Touré, Moustapha ¹ ; Kwadjo, Koffi Eric ² ; Doumbia, Mamadou ³ and Kreiter, Serge ⁴

¹Université Nangui Abrogoua, UFR-SN, Unité de Recherche en Entomologie Agricole du Pôle Production Végétale, 02 BP 801, Abidjan 02, Ivory Coast.
²Université Nangui Abrogoua, UFR-SN, Unité de Recherche en Entomologie Agricole du Pôle Production Végétale, 02 BP 801, Abidjan 02, Ivory Coast.
³Université Nangui Abrogoua, UFR-SN, Unité de Recherche en Entomologie Agricole du Pôle Production Végétale, 02 BP 801, Abidjan 02, Ivory Coast.
⁴✉ CBGP, Institut Agro Montpellier, INRAE, CIRAD, IRD, Univ Montpellier, Montpellier, France.

2025 - Volume: 65 Issue: 1 pages: 3-35

Original research

Keywords

Abstract

Introduction

Mites of the family Phytoseiidae are predatory species that feed on phytophagous mites and small insects such as thrips and whiteflies on both commercial plants and the endemic vegetation. Several species serve as biological control agents for the control of pest organisms in open field and protected crops worldwide (McMurtry and Croft 1997; McMurtry et al. 2013; Knapp et al. 2018). This family is widespread worldwide, present on all continents except Antarctica, and consists of about 2,500 valid species in 94 genera and three subfamilies (Demite et al. 2024). Biodiversity surveys in poorly investigated areas is still an urgent need, as they may lead to the discovery of additional species potentially useful for biological control, while also enhancing our understanding of biodiversity in these areas (Kreiter et al. 2018a, b, c, 2020a, b, c, 2021; Kreiter and Abo-Shnaf 2020a, b). From this perspective, areas with high biodiversity levels are particularly promising. Most of the African intertropical region constitutes one of the highest world biodiversity hotspots, a concept defined by Myers (1988) to identify regions of critical importance for biodiversity conservation. The common characteristics of these hotspots is their high levels of endemism, combined with the loss of at least 70% of their original natural vegetation (Myers et al. 2000). Knowledge of the phytoseiid diversity in these high interest areas is especially important in the context of global climate change, as it may help identify potential biological control agents (BCA) and support the future establishment of conservation programs.

Ivory Coast is an agricultural country where agriculture plays a crucial role in the national economy, contributing to one-third of the Gross Domestic Product (Gnago et al. 2010). In addition, in order to export crops such as coffee and cocoa, there are booming fruit crops such as papaya (Carica papaya L., Caricaceae). Papaya cultivation is more practiced in the southern and central regions of the country and has expanded significantly due to the Agricultural Export Promotion and Diversification Project (PPDEA). However, papaya production faces substantial challenges due to pressure from various pests, including phytophagous mites, which cause severe damage in papaya orchards. Biological control of these phytophagous mites required the use of predatory arthropods including mites of the family Phytoseiidae. Indeed, almost all species in this family are considered potential biological control agents (BCA) of mites and small phytophagous insects (McMurtry and Croft 1997; McMurtry et al. 2013).

Despite nearly 60 years of faunal studies conducted worldwide to search for phytoseiid species, many areas remain minimally explored or completely unexplored (Kreiter et al. 2020a, b, c, d). In Ivory Coast, very few surveys have focused on phytoseiids. Ragusa and Athias-Henriot (1983) recorded Amblyseius masiaka (Blommers & Chazeau, 1974), A. usitatus = N. usitatus (Van der Merwe), and Neoseiulus erugatus Ragusa and Athias-Henriot from soil samples. Athias-Henriot (1977) described N. lamticus (Athias-Henriot) from the Lamto region, in central Ivory Coast. Twenty kilometres from Yopougon, in the capital of Ivory Coast, Zannou et al. (2007) identified the species Amblyseius sundi Pritchard & Baker on Eucommia ulmoides Oliver (Eucommiaceae). Finally, Moraes et al. (1989a) recorded the species A. largoensis Muma. It should be noted that Amblyseius masiaka and N. usitatus are both junior synonyms of Neoseiulus barkeri Hughes (Ueckermann and Loots, 1988). Consequently, five species are currently recorded from Ivory Coast: N. barkeri, N. erugatus, N. lamticus, A. masiaka and A. sundi.

These results indicate that few studies have been carried out on the fauna of Phytoseiidae in Ivory Coast. This study was therefore carried out during years 2017 and 2018 on Carica papaya L. in order to contribute to a better knowledge of Phytoseiidae mites for biological control of crop pests in Ivory Coast. Here, we present the faunistic results.

Material and methods

Sampling of mites

The sampling of Phytoseiidae mites was conducted from 14/III/2017 to 17/IV/2018, as a part of the PhD thesis of the senior author (M. Touré) in Ivory Coast. The study covered three areas: Anyama, Yobouekro, and NGattakro, located in the departments of Abidjan, Toumodi, and Yamoussoukro, respectively. The average annual temperature and rainfall in these areas are 153.34 mm and 27.64 °C, 103.76 mm and 26.41 °C and 97.44 mm and 26.41 °C, respectively.

The sampling consisted of collecting papaya leaves from orchards at the different sites. The papaya leaves were cut, placed in transparent plastic bags, and transported to the laboratory for processing. Mites were collected from the papaya leaves using a binocular microscope. To facilitate observation under the microscope, the papaya leaves were cut into smaller lobes. A small brush soaked in alcohol was used to collect the mites, which were then transferred into microtubes containing 70% ethanol. These microtubes were labeled with information such as the date, collection area, plot, and sample block number.

Mounting of mites

The mites were mounted between the slide and coverslip using Hoyer's fluid (Walter and Krantz 2009). Slides were placed in an oven at 45 °C and subsequently sealed with a durable nail polish to ensure long-term preservation.

Taxonomical concepts

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, 2024) for distribution and information on descriptions and re-descriptions were used. World distribution indicated for each species is the world distribution prior to this study.

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, numbered I to IV depending on the position on legs I, II, III or IV) 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 the Results section should be considered as absent. Type of spermatheca or insemination apparatus are that of Denmark and Evans (2011). All species recorded are measured and data mentioned in the literature for those species are provided in this paper. All measurements are given in micrometres (µm) and presented with the mean in bold followed by the range in parenthesis.

Classification of plants follows the APG IV classification of 2016 (ex. Byng et al. 2018). Specimens of each species are deposited in the mite collections of Montpellier SupAgro conserved in UMR CBGP INRA/IRD/CIRAD/Institut Agro/University of Montpellier.

Abbreviations used in tables

The following abbreviations are used in this paper for morphological characters (Table 1 to 19) in addition to macrosetal notation (see above): dsl = dorsal shield length just above j1 to just below J5 in the middle line; dsw s4 = dorsal shield width at the level of s4; dsw R1 = dorsal shield width at the level of R1; gensl = genital shield length; gensw st5 = genital shield width at level of steae st5; gensw post. corn. = 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 shortest inguinal sigilla (= ''metapodal plate″) length; sisw = secondary or shortest inguinal sigilla (= ''metapodal plate″) width; vsl = ventrianal (or ventral for Iphiseius degenerans) shield length; asl = anal shield length; ; gv3 – gv3 = distance between centers of solenostomes gv3 on the ventrianal shield; vsw ant. corn. & vsw anus = ventrianal shield width at anterior corners level and at paranal setae level; asw = anal shield width; scl: calyx total length; scw = calyx widest width; fdl = fixed digit length; mdl = movable digit length; No teeth Fd = number of teeth on the fixed digit; No teeth Md = number of teeth on the movable digit; shaft = length of the shaft of spermatodactyl; branch = length of the toe; BCA = Biological control agent; aasl = altitude above sea level.

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; IA = Institut Agro; INRAE = Institut National de Recherche pour l′Agriculture, l′Alimentation et l′Environnement; IRD = Institut de Recherche pour le Développement; UMR = Unité Mixte de Recherche.

Results

We collected 12 species, 11 of which are new records for Ivory Coast.

All recorded species belong to the subfamily Amblyseiinae, with no representatives of the subfamilies Phytoseiinae and Typhlodrominae found on the papaya plant.

The species recorded belong to three Amblyseiinae tribes: Neoseiulini, Amblyseiini, Euseiini, and to five genera: Neoseiulus (2 species), Amblyseius (3 species), Proprioseiopsis (2 species), Iphiseius (1 species) and Euseius (4 species).

In this paper, we present taxonomical information, measurements, world distribution, list of field collections and general information (when available) on these twelve species: Neoseiulus longispinosus (Evans), N. teke Pritchard & Baker, Amblyseius sundi Pritchard & Baker, A. swirskii Athias-Henriot, A. tamatavensis Blommers, Proprioseiopsis mexicanus (Garman), P. ovatus (Garman), Iphiseius degenerans (Berlese), Euseius fustis (Pritchard & Baker), E. lokele (Pritchard & Baker), E. nyalensis (El-Badry), and E. ovaloides (Blommers).

Subfamily Amblyseiinae Muma

Amblyseiinae Muma, 1961: 273

Tribe Neoseiulini Chant & McMurtry

Neoseiulini Chant & McMurtry 2003: 6

Genus Neoseiulus Hughes

Neoseiulus Hughes, 1948: 141

Neoseiulus longispinosus (Evans)

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

Typhlodromus (Amblyseius) longispinosus, Chant 1959: 74.

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

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

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

This species is widely distributed in many countries of the world, mainly in tropical areas (Moraeset al. 2000; Mailloux et al. 2010; Kreiter et al. 2013, 2018 a, c; Demite et al. 2024). It was found rarely in surveys made in Guadeloupe, Martinique and La Réunion, except in studies on companion plants in citrus orchards (Mailloux et al. 2010; Kreiter et al. 2013, 2018c; Le Bellec et al., unpub. data). The species seems to be more common on weeds with populations of tetranychid mites. Neoseiulus longispinosus, a type II phytoseiid predatory mite, like N. californicus (McMurtry et al. 2013), has received increasing attention in Asia since 2010 for the control of different spider mites (of genera Eutetranychus, Oligonychus, and Tetranychus) (Nusartlert et al. 2011). The feeding, development, predation, cannibalism, intra-guild predation and behaviour have been extensively studied by several authors (see for exampleLuong et al. 2017) for pest control purposes. Neoseiulus longispinosus is well-known as a BCAused in many countries worldwide for spider mite management. A recent study by Huyen et al. (2017) have shown that, at least in controlled laboratory conditions, N. longispinosus is apotential biological control agent against the citrus red mite, P. citri.

Tetranychus urticae is a major pest of papaya in south Florida, where Neoseiulus longispinosus, a newly naturalized predator, has become the most abundant predator associated with this pest. Single releases of N. longispinosus significantly suppressed TSSM populations. That study demonstrated that N. longispinosus has the potential to control TSSM on papaya, but the level of control can be negatively affected by high levels of cannibalism at low prey densities, as well as intraguild predation by aggressive intraguild predators, like A. swirskii (Döker et al. 2021).

This is the first record of this species in Ivory Coast.

Specimens examined — 45 ♀♀ and 1 ♂ collected between 14/III/2017 and 10/III/2018 in Abidjan, Anyama Ahoue (aasl 42 m, Lat 5°26′00.87″N, Long 3°55′00.60″W), on Carica papaya L.

World distribution — Australia, China (Fujian, Guangdong, Guangxi, Hainan, Yunnan), Cuba, Dominican Republic, Egypt, Grande Comore Island (Comoros), Guadeloupe Island (France), Hawaii, Hong Kong, India (Andaman Islands, Andhra Pradesh, Bihar Pradesh, Kerala, Orissa, Pondicherry, Tamil Nadu, Uttar Pradesh), Indonesia, Japan, Les Saintes Islands (France), Malaysia, Marie-Galante Island (France), Martinique (France), New Zealand, Nicaragua, Pakistan, Papua New Guinea, Philippines, La Réunion Island (France), Rodrigues Island (Mauritius), Russia (Primorsky Territory), Saint-Barthélémy Island (France), South Korea, Sri Lanka, Taiwan, Thailand, USA (Florida), Vietnam.

Remarks — The measurements of the female and male specimens from Ivory Coast (Table 1 and 2) overlap with those obtained from populations in various countries. These measurements are close to those obtained for specimens from FCI and La Réunion Island.

Neoseiulus teke Pritchard & Baker

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

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

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

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

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

This species is found in sub-Saharan Africa often associated with Mononychellus tanajoa (Bondar), the cassava green mite (CGM). It has been studied for its potential as BCA against the CGM. Nwilene and Nachman (1996) studied its reproduction characteristics on M. tanajoa.

It was more efficient than Iphiseius degenerans (Berlese) but seems not efficient enough in field conditions (Nwilene and Nachman 1996). Quilici et al. (2000) and Kreiter et al. (2020c) have collected this specieson La Réunion Island. Measurements of specimens collected during this study are provided in table 3. This is the first record of that species for Ivory Coast.

Specimens examined — a single specimen (1 ♀) collected between 14/III/2017 and 10/III/2018 in Abidjan, Anyama Ahoue (aasl 42 m, Lat 5°26′00.87″N, Long 3°55′00.60″W), on Carica papaya L.

World distribution — Burundi, DR Congo, Ghana, Grande Comore Island (Comoros), Kenya, Madeira Island (Portugal), Malawi, Mayotte Island (France), Mohéli Island (Comoros), Mozambique, La Réunion Island (France), Rwanda, Sierra Leone, South Africa, Tanzania, Zimbabwe.

Remarks — The measurements of morphological characters for the single N. teke female specimen from Ivory Coast (Table 3) are very close to those of specimens from neighbouring countries, especially from various African countries and La Réunion. However, they differ from the holotype from Congo (Zannou et al. 2006) and specimens from South Africa, which are larger (van der Merwe 1965).

As this species is a potential BCA, and as it is very difficult to obtain type specimens from Museum for old described species, it would be very important to undertake molecular studies and / or crossbreeding experiments with population collected in the field to determine whether all specimens of this various countries / locations belong to the same species or represent several different cryptic species.

Tribe Amblyseiini Muma

Amblyseiini, Muma, 1961: 68

Subtribe Amblyseiina Muma

Amblyseiina Muma, 1961: 69.

Genus Amblyseius Berlese

Amblyseius Berlese, 1914: 143.

Amblyseius sundi Pritchard & Baker

Amblyseius (Amblyseius) sundi Pritchard & Baker, 1962: 244.

Amblyseius sundi, Moraes et al. 1989a: 97; Chant &McMurtry 2004: 212; Moraes et al. 2004b: 52.

Amblyseius (Proprioseiopsis) sundi, Matthysse & Denmark, 1981: 344.

Amblyseius sundi belongs to the sundi species group and the sundi species subgroup. This species has already been recorded from the Ivory Coast. Blommers (1976) observed some Tetranychus neocaledonicus André and Brevipalpus sp. on the lemon trees on which this species was collected. But the biology of this species remain unknown.

Specimens examined — a single specimen (1 ♀) collected between 14/III/2017 and 10/III/2018 in Abidjan, Anyama Ahoue (aasl 42 m Lat 5°26′00.87″N, Long 3°55′00.60″W), on Carica papaya L.

World distribution — Benin, Burundi, Cameroon, Cape Verde, Congo, Cuba, DR Congo, Ghana, Guadeloupe Island (France), Ivory Coast, Kenya, Madagascar Island, Malawi, Mozambique, Nigeria, Rwanda, Sierra Leone, Uganda, Zimbabwe.

Remarks — The measurements of the single specimen collected are very close to those of specimens from other African countries (Table 4).

Amblyseius swirskii Athias-Henriot

Amblyseius swirskii Athias-Henriot, 1962: 5; Swirski et al. 1998: 102; Chant & McMurtry 2004: 201.

Typhlodromips swirskii, Moraes et al. 2004b: 227.

Typhlodromips capsicum Basha, Youssef, Ibrahim & Mostafa 2001: 372 (synonymy according to Abo-Shnaf & Moraes 2014).

Amblyseius enab El-Badry 1967: 178 (synonymy according to Abo-Shnaf & Moraes 2014).

Amblyseius (Amblyseius) rykei Pritchard & Baker 1962: 249 (synonymy according to Zannou et al. 2007).

Amblyseius swirskii belongs to the obtusus species group, characterized by the presence of setae J2 and Z1, setae z4 are minute and a female ventrianal shield is neither vase-shaped nor divided. Within this group, it belongs to andersoni species subgroup (120 species), as its spermatheca has a cup-shaped calyx. The predatory mite Amblyseius swirskii Athias-Henriot, 1962 (Acari: Phytoseiidae) is one of the most efficient predators; it is currently released in more than 50 countries of the world. Originally described in 1962 from almond (Prunus amygdalus [Miller] D.A. Webb) in Bet Dagan, Israel, by Athias-Henriot. The species was reported along the coast of Israel, in the Middle East, Southern Europe, Sub-Saharan Africa, and the Americas (Demite et al. 2024).

This species is able to develop not only in the Mediterranean basin but also in subtropical and tropical areas (Zannou and Hanna 2011). Since it does not enter diapause, it is suitable for use throughout much of the growing season in regions where daytime temperatures regularly exceed 22 °C (Calvo et al. 2015). Amblyseius swirskii is commonly used to control whiteflies and thrips in greenhouse vegetables (especially cucumber, pepper and eggplant) and some ornamental crops, in Europe and North America (Calvo et al. 2015). The biology of this species and its importance for biocontrol were comprehensively reviewed by Calvo et al. (2015) and Buitenhuis et al. (2015). In 2015, a species ressembling to A. swirskii was discovered after a thrips outbreak on peppers and roses on La Réunion Island (Kreiter et al. 2016a, b), located thousands of kilometres from its supposed native area. Morphological and molecular analysis confirmed its identity as A. swirskii (Kreiter et al. 2016a, b). This is the first record of this species from Ivory Coast.

Specimens examined — 7 ♀♀ collected between 14/III/2017 and 10/III/2018 in Abidjan, Anyama Ahoue (aasl 42 m, Lat 5°26′00.87″N, Long 3°55′00.60″W), 3 ♀♀ and 1 ♂ collected between 05/IV/2017 and 17/IV/2018 in Toumodi, Yobouekro (aasl 184 m, Lat 6°31′19.196″N, Long 5°6′41.36″W) and 32 ♀♀ collected between 06/04/2017 au 18/03/2018 in Yamoussoukro, Ngattakro (aasl 158 m, Lat 6°49′39.443″N, Long 5°17′21.635″W) on Carica papaya L.

World distribution — Argentina, Azerbaijan, Benin, Burundi, Cape Verde, DR Congo, Egypt, Gaza Strip, Georgia, Ghana, Israel, Italy, Morocco, Reunion Island, Saudi Aarabia, Senegal, Slovenia, Spain, Syria, Tanzania, Türkiye, USA (California and Florida), Yemen.

Remarks — The measurement values (Tables 5 and 6) are very close to those reported in the literature, especially for specimens from Africa.

Amblyseius tamatavensis Blommers

Amblyseius tamatavensis Blommers 1974: 144; Moraes et al. 1986: 31; Denmark & Muma 1989: 13; Chant & McMurtry 2004: 203; Ehara & Amano 2004: 17; Moraes et al. 2004b: 52; Chant & McMurtry 2007: 81; Döker et al. 2018: 101.

Amblyseius (Amblyseius) tamatavensis, Ehara 2002: 33; Ehara & Amano 2002: 322.

Amblyseius aegyptiacus Denmark & Matthysse in Matthysse & Denmark 1981: 343 (synonymy according to Denmark & Muma 1989)

Amblyseius maai Tseng 1976: 123 (synonymy according to Denmark & Muma 1989).

Amblyseius tamatavensis, like the previous species, belongs to the obtusus species group, characterized by the presence of setae J2 and Z1, setae z4 are minute, and a female ventrianal shield that is neither vase-shaped nor divided. It is further classified in the aerialis species subgroup (46 species) due to the tubular calyx of the spermatheca (Chant and McMurtry 2004a).

It seems to fit the type III-b feeding habit (generalist predators living on glabrous leaves) as defined by McMurtry et al. (2013). Cavalcante et al. (2017) reported this species as a promising natural enemy of B. tabaci. Experimental releases on caged plants in a screen house have demonstrated its efficacy, reducing B. tabaci densities of on pepper plants by 60-80% (Massaro and Moraes 2019). Additionally, it can be easily mass-produced in large numbers using astigmatine mites as a food source, making it suitable for augmentative biological control programs (Massaro et al. 2021). This species has been reported in tropical areas from over 20 countries around the world including Africa, Asia, Americas, and Oceania. Measurements of specimens collected during this study are provided in table 7. This is the first record of this species from Ivory Coast.

Specimens examined — 4 ♀♀ and 1 ♂ collected between 14/III/2017 and 10/III/2018 in Abidjan, Anyama Ahoue (aasl 42 m, Lat 5°26′00.87″N, Long 3°55′00.60″W) on Carica papaya L.

World distribution — Australia (New South Wales, Queensland), Benin, Brazil (Alagoas, Amazonas, Bahia, Ceará, Esperíto Santo, Goiás, Minas Gerais, Pará, Pernambuco, Piauí, Rio de Janeiro, Rio Grande do Norte, Rio Grande do Sul, Roraima, Sergipe, São Paulo, Tocantins), Burundi, Cameroon, Cook Islands, Cuba, Dominican Republic, DR Congo, Easter Island/Isla de Pascua (Chile), Fiji, Ghana, Guadeloupe Island (France), Indonesia, Japan, Kenya, Madagascar Island, Malawi, Malaysia, Marie-Galante Island (France), Martinique (France), Mauritius Island, Mayotte Island (France), Mozambique, Nigeria, Papua New Guinea, Peru, Philippines, Réunion Island (France), Rodrigues Island (Mauritius), Rwanda, Singapore, South Africa, Sri Lanka, Taiwan, Thailand, Uganda, USA (Florida), Vanuatu, Venezuela, Vietnam, Western Samoa.

Remarks — The measurement values (Tables 7 and 8) are very close to that reported in the literature, especially for specimens from Africa. However, the current specimens exhibit r3, z2, Z5 and SgeIV being shorter compared to those of previous reports.

Subtribe Proprioseiopsina Chant & McMurtry

Proprioseiopsina Chant & McMurtry 2004: 219.

Genus Proprioseiopsis Muma

Proprioseiopsis Muma 1961: 277.

Proprioseiopsis mexicanus (Garman)

Amblyseiopsis mexicanus Garman 1958: 75.

Amblyseius mexicanus, Moraes & McMurtry 1983: 134.

Proprioseiopsis mexicanus, Muma & Denmark 1970: 48; Denmark & Muma 1973: 237; Moraes et al. 1986: 118; Kreiter & Moraes 1997: 379; Moraes et al. 2004b: 181; Chant & McMurtry 2005a: 13, 2007: 89.

Amblyseiulus amotus Zack 1969: 72 (Synonymy according to Denmark & Evans 2011).

Typhlodromus (Amblyseius) asetus Chant, 1959: 80 (Synonymy according to Denmark & Evans 2011).

Amblyseiulus clausae Muma 1962: 1 (Synonymy according to Denmark & Evans 2011).

Amblyseius kogi Chant & Hansell 1971: 713 (Synonymy according to Denmark & Evans 2011).

Typhlodromus (Amblyseius) putmani Chant 1959: 91 (Synonymy according to Denmark & Evans 2011).

Amblyseeiulus temperellus Denmark & Muma1967: 171 (Synonymy according to Denmark & Evans 2011).

Amblyseiopsis tropicanus Garman 1958: 77 (Synonymy according to Denmark & Evans 2011).

Amblyseius (Proprioseiopsis) tulearensis Blommers 1976: 100 (Synonymy according to Denmark & Evans 2011).

Amblyseiulus versutus Zack 1969: 74 (Synonymy according to Denmark & Evans 2011).

Proprioseiopsis mexicanus belongs to the belizensis species group as genu I lacks macrosetae. It is classified within the asetus species subgroup due to its spermatheca having a short, cup-shaped calyx (Chant and McMurtry 2005a). This species is known from all islands of French West Indies (Kreiter and Moraes 1997; Moraes et al. 2000, Kreiter et al. 2006; Mailloux et al. 2010; Kreiter et al. 2018c) but was found in large number only during a previous study on companion plant in Guadeloupe (Mailloux et al. 2010) and on La Réunion (Le Bellec, unpub. data). This species seems to be very abundant on weeds in the lower vegetation. Phytoseiid mites of the genus Proprioseiopsis are mainly found in ground surfaces, humus, litter, soil, moss or on grass (Muma and Denmark 1970; McMurtry et al. 2015). Populations of P. mexicanus increase when fed T. urticae eggs (Megevand et al. 1993), and this species has shown potential as a predator of thrips (Kreiter, unpub. data). It is one of the prevailing phytoseiid species in citrus orchards in Alabama (Fadamiro et al. 2009). Denmark and Evans (2011) mentioned that the species can be reared on T. urticae and Oligonychus pratensis (Banks) and is associated with Bryobia praetiosa Koch, Bryobia sp. and P. ulmi. It was also found in association with Tetranychus evansi Baker and Pritchard (Furtado et al. 2014), although it is mentioned as a poor predator of this species. The biology of this species remains, however, almost unknown. This is the first record of this species from the Ivory Coast.

Specimens examined — a single female (1 ♀) collected between 14/III/2017 and 10/III/2018 in Abidjan, Anyama Ahoue (aasl 42 m, Lat 5°26′00.87″N, Long 3°55′00.60″W) on Carica papaya L.

World distribution — Australia, Benin, Brazil (Alagoas, Bahia, Distrito Federal Maranhão, Mato Grosso do Sul, Paraná, Paraíba, Pernambuco, Piauí, Rondonia, São Paulo, Tocantins), Colombia, Cuba, Ghana, Guadeloupe Island (France), Hawaii, Kenya, Martinique Island, Mexico, New Zealand, Panama, Peru, Réunion Island (France), Rodrigues Island (Mauritius), USA (Alabama, Florida, Georgia, Iowa, Kansas, Louisiana, Maryland, Minnesota, Missouri, New Jersey, North Carolina, Ohio, Pennsylvania, West Virginia).

Remarks — The measurements of the single female specimen of the Ivory Coast (Table 9) fit well those obtained for populations of various countries. The number of setae on genu II of our single specimen is the same than the number given by Moraes et al. (2007) with the same formula: 2-2/0,2/0-1. This character may be of interest with some molecular taxonomy in the future, in the framework of an integrative taxonomy, in order to distinguish if we have one species or a species complex.

Proprioseiopsis ovatus (Garman)

Amblyseiopsis ovatus Garman 1958: 78.

Amblyseiulus ovatus, Muma 1961: 278.

Typhlodromus (Amblyseius) ovatus, Chant 1959: 90.

Typhlodromus ovatus, Hirschmann 1962: 19.

Proprioseiopsis (Proprioseiopsis) ovatus, Karg 1989: 208.

Proprioseiopsis ovatus, Moraes et al. 1986: 121; 2004b: 184; Chant & McMurtry 2005a: 15; 2007: 89.

Proprioseiopsis antonelli Congdon 2002: 15 (Synonymy according to Denmark & Evans 2011).

Amblyseiuluscannaensis Muma 1962: 4 (Synonymy according to Denmark & Evans 2011).

Amblyseius hundsonianus Chant & Hansell 1971: 723 (Synonymy according to Denmark & Evans 2011).

Amblyseius parapeltatus Wu & Chou 1981: 274 (synonymy according to Tseng 1983).

Amblyseius (Amblyseius) peltatus van der Merwe 1968: 119 (synonymy according to Tseng 1983).

Like P. mexicanus, P. ovatus belongs to the belizensis species (see above). As the spermatheca of this species is saccular, it belongs to the belizensis species subgroup (Chant and McMurtry 2005a).

This species is known from Guadeloupe, Marie-Galante and Martinique (Kreiter and Moraes 1997; Moraes et al. 2000; Mailloux et al. 2010; Kreiter et al. 2018c). It was found in large number only during a previous study on companion plants in Guadeloupe (Mailloux et al. 2010) and a recent study on La Réunion (Le Bellec, unpub. data). In other habitats, this species seems to be rare. Similar to P. mexicanus, P. ovatus seem to abundant on weeds in the lower vegetation. Denmark and Evans (2011) indicated that this species is associated with O. pratensis and Brevipalpus sp. It was also found in association with T. evansi (Furtado et al. 2014) but is considered a poor predator of that species. Despite this findings, the biology of this species remains largely unknown. This is the first record of this species from the Ivory Coast.

Specimens examined — a single female (1 ♀) collected between 14/III/2017 and 10/III/2018 in Abidjan, Anyama Ahoue (aasl 42 m, Lat 5°26′00.87″N, Long 3°55′00.60″W) on Carica papaya L.

World distribution — Argentina, Brazil (Alagoas, Bahia, Distrito Federal, Goías, Mato Grosso do Sul, Mato Grosso, Minas Gerais, Paraná, Pará, Rio Grande do Sul, São Paulo), Colombia, Costa Rica, Cuba, Ecuador, Egypt, French Guiana (France), Ghana, Grande Comore Island (Comoros), Hawaii, Honduras, Japan, Malaysia, Martinique Island (France), Mayotte Island (France), Mozambique, Peru, Philippines, Puerto Rico, La Réunion Island (France), Saudi Arabia, Sierra Leone, South Africa, Spain, Sri Lanka, Taiwan, Thailand, Türkiye, USA (Arizona, California, Florida, Kansas, Louisiana, Minnesota, Missouri, New Mexica, Texas, Utah, Washington), Venezuela.

Remarks– The measurements of the single female specimen from the Ivory Coast (Table 10) fit well those obtained for populations from various countries.

Tribe Euseiini Chant & McMurtry

Euseiini Chant & McMurtry 2005b: 191.

Genus Iphiseius Berlese

Iphiseius Berlese, 1916: 33, Chant & McMurtry 2005b: 217, 2007: 123.

Iphiseius degenerans (Berlese)

Seius degenerans Berlese 1889: 9.

Amblyseius (Iphiseius) degenerans, Muma 1961: 288.

Typhlodromus degenerans, Hirschmann 1962: 2.

Iphiseius (Iphiseius) degenerans, Pritchard & Baker 1962: 299.

Amblyseius degenerans, Zaher 1986: 99, Northcraft 1987: 521, Papadoulis & Emmanouel 1991: 36.

Iphiseius degenerans, Berlese 1921: 95, Evans 1954: 518, Moraes et al. 1986: 61, 2004b: 92, Chant & McMurtry 2005b: 215, 2007: 125.

Iphiseius martigellus El-Badry 1968: 325 (synonymy according to Chant & McMurtry 2005b).

The biological characteristics of this Ethiopian species have been well documented because of its use in controlling thrips on various cultivated plants in greenhouses. Iphiseius degenerans is a commercially available biological control agent of thrips and spider mites in greenhouse crops. Although it can feed on a wide range of foods, thrips larvae and sweet pepper pollen are less favourable for immature development (Vanthornhout et al. 2004, 2005), potentially hindering the establishment of this predator in sweet pepper crops. According to McMurtry et al. (2013), I. degenerans is classified as a type-IV generalist predator, it is one of the most common native phytoseiid mite species on cassava in southern Africa (Zannou et al. 2005) and feeds on Mononychellus tanajoa (Bondar) (Nwilene and Nachman 1996), a widely distributed neotropical mite pest of cassava in Africa, as well as insect larvae and pollen of many plants (Vantornhout et al. 2005). Another study concluded that I. degenerans can be considered a suitable biological control candidate based on its preference with Eutetranychus orientalis (Klein) in the Mediterranean region (Fantinou et al. 2012). Iphiseius degenerans preys on Oligonychus perseae Tuttle, Baker & Abbatiello outside the webbed nests. Although I. degenerans contributed towards the control of O. perseae, it is effectiveness is limited and needs further investigation, considering the inclusion of alternative food (e.g. Castor oil pollen) for predator population growth (Zappala et al. 2015). This is the first record of this species from the Ivory Coast.

Specimens examined — 34 ♀♀ and 21 ♂♂ collected between 05/IV/2017 and 17/IV/2018 14/III/2017 and 10/III/2018 in Abidjan, Anyama Ahoue (aasl 42 m, Lat 5°26′00.87″N, Long 3°55′00.60″W) on Carica papaya L.

World distribution — numerous countries in Northern and Southern Africa (Demite et al. 2024), Mediterranean area (Cyprus, Greece, Italy, Portugal), Near East or Middle East (Egypt, Israel, Lebanon, Saudi Arabia, Syria, Türkiye, Yemen), in Europe (Georgia), South America (Brazil) and in North America (USA in California, Florida, Georgia, New Hampshire), Indian Ocean (Grande Comore Island, Comoros).

Remarks — The measurements of the 10 ♀♀ + 1 ♂ (Tables 11 & 12) fit well with measurements of specimens reported in the literature.

Genus Euseius Wainstein

Amblyseius (Amblyseius) section Euseius, Wainstein, 1962: 15;

Euseius De Leon, 1967: 86.

Euseius fustis (Pritchard & Baker)

Amblyseius (Amblyseius) fustis Pritchard & Baker, 1962: 283; Ueckermann & Loots, 1988: 83.

Euseius fustis, Matthysse & Denmark, 1981: 348; Moraes et al. 1986: 45, 2004b: 69; Moraes & McMurtry, 1988: 15; Chant & McMurtry 2005b: 215, 2007: 121.

Bruce-Oliver et al. (1996) have tested various foods associated with cassava for their effect on the development, fecundity and longevity of Euseius fustis, the most common phytoseiid species found on cassava in Africa. Euseius fustis developed successfully to adulthood on the spider mite prey species Mononychellus tanajoa (Bondar) and Oligonychus gossypii (Zacher) and on pollen from maize, castor bean, and cassava. Euseius fustis also completed development on water-diluted phloem exudate from cassava, diluted honeydew from the cassava mealybug and on various pollen and prey combinations. When reared on Tetranychus urticae Koch prey or free water only, E. fustis did not develop after the deutonymphal stage. All larvae held on clean leaf discs on water-soaked cotton died without moulting. Diets of maize plus castor bean pollen and maize pollen plus M. tanajoa resulted in the highest rate of development, the highest fecundity and the greatest longevity. Castor bean pollen alone and maize pollen alone produced a higher fecundity and greater longevity than M. tanajoa tested alone. A colony of E. fustis reared continuously for seven generations on castor bean pollen produced nine times more adult females than a colony of E. fustis reared continuously on M. tanajoa. No negative effects on the development and fecundity of E. fustis were observed after seven generations were reared on pollen. On cassava, the leaf-dwelling Typhlodromalus manihoti and E. fustis occur on the middle leaves, whereas the apex-inhabiting T. aripo migrates from the apex to the top leaves only during the night (Magalhaes et al. 2002). We found that differential distributions of these predators allow prey to escape predation by vertical migration to other plant strata. We studied the role of odours in the underlying prey behaviour on predator-free plants placed downwind from plants with predators and prey or with prey only. Prey showed increased vertical migration in response to predator-related odours. Moreover, these responses were specific: when exposed to odours associated with T. manihoti, prey migrated upwards, irrespective of the plant stratum where they were placed. Odours associated with T. aripo triggered a flexible response: prey on the top leaves migrated downwards, whereas prey on the middle leaves migrated upwards. Odours associated with E. fustis, a low-risk predator, did not elicit vertical migration. Further experiments revealed that: (1) prey migrate up or down depending on the stratum where they are located, and (2) prey discrimination among predators is based upon the perception of predator species-specific body odours. Thus, at the scale of a single plant, odour-based enemy specification allows herbivorous mites to escape predation by vertical migration. This is the first record of this species from the Ivory Coast.

Specimens examined — a single specimen (1 ♀) collected between 05/IV/2017 and 17/IV/2018 14/III/2017 and 10/III/2018 in Abidjan, Anyama Ahoue (aasl 42 m, Lat 5°26′00.87″N, Long 3°55′00.60″W) and 107 ♀♀ and 39 ♂♂ collected between 06/IV/2017 au 18/III/2018 in Yamoussoukro, Ngattakro (aasl 158 m, Lat 6°49′39.443″N; Long 5°17′21.635″W) on Carica papaya L.

World distribution — Benin, Burundi, Cape Verde, Congo, DR Congo, Ghana, Malawi, Nigeria, Rwanda, Tanzania, Uganda, Zimbabwe.

Remarks — The measurements of specimens from the Ivory Coast (tables 13 and 14) are very close to those of specimens from other regions of Africa.