otagoensis Hammer 1966 S04
         |--H. plumosus Wallwork 1966 S04
         `--H. scotiae (Trouessart 1912) (n. d.) [=Notaspis scotiae] S04

*Ameronothrus lineatus (Thorell 1871) [=Eremaeus lineatus; incl. Scutovertex corrugatus Michael 1888, S. occidentalis Hull 1918] S04

Ameronothrus maculatus (Michael 1882) [=Scutovertex maculatus; incl. S. maculatus groenlandicus Trägardh 1904, S. pseudomaculatus insularis Hull 1914, S. pseudomaculatus Hull 1914] S04

Ameronothrus marinus (Banks 1896) [=Nothrus marinus; incl. A. fucicolum Brady 1875, Scutovertex spoofi Oudemans 1900] S04

Halozetes crozetensis (Richters 1908) S04 [=Scutovertex crozetensis S04, *Pertorgunia crozetensis CH98; incl. P. colobanthi Dalenius 1958 S04]

*Type species of generic name indicated

REFERENCES

Balogh, J. & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

[CH98] Colloff, M. J., & R. B. Halliday. 1998. Oribatid Mites: A catalogue of Australian genera and species. CSIRO Publishing: Collingwood.

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.

[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.

Liacaridae

Dorycranosus acutidens, photographed by David Porco.


Belongs within: Circumdehiscentiae.
Contains: Liacarus (Liacarus).

The Liacaridae are a family of oribatid mites with converging lamellae, and two pairs of humeral setae c1 and c2 placed near to each other. The lamellae remain well-separated in Planoristes acuspidatus but are fused apically in the remaining genera. Birsteinius has monodactyle legs whereas the legs of Opsioristes, Liacarus and Dorycranosus are tridactyle. Opsioristes eurhostus has five pairs of genital setae vs six in Liacarus and Dorycranosus (Balogh & Balogh 1992).

<==Liacaridae
    |--Opsioristes Woolley 1967 S04
    |    `--*O. eurhostus Woolley 1967 S04
    |--Planoristes Iturrondobeitia & Subías 1978 S04
    |    `--*P. acuspidatus Iturrondobeitia & Subías 1978 S04
    |--Birsteinius Krivolutsky 1965 S04
    |    |--*B. clavatus Krivolutsky 1965 S04
    |    |--B. krivolutskyi Rjabinin 1979 S04
    |    |--B. microchaetus Krivolutsky 1967 S04
    |    |--B. mongolicus (Mahunka 1964) [=Liacarus mongolicus] S04
    |    |--B. monticolus Wang & Shen 2001 S04
    |    |--B. perlongoides Hammer 1977 S04
    |    |--B. perlongus Krivolutsky 1965 S04
    |    `--B. trimucronatus Yamamoto & Aoki 2000 S04
    |--Liacarus Michael 1898 S04
    |    |  i. s.: L. globosus S22
    |    |--L. (Liacarus) S04
    |    |--L. (Procorynetes Woolley 1969) S04
    |    |    |--L. (*P.) nigerrimus Berlese 1916 S04
    |    |    |--L. (P.) andinus (Balogh 1984) [=Procorynetes andinus] S04
    |    |    |--L. (P.) breviclavatus Aoki 1970 S04
    |    |    |--L. (P.) clavatus Fujikawa & Aoki 1970 S04
    |    |    |--L. (P.) espeletiae (Balogh 1984) [=Procorynetes espeletiae] S04
    |    |    |--L. (P.) globifer (Kramer 1897) [=Leiosoma globiferum] S04
    |    |    `--L. (P.) tsendsureni (Bayartogtokh 1998) [=Procorynetes tsendsureni] S04
    |    `--L. (Rhaphidosus Woolley 1969 S04
    |         |--L. (*R.) carolinensis Banks 1906 S04
    |         |--L. (R.) acuminatus (Woolley 1969) [=Rhaphidosus acuminatus] S04
    |         |--L. (R.) alticola (Balogh 1984) [=Rhaphidosus alticola] S04
    |         |--L. (R.) bacillatus Fujikawa & Aoki 1970 S04
    |         `--L. (R.) carbonarius (Banks 1906) [=Notaspis carbonaria] S04
    `--Dorycranosus Woolley 1969 NB-P09, S04
         |--*D. abdominalis (Banks 1906) [=Liacarus abdominalis] S04
         |--D. acutidens (Aoki 1965) [=Liacarus (Dorycranosus) acutidens] S04
         |--D. acutus (Pschorn-Walcher 1951) (see below for synonymy) S04
         |--D. aequidentatus (Ewing 1918) [=Liacarus (Dorycranosus) aequidentatus] S04
         |--D. altaicus Krivolutsky 1974 [=Liacarus (Dorycranosus) altaicus] S04
         |--D. angustatus Weigmann 1976 [=Liacarus (Dorycranosus) angustatus] S04
         |--D. arcticus (Banks 1899) (n. d.) [=Oppia arctica, Liacarus (Dorycranosus) arcticus] S04
         |--D. badghysi (Krivolutsky 1966) [=Liacarus (Dorycranosus) badghysi] S04
         |--D. columbianus (Berlese 1908) [=Liacarus (Dorycranosus) columbianus] S04
         |--D. coronatus Golosova & Ljashchev 1984 [=Liacarus (Dorycranosus) coronatus] S04
         |--D. dickersoni Moraza 1990 [=Liacarus (Dorycranosus) dickersoni] S04
         |--D. floridensis (Berlese 1908) [=Liacarus (Dorycranosus) floridensis] S04
         |--D. frontinalis (Banks 1906) [=Liacarus (Dorycranosus) frontinalis] S04
         |--D. fusiformis (Ewing 1917) [=Liacarus (Dorycranosus) fusiformis] S04
         |--D. indentatus Aoki 1973 [=Liacarus (Dorycranosus) indentatus] S04
         |--D. keretinus (Nordenskiöld 1901) [=Liacarus (Dorycranosus) keretinus] S04
         |--D. latiusculus (Wen 1991) [=Liacarus (Dorycranosus) latiusculus] S04
         |--D. lingulatus Golosova & Ljashchev 1984 [=Liacarus (Dorycranosus) lingulatus] S04
         |--D. neonominatus (Subías 2004) (see below for synonymy) S04
         |--D. nitidus (Banks 1895) (see below for synonymy) S04
         |--D. paoliensis Woolley 1969 [=Liacarus (Dorycranosus) paoliensis] S04
         |--D. parallelus (Hammer 1967) [=Liacarus (Dorycranosus) parallelus] S04
         |--D. polychothomus (Wen 1991) [=Liacarus (Dorycranosus) polychothomus] S04
         |--D. pulcher Sergienko & Djaparidze 1981 [=Liacarus (Dorycranosus) pulcher] S04
         |--D. splendens (Coggi 1898) (see below for synonymy) S04
         |--D. tenuilamellatus (Hirauchi 1998) [=Liacarus (Dorycranosus) tenuilamellatus] S04
         |--D. triapicini Djaparidze 1983 [=Liacarus (Dorycranosus) triapicini] S04
         |--D. yezoensis (Fujikawa & Aoki 1970) [=Liacarus (Dorycranosus) yezoensis] S04
         `--D. zachvatkini (Kulijev 1962) [=Liacarus (Dorycranosus) zachvatkini; incl. D. ibericus Djaparidze 1973] S04

Dorycranosus acutus (Pschorn-Walcher 1951) [=Liacarus (Dorycranosus) acutus; incl. L. claviger Mihelčič 1956, Cultroribula grandis Mihelčič 1956, L. infissus Gunhold 1953] S04

Dorycranosus neonominatus (Subías 2004) [=Liacarus (Dorycranosus) neonominatus, Dorycranosus arcticus Grishina 1984 non Oppia arctica Banks 1899] S04

Dorycranosus nitidus (Banks 1895) [=Cepheus nitidus, Liacarus (Dorycranosus) nitidus; incl. L. nitidus fissuratus Jacot 1937, L. glaber Ewing 1909] S04

Dorycranosus splendens
(Coggi 1898) [=Cepheus splendens, Liacarus (Dorycranosus) splendens; incl. L. alatus Berlese 1904, L. alatus curtipilis Willmann 1935, L. diversus Mihelčič 1956, L. ghilarovi Bashkirova 1958, L. moraviacus Willmann 1954, Dorycranosus ovatus Djaparidze 1973, L. punctulatus Mihelčič 1956] S04

*Type species of generic name indicated

REFERENCES

[BB92] Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

[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.

[S22] Schweizer, J. 1922. Beitrag zur Kenntnis der terrestrischen Milbenfauna der Schweiz. Verhandl. Naturf. Ges. Basel 33: 23–112, 4 pls.

[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.

Tenuialidae

Hafenrefferia sp., photographed by Pfiegler Walter.


Belongs within: Circumdehiscentiae.

The Tenuialidae are a family of oribatid mites characterised by the presence of a long humeral process. They are most abundant in forest litter in temperate regions of the Holarctic. Depressions in the podosoma allow members of this family to retract their legs (Norton & Behan-Pelletier 2009).

Characters (from Norton & Behan-Pelletier 2009): Length 800–1200 µm. Subcapitulum diarthric; chelicerae chelate-dentate. Prodorsum and notogaster smooth, shiny. Prodorsum with true lamellae. Notogaster with longhumeral processes projecting anterolaterally to bothridium; octotaxic system absent; dorsophragmata and pleurophragmata present. Podosomal region with depressions for retracted legs; epimeral border IV with minitectum.

<==Tenuialidae
    |--Ceratotenuiala Aoki & Maruyama 1983 S04
    |    `--*C. echigoensis Aoki & Maruyama 1983 S04
    |--Tenuialoides Woolley & Higgins 1966 S04
    |    |--*T. medialis Woolley & Higgins 1966 S04
    |    `--T. fusiformis Aoki 1969 [incl. Hafenferrefia translamellata Aoki & Fujikawa 1979] S04
    |--Hafenrefferia Oudemans 1906 S04
    |    |--*H. gilvipes (Koch 1839) (see below for synonymy) S04
    |    `--H. acuta Aoki 1966 S04
    |--Peltenuiala Norton 1983 S04
    |    |--*P. pacifica Norton 1983 S04
    |    `--P. orbiculata (Aoki & Ohnishi 1974) [=Hafenferrefia orbiculata] S04
    |--Tenuiala Ewing 1913 S04
    |    |--*T. nuda Ewing 1913 S04
    |    |--T. crenulata Woolley & Higgins 1966 S04
    |    `--T. kurti Woolley & Higgins 1955 S04
    `--Hafenferrefia Jacot 1939 [incl. Hafenrefferiella Sellnick 1952] S04
         |--*H. nitidula (Banks 1906) [=Galumna nitidula] S04
         |--H. hyrcanica (Krivolutsky 1967) [=Hafenrefferiella hyrcanica] S04
         |--H. imperialis (Berlese 1910) (n. d.) [=Achipteria imperialis] S04
         |--H. lata Golosova 1980 S04
         `--H. nevesi (Sellnick 1952) [=Hafenrefferiella nevesi] S04

*Haffenrefferia gilvipes (Koch 1839) [=Oribata gilvipes; incl. Liacarus auritus Nordenskiöld 1901, L. pterotus Coggi 1900] S04

*Type species of generic name indicated

REFERENCES

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.

[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.

Carabodidae

Dorsal view of Carabodes dubius, from Murvanidze (2008).


Belongs within: Circumdehiscentiae.
Contains: Odontocepheus (Odontocepheus), Diplobodes, Yoshiobodes, Gibbicepheus (Gibbicepheus), Austrocarabodes (Austrocarabodes), Carabodes (Carabodes), Carabodes (Klappariches).

The Carabodidae are a family of oribatid mites with more or less parallel lamellae on the prodorsum (Balogh & Balogh 1992). Though diverse in adult biology, all species are probably endophagous as nymphs, burrowing in substrates such as decaying wood. Adults of many species are arboreal, feeding on fungi and lichens (Norton & Behan-Pelletier 2009). In a number of genera the notogaster bears longitudinal rows of large tubercles, such as in Gibbicepheus or Diplobodes. The genera Spathulocepheus and Trichocarabodes are distinctive in the high number of setae on their genital plates: eight pairs in Trichocarabodes, ten in Spathulocepheus. Other genera have at most seven pairs of genital setae (Balogh & Balogh 1992).

Characters (from Balogh & Balogh 1992): Prodorsum with true lamellae; lamellae marginal, almost parallel or weakly converging; interlamellar setae well developed. Notogaster usually pentagonal, with more or less parallel sides and large humeral tubercle.

<==Carabodidae
    |--Gibbicepheus Aoki 1958 S04 [Gibbicepheinae CH98]
    |    |--G. (Gibbicepheus) S04
    |    `--G. (Gibbibodes Mahunka 1986) S04
    |         `--G. (*G.) similis (Mahunka 1986) [=*Gibbibodes similis] S04
    `--Carabodinae CH98
         |--Austrocarabodes Hammer 1966 CH98
         |    |--A. (Austrocarabodes) S04
         |    |--A. (Baloghodes Mahunka 1986) S04
         |    |    |--A. (*B.) heterotrichosus (Mahunka 1986) [=*Baloghodes heterotrichosus] S04
         |    |    `--A. (B.) secundus (Mahunka 1993) [=Baloghodes secundus] S04
         |    `--A. (Uluguroides Mahunka 1983) [incl. Ngorongobodes Balogh & Balogh 1992] S04
         |         |--A. (*U.) trichosus (Mahunka 1983) [=*Uluguroides trichosus] S04
         |         |--A. (U.) grandis (Mahunka 1984) [=Ulugurozetes grandis] S04
         |         |--A. (U.) incrustatus (Wallwork 1977) [=Carabodes incrustatus] S04
         |         |--A. (U.) minitricha (Mahunka 1993) [=Uluguroides minitricha] S04
         |         `--A. (U.) pentatrichus (Balogh 1962) S04 (see below for synonymy)
         |              |--A. p. pentatrichus S04
         |              `--A. p. ‘clavatus’ (Wallwork 1977) (see below for synonymy) S04
         `--Carabodes Koch 1835 S04
              |  i. s.: C. elongatus S22
              |--C. (Carabodes) S04
              |--C. (Flexa Kulijev 1977) S04
              |    |--C. (*F.) dubius Kulijev 1968 S04
              |    |--C. (F.) bidens (Djaparidze 1990) [=Flexa bidens] S04
              |    |--C. (F.) breviclava Aoki 1970 S04
              |    |--C. (F.) horeo (Djaparidze 1990) [=Flexa horeo] S04
              |    |--C. (F.) intermedius Willmann 1951 S04
              |    `--C. (F.) scopulae Kulijev 1968 S04
              |--C. (Klapperiches) S04
              `--C. (Phyllocarabodes Balogh & Mahunka 1969) (see below for synonymy) S04
                   |--C. (*P.) octogonalis (Balogh & Mahunka 1969) [=*Phyllocarabodes octogonalis] S04
                   |--C. (P.) calcaratus (Mahunka 1985) [=Antillobodes calcaratus] S04
                   |--C. (P.) inopinatus (Mahunka 1985) [=Antillobodes inopinatus] S04
                   |--C. (P.) insolitus (Balogh 1984) S04 [=*Pentabodes insolitus BB92]
                   `--C. (P.) ‘ornatus’ (Balogh 1986) non C. ornatus Storkan 1925 [=Phyllocarabodes ornatus] S04

Carabodidae incertae sedis:
  Apotomocepheus Aoki 1965 S04
    `--*A. gressitti Aoki 1965 S04
  Meriocepheus Aoki 1973 S04
    `--*M. peregrinus Aoki 1973 S04
  Sagittabodes Balogh & Balogh 1992 S04
    `--*S. sagitta (Balogh & Mahunka 1966) [=Machadocepheus sagitta] S04
  Tansocepheus Mahunka 1983 S04
    `--*T. serratus Mahunka 1983 S04
  Tectocarabodes Mahunka 1988 S04
    `--*T. monstruosus Mahunka 1988 S04
  Tuberocepheus Balogh & Mahunka 1969 S04
    `--*T. longus (Balogh 1964) [=Machadocepheus longus] S04
  Singabodes Mahunka 1998 S04
    `--*S. rarus Mahunka 1998 S04
  Archegocepheus Aoki 1965 S04
    |--*A. imadatei Aoki 1965 S04
    `--A. macrofoliatus Wen 1997 S04
  Bathocepheus Aoki 1978 [incl. Philippobodes Balogh & Balogh 1992] S04
    |--*B. concavus Aoki 1978 S04
    `--B. manguiati (Corpuz-Raros 1979) S04 [=Machadocepheus manguiati S04, *Philippobodes manguiati BB92]
  Cavernocarabodes Mahunka 1974 S04
    |--*C. perreti Mahunka 1974 S04
    `--C. trigonosternum (Pérez-Íñigo 1976) [=Carabodes trigonosternum] S04
  Spathulocepheus Balogh & Mahunka 1969 S04
    |--*S. amazonicus Balogh & Mahunka 1969 S04
    `--S. plumiger Balogh & Mahunka 1977 S04
  Malgasodes Mahunka 2000 S04
    |--*M. curvisetus Mahunka 2000 S04
    `--M. hungarorum Mahunka 2000 S04
  Gymnobodes Balogh 1965 S04
    |--*G. fraterculus (Balogh 1963) [=Carabodes fraterculus] S04
    |--G. semengok Mahunka 1996 S04
    `--G. subnudus (Balogh 1963) [=Carabodes subnudus] S04
  Pseudocarabodes Mahunka 1991 [incl. Bakobodes Mahunka 1996, Yemenobodes Mahunka 2000] S04
    |--*P. xenus Mahunka 1991 S04
    |--P. improvisus (Mahunka 2000) [=Yemenobodes improvisus] S04
    `--P. orangutan (Mahunka 1996) [=Bakobodes orangutan] S04
  Trichocarabodes Balogh 1961 [incl. Ceylobodes Balogh & Balogh 1992] S04
    |--*T. celisi (Balogh 1958) [=Carabodes celisi] S04
    |--T. capillatus Balogh 1970 S04 [=*Ceylobodes capillatus BB92]
    |--T. costulatus Mahunka 1984 S04
    `--T. hettigei Balogh 1970 S04
  Hardybodes Balogh 1970 [incl. Carabodella Mahunka 1986] S04
    |--*H. mirabilis Balogh 1970 S04
    |--H. calcaratus (Mahunka 1986) S04 [=*Carabodella calcarata BB92]
    |--H. flabellatus Mahunka 1995 S04
    `--H. penicillatus Mahunka 1995 S04
  Cubabodes Balogh & Mahunka 1974 S04
    |--*C. hexagonalis Balogh & Mahunka 1974 S04
    |--C. confertus Balogh & Mahunka 1980 S04
    |--C. radiatus Balogh & Mahunka 1974 S04
    |--C. spathulatus Balogh & Mahunka 1980 S04
    `--C. verrucatus Balogh & Mahunka 1980 S04
  Machadocepheus Balogh 1958 S04
    |--*M. excavatus Balogh 1958 S04
    |--M. filiferus Mahunka 1986 S04
    |--M. pauliani Balogh 1962 S04
    |--M. phyllophorus Balogh 1970 S04
    |--M. taprobanicus Balogh 1970 S04
    `--M. tuberculosus Mahunka 1987 S04
  Odontocepheus Berlese 1913 S04
    |--O. (Odontocepheus) S04
    |--O. (Aokiella Balogh & Mahunka 1967) S04
    |    |--O. (*A.) florens (Balogh & Mahunka 1967) [=*Aokiella florens] S04
    |    `--O. (A.) rotundus (Hammer 1979) [=Aokiella rotunda] S04
    `--O. (Indotocepheus Mondal & Kundu 1999) S04
         `--O. (*I.) himalayensis (Mondal & Kundu 1999) [=*Indotocepheus himalayensis] S04
  Pasocepheus Aoki 1976 S04
    |--P. (Pasocepheus) [incl. Opisthocepheus Aoki 1976] S04
    |    |--*P. (P.) triarcuatus Aoki 1976 S04
    |    |--P. (P.) bako Mahunka 1996 S04
    |    |--P. (P.) eremaeozetoides Mahunka 1995 S04
    |    `--P. (P.) kirai (Aoki 1976) S04 [=*Opisthocepheus kirai BB92]
    `--P. (Guineobodes Mahunka 1987) S04
         `--P. (*G.) papuanus (Balogh 1970) [=Machadocepheus papuanus] S04
  Congocepheus Balogh 1958 S04
    |--*C. heterotrichus Balogh 1958 S04
    |--C. hauseri Mahunka 1989 S04
    |--C. involutus Mahunka 1997 S04
    |--C. latilamellatus Mahunka 1984 S04
    |--C. orientalis Mahunka 1989 S04
    |--C. ornatus Mahunka 1983 S04
    |--C. taurus Balogh 1961 S04
    `--C. velatus Mahunka 1986 S04
  Diplobodes S04
  Yoshiobodes S04

Austrocarabodes (Uluguroides) pentatrichus (Balogh 1962) S04 [=Carabodes pentatrichus S04, *Ngorongobodes pentatrichus BB92]

Austrocarabodes (Uluguroides) pentatrichus ‘clavatus’ (Wallwork 1977) [=Carabodes pentatrichus clavatus nec C. labyrinthicus clavatus Warburton 1912 nec C. gibbiceps clavatus Jacot 1938] S04

Carabodes (Phyllocarabodes Balogh & Mahunka 1969) [incl. Antillobodes Mahunka 1985, Pentabodes Balogh 1984] S04

*Type species of generic name indicated

REFERENCES

[BB92] Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

[CH98] Colloff, M. J., & R. B. Halliday. 1998. Oribatid Mites: A catalogue of Australian genera and species. CSIRO Publishing: Collingwood.

Murvanidze, M. 2008. Checklist and key to species of Carabodes (Acari, Oribatida, Carabodidae) of the Caucasian region, with description of a new species. Acarina 16 (2): 177–186.

[S22] Schweizer, J. 1922. Beitrag zur Kenntnis der terrestrischen Milbenfauna der Schweiz. Verhandl. Naturf. Ges. Basel 33: 23–112, 4 pls.

[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.