Licneremaeoidea

Micreremus sp., photographed by Scott Justis.


Belongs within: Circumdehiscentiae.
Contains: Bipassalozetes, Passalozetes, Licneremaeus, 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. It also includes 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).

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]
    |  i. s.: Glanderemaeus Balogh & Csiszár 1963 NB-P09, S04
    |           `--*G. hammerae Balogh & Csiszár 1963 S04
    |--Dendroeremaeidae NB-P09
    |--Passalozetidae S04
    |    |--Bipassalozetes S04
    |    `--Passalozetes S04
    |--Licneremaeidae S04
    |    |--Licneremaeus S04
    |    `--Huilicheremaeus Fernández, Marcangeli & Eguaras 1997 S04
    |         |--*H. michaii Fernández, Marcangeli & Eguaras 1997 S04
    |         `--H. atypicus (Mahunka 1984) [=Licneremaeus atypicus] S04
    |--Adhaesozetidae NB-P09
    |    |--Phylleremus NB-P09
    |    `--Adhaesozetes Hammer 1966 S04
    |         |--*A. barbarae Hammer 1966 S04
    |         `--A. polyphyllos Walter & Behan-Pelletier 1993 S04
    |--Charassobates Grandjean 1929 S94 [Charassobatidae NB-P09, Charassobatoidea]
    |    |--*C. cavernosus Grandjean 1929 S04
    |    |--C. baudi Mahunka 1984 S04
    |    |--C. incipatus Balogh & Mahunka 1974 S04
    |    |--C. minimus Balogh & Mahunka 1981 S04
    |    |--C. ornatus Balogh & Mahunka 1969 S04
    |    |--C. simplex Balogh & Mahunka 1969 S04
    |    `--C. tuberosus Balogh & Mahunka 1981 S04
    |--Lamellareidae S04
    |    |--Microlamellarea Coetzee 1987 S04
    |    |    `--*M. engelbrechti Coetzee 1987 S04
    |    |--Lamellarea Kok 1968 S04
    |    |    |--*L. ardua Kok 1968 S04
    |    |    |--L. digitata Kok 1968 S04
    |    |    |--L. forceps Kok 1968 S04
    |    |    `--L. quadrata Coetzee 1987 S04
    |    `--Tenuelamellarea Subías & Iturrondobeitia 1978 S04
    |         |--*T. hispanica Subías & Iturrondobeitia 1978 S04
    |         |--T. argentinensis Martínez, Velis et al. 1995 S04
    |         |--T. hawaiiensis Balogh 1985 S04
    |         |--T. inexpectata Balogh 1984 BB92b
    |         `--T. taprobanae (Oudemans 1915) [=Carabodes taprobanae] S04
    |--Eremellidae NB-P09
    |    |--Licnocepheus Woolley 1969 S04
    |    |    `--*L. reticulatus Woolley 1969 S04
    |    |--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 luisiae BB92a]
    |    |--Archeremella Balogh & Mahunka 1974 S04
    |    |    |--*A. leowae (Balogh & Mahunka 1974) [=Eremella (*Archeremella) leowae] S04
    |    |    `--A. africana Pérez-Íñigo 1982 S04
    |    `--Eremella Berlese 1913 [incl. Proteremella Balogh 1959] S04
    |         |--*E. vestita Berlese 1913 S04
    |         |--E. africana (Balogh 1966) [=Proteremella africana] S04
    |         |--E. ensifera Balogh & Mahunka 1968 S04
    |         |--E. induta Berlese 1913 S04
    |         |--E. matildebellae Mahunka & Palacios-Vargas 1995 S04
    |         `--E. pulchella (Balogh 1959) [=Proteremella pulchella] S04
    |--Micreremidae [Fenicheliidae] S04
    |    |--Mexiceremus Balogh & Balogh 1998 S04
    |    |    `--*M. rafaelinus (Mahunka & Palacios-Vargas 1995) [=Micreremus rafaelinus] S04
    |    |--Porofenichelia Mahunka 1985 NB-P09, S04
    |    |    `--*P. porosa Mahunka 1985 BB92a, S04 [=Fenichelia porosa S04]
    |    |--Phylloribatula Balogh & Mahunka 1978 S04
    |    |    |--*P. pulchella Balogh & Mahunka 1978 S04
    |    |    `--P. xanthoparmeliae Martínez & Palacios-Vargas 1998 S04
    |    |--Fenichelia Balogh 1970 S04
    |    |    |--*F. biroi Balogh 1970 S04
    |    |    `--F. latipilosa Mahunka 1982 S04
    |    `--Micreremus Berlese 1908 S04
    |         |--*M. brevipes (Michael 1888) (see below for synonymy) S04
    |         |--M. africanus Balogh 1963 S04
    |         |--M. faviger Mahunka 1983 S04
    |         |--M. florens Mahunka 1983 S04
    |         |--M. granulatus Balogh 1970 S04
    |         |--M. longisetus Mahunka 1986 S04
    |         |--M. macrofissura Hammer 1979 S04
    |         `--M. subglaber Ito 1982 S04
    |--Oribellidae S04
    |    |--Infernobates Karppinen & Poltavskaja 1990 S04
    |    |    `--*I. citelli Karppinen & Poltavskaja 1990 S04
    |    |--Kaszabobates Balogh 1972 [incl. Gobiella Balogh & Mahunka 1965 non Cienkowsky 1882] S04
    |    |    |--*K. kaszabi (Balogh & Mahunka 1965) [=Gobiella kaszabi] S04
    |    |    |--K. helveticus Mahunka & Mahunka-Papp 2000 S04
    |    |    `--K. olbiopolitanus Sergienko 1980 S04
    |    |--Pantelozetes Grandjean 1953 S04
    |    |    |--*P. paolii (Oudemans 1913) S04 (see below for synonymy)
    |    |    |--P. berlesei Fujikawa 1979 S04
    |    |    |--P. pannonicus (Mahunka 1969) [=Oribella pannonica] S04
    |    |    `--P. ‘tricuspidatus’ (Djaparidze 1986) [=Amazoppia tricuspidata non Balogh & Mahunka 1969] S04
    |    |--Oribella Berlese 1908 S04
    |    |    |--*O. pectinata (Michael 1885) (see below for synonymy) S04
    |    |    |--O. adelaidae Golosova & Karppinen 1985 S04
    |    |    |--O. canariensis Pérez-Íñigo 1986 S04
    |    |    |--O. fujikawae Mahunka 1982 S04
    |    |    |--O. matritensis Arillo, Bordel & Subías 1988 S04
    |    |    `--O. paolii L91
    |    |--Montizetes Kunst 1971 S04
    |    |    |--*M. alpestris (Willmann 1929) S04 [=Xenillus alpestris S04, Pantelozetes alpestris BB92a]
    |    |    |--M. abulensis Pérez-Íñigo 1984 S04
    |    |    |--M. arcticus (Pankov 1993) [=Gemmazetes arcticus] S04
    |    |    |--M. delamellatus Pérez-Íñigo 1991 S04
    |    |    |--M. mongolicus (Balogh & Mahunka 1965) [=Oribella mongolica] S04
    |    |    `--M. rarisetosus Bayartogtokh 1998 S04
    |    `--Oribellopsis Kunst 1971 S04
    |         |--*O. cavaticus (Kunst 1962) S04 [=Oribella cavatica S04, Gemmazetes cavatica BB92b]
    |         |--O. clavatus (Jacot 1937) [=Oribella crosbyi clavata] S04
    |         |--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
    `--Scutoverticidae S04
         |  i. s.: Scutoverticosus Kok 1968 S04
         |           |--*S. arcanus Kok 1968 S04
         |           `--S. insperatus Pérez-Íñigo & Peña 1996 S04
         |         Ethiovertex Mahunka 1982 S04
         |           |--*E. macrosetosus Mahunka 1982 S04
         |           |--E. ‘bidactylus’ (Mahunka 1989) [=Scutovertex bidactylus non Coggi 1900] S04
         |           |--E. elisae Pérez-Íñigo & Peña 1996 S04
         |           `--E. sculperens (Kok 1968) [=Scutovertex sculperens] S04
         |         Hypovertex Krivolutsky 1969 S04
         |           |--*H. mirabilis Krivolutsky 1969 S04
         |           |--H. africanus (Evans 1953) [=Scutovertex africanus] S04
         |           |--H. arcualis Hammer 1977 S04
         |           |--H. lenticulatus Kahwash, Ruiz & Subías 1990 S04
         |           |--H. saxicola Sitnikova 1975 S04
         |           `--H. transversalis (Balogh & Csiszár 1963) [=Scutovertex transversalis] S04
         |         Arthrovertex Balogh 1970 [incl. Argentinovertex Fernández & Cleva 2001] S04
         |           |--*A. segmentatus Balogh 1970 S04
         |           |--A. baloghi Mahunka 1978 S04
         |           |--A. bispinatus (Hammer 1961) [=Scutovertex bispinatus] S04
         |           |--A. coineaui (Fernández & Cleva 2001) [=Argentinovertex coineaui] S04
         |           |--A. hauseri Mahunka 1985 S04
         |           |--A. ilocanus Corpuz-Raros 1979 S04
         |           |--A. leucaena Corpuz-Raros 1991 S04
         |           `--A. xena Mahunka 1988 S04
         |--Scutovertex SK10
         `--+--Exochocepheus Wolley & Higgins 1968 SK10, S04 [incl. Latovertex Mahunka 1987 S04]
            |    |--*E. eremitus Wolley & Higgins 1968 S04
            |    |--E. borealis (Rjabinin 1984) [=Hypovertex borealis] S04
            |    |--E. contiguus (Balogh & Csiszár 1963) [=Scutovertex contiguus] S04
            |    |--E. hungaricus (Mahunka 1987) S04 [=*Latovertex hungaricus BB92a]
            |    |--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 1959 SK10, S04
               |  `--Lamellovertex Bernini 1976 SK10, S04
               |       `--*L. caelatus (Berlese 1895) [=Scutovertex caelatus] S04
               `--Provertex Mihelčič 1959 SK10, S04
                    |--*P. kuehnelti Mihelčič 1959 S04
                    |--P. delamarei Travé 1963 S04
                    |--P. forsslundi Krivolutsky 1969 S04
                    |--P. mailloli Travé 1965 S04
                    `--P. nepalensis Piffl 1971 S04

*Micreremus brevipes (Michael 1888) [=Eremaeus brevipes; incl. M. gracilior Willmann 1931, E. minimus Berlese 1891, E. minutipes Berlese 1902] S04

*Oribella pectinata (Michael 1885) [=Notaspis pectinata; incl. Dameosoma crinitum Berlese 1916, Xenillus limburgiensis Oudemans 1912] S04

*Pantelozetes paolii (Oudemans 1913) S04 [=Xenillus paolii S04; incl. Amazoppia caucasica Djaparidze 1986 S04, Oribella dentata Mihelčič 1956 S04, Pantelozetes dentatus BB92b, O. paolii multidentata Evans 1954 S04, P. paolii tuxeni Fujikawa 1979 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.

[L91] Luxton, M. 1991. Seasonal and spatial variation in food intake by the oribatid mites of beech woodland soil. In: Schuster, R., & P. W. Murphy (eds) The Acari: Reproduction, development and life-history strategies pp. 459–471. Chapman & Hall: London.

[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.

[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.

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