# Phytoseiidae of La Réunion Island (Acari: Mesostigmata): three new species and two males described, new synonymies, and new records

Serge Kreiter, Rose-My Payet, Martial Douin, Olivier Fontaine, Jacques

Fillâtre, Fabrice Le Bellec

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Serge Kreiter, Rose-My Payet, Martial Douin, Olivier Fontaine, Jacques Fillâtre, et al. Phytoseiidae of La Réunion Island (Acari: Mesostigmata): three new species and two males described, new synonymies, and new records. Acarologia, 2020, 60 (1), pp.111-195. 10.24349/acarologia/20204361. hal-02493724

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# Phytoseiidae of La Réunion Island (Acari: Mesostigmata): three new species and two males described, new synonymies, and new records

Serge Kreiter<sup>a</sup>, Rose-My Payet<sup>b</sup>, Martial Douin<sup>a</sup>, Olivier Fontaine<sup>c</sup>, Jacques Fillâtre<sup>d</sup>, Fabrice Le Bellec<sup>e</sup>

## Original research

### ABSTRACT

The Indian Ocean includes several Islands and Archipelagos and a survey of the fauna of Phytoseiidae was realized recently in these poorly known territories, despite La Réunion Island is located in an area being a hotspot of biodiversity. The Phytoseiidae fauna of La Réunion Island has been poorly investigated so far with only 33-recorded species including 24 Amblyseiinae, 5 Phytoseiinae and 4 Typhlodrominae, among which 8 species that had been described as new. All data concerning La Réunion have been published in nine papers until now. New results of surveys done recently (2015-2018) are presented in this 10th paper and add 19 newly recorded species among which 3 are new to Science and 21 already known species but with additional data. Among these 19 species, at least nine are already well known as biological control agents (BCA). Demonstration of the natural occurrence of already known efficient BCA in these territories is consequently of great agricultural and commercial interests, in addition of the contribution to fundamental knowledge of biodiversity of these poorly known territories.

**Keywords** survey; Phytoseiidae; collection; taxonomy; systematics; La Réunion Island

**Zoobank** http://zoobank.org/E7376941-8C9E-44B1-82F5-00D4A010E079

### Introduction

Several species in the family Phytoseiidae are important natural enemies of phytophagous mite and small insects in natural habitats, outdoor and protected crops all around the world (McMurtry and Croft 1997; McMurtry *et al.* 2013). However, despite the huge numbers of faunistic surveys carried out for more than 60 years, the fauna of some countries and particular ecosystems remain little explored. Consequently, it is important to survey phytoseiid faunas in poorly known areas in order to document the biodiversity of these areas, especially in biodiversity hotspots, as well as to discover new potential biological control agents (BCA). This is especially important given the context of new international and state regulations concerning import-export of BCA (Kreiter *et al.* 2020a, b).

The family Phytoseiidae is widespread all over the world and consists of 2,521 valid species dispatched in three sub-families and 94 genera (Demite *et al.* 2019).

Received 06 August 2019
Accepted 14 January 2020
Published 28 February 2020

Corresponding author
Serge Kreiter:
serge.kreiter@supagro.fr

Academic editor
Tsolakis, Haralabos

DOI
10.24349/acarologia/20204361

ISSN 0044-586X (print)
ISSN 2107-7207 (electronic)

Copyright
Kreiter S. *et al.*

Distributed under
Creative Commons CC-BY 4.0

OPEN ACCESS

<footnote><sup>a</sup> Montpellier SupAgro, UMR CBGP INRAE/IRD/CIRAD/SupAgro, 755 Avenue du Campus Agropolis (Baillarguet), CS 30016, 34988 Montferrier-sur-Lez cedex, France.</footnote>

<footnote><sup>b</sup> CIRAD, Hortsys, Station de Bassin-Plat, 97410, Saint-Pierre, Réunion, France.</footnote>

<footnote><sup>c</sup> SARL La Coccinelle, 6 Chemin Beaurivage, 97410 Saint-Pierre (La Réunion), France.</footnote>

<footnote><sup>d</sup> Armeflhor, 1 Chemin de l'IRFA, 97410 Saint-Pierre, Réunion, France.</footnote>

<footnote><sup>e</sup> HortSys, Université de Montpellier, CIRAD, TA B-103/C, Campus international de Baillarguet, 34398 Montpellier Cedex 5, France.</footnote>
Several terrestrial areas of the Indian Ocean constitute one of the world's hotspots of biodiversity. Myers (1988) defined the hotspot of biodiversity concept in order to identify the areas of biodiversity in most urgent need of conservation/protection. These world hotspots are characterized by high levels of endemism and have lost at least 70% of their original natural vegetation (Myers et al. 2000). The characterization of the phytoseiid mite diversity in these areas is thus contributing to this general topic of conservation. Located in the Indian Ocean at 700-800 km from the Eastern coast of Madagascar, La Réunion is the main island of the Mascareignes Archipelago (with the two other main islands being Mauritius and Rodrigues).

Ueckermann and Loots (1985) published the first paper concerning Phytoseiidae of La Réunion Island 35 years ago. It concerns a description of a new species to Science, *Phytoscutus reunionensis* (Ueckermann and Loots) found on *Prunus persica* (L.) in La Plaine des Cafres in 1983 by Dr Serge Quilici (Ueckermann and Loots 1985). In a study of the Tetranychidae and their predators, Guttierrez and Etienne (1986) mentioned two species of Phytoseiidae for La Réunion, *Euseius ovaloides* (Blommers) and *Phytoseiulus persimilis* Athias-Henriot, the last probably escaped in the neighbouring areas of greenhouses after releases. Quilici *et al.* (1988) mentioned then two species on litchi (*Litchi chinensis* Sonn.) in Bassin-Martin CIRAD research station, *Amblyseius largoensis* (Muma) and *E. ovaloides*, bringing to four the number of known species. In a further study focussing on mites of various crops of the Island, Quilici *et al.* (1997) mentioned six additional species and Quilici *et al.* (2000) added 14 species recorded for the first time in La Réunion. Just after, Kreiter *et al.* (2002) described seven new species from the Island, bringing to eight the number of new species described and to 31 the total number of species recorded from this island. Only one was collected in another place after its description, *Typhlodromus (Anthoseius) moraesi* Kreiter and Ueckermann, very far from La Réunion, in Guadeloupe and then also in Martinique, both in the Caribbean (Kreiter *et al.* 2013, 2018c). Moraes *et al.* (2012) added another species for La Réunion, *Neoseiulus recifensis* Gondim Jr. and Moraes, originally described from Brasil. Surprisingly, this species was found in samples from a survey for selection of a BCA against *Raoiella indica* Hirst in Brazil, along with two other common species, *A. largoensis* and *T. (A.) moraesi*. Finally, Kreiter *et al.* (2016a, b) mentioned the unexpected occurrence of *Amblyseius swirskii* Athias-Henriot in La Réunion, never collected and mentioned in previous studies, collected suddenly in high population in 2015 and 2016, bringing to 33 the number of the known species present in La Réunion Island.

We report in this 10th paper on La Réunion phytoseiids the results of additional surveys conducted from 2015 to 2018.

## Material and methods

The survey took place in La Réunion Island in 2015, 2016, 2017 and 2018. Plant-inhabiting mites were collected from various cultivated and wild plants in various locations of La Réunion. Mites were directly collected on leaves with a fine brush or by using the leaf "dipping-shaking-washing-filtering (dswf)" method of Boller (1984) or by beating the plants (mainly shrubs or trees) and collecting the mites in a black plastic rectangular saucer 45 x 30 cm (Ref. STR 45, BHR, 71370 Saint-Germain-du-Plain, France). The method selected was dependent on the plant investigated: large leaves of shrubs and trees were sampled using the direct collection method or with dswf; small leaves of shrubs and trees with the dswf or by beating and herbaceous plants with dswf.

An experiment was conducted at the Bassin-Plat CIRAD Research Station, Saint-Pierre, in La Réunion Island (altitude above sea level = aasl: 153 m, 55°29'18” E; 21°19'25” S). The experimental site was a 0.3 ha citrus orchard (*Citrus sinensis* x *C. reticulata* cv. Tangor grafted on Citrange Carrizo) with 149 meters spaced trees planted in March 2012, after a 2-year spontaneous fallow. Tree rows were planted six meters apart. The wet season spans from November to April and the dry season from June to October. The local average annual
precipitation for the period 2014 to 2016 was 1025 mm. From March 2012 to February 2014 (before the experiment started), weeds around the tree base and within 50-cm radii around them were controlled with the herbicide glyphosate (360 g.L-1 and 4 L.ha-1). Weeds in the 5-m-wide area between rows were controlled with a hammer mill.

The experiment began in March 2014. We used a complete bloc design with six replicates to test the effects of different weed management methods on the composition of the ground cover plant community. Four weed management treatments were compared: tillage (T), mowing (M), hammer mill (HM), and herbicide (H). These four treatments were distributed haphazardly within each replicate (four plots of 13 m x 5 m). Mowing and herbicide-spraying treatments were carried out in the same inter-rows so that they would not be disturbed by the tractors used for tillage or hammer mill. Hammer mill is the most commonly used engine for weed management in citrus orchards on the Réunion Island. Mowing was done with an adapted hedge-trimmer, which cuts up weeds at 10 cm above ground. A hammer mill [SML 155 SEPPI®, Caldaro (Bolzano), Italy] was used to crush weeds at the soil level. The herbicide treatment using glyphosate (360 g.L-1 at 4 L.ha-1) eliminated all weeds. A disk harrow (Grégoire and Besson®, Montigné-Montfaucon, France) was implemented once or twice in order to destroy the maximum of weeds. Weed management activities were activated when the ground cover was estimated by a farmer as being too high (70 to 80 cm height for at least one treatment). The timing for treatment was the same for all four treatments, and their timing was spaced out by a period of 93 + 27 days (mean ± SD) depending on the season.

Plots with different weed management (weed communities are variable in space and time and it is impossible to give a precise list of weed species for each date of sampling) were sampled in order to collect arthropods, including phytoseiid mites. Mites collected were then transferred with a brush into small plastic vials containing 70 % ethanol. Mites were then all mounted on slides using Hoyer's medium and identified using a phase or interferential contrast microscope (DMLB, Leica Microsystèmes SAS, Nanterre, France). Characters of specimens were measured using a graduate eyepiece (Leica, see above). We used Chant and McMurtry's (1994, 2007) concepts of the taxonomy of Phytoseiidae and the world catalogue database of Demite et al. (2019) for faunistical and biogeographical aspects. Only females were measured unless males were available. Immature will be measured and described in another paper.

In the (re)description of species, the setal nomenclature system adopted was that of Lindquist and Evans (1965) and Lindquist (1994), as adapted by Rowell et al. (1978) for the dorsum and by Chant and Yoshida-Shaul (1991) for the venter. The idiosomal setal pattern follows Chant and Yoshida-Shaul (1992). The notation for gland pores (solenostomes) or lyrifissures (poroids) is based on Athias-Henriot (1975).

Measurements of the main morphological characters were made as follows: dorsal shield length from the anterior to posterior shield margins along the midline; width between lateral margins at the level of setae *s4*; length of genital shield from the anterior margin of hyaline surface to the posterior margin of the shield; width of genital shield as the distance at the level of setae *st5* and between posterior corners of the shield; ventrianal shield length as the distance between anterior and posterior margins; ventrianal shield width between insertions of *ZV2* and at level of anus, of paranal setae between margins of the shield; cheliceral movable digit length was measured from basal articulation to tip of the digit; the fixed digit from the dorsal lyrifissure to the tip. Numbers of teeth on the fixed and movable cheliceral digits do not include the respective apical hook. Setae not referred to in the Results section should be considered as absent.

All measurements are given in micrometers (µm) and presented as the mean in bold followed by the range in parenthesis. New measurements for holotypes and paratypes of already described species are presented in tables in bold and underlined. Specimens of all species are deposited in the mite collections of Montpellier SupAgro conserved in UMR CBGP INRA/IRD/CIRAD/SupAgro. Specimens collected in fields in La Réunion within this survey were all identified. Very few single males collected alone within this study were not taken into account.
The following type or additional material have been borrowed and studied:

* The holotype of *Neoseiulus houstoni* (Schicha), from the reference collection of the Biosecurity Collections (NSW [= New South Wales] Department of Primary Industries), Orange NSW, Australia;

* The holotype, one paratype and additional material of *Neoseiulus recifensis* Gondim Jr and Moraes, housed in the mites reference collection of the Department Entomology and Acarology, Escuela Superior de Agricultura Luiz des Queiroz (ESALQ), University of Sao Paulo (USP), Piracicaba, Brasil.

We have also examined type specimens of *Amblyseius longipilus* Kreiter and Ueckermann (the holotype and 10 paratype females), *eckermannseius nesiotus* Ueckermann and Kreiter (one paratype female), *Phytoseius haroldi* Ueckermann and Kreiter (three paratype females), and *Neoseiulus barreti* Kreiter (one paratype female) of the mite collections of Montpellier SupAgro conserved in UMR CBGP, in order to complete descriptions and compared to specimens collected during this study.

The following abbreviations are used in this paper for morphological characters: **dsl** = dorsal shield length just under **j1** to just below **J5**; **dsw** = dorsal shield width at the level of **s4**; **Per. ext.**: peritreme extension; **gd** = solenostome; **Z4 ser.**, **Z5 ser.** = Z4, Z5 serrated (if Z4 and Z5 without ser. = not serrated); **knob.** = knobbed; **gensl** = genital shield length; **gensw st5** = genital shield width at the level of setae **st5**; **gensw post. corn.** = genital shield width between posterior corners; **lisl** = largest inguinal sigilla (= "metapodal plate") length; **lisw** = largest inguinal sigilla (= "metapodal plate") width; **sisl** = smallest inguinal sigilla (= "metapodal plate") length; **vsl** = ventrianal shield length; **gv3** = solenostome on ventrianal shield; **gv3 dist.** = distance between pre-anal solenostomes on the ventrianal shield; **vsw ZV2** and **vsw anus** = ventrianal shield width at ZV2 level and at paranal setae level; **asl** = anal shield length; **acw** = anal shield width at the level of paranal setae; **scl** = total spermatheca length (calyx + neck or cervix + atrium); **calyx l.**: calyx length; **scw** = calyx largest width; **FD** = fixed digit; **Fdl** = fixed digit length; **MD** = movable digit; **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. We also used the following abbreviation: **im.** = immatures; **BCA** = Biological control agents; **FCI** = French Caribbean Islands; **VCW**: various countries in the world.

The following abbreviations are used in this paper for institutions: **CBGP** = Centre de Biologie pour la Gestion des Populations; **CIRAD** = Centre International de Recherche Agronomique pour le Développement; **INRA** = Institut National de la Recherche Agronomique; **IRD** = Institut de Recherche pour le Développement; **MSA** = Montpellier SupAgro, France; **UMR** = Unité Mixte de Recherche; **UPR** = Unité Propre de Recherche, **Hortsys** = Name of the CIRAD research unit of Agroecology functioning and performances of horticultural systems; **PVBMT** = Plant Populations and Bio-aggressors in Tropical Ecosystems Joint Research Unit from University of La Réunion Island, CIRAD and INRA; **NSW** = New South Wales; **ESALQ** = Escola Superior de Agricultura Luiz de Queiroz; **USP** = Universidade de São Paulo.

## Results and discussion

A total of 44 species were found from the beginning of 2015 to the end of 2018 in our surveys.

* Four species were already well-known in the literature, very common in La Réunion and already recorded; *A. largoensis* was commonly found in the near past in different localities (Moraes *et al.* 2012) and several hundred specimens of this species as well as of the next species (see below), *A. swirskii*, were collected; *A. swirskii* (see Kreiter *et al.* 2016a) of which ten females and five males were measured.; *E. ovaloides* (mentioned in Quilici *et*
al. 1997, 2000), not measured in the present paper but this species has very characteristic features and is probably the most common species in La Réunion Island. Five hundred specimens were collected in the present study and everywhere in the Island; *T. (Anth.) moraesi* (see Kreiter *et al.* 2002), for which only eight females and one male collected in La Réunion were measured. This species is very characteristic and very common on weeds and low plants where several hundred specimens were collected during this study, mainly in experimental plots of Bassin-Plant CIRAD Research Station. Several specimens of this species was previously measured in the Caribbean islands (Kreiter *et al.* 2013, 2018c) and measurements of specimens of La Réunion agree well with measurements of these Caribbean specimens. These four species are very common and widespread in the island. Measurements of individuals of these four species are very largely overlap with those of original descriptions and of measurements published in other studies.

For the remaining 40 species, we give measurements and details in this paper:

* Nineteen have been already mentioned in previous papers (Quilici *et al.* 1997, 2000; Kreiter *et al.* 2002; Moraes *et al.* 2012) but they are rare and their occurrence have been given without any measurements which are still very interesting for further identifications: *Neoseiulus arkeri* Hughes, *N. bayviewensis* (Schicha), *N. scapilatus* (van der Merwe), *N. teke* (Pritchard and Baker), *Paraphytoseius orientalis* Narayanan, Kaur and Ghai, *Phytoseiulus persimilis* Athias-Henriot, *Scapulaseius reptans* (Blommers), *Amblyseius herbicolus* (Chant), *A. tamatavensis* (Blommers), *Proprioseiopsis mexicanus* (Garman), *Typhlodromalus spinosus* (Meyer and Rodrigues), *Ueckermannseius nesiotus* (Ueckermann and Kreiter), *Euseius hima* (Pritchard and Baker), *Phytoseius amba* Pritchard and Baker, *P. crinitus* Swirski and Shechter, *P. haroldi* Ueckermann and Kreiter, *P. intermedius* Evans and MacFarlane, *Kuzinellus scytinus* (Chazeau), *Typhlodromus (Anthoseius) transvaalensis* (Nesbitt).

* For one species, *A. longipilus*, 14 females have been measured in the original description of Kreiter *et al.* (2002) which is considered enough for a good estimate of the specific variability (Tixier 2012). Some character measures were however lacking in this original description and we provide here additional/complementary data on the species.

* For one species already known from the Island, *N. recifensis*, we were suspecting a synonymy with two other similar species of *Neoseiulus* (*N. barreti* and *N. houstoni*). These three species are compared thereafter and the male of one of this species is described for the first time.

* Seventeen species are new for the Island, namely: *Neoseiulus baraki* (Athias-Henriot), *N. californicus* (McGregor), *N. houstoni*, *N. longispinosus* (Evans), *N. lula* (Pritchard and Baker), *N. paspalivorus* (De Leon), *Paraphytoseius horrifer* (Pritchard and Baker), *Typhlodromips culmulus* (van der Merwe), *Transeius soniae* Zannou, Moraes and Oliveira, *Amblyseius neoankaratrae* (Ueckermann and Loots), *Proprioseiopsis ovatus* (Garman), *Ueckermannseius parahavu* Moraes, Zannou and Oliveira, *Amblydromalus nakuruensis* Moraes, Zannou and Oliveira, *Phytoseius punicae* Chinniah and Mohanasundaram, *P. woodburyi* De Leon, *Platyseiella eliahui* Ueckermann, *Typhlodromus (Anthoseius) ndibu* Pritchard and Baker. The male of *A. neoankaratrae* was previously unknown and is herein described.

* And finally, *three* species are new to Science, one belonging to the genus *Amblyseius* and two belonging to the genus *Transeius*. These three species are described in this paper.

All results concerning locations, measurements and some biological details for the 40 species are given thereafter.
**Subfamily Amblyseiinae Muma**

Amblyseiinae Muma, 1961: 273.

**Tribe Neoseiulini Chant & McMurtry**

Neoseiulini Chant & McMurtry 2003a: 6.

**Genus Neoseiulus Hughes**

*Neoseiulus* Hughes, 1948: 141.

**Neoseiulus baraki (Athias-Henriot)**

*Amblyseius baraki* Athias-Henriot 1966: 211.

*Amblyseius (Amblyseius) baraki*, Ehara & Bhandhufalck 1977: 54.

*Amblyseius (Neoseiulus) baraki*, Gupta 1986: 104.

*Neoseiulus baraki*, Moraes *et al*. 1986: 70; Chant & McMurtry 2003a: 27; Moraes *et al*. 2004a: 149; 2004b: 104; Zannou *et al*. 2006: 248; Chant & McMurtry 2007: 25.

*Amblyseius dhooriai* Gupta 1977: 30 (synonymy according to Gupta 1986).

This species belongs to the *paspalivorus* species group of the genus *Neoseiulus* as the female ventrianal shield is large, rectangular, rounded posteriorly, and the dorsal shield has marked “shoulders” at the level of setae *r3* (Chant and McMurtry 2003a). *Neoseiulus baraki* is a Mediterranean and subtropical species often found on monocotyledonous plants, and mainly on Poaceae. It is also a predatory mite associated with the coconut mite *Aceria guerreronis* Keifer in many parts of the world (Moraes *et al.* 2004b; Lawson-Balagbo *et al.* 2008). It is known to disperse from herbaceous weeds to the coconut “trees”. It has a flattened idiosoma with a small cross-sectional diameter (Moraes *et al.* 2004b), which enables it to reach the area underneath leaf bracts where the coconut mite feeds. Moreover, it shows a strong temporal relationship with the abundance of the coconut mite on palms (Fernando *et al.* 2003). Hence, *N. baraki* is considered a potential BCA against the coconut mite. However, in nature they are unable to maintain the coconut mite populations below the expected economic levels and so, additions of *N. baraki* to the environment to supplement natural populations for controlling the coconut mite has been considered. An essential pre-requisite to field augmentation is an effective mass rearing method. Use of coconut mites to mass rear *N. baraki* in the laboratory is expensive and time consuming. Eggs of *Tetranychus urticae* Koch, coconut pollen and maize pollen were found to be suitable alternative foods for rearing *N. baraki* but it can be more easily reared on *Tyrophagus putrescentiae* Shrank, which can be easily reared on cheap supports issued from agricultural products transformation.

This is the first mention of the occurrence of this species in La Réunion Island, the first mention in the Indian Ocean and the second mention from Sub-Saharan Africa *sensu lato*, in addition to Zannou *et al.* (2006).

**Specimens examined:** 16 ♀♀ + 8 ♂♂ in total, 12 ♀♀ + 5 ♂♂ measured. Saint-Pierre - Bassin-Plat CIRAD Research Station (altitude above sea level = aasl 153 m, Long 55°29'18" E, Lat 21°19'25" S), 2 ♀♀ + 1 ♂ on *Melinis repens* (Willd.), 20/2/2017; 6 ♀♀ + 3 ♂♂ in **CC** (Cover crop), **HM** (Hammer Mill), and **M** (Mowing) and on *Digitaria ramularis* (Trin.), 20/3, 3 and 6/4 and 20/6/2017; 1 ♂ on *Leucaena leucocephala* (Lam.), 30/03/2017; 3 ♀♀ + 2 ♂ + 1 im. on *Panicum maximum* Jacq., and 5 ♀♀ + 1 ♂ + 6 im. on *Bidens pilosa* L., 20/2, 30/3 and 20/6/2017.

**Remarks:** The measurements of characters of adult females (Table 1) are similar to those published in the literature, especially with those from specimens of Tanzania, except for the number of teeth on fixed digits of chelicerae that is higher in specimens of Tanzania.

The measurements of adult males (Table 1) are also close to those published, and closest to those obtained from specimens from Tanzania, except for the number of teeth (same remark for females).
Neoseiulus barkeri Hughes

Neoseiulus barkeri Hughes 1948: 141; Chant & McMurtry 2003a: 35; Moraes et al. 1986:70; Moraes et al. 2004: 104.

Typhlodromus (Neoseiulus) barkeri, Nesbitt 1951: 35.

Typhlodromus (Typhlodromus) barkeri, Chant 1959: 63.

Typhlodromus (Amblyseius) barkeri, Hughes 1961: 222.

Typhlodromus barkeri, Hirschmann 1962: 9.

Amblyseius barkeri, Athias-Henriot 1961: 440; Moraes et al. 1989: 95.

Amblyseius (Amblyseius) barkeri, van der Merwe 1968: 112.

Neoseiulus barkeri, Ryu et al. 2001: 8; Chant & McMurtry 2003a: 33; Moraes et al. 2004a: 104; Chant & McMurtry 2007: 25.

Amblyseius masiaka Blommers & Chazeau 1974: 308 (Synonymy according to Ueckermann & Loots 1988).

Amblyseius mckenziei Schuster & Pritchard 1963: 268 (Synonymy according to Ragusa & Athias-Henriot 1983).

<table><caption>**Table 1** Character measurements of *Neoseiulus baraki* adult females and males collected in this study with those of previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="2">Characters</th><th colspan="6">♀</th><th colspan="5">♂</th></tr><tr><th>La Réunion<br>12, this study</th><th>Africa<br>7</th><th>Sri Lanka<br>21</th><th>Taiwan<br>?</th><th>Tanzania<br>60</th><th>Thailand<br>2</th><th>Holotype<br>1</th><th>La Réunion<br>5, this study</th><th>Benin<br>20</th><th>Brazil<br>20</th><th>Tanzania<br>40</th><th>Thailand<br>1</th></tr></thead><tbody><tr><td>Dsl</td><td>358 (330-384)</td><td>353 (336-365)</td><td>350 (330-375)</td><td>365</td><td>361 (348-368)</td><td>345-358</td><td>338</td><td>280 (270-288)</td><td>276 (268-282)</td><td>288 (282-295)</td><td>278 (269-285)</td><td>350</td></tr><tr><td>Dsw</td><td>159 (150-170)</td><td>157 (142-168)</td><td>160 (146-175)</td><td>163</td><td>162 (158-168)</td><td>161-170</td><td>150</td><td>143 (138-150)</td><td>142 (136-143)</td><td>149 (146-152)</td><td>143 (133-155)</td><td>170</td></tr><tr><td>j1</td><td>15 (13-16)</td><td>15 (10-19)</td><td>14 (13-16)</td><td>17</td><td>13 (9-15)</td><td>11-15</td><td>13</td><td>13</td><td>9 (6-10)</td><td>13 (9-16)</td><td>11 (9-15)</td><td>13</td></tr><tr><td>j3</td><td>19 (16-20)</td><td>15 (13-18)</td><td>17 (13-19)</td><td>18</td><td>16 (13-19)</td><td>17-19</td><td>15</td><td>16 (13-18)</td><td>11 (9-13)</td><td>16 (13-19)</td><td>14 (10-16)</td><td>16</td></tr><tr><td>j4</td><td>12 (10-13)</td><td>9 (8-10)</td><td>11 (9-12)</td><td>11</td><td>10 (9-12)</td><td>11-12</td><td>11</td><td>9 (8-10)</td><td>8 (6-9)</td><td>9 (6-9)</td><td>9 (6-10)</td><td>11</td></tr><tr><td>j5</td><td>12 (10-14)</td><td>10 (8-11)</td><td>11 (10-18)</td><td>11</td><td>9 (6-10)</td><td>10-12</td><td>11</td><td>10 (9-11)</td><td>8 (6-9)</td><td>9 (6-10)</td><td>9 (6-10)</td><td>12</td></tr><tr><td>j6</td><td>14 (12-15)</td><td>11 (8-14)</td><td>13 (11-15)</td><td>14</td><td>12 (9-15)</td><td>13-15</td><td>13</td><td>13 (13-14)</td><td>9 (6-10)</td><td>10 (9-12)</td><td>10 (9-13)</td><td>13</td></tr><tr><td>J2</td><td>13 (11-14)</td><td>13 (11-14)</td><td>12 (10-13)</td><td>13</td><td>12 (9-15)</td><td>13-15</td><td>13</td><td>12 (10-13)</td><td>9 (6-9)</td><td>10 (9-12)</td><td>10 (10-13)</td><td>12</td></tr><tr><td>J5</td><td>12 (10-13)</td><td>10 (8-11)</td><td>11 (10-13)</td><td>12</td><td>10 (9-12)</td><td>13</td><td>11</td><td>10 (9-11)</td><td>9 (6-10)</td><td>9 (6-9)</td><td>8 (6-9)</td><td>11</td></tr><tr><td>r3</td><td>17 (14-20)</td><td>15 (11-18)</td><td>15 (13-16)</td><td>14</td><td>16 (12-19)</td><td>16</td><td>16</td><td>15 (13-18)</td><td>11 (9-13)</td><td>14 (13-16)</td><td>13 (9-15)</td><td>14</td></tr><tr><td>R1</td><td>17 (15-18)</td><td>12 (11-14)</td><td>13 (12-15)</td><td>9</td><td>16 (13-19)</td><td>15</td><td>10</td><td>13 (13-14)</td><td>12 (9-13)</td><td>15 (13-19)</td><td>12 (9-15)</td><td>16</td></tr><tr><td>s4</td><td>17 (15-20)</td><td>14 (13-18)</td><td>16 (14-17)</td><td>14</td><td>16 (13-19)</td><td>16-18</td><td>15</td><td>16 (13-18)</td><td>12 (10-13)</td><td>20 (16-22)</td><td>15 (13-16)</td><td>16</td></tr><tr><td>S2</td><td>18 (16-19)</td><td>16 (11-21)</td><td>15 (14-16)</td><td>17</td><td>16 (13-19)</td><td>16</td><td>15</td><td>16 (14-18)</td><td>13 (10-16)</td><td>19 (19-22)</td><td>14 (13-16)</td><td>16</td></tr><tr><td>S4</td><td>26 (20-30)</td><td>20 (16-29)</td><td>25 (23-27)</td><td>26</td><td>27 (22-28)</td><td>24-26</td><td>20</td><td>22 (20-23)</td><td>16 (15-19)</td><td>20 (16-22)</td><td>20 (19-22)</td><td>23</td></tr><tr><td>S5</td><td>24 (21-26)</td><td>25 (19-30)</td><td>25 (21-28)</td><td>22</td><td>26 (22-28)</td><td>23</td><td>20</td><td>19 (18-22)</td><td>16 (15-19)</td><td>19 (19-22)</td><td>20 (19-22)</td><td>21</td></tr><tr><td>z2</td><td>13 (10-15)</td><td>11 (6-16)</td><td>11 (10-13)</td><td>12</td><td>10 (9-12)</td><td>12-13</td><td>10</td><td>10</td><td>8 (6-9)</td><td>10 (9-12)</td><td>9 (6-12)</td><td>13</td></tr><tr><td>z4</td><td>15 (14-16)</td><td>13 (11-16)</td><td>14 (10-15)</td><td>12</td><td>13 (10-16)</td><td>15</td><td>13</td><td>13 (10-15)</td><td>10 (9-12)</td><td>11 (9-13)</td><td>12 (9-15)</td><td>15</td></tr><tr><td>z5</td><td>11 (10-13)</td><td>9 (8-10)</td><td>10 (9-15)</td><td>10</td><td>9 (6-12)</td><td>10-12</td><td>10</td><td>9 (8-10)</td><td>7 (6-9)</td><td>8 (6-9)</td><td>7 (6-9)</td><td>11</td></tr><tr><td>Z1</td><td>14 (11-16)</td><td>13 (11-16)</td><td>12 (11-14)</td><td>14</td><td>11 (9-15)</td><td>15</td><td>3</td><td>13</td><td>10 (7-13)</td><td>10 (9-12)</td><td>11 (9-13)</td><td>14</td></tr><tr><td>Z4</td><td>23 (21-28)</td><td>13 (18-24)</td><td>21 (19-25)</td><td>24</td><td>23 (19-28)</td><td>20-24</td><td>20</td><td>19 (14-20)</td><td>16 (10-19)</td><td>18 (13-22)</td><td>19 (16-25)</td><td>20</td></tr><tr><td>Z5</td><td>74 (65-78)</td><td>61 (53-67)</td><td>69 (60-73)</td><td>65</td><td>76 (70-79)</td><td>70-76</td><td>77</td><td>59 (55-63)</td><td>56 (51-63)</td><td>60 (57-63)</td><td>59 (54-63)</td><td>65</td></tr><tr><td>st1-st1</td><td>41 (38-43)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>39 (38-40)</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st2-st2</td><td>54 (50-58)</td><td>54 (51-59)</td><td>51 (49-53)</td><td>-</td><td>53 (51-57)</td><td>52-53</td><td>52</td><td>41 (39-43)</td><td>41 (38-41)</td><td>43 (41-44)</td><td>41 (38-44)</td><td>-</td></tr><tr><td>st3-st3</td><td>59 (56-61)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>45 (45-45)</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st1-st3 ♀ / st1-st5 ♂</td><td>87 (84-93)</td><td>80 (75-85)</td><td>83 (78-87)</td><td>-</td><td>84 (79-89)</td><td>80-81</td><td>80</td><td>125 (123-128)</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st4-st4</td><td>62 (50-76)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>58</td><td>33 (31-35)</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st5-st5</td><td>62 (58-67)</td><td>59 (56-62)</td><td>58 (55-60)</td><td>-</td><td>61 (57-66)</td><td>56-61</td><td>56</td><td>34 (32-35)</td><td>33 (32-35)</td><td>36 (34-38)</td><td>34 (32-38)</td><td>-</td></tr><tr><td>Lisl</td><td>39 (38-43)</td><td>-</td><td>-</td><td>41</td><td>-</td><td>32</td><td>-</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Lisw</td><td>5 (4-5)</td><td>-</td><td>-</td><td>4</td><td>-</td><td>-</td><td>-</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Sisl</td><td>9 (8-10)</td><td>-</td><td>-</td><td>7</td><td>-</td><td>10</td><td>-</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Vsl</td><td>112 (105-120)</td><td>117 (107-125)</td><td>112 (88-120)</td><td>111</td><td>116 (111-120)</td><td>110-115</td><td>112</td><td>103 (98-108)</td><td>105 (98-111)</td><td>110 (104-114)</td><td>104 (98-108)</td><td>-</td></tr><tr><td>Vsw ZV2</td><td>98 (90-103)</td><td>87 (82-93)</td><td>87 (71-118)</td><td>96</td><td>101 (95-105)</td><td>90-95</td><td>92</td><td>117 (98-130)</td><td>123 (114-130)</td><td>127 (120-133)</td><td>123 (111-133)</td><td>-</td></tr><tr><td>Vsw anus</td><td>84 (80-88)</td><td>86 (77-99)</td><td>83 (73-95)</td><td>-</td><td>85 (79-92)</td><td>-</td><td>87</td><td>90 (80-95)</td><td>79 (73-85)</td><td>89 (85-91)</td><td>83 (67-95)</td><td>-</td></tr><tr><td>JV5</td><td>30 (26-32)</td><td>-</td><td>-</td><td>26</td><td>-</td><td>26</td><td>-</td><td>21 (16-24)</td><td>-</td><td>-</td><td>-</td><td>26</td></tr><tr><td>StIV</td><td>39 (34-43)</td><td>34 (29-40)</td><td>38 (31-42)</td><td>42</td><td>37 (32-41)</td><td>35-40</td><td>35</td><td>36 (33-38)</td><td>26 (25-28)</td><td>35 (32-38)</td><td>31 (25-35)</td><td>34</td></tr><tr><td>Scl</td><td>10 (9-13)</td><td>12 (11-12)</td><td>10 (9-11)</td><td>14</td><td>10 (9-12)</td><td>11-12</td><td>10</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Scw</td><td>7 (5-8)</td><td>3</td><td>2 (2-3)</td><td>4</td><td>4 (3-6)</td><td>3-5</td><td>2</td><td>19 (18-23)</td><td>13 (13-16)</td><td>16 (13-16)</td><td>15 (12-19)</td><td>-</td></tr><tr><td>Fdl</td><td>28 (25-30)</td><td>25 (25-26)</td><td>31 (30-33)</td><td>-</td><td>25 (22-28)</td><td>25-28</td><td>-</td><td>4</td><td>-</td><td>-</td><td>-</td><td>8</td></tr><tr><td>No teeth Fd</td><td>8</td><td>7</td><td>10-11</td><td>11</td><td>10-11</td><td>8</td><td>8-9</td><td>21 (19-25)</td><td>18 (16-19)</td><td>20 (19-22)</td><td>19 (16-22)</td><td>-</td></tr><tr><td>Mdl</td><td>31 (28-33)</td><td>21 (21-22)</td><td>29 (27-31)</td><td>-</td><td>30 (28-32)</td><td>30-31</td><td>-</td><td>2</td><td>-</td><td>-</td><td>-</td><td>1</td></tr><tr><td>No teeth Md</td><td>2</td><td>1</td><td>2</td><td>-</td><td>2</td><td>1-2</td><td>2</td><td>13 (8-18)</td><td>9 (6-12)</td><td>10 (9-12)</td><td>9 (6-10)</td><td>-</td></tr><tr><td>Shaft</td><td colspan="6" style="text-align: center;">Not applicable</td><td></td><td colspan="5" style="text-align: center;">Not applicable</td></tr></tbody></table>

Sources of measurements - For ♀♀: Africa (Benin 4♀♀, Burundi 1♀ and Kenya 2♀♀): Zannou et al. (2006); Sri Lanka: Moraes et al. (2004b); Taiwan: Tseng (1983); Tanzania: Famah Sourassou et al. (2012); Thailand: Oliveira et al. (2012); Holotype from Algeria: Athias-Henriot (1966 in Famah Sourassou et al. 2012). For ♂♂: Benin, Brazil, Tanzania: Famah Sourassou et al. (2012); Thailand: Ehara & Bhandhufalck (1977). - : not provided.
*Amblyseius mycophilus* Karg 1970: 290 (Synonymy according to Ragusa & Athias-Henriot 1983).

*Amblyseius oahuensis* Prasad 1965: 1518 (Synonymy according to Ragusa & Athias-Henriot 1983).

*Amblyseius picketti* Specht 1968: 681 (Synonymy according to Ragusa & Athias-Henriot 1983).

*Amblyseius usitatus* van der Merwe 1965: 71 (Synonymy according to Ueckermann & Loots 1988).

This species belongs to the *barkeri* species group of the genus *Neoseiulus*, as the spermathecal atrium is large and forked at junction with major duct. It belongs to the *barkeri* species subgroup as the calyx is not markedly constricted at junction with the atrium, the atrium is deeply forked at the junction with major duct without vacuolated area, and the major duct, atrium and calyx are of approximately the same width (Chant and McMurtry 2003a).

*Neoseiulus barkeri* has a worldwide distribution (Moraes et al. 2004a; Demite et al. 2019). Various studies have shown its ability to control *Frankliniella occidentalis* Pergande (Rodriguez-Reina et al. 1992), *Thrips tabaci* (Lindeman) (Broodsgaard and Hansen 1992) and *T. urticae* in cucumber (Fan and Petitt 1994b). Fan and Petitt (1994a) showed that augmentative releases of *N. barkeri* provided control of broad mite, *Polyphagotarsonemus latus* (Banks), on peppers. *Neoseiulus barkeri* constitutes a potential BCA for several crops especially in vegetables greenhouses.

This species has been mentioned by Quilici et al., (2000) in La Réunion, with locations listed but without morphological measurements presented. Measurements of specimens collected during this study are provided in the table 2.

**Specimens examined:** 51 ♀♀ + 2 ♂♂ in total, 15 ♀♀ + 2 ♂♂ measured. St-Pierre - Ligne Paradis, La Coccinelle Inc. (aasl 164 m, Long 55°28'59" E, Lat 21°18'55" S), 5 ♀♀ in rearings of *Proprioseiopsis mexicanus* (Garman), 1/1/2017; Vincendo - Delaunay Jean-Max farm (aasl 110 m, Long 55°67'14" E, Lat 21°38" S), 45 ♀♀ + 2 ♂♂ on *Capsicum annuum* L., 11 and 18/1/2016 and 15/12/2016; Montvert-les-Hauts - EARL Le Mont Vert farm (aasl 582 m, Long 55°32'19" E, Lat 21°19'42" S), 1 ♀ on *C. annuum*, 19/9/2016.

**Remarks:** measurements of characters of females from La Réunion Island are only slightly different from female specimens from other countries, with less than 10 % differences (Table 2). All setae of specimens from South Africa (Ueckermann and Meyer 1988) are just slightly longer from those from other countries. Measurements of characters of males from La Réunion Island are also slightly different from male specimens from other countries, with < 10% differences (Table 2). In general, setae of specimens from La Réunion appears a bit smaller than those from other African countries but dorsal shied is larger.

Comparisons with *N. barkeri* measurements of female and male (Table 2) specimens of various origins in Beaulieu and Beard (2018) shows shorter dimensions of all characters of *La Réunion* specimens (all ranges of *La Réunion* specimens are in the lower parts of the ranges mentioned by these authors). These authors already mention in their paper the shorter dimensions of dorsal setae of African female and male specimens (lower part of observed ranges) compared to their own measurements (Beaulieu and Beard 2018).

## *Neoseiulus bayviewensis* (Schicha)

*Amblyseius bayviewensis* Schicha 1977b: 394; Schicha 1987: 107.

*Neoseiulus bayviewensis*, Moraes et al. 1986: 72; Quilici et al. 1997: 285, 2000: 101; Beard 2001: 97; Kreiter et al. 2002: 348; Chant & McMurtry 2003: 21; Moraes et al. 2004: 107; Chant & McMurtry 2007: 25.

This species belongs to the *cucumeris* species group of *Neoseiulus* as dorsocentral setae are not as short relative to dorsolateral setae, and the ratio s4/j6 is 1.3 to 3.5. The spermatheca is without a stalk between calyx and atrium, the atrium is differentiated, joined directly to calyx. The species belong to the *cucumeris* species subgroup (Chant and McMurtry 2003a).
This species was only known from Australia for a long time (Demite et al. 2019). Quilici *et al.* (1997, 2000) collected this species on *Hibiscus* sp., associated with populations of the eriophyid *Aceria hibisci* (Nalepa), which is common in La Réunion. However, the biology of this predator remains unknown.

Collection data were provided in previous papers (Quilici *et al.* 1997, 2000) but without measurements of specimens. Measurements of specimens collected during this study are provided in table 3.

**Specimens examined:** 2 ♀♀ in total, both measured. Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29′18″ E, Lat 21°19′25″ S), 2 ♀♀ in the plot H (Herbicide), 4/4/2017 (see Material and methods).

**Remarks:** measurements of characters of La Réunion Island female specimens largely overlap with those of female specimens from Australia (Table 3). Measurements of La Réunion specimens are in general just a few percent's shorter, except for some setae or character dimensions which are quite shorter in La Réunion specimens, for example macrosetae of leg IV and size of ventral shields.

<table><caption>**Table 2** Character measurements of adult females and males of *Neoseiulus barkeri* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="2">Characters</th><th colspan="6">♀</th><th colspan="6">♂</th></tr><tr><th>La Réunion</th><th>Africa 1</th><th>Africa 2</th><th>Madagascar</th><th>Senegal</th><th>South Africa</th><th>Thailand</th><th>Various origins</th><th>La Réunion</th><th>Japan</th><th>Madagascar</th><th>South Africa</th><th>Thailand</th><th>Various origins</th></tr></thead><tbody><tr><td>Dsl</td><td><strong>353</strong> (333 – 373)</td><td>332 (318 – 348)</td><td>340 (328 – 360)</td><td>355</td><td>355</td><td>330 – 340</td><td>358 (336 – 377)</td><td>350 – 398</td><td><strong>260 – 268</strong></td><td>270</td><td>280</td><td>246 – 271</td><td>258 – 260</td><td>282 – 301</td></tr><tr><td>Dsw</td><td><strong>198</strong> (180 – 223)</td><td>188 (180 – 204)</td><td>185 (173 – 203)</td><td>210</td><td>195</td><td>184 – 196</td><td>195 (185 – 210)</td><td>182 – 220</td><td><strong>160 – 163</strong></td><td>180</td><td>195</td><td>154 – 183</td><td>165 – 173</td><td>176 – 192</td></tr><tr><td>j1</td><td><strong>14</strong> (13 – 15)</td><td>16 (14 – 17)</td><td>16 (14 – 17)</td><td>16 – 18</td><td>15</td><td>17 – 20</td><td>18 (17 – 19)</td><td>18 – 23</td><td><strong>13 – 15</strong></td><td>14</td><td>12</td><td>13 – 16</td><td>11 – 15</td><td>-</td></tr><tr><td>j3</td><td><strong>17</strong> (13 – 20)</td><td>20 (18 – 23)</td><td>19 (18 – 21)</td><td>16 – 18</td><td>18</td><td>23 – 26</td><td>23 (23 – 24)</td><td>22 – 30</td><td><strong>13 – 15</strong></td><td>19</td><td>15</td><td>15 – 19</td><td>18</td><td>-</td></tr><tr><td>j4</td><td><strong>13</strong> (5 – 20)</td><td>16 (14 – 17)</td><td>15 (13 – 16)</td><td>16 – 18</td><td>15</td><td>17 – 20</td><td>17 (16 – 18)</td><td>17 – 23</td><td><strong>13</strong></td><td>15</td><td>12</td><td>12 – 13</td><td>13</td><td>-</td></tr><tr><td>j5</td><td><strong>13</strong> (10 – 15)</td><td>16 (14 – 17)</td><td>15 (13 – 16)</td><td>16 – 18</td><td>15</td><td>17 – 20</td><td>17</td><td>17 – 23</td><td><strong>13</strong></td><td>13</td><td>12</td><td>12 – 13</td><td>15</td><td>-</td></tr><tr><td>j6</td><td><strong>14</strong> (8 – 18)</td><td>18 (16 – 19)</td><td>16 (13 – 18)</td><td>16 – 18</td><td>18</td><td>17 – 20</td><td>20</td><td>18 – 25</td><td><strong>13</strong></td><td>15</td><td>15</td><td>13 – 16</td><td>13 – 16</td><td>-</td></tr><tr><td>j2</td><td><strong>16</strong> (10 – 20)</td><td>19 (19 – 20)</td><td>19 (16 – 22)</td><td>20</td><td>19</td><td>20 – 24</td><td>19 (17 – 21)</td><td>21 – 31</td><td><strong>13</strong></td><td>17</td><td>15</td><td>13 – 16</td><td>12 – 15</td><td>-</td></tr><tr><td>j5</td><td><strong>10</strong> (8 – 13)</td><td>11 (10 – 11)</td><td>10 (10 – 11)</td><td>-</td><td>10</td><td>19 – 23</td><td>12 (11 – 12)</td><td>12 – 15</td><td><strong>8</strong></td><td>7</td><td>10</td><td>9</td><td>8</td><td>-</td></tr><tr><td>r3</td><td><strong>16</strong> (10 – 18)</td><td>18 (17 – 20)</td><td>18 (16 – 19)</td><td>18</td><td>15</td><td>19 – 23</td><td>20</td><td>19 – 28</td><td><strong>13</strong></td><td>17</td><td>15</td><td>15 – 19</td><td>15 – 16</td><td>-</td></tr><tr><td>R1</td><td><strong>16</strong> (13 – 18)</td><td>16 (14 – 17)</td><td>16 (14 – 16)</td><td>18</td><td>13</td><td>19 – 23</td><td>20 (19 – 20)</td><td>19 – 28</td><td><strong>13</strong></td><td>15</td><td>15</td><td>13 – 16</td><td>15</td><td>-</td></tr><tr><td>s4</td><td><strong>18</strong> (8 – 23)</td><td>21 (19 – 24)</td><td>20 (18 – 22)</td><td>20</td><td>23</td><td>23 – 26</td><td>26 (25 – 26)</td><td>26 – 34</td><td><strong>15</strong></td><td>21</td><td>18</td><td>15 – 19</td><td>20 – 21</td><td>-</td></tr><tr><td>S2</td><td><strong>21</strong> (18 – 25)</td><td>23 (22 – 24)</td><td>20 (19 – 24)</td><td>22</td><td>23</td><td>20 – 24</td><td>24</td><td>25 – 37</td><td><strong>20</strong></td><td>18</td><td>18</td><td>15 – 19</td><td>18</td><td>-</td></tr><tr><td>S4</td><td><strong>19</strong> (15 – 23)</td><td>19 (17 – 22)</td><td>20 (18 – 22)</td><td>22</td><td>20</td><td>20 – 24</td><td>23 (22 – 23)</td><td>24 – 36</td><td><strong>15</strong></td><td>17</td><td>18</td><td>13 – 16</td><td>18 – 19</td><td>-</td></tr><tr><td>S5</td><td><strong>18</strong> (18 – 23)</td><td>18 (17 – 19)</td><td>17 (16 – 19)</td><td>22</td><td>18</td><td>17 – 20</td><td>21 (20 – 22)</td><td>20 – 34</td><td><strong>15</strong></td><td>15</td><td>15</td><td>13 – 16</td><td>15 – 17</td><td>-</td></tr><tr><td>c2</td><td><strong>16</strong> (10 – 18)</td><td>18 (17 – 19)</td><td>18 (18 – 21)</td><td>16</td><td>19</td><td>20 – 24</td><td>21 (20 – 21)</td><td>20 – 26</td><td><strong>13</strong></td><td>17</td><td>15</td><td>13 – 16</td><td>15 – 19</td><td>-</td></tr><tr><td>c4</td><td><strong>16</strong> (10 – 20)</td><td>18 (17 – 19)</td><td>17 (16 – 19)</td><td>20</td><td>20</td><td>20 – 24</td><td>21</td><td>20 – 28</td><td><strong>15</strong></td><td>18</td><td>15</td><td>13 – 16</td><td>16 – 17</td><td>-</td></tr><tr><td>c5</td><td><strong>13</strong> (10 – 15)</td><td>17 (16 – 19)</td><td>16 (13 – 18)</td><td>16</td><td>16</td><td>20 – 24</td><td>18</td><td>18 – 23</td><td><strong>15</strong></td><td>16</td><td>12</td><td>14</td><td>15 – 16</td><td>-</td></tr><tr><td>Z1</td><td><strong>17</strong> (15 – 20)</td><td>20 (19 – 22)</td><td>19 (18 – 21)</td><td>22</td><td>21</td><td>20 – 24</td><td>23 (22 – 23)</td><td>22 – 32</td><td><strong>10</strong></td><td>17</td><td>15</td><td>15 – 19</td><td>16 – 17</td><td>-</td></tr><tr><td>Z4</td><td><strong>33</strong> (30 – 38)</td><td>33 (31 – 34)</td><td>33 (29 – 38)</td><td>40</td><td>33</td><td>33 – 36</td><td>33 (32 – 33)</td><td>35 – 45</td><td><strong>25</strong></td><td>25</td><td>30</td><td>28 – 30</td><td>22 – 23</td><td>29 – 32</td></tr><tr><td>Z5</td><td><strong>50</strong> (45 – 53)</td><td>49 (46 – 54)</td><td>51 (46 – 56)</td><td>55</td><td>45</td><td>52 – 55</td><td>53 (52 – 55)</td><td>48 – 66</td><td><strong>30</strong></td><td>29</td><td>35</td><td>31 – 38</td><td>27 – 29</td><td>35 – 39</td></tr><tr><td>st1-st1</td><td><strong>52</strong> (48 – 55)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td><strong>45</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st2-st2</td><td><strong>65</strong> (63 – 68)</td><td>62 (60 – 67)</td><td>63 (61 – 67)</td><td>-</td><td>65</td><td>68 – 72</td><td>68 (65 – 70)</td><td>67 – 76</td><td><strong>55</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st3-st3</td><td><strong>73</strong> (70 – 75)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td><strong>58</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>86 – 93</td></tr><tr><td>st1-st3 ♀ / st1-st5 ♂</td><td><strong>67</strong> (63 – 70)</td><td>61 (58 – 62)</td><td>63 (61 – 65)</td><td>70</td><td>63</td><td>78 – 84</td><td>69 (67 – 72)</td><td>-</td><td><strong>108 – 113</strong></td><td>-</td><td>-</td><td>125</td><td>-</td><td>120 – 129</td></tr><tr><td>st4-st4</td><td><strong>73</strong> (65 – 80)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td><strong>45 – 48</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st5-st5</td><td><strong>64</strong> (60 – 68)</td><td>61 (60 – 62)</td><td>60 (56 – 63)</td><td>-</td><td>55</td><td>70 – 78</td><td>61 (60 – 64)</td><td>-</td><td><strong>35 – 38</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Lsl</td><td><strong>26</strong> (25 – 30)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>24 – 32</td><td colspan="6"></td></tr><tr><td>Lisw</td><td><strong>4</strong> (3 – 5)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="6">Not applicable</td></tr><tr><td>Ssl</td><td><strong>14</strong> (10 – 18)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="6"></td></tr><tr><td>Vsl</td><td><strong>120</strong> (113 – 128)</td><td>117 (115 – 120)</td><td>114 (109 – 120)</td><td>125</td><td>113</td><td>112 – 117</td><td>131 (122 – 139)</td><td>116 – 145</td><td><strong>105 – 110</strong></td><td>-</td><td>120</td><td>106 – 111</td><td>110</td><td>-</td></tr><tr><td>Vsw ZV2</td><td><strong>96</strong> (90 – 100)</td><td>105 (101 – 108)</td><td>102 (96 – 107)</td><td>110</td><td>90</td><td>98 – 104</td><td>106 (105 – 108)</td><td>99 – 120</td><td><strong>130 – 133</strong></td><td>-</td><td>145</td><td>85 – 91</td><td>133</td><td>133 – 146</td></tr><tr><td>Vsw anus</td><td><strong>29</strong> (28 – 33)</td><td>81 (72 – 86)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td><strong>63 – 68</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>JV5</td><td><strong>40</strong> (33 – 43)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>43 – 46</td><td>-</td><td>44 – 60</td><td><strong>30</strong></td><td>22</td><td>-</td><td>25 – 32</td><td>-</td><td>24 – 37</td></tr><tr><td>StIV</td><td><strong>57</strong> (48 – 68)</td><td>62 (60 – 67)</td><td>63 (58 – 69)</td><td>62</td><td>55</td><td>66</td><td>66 (62 – 70)</td><td>58 – 74</td><td><strong>42 – 50</strong></td><td>42</td><td>60</td><td>55 – 60</td><td>43 – 46</td><td>48 – 52</td></tr><tr><td>Scls</td><td><strong>40</strong> (20 – 50)</td><td>19 (17 – 22)</td><td>-</td><td>-</td><td>25</td><td>22</td><td>19 (18 – 20)</td><td>17 – 25</td><td colspan="6">Not applicable</td></tr><tr><td>Scw</td><td><strong>22</strong> (18 – 25)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>5 – 11</td><td colspan="6"></td></tr><tr><td>Fdl</td><td><strong>33</strong> (30 – 38)</td><td>-</td><td>30 (29 – 31)</td><td>29</td><td>-</td><td>30</td><td>28 (25 – 30)</td><td>30 – 34</td><td><strong>20</strong></td><td>-</td><td>24</td><td>-</td><td>22</td><td>20 – 23</td></tr><tr><td>No teeth Fd</td><td><strong>5</strong></td><td>-</td><td>5</td><td>-</td><td>-</td><td>3</td><td>-</td><td>4 – 6</td><td><strong>4</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>3 – 5</td></tr><tr><td>Mdl</td><td><strong>31</strong> (28 – 33)</td><td>-</td><td>33 (32 – 33)</td><td>33</td><td>-</td><td>34</td><td>38 (36 – 40)</td><td>33 – 37</td><td><strong>20</strong></td><td>-</td><td>24</td><td>-</td><td>23 – 26</td><td>21 – 23</td></tr><tr><td>No teeth Md</td><td><strong>1</strong></td><td>-</td><td>1</td><td>-</td><td>-</td><td>1</td><td>-</td><td>1</td><td><strong>1</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>1</td></tr><tr><td>Shaft</td><td colspan="6">Not applicable</td><td>-</td><td>-</td><td><strong>15</strong></td><td>-</td><td>15</td><td>-</td><td>13 – 21</td><td>15 – 17</td></tr></tbody></table>

Sources of measurements – For ♀♀: Africa 1 (Ghana = Nigeria); Moraes *et al.* (1989b); Africa 2 (Benin: ♀♀; Burundi: 1♀; Ghana: 1♀; Kenya: 2♀♀; Nigeria: 1♀; Mozambique: 1♀): Zannou *et al.* (2006); Madagascar: Blommers & Chazeau (1974); Senegal: Kade *et al.* (2011); South Africa (identified as *Amblyseius usitatus*, synonymized Ueckermann & Loots 1988): van der Merwe (1965); Thailand: Oliveira *et al.* (2012); Various origins (Canada: 7♀♀; Finland: 20♀♀; South Korea: 9♀♀; Spanish Morocco: 1♀; UK: 2♀♀; USA: 14♀♀): Beaulieu & Beard (2018). For ♂♂: Japan: Ehara (1972); Madagascar (identified as *Amblyseius masiaka*, synonymized by Ueckermann & Loots 1988): Blommers & Chazeau (1974); South Africa: Ueckermann & Loots (1988); Thailand: Oliveira *et al.* (2012); Various origins (Finland: 13♂♂; South Korea: 1♂; UK: 1♂; USA: 2♂♂): Beaulieu & Beard (2018). - : not provided.
Neoseiulus californicus (McGregor)

*Typhlodromus californicus* McGregor 1954: 89.

*Amblyseius californicus*, Schuster & Pritchard 1963: 271.

*Cydnodromus californicus*, Athias-Henriot 1977: 62.

*Amblyseius (Amblyseius) californicus*, Ueckermann & Loots 1988: 150; Ehara *et al.* 1994: 126.

*Amblyseius (Neoseiulus) californicus*, Ehara & Amano 1998: 33.

*Neoseiulus californicus*, Moraes *et al.* 1986: 73; Chant & McMurtry 2003a: 21; Moraes *et al.* 2004a: 109; Chant & McMurtry 2007: 25; Guanilo *et al.* 2008a: 27, 2008b: 19.

*Neoseiulus chilenensis* Dosse 1958: 55 (synonymy according to McMurtry & Badii 1989).

*Neoseiulus mungeri* McGregor 1954: 92 (synonymy according to Schuster & Pritchard 1963).

*Neoseiulus wearnei* Schicha 1987: 103 (Synonymy according to Tixier *et al.* 2014).

Like the previous species, *N. californicus* belongs also to the *cucumeris* species group of *Neoseiulus* (Chant and McMurtry 2003a).

This widespread species (Moraes *et al.* 2004; Demite *et al.* 2019) is considered by McMurtry and Croft (1997) to be a specialized predator, Type 2. Nevertheless, it has characteristics of both specialist and generalist predatory mites (Castagnoli and Simoni 2003). It prefers to feed on spider mites (Gomez *et al.* 2009), but can also consume other mite species like tarsonemid mites [*Phytonemus pallidus* (Banks)] (Easterbrook *et al.* 2001), small insects such as thrips (Rodriguez-Reina *et al.* 1992) and even pollen when the primary prey is unavailable (Rhodes and Liburd 2006). It can migrate from grasses to fruit trees or grapevines and vice versa (Auger *et al.* 1999). It is a specialist predator of *T. urticae* on annual plants and woody species, and of *Panonychus ulmi* (Koch) and various *Tetranychus* spp. (and perhaps eriophyid mites) on trees and less frequently on grapevines (Auger *et al.* 1999). *N. californicus* is well-known as a BCA sold in many countries around the world for the management of spider mites in greenhouses but also in outdoor crops such as fruit crops in Europe. This is the first mention of that species for La Réunion Island.

<table><caption>**Table 3** Character measurements of adult females of *Neoseiulus bayviewensis* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th>Characters</th><th>La Réunion<br>2, this study</th><th>Australia<br>7</th></tr></thead><tbody><tr><td>Dsl</td><td><strong>293 – 350</strong></td><td>349 – 357</td></tr><tr><td>Dsw</td><td><strong>143 – 150</strong></td><td>163 – 174</td></tr><tr><td>j1</td><td><strong>23</strong></td><td>21 – 25</td></tr><tr><td>j3</td><td><strong>33 – 38</strong></td><td>42 – 44</td></tr><tr><td>j4</td><td><strong>18 – 20</strong></td><td>23 – 25</td></tr><tr><td>j5</td><td><strong>16 – 20</strong></td><td>21 – 23</td></tr><tr><td>j6</td><td><strong>20</strong></td><td>21 – 24</td></tr><tr><td>J2</td><td><strong>18 – 25</strong></td><td>28 – 33</td></tr><tr><td>J5</td><td><strong>8</strong></td><td>11 – 12</td></tr><tr><td>r3</td><td><strong>28 – 30</strong></td><td>28 – 31</td></tr><tr><td>R1</td><td><strong>25 – 28</strong></td><td>28 – 31</td></tr><tr><td>s4</td><td><strong>43 – 48</strong></td><td>47 – 51</td></tr><tr><td>S2</td><td><strong>43 – 45</strong></td><td>43 – 49</td></tr><tr><td>S4</td><td><strong>40</strong></td><td>36 – 39</td></tr><tr><td>S5</td><td><strong>23 – 24</strong></td><td>26 – 29</td></tr></tbody></table><table><thead><tr><th>Characters</th><th>La Réunion<br>2, this study</th><th>Australia<br>7</th></tr></thead><tbody><tr><td>z2</td><td><strong>30 – 33</strong></td><td>35 – 38</td></tr><tr><td>z4</td><td><strong>30 – 35</strong></td><td>38 – 43</td></tr><tr><td>z5</td><td><strong>18 – 20</strong></td><td>19 – 20</td></tr><tr><td>Z1</td><td><strong>26 – 28</strong></td><td>32 – 38</td></tr><tr><td>Z4</td><td><strong>45 – 48</strong></td><td>47 – 50</td></tr><tr><td>Z5</td><td><strong>46 – 53</strong></td><td>57 – 61</td></tr><tr><td>st1-st1</td><td><strong>50</strong></td><td>-</td></tr><tr><td>st2-st2</td><td><strong>59 – 61</strong></td><td>-</td></tr><tr><td>st3-st3</td><td><strong>70 – 75</strong></td><td>96 – 101</td></tr><tr><td>st1-st3</td><td><strong>50 – 55</strong></td><td>68 – 71</td></tr><tr><td>st4-st4</td><td><strong>83 – 85</strong></td><td>-</td></tr><tr><td>st5-st5</td><td><strong>56 – 60</strong></td><td>73 – 78</td></tr><tr><td>Lisl</td><td><strong>20</strong></td><td>-</td></tr><tr><td>Lisw</td><td><strong>4</strong></td><td>-</td></tr><tr><td>Sisl</td><td><strong>6</strong></td><td>-</td></tr></tbody></table><table><thead><tr><th>Characters</th><th>La Réunion<br>2, this study</th><th>Australia<br>7</th></tr></thead><tbody><tr><td>Vsl</td><td><strong>90 – 125</strong></td><td>111 – 117</td></tr><tr><td>Vsw ZV2</td><td><strong>78 – 95</strong></td><td>75 – 85</td></tr><tr><td>Vsw <i>anus</i></td><td><strong>58 – 65</strong></td><td>-</td></tr><tr><td>JV5</td><td><strong>38 – 40</strong></td><td>-</td></tr><tr><td>SgeIV</td><td><strong>20 – 23</strong></td><td>28 – 31</td></tr><tr><td>StIV</td><td><strong>18</strong></td><td>21 – 28</td></tr><tr><td>StIV</td><td><strong>48</strong></td><td>61 – 65</td></tr><tr><td>Scls</td><td><strong>13</strong></td><td>14 – 16</td></tr><tr><td>Scw</td><td><strong>10</strong></td><td>-</td></tr><tr><td>Fdl</td><td><strong>23</strong></td><td>33 – 34</td></tr><tr><td>No teeth Fd</td><td><strong>3?</strong></td><td>4</td></tr><tr><td>Mdl</td><td><strong>25</strong></td><td>33 – 34</td></tr><tr><td>No teeth Md</td><td><strong>1?</strong></td><td>2</td></tr></tbody></table>

Sources of measurements – Australia: Schicha (1977); -: not provided.
**Specimens examined:** 18 ♀♀ + 2 ♂♂ in total, 14 ♀♀ + 2 ♂♂ measured. Ravine des Cabris - Ligne des Bambous, Lassay (aasl 221 m, Long 55°29'38" E, Lat 21°17'17" S), 1 ♀ on *Amaranthus viridis* L., 5/12/2016; Le Tampon - Grand Tampon (aasl 1100 m, Long 55°34'12" E, Lat 21°16'48" S), 6 ♀♀ + 1 ♂ on *Cyperus rotundus* L., 1 ♀ + 1 ♂ on *Plantago lanceolata* L. and 1 ♀ on *Solanum mauritianum* Scop., 18/1/2017; Montvert-les-Hauts - EARL Le Mont Vert farm (aasl 582 m, Long 55°32'19" E, Lat 21°19'42" S), 1 ♀ on *Fragaria* sp., 4/8/2015, 1 ♀ on *C. annuum*, 23/8/2016; Le 19ᵉ - Plaine des Caffres, JL Robert farm (aasl 1000 m, Long 55°32'9" E, Lat 21°14'16" S), 1 ♀ on *Physalis peruviana* L., and 2 ♀♀ on *Emilia sonchifolia* (L.) DC., 15/12/2015; Saint-Pierre - Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29'18" E, Lat 21°19'25" S), 4 ♀♀ in plots BM and CC, 4/4/2017.

**Remarks:** measurements of characters of the 14 female specimens (Table 4) fit well with those obtained for populations of other countries, such as those obtained from Tixier *et al.* (2008).

Values of measurements of La Réunion specimens are generally only slightly smaller (few percents variations). The same remark can be addressed for the adult male specimens (Table 4).

Comparisons with *N. californicus* measurements of a large number of female (Table 4) specimens of various origins in Beaulieu and Beard (2018) shows shorter dimensions of all characters of La Réunion specimens (all ranges of La Réunion specimens are in the lower parts of the ranges mentioned by these authors).

This is interesting to notice that it is not the case for males (Table 4), measurements of male specimens from La Réunion covering ranges or being in the middle of ranges mentioned by Beaulieu and Beard (2018) in their redescription.

## Neoseiulus houstoni Schicha

*Neoseiulus houstoni*, Schicha 1987: 111; Chant & McMurtry 2003a: 23; Moraes *et al.* 2004a: 123.

*Neoseiulus recifensis* Gondim Jr. & Moraes 2001: 77; Chant & McMurtry 2003a: 23; Moraes *et al.* 2004a: 140, **new synonymy**.

*Neoseiulus barreti* Kreiter, in Furtado *et al.* 2005: 135, **new synonymy**.

These three species belong also to the *cucumeris* species group of *Neoseiulus* like previous species. However, whereas Chant and McMurtry (2003a) classified *N. recifensis* in the *cucumeris* species subgroup, *N. houstoni* was placed in the *paraki* species subgroup (also in the *cucumeris* species group), despite the two species having identical spermathecae (Chant and McMurtry 2003a). *Neoseiulus barreti* was described later than 2003, is not mentioned in Chant and McMurtry (2007) but the spermatheca is also identical to that of the two former species.

*Neoseiulus houstoni* was the first species collected and described in 1987 on *Vigna unguiculata* (L.) Walp. in Queensland, Australia (Schicha 1987).

*Neoseiulus recifensis* was discovered for the first time in Brazil in 2001 (Gondim Jr. and Moraes 2001), collected on *Cocos nucifera* L. in Recife and Itamaraca, Pernambuco, Brazil. It was collected then in 2007 and 2008 in Brazil in the states of Alagoas, Bahia, Cera, Paraiba and Rio Grande do Norte (Fiaboe *et al.* 2007; Lawson-Balagbo *et al.* 2008). Finally, it was collected in La Réunion Island in 2011 within a survey investigating potential predators of *R. indica* on coconut (Moares *et al.* 2012).

Two females of *N. barreti* were collected, only one time, in Brazil in 2004 on *Solanum paniculatum* L. in Itapaje, Ceara, Brazil and described later (Furtado *et al.* 2005).

Biology of the three species remain unknown. Males of the three species are unknown.

**Specimens examined:** 37 ♀♀ + 8 ♂♂ + 4 im. in total, 24 ♀♀ + 8 ♂♂ measured. Langevin - Jacqueline Waterfall (aasl 5 m, Long 55°64'40" E, Lat 21°38'69" S), 4 ♀♀ + 2 ♂♂ on *Casuarina equisetifolia* L., and 3 ♀♀ + 1 im. on *Scaevola taccada* Vahl, 19/7/2017; Saint-Pierre - Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29'18" E, Lat 21°19'25" S), 1 ♀ in plot H, 12/4/2016; 1 ♀ on *B. pilosa*, 1 ♀ on *Euphorbia hypericifolia* Lam., and 1 ♀
on *P. maximum*, 20/2/2017; 1 ♀ on *Artocarpus heterophyllus* Lam., 5 ♀♀ on *C. equisetifolia*, 1 ♀ on *Acacia mearnsii* De Wild., and 2 ♂ on *Crotalaria retusa* L., 27/2/2017; 4 ♀♀ + 2 im. on *B. pilosa*, 2 ♀♀ on *L. leucocephala*, 1 ♀ on *Malvastrum coromandelianum* (L.), and 8 ♀♀ + 4 ♂♂ + 1 im. on *Parthenium hysterophorus* L., 30/3/2017; 1 ♀ in plot CC, 3/04/2017 and 1 ♀ in plot CC, 6/04/2017; 1 ♀ on *B. pilosa*, 19/6/2017; 1 ♀ on *A. viridis*, 20/6/2017.

We have also examined the following type or additional material:

* The holotype of *N. houstoni*;

* The holotype, paratypes female and additional female material of *N. recifensis*;

* One paratype female of *N. barreti*.

<table><caption>**Table 4** Character measurements of adult females and males of *Neoseiulus californicus* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="2">Characters</th><th colspan="6">♀</th><th colspan="5">♂</th></tr><tr><th>La Réunion<br>14, this study</th><th>Argentina<br>3</th><th>Australia<br>1</th><th>Peru<br>4</th><th>Senegal<br>5</th><th>VCW 1<br>300</th><th>VCW 2<br>99</th><th>La Réunion<br>2, this study</th><th>Argentina<br>1</th><th>Japan<br>1</th><th>Peru<br>1</th><th>VCW<br>34</th></tr></thead><tbody><tr><td>Dsl</td><td><b>374</b> (353 – 403)</td><td>345 (325 – 360)</td><td>370</td><td>378 (360 – 398)</td><td>350 (340 – 375)</td><td>367 (330 – 406)</td><td>340 – 406</td><td><b>288 – 293</b></td><td>280</td><td>280</td><td>288</td><td>265 – 316</td></tr><tr><td>Dsw</td><td><b>188</b> (168 – 205)</td><td>167 (150 – 175)</td><td>150</td><td>176 (160 – 185)</td><td>174 (168 – 187)</td><td>146 (130 – 189)</td><td>168 – 216</td><td><b>163 – 165</b></td><td>160</td><td>170</td><td>160</td><td>111 – 131</td></tr><tr><td>j1</td><td><b>21</b> (18 – 23)</td><td>21 (20 – 22)</td><td>21</td><td>23 (21 – 25)</td><td>22 (20 – 23)</td><td>22 (17 – 26)</td><td>20 – 26</td><td><b>15 – 18</b></td><td>16</td><td>17</td><td>18</td><td>-</td></tr><tr><td>j3</td><td><b>27</b> (20 – 30)</td><td>28 (25 – 30)</td><td>29</td><td>34 (31 – 38)</td><td>30 (25 – 33)</td><td>31 (22 – 39)</td><td>26 – 38</td><td><b>20 – 25</b></td><td>27</td><td>18</td><td>25</td><td>-</td></tr><tr><td>j4</td><td><b>21</b> (18 – 25)</td><td>21 (20 – 22)</td><td>19</td><td>24 (22 – 26)</td><td>25 (23 – 28)</td><td>22 (16 – 27)</td><td>18 – 30</td><td><b>15 – 18</b></td><td>20</td><td>14</td><td>18</td><td>-</td></tr><tr><td>j5</td><td><b>20</b> (15 – 23)</td><td>21 (20 – 24)</td><td>19</td><td>25 (22 – 26)</td><td>24 (20 – 27)</td><td>22 (16 – 28)</td><td>18 – 30</td><td><b>13 – 15</b></td><td>19</td><td>14</td><td>18</td><td>-</td></tr><tr><td>j6</td><td><b>26</b> (23 – 30)</td><td>25 (22 – 28)</td><td>25</td><td>30 (25 – 36)</td><td>28 (25 – 33)</td><td>27 (19 – 32)</td><td>24 – 36</td><td><b>18 – 20</b></td><td>23</td><td>17</td><td>25</td><td>-</td></tr><tr><td>j2</td><td><b>29</b> (28 – 30)</td><td>32 (30 – 35)</td><td>25</td><td>36 (31 – 40)</td><td>31 (28 – 33)</td><td>32 (25 – 40)</td><td>28 – 41</td><td><b>20 – 25</b></td><td>28</td><td>22</td><td>28</td><td>-</td></tr><tr><td>j5</td><td><b>12</b> (10 – 13)</td><td>11 (10 – 13)</td><td>10</td><td>13 (10 – 15)</td><td>13 (13 – 15)</td><td>13 (9 – 16)</td><td>11 – 16</td><td><b>8 – 10</b></td><td>10</td><td>9</td><td>10</td><td>-</td></tr><tr><td>r3</td><td><b>24</b> (20 – 28)</td><td>23 (20 – 25)</td><td>23</td><td>28 (25 – 30)</td><td>25 (24 – 25)</td><td>25 (18 – 32)</td><td>23 – 30</td><td><b>20 – 23</b></td><td>20</td><td>17</td><td>23</td><td>-</td></tr><tr><td>R1</td><td><b>23</b> (20 – 25)</td><td>21 (18 – 22)</td><td>19</td><td>26 (23 – 27)</td><td>23 (22 – 23)</td><td>23 (17 – 29)</td><td>20 – 28</td><td><b>20 – 23</b></td><td>19</td><td>17</td><td>23</td><td>-</td></tr><tr><td>s4</td><td><b>32</b> (28 – 38)</td><td>35 (30 – 39)</td><td>34</td><td>41 (36 – 45)</td><td>32 (30 – 38)</td><td>35 (27 – 44)</td><td>29 – 44</td><td><b>23 – 28</b></td><td>29</td><td>23</td><td>35</td><td>-</td></tr><tr><td>S2</td><td><b>38</b> (30 – 50)</td><td>36 (32 – 38)</td><td>38</td><td>46 (45 – 47)</td><td>40 (38 – 45)</td><td>40 (30 – 48)</td><td>33 – 47</td><td><b>33 – 35</b></td><td>34</td><td>27</td><td>35</td><td>-</td></tr><tr><td>S4</td><td><b>38</b> (33 – 43)</td><td>34 (31 – 37)</td><td>33</td><td>39 (36 – 42)</td><td>37 (35 – 38)</td><td>37 (27 – 49)</td><td>30 – 45</td><td><b>33 – 38</b></td><td>32</td><td>29</td><td>33</td><td>-</td></tr><tr><td>S5</td><td><b>27</b> (23 – 33)</td><td>28 (25 – 30)</td><td>25</td><td>32 (29 – 35)</td><td>31 (30 – 33)</td><td>32 (22 – 42)</td><td>26 – 39</td><td><b>25</b></td><td>22</td><td>26</td><td>28</td><td>-</td></tr><tr><td>z2</td><td><b>26</b> (20 – 33)</td><td>25 (20 – 30)</td><td>24</td><td>34 (32 – 35)</td><td>25 (23 – 28)</td><td>29 (20 – 29)</td><td>24 – 37</td><td><b>18 – 25</b></td><td>26</td><td>17</td><td>20</td><td>-</td></tr><tr><td>z4</td><td><b>25</b> (20 – 33)</td><td>28 (25 – 30)</td><td>25</td><td>34 (32 – 37)</td><td>28 (25 – 33)</td><td>29 (20 – 37)</td><td>24 – 37</td><td><b>18 – 23</b></td><td>23</td><td>17</td><td>25</td><td>-</td></tr><tr><td>z5</td><td><b>20</b> (15 – 23)</td><td>21 (19 – 23)</td><td>20</td><td>26 (23 – 27)</td><td>24 (23 – 28)</td><td>22 (17 – 27)</td><td>21 – 29</td><td><b>15</b></td><td>19</td><td>15</td><td>20</td><td>-</td></tr><tr><td>Z1</td><td><b>30</b> (23 – 35)</td><td>30 (28 – 32)</td><td>29</td><td>37 (34 – 40)</td><td>36 (33 – 40)</td><td>32 (20 – 41)</td><td>24 – 42</td><td><b>23 – 25</b></td><td>28</td><td>22</td><td>28</td><td>-</td></tr><tr><td>Z4</td><td><b>51</b> (48 – 55)</td><td>45 (40 – 48)</td><td>48</td><td>53 (51 – 55)</td><td>50 (48 – 53)</td><td>51 (42 – 59)</td><td>45 – 61</td><td><b>45 – 50</b></td><td>43</td><td>43</td><td>50</td><td>38 – 51</td></tr><tr><td>Z5</td><td><b>71</b> (65 – 75)</td><td>65 (62 – 67)</td><td>67</td><td>70 (65 – 75)</td><td>65 (55 – 73)</td><td>70 (59 – 97)</td><td>62 – 83</td><td><b>53 – 58</b></td><td>45</td><td>49</td><td>53</td><td>48 – 61</td></tr><tr><td>st1-st1</td><td><b>53</b> (50 – 58)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>50 (45 – 58)</td><td>-</td><td><b>48</b></td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st2-st2</td><td><b>63</b> (60 – 68)</td><td>57 (53 – 60)</td><td>-</td><td>63 (59 – 65)</td><td>62 (53 – 68)</td><td>60 (55 – 69)</td><td>-</td><td><b>53 – 55</b></td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st3-st3</td><td><b>73</b> (68 – 80)</td><td>-</td><td>70</td><td>-</td><td>-</td><td>71 (63 – 82)</td><td>-</td><td><b>55 – 58</b></td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st1-st3 ♀ / st1-st5 ♂</td><td><b>67</b> (63 – 70)</td><td>60 (58 – 62)</td><td>65</td><td>62 (59 – 65)</td><td>63 (53 – 68)</td><td>66 (60 – 75)</td><td>-</td><td><b>113 – 115</b></td><td>-</td><td>-</td><td>-</td><td>111 – 131</td></tr><tr><td>st4-st4</td><td><b>76</b> (65 – 88)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>86 (58 – 143)</td><td>-</td><td><b>48</b></td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st5-st5</td><td><b>67</b> (65 – 70)</td><td>62 (58 – 65)</td><td>72</td><td>60 (58 – 62)</td><td>67 (63 – 73)</td><td>69 (41 – 79)</td><td>-</td><td><b>40 – 43</b></td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Lisl</td><td><b>28</b> (23 – 30)</td><td>-</td><td>30</td><td>-</td><td>-</td><td>30 (20 – 36)</td><td>-</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Lisw</td><td><b>5</b> (3 – 5)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>5 (4 – 8)</td><td>-</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Sisl</td><td><b>12</b> (10 – 15)</td><td>-</td><td>13</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Vsl</td><td><b>123</b> (110 – 135)</td><td>118 (105 – 125)</td><td>119</td><td>130 (117 – 135)</td><td>119 (113 – 127)</td><td>117 (99 – 134)</td><td>110 – 137</td><td><b>113</b></td><td>155</td><td>-</td><td>113</td><td>105 – 122</td></tr><tr><td>Vsw ZV2</td><td><b>101</b> (95 – 108)</td><td>98 (88 – 105)</td><td>107</td><td>102 (95 – 105)</td><td>96 (95 – 105)</td><td>104 (88 – 102)</td><td>94 – 119</td><td><b>80 – 85</b></td><td>150</td><td>-</td><td>163</td><td>138 – 170</td></tr><tr><td>Vsw anus</td><td><b>84</b> (73 – 93)</td><td>68 (63 – 75)</td><td>-</td><td>66 (65 – 67)</td><td>-</td><td>73 (60 – 87)</td><td>-</td><td><b>23 – 25</b></td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>JV5</td><td><b>48</b> (43 – 53)</td><td>47 (45 – 50)</td><td>49</td><td>49 (48 – 50)</td><td>-</td><td>53 (40 – 70)</td><td>44 – 64</td><td><b>33 – 40</b></td><td>-</td><td>31</td><td>-</td><td>30 – 44</td></tr><tr><td>StIV</td><td><b>53</b> (45 – 58)</td><td>50 (45 – 54)</td><td>53</td><td>54 (51 – 60)</td><td>52 (48 – 60)</td><td>49 (30 – 62)</td><td>47 – 58</td><td><b>43</b></td><td>37</td><td>38</td><td>43</td><td>39 – 46</td></tr><tr><td>Scl</td><td><b>11</b> (8 – 15)</td><td>12</td><td>12</td><td>11 (10 – 13)</td><td>10 (9 – 10)</td><td>5 – 10</td><td>7 – 15</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Scw</td><td><b>11</b> (10 – 15)</td><td>-</td><td>14</td><td>-</td><td>-</td><td>-</td><td>8 – 18</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Fdl</td><td><b>29</b> (25 – 33)</td><td>26</td><td>27</td><td>30 (29 – 31)</td><td>-</td><td>28 – 35</td><td>23 – 26</td><td><b>20</b></td><td>-</td><td>-</td><td>-</td><td>19 – 21</td></tr><tr><td>No teeth Fd</td><td><b>3</b></td><td>3</td><td>5</td><td>2</td><td>3</td><td>-</td><td>4 – 5</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Mdl</td><td><b>27</b> (25 – 30)</td><td>25</td><td>29</td><td>25 (24 – 26)</td><td>-</td><td>23 – 30</td><td>26 – 31</td><td><b>20</b></td><td>-</td><td>-</td><td>-</td><td>20 – 23</td></tr><tr><td>No teeth Md</td><td><b>2</b></td><td>1</td><td>2</td><td>2</td><td>2</td><td>-</td><td>3</td><td colspan="5" style="text-align: center;">Not applicable</td></tr><tr><td>Shaft</td><td colspan="6">Not applicable</td><td><b>18</b></td><td>-</td><td>-</td><td>-</td><td>15 – 17</td></tr></tbody></table>

Sources of measurements – For ♀♀: Argentina: Guanilo *et al.* (2008b); Australia: Schicha (1987) identified as *N. wearnei*, synonymized by Tixier *et al.* (2014); Peru: Guanilo *et al.* (2008a); Senegal: Kade *et al.* (2011); VCW 1 (Various countries of the world: Brasil, Chile, France, Greece, Italia, Japan, Spain, Tunisia, California): Tixier *et al.* (2008); VCW2 (Australia: 28♀; Canada: 10♀; Chile: 7♀; Guatemala: 1♀; Kenya: 8♀♀; South Korea: 5♀♀; USA: 40♀♀): Beaulieu & Beard (2018). For ♂♂: Argentina: Guanilo *et al.* (2008b); Japan (identified as *N. chilenensis*, synonymized by McMurtry & Badii 1989): Ehara (1964); Peru: Guanilo *et al.* (2008a); VCW (Australia: 5♂♂; Canada: 6♂♂; Chile: 1♂; Kenya: 3♂♂; South Korea: 4♂♂; USA: 15♂♂): Beaulieu & Beard (2018); -: not provided.
**Remarks:** character measurements of the 24 females collected in La Réunion (Table 5) agree very well with those obtained from females of *N. barreti* or *N. recifensis* from Brazil and females of *N. houstoni* from Australia described previously (Gondim Jr. and Moraes 2001, Schicha 1987, Furtado *et al.* 2005). We consider so far that our specimens can be anyone of the three species and that examination of the specimens collected in this study can lead to anyone of the three species. Consequently, the morphometrics strongly suggest synonymy.

There are however some discrepancies between our measurements and observations and previous descriptions of the three species. In the three descriptions:

* Dorsal shield is reticulated in the description of *N. barreti* in the anterior lateral margins and starting to the back of *s4* and on all the posterior part of the dorsal shield except the center;

* VA shield has straight slightly convex margins in Schicha 1987 and not in the two other species as a concave part exists in this margin just after *ZV2* position;

* *Z4* and *Z5* are progressively tapered in drawings of *N. barreti* and *Z5* seems longer (Furtado *et al.* 2005) vs for other species they are blunt/rounded apically (or more regularly parallel-sided);

* Macrosetae *SgeIV*, *StiIV* and *StIV* are mentioned as setaceous for *N. recifensis* (Moraes and Gondim Jr 2001) but seem slightly knobbed on illustrations of the species. *SgeIV* and *StIV* are slightly knobbed for *N. barreti* and *N. houstoni* and *StiIV* is not mentioned for *N. houstoni*;

* two teeth on the fixed digit and no tooth on the movable digit are mentioned for *N. barreti* and *N. recifensis* but three teeth for fixed digit and one recurved tooth for the movable digit are mentioned for *N. houstoni*;

* Spermathecae of *N. barreti* and *N. recifensis* are mentioned as trumpet-shaped but as bell-shaped for *N. houstoni*;

* Setae *JV5* seem longer in *N. barreti* description and shorter in those of *N. houstoni* and *N. recifensis*;

* 6 poroids around genital/ventricular shield are drawn for *N. recifensis* but only 4 for *N. barreti* and 0 for *N. houstoni*;

* Occurrence of *JV3* is mentioned in the text of description of *N. barreti* but not illustrated (Furtado *et al.* 2005). These setae are not indicated for description of *N. houstoni* and *N. recifensis*.

* Chaetotactic formulae are not precised in the description of *N. houstoni* but mentioned only for genu II for *N. barreti* and for genua II and III for *N. recifensis*.

Our examination of the type material for *N. barreti* (one paratype ♀), *N. houstoni* (the single specimen found, the ♀ holotype) and *N. recifensis* (the ♀ holotype, one ♀ paratype and seven additional ♀) shows:

* dorsal shields of the three species present exactly the same reticulation as drawn in Furtado *et al.* (2005) for the description of *N. barreti*;

* VA shield present only a slight concavity after *ZV2* position in *N. houstoni*;

* *Z4* and *Z5* are progressively tapered regularly parallel-sided in the three species and *Z5* is of the same length (see table 8);

* Macrosetae *SgeIV*, *StiIV* and *StIV* are: rounded apically for *SgeIV*, pointed apically for *StiIV* and slightly knobbed for *StIV* for the three species;
* Three teeth on fixed digit (two strong anterior and one small tooth posterior to pilus dentilis) and one small recurved tooth in the anterior part of the movable digit for the three species, just as drawn by Schicha (1987);

* Spermathecae of *N. barreti* and *N. recifensis* are drawn exactly in the same way in original descriptions (Schicha 1987; Moraes and Gondim Jr. 2001; Furtado *et al.* 2005) and are identical in the three species after our examination (and “bell-shaped” seems more appropriate for the shape description);

* Setae *JV5* are of similar length for the three species (see table 5);

* 6 poroids are present around genital/ventrianal shield for the three species;

* The mention of *JV3* was an error in the text of the description of *N. barreti*. This seta is not present in the venter of that species and also not present in the two others;

* Chaetotactic formulae are identical for the three species: Genu II: 2-2/0 - 2/0-1 (seven setae); Genu III: 1-2/1 - 2/0-1 (seven setae).

Considering all these information, we can conclude that the three species are synonyms. Consequently, our specimens are identified as the first species described among the three species, *N. houstoni*. The valid species name is thus *Neoseiulus houstoni* (Schicha). Previous specimens collected in La Réunion Island and identified as *N. recifensis* are renamed *N. houstoni*. Consequently, this is the first report of that species in La Réunion Island.

The male (for the three species) being unknown, it is herein described for the first time, based on La Réunion specimens.

## Description of the adult male of *Neoseiulus houstoni* (Schicha)

n = 8 (Figs 1a-c)

**Diagnosis** — The following combination of characters indicated below in the description of the male of this species is quite similar to a lot of species of *Neoseiulus* belonging to the *cucumeris* and *paraki* species subgroups of the *cucumeris* species group. 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 the level between *j1* and *j3*, a limited reticulation compared to several species of these two subgroups, some dorsal setae length, especially *s4*, *S2*, *Z4* and *JV5* approximately of the same length (30 – 35) and longest setae after *Z5*, a spermatodactyl with a terminal part recurved as an open U (hook-like), no additional macrosetae on other legs than leg IV compared to several species of these two subgroups that have macrosetae on leg III, sometimes II, only three pairs of preanal setae instead of 4 pairs in males of several species of these subgroups, one pair of crateriform *gv3* very close to setae *JV2*.

**Dorsum** — (Fig. 1a). Dorsal shield fused with the peritremal shield at the level of *j1* position, with slight reticulations all around the edge and in the posterior part of the dorsal shield, 200 (183 – 213) long and 129 (115 – 140) wide, with six pairs of solenostomes (*gd* 1, 2, 5, 6, 8, 9). The dorsal shield bears 17 pairs of dorsal setae and 2 pairs of sub-lateral setae on the dorsal shield: *j1* not visible, *j3* 28, *j4* 13, *j5* 15, *j6* 18, *J2* 15, *J5* 5, *z2* 25, *z4* 28, *z5* 18, *Z1* 15, *Z4* 35, *Z5* 40, *s4* 38, *S2* 33, *S4* 18, *S5* 15, *r3* 20, *R1* 20. All setae smooth except *Z4* and *Z5* serrated.

**Peritreme** — (Fig. 1a). Extending between *j1* and *j3*. Peritremal shield fused with dorsal shield.

**Venter** — (Fig. 1b). Sternal shield smooth. Distances between *st1* – *st1* 43, *st2* – *st2* 50, *st3* – *st3* 53, *st1* – *st5* 98, *st4* – *st4* 43, *st5* – *st5* 33. Ventrianal shield with three pairs of pre-anal setae, *JV1*, *JV2*, and *ZV2*, and one pair crateriform *gv3*, mesad of the VAS, just a bit after the insertion line of setae *JV2* but very close from these setae. Three pairs of poroids discernible on specimen ventrianal shields examined. Soft cuticle surrounding ventrianal shield with one
<table><caption>**Table 5** Character measurements of adult females of Neoseiulus sp. collected in this study with those of N. barreti, N. houstoni and N. recifensis (species/localities followed by the number of specimens measured).</caption><thead><tr><th>Characters</th><th>La Réunion<br>24, this study</th><th>Neoseiulus barreti 2<br>Brazil<br>2, remeasured here*</th><th>N. houstoni 1<br>Australia<br>1</th><th>N. houstoni 2<br>Australia<br>1, remeasured here*</th><th>N. recifensis 1<br>Brazil<br>8</th><th>N. recifensis 2<br>Brazil<br>9*</th></tr></thead><tbody><tr><td>Dsl</td><td><strong>291</strong> (264 – 313)</td><td>295 – 298</td><td>285</td><td>288</td><td>279 (268 – 291)</td><td>290 (268 – 305)</td></tr><tr><td>Dsw</td><td><strong>150</strong> (138 – 165)</td><td>148 – 160</td><td>124</td><td>140</td><td>143 (142 – 145)</td><td>141 (113 – 150)</td></tr><tr><td>Peritreme reaching</td><td>j1</td><td>j1</td><td>j1</td><td>j1</td><td>j1</td><td>j1</td></tr><tr><td>Solenostomes</td><td>gd 1, 2, 5, 6, 8, 9</td><td>gd 1, 2, 5, 6, 8, 9</td><td>gd 1, 2, 5, 6, 8, 9</td><td>gd 1, 2, 5, 6, 8, 9</td><td>gd 1, 2, 5, 6, 8, 9</td><td>gd 1, 2, 5, 6, 8, 9</td></tr><tr><td>j1</td><td><strong>22</strong> (20 – 25)</td><td>23 – 25</td><td>19</td><td>18</td><td>21 (19 – 22)</td><td>21 (20 – 23)</td></tr><tr><td>j3</td><td><strong>34</strong> (28 – 38)</td><td>35 – 38</td><td>31</td><td>30</td><td>33 (32 – 35)</td><td>34 (28 – 38)</td></tr><tr><td>j4</td><td><strong>17</strong> (15 – 20)</td><td>15 – 18</td><td>13</td><td>15</td><td>16 (15 – 18)</td><td>17 (15 – 18)</td></tr><tr><td>j5</td><td><strong>17</strong> (15 – 20)</td><td>17</td><td>15</td><td>15</td><td>16 (15 – 18)</td><td>18 (15 – 20)</td></tr><tr><td>j6</td><td><strong>21</strong> (18 – 23)</td><td>20 – 23</td><td>17</td><td>18</td><td>20 (19 – 23)</td><td>21 (18 – 25)</td></tr><tr><td>J2</td><td><strong>24</strong> (20 – 28)</td><td>25</td><td>19</td><td>20</td><td>22 (21 – 24)</td><td>24 (20 – 28)</td></tr><tr><td>J5</td><td><strong>7</strong> (5 – 9)</td><td>7</td><td>8</td><td>7</td><td>7 (6 – 7)</td><td>8</td></tr><tr><td>r3</td><td><strong>29</strong> (25 – 33)</td><td>30</td><td>24</td><td>28</td><td>31 (30 – 34)</td><td>33 (28 – 35)</td></tr><tr><td>R1</td><td><strong>28</strong> (25 – 32)</td><td>30 – 33</td><td>29</td><td>30</td><td>28 (26 – 30)</td><td>31 (30 – 35)</td></tr><tr><td>s4</td><td><strong>43</strong> (40 – 48)</td><td>45 – 48</td><td>40</td><td>43</td><td>45 (43 – 47)</td><td>46 (43 – 48)</td></tr><tr><td>S2</td><td><strong>43</strong> (40 – 45)</td><td>43 – 48</td><td>40</td><td>43</td><td>42 (41 – 44)</td><td>43 (40 – 45)</td></tr><tr><td>S4</td><td><strong>23</strong> (20 – 25)</td><td>23 – 25</td><td>24</td><td>25</td><td>21 (21 – 24)</td><td>24 (23 – 25)</td></tr><tr><td>S5</td><td><strong>22</strong> (20 – 25)</td><td>23</td><td>22</td><td>20</td><td>21 (19 – 23)</td><td>22 (20 – 25)</td></tr><tr><td>z2</td><td><strong>33</strong> (30 – 38)</td><td>33</td><td>29</td><td>28</td><td>33 (30 – 35)</td><td>34 (28 – 38)</td></tr><tr><td>z4</td><td><strong>32</strong> (25 – 36)</td><td>33 – 38</td><td>31</td><td>30</td><td>34 (32 – 36)</td><td>34 (30 – 38)</td></tr><tr><td>z5</td><td><strong>18</strong> (18 – 20)</td><td>18</td><td>14</td><td>15</td><td>17 (16 – 19)</td><td>18 (15 – 20)</td></tr><tr><td>Z1</td><td><strong>26</strong> (23 – 30)</td><td>25 – 30</td><td>23</td><td>23</td><td>24 (22 – 26)</td><td>25 (20 – 28)</td></tr><tr><td>Z4 serrated</td><td><strong>46</strong> (43 – 53) ser.</td><td>45 – 50 ser.</td><td>41 ser.</td><td>45 ser.</td><td>47 (44 – 49) ser.</td><td>48 (45 – 50) ser.</td></tr><tr><td>Z5 serrated</td><td><strong>49</strong> (45 – 53) ser.</td><td>50 – 55 ser.</td><td>47 ser.</td><td>48 ser.</td><td>50 (47 – 53) ser.</td><td>52 (43 – 55) ser.</td></tr><tr><td>st1-st1</td><td><strong>50</strong> (48 – 53)</td><td><strong>54</strong>*</td><td>-</td><td>48</td><td>-</td><td>50 (43 – 53)</td></tr><tr><td>st2-st2</td><td><strong>60</strong> (55 – 73)</td><td>63</td><td>-</td><td>58</td><td>57 (54 – 60)</td><td>58 (55 – 63)</td></tr><tr><td>st3-st3</td><td><strong>71</strong> (65 – 75)</td><td><strong>75</strong>*</td><td>69</td><td>68</td><td>-</td><td>67 (58 – 73)</td></tr><tr><td>st1-st3</td><td><strong>52</strong> (48 – 55)</td><td>53 – 55</td><td>57</td><td>54</td><td>53 (49 – 53)</td><td>56 (50 – 60)</td></tr><tr><td>st4-st4</td><td><strong>79</strong> (68 – 93)</td><td><strong>88</strong>*</td><td>-</td><td>60</td><td>-</td><td>78 (48 – 93)</td></tr><tr><td>Gensl</td><td><strong>99</strong> (93 – 110)</td><td>100</td><td>-</td><td>100</td><td>-</td><td>97 (93 – 100)</td></tr><tr><td>Gensw st5</td><td><strong>61</strong> (55 – 65)</td><td>63</td><td>-</td><td>60</td><td>-</td><td>62 (60 – 63)</td></tr><tr><td>Gensw post. corn.</td><td><strong>69</strong> (65 – 75)</td><td>68</td><td>-</td><td>68</td><td>-</td><td>67 (63 – 70)</td></tr><tr><td>st5-st5</td><td><strong>57</strong> (53 – 60)</td><td>60</td><td>-</td><td>58</td><td>58 (57 – 60)</td><td>60 (58 – 60)</td></tr><tr><td>Lisl</td><td><strong>18</strong> (15 – 28)</td><td>17 – 22</td><td>19</td><td>20</td><td>-</td><td>20 (20 – 23)</td></tr><tr><td>Lisw</td><td><strong>4</strong> (3 – 5)</td><td>5</td><td>-</td><td>4</td><td>-</td><td>5 (4 – 5)</td></tr><tr><td>Sisl</td><td><strong>7</strong> (5 – 13)</td><td><strong>9</strong>*</td><td>8</td><td>8</td><td>-</td><td>9 (8 – 10)</td></tr><tr><td>Vsl</td><td><strong>92</strong> (85 – 98)</td><td>95 – 103</td><td>57</td><td>95</td><td>92 (89 – 95)</td><td>97 (88 – 100)</td></tr><tr><td>Vsw ZV2</td><td><strong>68</strong> (60 – 75)</td><td>73</td><td>69</td><td>73</td><td>69 (68 – 72)</td><td>76 (70 – 83)</td></tr><tr><td>Vsw anus</td><td><strong>61</strong> (58 – 65)</td><td>63 – 68</td><td>-</td><td>58</td><td>62 (59 – 64)</td><td>66 (62 – 68)</td></tr><tr><td>gv3 distance</td><td><strong>18</strong> (15 – 23)</td><td>18</td><td>-</td><td>18</td><td>-</td><td>18 (15 – 20)</td></tr><tr><td>JV5 ser. or not</td><td><strong>41</strong> (38 – 50) not</td><td>35 – 42 not</td><td>36 not</td><td>38 not</td><td>-</td><td>42 (30 – 48) not</td></tr><tr><td>SgeIV</td><td><strong>21</strong> (18 – 23) knob.</td><td>15 – 20 knob.</td><td>20 knob.</td><td>23 knob.</td><td>21 (19 – 21)</td><td>21 (20 – 23) knob.</td></tr><tr><td>StiIV</td><td><strong>15</strong> (13 – 18)</td><td>15 – 17</td><td>-</td><td>15</td><td>17 (16 – 19)</td><td>19 (18 – 20)</td></tr><tr><td>StIV</td><td><strong>29</strong> (25 – 33)</td><td>27 – 32</td><td>25 bl.</td><td>25 bl.</td><td>30 (27 – 32)</td><td>30 (28 – 33)</td></tr><tr><td>Scl</td><td><strong>22</strong> (13 – 40)</td><td><strong>13</strong>*</td><td>11</td><td>15</td><td>13 (12 – 14)</td><td>16 (13 – 18)</td></tr><tr><td>Calyx l.</td><td><strong>12</strong> (10 – 13)</td><td><strong>10</strong>*</td><td>-</td><td>10</td><td>-</td><td>12 (10 – 13)</td></tr><tr><td>Scw</td><td><strong>13</strong> (5 – 25)</td><td>12</td><td>8</td><td>10</td><td>-</td><td>10</td></tr><tr><td>Fdl, No teeth</td><td><strong>25</strong> (25 – 30), <strong>2+1</strong></td><td><strong>30, 2+1</strong>*</td><td>29 – 31, 2 + 1</td><td>30, 2 + 1</td><td>27 (26 – 27), 2</td><td>26 (20 – 28), 2 + 1</td></tr><tr><td>Mdl, No teeth</td><td><strong>24</strong> (23 – 28), <strong>1</strong></td><td><strong>30, 1</strong>*</td><td>29 – 31, 1</td><td>28, 1</td><td>28 (26 – 29), 0</td><td>25 (21 – 28), 1</td></tr></tbody></table>

Sources of measurements – Neoseiulus barreti Brazil: Furtado et al. (2005) (measured again by the senior author with * complementary measurements); N. houstoni Australia 1 (holotype): Schicha (1987); N. houstoni Australia 2: *measurements by the senior author; N. recifensis Brazil 1 (holo- and paratypes): Gondim Jr & Moraes (2001); N. recifensis Brazil 2 (holotype and additional material): *measurements by the senior author; -: not provided.
<figure><img src="image_7.png" /><figcaption>**Figure 1** Male of *Neoseiulus houstoni* Schicha: a – Dorsal shield and peritreme; b – Ventral shields; c – Spermatodactyl.</figcaption></figure>

pair of setae (JV5); ventrianal shield 83 long, 120 wide at anterior corners and 60 wide at level of paranal setae. JV5 smooth, 35 long. A pair of lyrifissures near JV5.

**Chelicera** — Fixed digit 20 long, no tooth discerned and movable digit 18 long with no tooth discerned because chelicerae are dorsoventrally oriented. Spermatodactyl with an open U-shaped foot, shaft (Fig. 1c) 13 long.

**Legs** — Legs IV with three macrosetae like in the female: SgeIV slightly knobbed 18, StiIV pointed 15, StIV very slightly knobbed 23. Chaetotactic formula of genu II and III similar to that of females.

**Specimens examined** — 8 ♂♂ collected, 8 ♂♂ measured. Langevin – Jacqueline Waterfall (aasl 5 m, Long 55°64'40" E, Lat 21°38'69" S), 2 ♂♂ on C. equisetifolia, 19/7/2017; Saint-Pierre – Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29'18" E, Lat 21°19'25" S), 2 ♂ on C. retusa, 27/2/2017; 4 ♂♂ + 1 im. on P. hysterophorus, 30/3/2017.

**Type material** — Eight paratype males in total. Three paratype males deposited in Montpellier SupAgro – INRA Acarology collection, Montpellier and five paratype males deposited in Bassin-Plat CIRAD Research station collection in La Réunion.

**Remarks** — This combination of characters of the male of this species are not unique and do not allow to distinguish it from all males of other species of *Neoseiulus* belonging to the *cucumeris* and *paraki* species sub-groups without collected females of the species. The male peritreme is shorter than the female peritreme. Other characters are very similar.
## Neoseiulus longispinosus (Evans)

*Typhlodromus longispinosus* Evans 1952: 413; Evans 1953: 465; Womersley 1954: 177; Ehara 1958: 55.

*Typhlodromus (Amblyseius) longispinosus*, Chant 1959: 74.

*Amblyseius longispinosus*, Corpuz & Rimando 1966: 129; Schicha 1975: 103.

*Neoseiulus longispinosus*, Moraes et al. 1986: 85; 2000: 245; Chant & McMurtry 2003a: 37; Moraes et al. 2004a: 129; Chant and McMurtry 2007: 29.

This species belongs to the *barkeri* species group like *N. barkeri* (see above). It belongs to the *womersleyi* species subgroup as the calyx is markedly constricted at the junction with the atrium, the atrium is deeply forked at the junction with the major duct, and the major duct, atrium and calyx are not of the same width (Chant and McMurtry 2003a).

This species is distributed in many countries of the world, mainly in tropical areas (Moraes et al. 2000; Mailloux et al. 2010; Kreiter et al. 2013, 2018 a, c; Demite et al. 2019). It was found rarely in surveys made in Guadeloupe, Martinique and La Réunion except in studies on companion plants in citrus orchards (Mailloux et al. 2010; Kreiter et al. 2013, 2018c; Le Bellec et al., unpub. data). This species seems actually to be more common on weeds with populations of tetranychid mites. *Neoseiulus longispinosus*, a type II phytoseiid predatory mite, as is *N. californicus* (McMurtry et al. 2013), has received increasing attention in Asia for the control of different spider mites (of *Eutetranychus*, *Oligonychus*, and *Tetranychus*) since 2010 (Nusartlert et al. 2011). The feeding, development, predation, cannibalism, intra-guild predation and behaviour have thus been extensively studied by several authors (see for example Luong et al. 2017) for pest control purposes. *Neoseiulus longispinosus* is well-known as a BCA sell in several countries in the world for the management of spider mites. The recent results of Huyen et al. (2017) show that at least in controlled laboratory conditions, *N. longispinosus* is a potential biological control agent against the citrus red spider mite *P. citri*.

This is the first record of this species for La Réunion Island.

**Specimens examined:** 37 ♀♀ + 1 ♂ + 4 im. in total, 18 ♀♀ + 1 ♂ measured. Saint-Paul - Savannah (aasl 61 m, Long 55°29'43" E, Lat 21°20'41" S), 8 ♀♀ in flowers of *Phaseolus vulgaris* L. 28/07/2015; Saint-Pierre - Eastern entrance of the city (aasl 61 m, Long 55°29'43" E, Lat 21°20'41" S), 1 ♀ on *Ricinus communis* L., 16/12/2015; Ravine des Cabris - Ligne des Bambous, Lassay (aasl 221 m, Long 55°29'38" E, Lat 21°17'17" S), 1 ♀ on *Solanum torvum* Swartz, 2/12/2016; 1 ♀ + 1 ♂ + 1 im. on *Mirabilis jalapa* L., 4/12/2016; Saint-Pierre - Bassin Martin, ARMEFLHOR Station (aasl 450 m, Long 55°31'9" E, Lat 21°18'14" S), 11 ♀♀ on *Fragaria* sp. + *A. viridis*, 18/1/2017; Saint-Gilles - Pépinières du Théâtre (aasl 70 m, Long 56°13'58" E, Lat 21°2'50" S), 1 ♀ on *A. viridis*, 14/2/2017; Saint-Pierre - Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29'18" E, Lat 21°19'25" S), 1 ♀ in plot H, 16/12/2016; 1 ♀ on *B. pilosa*, 2 ♀♀ + 1 im. on *M. repens*, 1 ♀ on *Ipomoea obscura* (L.), 1 ♀ on *Terannus labialis* (L.f.), 12/2/2017; 1 ♀ + 2 im. on *I. obscura*, 1 ♀ on *Digitaria radicosa* (J. Presl), 30/3/2017; 6 ♀♀ in plots CC, M, HM and H, 3 and 6/04/2017.

**Remarks:** measurements of specimens of La Réunion females and males (Table 6) overlap with those obtained for populations of various countries. Measurements are slightly greater than those obtained on specimens of F.C.I. except for setae *j4*, *J2*, *z5*, *StIV*. On Comoros specimens, setae are longer except sternal shield length (*st1-st3*), inguinal sigilla (metapodal plates) and of macrosetae of basitarsus IV.

## Neoseiulus lula (Pritchard & Baker)

*Amblyseius (Amblyseius) lula* Pritchard & Baker, 1962: 239.

*Neoseiulus lula*, Schicha 1981b: 212; Moraes et al. 1986: 87; Chant & McMurtry 2003a: 27; Moraes et al. 2004a: 130; Chant & McMurtry 2007: 29.

*Amblyseius (Amblyseius) insignitus* van der Merwe, 1968: 138 (synonymy according to Ueckermann & Loots, 1988).
Like *N. baraki* (see above), *N. lula* belongs to the *paspalivorus* species group (Chant and McMurtry 2003a).

The biology of this species remains unknown.

It is distributed in several countries and islands of sub-Saharan Africa but also in Cuba (Moraes *et al.* 2004a).

This is the first record of this species for La Réunion Island.

**Specimens examined:** a single ♀, measured. Saint-Pierre – Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29'18” E, Lat 21°19'25” S), 1 ♀ on *Chrysopogon zizanioides* (L.), 2/3/2017.

**Remarks:** the measurements of the single female of *Neoseiulus lula* (Table 7) fit well measurements of specimens of the original description from Central Africa (Schicha 1981b) and those of specimens from Africa (Zannou *et al.* 2006), except for *j6* and *S2* which are smaller (>20 %) in the single specimen of La Réunion. Specimens from Africa (Zannou *et al.* 2006) have longer *Z4* but a reduced ventral shield. Specimens from South Africa and Madagascar

<table><caption>**Table 6** Character measurements of adult females and one adult male of *Neoseiulus longispinosus* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="2">Characters</th><th colspan="7">♀</th><th colspan="3">♂</th></tr><tr><th>La Réunion<br>18, this study</th><th>FCI</th><th>Comoros</th><th>Martinique</th><th>Sri Lanka</th><th>Taiwan</th><th>Thailand</th><th>Holotype<br>Indonesia</th><th>La Réunion<br>1, this study</th><th>F.C.I.</th><th>Paratype<br>Indonesia</th></tr></thead><tbody><tr><td>Dsl</td><td><strong>320</strong> (278 - 348)</td><td>332 (308 - 398)</td><td>380</td><td>321 (295 - 340)</td><td>321 (313 - 338)</td><td>-</td><td>330 (315 - 340)</td><td>332</td><td><strong>248</strong></td><td>247 (224 - 269)</td><td>231</td></tr><tr><td>Dsw</td><td><strong>183</strong> (150 - 205)</td><td>179 (154 - 200)</td><td>192</td><td>168 (150 - 183)</td><td>187 (175 - 208)</td><td>-</td><td>186 (178 - 195)</td><td>173</td><td><strong>170</strong></td><td>160 (136 - 173)</td><td>138</td></tr><tr><td>j1</td><td><strong>18</strong> (15 - 20)</td><td>18 (16 - 22)</td><td>20</td><td>16 (13 - 18)</td><td>18 (17 - 19)</td><td>17</td><td>19 (18 - 21)</td><td>14</td><td><strong>15</strong></td><td>14 (13 - 16)</td><td>9</td></tr><tr><td>j3</td><td><strong>62</strong> (53 - 73)</td><td>59 (52 - 64)</td><td>69</td><td>58 (48 - 63)</td><td>62 (61 - 64)</td><td>58</td><td>61 (56 - 68)</td><td>51</td><td><strong>50</strong></td><td>44 (40 - 48)</td><td>43</td></tr><tr><td>j4</td><td><strong>55</strong> (48 - 60)</td><td>59 (52 - 65)</td><td>70</td><td>56 (50 - 65)</td><td>58 (56 - 60)</td><td>59</td><td>58 (54 - 62)</td><td>49</td><td><strong>45</strong></td><td>43 (42 - 43)</td><td>39</td></tr><tr><td>j5</td><td><strong>66</strong> (58 - 73)</td><td>69 (64 - 75)</td><td>78</td><td>66 (60 - 73)</td><td>70 (69 - 71)</td><td>-</td><td>69 (65 - 72)</td><td>59</td><td><strong>48</strong></td><td>52 (50 - 53)</td><td>43</td></tr><tr><td>j6</td><td><strong>73</strong> (65 - 83)</td><td>72 (68 - 75)</td><td>78</td><td>68 (60 - 83)</td><td>70 (68 - 72)</td><td>-</td><td>71 (68 - 73)</td><td>64</td><td><strong>53</strong></td><td>52 (50 - 53)</td><td>46</td></tr><tr><td>J2</td><td><strong>74</strong> (68 - 83)</td><td>76 (73 - 78)</td><td>88</td><td>76 (68 - 88)</td><td>77 (75 - 79)</td><td>-</td><td>74 (68 - 81)</td><td>66</td><td><strong>55</strong></td><td>56 (53 - 61)</td><td>54</td></tr><tr><td>J5</td><td><strong>8</strong> (6 - 10)</td><td>9 (8 - 11)</td><td>10</td><td>9 (8 - 10)</td><td>8</td><td>-</td><td>9 (7 - 12)</td><td>10</td><td><strong>8</strong></td><td>6 (5 - 6)</td><td>2</td></tr><tr><td>r3</td><td><strong>59</strong> (48 - 78)</td><td>57 (49 - 62)</td><td>75</td><td>54 (45 - 63)</td><td>55 (55 - 56)</td><td>-</td><td>61 (50 - 67)</td><td>54</td><td><strong>35</strong></td><td>32 (24 - 40)</td><td>32</td></tr><tr><td>R1</td><td><strong>61</strong> (54 - 80)</td><td>61 (57 - 65)</td><td>70</td><td>57 (50 - 63)</td><td>60 (59 - 62)</td><td>-</td><td>63 (55 - 70)</td><td>58</td><td><strong>38</strong></td><td>34 (32 - 37)</td><td>32</td></tr><tr><td>s4</td><td><strong>82</strong> (70 - 88)</td><td>77 (73 - 80)</td><td>-</td><td>78 (73 - 88)</td><td>82 (80 - 83)</td><td>72</td><td>81 (76 - 85)</td><td>75</td><td><strong>63</strong></td><td>62 (59 - 66)</td><td>57</td></tr><tr><td>S2</td><td><strong>75</strong> (68 - 85)</td><td>72 (68 - 76)</td><td>88</td><td>69 (63 - 76)</td><td>73 (70 - 79)</td><td>70</td><td>73 (68 - 80)</td><td>67</td><td><strong>55</strong></td><td>52 (50 - 54)</td><td>46</td></tr><tr><td>S4</td><td><strong>56</strong> (43 - 68)</td><td>57 (48 - 76)</td><td>63</td><td>52 (45 - 58)</td><td>59 (57 - 62)</td><td>58</td><td>61 (56 - 68)</td><td>49</td><td><strong>40</strong></td><td>30 (29 - 32)</td><td>22</td></tr><tr><td>S5</td><td><strong>17</strong> (15 - 18)</td><td>16 (14 - 16)</td><td>18</td><td>14 (13 - 15)</td><td>21 (19 - 23)</td><td>17</td><td>20 (18 - 25)</td><td>15</td><td><strong>15</strong></td><td>14 (13 - 16)</td><td>14</td></tr><tr><td>z2</td><td><strong>67</strong> (56 - 75)</td><td>65 (62 - 68)</td><td>75</td><td>64 (58 - 70)</td><td>69 (68 - 70)</td><td>62</td><td>66 (62 - 70)</td><td>58</td><td><strong>48</strong></td><td>44 (35 - 48)</td><td>45</td></tr><tr><td>z4</td><td><strong>73</strong> (65 - 80)</td><td>69 (67 - 73)</td><td>78</td><td>70 (63 - 87)</td><td>73 (73 - 75)</td><td>65</td><td>71 (68 - 75)</td><td>58</td><td><strong>50</strong></td><td>52 (48 - 54)</td><td>49</td></tr><tr><td>z5</td><td><strong>27</strong> (23 - 33)</td><td>35 (32 - 40)</td><td>38</td><td>31 (28 - 35)</td><td>32 (32 - 38)</td><td>-</td><td>31 (25 - 36)</td><td>-</td><td><strong>28</strong></td><td>27 (24 - 32)</td><td>-</td></tr><tr><td>Z1</td><td><strong>75</strong> (65 - 85)</td><td>75 (72 - 80)</td><td>83</td><td>74 (68 - 80)</td><td>77 (76 - 78)</td><td>-</td><td>75 (71 - 78)</td><td>67</td><td><strong>55</strong></td><td>56 (50 - 58)</td><td>46</td></tr><tr><td>Z4</td><td><strong>70</strong> (63 - 75)</td><td>71 (67 - 75)</td><td>78</td><td>69 (63 - 78)</td><td>72 (71 - 73)</td><td>65</td><td>71 (65 - 76)</td><td>68</td><td><strong>53</strong></td><td>55 (48 - 59)</td><td>45</td></tr><tr><td>Z5</td><td><strong>81</strong> (73 - 88)</td><td>80 (78 - 81)</td><td>85</td><td>77 (65 - 80)</td><td>80 (80 - 81)</td><td>70</td><td>78 (72 - 81)</td><td>72</td><td><strong>58</strong></td><td>57 (56 - 59)</td><td>57</td></tr><tr><td>st1-st1</td><td><strong>47</strong> (43 - 50)</td><td>-</td><td>48</td><td>46 (45 - 50)</td><td>-</td><td>-</td><td>-</td><td>-</td><td><strong>43</strong></td><td>-</td><td>-</td></tr><tr><td>st2-st2</td><td><strong>58</strong> (55 - 60)</td><td>55 (49 - 57)</td><td>55</td><td>54 (53 - 55)</td><td>53 (50 - 55)</td><td>-</td><td>59 (57 - 60)</td><td>-</td><td><strong>53</strong></td><td>-</td><td>-</td></tr><tr><td>st3-st3</td><td><strong>71</strong> (68 - 78)</td><td>60 (59 - 62)</td><td>60</td><td>58 (55 - 60)</td><td>-</td><td>-</td><td>-</td><td>77</td><td><strong>53</strong></td><td>-</td><td>-</td></tr><tr><td>st1-st3 ♀</td><td><strong>57</strong> (55 - 60)</td><td>-</td><td>73</td><td>70 (68 - 73)</td><td>55 (53 - 56)</td><td>-</td><td>63 (57 - 85)</td><td>62</td><td><strong>103</strong></td><td>-</td><td>-</td></tr><tr><td>st1-st5 ♀</td><td><strong>71</strong> (63 - 83)</td><td>-</td><td>85</td><td>72 (63 - 88)</td><td>-</td><td>-</td><td>-</td><td>-</td><td><strong>43</strong></td><td>-</td><td>-</td></tr><tr><td>st4-st4</td><td><strong>54</strong> (50 - 58)</td><td>56 (52 - 60)</td><td>58</td><td>53 (50 - 63)</td><td>53 (51 - 54)</td><td>-</td><td>61 (55 - 64)</td><td>-</td><td><strong>35</strong></td><td>-</td><td>-</td></tr><tr><td>st5-st5</td><td><strong>25</strong> (20 - 30)</td><td>-</td><td>23</td><td>28 (23 - 33)</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="3">Not applicable</td></tr><tr><td>Lisl</td><td><strong>3</strong> (3 - 6)</td><td>-</td><td>3</td><td>3</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="3">Not applicable</td></tr><tr><td>Lisw</td><td><strong>12</strong> (10 - 18)</td><td>-</td><td>18</td><td>13 (10 - 15)</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="3">Not applicable</td></tr><tr><td>Sisl</td><td><strong>114</strong> (93 - 125)</td><td>115 (94 - 121)</td><td>125</td><td>111 (103 - 120)</td><td>106 (103 - 111)</td><td>-</td><td>123 (120 - 125)</td><td>97</td><td><strong>100</strong></td><td>106 (99 - 112)</td><td>101</td></tr><tr><td>Vsl</td><td><strong>86</strong> (80 - 95)</td><td>86 (80 - 92)</td><td>90</td><td>84 (75 - 90)</td><td>91 (89 - 93)</td><td>-</td><td>97 (95 - 100)</td><td>87</td><td><strong>130</strong></td><td>135 (125 - 144)</td><td>130</td></tr><tr><td>Vsw ZV2</td><td><strong>69</strong> (67 - 73)</td><td>75 (67 - 83)</td><td>78</td><td>70 (65 - 75)</td><td>75 (73 - 77)</td><td>-</td><td>-</td><td>-</td><td><strong>65</strong></td><td>-</td><td>-</td></tr><tr><td>Vsw anus</td><td><strong>62</strong> (55 - 70)</td><td>-</td><td>73</td><td>60 (55 - 63)</td><td>-</td><td>-</td><td>-</td><td>-</td><td><strong>35</strong></td><td>-</td><td>-</td></tr><tr><td>JV5</td><td><strong>79</strong> (75 - 83)</td><td>80 (75 - 87)</td><td>75</td><td>81 (75 - 88)</td><td>68 (68 - 70)</td><td>74</td><td>74 (72 - 77)</td><td>80 - 87</td><td><strong>60</strong></td><td>66 (62 - 72)</td><td>-</td></tr><tr><td>Scl</td><td><strong>24</strong> (13 - 35)</td><td>28 (25 - 30)</td><td>25</td><td>20 (17 - 25)</td><td>21 (20 - 21)</td><td>17</td><td>19 (17 - 22)</td><td>30</td><td colspan="3">Not applicable</td></tr><tr><td>Scw</td><td><strong>4</strong> (3 - 10)</td><td>-</td><td>5</td><td>5</td><td>-</td><td>-</td><td>-</td><td>4</td><td colspan="3">Not applicable</td></tr><tr><td>Fdl</td><td><strong>24</strong> (19 - 28)</td><td>25 (22 - 27)</td><td>23</td><td>24 (23 - 25)</td><td>22 (21 - 22)</td><td>-</td><td>23 (22 - 25)</td><td>-</td><td><strong>18</strong></td><td>-</td><td>-</td></tr><tr><td>No teeth Fd</td><td><strong>6</strong></td><td>4 - 5</td><td>-</td><td>4</td><td>5</td><td>-</td><td>-</td><td>-</td><td><strong>-</strong></td><td>-</td><td>-</td></tr><tr><td>Mdl</td><td><strong>24</strong> (23 - 30)</td><td>25 (23 - 25)</td><td>23</td><td>24 (23 - 25)</td><td>25 (23 - 25)</td><td>-</td><td>26 (25 - 27)</td><td>-</td><td><strong>18</strong></td><td>-</td><td>-</td></tr><tr><td>No teeth Md</td><td><strong>2</strong></td><td>2</td><td>-</td><td>2</td><td>2</td><td>-</td><td>-</td><td>-</td><td><strong>15</strong></td><td>-</td><td>-</td></tr><tr><td>Shaft</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>12 (11 - 14)</td><td>21</td></tr></tbody></table>

Sources of measurements – For ♀♀: FCI (French Caribbean Islands): Moraes *et al.* (2000); Grande Comore: Kreiter *et al.* (2018b) (a mistake remains in the paper, 1 single female measured instead of 8 as indicated); Martinique: Kreiter *et al.* (2018c); Sri Lanka: Moraes *et al.* (2004b); Taiwan: Tseng (1983); Thailand: Oliveira *et al.* (2012); Holotype Indonesia: Schicha (1975). For ♂♂: FCI (French Caribbean Islands): Moraes *et al.* (2000); Paratype Indonesia: Schicha (1975). -: not provided.
(van der Merwe 1968; Ueckermann and Loots 1988) have in general greater dimensions than those obtained for specimens from La Réunion Island.

## Neoseiulus paspalivorus De Leon

*Typhlodromus paspalivorus* De Leon, 1957: 143.

*Amblyseius paspalivorus*, Schicha 1981b: 210.

*Neoseiulus paspalivorus*, Muma & Denmark 1970: 110; Moraes *et al.* 1986: 92; Chant & McMurtry 2003a: 27; Moraes *et al.* 2004b: 137.

Like *N. baraki* and *N. lula*, *N. paspalivorus* belongs to the *paspalivorus* species group (see above) (Chant and McMurtry 2003a).

*N. paspalivorus* was found only on coconut and on fruits, in association with *A. guerreronis* (Moraes *et al.* 2004b). This species is a promising candidate for the biological control of the coconut eriophyid (Lawson-Balagbo *et al.* 2008).

This is the first record of this species for La Réunion Island.

**Specimens examined:** 15 ♀♀ + 3 ♂♂ in total, 8 ♀♀ measured + 3 ♂♂ measured. Saint-Pierre - Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29'18" E, Lat 21°19'25" S), 1 ♀ on *D. ramularis*, 6 ♀♀ on *M. repens* + 2 ♂♂, 3 ♀♀ + 1 ♂ on *Cynodon dactylon* (L.), 1 ♀ on *Acanthospermum hispidum* DC., 1 ♀ on *L. leucocephala*, 20/2/2017; 3 ♀♀ in plots CC and HM, 3/04/2017.

**Remarks:** measurements of characters of the eight female and 3 male specimens (Table 8) fit well those obtained for populations of other countries, with only slight variations.

## Neoseiulus scapilatus (van der Merwe)

*Amblyseius (Amblyseius) scapilatus* van der Merwe, 1965: 71.

*Amblyseius scapilatus*, Meyer & Rodriguez, 1966: 28.

*Neoseiulus scapilatus*, Moraes *et al.* 1986: 95; McMurtry & Moraes 1991: 26; Chant & McMurtry 2003a: 37; Moraes *et al.* 2004a: 142; Chant & McMurtry 2007: 31; El-Banhawy & Knapp 2011: 12.

Like *N. barkeri* and *N. longispinosus*, *N. scapilatus* belongs to the *barkeri* species group (see above) (Chant and McMurtry 2003a).

Quilici et al. (2000) have collected before this species in La Réunion that is distributed in several countries of sub-Saharan Africa. Exact indications of locations were provided but without any measurements of specimens collected. Measurements of specimens collected during this study are provided in table 9. The biology of this species remains unknown.

**Specimens examined:** 11 ♀♀ + 1 ♂ in total, all measured. Le Tampon - Grand Tampon, Janick Bénard farm (aasl 861 m, Long 55°32'90" E, Lat 21°12'80" S), 1 ♀ on *Lantana camara* L., 9/12/2015; 1 ♀ on *Ageratum conyzoides* (L.) and 1 ♀ + 1 im. on *Raphanus raphanistrum* L.; 24/5/2016; 2 ♀♀ on *Bromus catharticus* Vahl, 1 ♀ on *Conyza sumatrensis* (S.F. Blake), 9/1/2017; Petite Île - Piton Bloc, Yébo Luguy farm (aasl 973 m, Long 55°34'64" E, Lat 21°18'64" S), 3 ♀♀ on *B. catharticus*, 18/10/2016; 1 ♀ + 1 ♂ on *Pteridium aquilinum* (L.), 9/1/2017; Le Tampon - Ligne des 400 (aasl 463 m, Long 55°30'36" E, Lat 21°17'24" S), 1 ♀ on *Ipomoea* sp., 10/1/2017.

**Remarks:** measurements of morphological characters of *N. scapilatus* female specimens from La Réunion (Table 9) are very close from measurements for specimens from neighbouring countries, except for specimens from South Africa that are larger (van der Merwe 1965). La Réunion specimens have slightly shorter macrosetae.

For the male (Table 9), some setae (j3, r3, S4, z5, JV5) or dimensions (st3-st3, st1-st5) are shorter.
<table><caption>**Table 7** Character measurements of an adult female of *Neoseiulus lula* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th>Characters</th><th>La Réunion</th><th>Africa</th><th>Central Africa</th><th>Holotype Congo</th><th>South-Africa Madagascar</th><th>South-Africa</th></tr><tr><td></td><td>1, this study</td><td>3</td><td>1</td><td>1</td><td>2?</td><td>1</td></tr></thead><tbody><tr><td>Dsl</td><td><strong>385</strong></td><td>386 (381 – 393)</td><td>381</td><td>389</td><td>397</td><td>377</td></tr><tr><td>Dsw</td><td><strong>188</strong></td><td>196 (182 – 211)</td><td>157</td><td>191</td><td>195</td><td>204</td></tr><tr><td>j1</td><td><strong>23</strong></td><td>21 (19 – 23)</td><td>21</td><td>19</td><td>24</td><td>21</td></tr><tr><td>j3</td><td><strong>23</strong></td><td>23 (22 – 24)</td><td>23</td><td>22</td><td>24</td><td>21</td></tr><tr><td>j4</td><td><strong>15</strong></td><td>15 (14 – 16)</td><td>15</td><td>14</td><td>16</td><td>12</td></tr><tr><td>j5</td><td><strong>15</strong></td><td>14 (13 – 16)</td><td>14</td><td>14</td><td>16</td><td>13</td></tr><tr><td>j6</td><td><strong>13</strong></td><td>18 (18 – 19)</td><td>17</td><td>17</td><td>16</td><td>17</td></tr><tr><td>J2</td><td><strong>18</strong></td><td>19 (19 – 20)</td><td>20</td><td>18</td><td>21</td><td>18</td></tr><tr><td>J5</td><td><strong>13</strong></td><td>11 (10 – 13)</td><td>14</td><td>13</td><td>16</td><td>11</td></tr><tr><td>r3</td><td><strong>20</strong></td><td>21 (19 – 22)</td><td>21</td><td>24</td><td>24</td><td>20</td></tr><tr><td>R1</td><td><strong>20</strong></td><td>22 (20 – 24)</td><td>21</td><td>21</td><td>24</td><td>20</td></tr><tr><td>s4</td><td><strong>25</strong></td><td>25 (23 – 27)</td><td>25</td><td>25</td><td>27</td><td>22</td></tr><tr><td>S2</td><td><strong>20</strong></td><td>27 (25 – 29)</td><td>27</td><td>25</td><td>29</td><td>25</td></tr><tr><td>S4</td><td><strong>25</strong></td><td>28 (26 – 30)</td><td>26</td><td>28</td><td>29</td><td>24</td></tr><tr><td>S5</td><td><strong>25</strong></td><td>27 (26 – 29)</td><td>28</td><td>25</td><td>29</td><td>22</td></tr><tr><td>z2</td><td><strong>18</strong></td><td>20 (19 – 21)</td><td>19</td><td>19</td><td>21</td><td>19</td></tr><tr><td>z4</td><td><strong>20</strong></td><td>21 (20 – 22)</td><td>19</td><td>20</td><td>21</td><td>17</td></tr><tr><td>z5</td><td><strong>15</strong></td><td>16 (15 – 16)</td><td>15</td><td>16</td><td>16</td><td>14</td></tr><tr><td>Z1</td><td><strong>19</strong></td><td>21 (19 – 22)</td><td>20</td><td>24</td><td>24</td><td>20</td></tr><tr><td>Z4</td><td><strong>28</strong></td><td>35 (30 – 43)</td><td>25</td><td>35</td><td>34</td><td>27</td></tr><tr><td>Z5</td><td><strong>65</strong></td><td>65 (64 – 66)</td><td>65</td><td>65</td><td>70</td><td>62</td></tr><tr><td>st1-st1</td><td><strong>48</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st2-st2</td><td><strong>63</strong></td><td>61 (58 – 64)</td><td>67</td><td>59</td><td>-</td><td>-</td></tr><tr><td>st3-st3</td><td><strong>70</strong></td><td>-</td><td>-</td><td>-</td><td>67</td><td>-</td></tr><tr><td>st1-st3</td><td><strong>75</strong></td><td>77 (77 – 78)</td><td>78</td><td>78</td><td>92</td><td>-</td></tr><tr><td>st4-st4</td><td><strong>68</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st5-st5</td><td><strong>70</strong></td><td>67 (62 – 70)</td><td>-</td><td>66</td><td>77</td><td>-</td></tr><tr><td>Lisl</td><td><strong>28</strong></td><td>-</td><td>28</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Lisw</td><td><strong>5</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Sisl</td><td><strong>10</strong></td><td>-</td><td>10</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Vsl</td><td><strong>138</strong></td><td>107 (102 – 110)</td><td>135</td><td>138</td><td>142</td><td>-</td></tr><tr><td>Vsw ZV2</td><td><strong>110</strong></td><td>85 (77 – 91)</td><td>109</td><td>107</td><td>107</td><td>-</td></tr><tr><td>Vsw <i>anus</i></td><td><strong>88</strong></td><td>-</td><td>-</td><td>86</td><td>-</td><td>-</td></tr><tr><td>JV5</td><td><strong>38</strong></td><td>-</td><td>35</td><td>-</td><td>40</td><td>-</td></tr><tr><td>StIV</td><td><strong>53</strong></td><td>56 (53 – 59)</td><td>54</td><td>55</td><td>65</td><td>50</td></tr><tr><td>Scl</td><td><strong>3</strong></td><td>5</td><td>2</td><td>-</td><td>5</td><td>-</td></tr><tr><td>Scw</td><td><strong>12</strong></td><td>11 (11 – 12)</td><td>12</td><td>-</td><td>14</td><td>-</td></tr><tr><td>Fdl</td><td><strong>38</strong></td><td>32</td><td>35</td><td>-</td><td>-</td><td>-</td></tr><tr><td>No teeth Fd</td><td><strong>8</strong></td><td>8</td><td>-</td><td>-</td><td>6 – 7</td><td>-</td></tr><tr><td>Mdl</td><td><strong>30</strong></td><td>35</td><td>35</td><td>-</td><td>-</td><td>-</td></tr><tr><td>No teeth Md</td><td><strong>1</strong></td><td>1</td><td>-</td><td>-</td><td>0?</td><td>-</td></tr></tbody></table>

Sources of measurements – Africa (Benin 2♀♀, Tanzania 1♀) & Holotype Congo: Zannou et al. (2006); Central Africa: Schicha (1981b); South Africa-Madagascar (identified as N. insignitus, synonymized by Ueckermann & Loots 1988): van der Merwe (1968); South Africa: Ueckermann & Loots (1988); - : not provided.
Neoseiulus teke Pritchard & Baker

*Amblyseius (Amblyseius) teke* Pritchard & Baker, 1962: 239.

*Amblyseius teke*, Meyer & Rodrigues 1966: 30; Moraes *et al.* 1989a: 83; Moraes *et al.* 1989b: 97.

*Neoseiulus teke*, Moraes *et al.* 1986: 98; Chant & McMurtry 2003a: 37; Moraes *et al.* 2004a: 147; Chant & McMurtry 2007: 31.

*Amblyseius (Amblyseius) bibens* Blommers 1973: 111 (synonymy according to Ueckermann & Loots 1988).

Like *N. barkeri*, *N. longispinosus* and *N. scapilatus*, *N. teke* belongs to the *barkeri* species group and like *N. longispinosus* and *N. scapilatus*, it belongs to the *womersleyi* species subgroup (see above) (Chant and McMurtry 2003a).

This species is found in sub-Saharan Africa often associated with *Mononychellus tanajoa* (Bondar), the cassava green mite (CGM). It has been studied for its potential as BCA against the CGM. Nwilene and Nachman (1996) studied its reproduction characteristics on *M. tanajoa*. It was more efficient than *Iphiseius degenerans* (Berlese), but seems not efficient enough in

<table><caption>**Table 8** Character measurements of adult females and males of *Neoseiulus paspalivorus* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="3">Characters</th><th colspan="5">♀</th><th colspan="5">♂</th></tr><tr><th>La Réunion</th><th>Benin</th><th>Ghana</th><th>Philippines</th><th>Sri Lanka</th><th>Holotype Florida</th><th>La Réunion</th><th>Benin</th><th>Brazil</th><th>Ghana</th><th>Holotype Florida</th></tr><tr><th>8, this study</th><th>20</th><th>20</th><th>1</th><th>7</th><th>1</th><th>3, this study</th><th>20</th><th>20</th><th>20</th><th>1</th></tr></thead><tbody><tr><td>Dsl</td><td><strong>343</strong> (328–375)</td><td>343 (336–352)</td><td>338 (333–342)</td><td>285</td><td>322 (307–340)</td><td>334</td><td><strong>271</strong> (268–275)</td><td>257 (250–269)</td><td>257 (241–263)</td><td>257 (253–263)</td><td>247</td></tr><tr><td>Dsw</td><td><strong>135</strong> (125–143)</td><td>138 (130–146)</td><td>139 (130–146)</td><td>134</td><td>134 (127–142)</td><td>138</td><td><strong>149</strong> (143–165)</td><td>125 (120–127)</td><td>124 (120–127)</td><td>118 (114–120)</td><td>95</td></tr><tr><td>j1</td><td><strong>12</strong> (10–13)</td><td>11 (10–12)</td><td>12 (10–12)</td><td>7–9</td><td>10 (9–11)</td><td>11</td><td><strong>10</strong></td><td>10 (8–10)</td><td>10 (9–10)</td><td>10 (9–11)</td><td>8</td></tr><tr><td>j3</td><td><strong>12</strong> (10–14)</td><td>11 (10–12)</td><td>11 (10–12)</td><td>8–13</td><td>10 (8–11)</td><td>11</td><td><strong>10</strong> (10–11)</td><td>10 (8–11)</td><td>10 (9–11)</td><td>10 (9–11)</td><td>8</td></tr><tr><td>j4</td><td><strong>9</strong> (8–10)</td><td>9 (8–10)</td><td>9 (8–11)</td><td>7–9</td><td>9 (8–10)</td><td>9</td><td><strong>10</strong></td><td>8 (6–9)</td><td>9 (7–9)</td><td>8 (6–9)</td><td>8</td></tr><tr><td>j5</td><td><strong>9</strong> (8–10)</td><td>9 (8–10)</td><td>10 (8–11)</td><td>7–9</td><td>9 (8–10)</td><td>10</td><td><strong>9</strong> (8–10)</td><td>8 (6–9)</td><td>8 (7–9)</td><td>8 (7–9)</td><td>8</td></tr><tr><td>j6</td><td><strong>9</strong> (8–13)</td><td>10 (9–11)</td><td>10 (9–11)</td><td>7–9</td><td>10 (9–15)</td><td>11</td><td><strong>10</strong> (9–10)</td><td>9 (7–10)</td><td>9 (7–10)</td><td>9 (8–11)</td><td>8</td></tr><tr><td>J2</td><td><strong>10</strong> (9–10)</td><td>11 (9–12)</td><td>11 (10–12)</td><td>7–9</td><td>10 (8–11)</td><td>11</td><td><strong>10</strong> (9–10)</td><td>9 (8–11)</td><td>10 (9–11)</td><td>10 (8–10)</td><td>8</td></tr><tr><td>J5</td><td><strong>11</strong> (10–13)</td><td>9 (8–10)</td><td>10 (8–11)</td><td>7–9</td><td>8 (7–9)</td><td>9</td><td><strong>8</strong> (8–9)</td><td>7 (6–8)</td><td>8 (6–9)</td><td>7 (6–8)</td><td>8</td></tr><tr><td>r3</td><td><strong>12</strong> (10–13)</td><td>12 (10–14)</td><td>14 (12–15)</td><td>8–10</td><td>10 (8–11)</td><td>11</td><td><strong>9</strong></td><td>10 (9–10)</td><td>11 (10–12)</td><td>10 (9–11)</td><td>8</td></tr><tr><td>R1</td><td><strong>11</strong> (9–13)</td><td>11 (9–12)</td><td>11 (10–12)</td><td>8–10</td><td>10 (9–10)</td><td>10</td><td><strong>12</strong> (11–13)</td><td>9 (8–10)</td><td>10 (9–11)</td><td>10 (8–10)</td><td>8</td></tr><tr><td>s4</td><td><strong>12</strong> (10–13)</td><td>13 (11–14)</td><td>12 (11–15)</td><td>8–13</td><td>12 (11–12)</td><td>12</td><td><strong>12</strong> (11–13)</td><td>11 (9–11)</td><td>11 (10–11)</td><td>11 (10–12)</td><td>10</td></tr><tr><td>S2</td><td><strong>12</strong> (10–13)</td><td>14 (12–15)</td><td>14 (12–15)</td><td>8–13</td><td>12 (10–13)</td><td>14</td><td><strong>11</strong> (10–13)</td><td>11 (10–12)</td><td>11 (10–12)</td><td>12 (11–13)</td><td>9</td></tr><tr><td>S4</td><td><strong>14</strong> (10–15)</td><td>15 (14–16)</td><td>15 (14–16)</td><td>8–13</td><td>14 (13–15)</td><td>15</td><td><strong>13</strong> (12–13)</td><td>12 (11–14)</td><td>12 (11–13)</td><td>13 (11–15)</td><td>12</td></tr><tr><td>S5</td><td><strong>17</strong> (15–20)</td><td>16 (15–18)</td><td>19 (17–20)</td><td>16</td><td>18 (16–20)</td><td>19</td><td><strong>10</strong> (9–10)</td><td>13 (11–14)</td><td>15 (13–16)</td><td>15 (13–16)</td><td>8</td></tr><tr><td>z2</td><td><strong>10</strong> (9–10)</td><td>9 (8–10)</td><td>11 (9–12)</td><td>8–13</td><td>10 (9–10)</td><td>11</td><td><strong>9</strong> (8–10)</td><td>8 (6–9)</td><td>9 (7–10)</td><td>9 (8–10)</td><td>8</td></tr><tr><td>z4</td><td><strong>10</strong> (8–10)</td><td>10 (9–11)</td><td>11 (9–12)</td><td>8–13</td><td>10 (9–11)</td><td>11</td><td><strong>10</strong></td><td>9 (8–10)</td><td>10 (9–11)</td><td>10 (9–11)</td><td>8</td></tr><tr><td>z5</td><td><strong>10</strong> (9–10)</td><td>9 (9–10)</td><td>10 (9–11)</td><td>8</td><td>9 (8–9)</td><td>9</td><td><strong>9</strong> (8–9)</td><td>7 (6–9)</td><td>8 (7–9)</td><td>8 (7–9)</td><td>8</td></tr><tr><td>Z1</td><td><strong>12</strong> (10–13)</td><td>11 (10–14)</td><td>11 (9–12)</td><td>8–13</td><td>10 (9–11)</td><td>11</td><td><strong>12</strong> (12–13)</td><td>10 (8–10)</td><td>10 (9–10)</td><td>9 (7–11)</td><td>7</td></tr><tr><td>Z4</td><td><strong>16</strong> (13–20)</td><td>17 (15–18)</td><td>17 (15–19)</td><td>15</td><td>15 (13–17)</td><td>17</td><td><strong>17</strong> (15–18)</td><td>14 (12–15)</td><td>15 (14–16)</td><td>14 (13–16)</td><td>12</td></tr><tr><td>Z5</td><td><strong>49</strong> (45–51)</td><td>55 (51–56)</td><td>50 (48–52)</td><td>52</td><td>48 (45–52)</td><td>52</td><td><strong>40</strong> (38–42)</td><td>40 (36–44)</td><td>40 (37–45)</td><td>41 (38–44)</td><td>39</td></tr><tr><td>st1-st1</td><td><strong>41</strong> (40–43)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td><strong>38</strong></td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st2-st2</td><td><strong>51</strong> (48–55)</td><td>52 (47–54)</td><td>51 (47–54)</td><td>-</td><td>48 (45–53)</td><td>-</td><td><strong>44</strong> (40–48)</td><td>40 (38–41)</td><td>42 (41–44)</td><td>40 (38–41)</td><td>-</td></tr><tr><td>st3-st3</td><td><strong>53</strong> (48–60)</td><td>-</td><td>-</td><td>55</td><td>-</td><td>55</td><td><strong>45</strong> (40–50)</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st1-st3 ♀ / st1-st5 ♂</td><td><strong>81</strong> (70–93)</td><td>81 (76–82)</td><td>81 (79–86)</td><td>78</td><td>74 (72–76)</td><td>84</td><td><strong>117</strong> (112–122)</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st4-st4</td><td><strong>63</strong> (58–70)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td><strong>36</strong> (31–40)</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st5-st5</td><td><strong>55</strong> (53–58)</td><td>60 (57–63)</td><td>58 (57–60)</td><td>67</td><td>50 (49–53)</td><td>-</td><td><strong>31</strong> (29–33)</td><td>34 (32–35)</td><td>33 (31–35)</td><td>31 (28–35)</td><td>-</td></tr><tr><td>List</td><td><strong>38</strong> (35–43)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="4" style="text-align: center;">Not applicable</td></tr><tr><td>Lisw</td><td><strong>3</strong></td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="4" style="text-align: center;">Not applicable</td></tr><tr><td>Sisl</td><td><strong>7</strong> (5–10)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>10</td><td colspan="4" style="text-align: center;">Not applicable</td></tr><tr><td>Vsl</td><td><strong>103</strong> (100–110)</td><td>104 (98–108)</td><td>108 (101–111)</td><td>105</td><td>103 (95–115)</td><td>170</td><td><strong>98</strong> (100–105)</td><td>93 (89–98)</td><td>93 (85–95)</td><td>94 (92–98)</td><td>90</td></tr><tr><td>Vsw ZV2</td><td><strong>86</strong> (80–90)</td><td>81 (79–82)</td><td>81 (79–85)</td><td>81</td><td>78 (70–85)</td><td>86</td><td><strong>143</strong> (138–160)</td><td>118 (111–127)</td><td>117 (108–120)</td><td>117 (111–123)</td><td>108</td></tr><tr><td>Vsw anus</td><td><strong>77</strong> (75–80)</td><td>74 (70–76)</td><td>61 (54–73)</td><td>-</td><td>73 (67–80)</td><td>-</td><td><strong>97</strong> (100–102)</td><td>77 (70–82)</td><td>79 (73–82)</td><td>75 (70–79)</td><td>-</td></tr><tr><td>JV5</td><td><strong>25</strong> (23–28)</td><td>-</td><td>-</td><td>24</td><td>22 (17–28)</td><td>-</td><td><strong>20</strong> (19–20)</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>SIIV</td><td><strong>18</strong> (15–20)</td><td>18 (16–19)</td><td>18 (16–20)</td><td>21</td><td>19 (17–21)</td><td>-</td><td><strong>14</strong> (15–17)</td><td>15 (14–16)</td><td>-</td><td>16 (15–17)</td><td><strong>15</strong></td></tr><tr><td>Scl</td><td><strong>9</strong> (8–10)</td><td>8 (6–9)</td><td>8 (6–9)</td><td>-</td><td>7</td><td>2</td><td colspan="4" style="text-align: center;">Not applicable</td></tr><tr><td>Scw</td><td><strong>7</strong> (7–8)</td><td>7 (6–9)</td><td>7 (5–7)</td><td>-</td><td>-</td><td>5</td><td colspan="4" style="text-align: center;">Not applicable</td></tr><tr><td>Fdl</td><td><strong>18</strong> (15–20)</td><td>18 (17–19)</td><td>18 (16–19)</td><td>-</td><td>25 (23–29)</td><td>15</td><td><strong>15</strong></td><td>12 (11–12)</td><td>12 (11–14)</td><td>13 (11–14)</td><td>-</td></tr><tr><td>No teeth Fd</td><td><strong>6</strong></td><td>-</td><td>-</td><td>-</td><td>6</td><td>6</td><td><strong>2</strong></td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Mdl</td><td><strong>22</strong> (20–25)</td><td>24 (23–25)</td><td>23 (22–25)</td><td>-</td><td>23 (20–26)</td><td>-</td><td><strong>18</strong></td><td>18 (16–19)</td><td>18 (16–19)</td><td>17 (16–19)</td><td>-</td></tr><tr><td>No teeth Md</td><td><strong>1</strong></td><td>-</td><td>-</td><td>-</td><td>1</td><td>-</td><td><strong>1</strong></td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Shaft</td><td colspan="5" style="text-align: center;">Not applicable</td><td><strong>18</strong> (15–20)</td><td>12 (11–12)</td><td>12 (11–12)</td><td>12 (11–12)</td><td>13</td></tr></tbody></table>

Sources of measurements – For ♀♀: Benin and Ghana: Famah Sourassou *et al.* (2011); Philippines: Schicha & Corpuz-Raros (1992); Sri Lanka: Moraes *et al.* (2004b); Hototype Florida (USA): De Leon (1957) and Schicha (1981b). For ♂♂: Benin, Brazil and Ghana: Famah Sourassou *et al.* (2011); Hototype Florida (USA): De Leon (1957) and Schicha (1981b); Philippines: Schicha & Corpuz-Raros (1992). - : not provided.
<table><caption>**Table 9** Character measurements of adult females and one adult male of *Neoseiulus scapilatus* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="2">Characters</th><th colspan="4">♀</th><th colspan="2">♂</th></tr><tr><th>La Réunion<br>11, this study</th><th>Africa<br>10</th><th>Kenya<br>1</th><th>South Africa<br>6</th><th>La Réunion<br>1, this study</th><th>South Africa<br>1?</th></tr></thead><tbody><tr><td>Dsl</td><td><strong>313</strong> (305 – 325)</td><td>330 (312 – 345)</td><td>280</td><td>322 – 336</td><td><strong>245</strong></td><td>270</td></tr><tr><td>Dsw</td><td><strong>197</strong> (163 – 213)</td><td>192 (188 – 203)</td><td>140</td><td>190 – 210</td><td><strong>170</strong></td><td>185</td></tr><tr><td>j1</td><td><strong>18</strong> (15 – 20)</td><td>20 (18 – 23)</td><td>16</td><td>18 – 22</td><td><strong>15</strong></td><td>15</td></tr><tr><td>j3</td><td><strong>26</strong> (18 – 33)</td><td>28 (25 – 30)</td><td>25</td><td>30 – 34</td><td><strong>18</strong></td><td>25</td></tr><tr><td>j4</td><td><strong>16</strong> (14 – 18)</td><td>17 (14 – 19)</td><td>12</td><td>18 – 22</td><td><strong>15</strong></td><td>17</td></tr><tr><td>j5</td><td><strong>14</strong> (12 – 16)</td><td>16 (14 – 18)</td><td>13</td><td>18 – 22</td><td><strong>13</strong></td><td>17</td></tr><tr><td>j6</td><td><strong>13</strong> (11 – 15)</td><td>16 (14 – 18)</td><td>12</td><td>18 – 22</td><td><strong>13</strong></td><td>15</td></tr><tr><td>J2</td><td><strong>15</strong></td><td>18 (16 – 21)</td><td>14</td><td>18 – 22</td><td><strong>17</strong></td><td>17</td></tr><tr><td>J5</td><td><strong>12</strong> (9 – 16)</td><td>13 (10 – 19)</td><td>15</td><td>10 – 14</td><td><strong>10</strong></td><td>11</td></tr><tr><td>r3</td><td><strong>22</strong> (20 – 24)</td><td>23 (20 – 28)</td><td>-</td><td>24 – 27</td><td><strong>15</strong></td><td>23</td></tr><tr><td>R1</td><td><strong>16</strong> (10 – 21)</td><td>22 (18 – 25)</td><td>14</td><td>22 – 25</td><td><strong>15</strong></td><td>15</td></tr><tr><td>s4</td><td><strong>42</strong> (38 – 48)</td><td>43 (38 – 46)</td><td>32</td><td>44 – 48</td><td><strong>33</strong></td><td>31</td></tr><tr><td>S2</td><td><strong>38</strong> (33 – 43)</td><td>37 (30 – 43)</td><td>32</td><td>40 – 44</td><td><strong>30</strong></td><td>31</td></tr><tr><td>S4</td><td><strong>28</strong> (23 – 33)</td><td>29 (23 – 33)</td><td>24</td><td>30 – 34</td><td><strong>20</strong></td><td>25</td></tr><tr><td>S5</td><td><strong>21</strong> (18 – 25)</td><td>20 (15 – 23)</td><td>16</td><td>22 – 26</td><td><strong>18</strong></td><td>18</td></tr><tr><td>z2</td><td><strong>24</strong> (21 – 28)</td><td>24 (22 – 26)</td><td>16</td><td>30 – 34</td><td><strong>18</strong></td><td>18</td></tr><tr><td>z4</td><td><strong>28</strong> (25 – 35)</td><td>27 (24 – 30)</td><td>23</td><td>30 – 34</td><td><strong>21</strong></td><td>23</td></tr><tr><td>z5</td><td><strong>14</strong> (13 – 18)</td><td>16 (14 – 20)</td><td>12</td><td>18 – 22</td><td><strong>13</strong></td><td>17</td></tr><tr><td>Z1</td><td><strong>20</strong> (18 – 23)</td><td>22 (19 – 26)</td><td>21</td><td>25 – 28</td><td><strong>18</strong></td><td>20</td></tr><tr><td>Z4</td><td><strong>56</strong> (50 – 64)</td><td>60 (56 – 63)</td><td>48</td><td>58 – 64</td><td><strong>43</strong></td><td>43</td></tr><tr><td>Z5</td><td><strong>61</strong> (55 – 71)</td><td>64 (54 – 73)</td><td>58</td><td>62 – 68</td><td><strong>43</strong></td><td>42</td></tr><tr><td>st1-st1</td><td><strong>54</strong> (53 – 58)</td><td>-</td><td>-</td><td>-</td><td><strong>45</strong></td><td>-</td></tr><tr><td>st2-st2</td><td><strong>64</strong> (58 – 68)</td><td>65 (62 – 70)</td><td>58</td><td>74</td><td><strong>53</strong></td><td>-</td></tr><tr><td>st3-st3</td><td><strong>74</strong> (68 – 78)</td><td>-</td><td>-</td><td>-</td><td><strong>58</strong></td><td>92</td></tr><tr><td>st1-st3 ♀ / st1-st5 ♂</td><td><strong>60</strong> (55 – 70)</td><td>59 (56 – 61)</td><td>58</td><td>70</td><td><strong>98</strong></td><td>131</td></tr><tr><td>st4-st4</td><td><strong>69</strong> (60 – 75)</td><td>-</td><td>-</td><td>-</td><td><strong>45</strong></td><td>-</td></tr><tr><td>st5-st5</td><td><strong>62</strong> (60 – 65)</td><td>60 (58 – 64)</td><td>60</td><td>75 – 80</td><td><strong>30</strong></td><td>-</td></tr><tr><td>Lisl</td><td><strong>23</strong> (18 – 28)</td><td>-</td><td>-</td><td>-</td><td colspan="2">Not applicable</td></tr><tr><td>Lisw</td><td><strong>4</strong> (3 – 6)</td><td>-</td><td>-</td><td>-</td><td colspan="2">Not applicable</td></tr><tr><td>Sisl</td><td><strong>10</strong> (5 – 13)</td><td>-</td><td>-</td><td>-</td><td colspan="2">Not applicable</td></tr><tr><td>Vsl</td><td><strong>111</strong> (100 – 125)</td><td>117 (110 – 125)</td><td>92</td><td>112 – 122</td><td><strong>100</strong></td><td>119</td></tr><tr><td>Vsw ZV2</td><td><strong>102</strong> (88 – 113)</td><td>108 (100 – 128)</td><td>87</td><td>100 – 108</td><td><strong>132</strong></td><td>132</td></tr><tr><td>Vsw anus</td><td><strong>88</strong> (70 – 100)</td><td>90 (83 – 95)</td><td>78</td><td>-</td><td><strong>75</strong></td><td>-</td></tr><tr><td>JV5</td><td><strong>50</strong> (38 – 67)</td><td>-</td><td>40</td><td>-</td><td><strong>23</strong></td><td>31</td></tr><tr><td>SgeIV</td><td><strong>32</strong> (26 – 38)</td><td>36 (30 – 42)</td><td>32</td><td>32 – 36</td><td><strong>23</strong></td><td>23</td></tr><tr><td>StIV</td><td><strong>61</strong> (53 – 68)</td><td>72 (63 – 80)</td><td>48</td><td>66 – 70</td><td><strong>53</strong></td><td>54</td></tr><tr><td>Scl</td><td><strong>32</strong> (26 – 38)</td><td>31 (30 – 35)</td><td>31</td><td>47</td><td colspan="2">Not applicable</td></tr><tr><td>Scw</td><td><strong>12</strong> (5 – 25)</td><td>-</td><td>10</td><td>-</td><td colspan="2">Not applicable</td></tr><tr><td>Fdl</td><td><strong>31</strong> (28 – 35)</td><td>28 (28 – 29)</td><td>-</td><td>28</td><td><strong>23</strong></td><td>-</td></tr><tr><td>No teeth Fd</td><td><strong>4</strong></td><td>4</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Mdl</td><td><strong>30</strong> (28 – 33)</td><td>32 (31 – 33)</td><td>-</td><td>33</td><td><strong>23</strong></td><td>-</td></tr><tr><td>No teeth Md</td><td><strong>1</strong></td><td>1</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Shaft</td><td colspan="4">Not applicable</td><td colspan="2">-</td></tr></tbody></table>

Sources of measurements – For ♀♀: Africa (Benin: 1♀; Burundi 5♀♀, Kenya 1♀, Malawi 1♀, Rwanda 1♀, Uganda 1♀): Zannou *et al.* (2006); Kenya: El-Banhawy and Knapp (2011a); South Africa: van der Merwe (1965). For ♂♂: South Africa: Ueckermann & Loots (1988). - : not provided.
field conditions (Nwilene and Nachman 1996). Quilici *et al.* (2000) have collected this species before in La Réunion. Exact indications of locations were provided in the paper but without any measurements of specimens collected. Measurements of specimens collected during this study are provided in table 10.

**Specimens examined:** 12 ♀♀ in total + 6 ♂♂ + 2 im., 10 ♀♀ + 2 ♂♂ measured. Saint-Pierre - Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29'18" E, Lat 21°19'25"

<table><caption>**Table 10** Character measurements of adult females and males of *Neoseiulus teke* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="2">Characters</th><th colspan="6">♀</th><th colspan="3">♂</th></tr><tr><th>La Réunion<br>10, this study</th><th>Africa<br>10</th><th>Kenya<br>2</th><th>Madagascar<br>1</th><th>South Africa<br>5</th><th>Holotype Congo<br>1</th><th>La Réunion<br>2, this study</th><th>Madagascar<br>5</th><th>South Africa<br>2</th></tr></thead><tbody><tr><td>Dsl</td><td><strong>307</strong> (293 – 328)</td><td>308 (293 – 320)</td><td>295</td><td>340</td><td>332 – 341</td><td>348</td><td><strong>225 – 230</strong></td><td>260</td><td>270 – 285</td></tr><tr><td>Dsw</td><td><strong>165</strong> (155 – 173)</td><td>172 (162 – 186)</td><td>160</td><td>200</td><td>190 – 203</td><td></td><td><strong>150</strong></td><td>170</td><td>197 – 200</td></tr><tr><td>j1</td><td><strong>15</strong> (13 – 18)</td><td>18 (16 – 19)</td><td>18</td><td>20</td><td>19 – 23</td><td></td><td><strong>10 – 13</strong></td><td>15</td><td>15 – 18</td></tr><tr><td>j3</td><td><strong>46</strong> (38 – 53)</td><td>44 (35 – 50)</td><td>39</td><td>50</td><td>62 – 67</td><td>61</td><td><strong>25 – 33</strong></td><td>34</td><td>40 – 46</td></tr><tr><td>j4</td><td><strong>25</strong> (23 – 28)</td><td>30 (23 – 36)</td><td>39</td><td>30</td><td>38 – 45</td><td>42</td><td><strong>18 – 20</strong></td><td>18</td><td>26 – 34</td></tr><tr><td>j5</td><td><strong>39</strong> (35 – 43)</td><td>42 (35 – 48)</td><td>39</td><td>45</td><td>47 – 54</td><td></td><td><strong>25 – 30</strong></td><td>28</td><td>34 – 40</td></tr><tr><td>j6</td><td><strong>47</strong> (45 – 53)</td><td>48 (43 – 53)</td><td>42</td><td>54</td><td>62 – 66</td><td>60</td><td><strong>28 – 35</strong></td><td>30</td><td>40 – 46</td></tr><tr><td>j2</td><td><strong>55</strong> (50 – 58)</td><td>53 (45 – 59)</td><td>48</td><td>65</td><td>68 – 75</td><td>68</td><td><strong>33 – 38</strong></td><td>35</td><td>40 – 46</td></tr><tr><td>j5</td><td><strong>11</strong> (10 – 13)</td><td>11 (10 – 12)</td><td>9</td><td>10</td><td>12 – 14</td><td></td><td><strong>8 – 13</strong></td><td>9</td><td>11</td></tr><tr><td>r3</td><td><strong>44</strong> (40 – 48)</td><td>40 (34 – 46)</td><td>37</td><td>50</td><td>54 – 66</td><td>61</td><td><strong>23 – 28</strong></td><td>30</td><td>26 – 34</td></tr><tr><td>R1</td><td><strong>42</strong> (40 – 45)</td><td>37 (27 – 48)</td><td>37</td><td>48</td><td>54 – 66</td><td>66</td><td><strong>23</strong></td><td>24</td><td>26 – 34</td></tr><tr><td>s4</td><td><strong>60</strong> (55 – 65)</td><td>60 (54 – 64)</td><td>55</td><td>66</td><td>75 – 82</td><td>71</td><td><strong>43 – 48</strong></td><td>45</td><td>54 – 59</td></tr><tr><td>S2</td><td><strong>64</strong> (60 – 68)</td><td>61 (56 – 67)</td><td>58</td><td>70</td><td>72 – 80</td><td></td><td><strong>40 – 45</strong></td><td>45</td><td>46 – 54</td></tr><tr><td>S4</td><td><strong>47</strong> (43 – 60)</td><td>43 (40 – 48)</td><td>39</td><td>50</td><td>56 – 63</td><td></td><td><strong>28 – 30</strong></td><td>27</td><td>31 – 34</td></tr><tr><td>S5</td><td><strong>36</strong> (28 – 38)</td><td>32 (23 – 40)</td><td>34</td><td>40</td><td>46 – 52</td><td>48</td><td><strong>20 – 23</strong></td><td>25</td><td>25 – 31</td></tr><tr><td>z2</td><td><strong>51</strong> (48 – 53)</td><td>49 (41 – 54)</td><td>46</td><td>56</td><td>66 – 71</td><td>62</td><td><strong>33 – 37</strong></td><td>32</td><td>42 – 49</td></tr><tr><td>z4</td><td><strong>50</strong> (48 – 50)</td><td>51 (43 – 56)</td><td>48</td><td>54</td><td>68 – 75</td><td>65</td><td><strong>33 – 35</strong></td><td>38</td><td>42 – 49</td></tr><tr><td>z5</td><td><strong>25</strong> (23 – 28)</td><td>29 (19 – 38)</td><td>35</td><td>25</td><td>33 – 44</td><td>42</td><td><strong>18</strong></td><td>15</td><td>20 – 28</td></tr><tr><td>Z1</td><td><strong>54</strong> (50 – 58)</td><td>53 (45 – 62)</td><td>44</td><td>55</td><td>71 – 77</td><td>65</td><td><strong>33 – 40</strong></td><td>37</td><td>40 – 46</td></tr><tr><td>Z4</td><td><strong>57</strong> (50 – 63)</td><td>60 (54 – 67)</td><td>55</td><td>66</td><td>66 – 74</td><td></td><td><strong>40 – 45</strong></td><td>42</td><td>46 – 54</td></tr><tr><td>Z5</td><td><strong>68</strong> (65 – 73)</td><td>65 (59 – 74)</td><td>58</td><td>76</td><td>80 – 90</td><td></td><td><strong>45 – 50</strong></td><td>50</td><td>54 – 59</td></tr><tr><td>st1-st1</td><td><strong>44</strong> (43 – 48)</td><td></td><td></td><td></td><td></td><td></td><td><strong>40</strong></td><td></td><td></td></tr><tr><td>st2-st2</td><td><strong>55</strong> (53 – 58)</td><td>58 (56 – 63)</td><td>53</td><td></td><td></td><td></td><td><strong>50</strong></td><td></td><td></td></tr><tr><td>st3-st3</td><td><strong>67</strong> (61 – 70)</td><td></td><td></td><td></td><td>63 – 67</td><td></td><td><strong>53</strong></td><td></td><td></td></tr><tr><td>st1-st3 ♀ / st1-st5 ♂</td><td><strong>60</strong> (55 – 63)</td><td>56 (53 – 58)</td><td>53</td><td></td><td>56 – 59</td><td></td><td><strong>95</strong></td><td></td><td>116 – 128</td></tr><tr><td>st4-st4</td><td><strong>66</strong> (53 – 70)</td><td></td><td></td><td></td><td></td><td></td><td><strong>40 – 43</strong></td><td></td><td></td></tr><tr><td>st5-st5</td><td><strong>54</strong> (50 – 58)</td><td>56 (51 – 63)</td><td>58</td><td></td><td>70 – 74</td><td></td><td><strong>35</strong></td><td></td><td></td></tr><tr><td>Lisl</td><td><strong>25</strong> (23 – 28)</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Lisw</td><td><strong>3</strong> (3 – 5)</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Sisl</td><td><strong>10</strong> (8 – 10)</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Vsl</td><td><strong>109</strong> (100 – 115)</td><td>111 (104 – 118)</td><td>105</td><td>125</td><td>115 – 122</td><td>108</td><td><strong>95 – 103</strong></td><td>115</td><td>112 – 131</td></tr><tr><td>Vsw ZV2</td><td><strong>90</strong> (83 – 95)</td><td>97 (93 – 102)</td><td>80</td><td>100</td><td>95 – 100</td><td>95</td><td><strong>125 – 140</strong></td><td></td><td>139 – 145</td></tr><tr><td>Vsw anus</td><td><strong>71</strong> (63 – 80)</td><td></td><td></td><td></td><td></td><td></td><td><strong>50</strong></td><td></td><td></td></tr><tr><td>JV5</td><td><strong>57</strong> (55 – 63)</td><td></td><td>55</td><td>64</td><td>66 – 72</td><td></td><td><strong>28 – 33</strong></td><td>33</td><td>34 – 40</td></tr><tr><td>StIV</td><td><strong>69</strong> (48 – 75)</td><td>66 (51 – 77)</td><td>65</td><td>75</td><td>75 – 78</td><td>72</td><td><strong>50 – 53</strong></td><td>54</td><td>54 – 63</td></tr><tr><td>Scl</td><td><strong>25</strong> (23 – 28)</td><td>24 (22 – 27)</td><td>16</td><td>24</td><td>27</td><td></td><td></td><td></td><td></td></tr><tr><td>Scw</td><td><strong>6</strong> (5 – 13)</td><td></td><td></td><td>7</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Fdl</td><td><strong>24</strong> (18 – 25)</td><td>24 (23 – 25)</td><td></td><td>24</td><td>24</td><td></td><td><strong>20</strong></td><td>19</td><td></td></tr><tr><td>No teeth Fd</td><td><strong>4</strong></td><td>4</td><td>7</td><td>3</td><td>4</td><td></td><td><strong>3-4?</strong></td><td>2</td><td></td></tr><tr><td>Mdl</td><td><strong>24</strong> (20 – 28)</td><td>26 (25 – 27)</td><td></td><td>26</td><td>27</td><td></td><td><strong>18 – 20</strong></td><td>19</td><td></td></tr><tr><td>No teeth Md</td><td><strong>2</strong></td><td>2</td><td>2</td><td>2</td><td>2</td><td></td><td><strong>2?</strong></td><td>1</td><td></td></tr><tr><td>Shaft</td><td></td><td></td><td></td><td></td><td>Not applicable</td><td></td><td><strong>13</strong></td><td>12</td><td></td></tr></tbody></table>

Sources of measurements – For ♀♀: Africa (Burundi 1♀, Ghana 2♀♀, Kenya 3♀♀, Malawi 1♀, Mozambique 1♀, Rwanda 1♀, Sierra Leone 1♀): Zannou *et al.* (2006); Kenya: El-Banhawy & Knapp (2011a); Madagascar (Identified as *Amblyseius bibens* but synonymized by Ueckermann & Loots 1988): Blommers (1973); South Africa: van der Merwe (1968); Holotype Congo: Zannou *et al.* (2006). For ♂♂: Madagascar (Identified as *Amblyseius bibens* but synonymized by Ueckermann & Loots 1988): Blommers (1973); South Africa: Ueckermann & Loots (1988). - : not provided.
S), 1 ♀ + 1 im. on *M. repens*, 20/2/2017; 1 ♀ on *A. hispidum*, 1 ♀ + 1 im. on *M. repens*, 1 ♀ + 1 ♂ on *P. maximum*, 30/3/2017; 4 ♀♀ + 1 ♂ on *M. repens*, 3/4/2017; 2 ♀♀ + in plots HM, 1 ♀ + 1 ♂ in plot CC, 1 ♀ in plot H, 4/4/2017 and 6/4/2017.

**Remarks:** measurements of morphological characters of *N. teke* female and male specimens from La Réunion (Table 10) are very close from measurements for specimens from neighbouring countries, especially from specimens from various countries in Africa, except for the holotype (Zannou *et al.* 2006) and specimens from South Africa which are larger (van der Merwe 1965).

## Tribe Kampimodromini Kolodochka

Kampimodromini Kolodochka 1998: 59; Chant & McMurtry, 2003b: 189; 2006b: 137; 2007: 33.

## Subtribe Paraphytoseiina Chant & McMurtry

Paraphytoseiina Chant & McMurtry 2003b: 211.

## Genus Paraphytoseius Swirskii & Shechter

*Paraphytoseius Swirski & Shechter 1961: 113; Moraes et al. 1986:104; Chant & McMurtry 2003b: 216; Moraes et al. 2004a: 160; Chant & McMurtry 2007: 49.*

*Amblyseius (Paraphytoseius), Ueckermann & Loots 1987: 221.*

*Amblyseius (Ptenoseius), Pritchard & Baker 1962: 295.*

*Proprioseius (Paraphytoseius), Karg 1983: 302.*

*Ptenoseius, Schuster & Pritchard 1963: 198.*

## Paraphytoseius horrifer (Pritchard & Baker)

*Amblyseius (Ptenoseius) horrifer Pritchard & Baker, 1962: 295.*

*Amblyseius horrifer, Meyer & Rodrigues 1966: 30.*

*Amblyseius (Paraphytoseius) horrifer, van der Merwe 1968: 169.*

*Proprioseius (Paraphytoseius) horrifer, Karg 1983: 302.*

*Paraphytoseius horrifer, Moraes et al. 1986: 105; Beard 2001: 84; Chant & McMurtry 2003a: 37; Moraes et al. 2004a: 152; Chant & McMurtry 2007: 53.*

In our specimens of this species, setae S5 are absent. So accordingly with Chant and McMurtry (2003b) all specimens belong to the *orientalis* species group.

Accordingly with these previous authors, and with Moraes *et al.* (2007), we consider that *P. horrifer* and *P. orientalis* are different valid species. Our specimens with longer setae s4, Z4, Z5, and lacking a distinctly short, thick, spatulate macroseta on genu I.

This species is widely distributed in Sub-Saharan Africa and Madagascar. The biology of *P. horrifer* remains totally unknown.

This is the first mention of this species from La Réunion Island.

**Specimens examined:** 13 ♀♀ + 1 ♂ + 1 im. in total, 12 ♀♀ + 1 ♂ measured. Le Tampon - Ligne des 400 (aasl 463 m, Long 55°30'36" E, Lat 21°17'24" S), 1 ♂ + 2 im. on *Ipomoea purpurea* (L.), 11/2/2017; Ravine Langevin - Grand-Galet Waterfall (aasl 850 m, Long 55°21'33" E, Lat 21°17'47" S), 7 ♀♀ on *Desmodium incanum* DC., 11/12/2016; Forêt de Bélouve - Gîte (aasl 1500 m, Long 55°33'36" E, Lat 21°6'0" S), 1 ♀ on *Fuchsia boliviana* Carrière, 20/12/16; Cilaos - Village (aasl 991 m, Long 55°27'0" E, Lat 21°8'24" S), 1 ♀ on *Acalypha hispida* Burm. f., 8/1/17; Saint-Pierre - Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29'18" E, Lat 21°19'25" S), 1 ♀ in plot BM, 23/8/2016; 1 ♀ on *M. coromandelianum*, 19/6/2017; 1 ♀ on *A. viridis*, 20/6/2017.

**Remarks:** even if lengths of setae seem very variable in *P. horrifer* in the literature (van der Merwe 1968; Moraes *et al.* 2007), measurements of morphological characters of La Réunion specimens (Table 11) agree well with measurements found in the literature, being however
slightly shorter for almost all characters compared with female and male specimens of South Africa (van der Merwe 1968).

**Paraphytoseius orientalis** (Narayanan, Kaur & Ghai)

*Typhlodromus (Amblyseius) orientalis* Narayanan, Kaur & Ghai 1960: 394.

*Paraphytoseius orientalis*, Moraes *et al*. 1986: 105; Chant & McMurtry 2003b: 220; Moraes *et al*. 2004a: 162, Chant & McMurtry 2007: 53.

*Amblyseius ipomeai*, Narayanan, Kaur & Ghai 1960: 394 (synonymy according to El-Banhawy 1984); *Paraphytoseius narayanani*, Ehara 1967: 67 (synonymy according to Ehara & Ghai, in Ehara 1967: 77); *Paraphytoseius multidentatus*, Swirski & Shechter 1961: 114 (synonymy according to Matthysse & Denmark 1981 in Denmark *et al*. 1999: 11).

As mentioned earlier, in our specimens of *Paraphytoseius* spp., setae S5 are absent, all our specimens belong to the *orientalis* species group (Chant and McMurtry 2003b) and we consider that *P. horrifer* and *P. orientalis* are distinct, valid species.

Our specimens with shorter s4, Z4 and Z5 setae, having a distinctly short, thick, spatulate macroseta on genu I belong to *P. orientalis*.

This species is widely distributed in tropical and subtropical areas in South America, Africa and Asia. This species belongs to a genus included in the large polyphagous generalist group named type III phytoseiid mites (McMurtry and Croft 1997; McMurtry *et al*. 2013). Navasero and Navasero (2016) have studied the life history of *P. orientalis* on the broad mite (*P. latus*) as prey. The authors reported high predation rates on the eggs of *P. latus*, suggesting good potential for the control of this pest. Quilici et al. (2000) have collected before this species in La Réunion but provided no measurements. We herein provide measurements of specimens collected in La Réunion (Table 12).

**Specimens examined:** 20 ♀♀ + 1 ♂ + 1 im. in total, 5 ♀♀ + 1 ♂ measured. Saint-Pierre - Bassin Martin, Armefhor Station (aasl 450 m, Long 55°31'9" E, Lat 21°18'14" S), 8 ♀♀ + 1 ♂ on *Borago officinalis* L., 17/12/2015; Saint-Pierre - Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29'18" E, Lat 21°19'25" S), 4 ♀♀ + 1 im. in plot BM, 2 ♀♀ in plot F, and 2 ♀♀ in plot H, 23/8/2016; 1 ♀ on *M. coromandelianum*, 19/6/2017; 1 ♀ on *A. viridis*, 20/6/2017; Ravine Langevin - Grand-Galet Waterfall (aasl 850 m, Long 55°21'33" E, Lat 21°17'47" S), 2 ♀♀ on *D. incanum*, 11/12/2016.

**Remarks:** measurements of morphological characters of La Réunion female and male specimens (Table 12) agree well with measurements of specimens from other countries found in the literature.

**Tribe Phytoseiulini Chant & McMurtry**

Phytoseiulini Chant & McMurtry 2006: 7.

**Genus Phytoseiulus Evans**

*Phytoseiulus* Evans 1952: 397.

**Phytoseiulus persimilis Athias-Henriot**

*Phytoseiulus persimilis* Athias-Henriot 1957: 347; Moraes *et al*. 1986: 109; Moraes *et al*. 2004a: 169; Chant & McMurtry 2006a: 20; 2007: 55.

*Phytoseiulus (Phytoseiulus) persimilis*, Wainstein 1962: 17.

*Typhlodromus persimilis*, Hirschmann 1962: 75.

*Phytoseiulus riegeli* Dosse 1958: 48 (synonymy according to Chant 1959); *Phytoseiulus tardi* Lombardini 1959: 166 (synonymy according to Kennett & Caltagirone 1968).
<table><caption>**Table 11** Character measurements of adult females and one adult male of *Paraphytoseius horrifer* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="2">Characters</th><th colspan="5">♀</th><th colspan="2">♂</th></tr><tr><th>La Réunion<br>12, this study</th><th>Africa<br>4</th><th>Senegal<br>2</th><th>South Africa<br>2</th><th>Holotype<br>Zaire, 1</th><th>La Réunion<br>1, this study</th><th>South Africa<br>2</th></tr></thead><tbody><tr><td>Dsl</td><td>295 (265-330)</td><td>300 (294-310)</td><td>300-304</td><td>318-339</td><td>298</td><td>250</td><td>260</td></tr><tr><td>Dsw</td><td>153 (128-175)</td><td>158 (149-166)</td><td>163-170</td><td>165-176</td><td>166</td><td>110</td><td>140</td></tr><tr><td>j1</td><td>33 (25-40)</td><td>38 (37-40)</td><td>35-38</td><td>39-42</td><td>38</td><td>15</td><td></td></tr><tr><td>j3</td><td>86 (78-100)</td><td>85 (80-88)</td><td>83-85</td><td>-</td><td>84</td><td>58</td><td>59</td></tr><tr><td>j4</td><td>5 (3-5)</td><td>3</td><td>3-5</td><td>-</td><td>3</td><td>4</td><td>-</td></tr><tr><td>j5</td><td>5 (3-5)</td><td>3</td><td>3-5</td><td>5</td><td>3</td><td>4</td><td>-</td></tr><tr><td>j6</td><td>8 (5-10)</td><td>7 (6-8)</td><td>5-6</td><td>9</td><td>6</td><td>7</td><td>-</td></tr><tr><td>j5</td><td>5 (3-8)</td><td>4 (3-5)</td><td>3-5</td><td>5</td><td>3</td><td>3</td><td>-</td></tr><tr><td>r3</td><td>48 (45-58)</td><td>43 (10-45)</td><td>43-45</td><td>52</td><td>45</td><td>30</td><td>33</td></tr><tr><td>R1</td><td>38 (30-45)</td><td>31 (27-41)</td><td>30-32</td><td>36</td><td>28</td><td>18</td><td>22</td></tr><tr><td>s4</td><td>126 (110-148)</td><td>131 (122-139)</td><td>118-125</td><td>136-141</td><td>138</td><td>80</td><td>103</td></tr><tr><td>z2</td><td>12 (9-13)</td><td>11 (10-12)</td><td>8-10</td><td>14</td><td>11</td><td>10</td><td>-</td></tr><tr><td>z4</td><td>11 (9-13)</td><td>8 (6-11)</td><td>8-10</td><td>9</td><td>6</td><td>10</td><td>-</td></tr><tr><td>z5</td><td>5 (3-8)</td><td>5 (4-5)</td><td>3-5</td><td>5</td><td>5</td><td>5</td><td>-</td></tr><tr><td>Z1</td><td>7 (5-8)</td><td>7 (6-8)</td><td>7-9</td><td>5</td><td>5</td><td>7</td><td>-</td></tr><tr><td>Z4</td><td>80 (70-88)</td><td>80 (77-84)</td><td>73-75</td><td>85-89</td><td>84</td><td>45</td><td>59</td></tr><tr><td>Z5</td><td>109 (93-133)</td><td>114 (112-118)</td><td>95-100</td><td>122-127</td><td>116</td><td>50</td><td>75</td></tr><tr><td>st1-st1</td><td>68 (55-73)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>53</td><td>-</td></tr><tr><td>st2-st2</td><td>71 (65-80)</td><td>73 (72-74)</td><td>68-69</td><td>78</td><td>69</td><td>55</td><td>-</td></tr><tr><td>st3-st3</td><td>79 (73-85)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>63</td><td>-</td></tr><tr><td>st1-st3 ♀ / st1-st5 ♂</td><td>68 (58-75)</td><td>66 (64-69)</td><td>68-70</td><td>92</td><td>63</td><td>98</td><td>-</td></tr><tr><td>st4-st4</td><td>88 (75-98)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>55</td><td>-</td></tr><tr><td>st5-st5</td><td>86 (75-95)</td><td>83 (82-86)</td><td>85-88</td><td>87</td><td>84</td><td>48</td><td>-</td></tr><tr><td>Lisl</td><td>30 (25-38)</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="2">Not applicable</td></tr><tr><td>Lisw</td><td>3</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="2">Not applicable</td></tr><tr><td>Sisl</td><td>11 (10-13)</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="2">Not applicable</td></tr><tr><td>Vsl</td><td>99 (53-118)</td><td>103 (96-110)</td><td>113-115</td><td>117</td><td>115</td><td>-</td><td>122</td></tr><tr><td>Vsw ZV2</td><td>59 (53-75)</td><td>68 (62-74)</td><td>68-70</td><td>70</td><td>75</td><td>105</td><td>120</td></tr><tr><td>Vsw anus</td><td>61 (55-68)</td><td>62 (59-67)</td><td>65</td><td>-</td><td>63</td><td>60</td><td>-</td></tr><tr><td>JV5</td><td>77 (70-85)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>50</td><td>28</td></tr><tr><td>SgeII</td><td>15 (13-15)</td><td>12 (11-13)</td><td>13-15</td><td>-</td><td>-</td><td>13</td><td>14</td></tr><tr><td>SgeIV</td><td>27 (23-45)</td><td>25 (24-27)</td><td>28-30</td><td>28</td><td>24</td><td>18</td><td>23</td></tr><tr><td>StiIV</td><td>36 (30-43)</td><td>37 (35-40)</td><td>32-38</td><td>47</td><td>34</td><td>23</td><td>34</td></tr><tr><td>StIV</td><td>43 (38-47)</td><td>40 (38-43)</td><td>43-45</td><td>47</td><td>36</td><td>30</td><td>37</td></tr><tr><td>SttIV</td><td>45 (43-48)</td><td>41 (39-42)</td><td>33-35</td><td>-</td><td>-</td><td>30</td><td>-</td></tr><tr><td>Scl</td><td>3 (3-5)</td><td>4 (3-5)</td><td>-</td><td>25</td><td>4</td><td colspan="2">Not applicable</td></tr><tr><td>Scw</td><td>10 (9-11)</td><td>12 (9-13)</td><td>-</td><td>10</td><td>-</td><td>20</td><td>22</td></tr><tr><td>Fdl</td><td>34 (23-50)</td><td>29</td><td>-</td><td>-</td><td>-</td><td>5?</td><td>8</td></tr><tr><td>No teeth Fd</td><td>11</td><td>10-11</td><td>-</td><td>-</td><td>-</td><td>20</td><td>22</td></tr><tr><td>Mdl</td><td>32 (25-38)</td><td>34 (33-34)</td><td>-</td><td>-</td><td>-</td><td>1</td><td>1</td></tr><tr><td>No teeth Md</td><td>2</td><td>2-3</td><td>-</td><td>-</td><td>-</td><td>15</td><td>18</td></tr><tr><td>Shaft</td><td colspan="5">Not applicable</td><td colspan="2"></td></tr></tbody></table>

Sources of measurements – For ♀♀: Africa (Benin 1♀, Kenya 1♀, Uganda 2♀♀): Moraes *et al.* (2007); Senegal: Kade *et al.* (2011); South Africa: van der Merwe (1968); Holotype Zaïre: Moraes *et al.* 1989). For ♂♂: South Africa: van der Merwe (1968). - : not provided.
*Phytoseiulus persimilis* is a Mediterranean/subtropical predatory mite, a type I species, i.e. a specialist predator of the *urticae* species group of the genus *Tetranychus* (McMurtry and Croft 1997; McMurtry *et al.* 2013). Considerable research has been conducted on this predator-prey interaction (see review by Kostiainen and Hoy 1996), and numerous biological control programs have used *P. persimilis* against *T. urticae* on a wide range of ornamental and vegetable crops. *Phytoseiulus persimilis* was the first greenhouse biological control agents available commercially and it is one of the most successful BCA in the world. It can also be used in temperate climates on open-field crops such as strawberries. Optimum conditions are 20-27 °C and relative humidity of 60-90 %. Cooler or warmer temperatures may have a negative effect on reproduction, development and efficiency of this predatory mite. This species is present in Mauritius (Kreiter *et al.* 2018a) and La Réunion probably because of its commercial introduction and uses in vegetable and ornamental greenhouses, dispersion of some specimens released and establishment in the environment. This species is actually reared and sold in La Réunion and commercialised in Mascareignes since a long time (Quilici, personal communication). *Phytoseiulus persimilis* was already known from La Réunion (Quilici *et al.* 1997, 2000). Exact indications of locations were provided in these papers but without any

<table><caption>**Table 12** Character measurements of adult females and one adult male of *Paraphytoseius orientalis* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="2">Characters</th><th colspan="6">♀</th><th colspan="5">♂</th></tr><tr><th>La Réunion<br>5, this study</th><th>Africa 1<br>?</th><th>Africa 2<br>5</th><th>Hong-Kong<br>7</th><th>Kenya<br>5</th><th>Madagascar<br>?</th><th>Mauritius<br>8</th><th>Paratyple<br>Hong-Kong</th><th>La Réunion<br>1, this study</th><th>Madagascar<br>?</th><th>Philippines<br>2</th><th>Taiwan<br>2</th><th>Thailand<br>4</th></tr></thead><tbody><tr><td>Dsl</td><td>266 (260-283)</td><td>291 (252-317)</td><td>291 (280-304)</td><td>259-305</td><td>250</td><td>280-295</td><td>290 (276-300)</td><td>306</td><td>250</td><td>220</td><td>197</td><td>220 (209-231)</td><td>230</td></tr><tr><td>Dsw</td><td>150 (145-150)</td><td>152 (127-173)</td><td>159 (149-168)</td><td>-</td><td>150</td><td>155-160</td><td>146 (140-149)</td><td>165</td><td>110</td><td>140</td><td>91</td><td>93 (90-96)</td><td>160</td></tr><tr><td>j1</td><td>31 (28-35)</td><td>35 (31-41)</td><td>32 (29-37)</td><td>36 (30-38)</td><td>40</td><td>30-36</td><td>35 (34-36)</td><td>36</td><td>23</td><td>27</td><td>27</td><td>23 (14-26)</td><td>25</td></tr><tr><td>j3</td><td>83 (81-88)</td><td>84 (72-91)</td><td>83 (72-90)</td><td>79 (76-81)</td><td>80</td><td>79-89</td><td>83 (78-89)</td><td>81</td><td>50</td><td>55</td><td>60</td><td>58 (49-64)</td><td>59</td></tr><tr><td>j4</td><td>5 (4-5)</td><td>2</td><td>3 (2-3)</td><td>4 (3-5)</td><td>2-3</td><td>&lt; 10</td><td>5</td><td>4</td><td>4</td><td>-</td><td>3-4</td><td>-</td><td>4</td></tr><tr><td>j5</td><td>5</td><td>2</td><td>3 (2-3)</td><td>4 (3-5)</td><td>2-3</td><td>&lt; 10</td><td>5</td><td>4</td><td>4</td><td>-</td><td>3-4</td><td>-</td><td>5</td></tr><tr><td>j6</td><td>8 (5-8)</td><td>6 (5-8)</td><td>6 (5-6)</td><td>4 (3-5)</td><td>2-3</td><td>&lt; 10</td><td>7 (6-7)</td><td>6</td><td>5</td><td>-</td><td>3-4</td><td>-</td><td>6</td></tr><tr><td>j5</td><td>5 (4-5)</td><td>3 (2-5)</td><td>4 (3-5)</td><td>4 (3-5)</td><td>9</td><td>&lt; 10</td><td>4 (4-5)</td><td>5</td><td>3</td><td>-</td><td>3-4</td><td>-</td><td>4</td></tr><tr><td>r3</td><td>38 (35-40)</td><td>44 (41-46)</td><td>42 (38-46)</td><td>43 (41-46)</td><td>46</td><td>36-48</td><td>44 (42-46)</td><td>45</td><td>30</td><td>30</td><td>29</td><td>32 (28-35)</td><td>27</td></tr><tr><td>R1</td><td>34 (33-35)</td><td>31 (24-38)</td><td>28 (26-30)</td><td>26 (20-33)</td><td>39</td><td>23-35</td><td>39 (36-41)</td><td>25</td><td>13</td><td>13</td><td>12</td><td>14 (14-15)</td><td>14</td></tr><tr><td>s4</td><td>120 (113-123)</td><td>120 (110-137)</td><td>117 (110-126)</td><td>117 (107-124)</td><td>127</td><td>118-130</td><td>115 (107-124)</td><td>117</td><td>78</td><td>-</td><td>81</td><td>83 (72-95)</td><td>81</td></tr><tr><td>z2</td><td>11 (10-15)</td><td>11 (7-14)</td><td>9 (8-10)</td><td>8 (5-11)</td><td>9</td><td>&lt; 10</td><td>12 (11-12)</td><td>9</td><td>13</td><td>-</td><td>6</td><td>-</td><td>9</td></tr><tr><td>z4</td><td>10 (5-10)</td><td>8 (7-10)</td><td>9 (8-10)</td><td>10 (8-13)</td><td>5</td><td>&lt; 10</td><td>9 (8-9)</td><td>11</td><td>16</td><td>-</td><td>8</td><td>-</td><td>8</td></tr><tr><td>z5</td><td>6 (6-8)</td><td>4 (2-5)</td><td>5</td><td>4 (3-5)</td><td>3</td><td>&lt; 10</td><td>5 (4-5)</td><td>3</td><td>4</td><td>-</td><td>3-4</td><td>-</td><td>5</td></tr><tr><td>Z1</td><td>9 (6-9)</td><td>7 (5-10)</td><td>7 (6-8)</td><td>8 (8-10)</td><td>6</td><td>&lt; 10</td><td>9 (8-10)</td><td>8</td><td>6</td><td>-</td><td>4</td><td>-</td><td>6</td></tr><tr><td>Z4</td><td>80 (70-80)</td><td>75 (60-86)</td><td>72 (67-77)</td><td>74 (69-81)</td><td>83</td><td>68-80</td><td>74 (71-80)</td><td>71</td><td>45</td><td>48</td><td>50</td><td>46 (43-48)</td><td>47</td></tr><tr><td>Z5</td><td>105 (99-107)</td><td>106 (84-130)</td><td>96 (90-101)</td><td>82 (76-91)</td><td>138</td><td>96-116</td><td>104 (102-107)</td><td>94</td><td>60</td><td>60</td><td>57</td><td>57 (52-61)</td><td>52</td></tr><tr><td>st1-st1</td><td>70</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>55</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st2-st2</td><td>68 (65-68)</td><td>68 (60-74)</td><td>65 (62-67)</td><td>-</td><td>60</td><td>-</td><td>69 (67-72)</td><td>66</td><td>55</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st3-st3</td><td>75 (75-78)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>63</td><td>-</td><td>-</td><td>63 (60-65)</td><td>-</td></tr><tr><td>st1-st3 ♀ / st1-st5 ♂</td><td>65 (64-65)</td><td>65 (58-72)</td><td>64 (61-66)</td><td>-</td><td>65</td><td>-</td><td>65 (62-68)</td><td>66</td><td>110</td><td>-</td><td>-</td><td>106 (101-111)</td><td>-</td></tr><tr><td>st4-st4</td><td>85 (85-93)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>55</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>st5-st5</td><td>85</td><td>73 (67-74)</td><td>82 (80-85)</td><td>-</td><td>96</td><td>-</td><td>-</td><td>79</td><td>48</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Lisl</td><td>18</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="5">Not applicable</td></tr><tr><td>Lisw</td><td>2</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="5">Not applicable</td></tr><tr><td>Vsl</td><td>80</td><td>97 (84-108)</td><td>99</td><td>93 (84-102)</td><td>100</td><td>102-108</td><td>102 (96-107)</td><td>97</td><td>108</td><td>-</td><td>76</td><td>95 (87-109)</td><td>-</td></tr><tr><td>Vsw ZV2</td><td>65</td><td>58 (48-62)</td><td>61 (56-64)</td><td>53</td><td>55</td><td>59-63</td><td>60 (57-64)</td><td>52</td><td>80</td><td>-</td><td>117</td><td>99 (95-108)</td><td>-</td></tr><tr><td>Vsw anus</td><td>63</td><td>52 (43-58)</td><td>59 (56-62)</td><td>53-55</td><td>-</td><td>-</td><td>56 (52-61)</td><td>55</td><td>60</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>JV5</td><td>66 (65-75)</td><td>-</td><td>69 (64-76)</td><td>70</td><td>63-82</td><td>78 (77-78)</td><td>-</td><td>-</td><td>23</td><td>22</td><td>-</td><td>16 (14-20)</td><td>21</td></tr><tr><td>Sgel</td><td>-</td><td>-</td><td>8</td><td>-</td><td>-</td><td>-</td><td>-</td><td>6</td><td>7</td><td>-</td><td>4</td><td>-</td><td>-</td></tr><tr><td>SgelII</td><td>13</td><td>13 (10-14)</td><td>12 (11-13)</td><td>-</td><td>10</td><td>Present</td><td>-</td><td>13</td><td>10</td><td>-</td><td>8</td><td>-</td><td>-</td></tr><tr><td>SgelIII</td><td>-</td><td>-</td><td>13 (11-14)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>13</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Still</td><td>-</td><td>-</td><td>13 (11-14)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>14</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>SgelIV</td><td>23 (21-28)</td><td>25 (24-29)</td><td>28 (24-32)</td><td>28 (25-33)</td><td>28</td><td>22-27</td><td>22 (21-24)</td><td>25</td><td>18</td><td>20</td><td>23</td><td>-</td><td>21</td></tr><tr><td>StIV</td><td>38 (30-38)</td><td>35 (29-41)</td><td>34 (33-37)</td><td>35 (33-38)</td><td>40</td><td>28-38</td><td>30 (26-33)</td><td>35</td><td>28</td><td>30</td><td>30</td><td>-</td><td>29</td></tr><tr><td>StV</td><td>41 (38-42)</td><td>37 (31-43)</td><td>41 (40-42)</td><td>44 (41-46)</td><td>40</td><td>38-45</td><td>38 (36-42)</td><td>43</td><td>33</td><td>33</td><td>35</td><td>-</td><td>35</td></tr><tr><td>StIV</td><td>33 (33-45)</td><td>-</td><td>38 (35-40)</td><td>33 (30-36)</td><td>-</td><td>34-42</td><td>38 (36-41)</td><td>36</td><td>33</td><td>30-35</td><td>29</td><td>-</td><td>-</td></tr><tr><td>Scl</td><td>3 (3-5)</td><td>-</td><td>3</td><td>-</td><td>5</td><td>20</td><td>5 (4-5)</td><td>-</td><td colspan="5">Not applicable</td></tr><tr><td>Scw</td><td>10 (8-11)</td><td>-</td><td>13 (13-14)</td><td>-</td><td>-</td><td>7</td><td>6</td><td>-</td><td>20</td><td>20</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Fdl</td><td>27 (27-30)</td><td>-</td><td>26</td><td>25</td><td>-</td><td>-</td><td>29 (27-31)</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td>No teeth Fd</td><td>7-8</td><td>-</td><td>7-8</td><td>7</td><td>8-9</td><td>8</td><td>9-11</td><td>-</td><td>-</td><td>8</td><td>-</td><td>7</td><td>8</td></tr><tr><td>Mdl</td><td>35 (31-35)</td><td>-</td><td>28</td><td>29 (28-30)</td><td>-</td><td>-</td><td>30 (28-31)</td><td>-</td><td>20</td><td>20</td><td>-</td><td>-</td><td>-</td></tr><tr><td>No teeth Md</td><td>2</td><td>-</td><td>2</td><td>2</td><td>-</td><td>3</td><td>2</td><td>-</td><td>-</td><td>1</td><td>-</td><td>1</td><td>1</td></tr><tr><td>Shaft</td><td colspan="6">Not applicable</td><td>-</td><td>-</td><td>15</td><td>17</td><td>11</td><td>-</td><td>-</td></tr></tbody></table>

Sources of measurements – For ♀♀: Africa 1 (Benin and Nigeria, no mention of respective number of specimens); Moraes *et al.* (1989); Africa 2 (Burundi 1♀, Kenya 2♀♀, Rwanda 2♀♀) and paratype from Hong-Kong: Moraes *et al.* (2007); Hong-Kong: Swirski & Shechter (1961); Kenya: El-Banhawy & Knapp (2011a); Madagascar (identified as *Paraphytoseius multidentatus*, synonymized by Mathysse & Denmark 1981); Blommers (1976); Mauritius: Ferragut & Baumann (2019). For ♂♂: Madagascar (identified as *Paraphytoseius multidentatus*, synonymized by Mathysse & Denmark 1981); Philippines: Schicha & Corpuz-Raros (1985); Taiwan: Ho & Lo (1989); Thailand (identified as *Paraphytoseius multidentatus*, synonymized by Mathysse & Denmark 1981): Ehara & Bhandhufalck (1977). - : not provided.
measurements of specimens collected. Measurements of specimens collected during this study are provided in Table 13.

**Specimens examined:** 5 ♀♀ in total, all measured. Montvert-les-Hauts - EARL Le Mont Vert farm (aasl 582 m, Long 55°32'19" E, Lat 21°19'42" S), 1 ♀ on *Fragaria* sp., 3 ♀♀ on *E. sonchifolia*, 4/8/2015; Le 19ᵉ - Plaine des Caffres, JL Robert farm (aasl 1000 m, Long 55°32'9" E, Lat 21°14'16" S), 1 ♀ on *Phytolacca americana* L., 15/12/2015.

**Remarks:** measurements of morphological characters (Table 13) of the 5 females fit well with measurements found in the literature, especially with those of specimens collected recently in Mauritius (Kreiter *et al.* 2018a), and particularly for setae *j4, J5, z2, z5* and *r3*. Some setae are slightly shorter in La Réunion specimens compared to specimens from other countries, mainly for the long setae of this species (*j6, s4, Z1, Z4, and Z5*). Some other shorter setae of this species, for example *R1* and macrosetae of the leg IV are also shorter in La Réunion specimens. Nevertheless, these findings are questionable given that only five females of *P. persimilis* have been measured compared to measurements of 14 females collected in Mauritius and to large numbers of specimens from other countries.

**Tribe Typhlodromipsini Chant & McMurtry**

**Typhlodromipsini Chant & McMurtry 2005c: 318.**

**Genus Typhlodromips De Leon**

Typhlodromipsini Chant & McMurtry 2005c: 318; 2006b: 137; 2007: 55.

**Typhlodromips culmulus (van der Merwe)**

*Amblyseius (Amblyseius) culmulus* van der Merwe 1968: 132; Ueckermann & Loots 1988: 157.

*Typhlodromips culmulus*, Moraes *et al.* 1986: 139; 2004a: 210; Chant & McMurtry 2005c: 327; Chant & McMurtry 2007: 61.

This species belongs to the *culmulus* species group as the spermatheca has a calyx shallow dish-shaped. This species group contains only 10 species (Chant and McMurtry 2005c).

*Typhlodromips culmulus* is mentioned from western and southern Africa (Demite *et al.* 2019). It was found recently in Mauritius (Kreiter *et al.* 2018a). Species of this quite large genus are supposed to all belong to the type III (McMurtry and Croft 1997; McMurtry *et al.* 2013), i.e. a polyphagous generalist predator. However, the biology of *T. culmulus* remains totally unknown. This is the first mention of this species from La Réunion Island.

**Specimens examined:** 105 ♀♀ + 18 ♂♂ + 6 im., 10 ♀♀ + 3 ♂♂ measured. Petite Île - Piton Bloc, Yébo Luguy farm (aasl 973 m, Long 55°34'64" E, Lat 21°18'64" S), 1 ♀ on *I. obscura*, 28/4/2016; 6 ♀♀ + 1 im. on *P. lanceolata*, 2 ♀♀ on *R. raphanistrum*, 1 ♀ on *I. obscura*, 18/10/2016; Le Tampon - Grand Tampon, Janick Bénard farm (aasl 861 m, Long 55°32'90" E, Lat 21°12'80" S), 1 ♀ on *Begonia cucullata*, 7/1/2016; 4 ♀♀ + 2 ♂♂ + 1 im. on *R. raphanistrum*, 1 ♀ on *B. pilosa*, 24/5/2016; 8 ♀♀ + 1 im. on *R. raphanistrum*, 2 ♀♀ on *B. pilosa*, 20/9/2016; Saint-Pierre - Bassin Martin, Armefhor Station (aasl 450 m, Long 55°31'9" E, Lat 21°18'14" S), 18 ♀♀ + 2 ♂♂ + 1 im. on various weeds and 62 ♀♀ + 13 ♂♂ + 2 im. on *C. rotundus*, 3/8/2017; Saint-Pierre - Bassin-Plat CIRAD Research Station (aasl 153 m, Long 55°29'18" E, Lat 21°19'25" S), 1 ♀ in plot BM, 23/8/2016; 1 ♀ on *A. viridis*, 20/6/2017.

**Remarks:** measurements of morphological characters of *T. culmulus* female and male specimens from La Réunion (Table 14) are very close from measurements for specimens from neighbouring countries, especially for specimens from Kenya and Mauritius and except for specimens from South Africa which are larger in van der Merwe (1968) but very close for the measurements of type material provided by Moraes *et al.* (2007).
<table><caption>**Table 13** Character measurements of adult females of *Phytoseiulus persimilis* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th>Characters</th><th>La Réunion<br>5, this study)</th><th>Mauritius<br>14</th><th>VCW 1<br>29</th><th>VCW 2<br>?</th><th>Types Algeria<br>3</th></tr></thead><tbody><tr><td>Dsl</td><td><strong>349</strong> (295 – 383)</td><td>349 (325 – 375)</td><td>339 (316 – 369)</td><td>322 (314 – 330)</td><td>300 – 330</td></tr><tr><td>Dsw</td><td><strong>245</strong> (245 – 245)</td><td>217 (178 – 245)</td><td>227 (196 – 256)</td><td>224 (215 – 232)</td><td>-</td></tr><tr><td>j1</td><td><strong>24</strong> (23 – 25)</td><td>25 (20 – 28)</td><td>26 (22 – 32)</td><td>28 (25 – 32)</td><td>25 – 35</td></tr><tr><td>j3</td><td><strong>44</strong> (43 – 45)</td><td>42 (37 – 50)</td><td>42 (31 – 51)</td><td>42 (38 – 46)</td><td>45 – 55</td></tr><tr><td>j4</td><td><strong>45</strong> (40 – 50)</td><td>46 (43 – 50)</td><td>52 (39 – 72)</td><td>50 (48 – 52)</td><td>45 – 55</td></tr><tr><td>j5</td><td><strong>57</strong> (53 – 60)</td><td>65 (55 – 75)</td><td>77 (62 – 92)</td><td>69 (65 – 74)</td><td>65 – 70</td></tr><tr><td>j6</td><td><strong>133</strong> (120 – 145)</td><td>145 (128 – 155)</td><td>150 (114 – 161)</td><td>152 (145 – 160)</td><td>145 – 160</td></tr><tr><td>J5</td><td><strong>5</strong></td><td>5 (4 – 6)</td><td>5 (4 – 8)</td><td>6 (5 – 6)</td><td>5</td></tr><tr><td>r3</td><td><strong>24</strong> (23 – 25)</td><td>24 (23 – 25)</td><td>24 (17 – 29)</td><td>23 (21 – 26)</td><td>-</td></tr><tr><td>R1</td><td><strong>19</strong> (15 – 23)</td><td>26 (25 – 31)</td><td>29 (22 – 33)</td><td>28 (25 – 32)</td><td>-</td></tr><tr><td>s4</td><td><strong>146</strong> (128 – 165)</td><td>165 (150 – 180)</td><td>163 (114 – 183)</td><td>165 (159 – 172)</td><td>145 – 160</td></tr><tr><td>S5</td><td><strong>25</strong> (23 – 28)</td><td>33 (25 – 40)</td><td>29 (20 – 37)</td><td>32 (25 – 38)</td><td>25 – 35</td></tr><tr><td>z2</td><td><strong>15</strong> (13 – 18)</td><td>14 (13 – 18)</td><td>12 (7 – 16)</td><td>12 (10 – 13)</td><td>10 – 15</td></tr><tr><td>z4</td><td><strong>55</strong> (50 – 60)</td><td>57 (53 – 63)</td><td>58 (39 – 68)</td><td>61 (57 – 65)</td><td>45 – 55</td></tr><tr><td>z5</td><td><strong>11</strong> (10 – 13)</td><td>11 (8 – 13)</td><td>10 (7 – 15)</td><td>9 (8 – 12)</td><td>10 – 15</td></tr><tr><td>Z1</td><td><strong>100</strong> (68 – 120)</td><td>105 (95 – 115)</td><td>107 (94 – 124)</td><td>110 (105 – 115)</td><td>80 – 90</td></tr><tr><td>Z4</td><td><strong>128</strong> (120 – 133)</td><td>129 (122 – 139)</td><td>135 (119 – 152)</td><td>134 (131 – 138)</td><td>115 – 125</td></tr><tr><td>Z5</td><td><strong>120</strong> (110 – 128)</td><td>121 (110 – 128)</td><td>125 (113 – 137)</td><td>126 (120 – 132)</td><td>115 – 125</td></tr><tr><td>st1-st1</td><td><strong>50</strong> (48 – 53)</td><td>53 (45 – 58)</td><td>54 (47 – 61)</td><td>-</td><td>-</td></tr><tr><td>st1-st3</td><td><strong>76</strong> (75 – 78)</td><td>71 (60 – 78)</td><td>74 (67 – 82)</td><td>-</td><td>73</td></tr><tr><td>st2-st2</td><td><strong>75</strong> (70 – 80)</td><td>79 (75 – 88)</td><td>80 (69 – 91)</td><td>-</td><td>-</td></tr><tr><td>st2-st3</td><td><strong>34</strong> (33 – 37)</td><td>33 (30 – 35)</td><td>33 (28 – 37)</td><td>-</td><td>32</td></tr><tr><td>st3-st3</td><td><strong>83</strong> (73 – 93)</td><td>90 (80 – 98)</td><td>94 (83 – 104)</td><td>-</td><td>93</td></tr><tr><td>st4-st4</td><td><strong>100</strong> (98 – 103)</td><td>101 (90 – 115)</td><td>99 (82 – 120)</td><td>-</td><td>-</td></tr><tr><td>st5-st5</td><td><strong>69</strong> (50 – 88)</td><td>82 (75 – 88)</td><td>86 (76 – 82)</td><td>-</td><td>86</td></tr><tr><td>Lisl</td><td><strong>33</strong></td><td>41 (35 – 50)</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Lisw</td><td><strong>5</strong></td><td>4 (3 – 5)</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Sisl</td><td><strong>20</strong></td><td>18 (13 – 20)</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Asl</td><td><strong>75</strong></td><td>76 (50 – 88)</td><td>81 (69 – 90)</td><td>93 (89 – 98)</td><td>-</td></tr><tr><td>Asw anus</td><td><strong>88</strong></td><td>76 (63 – 88)</td><td>77 (63 – 93)</td><td>-</td><td>-</td></tr><tr><td>JV5</td><td><strong>48</strong> (43 – 50)</td><td>46 (43 – 50)</td><td>45 (32 – 62)</td><td>40 (35 – 44)</td><td>-</td></tr><tr><td>SgeIV</td><td><strong>61</strong> (53 – 70)</td><td>84 (75 – 95)</td><td>83 (69 – 94)</td><td>84 (80 – 91)</td><td>90</td></tr><tr><td>StiIV</td><td><strong>47</strong> (43 – 50)</td><td>43 (30 – 50)</td><td>44 (38 – 48)</td><td>45 (40 – 48)</td><td>50</td></tr><tr><td>StIV</td><td><strong>116</strong> (100 – 133)</td><td>134 (125 – 140)</td><td>123 (108 – 132)</td><td>126 (110 – 135)</td><td>125</td></tr><tr><td>Scl</td><td><strong>30</strong></td><td>31 (25 – 38)</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Scw</td><td><strong>8</strong></td><td>9 (7 – 13)</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Fdl</td><td><strong>23</strong> (20 – 25)</td><td>26 (25 – 33)</td><td>-</td><td>-</td><td>-</td></tr><tr><td>No teeth Fd</td><td><strong>6</strong></td><td>6</td><td>-</td><td>-</td><td>-</td></tr><tr><td>Mdl</td><td><strong>23</strong> (20 – 25)</td><td>26 (25 – 30)</td><td>-</td><td>-</td><td>-</td></tr><tr><td>No teeth Md</td><td><strong>3</strong></td><td>3</td><td>-</td><td>-</td><td>-</td></tr></tbody></table>

Sources of measurements – Mauritius: Kreiter *et al.* (2018a); VCW 1 (Spain 7♀♀, Italy 4♀♀, Syngenta Bioline rearings 11♀♀, Tunisia 7♀♀): Okassa *et al.* (2010); VCW 2 (Sicily, Italy; Valparaiso, Chile; California, USA and Sydney, Australia): Takahashi & Chant (1993); Types Algeria: Athias-Henriot (1957); -: not provided.
<table><caption>**Table 14** Character measurements of adult females and males of *Typhlodromips culmulus* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="2">Characters</th><th colspan="5">♀</th><th colspan="3">♂</th></tr><tr><th>La Réunion<br>10, this study</th><th>Kenya<br>2</th><th>Mauritius<br>1</th><th>South Africa<br>1</th><th>Type South Africa<br>2</th><th>La Réunion<br>3, this study</th><th>South Africa 1<br>1</th><th>South Africa 2<br>1</th></tr></thead><tbody><tr><td>Dsl</td><td><strong>358</strong> (343 – 416)</td><td>300</td><td>313</td><td>355</td><td>334</td><td><strong>275</strong> (268 – 280)</td><td>289</td><td>268</td></tr><tr><td>Dsw</td><td><strong>233</strong> (198 – 332)</td><td>200</td><td>200</td><td>235</td><td>208</td><td><strong>193</strong> (157 – 218)</td><td>192</td><td>183</td></tr><tr><td>j1</td><td><strong>18</strong> (15 – 20)</td><td>18</td><td>18</td><td>21</td><td>19</td><td><strong>20</strong> (18 – 23)</td><td>23</td><td>19</td></tr><tr><td>j3</td><td><strong>20</strong> (17 – 23)</td><td>18</td><td>20</td><td>24</td><td>22</td><td><strong>21</strong> (20 – 22)</td><td>28</td><td>25</td></tr><tr><td>j4</td><td><strong>10</strong> (8 – 12)</td><td>9</td><td>10</td><td>11</td><td>9</td><td><strong>8</strong> (6 – 10)</td><td>10</td><td>9</td></tr><tr><td>j5</td><td><strong>11</strong> (9 – 12)</td><td>9</td><td>10</td><td>11</td><td>9</td><td><strong>11</strong> (10 – 14)</td><td>10</td><td>9</td></tr><tr><td>j6</td><td><strong>12</strong> (9 – 15)</td><td>10</td><td>10</td><td>11</td><td>13</td><td><strong>9</strong> (8 – 10)</td><td>10</td><td>9</td></tr><tr><td>J2</td><td><strong>14</strong> (13 – 15)</td><td>14</td><td>13</td><td>11</td><td>16</td><td><strong>10</strong> (8 – 13)</td><td>14</td><td>13</td></tr><tr><td>J5</td><td><strong>10</strong> (8 – 13)</td><td>7</td><td>10</td><td>11</td><td>9</td><td><strong>8</strong></td><td>10</td><td>9</td></tr><tr><td>r3</td><td><strong>20</strong> (15 – 27)</td><td>15</td><td>15</td><td>19</td><td>16</td><td><strong>19</strong> (15 – 23)</td><td>19</td><td>16</td></tr><tr><td>R1</td><td><strong>13</strong> (10 – 15)</td><td>12</td><td>13</td><td>15</td><td>13</td><td><strong>13</strong> (10 – 15)</td><td>15</td><td>11</td></tr><tr><td>s4</td><td><strong>29</strong> (25 – 35)</td><td>25</td><td>28</td><td>39</td><td>30 – 32</td><td><strong>22</strong> (20 – 23)</td><td>33</td><td>25</td></tr><tr><td>S2</td><td><strong>12</strong> (10 – 13)</td><td>12</td><td>13</td><td>11</td><td>13</td><td><strong>9</strong> (9 – 10)</td><td>10</td><td>9</td></tr><tr><td>S4</td><td><strong>11</strong> (9 – 12)</td><td>8</td><td>10</td><td>11</td><td>9</td><td><strong>10</strong></td><td>10</td><td>9</td></tr><tr><td>S5</td><td><strong>10</strong> (8 – 14)</td><td>8</td><td>8</td><td>11</td><td>9</td><td><strong>8</strong></td><td>10</td><td>9</td></tr><tr><td>z2</td><td><strong>13</strong> (11 – 15)</td><td>12</td><td>13</td><td>15</td><td>13</td><td><strong>10</strong></td><td>14</td><td>13</td></tr><tr><td>z4</td><td><strong>11</strong> (10 – 13)</td><td>12</td><td>13</td><td>15</td><td>13 – 14</td><td><strong>11</strong> (10 – 12)</td><td>14</td><td>13</td></tr><tr><td>z5</td><td><strong>9</strong> (8 – 10)</td><td>7</td><td>10</td><td>11</td><td>9</td><td><strong>9</strong></td><td>10</td><td>9</td></tr><tr><td>Z1</td><td><strong>15</strong> (13 – 18)</td><td>10</td><td>13</td><td>11</td><td>13 – 15</td><td><strong>11</strong> (10 – 11)</td><td>10</td><td>9</td></tr><tr><td>Z4</td><td><strong>38</strong> (33 – 42)</td><td>35</td><td>33</td><td>44</td><td>38 – 39</td><td><strong>34</strong> (32 – 35)</td><td>40</td><td>38</td></tr><tr><td>Z5</td><td><strong>73</strong> (69 – 78)</td><td>70</td><td>75</td><td>82</td><td>69 – 74</td><td><strong>51</strong> (47 – 55)</td><td>62</td><td>54</td></tr><tr><td>st1-st1</td><td><strong>52</strong> (46 – 58)</td><td>-</td><td>50</td><td>-</td><td>-</td><td><strong>47</strong> (46 – 49)</td><td>-</td><td>-</td></tr><tr><td>st2-st2</td><td><strong>63</strong> (60 – 66)</td><td>58</td><td>60</td><td>65</td><td>52 – 54</td><td><strong>54</strong> (53 – 55)</td><td>-</td><td>-</td></tr><tr><td>st3-st3</td><td><strong>72</strong> (68 – 77)</td><td>-</td><td>65</td><td>-</td><td>-</td><td><strong>60</strong> (58 – 63)</td><td>-</td><td>-</td></tr><tr><td>st1-st3 ♀ / st1-st5 ♂</td><td><strong>58</strong> (52 – 60)</td><td>52</td><td>55</td><td>65</td><td>47</td><td><strong>111</strong> (110 – 113)</td><td>-</td><td>-</td></tr><tr><td>st4-st4</td><td><strong>88</strong> (73 – 100)</td><td>-</td><td>60</td><td>-</td><td>-</td><td><strong>48</strong> (43 – 50)</td><td>-</td><td>-</td></tr><tr><td>st5-st5</td><td><strong>68</strong> (64 – 72)</td><td>66</td><td>68</td><td>85</td><td>60 – 62</td><td><strong>38</strong> (33 – 42)</td><td>-</td><td>-</td></tr><tr><td>Lisl</td><td><strong>20</strong> (18 – 21)</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="3">Not applicable</td></tr><tr><td>Lisw</td><td><strong>6</strong> (5 – 8)</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="3">Not applicable</td></tr><tr><td>Sisl</td><td><strong>10</strong> (8 – 11)</td><td>-</td><td>-</td><td>-</td><td>-</td><td colspan="3">Not applicable</td></tr><tr><td>Vsl</td><td><strong>118</strong> (113 – 125)</td><td>105</td><td>113</td><td>115</td><td>110</td><td><strong>113</strong> (110 – 118)</td><td>115</td><td>113</td></tr><tr><td>Vsw ZV2</td><td><strong>92</strong> (85 – 99)</td><td>82</td><td>85</td><td>90</td><td>82 – 85</td><td><strong>156</strong> (140 – 164)</td><td>90</td><td>151</td></tr><tr><td>Vsw anus</td><td><strong>86</strong> (80 – 95)</td><td>-</td><td>63</td><td>-</td><td>79 – 82</td><td><strong>70</strong> (65 – 75)</td><td>-</td><td>-</td></tr><tr><td>JV5</td><td><strong>39</strong> (35 – 43)</td><td>36</td><td>33</td><td>39</td><td>-</td><td><strong>29</strong> (25 – 38)</td><td>-</td><td>-</td></tr><tr><td>SgeI</td><td><strong>34</strong> (30 – 37)</td><td>-</td><td>30</td><td>-</td><td>30 – 32</td><td><strong>26</strong> (25 – 27)</td><td>-</td><td>24</td></tr><tr><td>SgeII</td><td><strong>30</strong> (28 – 34)</td><td>28</td><td>28</td><td>33</td><td>25 – 28</td><td><strong>28</strong> (25 – 30)</td><td>-</td><td>22</td></tr><tr><td>SgeIII</td><td><strong>32</strong> (28 – 39)</td><td>28</td><td>30</td><td>33</td><td>28</td><td><strong>28</strong> (27 – 30)</td><td>24</td><td>22</td></tr><tr><td>StiIII</td><td><strong>28</strong> (26–32)</td><td>28</td><td>-</td><td>-</td><td>22 – 26</td><td>- (broken)</td><td>24</td><td>25</td></tr><tr><td>SgeIV</td><td><strong>52</strong> (48 – 57)</td><td>46</td><td>48</td><td>54</td><td>47 – 49</td><td><strong>41</strong> (39 – 45)</td><td>42</td><td>35</td></tr><tr><td>StiIV</td><td><strong>39</strong> (28 – 46)</td><td>40</td><td>30</td><td>45</td><td>41</td><td><strong>37</strong> (35 – 39)</td><td>36</td><td>32</td></tr><tr><td>StIV</td><td><strong>58</strong> (50 – 65)</td><td>55</td><td>58</td><td>66</td><td>63</td><td><strong>50</strong> (50 – 51)</td><td>62</td><td>54</td></tr><tr><td>Scl</td><td><strong>3</strong></td><td>3</td><td>-</td><td>3</td><td>2</td><td colspan="3">Not applicable</td></tr><tr><td>Scw</td><td><strong>13</strong></td><td>14</td><td>-</td><td>12</td><td>-</td><td colspan="3">Not applicable</td></tr><tr><td>Fdl</td><td><strong>24</strong> (20 – 28)</td><td>-</td><td>25</td><td>-</td><td>22 – 25</td><td><strong>21</strong> (20 – 23)</td><td>20</td><td>-</td></tr><tr><td>No teeth Fd</td><td><strong>2 – 8</strong></td><td>7</td><td>5 – 6</td><td>8</td><td>11 – 12</td><td><strong>6</strong></td><td>8</td><td>-</td></tr><tr><td>Mdl</td><td><strong>27</strong> (25 – 30)</td><td>-</td><td>25</td><td>-</td><td>25 – 26</td><td><strong>19</strong> (18 – 23)</td><td>20</td><td>-</td></tr><tr><td>No teeth Md</td><td><strong>3 – 8</strong></td><td>3</td><td>3</td><td>3</td><td>3</td><td><strong>1</strong></td><td>1</td><td>-</td></tr><tr><td>Shaft</td><td colspan="5">Not applicable</td><td><strong>28</strong> (23 – 33)</td><td>20</td><td>19</td></tr></tbody></table>

Sources of measurements – For ♀♀: Kenya: El-Banhawy & Knapp (2011a); Mauritius: Kreiter *et al.* (2018a); South Africa: van der Merwe (1968); type material (the holotype and one paratype) collected in South Africa: Moraes *et al.* (2007a). For ♂♂: South Africa 1 (paratype male): van der Merwe (1968); South Africa 2: Moraes *et al.* (2007a); - : not provided.
## Genus Scapulaseius Karg & Oomen-Kalsbeek

*Scapulaseius* Karg & Oomen-Kalsbeek 1987: 132.

*Amblyseius* (*Scapulaseius*) Karg & Oomen-Kalsbeek 1987: 132.

*newsami* group of *Typhlodromus* (*Amblyseius*), Chant 1959: 95.

*markwelli* species group of *Amblyseius*, Schicha 1987: 25.

*japonicus* species group of *Amblyseius*, Schicha 1987: 26.

*oguroi* species group of *Amblyseius*, Wu & Ou 1999: 103.

*Scapulaseius*, Chant & McMurtry 2005c: 331; 2007: 65.

## Scapulaseius reptans (Blommers)

*Amblyseius* (*Amblyseius*) *reptans* Blommers 1974: 145.

*Typhlodromips* *reptans*, Moraes *et al.* 1986: 146; Moraes *et al.* 2004a: 222.

*Scapulaseius* *reptans*, Chant & McMurtry 2005c: 335; Chant & McMurtry 2007: 68.

This species belongs to the *ficilocus* species group of the genus *Scapulaseius* as the setae *R1* are inserted on lateral integument of adult female (Chant and McMurtry 2005c). This species group contains 40 species with setae *R1* inserted on the lateral integument of the adult female.

This species is mentioned only from the Indian Ocean area, Madagascar (Blommers 1974), La Réunion (Quilici *et al.* 2000) and recently Mauritius (Kreiter *et al.* 2018a). Species of this genus *Scapulaseius* are supposed to be of type III (McMurtry and Croft 1997; McMurtry *et al.* 2013), i.e. polyphagous generalist predators. However, the biology of *S. reptans* remains unknown. *S. reptans* was already mentioned from La Réunion by Quilici *et al.* (2000). Exact indications of locations were provided in this paper but without any measurements of specimens collected. They are provided for specimens collected during this study and listed in table 15.

**Specimens examined:** 2 ♀♀ + 1 ♂ in total, all measured. Ravine Langevin – Grand-Galet Waterfall (aasl 850 m, Long 55°21'33" E, Lat 21°17'47" S), 1 ♀ + 1 ♂ on *D. incanum*, 11/12/2016; Petite Île – Piton Bloc, Yébo Luguy farm (aasl 973 m, Long 55°34'64" E, Lat 21°18'64" S), 1 ♀ + 1 im. on *L. camara*, 9/1/2017.

**Remarks:** measurements of the two female and the male specimens in the table 15 show a great overlap with measurements mentioned in Kreiter *et al.* (2018a) on specimens from Mauritius, except for setae S2, S4 and S5 which are longer in La Réunion specimens. Setae of specimens from La Réunion are generally longer, except for *j1* and the macroseta of genu I that are nearly the same as for specimens from Mauritius. However, sternogenital shield is longer in Mauritius specimens. Both specimens from La Réunion and Mauritius have globally greater dimensions than type specimens from Madagascar, especially setae.

Ferragut and Baumann (2019) discussed a possible synonymy between *S. reptans* and *S. asiaticus* (Evans) recently. Our specimens of La Réunion Island markedly differ from specimens of *S. asiaticus* collected recently in Vietnam (Kreiter *et al.* in prep.) and we therefore disagree with this possible synonymy with arguments that will be developed in a future paper.

## Tribe Amblyseiini Muma

Amblyseiini, Muma, 1961: 68.

## Subtribe Amblyseiina Muma

Amblyseiina Muma, 1961: 69.

## Genus Transeius Chant & McMurtry

*Transeius* Chant & McMurtry, 2004a: 181.
<table><caption>**Table 15** Character measurements of adult females and one adult male of *Scapulaseius reptans* collected in this study with those in previous studies (localities followed by the number of specimens measured).</caption><thead><tr><th rowspan="2">Characters</th><th colspan="3">♀</th><th colspan="3">♂</th></tr><tr><th>La Réunion<br>2, this study</th><th>Madagascar<br>5</th><th>Mauritius<br>2</th><th>La Réunion<br>1, this study</th><th>Madagascar<br>3</th><th>Mauritius<br>1</th></tr></thead><tbody><tr><td>Dsl</td><td><strong>321</strong> (308 – 335)</td><td>290</td><td>303 – 305</td><td><strong>250</strong></td><td>250</td><td>255</td></tr><tr><td>Dsw</td><td><strong>210</strong> (198 – 223)</td><td>190</td><td>185 – 188</td><td><strong>150</strong></td><td>170</td><td>175</td></tr><tr><td>j1</td><td><strong>23</strong> (20 – 25)</td><td>21</td><td>23</td><td><strong>15</strong></td><td>16</td><td>18</td></tr><tr><td>j3</td><td><strong>25</strong> (25 – 25)</td><td>15</td><td>20 – 23</td><td><strong>20</strong></td><td>20</td><td>25</td></tr><tr><td>j4</td><td><strong>10</strong></td><td>8</td><td>8</td><td><strong>10</strong></td><td>7</td><td>10</td></tr><tr><td>j5</td><td><strong>9</strong> (8 – 10)</td><td>7</td><td>8</td><td><strong>8</strong></td><td>8</td><td>10</td></tr><tr><td>j6</td><td><strong>13</strong> (13 – 13)</td><td>10</td><td>10</td><td><strong>10</strong></td><td>8</td><td>10</td></tr><tr><td>J2</td><td><strong>13</strong> (13 – 13)</td><td>10</td><td>8 – 9</td><td><strong>13</strong></td><td>8</td><td>10</td></tr><tr><td>J5</td><td><strong>9</strong> (8 – 10)</td><td>7</td><td>8</td><td><strong>5</strong></td><td>3</td><td>6</td></tr><tr><td>z2</td><td><strong>21</strong> (20 – 23)</td><td>16</td><td>18 – 20</td><td><strong>23</strong></td><td>16</td><td>13</td></tr><tr><td>z4</td><td><strong>25</strong> (25 – 25)</td><td>16</td><td>23</td><td><strong>20</strong></td><td>16</td><td>13</td></tr><tr><td>z5</td><td><strong>10</strong></td><td>18</td><td>8</td><td><strong>8</strong></td><td>6</td><td>10</td></tr><tr><td>Z1</td><td><strong>13</strong> (13 – 13)</td><td>10</td><td>10</td><td><strong>10</strong></td><td>9</td><td>13</td></tr><tr><td>Z4</td><td><strong>61</strong> (60 – 63)</td><td>48</td><td>56 – 53</td><td><strong>40</strong></td><td>20</td><td>33</td></tr><tr><td>Z5</td><td><strong>79</strong> (75 – 83)</td><td>70</td><td>72 – 75</td><td><strong>58</strong></td><td>35</td><td>50</td></tr><tr><td>s4</td><td><strong>33</strong></td><td>25</td><td>28</td><td><strong>28</strong></td><td>20</td><td>25</td></tr><tr><td>S2</td><td><strong>24</strong> (23 – 25)</td><td>18</td><td>22 – 25</td><td><strong>20</strong></td><td>20</td><td>10</td></tr><tr><td>S4</td><td><strong>20</strong></td><td>15</td><td>18 – 23</td><td><strong>20</strong></td><td>15</td><td>10</td></tr><tr><td>S5</td><td><strong>20</strong> (18 – 23)</td><td>14</td><td>18 – 20</td><td><strong>18</strong></td><td>13</td><td>9</td></tr><tr><td>r3</td><td><strong>23</strong> (18 – 28)</td><td>15</td><td>18</td><td><strong>15</strong></td><td>14</td><td>15</td></tr><tr><td>R1</td><td><strong>14</strong> (13 – 15)</td><td>15</td><td>13</td><td><strong>13</strong></td><td>13</td><td>13</td></tr><tr><td>st1-st1</td><td><strong>50</strong></td><td>-</td><td>50</td><td><strong>30</strong></td><td>-</td><td>47</td></tr><tr><td>st2-st2</td><td><strong>60</strong> (58 – 63)</td><td>-</td><td>60</td><td><strong>53</strong></td><td>-</td><td>53</td></tr><tr><td>st1-st3 ♀ / st1-st5 ♂</td><td><strong>56</strong> (55 – 58)</td><td>-</td><td>53 – 56</td><td><strong>50</strong></td><td>-</td><td>55</td></tr><tr><td>st3-st3</td><td><strong>66</strong> (63 – 70)</td><td>-</td><td>65</td><td><strong>78</strong></td><td>-</td><td>105</td></tr><tr><td>st4-st4</td><td><strong>79</strong> (73 – 85)</td><td>-</td><td>65 – 78</td><td><strong>38</strong></td><td>-</td><td>40</td></tr><tr><td>st5-st5</td><td><strong>61</strong> (60 – 63)</td><td>-</td><td>60</td><td><strong>38</strong></td><td>-</td><td>35</td></tr><tr><td>Lisl</td><td><strong>20</strong> (18 – 23)</td><td>-</td><td>18 – 23</td><td colspan="3">Not applicable</td></tr><tr><td>Lisw</td><td><strong>5</strong></td><td>-</td><td>4 – 5</td><td colspan="3">Not applicable</td></tr><tr><td>Sisl</td><td><strong>11</strong> (10 – 13)</td><td>-</td><td>13 – 11</td><td colspan="3">Not applicable</td></tr><tr><td>Vsl</td><td><strong>115</strong> (115 – 115)</td><td>98</td><td>98 – 100</td><td><strong>100</strong></td><td>105</td><td>113</td></tr><tr><td>Vsw ZV2</td><td><strong>81</strong> (78 – 85)</td><td>78</td><td>85 – 87</td><td><strong>133</strong></td><td>-</td><td>135</td></tr><tr><td>Vsw anus</td><td><strong>75</strong> (75 – 75)</td><td>-</td><td>68 – 70</td><td><strong>75</strong></td><td>-</td><td>60</td></tr><tr><td>JV5</td><td><strong>34</strong> (33 – 35)</td><td>24</td><td>28</td><td><strong>18</strong></td><td>17</td><td>20</td></tr><tr><td>SgeI</td><td><strong>24</strong> (23 – 25)</td><td>-</td><td>23</td><td><strong>25</strong></td><td>-</td><td>25</td></tr><tr><td>SgeII</td><td><strong>15</strong></td><td>-</td><td>13 – 15</td><td><strong>15</strong></td><td>-</td><td>20</td></tr><tr><td>SgeIII</td><td><strong>19</strong> (18 – 20)</td><td>-</td><td>18</td><td><strong>18</strong></td><td>-</td><td>25</td></tr><tr><td>StiIII</td><td><strong>18</strong> (15 – 20)</td><td>-</td><td>18</td><td>-</td><td>-</td><td>20</td></tr><tr><td>SgeIV</td><td><strong>31</strong> (30 – 33)</td><td>27</td><td>28</td><td><strong>25</strong></td><td>19</td><td>35</td></tr><tr><td>StiIV</td><td><strong>28</strong> (28 – 28)</td><td>20</td><td>23 – 25</td><td><strong>23</strong></td><td>15</td><td>20</td></tr><tr><td>StIV</td><td><strong>59</strong> (55 – 63)</td><td>50</td><td>50 – 53</td><td><strong>45</strong></td><td>45</td><td>48</td></tr><tr><td>Scl</td><td><strong>30</strong></td><td>45</td><td>30 – 35</td><td colspan="3">Not applicable</td></tr><tr><td>Scw</td><td><strong>10</strong></td><td>2</td><td>2</td><td colspan="3">Not applicable</td></tr><tr><td>Fdl</td><td><strong>28</strong> (25 – 30)</td><td>26</td><td>25</td><td><strong>20</strong></td><td>-</td><td>20</td></tr><tr><td>No teeth Fd</td><td><strong>10</strong></td><td>8</td><td>9</td><td><strong>8</strong></td><td>8</td><td>8</td></tr><tr><td>Mdl</td><td><strong>23</strong> (20 – 25)</td><td>26</td><td>25</td><td><strong>23</strong></td><td>-</td><td>23</td></tr><tr><td>No teeth Md</td><td><strong>3</strong></td><td>3</td><td>3</td><td><strong>1</strong></td><td>1</td><td>1</td></tr><tr><td>Shaft</td><td colspan="3">Not applicable</td><td><strong>15</strong></td><td>15</td><td>17</td></tr></tbody></table>

Sources of measurements – For ♀♀: Madagascar: Blommers (1974); Mauritius: Kreiter *et al.* (2018a). For ♂♂: Madagascar: Blommers (1974); Mauritius: Kreiter *et al.* (2018a). - : not provided.
**Transeius maelliae Kreiter n. sp.**

Zoobank: C6EA34F5-9601-4208-8201-B14CE6C6FC5D

**Diagnosis** — *Transeius maelliae* Kreiter *n. sp.* belongs to the subfamily Amblyseiinae (absence of dorsolateral setae *z3* and *s6* and caudoventral seta *JV3*), to the tribe Amblyseiini (setae *j3*, *s4*, *Z4* and *Z5* longer than other setae, ratio *s4/Z1* > 3.1, many teeth on the fixed cheliceral digit and macrosetae on several legs in addition of the three on leg IV), to the subtribe Amblyseiina (sternal shield as long as wide, ventrianal shield longer than wide, seta *J2* present, genital shield almost as wide as ventrianal shield, ventral shields generally smooth, macrosetae on all legs, setae *j5*, *J2*, *S2*, *S4*, *S5* and *Z1* present), to the genus *Transeius* (ratio *s4/S2* < 2.7, Setae *S5* present, spermathecal with atrium not bifurcate) (Chant and McMurtry 2007). Seta *z4* is not as long as 2/3 the distance between its base and that of seta *s4* that allows to classify this new species in the species group *bellottii* (Chant and McMurtry 2004a). The bell-shaped spermatheca keys to the species subgroup *bellottii*.

*Transeius maelliae* Kreiter *n. sp.* is quite similar to the other new species described further, *T. mickaeli* Kreiter *n. sp.* with which it was at first confused early on during the identification process. *T. maelliae* *n. sp.* is different from *T. mickaeli* *n. sp.* by: having only five solenostomes instead of seven, a longer seta *z4*, almost double, the shape of the spermatheca also bell-shaped but with an undistinct atrium, longer setae *JV5*, less teeth on both digits of chelicerae and very slightly reticulated ventral shields (see table 27).

In the species subgroup *bellottii*, the species closest to *T. maelliae* *n. sp.* is *Transeius jujae* El-Banhawy and Knapp. *T. maelliae* *n. sp.* resembles to *T. jujae*, having similar lengths for setae *j3*, *s4*, *Z4*, *Z5*, and *JV5*, and the dimensions of the spermatheca. The new species can be distinguished however by the longer length of setae *z2*, above all by the length of setae *z4* which is 2.5 as long as that of *T. jujae*, and by the shape of the spermatheca with an undifferentiated atrium and walls of calyx slightly converging apically in the new species opposed to a distinct atrium and walls of calyx strictly parallel in *T. jujae*. In addition, a macroseta is existing in the genu of leg I of *T. maelliae* *n. sp.* and lacking in specimens of *T. jujae*, the ventrianal shield is wider and the dorsal shied is totally smooth in that species.

**Description of the adult female**

$n = 4$ (Figs. 2a – d)

**Dorsum** — (Fig. 2a). Dorsal shield fused with peritremal shield in the level of *j1* position, smooth except some slight reticulation near the anterolateral margins of the dorsal shield, 393 (368 – 418) long and 221 (203 – 243) wide, with five solenostomes (*gd1*, *gd5*, *gd6*, *gd8*, and *gd9*), 13 pairs of poroids or lyrifissures, 17 pairs of dorsal setae and two pairs of sub-lateral setae off the dorsal shield: *j1* 30 (25 – 35), *j3* 39 (38 – 40), *j4* 8, *j5* 8, *j6* 11 (10 – 13), *J2* 9 (8 – 10), *J5* 10, *z2* 16 (14 – 18), *z4* 30 (28 – 33), *z5* 7 (6 – 8), *Z1* 11 (10 – 11), *Z4* 53 (50 – 55), *Z5* 75 (70 – 81), *s4* 47 (43 – 51), *S2* 21 (20 – 21), *S4* 15 (13 – 15), *S5* 13 (10 – 15), *r3* 18 (18 – 20), *R1* 19 (18 – 20). All setae smooth except *Z5* which is very slightly serrated (with few barbs) in only two of the four specimens collected.

**Peritreme** — (Fig. 2a). Reaching the level of *j1*.

**Venter** — (Fig. 2b). All ventral shields smooth. Sternal shield with three pairs of setae and two pairs of lyrifissures; one pair of sternal setae on a small metasternal shield with one poroid; posterior margin of the sternal shield straight to slightly convex. Distances between *st1-st1* 66 (65 – 70), *st2-st2* 78, *st3-st3* 84 (80 – 85), *st1-st3* 73 (70-75), *st4-st4* 84 (80-95). Genital shield length 122 (113 – 128), width at the level of *st5* 83 (78 – 86), width at the level of the posterior corners 88 (83 – 93), distance *st5-st5* 75 (73 – 78). Two pairs of metapodal shields 22 (20 – 23) long and 5 (4 – 6) wide for the larger and 11 (10 – 11) long the slender shield. Ventrianal shield with three pairs of preanal setae (*JV1*, *JV2* and *ZV2*), two small oblong pre-anal solenostomes 37-40 apart. Cribrum spicules on three lines. Membrane surrounding ventrianal shield with four pairs of setae (*ZV1*, *ZV3*, *JV4* and *JV5*), and seven pairs of round to oblong poroids around