Eleutherengonides

Hoplocheylus sp., from Walter et al. (2009).

Belongs within: Anystina.
Contains: Cryptognathus, Eupalopsellidae, Pterygosomatidae, Eustigmaeus, Stigmaeus, Agistemus, Zetzellia, Tetranychoidea, Raphignathus, Caligonellidae, Neophyllobius, Cheyletoidea, Dolichocyboidea, Pygmephoroidea, Pyemotoidea, Tarsonemidae, Podapolipidae.

The Eleutherengonides are a group of mites in which the chelicerae have the fixed digit reduced or absent, with the movable digit hook-like or stylet;like and inserted apically or subapically on the closely contiguous or fused cheliceral bases. A palpal thumb-claw process is commonly present and the solenidia on leg tarsus I, if present, are erect (Walter et al. 2009).

Members of the Eleutherengonides are divided between the Raphignathina and Heterostigmata. The Raphignathina typically have stigmata opening between the bases of the chelicerae and lack bothridia on the prodorsum. The stigmata, if present, are associated with simple to highly convoluted peritremes. Members of the group vary from heavily armoured taxa such as the Homocaligidae and Cryptognathus to almost completely non-sclerotised taxa such as Barbutia and the Caligonellidae. Most Raphignathina other than Tetranychoidea lack tenent hairs on the leg claws (albeit possessing them on the associated empodia) but the claws are haired in the soil- and litter-dwelling Barbutia. Both stigmata and peritremes are absent in Homocaligidae, Mecognatha, Eupalopsellidae and Stigmaeidae. Most raphignathoids have two nymphal instars between larva and adult but a tritonymph is also present in Xenocalligonellididae (Walter et al. 2009).

Barbutia species are relatively elongate, soft-bodied mites known from seasonally arid regions of North America and Australia. The Homocaligidae are hemispherical mites found in association with fresh water. Camerobiidae are oval to subcircular mites with very long legs, with the tibia of each leg about three times the length of the tarsus. The Dasythyreidae and Xenocalligonellididae are neotrichous with long setae radiating outwards in a pincushion-like array many times the diameter of the body (Walter et al. 2009).

Eleutherengonides [Eleutherengona, Eleutherengonina]
    |--Raphignathina [Raphignathae, Raphignathoidea]ZF11
    |    |  i. s.: HomocaligidaeT01
    |    |           |--Annerosella Habeeb 1966S76
    |    |           |    `--A. pacifica Wood 1969S76
    |    |           `--Homocaligus Berl. 1910G44
    |    |                |--H. amphibius Wainstein 1975T01
    |    |                |--H. crassipesWL09
    |    |                |--H. muscorumWL09
    |    |                `--H. scapularis (Koch 1838)T01 [=Caligonus scapularisG46]
    |    |         Mecognatha Wood 1967H98 [MecognathidaeWL09]
    |    |           `--M. hirsuta Wood 1967H98
    |    |         CryptognathusH98
    |    |         EupalopsellidaeGFS97
    |    |--+--+--Barbutia [Barbutiidae]S-TK22
    |    |  |  |    |--B. anguineaWL09
    |    |  |  |    |--B. australisWL09
    |    |  |  |    `--B. perretaeWL09
    |    |  |  `--PterygosomatidaeS-TK22
    |    |  `--StigmaeidaeSS55
    |    |       |--EryngiopusWL09
    |    |       |    |--E. arboreum Wood 1967SL71
    |    |       |    |--E. bifidus Wood 1967SL71
    |    |       |    |--E. coimbatorensisG01
    |    |       |    `--E. similis Wood 1967SL71
    |    |       |--EustigmaeusKU02
    |    |       |--TacebiaKU02
    |    |       |--WooderiaKU02
    |    |       |--ChaudhriaKU02
    |    |       |--MendanaiaK90
    |    |       |--Summersiella ancydactylaWL09
    |    |       |--Mullederia arborea Wood 1964SL71
    |    |       |--PilonychiopusWL09
    |    |       |--Caligohomus durusWL09
    |    |       |--StigmaeusS-TK22
    |    |       |--ProstigmaeusE-VVM-M07
    |    |       |--Liostigmaeus Thor 1930G44
    |    |       |--AgistemusKU02
    |    |       |--Neilstigmaeus Gerson & Smith Meyer 1995H98
    |    |       |    `--N. lamingtoni Gerson & Smith Meyer 1995H98
    |    |       |--Podaia Oudemans 1923G44
    |    |       |    `--*P. rubens (Schrank 1781) [=Acarus rubens]G44
    |    |       |--Parastigmaeus Kuznetzov 1984KU02
    |    |       |    |--*P. capensis (Meyer 1969) [=Pseudostigmaeus capensis]KU02
    |    |       |    `--P. andreae Khanjani & Ueckermann 2002KU02
    |    |       |--Mediolata Canestrini 1890WL09, SB63
    |    |       |    |--*M. longirostris (Berlese 1887) [=Stigmaeus longirostris]SB63
    |    |       |    |--M. brevisetis Wood 1967SL71
    |    |       |    |--M. favulosa Wood 1967SL71
    |    |       |    |--M. mariaefrancaeWL09
    |    |       |    |--M. pini (Canestrini 1890) [=Eupalopsis pini]SB63
    |    |       |    |--M. robusta Gonzalez 1965SL71
    |    |       |    `--M. simplex Wood 1967SL71
    |    |       |--Ledermuelleriopsis Willmann 1953KU02
    |    |       |    |--*L. triscutata Willmann 1951 (n. d.)KU02
    |    |       |    |--L. incisa Wood 1967SL71
    |    |       |    |--L. medicae Khanjani & Ueckermann 2002KU02
    |    |       |    |--L. plumosa Willmann 1951 [=L. plumosus]KU02
    |    |       |    |--L. spinosa Wood 1967SL71
    |    |       |    `--L. zahiri Khanjani & Ueckermann 2002KU02
    |    |       |--Cheylostigmaeus Willmann 1952KU02
    |    |       |    |--*C. grandiceps Willmann 1951KU02
    |    |       |    |--C. iranensis Khanjani & Ueckermann 2002KU02
    |    |       |    |--C. luxtoni Wood 1968SL71
    |    |       |    |--C. pannonicusZ99
    |    |       |    |--C. scutatusK90
    |    |       |    `--C. torulusKU02
    |    |       |--Storchia Oudemans 1923KU02 [incl. Apostigmaeus Grandjean 1944G44]
    |    |       |    |--*S. robusta (Berlese 1885)KU02 (see below for synonymy)
    |    |       |    |--S. anuaeKU02
    |    |       |    |--S. coneataKU02
    |    |       |    |--S. pacifica (Summer 1964)KU02, C-RSV-S88 [=Apostigmaeus pacificusC-RSV-S88]
    |    |       |    |--S. shanghaiansisKU02
    |    |       |    `--‘Raphignathus’ striatus Trägårdh 1904G44
    |    |       |--ZetzelliaKU02
    |    |       `--PseudostigmaeusSL71
    |    |            |--P. collyerae Wood 1967SL71
    |    |            |--P. longisetis Wood 1970SL71
    |    |            `--P. striatus Wood 1967SL71
    |    `--+--TetranychoideaS-TK22
    |       `--+--+--RaphignathidaeSS55
    |          |  |    |--Neoraphignathus howeiWL09
    |          |  |    |--Syncaligus Berl. 1910G44
    |          |  |    |--RaphignathusS-TK22
    |          |  |    |--Acheles Oudemans 1903SS55
    |          |  |    |    `--*A. mirabilisSS55
    |          |  |    `--Pimeliaphilus Trägårdh 1905J61
    |          |  |         |--*P. podapolipophagus Trägårdh 1905J61
    |          |  |         |--P. cunliffei Jack 1961J61
    |          |  |         |--P. gloriosusNK94
    |          |  |         |--P. isometri Cunliffe 1949J61
    |          |  |         |--P. rapax Beer 1960B60
    |          |  |         `--P. triatomaeWL09
    |          |  `--+--CaligonellidaeS-TK22
    |          |     `--+--+--DasythyreidaeWL09
    |          |        |  |    |--Xanthodasythyreus Walter & Gerson 1998WG98
    |          |        |  |    |    `--*X. toohey Walter & Gerson 1998WG98
    |          |        |  |    `--Dasythyreus Atyeo 1961WG98
    |          |        |  |         `--*D. hirsutus Atyeo 1961WG98
    |          |        |  `--XenocaligonellididaeWL09
    |          |        |       |--Apocaligonellidus [incl. Echinopsis Fan & Chen 1996 non Agassiz 1840]WL09
    |          |        |       |    `--A. fukiensis (Fan & Chen 1996) [=*Echinopsis fukiensis]WL09
    |          |        |       `--Xenocaligonellidus DeLeon 1959WG98
    |          |        |            |--*X. ovaerialis DeLeon 1959WG98
    |          |        |            |--X. galapagus González 1978WL09
    |          |        |            |--X. smileyi Hu & Liang 1995WG98
    |          |        |            `--X. teclae Meyer & Ueckermann 1989WL09
    |          |        `--Camerobiidae [Neophyllobiidae]FW06
    |          |             |--NeophyllobiusFW06
    |          |             |--Tycherobius Bolland 1986S96
    |          |             |    |--T. acicula Fan & Walter 2006FW06
    |          |             |    |--T. australis Fan & Walter 2006FW06
    |          |             |    |--T. elongata Fan & Walter 2006FW06
    |          |             |    |--T. lombardiniiWG98
    |          |             |    `--T. stipula Fan & Walter 2006FW06
    |          |             `--Camerobia Southcott 1957WG98, FW06
    |          |                  |--*C. australis Southcott 1957FW06
    |          |                  |--C. fissusWL09
    |          |                  |--C. monspeliensis Bolland 1986FW06
    |          |                  |--C. pistaciae Bolland 1986FW06
    |          |                  |--C. rugulosa Fan & Chen 1997FW06
    |          |                  |--C. southcotti Gerson 1972FW06
    |          |                  `--C. turcicus Koç & Kara 2005FW06
    |          `--CheyletoideaS-TK22
    `--Heterostigmata [Heterostigmae, Heterostigmatina, Tarsocheylina, Tarsonemida, Trachelostigmata]ZF11
         |  i. s.: Nasutiacarus Sidorchuk & Lindquist in Sidorchuk, Perrichot & Lindquist 2016 (see below for synonymy)SPL16
         |           `--*N. perplexus Sidorchuk & Lindquist in Sidorchuk, Perrichot & Lindquist 2016SPL16
         |--Tarsocheylidae [Tarsocheyloidea]L86
         |    |--Tarsocheylus paradoxusWL09
         |    `--Hoplocheylus Atyeo & Baker 1964S76
         |         |--H. atomariusNK94
         |         |--H. discalis Atyeo & Baker 1964S76
         |         `--H. pickardiWL09
         `--+--Heterocheylus Lombardini 1926H98 [Heterocheyletinae, HeterocheylidaeL86, Heterocheyloidea]
            |    `--H. lombardinii Womersley 1960H98
            `--TarsoneminaL86
                 |--DolichocyboideaL86
                 `--+--+--PygmephoroideaL86
                    |  `--+--PyemotoideaL86
                    |     `--TarsonemoideaZF11
                    |          |--TarsonemidaeL86
                    |          `--PodapolipidaeZF11
                    `--TrochometridioideaZF11
                         |--Trochometridium Cross 1965 [Trochometridiidae]L86
                         |    |--T. chinensisWL09
                         |    `--T. tribulatumWP99
                         `--AthyreacaridaeZF11
                              |--Athyreacarus pleiotretusWL09
                              `--Bembidiacarus [Bembidiacaridae]WL09

Eleutherengonides incertae sedis:
  CloacaroideaZF11
    |--EpimyodexWL09 [EpimyodicidaeZF11, Epimyodicinae]
    |    `--E. soricisWL09
    `--CloacaridaeZF11
         |--Pneumophagus bubonisWL09
         |--Caminacarus theodoriWL09
         |--Theodoracarus testudinisWL09
         |--Cloacarus fainiWL09
         `--Emyduracarus Fain 1968H98
              `--E. australis Fain 1968H98

Nasutiacarus Sidorchuk & Lindquist in Sidorchuk, Perrichot & Lindquist 2016 [Nasutiacaridae, Nastuiacaroidea]SPL16

*Storchia robusta (Berlese 1885)KU02 [=Caligonus robustusKU02, Stigmaeus robustaKU02; incl. *Apostigmaeus navicella Grandjean 1944G44, KU02, Storchia navicellaNK94]

*Type species of generic name indicated

References

[B60] Beer, R. E. 1960. A new species of Pimeliaphilus (Acarina: Pterygosomidae) parasitic on scorpions, with discussion of its postembryonic development. Journal of Parasitology 46 (4): 433–440.

[C-RSV-S88] Corpuz-Raros, L. A., G. C. Sabio & M. Velasco-Soriano. 1988. Mites associated with stored products, poultry houses and house dust in the Philippines. Philippine Entomologist 7 (3): 311–321.

[E-VVM-M07] Estrada-Venegas, E. G., I. M. Vázquez & J. A. Moreno-Moreno. 2007. Prostigmatid mites (Acarida: Prostigmata) in decaying wood from “La Mancha” Veracruz, Mexico. 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. 691–695. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[FW06] Fan, Q.-H., & D. E. Walter. 2006. Camerobia and Neophyllobius (Acari: Prostigmata: Camerobiidae) from Australia, with the descriptions of two new species. Zootaxa 1309: 1–23.

[GFS97] Gerson, U., W. E. Frost & S. F. Swift. 1997. A new genus of the family Eupalopsellidae from Australia (Acari: Prostigmata). International Journal of Acarology 23 (3): 185–189.

[G44] Grandjean, F. 1944. Observations sur les acariens de la famille des Stigmaeidae. Archives des Sciences Physiques et Naturelles, 5me Période 26: 103–131.

[G46] Grandjean, F. 1946. Au sujet de l’organe de Claparède, des eupathidies multiples et des taenidies mandibulaires chez les acariens actinochitineux. Archives des Sciences Physiques et Naturelles, 5me Période 28: 63–87.

[G01] Gupta, S. K. 2001. A conspectus of natural enemies of phytophagous mites and mites as potential biocontrol agents of agricultural pests in 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. 484–497. CSIRO Publishing: Melbourne.

[H98] Halliday, R. B. 1998. Mites of Australia: A checklist and bibliography. CSIRO Publishing: Collingwood.

[J61] Jack, K. M. 1961. A re-examination of the genera Pimeliaphilus Trägårdh 1905 and Hirstiella Berlese 1920 (Acari: Prostigmata). Annals and Magazine of Natural History, series 13, 4: 305–314.

[K90] Kethley, J. 1990. Acarina: Prostigmata (Actinedida). In: Dindal, D. L. (ed.) Soil Biology Guide pp. 667–756. John Wiley & Sones: New York.

[KU02] Khanjani, M., & E. A. Ueckermann. 2002. The stigmaeid mites of Iran (Acari: Stigmaeidae). International Journal of Acarology 28 (4): 317–339.

[L86] Lindquist, E. E. 1986. The world genera of Tarsonemidae (Acari: Heterostigmata): a morphological, phylogenetic, and systematic revision, with a reclassification of family-group taxa in the Heterostigmata. Memoirs of the Entomological Society of Canada 118 (S136): 1–517.

[NK94] Norton, R. A., & J. B. Kethley. 1994. Ecdysial cleavage lines of acariform mites (Arachnida, Acari). Zoologica Scripta 23 (3): 175–191.

[SB63] Schweizer, J., & C. Bader. 1963. Die Landmilben der Schweiz (Mittelland, Jura und Alpen): Trombidiformes Reuter, mit 217 Arten und Unterarten und 193 Originalzeichnungen. Denkschriften der Schweizerischen Naturforschenden Gesellschaft [Mémoires de la Société Helvétique des Sciences Naturelles] 84 (2): i–vi, 209–378.

[S76] Shiba, M. 1976. Taxonomic investigation on free-living Prostigmata from the Malay Peninsula. Nature and Life in Southeast Asia 7: 83–229.

[SPL16] Sidorchuk, E. A., V. Perrichot & E. E. Lindquist. 2016. A new fossil mite from French Cretaceous amber (Acari: Heterostigmata: Nasutiacaroidea superfam. nov.), testing evolutionary concepts within the Eleutherengona (Acariformes). Journal of Systematic Palaeontology 14 (4): 297–317.

[SL71] Spain, A. V., & M. Luxton. 1971. Catalog and bibliography of the Acari of the New Zealand subregion. Pacific Insects Monograph 25: 179–226.

[SS55] Summers, F. M., & E. I. Schlinger. 1955. Mites of the family Caligonellidae (Acarina). Hilgardia 23 (12): 539–561.

[S96] Swift, S. F. 1996. Biodiversity of raphignathoid mites in the Hawaiian islands (Acariformes: Prostigmata). In: Mitchell, R., D. J. Horn, G. R. Needham & W. C. Welbourn (eds) Acarology IX vol. 1. Proceedings pp. 339–341. Ohio Biological Survey: Columbus (Ohio).

[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.

[T01] Tolstikov, A. V. 2001. Reproductive behaviour of the semi-aquatic mite Homocaligus cf. amphibius Wainstein, 1975 (Acari: Homocaligidae). 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. 180–182. CSIRO Publishing: Melbourne.

[WG98] Walter, D. E., & U. Gerson. 1998. Dasythyreidae, new family, and Xanthodasythyreus n. g. (Acari: Prostigmata: Raphignathoidea) from Australia. International Journal of Acarology 24: 189–197.

[WL09] Walter, D. E., E. E. Lindquist, I. M. Smith, D. R. Cook & G. W. Krantz. 2009. Order Trombidiformes. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 233–420. Texas Tech University Press.

[WP99] Walter, D. E., & H. C. Proctor. 1999. Mites: Ecology, Evolution and Behaviour. CABI Publishing: Wallingford (UK).

[ZF11] Zhang, Z.-Q., Q.-H. Fan, V. Pesic, H. Smit, A. V. Bochkov, A. A. Khaustov, A. Baker, A. Wohltmann, T. Wen, J. W. Amrine, P. Beron, J. Lin, G. Gabryś & R. Husband. 2011. Order Trombidiformes Reuter, 1909. In: Zhang, Z.-Q. (ed.) Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 129–138.

[Z99] Zinkler, D. 1999. Ecophysiological adaptations of soil mites to oxygen deficiency. In: Needham, G. R., R. Mitchell, D. J. Horn & W. C. Welbourn (eds) Acarology IX vol. 2. Symposia pp. 47–51. Ohio Biological Survey: Columbus (Ohio).

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