Tsolakis, Haralabos ¹ and Ranja-William, François Antoine ²

¹✉ University of Palermo, Department of Agricultural Food and Forest Sciences, Laboratory of Applied Acarology "Eliahu Swirski", Edifice 5A, Viale delle Scienze, 13 – 90128 Palermo, Italy.
²University of Palermo, Department of Agricultural Food and Forest Sciences, Laboratory of Applied Acarology "Eliahu Swirski", Edifice 5A, Viale delle Scienze, 13 – 90128 Palermo, Italy.

2025 - Volume: 65 Issue: 2 pages: 448-460

ZooBank LSID: F6B43CD1-A02E-49A5-8505-97A841408B57

Original research

Keywords

Abstract

Introduction

Predatory mites of the family Phytoseiidae (Acari, Parasitiformes) are considered important biocontrol agents in agricultural and natural ecosystems (McMurtry et al. 2013, 2015; Heyler et al. 2014; Lorenzon et al. 2018). The family includes at present more than 2,500 valid species (Demite et al. 2024) and many new species are added in the family every year (Kreiter et al. 2021; Döker et al. 2021; Arjmandi-Nezhad et al. 2022; Biswas and Karmakar 2024). Some geographic areas, i.e. Europe or single countries, i.e. India, China, Japan, U.S.A., Canada, Brazil, Israel, Egypt, South Africa, Australia, were widely investigated during the last decades because of specialists' presence and research groups created for that purpose. In other parts of the world, information on the phytoseiid fauna is limited and fragmented. However, investigation for discovering indigenous phytoseiid species, in inadequately investigated areas is useful to enrich our knowledge on the third trophic level in natural and agricultural ecosystems worldwide and to discover new biocontrol agents able to control infestations of mite pests (Kreiter et al. 2018; Novljan et al. 2023; Abo-Shnaf and Moraes 2014).

Madagascar is the fourth largest island of the world located at about 400 km of the southeastern coast of Africa. It is identified as one of the most important biotopes on Earth and it is one of the 25 ''biodiversity hotspots″, i.e biogeographic areas with a high level of species endemism suffering a remarkable habitat loss (Myers et al. 2000; Vences et al. 2009). Several studies in recent decades have dealt with the specific composition of arthropods in Madagascar, particularly insects (Biondi 2001; Huber 2003; Fisher 2005; Wesener et al. 2011; Biondi and D'Alessandro 2013, 2016), but studies on Phytoseiidae dates back to the 1980s. Currently, 41 species are known from Madagascar, eight of which endemic (Demite et al. 2024). About 76% of records are attributed to Blommers (1973, 1974, 1976) and the remaining to Chazeau (1970), Blommers and Chazeau (1974), Blommers and Gutierrez (1975) and Schicha (1983, 1987) (Demite et al. 2024).

The objective of this paper is to present the results of a new survey for phytoseiid mites in Madagascar, with the descriptions of new species.

Material and methods

Mites were collected in October and November 2023 in the central and northern areas of Madagascar on cultivated and spontaneous arboreal plants (Table 1).

Phytoseiid mites were collected using the branch-shaking method (Tsolakis and Ragusa 1999). Mites were collected using a micro–aspirator, preserved in alcohol 70%, cleared in Nesbitt solution and mounted in Hoyer's medium. Identification and measurements were done using a differential interference contrast microscope (DIC) Zeiss Axioplan. Illustrations and measurements were made using the Axiovision 40V 4.6.1.0 application (Zeiss, 2002–2007); measurements are given in micrometres (μm), reporting mean (in bold), minimum and maximum (in brackets). Drawings were hand–made using the Zeiss Axioplan Camera lucida and defined in vectorial format by Affinity Designer 1.10.8. In the description of the new species, we followed the setal nomenclature proposed by Lindquist and Evans (1965), as adapted to the Phytoseiidae by Rowell et al. (1978). For dorsal and ventral patterns, we followed Chant and Yoshida-Shaul (1989, 1991). For the terminology of various sections of the insemination apparatus (spermathecal apparatus), we followed Wainstein (1973), Athias-Henriot (1975; 1977) and Beard (2001), with some additions and changes. For setal patterns of legs podomeres (genu, tibia, tarsus), the formulae proposed by Evans and Till (1979) were adopted. For the macrosetae, we considered the concept defined by Beard (2001). Nomenclature of the adenotaxy was that suggested by Athias-Henriot (1975). Dorsal shield length was measured along the midline from j1 setae level to the end of the shield. The apical tooth of each cheliceral digit was not included in the determination of the number of teeth. For descriptive terms of morphological characters, see Tsolakis and Ragusa (2020). The world distribution of species found during our surveys is based on Demite et al. (2024).

Type materials and specimens of all identified species are kept in the Acari collection of the laboratory of Agricultural and Applied Acarology ''Eliahu Swirski″, Department of Agricultural, Food and Forestry Sciences, University of Palermo (Italy).

Results

During surveys carried out on 10 species of arboreal plants, 228 specimens of phytoseiid mites (217 females and 11 males) were collected and identified (Table 1). Nine species belonging to seven genera within the sub-families Amblyseiinae and Typhlodrominae have been identified; two of them are new records for Madagascar and other two are new for science and described herein.

Amblyseiinae Muma 1961

Euseiini Chant and McMurtry 2005

Subtribe Typhlodromalina

Typhlodromalus Muma, 1961

Typhlodromalus andramasinus sp. nov.

ZOOBANK: 4193DE30-AEEE-4A10-B79D-9AD6AD03D3DF

Diagnosis of female

(Figure 1)

Peritrematal shield is fused with the dorsal one at level of setae j1. A caudal lobe after the level of seta Z5, which is a characteristic of many species of this genus is present. Setae J5 are present on this lobe, below of level of setae Z5. Dorsal shield slightly reticulate, all dorsal setae slender and smooth, except Z4 and Z5 which are slightly serrated. Adenotaxy complete (holoadenic), all solenostomes small and crateriform. Solenostomes gd3 and poroids id3 well visible on peritrematal shield. Peritreme reaching the base of j1. Sternal shield smooth with three pairs of setae (ST1–ST3). Setae ST4 and poroids iv3 tylochorous (on metasternal platelets). Genital shield flask shaped with posterior margin almost transversally straight; Ventrianal shield smooth with three pairs of preanal setae and a pair of large crateriform solenostomes (gv3). Setae JV5 smooth. Calyx of the spermathecal apparatus vase shaped. Atrium differentiated, minor duct well visible. Major duct short. Fixed digit of chelicera with nine teeth, movable digit with three teeth. Seven setae on genu II, with three smooth and pointed macrosetae on leg IV, of which the shortest on the tibia and the longest on the basitarsus.

Description of female

Dorsum — (Fig. 1A) (Eight females measured)

Dorsal setal pattern 10A:9B. Seventeen pairs of setae on dorsal shield, two pairs of lateral setae (r3, R1) on the interscutal membrane. Dorsal shield 349 (334–363) long, 209 (200–219) wide at level of s4, 195 (184–207) at level of seta Z1, 197 (188–207) at level of seta S2 and 193 (175–202) at level of seta S4. Measurements of setae: j1 27 (23–31); j3 33 (28–35); j4 18 (14–21); j5 17 (14–20); j6 26 (21–29); J2 25 (17–28); J5 9 (9–13); z2 25 (17–28); z4 33 (28–36); z5 21 (18–24); s4 45 (41–47); Z1 27 (23–30); Z4 47 (42–52); Z5 67 (60–72); S2 36 (31–40); S4 21 (17–25); S5 15 (11–19); r3 23 (20–25); R1 21 (19–26). Seven pairs of solenostomes are present on dorsal shield: gd1 posteroantiaxial to j3, gd2 posteroantiaxial to j4, gd4 posteroantiaxial to s4, gd5 posteroparaxial to z5, gd6 anteroparaxial to Z1, gd8 anteroantiaxial to Z4, gd9 paraxial to S5.

Peritreme — (Fig. 1A). The apex of peritreme reaching the basis of j1. Length of peritreme 213 (201–222). Solenostome gd3 and poroid id3 on peritrematal shield, posteriad to r3. Solenostome gdp crateriform.

Ventral idiosoma — (Fig. 1B). Sternal shield smooth; anterior arms well visible. Poroids iv1 and iv2 well visible on the shield. Two small projections of the shield visible at level of poroids iv2. Posterior margin of the shield with a median lobe, whose tip is not discernible. Distance between bases of setae ST1–ST1 62 (59–65), ST2–ST2 68 (64–70), ST3–ST3 80 (76–84), ST1–ST3 68 (64–72). Distance between the genital seta ST5 73 (68–78). First to third pairs genital sigilla (gs) well visible. Fourth and fifth pairs of sigilla well visible on the interscutal membrane between genital shield and setae ZV1; sixth pair of sigilla posteroparaxial to ZV1. Ventrianal shield narrow, flask–shaped, with a pronounced constriction posteriad gv3, 112 (103–120) long, 59 (53–62) at level of ZV2, 56 (51–60) at level of JV2, 63 (58–67) at level of paranal setae. Distance between solenostomes gv3 24 (22–26). Setae JV1 insert on anterior margin of ventrianal shield. Seta JV5 smooth, 25 (20–29) long. Inguinal sigilla (metapodal platelets) 19 (18–19) and 7 (7–8) long.

Spermathecal apparatus — (Fig. 1C)

The proximal part of the major duct differentiated (Receptaculum); 13 (12–14) long, cylindrical and simple (2 wide). Atrium almost bulbous, slightly differentiated from the major duct (3 long), inserted at the basis of calyx; accessus absent. Minor duct well visible in most specimens. Calyx 18 (15–21) long, enlarged near atrium, narrow afterwards, forming a funnel towards vesicle 10 (8–12 wide).

Chelicerae — (Fig. 1 D). Fixed digit 34 (32–36) long with 9 teeth. Pilus dentilis visible in some specimens. Movable digit 33 (30–38) long, with three teeth.

Legs (Fig. 1E). Macrosetae sensu Beard (2001) present on all legs: Sge I 18 (16–19), Sge II 19 (18–19), Sge III 24 (23–24), Sge IV 38 (36–42), Sti IV 26 (24–29) and St IV 62 (57–65). All macrosetae pointed except Sge IV which is slightly knobbed. Chaetotaxy of legs: Leg I, coxa 0 0/1 0/1 0, trochanter 1 0/1 0/2 1, femur 2 3/2 2/2 2, genu 2 2/1 2/1 2, tibia 2 2/1 2/1 2. Leg II, coxa 0 0/1 0/1 0, trochanter 1 0/1 0/2 1, femur 2 3/1 2/1 1, genu 2 2/0 2/0 1, tibia 1 2/1 1/1 1, tarsus 1 1/0 1/0 1. Leg III, coxa 0 0/1 0/1 0, trochanter 1 0/1 0/2 1, femur 1 2/1 1/0 1, genu 1 2/1 2/0 1, tibia 1 1/1 2/1 1, tarsus 1 1/0 1/0 1. Leg IV, coxa 0 0/1 0/0 0, trochanter 1 0/1 0/2 1, femur 1 2/0 1/0 1, genu 1 2/0 2/1 1, tibia 1 1/0 2/1 1, tarsus 1 1/1 0/0 1.

Comparative notes

Typhlodromalus andramasinus sp. nov. mostly resembles the following Typhlodromalus species: T. araucariae Gonçalves & Ferla, T. olombo (Pritchard & Baker), T. mangiferae (Chaterjee & Gupta), T. aripo De Leon, T. serengati El-Banhawy & Abou-Awad, T. spinosus (Meyer & Rodrigues) and T. sinespinosus Moraes, Zanou & Oliveira, differing as subsequently described. Typhlodromalus araucariae has shorter j3, j5, j6, z4, z5, s4, Z1, S2, S4 and Z4; respectively seven and two teeth on fixed and movable cheliceral digits; a single macroseta on leg IV, which is shorter (37μm) and knobbed; spermathecal apparatus of different shape. Typhlodromalus olombo has some setae shorter (z2, z5, Z1, R1) and others longer (j3, s4, S2); four teeth on the movable cheliceral digit; macrosetae of leg IV shorter; macroseta of basitarsus IV knobbed; spermathecal apparatus of different shape. Typhlodromalus aripo has j4, j5, j6 and J2 shorter and j3 longer; macroseta of genu and tarsus of leg IV longer (53 and 73, respectively), macroseta of tarsus IV knobbed; spermathecal apparatus of different shape. Typhlodromalus mangiferae has j3, j6, z5, Z1, S2, S5, R1 and macroseta on tarsus IV shorter. Typhlodromalus spinosus, T. sinespinosus and T. serengati differ from the new species by the length of several dorsal shield setae, the absence of S5 and the different shapes of the spermathecal apparatus (Table 2).

Specimens examined

Holotype female (serial number M-019) collected from Mangifera indica L. (Anacardiaceae) in the Ambohimasina forest in the Andramasina district, 19°12′23″S, 47°38′33″E, 1354 m a.s.l., November 4, 2023. Two paratype females (serial number M-016) collected on Psidium guajava L. (Myrtaceae), November 2, 2023, at the same locality as holotype. Four paratype females (serial number M-008 and M-009) collected on Dombeya reclinata Cordem. (Malvaceae) along the edge of the river Analasarotra (15°29′31.48″S, 47°44′0.64″E, m 19 a.s.l.), October 14, 2023. One paratype female (serial number M-042) collected on M. indica in Mahabibo forest in the Mahajanga district, (15°42′43.15″S, 46°23′34.18″E), November 19, 2023.

Etymology

The specific name andramasinus refers to the district Andramasina where the holotype and two of the paratypes were collected.

Euseiini Chant & McMurtry 2005

Subtribe Euseiina

Moraeseius Chant & McMurtry 2005

Moraeseius papayana (Van der Merwe 1965)

Amblyseius (Amblyseius) papayana — Van der Merwe (1965)

Amblyseius (Proprioseiospsis) papayana — Van der Merwe (1968)

Euseius papayana — Moraes et al. (1986, 2004)

Originally described from specimens collected on Carica papaya L in Mbombela, Mpumalanga Province, South Africa. This is the first report of M. papayana from Madagascar.

World distribution — Kenya; Mayotte Island; Mozambique; South Africa, Madagascar.

Remarks — Measurements of setae of the three females collected, correspond to those reported by van der Merwe (1965) and by Moraes et al. (2001). However, the above-mentioned authors reported an entire ventrianale shield, while in our specimens it is divided into ventral and anal shield, being the solenostome gv3 on the interscutal membrane. However, the posterior margin of ventral shield and the anterior one of the anal shield are evanescent and further collections are needed to clarify if it is an inter population variation and to define the systematic weight of this character. With a subulate seta (al) on trochanter I, as also reported for other Euseius species (Döker et al. 2024, 2025) and as observed in our examination of specimens of Iphiseius degenerans (Berlese) from Citrus x limon (L.) Osbeck collected in Sicily (Fig. 2). Basing on molecular analyses, the latter phytoseiid is now included within the genus Euseius (Döker et al. 2025), as already hypothesized by Santos and Tixier (2018). With E. degenerans, M. papayana shares also a similar spermathecal apparatus. Further investigation within these genera, could be in searching for systematic characters to evaluate their possible relationship as also suggested by Döker et al. (2025).

Specimens examined — Mahajanga, natural park of Ankarafantsika, 16°16′26″S, 46°48′39″E, 67 m a.s.l., 3 ♀♀ on Tamarindus indica L. (Fabaceae).

Typhlodrominae Wainstein 1962

Paraseiulini Wainstein 1976

Kuzinellus Wainstein 1976

Kuzinellus querellus (Ueckermann & Loots, 1988)

Typhlodromus (Anthoseius) querellus Ueckermann & Loots, 1988

Originally described from specimens collected on Hemizygia parvifolia Codd at Motlatse Canyon (Mpumalanga Province) South Africa. It was also reported from Zimbabwe (Ueckermann and Loots 1988) and afterwards from Burundi and Sierra Leone (Moraes et al. 2008) and from Kenya (Santos et al. 2021). This is the first record of K. querellus from Madagascar.

Specimens examined — Andramasina, Abohimasina, 19°12′23″S, 47°38′33″E, 1354 m a.s.l., 1 ♀ on Pinus pinea L. (Fabaceae).

Typhlodrominae Wainstein 1962

Typhlodromini Wainstein 1962

Typhlodromus Scheuten 1857

Subgenus Anthoseius De Leon 1959

Typhlodromus (Anthoseius) amphoraeformis sp. nov.

ZOOBANK: 135E1860-795A-4E3A-8EC8-FCAF68017B24

Diagnosis of female

(Figure 3)

Dorsal shield slightly reticulated. Peritreme reaching the base of seta j1. All dorsal setae are slender, smooth and sharp-tipped, except Z4 and Z5 which are slightly serrated and the latter is also spatulate. Adenotaxy incomplete (meriadenic), solenostomes gd1 and gd5 absent. Sternal shield smooth with two pairs of setae (ST1–ST2) and two pairs of poroids (iv1 and iv2). Seta ST3 gymnochorous (situated on the interscutal membrane); seta ST4 and poroids iv3 tylochorous (situated on a sclerite, often indicated as metasternal platelet). Genital shield almost smooth with the posterior margin straight; first to third pairs of sigilla well visible. Ventrianal shield smooth with four pairs of preanal setae and a pair of small crateriform solenostomes (gv3) between setae JV3. Seta JV5 smooth and spatulate. Calyx of the spermathecal apparatus distinctly constricted shortly before fusion with vesicle. Atrium nodular, accessus present; spermatic channel (minor duct) visible. Major duct cylindrical and narrow. Fixed digit of chelicera with five teeth, movable digit with three teeth. Seven setae on genu II; three smooth and spatulate macrosetae on leg IV and none on other legs; macrosetae not much longer than other setae of the respective segments but distinguished for being slightly stout and spatulate.

Description of female

Dorsum — (Fig. 3A) (Sixteen females measured)

Dorsal setal pattern 12A:8A. Eighteen pairs of setae on dorsal shield, two pairs of lateral setae on the interscutal membrane (r3, R1). Dorsal shield 315 (297–337) long, 173 (165–181) wide at level of seta s4, 165 (161–170) at level of R1, 189 (181–202) at level of S4. Measurements of setae: j1 18 (16–20); j3 20 (16–22); j4 14 (11–16); j5 15 (13–16); j6 19 (15–21); J2 21 (19–23); J5 11 (8–16); z2 17 (15–19); z3 19 (17–21); z4 20 (18–22); z5 17 (13–23); s4 22 (18–25); s6 23 (20–25); Z4 32 (27–35); Z5 46 (42–50); S2 26 (23–28); S4 24 (21–27); S5 17 (13–19); r3 15 (13–17); R1 18 (16–20). Five pairs of solenostomes on dorsal shield: gd2 posteroparaxial to z3, gd4 posteroantiaxial to s4, gd6 posteroparaxial to s6, gd8 anteroantiaxial to Z4, gd9 paraxial and very close to the basis of S5.

Peritreme — (Fig. 3A). Length of peritreme 183 (174–208). The apex reaches the basis of j1. Solenostome gd3 and poroid id3 on peritrematal shield, posteriad to r3. Solenostome gdp crateriform.

Ventral idiosoma — (Fig. 3B). Sternal shield smooth, anterior arms visible. Poroids iv1 and iv2 well visible on the shield; posterior margin almost convex. Setae ST3 on the interscutal membrane. Distance between bases of setae ST1–ST1 46 (42–48), ST2–ST2 54 (52–57), ST3–ST3 66 (63–69), ST1–ST2 32 (30–33). Distance between the genital setae ST5 61 (56–64). Two arcuated arms of the membranous cover of the genital opening are well visible. Sigilla of fourth and fifth pairs well visible on the interscutal membrane anteroparaxially to setae ZV1; sigilla of sixth pair posteroparaxial to setae ZV1. Ventrianal shield subpentagonal, 107 (101–110) long, 86 (81–92) wide at level of setae ZV2, 84 (81–87) at level of setae JV3, 76 (72–80) at level of paranal setae. Distance between solenostomes gv3 19 (18–21). Seta JV5 smooth, spatulate 31 (28–33) long. Inguinal sigilla well visible, 22 (20–25) and 7 (7–8) long.

Spermathecal apparatus — (Fig. 3C). Major duct 7 (6–8) long, cylindrical, simple (2 wide) (receptaculum not visible). Atrium almost bulbous, inserted at the basis of calyx, accessus into the major duct visible. Spermatic channel discernible in many specimens. Calyx 17 (16–18 long, it enlarged near atrium (8 wide), with a pronounced constriction just before connection with the vesicle, resembling an amphora-shaped vase.

Chelicerae — (Fig. 3 D). Fixed digit of the chelicera 22 (22–23) long with 5 teeth. Pilus dentilis well visible. Movable digit 21 (20–22) long, with three teeth plus the apical tooth the basal of which vestigial.

Legs — (Fig. 3E). Chaetotaxy of genu II 2 2/0 2/0 1. Three spatulate macrosetae sensu Beard (2001) present on leg IV: Sge 12 (11–13), Sti 14 (12–15) and St 20 (18–21). Absence of macrosetae on the other legs. Chaetotaxy of legs: Leg I, coxa 0 0/1 0/1 0, trochanter 1 0/1 1/2 1, femur 2 3/1 2/2 2, genu 2 2/1 2/1 2, tibia 2 2/1 2/1 2. Leg II, coxa 0 0/1 0/1 0, trochanter 1 0/1 0/2 1, femur 2 3/1 2/1 1, genu 2 2/0 2/0 1, tibia 1 2/1 1/1 1, tarsus 1 1/0 1/0 1. Leg III, coxa 0 0/1 0/1 0, trochanter 1 1/1 0/2 0, femur 1 2/0 1/0 1, genu 1 2/1 2/0 1, tibia 1 1/1 2/1 1, tarsus 1 1/0 1/0 1. Leg IV, coxa 0 0/1 0/0 0, trochanter 1 1/1 0/1 1, femur 1 2/ 1/0 1, genu 1 2/0 2/1 1, tibia 1 1/0 2/1 1, tarsus 1 1/0 1/0 1.

Comparative notes

Typhlodromus (Anthoseius) amphoraeformis sp. nov. resembles 12 African species belonging to the subgenus Anthoseius, eight of which described by van der Merwe (1968) and four, by Ueckermann and Loots (1988): T. (A.) terrulentis van der Merwe, T. (A.) saevus van der Merwe, T. (A.) incisivus van der Merwe, T. (A.) apoxys van der Merwe, T. (A.) capparidis van der Merwe, T. (A.) muliebris van der Merwe, T. (A.) microbullatus van der Merwe, T. (A.) februs van der Merwe, T. (A.) grastis Ueckermann & Loots, T. (A.) michaeli Ueckermann & Loots, T. (A.) umbraculus Ueckermann & Loots and T. (A.) drymis Ueckermann & Loots. The new species differs from all the abovementioned species in the unique shape of the spermathecal apparatus, the number of teeth on the movable cheliceral digit, the number and lengths of macrosetae on leg IV, as well as in the length of various setae of the dorsal shield (Table 3).

Specimens examined

Holotype female (serial number M-051) and 2 paratype females collected on Anacardium occidentale L. at Mahajanga, Mahabibo forest, 15°41′27.81″S, 46°25′34.17″E, 10 m a.s.l., November 22, 2023. Seven paratype females (serial numbers M-033, M-034, M-037, M-038, M-040 and M-041) collected on Mangifera indica on the same date and at the same locality as holotype. Six paratype females (serial number M-056 and M-057) collected on Tamarindus indica L. (Fabaceae) in Natural parks of Ankarafantsika (Mahajanga), 16°16′21.14″S, 46°48′35.71″E, 13m a.s.l., November 24, 2023. Two paratype females (serial number M-054) collected on Terminalia mantaly Perrier at Tsaramandroso (Mahajanga), 16°22′37.41″S, 47° 4′53.51″E, November 21, 2023.

Etymology

The specific name amphoraeformis is a composite word from the ancient Greek ἀμφορεύς transcribed in Latin as Amphora + the Latin word Forma = shape. It refers to the characteristic shape of the spermathecal apparatus that resembles an ancient Greek vase.

Acknowledgements

Authors are deeply indebted to the three anonymous reviewers for their precious and constructive comments. This work was supported by funding from ''Fondo di Finanziamento per la Ricerca 2023″, University of Palermo (FFRD13).

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