Exochocepheus sp., copyright Centre for Biodiversity Genomics.

Belongs within: Poronoticae.
Contains: Bipassalozetes, Passalozetes, Licneremaeus, Micreremus, Scutovertex.

The Licneremaeoidea are a group of fungi-feeding oribatid mites found in periodically or permanently dry environments such as deserts or tree canopies (Norton & Behan-Pelletier 2009) The octotaxic system of glandular openings is reduced or absent in some families, making them potentially of interest for determining whether this system is monophyletic in oribatids. In Scutoverticidae, for instance, most genera lack the octotaxic system, but one pair of sacculi is present in Scutovertex and three pairs in Exochocepheus. As recognised by Norton & Behan-Pelletier (2009), the Licneremaeoidea includes the family Eremellidae, previously included in a distinct superfamily Eremelloidea with a number of other families that they placed in the Oppioidea and Ameroidea. Eremellidae have H-shaped costulae on the prodorsum, and seven pairs of notogastral setae arranged in a submarginal longitudinal row (Norton & Behan-Pelletier 2009). Licneremaeoidea also include the Neotropical Charassobates whose affinities have been subject to some debate.

A number of families in the Licneremaeoidea are primarily arboreal, including the Licneremaeidae, Dendroeremaeidae, Micreremidae and Adhaesozetidae. The Adhaesozetidae, found in east Asia and Australasia, have an expanded pulvillus under the tarsal claw that is used to grip the surfaces of leaves (Norton & Behan-Pelletier 2009). A lenticulus is commonly present at the front of the notogaster; in most Passalozetidae, this is oval and particularly sharply defined, sitting at the center of a distinct concavity (Norton & Behan-Pelletier 2009).

South American mites
Published 31 May 2011


Diagnostic views of Charassobates cavernosus (clockwise from left: dorsum; prodorsum with covering lamellae removed; venter) from Schuster (1969).


For this week’s random taxon, I drew the oribatid mite family Charassobatidae. This has proved to be something of a challenge: info on the charassobatids seems a little hard to come by. It doesn’t help that the ‘Charassobatidae’ has haemorrhaged taxa somewhat. Balogh & Balogh (1992), in their invaluable (if not entirely unproblematic) identification guide to oribatids, listed three genera in the Charassobatidae: Charassobates, Topalia and Ametroproctus. The main reason for associating these genera appears to have been the presence of massively expanded lamellae on the prodorsum (the ‘head’ part of the mite). However, Behan-Pelletier (1988), in transferring Ametroproctus to the family Cymbaeremaeidae, argued that this feature had probably arisen convergently as similar large lamellae are known from other oribatid families. Topalia has also since been removed from Charassobatidae, which is now redundant with Charassobates. Norton & Behan-Pelletier (2009) placed Charassobates in the superfamily Licneremaeoidea, but the monophyly of that group has been questioned (Schäffer et al. 2010).

Charassobates is a strictly South American group of mites (with an outlying species in the Galapagos). They have pelopsiform (very long and slender) chelicerae, possibly indicating a liquid diet, that are also somewhat unusual among oribatids in lacking an articulation between the chelicerae and the underside of the ‘head’. Some species, such as the type Charassobates cavernosus, also possess deep fossae on the dorsal surface of the main body. The nymphs are wrinkly (unlike humans, some oribatids become less wrinkly as they get older) and apheredermous (i.e. they do not carry the shed skins of former instars as protective scalps) according to Norton & Behan-Pelletier (2009) and Behan-Pelletier & Walter (2007), indicating that their description as eupheredermous (carrying scalps) by Behan-Pelletier (1988) was probably an error.

Systematics of Licneremaeoidea

Characters (from Norton & Behan-Pelletier 2009): Prodorsum with or without lamellae or costulae; lamellar cusps and translamella present or absent; tutorium absent. Genal notch present or absent. Dorsophragmata and pleurophragmata present. Chelicerae chelate-dentate; palpal eupathidium acm attached to, or separate from, solenidion; subcapitular mentum with or without complete tectum. Axillary saccule of the subcapitulum present or absent. Discidium, custodium, and postanal porose area absent. Octotaxic system present as 1-4 pairs of porose areas or saccules, or absent. Integument often tuberculate, nodulate, or reticulate. Pedotectum I present; pedotectum II present or absent. Notogaster with or without lenticulus, posterior notogastral tectum present or absent, if present, overlapping medially or not; pteromorph absent. Notogastral setation 10-15 pairs. Coxisternal setation either 3-1-3-3 or 3-1-2-2. Porose organs of legs variously developed as porose areas, saccules, or tracheae. Genital setation 4-6 pairs. Genital plate proportionally large, subtriangular, with posterior margin aligned with posterior border of epimere IV. Anal setation 1-2 pairs. Solenidion δ1 of tibia I inserted on distinct cylindrical tubule. Immatures apheredermous, plicate. Larva unideficient; nymphs unideficient, bideficient or trideficient. Humeral organ (small, usually papilliform porose organ located anterolateral to seta c2) absent. Coupled seta d present to the tritonymph or absent from immatures.

<==Licneremaeoidea [Eremelloidea, Passalozetoidea]
    |--Dendroeremaeus Behan-Pelletier, Eamer & Clayton 2005 [Dendroeremaeidae]B-PEC05
    |    |--*D. krantzi Behan-Pelletier, Eamer & Clayton 2005B-PEC05
    |    `--D. foveolatus Behan-Pelletier, Eamer & Clayton 2005B-PEC05
    |    |--BipassalozetesNB-P09
    |    `--PassalozetesNB-P09
    |    |--LicneremaeusNB-P09
    |    `--Huilicheremaeus Fernández, Marcangeli & Eguaras 1997S04
    |         |--*H. michaii Fernández, Marcangeli & Eguaras 1997S04
    |         `--H. atypicus (Mahunka 1984) [=Licneremaeus atypicus]S04
    |    |--PhylleremusNB-P09
    |    `--Adhaesozetes Hammer 1966S04
    |         |--*A. barbarae Hammer 1966S04
    |         `--A. polyphyllos Walter & Behan-Pelletier 1993S04
    |--Charassobates Grandjean 1929S04 [CharassobatidaeNB-P09, Charassobatoidea]
    |    |--*C. cavernosus Grandjean 1929S04
    |    |--C. baudi Mahunka 1984S04
    |    |--C. incipatus Balogh & Mahunka 1974S04
    |    |--C. minimus Balogh & Mahunka 1981S04
    |    |--C. ornatus Balogh & Mahunka 1969S04
    |    |--C. simplex Balogh & Mahunka 1969S04
    |    `--C. tuberosus Balogh & Mahunka 1981S04
    |    |--Microlamellarea Coetzee 1987S04
    |    |    `--*M. engelbrechti Coetzee 1987S04
    |    |--Lamellarea Kok 1968S04
    |    |    |--*L. ardua Kok 1968S04
    |    |    |--L. digitata Kok 1968S04
    |    |    |--L. forceps Kok 1968S04
    |    |    `--L. quadrata Coetzee 1987S04
    |    `--Tenuelamellarea Subías & Iturrondobeitia 1978S04
    |         |--*T. hispanica Subías & Iturrondobeitia 1978S04
    |         |--T. argentinensis Martínez, Velis et al. 1995S04
    |         |--T. hawaiiensis Balogh 1985S04
    |         |--T. inexpectata Balogh 1984BB92b
    |         `--T. taprobanae (Oudemans 1915) [=Carabodes taprobanae]S04
    |--Micreremidae [Fenicheliidae]NB-P09
    |    |--MicreremusS04
    |    |--Mexiceremus Balogh & Balogh 1998S04
    |    |    `--*M. rafaelinus (Mahunka & Palacios-Vargas 1995) [=Micreremus rafaelinus]S04
    |    |--Porofenichelia Mahunka 1985NB-P09, S04
    |    |    `--*P. porosa Mahunka 1985BB92a, S04 [=Fenichelia porosaS04]
    |    |--Glanderemaeus Balogh & Csiszár 1963B-PEC05, S04
    |    |    `--*G. hammerae Balogh & Csiszár 1963S04
    |    |--Phylloribatula Balogh & Mahunka 1978S04
    |    |    |--*P. pulchella Balogh & Mahunka 1978S04
    |    |    `--P. xanthoparmeliae Martínez & Palacios-Vargas 1998S04
    |    `--Fenichelia Balogh 1970S04
    |         |--*F. biroi Balogh 1970S04
    |         `--F. latipilosa Mahunka 1982S04
    |    |--Licnocepheus Woolley 1969S04
    |    |    `--*L. reticulatus Woolley 1969S04
    |    |--Triteremella Kunst 1971 [incl. Afreremella Mahunka 1974]S04
    |    |    |--*T. kaszabi (Csiszár 1962) [=Eremella kaszabi; incl. Licneremaeus tuberculatus Paoli 1908 (n. d.)]S04
    |    |    `--T. luisiae (Mahunka 1974)S04 [=*Afreremella luisiaeBB92a]
    |    |--Archeremella Balogh & Mahunka 1974S04
    |    |    |--*A. leowae (Balogh & Mahunka 1974) [=Eremella (*Archeremella) leowae]S04
    |    |    `--A. africana Pérez-Íñigo 1982S04
    |    `--Eremella Berlese 1913 [incl. Proteremella Balogh 1959]S04
    |         |--*E. vestita Berlese 1913S04
    |         |--E. africana (Balogh 1966) [=Proteremella africana]S04
    |         |--E. ensifera Balogh & Mahunka 1968S04 [=E. (Proteremella) ensiferaBM74]
    |         |--E. induta Berlese 1913S04
    |         |--E. matildebellae Mahunka & Palacios-Vargas 1995S04
    |         `--E. pulchella (Balogh 1959)S04 [=*Proteremella pulchellaB65]
    |    |--Infernobates Karppinen & Poltavskaja 1990S04
    |    |    `--*I. citelli Karppinen & Poltavskaja 1990S04
    |    |--Kaszabobates Balogh 1972 [=Gobiella Balogh & Mahunka 1965 non Cienkowsky 1882]S04
    |    |    |--*K. kaszabi (Balogh & Mahunka 1965)S04 [=*Gobiella kaszabiBM65]
    |    |    |--K. helveticus Mahunka & Mahunka-Papp 2000S04
    |    |    `--K. olbiopolitanus Sergienko 1980S04
    |    |--Pantelozetes Grandjean 1953S04
    |    |    |--*P. paolii (Oudemans 1913)S04 (see below for synonymy)
    |    |    |--P. arcuata (Hammer 1958)BC63
    |    |    |--P. berlesei Fujikawa 1979S04
    |    |    |--P. pannonicus (Mahunka 1969) [=Oribella pannonica]S04
    |    |    `--P. ‘tricuspidatus’ (Djaparidze 1986) [=Amazoppia tricuspidata non Balogh & Mahunka 1969]S04
    |    |--Oribella Berlese 1908S04
    |    |    |--*O. pectinata (Michael 1885)S04 (see below for synonymy)
    |    |    |--O. adelaidae Golosova & Karppinen 1985S04
    |    |    |--O. canariensis Pérez-Íñigo 1986S04
    |    |    |--O. fujikawae Mahunka 1982S04
    |    |    `--O. matritensis Arillo, Bordel & Subías 1988S04
    |    |--Montizetes Kunst 1971S04
    |    |    |--*M. alpestris (Willmann 1929)S04 [=Xenillus alpestrisS04, Pantelozetes alpestrisBB92a]
    |    |    |--M. abulensis Pérez-Íñigo 1984S04
    |    |    |--M. arcticus (Pankov 1993) [=Gemmazetes arcticus]S04
    |    |    |--M. delamellatus Pérez-Íñigo 1991S04
    |    |    |--M. mongolicus (Balogh & Mahunka 1965)S04 [=Oribella mongolicaS04, Gemmazetes mongolicusB80]
    |    |    `--M. rarisetosus Bayartogtokh 1998S04
    |    `--Oribellopsis Kunst 1971S04
    |         |--*O. cavaticus (Kunst 1962)S04 [=Oribella cavaticaS04, Gemmazetes cavaticusB80]
    |         |--O. clavatus (Jacot 1937)S04 [=Oribella crosbyi clavataS04, Gemmazetes crosbyi clavatusB80]
    |         |--O. clavigerus (Mihelčič 1958) [=Oribella clavigera]S04
    |         |--O. etruscus (Bernini 1980) [=Gemmazetes etruscus]S04
    |         |--O. forsslundi (Moritz 1965) [=Oribella forsslundi]S04
    |         |--O. kushiroensis (Aoki 1992) [=Gemmazetes kushiroensis]S04
    |         `--O. tianshanensis (Wen 1989) [=Gemmazetes tianshanensis]S04
         |  i. s.: Scutoverticosus Kok 1968S04
         |           |--*S. arcanus Kok 1968S04
         |           `--S. insperatus Pérez-Íñigo & Peña 1996S04
         |         Ethiovertex Mahunka 1982S04
         |           |--*E. macrosetosus Mahunka 1982S04
         |           |--E. ‘bidactylus’ (Mahunka 1989) [=Scutovertex bidactylus non Coggi 1900]S04
         |           |--E. elisae Pérez-Íñigo & Peña 1996S04
         |           `--E. sculperens (Kok 1968) [=Scutovertex sculperens]S04
         |         Hypovertex Krivolutsky 1969S04
         |           |--*H. mirabilis Krivolutsky 1969S04
         |           |--H. africanus (Evans 1953) [=Scutovertex africanus]S04
         |           |--H. arcualis Hammer 1977S04
         |           |--H. hispanicusSC19
         |           |--H. lenticulatus Kahwash, Ruiz & Subías 1990S04
         |           |--H. saxicola Sitnikova 1975S04
         |           `--H. transversalis (Balogh & Csiszár 1963) [=Scutovertex transversalis]S04
         |         Arthrovertex Balogh 1970S04
         |           |--*A. segmentatus Balogh 1970S04
         |           |--A. baloghi Mahunka 1978S04
         |           |--A. bispinatus (Hammer 1961) [=Scutovertex bispinatus]S04
         |           |--A. hauseri Mahunka 1985S04
         |           |--A. hurdi (Woolley 1971) [=Oppia hurdi]NP93
         |           |--A. ilocanus Corpuz-Raros 1979S04
         |           |--A. leucaena Corpuz-Raros 1991S04
         |           `--A. xena Mahunka 1988S04
         |         Argentinovertex Fernández & Cleva 2001S04
         |           `--A. coineaui Fernández & Cleva 2002B-PEC05 [=Arthrovertex coineauiS04]
         `--+--Exochocepheus Wolley & Higgins 1968SK10, S04 [incl. Latovertex Mahunka 1987S04]
            |    |--*E. eremitus Wolley & Higgins 1968S04
            |    |--E. borealis (Rjabinin 1984) [=Hypovertex borealis]S04
            |    |--E. contiguus (Balogh & Csiszár 1963) [=Scutovertex contiguus]S04
            |    |--E. hungaricus (Mahunka 1987)S04 [=*Latovertex hungaricusBB92a]
            |    |--E. laticuspis (Balogh & Mahunka 1965) [=Scutovertex laticuspis]S04
            |    |--E. stenolamellatus (Golosova 1984) [=Hypovertex stenolamellatus]S04
            |    `--E. tuberculatus (Golosova 1984) [=Hypovertex tuberculatus]S04
            `--+--+--‘Scutovertex pictus Kunst 1959SK10, S04
               |  `--Lamellovertex Bernini 1976SK10, S04
               |       `--*L. caelatus (Berlese 1895) [=Scutovertex caelatus]S04
               `--Provertex Mihelčič 1959SK10, S04
                    |--*P. kuehnelti Mihelčič 1959S04
                    |--P. delamarei Travé 1963S04
                    |--P. forsslundi Krivolutsky 1969S04
                    |--P. mailloli Travé 1965S04
                    `--P. nepalensis Piffl 1971S04

*Oribella pectinata (Michael 1885)S04 [=Notaspis pectinataS04, Autogneta pectinataL87, Belba pectinataM98, Dameosoma pectinatumM98; incl. Damoeosoma crinitum Berlese 1916S04, B16, Xenillus limburgiensis Oudemans 1912S04]

*Pantelozetes paolii (Oudemans 1913)S04 [=Xenillus paoliiS04, Oribella paoliiB63; incl. Amazoppia caucasica Djaparidze 1986S04, Oribella dentata Mihelčič 1956S04, Pantelozetes dentatusBB92b, O. paolii multidentata Evans 1954S04, P. multidentataL96, P. paolii tuxeni Fujikawa 1979S04]

*Type species of generic name indicated


[B63] Balogh, J. 1963. Identification keys of Holarctic oribatid mites (Acari) families and genera. Acta Zoologica Academiae Scientiarum Hungaricae 9 (1–2): 1–60.

[B65] Balogh, J. 1965. A synopsis of the world oribatid (Acari) genera. Acta Zoologica Academiae Scientiarum Hungaricae 11 (1–2): 5–99.

[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.

[BC63] Balogh, J., & J. Csiszár. 1963. The zoological results of Gy. Topál’s collectings in south Argentina. 5. Oribatei (Acarina). Annales Historico-Naturales Musei Nationalis Hungarici 55: 463–485.

[BM65] Balogh, J., & S. Mahunka. 1965. Ergebnisse der zoologischen Forschungen von Dr. Z. Kaszab in der Mongolei 34. Acarina: Oribatei. Annales Historico-Naturales Musei Nationalis Hungarici 57: 451–465.

[BM74] Balogh, J., & S. Mahunka. 1974. Oribatid species (Acari) from Malaysian soils. Acta Zoologica Academiae Scientiarum Hungaricae 20 (3–4): 243–264.

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.

[B-PEC05] Behan-Pelletier, V. M., B. Eamer & M. Clayton. 2005. Dendroeremaeidae n. fam., from forest trees in western North America (Acari: Oribatida: Licneremaeoidea). Acarologia 46 (4): 321–339.

Behan-Pelletier, V. M., & D. E. Walter. 2007. Phylleremus n. gen., from leaves of deciduous trees in eastern Australia (Oribatida: Licneremaeoidea). Zootaxa 1386: 1–17.

[B16] Berlese, A. 1916. Centuria terza di Acari nuovi. Redia 12: 289–338.

[B80] Bernini, F. 1980. Notulae oribatologicae XXIV. Gli Acari Oribatei di alcune piccole grotte del Senese. Redia 63: 359–405.

[L87] Luxton, M. 1987. The oribatid mites (Acari: Cryptostigmata) of J. E. Hull. Journal of Natural History 21: 1273–1291.

[L96] Luxton, M. 1996. Oribatid mites of the British Isles: a check-list and notes on biogeography (Acari: Oribatida). Journal of Natural History 30 (6): 803–822.

[M98] Michael, A. D. 1898. Oribatidae. In: H. Lohmann (ed.) Das Tierreich. Eine Zusammenstellung und Kennzeichnung der rezenten Tierformen vol. 3. Acarina pp. 1–93. R. Friedländer und Sohn: Berlin.

[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.

[NP93] Norton, R. A., & G. O. Poinar. 1993. Reassessment and new records of oribatid mite fossils from Tertiary Neotropical amber. Acarologia 34 (1): 57–68.

[SC19] Schaefer, I., & T. Caruso. 2019. Oribatid mites show that soil food web complexity and close aboveground-belowground linkages emerged in the early Paleozoic. Communications Biology 2: 387.

[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.

Schuster, R. 1969. Die terrestrische Milbenfauna Südamerikas in zoogeographischer Sicht. In: Fittkau, E. J. (ed.) Biogeography and Ecology in South America vol. 2 pp. 741–763. Dr W. Junk N. V.: The Hague.

[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.

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