Döker, Ismail ¹ ; Atchia, Isabelle ² ; Jose, Anna ³ and Bolton, Samuel J. ⁴

¹✉ Cukurova University, Agricultural Faculty, Department of Plant Protection, Acarology Laboratory, 01330, Adana, Türkiye.
²Florida Department of Agriculture and Consumer Services, 32608, Gainesville, FL, USA.
³Department of Agricultural Entomology, University of Agricultural Sciences, Bangalore, India.
⁴Florida Department of Agriculture and Consumer Services, 32608, Gainesville, FL, USA.

2024 - Volume: 64 Issue: 1 pages: 192-201

ZooBank LSID: 1911F24F-A40B-414D-8C96-BB4F16B0D4E7

Original research

Keywords

Abstract

Introduction

The phytoseiids (Acari: Phytoseiidae) constitute a group of predatory mites of the superorder Parasitiformes, most widely studied due to their potential as biological control agents (McMurtry et al. 2013). Their accurate identification is considered of utmost importance for achieving effective biological control of target pests (Gerson 2014).

The subgenus Anthoseius De Leon of the genus Typhlodromus Scheuten is the largest group of predators in the subfamily Typhlodrominae, with about 400 nominal species (Ueckermann et al. 2008; Demite et al. 2014; Ferragut and Baumann 2019; Kreiter et al. 2021). Studies on the taxonomy of the Asian species of the subgenus, including India and Pakistan, date back to early 1960s, with a series of new species being described by various authors (Chant 1960; Chaudhri 1965; Gupta 1970). Yet, most of these descriptions did not include crucial taxonomical characters currently used for species delimitation, such as dorsal setal lengths, dorsal solenostomes, shape of spermatheca, cheliceral dentition, and leg chaetotaxy. The inadequate descriptions and the absence of redescriptions of Typhlodromus (Anthoseius) species have made accurate species identifications a challenge and caused confusion among taxonomists.

Among species of Typhlodromus (Anthoseius), Muma (1967) described both T. (A.) ghanii (Muma) and T. (A.) loralaiana (Muma) based on mites collected from Pakistan. Those descriptions were not very detailed, being based on simple drawings and an absence of diagnostic characters mentioned earlier. We suspected that most likely due to its inadequate description, T. (A.) ghanii was not properly recognized by acarologists and it was therefore described as a new species under different names. Indeed, based on literature information, we were unable to find reliable morphological differences between this and several other species described from India and Pakistan, namely T. (A.) homalii Gupta 1970, T. (A.) plebeius (Chaudhri et al. 1974), T. (A.) incasus (Chaudhri, 1975), and T. (A.) sagaricus Karmakar et al. (in Molla et al. 2021).

Therefore, in this study, we tried to access type specimens of the aforementioned species to confirm our hypothesis, with success only for T. (A.) homalii and T. (A.) sagaricus. The holotype of T. (A.) kuznetsovi (Denmark & Welbourn, 2002) was also examined due to its great morphological similarity with T. (A.) loralaiana (Khaustov et al. 2022). Hence, complementary descriptions of T. (A.) ghanii and T. (A.) loralaiana are provided, based on their holotypes, to improve their diagnosis and facilitate species identification.

Material and methods

The examination of mite specimens was undertaken with a Zeiss Axio Imager.M2 compound microscope equipped with differential interference contrast optical systems and a AxioCam 820 mono camera, as well as with a Zeiss Axioscope 5 compound microscope equipped with a phase contrast optic system and a Canon EOS 90D camera. Most images were captured in stacks and combined using Helicon Focus 7.6.4 Pro (Helicon Soft Ltd., 2000). The drawings were prepared using Adobe Photoshop CS2 software. Measurements were taken via ZEN 2012 software (version 8.0) and given in micrometers. The taxonomic system follows that of Chant and McMurtry (2007). Dorsal setal nomenclature is based on Lindquist and Evans (1965), as adapted by Rowell et al. (1978); ventral setal nomenclature is based on Chant and Yoshida-Shaul (1991). Nomenclature of dorsal solenostomes (gland pores) is based on Athias-Henriot (1975). Leg chaetotaxy follows that of Evans (1963).

The holotypes examined in this study are deposited at: T. (A.) ghanii and T. (A.) loralaiana, Florida Department of Agriculture and Consumer Services 1911 SW 34^th St, Gainesville, FL 32608, USA; T. (A.) homalii and T. (A.) sagaricus, National Zoological Collections, Prani Vigyan Bhawan, Zoological Survey of India, New Alipore, Kolkata, India; and T. (A.) kuznetsovi, Tyumen State University, Museum of Zoology, Tyumen, Russia.

Results

Typhlodromus (Anthoseius) ghanii Muma

(Figures 1–2)

Amblydromella ghanii Muma, 1967: 279.

Typhlodromus homalii Gupta, 1970: 188. New synonym.

Amblydromella plebeius Chaudhri, Akbar & Rasool, 1974: 209. Suspected junior synonym.

Amblydromella incasus Chaudhri, 1975: 206. Suspected junior synonym.

Typhlodromus (Anthoseius) sagaricus Karmakar et al., in Molla et al. 2021: 550. New synonym.

Material examined

Holotype female, Amblydromella ghanii, on Punica granatum L. (Punicaceae), 1 November 1965, Multan, Pakistan, coll. M.A. Ghani.

Holotype female, Typhlodromus homalii, on Homalium tomentosum (Vent.) Benth. (Salicaceae), February 1967, Botanical Garden, Howrah, West Bengal, India, coll. S.K. Gupta.

Holotype female, Typhlodromus (Anthoseius) sagaricus, on Phyllanthus emblica L. (Phyllanthaceae), 15 November 2019, Sagar Island, India, coll. unknown.

Diagnosis

Idiosomal setal pattern 12A:8A/JV:ZV (r3 and R1 off shield). Dorsal shield strongly reticulate with strong waist at level of seta R1; with five pairs of solenostomes (gd2, gd4, gd6, gd8 and gd9). Dorsal setae smooth and acuminate, except Z4 and Z5, serrate, and Z5, with large apical knob. Peritremes extending to base of j1. Sternal area striated anteriorly. Ventrianal shield pentagonal, smooth, with four pairs of preanal setae and with crescentic preanal solenostomes. Seta JV5 smooth with large apical knob. Spermatheca with bell-shaped calyx and nodular atrium without neck. Fixed digit of chelicera with five teeth and movable digit with three teeth. Genu II with seven setae (2 2/0 2/0 1). Leg IV with one knobbed macroseta on tarsus; other legs without macrosetae.

Complementary description

Holotype Female — (The specimen described by Muma 1967).

Dorsal idiosoma – (Figure 1A). Dorsal setal pattern 12A:8A (r3 and R1 off shield). Dorsal shield much longer than wide, entire, strongly reticulated with strong waist at level of seta R1; with five pairs of solenostomes (gd2, gd4, gd6, gd8 and gd9), and 13 pairs of visible poroids (id2, id4, id5, id6, idm2, idm3, idm4, idm6, idx, is1, idl1, idl3, idl4). Muscle-marks (sigillae) visible mostly on podosoma; length of dorsal shield 335, width at level of s4 184, width at level of S2 207. Dorsal setae smooth and acuminate, except Z4 and Z5, serrate, and Z5, with large apical knob. Measurements of dorsal setae as follows: j1 19, j3 23, j4 18, j5 19 j6 21, J2 24, J5 11, z2 21, z3 21, z4 23, z5 20, Z4 33, Z5 46, s4 25, s6 27, S2 28, S4 30, S5 20, r3 23 and R1 22. Peritremes extending to base of j1.

Ventral idiosoma – (Figure 1B). Ventral setal pattern 15:JV:ZV. Sternal shield lightly sclerotized, sternal area striated anteriorly; with four pairs of setae (ST1, ST2, ST3 and ST4), two pairs of poroids (iv1 and iv2), metasternal platelet and poroid iv3 not visible; distance between ST1–ST3 66, ST2–ST2 58. Genital shield smooth; one pair of para-genital poroids iv5 on soft cuticle; width at level of setae ST5 62. Ventrianal shield pentagonal, smooth; with four pairs of pre-anal setae (JV1, JV2, JV3 and ZV2), one pair of paranal (Pa) and one post-anal seta (Pst); with one pair of crescentic solenostomes located posteromesad JV2; distance between gv3 pores 20. Length of ventrianal shield 110, width at level of setae ZV2 79, width at level of paranal setae 79. Setae ZV1, ZV3, JV4 and JV5 and five pairs of poroids (ivo, ivo, ivo, ivo, ivp) on integument surrounding ventrianal shield. Seta JV5 smooth, with large apical knob, 35 in length.

Chelicera – (Figure 1C). Fixed digit 22 long with five teeth and pilus dentilis; movable digit 23 long with three teeth.

Spermatheca – (Figure 1D). Calyx bell-shaped, flaring slightly distally, 12 long; atrium nodular incorporated with calyx without neck.

Legs – (Figures 2A–E). Length of legs (excluding pretarsus): I, 284; II, 244; III, 237; IV, 358. Chaetotactic formulae as follows; 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 1/1 2/1 1. Leg III: coxa 0 0/1 0/1 0, trochanter 1 1/1 0/2 0, femur 1 2/1 1/0 1, genu 1 2/1 2/0 1, tibia 1 1/1 2/1 1. Leg IV: coxa 0 0/1 0/0 0, trochanter 1 1/1 0/2 0, femur 1 2/1 1/0 1, genu 1 2/1 2/0 1, tibia 1 1/1 2/0 1. Leg IV with one macroseta knobbed apically, StIV (pd3) 22 in length. Other legs without macroseta.

Remarks

Typhlodromus (A.) ghanii is closely related to four other species namely T. (A.) ndibu Pritchard & Baker 1962, T. (A.) fleschneri Chant 1960, T. (A.) zafari Chaudhri 1965, and T. (A.) yasumatsui Ehara 1966. However, it can be easily separated from these four species by having only one bulbous macroseta on leg IV. In contrast, all other species have three bulbous macrosetae on this leg, and T. (A.) ndibu also has bulbous setae on leg III in addition to leg IV.

According to the original description of T. (A.) homalii, the cheliceral movable digit is toothless. However, our examination of the holotype showed the presence of three teeth on this digit. After confirming a complete similarity to all other morphological and morphometric characters, we conclude that T. (A.) homalii is a new junior synonym of T. (A.) ghanii.

In the original description of T. (A.) sagaricus, gd5 is mentioned to be present and gd4 absent. So far, the presence of gd5 in Typhlodromus (Anthoseius) has only been reported in recent descriptions of Indian mites (Molla et al. 2021; Kar and Karmakar 2022). Moreover, the location of gd5 shown in the illustration of the original description of T. (A.) sagaricus, in transverse alignment with z5, is unusual, contrasting with its usual placement, about in line with z5 and j6, as shown in phytoseiids by Athias-Henriot (1975). Our examination of the holotype of T. (A.) sagaricus confirmed the presence of gd4, but we could not distinguish gd5. In addition, we also confirmed a complete similarity between T. (A.) ghanii and T. (A.) sagaricus, in relation to other morphometric and morphological characters, including leg chaetotaxy. Therefore, we here proposed the latter as a new junior synonym of the former.

As we were unable to examine type specimens of T. (A.) plebeius and T. (A.) incasus, we considered them as suspected junior synonyms of T. (A.) ghanii. Because the type materials maybe related to some other species mentioned earlier that differ in number of bulbous setae on legs. In addition, the original descriptions of both T. (A.) plebeius and T. (A.) incasus, are not sufficiently detailed to conclude their synonymy. Therefore, further examination on the type series of these two species may confirm our hypothesis.

Typhlodromus (Anthoseius) loralaiana (Muma)

(Figures 3–4)

Amblydromella loralaiana Muma, 1967: 278.

Amblydromella (Aphanoseia) kuznetsovi Denmark & Welbourn, 2002: 297. replacement name for Anthoseius richteri Kuznetsov, 1984: 385.

Typhlodromus (Anthoseius) kuznetsovi (Denmark & Welbourn, 2002), Khaustov et al. 2022: 226. New synonym.

Amblydromella dalfardica Daneshvar, 1987: 23. Suspected junior synonym.

Typhlodromus (Anthoseius) dalfardicus (Daneshvar), Kazemi et al. 2022: 724.

Material examined

Holotype female, Amblydromella loralaiana, on Punica granatum L. (Punicaceae), 27 September 1964, Loralai, Pakistan, coll. M.A. Ghani.

Holotype female, Anthoseius richteri, on Cydonia oblonga Mill. (Rosaceae), 2 September 1976, Botanical Garden, Dushanbe, Tajikistan, coll. N. N. Kuznetsov.

Diagnosis

Idiosomal setal pattern 12A:8A/JV:ZV (r3 and R1 off shield). Dorsal shield reticulate, except for smooth marginal area posteriad s6 and central between setae S4; with strong waist at level of seta R1; with five pairs of solenostomes (gd2, gd4, gd6, gd8 and gd9). Dorsal setae smooth and acuminate, except Z4 and Z5, serrate. Setae S5 short, subequal to J5 in length, and at least four times shorter than other setae in s-S series. Peritremes extending almost to base of j1. Sternal shield smooth, lightly sclerotized, with two pairs of setae. Ventrianal shield vase-shaped, much longer than wide, smooth, with four pairs of preanal setae and with rounded preanal solenostomes. Seta JV5 smooth and acuminate. Spermatheca with broad tubular calyx, flaring slightly next to vesicle, and large atrium without neck. Fixed digit of chelicera with four apical teeth and movable digit with three teeth. Genu II with eight setae (2 2/1 2/0 1). Leg IV with two macrosetae on tarsus; other legs without macroseta.

Complementary description

Holotype Female — (The specimen described by Muma 1967).

Dorsal idiosoma – (Figure 3A). Dorsal setal pattern 12A:8A (r3 and R1 off shield). Dorsal shield much longer than wide, entire, reticulate, except for smooth marginal area posteriad s6 and central between setae S4; with strong waist at level of seta R1; with five pairs of solenostomes (gd2, gd4, gd6, gd8 and gd9). Muscle-marks (sigillae) visible mostly on podosoma; length of dorsal shield 323, width at level of s4 172, width at level of S2 196. Dorsal setae smooth and acuminate, except Z4 and Z5, serrate. Measurements of dorsal setae as follows: j1 23, j3 29, j4 23, j5 23, j6 31, J2 38, J5 9, z2 20, z3 30, z4 33, z5 23, Z4 56, Z5 57, s4 37, s6 44, S2 49, S4 53, S5 8, r3 27 and R1 31. Peritremes extending almost to base of j1.

Ventral idiosoma – (Figure 3B). Ventral setal pattern 15:JV:ZV. Sternal shield smooth, lightly sclerotized, posterior margin not visible, smooth with two pairs of setae (ST1 and ST2) and two pairs of poroids (iv1 and iv2); seta ST3 on soft integument, seta ST4 and poroid iv3 on metasternal platelets; distance between ST1–iv2 55, ST2–ST2 52. Genital shield smooth; one pair of para-genital poroids iv5 on soft cuticle; width at level of setae ST5 60. Ventrianal shield elongated, vase-shaped, smooth; with four pairs of pre-anal setae (JV1, JV2, JV3 and ZV2); one pair of paranal (Pa) and a post-anal seta (Pst); with one pair of rounded solenostomes located posteromesad of JV2; distance between gv3 pores 17. Length of ventrianal shield 114, width at level of setae ZV2 59, width at level of paranal setae 61. Setae ZV1, ZV3, JV4 and JV5 and five pairs of poroids (ivo, ivo, ivo, ivo, ivp) on integument surrounding ventrianal shield. Seta JV5 smooth, acuminate, 44 in length.

Chelicera – (Figure 3C). Fixed digit 23 long with four apical teeth and pilus dentilis; movable digit 23 long with three teeth. The unusual direction of pilus dentilis in the illustrated chelicera is probably because of mounting as its position is normal in the other chelicera.

Spermatheca – (Figure 3D). Calyx broad, elongate, tubular, flaring slightly next to vesicle, 28 long; atrium large and incorporated into base of calyx without neck.

Legs – (Figures 4A–D). Length of legs (excluding pretarsus): I, 282; II, 235; III, 218; IV, 300. Chaetotactic formulae as follows; 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 2/1 2/0 1, tibia 1 1/1 2/1 1. Leg III: coxa 0 0/1 0/1 0, trochanter 1 1/1 0/2 0, femur 1 2/1 1/0 1, genu 1 2/1 2/0 1, tibia 1 1/1 2/1 1. Leg IV: coxa 0 0/1 0/0 0, trochanter 1 1/1 0/2 0, femur 1 2/1 1/0 1, genu 1 2/1 2/0 1, tibia 1 1/1 2/0 1. Leg IV with two acuminate macrosetae, StIV (pd3) 36 in length, and that on telotarsus 32 in length. Other legs without macroseta.

Remarks

We herein confirm that Typhlodromus (A.) kuznetsovi (Denmark and Welbourn), a species previously described and known only from Tajikistan, is a junior synonym of T. (A.) loralaiana, based on the examination of their holotypes. A full redescription of T. (A.) kuznetsovi was recently provided by Khaustov et al. (2022). In addition, T. (A.) dalfardicus, described by Daneshvar (1987) based on specimens collected from Citrus sp. (Rutaceae) in Iran, shows affinity to T. (A.) loralaiana. Daneshvar (1987) separated these two species based on setal lengths and cheliceral movable digit dentition. We were unable to find reliable morphometric differences between the holotype T. (A.) loralaiana and the original description of T. (A.) dalfardicus. In addition, Daneshvar (1987) implied that T. (A.) dalfardicus has two teeth on the movable digit of the chelicera as compared with no teeth in T. (A.) loralaiana. Our examination of the holotype of T. (A.) loralaiana shows that cheliceral digits are almost totally closed, but it was still possible to observe a movable digit and count its denticles (three teeth) by using a plan-apochromatic oil lens (1.4 NA) with a DIC microscope. This finding is still different from what was reported in the original description of T. (A.) dalfardicus. In conclusion, until further examination of the type specimens of the latter or until new specimens are examined, we prefer considering T. (A.) dalfardicus a suspected junior synonym of T. (A.) loralaiana.

Acknowledgements

We are grateful to Dr. Shelley Acharya for help in examination of the type materials of Anthoseius species deposited in National Zoological collections, Prani Vigyan Bhawan, New Alipore, Kolkata, India. We are also grateful to Vladimir A. Khaustov (Tyumen State University, Russia) for examining the holotype T. (A.) kuznetsovi. Study of Ismail Döker was supported by Cukurova University Scientific Projects Foundation Units, grant number, FAY-2022-14495.

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instructions