New Phytoseiidae (Acari: Mesostigmata) of Mascareignes and Comoros Archipelagos (Indian Ocean): one new record, three new species groups and description of six new species and of six unknown males

Faunas of Phytoseiidae of the Mascareignes Archipalago (Réunion, Mauritius and Rodrigues Islands) and of the Comoros Archipelago (Mayotte, Anjouan, Mohéli and Grande Comore Isands) were recently investigated by authors of this paper and results were published in seven already published papers. We described in this eighth paper six species new to science and six unknown males collected during these surveys.


Introduction
Mites of family Phytoseiidae are all predatory species on phytophagous mites and small insects like thrips and whiteflies, on commercial plants and the wild vegetation, many of these arthropods being important pests for agriculture. Several species are biological control agents for the control of these pest organisms in both open and protected crops all around the world (McMurtry and Croft 1997; McMurtry et al. 2013; Knapp et al. 2018. This family is widespread around the world, present on all continents except Antarctica, and consists of about 2,521 valid species in 95 genera, 15 tribes and three subfamilies (Demite et al. 2021).
Despite several interests of this family and its large distribution, many areas of the world are very poorly investigated or not investigated, some areas remaining white spots concerning the fauna of Phytoseiidae.
Thus, biodiversity surveys in these poorly investigated areas are still an urgent need and might result in the discovery of additional species potentially useful for biological control as well as having more information on the biodiversity of these areas for biodiversity practical purposes.
In these perspectives, the more interesting areas are probably those with a high level of biodiversity. Most of the Indian Ocean constitutes one of the highest world biodiversity areas, those areas called hotspots, concept defined by Myers (1988) in order to identify the most immediately important areas for biodiversity conservation. The common characteristics of these hotspots are that they hold high endemism levels and have lost at least 70% of their original natural vegetation (Myers et al. 2000). Knowledge of the phytoseiid diversity in these high interest areas in the context of global climate changes may contribute to identify potential biological control agents (BCAs) and future establishment of conservation programs.
Several Islands are located in the Indian Ocean, especially in two archipelagos, Mascareignes and Comoros. The former is constituted of several small Islands and three main Islands: La Réunion, Mauritius and Rodrigues. The later is constituted of some small Islands and four main Islands: Mayotte, Anjouan, Mohéli and Grande Comore.
Although these Islands, especially Mascareignes Islands, are a top destination for tourism and attracted the interest of many European naturalists, the fauna of phytoseiid mites remains poorly known (Ferragut and Baumann 2019).
These main Islands of the two Archipelagos (except La Réunion which was investigated before, see  were investigated from October 25 th to December 12 th , 2018. Results of Phytoseiidae records were already published in six papers; Kreiter and Abo-Shnaf 2020a, b for Rodrigues and Mauritius (in addition to Mauritius, see Kreiter et al. 2018a; Kreiter et al. 2020a, 2021b for Mayotte, Anjouan, Mohéli, and Grande Comore (in addition to Grande Comore, see Kreiter et al. 2018b), respectively. This paper aims to give the description of six species new to science and six unknown males along with one new record collected during this survey.
Mites were directly collected on leaves with a fine brush with or without a pocket lens or a stereo-microscope when available (large leaves and herbaceous plants) or by beating the plants (mainly shrubs and trees with very small or spiny leaves) and collecting the mites in a black plastic rectangular saucer 45 x 30 cm (Ref. STR 45, BHR, 71370 Saint-Germain-du-Plain, France). Collected mites were then transferred with a fine brush into small plastic vials containing 1.5 ml of 70% ethanol.
The mites were then all slide-mounted in Hoyer's medium (Walter and Krantz 2009), the slides were dried at 45-50 o C for at least two weeks and then all examined and identified using a phase and interferential contrast microscope (DMLB, Leica Microsystèmes SAS, Nanterre, France). Characters of specimens were measured using a Leica graded eyepiece. Chant andMcMurtry's (1994, 2007) concepts of the taxonomy of the family Phytoseiidae for identification and the world catalogue database of Demite et al. (2014Demite et al. ( , 2021 for distribution and information on descriptions and re-descriptions were used. The setal nomenclature system adopted was that of Lindquist and Evans (1965) and Lindquist (1994) as adapted by Rowell et al. (1978) and Chant and Yoshida-Shaul (1989) for the dorsal surface and by Chant and Yoshida-Shaul (1991) for the ventral surface. Pore (= solenostome) and poroid (= lyrifissure) notations are that of Athias-Henriot (1975). Macrosetal notation (Sge = genual macroseta; Sti = tibial macroseta; St = tarsal macroseta) are that of Muma and Denmark (1970). Numbers of teeth on the fixed and movable cheliceral digits do not include the respective apical teeth. Setae not referred to in results section should be considered as absent. All measurements are given in micrometres (µm) and presented with the mean in bold followed by the range in parenthesis. Type of spermatheca or insemination apparatus is that of Denmark and Evans (2011).
Classification of plants follows the APG IV classification of 2016 (ex. Byng et al. 2018). Specimens of each species are deposited in the mite collections of Institut Agro (Montpellier SupAgro) conserved in UMR CBGP INRAE/IRD/CIRAD/SupAgro/University of Montpellier.
The following abbreviations are used in Tables (1-3) for morphological characters: n = number of individuals measured; dsl = dorsal shield length just above j1 to just below J5 in the mid line; dsw = dorsal shield width at the level of s4; Peritreme = level of the peritreme extension; gd = number of solenostomes; gensl = genital shield length; gensw st5 = genital shield width at level of setae st5; gensw post. cor. = genital shield width at level of posterior corners; lisl = primary or largest inguinal sigilla (= "primary metapodal plate") length; lisw = primary or largest inguinal sigilla (= "primary metapodal plate") width; sisl = secondary or smallest inguinal sigilla (= "secondary metapodal plate") length; vsl = ventrianal shield length; gv3-gv3 = distance between centres of each solenostome gv3 on the ventrianal shield; vsw ZV2 & vsw anus = ventrianal shield width at ZV2 level and at para-anal setae level; scl: largest calyx length; scw = calyx widest width; Fdl = fixed digit length; Mdl = movable digit length; Nb teeth Fd = number of teeth on the fixed digit; Nb teeth Md = number of teeth on the movable digit; Shaft = length of the shaft of spermatodactyl; branch = length of the branch; BCA = Biological Control Agent; aasl = altitude above sea level; imm. = immature.

Results and Discussion
During the survey in Indian Ocean Islands, we found six new species to science and other six unknown males and one new record (in the chronological order of the survey: Mauritius, Rodrigues, Mayotte, Anjouan, Mohéli, and Grande Comore): • One unknown male of Amblyseius haleakalus Prasad  • the subfamily Amblyseiinae Muma (absence of dorsolateral setae z3 and s6 and the caudoventral seta JV3), • to the tribe Neoseiulini Chant & McMurtry (seta S4 present, ratio s4/Z1 < 3.0, setae s4, Z4 and Z5 not greatly longer than other dorsal setae, usually slightly sclerotized, never with wide sternal shield, seta J2 always present), • to the genus Paragigagnathus Amitai & Grinberg (female ventrianal shield reduced and/or markedly wider at the level of anus, with a prominent waist, chelicerae with teeth only on apical region, fixed digit with one to three teeth, movable digit with a single tooth, primary metapodal plate or unguinal sigillum elongate (Chant and McMurtry 2007). There are 12 species within this genus, • Seta st3 is inserted off sternal shield of female on separate platelets (see below), which allows to classify this new species in the species group strunkhovae (Chant and McMurtry 2003). This species group contains four species (Chant and McMurtry 2003). The following list of characters of this new species is very different of all species of the genus and the species group. Despite the fact that we collected a single specimen, we consider this very original specimen as belonging to a new original species to science and we describe it thereafter.
Peritreme and peritremal plate (Fig. 1a) -Extending to level of j1; peritremal plate fused with dorsal shield at level between j1 and j3.
Venter (Fig. 1b) -All ventral shields smooth. Sternal shield with two pairs of setae (st1 and st2) and a pair of poroids (iv1); two pairs of setae (st3 and st4) on two separate metasternal plates (no discernible pores on both of them); posterior margin of the sternal shield apparently straight; a pair of poroids (iv2) off sternal shield. Distances st1-st1 40, st2-st2 46, st3-st3 64, st1-st3 53, st4-st4 82. Genital shield length 108, width at level of st5 58, width at level of posterior corners 58, distance st5-st5 54. Two pairs of metapodal plates, primary metapodal plate moderately long compared to some other species (Table 1), 29 long and 3 wide and secondary short, 6 long and 2 wide. Ventrianal shield 93 long, 55 wide at level of anterior corners (ZV2), and 56 wide at level of para-anal setae. Ventrianal shield with three pairs of pre-anal setae (JV1, JV2 and ZV2), and a pair of small crateriform gv3, 13 apart. Unsclerotized cuticle arround ventrianal shield with four pairs of setae (JV4, JV5, ZV1 and ZV3), and apparently five pairs of round to oblong poroids difficult to see on our preparation, except for ivp on posterior part of the ventrianal shield. Seta JV5 short, thick and probably smooth (impossible to confirm on the single specimen), 12 long.
Etymology. The name "philippei" refers to the first name of the senior author's second brother, Philippe Luc Kreiter, Engineer-Researcher in INRAE and specialist of biological control of mealybugs. The species is named in his honour.
Differential diagnosis and remarks. This species is unique in the genus Paragigagnathus by a set of unique characters (Table 1) and especially the small size of the body, the setae all plumose, thick and serrate, the reduce size of metapodal plates, the reduce size of ventrianal shield and the occurrence of macrosetae on all legs, the sternal shield with only two setae, along with an assemblage of specific setae lengths. No other species are closed to the new species, especially within the strunkhovae species group to which this new species belongs. For this reason, this species is described despite the single specimen collected. New surveys on Mohéli Island must occur in order to recover the species and to increase the description. Paragigagnathus philippei Kreiter n. sp. is the 13 th species of the genus Paragigagnathus and the fifth species of the strunkhovae species group (the eight other species belonging to the desertorum species group).  Moraes et al. 1986: 139, 2004b: 210; Chant & McMurtry 2005c: 327, 2007 2, 6, 8 , 9 1, 2, 6, 8, 9 1, 2, 6, 8, 9 1, 2, 6, 8, 9 1, 2, 5, 8, 91, 2, 5, 6, 7, 8, 9 -1, 2, 6, 8 , 9 1, 2, 6, 8 , 9 1, 2, 6, 8 , 9 S     This species belongs to the culmulus species group of the genus Typhlodromips with nine other species. It is also probably a type III species Croft 1997; McMurtry et al. 2013), i.e., a polyphagous generalist predator. However, its biology remains totally unknown. It was already recorded on Mauritius Island, but only one record based on a single female and a single location (Kreiter et al. 2018a). It was also recorded in La Réunion Island, but with few specimens collected after intensive surveys ). This species seems rather rare.
Peritreme and peritremal plate (Fig. 2a) -Extending to level of j1; peritremal plate fused with dorsal shield at level of j3.
Peritreme and peritremal plate ( Fig. 3a) -Extending to level of j1; peritremal plate fused with dorsal shield at level of j1.
Legs (Fig. 3d) -One macroseta on legs I and II, two macrosetae on leg III and three macrosetae on legs IV similar to adult female. All macrosetae sharp-tipped. Measurements: Type material. One holotype ♀ on one slide, one paratype ♀ and one paratype ♂ on another slide are deposited in Institut Agro (Montpellier SupAgro) -INRAE Acarology collection, Montpellier, France.
Etymology. The name "erici" refers to the first name of the senior author's youngest and third brother, Eric Kreiter. The species is named in his honour.
Differential diagnosis and remarks. None of the females of species of Amblyseius (of the obtusus species group and of the andersoni species subgroup) included in Table 2 share similar characters with females of Amblyseius erici Kreiter n. sp. The two closest species concerning setae length are A. angulatus Karg and A. compositus Denmark & Muma, but several other details are different: macrosetae lengths and number of teeth of these two species compared to the new species. But descriptions of these two new species are old and very poor and lacking information for a complete description. The shape of the spermatheca of the new species is unique and allows distinguishing this new species from all others in Table 2 and all species of the andersoni species subgroup. The following combination of characters, of the male indicated in the description of the male of this new species, is quite similar to that of the few described males of species of Amblyseius belonging to the obtusus species group and to the andersoni species subgroup.
No teeth Fdl 9 9 9 9-11 -9 8 9 10 In each upper boxes of the first line, the first name is the name of the species (for example andersoni ), the second line of name(s) is name(s) of describer(s) (for examaple Chant for andersoni) and the third line is the source of measurements (for example Denmark & Muma 1989 for andersoni). Differential diagnosis and remarks. The male of this species was mentioned in El-Banhawy and Knapp (2011), but it is not indicated that this is the first mention of the male of this species, the male was illustrated, but the description lacks detail (El-Banhawy and Knapp 2011). We thus decide on a more detailed description of the male of this species.
This species belongs to the largoensis species group as setae J2 and Z1 are present, seta s4 is minute and the ventrianal shield of the female is vase-shaped. It belongs to the largoensis species subgroup as seta Z4 is long, spermatheca has the calyx elongate mostly tubular and the female ventrianal shield is entire (Chant and McMurtry 2004).
The following combination of characters, indicated in the description of the male of this species, is quite similar to the few described males of species of Amblyseius belonging to the largoensis species group and to the largoensis species subgroup.
Not many characters allow to distinguish it from all males of other species if no females are collected in the same time (all the males used for description were collected with females of this species): the peritreme reaching level of j1, absence of reticulation of the dorsal shield, all dorsal setae including J5 length approximately of the same length (8-11), except for j1, j3, s4, Z4, Z5 longer and z5 shorter, additional macrosetae on all other legs than leg IV, macrosetae of leg IV not sub-equal, a sternogenital shield mostly smooth, a ventrianal shield reticulate, only three pairs of pre-anal setae, a pair of crateriform gv3 between JV2.
All described males of the large species subgroup largoensis have very similar ventrianal shield reticulate with three pairs of pre-anal setae.
Peritreme and peritremal plate (Fig. 5a) -Extending to level of j1; peritremal plate fused with dorsal shield at level of z2.
Legs ( Differential diagnosis and remarks. This species belongs to the obtusus species group as seta z4 is minute and female ventral shield is not vase-shaped or divided. It belongs to the andersoni species subgroup as the spermatheca has a differentiated atrium, a calyx not dotted or annulated, not swollen basally and calyx dish-, cup-, bell-or V-shaped. The following combination of characters, indicated below in the description of the male of this species, is quite similar to the few described males of species of Amblyseius belonging to the obtusus species group and to the andersoni species subgroup. Not many characters allow to distinguish it from all males of other species if no females are collected in the same time: the peritreme reaching level of j1, absence of reticulation of the dorsal shield, some dorsal setae lengths, especially z2, z4, r3 and S2 approximately of the same length (12-15), additional macrosetae on all other legs than leg IV, macrosetae of leg IV not subequal, a sternogenital shield smooth, ventrianal shield reticulate, only three pairs of pre-anal setae, a pair of crateriform gv3 between JV2. All described males of the large species subgroup andersoni have similar ventrianal shield reticulate with three pairs of pre-anal setae.
Characters of males are very similar to that of adult females, except of course for length of setae and few other characters. The only difference is that ventrianal shield of the male is moderately reticulate, while the ventrianal shield of the female is not.
Peritreme and peritremal plate (Fig. 6a) -Extending to level of j1; peritremal plate fused with dorsal shield at level between z2 and z4.
Venter (Fig. 6b) -Sternogenital shield smooth with very few striae, five pairs of setae (st1-st5) and two pairs of poroids (iv1 and iv2). Differential diagnosis and remarks. This species has no seta Z1 and consequently belongs to the sundi species group and having the spermatheca elongate, tube-like, it belongs to the sundi species subgroup. The following combination of characters indicated below in the description of the male of this species is quite similar to the unique described males of species of Amblyseius belonging to the sundi species group and to the sundi species subgroup. Not many characters allow distinguishing it from the single described male of this sundi subgroup, the male of A. sundi Pritchard & Baker. If no females are collected in the same time, the identification will be impossible. These characters are: the peritreme reaching level of j1, absence of reticulation of the dorsal shield, some dorsal setae length, especially j-J serie starting to j4, z2 to z5, R1 and S series (after s4) approximately of the same length (4-8), additional macrosetae on all other legs than leg IV, macrosetae of leg IV not subequal and long, sternogenital shield smooth, ventrianal shield reticulate, only three pairs of pre-anal setae, a pair of round gv3 between JV2, a macroseta present also on genu lI, only three teeth on the movable digit and 11 on the fixed digit of chelicera instead of one and six in the male chelicera of A. sundi, respectively.
Characters of males are very similar to adult females, except of course for lengths of setae and other few characters. The only difference is that the ventrianal shield of the male is lightly reticulate in the anterior part and the ventrianal shield of the female is not. Blommers and Gutierrez (1975) found this species very abundant on fruit trees preying on several species of tetranychid mites. Amblyseius sundi is reported by Blommers (1974) as being a thelytokous species in mass-rearing and field collected specimens and similar information is also mentioned by Denmark and Muma (1989). In nature, reproduction of A. parasundi seems more complicated. Males were not so rare in fields of the two Islands where they were found (Mayotte and Mohéli). This suggests further fundamental studies on the biology of this species. Classification. Typhlodromalus baillodi Kreiter n. sp. belongs to:

Tribe Euseiini Chant & McMurtry
• the subfamily Amblyseiinae (absence of dorsolateral setae z3 and s6 and the caudoventral seta JV3), • to the tribe Euseiini (sternal shield with median posterior projection, deutosternal groove > 5 µm in width, forward migration of pre-anal setae JV2 and ZV2), • to the subtribe Typhlodromalina (chelicera of normal size and shape, with prominent teeth evenly distributed along fixed digit, peritreme usually extending to level of j1, deutosternal groove narrow, 4-7 µm width), • to the genus Typhlodromalus (female ventrianal shield with more than one pair of preanal setae, GeI usually with a macroseta, GeII and III with macrosetae, leg IV with three macrosetae usually stout, often knobbed or blunt, male ventrianal shield with three pairs of pre-anal setae, most dorsal setae either setiform or thickened, thorn like, tapering distally, without terminal knobs, fixed digit with 6-12 teeth evenly distributed along the digit, BtI without erected seta, female ventrianal shield with three pairs of pre-anal setae, ratio s4/Z1 < 3.0 : 1.0, dorsal setae of medium length subequal, dorsal shield ornamented in addition to anterolateral striations, seta Z4 longer than distance between its base and that of S4, • to the peregrinus species group as seta S5 is present (Chant and McMurtry 2007)  Description of adult female (n = 15 of 44 collected during this study, Figs. 7 a-e) Dorsum (Fig. 7a) -Dorsal shield strongly ornamented and reticulate, with margins of posterior part slightly indented at level of S5 creating a slight "trilobite appearance", with an expansion on each lateral side at level of s4-Z1 and with a constriction at level of R1, 310 (283-333) long and 187 (165-210) wide at level of s4, with seven solenostomes (gd1, gd2, gd4,  gd5, gd6, gd8 and gd9) (16-23). All setae thickened and smooth, except for Z5 strongly serrate.
Peritreme and peritremal plate (Fig. 7a) -Extending to level of j1; peritremal plate fused with dorsal shield at level between j1 and j3, much closer to the former.
Venter (Fig. 7b) -All shields smooth. Sternal shield with three pairs of setae (st1-st3) and two pairs of rounded poroids (iv1 and iv2); a pair of st4 and a pair of rounded pores (iv3) on a metasternal plate; posterior margin of the sternal shield convex, with a posterior projection. Chelicerae (Fig. 7c) -Fixed digit 26 (25-28) long, with five teeth in row and one subapical tooth; and movable digit 27 (25-28) long, with two teeth. Pilus dentilis not visible. Spermatheca (Fig. 7d) -Resembles that of Ueckermannseius payetae Kreiter n. sp. in the new species group havu of the genus Ueckermannseius, with the atrium bulbous and elongate, the calyx basally swollen, bladder-like and then elongate and slender, 36 (30-45) long and 9 (8-11) wide at the widest of the calyx, small minor duct visible.
Material examined. Fourty-four ♀♀, nine ♂♂ and two imm. collected during this study, fifteen ♀♀ and nine ♂♂ measured, 43 ♀♀, nine ♂♂ and two imm. as type material. Etymology. The name "baillodi" refers to the family name of the researcher Dr Marc Baillod, who has worked during his career at the Station Fédérale de Recherche Agronomique de Changins in Switzerland (now called Agroscope) and has published many useful papers on plant inhabiting mites in agrosystems. He contributed towards the senior author's knowledge of the Phytoseiidae (taxonomy, biology, ecology, side effects of pesticides, etc.) more than 35 years ago. Marc Baillod was a real Master and deserves billions of billions of thanks! This new species is named in his honour.
Differential diagnosis and remarks. This species is very original by the set of characters described above. Lengths of most of the major setae are very similar to those obtained by Yoshida-Shaul and Chant (1991) for T. fragosoi Yoshida-Shaul & Chant and by Kreiter et al. (2002) for T. etiennei Kreiter & Ueckermann. However, the unique shape of the spermatheca not only distinguishes if from the latter two species, but also from all known species of the genus Typhlodromalus. The spermatodactyl also distinguishes it from that of T. spinosus Meyer and Rodrigues, allowing an easy distinction between the two species mentioned from this region.

Ueckermannseius gutierrezi Kreiter n. sp.
Zoobank: A56A4712-1C00-4F5B-84BE-2B9592180D24 Classification. Ueckermannseius gutierezzi Kreiter n. sp. belongs to: • the subfamily Amblyseiinae (absence of dorsolateral setae z3 and s6 and the caudoventral seta JV3), • to the tribe Euseiini (sternal shield with median posterior projection, deutosternal groove > 5 µm in width, forward migration of pre-anal setae JV2 and ZV2), • to the subtribe Typhlodromalina (chelicera of normal size and shape, with prominent teeth evenly distributed along fixed digit, peritreme usually extending to level of j1, deutosternal groove narrow, 4-7 µm width), • to the genus Ueckermannseius (dorsal setae short/minute, shorter than distances between their bases, seta Z4 not as long as distance between its base and that of S4, dorsal shield smooth, except for anterolateral striation) ( The two other new species groups proposed are: • the species group ultimus Kreiter, with spermatheca elongate, tubular, flared distally with an atrium prominent, but small. This kind of spermatheca is shared by six African species of Ueckermannseius we proposed to include in the ultimus species-group: U. aequidens Blommers, U. bunyalae El-Banhawy and Knapp, U. kiminini El-Banhawy and Knapp, U. munsteriensis (van der Merwe), U. tenuiscutus McMurtry and Moraes and U. ultimus (Chant and Baker), • the species group danhomensis Kreiter, with spermatheca with calyx short, funnelshaped, with an atrium distinctly bulbous. This kind of spermatheca is shared by only two species of Ueckermannseius we proposed to include in the danhomensis species-group: U. danhomensis Moraes, Zannou and Oliveira and U. musoli El-Banhawy and Knapp.
Peritreme and peritremal plate (Fig. 9a) -Extending to level between j3 and z2; peritremal plate fused with dorsal shield at a level between j3 and z2.
Peritreme and peritremal plate (Fig. 10a) -Extending to level between j1 and j3; peritremal plate fused with dorsal shield at level between j3 and z2.
Chelicerae -Fixed digit 20 long, no discernible teeth; and movable digit 20 long, with no discernible teeth. Spermatodactyl shaft renders measurement and illustration impossible.
Legs (Fig. 10c)  Etymology. The name "gutierrezi" refers to the family name of the researcher Dr Jean Gutierrez, who has worked during his career at ORSTOM (= IRD for now) and have published many papers on plant inhabiting mites, mainly tetranychid mites, from Indian Ocean among many other sites. He has helped the senior author in many aspects at the beginning of his career, especially with exciting and stimulating scientific discussions on mites and many other subjects. This species is named in his honour. Differential diagnosis and remarks. This species closely resembles U. neohavu concerning length of setae on dorsal shield. However, it differs from the latter in having: setae j4-j6, R1, s4, z2, z4 and Z5 shorter with Z5 serrate, ventrianal shield and calyx of spermatheca shorter, cheliceral digits also shorter with less teeth (5/1 in the new species compared to 11/4 in U. neohavu) (Table 3). It is also close to U. macrosetosus, but differs in shorter dorsal setae especially s4, z2, z4, Z5 and all macrosetae, except for StIV longer, by fewer teeth on both digits of chelicera and the shape of macrosetae that are all pointed and not knobbed as in U. macrosetosus. Table 3 Comparison of characters of the 13 species of Ueckermannseius of the havu Kreiter new species group with those of the three new species of Ueckermannseius described in this paper. Sisw In each upper boxes of the first line, the first name is the name of the species (for example bundibugyoensis ), the second line of name(s) is name(s) of describer(s) (for example Moraes, Zannou & Oliveira for bundibugyoensis ) and the third line is the source of measurements (for example Moraes et al. 2006 for bundibugyoensis ) It also closely resembles U. eastafricae Moraes, Zannou & Oliveira, but differs by shorter setae z2 and z4 and a longer Z5, a longer StIV, by the fewer teeth on both digits of chelicera and the shape of macrosetae that are all pointed in the new species and not knobbed as in U. eastafricae, the shape of macrosetae being considered as a diagnostic character in all previous descriptions.
This species was identified as U. eastafricae in two previous papers (Kreiter et al. 2021a, c) for fauna of Anjouan and Mohéli Islands, but here it is considered a new species in the new havu species group Kreiter (Table 3) and named U. gutierezzi Kreiter n. sp.
Classification. Ueckermannseius jean-mariei Kreiter n. sp. belongs to: • the subfamily Amblyseiinae (absence of dorsolateral setae z3 and s6 and the caudoventral seta JV3), • to the tribe Euseiini (sternal shield with median posterior projection, deutosternal groove > 5 µm in width, forward migration of pre-anal setae JV2 and ZV2), • to the subtribe Typhlodromalina (chelicera of normal size and shape, with prominent teeth evenly distributed along fixed digit, peritreme usually extending to level of j1, deutosternal groove narrow, 4-7 µm width), • to the genus Ueckermannseius (dorsal setae short/minute, shorter than distances between their bases, seta Z4 not as long as distance between its base and that of S4, dorsal shield smooth, except for anterolateral striation) (Chant and McMurtry 2007), • Like the two previous species and for the same reasons, to the species-group havu Kreiter new species group (see text for U. gutierrezi Kreiter n. sp.).
The following list of characters of this new species is very different from all other species of the genus and the species group. So, despite the fact that we collected a single specimen, we still consider to describe this very original specimen as belonging to a very original new species.
Peritreme and peritremal plate (Fig. 11a) -Extending to level between j1 and j3, but much closer to j1; peritremal plate fused with dorsal shield at level of j1.
Venter (Fig. 11b) -Sternal shield smooth with few lateral striae, with three pairs of setae (st1-st3) and two pairs of poroids (iv1 and iv2); a pair of seta (st4) and a pair of pores (iv3) on a small metasternal plate; posterior margin of the sternal shield with a central posterior projection. Distances st1-st1 59, st2-st2 63, st3-st3 73, st1-st3 59, st4-st4   Chelicerae (Fig. 11c) -Fixed digit 25 long, with one tooth not well visible because digit not well positioned; and movable digit 26 long, edentate, but digit also not well positioned. Pilus dentilis not visible. Spermatheca (Fig. 11d) -Like the two other new species of Ueckermannseius, spermatheca is with the atrium bulbous, the calyx basally swollen, bladder-like and then elongate and slender, this shape of the spermatheca is shared by ten African species of Ueckermannseius of the species group havu Kreiter new species group. Spermatheca 30 long and 3 wide at the widest base of calyx.
Legs (Fig. 11e)  Etymology. The name "jean-mariei" refers to the first name of the eldest brother of the senior author after him, Jean-Marie Kreiter. The species is named in his honour.
Differential diagnosis and remarks. This species is very similar to U. quilicii concerning length of setae (Table 3). However, comparison with the available characters listed in Table  (3) shows that the new species has: macrosetae all pointed (against knobbed in U. quilicii), a longer seta JV5, longer macrosetae on leg IV, both digits of chelicera longer and with less teeth (1/0 in the new species compared to 6-8/1 in U. quilicii). Other species of the species group havu Kreiter new species group are very different in many aspects concerning measurements and shape of characters (Table 3).
Zoobank: 467E989B-C03A-4997-8E25-823EECB3A126 Classification. Ueckermannseius payetae Kreiter n. sp. belongs to: • the subfamily Amblyseiinae (absence of dorsolateral setae z3 and s6 and the caudoventral seta JV3), • to the tribe Euseiini (sternal shield with median posterior projection, deutosternal groove > 5 µm in width, forward migration of pre-anal JV2 and ZV2), • to the subtribe Typhlodromalina (chelicera of normal size and shape, with prominent teeth evenly distributed along fixed digit, peritreme usually extending to level of j1, deutosternal groove narrow, 4-7 µm width), • to the genus Ueckermannseius (dorsal setae short/minute, shorter than distances between their bases, seta Z4 not as long as distance between its base and that of S4, dorsal shield smooth, except for anterolateral striation) (Chant and McMurtry 2007), • it also belongs to the species-group havu Kreiter new species group for the same reasons as the previous species (see text for previous species).
Peritreme and peritremal plate (Fig. 12a) -Extending to level of j1; peritremal plate fused with dorsal shield at level between j1 and j3.
Peritreme and peritremal plate (Fig. 13a) -Extending almost to level of j1; peritremal plate fused with dorsal shield at level between j1 and j3.
Legs (Fig. 13d)   Etymology. The name "payetae" refers to the family name of the researcher Rose-My Payet, co-author of this paper, who has worked during her career at CIRAD. She has helped the senior author in many aspects concerning fauna of Phytoseiidae of Indian Ocean Island. This species is named in her honour.
Differential diagnosis and remarks. This species is very similar to U. saltus concerning length of setae (Table 3). The description of U. saltus is however quite incomplete. But comparison with available characters listed in Table (3) allows that the new species can be distinguished by having: peritreme ending at level of j1 (and not between j1 and j3 as illustrated by Mathysse and Denmark in 1981, or very close, but anteriorly to j1 as illustrated by Moraes et al. in 2006 for U. saltus), setae r3 and s4 slightly shorter, occurrence of clear macrosetae on genu I and tibia III, a shorter calyx of spermatheca and both digits of chelicera with less teeth (8/3 in the new species compared to 10/4 in U. saltus). However, the other species of the species group havu Kreiter new species group can be clearly distinguished (Table 3). Diagnosis. The male of this species has five solenostomes (gd2, gd4, gd6, gd8 and gd9) similar to adult female, all dorsal setae lanceolate, strongly serrate and inserted on tubercules, except for J5 smooth, setiform and sharp-tipped, peritreme extending to level of j1, three setae on the ventrianal shield with small punctiform pre-anal solenostomes, three thick macrosetae strongly knobbed. This is a unique combination of characters which make specimens of this species very different from all other species within the genus Typhlodromus, subgenus Anthoseius.
Peritreme and peritremal plate (Fig. 15a) -Extending to level of z2; peritremal plate fused with dorsal shield at level of z2.
Voucher material. One male on one slide is deposited in Institut Agro (Montpellier SupAgro) -INRAE Acarology collection, Montpellier, France.
Remarks. Characters of males are very similar to that of females, except of course for length of setae and few other characters. Ventrianal shield of the male is reticulate as that of female, the sternogenital shield is almost reticulate, of normal size, macroseta StIV of leg IV in the male is the longer followed by SgeIV and StiIV as in adult female, seta JV5 smooth and sharp-tipped as in adult female. The only difference concerns dorsal setae: setae j1, z2, S5 and r3 are sharp-tipped in adult female, while also j3, j5, j6, z3 and z4 in the male.

Typhlodromus (Anthoseius) lobatus Zannou, Moraes & Oliveira
Typhlodromus (Anthoseius) lobatus Zannou, Moraes & Oliveira in Ueckermann et al. 2008: 59. Diagnosis. The male of this species has four solenostomes (gd2, gd4, gd6 and gd9), all dorsal setae sub-equal in length, smooth, except for Z4 and Z5 serrate and knobbed, peritreme extending to level between j3 and z2, four setae on ventrianal shield, one macroseta on basitarsus of leg IV, knobbed. This is a unique combination of characters that makes specimens of this species very different from all other species within the genus Typhlodromus, subgenus Anthoseius.
Peritreme and peritremal plate (Fig. 16a) -Extending to level between j3 and z2, much closer to z2; peritremal plate fused with dorsal shield at level between z4 and s4.