A new genus and species in the mite family Eupodidae (Acari, Eupodoidea) from Crimea

Abstract A new genus Pseudoeupodes Khaustov, gen. n. and new species Pseudoeupodes porosus sp. n. are described from moss in Crimea. The taxonomy of the Eupodidae and some other families and genera of Eupodoidea is reviewed. The genus Turanopenthalodes Barilo, 1988 is transferred from Penthalodidae to Penthaleidae. The family Cocceupodidae Jesionowska, 2010 and the genus Filieupodes Jesionowska, 2010 are considered as junior synonyms of Eupodidae Koch, 1842 and Cocceupodes Thor, 1934, respectively. A key to genera of the family Eupodidae is provided.

The family Eupodidae includes two subfamilies: Benoinyssinae Fain, 1958 andEupodinae Koch, 1842, although this subdivision is not followed by most workers. The genera Cocceupodes Thor, 1934 andLinopodes Koch, 1835, which were previously placed in the family Eupodidae, were recently transferred to the separate family Cocceupodidae (Jesionowska 2010); and the genus Hawaiieupodes Strandtmann & Goff, 1978 was transferred to the family Penthalodidae (Jesionowska 2008).
This paper presents a description of a new genus and species of eupodid mite, Pseudoeupodes porosus gen. n., sp. n., collected from moss in Crimea, and discusses the taxonomy of some families and genera of Eupodoidea.

Materials and methods
Mites were collected from moss using Berlese funnels and mounted in Hoyer's medium. Notations for the prodorsal and leg setae follow Lindquist and Zacharda (1987) and Baker (1995), and the remaining nomenclature is as applied to eupodoids by Baker (1990). All measurements are given in micrometres (μm) for the holotype and for five paratypes (in parentheses). In descriptions of leg setation the number of solenidia is given in parentheses. Photographs were taken with a digital camera Tucsen 3.0 via the ocular of light microscope MBI-11 with phase contrast device. The type material is deposited in the mite collection of the Tyumen State University, Tyumen, Russia.
Male and immatures. Unknown. Etymology. The genus name is derived from the related genus Eupodes and prefixed pseudo-(from Greek ψευδής) meaning false.
Etymology. The name of the new species refers to the presence of unusual pore-like structures in the soft cuticle between the coxisternal plates and the trochanters of all legs.

Discussion
The present systematic organisation of the family Eupodidae and the superfamily Eupodoidea is highly unstable. Preliminary morphological cladistic analyses (Qin 1996, Qin andHalliday 1997) suggested that only two families (the Eriorhynchidae and Penthalodidae) are monophyletic. Of the other four, the Rhagidiidae plus Strandtmanniidae formed a monophyletic group, the Penthaleidae was paraphyletic, and the status of the Eupodidae was not resolved (Baker and Lindquist 2002). Jesionowska (1989) described the genus Protopenthalodes Jesionowska, 1989 in the family Penthalodidae and moved Hawaiieupodes Strandtmann & Goff, 1978 from Eupodidae to Penthalodidae (Jesionowska 2008). According to Jesionowska (2008), this new concept of the Penthalodidae will be published in a separate paper. Qin (1997) reconsidered the taxonomic position of Protopenthalodes and suggested that this genus is more appropriately placed in the family Eupodidae because of soft body integument. Barilo (1988) described Turanopenthalodes Barilo, 1988, which is another problematic genus in the family Penthalodidae. The key characters of the family Penthalodidae are the fully sclerotized body and the presence of an epirostrum projecting over the gnathosoma (Qin and Halliday 1997;Walter et al. 2009), but other characters such as the idiosomal setation have never been used to separate Penthalodidae from other families of Eupodoidea. In my opinion the genera Hawaiieupodes and Protopenthalodes are more closely related to Penthalodidae than to Eupodidae, in agreement with the suggestion by Jesionowska (2008). Like Penthalodes Murray, 1877, the type genus of the family Penthalodidae, both Hawaiieupodes and Protopenthalodes have the following synapomorphies: sejugal furrow not developed, lens-like eyes present near the setae sc 2 , setae h 2 absent, naso usually very small, oval with minute setae v 1 . This combination of characters is not found in the closely related families Penthaleidae and Eupodidae, and following Jesionowska (2008), I retain Hawaiieupodes and Protopenthalodes in the Penthalodidae. The position of the genus Turanopenthalodes in the family Penthalodidae is doubtful. Barilo (1988) placed Turanopenthalodes in Penthalodidae based on a single character, the presence of epirostral processes lateral to the naso, similar to those found in the penthalodid genus Stereotydeus Berlese, 1901, which he considered the main differential character of the family Penthalodidae. Other apomorphic characters of this genus, such as neotrichy of the idiosoma, a small anal opening situated dorsally, short and truncated palptarsus and very characteristic "trident" at the distal end of the fixed digit of the chelicera, are similar to those found in Penthaleus Dugès, 1834, the type genus of the family Penthaleidae, and I currently place Turanopenthalodes in the family Penthaleidae. Undoubtedly some characters that are now used to separate some families in the Eupodoidea should be re-evaluated. Such an attribute as more conspicuously sclerotized dorsal body surfaces, which is characteristic of the family Penthalodidae (Walter et al. 2009), is highly variable. There is an undescribed species of Protopenthalodes in my collection with a soft body, but having subcuticular reticulate ornamentation throughout the body surface, which I consider as intermediate in the extent of body sclerotization.
The monotypic family Dendrochaetidae (Olivier , 2009, which includes only the genus Dendrochaetus Olivier, 2009, shares some synapomorphic characters with Hawaiieupodes and Protopenthalodes. All these genera lack setae h 2 , the sejugal furrow is absent, the naso is small, almost round, and well separated from the anterior margin of the prodorsum. The only difference between Dendrochaetidae and soft-bodied Penthalodidae (Hawaiieupodes and Protopenthalodes) is the presence of an additional transverse furrow at the level of setae v 1 . Dendrochaetes acarus  needs to be restudied and redescribed to clarify the status of the family Dendrochaetidae. Jesionowska (2010) erected the family Cocceupodidae, in which she included three genera: Cocceupodes Thor, 1934, Linopodes Koch, 1835and Filieupodes Jesionowska, 2010. According to Jesionowska (2010) the family Cocceupodidae differs from Eupodidae by two main characters: setae v 1 situated posterior to naso and the presence of only two pairs of circumanal setae (ps 1 and ps 3 ). In my opinion, the decision to create the family Cocceupodidae is groundless. The similar location of setae v 1 on dorsal part of naso near anterior margin of prodorsum is also found in the eupodid genera Caleupodes, Niveupodes and Pseudoeupodes, but in these genera the naso is not directed anteriorly, but folded to the ventral side of the prodorsum. The presence of only two pairs of pseudanal setae (ps 2 absent, according to Baker 1990) is a variable character in the family Eupodidae. Setae ps 2 are absent in the genera Caleupodes, Niveupodes, Pseudoeupodes and Benoinyssus Fain, 1958. The absence of setae ps 2 is a reduction and could happen independently in different lineages of eupodoid mites (homoplasy). On the other hand, Jesionowska (2010) did not mention some synapomorphic characters of Cocceupodes and Eupodes Koch, 1842. Both genera have characteristic swollen femora IV adapted for jumping. Another synapomorphic attribute is the relatively long and thin legs I, which are usually subequal to or longer than the idiosoma, and much longer than legs II. In early derivative genera of Eupodidae, such as Neoprotereunetes Fain & Camerik, 1994, Caleupodes and Niveupodes, femora IV are not swollen and legs I are not so long and thin. Thus, the characters separating Cocceupodidae and Eupodidae proposed by Jesionowska (2010) are variable within the family Eupodidae. I therefore consider the family Cocceupodidae as a junior synonym of Eupodidae.
Jesionowska (2010) created a new genus Filieupodes, which differs from Cocceupodes by a single character, the filiform setae v 1 (clavate or capitate in Cocceupodes). Filiform setae v 1 is a plesiomorphic character state and should not be used for recognition of a new taxon. I therefore consider Filieupodes as a junior synonym of Cocceupodes.