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. 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).

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 DC91
    |--Parantricola marginatus DC91
    |--Nothoaspis reddelli Ke09
    |--Otobius Ke09
    |    |--O. lagophilus Ke09
    |    `--O. megnini OW01
    |--Ornithodoros OW01
    |    |--O. amblus Ke09
    |    |--O. capensis R01
    |    |--O. coriaceus Ke09
    |    |--O. dugesi Mazzotti 1943 M-PV-G01
    |    |--O. gurneyi R01
    |    |--O. hermsi Wheeler et al. 1935 M-PV-G01
    |    |--O. kelleyi Kr09
    |    |--O. macmillani R01
    |    |--O. maritimus NJ91
    |    |--O. (Ornithodoros) moubata (Murray 1877) DJS91, P91
    |    |--O. muesebecki Ke09
    |    |--O. papillipes (Birula 1895) P91
    |    |--O. parkeri Cooley 1936 M-PV-G01
    |    |--O. (Pavlovskyella) tartakovskyi DJS91
    |    |--O. tholozani WP99
    |    `--O. transversus WP99
    `--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 KH75
         |    |--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.) vulgaris LD91
         |--A. (Carios) E-PAG91
         |    |--‘Carios’ jerseyi GE05
         |    `--A. (C.) vespertilionis Latreille 1802 E-PAG91
         `--A. (Persicargas) KH75
              |--A. (P.) arboreus DJS91
              |--A. (P.) nullaborensis Hoogstraal & Kaiser 1973 KH75
              |--A. (P.) persicus KH75
              |--A. (P.) robertsi Hoogstraal et al. 1968 KH75
              `--A. (P.) walkerae DJS91

*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 202-207.

[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.

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

[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, 3rd 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, 3rd 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.

[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 Acarology: Proceedings of the 10th International Congress (R. B. Halliday, D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff, eds) 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 Acarology: Proceedings of the 10th International Congress (R. B. Halliday, D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff, eds) 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.

[R01] Russell, R. C. 2001. The medical significance of Acari in Australia. In Acarology: Proceedings of the 10th International Congress (R. B. Halliday, D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff, eds) pp. 535-546. CSIRO Publishing: Melbourne.

[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).

Parasitiformes

Venter (left) and dorsum of Heatherella acanthocharis, from here.


Belongs within: Arachnida.
Contains: Opilioacaridae, Argasidae, Ixodidae, Sejoidea, Trigynaspida, Gamasina, Uropodina.

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. Lindquist et al. (2009a) expanded the Parasitiformes to include the Opilioacaridae, commonly treated as the sister taxon to Parasitiformes by other authors. The majority of Parasitiformes are included in the Gamasida or Mesostigmata, in which the body often has well-developed dorsal and ventral shields. The gamasidans are also often characterised by a well-developed tritosternum arising behind the gnathosoma, bearing one to three elongate laciniae, that functions together with the underside of the gnathosoma to form a fluid transport mechanism (Lindquist et al. 2009b). The Parasitiformes also include the ticks, Ixodidae, very large mites with the hypostome modified into a piercing organ with rear-facing teeth, in order to be able to feed on blood from vertebrate hosts.

Characters (from Lindquist et al. 2009a): One to four pairs of ventrolateral stigmata posterior to coxae II; coxae of legs free, usually movable; tarsi of legs II-IV with peripodomeric fissure associated with slit organs; tarsus of leg I with dense dorsal cluster of solenidiform setae subdistally (may be further elaborated into receptor organ complexes). Excluding Opilioacaridae: Tarsus of palp without terminal claws, at most with a tined clawlike structure on inner basal surface; trochanters III and IV undivided; opisthosoma with one pair of ventrolateral stigmata in region lateral to coxae II-IV or posterior to coxae IV; anus ventral or ventro-subterminal.

<==Parasitiformes [Anactinotrichida, Parastigmata] LKW09a
    |  i. s.: Plesiosejus S86
    |--Opilioacaridae S07
    `--+--Ixodida [Ixodoidea] L91
       |    |  i. s.: Alveonasus lahorensis (Neumann 1908) K91
       |    |--Argasidae S07
       |    |--Ixodidae S07
       |    `--Nuttalliella [Nuttalliellidae] K09
       |         `--N. namaqua K09
       `--+--Holothyrida [Holothyrina, Holothyroidea, Tetrastigmata] L91
          |    |  i. s.: Haplothyrus W09
          |    |         Leiothyrus W09
          |    |--Allothyridae L91
          |    |    |--Allothyrus constrictus S07
          |    |    `--Australothyrus ocellatus Hammen 1983 W09, L91
          |    `--+--Neothyrus W09 [Neothyridae L91]
          |       |    `--N. ana W09
          |       `--Holothyridae L91
          |            |--Holothyrus coccinella L91
          |            |--Sternothyrus braueri DW10
          |            |--Lindothyrus W09
          |            |--Thonius W09
          |            `--Hammenius W09
          `--Gamasida [Gamasini, Gamasoidea, Mesostigmata] L91
               |  i. s.: Paraamblyseius W01
               |         Caliphis W01
               |         Streptolaelaps heteromys Furman 1955 QV01
               |         ‘Acarus’ coleoptratorum Linnaeus 1758 [incl. A. parasitus Latreille 1795] S61
               |         Willmannia sejiformis QA91
               |         Tricholaelaps myonysognathus Grochovskaya & Mguen-Xuan-Hoe 1961 MZ91
               |         Echinolaelaps echidninus WP99
               |         Trigonuropoda [Trigonuropodidae] DW10
               |         Spelaeorhynchus [Spelaeorhynchidae] LKW09b
               |           `--S. praecursor LKW09b
               |         ‘Nicoletia’ Canestrini & Fanzago 1877 non Gervais 1842 CF77
               |           `--*N. cornuta Canestrini & Fanzago 1877 CF77
               |         ‘Acarus’ denticulatus (n. d.) CF77
               |         Trachynotus CF77
               |           |--T. cadaverinus S22
               |           |--T. pyriformis CF77
               |           `--T. troguloides [incl. T. elongatus] CF77
               |--Sejoidea L91
               `--+--Trigynaspida L91
                  `--Monogynaspida [Monogynaspides] L91
                       |--Gamasina KA90
                       |--Uropodina KA90
                       |--Heatherella WP99 [Heatherellidae, Heatherellina LKW09a, Heatherelloidea]
                       |    |--H. acanthocharis LKW09b
                       |    `--H. callimaulos WP99
                       |--Microgynioidea [Microgyniina] LKW09b
                       |    |--Nothogynus [Nothogyniidae] LKW09b
                       |    |    `--N. klompeni LKW09b
                       |    `--Microgyniidae LKW09b
                       |         |--Microgynium incisum KA90
                       |         `--Microsejus truncicola LKW09b
                       `--Heterozerconoidea [Heterozerconina] LKW09a
                            |--Discozerconidae LKW09b
                            |    |--Discomegistus pectinatus LKW09b
                            |    `--Discozercon mirabilis LKW09b
                            `--Heterozerconidae LKW09b
                                 |--Heterozercon latus LKW09b
                                 |--Afroheterozercon ancoratus LKW09b
                                 |--Amheterozercon amphisbaenae [=Zeterohercon amphisbaenae] LKW09b
                                 `--Narceoheterozercon ohioensis LKW09b

*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 202-207.

[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.

[K09] Keirans, J. E. 2009. Order Ixodida. In: Krantz, G. W. & D. E. Walter (eds) A Manual of Acarology, 3rd ed., pp. 111-123. Texas Tech University Press.

[KA90] Krantz, G. W., & B. D. Ainscough. 1990. Acarina: Mesostigmata (Gamasida). In: Dindal, D. L. (ed.) Soil Biology Guide pp. 583-665. John Wiley & Sones: New York.

[K91] Kryuchechnikov, V. N. 1991. Protective responses of Ixodoidea hemocytes. 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. 331-334. SPB Academic Publishing: The Hague.

[L91] Lehtinen, P. T. 1991. Phylogeny and zoogeography of the Holothyrida. 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. 101-113. SPB Academic Publishing: The Hague.

[LKW09a] Lindquist, E. E., G. W. Krantz & D. E. Walter. 2009a. Classification. In: Krantz, G. W. & D. E. Walter (eds) A Manual of Acarology, 3rd ed., pp. 97-103. Texas Tech University Press.

[LKW09b] Lindquist, E. E., G. W. Krantz & D. E. Walter. 2009b. Order Mesostigmata. In: Krantz, G. W. & D. E. Walter (eds) A Manual of Acarology, 3rd ed., pp. 124-232. Texas Tech University Press.

[MZ91] Meng Y.-C., Zhuge H.-X., Lan M.-Y. & Zhou H.-F. 1991. Experimental study on transmission of haemorrhagic fever with remal syndrome virus by mites, Ornithonyssus bacoti (Hirst). 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. 35-39. SPB Academic Publishing: The Hague.

[QA91] Quintero, M. T., & H. A. Acevedo. 1991. Studies on deep litter mites on farms in Mexico. 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. 443-448. SPB Academic Publishing: The Hague.

[QV01] Quintero, M. M. T., M. Vargas, B. S. Hernández, P García & N. J. Otero. 2001. Ectoparasitic mites on Heteromys gaumeri in the south of Yucatan, Mexico. In Acarology: Proceedings of the 10th International Congress (R. B. Halliday, D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff, eds) pp. 583-585. CSIRO Publishing: Melbourne.

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

[S86] Schwoerbel, J. 1986. Acari: “Hydrachnellae”. In Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) (L. Botosaneanu, ed.) pp. 652-696. E. J. Brill / Dr. W. Backhuys: Leiden.

[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.

[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.

[W01] Walter, D. E. 2001. Achilles and the mite: Zeno’s paradox and rainforest mite diversity. In Acarology: Proceedings of the 10th International Congress (R. B. Halliday, D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff, eds) pp. 113-120. CSIRO Publishing: Melbourne.

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

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

Prostigmata

Labidostomma (Nicoletiella) denticulata, copyright Walter Pfiegler.


Belongs within: Arachnida.
Contains: Bdellidae, Cunaxidae, Halacaridae, Eupodoidea, Eriophyoidea, Tydeidae, Anystae, Hydrachnidia, Trombidia, Raphignathina, Pygmephoroidea, Pyemotoidea, Dolichocyboidea, Tarsonemidae, Podapolipidae.

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 Anystae, Parasitengona and Eleutherengona, but not in the Eupodina or Labidostommatidae, 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).

The Eupodina or Eupodides are a diverse group of mites of uncertain support; they are largely united by plesiomorphic characters relative to other Prostigmata. Most are soft-bodied forms with striated integument, weak subcutaneous sclerites, and reduced opisthosomatic chaetotaxy (Walter et al. 2009). Their palps lack the thumb-claw complex found in other Prostigmata. Members of the Halacaroidea are aquatic; the remaining Eupodides are terrestrial. The Bdelloidea, snout mites, are predators of small invertebrates with the chelicerae inserted on a long subcapitular 'snout'.

The Parasitengona 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. The Eleutherengona are an ecologically diverse group in which the cheliceral bases are closely contiguous or fused, with the fixed cheliceral digits reduced or absent and the movable digits hook-like or stylet-like. They are divided between two main lineages, the Raphignathina and Heterostigmata, with females of the latter having the tracheal system opening by prolateral stigmata on the prodorsum; these are usually absent in males and juveniles (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 [Anystina, Promatides]
    |  i. s.: Eupalus B16
    |           |--E. sternalis Berlese 1916 B16
    |           `--E. subterraneus Berlese 1916 B16
    |--Eupodina [Bdellina, Eupodides] DW10
    |    |  i. s.: Caleupodes Baker 1987 AF00
    |    |--+--Bdelloidea K90
    |    |  |    |--Bdellidae OW01
    |    |  |    `--Cunaxidae K90
    |    |  `--Halacaroidea OW01
    |    |       |  i. s.: Veladeadcarus gasconi WP99
    |    |       |         Pelacarus WL09
    |    |       |         Corallihalacarus chilcottensis WL09
    |    |       |--Halacaridae S86
    |    |       `--Peza Harvey 1990 H98 [Pezidae WP99]
    |    |            |--P. daps Harvey 1990 H98
    |    |            `--P. ops Harvey 1990 H98
    |    `--+--Eupodoidea OW01
    |       `--+--Eriophyoidea OW01
    |          `--Tydeoidea OW01
    |               |--Tydeidae AF91
    |               `--Triophtydeidae [Edbakerellidae, Meyerellidae] WL09
    |                    |--Edbakerellinae [Meyerellinae] WL09
    |                    |    |--Pseudotriophtydeus AF00
    |                    |    `--Edbakerella André 2004 [=Meyerella Baker 1968 (preoc.)] WL09
    |                    |         `--‘Meyerella’ marshi AF00
    |                    `--Triophtydeinae AF00
    |                         |--Pretriophtydeus tilbrooki AF00
    |                         `--Triophtydeus Thor 1932 H98
    |                              |--T. bakeri WL09
    |                              |--T. immanis WL09
    |                              `--T. pinicolus (Oudemans 1929) [=Tydeus pinicolus] H98
    `--+--Anystae DW10
       `--+--Labidostommatidae (see below for synonymy) DW10
          |    |--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. integrum G41
          |         |         L. mamillatus (Say 1821) [=Erythraeus mamillatus, Smaris mamillatus] S61
          |         |         L. nepalense WL09
          |         |--*L. (Labidostomma) luteum Kramer 1879 SB63 [=Nicoletiella lutea WP99]
          |         `--L. (Nicoletiella) WL09
          |              |--L. (N.) cornuta WL09
          |              |--L. (N.) denticulata WL09
          |              |--L. (N.) jaquemarti WL09
          |              `--L. (N.) pacifica WL09
          `--+--Parasitengona [Engonostigmata, Parasitengonae, Parasitengonina, Parasitenini, Trombidina] DW10
             |    |  i. s.: Allotanaupodidae [Allotanaupodoidea] ZF11
             |    |--Hydrachnidia W91
             |    `--Trombidia W91
             `--Eleutherengona [Eleutherengonides, Eleutherengonina] DW10
                  |  i. s.: Cloacaroidea ZF11
                  |           |--Epimyodex WL09 [Epimyodicidae ZF11, Epimyodicinae]
                  |           |    `--E. soricis WL09
                  |           `--Cloacaridae ZF11
                  |                |--Pneumophagus bubonis WP99
                  |                |--Caminacarus theodori WL09
                  |                |--Theodoracarus testudinis WL09
                  |                |--Cloacarus faini WL09
                  |                `--Emyduracarus Fain 1968 H98
                  |                     `--E. australis Fain 1968 H98
                  |--Raphignathina LKW09
                  `--Heterostigmata [Heterostigmae, Heterostigmatina, Tarsonemida, Tarsonemina, Tarsonemini] ZF11
                       |--Pygmephoroidea CC01
                       |--Pyemotoidea K90
                       |--Dolichocyboidea WL09
                       |--Heterocheylus Lombardini 1926 H98 [Heterocheyletinae, Heterocheylidae, Heterocheyloidea WL09]
                       |    `--H. lombardinii Womersley 1960 H98
                       |--Tarsonemoidea WL09
                       |    |--Tarsonemidae K90
                       |    `--Podapolipidae WL09
                       |--Tarsocheylidae [Tarsocheyloidea] WL09
                       |    |--Tarsocheylus paradoxus WL09
                       |    `--Hoplocheylus Atyeo & Baker 1964 S76
                       |         |--H. discalis Atyeo & Baker 1964 S76
                       |         `--H. pickardi WL09
                       `--Trochometridioidea WL09
                            |--Trochometridium WP99 [Trochometridiidae]
                            |    |--T. chinensis WL09
                            |    `--T. tribulatum WP99
                            `--Athyreacaridae WL09
                                 |--Athyreacarus pleiotretus WL09
                                 `--Bembidiacarus [Bembidiacaridae] WL09

Labidostommatidae [Labidostomatidae, Labidostomatides, Labidostomidae, Labidostommatina, Labidostommatoidea, Labidostommidae, Labidostommina, Nicoletiellidae] DW10

*Type species of generic name indicated

REFERENCES

[AF91] André, H. M., & A. Fain. 1991. Ontogeny in the Tydeoidea (Ereynetidae, Tydeidae and Iolinidae). 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. 297–300. SPB Academic Publishing: The Hague.

[AF00] André, H. M., & A. Fain. 2000. Phylogeny, ontogeny and adaptive radiation in the superfamily Tydeoidea (Acari: Actinedida), with a reappraisal of morphological characters. Zoological Journal of the Linnean Society 130: 405–448.

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

[CC01] Coetzee, S. H., & A. M. Camerik. 2001. The use of autofluorescence of the pharyngeal pump complex in Pygmephoridae (Acari: Heterostigmata) as a new taxonomic aid. 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. 213–216. CSIRO Publishing: Melbourne.

[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.

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

[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.

[LKW09] Lindquist, E. E., G. W. Krantz & D. E. Walter. 2009. Classification. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology 3rd ed. pp. 97–103. Texas Tech University Press.

[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 10th 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.

[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).

[W91] Witte, H. 1991. The phylogenetic relationships within the Parasitengonae. 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. 171–182. SPB Academic Publishing: The Hague.

[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.

Trombidia

Johnstoniana parva, photographed by Andreas Wohltmann.


Belongs within: Prostigmata.
Contains: Erythraeoidea, Trombidioidea, Trombiculidae, Trombellidae.

The Trombidia include the terrestrial representatives of the Parasitengona mites. Adults of this group are hairy, often relatively large predatory mites that are often bright red in colouration. In members of the Erythraiae, the movable digit of the chelicera is styletiform and retractable; members of the Trombidiae retain a blade-like, non-retractable cheliceral digit (Walter et al. 2009).

Characters (from Witte 1991, Walter et al. 2009 [as Trombidiae + Erythraiae]): Terrestrial. Adults with idiosoma and appendages hypertrichous; idiosoma without glandularia, subcuticular glands present that release secretions in a space between epidermis and cuticle; crista metopica located within prodorsal shield; capitulum retractor 3 divided into two rami, which insert at capitulum base and orginate on the endosternite; two pairs of cheliceral retractors, with one inserting at posterior end of chelicera; intercalary segment of coxal glands located in longitudinal furrow of proximal tubercle; copulatory organ provided with a furca, with furca-proctractor inserting at proximal end and originating on anterior keel, furca-retractor originating on apodemes and on proximal chamber sclerite adjacent to apodemes; stalk of spermatophore provided with three apical rami, basal part of spermatophore droplet containing a dome of large secretion droplets which are provided with peripheral secretion filaments. Larvae with palpgenu usually bearing one seta, rarely two to four setae.

<==Trombidia We91
    |--Erythraiae [Apobolostigmata, Erythraeina] LKW09
    |    |--Erythraeoidea We91
    |    `--Calyptostoma Cambridge 1875 S61 (see below for synonymy)
    |         |--C. expalpe (Hermann 1804) S61 (see below for synonymy)
    |         |--C. lyncaea (Berlese 1887) [=Smaris lyncaea; incl. S. impressa Canestrini 1882 non Koch 1837] S61
    |         `--C. velutinum (Müller 1776) [=Acarus velutinus] S61
    `--Trombidiae [Trombidiina] LKW09
         |--Tanaupodidae [Tanaupodoidea] We91
         |    |--Lassenia Newell 1957 [Lasseniinae] We91
         |    |--Polydiscia Methlagl 1928 [Polydisciinae] We91
         |    |--Rhinothrombium nemoricola WL09, S51
         |    |--Amphotrombium jenseni WL09
         |    |--Tanaupodus Haller 1882 [Tanaupodinae] We91
         |    |    |--*T. steudeli Haller 1882 SB63
         |    |    `--T. passimpilosus Berlese 1910 SB63
         |    `--Eothrombium WL09
         |         |--E. lasseni WL09
         |         `--E. nemoricola WL09
         `--+--Chyzeriidae [Chyzerioidea] We91
            |    |--Cretessenia leoni WL09
            |    |--Napassenia aginapaica WL09
            |    |--Perumaropta [Perumaroptinae] WL09
            |    |    `--P. mirsadi WL09
            |    |--Pteridopus Newell & Vercammen-Grandjean 1964 [Pteridopodinae] We91
            |    |    |--P. auditor WL09
            |    |    `--P. pseudhannemania WL09
            |    |--Ralphaudyana Vercammen-Grandjean et al. 1974 [Ralphaudyaninae] We91
            |    |    |--R. amamiensis WL09
            |    |    `--R. iranensis WL09
            |    `--Chyzeria Canestrini 1897 S61 [Chyzeriinae We91]
            |         |--C. derricki WL09
            |         |--C. hirsti WL09
            |         |--C. novaezealandiae Hirst 1924
            |         `--C. poivrei WL09
            `--+--Trombidioidea We91
               `--Trombiculoidea We91
                    |--Trombiculidae We91
                    |--Trombellidae We91
                    |--Audyana [Audyanidae] WL09
                    |    `--A. thompsoni WL09
                    |--Leeuwenhoekiidae We91
                    |    |  i. s.: Shunsennia K90
                    |    |         Hannemania K90
                    |    |--Apoloniinae We91
                    |    `--Leeuwenhoekia Oudemans 1911 S61 [Leeuwenhoekiinae We91]
                    |         `--L. americana WL09
                    |--Neotrombidiidae We91
                    |    |  i. s.: Discotrombidium villasiense Feider 1977 WL09, DC91
                    |    |         Monunguis streblida WL09
                    |    |--Anomalothrombiinae WL09
                    |    `--Neotrombidium Leonardi 1901 S61 [Neotrombidiinae]
                    |         |--N. anuroporum WL09
                    |         |--N. barringunense WL09
                    |         |--N. beeri WWO01
                    |         |--N. neptunium Southcott 1961 (see below for synonymy) S61
                    |         |--N. samsinaki WL09
                    |         `--N. tridentifer (Ewing 1909) [=Rhyncholophus tridentifer] S61
                    `--Johnstonianidae We91
                         |--Charadracarus K90 [Charadracarinae We91]
                         |--Polydiscus [Polydiscinae] K90
                         `--Johnstonianinae SB63
                              |--Centrotrombidium Kramer 1896 SB63
                              |    `--*C. schneideri Kramer 1896 (see below for synonymy) SB63
                              |--Diplothrombium Berlese 1910 SB63
                              |    |--*D. longipalpe (Berlese 1887) [=Trombidium longipalpe] SB63
                              |    |--D. eximium S22
                              |    |--D. micidium WL09
                              |    `--D. monoense WL09
                              `--Johnstoniana George 1909 S61 [incl. Rohaultia Oudemans 1911 SB63]
                                   |--*J. errans (Johnston 1852) [=Rhyncholophus errans] SB63
                                   |--J. dubiosa Schweizer 1951 SB63
                                   |--J. insignia (Berlese 1916) (see below for synonymy) SB63
                                   |--J. parva WWO01
                                   |--J. rapax WWO01
                                   |--J. tuberculata Schweizer 1951 [=J. tuberculatua] SB63
                                   `--J. ventripilosa Wi91

Calyptostoma Cambridge 1875 S61 [Calyptostomatidae We91, 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]

*Centrotrombidium schneideri Kramer 1896 [incl. Diplothrombium walteri Schweizer 1922, C. walteri] SB63

Johnstoniana insignia (Berlese 1916) [=Diplothrombium eximium insignius; incl. D. longipes Schweizer 1922] SB63

Neotrombidium neptunium Southcott 1961 [=N. tridentifer Southcott 1957 non Rhyncholophus tridentifer Ewing 1909] S61

*Type species of generic name indicated

REFERENCES

[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.

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

[LKW09] Lindquist, E. E., G. W. Krantz & D. E. Walter. 2009. Classification. In: Krantz, G. W. & D. E. Walter (eds) A Manual of Acarology, 3rd ed., pp. 97-103. Texas Tech University Press.

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

[S51] Schweizer, J. 1951. Die Landmilben des schweizerischen Nationalparkes. 2. Teil: Trombidiformes. Ergebnisse der wissenschaftlichen Untersuchung des schweizerischen Nationalparks [Résultats des recherches scientifiques entreprises au Parc National Suisse] 3 (Neue Folge): 51-172.

[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.

[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. Band IV, Lieferung I. 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, 3rd ed., pp. 233-420. Texas Tech University Press.

[We91] Welbourn, W. C. 1991. Phylogenetic studies of the terrestrial Parasitengona. 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. 163-170. SPB Academic Publishing: The Hague.

[Wi91] Witte, H. 1991. The phylogenetic relationships within the Parasitengonae. 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. 171-182. SPB Academic Publishing: The Hague.

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

Erythraeoidea

Adult male Sphaerotarsus allmani, from Southcott (1961).


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

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. The 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 (Southcott 1961).

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
    |--Smarididae [Smaridae, 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
    |    |--Smaridinae S61
    |    `--Hirstiosomatinae S61
    |         |--Trichosmaris jacoti K90, WL09
    |         |--Clipeosoma Southcott 1948 S61
    |         |    |--*C. copiolarum Southcott 1948 S61
    |         |    `--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
    |         `--Sphaerotarsus Womersley 1936 S61
    |              |--S. leptopilus Womersley & Southcott 1941 S61
    |              |--S. allmani Womersley 1936 S61
    |              `--S. womersleyi Southcott 1946 [=S. ripicolus Womersley & Southcott 1941 (nom. inv.)] S61
    `--Erythraeidae [Erythreidae, Leptidae, Leptides, Leptoidae, Rhyncholophini, Rhyncholophoidae] G91
         |  i. s.: Morieria Oudemans 1941 (n. d.) (see below for synonymy) S61
         |           |--*M. novaehollandiae Oudemans 1941 (n. d.) (see below for synonymy) S61
         |           `--‘Erythraeus (Erythrolophus)’ mollis Berlese 1923 S61
         |         Cloquetia Oudemans 1941 S61
         |           `--*C. medioareolata (Kramer 1898) [=Rhyncholophus medioareolatus] S61
         |         Pseudoerythraeus Haitlinger 2004 F05
         |           `--*P. ernesti Haitlinger 2004 F05
         |         Proterythraeus [Proterythraeidae] WL09
         |           `--P. southcotti WL09
         |         Phanolophus André 1927 S61 (see below for synonymy)
         |           |--P. oedipodarum (Frauenfeld 1868) (see below for synonymy) S61
         |           `--*Platylophus’ argasoides Berlese 1910 [=*Euplatylophus argasoides] S61
         |         Augustsonella Southcott 1961 G91 [=Augustsonia Southcott 1961 S61; Augustsonellinae WL09]
         |           `--*A. tuberculata (Augustson 1940) [=Erythraeus tuberculatus, *Augustsonia tuberculata] S61
         |         Actineda S22
         |           |--A. pallescens CF77
         |           `--A. vitis S22
         |         Scyphius cerinus [incl. S. pratensis, S. pyrroleucus] CF77
         |--Erythraeinae G91
         `--+--Balaustiinae G91
            |--Callidosomatinae G91
            `--+--Leptus G91
               `--Myrmicotrombium Womersley 1934 S61 [=Myrmicothrombium (l. c.) S61; Myrmicotrombiinae G91]
                    `--*M. brevicristatum Womersley 1934 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]

Morieria Oudemans 1941 (n. d.) [=Coerythrolophus Radford 1950, Erythrolophus Berlese 1920 non Swinhoe 1892] S61

*Morieria novaehollandiae Oudemans 1941 (n. d.) [=Erythraeus (*Erythrolophus) froggatti Berlese 1920 non E. froggatti Oudemans 1910, *Coerythrolophus froggatti] S61

Phanolophus André 1927 S61 [=Phanalophus (l. c.) S61; incl. Euplatylophus Radford 1950 S61, Euplathylophus (l. c.) S61, Platylophus Berlese 1910 nec Swainson 1832 nec Koch 1839 S61, Platerythraeus (l. c.) S61; Phanolophinae WL09]

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 202-207.

[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.

[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.

[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, 3rd ed., pp. 233-420. Texas Tech University Press.

Leptus

Leptus sp., photographed by Tom Murray.


Belongs within: Erythraeoidea.

Leptus is a cosmopolitan genus of erythraeid mites in which the single pair of eyes is placed relatively close to the front of the main body. Larvae are found on a wide range of hosts, including almost all terrestrial arachnids and insects.

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 H98 [incl. Achorolophus Berlese 1881 H98, Ritteria Kramer 1877 S61; Leptinae]
    `--L. (Leptus) H09
         |--*L. (L.) phalangii (de Geer 1778) [=Acarus phalangii] S61
         |--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 S61
  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 JM11
  L. chibai Shiba 1976 S76
  L. clethrionomydis Haitlinger 1987 S91a
  L. coccineus (Scopoli 1763) (see below for synonymy) S61
  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 CM07
  L. hirtipes (Berlese 1910) [=Abrolophus hirtipes] V25
  L. hozumii Shiba 1976 S76
  L. ignotus (Oudemans 1903) [=Erythraeus ignotus, 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) MF11 [=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
  ‘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. 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. warregense (Hirst 1928) [=Belaustium (l. c. for Balaustium) warregense] H98
  L. yoshimii Shiba 1976 S76

Leptus coccineus (Scopoli 1763) [=Pediculus coccineus, L. coccineous (l. c.), 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, 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).

[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.

[JM11] Judson, M. L. I., & J. Mąkol. 2011. Pseudoscorpions (Chelonethi: Neobisiidae) parasitized by mites (Acari: Trombidiidae, Erythraeidae). Journal of Arachnology 39 (2): 345–348.

[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. Band IV Lieferung I. 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 3rd 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 10th International Congress pp. 83–99. CSIRO Publishing: Melbourne.