A revision of the family Ameroseiidae (Acari, Mesostigmata), with some data on Slovak fauna

Abstract The family Ameroseiidae Evans, 1961 (Acari: Mesostigmata) includes a total of 12 valid and adequately described genera, namely Afrocypholaelaps Elsen, 1972, Ameroseiella Bregetova, 1977, Ameroseius Berlese, 1904, Asperolaelaps Womersley, 1956, Brontispalaelaps Womersley, 1956, Epicriopsis Berlese, 1916, Hattena Domrow, 1963, Kleemannia Oudemans, 1930, Neocypholaelaps Vitzthum, 1942, Pseudoameroseius gen. n., Sertitympanum Elsen & Whitaker, 1985 and Sinoseius Bai & Gu, 1995. One of these genera includes subgenera, namely Kleemannia (Primoseius) Womersley, 1956. All genera are reviewed and re-diagnosed, and a dichotomous key is provided for their identification. Ameroseius (50 species), Kleemannia (28 species) and Neocypholaelaps (22 species) are the largest genera in the family. Ameroseiella, Kleemannia, Kleemannia (Primoseius) and Sinoseius are considered to be valid taxa and, in presented systematic classification, they are removed from synonymy with Ameroseius. The genus Pseudoameroseius gen. n., with type species Ameroseius michaelangeli Moraza, 2006 (from Canary Islands), is newly erected to further refine broad primary concept of Ameroseius as understood by some former authors (Karg, Bregetova). Asperolaelaps is removed from synonymy with Neocypholaelaps. Three new species are here described, namely Ameroseius renatae sp. n. (based on specimens from Slovakia), Kleemannia dolichochaeta sp. n. (from Spain) and Kleemannia miranda sp. n. (from U.S.A.). The following new junior synonymies are proposed: Ameroseius apodius Karg, 1971 = Ameroseiella macrochelae (Westerboer, 1963); Ameroseius bregetovae Livshits & Mitrofanov, 1975 = Neocypholaelaps favus Ishikawa, 1968; Ameroseius chinensis Khalili-Moghadam & Saboori, 2016 = Ameroseius guyimingi Ma, 1997; Ameroseius crassisetosus Ye & Ma, 1993, Ameroseius qinghaiensis Li & Yang, 2000 and Ameroseius norvegicus Narita, Abduch & Moraes, 2015 = Ameroseius corbiculus (Sowerby, 1806); Ameroseius dubitatus Berlese, 1918 = Kleemannia plumosa (Oudemans, 1902); Ameroseius eumorphus Bregetova, 1977 and Kleemannia potchefstroomensis Kruger & Loots, 1980 = Kleemannia pseudoplumosa (Rack, 1972); Ameroseius gilarovi Petrova, 1986 = Kleemannia plumigera Oudemans, 1930; Ameroseius imparsetosus Westerboer, 1963 = Ameroseius georgei (Turk, 1943); Ameroseius lanatus Solomon, 1969 and Ameroseius fimetorum Karg, 1971 = Kleemannia tenella (Berlese, 1916); Ameroseius lanceosetis Livshits & Mitrofanov, 1975 = Kleemannia pavida (C. L. Koch, 1839); Ameroseius marginalis Fan & Li, 1993 and Ameroseius sichuanensis Fan & Li, 1993 = Kleemannia insignis (Bernhard, 1963); Ameroseius pseudofurcatus Livshits & Mitrofanov, 1975 = Ameroseius furcatus Karg, 1971; Ameroseius stramenis Karg, 1976 and Lasioseius (Lasioseius) gracilis Halbert, 1923 = Kleemannia delicata (Berlese, 1918); Epicriopsis langei Livshits & Mitrofanov, 1975 and Epicriopsis baloghi Kandil, 1978 = Epicriopsis palustris Karg, 1971; Epicriopsis rivus Karg, 1971 = Epicriopsis mirabilis Willmann, 1956; Neocypholaelaps ewae Haitlinger, 1987 = Neocypholaelaps indicus Evans, 1963; Neocypholaelaps lindquisti Prasad, 1968, Afrocypholaelaps ranomafanaensis Haitlinger, 1987 and Afrocypholaelaps analicullus Ho, Ma, Wang & Severinghaus, 2010 = Afrocypholaelaps africanus (Evans, 1963); Sinoseius pinnatus Huhta & Karg, 2010 = Sinoseius lobatus Bai, Gu & Fang, 1995. Ameroseius womersleyi Mašán, is a replacement name proposed for Ameroseius ornatus Womersley, 1956, a junior secondary homonym of Cornubia ornata Turk, 1943 [= Ameroseius corbiculus (Sowerby, 1806)]. Cornubia georgei Turk, 1943 is removed from synonymy with Ameroseius corbiculus (Sowerby, 1806). An annotated catalogue of the world species of Ameroseiidae is provided, partly based on type (in more than 60 species) and non-type specimens from various museum deposits and personal collections, including new or revised material from Slovakia. It contains 206 named species (138 valid species, 37 synonyms, nine unrecognizable species, 15 species previously excluded from Ameroseiidae, and seven “nomina nuda”), with details of their authorship, synonyms, nomenclatural and bibliographic details, generic placement, and morphology. Altogether 23 new combinations are proposed. The genus Sertitympanum with Sertitympanum nodosum (Sheals, 1962) and two further species, namely Kleemannia kosi El-Badry, Nasr & Hafez, 1979 and Kleemannia parplumosa Nasr & Abou-Awad, 1986, are reported from Europe for the first time. New keys are given for identification of 37 species belonging to eight genera which have been found in Europe to date (Ameroseiella, Ameroseius, Epicriopsis, Kleemannia, Neocypholaelaps, Pseudoameroseius gen. n., Sertitympanum and Sinoseius). All of these genera except Pseudoameroseius gen. nov. and Sertitympanum occur in Slovakia. So, the fauna of Slovakia includes six genera and 27 species, including ten first reports for the country.


Introduction
Mites of the family Ameroseiidae Evans, 1961 are commonly found in greatly heterogeneous substrates and habitats. A large and diverse ameroseiid fauna is saproxylic and mycophagous (fungivorous), thereby depending upon dead and decaying wood and wood-inhabiting fungi (including ambrosia fungi), or the presence of fresh fruiting bodies (sporocarps) of wood-destroying bracket fungi or other saproxylic animals such as bark and wood-boring beetles. It uses snags, stumps, logs, subcortical area, insect galleries, stems and other forms of woody material for food, foraging, shelter, development, reproduction and other life activities (Westerboer and Bernhard 1963, De Leon 1964, Flechtmann 1985, Mašán 1998. Some ameroseiid species in tropical areas are nectar-and pollen-feeders, associated with various flowers and flower-visiting animals, namely bees and wasps (Hymenoptera, Apoidea), butterflies (Lepidoptera), and rarely birds (Evans 1963a;Elsen 1972aElsen , 1972bElsen , 1973Klimov et al. 2016;Allred 1970). Along with saproxylic and plant-inhabiting ameroseiids, there is a large group of aerial and potentially cosmopolitan species that scavenge for fungal spores and hyphae in various synanthropic habitats such as inside of human houses (especially new buildings and those under construction), office blocks, cellars, farms, granaries, food and product storages, haystacks, and arable lands (Rack 1963(Rack , 1972Hughes 1976). A rather small group of ameroseiid mites contains edaphic species associated with humid soils with high content of raw humus. Phoretic associations with insects and other animals have been established for many different species of the family, but they are not common among the species of the European fauna. It is assumed that many ameroseiids are non-phoretic species widely distributed by wind and air currents.

Materials and methods
The essential part of this paper consists of a revision and catalogue of supraspecific and specific taxa arranged in alphabetical order with basic bibliographical references to taxonomic nomenclature and previously published records on each species. For many species short taxonomic notes are given. I conducted a comprehensive search for data on Ameroseiidae in the world literature, up to April 2017, using a range of electronic and print resources. In addition, I examined type specimens of more than 60 species treated herein (see "Comparative material" in individual species), including published or unpublished non-type material from various European countries and Iran, and borrowed from the existing slide collections in a number of zoological and museum institutions (see abbreviations below). I tried to base my conclusion on the direct examination of type specimens wherever possible. In some cases (especially in China), I was unable to contact the relevant curators, and in other cases the type material has been destroyed or lost. The available information on some species and genera is still incomplete, and many of the existing species cannot be properly identified. However, the revision presented here should provide a basis for further study of the family in the future. Altogether almost 1,250 specimens found in 130 samples are newly reported from Slovakia. If accessible, older material, pub lished by Slovak authors in the past, was re-examined and revised, particularly for rare species. Names of incorrectly identified species are also given, within the survey of revised data. A variety of quantitative methods (mainly a core sampler or sifting) and individual collecting were used. Quantitative collection methods include extraction of heterogeneous samples and organic substrates including grassland soils, soils from wet alluvial or littoral habitats, the rhizosphere of herbs, fresh or old dung, compost, and decaying plant or animal remains, etc. Sifting was used to obtain living mites from leaf and needle litter, soil detritus, raw humus, alluvial litter deposits, moss and rotting wood. Mites were extracted from soil samples and sifted detritus by means of a modified Berlese-Tullgren funnel provided with a 40-watt bulb. The extraction lasted 48-72 hours. Individual collecting was especially used to obtain mites living on bracket fungi and decaying wood, and under bark of dying trees. The mites were transferred to tubes with 70% ethanol using fine forceps. The type material of the new species is deposited at the Institute of Zoology, Slovak Academy of Sciences, Bratislava.
For identification, specimens were mounted on permanent microscope slides using Swan's medium (gum arabic/chloral hydrate). Illustrations were made using photographed specimens. A Leica DM 1000 light microscope equipped with a stage-calibrated ocular micrometer and a Leica EC3 digital camera was used to obtain measurements and photos. Measurements were made from slide-mounted specimens. Some multiple images were combined using the CombineZP software program (Hadley, 2010). Lengths of shields were measured along their midlines, and widths at their widest point (if not otherwise specified in the description). Dorsal setae were measured from the bases of their insertions to their tips. Measurements are mostly presented as ranges (minimum to maximum).
The nomenclature used for the dorsal idiosomal chaetotaxy is based on that of Lindquist and Evans (1965); leg chaetotaxy is that of Evans (1963b), and other anatomical structures follow Evans and Till (1979). The chaetotaxy symbols used here are illustrated in Fig. 1. Some dorsal shield setae are difficult to recognise, and I have identified them on the basis of their relative positions, not as first appears during ontogenetic development in the developmental stages. In some cases I have amended specific names to comply with the International Code of Zoological Nomenclature.

On leg chaetotaxy in the Ameroseiidae
I have examined the leg chaetotaxy patterns in all ameroseiid species found in Slovakia (27 species), and 11 further species from outside Slovakia, and the results are summarised in Tables 1-3. The examination was motivated by the existence of many dissonant and inconsistent data on leg setation available in world literature on Ameroseiidae. In addition, chaetotaxy of genu III has been checked in all species available for this study.

On ecology and habitat preference of Slovak Ameroseiidae
According to specific microhabitat requirements, the Slovak species of Ameroseiidae can be classified into the following ecological groups: (1) Edaphic species (eight species, 29.6% of the total). This group includes all representatives of Epicriopsis, and several species of Ameroseius: Ameroseius cavernosus, Ameroseius corbiculus, Ameroseius lidiae and Ameroseius sculptilis. With the excep-tion of A. cavernosus, A. corbiculus and Epicriopsis horridus, all of these representatives are apparently hygrophilous, inhabiting permanently moist places of wetlands or periodically flooded areas like littoral reed stands of shores, bogs, flood plains, riverine inundation zones and swamps (they can apparently survive in soaked substrates). Edaphic ameroseiids are bound to various kinds of soil substrates, such as highly humic soils, raw humus, decomposing plant substances, leaf and needle litter, moss, etc. Epicriopsis horridus is the only Slovak Epicriopsis species that can be considered as mesohygrophilous. It colonises also relatively drier but often umbriferous woodland stands. Epicriopsis species occur mostly in lowlands, only Epicriopsis mirabilis colonises cold mountaineous areas. Ameroseius corbiculus is the most common and most widely distributed ameroseiid species in Slovakia, although not in cold stands in montane zone. It occurs in various microhabitats but it prefers substrates with high content of raw humus and decaying plant parts, mainly in lowlands and low highlands, at altitudes with an optimum up to 600 m. It can also penetrate into cultivated landscape habitats (orchards, gardens and degraded stands). A. cavernosus seems to be the only xerotolerant species exclusively found in relatively warmer and drier localities of the southern part of Slovakia.
(2) Saproxilic species (eight species, 29.6% of the total). They are adapted to living in rotting wood with growth of wood-inhabiting fungi, including sporocarps of wood-destroying fungi, snags, stumps, stems, subcortical area of dying trees, and galleries of xylophagous insects. This group in cludes several Ameroseius species, namely Ameroseius callosus, Ameroseius corniculus, Ameroseius fungicola, Ameroseius furcatus, Ameroseius georgei, Ameroseius longitrichus, Ameroseius renatae sp. n. and Ameroseius ulmi. Interesting cases of saproxilic detriticoles appear to be the strictly specialised mycetobionts A. callosus and A. fungicola exclusively associated with the fruiting bodies of polypore shelf fungi (Holobasidiomycetidae, Polyporales), permanently dwelling fungal sporocarps. (3) Saprophilous species (two species, 7.4% of the total). They include species showing affinity to strongly decaying organic matter in dunghills, excrements and compost heaps. They facultatively occur in other substrates consisting of a portion of excrements or decomposing plant and animal substances, such as manured arable soils, bird nests, heterogeneous organic refuses, etc. This ecological group includes only Ameroseiella macrochelae and Kleemannia tenella. (4) Aerial species (eight species, 29.6% of the total). Especially in species with foliately expanded dorsal setae, spreading by air currents (passive air-borne dispersal) seems to be a very important part of the life history strategy of these mites (real evidence for this phenomenon has not been confirmed up to now), and ensures the continued existence of species that occur in ephemeral and scattered habitats such as mould growths living on diverse organic substrates. A relatively larger group of species that colonise manifold natural and synanthropic microhabitats such as interior of human buildings, cellars, farms, stored food, granaries, dumping grounds, haystacks, vertebrate nests and agrocoenoses. I incorporate here several species as follows: Kleemannia delicata, Kleemannia insignis, Kleemannia pavida, Kleemannia plumea, Kleemannia plumigera, Kleemannia plumosa, Kleemannia pseudoplumosa and Sinoseius lobatus. Occasionally, some aerial species can be found in caves. For example, Kleemannia plumigera and Sinoseius lobatus were found in the guano, baited straw or soil detritus in Slovakian caves. (5) Insecticolous species (one species, 3.7% of the total). This ecological group includes species with well-developed phoretic activity and that spend all or part of their life cycle on a host insect or in its nest. Neocypholaelaps favus, as the only representative of this group in Slovak fauna, lives in association with European honey bee Apis mellifera Linnaeus, 1758 and its nests (hives), and feeds probably on pollen of flowers visited by their bees carriers.

Catalogue of world species of Ameroseiidae
I present below an alphabetic list of all genera and lower taxa of Ameroseiidae that I am aware of, together with original identification keys to world genera and the species occurring in Europe. For each species (except three newly described species), the following data are provided: (1) current generic combination of the species, author(s) and year of the original description, (2) name of the species in its original combination, with reference to the author, year and the page on which the original description begins, (3) references to subsequent literature on the species, including different names used in diverse systematic concepts, (4) synonyms if any, each followed by its author, year and page number on which the description of the synonymised species begins, and the reference in which the synonymy is established, (5) extra information on synonymy and other nomenclatural problems where necessary, (6) type depository, name of the institution where the type material is deposited, (7) type locality and habitat from which the name-bearing type specimens are collected, (8) comparative material, type or non-type specimens which were available for comparative purpose of this study, (9) material from Slovakia, references to the older faunistic records, together with complete collection data on revised and newly reported specimens if found in Slovakia, (10) occasional remarks are provided to explain some complicated taxonomic or morphological problems. There are also similar data provided for each genus (see previous points 2, 4, 5 and 10), including newly elaborated morphological diagnosis for each genus. Some species are not included in the genus classification because they are not described in enough detail, or unavailable for examination, and these are listed separately, after the presentation of the valid species, as (1) unrecognizable species (species inquirendae), by the lack sufficient information about important characters needful for their specific identification, (2) species excluded from Ameroseiidae that have been placed in Ameroseius in the past, but are now placed in the Ascidae, Blattisociidae, Phytoseiidae and Ologamasidae, (3) nomina nuda, names of the species which were apparently never described in any of published papers, and unavailable according to the International Code of Zoological Nomenclature, and (4) other unavailable names.
Diagnosis (adults). Idiosoma oblong and dorsoventrally flattened. Dorsal shield entire, holotrichous, having at most 30 pairs of setae; setae J1 and J5 absent. Sternal region with separate sternal and epigynal shields in female, sternitogenital shield in male, and five pairs of sternal setae in both sexes (st1-st5). Sternal shield reduced in size, but entire, with 1-3 pairs of sternal setae (normally two pairs); presternal platelets absent. Metasternal setae (st4) on soft integument or endopodal platelets. Epigynal shield with a pair of genital setae (st5); genital opening of male with presternal position. Peritremes well developed, long. Posterior ventral surface of adults with anal or ventrianal shield, and with at least four pairs of opisthogastric setae. Three circum-anal setae present. Cheliceral digits short; arthrodial membrane at base of movable digit smooth, without brush-like or filamentous processes. Palptarsal apotele with two or three tines. Hypostome with internal malae usually unmodified and lightly fringed. Tibia I with five dorsal setae; legs II of males without spur-like structures; tarsus II in female usually without enlarged spine-like distal setae; genu IV with normally nine setae, including one ventral (but aberrations exist). Mites with podospermal insemination, spermatodactyl of male present and placed on movable digit of chelicera. Female monogynaspid.
Key to world genera of Ameroseiidae (females) 1 Ventrianal shield with three pairs of opisthogastric setae (JV1, JV2, ZV2) and three circum-anal setae; endopodal platelets II-III and III-IV connected, bearing metasternal setae (st4) Diagnosis (female). Dorsal shield weakly sclerotised, smooth medially and reticulate laterally, oblong, with 29 pairs of subequal setae. Dorsal setae including j1 short and needle-like, mostly smooth, sometimes delicately pilose to serrate. Setae st1 and st2 on sternal shield, st3 on small pseudo-metasternal platelets and st4 on soft integument. Sternal and epigynal shield relatively long and narrow, smooth and unornamented on surface. Epigynal shield with anterior hyaline portion produced into a prominent cusp reaching the level of st2; genital poroids on soft integument, outside the shield. Metapodal platelets absent. Anal shield relatively small, subcircular, bearing three circum-anal setae. Peritrematal shields or peritremes with anterior ends free, not fused to dorsal shield. Opisthogastric soft integument with six pairs of setae (JV1-JV5, ZV2). Soft striate integument smooth, not incrusted with sclerotic denticles or tubercles. Corniculi slender and convergent, surrounded by hyaline membranes, with undivided and pointed apex. Fixed digit of chelicera unidentate, having hyaline lobed appendage; movable digit edentate, well hooked distally. Epistome with curved and denticulate anterior margin. Palptarsal apotele two-tined. Femur II with two ventral setae, genua II-III and tibiae II-III each with one anterolateral and one posterolateral seta. Tarsi I-IV each with well developed empodium but reduced claws (the claws normal in males and developmental stages). Insemination apparatus with spermathecal ducts entering sacculus foemineus through a common neck-like process of the sacculus. Remark. The genus Afrocypholaelaps is distributed in the tropical and subtropical areas of the Old World and Oceania (Australia, Papua New Guinea, Hawaii Islands, Japan, Taiwan, Saint Helena Island, Angola and Madagascar). Mites of this monobasic genus live on flowers of various plants where they probably feed on pollen and nectar. Phoresy of females is reported from European honey bee (Apis mellifera), bees and wasps of various apoid genera (Meliponula, Ceratina, Hylaeus and Chlorion), and other flower-visiting insects (Lepidoptera). (Evans, 1963) Hypoaspis sp. -Simmonds 1949: 42. By Halliday 1997. Neocypholaelaps sp. -Brimblecombe and Roff 1960: 447. By Halliday 1997. Neocypholaelaps africana Evans, 1963a: 224. Neocypholaelaps lindquisti Prasad, 1968. Syn. n. Afrocypholaelaps africana. -Elsen, 1972b: 159;Seeman and Walter 1995: 45;Halliday 1997: 181;Ho et al. 2010: 90. Afrocypholaelaps lindquisti. -Haitlinger 1987aHalliday 1997: 181. Neocypholaelaps africanus. -Domrow 1979: 105. Neocypholaelaps africana. -Ishikawa 1979Baker and Delfinado-Baker 1985: 232;Delfinado-Baker et al. 1989: 610;Karg 1993: 233. Neocypholaelaps lindquisti. -Treat 1975Baker and Delfinado-Baker 1985: 232. Afrocypholaelaps ranomafanaensis Haitlinger, 1987b: 531 Remarks. There are some contradictory statements on the leg chaetotaxy of this species in some available papers on the genus Afrocypholaelaps. The leg chaetotaxy for Afrocypholaelaps africanus in Evans (1963a) is incomplete. His figures (1-4, page 211) show the chaetotaxy of Neocypholaelaps, with three ventral setae on femur II, and two anterolateral setae on genu II, tibia II, genu III and tibia III. He then mentioned that A. africanus has only one anterolateral seta on genu III and tibia III (page 213, and 225). He did not mention the deficiency of a ventral seta on femur II, and one anterolateral seta on genu II and tibia II. Prasad (1968) misunderstood this. He thought A. africanus had three ventral setae on femur II and two anterolateral setae on genu II and tibia II. That is why he thought A. africanus had more leg setae than the species he then described as Afrocypholaelaps lindquisti. These two species actually have the same chaetotaxy, and they are here considered to be conspecific. Later, in his original generic diagnosis of Afrocypholaelaps, Elsen (1972b) re-examined the type species A. africanus, stating the correct number of two ventral setae on femur II, and one anterolateral and one posterolateral seta on genu II and tibia II. This chaetotactic pattern was confirmed also by Halliday (personal communication), who additionally checked his Australian specimens of A. africanus, although his previous chaetotactic data (Halliday 1997) showed the same misinterpretation of Prasad (1968) and Ho et al. (2010).

Afrocypholaelaps africanus
Afrocypholaelaps ranomafanaensis was quite briefly and inadequately described on the basis of the single female specimen by Haitlinger (1987b), and his description needs various amendments because it contains several morphological features inconsistent with Afrocypholaelaps (dorsal shield with 28 instead of 29 pairs of setae, soft integument with four instead of six pairs of opisthogastric setae, st3 on soft integument instead of on small shields). The only further differences to the other congeners of Afrocypholaelaps that he noted were the relatively larger dorsal shield (488 × 280 vs 350-436 × 216-279 μm respectively) and the shields situated on ventral surface (sternal shield 120 × 90 vs 68-84 × 68-78 μm, epigynal shield 134 × 86 vs 100-173 × 53-60 μm, and anal shield 84 × 92 vs 52-69 × 62-78 μm) in his own specimen from Madagascar. I examined the holotype specimens of A. ranomafanaensis and found no important morphological differences with specimens described by various authors under the name Afrocypholaelaps (Evans 1963a, Prasad 1968, Elsen 1972b, Domrow 1979, Ishikawa 1979, Seeman and Walter 1995, Halliday 1997, and Ho et al. 2010. Despite the fact that the holotype is not in perfect condition for observation, I could detect full complement of 29 pairs of dorsal shield setae typical of Afrocypholaelaps. Setae s1 and s2 are asymmetrically expressed in right side of the dorsal shield, as their bases are markedly adjacent each other. Dorsal setae are quite short and similar in length (anterior and marginal dorsal setae 12-14 μm, posteriormost dorsal setae 10-12 μm, Z5 11 μm), with otherwise position and relative length as in original illustration. Posteroventral region possesses the complete set of four pairs of slightly asymmetrically situated opisthogastric setae (JV1-JV3, ZV2) of which left JV2 displaced somewhat posteriorly and left JV3 abnormally placed on anal shield. Remaining two pairs of opisthogastric setae can be found on dorsal marginal surface, but I have detected only two of them (JV4, JV5) on the left side of the holotype. Ventrally placed setae clearly longer than those on dorsum (JV1 27-29 μm; JV2, JV3 and ZV2 16-23 μm). Setae st3 inserted on very small suboval pseudo-metasternal platelets. Anal shield not regularly rounded, but with posterior portion slightly expanded. I have checked the size of the all above mentioned shields, and can report holotype specimen with lower values for these measurements: dorsal shield 488 × 280 vs 428 × 248 μm respectively, sternal shield 120 × 90 vs 74 × 77,5 μm (at the level of st2), width of epigynal shield 86 vs 80 or 69 (at the level of genital setae), and anal shield 84 × 92 vs 78 × 89 μm. I believe that slight differences in size of scutal structures and setal length, found in different world populations of Afrocypholaelaps and documented by various above cited authors can be interpreted as variation within a widespread species. Seeman and Walter (1995) examined the holotype and paratype of Afrocypholaelaps africanus and a rich material of the specimens from Australia, finding no morphological differences between them. I believe that also A. ranomafanaensis and A. africanus cannot be distinguished morphologically, and are considered here to be conspecific.
Afrocypholaelaps analicullus was characterised by the features of hypostome, anal shield and adjacent soft integument (Ho et al. 2010), and introduced as a species closely related with Afrocypholaelaps lindquisti. The most important diagnostic characters of A. analicullus were mentioned to be (1) setae h2 and h3 similar in length; (2) ventral hypostome with five short rows of denticles on each side; (3) anal shield expanded posteriorly, with V-shaped cribrum; (4) setae JV4 and JV5 inserted close to the anal shield; in A. lindquisti, these characteristics were mentioned as (1) setae h2 longer than h3; (2) ventral hypostome with one short and two long rows of denticles; (3) anal shield rounded, with U-shaped cribrum; (4) setae JV4 and JV5 distant from the anal shield. Undoubtedly, Ho's interpretation of A. lindquisti was based on the original descriptions and illustrations, and not on study of the type material. The above enumerated distinguishing characters do not provide a useful basis for establishment of a new species, so I relegate A. analicullus into synonymy with Afrocypholaelaps africanus. For example, the position of JV4 and JV5, and their distance from the anal shield, depends on an expansion of soft integument. The soft integument is well striated in Afrocypholaelaps, and it could conceivably increase its surface considerably in gravid females, as it is originally depicted in A. lindquisti by Prasad (1968). Also anal shield may vary in size and shape, as documented in A. africanus by illustrations of Evans (1963a) and Ishikawa (1979). Except for the leg chaetotaxy, the only reliable difference between A. africanus and other two species, A. lindquisti and A. analicullus, appeared the number of serrated setae on the posteriormost surface of the dorsal shield. Evans (1963a) stated three pairs of serrate setae in A. africanus, whereas Prasad (1968) and Ho et al. (2010) reported six pairs of such setae for A. lindquisti and A. analicullus, respectively. But according to Elsen (1972b), the former species actually has the same number of serrate posterior setae as the two latter ones.
Diagnosis (adults). Dorsal shield well sclerotised and coarsely ornamented, with 29 pairs of setae, including z6, similar in length and form, thickened, often flattened, with longitudinal vein and smooth or sparsely serrate lateral margins; setae j1 differently formed, leaf-like to fan-shaped, with distinctive longitudinal midrib and regularly denticulate anterolateral margin. In female, sternal setae on sternal shield (st1, st2), soft integument (st3, st4) and epigynal shield (st5); opisthogastric region with six pairs of setae (JV1-JV5, ZV2), all on soft integument (anal shield with only three circum-anal setae). In male, ventrianal shield with 3-4 pairs of opisthogastric setae (JV1 on or off the shield, JV5 always off the shield); setae JV4 absent. Opisthogastric setae mostly smooth, short and needle-like; setae JV4 pilose; setae JV5 modified and similar to those on dorsal shield. Corniculi normally sclerotised, relatively slender, with splitted apex; setae h1 markedly thickened. Cheliceral digits relatively large, fixed digit of chelicera with three well developed teeth on proximal masticatory area (two proximal teeth somewhat adjacent). Epistome with anterior margin produced into long and narrow central projection. Palptarsal apotele two-tined. Genu III, and tibiae III-IV with two anterolateral and two posterolateral setae. Tarsi of legs I with empodium and claws not developed, terminating with four conspicuously lengthened sensory setae and some shorter ones; tarsi II-IV each with normal empodium and claws. Remarks. The genus Ameroseiella Bregetova, 1977 was originally diagnosed from other ameroseiid genera by the absence of pretarsal empodium and claws on legs I. This genus was considered as a synonym of Ameroseius Berlese, 1904(Karg 1993. In the meantime, Evans and Till (1979) and Halliday (1997) accepted Ameroseiella as a distinct genus, and their concept is adopted in the present paper because the genus exhibits some character states presented by Kleemannia and others by Ameroseius. For example, the presence of anal shield and three teeth on fixed cheliceral digit is not consistent with Kleemannia, whereas the presence of two posterolateral setae on genu III and tibiae III-IV, pointed epistome, thickened h1 are the features not consistent with those found in Ameroseius.
Ameroseiella is distributed exclusively throughout the Palaearctic region, and it currently comprises two species reported from Europe and Asia. They occur in various decomposing organic materials like compost and dung, and in the nests of mammals and birds (Westerboer and Bernhard 1963, Karg 1971a, Bregetova 1977. Town, Ostrov Orliaka Morského Forest; Vojka Nad Dunajom Village, Hajóšok Forest (Fenďa and Lengyel 2007, cited as Ameroseius apodius). Svätý Jur Town, Šúr Forest (Fenďa et al. 2011, cited as Ameroseius apodius).
Remarks. Ameroseius is the most speciose genus of Ameroseiidae. In this paper, it comprises 50 valid species having their type specimens reported from almost all continents: 13 species each from Africa and Asia, 12 species from Europe, five species each from North and South America, and two species from Australia. In Slovakia, this genus is represented by 13 recorded species. Mites of this genus are apparently fungivorous living in a wide variety of habitats such as wood substrates, wood-destroying fungi, decomposing plant material and humic soils. Most of them belong to highly specialised species, each adapted to a particular environment. There are species associated with bark beetle galleries, or wood-boring beetle galleries, in subcorticolous habitats, and feeding on a specific diet of ambrosia fungi, and phoretically active on xylophagous insects (mostly Cerambycidae). Some African species are phoretically associated with wasps and bees. In Slovakia, most species can be easily found on bracket fungi, especially on a lower fertile surface of the sporocarp (fruiting body).
Some authors (Bregetova 1977, Evans andTill 1979) attempted to clarify the general concept of Ameroseius by removing some species that obviously belong to other genera (Ameroseiella, Kleemannia). In this paper I refine the concept of the genus further, by establishing a new genus based on Ameroseius michaelangeli Moraza, 2006, removing all species that belong to Kleemannia and Asperolaelaps and that were previously placed in Ameroseius, and introducing two species groups based on Ameroseius magnisetosus and Ameroseius fungicola (see below). The process of clarifying the genus should continue, especially with regard to some characters inconsistent with Ameroseius in the species from Africa described by Elsen (1973). His African species show some atypical characters for Ameroseius, for example: (1) cheliceral digits distally curved, fixed digit with five proximal denticles in Amerosieus megatritosternum, or with only one very robust medial tooth in Ameroseius bembix, Ameroseius gabonensis and Ameroseius leclercqi; (2) peritremes and peritrematal shields densely spinate, especially the outer posterior margin of peritrematal shields with large spines; (3) dorsal shield setae relatively short and stout, brush-shaped; (4) anterior margin of epistome deeply dentate; (5) setae JV5 similar to those on dorsal shield, brush-shaped; (6) strong sclerotic incrustation of soft integument on opisthogastric surface. Generally, the border between Ameroseius and Neocypholaelaps is weak, based on a few diagnostic features of gnathosoma. The species of Elsen (1973) appear to have an intermediate position between these two genera because, in some respects, they exhibit certain similarities with Neocypholaelaps, whose members are also associated with bees and wasps (form of cheliceral digits and dorsal setae, and additional sclerotization of soft integument).
There are three Asian Ameroseius species (denticulatus, magnisetosus and submagnisetosus) representing a specific group of closely related (if not identical) congeners characterised especially by a combination of the following character states: (1) dorsum with deficient chaetotaxy, having only 26-27 pairs of setae; (2) dorsal shield lacking coarse sculpture, with only delicate reticulate pattern; (3) in female, st3 on soft integument due to the absence of pseudo-metasternal platelets; (4) in male, st4 on soft integument, outside the sternogenital shield (5) five pairs of opisthogastric setae present (JV4 absent), of which JV3 on (ventri)anal shield, and JV5 similar to other setae on ventral surface. The above enumerated species are here referred to as the Ameroseius magnisetosus group. Unfortunately, I have examined no representative of this peculiar group to confirm reduced number of dorsal setae and other features as stated above.
There is a combination of diagnostic characters to recognise the newly designated Ameroseius fungicola species group, namely (1) fixed digit of chelicera bidentate, the two teeth small and similar in size; (2) corniculi unsclerotised, membranous, hyaline, medially curved, with tapered and undivided apex directed anterolaterally; (3) conspicuous dimorphism of dorsal chaetotaxy: in males, centrally situated setae strongly reduced in length when compared with those in females; (4) male with anal shield bearing only three circum-anal setae; (5) setae z5 minute; (6) absence of postgenital slit-like sclerites; (7) anus close to anterior margin of anal shield; (8) cheliceral digits relatively small; (9) legs I relatively short and thick (especially tarsi); (10) in male, legs I and palptrochanters with some setae thicker and spiniform when compared with those in female; and (11) empodium and claws of tarsi I-IV well developed, relativelly large; (12) in males, tarsal claws of legs II apparently larger than in other legs. This group contains only two described species (fungicola, callosus), and it can be characterised by several peculiar characters (see points [1][2][3][4]12), unique or rarely expressed in Ameroseiidae. From a phylogenetic point of view, presence of these characters might support the idea of the justified existence of a separate ameroseiid taxon based on A. fungicola and A. callosus. Narita et al. (2015) constituted a separate species group for 21 species of Ameroseius, referred to as the sculptilis group, with the following character states: (1) dorsal shield with ridges and pit-like depressions combined with a reticulate pattern of simple lines; (2) dorsal shield with 29 pairs of mostly stout and serrate setae; (3) posterior ventral surface with five or six pairs of opisthogastric setae; (4) ventrianal shield with 0-2 pairs of setae (in addition to three circum-anal setae). However, all these character-state arguments are weak, based on greatly variant and vague features, and not suitable for correct separation of the species, even at least at the level of Ameroseius/Kleemannia species as understood in this paper. So, I here did not follow Narita et al. (2015) in their concept of sculptilis as a separate and reasonably derived species group of Ameroseius.

Key to species of Ameroseius occurring in Europe (females)
European species of Ameroseius can be identified using keys from Westerboer and Bernhard (1963), Karg (1971aKarg ( , 1993 and Bregetova (1977). In their keys, these authors included six, eight or nine species, respectively. A smaller part of the world species of Ameroseius (17 species) can be identified using keys of Narita et al. (2015), but better portion of congeners should be exclusively identified using the primary species descriptions. A new key presented here contains 13 species considered to be of the European origin, including recently described species (Ameroseius callosus, Ameroseius fungicola and Ameroseius lehtineni), and a new species described in this study.

1
Setae JV3 on (ventri)anal shield close to its anterolateral margins; soft integument between pseudo-metasternal platelets with a pair of rounded platelets; lateral margins of epistome produced into elongate, narrow, rounded and deeply dentate apex; movable digit of chelicera terminally with short and rounded hyaline appendage .  Bregetova, 1977, if not identical. I have seen a series of photos displaying a specimen of this species from Egypt (kindly sent by R. Abo-Shnaf ). Some differential characters for separating Ameroseius aegypticus and A. lidiae could be based on dorsal chaetotaxy (in A. aegypticus, dorsal setae slightly longer and more slender when compared with A. lidiae, the distance between left and right base in setal pairs j5, J2 and J4 relatively shorter; setae Z2 only slightly longer than Z1 and Z3). Karg, 1994 Ameroseius asper Karg, 1994a: 117. Ameroseius asper. -Narita et al. 2015 Type depository. Museum für Naturkunde, Berlin, Germany.
Comparative material. Ecuador: 2 ♀♀ (ZMB: 45134, holotype; ZMB: 45135, paratype) - 6. 4. 1985, Galapagos I., Floreana, Uferzone, 85-335, 6612-6613. Remarks. In the holotype, fixed digit of chelicerae with normal set of three denticles on proximal masticatory area (originally stated five to six teeth on fixed digit). Karg, 1976 Ameroseius avium Karg, 1976 Remarks. I examined the holotype (♀) and three paratype females of the original series of Karg (1976) who has not specified the number of type specimens. All specimens examined are in poor condition for study, and only dorsal shield in those specimens can be examined with some difficulty. I was unable to check most of the characters on the venter of the idiosoma satisfactorily, especially the posterior margin of the sternal shield and the placement of st3. This species is very similar to those in the genus Sinoseius in that it has pinnate dorsal setae, curved epistome, oblong and anteriorly bilobed sternal shield, five pairs of opisthogastric setae, enlarged cheliceral digits and subcircular anal shield. Therefore, the systematic placement of Ameroseius avium in Ameroseius should be considered provisional and questionable. Karg (1976) did not state the number of dorsal shield setae; 26 pairs of setae were depicted in the original illustration, but I observed the normal complement of 29 pairs of the setae in the holotype. Karg overlooked some marginally inserted setae in medial part of the dorsal shield, and also overestimated the length of some very short setae inserted in the marginal region. Elsen, 1973 Ameroseius bembix Elsen, 1973 Remarks. I consider both subspecies to be identical although no type material of Ameroseius bembix bembix was examined for the purpose of this study. Given that the characteristics of the type specimens of Ameroseius bembix ealensis are within the range of variability found in Ameroseius leclercqi Elsen, 1973, I suspect A. bembix ealensis to be a synonym of A. leclercqi.

Ameroseius benoiti Elsen, 1973
Ameroseius benoiti Elsen, 1973 Remarks. No fundamental differences could be found between the type specimens of the four species described by Elsen (1973), namely Ameroseius benoiti, Ameroseius gillardinae and Ameroseius longitarsis. These species were erected in my opinion on untenable characters. For example, I could not detect any of the illustrated structures of sperm induction system as they were presented in original figures 1-18 given by Elsen (1973).
Remarks. Ameroseius callosus shows a close resemblance to Ameroseius fungicola, except for some characters of the dorsal shield (sculpture and relative length of setae). Although A. callosus may be easy and reliably separated from A. fungicola, several pieces of evidence show that this species could be only a polymorphic form and synonym of A. fungicola. This hypothesis should be confirmed by further comparative studies, based also on laboratory rearing and molecular analyses, to establish their correct systematic status. Polymorphism has been mentioned in a few mesostigmatic mites of the families Ascidae, Laelapidae, Macrochelidae and Parasitidae.
The dorsal shield of Ameroseius corbiculus was illustrated by Westerboer and Bernhard (1963) as having 30 pairs of quite slim setae instead of 29 pairs of apparently stout and robust setae. In addition, Karg (1971a) erroneously illustrated a male specimen of this species with six instead of five pairs of opisthogastric setae (see his page 15). These facts could lead to misinterpretation of the morphological characters resulting in an incorrect identification of this easily recognizable species.
I have examined three type specimens of Ameroseius norvegicus, a species decribed by Narita et al. (2015) from Norway. However, the slide labelled "Holotype" contains two individuals. They are both females, and are clearly conspecific, in a perfect agreement with the primary description and original illustrations. I here designate as lectotype the specimen nearer to the left-hand edge of the slide when the labels are in the upright position. I have ringed this specimen with black ink and labelled the slide with the words "Lectotype ringed". There is no doubt that these type specimens from Norway are conspecific with those widely distributed in Slovakia and other European countries (including those from Asia, Iran), and known under the name Ameroseius corbiculus. Narita et al. (2015) added complete leg chaetotaxy to the description of A. norvegicus. According to their original data, genu I and femur IV would have respectively 11 and 5 setae, but in the type specimens I found typical 12 and 6 setae instead (including two anterolateral setae on genu I, and a ventral seta on femur IV).
I have not examined type specimens of Ameroseius corbiculus (no type material exists for this species), Ameroseius crassisetosus and Ameroseius qinghaiensis, but the distinctions made in the original descriptions are based on characteristics that vary considerably, and I do not hesitate to propose the above synonymy. Karg, 1971 Plates 9, 10 Ameroseius corniculus Karg, 1971a: 233. Ameroseius corniculus. -Vasilyeva et al. 1976Bregetova 1977: 163;Karg 1993
Type locality and habitat. Vietnam, Cha-Pa, SW from Lao-Kay (at Geophysical Station), in mould under wood trunk.
Remarks. Ameroseius fungicola appears to be a specialised mycetobiont primarily colonising the fruiting bodies of the willow bracket fungus, Phellinus igniarius, where it occurs on the active hymenophore. This polypore fungus is common mostly throughout the temperate Northern Hemisphere, where it grows mainly on living willow, poplar and aspen. A. fungicola is a very common and abundant species, often with several hundred mites in a single fungus, occasionally including all post-embryonic stages.

Ameroseius furcatus Karg, 1971
Plates 14-16 Ameroseius furcatus Karg, 1971a Remarks. Ameroseius furcatus was originally described from Germany, where it was collected in a tussock of grass with roots (Karg 1971a). Karg included this species to an identification key, without adequate morphological description. The species was only very briefly diagnosed by him, and based only on two accompanying illustrations, the epistome and dorsal shield. These illustrations were partly confusing because Karg drew the epistome as a flat structure having smooth anterior margin, and the dorsal shield bearing only 24 pairs of setae. The examination of the type material of A. furcatus showed that the epistome is elongate and marginally serrate, and the dorsal shield possesses the normal 29 pairs of setae. Distinctive features of A. furcatus females include the peculiar presence of a pair of small rounded platelets between pseudo-metasternal platelets, short and rounded hyaline appendage on distal apex of movable cheliceral digit, and the insertion of JV3 on (ventri)anal shield, but they were neglected by Karg (1971a).
I believe that the inconsistent original description of Ameroseius furcatus, especially the absence of comments on specific ventral features and a misinterpretation of the dorsal chaetotaxy (Karg 1971a), led to description of the same species under the name Ameroseius pseudofurcatus. The latter has been thoroughly described and illustrated by Livshits and Mitrofanov (1975) from specimens from the Nikita Botanical Gardens, Crimea, from leaf litter under common hazel. I have not examined type specimens of that species, but I have a slide with numerous specimens (ten females, four males) from a mite collection of the Nikita Botanical Gardens, National Scientific Center, Yalta, Crimea (received from Alex Khaustov), collected in the type locality, and labelled as follows: Ameroseius pseudofurcatus, дуб пушистый, подстилка, ГНБС (Госудaрсенный Никитский ботанический сaд), 9. 1. 1973 (leaf litter under downy oak, Quercus pubescens, State Nikita Botanical Gardens). From previously pub-lished descriptions (Karg 1971a, Livshits andMitrofanov 1975), observations of the primary type of A. furcatus, and examination of conspecific material from Germany, Crimea and Slovakia I have concluded that the both above mentioned species are identical and should be considered as synonymous.

Ameroseius gabonensis Elsen, 1973
Ameroseius gabonensis Elsen, 1973 Remarks. In his paper on the genus Epicrius from Lincolnshire in England, George (1906) re-described and illustrated three species of which two are now regarded to be members of Ameroseiidae. According to Evans (1955a), the species George believed to be Epicrius mollis (Kramer, 1876) is actually Epicriopsis horridus and what George identified as Epicrius canestrinii Haller, 1881 has formerly been given the new name Cornubia georgei by Turk (1943). Moreover, Turk (1943) proposed the new genus Cornubia with his new species Cornubia ornata as type species. Later,  regarded Cornubia as being synonymous with Ameroseius. At the same time, he synonymised the two apparently different species, originally included in Cornubia (C. ornata and C. georgei), with Ameroseius corbiculus. This interpretation is accepted here only in part because C. georgei is considered to be a distinct and reliably distinguishable species. Therefore, it is removed from a synonymy with A. corbiculus. George's original illustration in his paper (1906) is sufficiently detailed for the recognition of C. georgei. This species can be characterised as having dorsal setae extremely long (for example, j6 almost reaching posterior margin of idiosoma, and J2 reaching apparently beyond posterior margin of idiosoma). Ameroseius imparsetosus, based on a female from Spain and described by Westerboer (in Westerboer and Bernhard 1963), is in perfect agreement with what is reported by George (1906). A. georgei resembles also Ameroseius elegantissimus by Ishikawa (1984) from Japan. These two species can be easily distinguished by the number of elongate setae (11 pairs in A. georgei, only eight pairs in A. elegantissimus).
I examined three slides (BMNH.E.010147154-56) of Michael Collection deposited at the Natural History Museum, London, each bearing a specimen originally labelled Epicrius and additionally specified as Cornubia georgii (sic). The specimen from Port Garrah (on the slide BMNH.E.010147154) labelled "nymph" is un suitable for adequate study, and definitely not conspecific with those on the remaining two slides. Other two female specimens from Ranmore are identical to one another, in very good condition to be easily recognised as Ameroseius imparsetosus. Unfortunately, none of available slides bears a type designation and more specific collection data, but I believe that one female belongs to the authentic specimen of George (1906) because the original illustration of the dorsum with legs given by George (1906), for his Epicrius canistrinii (sic), match perfectly one of the two available mounted specimens (on slide E010147156). Based on the above mentioned circumstances, a new synonymy is established between C. georgei and A. imparsetosus in present study.

Ameroseius haplocosmus Elsen, 1973
Ameroseius haplocosmus Elsen, 1973 Remarks. The dorsal chaetotaxy shown in the original description is partly confused by depicting a non existing setae next to vertex (see depicted s1), and by omitting two pairs of setae on mediolateral and marginal surface (z5, r4). What was shown as s1 is apparently part of the palps, strongly pressed under the vertex. Thus, I could detect the normal set of 29 pairs of the dorsal shield setae for this species.

Ameroseius longitarsis Elsen, 1973
Ameroseius longitarsis Elsen, 1973 Moser and Roton (1971), Furniss et al. (1972) and Moser (1975), has revealed that these specimens represents at least two species different from A. longitrichus, including Ameroseius peniophorae De Leon, 1964, previously described from Tennessee. These two species seem to be very similar to A. longitrichus, but can be distinguished by some characters of dorsal and leg chaetotaxy (in A. peniophorae, all legs are notably thickened and shortend, and femur I and genu I bear a robust and smooth seta on medial ventral surface).

Ameroseius macropilis De Leon, 1964
Ameroseius macropilis De Leon, 1964: 215. Remarks. Ameroseius macropilis is very similar to Ameroseius corbiculus (Sowerby, 1806), but they can be easily distinguished by the number of opisthogastric setae expressed on soft integument (six pairs in A. macropilis; five pairs in A. corbiculus, having its JV4 absent). The reports of A. corbiculus given by Oatman (1971), Freitag and Ryder (1973), and Goh and Lange (1989) seem to be a misidentification of A. macropilis, and they should be revised.
Etymology. The new species is named in honour of my dear older sister.
Remarks. There are two slides of this species in the Berlese Collection in Florence, only one of them (169/3) is labelled "tipico". Unfortunately, the holotype specimen is wholly unsuitable for study, and practically the same can be stated for the female mounted onto the second slide (190/19), reported from Florence. Bregetova (1977) considered Ameroseius sculptilis to be identical to Ameroseius pulcher Westerboer, 1963, and her interpretation is accepted in this paper. I examined one quite well preserved female on a slide in the Hirschmann/Willmann Collection in Munich labelled "Ameroseius pulcher n. sp.". Unfortunately, this available specimen (slide) bears no type designation and specific collection data, but without doubt it belongs to the original series of Westerboer (see remarks under Ameroseiella macrochelae).
Type locality and habitat. India, West Bengal, Sitala, nest of lesser banded hornet, Vespa affinis (Hymenoptera). Karg, 1996 Ameroseius stultus Karg, 1996 Remarks. Karg (1996) illustrated this species as having only 28 pairs of dorsal setae on the dorsal shield. I could detect 29 setal pairs (including r4, missing in the illustration of the original description), leg chaetotaxy typical of the genus, a prominent poroid structure on each peritrematal shield, and slightly smaller sternal shield than depicted in the original description.
I examined the holotype female of this species and could detect the normal complement of 29 pairs of setae on dorsal shield (not stated in original description), including j1 which are missing in the holotype (together with a small part of vertex). In Ameroseius womersleyi, st3 are on soft integument, opisthogastric soft integument presumably has six pairs of setae (JV4 present), and fixed digit of chelicera possesses three well observable teeth on proximal masticatory area of which two proximal are slightly smaller and somewhat separate from the most distal tooth. The recent diagnosis given by Halliday (1997) does not require additional amendment.

Diagnosis (adults).
Soft striate integument delicately incrusted with sclerotic tubercles. Dorsal shield widely oval, not completely covering dorsal surface, reticulate or striate, with 6-7 pairs of distinct protuberances (each bearing a seta), and 29 pairs of setae differing in length and form; setae mounted on protuberances (z4, z5, s5, Z1, Z3, S5, and sometimes z2) markedly longer, thicker, and more heavily pilose than the other setae on the shield. Dorsal shield setae needle-like to lanceolate, smooth, pilose or serrate, the thicker setae densely plumose; sexual dimorphism of dorsal chaetotaxy not developed. In female, sternal setae on sternal shield (st1, st2), soft integument (st3, st4) and epigynal shield (st5). Female with anal shield bearing only three circum-anal setae, male with expanded ventrianal shield capturing some opisthogastric setae. Peritrematal shields and peritremes well developed: anterior end of peritremes reaching or overlapping bases of j1; in male, peritrematal shields well expanded beyond coxae IV and abutting the ventrianal shield. Opisthogastric soft integument with six pairs of setae in female. Corniculi stout, weakly sclerotised, well separate and parallel, with undivided apex having one or two denticles. In female, cheliceral shafts relatively elongate, slender; fixed digit with three prominent sharp teeth (two proximal teeth slightly separated from the other, medial tooth), and a special bilobed tooth close to terminal hook (Plate 76F); movable digit only with tiny subapical denticle; cheliceral digits almost straight, not conspicuously hooked distally, without hyaline petal-like appendages. Male spermatodactyl relatively short, directed forward. Palptarsal apotele three-tined. Tibia IV with two posterolateral setae. Tarsi I-IV each with well developed empodium and claws. Insemination apparatus with spermathecal ducts separated.

Remarks.
The genus Asperolaelaps was proposed by Womersley (1956a), with Asperolaelaps rotundus as its type species, but it has been considered a synonym of Neocypholaelaps Vitzthum, 1942, for example by Domrow (1979), Halliday (1997), and Moraes and Narita (2010). Womersley (1956) originally placed it in the family Neoparasitidae but Domrow (1979) re-examined the type species observing the normal complement of 29 pairs of dorsal shield setae instead of the reduced number illustrated by Domrow, and some further features typical of ameroseiid species.
In this paper, the genus is removed from synonymy with Neocypholaelaps, re-diagnosed, and ressurected to accommodate one further species from Australasian Region, namely Asperolaelaps sextuberculi (Karg, 1996). None of these species have been found in association with flowers and their pollinators, as opposed to species of Neocypholaelaps, the closest genus. Separate systematic position of Asperolaelaps species can better clarify the concept of Neocypholaelaps. Asperolaelaps rotundus and A. sextuberculi are considered not to belong to Neocypholaelaps because their diagnostic character states are based on the features inconsistent with Neocypholaelaps: (1) fixed digit of chelicera with three sharp teeth in proximal-medial part and a wide bilobed subapical tooth (edentate in Neocypholaelaps); (2) chelicera without hyaline appendages (appendages developed in Neocypholaelaps); (3) corniculi well separate and parallel to each other (adjacent and convergent in Neocypholaelaps); (4) palptarsal claw three-tined (two-tined in Neocypholaelaps); (5) some dorsal setae mounted on protuberances, and some with unusual position on the dorsal shield: e.g., Z-series setae with insertions well separate from those in central J-rows and more adjacent to those in marginal S-rows (dorsal shield smooth, with dorsal setae otherwise situated in Neocypholaelaps); (6) sexual dimorphism of dorsal setation absent (present in Neocypholaelaps); (7) tibia IV with two posterolateral setae (only one posterolateral seta in Neocypholaelaps). The dentation of cheliceral digits (together with other characters of gnathosoma) shows important generic difference between Asperolaelaps and Neocypholaelaps, probably due to their specific mode of life and different feeding habits. On the contrary, except for items 1, 5 and 7, the above mentioned character states are well consistent with the genus Ameroseius. Asperolaelaps represents a group of species with Neocypholaelaps-like venter, Ameroseius-like gnathosoma, and specific dorsal chaetotaxy, cheliceral dentation and tibial setation of legs IV.
Examination of the type specimens of Asperolaelaps rotundus and Asperolaelaps sextuberculi confirmed their validity and specific features. They are quite similar species originally reported from the same biogeographic realm, namely from Australia and New Caledonia, respectively. Female of A. rotundus differs from A. sextuberculi by having three setae on femur of leg I conspicuously thickened and shortened, conical (pl2, pv1 and pv2; pl2 unusually displaced close to pv2), and by the peritremes with anterior tips touching each other. In A. sextuberculi, femur I is relatively narrower and shorter than in previous species, having only normally formed setae, and anterior ends of peritremes separated by insertions of j1.
Asperolaelaps sextuberculi (Karg, 1996), comb. n. Karg, 1996 Remarks. In the original description, Karg (1996) did not comment the number of the dorsal shield setae. There are 29 pairs of the dorsal shield setae on his quite detailed illustration of the dorsum. Examination of the type specimens showed that the dorsal chaetotaxy is partly confused by depicting a non existing pair between r3 and r4, and by omitting one setal pair on medial surface (z6). In addition, I could find six pairs of opisthogastric setae (only five pairs shown in the original illustration). Genu III with one posterolateral seta (as in Ameroseius), but tibiae III and IV with two posterolateral setae (as in Kleemannia). Diagnosis (adults). Dorsal shield well sclerotised and ornamented, with 27-29 pairs of setae. Dorsal setae not conspicuously thickened or otherwise modified. In female, sternal setae on sternal shield (st1, st2, st3), complex metasternal-endopodal platelet (st4), and epigynal shield (st5). Epigynal shield relatively large, well oblong, and with posterior margin widely abutting ventrianal shield. In both sexes, posteroventral region with reduced number of four pairs of opisthogastric setae on ventrianal shield (JV2, JV3, ZV2) or on soft integument (JV5); setae JV1 and JV4 absent. Ventrally inserted setae including JV5 short, smooth and needle-like. Corniculi apically bifid. Fixed digit of chelicera with eight subequal denticles on masticatory area; spermatodactyl relatively long, apparently longer than movable digit. Epistome subtriangular, with one or two sharply pointed cusps. Palptarsal apotele three-tined. Genua II-III without ventral setae, genu III with two anterolateral and one posterolateral setae, tibia IV with two anterolateral and two posterolateral setae. Tarsi I-IV each with normal empodium and claws.

Genus
Remarks. The genus comprises only two described species. They are reported from the Solomon Islands, Papua New Guinea, Australia (Queensland) and Thailand (Womersley 1956a, Halliday 1997, Silva et al. 2014, as phoretic associates of phytophagous insects (see below).
Remarks. Some important amendments to the original description of Brontispalaelaps leveri were carried out by Halliday (1997) and Silva et al. (2014). This species was described from specimens associated with the chrysomelid beetle Brontispa froggatti Sharp, 1904[= Brontispa longissima (Gestro, 1885] from the Solomon Islands and Papua New Guinea. It is remarkable that the type specimens of Ameroseius crassipes were collected from the same host beetle in Australia. For that reason there may be some speculation about conspecificity of A. crassipes and B. leveri, the type species of the genus Brontispalaelaps. The conspecificity of both species should be carefully checked by further study once more, despite of the fact that Lindquist's examination of the type specimens has shown that A. crassipes belongs to an unspecified genus of Ologamasidae (Halliday 1997).
In the checklist below, ten species of this genus are recognised as valid. Among them, now I recognise five species occurring in Europe (horridus = berlesei, hungaricus, mirabilis = rivus syn. n., palustris = baloghi syn. n. and langei syn. n., and suedus), with the new synonymies proposed after my examination of the type specimens and speci-mens collected in Slovakia. For specific remarks to the individual Epicriopsis species and their synonyms see the checklist below.
The dorsal chaetotaxy of individual species occurring in Europe is not adequately described up to now. The two species (rivus and suedus) described by Karg (1971a) are known only on the base of illustrations and very short description as a part of an identification key. The dorsal shield bears an uncomplete set of setae in the illustrations of Epicriopsis horridus, Epicriopsis rivus and Epicriopsis suedus by Karg (1971a). Therefore, I carefully checked the number of dorsal shield setae in available Epicriopsis species. Epicriopsis hungaricus, Epicriopsis palustris and Epicriopsis mirabilis (= E. rivus) possess 24 pairs of setae (j1-j6, J2, J4, z2, z4, z5, Z1, Z3, Z5, s1, s2, s4-s6, S3-S5), E. horridus 23 pairs of setae (j6 absent in comparison with previous three species), and E. suedus 28 pairs of setae (s6 absent in comparison with normal complement of dorsal setae found in most of Ameroseiidae).
Dubious published material from Slovakia. Malé Karpaty Mts.: Bratislava Capital, Devín Settlement, Devínska Lesostep Forest (Kalúz 2005). [Notes: Epicriopsis horridus belongs to the common species in Slovakia but the published specimens from Malé Karpaty Mountains are excluded from the list of valid findings of Epicrius horridus because accompanying "original" figure available in Kalúz (2005) is obviously not belonging to this species, and it is only a true copy of illustration of Epicriopsis langei given to the original description of the new species from Central Asia by Livshitz and Mitrofanov (1975). Moreover, in a previous paper on mites collected in the Abrod Meadows by Kalúz (2003), the same illustration was used although no record of any Epicriopsis species from that locality is reported by the author.
New  (4061-4063), all with no type designation. The revised specimens belong at least to the two different species: (1) a deutonymph from San Remo, labelled 4061, is weakly sclerotised and unidentifiable, (2) a female from Sucy-en-Brie, labelled 4062, is hardly observable and very similar if not identical with Epicriopsis palustris, (3) a "deutonymph" from Borkum Island, labelled 4063, is a female of E. horridus. Karg (1971aKarg ( , 1971b) considered E. berlesei to be a junior synonym of E. horridus giving no supporting information. Here I have adopted his synonymy although I have not examined the Italian specimens of E. horridus deposited in Berlese Collection in Firenze.

Epicriopsis hungaricus Kandil, 1978 Plate 30
Epicriopsis hungarica Kandil, 1978 Remarks. There are seven slides labelled Epicriopsis hungarica in the Budapest Museum Collection, none with type designation. Except for one female from Dobogókő (see comparative material above), all of these specimens belong to the original type series. The same is true for the five available slides labelled Epicriopsis baloghi Kandil, 1978. One of the slides, labelled Meso-1350, bears a female of E. hungaricus, misidentified as E. baloghi.

Epicriopsis jilinensis Ma, 2002
Epicriopsis jilinensis Ma, 2002 Remarks. There are two apparently original slides of Epicriopsis mirabilis with a dissected male in comparatively good condition in the Hirschmann/Willmann Collection in München (gnathosoma, chelicera and leg I are separately mounted on a slide), labelled No. 42/8 and 43/8. The male may be considered as holotype by monotypy. My examination of this Willmann's specimen confirmed the conspecificity of E. mirabilis with the species Epicriopsis rivus Karg 1971. According to Karg (1971a) and the followers, the only reliable diagnostic feature for distinguishing E. mirabilis from E. rivus seemed to be the form of the epistome. In E. mirabilis, the epistome was misinterpretated by Willmann (1956) and illustrated as a trilobate structure while the epistome of E. rivus was correctly understood to be a sharply pointed structure. There is a relatively clearly visible epistome in dissected gnathosoma on slide No. 43/8, with the anterior margin produced into a medial acute process, as in E. rivus. The type specimens of both species are also identical in relation to all other morphological features, including the peculiar sculpture composed of polygonal instead of starshaped tubercles. Remarks. Despite of the fact that Epicriopsis palustris was described in 1971 by Karg, it was not included in Karg's widely used monograph focused on identification of the European mesostigmatans, published in the same year (Karg 1971a). The original description of this species contains an incorrect statement about the setation of ventrianal shield in female. I have carefully checked the ventrianal shield in the paratype female, and I could detect only three circum-anal setae standard for the genus, but not the two additional opisthogastric setae inserted on the lateral parts of the shield, as illustrated in Fig. 7a (page 6) of Karg (1971b). Karg's confusion is still respected by some authors, and it can be found in the most recent studies on the genus Epicriopsis up to now (Narita and Moraes 2016), although E. palustris was adequately redescribed by Călugăr (2008), based on females and males from Romania. Epicriopsis palustris was originally described based on the type specimens from Germany; subsequent findings of this species are reported from various countries in central and northern Europe (Hungary, Latvia, Poland, Romania, Slovakia). I have collected this species in the Mediterranean area of Greece (Sarti Village, Chalkidiki Peninsula), and checked relatively rich material from various parts of Iran. Under the name horridus, the species is reported from Israel by Iavorschi (1995). Therefore, the species has clearly more expanded distribution area than previously expected. The specimens of E. palustris from Ukraine and Hungary were treated as a new species by Livshits and Mitrofanov (1975) and Kandil (1978), namely Epicriopsis langei and Epicriopsis baloghi. These authors apparently neglected the existence of E. palustris; only Epicriopsis horridus and Epicriopsis rivus were compared with their newly described species in the differential diagnose, and no reference to E. palustris can be found there.
Tracking of the dorsal setae in European species of Epicriopsis, especially the shortest ones, may be quite tricky task due to heavy sclerotization and coarse sculpture of the dorsal shield. In published papers, the full complement of the dorsal shield setae has been correctly depicted only exceptionally and in limited number of illustrations. For instance, also in Epicriopsis palustris, we can see incomplete numbers of setae on the dorsal shield, given by various authors, namely 18 pairs by Karg (1971b) and Livshits and Mitrofanov (1975), 17 pairs by Kandil (1978), 16 pairs by Iavorschi (1995), and 21 pairs by Călugăr (2008). I have found 24 pairs of the dorsal shield setae, and the same number is depicted on a figure of misidentified female published by Evans and Till (1979). Livshits and Mitrofanov (1975) described Epicriopsis langei on the base of a single female reportedly deposited at the Nikita Botanical Gardens, National Scientific Center, Yalta, Crimea. The species seems to be apparently conspecific with Epicriopsis palustris, when compared with the types from Germany and my own specimens from Slovakia, except the form of epistome (illustrated as pointed in E. langei, rounded in E. palustris). I dare to say the epistome of E. langei is not correctly illustrated in the original illustrations, as in Epicriopsis mirabilis. I tried to obtain the type specimen of E. langei for examination, asking Alex Khaustov (at that time an acarologist of the National Scientific Center in Yalta) for help. Despite his great effort, the types were not found and are presumably lost.

Epicriopsis stellata Ishikawa, 1972
Epicriopsis stellata Ishikawa, 1972 Remarks. I examined a paratype female of this species. I could detect 24 pairs of dorsal shield setae instead of 23 pairs as stated by Ishikawa (1972). Epicriopsis stellata is closely related to Epicriopsis hungaricus and Epicriopsis palustris, and it can be distinguished by the number of opisthogastric setae inserted on soft integument (six pairs in E. stellata, five pairs in E. palustris and E. hungaricus), and the relative length of S4 and S5 (in E. stellata, both similar; in E. palustris and E. hungaricus, S5 thicker, delicately pilose and at least twice longer than S4). Karg, 1971 Plates 34, 76C

Epicriopsis suedus
Epicriopsis suedus Karg, 1971a Remarks. This species bears almost full complement of dorsal shield setae. I found 28 pairs in the type specimens from Sweden (s6 presumably absent). Karg (1971a) depicted only 22 pairs in his original illustration. I was unable to check the number of opisthogastric setae due to the position of the specimens on the slide.

Diagnosis (adults).
In female, dorsal shield lightly sclerotised, weakly reticulated or unornamented, often irregular in outlines (with 1-3 pairs of lateral incisions), narrowed and not covering whole dorsal surface; dorsal setae short, needle-shaped, similar in length, inserted on dorsal shield (18-25 pairs) and soft integument beside the dorsal shield (3-20 pairs); sometimes dorsolateral and ventrolateral soft integument hypertrichous, with up to 35 setal pairs (non-hypertrichous forms usually with standard number of 28-29 pairs of dorsal setae on dorsal shield and soft integument). Male often with some dorsal shield setae (8-10 pairs) conspicuous, thickened, extremely long and much longer than some of the shortest ones; with slightly expanded dorsal shield capturing some additional pairs of setae inserted on soft integument in female. Female with st1 on weakly sclerotised and inconspicuous sternal shield; setae st2 on sternal shield, small sternal shield fragments or soft integument; setae st3 associated with small endopodal platelets II-III or on soft integument; and st4 on soft integument; sternal shield usually markedly reduced; epigynal shield widely separated from anal shield, with parallel lateral margins, and usually with genital poroids on its surface. Female with anal shield, male with anal shield or ventrianal shield bearing 1-3 pairs of opisthogastric setae. Peritrematal shields or peritremes with anterior ends free and not connected to dorsal shield. Opisthogastric soft integument with five to six pairs of setae (JV3 usually absent in male). Corniculi slender and pointed, with undivided apex. Fixed digit of female chelicera usually edentate, with bifid terminal hook, sometimes with one or two medial denticles; movable digit edentate, with strongly developed terminal hook; pilus dentilis usually not modified into a membranous structure; male with well elongated spermatodactyl directed downward. Palptarsal apotele two-tined. Genu III and tibia III with two anterolateral and one posterolateral setae; tibia IV with two anterolateral and two posterolateral setae. Tarsi I-IV each with empodium and slightly to fully reduced claws. Insemination apparatus with spermathecal ducts unfused.

Remarks.
Generally, the genus Hattena as a whole shows relatively high diversity in external morphology, and it represents a conglomerate of morphologically heterogeneous species. The following character states were used to separate the Hattena species into five species groups (see the identification key below): (1) expression of z6 (absent in cometis group); (2) expression of J3 (present in panopla group); (3) position of Z5 (on soft integument in panopla group); (4) setation of soft integument (hypertrichy in cometis and erosa groups); (5) form of posterior part of dorsal shield (incised in dalyi and panopla groups); (6) sexual dimorphism of dorsal shield setae (markedly developed in cometis group); (7) position of genital poroids (outside the epigynal shield in erosa and senaria groups); (8) sexual dimorphism of anal shield (absent in dalyi group); (9) character of tritosternal laciniae (fused for most of their length in panopla group); (10) setation of ventrianal shield in male (with a pair of opisthogastric setae in panopla group and three pairs in cometis group); (11) setation of palptrochanter (with only one ventral seta in cometis and erosa groups); (12) legs IV in male (spurred in cometis group).
In this genus, the dorsal shield of males is more expanded than in females, thus bearing up to ten pairs of the setae that in females are inserted on the soft integument. Hattena cometis, Hattena erosa and Hattena floricola have polytrichous soft integument on dorso-and ventrolateral idiosomal surface. Polytrichy has been mentioned in the literature only for females; it was not reported in the descriptions of males of H. cometis and H. floricola. Therefore, the polytrichy of male soft integument may be neglected by Domrow (1979) and Halliday (1997), if not considerably reduced or absent.
It is remarkable that there is an additional setal pair of the J-series (J3) expressed on the dorsal shield in Hattena panopla and Hattena tongana. This seta is also present in Neocypholaelaps novaehollandiae.
Hattena clemmys was not included in the species group classification scheme introduced in this paper because there are several evidences that it is incorrectly placed in this genus: (1) dorsal shield divided into six fragments; (2) sternal shield not strongly reduced but without sternal setae and poroids; (3) setae st5 on soft integument, off the epigynal shield; (4) peritremes strongly abbreviated; (5) movable digit of chelicera with elongate and fringed process; (6) palptarsal apotele three-tined; (7) tarsus of legs I-IV with normally developed claws on empodium. It was placed in Hattena because of the extent of the dorsal shield and the number of dorsal shield setae, but it probably represents a new genus. Therefore the above mentioned specific characters of H. clemmys are not included in the diagnosis of Hattena.
The genus Hattena now comprises ten species distributed in the tropical areas of the Old and New Worlds (Australia, Ecuador, Kenya, Malaysia, Pacific Ocean Region, Papua New Guinea, Taiwan and Vietnam), in association with flowers and flower-visiting animals (birds, bees and bats). Hattena dalyi and Hattena tonganus are known to be phoretically associated with the Euroepean honey bee, Apis mellifera (from tropics).
Partial keys to the known species of Hattena may be found only in Halliday (1997), for four species reported from Australia (cometis, floricola, incisa and panopla), and in Faraji and Cornejo (2006), based on six species known up to that time (cometis, erosa, floricola, incisa, panopla and rhizophorae). A new and more complete key to the genus is needed to include other species, namely Hattena dalyi (transferred from Afrocypholaelaps by Klimov et al. in 2016) and Hattena senaria comb. n.
Type locality and habitat. Australia, Northern Territory, Elizabeth River near Palmerston, on flowers of spotted mangrove, Rhizophora stylosa (Rhizophoraceae).
Remarks. Hattena dalyi (Elsen, 1974) is exceedingly similar to Hattena incisa. The position of st2 appears to be the only important difference between these species; these are on the sternal shield margin in H. dalyi and on separate and small shield fragments close to lateral margins of the sternal shield in H. incisa. Halliday (1997) may have overlooked the existence of H. dalyi because it was described as a species of Afrocypholaelaps, until Klimov et al. (2016) transferred it to Hattena. In addition, Halliday (1997) did not cite Elsen (1974). Thus, a thorough morphological comparison between these species is necessary. Domrow, 1966 Hattena panopla Domrow, 1966: 208. Hattena panopla. -Fain andLukoschus 1979: 20;Domrow 1979: 105;Seeman 1996: 193;Halliday 1997: 195 Remarks. Faraji and Cornejo (2006) reported this species as having only four instead of six pairs of setae on opisthogastric soft integument, taking no account of the relative positions of all these setae on an expanded striate integument and their dimorphic expression common in Ameroseiidae (a pair of opisthogastric setae usually absent in male). For that reason, they interpreted JV4 and JV5 (designated in their illustration as R3 and R4) to belong to a complement of posterior marginal dorsal setae. Later, in their description of male and deutonymph, these setae are not referred to. Their JV5 in female and deutonymph are interpreted here as JV3, and they seem to be absent in male. I suspect JV4 to be omitted in the male illustrations of Faraji and Cornejo (2006), and JV5 not to be homologous in adults of both sexes. (Allred, 1970), comb. n. Allred, 1970: 101. Hattena tongana (Manson, 1974) Edbarellus tonganus Manson, 1974: 116. Hattena tongana. -Halliday 1997: 188. Hattena tonganus. -Klimov et al. 2016. Type locality and habitat. Pacific Ocean Region, Kingdom of Tonga, Tongatapu Island, in a hive of European honey bee, Apis mellifera (Hymenoptera).

Ameroseius senarius
Remarks. I suspect this species to be a junior synonym of Hattena panopla. Halliday (1997) reported many extreme similarities between Hattena tongana and H. panopla, but he did not consider them to be conspecific. Differences in shape of sternal shields described by Halliday (1997) could be questionable due to the weak sclerotization of the sternal shield and its margins, characterstic of Hattena. However, I adopted in this paper the interpretation of Halliday (1997), considering them distinct species.

Diagnosis (adults).
Dorsal shield variously sclerotised and ornamented, normally with 28-29 pairs of setae (z6 present or absent). Dorsal shield setae similar in adults of both sexes, well developed and relatively long, thickened, tubiform or flattened, mostly densely pilose or serrate. In female, st1 and st2 on sternal shield, st3 on small suboval or subcircular pseudo-metasternal platelets or sometimes on soft integument (in Kleemannia mirabilis sp. n. on sternal shield), and st4 on soft integument; endopodal platelets II-III present, subtriangular, and relatively small; anterior margin of epigynal shield convex, genital poroids on soft integument. Female with five or six pairs of opisthogastric setae (JV1-JV3, JV5, ZV2; JV4 sometimes absent), two pairs of which (JV2, JV3) on ventrianal shield (in Kleemannia insignis, JV2 occasionally on soft integument but contiguous to anterior margin of ventrianal shield), other on soft integument; male with five pairs of opisthogastric setae (JV4 always absent), 3-5 pairs of which on expanded ventrianal shield (JV1 and/or JV5 sometimes on soft integument). In both sexes, JV5 usually similar to those on dorsal shield, other ventrally inserted setae simple, smooth and needle-like. Peritrematal shields anteriorly connected to dorsal shield, and often with enlarged cavity-like poroid structure at level of coxae III. Metapodal platelets well developed, elongate. Corniculi well sclerotised and spaced, usually with bifid or trifid apex; setae h1 thickened, straight or slightly and regularly curved, progressively taperred and pointed. Fixed digit of chelicera with four subequal teeth on proximal masticatory area; male spermatodactyl relatively small, usually as long as movable cheliceral digit (if not shorter). Epistome with anterior margin produced into narrow and pointed central projection or, sometimes, short obtuse cusp; lateral margins mostly smooth. Palptarsal apotele usually two-tined. Coxae I-IV with delicate sculptural ornamentation, striate and reticulate. Genu III and tibiae III-IV with two anterolateral and two posterolateral setae. Tarsi I-IV each with empodium and claws; tarsi I usually well pigmented, brown in colour. Insemination apparatus with papilla-like sperm induction pores (solenostomes) associated with inner margin of coxae III. Remarks. Oudemans (1930) proposed the new genus Kleemannia with Zercon pavidus C. L. Koch, 1839 as type species, and added three further species in the genus (plumosa, plumea and plumigera). Since then, this taxon has been variously treated as a genus (Vitzthum 1942;Hughes , 1976Evans 1963a;Ishikawa 1972;Domrow 1974Domrow , 1979Evans and Till 1979), a subgenus of Ameroseius (Athias-Henriot 1959, or a junior synonym of Ameroseius (Westerboer and Bernhard 1963;Bregetova 1977;Karg 1993Karg , 2005Halliday 1997;Karg and Schorlemmer 2009;Narita et al. 2013b), although it may be easily and reliably separated from Ameroseius by the presence of two posterolateral setae on genu III and tiabiae III-IV (one in Ameroseius), four proximal denticles on fixed digit of chelicerae (at most three denticles in Ameroseius), two pairs of opisthogastric setae on ventrianal shield (at most one pair in Ameroseius, but never JV2), two-tined claw on palptarsus (three-tined in Ameroseius), thickened h1 (similar if compared to other hypostomal setae in Ameroseius), epistome usually with sharply or obtusely pointed apex (never produced into a single pointed process in Ameroseius). Therefore, the genus Kleemannia is considered to be a valid genus, and is removed from synonymy with Ameroseius.
The related genus Primoseius was proposed and very briefly characterised by Womersley (1956b). He included two species in the genus, the type species, mentioned as Lasioseius (Zercoseius) macauleyi Hughes, 1948 (sic, should be Zercoseius macauleyi), and Lasioseius (Zercoseius) gracei Hughes, 1948 (sic, should be Zercoseius gracei), which were subsequently synonymised with Kleemannia plumosa and Kleemannia plumigera, respectively (Evans 1954). Later, the separate taxonomic status of this genus was rejected by , Evans and Till (1979), Karg (1971aKarg ( , 1993Karg ( , 2005, Bregetova (1977), and many others, because the original concept of Primoseius was defined with the help of the same characters as they can be found in the most relative genus Ameroseius, or Kleemannia s. l. Womersley (1956b) based the diagnosis of his new genus especially on the possession of two pairs of setae (st1, st2) on sternal shield, a pair of pseudo-metasternal platelets each bearing a seta (st3), and metasternal setae (st4) on soft integument. Despite these facts, Primoseius is included as a subgenus of Kleemannia in the list of valid taxa of Ameroseiidae in this paper. It may be reliably distinguished from Kleemannia s. str. mainly by the absence of z6, and presence of specifically modified dorsal setae (conspicuously flattened, leaf-to feather-shaped). Primoseius currently comprises 12 described species (bisetae, dipankari, dubitatus, eumorphus, macauleyi, mineiro, parplumosa, plumosoides, plumosa, potchefstroomensis, pseudoplumosa and wahabi), of which only eight species are considered here to be valid (bisetae, dipankari, mineiro, parplumosa, plumosoides, plumosa, pseudoplumosa and wahabi). Narita et al. (2013b) classified eight species of "Ameroseius" into a species group named plumosus-group (dipankari, eumorphus, mineiro, parplumosa, plumosa, potchefstroomensis, reticulatus and wahabi). They defined the group by the following character states, of which some are vague and applicable also for other groups inside the genus: corniculus bifid or trifid; dorsal shield reticulate and without pit-like depressions; 26 or 28 pairs of dorsal shield setae, most of which are lanceolate to leaf-shaped; and five or six pairs of opisthogastric setae, two pairs of which are on ventrianal shield. The plumosus-group of Narita et al. (2013b) could be considered to be identical with Primoseius, and it omits three species described before 2013, namely Kleemannia bisetae, Kleemannia plumosoides and Kleemannia pseudoplumosa, and inexplicably includes Kleemannia reticulata, which has z6 clearly developed and present on the dorsal shield.
Kleemannia currently comprises 28 valid species described mostly from the Palaearctic, and only rarely from Neotropics (four species), Nearctic (one species), Afrotropics (one species) and Oriental Region (two species). Specimens reported as plumosa and plumigera have been reported from many countries around the world, from various natural and synanthropic habitats (stored grain and food, litter in sheds and stalls, mould in buildings). In Slovakia, this genus is represented by eight recorded species.

Key to species of Kleemannia occurring in Europe (females)
Partial keys to species of Kleemannia (classified in Ameroseius) from Europe and former Soviet Union may be found in Karg (1971aKarg ( , 1993 and Bregetova (1977), and they include eight and 13 species, respectively. The world species can be identified using keys from Narita et al. (2013bNarita et al. ( , 2015. A new key to the genus is needed to include a species here firstly reported from Europe (Kleemannia kosi and Kleemannia parplumosa), the new species described in this study (Kleemannia dolichochaeta sp. n.), and a revised Berlese species previously considered to be dubious (Kleemannia tenella). The identification key for 12 species presented below is based on direct examination of type specimens or specimens considered to be conspecific with the type material, except for Kleemannia elegans.
Several diagnostic features can be useful in recognising of the Kleemannia species occurring in Europe, namely (1) presence/absence of z6 (absent in dolichochaeta, parlplumosa, plumosa and pseudoplumosa); (2) placement of st3 (on soft integument in plumea and tenella); (3) number of opisthogastric setae (with reduced number of five pairs in insignis, plumea and tenella); (4) presence/absence of cavity-like poroid structure on peritrematal shield (absent in plumea, plumigera and tenella); (5) placement of JV2 and JV3 on anterior portion of ventrianal shield (both setae on medial surface in elegans, kosi, pavida and plumigera; JV2 on anterior margin or surface in dolichochaeta, insignis, parplumosa, plumosa and pseudoplumosa; both setae on anterior margin or surface in delicata and tenella; and (6) relative length of Z-and S-setae (Z-setae apparently shorter than S-setae in tenella).

1
Dorsal shield with 28 pairs of setae (z6 absent); dorsal setae flattened, feather-to leaf-shaped (subgenus Primoseius Womersley, 1956  Remarks. There are six slides of this species in the Berlese 194/18,. These slides contain at least 12 specimens, including those of Ameroseius molliculus (nomen nudum) and Ameroseius dubitatus, all collected in Italy (Firenze and Padova), and not labelled as "tipico". Without doubt, those from Firenze belong to the original type specimens of Berlese. They are mostly in bad condition to be examined. However, an examination of the specimens showed specific features also detected in well preserved specimens available for study from Slovakia and other regions of Europe.
Based on an examination of the type specimens, Ameroseius stramenis Karg, 1976 andLasioseius gracilis Halbert, 1923 are considered new junior synonyms of Kleemannia delicata (Berlese, 1918). The two former species were briefly described in their original descriptions. Halbert (1923) did not inform the number of setae on the dorsum, but his illustration shows 26-27 setae, lacking z6. Karg (1971aKarg ( , 1993 slightly modified Halbert's illustration to show 27 setal pairs. I could detect the full complement of 29 pairs of setae in type series of both, delicatus and gracilis. The re-description and illustrations given by Westerboer and Bernhard (1963) for delicatus do not require serious amendment, except for the shape of ventrianal shield. This shield is usually not so regularly subcircular as illustrated, but almost truncate and slightly concave anteriorly. In A. stramenis, Karg's (1976) interpretation of curved epistome seems to be apparently atypical for the genus, but I could not confirm it in the single available type specimen loaned from Berlin Collection (the epistome is obscure and not observable). In addition, the female of A. stramenis was illustrated by Karg as lacking metapodal platelets and one pair of opisthogastric setae (JV4), although both structures were reliably detected in an axamination of the holotype. Examination of numerous specimens from various mueum collections showed that Kleemannia delicata is fairly common in buildings in Europe and has often been misidentified, mainly as Kleemannia plumigera.

Ameroseius dipankari
Remarks. Based on the illustrations given by Nasr and Abou-Awad (1986) for Kleemannia parplumosa and Bhattacharyya (2004) for Kleemannia dipankari, both species are remarkably similar in having nearly identical inverted U-shaped sculptural structure on the anterior portion of the epigynal shield. Adult females of only a few Kleemannia species (namely dipankari, plumea and parplumosa) have been described as having this specific ornamentation structure. Other similarities include an elongate idiosoma; form and relative length of dorsal setae; relative size, shape and arrangement of ventral shields; and form of epistome, with lateral margins straight. Therefore, I suspect these two species to be conspecific. However, the illustration of Bhattacharyya (2004) shows the dorsal and ventral chaetotaxy to be quite dissimilar to those of K. parplumosa in lacking two pairs of dorsal shield setae (s1 or z2, and s4), and a pair of opisthogastric setae (JV4), usually placed on soft integument close to lateral margins of ventrianal shield. However, I believe the complement of dorsal and opisthogastric setae to be the same as in K. parplumosa, some setae being inadvertently omitted in its original description. Diagnosis. Dorsal shield with 28 pairs of setae (z6 absent); setae relatively narrow, sword-shaped. Setae j5 and j6 conspicuously lengthened, subequal to J2 and J4, and with tips reaching beyond bases of j6 and J2, respectively. Ventrianal shield with two pairs of opisthogastric setae (JV2, JV3); setae JV2 close to anterior margin of the shield. Setae JV4 present.
Remarks. Based on primary description of Bernhard (in Westerboer and Bernhard 1963), and re-description of Vasilyeva et al. (1976), females of Kleemannia insignis may be distinguished from those of all other members of the genus by the insertion of JV2 on soft integument; these setae are illustrated as being out but adjacent to the anterior edge of ventrianal shield. In the only Slovak specimen available for study, my examina-tion showed that there can be an alternative position of these setae because they are both clearly on the shield, adjacent to its anterior margin.
Based on the original illustrations of Fan and Li (1993), Ameroseius marginalis perfectly fits to Kleemannia insignis from Europe (dorsal and ventral chaetotaxy, specific sculptural granulation of dorsal shield, sculpture of metapodal platelets, shape and arrangement of ventral shields, etc.). Unfortunately, differential characters for separating A. marginalis (and also A. sichuanensis) from K. insignis given by Fan and Li (1993) are founded on variable (shape of pseudo-metasternal platelets, relative length of J2) or misinterpreted characters (in the original description of insignis, the number of teeth on fixed digit is stated to be five instead of four).
Remarks. El-Badry et al. (1979) and Zaher (1986) gave only a brief description of Kleemannia kosi. So, my identification of this species was initially based on insufficiently detailed illustrations available in their papers, later also on detailed photos of specimens stated to be kosi and originated from Egypt (kindly sent by R. Abo-Shnaf ).
Together with other ameroseiid species such as Kleemannia parplumosa and Sertitympanum nodosum, it can represent a group of stored product mites unintentionally introduced to Europe by various human activities (see comparative material above).

Kleemannia pinicola
New Remarks. The presence of a pair of pseudo-metasternal platelets anteriorly connected to the posterolateral margins of the sternal shield in female of Kleemannia plumigera distinguishes this species from females of all other species of Kleemannia and of other taxa of the Ameroseiidae. The female of K. plumigera was described by Oudemans (1930) and recently re-described by Narita et al. (2013b), as having pseudo-metasternal platelets separate and free from sternal shield, although this was not found by me in type female (numbered 4301) available in Oudemans Collection. Only Petrova (in Petrova and Koshchanova 1986) noted the partial fusion of pseudo-metasternal platelets with sternal shield in the specimens she described as Ameroseius gilarovi; other morphological characters mentioned in her description and detailed illustrations of A. gilarovi are in good agreement with those found in the type of K. plumigera (except for epistome, that seems to be misinterpreted). Therefore, a new synonymy is established between these two species in the present study. In one of the illustrations of specimens reported as Kleemannia delicata from Switzerland (here considered to be a misidentified K. plumigera), Schweizer (1961) only indistinctly indicated a connection between pseudo-metasternal platelets and sternal shield, but clearly and correctly depicted a pair of well sclerotised and prominent sperm induction pores associated with coxae III (another specific character of K. plumigera which is often overlooked in its desriptions). Oudemans (1930) described the dorsal chaetotaxy of Kleemannia plumigera as having 30 pairs of setae. However, as pointed out by Narita et al. (2013b), this is an error, as the dorsal shield actually has a normal number of 29 pairs of setae. The identity of K. plumigera sensu Rack (Rack 1963(Rack , 1964(Rack , 1971(Rack , 1972 cannot be established with any confidence because of the absence of some information about the ventral idiosoma in her re-description of this species (Rack 1963), and incorrect identity of most of the revised specimens available in Hamburg Collection, labelled K. plumigera and identified by this author. A re-examination of the specimens described by  as Zercoseius gracei, and later synonymised with K. plumigera (Evans 1954, would be necessary to confirm the conspecificity with K. plumigera or other related species, including Kleemannia delicata. Also  illustrated the dorsal chaetotaxy of putative K. plumigera incorrectly, as having 25 and 28 pairs of setae, respectively. Kleemannia maculeyi (sic). -Bhattacharyya 1972: 133. Kleemannia plumosa. -Ishikawa 1972Domrow 1979: 114. Ameroseius dubitatus. -Castagnoli andPegazzano 1985: 118. Ameroseius (Kleemannia) plumosus. -Hajizadeh et al. 2013a: 150. Type depository. Of Seiulus plumosus -National Museum of Natural History, Naturalis Biodiversity Center, Leiden, Netherlands; of Ameroseius dubitatus -Istituto Sperimentale per la Zoologia Agraria, Firenze, Italy; of Zercoseius macauleyi -not stated (holotype not designated).

Dubious published material from Slovakia. Ondavská vrchovina Highland:
Ruská Poruba Village (Mrciak 1963, cited as Ameroseius dubiatus). [Notes: The original description of A. dubitatus is based only on a brief enumeration of unclear or vague diagnostic characters. Therefore it was not included in the identification keys of Westerboer and Bernhard (1963), Karg (1971aKarg ( , 1993 or Bregetova (1977). According to Bregetova (1977), A. dubitatus seems to be a species very closely related with Ameroseius elegans Bernhard, 1963. Now we know that A. dubitatus is a synonym of Kleemannia plumosus. Slovak voucher material labelled Ameroseius dubiatus (sic) is not available for revision, so it is excluded from the list of valid published records.
New material from Slovakia. Malé Karpaty Mts.: 2 ♀♀ -9. 5. 2009, Bratislava Capital, zoological garden, dung-hill, manure, altitude 180 m, leg. P. Mašán. Remarks. When Oudemans (1902 described this species, as a member of Seiulus, he inadvertently included three different congeneric species to the type specimen series (pavida as a protonymph, plumosa as a deutonymph, and plumea as a female), but did not designate the holotype. Oudemans tried to correct his mistake in 1930, referring the name plumosus to a female specimens collected from Vespertilio dasycneme (Chiroptera), originally described as a deutonymph and illustrated in figures 18-20 (on plates 1 and 2) of Oudemans (1902). In the re-description provided in that paper, he was ambiguous in relation to the number of dorsal shield setae, stating 28 as well as 29 setal pairs (Oudemans 1930). The specimen shown in figure 19 of that publication apparently has z6, differing from the single type female of Kleemannia plumosa available in the Oudemans Collection in Leiden (numbered 4303); thus, the identity of that illustrated specimen is uncertain. Another slide in Oudemans Collection, numbered 4302, and labelled "Kleemannia plumosus, ♀ dors., Arnhem, 1919, Oudemans" bears only part of a sarcoptifom mite, with gnathosoma and the anteriormost part of idiosoma.
Kleemannia plumosa is easily recognisable by the specific inverted U-shaped sculptural structure on anterior surface of sternal shield having its anterior margin scalloped and heavily sclerotised, but it has been referred to a variety of names, including some synonyms and misidentifications. A new synonymy is here proposed for K. plumosa and Ameroseius dubitatus, based on direct examination of their type specimens. There is a relatively large series of slides in the Berlese Collection in Firenze, but none of them bears a type designation. Only specimens from Firenze may belong to the original type material of Berlese. Castagnoli and Pegazzano (1985) considered the female on slide No. 27/21 (from Castions di Strada, Udine Region) to be the holotype specimen but I could not validate their statement here.

Plates 57-60
Primoseius macauleyi. -Womersley 1956b: 116. Misidentification. Ameroseius pseudoplumosus Rack, 1972: 249. Ameroseius eumorphus Bregetova, 1977 Remarks. It appears likely that the names plumosa and macauleyi have been widely used for Kleemannia pseudoplumosa in the literature about mites associated with synanthropic habitats throughout the world. Based on my examination, the Australian misidentified specimens of Womersley (1956b), originally named macauleyi and classified in the newly erected genus Primoseius, are in excellent agreement with the type specimens of K. pseudoplumosa, collected in new buildings in Germany, and understood here to be conspecific with the specimens from Australia. In the illustration of the specimens identified as Primoseius macauleyi, Womersley (1956b) did not comment on the absence of the semi-circular scalloped sculptural pattern on the sternal shield (typical character of the species). Together with Halliday (1997) who revised the Australian fauna of Ameroseiidae, he did not consider this absence to indicate the existence of a separate species.
Further literature records that are consistent with the morphological features of the specimens studied by Womersley (1956b) and Rack (1972) include those from Israel (Costa 1966), Murmansk Region in Russia, Azerbaijan and Uzbekistan (Bregetova 1977), South Africa (Kruger and Loots 1980), Egypt (Zaher 1986), and Iran (Khalili-Moghadam and Saboori 2016). These records are supported by specimens examined here from Slovakia, Greece and Israel, and some photographs of the holotype of Ameroseius eumorphus (made by M. Stanyukovich, Russian Academy of Sciences, St. Petersburg, Russia, and obtained from A. Khalili-Moghadam, Iran). Accordingly, A. eumorphus Bregetova, 1977 andKleemannia potchefstroomensis Kruger &Loots, 1980 are placed in synonymy with Kleemannia pseudoplumosa (Rack 1972).
The description of Kleemannia pseudoplumosa by Rack (1972) was apparently overlooked by the followers. Bregetova (1977), Kruger and Loots (1980), Ibrahim and Abdel-Samed (1992) and Karg (1993) have not cited this species in their identification keys and/or differential diagnoses. The same can be stated for the descriptive paper of Narita et al. (2013b), on a new species Kleemannia mineiro from Brazil that is closely related with K. pseudoplumosa. In their identification key elaborated for a comparative purpose, Narita et al. (2013b) included mineiro close to eumorphus and potchefstroomensis, relying on the detailing the discrepancies in the original drawings. They stated that eumorphus differs from potchefstroomensis only in the sculpture of the sternal shield. This shield being unevenly ornamented, and with central portion smooth in eumorphus (in potchefstroomensis, whole surface with even but weak sculptural reticula). This difference is well within the range of variability found in European specimens of K. pseudoplumosa, and I regard it to be untenable. Moreover, there are some contradictory data on chaetotaxy of some leg segments as given by Kruger and Loots (1980) for potchefstroomensis, Narita et al. (2013b) for mineiro, and Khalili-Moghadam and Remarks. This species was only briefly and inadequately described by Berlese (1916a). There are seven slides of this species in the Berlese Acaroteca (21/8,168/50,, from moss, hay and litter from Italy (Firenze, Tiarno -Trentino, Palermo). Only one slide bearing a female from Firenze is labelled "tipico" (171/12). I have examined six females mounted onto five slides of the collection (slides 219/48-49 were not available for my study). Despite such a sufficient series of slides, only two of them contains at least partially examinable specimens in which some limited observation of features is possible. Some characters on the dorsum can be examined in the holotype specimen, and some ventral characters in a paratype specimen on slide 168/50. Other examined specimens are in poor condition and barely suitable for thorough observation. As a critical examination of the type specimens has revealed that Kleemannia tenella and Kleemannia fimetorum (Karg 1971) are identical, the latter must be placed in synonymy with the former. This comparison was extended to specimens of K. tenella collected in Slovakia and the agreement was found to be complete. In the original illustrations of K. fimetorum, Karg (1971a) asymmetrically depicted the setal complement of 29 pairs of setae on the dorsal shield (30 setae on left portion, and 27 setae on right portion), and mistakenly concluded that only three instead of four teeth were present on proximal masticatory area of the fixed cheliceral digit. On the basis of my examinations of description and illustrations, as the type material of Ameroseius lanatus Solomon, 1969 is probably no longer available (Adina Cǎlugǎr made a notable effort to find the type specimens), this species is also placed in synonymy with K. tenella. I am not able to detect any consistent morphological differences in the material from Italy, Romania, Germany and Slovakia, and thus all are considered to belong to a single species.
Kleemannia tenella and the closely related Chinese species Kleemannia guyimingi (Ma, 1997) and Kleemannia longisetosa (Ye & Ma, 1993) differ so markedly from the other members of this genus in the length of the dorsal shield setae (J-setae lengthened, Z-setae shortened), and in the arrangement of the ventral shields (pseudo-metasternal platelets absent; setae st3 on soft integument; ventrianal shield reduced and rounded, concave anteriorly, with two pairs of setae very close to its anterior margin), that they might be considered as belonging to a separate species group.
Remarks. This genus now comprises 22 species based from tropical and subtropical areas of Africa (eight species), Asia (ten species), Australia and Oceania (three species) and South America (one species). Neocypholaelaps favus and Neocypholaelaps apicola seem to be the only two species distributed in temperate zone of the Palaearctic region (see remark under N. favus), whereas Neocypholaelaps novaehollandiae was originally reported from temperate region of the southern hemisphere, New Zealand. Neocypholaelaps species belong to the nectar-and pollen-feeders associated with various flowers and flower-visiting animals (Evans 1963a, Klimov et al. 2016, namely apid bees (Hymenoptera) and butterflies (Lepidoptera). Neocypholaelaps wilsoni is reported from nasal cavities of a psittacid bird in Papua New Guinea (Allred 1970).
Neocypholaelaps favus, originally known from Japan (Ishikawa 1968), was firstly reported from Europe by Haragsim et al. (1978), based on findings in apiaries in the Czech Republic. There are several other reports of the species of Neocypholaelaps from Europe (all in association with Apis mellifera), namely from Greece by Emmanouel et al. (1983), from Denmark by Schousboe (1986), from Belgium by Fain and Hosseian (2000), from Slovakia by Fenďa and Lukáš (2014), and from Hungary by Kontschán et al. (2015). The specimens from Belgium and Hungary are reported as Neocypholaelaps apicola, remaining as N. favus. There is one further finding of Neocypholaelaps reported from Europe, a species described as a new Ameroseius from Crimean Peninsula by Livshits and Mitrofanov (1975), namely Ameroseius bregetovae. It apparently belongs to the genus Neocypholaelaps although it was not found in association with flowers or flower-visiting animals.
Remarks. I suspect this species to be a junior synonym of Neocypholaelaps stridulans (Evans, 1963). Differential characters for separating Neocypholaelaps ceylonicus from N. stridulans given by Narita and Moraes (2011) and Silva et al. (2014) are questionable and not well founded.
Remarks. In the original descriptive paper on Neocypholaelaps apicola from Pakistan by Delfinado- Baker and Baker (1983), the main character for distinguishing this species from the other congeners was based on the reduced length of some setae in central region of the dorsal shield (j4-j6 and z5 in female, and j5 in male and deutonymph), when they are compared with other dorsal shield setae. Unfortunately, those authors did not provide any metric data for these setae and a comparison with the most related species, Neocypholaelaps favus. Later, in their identification key to the species of Neocypholaelaps, Baker and Delfinado-Baker (1985) misinterpreted the form of dorsal setae in N. favus. They separated N. favus from N. apicola (see their couplet 3) by the presence of leaf-like dorsal setae in N. favus, whereas those setae are regularly thickened in this species.
A relatively large number of Neocypholaelaps favus collected from debris on the bottom of bee hives in Slovakia enabled an adequate examination of the morphology of this species. The length and form of some medially inserted setae (except j5) showed a relatively high degree of variability. Setae j4, j6 and z5 are more or less abbreviated and narrowed in almost all Slovakian females of N. favus. But mostly in freshly moulted, albescent and weakly sclerotised specimens, these setae are usually better developed, relatively larger and thicker than in the older, brown and strongly sclerotised adult females (rarely j4 and j6 are subequal with j3 and J2). I have compared the Slovakian specimens with a paratype female of N. favus (kindly sent me by Kazuo Ishikawa), and I could not find any important differences. In N. favus, j5 are the shortest and often subequal in adults of both sexes, and they can be smooth or with fine pilosity. Based on the chaetotaxy described above, and a series of photos taken from holotype of closely related Neocypholaelaps apicola received from Debbie Creel and Ronald Ochoa (USDA ARS, Beltsville Agricultural Research Center, Systematic Entomology Laboratory), I consider Neocypholaelaps favus and N. apicola as two distinct species, and Neocypholaelaps bregetovae n. comb. as a synonym of N. favus. In addition, Neocypholaelaps apicola sensu Kontschán et al. (2015) is considered here as a misidentification of N. favus.
The sperm induction structures of the female, especially the structure of sacculus foemineus and associated tubuli annulati, were considered to be important for separation of some Neocypholaelaps species (ampullula, cocos, indicus, novaehollandiae and stridulans) in taxonomic work of Evans (1963a). Because of the weak sclerotization of these structures, it is not easy to discern the sacculus in Neocypholaelaps favus, especially in dark brown mature specimens (Plate 66A). In some freshly moulted and weakly sclerotised immature females of N. favus I could detect the sacculus foemineus as small, conspicuous and well outlined pear-shaped structure having well developed neck-like process that is connected with tubuli annulati (Plate 66B); the tubuli are free and enter the narrowed part of the sacculus separately. In adult females with egg, or in females after oviposition, the sacculus foemineus with neck-like process was not detectable in any of examined specimens (probably due to its conspicuous expansion), and the tubuli annulati had their bases well distant from each other, and much more separated when compared with immature females of N. favus.

Neocypholaelaps indicus Evans, 1963
Neocypholaelaps indica Evans, 1963a: 217. Neocypholaelaps indica. -Teng andPan 1964: 772;Mo 1971: 97;Mo 1972: 15;Treat 1975: 116;Baker and Delfinado-Baker 1985: 232;Haitlinger 1987a Remarks. The original description and illustrations of Neocypholaelaps ewae are not detailed and consistent enough to allow it to be correctly recognised and classified in a genus with any confidence. Therefore, Narita et al. (2013a) and Klimov et al. (2016) transferred this species to Hattena, without providing any explanation for the new systematic position. The number of 19 pairs of dorsal shield setae stated for the adults (and developmental stages, protonymph and deutonymph) by Haitlinger (1987a) perfectly fits to the setal patterns of Hattena. But many other characters resembling those in Neocypholaelaps, especially Neocypholaelaps indicus: large subrectangular epigynal shield with slight posterior expansion, dorsal setae smooth (except j1), anal shield with only three circum-anal setae in adults of both sexes, cheliceral spermatodactyl thin and long, J2 of male conspicuously thickened and lengthened, protonymph with S4 thickened and spur-like. My examination of the type specimens of N. ewae has shown that the statement on decreased complement of the dorsal setae is misinterpreted and in error. I could found all 29 pairs of setae on the dorsal shield in examined adults of both sexes. Using the key to species of Neocypholaelaps by Evans (1963a) and observing the type female from Vietnam, it can be identified as N. indicus by the following characteristics: dorsal shield has 29 pairs of setae (entry 1), most of which setiform and at the most weakly serrated (entry 2), all leg ambulacra with two claws and genu and tibia III each with two anterolateral setae (entry 3), Z5 short (about 20 μm long in type female), dorsal setae of tibia I smooth, and anal shield approximately 104 × 135 μm in size in type female (entry 4). Accordingly, a new synonymy is established between N. ewae and N. indicus in the present study.
Type locality and habitat. Indian Ocean Region, Réunion Island, on inflorescence of coconut palm, Cocos nucifera (Arecaceae).

Neocypholaelaps leopoldi Elsen, 1972
Neocypholaelaps leopoldi Elsen, 1972a Gupta, 1969 Neocypholaelaps pradhani Gupta, 1969: 335. Neocypholaelaps pradhani. -Gupta 1985 Remarks. I examined a well preserved paratype of Ameroseius wilsoni deposited at the Natural History Museum in London, concluding that it should be included in Neocypholaelaps. The original description and illustrations given by Allred (1970) are quite adequate and do not require amendment, except for the sperm induction apparatus. In the studied specimen, spermathecal ducts (tubuli annulati) are well indicated, and they enter the sacculus foemineus independently.
Remarks. Pseudoameroseius gen. n. seems to be monotypic, with Pseudolaelaps michaelangeli described from soil and leaf litter in Tenerife Island, Canary Islands (Moraza 2006 Remarks. Moraza (2006) added complete leg chaetotaxy to the description of michaelangeli. According to her original data, genu and tibia of legs IV would have one anterolateral seta and two posterolaterals, but in the type specimens I found the typical two anterolaterals and one posterolateral seta instead (including some other slight differences in leg chaetotaxy, as detected by a comparison with data presented in Table 1 and in the original description).
Diagnosis (adults). Dorsal shield variously sclerotised and ornamented, with 28-29 pairs of setae, including z6; setae Z2 absent in some species. Dorsal shield setae similar in adults of both sexes, relatively short (with tips never reaching bases of the following setae of the same longitudinal rows), mostly smooth and spatulate, sometimes crenelated, paddle-shaped or spoon-shaped, with a broad flat end tapering to the base; setae j1 otherwise modified, smooth or serrate, usually with pointed apex, and often with ventral position on vertex. In female, st1 and st2 on sternal shield, st3 on small suboval or subtriangular pseudo-metasternal platelets, and st4 on soft integument; endopodal platelets II-III well developed, relatively large, subtriangular and close to posterolateral angles of sternal shield; anterior margin of epigynal shield usually deeply concave to form two cusps, genital poroids on soft integument. Opisthogastric region with five pairs of setae in females and males (JV1-JV3, JV5, ZV2; JV4 absent); setae JV5 usually similar to those on dorsal shield, other ventrally inserted setae simple, smooth and needle-like; females with three pairs of opisthogastric setae (JV1, JV5, ZV2) on soft integument and two pairs (JV2, JV3) on ventrianal shield; male with more expanded ventrianal shield bearing all five pairs of opisthogastric setae. Peritrematal shields anteriorly fused to dorsal shield, with enlarged cavity-like poroid structure at the level of coxa III and well developed parapodal portion. Metapodal platelets relatively large, longitudinally elongate. Soft striate integument simple, not with additional sclerotic granulation. Corniculi normally sclerotised, well spaced and trifid; setae h1 markedly thickened proximally, curved and abruptly tapered medially, attenuate distally, and sharply pointed apically. Fixed digit of chelicera with three subequal teeth on proximal masticatory area; male spermatodactyl small, shorter than movable cheliceral digit. Epistome with anterior margin produced into narrow and bifid central projection; lateral margins with several minute spines. Palptarsal apotele two-tined. Coxae I-IV with rows of minute spines, coxae II with spine-like process on anterior surface; coxae I-IV and femora I-IV with lamellar and/or papular sclerotic structures. Genu III, and tibiae III-IV with two anterolateral and two posterolateral setae. Tarsi I-IV each with empodium and rudimentary claws, or claws absent. Insemination apparatus with papilla-like sperm induction pores (solenostomes) associated with inner margin of coxae III; spermathecal ducts free and elongated. Remarks. Elsen and Whitaker (1985) proposed the genus Sertitympanum to accommodate three species based on females and found in North America, the type species Sertitympanum separationis, as well as Sertitympanum exarmatum and Sertitympanum contiguum. They characterised this genus by the presence of leg excrescences, peculiar cog-wheel-like sculpture on sternal shield and presence of a pair of large ventrolateral lenticular metapodal platelets (and by many other characters). These characters were considered by them to be the most important and distinct for their newly established genus, although they can be detected also in the other ameroseiid species, especially in the genus Kleemannia (for example, rounded and well scalloped circular structure on the sternal shield can be found in Kleemannia plumosa).
The two further species of the genus, described from the North Africa (Egypt) by Nasr and Abou-Awad (1986), are lacking the cog-wheel-like sculptural pattern on the sternal shield, but despite this and slightly different appearance they are congeneric with the species from the American continent in many character states as follows: (1) form of dorsal shield setae; (2) dorsal chaetotaxy (z6 present and Z2 absent, as in Sertitympanum separationis); (3) general arrangement of ventral shields; (4) bicuspid anterior margin of epigynal shield; (5) chaetotaxy of opisthogastric soft integument and ventrianal shield; (6) presence of cavity-like poroid structure on peritrematal shield; (7) presence of papilla-like sperm induction pores close to coxae III; (8) sclerotic sculpture and chaetotaxy of legs; (9) reduction of tarsal claws in legs I-IV (not stated in an original description of the genus, but found in S. separationis in which the claws are fully reduced); (10) special form of h1; (11) dentation of chelicera; (12) trifid form of corniculi; (13) bifurcate form of epistome (not stated in an original description of the genus but found in S. separationis); and (14) two-tined form of palptarsal apotele.
Sertitympanum is similar in some respects to Kleemannia. Both genera have the same leg chaetotaxy, two pairs of opisthogastric setae on ventrianal shield, and arrangement and shape of some ventral shields. As a result of this similarity and the absence of a cog-wheel-like sculpture on sternal shield in some species, three Sertitympanum species were initally described in Kleemannia (bassolsae, nodosum and zaheri) and classified in this genus up to now. Sertitympanum can be distinguished from Kleemannia by the following characters: (1) dentation of fixed cheliceral digit (three teeth in Sertitympanum, four in Kleemannia); (2) structure of ambulacral apparatus (claws rudimentary or absent in Sertitympanum, claws normally developed in Kleemannia); (3) form of central process of epistome (furcate in Sertitympanum, undivided in Kleemannia); (4) form of h1 (thickened proximally, curved and attenuate medially, subfalcate in Sertitympanum; thin or thickened, always straight and regularly tapered in Kleemannia); (5) dorsal setation in forms with 28 pairs of dorsal shield setae (z6 present and Z2 absent in Sertitympanum, z6 absent and Z2 present in Kleemannia); (6) length and form of dorsal setae (shorter, with their apices and bases well separated, leaf-shaped, oblanceolate, spatulate or obovate, with rounded apex in Sertitympanum; longer, with their apices and bases adjacent or overlapping, tubiform or feather-shaped, with pointed apex in Kleemannia).
There were five very similar species described to this time (aegyptiacum, bassolsae, nodosum, palmatum and zaheri), with only simple reticulation on the sternal shield, although some curved sculptural lines, and perhaps also shallow depression, are present and better defined. Sertitympanum nodosum was introduced first (Sheals, 1962), based only on a single specimen, a weakly sclerotised and medially distorted female, found in Argentina. Unfortunately, the followers overlooked the existence of S. nodosum in a differential diagnosis to their new species (El-Badry et al. 1979, Nasr and Abou-Awad 1986, Vargas and Polaco 2001. Sertitympanum needs a thorough revision, and it comprises eight species considered here to be valid, most of which are widespread in nests of small mammals (species from North America), soil detritus and litter (species from Egypt). In Iran, Sertitympanum aegyptiacum is reported from honey bee hives, stored rice, rice dust and debris, and soil of vineyards (Kazemi and Rajaei 2013). The genus Sertitympanum is here firstly recorded from Europe (see remarks in Sertitympanum nodosum). Remarks. I have seen two females of this species from the British Museum Collection in London, both collected from small mammals in Utah in 1955, and additionaly identified by H. Hurlbutt as Kleemannia plumosa. These specimens are most probably a part at least of the material forming the record by Allred and Beck (1966) of Kleemania (sic) sp., who added also some fragmentary illustrations to this unidentified species. The original description given by Elsen and Whitaker (1985) requires a small amendments: vertex simple, not produced into bifid spur supporting j1 as stated in the original generic diagnosis; setae j1 smooth, relatively short and thick, wedge-shaped, with ventral position on vertex; epistome with bifurcate central process and spinate lateral margins; tarsi of legs I-IV with empodium but no claws).
Type depository. United States National Museum, Washington, D.C., USA.

Sertitympanum mexicanum Villegas-Guzmán, Montiel-Parra, Vargas & Polaco, 2004
Sertitympanum mexicanum Villegas-Guzmán, Montiel-Parra, Vargas & Polaco, 2004 Evans in 1964or G. Rack in 1964, 1973 Here I regard their identification as correct. The first record of Sertitympanum in Europe is perhaps unexpected and surprising from a biogeographic and ecological point of view, but specimens from Germany agree in detail with the specimens from South America and Asia. I expect S. nodosum to be cosmopolitan species distributed by human activities and associated with various synanthropic habitats, including stored grain and other food products, especially in the temperate climatic regions.
rounded. However, I suspect this species to be a junior synonym of Sertitympanum nodosum. Photographed specimens stated to be zaheri and originated from Egypt (kindly sent by R. Abo-Shnaf ) apparently belong to S. nodosum.
Diagnosis (female). Dorsal shield weakly sclerotised but coarsely ornamented, with 29 pairs of setae (z6 present). Dorsal setae including j1 similar in length and form, odd-pinnate. Sternal setae on sternal shield (st1, st2, st3), soft integument (st4) and epigynal shield (st5); opisthogastric soft integument with five pairs of setae (JV1-JV3, JV5, ZV2), all on soft integument (anal shield only with three circum-anal setae). Opisthogastric setae mostly short, smooth and needle-like; setae JV5 short or long, thickened and densely pilose to plumose; postanal seta smooth or pilose. Corniculi well sclerotised, relatively slender, with incised apex and subdistal tubercle; setae h1 slightly thickened. Cheliceral digits relatively large, fixed digit with three well developed teeth on proximal masticatory area (two proximal teeth somewhat adjacent). Epistome curved, smooth or with minute serrations. Palps relatively small, palptarsal apotele three-tined. Genu III and tibia III each with two anterolateral and one posterolateral setae, genu IV with two ventral setae, tibia IV with two anterolateral and two posterolateral setae. Tarsi I-IV each with normal empodium and claws. Remarks. Bai et al. (1995) established their originally monotypic genus Sinoseius on the basis of the type species Sinoseius lobatus collected in a mammal nest in China. The genus was characterised especially by the form of the dorsal shield setae and three pairs of setae on sternal shield. Karg (2005) and Karg and Schorlemmer (2009) synonymised Sinoseius with Ameroseius Berlese, 1904. Later, Karg as a collaborating author validated the genus by the description of a congeneric species, Sinoseius pinnatus, found in detritus from a straw shed in Finland (Huhta and Karg 2010). Barilo (1986) included one further new species to the genus, collected from soil and wood substrate in Central Asia (Uzbekistan, Tajikistan). The species he named as Ameroseius fossatus is here transferred to Sinoseius.
The genus Sinoseius is considered here to be a valid genus, based on the following combination of female characters (male is unknown): (1) sternal shield with three pairs of setae; (2) unusual chaetotaxy of some leg segments: genu III and tibia III each with two anterolateral and one posterolateral setae (as in Ameroseius), genu IV with two ventral setae (not found in Ameroseius and Kleemannia), tibia IV with two anterolateral and two posterolateral setae (as in Kleemannia); (3) dorsal shield setae pinnate; (4) flat curved epistome; (5) cheliceral digits robust, fixed digit with three large teeth; (6) all opisthogastric setae on soft integument; (7) palps relatively small, palptarsal apotele three-tined. Some of these character states occur in other genera of Ameroseiidae, but infrequently, and not in combination. For example, three pairs of setae on sternal shield can be found also in Kleemannia miranda sp. n., while specific pinnate setae are typical of Ameroseius avium and Kleemannia bella.
Remarks. Sinoseius fossatus cannot be distinguished on morphological grounds from Sinoseius lobatus, except for the form of JV5. These setae are considerably lengthened and similar in length and form to those on dorsal shield, as depicted in the original illustration of Barilo (1986), and also in the illustration of a Sinoseius specimen found in Iran (based on personal communication from Shahrooz Kazemi).

Sinoseius lobatus Bai, Gu & Fang, 1995
Plates 73-75 Sinoseius lobatus Bai, Gu & Fang, 1995 Remarks. The first European finding of the genus Sinoseius was that of Huhta and Karg (2010) from Finland (bottom of straw shed and grass in garden of old farm). Fenďa and Lukáš (2014) found the species in a frozen nest of Sitta europaea Linnaeus, 1758 (Aves, Passeriformes), in a nest box, and in a soil sample from the dysphotic zone of a cave. I have compared the holotype and one paratype of Sinoseius pinnatus with the specimens reported from Slovakia by Fenďa and Lukáš (2014), and found that they are clearly conspecific. According to Huhta and Karg (2010), Sinoseius lobatus and S. pinnatus can be distinguished by the features presented as follows: in S. lobatus, dorsal setae "remarkably long", j5 = j5-j6, j6 > j6-J2, tines of dorsal setae strong (depicted seven pairs of tines), anal shield distinctly wider than long (length/width = 2:3), corniculi bifid; in S. pinnatus, dorsal setae "moderate", j5 = 2/3 × j5-j6, j6 < j6-J2, tines of dorsal setae very thin (depicted up to 19 instead of 9-10 pairs of tines), anal shield only a little wider than long, corniculi distally trifid. It is clear that the distinctions made in the original descriptions and differential diagnosis of S. pinnatus by Huhta and Karg (2010) are based on characteristics that are misinterpreted (form of setae and corniculi) or vary considerably (relative length of dorsal setae, proportion of anal shield), and I do not hesitate to propose the synonymy of both mentioned species, although no types of S. lobatus were examined in this study. When compared the Slovak specimens of Sinoseius with quite adequate original illustrations of S. lobatus from China, it was not possible to find reliable distinguishing characters between them.

Unrecognizable species (species inquirendae)
The following species are temporarily relegated to the species "incertae sedis" because the descriptions do not include information about some important characters needful for their specific identification. If some of them are correctly placed in Ameroseiidae, then they cannot be included in the above classification. I dare to say that many species in this list can be excluded from Ameroseiidae after their revision (if their type specimens are available for a study).
Type locality and habitat. Argentina, La Plata, habitat not stated. Remarks. Actinoseius Berlese, 1916 was originally introduced as a subgenus of the genus Epicriopsis Berlese, 1916, considering Epicriopsis (Actinoseius) terrificans, then described, as the type species. Castagnoli and Pegazzano (1985) did not include E. (A.) terrificans in their Catalogue of the Berlese Acaroteca. Hallan (2005) included Actinoseius in his compiled list of pachylaelapid genera. Original description of this taxon and of its type species is absolutely useless for species and subgenus recognition.
Thus, the reason leading Hallan (2005) to include it in Pachylaelapidae is not clear. Most likely, this species could be based on a specimen mounted onto slide No. 162/9 of the Berlese Collection in Florence, and labelled Ameroseius terrificans (nomen nudum). That specimen and, according to the original description, the type specimen of E. (A.) terrificans were collected in La Plata, Argentina. Berlese, 1916 Ameroseius epicrioides Berlese, 1916a: 46. Ameroseius epicrioides. -Castagnoli andPegazzano 1985: 130. Type depository. Istituto Sperimentale per la Zoologia Agraria, Firenze, Italy.

Ameroseius epicrioides
Type locality and habitat. Argentina, Rio Santiago, La Plata, under bark of tree.
Remarks. None of the two slides in the Berlese Collection bears a type designation, but without doubt they belong to the original type series of Berlese. Unfortunately, a female on slide 208/33 is un suitable for study and will require remounting for definitive study, whereas the female on slide 210/39 must be lost because I was unable to find any specimen or dissected structures on that slide (both slides are opaque and essentially unusable).

Remarks.
Hull's description and illustration of this species are not detailed enough to allow it to be correctly recognised and classified in a genus with any confidence. The illustrated specimen resembles a Cheiroseius species.
Remarks. Hull's description and illustration of this species are not detailed enough to allow it to be identified to the generic level with any confidence.

Species excluded from Ameroseiidae
Below is a list of species that have been incorrectly classified in Ameroseiidae at some time, but are now placed in other families. The species are listed in alphabetical order of original species names, with current valid name and systematic placement of each species. The species transferred to Ascidae, Blattisociidae or Melicharidae were checked using recent catalogue for these families by Moraes et al. (2016) to confirm their current systematic placement. Berlese, 1910b: 253. Type locality: Italy -Sicily. = Zerconopsis remiger (Kramer, 1876), Ascidae -synonymy by Evans and Hyatt (1960) Ameroseius borealis Berlese, 1904: 259. Type locality: Norway. = Cheiroseius borealis (Berlese, 1904), Blattisociidae -transferred by Westerboer (1963) Ameroseius crassipes Berlese, 1910c: 370. Type locality: Australia. = a species of unspecified genus of Ologamasidae -by Halliday (1997) Remarks. Berlese (1916a) transferred this species from Ameroseius to the subgenus Lasioseius (Leioseius). Bernhard (1963) did not consider Ameroseius crassipes to belong to Leioseius, but probably to Ameroseius. Halliday (1997) transferred it to Ologamasidae, based on his personal communication with Evert E. Lindquist who examined the type material of A. crassipes in the Berlese Collection in Florence. Berlese, 1910b: 254. Type locality: Tasmania. = a species of an unspecified genus of Phytoseiidae -by Halliday (1997), based on his personal communication with Evert E. Lindquist

Synopsis
This catalogue of the family Ameroseiidae includes 12 genera and 138 valid species. Another nine species are temporarily relegated to the unrecognizable species (species inquirendae), because the descriptions do not include information about important characters required for identification, and are here regarded as of unknown systematic position. In summary, altogether 206 named species (including 37 synonyms, 15 species previously excluded from the family, and seven "nomina nuda") are mentioned in this paper. The most diverse genera are Ameroseius, Kleemannia and Neocypholaelaps, which include 50 (36%), 28 (20%) and 22 (16%) species, respectively. Two genera are monotypic, Afrocypholaelaps and Pseudoameroseius gen. n. The objective of this paper was not to present a detailed historical-taxonomic review of the family Ameroseiidae, but to analyse and revise the current state of knowledge of the family and draw attention to some unresolved problems. This review and the identification keys should provide a useful basis for identification of world genera and the species from Europe in future taxonomic research. However, the research work of the ameroseiid mites is apparently far from complete and should continue, especially with detailed study of taxa from tropical areas of the world.