1Laboratory of Biotechnology and Valorisation of Natural Resources, Faculty of Sciences, 80000 Agadir, Ibn Zohr University (UIZ), Agadir, Morocco.
2Laboratory of Biotechnology and Valorisation of Natural Resources, Faculty of Sciences, 80000 Agadir, Ibn Zohr University (UIZ), Agadir, Morocco
3✉ CBGP, Institut Agro, CIRAD, INRAE, IRD, Univ Montpellier, Montpellier, France.
4CBGP, Institut Agro, CIRAD, INRAE, IRD, Univ Montpellier, Montpellier, France.
5Laboratory of Biotechnology and Valorisation of Natural Resources, Faculty of Sciences, 80000 Agadir, Ibn Zohr University (UIZ), Agadir, Morocco.
6Laboratory of Biotechnology and Valorisation of Natural Resources, Faculty of Sciences, 80000 Agadir, Ibn Zohr University (UIZ), Agadir, Morocco.
2023 - Volume: 63 Issue: 3 pages: 691-724
https://doi.org/10.24349/v5of-5oe1In Morocco, citrus sector development remains confronted to several constraints such as phytosanitary problems associated to damage caused by pests (both indigenous and exotic) (Benziane et al. 2003; Mazih 2008; Lamharchi et al. 2020a). The main pests in citrus orchards in Morocco and in Mediterranean countries are the Mediterranean fruit fly [Ceratitis capitata (Wiedemann), Tephritidae], scale insects including the California red scale [Aonidiella aurantii Maskell, Diaspididae] and the citrus leaf miner [Phyllocnistis citrella Stainton, Gracillariidae] (Abbassi 1990; Mazih 2008, 2015; Lamharchi et al. 2016). To control these pests, producers use large amount of pesticides. As well as phytophagous mites (Tetranychidae, Tenuipalpidae, Tarsonemidae, and Eriophyidae) have started to appear althougth they were considered secondary pests for 20 years (Saba 1978; Kreiter et al. 2002; Grissa and Khoufi 2012; Sahraoui et al. 2012). To limit pesticide used for controlling these mites, biological control with predators of phytophagous species such as phytoseiid mites, is an innovative alternative.
Phytoseiid mites (Acari: Mesostigmata) have been extensively studied for their potential as biological control agents of phytophagous mites in several countries (McMurtry and Croft 1997; Swirski et al. 1998; Tixier et al. 2008; Kreiter et al. 2010; McMurtry et al. 2013). Some species are of great importance for Integrated Pest Management (IPM) programs in Mediterranean orchards (Kasap and Sekeroglu 2004; Tixier et al. 2009; McMurtry et al. 2013).
The Phytoseiidae family accounts for more 2,557 species (Demite et al. 2014, 2023). However, the fauna of some countries such as Morocco has not been completely surveyed (Tixier et al. 2008). Until now, about 50 species are reported from Morocco (McMurtry and Bounfour 1989; Kreiter et al. 2007; Tixier et al. 2016). Knowledge regarding the mite fauna, especially in citrus orchards of central west Morocco, is of considerable importance to establish effective programs for the biological control of pests. The aim of this study is to evaluate the phytoseiid species associated to plants both in citrus orchards, and on uncultivated plants in the argan forest in central western of Morocco, sampled at different seasons.
A survey of phytoseiid diversity was carried out in different citrus orchards in Taroudant region between April and October for two consecutive years 2016–2017 and 2017–2018. Orchard's names and geographic location are shown Table 1. All are conventionally conducted. Some pesticides were applied to control several pests and diseases (sulphur, abamectin, spirodiclofen, fenpropathrin, propargite, benzoxamate, clofentezine, tetradifon, phosmet, malathion, benomyl, deltamethrin, spirotetramat...). Herbicides (glyphosate, oxyfluorfen) were also applied from spring until autumn, and ground was ploughed several times. Additional sampling was undertaken on uncultivated plants in April 2019 between Agadir and Imouzzar Ida or Tanane, area of the argan Forest (Table 1).
Plants (citrus leaves, weeds, and windbreaks) were collected in all orchards in two seasons (spring and autumn). Each sample contains: 150 leaves of citrus varieties/ plot/ date/ year; the aerial part of 50 plants of weeds/ plot/ date/ year and 50 branches of windbreaks/ plot/ date/ year. Mites were collected using the dipping-checking-washing-filtering of leaves method (Boller 1984; Tixier et al. 2016). The collected individuals were preserved in 70% ethanol.
Mites were also collected from uncultivated plants in the area of the argan forest between Agadir city and Imouzzar Ida ou Tanane. This additional sampling was carried out as follows: branches of each plant (perennial woody plant: tree or shrub), at a level between 1 m and 1.5 m, were subjected to between 10 to 15 light taps with a stick during five to ten minutes/ plant. Among the individuals fallen on a black background, only the specimens considered to be phytoseiids were isolated, collected with a fine brush imbibed with alcohol. The collected individuals were kept in ethanol for identification. All phytoseiid mites were counted and were mounted on slides using Hoyer's medium (Gutierrez 1985; André 1986). Slides were then dried in an oven at 47 °C for four days for thinning. Once dried, slides were sealed for conservation. Identifications were carried out with a phase and interference contrast microscope (Leica DM LB 2 phase contrast microscope, 10 x 25 HC Plan, Fluotar 100) (400x magnification). The generic classification of Chant and McMurtry (2007) was used for identification. Other more specific literature was used for species determination (Swirski et al. 1998; Kreiter et al. 2007, 2016a; Papadoulis et al. 2009; Ferragut et al. 2010; Tixier et al. 2013, 2016).
A total of 248 individuals of phytoseiid mites were identified from citrus leaves, weeds, and associated windbreaks (Cupressus, other conifers, Acacia) in nine orchards in Taroudant region and in the argan forest in central west of Morocco (Table 1). Twenty-two species found were already reported and seven species are new for the Moroccan fauna. The list of these species is presented below, with some biological information and uses in biological control when available. Geographical distributions information is issued from Demite et al. (2023).
A key of all recorded Moroccan species is given after.
Amblyseiinae Muma 1961: 273.
Neoseiulini Chant & McMurtry 2003a: 6.
Neoseiulus Hughes 1948: 141.
Amblyseius obtusus var. alpinus Schweizer 1922: 41.
Amblyseius alpinus Schweizer 1949: 79.
Typhlodromus (Amblyseius) alpinus, Chant 1959: 105.
Typhlodromus (Typhlodromus) alpinus, Westerboer & Bernhard 1963: 651.
Typhlodromips alpinus, Moraes et al. 1986: 136.
Amblyseius (Neoseiulus) alpinus, Karg 1993: 189.
Neoseiulus alpinus, Evans 1987: 1461; Moraes et al. 2004a: 100; Chant & McMurtry 2003: 37, 2007: 25.
Neoseiulus polonicus (Willmann 1949): 117 (Synonymy according to Evans 1987).
Neoseiulus aurescens (Athias-Henriot 1961): 441 (Synonymy according to Evans 1987).
Neoseiulus polyporus Wainstein 1962: 143 (Synonymy according to Evans 1987).
World distribution — Algeria, Austria, Australia, Belgium, Chile, Cuba, Czech Republic, England, Finland, France, Georgia, Germany, Greece, Hawaii, Hungary, Italy, Jordan, Latvia, Morocco, Norway, Portugal, Poland, Russia, Slovakia, Spain, Switzerland, Tunisia, Turkey, Ukraine, and USA (Arizona, California, and Washington).
Previous records from Morocco — This species has been reported for the first time by Tixier et al. (2016), one specimen was collected on Echium vulgare L. (Boraginaceae) in Larache Beach.
Specimens examined — 2 ♀♀ on Solanum nigrum L. (Solanaceae) in July 25, 2018 at Taoufiqui 1 domain; 1 ♀ and 2 ♀♀ on the same previous host plant respectively in September 2017 and 2018.
Remarks — Measurements and description of the adult females agree with those provided by Athias-Henriot (1961) and Chant and McMurtry (2003a). This species is considered a synonym of Neoseiulus aurescens (Athias-Henriot) in Chant and McMurtry (2003a). It is essentially recorded from Europe and it is observed rarely in crops, sometimes in vineyards and apple orchards (Kreiter et al. 2000; Tixier et al. 2013).
Typhlodromus californicus McGregor, 1954: 89.
Amblyseius californicus, Schuster & Pritchard 1963: 271.
Cydnodromus californicus, Athias-Henriot 1977: 64.
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, 2004a: 109; Chant & McMurtry 2003a: 21, 2007: 25; Guanilo et al. 2008a: 27, 2008b: 19; sensu Athias-Henriot 1977, Beaulieu & Beard 2018: 469.
Typhlodromus mungeri McGregor, 1954: 92 (Synonymy according to Chant & McMurtry, 2003a).
Typhlodromus chilenensis Dosse, 1958: 55 (Synonymy according to Chant & McMurtry, 2003a).
Amblyseius wearnei Schicha 1987: 103 (Synonymy according to Tixier et al. 2014).
World distribution — Algeria, Argentina, Azores, Brazil, Canada, Canary Islands, Chile, Colombia, Cuba, Cyprus, France, Greece, Guadeloupe, Guatemala, Italy, Japan, Mexico, Morocco, Peru, Portugal, Reunion Island, Senegal, Serbia, Slovenia, South Africa, South Korea, Spain, Syria, Taïwan, Tunisia, Turkey, Uruguay, USA, Venezuela, and Vietnam.
Previous records from Morocco — This species has been reported from the Moroccan fauna by Kreiter et al. (2007). Several specimens were collected on many plants belonging to the families Malvaceae (Hibiscus rosa-sinensis L.), Rhamnaceae (Ziziphus lotus L. (Lam)), Anacardiaceae (Pistacia lentiscus L.) and Asteraceae (Calendula arvensis (M.) Bieb) and Convolvulaceae (Convolvulus althaeoides L.) (Tixier et al. 2016).
Specimens examined — 8 ♀♀ + 3 ♂♂ on Convolvulus althaeoides L. (Convolvulaceae) in May 2017 and in July 2017; 4 ♀♀ on citrus leaves at Stah Al Madina orchard in 2018.
Remarks — Measurements and description of the adult females collected agree with those provided by Tixier et al. (2008) for specimens collected worldwide, by Ferragut et al. (2010) for specimens from Spain and by Kreiter et al. (2020b) for specimens from La Réunion and various regions in the world. This is a very widespread species (Demite et al. 2023), reported as generalist predatory mite (Castagnoli and Simoni 2003). It can feed on Tetranychidae (Ferragut et al. 2010; Gomez-Moya and Ferragut 2009), tarsonemid mites (Easterbrook et al. 2001), or small insects (Rodriguez-Reina et al. 1992; Fraulo et al. 2008) and even pollen when the primary prey is unavailable (Rhodes and Liburd 2005). Akchour et al. (2016, 2021) reported that N. californicus is considered as a potential agent to control Eutetranychus orientalis (Klein) in citrus orchards in Souss region in central western of Morocco.
Typhlodromus cucumeris Oudemans 1930: 69.
Amblyseius cucumeris, Athias-Henriot 1957a 336.
Amblyseius (Typhlodromopsis) cucumeris, De Leon 1959a.
Typhlodromus (Amblyseius) cucumeris, Chant 1959: 78.
Amblyseius (Amblyseius) cucumeris, Wainstein 1962.
Typhlodromus (Typhlodromus) cucumeris, Westerboer & Bernhard 1963.
Amblyseius (Neoseiulus) cucumeris, De Leon 1965: 23.
Neoseiulus cucumeris, Moraes et al. 1986: 76, 2004a: 115; Chant & McMurtry 2003: 21, 2007: 25.
Neoseiulus thripsi (McGill 1939): 310 (Synonymy according to Chant 1959).
Neoseiulus bellinus (Womersley 1954): 177 (Synonymy according to Athias-Henriot 1960a).
World distribution — Algeria, Armenia, Australia, Austria, Azerbaijan, Azores, Belarus, Belgium, Canada, Canary Islands, Caucasus Region, Chile, Cyprus, Egypt, England, Finland, France, Georgia, Germany, Greece, Hungary, India, Iran, Israel, Italy, Latvia, Mexico, Moldova, Morocco, Netherlands, New Zealand, Norway, Poland, Portugal, Russia, Saudi Arabia, Slovakia, Slovenia, Spain, Sweden, Switzerland, Tunisia, Turkey, Ukraine, and USA.
Previous records from Morocco — This species has been reported in Morocco by McMurtry and Bounfour (1989). In addition, some females were collected on Juncus articulatus L. (Juncaceae) at Larache, on Lavandula dentata L. (Lamiaceae) at Gorges of Zegzel (Tixier et al. 2016).
Specimens examined — 3 ♀♀ collected at Taoufiqui 1 domain on citrus leaves in June 2018.
Remarks — Measurements and description of the adult females collected agree with those provided by Beard (1999) and Schicha (1976) for specimens from Australia. This species is commercialised and used in greenhouses to control phytophagous mites and thrips (Messelink et al. 2006). It seems able to control Tetranychus urticae Kock and Phytonemus pallidus (Banks) in field-grown strawberry (Fitzgerald et al. 2008). It was also known to feed on tarsonemid mites (Tuovinen and Lindquist 2010), eggs of Bemisia tabaci (Gennadius) on vegetables (cucumber, sweet pepper, tomato) in laboratory conditions as reported by Kim et al. (2008).
Typhlodromus longilaterus Athias-Henriot, 1957b: 218.
Typhlodromus (Amblyseius) longilaterus, Chant 1959: 79.
Cydnodromus longilaterus, Muma 1961: 290.
Amblyseius (Amblyseius) longilaterus, Ueckermann & Loots 1988: 128.
Amblyseius longilaterus, Swirski & Amitai 1997a: 14.
Neoseiulus longilaterus, Moraes et al. 1986: 85, 2004a: 128; Chant & McMurtry 2003: 23, 2007: 29.
World distribution — Algeria, Israel, Morocco, and Tunisia.
Previous records from Morocco — This species has been reported for the first time in Morocco on Echium vulgare (L.) (Boraginaceae) at Larache Beach (Tixier et al. 2016).
Specimens examined — 2 ♀♀ collected in June 2018 at Stah Al Madina on Convolvulus arvensis M. (Bieb) (Convolvulaceae).
Remarks — Measurements and description of the adult females collected agree with those provided by Evans (1987). Tsolakis et al. (2013) showed the ability of this species to feed on T. urticae, Panonychus citri (McGregor), P. ulmi (Koch). This species is only known from the Mediterranean basin and has been observed on Convolvulus spp. (Convolvulaceae) and on Cynodon dactylon (L.) (Poaceae) (Sahraoui et al. 2012). In addition, as reported by Tsolakis et al. (2019), N. longilaterus have similar traits with some phytoseiid species belonging to type II selective predators, but also with some others included within type III generalist predators. As a matter of fact, it can be considered an interesting biocontrol agent towards T. urticae, able also to develop on alternative foods as the pollen of Oxalis spp. L. (Oxalidaceae).
Typhlodromus umbraticus Chant, 1956: 26.
Typhlodromus (Typhlodromus) umbraticus, Beglyarov 1958: 107.
Amblyseius umbraticus, Athias-Henriot 1959: 138.
Typhlodromus (Amblyseius) umbraticus, Chant 1959: 75.
Amblyseius (Typhlodromopsis) umbraticus, Muma 1961: 287.
Amblyseius (Amblyseius) umbraticus, Wainstein & Vartapetov 1973: 103.
Amblyseius (Neoseiulus) umbraticus, Karg 1991: 23.
Neoseiulus umbraticus, Moraes et al. 1986: 99, 2004a: 149; Chant & McMurtry 2003a: 23, 2007: 31.
World distribution — Armenia, Azerbaijan, Azores, Belarus, Caucasus region, Denmark, England, France, Georgia, Germany, Hungary, Iran, Italy, Jamaica, Latvia, Madeira Island, Mexico, Moldova, Montenegro, Morocco, Norway, Poland, Russia, Slovakia, Slovenia, Spain, Switzerland, Turkey, Ukraine, and USA.
Previous records from Morocco — The first report of this species in Morocco has been published by Tixier et al. (2016). It was collected on many plants such as Echium spp. L. (Boraginaceae), Mentha pulegium L. (Lamiaceae), Dittrichia viscosa L. Greuter, (Asteraceae), Juncus articulatus L. (Juncaceae), and Lavandula dentata L. (Lamiaceae).
Specimens examined — 2 ♀♀ on Convolvulus althaeoides L. (Convolvulaceae) in July 2017; 1 ♀ and 1 ♂ on Solanum nigrum L. (Solanaceae) collected at Chbaika domain in July 2017.
Remarks — Measurements and description of adult females collected in this survey agree with those provided by Ferragut et al. (2010) for specimens from Spain and by Tixier et al. (2016) for specimens from Morocco. Few studies exist on its biology; it has the ability to develop feeding on T. urticae (Kazak et al. 2002) and on Thrips tabaci Lindeman on leek (Allium porrum L.) (Alliaceae) in laboratory conditions (Sengonca and Dresher 2001).
Paragigagnathus Amitai & Grinberg 1971: 327; Karg 1983: 300; Kolodochka 1994: 3; Chant & McMurtry 2003: 39.
Pamiroseius molestus Kolodochka 1989: 228;
Paragigagnathus molestus, Moraes et al. 1986: 103, 2004a: 159; Chant & McMurtry 2003: 44, 2007: 33.
World distribution — Iran and Turkmenistan.
Specimens examined — 3 ♀♀ + 1 im. associated with tetranychid mites on Ricinus communis L. (Euphorbiaceae), specimens collected in April 2019, at the Bottom of Aourir river, between Alma and Aourir villages, Agadir Ida Outanan Province, in the argan forest. This is the first record of this species in Morocco.
Remarks — The description and measurements of the adult females collected agree with those provided by Hajizadeh et al. (2010) for specimens from Iran. The type specimens were collected from Tamarix spp. L. (Tamaricaceae) in Turkmenistan, Kalinin (Kolodochka 1989), and additional reports from Iran were observed on Tamarix spp. (Hajizadeh et al. 2010).
Paragigagnathus tamaricis Amitai & Grinberg 1971: 327; Moraes et al. 2004a: 159; Chant & McMurtry 2003: 44, 2007: 33.
Paragigagnathus tawfiki (Yousef 1974): 381 (Synonymy according to Chant & McMurtry 2003).
World distribution — Egypt, Iran, Israel, Italy, Jordan, Morocco, Saudi Arabia, and Tunisia.
Previous records from Morocco — This species is reported for the first time by Tixier et al. (2016) for Morocco and the Maghreb region. It was collected in the Oasis of Zagora on Tetraclinis articulata (Vahl) Mast. (Cupressaceae), at the Damon Oum Er Bia River on Tamarix africana Poir. (Tamaricaceae), in the Gorges of Zegzel on Tamarix communis L. (Tamaricaceae) (Tixier et al. 2016). These authors suggest a narrow relationship between this species and plants of Tamarix genus.
Specimens examined — 7 ♀♀ and 1 ♂ on citrus leaves in July 2017, at Nour domain.
Remarks — The description and measurements of the adult females collected agree with those provided by Hajizadeh et al. (2010) for specimens from Iran.
Phytoseiulini Chant &McMurtry 2006: 17.
Phytoseiulus Evans 1952: 397.
Phytoseiulus persimilis Athias-Henriot 1957a: 347.
Typhlodromus persimilis, Hirschmann 1962: 75.
Phytoseiulus (Phytoseiulus) persimilis, Wainstein 1962: 17.
Phytoseiulus persimilis, Chant 1959: 109; Moraes et al. 1986: 109, 2004a: 169; Chant & McMurtry 2006: 20, 2007: 53.
Phytoseiulus riegeli Dosse, 1958: 48 (Synonymy according to Kennett & Caltagirone 1968).
Phytoseiulus tardi Lombardini, 1959: 166 (Synonymy according to Kennett & Caltagirone 1968).
World distribution — Algeria, Australia, Canada, Canary Islands, Chile, China, Costa Rica, Cyprus, Egypt, Finland, France, Greece, Guatemala, Hungary, Iran, Israel, Italy, Japan, Jordan, Kenya, Latvia, Lybia, Martinique, Mauritius, Morocco, Netherlands, New Caledonia, New Zealand, Peru, Philippines, Portugal, Reunion Island, Rodrigues Island (Mauritius), Serbia, Slovenia, South Africa, South Korea, Spain, Syria, Tunisia, Turkey, USA, and Venezuela.
Previous records from Morocco — The presence of this species has been reported on wild plants and crops in various regions of Morocco by McMurtry and Bounfour (1989) and Tixier et al. (2003). It was observed on Ricinus communis L. (Euphorbiaceae), Stachys ocymastum (L.) Briq. (Lamiaceae), Vicia benghalensis L. (Fabaceae), Solananum sodomeum L. (Solanaceae), and Malva spp. (Malvaceae).
Specimens examined — 2 ♀♀ on citrus leaves in June 2017; 4 ♀♀ + 2 ♂♂ on Malva parviflora L. (Malvaceae); 3 ♀♀ on Convolvulus althaeoides L. (Convolvulaceae) at Stah Al Madina. 4 ♀♀ on the same previous substrate at Saouda in June 2017; 4 ♀♀ and 1 ♂ at Taoufiqui 1 domain in the same date. 4 ♀♀ at Chbaika orchards on the same previous substrate in June and September 2018.
Remarks — Description and measurements of the adult females agree with those provided by Ferragut et al. (2010) for specimens from Spain and by Kreiter et al. (2018, 2020b) for specimens from Mauritius, La Réunion and various regions in the world. This species was mainly known from Mediterranean climates around the world (Takahashi and Chant 1993; Kreiter et al. 2010). It is one of the best-known phytoseiid species as specialist predator, since its use to control T. urticae in greenhouses (McMurtry and Croft 1997; Escudero and Ferragut 2004; McMurtry et al. 2013).
Amblyseiini Muma Wainstein 1962: 26.
Amblyseiina Muma Chant & McMurtry 2004a: 179.
Transeius Chant & McMurtry 2004: 181.
Transeius audeae Kreiter, Allam & Tixier in Tixier et al. 2016: 532.
World distribution — Morocco.
Previous records from Morocco — The first report from Morocco of this recently described species was mentioned in Tixier et al. (2016). One specimen (1 ♀) has been collected on Vitis vinifera L. (Vitaceae) in Meknes region.
Specimens examined — 2 ♀♀ on citrus leaves in July 2018 at Nour domain; 5 ♀♀ on Withania somnifera L. (Solanaceae) in July 2018 at Nour domain.
Remarks — The morphological features of the specimens here in collected sample agree with those provided by Tixier et al. (2016) for specimens from Morocco.
Amblyseiella Muma 1955: 266.
Amblyseiella setosa Muma 1955: 266.
Phytoseiulus setosa, Garman 1958: 71.
Typhlodromus (Amblyseius) setosus, Chant 1959: 70.
Typhlodromus setosus, Hirschmann 1962: 128.
Amblyseius (Amblyseiellus) setosus, Wainstein 1962: 15.
Amblyseius setosus, Porath and Swirski 1965: 97.
Amblyseius (Amblyseiella) setosus, Ehara 1966a: 24.
Amblyseius (Amblyseiellus) setosus, Arutunjan 1977: 40.
Amblyseius setosa, Rowell et al. 1978.
Amblyseiella setosa, Moraes et al. 1986: 4, 2004a: 9; Chant & McMurtry 2004: 186, 2007: 71.
Amblyseiella denmarki (El-Borolossy in Nasr & Abou-Awad 1985): 245 (Synonymy according to Chant & McMurtry 2004).
Amblyseiella rusticana (Athias-Henriot, 1960a): 292 (Synonymy according to Muma et al. 1970).
World distribution — Algeria, Austria, Brazil, Egypt, France, Georgia, Greece, Israel, Peru, Spain, Turkey, and USA.
Specimens examined — 2 ♀♀ and 1 ♀ respectively on Convolvulus althaeoides L. and C. arvensis M. (Bieb) (Convolvulaceae) in May 2017 at Errahim domain in 2017. This is the first record of this species for the Moroccan fauna.
Remarks — The description and measurements of the adult females collected agree with those provided by Swirski et al. (1998) for specimens from Israel and Muma (1955) for specimens from Florida (USA).
Amblyseius Berlese 1914: 143.
Typhlodromus andersoni Chant 1957: 296.
Typhlodromus (Amblyseius) andersoni, Chant 1959: 92.
Amblyseius (Amblyseius) andersoni, Muma 1961: 287.
Typhlodromus (Typhlodromus) andersoni, Westerboer & Bernrhard 1963: 682.
Amblyseius (Multiseius) andersoni, Denmark & Muma 1989: 84.
Amblyseius andersoni, Athias-Henriot 1958: 33; Moraes et al. 1986: 7, 2004a: 14; Chant & McMurtry 2004: 199, 2007: 75.
Amblyseiopsis potentillae Garman, 1958: 7 (Synonymy according to Chant & Yoshida-Shaul 1990).
Typhlodromus (Amblyseius) britannicus Chant, 1959: 87 (Synonymy according to Chant & Yoshida-Shaul 1990).
Amblyseius charui Gupta 1969: 126 (Synonymy according to Gupta 1985).
Amblyseius reflexus Knisley & Denmark 1978: 8 (Synonymy according to Chant and Yoshida-Shaul 1990).
World distribution — Algeria, Austria, Azerbaijan, Canada, Cyprus, Czechoslovakia, Czech Republic, Denmark, England, France, Georgia, Germany, Greece, Hungary, Italy, Japan, Latvia, Moldova, Morocco, Netherlands, Poland, Portugal, Serbia, Slovakia, Slovenia, Spain, Sweden, Switzerland, Syria, Turkey, Ukraine, and USA.
Previous records from Morocco — This species was reported from Morocco on citrus (Tixier et al. 2003). It has been observed in many regions on many other plants such as Malva spp. (L.) (Malvaceae), Quercus suber L. (Fagaceae), Erica arborea L. (Ericaceae), and Cistus parviflorus L. (Cistaceae) (Tixier et al. 2016).
Specimens examined — 6 ♀♀ + 2 ♂♂ on Malva parviflora L. (Malvaceae) collected in May 2017, at Ouled Abdellah orchard (Taroudant); 3 ♀♀ + 1 ♂ on citrus leaves (Washington sanguine variety) (Rutaceae) collected at Stah Al Madina (Taroudant) in September 2018.
Remarks — The description and measurements of adult females collected agree with those provided by Denmark and Muma (1989). This species is cosmopolitan and considered a polyphagous predatory mite. It has been reported on a wide range of plants, particularly in vineyards (Kreiter et al. 2000) and on fruit trees like in apple orchards as reported by Tixier et al. (2013). In addition, it feeds on many species of mites, such as Aculops lycopersici (Tryon) (Eriophyidae) responsible for tomato russet, key pests of ornamental coniferous plants, i.e., Oligonychus ununguis (Jacobi) and Pentamerismus taxi (Haller), and pollen of Pinus sylvestris L. (Pinaceae) (Lara et al. 2012; Puchalska et al. 2021).
Typhlodromus (Amblyseius) italicus Chant 1959: 70.
Amblyseius (Amblyseialus) italicus, Muma 1961: 287.
Amblyseius (Amblyseius) italicus, Wainstein 1962: 15.
Amblyseius (Multiseius) italicus, Denmark & Muma 1989: 82.
Amblyseius italicus, Chant 1959: 70; Athias-Henriot 1966; Moraes et al. 1986: 16, 2004a: 31; Chant & McMurtry, 2004: 193, 2007: 78.
World distribution — Italy, Kenya, and Morocco.
Previous records from Morocco — This species was reported by McMurtry and Bounfour (1989) in El Jadida, 2 ♀♀ on cucumber (Cucurbitaceae) leaves in greenhouse.
Specimens examined — 3 ♀♀ on Cupressus sp. L. (Cupressaceae) used as windbreaks at Nour orchard (Ouled Teima) in September 2018; 1 ♀ collected on citrus leaves at Stah Al Madina domain in September 2018.
Remarks — The description and measurements of the adult females collected agree with those provided by McMurtry and Bounfour (1989) for specimens from Morocco.
Amblyseiopsis largoensis Muma 1955: 266.
Typhlodromus (Amblyseius) largoensis, Chant 1959: 96.
Amblyseius (Amblyseialus) largoensis, Muma 1961: 287.
Typhlodromus largoensis, Hirschmann 1962.
Amblyseius (Amblyseius) largoensis, Ehara 1966: 22; Ehara & Bhandhufalck 1977: 67; Denmark & Evans 2011: 69.
Amblyseius largoensis, Ehara 1959: 293; Muma & Denmark 1970: 69; McMurtry & Moraes 1984: 29; Moraes et al. 1986: 17, 2000: 239, 2004a: 143, 2004b: 33; Chant & McMurtry 2004: 208, 2007: 78.
Amblyseius sakalava Blommers, 1976: 96 (Synonymy according to Ueckermann & Loots 1988).
Amblyseius amtalaensis Gupta, 1977: 53 (Synonymy according to Gupta 1986).
Amblyseius magnoliae Denmark & Evans, 2011: 69 (Synonym according to Denmark & Evans 2011).
World distribution — Angola, Anjouan Island, Australia, Benin, Brazil, Cambodia, China, Colombia, Cook Islands, Costa Rica, Cuba, Dominican Republic, Figi, French Guiana, Futuna Island, Georgia, Gilbert Islands, Grande Comore Island, Guadeloupe, Guatemala, Guyana, Hawaii, Honduras, India, Indonesia, Iran, Israel, Ivory Coast, Jamaica, Japan, Kenya, La Désirade Island, Madagascar Island, Madeira Island, Malaysia, Marie-Galante Island, Martinique, Mauritius, Mayotte Island, Mexico, Mohéli Island, Mozambique, New Caledonia, New Zealand, Oman, Papua New Guinea, Peru, Philippines, Puerto Rico, Reunion Island, Rodrigues Island, Saint Barthélémy, Saint Kitts, Saint Martin, Saudi Arabia, Sierra Leone, Singapore, Sri Lanka, Tahiti, Taiwan, Tanzania, Thailand, Trinidad, Turkey, USA, US Samoa, Vanuatu, Venezuela, and Vietnam.
Specimens examined — 3 ♀♀ collected on Cupressus spp. L. (Cupressaceae) used as windbreaks at Ouled Abdellah in September 2018. This is the first record of this species in Morocco.
Remarks — The description and measurements of the adult females collected agree with those provided by McMurtry and Moraes (1984) for specimens from South Pacific, by Karmakar et al. 2017 for specimens from West Bengal (India) and by Fang et al. (2020) for specimens from Vietnam. This species is an important natural enemy of mite pests in China (Wu et al. 2009). It prefers high humidity environment, and was found the dominant native species in surveyed citrus orchards in Vietnam (Fang et al. 2020). Amblyseius largoensis is a generalist predator that feeds on a diversity of phytophagous mites, mainly those of the families Eriophyidae, Tenuipalpidae, and Tetranychidae, small insects and other food types, including pollen (Rodriguez and Ramos 2004; Navia et al. 2007).
Amblyseius swirskii Athias-Henriot 1962: 5.
Amblyseius (Amblyseius) swirskii, Ehara 1966: 23.
Typhlodromips swirskii, Moraes et al. 1986: 149, 2004a: 227.
Amblyseius swirskii, Chant & McMurtry 2004: 201, 2007: 81.
Amblyseius rykei, Pritchard & Baker 1962: 249 (Synonymy according to Zannou & Hanna 2011).
Amblyseius enab, El-Badry 1967a: 178 (Synonymy according to Ramadan et al. 2009).
Amblyseius capsicum (Basha, Yousef, Ibrahim & Mostafa 2001): 372 (Synonymy according to Abo-Shnaf & Moraes 2014).
World distribution — Argentina, Azerbaijan, Benin, Burundi, Cape Verde, DR Congo, Egypt, Gaza strip, Georgia, Ghana, Israel, Italy, Kenya, Reunion Island, Saudi Arabia, Senegal, Slovenia, Spain, Syria, Tanzania, Turkey, USA, and Yemen.
Specimens examined — 2 ♀♀ collected on citrus leaves at Stah Al Madina orchard in September 2018; 5 ♀♀ on Solanum nigrum L. (Solanaceae) at Saouda orchard (Ouled Teima) in September 2018. This is the first record of that species in Morocco, naturally occurring or originating from dispersion in the environment after commercial releases in greenhouses.
Remarks — The description and measurements of the adult females collected agree with those provided by Ferragut et al. (2010) for specimens from Spain, by Zannou and Hanna (2011) for specimens from sub-Saharan Africa and by Kreiter et al. (2016a, b) for specimens from Reunion Island and from various countries in the world. Amblyseius swirskii is one of the most efficient phytoseiid species; it is currently released in more than 50 countries of the world (Calvo et al. 2015). It originates from the East Mediterranean coast and has been described in 1962 from almond [Prunus dulcis (Mill.) D.A. Webb.] (Rosaceae) in Bet Dagan, Israel by Athias-Henriot. This species is able to develop not only in the Mediterranean basin, but also in subtropical and tropical areas in Africa (Zannou and Hanna 2011). It is commonly used to control whiteflies and thrips in greenhouse vegetables (especially cucumber and pepper) and some ornamental crops in Europe and North America (Buitenhuis et al. 2014; Calvo et al. 2015).
Graminaseius Chant & McMurtry 2004: 215.
Amblyseius graminis Chant 1956: 34.
Typhlodromus (Amblyseius) graminis, Chant 1959: 89; Arutunjan 1970.
Amblyseius (Typhlodromopsis) graminis, Muma 1961: 287.
Typhlodromus (Typhlodromus) graminis, Westerboer & Bernhard, 1963: 636.
Amblyseius (Amblyseius) graminis, Wainstein 1973: 178, 1975: 920, 1977: 1415; Ueckermann & Loots 1988: 132.
Neoseiulus graminis, Moraes et al. 1986: 81.
Amblyseius (Typhlodromips) graminis, Karg 1991.
Amblyseius graminis, Moraes et al. 2004a: 26.
Graminaseius graminis, Chant & McMurtry 2004: 219, 2007: 85.
Graminaseius collyerae (Chant, 1959): 87 (Synonymy according to Athias-Henriot 1966).
Graminaseius exiguus (Hirschmann, 1962): 26 (Synonymy according to Athias-Henriot 1966).
World distribution — Algeria, Armenia, Australia, Azerbaijan, Azores, Chile, Denmark, England, France, Germany, Greece, Hungary, Iran, Italy, Latvia, Mexico, Moldova, Montenegro, Morocco, Norway, Poland, Portugal, Russia, Spain, Syria, Tunisia, Turkey, Ukraine, and USA.
Previous records from Morocco — This species was already reported by McMurtry and Bounfour (1989) and Tixier et al. (2016) from several regions on many plant families such as Apiaceae, Asteraceae, Brassicaceae, and Lamiaceae.
Specimens examined — 3 ♀♀ on Malva parviflora L. (Malvaceae) in May 2017 at Chabaika orchard; 2 ♀♀ on the same plant in July 2017 at the same domain.
Remarks — The description and measurements of the adult females collected agree with those provided by Döker et al. (2019) for specimens from Turkey. This species is distributed worldwide and has been collected on a great variety of plants including citrus (Kreiter et al. 2010; Sahraoui et al. 2012) and horticultural crops (Tixier et al. 2016).
Euseiini Chant & McMurtry 2005: 191.
Euseiina Chant & McMurtry 2005: 209.
Amblyseius (Amblyseius) section Euseius Wainstein 1962: 15.
Amblyseius (Amblyseius) batus Ueckermann & Loots 1988: 92.
Euseius batus, Moraes et al. 2004a: 62.
World distribution — Kenya, Saudi Arabia, South Africa, and Yemen.
Specimens examined — 4 ♀♀ on Malva parviflora L. (Malvaceae) in July 2017 and in September 2018 at Nour; 3 ♀♀ on the same plant in association with Thrips spp. (Thripidae). This is the first record of this species in Morocco.
Remarks — The description and measurements of the adult females collected agree with those provided by Moraes et al. (2001) for specimens from Sub-Saharan Africa by Basahih et al. (2015) for specimens from Saudi Arabia.
Amblyseius (Amblyseius) dossei Pritchard & Baker, 1962: 286; Ueckermann & Loots 1988: 110.
Euseius dossei, Moraes et al. 1986: 40, 1989: 83, 2004a: 65; Chant & McMurtry 2005: 215, 2007: 120.
World distribution — Benin, Burundi, DR Congo, Ghana, Kenya, Mozambique, Nigeria, Rwanda, Sierra Leone, and Uganda.
Specimens examined — 8 ♀♀ on Malva parviflora L. (Malvaceae) in September 2018 at Ouled Abdellah. This is the first record of this species in Morocco.
Remarks — The description and measurements of the adult females collected agree with those provided by Moraes et al. (2001) for specimens from Sub-Saharan Africa.
Typhlodromus scutalis Athias-Henriot 1958: 183.
Amblyseius scutalis, Athias-Henriot 1960b.
Amblyseius (Typhlodromalus) scutalis, Muma 1961.
Amblyseius (Amblyseius) scutalis, Ueckermann & Loots 1988: 109
Amblyseius scutalis, Ueckermann 1992: 149.
Amblyseius (Euseius) scutalis, Gupta 2003.
Euseius scutalis, Ferragut & Escudero 1997: 233; Swirski et al. 1998: 107; Moraes et al. 1986: 52, 2004a: 82; Chant & McMurtry 2005: 216, 2007: 123.
Euseius delhiensis (Narayanan & Kaur, 1960: 5) (Synonymy according to Wysoki & Bolland 1983).
Euseius rubini (Swirski & Amitai 1961): 196 (Synonymy according to Amitai & Swirski 1966).
Euseius gossipi (El-Badry, 1967a): 177; 1970: 503 (Synonymy according to Wysoki & Bolland 1983).
Euseius libanesi (Dosse, 1967): 30 (Synonymy according to Wysoki & Bolland 1983).
Euseius plumerii Romeih et al. 2010: 28 (Synonymy according to Abo-Shnaf & Moraes 2014).
World distribution — Algeria, Canary Islands, Cape Verde, Cyprus, Egypt, Ghana, Greece, India, Iran, Israel, Italy, Jordan, Madeira Island (Portugal), Morocco, Oman, Pakistan, Peru, Saudi Arabia, Spain, Syria, Tunisia, Turkey, and Yemen.
Previous records from Morocco — This species was reported by McMurtry and Bounfour (1989) and Tixier et al. (2003, 2016).
Specimens examined — 4 ♀♀ on Solanum nigrum L. (Solanaceae) collected at Ouled Abdellah in May 2018; 9 ♀♀ + 5 ♂♂ + 4 im. on Ceratonia siliqua L. (Fabaceae) collected in April 2019 in Immouzer region at the argan forest.
Remarks — The description and measurements of the adult females collected agree with those provided by Moraes et al. (2001) for specimens from Sub-Saharan Africa. This species is mainly reported from the South of the West Palaearctic region and is found in dry climates. It is very common in Maghreb and South of Spain (Kreiter et al. 2004) and recorded in Tunisia on many plants including citrus (Kreiter et al. 2010; Sahraoui et al. 2012). This species is able to feed on T. urticae and P. citri (Abbassy et al. 2012). It can be reared on pollen and is considered as a biological control agent against phytophagous mites, scale insects, and whiteflies on greenhouse cucumber (Nomikou et al. 2002).
Amblyseius stipulatus Athias-Henriot 1960a: 294.
Typhlodromus stipulatus, Hirshmann 1962.
Amblyseius (Amblyseius) stipulatus, Ueckermann & Loots 1988: 110.
Euseius stipulatus, Ferragut et al. 1985: 225; Moraes et al. 1986: 55, 2004a: 84; Chant & McMurtry 2005: 216. 2007: 123.
World distribution — Algeria, Azores, Canary Islands, France, Greece, Hungary, Iran, Italy, Lebanon, Madeira Island, Montenegro, Morocco, Peru, Portugal, Slovenia, Spain, Syria, Tunisia, Turkey, and USA.
Previous records from Morocco — This species was recorded in many regions in Morocco and collected on several plants as reported by Tixier et al. (2016).
Specimens examined — 6 ♀♀ + 2 ♂♂ on Bryonia dioica Jacq. (Cucurbitaceae) collected in Ouled Abdellah citrus orchard in May 2017; 1 ♀ + 1 ♂ were collected on citrus leaves at Saouda orchard in September 2018.
Remarks — The description and measurements of the adult females collected agree with those provided by Ferragut and Escudero (1997) and by Ferragut et al. (2010) for specimens from Spain. Euseius stipulatus was classified by McMurtry and Croft (1997) as a specialized pollen feeder. This species feeds also on pest mites such as P. citri (Ferragut et al. 1988, 1992), T. urticae (Moyano et al. 2009) and eriophyid mites (Ferragut et al. 1987) on many crops such as peach, avocado orchards, and vineyards (Ferragut et al. 1983; Papaioannou-Souliotis et al. 1994; Ragusa 2006) and citrus (Kreiter et al. 2010; Sahraoui et al. 2012).
Iphiseius Berlese 1916: nomen nudum; 1921: 95.
Amblyseius (Iphiseius) Muma 1961: 288.
Iphiseius (Iphiseius) Pritchard & Baker 1962: 298.
Seius degenerans Berlese 1889: 9.
Seiulus degenerans, Berlese 1887: 9.
Amblyseius (Iphiseius) degenerans, Muma 1961: 288.
Iphiseius (Iphiseius) degenerans, Prichard & Baker 1962: 299.
Typhlodromus degenerans, Hirschmann 1962: 2.
Amblyseius degenerans, Zaher 1986: 99.
Iphiseius degenerans, Berlese 1921: 95; Chant 1959: 110; Moraes et al. 1986: 61, 2004a: 92; Chant & McMurtry 2005: 218, 2007: 12.
Iphiseius martigellus El-Badry 1968: 325 (Synonymy according to Chant & McMurtry 2005: 217).
World distribution — Algeria, Azores, Benin, Brazil, Burundi, Cameroon, Canary Islands, Cape Verde, Cyprus, DR Congo, Egypt, Georgia, Ghana, Grande Comore Island, Greece, Israel, Italy, Kenya, Lebanon, Madagascar Island, Madeira Island, Malawi, Morocco, Nigeria, Portugal, Rwanda, Saudi Arabia, Sierra Leone, South-Africa, Syria, Tanzania, Tunisia, Turkey, Uganda, USA, Yemen, Zambia, and Zimbabwe.
Previous records from Morocco — It was reported from Morocco by McMurtry and Bounfour (1989) and by Tixier et al. (2003) on Nerium oleander L. (Apocynaceae) in Gorges of Zegzel.
Specimens examined — 3 ♀♀ on Cupressus spp. (Cupressaceae) collected at Errahim domain in July 2017; 2 ♀♀ and 1 ♀ on the same previous substrate in May 2017 respectively at Taoufiqui 1 and 2 orchards.
Remarks — The description and measurements of the adult females collected agree with those provided by Döker et al. (2018) for specimens from Turkey. This species has a wide distribution in Africa and South Europe and around the Mediterranean Sea (Swirski and Amitai, 1997b; Moraes et al, 2004a; Sahraoui et al, 2012). It is described by McMurtry and Croft (1997) as a generalist (type III predator), able to feed on a wide range of food such as thrips larvae, spider mites, and pollen (Nwilene and Nachman 1996; Van Rijn and Tanigoshi 1999; Vantornhout et al. 2004; Messelink et al. 2006). This species is commercialised for biological control of the western flower thrips Frankliniella occidentalis (Pergande) in greenhouses (Vantornhout et al. 2005).
Phytoseiini Berlese 1913: 3.
Phytoseius Ribaga 1904: 177
Phytoseius finitimus Ribaga 1904: 178; Chant 1959: 108; Moraes et al. 2004a: 252; Chant & McMurtry 2007: 129.
Phytoseius (Dubininellus) finitimus, Wainstein 1959: 1365.
Phytoseius (Pennaseius) finitimus, Pritchard & Baker 1962: 223; Moraes et al. 1986: 214.
Pennaseius finitimus, Schuster & Pritchard 1963: 279.
Phytoseius (Phytoseius) finitimus, Denmark 1966: 16.
Phytoseius dubinini (Beglyarov 1958): 116 (Synonymy according to Pritchard & Baker 1962).
World distribution — Algeria, Azores, Egypt, France, Greece, Iran, Israel, Italy, Montenegro, Morocco, Portugal, Slovenia, Spain, Syria, Tunisia, Turkey, and USA.
Previous records from Morocco — This species was reported from Morocco for the first time by Tixier et al. (2016). It was collected on several regions and plants such as Mentha spp. L. (Lamiaceae), Ficus carica L. (Moraceae), Salix pedicellata Desf. (Salicaceae), and Coleostephus myconis (L.) Rchb.f. (Asteraceae).
Specimens examined — 6 ♀♀ on citrus leaves in July 2017at Taoufiqui 1 domain.
Remarks — The measurements of adult females collected agree with those provided by Tixier et al. (2016) for specimens from Morocco. Phytoseius finitimus is often recorded on plants with highly pubescent leaves (Walter 1992; McMurtry and Croft, 1997). It is found in higher densities on such host plants (Tixier et al. 1998; Duso and Vettorazzo 1999), suggesting that it may be promising candidates for biological control of pests. Hairy structures may offer shelters and domatia with favourable microclimatic conditions (increased relative humidity, reduced temperature) for the development and reproduction of this mite (Walter 1996; McMurtry and Croft 1997; Tixier et al. 1998; Kreiter et al. 2003). Phytoseius finitimus is a generalist phytoseiid mite mainly recorded in the Mediterranean region on a variety of both cultivated and non-cultivated plants, such as grapevine, hazelnut, citrus, walnut, mulberry, peach, plum, pomegranate, and cotton (Swirski and Ragusa 1977; Duso and Vettorazzo 1999; Papaioannou Souliotis et al. 1999; Tsolakis et al. 2000; Nomikou et al. 2001). It is a natural enemy of tetranychid and eriophyid mites, but it also feeds on small insect and pollen (Nomikou et al. 2001; Momen and El-Borolossy 2010).
Typhlodromini Wainstein 1962: 26.
Chant & McMurtry 1994: 240; 2007: 137.
Cydnoseius Muma 1967: 274.
Typhlodromus (Typhlodromus) negevi Swirski & Amitai, 1961: 194.
Typhlodromus negevi, Amitai & Swirski 1966: 21.
Typhlodromus (Neoseiulus) negevi, Ehara 1966a: 19.
Cydnodromella negevi, Chant & Yoshida-Shaul 1986: 2815.
Amblydromella negevi, Moraes et al. 1986: 168.
Cydnoseius negevi, Moraes et al. 2004a: 263; Chant & McMurtry 2007: 137; Abo-Shnaf & Moraes 2014: 39.
Cydnoseius cordiae (Muma 1967): 276 (Synonymy according to Chant & Yoshida-Shaul 1986).
Cydnoseius medanicus (El-Badry 1967b): 108 (Synonymy according to Chant & Yoshida-Shaul 1986).
Cydnoseius zaheri (El-Badry 1967a): 182 (Synonymy according to Chant & Yoshida-Shaul 1986).
Cydnoseius africanus (Yousef 1980): 122 (Synonymy according to Chant & Yoshida-Shaul 1986).
Cydnoseius schusteri (Yousef & El-Borolossy in Zaher 1986): 129 (Synonymy according to Kanouh et al. 2012).
World distribution — Egypt, Israel, Oman, Saudi Arabia, and United Arab Emirates.
Specimens examined — 14 ♀♀ + 2 ♂♂ on Withania somnifera (L.) Dunal. (Solanaceae); 7 ♀♀ on the same plant in July 2017 respectively at Stah AlMadina and Errahim orchards. This is the first record of this species in Morocco.
Remarks — The measurements of adult females fit well with those provided by Alatawi et al. (2017) for specimens from Saudi Arabia.
Paraseiulini Wainstein 1976: 697.
Paraseiulus Muma, 1961: 299.
Typhlodromus talbii Athias-Henriot 1960b: 75.
Typhlodromus (Neoseiulus) talbii, Ehara 1966a: 17.
Paraseiulus (Bawus) talbii, Wainstein, 1976: 699.
Bawus talbii, Moraes et al. 1986: 180; Swirski & Amitai 1990: 117.
Seiulus talbii, Nicotina & Cioffi 1999: 11.
Paraseiulus talbii, Abbasova 1972: 11; Moraes et al. 2004a: 301; Chant & McMurtry 2007: 143.
Paraseiulus subsoleiger, Wainstein 1962: 139 (Synonymy according to Chant & Yoshida-Shaul 1982).
Paraseiulus tetramedius, Zaher & Shehata 1970: 117 (Synonymy according to Chant & Yoshida-Shaul 1982).
Seiulus amaliae, Ragusa & Swirski 1976: 183 (Synonymy according to Chant & Yoshida-Shaul 1982).
Paraseiulus ostiolatus, Athias-Henriot 1978: 699 (Synonymy according to Chant & Yoshida-Shaul 1982).
World distribution — Algeria, Argentina, Austria, Azerbaijan, China, Croatia, Cyprus, Czechoslovakia, Czech Republic, Denmark, Egypt, Finland, France, Georgia, Greece, Hungary, Iran, Israel, Italy, Morocco, Netherlands, Poland, Portugal, Serbia, Slovakia, Slovenia, Spain, Sweden, Syria, Tunisia, and Turkey.
Previous records from Morocco — This species has been reported for the first time by Tixier et al. (2016). It has been collected on Convolvulus althaeoides L. (Convolvulaceae) and Vinca major L. (Apocynaceae).
Specimens examined — 3 ♀♀ + 1 ♂ on citrus leaves in June 2017 at Ouled Abdellah orchard; 2 ♀♀ on Acacia spp. (Fabaceae) used as windbreaks at the same domain.
Remarks — The measurements of adult females collected agree with those provided by Chant and McMurtry (1994).
Typhlodromus triporus Chant & Yoshida-Shaul 1982: 3029.
Paraseiulus triporus, Moraes et al. 1986: 208, 2004a: 303; Hajizadeh et al. 2002: 376; Chant & McMurtry 2007: 143.
World distribution — Czechoslovakia, Czech Republic, Denmark, Finland, France, Georgia, Germany, Greece, Hungary, Iran, Italy, Kazakhstan, Moldova, Morocco, Netherlands, Poland, Portugal, Russia, Serbia, Slovakia, Slovenia, Spain, Sweden, Syria, Turkey, Ukraine, and USA.
Previous records from Morocco — This species has been reported for the first time by Tixier et al. (2003). It has been collected on Ulmus spp. L. (Ulmaceae) at Ifrane city.
Specimens examined — 5 ♀♀ and 4 ♀♀ on citrus leaves at Stah Al Madina and Nour orchards, respectively; 3 ♀♀ on Malva parviflora L. (Malvaceae) at Taoufiqui 1 domain in June 2018.
Remarks — The measurements of adult females collected agree with those provided by Hajizadeh et al. (2002) and by Faraji et al. (2007).
Typhlodromini Wainstein 1962: 26.
Typhloseiulus Chant & McMurtry 1994: 246.
Typhloseiulus eleonorae (Ragusa & Swirski)
Seiulus eleonorae Ragusa & Swirski 1981: 269; Moraes et al. 1986: 231.
Typhlodromus eleonorae, Chant & Yoshida-Shaul 1983: 1150.
Typhloseiulus eleonorae, Chant & McMurtry 1994: 247, 2007: 145; Moraes et al. 2004a: 374.
World distribution — France, Italy, and Morocco.
Previous records from Morocco — The first report of this species was observed by Tixier et al. (2016). Specimens were collected near Larache on Quercus suber L. (Fagaceae), Knautia purpurea Vill. (Caprifoliaceae), Road to Souk El Arba on Quercus suber L. (Fagaceae), Daphne gnidium L. (Thymelaeaceae).
Specimens examined — 5 ♀♀ and 3 ♀♀ were collected on citrus leaves, at Saouda and Taoufiqui 1 domains, respectively in July 2017.
Remarks — The measurements of adult females collected agree with those provided by Ferragut (2018) from Spain.
Neoseiulella Muma 1961: 295.
Neoseiulella litoralis (Swirski & Amitai)
Typhloctonus litoralis Swirski & Amitai 1984: 73-76.
Neoseiulella (Typhloctona) litoralis, Denmark & Rather 1996: 71-72.
Neoseiulella litoralis, Swirski & Amitai 1997b: 37; Moraes et al. 2004a: 294; Chant & McMurtry 2007: 147.
World distribution — Cape Verde, Greece, Israel, Morocco, and Spain.
Previous records from Morocco — This species was reported in Morocco and was mainly found near the Atlantic Ocean coast. Tixier et al. (2016) reported that this species was observed on several plants such as Carlina racemosa L. (Asteraceae), Emex spinosa L. (Polygonaceae), Eryngium maritimum L. (Apiaceae), Euphorbia pithyusa L. (Euphorbiaceae), and Medicago marina L. (Fabaceae).
Specimens examined — 2 ♀♀ on Withania somnifera (L.) Dunal. (Solanaceae) at Saouda (Ouled Teima) in September 2018.
Remarks — The measurements of adult females collected agree with those provided by Kanouh et al. (2010) for specimens from France.
Typhlodromus Scheuten 1857: 111.
Typhlodromus (Anthoseius) De Leon 1959: 258.
Typhlodromus rhenanoides Athias-Henriot 1960b: 85.
Neoseiulus rhenanoides, Schuster & Pritchard 1963: 205.
Anthoseius rhenanoides, Charlet & McMurtry 1977: 186.
Amblydromella rhenanoides, Moraes et al. 1986: 174.
Amblydromella (Aphanoseia) rhenanoides, Denmark & Welbourn 2002: 308.
Typhlodromus (Anthoseius) rhenanoides, Moraes et al. 2004a: 347, Chant & McMurtry 2007: 155.
World distribution — Algeria, Canary Islands, France, Greece, Hawaii, Italy, Les Saintes, Madeira Island (Portugal), Morocco, Portugal, Spain, Tunisia, and USA.
Previous records from Morocco — This species was reported from Morocco by Kreiter et al. (2004), McMurtry and Bounfour (1989) on Cupressus spp. L. (Cupressaceae) and Tixier et al. (2016) on a great variety of plants and locations, such as on Lupinus humulus L. (Fabaceae), Solananum sodomeum L. (Solanaceae), Nicotiana glauca Grah. (Solanaceae), Urtica dioica L. (Urticaceae), and Solanum nigrum L. (Solanaceae).
Specimens examined — 6 ♀♀ + 2 ♂ + 4 im. on Tamarix aphylla L. (Tamaricaceae) in April 2019; 1 ♀ + 1 ♂ + 1 im. on Ficus carica L. (Moraceae) in the argan fores in the Road to Aksri and Imouzzar Ida ou Tenane villages in tha same date.
Remarks — The measurements of adult females agree with redescription provided by Tixier et al. (2016) for specimens from Morocco. This species has been reported on many uncultivated plants and sometimes on some orchards such as peach, plum, olive, citrus, and vineyards (Ragusa 2006). Nothing is known of its ability to control pests (Tixier el al. 2016).
Typhlodromus Scheuten 1857: 111.
Typhlodromus (Typhlodromus) octogenipilus Kreiter et al. 2010: 172.
World distribution — Croatia, Greece, Morocco, and Turkey.
Previous records from Morocco — The first report from Morocco of this recently was provided by Tixier et al. (2016). Specimens have been collected on Cistus libanotis L. (Cistaceae) in Road to Souk El Arba and Mazari Cape.
Specimens examined — 4 ♀♀ on citrus leaves were collected in July 2017 at Taoufiqui 2 domain; 5 ♀♀ on citrus leaves in September 2018 at the same domain.
Typhlodromus setubali Dosse 1961: 313.Typhlodromus (Typhlodromus) setubali, Moraes et al. 1986: 246, 2004a: 369; Chant & McMurtry 2007: 157.Typhlodromus laurentii, Ragusa & Swirski 1978: 213 (Synonymy according to Chant & Yoshida-Shaul 1987).
World distribution — Germany, Israel, Italy, Jordan, Madeira Island (Portugal), Morocco, Portugal, Slovakia, Spain, and Tunisia.
Previous records from Morocco — specimens were collected on several plants such as Quercus suber L. (Fagaceae), Nerium oleander L. (Apocynaceae), Rosmarinus officinalis L. (Lamiaceae), Marrubium incanum L. (Lamiaceae), Cistus spp. L. (Cistaceae), and Thuya spp. L. (Cupressaceae) in Damon Oum Er Bia, Beni Slimane, and Ras El Ma regions (Tixier et al. 2003, 2016). It was also already reported by McMurtry and Bounfour (1989) on Olea europea L. (Oleaceae).
Specimens examined — 4 ♀♀ + 2 ♂♂ on citrus leaves at Nour orchard in July 2018; 2 ♀♀ on Rhamnus spp. L. (Rhamnaceae) and 2 ♀♀ on Zyziphus lotus L. (Rhamnaceae) in April 2019 at Road to Alma village in the argan forest.
Remarks — The measurements of adult females fit well with those provided by Tixier et al. (2016) for specimens from Morocco. The measurements of adult females agree with those provided by Tixier et al. (2010) for specimens from Croatia on Cornus sanguinea L. (Cornaceae) and by Tixier et al. (2016) for specimens from Morocco.
This key contains the 59 species presently recorded from Morocco.
1. Podonotal region of the dorsal shield (anterior to setae R1) of the female with five or six pairs of ''lateral'' setae j3, z2, z4 and s4 always present and z3 and/or s6 present
...... 2
— Podonotal region of the dorsal shield (anterior to setae R1) of the female with four pairs of ''lateral'' setae j3, z2, z4 and s4 present, z3 and s6 absent
...... Sub-family Amblyseiinae: 3
2. Posterior ''lateral'' setae Z1, S2, S4 and S5 absent. Setae r3 usually inserted on the dorsal shield
...... Sub-family Phytoseiinae: Phytoseius finitimus
— At least one of setae Z1, S2, S4 and S5 present. Setae r3 usually inserted on the interscutal soft cuticule (rarely on the shield)
...... Sub-family Typhlodominae: 36
3. Sternal shield with median posterior projection, some forward ''migration'' of pre-anal setae ZV2 and/or JV2
...... 4
— Sternal shield without posterior projection, without forward ''migration'' of pre-anal setae ZV2 and/or JV2
...... 8
4. Heavily sclerotised and brown body with separate anal shield and subrectangular ventral shield
...... Iphiseius degenerans
— Slightly sclerotised and ventri-anal shield entire
...... Genus Euseius: 5
5. Cervix of the spermatheca tubular
...... 6
— Cervix of the spermatheca of a different shape
...... 7
6. Peritreme long, extending to level of j3 or between j3 and z2. stIV relatively short (54 μm). Cervix of spermatheca tubular with a long cervix of 20 to 25 μm long
...... Euseius stipulatus
— Peritreme short, extending at maximum to level of z2. Dorsal setae long. stIV long (77 μm). Cervix of spermatheca tubular, thin, long and sinuous (43 μm)
...... Euseius scutalis
7. All setae very short, peritreme extending to the level of j1. Macrosetae of the leg IV sharp-tipped. Distal half of the cervix thick-walled with the one side curved distally to house the atrium
...... Euseius batus
— setae z2, z4 and s4 long, peritreme extending to the level of j3, macrosetae of the leg IV knobbed, stIV very long. Spermatheca with broad cervix, about twice as long, as wide and slightly narrowing
...... Euseius dossei
8. Setae S4 absent
...... 9
— Setae S4 present
...... 12
9. Setae J2, S2 absent, female ventri-anal shield reduced, setae j6 very long: 2–3 times longer than distance between their bases, female ventri-anal shield with one pair of pre-anal setae
...... Phytoseiulus persimilis
— Setae J2, S2 present, female ventri-anal shield elongated with three pre-anal setae, setae j6 not long
...... 10
10. Striated dorsal shield, ZV3 absent
...... Eharius chergui
— Not striated dorsal shield, ZV3 present
...... 11
11. Presence of six solenostomes on the dorsal shield (gd1, gd2, gd4, gd6, gd8, gd9); ventri-anal shield reduced with two pairs of pre-anal setae
...... Kampimodromus hmiminai
— Presence of four solenostomes on the dorsal shield (gd1, gd2, gd6, gd9); ventri-anal shield not reduced, elongated with threepairs of pre-anal setae
...... Kampimodromus aberrans
12. Z2 present, ventri-anal shield reduced with one pair of pre-anal seta
...... Typhloseiella isotricha
— Z2 absent, ventri-anal shield not reduced
...... 13
13. Absence of J2
...... 14
— Presence of J2
...... 15
14. Differences in the length of the setae Z5 = 170–196, Z4 = 155
...... Proprioseiopsis messor
— Differences in the length of the setae Z5 = 203–225, Z4 = 125–127
...... Proprioseiopsis bordjelaini
15. Macrosetae usually present only on leg IV (rarely missing on this leg) but sometimes also on leg III. Lateral dorsal setae except Z5 usually subequal. J2, Z1, S2, S4 and S5 always present
...... 16
— Macrosetae usually present on legs II, III and IV, and sometimes also on leg I. Lateral dorsal setae often of quite differentlengths. J2, Z1, S2, S4 or S5 may be missing
...... 28
16. Ventri-anal shield reduced and/or markedly wider at anus level, with a marked waist
...... Genus Paragigagnathus: 17
— Female ventri-anal shield not reduced and/or markedly wider at anus level, without a marked waist
...... Genus Neoseiulus: 18
17. Seta st3 inserted on sternal shield of female
...... Paragigagnathus tamaricis
– Seta st3 inserted off sternal shield of female
...... Paragigagnathus molestus
18. Absence of ZV3, dorsal shield longitudinally striated
...... Neoseiulus thymeleae
— Presence of ZV3, dorsal shield not longitudinally striated
...... 19
19. Spermatheca with atrium forked for at least half its length at juncture with major duct, or atrium appearing thick-walled, vacuolated
...... 20
— Spermatheca with atrium not deeply forked at juncture with major duct, nor appearing thick-walled, vacuolated
...... 21
20. Punctuated pores on the ventri-anal shield
...... N. alpinus
— Crescenticpores on the ventri-anal shield
...... N. barkeri & N. stolidus
NB: impossible to distinguish between these two latter species already suspected to be synonyms by Ragusa & Athias-Henriot 1983
21. Spermatheca with elongated cervix
...... 22
— Spermatheca with bell shape cervix
...... 23
22. Very elongated cervix (38 µm)
...... N. pseudotauricus
— Not very elongated cervix
...... 24
23. Non crescentic pores on the ventrianal shield
...... 24
— Crescentic pores on the ventrianal shield
...... 25
24. Z5 = 47, Z4 =38, j3 = 22, s4 =27
...... N. cucumeris
— Z5 = 60–75, Z4 = 23–28, j3 = 18–21, s4 = 23–26
...... N. longilaterus
25. Crescenticpores on the ventri-anal shield
...... N. californicus
— Minute pores on the ventri-anal shield
...... 26
26. Z4 = 75, j3 = 45–50, s4 = 45–60
...... N. ornatus
— Z4 = 32, j3 =17, Z5 = 42–50, s4 = 27–30
...... 27
27. Z4 = 32, j3 =17, Z5 = 42–50, s4 = 27–30
...... N. marinus
— Z4 = 53, j3 =38, Z5 =64, s4 = 52
...... N. umbraticus
28. Ratio seta s4/S2 < 2.7:1
...... 29
— Ratio seta s4/S2 < 3.0:1.0
...... 30
29. Seta S5 present
...... Transeius audeae
— Seta S5 absent
...... Amblyseiella setosa
30. Spermatheca with atrium bifurcate/vacuolated at juncture with major duct
...... Graminaseius graminis
— Spermateca with atrium not bifurcate/vacuolated at the juncture with major duct
...... Genus Amblyseius: 31
31. Setae ZV2 off the ventri-anal shield
...... A. italicus
— Setae ZV2 on the ventri-anal shield
...... 32
32. Ventri-anal pores punctiform
...... 33
— Ventri-anal pores crescentic
...... 34
33. Setae Z5 315; Z4 162
...... A. obtusus
— Setae Z5 180–210; Z4 110–130
...... A. meridionalis
34. Female ventrianal shield vase-shaped, wider at level of anus than at level of setae ZV2 present
...... Amblyseius largoensis
— Female ventrianal shied not vase-shaped, not wider at level of anus than at level of setae ZV2
...... 35
35. Spermatheca with short neck; seta Z5, around 145 µm long
...... A. andersoni
— Spermatheca with calyx cup-shaped; setae Z5, maximum 100-120 µm long
...... A. swirskii
36. Setae z3 absent
...... Cydnoseius negevi
— Seta z3 present
...... 37
37. Setae z6 present
...... 38
— Setae z6 absent
...... 41
38. JV3 Present
...... Kuzinellus saharae
— JV3 absent
...... 39
39. Seta Z3 present
...... Paraseiulus talbii
— Seta Z3 absent
...... 40
40. Genu II 1 2/0, 2/0 1; light vertical reticulation on dorsal shield, dorsal shield elongated
...... Paraseiulus minutus
— Genu II 2 2/1, 2/0 1; distinct to strong squarish reticulation on dorsal shield, rounded shape of the dorsal shield
...... Paraseiulus triporus
41. Setae Z1 absent
...... Genus Typhlodromus: 42
— Setae Z1 present
...... 58
42. Seta S5 present
...... Typhlodromus (Anthoseius): 43
— Seta S5 absent
...... Typhlodromus (Typhlodromus): 49
43. Two pairs of setae on the ventri-anal shield
...... T. (Anthoseius) ilicis
— Four pairs of setae on the ventri-anal shield
...... 44
44. Presence of three pairs (gd2, gd6, gd9) of solenostomes on a strongly reticulated dorsal shield
...... Typhlodromus (A.) recki
— Presence five pairs of solenostomes on a less reticulate dorsal shield
...... 45
45. Ventri-anal shield without pre-anal pores
...... 46
—- Ventri-anal shield with pre-anal pores
...... Typhlodromus (A.) rhenanoides
46. Spermatheca with cylindrical calyx with a neck between atrium and cervix, three macrosetae on leg IV
...... 47
— Spermatheca without a neck between atrium and cervix and a long cylindrical major duct, 1 macrosetae on the leg IV (on the basitarsus)
...... 48
47. Seta j1 23–34, peritreme reaching j1
...... Typhlodromus (A.) foenilis
— Seta j1 shorter 14–22, peritreme extending between j1 and j3
...... Typhlodromus (A.) maspalomensis
48. Peritreme extending between j1 and j3, elongated and very thin metapodal plate 1
...... Typhlodromus (A.) clairathiasae
— Peritreme extending between j1 and j3, not so elongated and thin metapodal plate 1
...... Typhlodromus (A.) athenas
49. Three solenostomes on the dorsal shield
...... 50
— More than three solenostomes on the dorsal shield
...... 53
50. Three pairs of setae on the ventri-anal shield
...... T. (T.) thuriferus
— Four pairs of setae on the ventri-anal shield
...... 51
51. Seven setae on the genu II
...... 52
— Eight setae on the genu II
...... T. (T.) ballotae
52. Z4 length 27 (21–33), Z5 length 40 (32–48)
...... T. (T.) baccettii
— Z4 length 43 (40–48), Z5 length 61 (53–68)
...... .T. (T.) leclanti
53. Six setae on the genu II
...... 54
— More than six setae on the genu II
...... 55
54. Peritreme extending to z2
...... Typhlodromus (T.) setubali
— Peritreme extending to j3
...... Typhlodromus (T.) moroccoensis
55. Genu II with eight setae
...... 56
— Genu II with seven setae
...... 57
56. Z4 length 38 (30–46), Z5 length 59 (49–69), stIV length 33 (26–40)
...... Typhlodromus (T.) octogenipilus
— Z4 length 70 (65–75), Z5 length 91 (85–95), stIV length 45 (40–48)
...... Typhlodromus (T.) mazarii
57. Calyx of the spermatheca squared basally, with a short neck
...... Typhlodromus (T.) exhilarates
— Calyx of the spermatheca rounded basally, without neck
...... Typhlodromus (T.) phialatus
58. No thick setae not araising from tubercules, six small round solenostomes on the dorsal shield (gd1, gd2, gd5, gd6, gd8, gd9)
...... Neoseiulella litoralis
— Thick setae raising from tubercules
...... Typhloseiulus eleonorae
Predacious mites of the family Phytoseiidae are small mesostigmatid mites whose habits are very diverse (Chant 1959; McMurtry et al. 1970, 2013; Kreiter et al. 2001; Tixier et al. 2016). Some species of predatory mites rank among the most important biological control agents used in populations control of various pests (whiteflies, thrips, and mites), such as: Phytoseiulus persimilis, Neoseiulus californicus, Amblyseius andersoni and A. swirskii. Fifty-two species were known from Morocco until 2016 (McMurtry and Bounfour 1989; Kreiter et al. 2001; Tixier et al. 2003; 2016). In this study, we report 29 species of Phytoseiidae in surveys carried out in nine citrus orchards covering an area of over 3000 hectares and from uncultivated plants in the argan forest between Agadir city and Imouzzar Ida ou Tanane. Among the 29 species herein reported (belonging to 15 genera), seven are new for the Moroccan fauna: Amblyseius largoensis, A. swirskii, Amblyseiella setosa, Euseius dossei, E. batus, Paragigagnathus molestus, and Cydnoseius negevi. Some genera were found on citrus leaves such as: Amblyseius, Neoseiulus, Transeius, Typhlodromus, Phytoseiulus, Paraseiulus, and Phytoseius. Phytoseiulus persimilis is collected from four domains. Cydnoseius negevi is observed in two domains, while Amblyseius largoensis and Transeius audeae were observed in only one domain. The Taoufiqui 3 orchard does not contain phytoseiid mites. These results could probably be explained by the importance of chemical applications and their detrimental effect on natural enemy fauna (Howarth 1991; Naranjo 2001; Cock et al. 2010; El Wakeil et al. 2013).
Some species have been mainly found on weeds (Malva parviflora and Convolvulus althaeoides) such as Phytoseius finitimus, Neoseiulella litoralis, Typhlodromus (Typhlodromus) setubali, Typhlodromus (T.) octogenipilus and Paraseiulus talbii. Some weeds such as Malva parviflora and Convolvulus althaeoides show quite high numbers of Phytoseiidae. This can be explained by leaf characteristics, food occurrence or to biological preferences of mite species as reported by Tixier el al. (2016). The new-recorded species were found in Stah Al Madina, Saouda, Ouled Abdellah, Errahim and Nour orchards (Table 1). The global diversity of Phytoseiidae in these four orchards represented more than 70% of species recorded. Results of the present survey will improve phytoseiid fauna knowledge especially in this region of Morocco area with more than 70% of the citrus production intended for export, and will be helpful in biological pest management programs against T. urticae, P. citri and E. orientalis. These species are serious pests, especially species of many fruit trees and grapevines in many countries (Bolland et al. 1998; Hare and Phillips 1992; Meck et al. 2009; Ledesma et al. 2011; González-Domínguez et al. 2015). Among phytoseiid species collected in orchards, some Amblyseiinae such as Neoseiulus californicus, Neoseiulus longilaterus, Neoseiulus cucumeris, Amblyseius andersoni, Amblyseius italicus, Phytoseiulus persimilis, Euseius stipulatus, E. sculatis, Euseius batus, Iphiseius degenerans and some Typhlodrominae can constitute potential biological control agents since some are considered as general predators which feed on a wide variety of food sources including mites, insects, and pollen. Malva parviflora, Convolvulus arvensis, C. althaeoides, Solanum nigrum, Bryonia dioica, Withania somnifera, Tecoma spp. would be among the best plant species to plant in the inter-row in citrus orchards in this region of Morocco to ensure sustainability, high densities and diversity of Phytoseiidae and insect predators species. It would be interesting to complete this survey by studying extent of exchanges of Phytoseiidae between citrus and the ground cover, as well as the influence of the weed management practices currently adopted in orchards on the diversity and density of native fauna, especially to determine how these plants that shelter phytoseiid mites can not also be host plants for mite pests.
We are deeply indebted to presidency of Ibn Zohr University for financial support and to Administration of Agricultural Domains for helpful and for assisting us in monitoring and collecting mite samples and for their important help given during the surveys. Our sincere thanks are also to Mr. Martial Douin for his unconditional involvement in the preparation of the slides for microscope observation at CBGP. We are grateful to anonymous reviewers and office journal which provided helpful comments that greatly improved the manuscript.
We are sincerely very grateful to the two reviewers of this paper for huge improvements of a previous version of this manuscript and especially to Dr Reham Abo-Shnaf, Senior Researcher at Plant Protection Research Institute in Egypt.
The authors declare that they have no conflicts of interest.
Not applicable, this article does not contain any studies with human participants or animals performed by any of the authors.
The datasets used during the current study are available from the corresponding author on reasonable request.