Revision of the genus Neoseiulella Muma (Acari: Phytoseiidae). Re-description of species, synonymies assessment, biogeography, plant supports and key to adult females

Identification of species within the family Phytoseiidae is of primary importance for biological control programs. However, considerable disagreements between authors are often observed concerning the placement of species in genera. Moreover, considering the small number of available morphological characters and the poor knowledge of their reliability for species diagnostics, synonymies between several species could be questioned. This study aims to provide a taxonomic revision of a genus belonging to the sub-family Typhlodrominae: Neoseiulella Muma. Whereas the last revision of this genus (carried out in 1996) took into account 26 valid species, we propose now to include 47 species in the genus Neoseiulella based on type material observations. At first, three species (Neoseiulella eiko, N. eleglidus and N. schusteri) have been excluded from this genus, as their morphological attributes do not match the morphological characters which define the genus. Secondly, we also think that another species placed in the genus Typhlodromus (Anthoseius): T. (A.) elisae, has to be included in the genus Neoseiulella. Finally, several synonymies have been analysed and we conclude that: (i) N. aceri is a synonym of N. squamiger and provisionally of N. aceris, (ii) N. tiliarum is a synonym of N. formosa, (iii) N. transitans is a synonym of N. prunus, and provisionally of N. vollsella, (iv) N. manukae is a synonym of N. glenfieldensis and (v) Neoseiulella tuberculata, N. sexapori and N. arutunjani are considered valid specific entities. A new synonymy between Neoseiulella nesbitti and N. armidalensis is also proposed. A standardised re-description of all the species of the genus is proposed and a key to the species for the adult females is provided. This study also gives an exhaustive compilation of all available information concerning the geographical distribution of species of Neoseiulella and their plant supports.


INTRODUCTION
Over the past fifty years, mites of the family Phytoseiidae Berlese (Acari: Mesostigmata) have been widely studied because of the ability of several species to control phytophagous mites and small insect pests in various crops worldwide (Kostiainen and Hoy 1996;McMurtry and Croft 1997). Tax-onomy studies, especially those dealing with diagnosis, are thus of first importance to ensure biological control program successes. Furthermore, knowledge of Phytoseiidae diversity can open new insights for the control of new pests. McMurtry (1994, 2007) recognized 2,280 species of Phytoseiidae dispatched in three sub-families (Amblyseiinae, Phytoseiinae, and Typhlodrominae) and 84 genera. Species identification is mainly based on the morphological characters of females, as dorsal chaetotaxy, the shape and the setation of the ventral shields, the spermatheca shape, the setation of legs and the morphology and dentition of the chelicerae (Chant and McMurtry 2007). Identification of species is difficult, essentially because of the small number of available visible characters and of the poor knowledge on their reliability for species diagnostic. Thus, species validity (synonymy) is often questioned and disagreements between authors are noted in literature. Furthermore, as new taxa are discovered, as phylogenetic analyses are developped (Kanouh et al. 2010a), genera description and attributes could be modified, as well as the number of species belonging to them.
The aim of the present study is to provide an updated and complete revision of one genus belonging to the sub-family Typhlodrominae, tribe Typhlodromini: Neoseiulella Muma. The last revision of this genus carried out by Denmark andRather in 1996 included 26 valid species, whereas in 2004, in the last world catalogue of the family Phytoseiidae, Moraes et al. reported 43 species, 35 being considered valid. In 2007, Chant and McMurtry in their world revision of the family have included 46 species and did not consider the previous synonymies. Furthermore, these latter authors did not take into account one additional species described in 2006. The genus Neoseiulella is thus currently assumed to include 46 species. However, doubts on some synonymies still exist as authors did not agree (Denmark and Rather 1984;Chant and Yoshida-Shaul 1989;Denmark and Rather 1996;Kolodochka 2009). The revision presently provides complete morphological descriptions of females based on the examination of type material and includes all the species reported in the genus Neoseiulella. Males are also described for some species for the first time. Moreorer, this paper presents an exhaustive compilation of all available information on geographical distribution of species of the genus and their plant supports. The present data set constitutes thus not only a taxonomic revision but also an actualised catalogue of the genus Neoseiulella.
The first part of the paper focuses on an histor-ical review of the genus Neoseiulella. The morphological characters defining the genus Neoseiulella, and validity of species presently included in this genus are consequently analysed. Six cases of synonymy within this genus are then studied and discussed. Finally, redescriptions of the species that we consider valid, and a key to adult females, are provided.

HISTORICAL REVIEW OF THE GENUS Neoseiulella
A summary of the complex history of the genus Neoseiulella is given in the Table 1.
In 1959, Chant proposed the tiliarum species group in the genus Typhlodromus and included in it these three species. He characterized this species group by the presence of 11 pairs of lateral setae on the dorsal shield, and three or four pairs of preanal setae on the ventrianal shield.
In 1961, Muma proposed two new genera, Neoseiulella including N. nesbitti, and Typhloctonus including N. aceri and N. tiliarum. These two genera differed by the number of setae on the sternal shield, and by the occurrence of the preanal seta JV3 and macrosetae on the leg IV.
In 1962, Wainstein proposed two new subgenera differentiated according to the number of preanal setae: Nesbitteius including N. nesbitti and Kallistoseius including N. aceri and N. tiliarum. At the same time, Pritchard and Baker (1962) included these three species in the genus Typhlodromus (Seiulus) Berlese. In 1966, Ehara also used the sub-genus Typhlodromus (Seiulus) to designate the tiliarum species group.
In 1989, in their revision of the tiliarum group within the genus Typhlodromus, Chant and Yoshida-Shaul considered 26 species and synonymised the genera Neoseiulella, Typhloctonus, Shiehia, Pegodromus, and Heteroseiulus.
Then in 1994, in the revision of the sub-families Phytoseiinae and Typhlodrominae, Chant and Mc-Murtry divided the genus Neoseiulella in two species groups: the tiliarum species group (JV3 present) and the cottieri species group (JV3 absent).
In 1996, Denmark and Rather revised the genus Neoseiulella and also divided it into two sub-genera with three or four pairs of preanal setae: JV1, JV2, JV3 (present: N. [Neoseiulella], / absent: N. [Typhloctona]) and ZV2. Then, Kolodochka (2009) used the genus Typhloctonus for the species of the genus Neoseiulella occurring in Ukraine.
Still today, the definition of the genus Neoseiulella is not thus very clear, and species of the genus Neoseiulella could be included in other genera or in different sub-genera, depending on the author's opinions. In the present revision, as Mc-Murtry (1994, 2007), we considered the genera Typhloctonus, Heteroseiulus, Pegodromus, and Shiehia junior synonyms of the genus Neoseiulella.
Ventral shields -The female caudoventral pattern is JV: ZV; JV-3/ ZV; JV-3, 4/ ZV (Chant and McMurtry 1994). Sternal shield (35 to 95 µm long; 50 to 100 µm wide, at level of ST2), is smooth with two or three pairs of setae (ST1, ST2, ST3) and two pairs of poroids. A pair of metasternal setae (ST4) is inserted on separate platelets with a pair of small poroids. Genital shield smooth (95 to 170 µm long; 40 to 90 µm wide, at level of ST5), except for N. crassipilis that has genital shield distinctly reticulated. Three, four or six elongate platelets (for some authors genital sigilla in three pairs maximum) are situated between genital and ventrianal shields. For some species, these latter platelets or genital sigilla are fold under the genital shield. Ventrianal shield smooth, faintly striate, or reticulated (75 to 165 µm long; 45 to 170 µm wide, at level of ZV2), with three or four pairs of preanal setae: JV1, JV2, JV3 (present/ absent) and ZV2, usually with a pair of solenostomes gv3. Four pairs of caudoventral setae (ZV1, ZV3, JV4, and JV5) are situated on the integument surrounding the ventrianal shield. The caudoventral seta JV4 is absent for one species, N. oleariae (Collyer). This absence is also found in the species of the tribes Typhloseiopsini and Metaseiulini in the sub-family Typhlodrominae (Chant and McMurtry 1994). However, N. oleariae differs from these species by the presence of both Z1 and S4. The primary metapodal plate or inguinal sigilla for some authors (18 to 40 µm long; 2 to 11 µm wide) is elongate. The secondary metapodal plate (or inguinal sigilla) is of about 1/3 as long as the primary one. Some poroids surround the ventrianal shield.
Spermatheca -The cervix (2 to 16 µm long) has a cup-or U-shaped type, with atrium nodular to indistinct.
Chelicera -One to 13 teeth, without or usually with a pilus dentilis, are observed on the fixed digit of the chelicera. The movable digit (20 to 60 long) is edentate in some species but usually with 1 to 4 teeth.

MATERIAL AND METHODS
Type materials of the species of the genus Neoseiulella have been examined (whenever possible) and measured with a phase and differential interference contrasts microscope (Leica DMLB, Leica Microsystèmes SAS, Rueil-Malmaison, France) (40x magnification). All the measurements are given in micrometers (µm) and all measurements were performed with Perfect Image v. 7.6 software (Clara Vision). When more than one specimen was measured, mean and ranges (minimum -maximum) 263  Denmark and Rather 1996); N. compta (after Chant and Yoshida-Shaul 1989); N. multispinosa (after Tseng 1975); N. vollsella (after Denmark and Rather 1984)   were provided. The kind of material (holotype, paratype, syntype, lectotype and/or paralectotype) and the institution or the museum where this material is deposited are provided in the redescription of each species. Setal nomenclature used in this paper follows Lindquist and Evans (1965) as adapted by Rowell et al. (1978) for the Phytoseiidae, with modifications for the caudal region as given by Lindquist (1994). Idiosomal gland nomenclature follows Athias-Henriot (1975) and .
The type materials of 41 species were examined. Despite repeated requests, it has been impossible to borrow the type materials of the four following species: N. celtis (Denmark and Rather) from India, N. compta (Corpuz-Raros) from Philippines, N. multispinosa (Tseng) from Taiwan, and N. vollsella (Chaudhri, Akbar and Rasool) from Pakistan. Detailed characteristics of these latter species were thus taken from the original descriptions and re-descriptions (Table 2). All other species were redescribed, even for synonyms in order to have objective arguments based on the real observation and study of morphological characters. This was done in order to establish synonymies on scientific bases and not on claimed bases. Data on the geographical distribution and plant supports were mainly obtained from the world catalogue of the family Phytoseiidae (Moraes et al. 1986(Moraes et al. , 2004, from the original descriptions and from all the other publications reporting species of the genus Neoseiulella.

RESULTS
The result section is divided in five parts. The first part presents species that have been eliminated from the genus and those newly included. The second part presents a discussion on synonymies emphasized in literature. The third section deals with new suspected synonymies. The fourth part provides a re-description of the species we consider valid. Lastly, a key for female identification is proposed in the fifth part.

The species considered as belonging to the genus Neoseiulella
Are all the species presently known as Neoseiulella included into this genus?
Among the 46 species presently considered, three have been excluded from the genus Neoseiulella, as their attributes do not fit the morphological characters defining it. These species are N. schusteri, N. eiko and N. eleglidus.
Neoseiulella schusteri (Youssef and El-Brollosy) Moraes et al. 2004: 295;Chant and McMurtry 2007: 147. This species was not included in the genus Neoseiulella in the previous revisions of Chant and Yoshida-shaul (1989) and Denmark andRather (1984, 1996). However, it was considered a member of the genus Neoseiulella in the last world catalogue of the family Phytoseiidae (Moraes et al. 2004), and then in the revision of the family (Chant and Mc-Murtry 2007). As it has been impossible to borrow and thus to observe the type specimens (deposited in Plant Protection Department, Faculty of Agriculture, Cairo University, Egypt), an attentive examination of the dorsal chaetotaxy of the female, based on the original description, was carried out. We observe that the dorsal seta z3 is absent. This latter character with the presence of nine pairs of lateral setae on the dorsal shield (j3, z2, z4, s4, s6, S2, S4, S5, Z5) keys to the species Cydnoseius negevi (Swirski and Amitai), tribe Galendromimini (sub-family Typhlodrominae). El-Brollosy (personal comm. 2008) confirmed that N. schusteri had been misidentified, and that it is a junior synonym of C. negevi.

(iii). Neoseiulella eiko Walter
Neoseiulella eiko Walter 1997: 335; Moraes et al. 2004: 293;Chant and McMurtry 2007: 147. The examination of the type materials of this species shows that Z1 is absent, and Z2 is present. These two characters with the presence of z3, s6 and S5, and the absence of z6 clearly constitute a new dorsal chaetotaxic pattern within the family Phytoseiidae. We therefore propose species to be included in a new genus in the tribe Typhlodromini (sub-family Typhlodrominae). The description of this new genus will be on the scope of another publication. Material examined. The female holotype and three female paratypes, collected by Walter (1995) on leaves of rainforest trees near the Boulders (Babinda, Queensland, Australia). Type materials are deposited in the UQIC, Department of Entomology, University of Queensland, St Lucia, Australia.
Are there species currently placed in other genera that in fact belong to the genus Neoseiulella?
Working on the genus Typhlodromus (Anthoseius), we observed that the species Typhlodromus (Anthoseius) elisae (Schicha and McMurtry 1986) might be included in the genus Neoseiulella. Schicha and Mc-Murtry (1986), when described this species in the genus Typhlodromus, stated the absence of S5 on the dorsal shield. Furthermore, they drew nine pairs of ventrianal setae (4 preanals and 5 caudoventrals) whereas Phytoseiidae only could have eight (Chant and McMurtry 2007). As no species of Phytoseiidae presently described have the combination of characters beared by T. (A.) elisae: absence of S5, presence of both Z1 and S2, and five pairs of caudoventral setae, one of the hypotheses to explain such an 266 aberration would be that one of the caudoventral setae might correspond to the seta S5. If S5 is effectively present, T. (A.) elisae would key to the genus Neoseiulella. Chant and Yoshida-Shaul (1989) also considered this hypothesis as they thought that S5 might be present (despite the poor conditions of the holotype). They thus included this later species in the tiliarum group. However, Denmark and Rather (1996) did not agree with this hypothesis, and consequently, excluded it from the genus Neoseiulella. This species was then cited in the sub-genus Typhlodromus (Anthoseius) De Leon in the world catalogue of the family Phytoseiidae (Moraes et al. 2004) and in the last revision of the family (Chant and McMurtry 2007). However, the examination of the holotype ( Figure 1) clearly shows that S2 and Z1 (in the original description) are both present. This species thus does not belong to the genus Typhlodromus (Anthoseius) but to the genus Neoseiulella.
The questionements concerning T. (A.) elisae should alert us on the fact that in the literature other similar cases could exist, i.e. species placed in genera but clearly not belonging to them. In a further work, it would be worth to check for this.
Material examined -The female holotype deposited in the New South Wales Department of Primary Industries, Agricultural Scientific Collections Unit (Acarology), Orange Agricultural Institute, Forest Road, Orange NSW 2800, Australia.
Adult male -The male of this species is unknown.
Previous reports -N. elisae is only known from Western Australia on Eucalyptus sp. (Myrtaceae).
We thus consider 44 valid species in the genus Neoseiulella in the followings parts of this paper.

Are the synonyms previously suspected valid or not?
Six synonymies (including 15 species) have been advanced by different authors (Denmark and Rather 1984;Chant and Yoshida-Shaul 1989;Denmark and Rather 1996;Moraes et al. 2004;Kolodochka 2009). In the present paper, we will 267 Kanouh M. et al. 268 only focus on five of them, as it has been impossible to borrow and examine the type materials of Neoseiulella compta, N. multispinosa, and N. vollsella despite repeated requests during three years. For each synonymy, we present a complete description of the species based on the type materials.
97 long and 138 wide, distinctly reticulated and with five pairs of preanal setae and three pairs of poroids. Spermatodactyl with a terminal foot, and with a non-enlarged toe. This description is based on one male syntype.
Neoseiulella aceri was collected from a wide range of plant supports: Acer campestre L., A. macrophyllum Pursh, A. platanoides L., A. pseudoplatanus L., Acer sp.    Ventral shields (Figure 3b) -Sternal shield 37 long and 53 wide (at level of ST2), smooth with three pairs of setae (ST1, ST2 and ST3) and two pairs of poroids (posterior to ST1; anterior to ST3). ST3 on an elongate projection of the sternal shield. A pair of metasternal setae (ST4) on separate platelets with a pair of small poroids. Genital shield 130 long and 60 wide (at level of ST5), smooth. Four elongate platelets or sigilla between genital and ventrianal shields. One pair of poroids close to the genital shield (ST 5) and 3 pairs of poroids around the genital shield. Ventrianal shield subquadrateshaped 103 long and 86 wide (at level of ZV2), distinctly reticulated, with four pairs of preanal setae (JV1, JV2, JV3 and ZV2) and a pair of solenostomes gv3 posterior to JV3. Four pairs of caudoventral setae (ZV1, ZV3, JV4 and JV5) on the integument surrounding the ventrianal shield. JV5 17 long, smooth. Primary metapodal plate or inguinal sigillum 30 long and 3 wide. Spermatheca ( Figure 3c) -Cervix 4 long, cupshaped, with an enlarged atrium.
Chelicera ( Figure 3d) -Two teeth and a pilus dentilis on the fixed digit. Movable digit 25 long, is edentate.
Material examined -The female holotype deposited in the ASU, Institute of Zoology, Academy of Sciences, Kiev, Ukraine.
Dorsal shield chaetotaxy is similar to the female. Ventrianal shield 103 long and 142 wide, distinctly reticulated and with five pairs of preanal setae and three pairs of poroids. Spermatodactyl with a terminal foot, and a slightly enlarged toe.
This description is based on one of the male specimens of our collection (Montpellier SupAgro collection, UMR CBGP), collected in Kiev (Ukraine) on Norway maple (Acer platanoides L.).
Previous reports: N. aceris is only known from Pennsylvania (USA; Nearctic area). Plant supports on which this species was collected are: A. platanoides (Aceraceae); Aesculus hippocastanum (Hippocastanaceae); Ilex crenata 'rotundifolia' Thunb. The examination of the type materials of N. aceri, N. squamiger and N. aceris shows similar measurements. However, N. aceri differs from both N. squamiger and N. aceris by the position of the sub-lateral seta R1. Recent molecular experiments (Kanouh et al. 2010) showed that the position of sub-lateral seta (R1) is not a valid diagnostic criteria to distinguish between these species and that N. squamiger and N. aceri are synonyms (Kanouh et al. 2010). This agrees with the conclusions of Chant and Yoshida-Shaul (1989) and Kolodochka (1986). On the other hand, only one pair of solenostomes (gd9) is observed on N. aceris, whereas five pairs (gd1, gd2, gd6, gd8, gd9) are observed on N. aceri and N. squamiger. Lehman (1982) drew three pairs of solenostomes (gd2, gd6, gd8) that we did not observe on the specimen examined. As some authors have shown the importance of such characters for species differentiation (Chant and Yoshida-  Shaul 1987; Ragusa and Tsolakis 1994;Tixier et al. 2006a, b), but as we do not know if this reliability is universal for species diagnostic, the synonymy of N. aceris and N. aceri / N. squamiger is still questioned. Examination of other female paratypes of N. aceris, as well as molecular analyses or cross breedings, would thus be required. We thus consider that N. aceri is a senior synonym of N. squamiger, and we propose, at this time, that N. aceris as a provisional junior synonym of N. aceri.
Material examined -The female holotype deposited in the Rijksmuseum van Natuurlijke Historie, Leiden, the Netherlands. As this specimen was in bad conditions, the leg chaetotaxy and lengths were observed on one specimen of our collection (Montpellier SupAgro, UMR CBGP), collected in Valleraugues (Gard, South of France) on small leaved linden (Tilia cordata Miller).
Material examined -The female lectotype deposited in the ASU, Institute of Zoology, Academy of Sciences, Kiev, Ukraine.
Adult male (Figure 6e, f) Dorsal shield chaetotaxy similar to the female, but sub-lateral seta R1 on the dorsal shield. Ventrianal shield 99 long and 139 wide, distinctly reticulated and bearing five pairs of preanal setae and four pairs of poroids. Spermatodactyl L-shaped with a terminal foot and a toe enlarged. This description is based on one of the paralectotype specimens (deposited in the ASU, Ukraine).
(iii). Neoseiulella tuberculata (Wainstein), N. sexapori (Karg and Edland) and N. arutunjani (Kuznetsov) Karg and Edland (1987) differentiated N. sexapori and N. tuberculata by the following characters: the shape of the ventrianal shield, the length of the peritreme, the surface of the dorsal shield and the number and the shape of dorsal solenostomes. Chant and Yoshida-Shaul (1989) synonymised N. sexapori and N. tuberculata. In 1996, Denmark and Rather did not mention N. sexapori in their revision. Kuznetsov (1984) then described N. arutunjani, similar to N. tuberculata but lacking the caudoventral seta JV4. However, N. arutunjani was not included in revisions of the genus Neoseiulella carried out by Chant and Yoshida-Shaul (1989) and Denmark and Rather (1996). Kolodochka (2009) discussed the presence of JV4 on N. arutunjani, and then synonymised this species with N. tuberculata.

Adult male
Unknown from the type materials. However, it has been reported on maple (Acer pseudoplatanus) in Kremenets region, Ternopol district (Ukraine), collected and described, for the first time, by Kolodochka (2009).
Material examined -The female holotype and three female paratypes deposited in the Museum für Naturkunde, Arthropod collection, Berlin, Germany.

Adult male
The male of this species is unknown.
Previous reports -N. sexapori is only known from Norway on Rubus fruticosus L. (Rosaceae).  Ventral shields (Figure 9b) -Sternal shield 63 long and 51 wide (at level of ST2), smooth with two pairs of setae (ST1 and ST2) and two pairs of poroids. The third and fourth pairs of sternal setae (ST3, ST4) on separate platelets. A pair of small poroids accompanying ST4. Genital shield 105 long and 49 wide (at level of ST5), smooth. Four elongate platelets or genital sigilla between genital and ventrianal shields. One pair of poroids close to the genital shield (ST 5) and 3 pairs of poroids around the genital shield. Ventrianal shield subquadrateshape, 110 long and 78 wide (at level of ZV2), distinctly reticulated throughout, with four pairs of preanal setae (JV1, JV2, JV3 and ZV2), with a pair of circular solenostomes gv3 posteromediad to JV3. Four pairs of caudoventral setae (ZV1, ZV3, JV4 and JV5) on the integument surrounding the ventrianal shield. NB: one seta of the JV4 pair is present (reported absent on the original description) supporting Kolodochka (2009). JV5 13 long, smooth. Primary metapodal plate or inguinal sigillum 35 long and 4 wide.
Material examined -The female holotype deposited in the collection of Nikita State Botanic Garden, Yalta, Crimea, Ukraine. This material was examined in the ASU, Institute of Zoology, Academy of Sciences, Kiev, Ukraine.

Adult male
The male of this species is unknown.
Previous reports -N. arutunjani is only known from Ukraine, on Artemisia sp. (Asteraceae).
Remarks -It is noteworthy that in the original description, two females mounted on one slide were mentioned. Kolodochka (2009) showed that only one female actually belonged to this species. The other female was Paraseiulus incognitus Wainstein and Arutunjan. Our examination of these two females supports Kolodochka's observations. The present examination of the type materials of these three latter species could emphasize the following conclusions: Neoseiulella tuberculata does not differ from N. sexapori in setal and body measurements. Even if the shape of the spermatheca of these two species is slightly different (cup-shaped cervix in N. sexapori and a disc-shaped [shallowly cup-shaped] cervix in N. tuberculata), these two types of shapes are very close and such differences could be simply due to specimen mounting artefacts. The pair of solenostomes gd5 is observed on the four examined specimens of N. sexapori, and only on one of the three type specimens of N. tuberculata. At last, we observe other differences not mentioned by Chant and Yoshida-Shaul (1989): all dorsal setae are arising from tubercles in N. sexapori but not in N. tuberculata; the peritreme is shorter (at level between z2-j3) for N. sexapori than for N. tuberculata (at level of j1 or between j1-j3); the number of teeth on the fixed digit is different (three for N. sexapori, five for N. tuberculata); the ventrianal shield is heavily reticulated in N. sexapori, and lightly in N. tuberculata. As some 283 Kanouh M. et al. FIGURE 9: Neoseiulella arutunjani (Kuznetsov). Female (holotype): a -dorsal shield; b -ventral shields; c -spermatheca; d -chelicera.
authors have already shown the importance of such morphological characters for species differentiation (Chant and McMurtry 1994;Tixier et al. 2006a, b;Chant and McMurtry 2007;Okassa et al. 2009), we conclude that N. tuberculata and N. sexapori seem to be separate specific entities. These results agree with the conclusions of Karg and Edland (1987), but not with Chant and Yoshida-Shaul (1989).
Small differences in setal and body measurements are observed between N. tuberculata and N. arutunjani. According to intraspecific variations already observed within the family Phytoseiidae for continued characters, these differences seem to be too tiny for separating N. tuberculata from N. arutunjani. However, other differences are observed: the peritreme is shorter in N. arutunjani (reaching between z2-j3) than in N. tuberculata (reaching j1 or between j1-j3); the ventrianal shield is heavily reticulated on N. arutunjani, and lightly on N. tuberculata; fixed digit of chelicera bear seven teeth on N. arutunjani, and five on N. tuberculata; sub-lateral seta r3 is inserted on the dorsal shield on N. arutunjani and on lateral margin on N. tuberculata. However, as the position of sub-lateral seta (R1) was de-montrasted to be a non reliable diagnostic character to differentiate between N. aceri and N. squamiger (Kanouh et al. 2010), the position of r3 could be also assumed to be a non reliable diagnostic character as well. Though, the other morphological differences between N. tuberculata and N. arutunjani seem sufficiently discriminant to conclude that these two species are not synonyms. These results do not agree with the conclusion of Kolodochka (2009).
Small differences in setal and body measurements are observed between N. arutunjani and N. sexapori. However, all dorsal setae are arising from tubercles on N. sexapori, but not on N. arutunjani; the number of teeth on the fixed digit is different (three on N. sexapori, seven on N. arutunjani); spermatheca has a cup-shaped cervix in N. sexapori, but a discshaped (shallowly cup-shaped) cervix in N. arutunjani. These morphological differences between N. sexapori and N. arutunjani seem also sufficiently discriminant to conclude that these two species are not synonyms.
It seems thus that N. tuberculata, N. arutunjani and N. sexapori present sufficiently different characters to support their specific validity. These conclusions did not agree with Chant and Yoshida-Shaul (1989) and Kolodochka (2009).
(iv). Neoseiulella vollsella (Chaudhri, Akbar and Rasool), N. transitans (Gupta) and N. prunus (Denmark and Rather) Denmark and Rather (1984) differentiated N. prunus from N. vollsella by the length of both dorsal setae j3 and S4. Then, Chant and Yoshida-Shaul (1989) considered N. prunus a junior synonymy of N. transitans, and N. vollsella a provisional senior synonym to these two latter species. Denmark and Rather (1996) supported this conclusion. ( Material examined -The female holotype and two female paratypes deposited in the collection of the Zoological Survey of India (ZSI), Kolkata, India.

Adult male
The male of this species is unknown.  Ventral shields (Figure 11b) -Sternal shield 73 long and 68 wide (at level of ST2), smooth with three pairs of setae (ST1, ST2 and ST3) and two pairs of poroids. A pair of metasternal setae (ST4) on separate platelets accompanied by a pair of small poroids. Genital shield 100 long and 64 wide (at level of ST5), smooth. Elongate platelets or genital sigilla separating genital and ventrianal shields folded under the genital shield. One pair of poroids close to the genital shield (ST 5) and 2 pairs of poroids around the genital shield. Ventrianal shield much longer than wide, with a distinct waist, 100 long and 67 wide (at level of ZV2), smooth, with four pairs of preanal setae (JV1, JV2, JV3 and ZV2), and a pair of small circular solenostomes gv3 posteromediad to JV2. Four pairs of caudoventral setae (ZV1, ZV3, JV4 and JV5) on the integument surrounding the ventrianal shield. JV5 40 long, smooth. Primary metapodal plate or inguinal sigillum 22 long and 3 wide. No difference in all the morphological characters considered was found between N. transitans and N. prunus. Consequently, we consider that N. prunus is a junior synonym of N. transitans. Denmark (personal comm. 2010) confirmed this conclusion. As we were not able to borrow the type materials of N. vollsella, an attentive examination of the original description of this latter species was conducted. No morphological difference was observed between N. vollsella, N. transitans and N. prunus. Moreover, the geographical distributions of these three species are very close. Thus, we support the suggestion of Chant and Yoshida-Shaul (1989) concerning the synonymy of these three species.
Previous reports -N. manukae is only known from the Australasian area. The type specimens were collected at Waitakeres, near Auckland, New Zealand, on Leptospermum scoparium J.R. Forst. and G. Forst. (Myrtaceae) (Collyer 1964). This species has been also found on L. ericoides A. Rich. at Awanui Inlet, New Zealand (Collyer 1982); and on Eucalyptus parivolia Cambage (Myrtaceae) in New South Wales, Australia (Schicha 1987). (

Adult male
Described by Schicha (1980). We were not able to borrow the male type specimen of this species. The present examination of the type materials of N. manukae and N. glenfieldensis shows two main differences. The first one concerns the dentition of the fixed digit (11 teeth in N. manukae; 12-13 teeth in N. glenfieldensis). However, Schicha (1980) mentioned that N. glenfieldensis has 11-13 teeth on the fixed digit; we do not thus consider this difference to be significant. The second difference relates to the length of the seta Z5 (Z5 134 on N. manukae; Z5 171 on N. glenfieldensis). Even if this difference is quite high, high intraspecific vatiations have been previously observed for idiosomal seta lengths in the family Phytoseiidae (Tixier et al. 2003(Tixier et al. , 2008. Furthermore, it would be the unique diagnostic characters to separate these two species. We thus propose that N. manukae is a senior synonym of N. glenfieldensis, in accordance with the conclusions of Denmark and Rather (1996), but not with those of Schicha (1980) and Chant and Yoshida-Shaul (1989). However, an examination of other female paratype specimens of both species and/or molecular studies would be useful to determine the reliability of the length of Z5 for species diagnosis.

Are there new cases of synonymies within the genus Neoseiulella?
The examination of the type materials allows to highlight similarities between two species, and thus we propose a new synonymy within this genus: Neoseiulella nesbitti (Womersley) and N. armidalensis (Schicha and Elshafie). These two species were considered in the previous revisions as two differ-ent entities (Chant and Yoshida-Shaul 1989;Denmark and Rather 1996). Schicha and Elshafie (1980) when describing N. armidalensis have noted the similarity with N. nesbitti but stated that these species differ in the presence/absence of JV3 (present on N. armidalensis, absent on N. nesbitti). However, Chant and Yoshida-Shaul (1989) showed the absence of JV3 on the holotype specimens of these two species, but distinguished them by the presence of macrosetae on leg IV (one macroseta on N. nesbitti; three macrosetae on N. armidalensis), and the occurrence of pilus dentilis on the fixed digit (present on N. nesbitti; absent on N. armidalensis).

Adult male
Described by Schicha (1978). We were not able to borrow the male type specimens of this species.
Material examined -One female paratype specimen (deposited in NSW Department of Primary Industries, Agricultural Scientific Collections Unit (Acarology), Orange Agricultural Institute, Australia.

Adult male
The male of this species is unknown.
Previous reports -N. armidalensis is only known from New South Wales (Australia), on Malus sp. (Rosaceae) and Eucalyptus sp. (Myrtaceae).
The present examination of the type materials of N. nesbitti and N. armidalensis shows that JV3 is absent; the fixed digit has a pilus dentilis, and that leg IV bears three macrosetae for the two species. Moreover, no difference in setal and body measurements between these two species is observed. We thus propose that N. nesbitti is a senior synonym of N. armidalensis. This conclusion does not agree with those of Chant and Yoshida-Shaul (1989) and Denmark and Rather (1996). The unique difference we observe between N. nesbitti and N. armidalensis is the nature of macrosetae on leg IV (pointed on N. nesbitti, knobbed on N. armidalensis). Further experiments would be consequently interesting to carry out in order to determine the reliabilty of such a difference for species diagnosis.

Adult male
Described by Moraza et al. (2005). We were not able to borrow the male type specimen of this species.
Previous reports -N. arinoi is only known from the Canary Islands. It was collected from lichens on dead log, from soil of "lapilli" and litter of Ficus carica (Moraceae), Castanea sativa (Fagaceae) and Echium virescens D. C. (Boraginaceae) (Moraza et al. 2005).

Neoseiulella ashleyae (Chant and Yoshida-Shaul) (Figure 17)
Typhlodromus oleariae sensu Schicha (1987, 187 Adult female (Figure 17a -e) Dorsal shield (Figure 17a) -Dorsal shield smooth: length 464 (458 -469); width 302 (299 -304) (at level of s4) and 285 (280 -290) (at level of Z1). Six pairs of small solenostomes: gd2, gd4, gd5, gd6, gd8 and gd9. Four pairs of poroids. In the original description of this species, Chant and Yoshida-Shaul (1989) noted the presence of only five pairs of solenostomes (gd2, gd4, gd5, gd6, and gd9). However, on their drawings a large pair of solenostomes corresponding to the position of gd8 (according to Athias-Henriot [1975] and Swirski et al. [1998]) is present. We observed this pair of solenostomes on the four type specimens examined, we thus considered that gd8 is present and that N. ashleyae bears six solenostomes on the dorsal shield. Material examined -The female holotype and three female paratypes deposited in the Canadian National Collection, Department of Zoology, University of Toronto, Toronto, Canada. A specimen of this species labeled as Typhlodromus oleariae (Schicha 1987) (deposited in the New Zealand Arthropod Collection [NZAC], Landcare Research, Auckland, New Zealand) was also presently examined. Our examination of this specimen shows that, as stated by Chant and Yoshida-Shaul (1989), it belongs to N. ashleyae, but not to N. oleariae.

Adult male
The male of this species is unknown.
Previous reports -N. ashleyae is only known from New Zealand (Nelson Province) (Chant and Yoshida-Shaul 1989). Plants on which this species was collected are: Elaeocarpus hookerianus Raoul

Adult male
The male of this species is unknown.
This species is easily distinghuished from the other species in the genus Neoseiulella by the tuberculous ornements on the dorsal shield, and by the small size and the shape of the ventrianal shield, which is narrowly vase-shaped.
Material examined -The female holotype and five female paratypes deposited in the UQIC, Department of Entomology, University of Queensland, St Lucia, Australia.

Adult male
Described by Walter (1997). We were not able to borrow any male type specimens of this species.
Previous reports -N. coreen is only known from Queensland (Australia) on rainforest trees. smooth. Primary metapodal plate or inguinal sigillum 29 long and 8 wide.
Material examined -The female holotype deposited in NSW Department of Primary Industries, Agricultural Scientific Collections Unit (Acarology), Orange Agricultural Institute, Australia.

Adult male
The male of this species is unknown. Previous reports -N. corrugata is only known from Australia on Malus domestica (Rosaceae).
Ventral shields (Figure 23b) -Sternal shield 77 long and 70 wide (at level of ST2), smooth with three pairs of setae (ST1, ST2 and ST3) and two pairs of poroids. A pair of metasternal setae (ST4) on separate platelets accompanied by a pair of small poroids. Genital shield 120 long and 68 wide (at level of ST5), smooth. Four elongate platelets or genital sigilla situated between genital and ventrianal shields. One pair of poroids close to the genital shield (ST 5) and 2 pairs of poroids around the genital shield. Ventrianal shield, longer than wide 124 long and 101 wide (at level of ZV2), smooth, with three pairs of preanal setae (JV1, JV2 and ZV2; JV3 absent), with a pair of circular solenostomes Material examined -The female lectotype deposited in the BMNH, the British Museum of Natural History, Cromwell Road, London, UK.

Adult male
Described by Collyer (1964) and Schicha (1980). We were not able to borrow the male type specimens of this species.
Previous reports -The type specimens of N. cottieri were collected from New Zealand on thistle (Asteraceae). This species is only known from the Australasian area: Australia (Wainstein 1977;Schicha 1980Schicha , 1987 and New Zealand (Collyer 1964(Collyer , 1982. Plants on which this species was collected are: Astelia sp. Previous reports -N. crassipilis is only known from the West-Palaearctic area. Countries from which this species was reported are: France (Athias-Henriot and Fauvel 1981;Viollier and Fauvel 1984) and Greece (Ragusa and Tsolakis 1998;Ragusa 2006). It was collected on Pyrus amygdaliformis Vill. and Pyrus sp. (Rosaceae).

Adult male
Described by Collyer (1964) and Schicha (1980). We were not able to borrow the male type specimen of this species.

Adult male
The male of this species is unknown.
Material examined -The female holotype deposited in Museum of Zoology of University of Navarra (MZUNAV), Spain.
Adult male (Figure 28f, g) Male specimen of this species similar to but smaller than the female. Moraza and Peña-Estevez (2006) described the male of N. ferraguti. However, these authors did not give a complete setal and body measurements. We thus present a complete description of this male.
The male specimen of this species is different from the female by the position of both sub-lateral setae (r3 and R1) (on dorsal shield on the male; on lateral margin on the female), and the number of solenostomes on the ventrianal shield (two pairs on the male; one pair on the female).
Material examined -A male paratype specimen.
Previous reports -N. ferraguti is only known from Tenerife (Canary Islands). It was reported from lichens on dead log, and from soil and litter of Cheirolophus canariensis var. subexpinnatus (Burch.) A. Hansen and Sunding (Asteraceae). Adult male (Figure 29f, g) Dorsal shield chaetotaxy similar to but smaller than the female. Sub-lateral seta R1 inserted on the dorsal shield (different from the female, on which R1 on lateral margin). Six pairs of solenostomes on the dorsal shield. Ventrianal shield 124 long and 142 wide, distinctly reticulated, bearing six pairs of preanal setae and a pair of poroids. Spermatodactyl L-shaped. This description is based on a paratype specimen.

Neoseiulella litoralis (Swirski and
Previous reports -N. litoralis is only known from the West-Palaearctic area. Countries from which this species was reported are: Israel (Swirski and Amitai 1984;Klein et al. 1994

Adult male
Described by Moraza et al. (2005). We were not able to borrow the male type specimen of this species.
Previous reports -N. longiseta is only known from the Canary Islands. It was collected from dry soil, moss and litter under the following plants:
Chelicera (Figure 31c) -Two or three teeth and a pilus dentilis on the fixed digit. Movable digit 39 (38 -40) long, bears one or two teeth.
Material examined -Two female paratypes deposited in the Division of Entomology, Agricultural Research Organization, Bet Dagan, Israel.

Adult male
The male of this species is unknown.
Material examined -Two female paratypes deposited in the New Zealand Arthropod Collection (NZAC), Landcare Research, Auckland, New Zealand.

Adult male
The male of this species is unknown.
Previous reports -N. myopori is only known from New Zealand. The female holotype and six female paratypes were collected on Myoporum laetum G. Forst. (Myoporaceae) at Clifton Coast, South of Napier. One additional report (1 female and a deutonymph), excluded from the type series, was from Dun Mountain (Nelson Province), on Holocarpus bidwillii (Hook. f.ex T.Kirk.) Quinn (Podocarpaceae) (Collyer 1982).

Adult male
The male of this species is unknown.
Remarks -N. neoviniferae is similar to N. montforti but differs by the presence/ absence of gd5 (present on N. neoviniferae; absent on N. montforti), the shape of the spermatheca (c-shaped on N. neoviniferae; U-shaped on N. montforti) and measurements of some dorsal setae (j6, J2, z2, Z1 and R1).
Material examined -One female paralectotype deposited in the New Zealand Arthropod Collection (NZAC), Landcare Research, Auckland, New Zealand.

Adult male
Described by Collyer (1964) and Schicha (1980) and Denmark and Rather (1996). We were not able to borrow male type specimens of this species.
(Fagaceae); Eucalyptus sp., Leptospermum scoparium Remarks -N. novaezealandiae is very similar to N. nesbitti in setal and body measurements. However, differences are observed for the ornementation of dorsal shield (lightly reticulated on N. nesbitti, heavily reticulated on N. novaezealandiae) and the position and size of prenanal solenostomes (small and posterior to JV2 in N. nesbitti, large and mediad to JV2 in N. novaezealandiae). As several authors have showed the importance of such characters for species identification (Shicha 1980;Chant and Mc-Murtry 1994;Swirski et al. 1998;Chant and Mc-Murtry 2007;Okassa et al. 2009), we consider that these two species are separate specific entities.
Material examined -Two female paratypes deposited in the New Zealand Arthropod Collection (NZAC), Landcare Research, Auckland, New Zealand.

Adult male
Described by Collyer (1982) and Denmark and Rather (1996). We were not able to borrow male type specimens of this species.
Previous reports -N. oleariae is only known from New Zealand. The female holotype and two paratype specimens (1 female and 1 male) of N. oleariae were collected on Olearia colensoi Moench. (Asteraceae) at Ruahine Range. Other paratype specimens (2 females and 1 male) were also collected on O. colensoi at Magister Ridge (Westland), Strachan Ridge (South Westland), and Pillans Pass (Manapouri). Other specimens (excluded from the type series) were collected from both North and South islands of New Zealand. Plants on which this species was collected are: Carmichaelia sp. (Fagaceae); Carpodetus serratus J.R. and G.Forst. (Car-  Ventral shields (Figure 37b) -Sternal shield 51 long and 64 wide (at level of ST2), smooth, with two pairs of setae (ST1 and ST2) and two pairs of poroids. ST3 and ST4 are inserted on separates Chelicera (Figure 37d) -Four teeth and a pilus dentilis on the fixed digit. Movable digit 23 long, bidentate.
Material examined -The female holotype deposited in the ASU, Institute of Zoology, Academy of Sciences, Kiev, Ukraine.
Adult male (Figure 37e-f) Dorsal shield similar to but smaller than female. Ventrianal shield 120 long and 142 wide, distinctly reticulated and bearing 5 pairs of preanal setae and three pair of poroids. Spermatodactyl L-shaped. This description is based on a paratype specimen.

Adult male
Described by Collyer (1982) and Denmark and Rather (1996). We were not able to borrow the male type specimen of this species.
Previous reports -N. spaini is only known from New Zealand (Collyer 1982;Schicha 1987). The female holotype, paratypes and other specimens of this species are reported on Olearia colensoi (Asteraceae) at Ruahine Range and Urewera National Park. Adult male (Figure 39f, g) Dorsal shield -dorsal shield similar to but smaller than the female. Sub-lateral seta R1 inserted on the dorsal shield (different from the female, on which R1 inserted on lateral margin). Ventrianal shield 140 long and 135 wide, distinctly reticulated, bearing seven pairs of preanal setae with a pair of poroids (Figure 39 f). Spermatodactyl Lshaped (Figure 39 g). This description is based on a paratype specimen. Remarks -N. splendida is similar to N. litoralis. However these two species differ by the following characters: gd4 presence/ absence (present on N. splendida; absent on N. litoralis); j3, z3, Z4, Z5, s4, and s6 lengths; the length of the peritreme (extending between j1-j3 on N. splendida; extending between z2-j3 on N. litoralis). Moreover, the males of these two species differ in the number of setae on the ventrianal shield (seven pairs on N. splendida, six on N. litoralis). These two species seem thus to be separate entities. However, further experiments would be useful in order to conclude on the reliability of these morphological characters in species identification.
Ventral shields (Figure 40b) -Sternal shield 82 long and 85 wide (at level of ST2), smooth, with three pairs of setae (ST1, ST2 and ST3) and two pairs of poroids. A pair of metasternal setae (ST4) on separate platelets accompanied by a pair of small poroids. Genital shield 139 long and 86 wide (at level of ST5), smooth. Four elongate platelets or genital sigilla separating genital and ventrianal shields folded under the genital shield. One pair of poroids close to the genital shield (ST 5) and 1 pair of poroids around the genital shield. Ventrianal shield subtriangular-shaped, 144 long and 113 wide (at level of ZV2), smooth, with four pairs of preanal setae (JV1, JV2, JV3 and ZV2) and a pair of small and circular solenostomes gv3 posteromediad to JV3. Four pairs of caudoventral setae (ZV1, ZV3, JV4 and JV5) on the integument surrounding the ventrianal shield. JV5 28 long, smooth. Primary metapodal plate or inguinal sigillum 30 long and 7 wide. Spermatheca ( Figure 40c) -Cervix 9 long, cupshaped.
Chelicera -Dentition of the chelicera not discernible on the specimen examined. Schicha and McMurtry (1986) mentioned the presence of ten teeth and a pilus dentilis on the fixed digit of the chelicera, and of three teeth on the movable digit (32 long).
Material examined -The female holotype deposited in NSW Department of Primary Industries, Agricultural Scientific Collections Unit (Acarology), Orange Agricultural Institute, Australia.

CONCLUSION
The present paper provides a homogenous taxonomic revision of nearly the totality of the species belonging to the genus Neoseiulella with an identification key of the adult females of valid species. This revision allows to redefine this genus, excluding three species, including an additional species T. ( A.) elisae and discussing six synonymies. Further experiments, especially cross-breedings and molecular analyses are required to determine the relia-bility of some morphological characters considered to be important by some authors to discriminate between species (ex. ornamentation of dorsal and ventral shields; presence of certain dorsal and ventral solenostomes; length of the peritreme; chelicera dentition; the position and size of preanal solenostomes). This paper also provides complete biogeographic data sets (distribution and host plants) of all the species of the genus Neoseiulella. Eighteen species are reported from the West-Palearctic area; two of them (N. aceri and N. tiliarum) being also present in the Nearctic area; 15 species are reported from the Australasian area; one of them (N. nesbitti) being also present in the Oriental area. Three other species are reported from this latter area. These data associated to the ecological distribution (plant supports) could constitute the starting point for further analyses, especially biogeographic and phylogenetic analyses. The disjoint distribution of the species of the genus Neoseiulella questions actually the monophyly of this genus.