Argasidae

Pigeon tick Argas reflexus, photographed by Marcello Romano.

Belongs within: Parasitiformes.

The Argasidae, soft ticks, are a group of ticks lacking a well-developed scutum. Most feed on hosts returning to regular shelters such as nests, burrows or caves. Rather than permanently attaching to a single host like the hard ticks of the Ixodidae, females of Argasidae feed multiple times following maturity. Most argasids feed on their hosts for only short periods at a time, and may survive for extraordinarily long periods (up to several years) without feeding. However, the spinose ear tick Otobius megnini feeds deep within the ears of its mammalian host, causing damage that in extreme cases may even be fatal (Keirans 2009).

Species of Argas possess a distinct sutural line separating dorsal and ventral body surfaces that is absent in other genera. In Nothoaspis the anterior part of the dorsal integument is smooth and leathery (Keirans 2009).

Characters (from Keirans 2009): Capitulum situated anteriorly in larva, situated ventrally and not visible from above in postlarval stages; dorsal plate present or absent in larvae, leathery pseudoscutum rarely present in nymphs and adults. Paired spiracular plates situated dorsolaterally between coxae III-IV.

<==Argasidae [Argasini]
    |--Antricola Ke09
    |--Parantricola marginatus DC91
    |--Nothoaspis reddelli Ke09
    |--Otobius Banks 1912 H98
    |    |--O. lagophilus Ke09
    |    `--O. megnini (Dugès 1884) [=Argas megnini] H07
    |--Ornithodoros F-M91
    |    |--O. amblus Ke09
    |    |--O. coriaceus Koch 1844 H07
    |    |--O. dugesi Mazzotti 1943 M-PV-G01
    |    |--O. gurneyi Warburton 1926 [=Ornithodorus (l. c.) gurneyi] H98
    |    |--O. hermsi Wheeler et al. 1935 M-PV-G01
    |    |--O. kelleyi Kr09
    |    |--O. macmillani Hoogstraal & Kohls 1966 H98
    |    |--O. maritimus NJ91
    |    |--O. (Ornithodoros) moubata (Murray 1877) DJS91, P91
    |    |--O. muesebecki Ke09
    |    |--O. nicollei Mooser 1932 H07
    |    |--O. papillipes (Birula 1895) P91 [incl. O. tholozani (Laboulbène & Mégnin 1882) A56]
    |    |--O. parkeri Cooley 1936 M-PV-G01
    |    |--O. porcinus LL07
    |    |--O. savignyi F-M91
    |    |--O. talaje H07
    |    |--O. (Pavlovskyella) tartakovskyi DJS91
    |    |--O. tholozani F-M91
    |    |--O. transversus WP99
    |    |--O. turicata (Dugès 1876) [=Argas turicata] H07
    |    `--O. verrucosus B96a
    `--Argas Latreille 1795 S61
         |  i. s.: A. erraticus Lucas 1847 E12
         |         A. lahorensis OW01
         |--A. (Argas) KH75
         |    |--A. (A.) cooleyi KH75
         |    |--A. (A.) falco Kaiser & Hoogstraal 1974 KH75
         |    |--A. (A.) japonicus KH75
         |    |--A. (A.) lagenoplastis Froggatt 1906 KH75, H98
         |    |--A. (A.) latus LD91
         |    |--A. (A.) lowryae Kaiser & Hoogstraal 1975 KH75
         |    |--A. (A.) polonicus DJS91
         |    |--A. (A.) reflexus DJS91 [=Acarus reflexus CF77; incl. Rhynchoprion columbae CF77]
         |    |--A. (A.) tridentatus LD91
         |    |--A. (A.) radiatus B96b
         |    `--A. (A.) vulgaris LD91
         |--A. (Carios Latreille 1796) E-PAG91, H98
         |    |--A. (C.) australiensis Kohls & Hoogstraal 1962 H98
         |    |--‘Carios’ capensis (Neumann 1901) [=Ornithodoros talaje var. capensis] H98
         |    |--A. (C.) daviesi Kaiser & Hoogstraal 1973 H98
         |    |--A. (C.) dewae Kais & Hoogstraal 1974 H98
         |    |--‘Carios’ jerseyi GE05
         |    |--A. (C.) macrodermae Hoogstraal et al. 1977 H98
         |    `--A. (C.) vespertilionis Latreille 1802 E-PAG91
         `--A. (Persicargas) KH75
              |--A. (P.) arboreus DJS91
              |--A. (P.) nullaborensis Hoogstraal & Kaiser 1973 KH75
              |--A. (P.) persicus (Oken 1818) KH75, H98 (see below for synonymy)
              |--A. (P.) robertsi Hoogstraal et al. 1968 KH75
              `--A. (P.) walkerae DJS91

Argas (Persicargas) persicus (Oken 1818) KH75, H98 [=Rhynchoprion persicum H98; incl. A. americana Packard 1873 H98, A. americanus H98, A. victoriensis Sweet 1910 H98]

*Type species of generic name indicated

REFERENCES

[A56] Anastos, G. 1956. The ticks (Acarina: Ixodoidea) of the J. Klapperich Afghanistan Expedition, 1952 and 1953. Journal of the Washington Academy of Sciences 46 (1): 18–19.

[B96a] Buczek, A. 1996a. Anomalies in Argas reflexus (Fabr.) (Argasidae) collected from nature. In: Mitchell, R., D. J. Horn, G. R. Needham & W. C. Welbourn (eds) Acarology IX vol. 1. Proceedings pp. 319–324. Ohio Biological Survey: Columbus (Ohio).

[B96b] Buczek, A. 1996b. Regeneration of legs in Argas (Argas) reflexus (Fabricius) (Argasidae). In: Mitchell, R., D. J. Horn, G. R. Needham & W. C. Welbourn (eds) Acarology IX vol. 1. Proceedings pp. 709–711. Ohio Biological Survey: Columbus (Ohio).

[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.

[DC91] Daniel, M., V. Černý, J. Štěrba & J. de la Cruz. 1991. Morphological study of adults of trombiculid mites from Cuban hot caves. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 261–263. SPB Academic Publishing: The Hague.

[DJS91] Dusbábek, F., A. Jegorov & P. Šimek. 1991. Artificial assembly pheromone of argasid ticks (Ixodoidea: Argasidae). In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 59–68. SPB Academic Publishing: The Hague.

[E-PAG91] Estrada-Peña, A., A. Albó & J. Gutiérrez. 1991. Phenotypic variation of larval Argas (Carios) vespertilionis (Argasidae) in Spain and northern Africa. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 147–153. SPB Academic Publishing: The Hague.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[F-M91] Feldman-Muhsam, B. 1991. The role of Adlerocystis sp. in the reproduction of argasid ticks. In: Schuster, R., & P. W. Murphy (eds) The Acari: Reproduction, development and life-history strategies pp. 179–190. Chapman & Hall: London.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

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

[H07] Hoffmann, A. 2007. Acarology in Mexico: prehispanic to modern times. 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. 21–26. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[KH75] Kaiser, M. N., & H. Hoogstraal. 1975. Observations on the subgenus Argas (Ixodoidea: Argasidae: Argas). A. (A.) lowryae, new species, parasitizing cave-dwelling kestrels in the Nullarbor region of Western Australia. Annals of the Entomological Society of America 68 (3): 585–590.

[Ke09] Keirans, J. E. 2009. Order Ixodida. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3^rd ed. pp. 111–123. Texas Tech University Press.

[Kr09] Krantz, G. W. 2009. Form and function. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3^rd ed. pp. 5–53. Texas Tech University Press.

[LD91] Leonovich, S. A., & F. Dusbábek. 1991. Pheromone sensory subsystem in ticks: correlation between structure of sensilla and evolution of behaviour. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 53–58. SPB Academic Publishing: The Hague.

[LL07] Liu, J., L. Liu, Y. Zhang, Z. Wei & J. Liu. 2007. Preliminary studies on the juvenile hormone of Haemaphysalis longicornis Neumann (Acari: Ixodidae). 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. 205–208. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[M-PV-G01] Montiel-Parra, G., G. A. Villegas-Guzman, M. Vargas & O. J. Polaco. 2001. Mites associated with nests of Neotoma albigula Hartley, 1894 (Rodentia: Muridae) in Durango, México. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 586–593. CSIRO Publishing: Melbourne.

[NJ91] Nuttall, P. A., & L. D. Jones. 1991. Non-viraemic tick-borne virus transmission: mechanism and significance. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 3–6. SPB Academic Publishing: The Hague.

[OW01] Otto, J. C., & K. J. Wilson. 2001. Assessment of the usefulness of ribosomal 18S and mitochondrial COI sequences in Prostigmata phylogeny. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 100–109. CSIRO Publishing: Melbourne.

[P91] Podboronov, V. M. 1991. Antibacterial protective mechanisms of ixodoid ticks. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 375–380. SPB Academic Publishing: The Hague.

[S61] Southcott, R. V. 1961. Studies on the systematics and biology of the Erythraeoidea (Acarina), with a critical revision of the genera and subfamilies. Australian Journal of Zoology 9: 367–610.

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

Last updated: 21 May 2019.

Parasitiformes

Neocarus texanus, copyright Robert Deans.

Belongs within: Euchelicerata.
Contains: Mesostigmata, Ixodida.

The Parasitiformes are a group of usually well-sclerotised mites, many species of which are predatory or otherwise feeders on animal matter (scavengers or parasites). They are characterised by having the respiratory openings at the side of the body, close to the hind pairs of legs; four pairs of dorsolateral stigmata are present in Opilioacaridae but a single ventrolateral pair is present in other Parasitiformes. Earlier authors often treated the Opilioacaridae as the sister taxon to Parasitiformes rather than a member of the Parasitiformes proper, but they are commonly included by recent authors.

Opilioacarids are relatively large, long-legged mites with a leathery integument known from tropical and warm temperate parts of the world. The prodorsum bears two or three pairs of lateral ocelli whereas at most one pair is present in other Parasitiformes, and the opisthosoma bears indications of the ancestral segmentation. Opilioacarids also differ from other Parasitiformes in the presence of a large pair of claws on the palptarsi and the division of trochanters III and IV in tritonymphs and adults (Walter & Harvey 2009).

The remaining Parasitiformes are divided between three lineages whose relationships are uncertain. Recent studies, both morphological and molecular, have suggested that the Holothyrida are more closely related to the Ixodida than to the Mesostigmata but the support for this position is weak (Walter 2009). The Holothyrida are large, heavily sclerotised mites with a relictual distribution in South and Central America and Australasia. Members of the family Neothyridae are found in the Americas whereas Allothyridae and Holothyridae are found in Australasia; Holothyridae are also found on islands in the Indian Ocean (Walter 2009).

<==Parasitiformes [Anactinochaeta, Anactinochitinosi, Anactinotrichida, Gamasiformes, Parastigmata] WH09
    |--Opilioacaridae [Notostigmata, Opilioacarida, Opilioacariformes, Opilioacarina, Opilioacaroidea] WH09
    |    |  i. s.: Phalangiacarus brosseti WH09, WP99
    |    |         Panchaetes WH09
    |    |         Salfacarus WH09
    |    |         Adenacarus arabicus WH09
    |    |         Vanderhammenacarus WH09
    |    |         Gibocellum Stecker 1876 [Gibocellinae] K18
    |    |           `--*G. sudeticum Stecker 1876 K18
    |    |         Neocarus texanus (Chamberlin & Mulaik 1942) WH09, PL17 [=Opilioacarus texanus PL17]
    |    |--+--Paracarus Chamberlin & Mulaik 1942 H02
    |    |  |    `--P. hexophthalmus WH09
    |    |  `--Siamacarus Leclerc 1989 H02
    |    |       `--S. withi S07
    |    `--+--Eucarus With 1904 H02
    |       |    |--E. arabicus With 1904 H02 [=Opilioacarus arabicus S04]
    |       |    `--E. italicus With 1904 H02 [=Opilioacarus italicus S04]
    |       `--Opilioacarus With 1902 H02
    |            |--O. platensis Silvestri 1904 S04
    |            `--O. segmentatus With 1902 [=Eucarus segmentatus] H02
    `--Archicotylea AL37
         |--Mesostigmata W07
         `--+--Ixodida W07
            `--Holothyrida [Holothyriformes, Holothyrina, Holothyroidea, Tetrastigmata] W07
                 |  i. s.: Haplothyrus W09
                 |--Allothyridae W09
                 |    |--Australothyrus van der Hammen 1983 H98
                 |    |    `--*A. ocellatus van der Hammen 1983 H83
                 |    `--Allothyrus Womersley 1961 H98
                 |         |--A. australasiae (Womersley 1935) [=Holothyrus australasiae] H98
                 |         `--A. constrictus (Domrow 1955) [=Holothyrus constrictus] H98
                 `--+--Neothyrus [Neothyridae] W09
                    |    `--N. ana Lehtinen 1981 H83
                    `--Holothyridae W09
                         |--Holothyrus H83
                         |    |--H. armatus Canestrini 1897 H83
                         |    |--H. coccinella Gervais 1842 H83
                         |    |--H. gardineri Warburton 1913 H83
                         |    |--H. legendrei Van der Hammen 1983 H83
                         |    |--H. seychellensis Thon 1906 H83
                         |    `--H. testudineus Butler 1876 H83
                         |--Sternothyrus braueri DW10
                         |--Lindothyrus W09
                         `--Hammenius W09
                              |  i. s.: H. braueri (Thon 1905) [=Holothyrus braueri, Thonius braueri] H83
                              |--H. (Hammenius) H83
                              |    |--H. (H.) fujuge Lehtinen 1981 H83
                              |    |--H. (H.) ingii Lehtinen 1981 H83
                              |    `--H. (H.) niger (Thon 1905) H83
                              |--H. (Leiothyrus van der Hammen 1983) H83
                              |    |--H. (*L.) holthuisi Van der Hammen 1983 H83
                              |    |--‘Holothyrus’ expolitissimus Berlese 1923 H83
                              |    `--H. (L.) nitidissimus (Thorell 1882) (see below for synonymy) H83
                              `--H. (Thonius) H83
                                   |--H. (T.) grandjeani (Van der Hammen 1961) H83
                                   |--H. (T.) longipes (Thorell 1882) (see below for synonymy) H83
                                   `--H. (T.) montanus Van der Hammen 1983 H83

Nomen nudum: Holothyrus gervaisi Thon in Hughes 1959 H83

Hammenius (Leiothyrus) nitidissimus (Thorell 1882) [=Holothyrus nitidissimus; incl. Ho. scutifer Thorell 1882] H83

Hammenius (Thonius) longipes (Thorell 1882) [incl. Holothyrus longipes var. ferrugineus Thorell 1882] H83

*Type species of generic name indicated

REFERENCES

[AL37] André, M., & É. Lamy. 1937. Les Idées actuelles sur la Phylogénie des Acariens. Published by the author: Paris.

[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.

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

[H83] Hammen, L. van der. 1983. New notes on Holothyrida (anactinotrichid mites). Zoologische Verhandelingen 207: 1–48.

[H02] Harvey, M. S. 2002. The neglected cousins: what do we know about the smaller arachnid orders? Journal of Arachnology 30 (2): 357–372.

[K18] Kury, A. B. 2018. Familial nomina in harvestmen (Arachnida, Opiliones). Bionomina 13: 1–27.

[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.

[S07] Shultz, J. W. 2007. A phylogenetic analysis of the arachnid orders based on morphological characters. Zoological Journal of the Linnean Society 150 (2): 221–265.

[S04] Silvestri, F. 1904. Note Aracnologiche I–III. Redia 2: 239–261.

[W07] Walter, D. E. 2007. A plethora of questions, but only a modicum of answers: Australian Parasitiformes of tropical forests. 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. 51–57. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[W09] Walter, D. E. 2009. Order Holothyrida (Holothyrina, Tetrastigmata). In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3^rd ed. pp. 107–110. Texas Tech University Press.

[WH09] Walter, D. E., & M. S. Harvey. 2009. Order Opilioacarida. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3^rd ed. pp. 104–106. Texas Tech University Press.

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

Last updated: 14 March 2020.

Prostigmata

Labidostomma denticulata, copyright Walter Pfiegler.

Belongs within: Acariformes.
Contains: Eupodoidea, Eriophyoidea, Tydeoidea, Halacaridae, Trombidiae, Erythraeoidea, Hydrachnidia, Bdelloidea, Anystina, Eleutherengonides.

The Prostigmata are a diverse group of mites exhibiting a wide range of morphologies. The name of the group refers to the tracheal system which, if present, opens at the base of the chelicerae or on the anterior prodorsum (Lindquist et al. 2009). Many members of the group are more or less soft-bodied, but the Labidostommatidae are a group of heavily armoured predatory mites with an undivided dorsal shield and a complete ventral armature (Walter et al. 2009). In many members of the subgroups Anystina, Parasitengonina and Eleutherengonides, but not in the remaining subgroups, a terminal seta on the palptibia has become large and claw-like, and together with the reduced, laterally displaced tarsus forms a distinct arrangement referred to as the 'thumb-claw process' (Walter et al. 2009).

Walter et al. (2009) divided the Prostigmata between four major groups, the Labidostommatidae, Eupodides, Anystides and Eleutherengonides. Phylogenetic analysis supports the Labidostommatidae as the sister lineage of the remaining Prostigmata (Dabert et al. 2016). As well as their heavy armature, labidostommatids differ from other prostigmatans in having six pairs of prodorsal setae; the remaining Prostigmata all possess fewer. Other distinctive (though not unique) features of the Labidostommatidae include the presence of an unpaired median ocellus, and two pairs of prodorsal trichobothria. They are predators of Collembola and other small arthropods, and are found in soil or associated habitats (Walter et al. 2009).

The Eupodides were reocgnised by Walter et al. (2009) as a diverse group of mites largely united by plesiomorphic characters relative to other Prostigmata, including the Eupodoidea, Eriophyoidea, Tydeoidea, Halacaroidea and Bdelloidea. Most are soft-bodied forms with striated integument, weak subcutaneous sclerites, and reduced opisthosomatic chaetotaxy. Members of the Halacaroidea are aquatic; the remaining lineages are terrestrial. Halacaroidea also lack prodorsal trichobothria (present in other Eupodides) and possess large, often serrate, lateral claws on the leg tarsi that function as holdfasts for clinging to the substrate. Most species are included in the family Halacaridae, characterised by an idiosomal dorsum bearing one to four platelets and five to ten pairs of setae; the Australian genus Peza, in which the idiosomal dorsum has twelve to sixteen poorly defined platelets and sixteen to twenty pairs of setae, is placed in its own family. Phylogenetic analyses of the Prostigmata (e.g. Dabert et al. 2016) have not supported monophyly of the Eupodides.

The Anystides of Walter et al. (2009) included two subgroups, the Anystina and Parasitengonina. Again, Dabert et al. (2016) did not recover monophyly of this grouping but instead placed Anystina and Parasitengonina as distinct lineages. The Parasitengonina are a diverse group of mostly predatory mites with parasitic larvae, in which the larval instar is highly divergent in morphology from later instars, and the protonymphal and tritonymphal instars have become inactive calyptostases. Many authors have proposed dividing the Parasitengonina between primarily terrestrial (Trombidia) and aquatic (Hydrachnidia) lineages but Dabert et al. (2016) identified the former as paraphyletic to the latter. Terrestrial Parasitengonina are hairy and often relatively large, and often bright red in coloration. In members of the Erythraeoidea and Calyptostoma, the movable digit of the chelicera is styletiform and retractable. Calyptostoma species retain urstigmata between coxal plates I and II, lost in Erythraeoidea; they also bear fifteen or more setae on each of the coxal plates (Walter et al. 2009).

See also: The Prostigmata: endless forms.

Characters (from Lindquist et al. 2009): Tracheal system usually present with one pair of stigmata opening between bases of chelicerae or on anterior prodorsum (sometimes absent), sometimes associated with peritremes dorsally on cheliceral bases or on anterior margin of prodorsum; prodorsum usually with four or fewer pairs of setae or hypertrichous, sometimes including one or two pairs of bothridial sensilla; subcapitulum without rutella; chelicerae rarely chelate, usually with fixed digit sheathlike or completely regressed, movable digit usually a hook, knife, needle or stylet-like structure; cheliceral bases sometimes fused medially; palpi simple or modified into a thumb-claw process, sometimes reduced; coxae of legs integrated with venter of podosoma, coxal fields contiguous or II-III separated; ambulacra of at least legs II and III usually with two lateral claws and with or rarely without a median empodium that may be padlike or rayed and often armed with tenent hairs, or occasionally claw- or suckerlike; opisthosoma lacking paired lateral glands; opisthosomatic setal row c usually with two pairs of setae, rarely with three pairs or hypertrichous.

<==Prostigmata [Bdellina, Eupodides, Eupodina, Promatides]
    |  i. s.: Eupalus B16
    |           |--E. sternalis Berlese 1916 B16
    |           `--E. subterraneus Berlese 1916 B16
    |         Paratetranychus indicus Hirst 1923 H23
    |         Anychus latus H23
    |--Labidostommatidae (see below for synonymy) DPD16
    |    |--Eunicolina tuberculata WL09
    |    |--Akrostomma WL09
    |    |--Sellnickiella Feider & Vasilliu 1969 H98
    |    |    |--S. adelaideae (Womersley 1935) [=Labidostomma adelaideae, S. adelaidae (l. c.)] H98
    |    |    |--S. brasiliense WL09
    |    |    `--S. womersleyi (Atyeo & Crossley 1961) [=Labidostomma womersleyi, S. wommersleyi (l. c.)] H98
    |    `--Labidostomma Kramer 1879 S61
    |         |  i. s.: L. glumma Grandjean 1942 G42a
    |         |         L. hoegi G42a
    |         |         L. integrum G41
    |         |         L. lyra G42a
    |         |         L. mamillatus (Say 1821) [=Erythraeus mamillatus, Smaris mamillatus] S61
    |         |         L. nepalense WL09
    |         |         ‘Raphignathus’ ruber G42a
    |         |--*L. (Labidostomma) luteum Kramer 1879 SB63 [=Nicoletiella lutea WP99]
    |         |    |--L. l. luteum G42b
    |         |    |--L. l. elongatum Grandjean 1942 G42b
    |         |    `--L. l. repetitor Grandjean 1942 G42a
    |         `--L. (Nicoletiella Can. 1882) WL09, G42a
    |              |--L. (N.) cornuta WL09
    |              |--L. (N.) denticulata WL09 [=Acarus denticulatus G42a]
    |              |--L. (N.) jaquemarti WL09
    |              `--L. (N.) pacifica WL09
    `--+--+--Eupodoidea DPD16
       |  `--+--Eriophyoidea OW01
       |     `--Tydeoidea OW01
       `--+--+--Halacaroidea DPD16
          |  |    |  i. s.: Veladeadcarus gasconi WP99
          |  |    |         Pelacarus WL09
          |  |    |         Corallihalacarus chilcottensis WL09
          |  |    |--Halacaridae S86
          |  |    `--Peza Harvey 1990 H98 [Pezidae WL09]
          |  |         |--P. daps Harvey 1990 H98
          |  |         `--P. ops Harvey 1990 H98
          |  `--Parasitengonina (see below for synonymy) DPD16
          |       |  i. s.: Allotanaupodidae [Allotanaupodoidea] ZF11
          |       |--Trombidiae DPD16
          |       `--+--Erythraeoidea DPD16
          |          `--+--Hydrachnidia DPD16
          |             `--Calyptostoma Cambridge 1875 S61 (see below for synonymy)
          |                  |--C. expalpe (Hermann 1804) S61 (see below for synonymy)
          |                  |--C. caelatum (Berlese 1904) [=Smaris caelata, C. caelata] S76
          |                  |--C. extranea (Koch 1867) [=Smaridia extranea] H98
          |                  |--C. latiseta Shiba 1976 S76
          |                  |--C. lyncaea (Berlese 1887) (see below for synonymy) S61
          |                  |--C. simplexa Shiba 1976 S76
          |                  `--C. velutinus (Müller 1776) [=Acarus velutinus] H98
          `--+--Bdelloidea DPD16
             `--+--Anystina DPD16
                `--Eleutherengonides DPD16

Calyptostoma Cambridge 1875 S61 [Calyptostomatidae DPD16, Calyptostomatoidea, Calyptostomidae, Calyptostomidoidea]

Calyptostoma expalpe (Hermann 1804) S61 [=Trombidium expalpe S61, Smaris expalpis S61; incl. *C. hardii Cambridge 1875 S61, C. hardyi S61, Smaris impressa Koch 1837 S61, S. plana Koch 1879 T04]

Calyptostoma lyncaea (Berlese 1887) [=Smaris lyncaea; incl. S. impressa Canestrini 1882 non Koch 1837] S61

Labidostommatidae [Labidostomatidae, Labidostomatides, Labidostomidae, Labidostommatina, Labidostommatoidea, Labidostommidae, Labidostommina, Nicoletiellidae] DPD16

Parasitengonina [Apobolostigmata, Engonostigmata, Erythraeina, Erythraiae, Parasitengona, Parasitengonae, Parasitenini, Trombidia, Trombidina] DPD16

*Type species of generic name indicated

REFERENCES

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

[DPD16] Dabert, M., H. Proctor & J. Dabert. 2016. Higher-level molecular phylogeny of the water mites (Acariformes: Prostigmata: Parasitengonina: Hydrachnidiae). Molecular Phylogenetics and Evolution 101: 75–90.

[G41] Grandjean, F. 1941. Observations sur les acariens (6^e série). Bulletin du Muséum National d’Histoire Naturelle, 2^e Série 13 (6): 532–539.

[G42a] Grandjean, F. 1942a. Observations sure les Labidostommidae (1^re série). Bulletin du Muséum d’Histoire Naturelle, 2^e série 14 (2): 118–125.

[G42b] Grandjean, F. 1942b. Observations sur les Labidostommidae (2^e série). Bulletin du Muséum d’Histoire Naturelle, 2^e série 14 (3): 185–192.

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

[H23] Hirst, S. 1923. On some new or little-known species of Acari. Proceedings of the Zoological Society of London 1923: 971–1000.

[OW01] Otto, J. C., & K. J. Wilson. 2001. Assessment of the usefulness of ribosomal 18S and mitochondrial COI sequences in Prostigmata phylogeny. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 100–109. CSIRO Publishing: Melbourne.

[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.

[S86] Schwoerbel, J. 1986. Acari: Limnohalacaridae and Hydrovolziidae. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 643–647. E. J. Brill/Dr W. Backhuys: Leiden.

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

[S61] Southcott, R. V. 1961. Studies on the systematics and biology of the Erythraeoidea (Acarina), with a critical revision of the genera and subfamilies. Australian Journal of Zoology 9: 367–610.

[T04] Trägårdh, I. 1904. Fauna Arctica: Eine Zusammenstellung der arktischen Tierformen, mit besonderer Berücksichtigung des Spitzbergen-Gebietes auf Grund der Ergebnisse der Deutschen Expedition in das Nördliche Eismeer im Jahre 1898 vol. 4 pt 1. Monographie der arktischen Acariden. Gustav Fischer: Jena.

[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 3^rd 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 Trombiformes Reuter, 1909. In: Zhang, Z.-Q. (ed.) Animal biodiversity: an outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 129–138.

Last updated: 24 February 2019.

Erythraeoidea

Adult male Sphaerotarsus allmani, from Southcott (1961).

Belongs within: Parasitengonina.
Contains: Erythraeinae, Balaustiinae, Callidosomatinae, Leptus, Smaridinae.

The Erythraeoidea are a group of mites in which the adults and nymphs are predatory. The larvae are generally parasitic on other arthropods, though a few species are known to feed on pollen. Living erythraeoids are divided between the two families Smarididae and Erythraeidae. Adults of the Smarididae generally have flattened bodies, and the gnathosoma is placed on an extendable collar that can be fully retracted into the idiosoma. The Erythraeidae are generally less flattened, and the gnathosoma is not fully retractable. Two pairs of eyes are present in the smaridid taxa Smaridinae and Phanolophus and the erythraeid Erythraeinae. Phanolophus has the eyes placed between the levels of the anterior and posterior sensillary areas on the crista. A single pair of eyes is present in the smaridid Hirstistomatinae and the other erythraeid subfamilies (Southcott 1961). A third family has been recognised for Proterythraeus southcotti, described from a larva found in Cretaceous Canadian amber.

Characters (from Southcott 1961): Eyes always present, sessile, one or two on each side. Thumb-complex present to palp. Two pairs of idiosomal sensillae (bothrialae) present: in the adult these are usually placed on a dorsal crista metopica; in the larva usually placed on a dorsal scutum. Chelicerae of adult styliform. Adult peritremes placed dorsally on mouth-cone, longitudinal. Larvae without external genital aperture or anus. Larval coxae always separated. Larva without urstigma.

<==Erythraeoidea [Leptoidea, Rhyncholophidae, Rhyncholophides, Rhyncholophinae]
    |  i. s.: Erythrombium Berlese 1910 (n. d.) S61
    |           `--*E. eusisyrum Berlese 1910 (n. d.) S61
    |         Pararainbowia martilli Dunlop 2007 ZF11
    |--Proterythraeus WL09 [Proterythraeidae ZF11]
    |    `--P. southcotti WL09
    |--Erythraeidae [Erythreidae, Leptidae, Leptides, Leptoidae, Rhyncholophini, Rhyncholophoidae] G91
    |    |  i. s.: Cloquetia Oudemans 1941 S61
    |    |           `--*C. medioareolata (Kramer 1898) [=Rhyncholophus medioareolatus] S61
    |    |         Pseudoerythraeus Haitlinger 2004 F05
    |    |           `--*P. ernesti Haitlinger 2004 F05
    |    |         Actineda S22
    |    |           |--A. pallescens CF77
    |    |           `--A. vitis S22
    |    |         Scyphius cerinus [incl. S. pratensis, S. pyrroleucus] CF77
    |    |         Harpagella Southcott 1996 H98
    |    |           `--H. moxonae Southcott 1996 H98
    |    |         Neabrolophus S76
    |    |           |--N. bisetis (Evans 1953) S76, S61 [=Balaustium bisetis S61]
    |    |           |--N. brevisetorum S76
    |    |           |--N. hirsutus Shiba 1976 S76
    |    |           `--N. longisetorum S76
    |    |         Arythaena troguloides Menge 1854 S93
    |    |--Erythraeinae G91
    |    `--+--Balaustiinae G91
    |       `--+--Callidosomatinae G91
    |          `--+--Leptus G91
    |             `--Myrmicotrombium Womersley 1934 S61 [=Myrmicothrombium (l. c.) S61; Myrmicotrombiinae G91]
    |                  `--*M. brevicristatum Womersley 1934 S61
    `--Smarididae [Smaridae, Smarididen, Smaridiidae, Smarisidae] G91
         |  i. s.: Hehlenia Oudemans 1941 (n. d.) S61
         |           `--*H. papillosa (Koch 1837) (n. d.) (see below for synonymy) S61
         |         Pilosoma Southcott 1961 S61
         |           `--*P. pluto Southcott 1961 S61
         |         Surasmaris longirostris WL09
         |         Neophanolophus Shiba 1976 S76
         |           `--*N. morii Shiba 1976 S76
         |--Smaridinae S61
         |--Phanolophus André 1927 (see below for synonymy) S61
         |    |--P. oedipodarum (Frauenfeld 1868) (see below for synonymy) S61
         |    `--‘*Platylophus’ argasoides Berlese 1910 [=*Euplatylophus argasoides] S61
         `--Hirstiosomatinae S61
              |--Trichosmaris jacoti K90, WL09
              |--Clipeosoma Southcott 1948 S61
              |    |--*C. copiolarum Southcott 1948 S61
              |    |--C. furtadoi Shiba 1976 S76
              |    `--C. jupiter Southcott 1961 S61
              |--Hirstiosoma Womersley 1934 [incl. Niemannia Oudemans 1941] S61
              |    |--*H. scalare Womersley 1934 [=H. scalaris] S61
              |    |--H. ampulligera (Berlese 1887) S61 (see below for synonymy)
              |    |--H. latreillei (Grandjean 1947) S61
              |    |--H. novaehollandiae Womersley 1936 SL71
              |    `--H. tasmaniensis Womersley & Southcott 1941 H98
              `--Sphaerotarsus Womersley 1936 S61
                   |--S. allmani Womersley 1936 S61
                   |--S. claviger Womersley & Southcott 1941 H98
                   |--S. leptopilus Womersley & Southcott 1941 S61
                   |--S. monticolus Southcott 1997 H98
                   `--S. womersleyi Southcott 1946 S61

*Hehlenia papillosa (Koch 1837) (n. d.) [=Rhyncholophus papillosus non Trombidium papillosum Hermann 1804] S61

Hirstiosoma ampulligera (Berlese 1887) S61 [=Smaridia ampulligera S61, *Niemannia ampulligera S61, Smaris ampulligera SB63]

Phanolophus André 1927 [=Phanalophus (l. c.); incl. Euplatylophus Radford 1950, Euplathylophus (l. c.), Platylophus Berlese 1910 nec Swainson 1832 nec Koch 1839, Platerythraeus (l. c.); Phanolophinae] S61

Phanolophus oedipodarum (Frauenfeld 1868) [=Rhyncholophus oedipodarum; incl. Smaridia magnifica Berlese 1918, Smaris magnifica, *Phanolophus nasica André 1927, Smaris nasica] S61

*Type species of generic name indicated

REFERENCES

[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.

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261–282.

[G91] Gabryś, G. 1991. Intergeneric relationships within Erythraeidae (Acari, Actinedida). In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 207–212. SPB Academic Publishing: The Hague.

[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.

[S22] Schweizer, J. 1922. Beitrag zur Kenntnis der terrestrischen Milbenfauna der Schweiz. Verhandl. Naturf. Ges. Basel 33: 23–112, 4 pls.

[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.

[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.

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

[S61] Southcott, R. V. 1961. Studies on the systematics and biology of the Erythraeoidea (Acarina), with a critical revision of the genera and subfamilies. Australian Journal of Zoology 9: 367–610.

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

[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 3^rd ed. pp. 233–420. Texas Tech University Press.

[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.

Last updated: 2 July 2022.

Leptus

Leptus sp., photographed by Tom Murray.

Belongs within: Erythraeoidea.

Leptus is a cosmopolitan genus of erythraeid mites in which adults have the single pair of eyes placed relatively close to the front of the main body. The idiosoma is distinctly bushy and usually darkly coloured (Walter et al. 2009). Larvae have a triangular scutum that is widest anteriorly. Larvae are found on a wide range of hosts, including almost all terrestrial arachnids and insects. The type species Leptus phalangii was, as its name indicates, first described as a larva parasitising a harvestman in Sweden. Adults were initially described as belonging to a genus Achorolophus, until rearing experiments established the association of larvae and adults (Southcott 1961).

Characters (from Southcott 1961): Idiosoma with one eye on each side, placed in the adult and nymph alongside the anterior half of the crista. Crista of adult and nymph with two sensillary areas, the anterior at the apex of the propodosoma, and the posterior at the posterior end of the crista. Larva with a triangular dorsal scutum, the apex posterior, and with two pairs of scutalae placed anterolaterally; two pairs of scutal sensillae present, one pair anteromedian, at about the level of the AL scutalae; posterior pair at or near posterior pole of scutum; larval lateral (neolateral) tarsal claws dissimilar.

<==Leptus Latreille 1796 S91b [incl. Achorolophus Berlese 1891 S91b, Ritteria Kramer 1877 S61; Leptinae]
    `--L. (Leptus) H09
         |--*L. (L.) phalangii (de Geer 1778) [=Acarus phalangii] S91b
         |--L. (L.) auliacus Haitlinger 1998 H09
         |--L. (L.) barmeedius Southcott 1999 H09
         |--L. (L.) batoricus Haitlinger 1998 H09
         |--L. (L.) boggohoranus Haitlinger 1990 H09
         |--L. (L.) esmaili Saboori & Ostovan 2000 H09
         |--L. (L.) heleus Southcott 1993 H09
         |--L. (L.) josifovi Beron 1975 H09
         |--L. (L.) kamalii Karini Irvanlou & Saboori 2001 H09
         |--L. (L.) korematus Southcott 1999 H09
         |--L. (L.) korneli Haitlinger 2009 H09
         |--L. (L.) mirenae Haitlinger 1994 H09
         |--L. (L.) polythrix Fain & Elsen 1987 H09
         |--L. (L.) rosellae Haitlinger 1999 H09
         `--L. (L.) salicus Haitlinger 2009 H09

Leptus incertae sedis:
  L. acuminatus Shiba 1976 S76
  L. agrotis Southcott 1993 H98
  L. alopecurus Southcott 1991 S91b
  L. alvimordax Southcott 1991 S91b
  L. americanus Riley 1873 (n. d.) S61
  L. anomalus Southcott 1946 S91b
  L. ariel WL09
  L. atticolus WY88
  L. bathypogonus Womersley 1934 S91b
  L. belicolus Southcott 1993 H98
  L. beroni Fain 1991 GF11
  L. bicristatus CM07
  L. calidus Shiba 1976 S76
  L. cameronensis Shiba 1976 S76
  L. cerambycius Southcott 1993 H98
  L. charon Southcott 1991 S91b
  L. chelonethus Womersley 1934 S91b
  L. chibai Shiba 1976 S76
  L. clethrionomydis Haitlinger 1987 S91a
  L. coccineus (Scopoli 1763) SB63 (see below for synonymy)
  L. comosus Southcott 1991 S91b
  L. curtipes Schweizer 1951 SB63
  L. debeauforti (Oudemans 1905) (n. d.) [=Erythraeus debeauforti, Achorolophus debeauforti] S91b
  L. draco S91b
  L. dubius (Paoli 1937) [=Achorolophus dubius] S61
  L. faini Southcott 1993 H98
  L. fernandezi WWO01
  L. fortei Southcott 1991 S91b
  L. francesi Southcott 1991 S91b
  L. gagrellae CM07
  L. gauphalus Haitlinger 1990 H98
  L. georgeae Southcott 1993 H98
  L. gifuensis WY88
  L. gyas CM07
  L. halli Southcott 1993 H98
  L. hidakai WY88
  L. hirtipes (Berlese 1910) [=Abrolophus hirtipes] V25
  L. hozumii Shiba 1976 S76
  L. ignotis (Oudemans 1903) WY88, S61 [=Erythraeus ignotus S61, Achorolophus ignotus S61]
  L. incertus (Koch 1854) [=Rhyncholophus incertus] S61
  L. indianensis CM07
  L. jenseni Southcott 1993 H98
  L. jocquei CM07
  L. kalaallus CM07
  L. killingtoni Turk 1945 S61
  L. lomani [=Erythraeus lomani] S61
  L. maculatus (Banks 1894) (see below for synonymy) S61
  L. mariae Haitlinger 1987 MF11
  ‘Abrolophus’ marmoratus S61
  L. molochinus (Koch 1837) [=Rhyncholophus molochinus] SB63
  L. monteithi Southcott 1993 H98
  L. murteri Schweizer 1951 SB63
  L. muscarum (Linnaeus 1758) [=Acarus muscarum] H98
  L. nearcticus CM07
  L. nemorum (Koch 1835) SB63 (see below for synonymy)
  ‘Ritteria’ norvegica T04
  L. ornatus Womersley 1934 H98
  L. orthrius Southcott 1993 H98
  L. oudemansi CM07
  L. ovatus S76
  L. phuketicus CM07
  L. puylaerti CM07
  L. pyrenaeus André 1953 F91
  ‘Rhyncholophus’ quinquemaculatus [=Trombidium quinquemaculatum, R. (Abrolophus) quinquemaculatus] S61
  L. repens S76
  L. rubricatus (Koch 1837) [=Rhyncholophus rubricatus] SB63
  L. rupestris W91
  L. schedingi (Oudemans 1911) S91b
  L. siculus [=Rhyncholophus siculus, R. (Abrolophus) siculus] S61
  L. spinalatus Southcott 1993 H98
  L. stieglmayri (Oudemans 1905) S91b [=Erythraeus stieglmayri S61]
  L. swani Southcott 1991 S91b
  L. tarranus Southcott 1993 H98
  L. terebrans W34
  L. tetrigius Southcott 1988 S91b
  L. titinius Southcott 1993 H98
  L. torresianus Southcott 1988 H98
  L. trimaculatus (Hermann 1804) SB63 (see below for synonymy)
  L. trucidatus (Hull 1923) [=Achorolophus trucidatus] H98
  L. truncatus Southcott 1993 H98
  L. utheri Southcott 1993 H98
  L. vertex (Kramer 1881) SB63 (see below for synonymy)
  L. villosus S76
  L. waldockae Fain 1991 F91
  L. warregense (Hirst 1928) [=Belaustium (l. c. for Balaustium) warregense] H98
  L. yoshimii Shiba 1976 S76

Leptus coccineus (Scopoli 1763) SB63 [=Pediculus coccineus SB63, L. coccineous (l. c.) S61, Pediculeus (l. c.) coccineus S61]

Leptus maculatus (Banks 1894) [=Rhyncholophus maculatus, Rhyncolophus (l. c.) maculatus, Erythraeus (Atomus) maculatus] S61

Leptus nemorum (Koch 1835) SB63 [=Rhyncholophus nemorum SB63, Rh. (Abrolophus) nemorum S61, Rh. (Achorolophus) nemorum S61, *Ritteria nemorum S61]

Leptus trimaculatus (Hermann 1804) SB63 [=Trombidium trimaculatum SB63, Rhyncholophus trimaculatus S61, R. (Abrolophus) trimaculatus S61, R. (Achorolophus) trimaculatus S61]

Leptus vertex (Kramer 1881) SB63 [=Rhyncholophus nemorum vertex SB63, Erythraeus vertex T04, Ritteria vertex T04; incl. Rh. tonsus Koch 1879 (n. d.) T04]

*Type species of generic name indicated

REFERENCES

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[F91] Fain, A. 1991. Notes on mites parasitic or phoretic on Australian centipedes, spiders and scorpion. Records of the Western Australian Museum 15: 69–82.

[GF11] Gabryś, G., M. Felska, A. Kłosińska, W. Staręga & J. Mąkol. 2011. Harvestmen (Opiliones) as hosts of Parasitengona (Acari: Actinotrichida, Prostigmata) larvae. Journal of Arachnology 39 (2): 349–351.

[H09] Haitlinger, R. 2009. Three new species of mites (Acari: Prostigmata: Erythraeidae) from the Republic of Cape Verde. Biologia 64 (6): 1150–1156.

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

[MF11] Mąkol, J., & M. Felska. 2011. New records of spiders (Araneae) as hosts of terrestrial Parasitengona mites (Acari: Actinotrichida: Prostigmata). Journal of Arachnology 39 (2): 352–354.

[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.

[S61] Southcott, R. V. 1961. Studies on the systematics and biology of the Erythraeoidea (Acarina), with a critical revision of the genera and subfamilies. Australian Journal of Zoology 9: 367–610.

[S91a] Southcott, R. V. 1991a. A further revision of Charletonia (Acarina: Erythraeidae) based on larvae, protonymphs and deutonymphs. Invertebrate Taxonomy 5: 61–131.

[S91b] Southcott, R. V. 1991b. Descriptions of larval Leptus (Acarina: Erythraeidae) ectoparasitic on Australian Diptera, and two earlier described Australian larvae. Invertebrate Taxonomy 5: 717–763.

[T04] Trägårdh, I. 1904. Fauna Arctica: Eine Zusammenstellung der arktischen Tierformen, mit besonderer Berücksichtigung des Spitzbergen-Gebietes auf Grund der Ergebnisse der Deutschen Expedition in das Nördliche Eismeer im Jahre 1898 vol. 4 pt 1. Monographie der arktischen Acariden. Gustav Fischer: Jena.

[V25] Vitzthum, H. 1925. Fauna sumatrensis. (Beitrag no. 5). Acarinae. Supplementa Entomologica 11: 1–79.

[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 3^rd ed. pp. 233–420. Texas Tech University Press.

[WY88] Welbourn, W. C., & O. P. Young. 1988. Mites parasitic on spiders, with a description of a new species of Eutrombidium (Acari, Eutrombidiidae). Journal of Arachnology 16: 373–385.

[W91] Witte, H. 1991. Indirect sperm transfer in prostigmatic mites from a phylogenetic viewpoint. In: Schuster, R., & P. W. Murphy (eds) The Acari: Reproduction, development and life-history strategies pp. 137–176. Chapman & Hall: London.

[WWO01] Wohltmann, A., H. Witte & R. Olomski. 2001. Organismal patterns causing high potential for adaptive radiation in Parasitengonae (Acari: Prostigmata). In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10^th International Congress pp. 83–99. CSIRO Publishing: Melbourne.

[W34] Womersley, H. 1934. A revision of the trombid and erythraeid mites of Australia with descriptions of new genera and species. Records of the South Australian Museum 5 (2): 179–254.

Last updated: 2 July 2022.

Erythraeinae

Eatoniana plumipes, photographed by Cor Zonneveld.

Belongs within: Erythraeoidea.
Contains: Erythraeus.

The Erythraeinae are a group of often large erythraeid mites that possess two pairs of eyes, rather than one pair as in other erythraeids. In members of the genera Augustsonella to Kakamasia in the tree below, the palp bears distinctly thickened, peg- or cone-like setae (conalae) on the genu and tibia; these modified setae are absent in other genera (Gabryś 1991). The palpal conalae are present dorsolaterally on the appendage in the North American Augustsonella tuberculata whereas in other genera with palpal conalae are present ventrally only. The New Zealand species Neosmaris novaezealandiae has the crista obsolescent with only the anterior and posterior sensillary areas remaining (Southcott 1961).

Characters (from Southcott 1961): Erythraeidae with two eyes on each side. Adult and nymph normally with a crista (absent in Neosmaris); anterior sensillary area of crista placed dorsally at the anterior pole of the propodosoma. Larvae with two eyes on each side; dorsal scutum rounded or squarish with two or three pairs of scutalae (except teratologically); lateral (neolateral) tarsal claws asymmetrical.

<==Erythraeinae S61
    |  i. s.: Forania Southcott 1961 S61
    |           `--*F. mentonensis (André 1929) [=Bochartia mentonensis] S61
    |         Grognieria Oudemans 1941 (n. d.) S61
    |           `--*G. agilis (Canestrini 1884) [=Rhyncholophus agilis] S61
    |         Bochartia Oudemans 1910 [=Bockartia (l. c.), Brochartia (l. c.)] S61
    |           |--*B. kuyperi Oudemans 1910 S61
    |           `--B. adrastus Southcott 1961 S61
    |--+--Augustsonella Southcott 1961 G91 [=Augustsonia Southcott 1961 S61; Augustsonellinae]
    |  |    `--*A. tuberculata (Augustson 1940) [=Erythraeus tuberculatus, *Augustsonia tuberculata] S61
    |  `--+--Lasioerythraeus Welbourn & Young 1987 G91
    |     |    |--L. johnstoni WL09
    |     |    `--L. whitcombi WL09
    |     `--+--+--Tepoztlana Hoffmann & Mendez 1973 G91
    |        |  `--Erythraeus G91
    |        `--+--Podosmaridia Trägårdh 1937 G91
    |           |    `--*P. longipes Trägårdh 1937 S61
    |           `--+--Claverythraeus Trägårdh 1937 G91
    |              |    `--*C. mongolicus Trägårdh 1937 S61
    |              `--Kakamasia Lawrence 1944 G91
    |                   |--*K. polygona Lawrence 1944 S61
    |                   `--K. pilosa Lawrence 1944 S61
    `--+--Neosmaris Hirst 1926 G91
       |    `--*N. novaezealandiae Hirst 1926 S61
       `--+--Erythrellus Southcott 1946 G91
          |    `--*E. imbricatus Southcott 1946 S61
          `--+--+--+--Paraphanolophus Smiley 1968 G91
             |  |  `--Erythraxus Southcott 1961 G91
             |  |       `--*E. bisetosus (Wharton 1938) [=Erythraeus bisetosa, Erythraeus bisetosus] S61
             |  `--+--Curteria Southcott 1961 G91 [incl. Pedroerythraeus Haitlinger 2004 WMG07]
             |     |    |--*C. curticristata (Willmann 1950) [=Morieria curticristata] WMG07
             |     |    |--C. episcopalis (Koch 1837) WMG07
             |     |    |--C. ernesti (Haitlinger 2004) [=Pedroerythraeus ernesti] WMG07
             |     |    `--C. southcotti Gabryś 1992 WMG07
             |     `--Eatoniana Cambridge 1898 G91 (see below for synonymy)
             |          |--E. plumipes (Koch 1856) S61 (see below for synonymy)
             |          |--‘Erythraeus (*Ptilolophus)’ claviger Berlese 1916 B16
             |          |--E. namaquensis (Lawrence 1937) S61
             |          `--E. plumifer Birula 1893 (see below for synonymy) S61
             `--+--Erythroides Southcott 1946 G91
                |    |--*E. serratus (Womersley 1936) S61 [=Erythraeus serratus H98]
                |    |--E. clavatus Southcott 1946 H98
                |    |--E. darwini Southcott 1946 H98
                |    |--E. macdonnelli Southcott 1946 H98
                |    `--E. neoserratus Southcott 1946 H98
                `--+--Rainbowia Southcott 1961 G91
                   |    |--*R. imperator (Hirst 1928) S61 (see below for synonymy)
                   |    `--R. celeripes (Rainbow 1906) [=Rhyncholophus celeripes, Erythraeus celeripes] H98
                   `--Erythrites Southcott 1961 G91
                        |  i. s.: E. jacksoni WL09
                        |--E. (Erythrites) S61
                        |    |--*E. (E.) reginae (Hirst 1928) S61 (see below for synonymy)
                        |    `--E. (E.) stuarti (Southcott 1946) S61 [=Erythraeus stuarti H98]
                        |--E. (Gerites Southcott 1961) S61
                        |    |--E. (*G.) guttatus (Southcott 1946) S61 [=Erythraeus guttatus H98]
                        |    `--E. (G.) osmondensis (Southcott 1946) S61 [=Erythraeus osmondensis H98]
                        |--E. (Terebrites Southcott 1961) S61
                        |    `--E. (*T.) womersleyi (Southcott 1946) S61 [=Erythraeus womersleyi H98]
                        `--E. (Urrbites Southcott 1961) S61
                             |--E. (*U.) urrbrae (Womersley 1934) S61 (see below for synonymy)
                             `--E. (U.) pilosus (Hirst 1928) S61 [=Leptus pilosus H98, Erythraeus pilosus H98]

Eatoniana Cambridge 1898 G91 [=Eatonia Cambridge 1897 nec Hall 1857 nec Smith 1875 nec Meunier 1905 S61; incl. Lucasiella Banks 1900 SB63, Macropus Birula 1893 nec Shaw 1790 nec Latreille 1802 nec Thunberg 1805 nec Spix 1824 S61, Ptilolophus Berlese 1916 B16, S61]

Eatoniana plumifer Birula 1893 [=Rhyncholophus plumifer; incl. E. georgei Oudemans 1941, E. plumifer f. transcaspica Oudemans 1941 (n. n.)] S61

Eatoniana plumipes (Koch 1856) S61 [=Rhyncholophus plumipes S61, *Lucasiella plumipes SB63; incl. R. plumipes Lucas 1864 non Koch 1856 S61, *Eatonia scopulifera Cambridge 1897 S61, *Eatoniana scopulifera S61]

*Erythrites (Erythrites) reginae (Hirst 1928) S61 [=Leptus reginae H98, Erythraeus regina (l. c.) H98, Erythraeus reginae H98; incl. L. antepodianus Hirst 1928 H98, Erythraeus antepodianus H98, Erythrites antepodianus H98]

Erythrites (*Urrbites) urrbrae (Womersley 1934) S61 [=Erythraeus urrbrae H98; incl. Bockartia (l. c.) longipes Womersley 1934 H98, Bochartia longipes H98]

*Rainbowia imperator (Hirst 1928) S61 [=Leptus imperator H98, Erythraeus imperator H98; incl. Bochartia oudemansi Womersley 1936 H98, E. oudemansi H98]

*Type species of generic name indicated

REFERENCES

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

[G91] Gabryś, G. 1991. Intergeneric relationships within Erythraeidae (Acari, Actinedida). In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 207–212. SPB Academic Publishing: The Hague.

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

[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.

[S61] Southcott, R. V. 1961. Studies on the systematics and biology of the Erythraeoidea (Acarina), with a critical revision of the genera and subfamilies. Australian Journal of Zoology 9: 367–610.

[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 3^rd ed. pp. 233–420. Texas Tech University Press.

[WMG07] Wohltmann, A., J. Mąkol & G. Gabryś. 2007. Description of the larva of Curteria Southcott, 1961 (Acari, Parasitengona, Erythraeidae) with notes on biology and life cycle. Biologia 62 (5): 573–580.

Last updated: 2 July 2022.