Dorsal view of Molothrognathus leptostylus (cuticle texture shown on left side only), from Summers & Schlinger (1955).

Belongs within: Eleutherengonides.
Contains: Cryptognathus, Raphignathus, Neophyllobius, Eustigmaeus, Stigmaeus, Agistemus.

The Raphignathoidea are a group of often predatory mites, many species of which are found in dry habitats. Members of the group vary from heavily armoured taxa such as the Homocaligidae and Cryptognathidae to almost completely non-sclerotised taxa such as Barbutia and the Caligonellidae. Most raphignathoids 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 Barbutiidae. Members of the Cryptognathidae are heavily armoured mites with the dorsal shield produced anteriorly into a distinct collar into which the gnathosoma may be withdrawn. Homocaligidae are hemispherical mites found in association with fresh water. The 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).

Characters (from Walter et al. 2009): Soft-bodied or well-sclerotised, with a holonotal shield or a series of smaller weakly tanned, reticulate or dimpled dorsal sclerites, rarely without shields; idiosomatic neotrichy present in some families; trichobothria absent; prodorsum usually with 3-4 pairs of setae (si may be absent), rarely neotrichous; 0-2 pairs of ocelli, second pair (postocular organ) sometimes stalked. Cheliceral bases usually swollen, free, adnate, partially fused, or amalgamated into a stylophore, each with a styliform movable digit encased in the sheathlike remnant of the fixed digit; chelicerae not fused to underlying rostral elements, retractile in Cryptognathidae. Palpi with five free segments, palptibial claw (thumb-claw process) usually present but often weakly developed or obsolete, occasionally absent. Stigmatic openings present at base of chelicerae or absent; peritremes absent or simple to highly convoluted on cheliceral bases, or located on anterior margin of prodorsum and distally emergent (Raphignathidae). Empodial elements of tarsi commonly with tenent hairs, claws usually present on all legs (replaced by padlike structures in some Stigmaeidae), tenent hairs lacking on claws except in Barbutiidae. Males with an aedeagus, genital opening typically terminal, rarely dorsal; typically with solenidia on all tarsi. Generally with four active postembryonic instars: larva, protonymph, deutonymph and adult (tritonymph present in Xenocaligonellidae).

    |--Dasythyreidae WL09
    |    |--Xanthodasythyreus tooheyi Walter & Gerson 1998 WL09
    |    `--Dasythyreus hirsutus WL09
    |--Barbutia [Barbutiidae] WL09
    |    |--B. anguinea WL09
    |    |--B. australis WL09
    |    `--B. perretae WL09
    |--Homocaligidae T01
    |    |--Annerosella Habeeb 1966 S76
    |    |    `--A. pacifica Wood 1969 S76
    |    `--Homocaligus Berl. 1910 G44
    |         |--H. amphibius Wainstein 1975 T01
    |         |--H. crassipes WL09
    |         |--H. muscorum WL09
    |         `--H. scapularis (Koch 1838) T01 [=Caligonus scapularis G46]
    |--Mecognatha Wood 1967 H98 [Mecognathidae WL09]
    |    `--M. hirsuta Wood 1967 H98
    |--Xenocaligonellididae WL09
    |    |--Apocaligonellidus [incl. Echinopsis Fan & Chen 1996 non Agassiz 1840] WL09
    |    |    `--A. fukiensis (Fan & Chen 1996) [=*Echinopsis fukiensis] WL09
    |    `--Xenocaligonellidus WL09
    |         |--X. galapagus Gonzalez 1978 WL09
    |         |--X. ovaerialis WL09
    |         |--X. smileyi WL09
    |         `--X. teclae Meyer & Ueckermann 1989 WL09
    |--Cryptognathidae S01
    |    |--Favognathus S01
    |    |    |--F. distinctus S01
    |    |    |--F. favus (Summers & Chaudri 1965) [=Cryptognathus (Favognathus) favus] E-VVM-M07
    |    |    `--F. pictus (Summers & Chaudhri 1965) S96
    |    `--Cryptognathus H98
    |--Raphignathidae S01
    |    |--Neoraphignathus howei WL09
    |    |--Syncaligus Berl. 1910 G44
    |    |--Acheles Oudemans 1903 SS55
    |    |    `--*A. mirabilis SS55
    |    `--Raphignathus SB63
    |--Eupalopsellidae GFS97
    |    |--Eupalopsellus olandicus WL09
    |    |--Peltasellus WL09
    |    |--Saniosulus Summers 1960 H98
    |    |    `--S. nudus Summers 1960 H98
    |    |--Exothorhis Summers 1960 H98
    |    |    |--E. armata Summers 1960 H98
    |    |    |--E. caudata WL09
    |    |    `--E. echinocactii Gerson 1994 H98
    |    |--Eupalopsis Canestrini 1886 WL09, SB63
    |    |    |--*E. maseriensis (Canestrini & Fanzago 1876) [=Eupalus maseriensis] SB63
    |    |    |--E. jamesi Gerson 1994 H98
    |    |    |--E. pinicola WL09
    |    |    `--E. reticulatus Berlese 1910 SB63
    |    |--Acaciacarus Gerson, Frost & Swift 1997 GFS97
    |    |    `--*A. paradoxus Gerson, Frost & Swift 1997 GFS97
    |    `--Paraeupalopsellus Smiley & Moser 1968 GFS97
    |         |--P. aquatilis GFS97
    |         |--P. clareae WL09
    |         `--P. hodgesi GFS97
    |--Camerobiidae [Neophyllobiidae] S01
    |    |--Neophyllobius FW06
    |    |--Tycherobius Bolland 1986 S96
    |    |    |--T. acicula Fan & Walter 2006 FW06
    |    |    |--T. australis Fan & Walter 2006 FW06
    |    |    |--T. elongata Fan & Walter 2006 FW06
    |    |    `--T. stipula Fan & Walter 2006 FW06
    |    `--Camerobia Southcott 1957 FW06
    |         |--*C. australis Southcott 1957 FW06
    |         |--C. fissus WL09
    |         |--C. monspeliensis Bolland 1986 FW06
    |         |--C. pistaciae Bolland 1986 FW06
    |         |--C. rugulosa Fan & Chen 1997 FW06
    |         |--C. southcotti Gerson 1972 FW06
    |         `--C. turcicus Koç & Kara 2005 FW06
    |--Caligonellidae S01
    |    |--Namibarbutia WL09
    |    |--Paraneognathus [incl. Sinognathus] WL09
    |    |--Acheleopsis Willman 1951 SS55
    |    |    `--*A. quadrioculata SS55
    |    |--Caligonella Berlese 1910 SB63
    |    |    |--*C. humilis (Koch 1838) SB63 [=Stigmaeus humilis SB63, Caligonus humilis G46]
    |    |    `--C. afroensis S01
    |    |--Stigmagnathus Summers & Schlinger 1955 SS55
    |    |    |--*S. spectabilis Summers & Schlinger 1955 SS55
    |    |    `--S. terrestris Summers & Schlinger 1955 SS55
    |    |--Neognathus S01
    |    |    |--N. oblongus E-KO07
    |    |    |--N. spectabilis S01
    |    |    `--N. terrestris WL09
    |    |--Coptocheles Summers & Schlinger 1955 SS55
    |    |    |--*C. triscutatus Summers & Schlinger 1955 SS55
    |    |    |--C. boharti Summers & Schlinger 1955 SS55
    |    |    |--C. grandjeani Robaux 1975 S01
    |    |    |--C. kalapanaensis S01
    |    |    |--C. nakaharai S01
    |    |    `--C. solanii S01
    |    `--Molothrognathus Summers & Schlinger 1955 SS55
    |         |--*M. leptostylus Summers & Schlinger 1955 SS55
    |         |--M. colei S01
    |         |--M. conantae S01
    |         |--M. crucis Summers & Schlinger 1955 SS55
    |         |--M. fulgidus Summers & Schlinger 1955 SS55
    |         |--M. minutus E-KO07
    |         |--M. platlettus E-KO07
    |         |--M. rosei WL09
    |         `--M. tumipalpus S01
    `--Stigmaeidae S01
         |--Eryngiopus coimbatorensis WL09, G01
         |--Eustigmaeus KU02
         |--Tacebia KU02
         |--Wooderia KU02
         |--Chaudhria KU02
         |--Mendanaia K90
         |--Summersiella ancydactyla WL09
         |--Mullederia WL09
         |--Pilonychiopus WL09
         |--Caligohomus durus WL09
         |--Stigmaeus KU02
         |--Prostigmaeus E-VVM-M07
         |--Liostigmaeus Thor 1930 G44
         |--Agistemus KU02
         |--Neilstigmaeus Gerson & Smith Meyer 1995 H98
         |    `--N. lamingtoni Gerson & Smith Meyer 1995 H98
         |--Podaia Oudemans 1923 G44
         |    `--*P. rubens (Schrank 1781) [=Acarus rubens] G44
         |--Parastigmaeus Kuznetzov 1984 KU02
         |    |--*P. capensis (Meyer 1969) [=Pseudostigmaeus capensis] KU02
         |    `--P. andreae Khanjani & Ueckermann 2002 KU02
         |--Mediolata Canestrini 1890 WL09, SB63
         |    |--*M. longirostris (Berlese 1887) [=Stigmaeus longirostris] SB63
         |    |--M. mariaefrancae WL09
         |    `--M. pini (Canestrini 1890) [=Eupalopsis pini] SB63
         |--Ledermuelleriopsis Willmann 1953 KU02
         |    |--*L. triscutata Willmann 1951 (n. d.) KU02
         |    |--L. medicae Khanjani & Ueckermann 2002 KU02
         |    |--L. plumosa Willmann 1951 [=L. plumosus] KU02
         |    `--L. zahiri Khanjani & Ueckermann 2002 KU02
         |--Cheylostigmaeus Willmann 1952 KU02
         |    |--*C. grandiceps Willmann 1951 KU02
         |    |--C. iranensis Khanjani & Ueckermann 2002 KU02
         |    |--C. scutatus K90
         |    `--C. torulus KU02
         |--Storchia Oudemans 1923 KU02 [incl. Apostigmaeus G44]
         |    |--*S. robusta (Berlese 1885) KU02 (see below for synonymy)
         |    |--S. anuae KU02
         |    |--S. coneata KU02
         |    |--S. pacifica (Summer 1964) KU02, C-RSV-S88 [=Apostigmaeus pacificus C-RSV-S88]
         |    |--S. shanghaiansis KU02
         |    `--‘Raphignathus’ striatus Trägårdh 1904 G44
         `--Zetzellia Oudemans 1927 KU02
              |--*Z. methlagi Oudemans 1927 KU02
              |--Z. alni SS55
              |--Z. australis González-Rodríguez 1965 H98
              |--Z. ikusimai Shiba 1976 S76
              |--Z. longiseta S76
              |--Z. mali (Ewing 1917) [=Caligonella mali] KU02
              `--Z. maori W99

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

*Type species of generic name indicated


[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-KO07] El-Kammah, K. M., & L. M. I. Oyoun. 2007. Taxonomic records of mite species associated with animal and poultry farms in the Nile delta, vally and N. Sinai Egypt. 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. 223–232. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[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.

[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.

[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.

[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).

[S01] Swift, S. F. 2001. The leg chaetotaxy of Caligonellidae (Prostigmata: Raphignathoidea). 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. 242–249. CSIRO Publishing: Melbourne.

[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.

[W99] Walter, D. E. 1999. Cryptic inhabitants of a noxious weed: mites (Arachnida: Acari) on Lantana camara L. invading forests in Queensland. Australian Journal of Entomology 38: 197–200.

[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.

Last updated: 16 February 2019.

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