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: Brachypylina.

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. Cepheus latus is a borer in birch twigs (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 S04
    |--Protocepheus Jacot 1928 S04
    |    `--*P. hericius (Michael 1887) S04 [=Tegeocranus hericius S04, Cepheus hericius BN82]
    |--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 S04 (see below for synonymy)
    |    `--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 PL17, S04 [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 BB92a]
    |    |--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 [=Tegeocranus latus BN82]
         |--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
         |--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 G36; 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

*Oribatodes mirabilis Banks 1895 S04 [=Cepheus mirabilis BN82; incl. C. lamellatus Banks 1906 S04, Tegeocranus longisetus Berlese 1910 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

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

[BN82] Bernini, F., & R. Nannelli. 1982. Notulae oribatologicae XXVI. Contribution to the knowledge of the genus Cepheus (Oribatida, Acarida) in Italy. Redia 45: 155–188.

[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 2–7.

[G36] Grandjean, F. 1936. Observations sur les oribates (10e série). Bulletin du Muséum, 2e série, 8 (3): 246–253.

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.

[PL17] Pachl, P., A. C. Lindl, A. Krause, S. Scheu, I. Schaefer & M. Maraun. 2017. The tropics as an ancient cradle of oribatid mite diversity. Acarologia 57 (2): 309–322.

[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.

Last updated: 30 June 2017.

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.

Licneremaeoidea

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).

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 2005 B-PEC05
    |    `--D. foveolatus Behan-Pelletier, Eamer & Clayton 2005 B-PEC05
    |--Passalozetidae NB-P09
    |    |--Bipassalozetes NB-P09
    |    `--Passalozetes NB-P09
    |--Licneremaeidae NB-P09
    |    |--Licneremaeus NB-P09
    |    `--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 S04 [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 NB-P09
    |    |--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) S04 [=*Proteremella pulchella B65]
    |--Micreremidae [Fenicheliidae] NB-P09
    |    |--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
    |    |--Glanderemaeus Balogh & Csiszár 1963 B-PEC05, S04
    |    |    `--*G. hammerae Balogh & Csiszár 1963 S04
    |    `--Micreremus 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) S04 (see below for synonymy)
    |    |    |--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 NB-P09
         |  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 S04
         |           |--*A. segmentatus Balogh 1970 S04
         |           |--A. baloghi Mahunka 1978 S04
         |           |--A. bispinatus (Hammer 1961) [=Scutovertex bispinatus] S04
         |           |--A. hauseri Mahunka 1985 S04
         |           |--A. ilocanus Corpuz-Raros 1979 S04
         |           |--A. leucaena Corpuz-Raros 1991 S04
         |           `--A. xena Mahunka 1988 S04
         |         Argentinovertex Fernández & Cleva 2001 S04
         |           `--A. coineaui Fernández & Cleva 2002 B-PEC05 [=Arthrovertex coineaui 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

*Oribella pectinata (Michael 1885) S04 [=Notaspis pectinata S04; incl. Damoeosoma crinitum Berlese 1916 S04, B16, 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

[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.

[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.

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

[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.

Last updated: 6 May 2019.

Scheloribatidae

Dometorina plantivaga, copyright R. Penttinen.


Belongs within: Oripodoidea.
Contains: Perscheloribates (Perscheloribates), Scheloribates (Scheloribates), Scheloribates (Topobates).

The Scheloribatidae are a diverse family of oribatid mites with the octotaxic system commonly developed as sacculi, and immovable pteromorphs usually present. The Tongan species Planobates circumalatus has a single pair of setae on the genital plates; other genera have three to five pairs. The notogastral setae are often reduced with only the setal alveoli visible; in the African genus Similobates, a single pair p2 remains developed. Similobates also has five pairs of genital setae. The rostrum bears a pair of incisions in Rhabdoribates siamensis and Euscheloribates subgenus Nanobates (Balogh & Balogh 1992). Members of the genus Scheloribates are among the most abundant mites in soil and litter, including in anthropogenic habitats (Norton & Behan-Pelletier 2009).

Characters (from Norton & Behan-Pelletier 2009): Length 200-650 µm. Prodorsum with prolamella often present; sublamella present; tutorium and pleural carinae absent. Prodorsum curving over gnathosoma, covering chelicerae dorsally; rostrum rounded. Chelicerae with two setae. Bothridium inserted anterior to notogastral margin. Distal eupathidia on palptarsus in same plane. Notogaster with 2-4 pairs of porose areas or saccules. Pteromorphs or humeral projections usually present. Posterior notogastral tectum absent. Sternal furrow extending from genital plates anteriorly on coxisternum often present. 1-4 pairs of genital setae. Pedotectum II present. Femora III-IV each with one porose area; solenidion σ present on genu III.

<==Scheloribatidae [Parapirnodidae, Scheloribatinae, Scheloribatini]
    |--Planobates Hammer 1973 S04
    |    `--*P. circumalatus Hammer 1973 S04
    |--Rhabdoribates Aoki 1967 S04
    |    `--*R. siamensis Aoki 1967 S04
    |--Fissurobates Balogh & Mahunka 1969 S04
    |    `--*F. spectabilis Balogh & Mahunka 1969 S04
    |--Striatobates Hammer 1973 S04
    |    `--*S. tuberculatus Hammer 1973 S04
    |--Samoabates Hammer 1973 S04
    |    `--*S. acutirostrum Hammer 1973 S04
    |--Scheloribatoides Mahunka 1988 S04
    |    `--*S. lamellatus Mahunka 1988 S04
    |--Pachygena Hammer 1972 S04
    |    `--*P. falcata Hammer 1972 S04
    |--Coronibatula Mahunka 1988 S04
    |    `--*C. lienhardi Mahunka 1988 S04
    |--Cosmobates Balogh 1959 S04
    |    `--*C. tunicatus Balogh 1959 S04
    |--Pontiobates Luxton 1989 S04
    |    `--*P. denigratus Luxton 1989 S04
    |--Similobates Mahunka 1982 S04
    |    |--*S. demetororum Mahunka 1982 S04
    |    `--S. deficiens (Wallwork 1977) [=Scheloribates deficiens] S04
    |--Aellenobates Mahunka 1978 S04
    |    |--*A. cryptacus Mahunka 1978 [=Grandjeanobates cryptacus] EA14
    |    `--A. sculpturatus Mahunka 1988 [=Grandjeanobates sculpturatus] EA14
    |--Nannerlia Coetzer 1968 S04
    |    |--*N. longiqua Coetzer 1968 S04
    |    |--N. bombretensis Hammer 1977 S04
    |    `--M. elongatissima Mahunka 1984 S04
    |--Annobonzetes Pérez-Íñigo 1983 S04
    |    |--*A. sphaericus Pérez-Íñigo 1983 S04
    |    |--A. latissimus (Berlese 1916) [=Podoribates latissimus] S04
    |    `--A. latus (Warburton 1912) [=Oribata lata] S04
    |--Gerloubia Coetzer 1968 [incl. Fuerteventuria Pérez-Íñigo & Peña 1996] S04
    |    |--*G. bicuspidata (Hammer 1958) [=Eporibatula bicuspidata] S04
    |    |--G. chinensis Aoki & Yamamoto 2000 S04
    |    `--G. hispanica Subías 1977 (see below for synonymy) S04
    |--Fijibates Hammer 1971 S04
    |    |--*F. rostratus Hammer 1971 S04
    |    |--F. aelleni (Mahunka 1988) [=Scheloribates aelleni] S04
    |    |--F. anterostratus Corpuz-Raros 1980 S04
    |    `--F. dlouhyi (Mahunka 1984) [=Scheloribates dlouhyi] S04
    |--Parapirnodus Balogh & Mahunka 1968 NB-P09, S04
    |    |--*P. longus Balogh & Mahunka 1968 S04
    |    |--P. coniferinus Behan-Pelletier, Clayton & Humble 2001 S04
    |    |--P. hexaporosus Behan-Pelletier, Clayton & Humble 2001 S04
    |    `--P. prosopis Martínez, Fernández & Monetti 1996 S04
    |--Perscheloribates Hammer 1973 S04
    |    |--P. (Perscheloribates) S04
    |    |--P. (Makischeloribates Corpuz-Raros 1980) S04
    |    |    `--P. (*M.) tripartitus (Corpuz-Raros 1980) [=*Makischeloribates tripartitus] S04
    |    `--P. (Oxyscheloribates Balogh & Balogh 1990) S04
    |         `--P. (*O.) aculeatus (Hammer 1961) [=Scheloribates aculeatus] S04
    |--Hammerabates Balogh 1970 [incl. Otaheitea Hammer 1972, Yoronoribates Aoki 1987] S04
    |    |--*H. trisetosus Balogh 1970 S04
    |    |--H. elongatus Tseng 1984 S04
    |    |--H. minusculus (Aoki 1987) S04 [=*Yoronoribates minusculus BB92]
    |    |--H. nasalis Mahunka 1983 S04
    |    |--H. rotundus Mahunka 1983 S04
    |    `--H. sulcatus (Hammer 1972) S04 [=*Otaheites sulcata BB92]
    |--Megascheloribates Lee & Pajak 1990 LP90
    |    |--*M. microsetosus Lee & Pajak 1990 CH98, LP90 (see below for synonymy)
    |    |--M. anzacensis (Hammer 1967) [=Scheloribates anzacensis] LP90
    |    |--M. calcaratus Lee & Pajak 1990 LP90 (see below for synonymy)
    |    |--‘Scheloribates’ iteratus Subías 2004 S04 (see below for synonymy)
    |    |--M. maximus (Balogh 1962) LP90
    |    `--M. rostrodentatus (Hammer 1977) [=Scheloribates rostrodentatus] LP90
    |--Muliercula Coetzer 1968 S04
    |    |--*M. muliercula Coetzer 1968 S04
    |    |--M. bilineata Mahunka 1986 S04
    |    |--M. discrepans (Balogh 1959) [=Scheloribates discrepans] S04
    |    |--M. inexpectata Badejo, Woas & Beck 2002 S04
    |    |--M. longisacculus (Mahunka 1984) [=Incabates longisacculus] S04
    |    |--M. ngoyensis Coetzer 1968 S04
    |    `--M. spora Coetzer 1968 S04
    |--Scheloribates Berlese 1908 S04
    |    |  i. s.: S. areatus Sellnick 1931 S93
    |    |         S. azumaensis NB-P09
    |    |         S. madrasensis H01
    |    |         S. setatus Sellnick 1931 S93
    |    |--S. (Scheloribates) S04
    |    |--S. (Grandjeanobates Ramsay 1967) EA14
    |    |    |--S. (*G.) novazelandicus Hammer 1967 [=*Grandjeanobates novazelandicus] EA14
    |    |    `--‘Grandjeanobates’ australis Ramsay 1967 S04
    |    `--S. (Topobates) EA14
    |--Euscheloribates Kunst 1958 S04
    |    |--E. (Euscheloribates) [incl. Corynobates Balogh & Balogh 1992, Tribates Mahunka 1978] S04
    |    |    |--*E. (E.) samsinaki Kunst 1958 S04
    |    |    |--E. (E.) clavisetus Mahunka 1988 S04 [=*Corynobates clavisetus BB92]
    |    |    |--E. (E.) cryptus (Mahunka 1978) [=Tribates cryptus] S04
    |    |    |--E. (E.) mixtus (Mahunka 1978) S04 [=*Tribates mixtus BB92]
    |    |    `--E. (E.) pembertoni (Jacot 1934) [=Protoschelobates pembertoni] S04
    |    `--E. (Nanobates Balogh & Balogh 1984) S04
    |         |--E. (*N.) fenicheli (Balogh 1970) [=Birobates fenicheli] S04
    |         `--E. (N.) clavatus (Mahunka 1988) [=Nanobates clavatus] S04
    `--Dometorina Grandjean 1951 NB-P09, S04 [incl. Cryptozetes Hammer 1962 S04]
         |--*D. plantivaga (Berlese 1895) S04 [=Oribatula plantivaga S04, O. (Hemileius) plantivaga B16]
         |    |--D. p. plantivaga S04
         |    |--D. p. brasiliana Pérez-Íñigo & Pérez-Íñigo 1993 S04
         |    |--D. p. insularis Travé 1969 S04
         |    `--D. p. saxicola Grandjean 1951 S04
         |--D. chilensis (Hammer 1962) S04 [=*Cryptozetes chilensis BB92]
         |--D. crystallina Tseng 1984 S04
         |--D. elongata Balogh & Csiszár 1963 S04
         |--D. inflata (Berlese 1916) [=Oribatula (Hemileius) inflata] S04
         |--D. lata Mahunka 1986 S04
         |--D. limpida Tseng 1984 S04
         |--D. marionensis Pletzen & Kok 1971 S04
         |--D. praedatoria Wu, Xin & Aoki 1986 S04
         |--D. rostrata Luxton 1993 S04
         |--D. sicata Mahunka 1983 S04
         |--D. suramericana (Hammer 1958) S07
         |--D. taiwanica Tseng 1984 S04
         |--D. tropica (Balogh 1958) [=Hemileius tropicus] S04
         |--D. tuberculata Aoki 1984 S04
         `--D. usnea (Norton & Palacios-Vargas 1987) [=Cryptozetes usnea] S04

Gerloubia hispanica Subías 1977 [incl. Liebstadia ahumerata Mahunka 1987, Fuerteventuria mirabilis Pérez-Íñigo & Peña 1996] S04

Megascheloribates calcaratus Lee & Pajak 1990 nec Scheloribates fimbriatus calcaratus Jacot 1934 nec S. calcaratus Wallwork 1977 (not preoc. if in dif. gen.) LP90 [=Scheloribates praeoccupatus Subías 2004 S04]

*Megascheloribates microsetosus Lee & Pajak 1990 CH98, LP90 [=Scheloribates microsetosus S04; incl. M. pajaki Balogh & Balogh 2002 S04]

‘Scheloribates’ iteratus Subías 2004 S04 [=S. microclava Balogh 1962 non Hammer 1961 S04, Megascehloribates microclava LP90]

*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.

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

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

[EA14] Ermilov, S. G., & A. E. Anichkin. 2014. A new species of Scheloribates (Scheloribates) from Vietnam, with notes on taxonomic status of some taxa in Scheloribatidae (Acari, Oribatida). International Journal of Acarology 40 (1): 109–116.

[H01] Haq, M. A. 2001. Potential oribatid mite vectors of cestode parasites in Kerala, India. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 569–575. CSIRO Publishing: Melbourne.

[LP90] Lee, D. C., & G. A. Pajak. 1990. Scheloribates Berlese and Megascheloribates gen. nov. from southeastern Australia, with comments on Scheloribatidae (Acarida: Cryptostigmata: Oriopodoidea). Invertebrate Taxonomy 4: 205–246.

[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.

[S07] Schatz, H. 2007. Biogeography of oribatid mites (Acari, Oribatida) from the Cordillera de Talamanca, Costa Rica and Panama. In: Morales-Malacara, J. B., V. M. Behan-Pelletier, E. Ueckermann, T. M. Pérez, E. G. Estrada-Venegas & M. Badii (eds) Acarology XI: Proceedings of the International Congress pp. 151–167. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[S93] Selden, P. A. 1993. Arthropoda (Aglaspidida, Pycnogonida and Chelicerata). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 297–320. Chapman & Hall: London.

[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.

Last updated: 24 February 2019.