1✉ CBGP, Institut Agro (SupAgro), INRAE, CIRAD, IRD, Univ Montpellier, Montpellier, France.
2CIRAD, Université de Montpellier, Unité Hortsys, Station de Bassin-Plat, 97410 Saint-Pierre, La Réunion, France.
3INRAPE, Moroni, Grande Comore, Union des Comores.
2021 - Volume: 61 Issue: 1 pages: 94-114
https://doi.org/10.24349/acarologia/20214419Mites of the family Phytoseiidae are all predatory species on phytophagous mites and small insects like thrips and whiteflies, on commercial plants and the wild vegetation. Several species are biological control agents for the control of pest organisms in both open and protected crops all around the world (McMurtry and Croft 1997; McMurtry et al. 2013).
This family is widespread around the world, presents on all continents except Antarctica, and consists of 2,521 valid species in 94 genera belonging to three subfamilies (Demite et al. 2020).
Biodiversity surveys in poorly investigated areas is still urgently needed and might result in the discovery of additional species potentially useful for biological control as well as having more information on the biodiversity of these areas (Kreiter et al. 2018a, b, c, 2020a, b, c, d; Kreiter and Abo-Shnaf 2020a, b, Kreiter et al. 2021).
In these perspectives, the more interesting area are probably those with a high level of biodiversity. Most of the Indian Ocean constitutes one of the highest world biodiversity area, those area being called hotspots, concept defined by Myers (1988) in order to identify the most immediately important areas for biodiversity conservation. The common characteristics of these hotspots is that they hold high endemism levels and have lost at least 70% of their original natural vegetation (Myers et al. 2000). Knowledge of the phytoseiid diversity in these high interest areas in the context of global climate changes may contribute to identify potential biological control agents (BCA) and future establishment of conservation programs.
Located in the Indian Ocean at around 1,000 km from the northern coast of Madagascar, 165 km from Mayotte and about only 70 km from Anjouan and Grande Comore Islands, Mohéli (Mwali in Shicomori language) Island is one of the four main islands constituting Comoros Archipelago, with Mayotte, Anjouan and Grande Comore. No phytoseiid species are known from this island.
The objective of this paper is to present the phytoseiid species reported in a survey conducted in early December 2018 in Mohéli Island.
The survey took place in Mohéli in the very beginning of December 2018. Plant inhabiting mites were collected from cultivated and wild plants in few locations mainly in the southern part of the island.
Mites were directly collected on leaves with a fine brush with or without a pocket lens or a stereoscopic microscope when available (large leaves and herbaceous plants) or by beating the plants (mainly shrubs and trees with very small or spiny leaves) and collecting the mites in a black plastic rectangular saucer 45 x 30 cm (Ref. STR 45, BHR, 71370 Saint-Germain-du-Plain, France). Collected mites were then transferred with a fine brush into small plastic vials containing 1.5 ml of 70% ethanol.
The mites were then all slide-mounted in Hoyer's medium (Walter and Krantz 2009), the slides were dried at 45-50 oC for at least two weeks and then all examined and identified using a phase and interferential contrast microscope (DMLB, Leica Microsystèmes SAS, Nanterre, France). Characters of specimens were measured using a graded eyepiece (Leica, see above).
Chant and McMurtry's (1994, 2007) concepts of the taxonomy of the family Phytoseiidae for identification and the world catalogue database of Demite et al. (2014, 2020) for distribution and information on descriptions and re-descriptions were used. The setal nomenclature system adopted was that of Lindquist & Evans (1965) and Lindquist (1994) as adapted by Rowell et al. (1978) and Chant & Yoshida-Shaul (1989) for the dorsal surface and by Chant & Yoshida-Shaul (1991) for the ventral surface. Pore (= solenostome) and poroid (= lyrifissure) notations are that of Athias-Henriot (1975). Macrosetal notation (Sge = genual macroseta; Sti = tibial macroseta; St = tarsal macroseta) are that of Muma and Denmark (1970). Numbers of teeth on the fixed and movable cheliceral digits do not include the respective apical teeth. Setae not referred to in Results section should be considered as absent. All measurements are given in micrometres (µm) and presented with the mean in bold followed by the range in parenthesis. Type of spermatheca or insemination apparatus are that of Denmark and Evans 2011. Only some species with only few measurements mentioned in the literature are provided in this paper. Classification of plants follows the APG IV classification of 2016 (ex. Byng et al. 2018). Specimens collected in fields in Mohéli Island within these surveys were all identified. Only very few single males or immatures collected during this study are not taken into account.
Specimens of each species are deposited in the mite collections of Montpellier SupAgro conserved in UMR CBGP INRA/IRD/CIRAD/SupAgro/University of Montpellier.
The following abbreviations are used in this paper for morphological characters: dsl = dorsal shield length just above j1 to just below J5; dsw = dorsal shield width at the level of s4; Z4 ser., Z5 ser. = Z4, Z5 serrated (if Z4 and Z5 without ser. = not serrated); gensl = genital shield length; genswst5 = genital shield width at level of steae st5; gensw post. cor. = genital shield width at level of posterior corners; lisl = primary or largest inguinal sigilla (= ''metapodal plate'') length; lisw = primary or largest inguinal sigilla (= ''metapodal plate'') width; sisl = secondary or smallest inguinal sigilla (= ''metapodal plate'') length; vsl = ventrianal shield length; gv3 – gv3 = distance between solenostomes gv3 on the ventrianal shield; vsw ZV2 & vsw anus = ventrianal shield width at ZV2 level and at paranal setae level; scl: calyx total length; scw = calyx widest width; Fdl = fixed digit length; Mdl = movable digit length; Nb teeth Fd = number of teeth on the fixed digit; Nb teeth Md = number of teeth on the movable digit; Shaft = length of the shaft of spermatodactyl; toe = length of the toe; BCA = Biological control agent; aasl = altitude above sea level; imm.: immature.
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; INRAE = Institut National de Recherche pour l'Agriculture, l'Alimentation et l'Environnement; INRAPE = Institut National de Recherche pour l'Agriculture, la Pêche et l'Environnement; IRD = Institut de Recherche pour le Développement; MSA = Montpellier SupAgro, France; UMR = Unité Mixte de Recherche; UR = Unité de Recherche.
A total of 18 species had been found during this study, 16 presented thereafter (two new species, one Paragigagnathus n. sp. (Tribe Neoseiulini) and one Typhlodromalus n. sp. (Tribe Euseiini) found also in other islands will be published later on, five of them with new measurements compared to the only few references already available in the literature.
Amblyseiinae Muma 1961: 273.
Neoseiulini Chant & McMurtry 2003a: 6.
Neoseiulus Hughes 1948: 141.
Amblyseius (Amblyseius) teke Pritchard & Baker 1962: 239.
Amblyseius teke, Meyer & Rodrigues 1966: 30; Moraes et al. 1989a: 83, 1989b: 97.
Neoseiulus teke, Moraes et al. 1986: 98, 2004: 147; Chant & McMurtry 2003a: 37, 2007: 31.
Amblyseius (Amblyseius) bibens Blommers 1973: 111 (synonymy according to Ueckermann and Loots 1988).
Neoseiulus teke belongs to the barkeri species group and the womersleyi species subgroup (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 I. degenerans, but seems not efficient enough in field conditions (Nwilene and Nachman 1996). Quilici et al. (2000) had collected this species before in La Réunion Island and it was recovered recently by Kreiter et al. (2020d).
World distribution: Burundi, DR Congo, Ghana, Kenya, Malawi, Mozambique, La Réunion Island, Rwanda, Sierra Leone, South Africa, Tanzania, Zimbabwe.
Specimens examined: 2 specimens (2 ♀♀) in total. Fomboni, inside the town (15 m aasl, 12°17'29'' S, 43°44'35'' E), 1 ♀ on Abutilon hirtum (Lamarck) Sweet (Malvaceae), 2/XII/2018; Fomboni, University (25 m aasl, 12°17'3'' S, 43°44'34'' E), 1 ♀ on Zyzyphus mauritiana Lamarck (Malvaceae), 3/XII/2018.
Remarks: measurements of morphological characters of Neoseiulus teke female and male specimens from Mohéli Island are very close to measurements for specimens from neighbouring countries, especially from specimens from Mayotte (Kreiter et al. 2020a), La Réunion Island (Keiter et al. 2020d) and various countries in Africa, except for the holotype (Zannou et al. 2006) and specimens from South Africa which are larger (van der Merwe 1965).
Kampimodromini Kolodochka 1998: 59; Chant & McMurtry 2003b: 189, 2006: 137, 2007: 33.
Paraphytoseiina Chant & McMurtry 2003b: 211.
Paraphytoseius Swirski & Shechter 1961: 113; Moraes et al. 1986: 104, 2004a: 160; Chant & McMurtry 2003b: 216, 2007: 49.
Typhlodromus (Amblyseius) orientalis Narayanan, Kaur & Ghai 1960: 394.
Paraphytoseius orientalis, Moraes et al. 1986: 105, 2004a: 162; Chant & McMurtry 2003b: 220, 2007: 53.
Amblyseius ipomeai, Narayanan, Kaur & Ghai 1960: 394 (synonymy according to El-Banhawy 1984).
Paraphytoseius multidentatus, Swirski & Shechter 1961: 114 (synonymy according to Matthysse & Denmark 1981 in Denmark et al. 1999).
Paraphytoseius narayanani, Ehara 1967: 67 (synonymy according to Ehara & Ghai, in Ehara 1967).
This species belongs to the orientalis species group (Chant and McMurtry 2003b). Our specimens with relatively shorter setae s4, Z4 and Z5, having a distinctly short, thick, spatulate macroseta on genu I belong to the species P. orientalis. This species is widely distributed in tropical and subtropical areas in South America, Africa and Asia. It 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) had studied the life history of P. orientalis on the broad mite (Polyphagotarsonemus latus) (Banks) as prey and reported high predation rates on the eggs of P. latus, suggesting good potential for the control of this pest.
World distribution: Argentina, Brazil, Burundi, India, Japan, Kenya, La Réunion Island, Madagascar Island, Martinique Island, Mauritius Island, Mozambique, Rwanda.
Specimens examined: 3 specimens (2 ♀♀ and 1 ♂) in total. Bandar-Es-Salam, Les Abous Inn (23 m aasl, 12°17'37'' S, 43°45'27'' E), 2 ♀♀ and 1 ♂ on Solanum pyracanthos Lamarck (Solanaceae), 2/XII/2018.
Remarks: morphological and morphometric characters and all measurements fit well measurements in Ferragut and Baumann (2019) and Kreiter et al. (2020d). This species was described from Asia (Narayanan et al. 1960) and presented also in Mauritius (Ferragut and Baumann 2019; Kreiter et al. 2018a; Kreiter and Abo-Shnaf 2020b), Mayotte (Kreiter et al. 2020a), Vietnam (Kreiter et al. 2020b), and in La Réunion Island (Kreiter et al. 2020d).
Amblyseiinae Muma 1961: 273 and Amblyseiini Muma, Wainstein 1962: 26.
Amblyseiina Muma, Chant & McMurtry 2004: 179.
Amblyseius Berlese 1914: 143.
Amblyseius duplicesetus Moraes & McMurtry 1988: 13; Moraes et al. 2004a: 143, 2004b: 22; Zannou et al. 2007: 10; El-Banhawy & Knapp 2011: 25.
Amblyseius duplicisetus [sic], Chant & McMurtry 2004: 208, 2007: 78.
Like the previous species and for the same reasons, A. duplicesetus belongs to the largoensis species group and to the largoensis species subgroup (Chant and McMurtry 2004a, 2007). First of all found only in Kenya (Moraes and McMurtry 1988; Zannou et al. 2007; El-Banhawy and Knapp 2011) and more recently in Sri Lanka (Moraes et al. 2004a), its biology is totally unknown. This is the second mention of that species in Indian Ocean Islands after Anjouan (Kreiter et al. 2021).
World distribution: Kenya, Sri Lanka.
Specimens examined: 55 specimens (30 ♀♀, 11 ♂♂ and 14 imm) collected during this study. Fomboni, inside the town (15 m aasl, 12°17'29'' S, 43°44'35'' E), 1 ♀ and 1 ♂ on Annona muricata L. (Annonaceae), 2/XII/2018; Fomboni, Les-Hauts (60 m aasl, 12°17'29'' S, 43°44'35'' E), 2 ♂♂ and 3 imm. on an unknown host plant, 2/XII/2018; Fomboni, University (25 m aasl, 12°17'3'' S, 43°44'34'' E), 1 ♀ on Ricinus communis L. (Euphorbiaceae), 3/XII/2018; Hoani, inside village (38 m aasl, 12°17'3'' S, 43°44'34'' E), 5 ♀♀, 1 ♂ and 2 imm. on the same unknown host plant than above, 1 ♀, 1 ♂ and 2 imm. on A. muricata, 10 ♀♀, 2 ♂♂ and 1 imm. on Artocarpus altilis J.R. Forster and G. Forster (Moraceae) and 4 ♀♀, 1 ♂ and 3 imm. on Theobroma cacao L. (Malvaceae), 3/XII/2018; Bangoma, Les Hauts (137 m aasl, 12°17'18'' S, 43°43'41'' E), 1 ♂ on Cinnamomum odoratum Schäffer (Lauraceae), 3 ♀♀ and 2 imm. on Annona senegalensis Persoon (Annonaceae), 1 ♀ on A. muricata, 1 ♀ on Mangifera indica L. (Anacardiaceae), 2 ♀♀ and 1 ♂ on A. altilis and 1 ♀, 1 ♂ and 1 imm. on Persea americana Miller (Lauraceae), 4/XII/2018.
Remarks: this is the more abundant species found in several sites. All measurement values fit well those already published on this species with only very slight variations (Moraes and McMurtry 1988; Moraes et al. 2004a; Zannou et al. 2007; El-Banhawy and Knapp 2011; Kreiter et al. 2021). Measurement values of female specimens of Mohéli are very similar with values for specimens from Kenya and Sri Lanka, however with some shorter setae just like specimens of Anjouan (Kreiter et al. 2021). The male of that species will be redescribed in following papers.
Typhlodromus (Amblyseius) herbicolus Chant 1959: 84.
Amblyseius (Amblyseius) herbicolus, Muma 1961: 287.
Typhlodromus herbicolus, Hirschmann 1962: 23.
Amblyseius herbicolus, Moraes et al. 1986: 14, 1989: 79, 2004b: 27; Chant & McMurtry 2004: 208, 2007: 78.
Amblyseius impactus Chaudhri 1968: 553 (synonymy according to Daneshvar & Denmark 1982).
Typhlodromus (Amblyseius) amitae Bhattacharyya 1968: 677 (synonymy according to Denmark & Muma 1989).
Amblyseius deleoni Muma & Denmark 1970: 68 (synonymy according to Daneshvar & Denmark 1982).
Amblyseius giganticus Gupta 1981: 33 (synonymy according to Gupta 1986).
Amblyseius (Amblyseialus) thermophilus Karg 1991: 12 (synonymy according to El-Banhawy & Knapp 2011).
This species belongs to the largoensis species group as setae J2 and Z1 are present, setae s4 are minute and the ventrianal shield of the female is vase-shaped. It belongs to the largoensis species subgroup as setae Z4 are long, spermatheca has the calyx elongate and the female ventrianal shield is entire (Chant and McMurtry 2004).
Amblyseius herbicolus is widespread in all tropical and subtropical regions of the world. It is the second most abundant phytoseiid mite on Coffea arabica L. in Brazil, associated with Brevipalpus phoenicis (Geijskes), vector of the coffee ring spot virus and it was found to be an efficient predator (Reis et al. 2007). Amblyseius herbicolus is also found associated with the broad mite, P. latus, in crops such as chili pepper (Capsicum annuum L.) in Brazil and has also a good potential for controlling the pest. Rodriguez-Cruz et al. (2013) had studied biological, reproductive and life table parameters of A. herbicolus on three different diets: broad mites, castor bean pollen (Ricinus communis L.) and sun hemp pollen (Crotalaria juncea L.). The predator was able to develop and reproduce on all three these diets. However, its intrinsic growth rate was higher on broad mites and castor bean pollen. Feeding on alternative food such as pollen can facilitate the predator's mass rearing and maintain its population on crops when prey is absent or scarce. Many polyphagous generalist phytoseiid mites are important natural enemies because they can feed on plant provided pollen and various prey species, and thus persist in crops even in the absence of target pests (McMurtry et al. 2013). Hence, populations of these predators can be established in a crop by providing alternative food, thus increasing biological control. Alternative food affects P. latus control on chilli pepper plants by predatory mites (Duarte et al. 2015). Amblyseius herbicolus had high oviposition and population growth rates when fed with cattail pollen (Typha latifolia L.), chilli pepper pollen and bee-collected pollen, and a low rate on the alternative prey (Tetranychus urticae Koch). Supplementing pepper plants with pollen resulted in better control of broad mite populations (Duarte et al. 2015). Release of A. herbicolus on young plants with weekly addition of honeybee pollen or cattail pollen until plants produce flowers seems a viable strategy to sustain populations of this predator (Duarte et al. 2015).
World distribution: Argentina, Australia, Azores, Benin, Brazil, Burundi, Canary Islands, China, Colombia, Grande Comore Island, Costa Rica, Dominican Republic, Dr Congo, El Salvador, Ghana, Guadeloupe Island, Guatemala, Hawaii, Honduras, India, Iran, Kenya, Les Saintes, La Réunion and Madagascar Islands, Malawi, Malaysia, Martinique Island, New Caledonia Island, Papua New Guinea, Peru, Philippines, Portugal, Puerto Rico, Rwanda, Senegal, Singapore, South Africa, Spain, Taiwan, Thailand, Turkey, USA, Venezuela, West Indies.
Specimens examined: 18 specimens (10 ♀♀, 5 ♂♂ and 3 imm.) collected during this study. Hoani, inside village (38 m aasl, 12°17'3'' S, 43°44'34'' E), 1 ♂ on Theobroma cacao L. (Malvaceae), 3/XII/2018; Bangoma, top of the village (42 m aasl, 12°17'15'' S, 43°43'40'' E), 2 ♀♀ and 1 imm. on Morinda citrifolia L. (Rubiaceae), 1 ♂ on Dendrocnide moroides (Weddel) Chew (Urticacae), 4/XII/2020; Bangoma, Les Hauts (137 m aasl, 12°17'18'' S, 43°43'41'' E), 1 ♂ and 1 imm. on Asplenium petiolulatum Mettenius (Aspleniaceae), 1 ♀ and 1 imm. on Cinnamomum odoratum Schäffer (Lauraceae), 1 ♀ and 1 ♂ on Annona muricata L. (Annonaceae), 3 ♀♀ on Mangifera indica L. (Annacardiaceae), 1 ♂ on Litchi chinensis Sonnerat (Sapindaceae), 2 ♀♀ on Artocarpus altilis J.R. Forster and G. Forster (Moraceae) and 1 ♀ on an unknown host plant, 4/XII/2018.
Remarks: morphological and morphometric characters and all measurements fit well measurements provided in Kreiter et al. (2018b, 2020c, d). Amblyseius herbicolus was previously recorded in a lot of countries world-wide and especially in French West Indies (Moraes et al. 2000, Kreiter et al. 2006). It was first reported by Kreiter et al. (2018b) in the Comoros Archipelago in Grande Comore Island with two females collected. Amblyseius herbicolus was reported in the past from La Réunion Island from few specimens (Quilici et al. 1997, 2000) and more recently from a lot of specimens (Kreiter et al. 2020d). It is also reported recently from Vietnam (Kreiter et al. 2020c), Rodrigues and Maurice Islands (Kreiter and Abo-Shnaf 2020a, b) but only from females.
The five male specimens of that species collected in this study will be redescribed in a following paper.
Amblyseiopsis largoensis Muma 1955: 266.
Typhlodromus (Amblyseius) largoensis, Chant 1959: 96.
Amblyseius (Amblyseialus) largoensis, Muma 1961: 287.
Typhlodromus largoensis, Hirschmann 1962: 2.
Amblyseius (Amblyseius) largoensis, Ehara 1966: 22.
Amblyseius largoensis, Swirski & Golan 1967: 225; Moraes et al. 1986: 17, 2004b: 33; Chant & McMurtry 2004: 208, 2007: 78.
Amblyseius magnolia Muma 1961: 289 (synonymy by Denmark & Evans 2011).
Amblyseius sakalava Blommers 1976: 96 (synonymy by Ueckermann & Loots 1988).
Amblyseius amtalaensis Gupta 1977: 53 (synonymy by Gupta 1986).
Amblyseius largoensis belongs to the largoensis species group and to the largoensis species subgroup. It is widespread in all tropical and subtropical regions of the world and was the most abundant species collected by Moraes et al. (2000) in French Caribbean Islands and as a potential BCA of Raoiella indica Hirst in La Réunion Island (Moraes et al. 2012). Using morphometric analyses of 36 characters, molecular analyses and crossing tests, Navia et al. (2014) studied specimens collected in Brazil, La Réunion Island and Trinidad and Tobago to determine whether A. largoensis populations from different geographic origins belong to the same taxonomic entity. Though differences in the lengths of some setae were observed, molecular analyses and crossing experiments indicated that populations from Indian Ocean and America were conspecific. This species was previously recorded from Rodrigues Island by Kreiter and Abo-Shnaf (2020a), from Mauritius Island by Ferragut and Baumann (2019) and Kreiter and Abo-Shnaf (2020b) and from Mayotte by Kreiter et al. (2020a).
World distribution: this species is widely distributed in the tropical and subtropical regions of Africa, America, Asia and the Pacific Islands.
Specimens examined: 24 specimens (16 ♀♀, 5 ♂♂ and 3 imm.) collected during this study.Fomboni, inside the town (15 m aasl, 12°17'29'' S, 43°44'35'' E), 11 ♀♀, 4 ♂♂ and 1 imm. on Terminalia mantaly H. Perrier (Combretaceae), 1 imm. on Pentas lanceolata (Forskål) Deflers (Rubiaceae) and 1 ♀ on Artocarpus altilis J.R. Forster and G. Forster (Moraceae), 2/XII/2018; Bandar-Es-Salam, Les Abous Inn (23 m aasl, 12°17'37'' S, 43°45'27'' E), 2 ♀♀, 1 ♂ and 1 imm. on Citrus limon (L.) Burman (Rutaceae), 2/XII/2018; Hoani, inside village (38 m aasl, 12°17'3'' S, 43°44'34'' E), 2 ♀♀ on Amaranthus viridis L. (Amaranthaceae), 3/XII/2018.
Remarks: morphological and morphometric characters and all measurements of the Mohéli specimen fit well with those given in Zannou et al. (2007) for specimens from Africa, Navia et al. (2014) for specimens from Brazil, La Réunion and Trinidad & Tobago, by Ferragut and Baumann (2019) and Kreiter and Abo-Shnaf (2020b) for specimens from Mauritius Island and by Kreiter and Abo-Shnaf (2020a) for specimens from Rodrigues Island.
Euseiini Chant & McMurtry 2005a: 191.
Euseiina Chant & McMurtry 2005a: 209.
Amblyseius (Amblyseius) section Euseius Wainstein 1962: 15; Euseius De Leon 1966: 86.
Amblyseius baetae (Meyer & Rodrigues, 1966): 28.
Euseius baetae (Meyer & Rodrigues), Moraes et al. 1986: 37, 2001: 11, 2004b: 62; Chant & McMurtry 2005b: 215; 2007: 120; El Banhawy & Knapp, 2011: 36.
Amblyseius (Amblyseius) kangwanensis Ueckermann & Loots 1988: 85 (synonymy according to Ueckermann & Loots, 1988).
The 200 species of the genus Euseius are considered as Type IV species, polliniphagous generalists (McMurtry and Croft 1997; McMurtry et al. 2013). Like the previous Euseius species, the biology of E. baetae is however totally unknown.
Specimens examined: seven specimens (5 ♀♀, 1 ♂ and 1 imm.) collected during this study. Fomboni, inside the town (15 m aasl, 12°17'29'' S, 43°44'35'' E), 3 ♀♀ on Senna siamea (Lamarck) Irwin & Barneby (Fabaceae), 2/XII/2018 and 1 ♂ and 1 imm. on Pterocarpus indicus Willdenow (Fabaceae, 4/XII/2018; Bangoma, Bridge (11 m aasl, 12°17'2'' S, 43°45'5'' E), 3 ♀♀ on Averroha carambola L. (Oxalidaceae), 4/XII/2018.
Previous record: Congo, Grande Comore, Kenya, Malawi, Mozambique, South Africa.
Remarks: measurements of the five females (Table 1) and one male (Table 2) fit well with the measurements from the literature with slightly shorter dimensions in general, except concerning specimens from Grande Comore (Kreiter et al. 2018b), the neighboring island in the West less than 40 km from coasts of Mohéli. These specimens have almost identical measurements.
Amblyseius (Amblyseius) hima Pritchard & Baker 1962: 257; Blommers 1976: 89.
Euseius hima, Moraes et al. 1986: 46, 2004b: 71; Quilici et al. 2000: 99; Chant & McMurtry 2005a: 215, 2007: 121.
The 200 species of the genus Euseius are considered as Type IV species, polliniphagous generalists (McMurtry and Croft 1997; McMurtry et al. 2013). The biology of this species, however, remains totally unknown.
World distribution: Cameroon, Equatorial Guinea, La Réunion Island, Madagascar Island.
Specimens examined: six specimens (6 ♀♀) collected during this study. Fomboni, inside the town (15 m aasl, 12°17'29'' S, 43°44'35'' E), 6 ♀♀ on Pentas lanceolata (Forskål) Deflers (Rubiaceae), 2/XII/2018.
Remarks: this species was recorded from several countries of Sub-Saharan Africa, but also from India (Demite et al. 2020), La Réunion (Quilici et al. 2000; Kreiter et al. 2020d), Madagascar (Blommers 1976), Mauritius (Kreiter and Abo-Shnaf 2020b) and Anjouan (Kreiter et al. 2021). Morphological and morphometric characters and all measurements of our specimens fit well measurements published in Kreiter et al. (2020d) and with measurements of specimens from Mauritius Island (Kreiter and Abo-Shnaf 2020b).
Amblyseius (Amblyseius) rhusi van der Merwe 1965: 63.
Euseius rhusi Moraes et al. 1986: 52.
Typhlodromalus rhusi Moraes et al. 2004b: 203.
Euseius rhusi Chant & McMurtry 2005a: 215, 2007: 123.
Species of Euseius are classified as type IV = polliniphagous generalists (McMurtry and Croft 1997; McMurtry et al. 2013). The biology of this species, however, is totally unknown.
World distribution: Burundi, Colombia, Kenya, Mozambique, La Réunion Island, Rwanda, South Africa.
Specimens examined: A single female was collected during this study. Fomboni, University (25 m aasl, 12°17'3'' S, 43°44'34'' E), 1 ♀ on Zyzyphus mauritiana Lamarck (Malvaceae), 3/XII/2018.
Remarks: this species was described from Amanzimtoti Kwazulu-Natal Province in South Africa on Rhus rehmanniana Engler (Anacardiaceae) (van der Merwe 1965) and recorded in several countries of Africa (Ueckermann and Loots 1998; Moraes et al. 2006; El-Banhawy and Knapp 2011) and La Réunion (Quilici et al. 2000).
Measurements of the single female (Table 3) fit well with the measurements from the literature (Ueckermann and Loots 1988; Moraes et al. 2006; El-Banhawy and Knapp 2011), with however some slightly shorter dimensions, especially for Z5, and some longer dimensions, especially of j3.
Typhlodromalina Chant & McMurtry 2005a: 195.
Amblyseius (Typhlodromalus) Muma 1961: 288; Typhlodromalus De Leon 1966: 87.
Amblyseius spinosus Meyer & Rodrigues 1966: 30; Moraes et al. 1986: 31.
Kampimodromus spinosus, Quilici et al. 2000: 100.
Typhlodromalus spinosus, Moraes et al. 2004b: 204; Chant & McMurtry 2005a: 199, 2007: 111.
This species belongs to the athiasae species group as setae J1 and S5 are absent. This species group contains six species (Chant and McMurtry 2005a, Moraes et al. 2006).
Typhlodromalus spinosus was collected from Eastern, Western, but mainly Southern Africa and La Réunion (Demite et al. 2020). The rapid multiplication of this species on the western flower thrips (WFT), Frankliniella occidentalis Pergande, was confirmed and clear evidence that T. spinosus predates on WFT under laboratory and field conditions, but not on T. urticae (Mwangi et al. 2015). It seems abundant in low vegetation as it was found in high populations in a study of companion plants in a citrus orchard (Le Bellec et al. unpub. data).
This species have never been record from Guadeloupe or Martinique in similar studies, but it is interesting to notice that in those islands, another Typhlodromalus was collected, T. peregrinus (Muma) (Mailloux et al. 2010; Kreiter et al. 2013, 2018c). Typhlodromalus spinosus was recorded from La Réunion by Quilici et al. (2000) and was then find in quite high numbers by Kreiter et al. (2020d) and in lower numbers from Mauritius Island (Kreiter and Abo-Shnaf 2020b).
World distribution: Benin, Burundi Dr Congo, Kenya, Malawi, Mozambique, La Réunion Island.
Specimens examined: four specimens (2 ♀♀ and 2 ♂♂) collected during this study. Hoani, inside village (38 m aasl, 12°17'3'' S, 43°44'34'' E), 2 ♀♀ and 2 ♂♂ on Amaranthus viridis L. (Amaranthaceae), 3/XII/2018.
Remarks: morphological and morphometric characters and all measurements of our specimens fit well measurements in Kreiter et al. (2020d). This species was described from Mozambique (Meyer and Rodrigues 1966), then mentioned in the Indian Ocean from la Réunion (Quilici et al. 2000; Kreiter et al. 2020d), Mauritius (Kreiter and Abo-Shnaf 2020b) and more recently from Anjouan (Kreiter et al. 2021).
Ueckermannia Chant & McMurtry 2005a: 201. Preoccupied by Ueckermannia Kaźmierski, 1996 (Tydeidae).
Ueckermannseius Chant & McMurtry 2005b: 337, 2007: 115.
Ueckermannseius eastafricae Moraes et al. 2006: 30.
This species was described from Uganda and Kenya (Moraes et al. 2006) and recovered only once in Kenya (El-Banhawy et al. 2009). Its biology is totally unknown.
World distribution: Kenya, Uganda.
Specimens examined: 12 specimens (11 ♀♀ and 1 ♂) collected during this study. Bandar-Es-Salam, Les Abous Inn (23 m aasl, 12°17'37'' S, 43°45'27'' E), 11 ♀♀ and 1 ♂ on Carica papaya L. (Caricaceae), 2/XII/2018.
Remarks: morphological and morphometric characters and all measurements of our specimens (Table 4) fit well measurements in Moraes et al. (2006) for specimens from Africa and above all that of Kreiter et al. (submitted) for specimens from Anjouan. But specimens from Mohéli have shorter setae j4, j5, j6, R1, s4, S2, z2, z4, z5, SgeII, SgeIII and smaller ventrianal shield and cheliceral digits but have longer macrosetae StIV compared to specimens from Africa.
Above all, our specimens have pointed macrosetae or slightly rounded macrosetae on leg IV and the original description (Moraes et al. 2006) mentioned three evidently knobbed macrosetae. As it is the only main difference, we consider this difference as a variation of the character and not enough for considering that our specimens belong to a new species. The male of U. eastafricae was unknown until our collection. The single male collected will be described in a following paper.
Phytoseiini Berlese 1913: 3; Phytoseiinae Vitzthum 1941: 767.
Phytoseius Ribaga 1904: 177.
Phytoseius (Phytoseius) hongkongensis Swirski & Shechter 1961: 99; Amitai & Swirski 1966: 22.
Phytoseius (Pennaseius) hongkongensis Ehara 1966: 25; Ehara 1972: 169; Moraes et al. 1986: 211.
Phytoseius hongkongensis Moraes et al. 2004a: 240; Chant & McMurtry 2007: 129.
This species belongs to the plumifer species group (Chant and McMurtry 1994) as setae R1 and J2 are present. This species was described from specimens collected on Heterosmilax gaudichaudiana (Kunth) Maximovich (Smilacaceae), and Urena lobata L. (Malvaceae) in Victoria Mount Forest, Hong Kong Island, Hong Kong. Although species of the genus Phytoseius are considered to belong to the type III (polyphagous generalist predators) of McMurtry and Croft (1997) and McMurtry et al. (2013), its specific biology is totally unknown.
World distribution: Benin, Burundi, Cameroon, Cape Verde, DR Congo, Kenya, Madagascar Island, Malawi, Mozambique, Nigeria, Reunion Island, Rwanda, Senegal, South Africa, Zambia, Zimbabwe.
Specimens examined: 22 specimens (16 ♀♀, 2 ♂♂ and 4 imm.) collected during this study. Fomboni, Les-Hauts (60 m aasl, 12°17'29'' S, 43°44'35'' E), 7 ♀♀ and 2 imm. on Psidium cattleianum Afzelius ex Sabine (Myrtaceae), 7 ♀♀ and 2 imm. on Annona senegalensis Persoon (Annonaceae) and 4 ♀♀, 2 ♂♂ and 1 imm. on Adansonia digitata L. (Malvaceae), 2/XII/2018.
Remarks: Measurements of the three adult females agree well with measurements of the literature, especially with those of Ueckermann et al. (2007) obtained with a great number of specimens (29) from various countries in Africa with those of Kreiter et al. (2020b) for specimens from Vietnam.
Typhlodromini Wainstein 1962: 26; Typhlodrominae Chant & McMurtry 1994: 235.
Chanteiini Chant & McMurtry 1994: 237, 2007: 132.
Chanteius Wainstein 1962: 19.
Typhlodromus (Typhlodromus) contiguus Chant 1959: 29.
Typhlodromus (Diadromus) contiguus, Athias-Henriot 1960: 62.
Typhloseiopsis contiguus, Muma 1961: 294.
Chanteius (Chanteius) contiguus, Wainstein 1962: 9.
Typhlodromus contiguus, Hirshmann 1962: 2.
Typhlodromus (Typhloseiopsis) contiguus, Pritchard & Baker 1962: 222.
Diadromus contiguus, Chant & Yoshida-Shaul 1986: 2030, Moraes et al. 1986: 184.
Chanteius contiguus, Moraes et al. 2004b: 261; Chant & McMurtry 1994: 239.
Chanteius lieni (Tseng 1976): 97 (synonymy according to Chant & Yoshida-Shaul 1986).
This species belongs to the contiguus species group (Chant and McMurtry 1994) and its biology remains totally unknown.
World distribution: China, Hong-Kong, Japan, Madagascar, Philippines, Singapore.
Specimens examined: 31 specimens (10 ♀♀, 15 ♂♂ and 6 imm.) collected during this study. Fomboni, inside the town (15 m aasl, 12°17'29'' S, 43°44'35'' E), 1 ♂ on Senna siamea (Lamarck) Irwin & Barneby (Fabaceae), 2/XII/2018; Fomboni, Les-Hauts (60 m aasl, 12°17'29'' S, 43°44'35'' E), 1 imm. on Lawsonia inermis L. (Lythraceae), 1 ♀ and 1 ♂ on Psidium cattleianum Afzelius ex Sabine (Myrtaceae) and 1 ♀ on Adansonia digitata L. (Malvaceae), 2/XII/2018; Bandar-Es-Salam, Les Abous Inn (23 m aasl, 12°17'37'' S, 43°45'27'' E), 2 ♀♀, 5 ♂♂ and 2 imm. on Cajanus cajan L. (Fabaceae), 2/XII/2018; Fomboni, University (25 m aasl, 12°17'3'' S, 43°44'34'' E), 2 ♀♀, 5 ♂♂ and 2 imm. on Zyzyphus mauritiana Lamarck (Malvaceae), 3/XII/2018; Hoani, inside village (38 m aasl, 12°17'3'' S, 43°44'34'' E), 1 ♂ on Amaranthus viridis L. (Amaranthaceae), 1 ♀ + 1 imm. on Citrus sp., 3/XII/2018; Bangoma, top of the village (42 m aasl, 12°17'15'' S, 43°43'40'' E), 1 ♀ on Dendrocnide moroides (Weddel) Chew (Urticacae), 1 ♀ and 1 ♂ on Anacardium occidentale L. (Anacardiaceae) 4/XII/2020; Bangoma, Les Hauts (137 m aasl, 12°17'18'' S, 43°43'41'' E), 1 ♂ on Asplenium petiolulatum Mettenius (Aspleniaceae) and 1 ♀ on Abutilon hirtum (Lamarck) Sweet (Malvaceae), 4/XII/2018.
Remarks: morphological and morphometric characters and all measurements of our specimens fit well measurements in numerous descriptions and redescriptions available in the literature, especially those of Blommers (1976) for specimens from Madagascar. Mentioned only from South-East Asia and Madagascar, it is the second mention in the Indian Ocean of this species outside Madagascar after Mayotte Island (Kreiter et al. 2020a).
Typhlodromini Wainstein 1962: 26.
Typhlodromus Scheuten 1857: 111.
Typhlodromus (Anthoseius) De Leon 1959: 258; van der Merwe 1968: 20; Karg 1982: 194; Chant & McMurtry 1994: 250, 2007: 149.
Typhlodromus (Anthoseius) grewiae Zannou, Moraes & Oliveira in Ueckermann et al. 2008: 48.
This species belongs to the singularis species group as setae JV3 are absent and dorsal shield setae are short (Chant and McMurtry 1994). The biology of that species is totally unknown. It was mentioned only from Kenya (Ueckermann et al. 2008) only from one female.
World distribution: Kenya, Mayotte Island.
Specimens examined: 5 specimens (2 ♀♀ and 3♂♂) collected during this study. Fomboni, Les-Hauts (60 m aasl, 12°17'29'' S, 43°44'35'' E), 1 ♀ on Annona senegalensis Persoon (Annonaceae), 2/XII/2018; Bangoma, top of the village (42 m aasl, 12°17'15'' S, 43°43'40'' E), 1 ♀ and 3 ♂♂ on Syzygium aromaticum (L.) Merrill & Perry, 4/XII/2018.
Remarks: the species was mentioned only once from Kenya and described from only one female specimen (Ueckermann et al. 2008). Morphological and morphometric characters and all measurements of our specimens (Table 5) fit well measurements of the original description of Zannou, Moraes & Oliveira in Ueckermann et al. (2008) and with measurements of specimens from Mayotte Island (Kreiter et al. 2020a)
Typhlodromus (Typhlodromus) hartlandrowei Evans, 1958: 580-581; Chant 1959: 60.
Clavidromus hartlandrowei, Muma, 1961: 296.
Typhlodromus (Neoseiulus) hartlandrowei, Pritchard & Baker, 1962: 222.
Typhlodromus (Anthoseius) hartlandrowei, Moraes et al. 2004b: 328; Chant & McMurtry, 2007: 155; Ueckermann et al. 2008: 50.
This species belongs to the bergi species group (Chant and McMurtry 1994). The biology of that species is totally unknown. This is the first mention of that species outside the African continent.
World distribution: Democratic Republic of Congo, Nigeria, Uganda.
Specimens examined: two specimens (2 ♀) collected during this study. Fomboni, University (25 m aasl, 12°17'3'' S, 43°44'34'' E), 1 ♀ on Persea americana Miller (Lauraceae), 4/XII/2018; Bangoma, top of the village (42 m aasl, 12°17'15'' S, 43°43'40'' E), 1 ♀ on Anacardium occidentale L. (Anacardiaceae) 4/XII/2020.
Remarks: morphological and morphometric characters and all measurements of our specimens (Table 6) fit well with measurements of the original description given by Evans (1958), those given by Ueckermann et al. (2008) concerning specimens from Africa and those of specimens from Anjouan Island (Kreiter et al. 2021).
Typhlodromus (Anthoseius) lobatus Zannou, Moraes & Oliveira in Ueckermann et al. 2008: 59.
This species belongs to the large rhenanus species group (Chant and McMurtry 1994). The biology of this species is totally unknown.
World distribution: Ghana, Mauritius Island, Mayotte Island, Rodrigues Island.
Specimens examined: ten specimens (9 ♀♀ and 1 imm.) collected during this study. Fomboni, inside the town (15 m aasl, 12°17'29'' S, 43°44'35'' E), 2 ♀♀ + 1 imm. on Abutilon hirtum (Lamarck) Sweet (Malvaceae) and 4 ♀♀ on Hibiscus tiliaceus L. (Malvaceae), 2/XII/2018; Bandar-Es-Salam, Les Abous Inn (23 m aasl, 12°17'37'' S, 43°45'27'' E), 1 ♀ on Cajanus cajan L. (Fabaceae), 2/XII/2018; Bangoma, Les Hauts (137 m aasl, 12°17'18'' S, 43°43'41'' E), 2 ♀♀ on Cananga odorata (Lamarck) Hooker & Thomson (Malvaceae), 4/XII/2018.
Remarks: morphological and morphometric characters and all measurements of our specimens fit well with measurements of the original description by Zannou, Moraes and Oliveira in Ueckermann et al. (2008) concerning specimens from Ghana, Western Africa. And with measurements of specimens from Rodrigues (Kreiter and Abo-Shnaf 2020a), Mauritius (Kreiter and Abo-Shnaf 2020b), Mayotte (Kreiter et al. 2020a) and Anjouan (Kreiter et al. 2021). This species seems rather common in the Indian Ocean Islands, except in La Réunion Island.
Typhlodromus (Anthoseius) moraesi Kreiter & Ueckermann in Kreiter et al. 2002: 338.
The biology of this species found in La Réunion Island by Kreiter et al. (2002) on various host plants (Kreiter et al. 2002) and then in French Caribbean Islands (Mailloux et al. 2010; Kreiter et al. 2013) remains unknown.
World distribution: La Réunion Island.
Specimens examined: three specimens (3 ♀♀) collected during this study. Bandar-Es-Salam, Les Abous Inn (23 m aasl, 12°17'37'' S, 43°45'27'' E), 1 ♀ on Punica granatum L. (Lythraceae), 4/XII/2018/
Remarks: several species are found both in La Réunion Island (in the Indian Ocean) and in the West Indies, probably because of reciprocal introductions certainly a long time ago with slave-trade and commercial exchanges between the two areas or because of introduction of plants from Antilles into La Réunion coming from the same African countries than slaves. The measurements and description of the specimens collected fit very well those given by Kreiter et al. (2002) for the original description and measurements of specimens from La Réunion (Kreiter et al. 2020d) and Rodrigues Islands (Kreiter and Abo-Shnaf 2020a).
The results of a survey carried out in 2018 in Mohéli Island is presented in this paper. A total of 18 new records, 16 of which being documented in this paper: 12 Amblyseiinae, 1 Phytoseiinae and 5 Typhlodrominae, have been obtained, namely: Neoseiulus teke, Paraphytoseius orientalis, Amblyseius duplicesetus, A. herbicolus, A. largoensis, Euseius hima, E. baetae, E. rhusi, Typhlodromalus spinosus, Ueckermannseius eastafricae, Phytoseius hongkongensis, Chanteius contiguus, Typhlodromus (Anthoseius) grewiae, T. (A.) hartlandrowei, T. (A.) lobatus, T. (A.) moraesi. Two new species (Paragigagnathus sp. and Typhlodromalus sp.) found in several islands will be described in a next paper
Among the 18 recorded species, at least five species (P. orientalis, A. largoensis, A. herbicolus, T. spinosus and T. [A.] transvaalensis) are already known as biological control agents (BCAs). In addition to the intrinsic value of phytoseiid mite biodiversity in tropical environments, demonstration of the natural occurrence of efficient BCAs in a developing country such as Mohéli Island is of great agricultural, commercial and strategic interests for the country.
Thanks are first due to the Department to which the senior author belong for funds that have granted travels and accommodations to him in Mohéli Island: UMR CBGP (Internal call for proposals 2018). A grateful thanks to UR Hortsys and to the head Dr Fabrice Le Bellec that have allowed Rose-My Payet to join the senior author for collections and partly granted field collecting trips. Field collections had been made with authorizations of the government of Union des Comores by letters of the head of INRAPE, Union des Comores (ref. n°18/193/INRAPE/DG and n°18/210/INRAPE/DG, respectively).
We also would like to warmly thank Professor Marie-Stéphane Tixier for valuable discussion on taxonomy of some species and especially Ueckermannseius eastafricae Moraes, Zannou and Oliveira.
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