Holonota

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

Belongs within: Oribatida.
Contains: Epilohmanniidae, Ptyctima, Perlohmanniidae, Hermanniidae, Astigmatina, Malaconothridae, Trhypochthoniidae, Nanhermanniidae, Crotoniidae, Nothridae, Microzetidae, Polypterozetoidea, Amerobelbidae, Damaeolidae, Eremobelba, Ameridae, Ctenobelba, Eremulidae, Basilobelbidae, Zetorchestidae, Cepheidae, Thyrisomidae, Peloppiidae, Plateremaeoidea, Eutegaeidae, Anderemaeidae, Carabodoidea, Rhynchoribatidae, Oppioidea, Eremaeoidea, Neoliodidae, Hermannielloidea, Cymbaeremaeidae, Gustavioidea, Damaeidae.

The Desmonomata have been recognised as a group of oribatid mites with a holoid body form, in which epimera II and III are fused and the sejugal furrow is absent ventrally, in combination with an undivided notogaster and legs in which the genua are not reduced to a knee-like segment. With the exception of the Hermanniidae, most Desmonomata in this sense also have a macropyline venter, in which the large genital and anal plates occupy almost the entire venter posterior to the epimeres. However, phylogenetic analyses have established that the Desmonomata sensu stricto is paraphyletic with regard to the Brachypylina (or Circumdehiscentiae; ‘higher’ oribatid mites) and the Astigmatina (previously recognised as a separate group from the Oribatida), both of which share the holoid body form. The dichoid genus Collohmannia, including large mites characterised a dorsally and ventrally convex hysterosoma and flagellate and particularly elongate setae d2, h2 and p1, was also placed within the desmonomatan clade by Pachl et al. (2017). Many Desmonomata, including the families Malaconothridae, Camisiidae, Trhypochthoniidae and the genus Nothrus, are parthenogenetic, though the Crotoniidae and the other genera of Nothridae are sexual (Norton & Behan-Pelletier 2009).

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

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

Mite-in-a-box
Published 22 September 2008
Nothrus truncatus, a member of the holoid Desmonomata, from Agriculture and Agri-Food Canada.

Mites are pretty remarkable creatures. I don’t know if any other group of animals can rival mites for ecological diversity. There are mites burrowing in the leaf litter of forests, there are mites living in the sediment at the bottom of the sea, there are mites living off the secretions in your hair follicles, there are mites that live as parasites of other animals. Whatever freakish thing you can think of an animal doing, odds are that there is a mite doing it right now. Mites also include one of the few groups of terrestrial arthropods to have developed a mineralised exoskeleton—the oribatids or beetle mites. The subject of this page is one of the subgroups of oribatids, the Holonota.

The Holonota include the most heavily armoured of the oribatids, with most of the body encased in hardened plates. Morphologically, Holonota are distinguished from other oribatids by having the entire body covered by two dorsal plates, with the division between the two plates between the positions of the second and third pairs of legs. Technically speaking, various Holonota may be dichoid, holoid or ptychoid (Norton 2001). In dichoid forms, a non-hardened zone runs around the body between the second and third pairs of legs, allowing the body to bend at that position. Holoid forms only have this articulation dorsally, with the ventral surface fused to a solid plate. Ptychoid forms, on the other hand, have reduced the ventral hardening and are actually able to withdraw the legs and close the anterior dorsal plate over them—the mites’ answer to ostracods.

Norton (2001) suggests that the development of heavy armour in the oribatids may be related to their lifestyle. Most oribatids are long-lived (at least for mites) and have low reproductive rates, a situation that may result from their usual diet of low-nutrient decaying vegetation and fungi. Slow growth and replacement rates may result in the selective favouring of features that extend the life expectancies of individuals.

The ptychoid mite Atropacarus striculus (Phthiracaridae), showing how the legs can be withdrawn and covered over, from Oribatid Mites of Alberta.

Morphologically, Holonota have been divided between three groups, the Mixonomata, Desmonomata and Circumdehiscentiae (Maraun et al. 2004). The Mixonomata include dichoid and ptychoid forms, and have been suggested to be paraphyletic to a holoid clade formed by the other two taxa. In turn, the Desmonomata are probably paraphyletic with regards to the Circumdehiscentiae. The holoid Circumdehiscentiae are one of most speciose groups of oribatids, and show the highest degree of plate fusion, with the armour of almost the entire underside fused with the anterior dorsal plate (Norton 2001). Molecular analyses, in contrast, have been divided in their support for this arrangement. Maraun et al. (2004) failed to support the morphological view, and did not even recover monophyly for the Holonota as a whole or for the Circumdehiscentiae. More recently, however, the morphological phylogeny with monophyletic Holonota and serially paraphyletic Mixonomata and Desmonomata was supported by the results of Domes et al. (2007).

In light of the repeated evolution of ever-greater degrees of sclerotisation within the Holonota, it might seem surprising if one lineage was to do a complete volte-face and lose all trace of armour, but exactly this possibility has been suggested (Norton 2001). The Astigmata are a lineage of mites related to the Oribatida, but ecologically distinct. While oribatids are armoured, slow-living, litter feeders, astigmatans are unarmoured, fast-breeding and mostly live in close association with other animals, often as parasites. Astigmata include such luminaries as Sarcoptes scabiei, the skin-burrowing monstrosity that causes scabies*. Despite these differences, it has been suggested that Astigmata are actually derived from oribatids through paedomorphosis (retention of juvenile characters as adults)—and specifically from Desmonomata, one of the most heavily armoured groups of oribatids. However, support for Astigmata as derived desmonomates remains equivocal. While supported by some morphological characters and gland chemistry, the suggestion has not garnered molecular support. Mauran et al. (2004) supported an oribatid ancestry for Astigmata, though not necessarily from Desmonomata (they also only included a single species of Astigmata in their analysis). Domes et al. (2007), analysing a larger selection of astigmates, rejected a position for Astigmata within Oribatida.

*Offhand, if you had to invent a name for a revolting skin condition, could you ever come up with a more appropriate-sounding term than ‘scabies’?

Systematics of Holonota
Holonota [Mixonomata]N01
|--+--EpilohmanniidaePL17
| `--PtyctimaPL17
`--+--Nehypochthonius Norton & Metz 1980S-TK22, S04 [Nehypochthoniidae, Nehypochthonoidea]
| |--*N. porosus Norton & Metz 1980S04
| `--N. yanoi Aoki 2002S04
`--+--PerlohmanniidaePL17
`--Desmonomata (see below for synonymy)PL17
|--+--HermanniidaeDW10
| `--AstigmatinaDW10
`--+--MalaconothridaeS-TK22
`--+--TrhypochthoniidaeS-TK22
`--+--+--+--NanhermanniidaeS-TK22
| | `--CrotoniidaeSC19
| `--+--NothridaeS-TK22
| `--Collohmannia Sellnick 1922S-TK22, S04 (see below for synonymy)
| |--*C. gigantea Sellnick 1922 [incl. C. nova Sellnick 1932]S04
| |--C. asiatica Krivolutsky & Christov 1970S04
| |--C. johnstoni Norton & Sidorchuk 2014PH22
| `--C. schusteriSC19
`--Brachypylina (see below for synonymy)S-TK22
| i. s.: MicrozetidaeS04
| PolypterozetoideaS04
| AmerobelbidaeS04
| DamaeolidaeS04
| EremobelbaS04
| AmeridaeS04
| CtenobelbaS04
| EremulidaeS04
| BasilobelbidaeS04
| StaurobatidaeS04
| |--Stauroma Grandjean 1966S04
| | `--*S. cephalotum Grandjean 1966S04
| `--Staurobates Grandjean 1966S04
| `--*S. schusteri Grandjean 1966S04
| |--S. s. schusteriS04
| `--S. s. cordobensis Balogh & Mahunka 1968S04
| Oxyamerus Aoki 1965NB-P09, S04 [OxyameridaeS04, Oxyamerinae]
| |--*O. spathulatus Aoki 1965S04
| |--O. aokii Balogh 1968S04
| |--O. hauserorum Mahunka 1987S04
| |--O. hyalinus Hammer 1979S04
| |--O. latirostris Balogh 1968S04
| `--O. truncatus Hammer 1979S04
| SpinozetidaeNB-P09
| |--Iberoppia Pérez-Íñigo 1986S04
| | `--*I. paradoxa Pérez-Íñigo 1986S04
| |--Grypoceramerus Suzuki & Aoki 1970S04
| | `--*G. acutus Suzuki & Aoki 1970S04
| `--Spinozetes Piffl 1966 (see below for synonymy)S04
| |--*S. inexspectatus Piffl 1966S04
| `--S. pectinatus (Kulijev 1967) [=Mirus pectinatus]S04
| HungarobelbidaeS04
| |--Hungarobelba Balogh 1943S04
| | |--*H. visnyai (Balogh 1938) [=Belba visnyai]S04
| | |--H. baloghi Bulanova-Zachvatkina 1967S04
| | `--H. pyrenaica Miko & Travé 1996S04
| `--Costeremus Aoki 1970S04
| |--*C. ornatus Aoki 1970S04
| |--C. barbatus Choi 1997S04
| |--C. cornutus Wang 1996S04
| `--C. yezoensis Fujikawa & Fujita 1985S04
| MultoribulidaeS04
| |--Peloptoribula Mahunka 1984S04
| | `--*P. spinulosa Mahunka 1984S04
| `--Multoribula Balogh & Mahunka 1966S04
| |--*M. multipunctata Balogh & Mahunka 1966S04
| `--M. suramericana Balogh & Mahunka 1977S04
| Kodiakella Hammer 1967S04 [KodiakellidaeNB-P09]
| |--*K. lutea Hammer 1967S04
| `--K. dimorpha Pérez-Íñigo & Subías 1978S04
| Pterobates Balogh & Mahunka 1977 [Pterobatidae, Pterobatinae]S04
| `--*P. incertus Balogh & Mahunka 1977S04
| CerocepheidaeS04
| |--Cerocepheus Trägårdh 1931S04
| | `--*C. mirabilis Trägårdh 1931S04
| |--Dicrotegaeus Luxton 1988S04
| | `--*D. mirabilis Luxton 1988S04
| `--Bornebuschia Hammer 1966S04
| |--*B. peculiaris Hammer 1966S04
| `--B. binodosa Luxton 1988S04
| NosybeidaeNB-P09
| |--Lamellocepheus Balogh 1961 (see below for synonymy)S04
| | |--*L. personatus (Berlese 1910)S04 (see below for synonymy)
| | `--L. genavensis (Mahunka 1993) [=Nosybea genavensis]S04
| `--Topalia Balogh & Csiszár 1963S04
| |--*T. problematica Balogh & Csiszár 1963S04
| |--T. africana Mahunka 1985S04
| |--T. clavata Hammer 1966S04
| |--T. granulata Hammer 1966S04
| `--T. velata Hammer 1966S04
| ZetorchestidaeSK10
| CepheidaeSC19
| ThyrisomidaeSC19
| PeloppiidaeSC19
|--PlateremaeoideaS-TK22
`--+--+--EutegaeidaePL17
| `--+--+--AnderemaeidaePL17
| | `--CarabodoideaS-TK22
| `--+--RhynchoribatidaePL17
| `--OppioideaSC19
`--+--+--EremaeoideaS-TK22
| `--+--NeoliodidaeS-TK22
| `--HermannielloideaSK10
`--+--+--CymbaeremaeidaeSC19
| `--GustavioideaS-TK22
`--+--Caleremaeidae [Caleremaeoidea]NB-P09
| |--Luxtoneremaeus Balogh & Balogh 1922S04
| | `--*L. forsteri (Balogh & Balogh 1985) [=Anderemaeus forsteri]S04
| |--Veloppia Hammer 1955S04
| | |--*V. pulchra Hammer 1955S04
| | `--V. kananaskis Norton 1978S04
| `--Caleremaeus Berlese 1910SC19, S04
| |--*C. monilipes (Michael 1882)S04 (see below for synonymy)
| |--C. divisus Mihelčič 1952S04
| `--C. retractus (Banks 1947) [=Carabodoides retractus]S04
`--DamaeidaeSC19

Brachypylina [Amerobelboidea, Ameroidea, Apheredermata, Apterogasterina, Brachypylides, Carinotae, Cepheoidea, Circumdehiscentiae, Eremuloidea, Euoribatida, Eupheredermata, Gymnonota, Holosomatina, Liacaroidea, Notaspidinae, Planofissurae, Poronota, Poronotae, Pterogasterina, Pycnonota, Pycnonoticae]SK10

*Caleremaeus monilipes (Michael 1882)S04 [=Damaeus monilipesS04, Cymbaeremaeus monilipesM98, Notaspis monilipesBB92]

Collohmannia Sellnick 1922S-TK22, S04 [incl. Phthiracaroides Storkán 1923B65; Collohmanniidae, Collohmannioidea]

Desmonomata [Comalida, Desmonomatides, Holosomata, Holosomatina, Malaconothroidea, Nothrina, Nothrinae, Nothronata, Trhypochthonioidea]PL17

Lamellocepheus Balogh 1961 [incl. Nosybea Mahunka 1993, Tessacarus Grandjean 1962]S04

*Lamellocepheus personatus (Berlese 1910)S04 [=Tectocepheus personatusS04, Tegeocranus personatusBB92; incl. Lamellocepheus ambitus Kulijev 1966S04]

Spinozetes Piffl 1966 [incl. Mirus Kulijev 1967 nec Albers 1850 nec Saulcy 1877]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.

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

[DW10] Dabert, M., W. Witalinski, A. Kazmierski, Z. Olszanowski & J. Dabert. 2010. Molecular phylogeny of acariform mites (Acari, Arachnida): strong conflict between phylogenetic signal and long-branch attraction artifacts. Molecular Phylogenetics and Evolution 56 (1): 222–241.

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.

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

[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.

[N01] Norton, R. A. 2001. Systematic relationships of Nothrolohmanniidae, and the evolutionary plasticity of body form in Enarthronota (Acari: Oribatida). 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. 58–75. CSIRO Publishing: Melbourne.

[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.

[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.

[PH22] Pfingstl, T., S. F. Hiruta, I. Bardel-Kahr, Y. Obae & S. Shimano. 2022. Another mite species discovered via social media—Ameronothrus retweet sp. nov. (Acari, Oribatida) from Japanese coasts, exhibiting an interesting sexual dimorphism. International Journal of Acarology 48 (4–5): 348–358.

[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.

[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.

[S-TK22] Szudarek-Trepto, N., A. Kaźmierski, A. Skoracka, M. Lewandowski & J. Dabert. 2022. Molecular phylogeny supports the monophyly of the mite supercohort Eupodides (Acariformes: Trombidiformes) and greatly coincides with traditional morphological definition of the taxon. Annales Zoologici 72 (4): 757–786.

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