Circumdehiscentiae

Dorsum and venter of Niphocepheus nivalis delamarei, from Balogh & Balogh (1992 vol. 2).

Belongs within: Desmonomata.
Contains: Microzetidae, Polypterozetoidea, Oppiidae, Autognetidae, Machuellidae, Quadroppiidae, Sternoppia, Granuloppiidae, Teratoppiidae, Suctobelbidae, Platyameridae, Banksinoma, Machadobelba, Damaeidae, Amerobelbidae, Damaeolidae, Eremobelba, Ameridae, Ctenobelba, Eremulidae, Heterobelbidae, Basilobelbidae, Rhynchoribatidae, Eremaeozetoidea, Tenuialidae, Astegistidae, Gustavia, Oribatellidae, Neoliodidae, Hermanniellidae, Plasmobatidae, Zetorchestidae, Liacaridae, Eremaeidae, Arceremaeidae, Peloppiidae, Cepheidae, Eutegaeidae, Microtegeidae, Anderemaeidae, Cymbaeremaeidae, Carabodidae, Nippobodidae, Otocepheidae, Dampfiellidae, Oripodoidea, Plateremaeidae, Licnodamaeidae, Pheroliodidae, Gymnodamaeidae, Pedrocortesellidae, Xenillidae, Hydrozetes, Limnozetidae, Austrachipteria, Achipterioidea, Tectocepheidae, Licneremaeoidea, Scapheremaeus, Ameronothridae, Fortuyniidae, Selenoribatidae, Tegeocranellus, Galumnidae, Galumnellidae, Ceratozetoidea, Phenopelopidae.

The Circumdehiscentiae, or 'higher oribatid mites' (also known as Brachypylina), are a major clade within the oribatids. They are characterised by having the genital and anal plates usually well separated on the ventral plate, and by having the tibiae and genua of the legs differing from each other in length and shape (Balogh & Balogh 1992). Relationships within the Circumdehiscentiae are, as yet, poorly understood. A number of authors have divided them between the Pycnonoticae and Poronoticae, based primarily on the presence (Poronoticae) or absence (Pycnonoticae) of the octotaxic system, a series of glandular openings on the notogaster. Alternatively, Pycnonoticae has been labelled as Apterogastrina and Poronoticae as Pterogastrina, in reference to the absence or presence, respectively, of pteromorphs, lateral extensions of the notogaster that may or may not be movable. However, the Pycnonoticae are undoubtedly paraphyletic, while the Poronoticae may possibly be polyphyletic. Conversely, characters of the nymphs have been considered significant in classification, such as whether they retain scalps (exuviae of previous instars) for protection (scalps are retained in 'Eupheredermata', lost in 'Apheredermata'), and whether the nymphs have a plicate (wrinkled) integument. However, immature Circumdehiscentiae are often very different in appearance from their adults, and the nymphs for many species remain undescribed. Molecular phylogenies of Circumdehiscentiae published to date have only included a small subsection of brachypyline diversity, with many major groups as yet unrepresented.

Eupheredermous taxa include the Plasmobatidae, Neoliodidae, Plateremaeoidea, Damaeidae, Cepheoidea, Polypterozetoidea, Microzetidae and most Ameroidea (Norton & Behan-Pelletier 2009). Some apheredermous taxa, such as Liacaridae and Peloppiidae, show a loss of dorsocentral setae on the notogaster similar to that found in eupheredermous taxa (Schäffer et al. 2010) which may indicate eupheredermous relationships. It seems likely that the Neoliodidae and Hermannielloidea are relatively basal in the brachypyline clade. Members of the Hermannielloidea are united by having the openings of the opisthonotal glands protruding from the notogaster, either on distinct funnel-shaped tubes or on large apophyses. Both Neoliodidae and Hermanielloidea also share precocious development of apodemato-acetabular brachytracheae in the immatures with members of the Eremaeoidea and Plateremaeoidea. The Eremaeoidea commonly have costulae on the prodorsum, and the notogastral cerotegument forming large tubercles. The Plateremaeoidea also have a well-developed cerotegument, and their notogaster is generally flattened in lateral view (Norton & Behan-Pelletier 2009).

<==Circumdehiscentiae (see below for synonymy) SK10
    |--+--Neoliodidae DA07
    |  `--Hermannielloidea S04
    |       |--Hermanniellidae SK10
    |       `--Plasmobatidae S04
    `--+--Zetorchestidae SK10
       `--+--+--Liacaridae SK10
          |  `--+--Eremaeoidea S04
          |     |    |--Eremaeidae DW10
          |     |    |--Arceremaeidae NB-P09
          |     |    |--Aribates Aoki, Takaku & Ito 1994 [Aribatidae] S04
          |     |    |    `--*A. javensis Aoki, Takaku & Ito 1994 S04
          |     |    |--Megeremaeus Higgins & Woolley 1965 [Megeremaeidae] S04
          |     |    |    |--*M. montanus Higgins & Woolley 1965 S04
          |     |    |    |--M. ditrichosus Woolley & Higgins 1968 S04
          |     |    |    |--M. expansus Aoki & Fujikawa 1971 S04
          |     |    |    |--M. hylaius Behan-Pelletier 1990 S04
          |     |    |    |--M. keewatin Behan-Pelletier 1990 S04
          |     |    |    |--M. kootenai Behan-Pelletier 1990 S04
          |     |    |    |--M. rameus Wen & Zhao 1993 S04
          |     |    |    `--M. spinosus Aoki & Yamamoto 2000 S04
          |     |    `--Niphocepheus Balogh 1943 S04 [Niphocepheidae SK10, Niphocepheoidea]
          |     |         |--*N. nivalis (Schweizer 1922) [=Cepheus nivalis] S04
          |     |         |    |--N. n. nivalis S04
          |     |         |    |--N. n. baloghi Travé 1959 S04
          |     |         |    |--N. n. delamarei Travé 1959 S04
          |     |         |    `--N. n. grandjeani Travé 1959 S04
          |     |         |--N. aborigensis Behan-Pelletier 1982 S04
          |     |         |--N. guadarramicus Subías 1977 S04
          |     |         `--N. travei Bulanova-Zavchvatkina 1967 S04
          |     `--+--Peloppiidae SK10
          |        `--Cepheoidea [Eutegeoidea] S04
          |             |--Cepheidae MH04
          |             |--Eutegaeidae SK10
          |             |--Microtegeidae NB-P09
          |             |--Anderemaeidae NB-P09
          |             |--Pterobates Balogh & Mahunka 1977 [Pterobatidae] S04
          |             |    `--*P. incertus Balogh & Mahunka 1977 S04
          |             |--Cerocepheidae S04
          |             |    |--Cerocepheus Trägardh 1931 S04      
          |             |    |    `--*C. mirabilis Trägardh 1931 S04
          |             |    |--Dicrotegaeus Luxton 1988 S04
          |             |    |    `--*D. mirabilis Luxton 1988 S04
          |             |    `--Bornebuschia Hammer 1966 S04
          |             |         |--*B. peculiaris Hammer 1966 S04
          |             |         `--B. binodosa Luxton 1988 S04
          |             `--Nosybeidae NB-P09
          |                  |--Lamellocepheus Balogh 1961 [incl. Nosybea Mahunka 1993, Tessacarus Grandjean 1962] S04
          |                  |    |--*L. personatus (Berlese 1910) S04 (see below for synonymy)
          |                  |    `--L. genavensis (Mahunka 1993) [=Nosybea genavensis] S04
          |                  `--Topalia Balogh & Csiszár 1963 S04
          |                       |--*T. problematica Balogh & Csiszár 1963 S04
          |                       |--T. africana Mahunka 1985 S04
          |                       |--T. clavata Hammer 1966 S04
          |                       |--T. granulata Hammer 1966 S04
          |                       `--T. velata Hammer 1966 S04
          `--+--Cymbaeremaeidae SK10
             `--+--+--Carabodoidea [Otocepheoidea] S04
                |  |    |--Carabodidae SK10
                |  |    |--Nippobodidae S04
                |  |    |--Otocepheidae NB-P09
                |  |    |--Dampfiellidae S04
                |  |    |--Tokunocepheus Aoki 1966 [Tokunocepheidae] S04
                |  |    |    `--*T. mizusawai Aoki 1966 S04
                |  |    `--Carabocepheus Berlese 1910 [Carabocepheidae] S04
                |  |         `--*C. lounsburyi (Berlese 1910) [=Carabodes (*Carabocepheus) lounsburyi] S04
                |  |              |--C. l. lounsburyi S04
                |  |              `--C. l. latior Balogh & Mahunka 1966 S04
                |  `--+--Oripodoidea SK10
                |     `--Plateremaeoidea [Gymnodamaeoidea, Pheronotae] NB-P09
                |          |--Plateremaeidae S04
                |          |--Licnodamaeidae S04
                |          |--Pheroliodidae S04
                |          |--Gymnodamaeidae SK10
                |          |--Pedrocortesellidae NB-P09
                |          |--Nacunansella Fernández & Cleva 1998 S04 [Nacunansellidae NB-P09]
                |          |    `--*N. diminuta Fernández & Cleva 1998 S04
                |          |--Lyrifissella Paschoal 1989 NB-P09, S04 [Lyrifissellidae]
                |          |    |--*L. latoclava (Hammer 1966) [=Pedrocortesella latoclava] S04
                |          |    |--L. cryptonota (Hammer 1966) [=Pedrocortesella cryptonota] S04
                |          |    `--L. vestita (Trägardh 1931) [=Plateremaeus vestitus] S04
                |          |--Idiodamaeidae NB-P09
                |          |    |--Austrodamaeus Balogh & Mahunka 1981 S04
                |          |    |    `--A. elegantulus (Hammer 1958) S04 (see below for synonymy)
                |          |    `--Idiodamaeus Paschoal 1984 NB-P09, S04
                |          |         `--*I. illecebrosus Paschoal 1984 NB-P09, S04
                |          |--Licnobelbidae S04
                |          |    |--Flabellobelba Pérez-Íñigo 1995 S04
                |          |    |    `--*F. almeriensis (Ruiz, Kahwash & Subías 1990) [=Licnobelba almeriensis] S04
                |          |    `--Licnobelba Grandjean 1931 S04
                |          |         |--*L. alestensis Grandjean 1931 S04, NB-P09, S04
                |          |         |--L. caesarea (Berlese 1910) [=Licneremaeus caesareus] S04
                |          |         `--L. latiflabellata (Paoli 1908) (see below for synonymy) S04
                |          `--Aleurodamaeus Grandjean 1954 [Aleurodamaeidae] S04
                |               |  i. s.: A. australis Woas 1992 NB-P09, S04 [=Austrodamaeus australis S04]
                |               |--A. (Aleurodamaeus) S04
                |               |    |--*A. (A.) setosus (Berlese 1883) (see below for synonymy) S04
                |               |    |--A. (A.) cephalotes (Berlese 1916) [=Gymnodamaeus cephalotes] S04
                |               |    |--A. (A.) hungaricus Paschoal & Johnston 1985 S04
                |               |    `--A. (A.) trichosus (Kulijev 1979) [=Allodamaeus trichosus] S04
                |               `--A. (Trichodamaeus Mahunka 1984) S04
                |                    `--A. (*T.) africanus Mahunka 1984 S04
                `--+--Xenillidae DW10
                   |--Hydrozetoidea [Limnozetoidea]
                   |    |--Hydrozetes SK10
                   |    |--Limnozetidae NB-P09
                   |    `--Austrachipteriidae S04
                   |         |--Austrachipteria S04
                   |         |--Granizetes Hammer 1961 S04
                   |         |    `--*G. curvatus Hammer 1961 S04
                   |         |--Palmitalia Pérez-Íñigo & Peña 1997 S04
                   |         |    `--*P. latilamellata Pérez-Íñigo & Peña 1997 S04
                   |         |--Patagonozetes Balogh & Balogh 1990 S04
                   |         |    `--*P. exornatus (Balogh & Csiszár 1963) [=Sphaerozetes exornatus] S04
                   |         |--Punctizetes Hammer 1971 S04
                   |         |    `--*P. penicillifer Hammer 1971 S04
                   |         |--Sagittazetes Balogh & Balogh 1983 S04
                   |         |    `--*S. agressor Balogh & Balogh 1983 S04
                   |         |--Ceratobates Balogh & Mahunka 1969 S04
                   |         |    |--*C. pontiger Balogh & Mahunka 1969 S04
                   |         |    |--C. fornerisae Pérez-Íñigo & Baggio 1985 S04
                   |         |    `--C. spathulatus Balogh & Mahunka 1981 S04
                   |         `--Allozetes Berlese 1913 S04
                   |              |--*A. pusillus (Berlese 1913) [=Ceratozetes (*Allozetes) pusillus] S04
                   |              |--A. africanus Balogh 1958 S04
                   |              |--A. alas Behan-Pelletier 1998 S04
                   |              |--A. lacandonicus Mahunka & Palacios-Vargas 1996 S04
                   |              |--A. latus Tseng 1984 S04
                   |              |--A. levis Ohkubo 1981 S04
                   |              `--A. translamellatus Hammer 1973 S04
                   `--+--+--Achipterioidea SK10
                      |  `--Tectocepheidae SK10
                      `--+--+--Licneremaeoidea SK10
                         |  |--Scapheremaeus SK10
                         |  `--Ameronothroidea S04
                         |       |--Ameronothridae SK10
                         |       |--Fortuyniidae S04
                         |       |--Selenoribatidae S04
                         |       `--Tegeocranellus NB-P09
                         `--+--Galumnoidea B-P01
                            |    |--Galumnidae SK10
                            |    `--Galumnellidae S04
                            `--+--Podacarus Grandjean 1955 SK10, S04 [Podacaridae]
                               |    `--*P. auberti Grandjean 1955 S04
                               |         |--P. a. auberti S04
                               |         `--P. a. occidentalis Wallwork 1966 S04
                               `--+--Ceratozetoidea SK10
                                  `--Phenopelopoidea [Pelopoidea] NB-P09
                                       |  i. s.: Plakoribates Popp 1960 NB-P09, S04
                                       |           |--*P. multicuspidatus Popp 1960 S04
                                       |           |--P. confluens Balogh 1970 S04
                                       |           |--P. neotropicus Balogh & Mahunka 1978 S04
                                       |           `--P. scutatus Hammer 1979 S04
                                       |--Phenopelopidae SK10
                                       `--Unduloribatidae [Unduloribatoidea] NB-P09
                                            |--Koreoribates Choi 1994 S04
                                            |    `--*K. foliatus Choi 1994 S04
                                            `--Unduloribates Balogh 1943 S04
                                                 |--*U. undulatus (Berlese 1914) (see below for synonymy) S04
                                                 |--U. brevisetosus Nübel-Reidelbach & Woas 1992 S04
                                                 |--U. hebes Aoki 1965 S04
                                                 |--U. medusa Piffl 1972 S04
                                                 `--U. parvus NB-P09

Circumdehiscentiae incertae sedis:
  Microzetidae S04
  Polypterozetoidea S04
  Oppioidea [Trizetoidea] NB-P09
    |--Oppiidae S04
    |--Autognetidae S04
    |--Machuellidae S04
    |--Quadroppiidae S04
    |--Sternoppia S04
    |--Granuloppiidae S04
    |--Teratoppiidae S04
    |--Suctobelbidae S04
    |--Platyameridae NB-P09
    |--Trizetes Berlese 1904 [Trizetidae] S04
    |    `--*T. pyramidalis Berlese 1904 S04
    |--Nosybelba Mahunka 1994 [Nosybelbidae] S04
    |    `--*N. oppiana Mahunka 1994 S04
    |--Enantioppia Balogh & Mahunka 1969 S04 [Enantioppiidae NB-P09]
    |    `--*E. multituberculata Balogh & Mahunka 1969 S04
    |--Hexoppia Balogh 1958 S04 [Hexoppiidae NB-P09]
    |    `--*H. heterotricha Balogh 1958 S04
    |--Luxtonia Mahunka 2001 S04 [Luxtoniidae NB-P09]
    |    `--*L. hauseri Mahunka 2001 S04
    |--Tuparezetes Spain 1969 [Tuparezetidae] S04
    |    |--*T. christineae Spain 1969 S04
    |    `--T. philodendrus Spain 1969 S04
    |--Cuneoppia Balogh & Mahunka 1969 S04 [Cuneoppiidae NB-P09]
    |    |--*C. laticeps Balogh & Mahunka 1969 S04
    |    `--C. dogmai Corpuz-Raros 1979 S04
    |--Chavinia Hammer 1961 S04 [Chaviniidae NB-P09, Chaviniinae]
    |    |--*C. paradoxa Hammer 1961 S04
    |    `--C. similis Balogh 1984 S04
    |--Papillonotus Wallwork 1961 [Papillonotidae] S04
    |    |--*P. maculatus Walwork 1961 S04
    |    |--P. granulosus Wallwork 1961 S04
    |    |--P. hauseri Mahunka 1988 S04
    |    `--P. tricarinatus Sarkar & Subías 1983 S04
    |--Thyrisomidae NB-P09
    |    |--Banksinoma S04
    |    `--Gemmazetes Fujikawa 1979 NB-P09, S04
    |         `--*G. crosbyi (Berlese 1908) BB92, S04 [=Oribella crosbyi S04, Oribellopsis crosbyi S04]
    |              |--G. c. crosbyi S04
    |              `--‘Oribellopsis’ c. maoershanensis Wen 1992 S04
    |--Lyroppia Balogh 1961 [Lyroppiidae] S04
    |    |--*L. scutigera Balogh 1961 S04
    |    |--L. anareolata Balogh & Mahunka 1981 S04
    |    |--L. delicata Wang & Lu 1995 S04
    |    |--L. neotropica Balogh & Mahunka 1974 S04
    |    `--L. similis Balogh & Mahunka 1977 S04
    |--Machadobelbidae [Rioppiidae] NB-P09
    |    |--Machadobelba S04
    |    |--Ramogneta Karppinen 1966 S04
    |    |    `--*R. lamottei Karppinen 1966 S04
    |    `--Rioppia Balogh & Mahunka 1977 S04
    |         |--*R. nodulifera Balogh & Mahunka 1977 S04
    |         `--R. comteae Mahunka 1985 S04
    `--Epimerellidae S04
         |--Enisella Ayyildiz & Luxton 1989 S04
         |    `--*E. turcica Ayyildiz & Luxton 1989 S04
         `--Epimerella Kulijev 1967 S04
              |--*E. smirnovi (Kulijev 1962) [=Oppia smirnovi] S04
              |    |--E. s. smirnovi S04
              |    `--E. s. longisetosa Kulijev 1967 S04
              |--E. distenta Ayyildiz & Luxton 1989 S04
              |--E. puzanovi Gordeeva & Karppinen 1988 S04
              `--E. rubeni Khanbekjan & Gordeeva 1991 S04
  Damaeidae MH04
  Ameroidea [Amerobelboidea, Eremuloidea] NB-P09
    |--Amerobelbidae S04
    |--Damaeolidae S04
    |--Eremobelba S04
    |--Ameridae S04
    |--Ctenobelba S04
    |--Eremulidae S04
    |--Heterobelbidae S04
    |--Basilobelbidae S04
    |--Rhynchoribatidae NB-P09
    |--Staurobatidae S04
    |    |--Stauroma Grandjean 1966 S04
    |    |    `--*S. cephalotum Grandjean 1966 S04
    |    `--Staurobates Grandjean 1966 S04
    |         `--*S. schusteri Grandjean 1966 S04
    |              |--S. s. schusteri S04
    |              `--S. s. cordobensis Balogh & Mahunka 1968 S04
    |--Oxyamerus Aoki 1965 NB-P09, S04 [Oxyameridae S04]
    |    |--*O. spathulatus Aoki 1965 S04
    |    |--O. aokii Balogh 1968 S04
    |    |--O. hauserorum Mahunka 1987 S04
    |    |--O. hyalinus Hammer 1979 S04
    |    |--O. latirostris Balogh 1968 S04
    |    `--O. truncatus Hammer 1979 S04
    |--Spinozetidae NB-P09
    |    |--Iberoppia Pérez-Íñigo 1986 S04
    |    |    `--*I. paradoxa Pérez-Íñigo 1986 S04
    |    |--Grypoceramerus Suzuki & Aoki 1970 S04
    |    |    `--*G. acutus Suzuki & Aoki 1970 S04
    |    `--Spinozetes Piffl 1966 [incl. Mirus Kulijev 1967 nec Albers 1850 nec Saulcy 1877] S04
    |         |--*S. inexspectatus Piffl 1966 S04
    |         `--S. pectinatus (Kulijev 1967) [=Mirus pectinatus] S04
    |--Hungarobelbidae S04
    |    |--Hungarobelba Balogh 1943 S04
    |    |    |--*H. visnyai (Balogh 1938) [=Belba visnyai] S04
    |    |    |--H. baloghi Bulanova-Zachvatkina 1967 S04
    |    |    `--H. pyrenaica Miko & Travé 1996 S04
    |    `--Costeremus Aoki 1970 S04
    |         |--*C. ornatus Aoki 1970 S04
    |         |--C. barbatus Choi 1997 S04
    |         |--C. cornutus Wang 1996 S04
    |         `--C. yezoensis Fujikawa & Fujita 1985 S04
    `--Caleremaeidae [Caleremaeoidea] NB-P09
         |--Luxtoneremaeus Balogh & Balogh 1922 S04
         |    `--*L. forsteri (Balogh & Balogh 1985) [=Anderemaeus forsteri] S04
         |--Veloppia Hammer 1955 S04
         |    |--*V. pulchra Hammer 1955 S04
         |    `--V. kananaskis Norton 1978 S04
         `--Caleremaeus Berlese 1910 S04
              |--*C. monilipes (Michael 1882) S04 [=Damaeus monilipes S04, Notaspis monilipes BB92]
              |--C. divisus Mihelčič 1952 S04
              `--C. retractus (Banks 1947) [=Carabodoides retractus] S04
  Eremaeozetoidea NB-P09
  Tenuialidae S04
  Astegistidae S04
  Gustavia S04
  Multoribulidae S04
    |--Peloptoribula Mahunka 1984 S04
    |    `--*P. spinulosa Mahunka 1984 S04
    `--Multoribula Balogh & Mahunka 1966 S04
         |--*M. multipunctata Balogh & Mahunka 1966 S04
         `--M. suramericana Balogh & Mahunka 1977 S04
  Oribatellidae B-P01
  Kodiakella Hammer 1967 S04 [Kodiakellidae NB-P09]
    |--*K. lutea Hammer 1967 S04
    `--K. dimorpha Pérez-Íñigo & Subías 1978 S04

*Aleurodamaeus (Aleurodamaeus) setosus (Berlese 1883) [=Damaeus setosus; incl. Gymnodamaeus gibbus Mihelčič 1963, G. nitidus Mihelčič 1956] S04

Austrodamaeus elegantulus (Hammer 1958) S04 [=Gymnodamaeus elegantulus S04; incl. *Au. rimosus Balogh & Mahunka 1981 S04, Allodamaeus trisetosus Balogh & Mahunka 1969 S04]

Circumdehiscentiae [Apheredermata, Apterogasterina, Brachypylides, Brachypylina, Carinotae, Euoribatida, Eupheredermata, Gustavioidea, Gymnonota, Holosomatina, Liacaroidea, Notaspidinae, Planofissurae, Poronota, Poronotae, Poronoticae, Pterogasterina, Pycnonota, Pycnonoticae] SK10

*Lamellocepheus personatus (Berlese 1910) S04 [=Tectocepheus personatus S04, Tegeocranus personatus BB92; incl. Lamellocepheus ambitus Kulijev 1966 S04]

Licnobelba latiflabellata (Paoli 1908) [=Licneremaeus latiflabellatus; incl. Licno. montana Mihelčič 1957] S04

*Unduloribates undulatus (Berlese 1914) [=Sphaerozetes (Tectoribates) undulatus] 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.

[B-P01] Behan-Pelletier, V. M. 2001. Phylogenetic relationships of Hypozetes (Acari: Tegoribatidae). In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 50–57. CSIRO Publishing: Melbourne.

[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.

[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.

[MH04] Maraun, M., M. Heethoff, K. Schneider, S. Scheu, G. Weigmann, J. Cianciolo, R. H. Thomas & R. A. Norton. 2004. Molecular phylogeny of oribatid mites (Oribatida, Acari): evidence for multiple radiations of parthenogenetic lineages. Experimental and Applied Acarology 33 (3): 183–201.

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

[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.

[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.