1✉ Department of Plant Protection, College of Food and Agriculture Sciences, King Saud University, Riyadh, Saudi Arabia.
2Department of Plant Protection, College of Food and Agriculture Sciences, King Saud University, Riyadh, Saudi Arabia.
2024 - Volume: 64 Issue: 3 pages: 925-940
https://doi.org/10.24349/hr32-1gg9Amblyseius Berlese (Phytoseiidae) is one of the largest genera in the subfamily Amblyseiinae Muma, with 434 nominal species worldwide (Demite et al. 2024). It was erected by Berlese (1914) based on the type species, Amblyseius obtusus (Koch, 1839). It is distinguished from the other genera belonging to the subtribe Amblyseiina by combination of following characters: setae s4, Z4, and Z5 greatly longer (ratio of setae s4: S2 \textgreater3:1); both cheliceral digits of normal in size, setae JV2 present; ventrianal shield well-developed, rarely divided; genu and tibia III each with one macroseta or rarely none (Chant and McMurtry 2003, 2004).
Denmark and Muma (1989) reviewed the genus Amblyseius and proposed three subgenera based on number of teeth on fixed digit of chelicerae and length of macrosetae on leg IV, subgenus Amblyseius Berlese with 16 species groups (72 species), subgenus Pauciseius Denmark & Muma with three species groups (12 species) and subgenus Multiseius Denmark & Muma with 12 species groups (51 species). Chant and McMurtry (2004, 2007) ignored this sub generic division and species groups and suggested only five species groups in Amblyseius; pusillus species group Chant & McMurtry, sundi species group Denmark & Muma, obtusus species group Chant, americanus species group Denmark & Muma, and largoensis species group McMurtry & Moraes. These species groups were based on relative length of setae z4, shape of spermathecae and venterianal shield, and presence/absence of setae Z1. Later, Denmark and Evans (2011) followed sub generic division of Amblyseius proposed by Denmark and Muma (1989) but ignored species grouping in Amblyseius.
Until now, 25 Amblyseius species have been reported from Pakistan (Chaudhri et al. 1979; Halliday et al. 2018). The species, A. bajulus Chaudhri, Akbar & Rasool 1979 was synonymized with A. obtusus (Koch, 1839) based on original descriptions and illustrations (Daneshvar and Denmark 1982; Denmark and Muma 1989). In the present study, A. excelsus Chaudhri, Akbar & Rasool, 1979, A. adhatodae Muma, 1967, and A. isuki Chant & Hansel, 1971 are re-described based on new material collected from different localities in Pakistan. In addition, keys to the world species of duncansoni and largoensis species subgroups of Amblyseius are presented.
The phytoseiid mites were collected from a diversity of host plants in different localities across Pakistan during the year 2023. The mite specimens were collected by shaking the plant foliage over a white piece of paper and preserved in 70% ethanol. Mite specimens were mounted in Hoyer's medium and identified under a phase contrast microscope (BX51, Olympus®, Japan) using diagnostic keys and available literatures. The taxonomic literature of different species groups of the genus Amblyseius was collected to develop world species diagnostic keys of duncansoni and largoensis, species subgroup. Different mite body parts were imaged using an Auto-montage software system (Syncroscopy, Cambridge, UK) attached with phase contrast microscope (DM2500, Leica®, Germany). The illustrations were drawn with Adobe Illustrator (Adobe System Inc., San Jose, CA, USA). All measurements are given in micrometers (μm). The morphological terminology used in this study follows that of Lindquist and Evans (1965), Rowell et al. (1978), and Chant and Yoshida-Shaul (1992). All the collected specimens have been deposited in the Museum of Arthropods (KSMA, Acarology section), Department of Plant Protection, College of Food and Agriculture Sciences, King Saud University, Riyadh, Saudi Arabia.
Amblyseius Berlese, 1914: 143; Chant and McMurtry, 2004: 188
Amblyseius excelsus Chaudhri, Akbar and Rasool, 1979: 68.
Two females, one male, grasses (Poaceae), Kaghan, (34.7840417° N, 73.5207167° E), 19 July 2023, three females, one male, grasses (Poaceae), Mansehra, Khyber Pukhtun Khawa (34.4062372° N, 73.3375557° E), 20 July 2023; two females, one male, grasses (Poaceae), Kuttan, Neelam valley, Azad Jammu and Kashmir (AJK) (34.5593598° N, 73.7921257° E), 18 July 2023; two females, two males, grasses (Poaceae) Shahkot, Neelam valley, AJK (34.5671687° N, 73.8912957° E), 18 July 2023; three females, grasses (Poaceae), Kiran, Neelam Valley, AJK (34.6318277° N, 73.9191069° E), 18 July 2023; all collected by M. Kamran.
Balakot, Abbotabad, Muzaffarabad (Pakistan) (Chaudhri et al. 1979); Jammu Kashmir, India (Rather 1985).
Dorsum –(Figure 1A, B). Dorsal shield mostly smooth with few striations anterolaterally from setae j3 to R1, 330–349 long and 215–231 wide at the level of s4; with seven pairs of solenostomes (gd1, gd2, gd4, gd5, gd6, gd8, gd9) and 16 pairs of poroids (id1, id2, id4, id5, id6, idm1, idm2, idm3, idm4, idm5, idm6, isl, idl1, idl3, idl4, idx). Setae length: j1 25–28, j3 45–56, j4 5–7, j5 6–7, j6 7–8, J2 7–8, J5 7–8, z2 19–22, z4 21–26, z5 4–5, Z1 8–9, Z4 82–90, Z5 155–175, s4 75–84, S2 12–13, S4 8–9, S5 8 7–8, r3 21–23, R1 11–13. All setae smooth, except for Z4 and Z5, finely serrate near tips. Peritreme extending to the level of j1.
Venter — (Figure 2A). Sternal shield smooth, posteriorly almost flat, with three pairs of setae (st1-st3) and two pairs of poroids (iv1, iv2); distance st1-st3 65–70, st2-st2 70–78, st4 and poroid iv3 tylochorous; genital shield smooth, distance st5–st5 70–75; a transversely elongate platelet between genital and ventrianal shields. Ventrianal shield subpentagonal, slightly constricted at the level of solenostomes, with sparsely striae, about 6–7 transverse lines, 125–135 long, 90–98 wide at ZV2 level, and 85–89 wide at anus level; with three pairs of pre-anal setae (JV1, JV2, and ZV2) along with para-anal and post-anal setae, a pair of solenostomes (gv3), crescentic shape, posteromesad JV2; four pairs of setae (JV4, JV5, ZV1, and ZV3), four pairs of poroids (ivo, ivp), and two pairs of metapodal platelets on the membrane surrounding ventrianal shield, primary metapodal plate 22–24, secondary 9–10 long. Setae ZV1 23–25, ZV3 = JV4 12–13, JV5 70–75, smooth.
Chelicera — (Figure 2B). Movable cheliceral digit 29–32 long, with two teeth; fixed digit 31–35 long, with nine teeth and a pilus dentilis.
Spermatheca — (Figure 2C). Calyx bell shaped, 15–18 long and 12–13 wide at apex, atrium visible, minute not bulbous, major duct narrow, tube-like.
Legs — (Figure 2D). Genu II with eight setae; genua III-IV each with seven setae; genua I-III each with a macroseta: 28–31, 28–30, and 35–39 long, respectively; tibiae IIII without macrosetae; leg IV with three macrosetae: SgeIV 68–77, StiIV 55–65, StIV 76–80, all macrosetae smooth with pointed tip (Figure 2D).
Dorsum — (Figure 3A). Dorsal shield smooth, with few striae laterally, eight pairs of solenostomes, gd3 present, poroids indistinct, 260–265 long and 190–195 wide. Setae length: j1 22–23, j3 45–48, j4 6–7, j5 6–7, j6 = J2 6, J5 6–7, z2 17–18, z4 19–20, z5 5–6, Z1 12, Z4 64–66, Z5 115–120, s4 62–65, S2 10–11, S4 10, S5 9, r3 18–19, R1 11–12. All setae smooth, except for Z4 and Z5, finely serrated near tips. Peritreme extending to the level behind j1.
Venter — (Figure 3B). Sternogenital shield almost smooth with lateral striae, st-st5 113–118, st2-st2 60–63. Ventrianal shield entirely with transverse lines/striae, wider than long, 114–118 long and 150–155 wide at the anterior corners level; with three pairs of pre-anal setae (JV1, JV2, and ZV3), a pair of solenostomes (gv3) posteromesad JV2, almost oval in shape, four pairs of poroids (ivo); JV5 43–45 smooth, a pair of poroids (ivp) mesad JV5.
Chelicerae — (Figure 4A). Spermatodactyl T-shaped with slightly developed toe, but the axes of shaft and toe make almost right angle, shaft 18–19 long, toe 4–5 long. Movable cheliceral digit 17–19 long, fixed digit 19–21 long.
Legs — (Figure 4B). Three macrosetae on leg IV: SgeIV 48–50, StiIV 43–45, StIV 70–72. One macroseta each on genu of legs I–III: 23–25, 21–24, and 24–28 long, respectively. All macrosetae smooth with pointed tips.
Amblyseius excelsus belong to the duncansoni species subgroup Chant & McMurtry, 2004 of americanus species group Denmark & Muma, 1989 (Chant and McMurtry 2004). This species subgroup includes 14 species worldwide that having spermathecal calyx cup-dish, or bell-shaped. Morphological characters and measurements of the specimens of A. excelsus collected in the present study, are similar to its original description (Chaudhri et al. 1979), except for small differences in length of some dorsal setae (z4 16–26 vs. 21) and number of teeth on movable and fixed digits of chelicerae (M.D. 2 vs. 4; F.D. 9 vs. 6) in the original description of A. excelsus. This might be due to chelicerae in the type specimen were not properly opened as shown in the original illustrations and four teeth were considered on the movable digit that was originally on the fixed digit. Rather (1985) also reported this species from Azad Jammu and Kashmir (India) and provided a brief description without details of teeth on cheliceral digits. However, Denmark and Muma (1989) redescribed A. excelsus from type specimens with nine teeth on fixed digit and one tooth on movable digit of chelicerae vs. (F.D 6; M.d. 2) and length of z4 (19 vs. 21) in the original description (Chaudhri et al. 1979).
1. Setae Z5 ≥ 210
...... 2
— Setae Z5 ≤ 190
...... 4
2. Setae Z5 260–280
...... A. fennicus Karg & Huhta, 2012
— Setae Z5 ≤ 230
...... 3
3. Setae z4 51, s4 70, fixed digit of chelicerae with nine teeth, gv3 solenostomes on the ventrianal shield posteroletrad JV2
...... A. williamsi Schicha, 1983
— Setae z4 31, s4 106, fixed digit of chelicerae with 13 teeth, gv3 solenostomes on the ventrianal shield posteromesad JV2
...... A. duncansoni Specht & Rasmy, 1970
4. Setae Z5 ≥ 150
...... 5
— Setae Z5 ≤ 140
...... 8
5. Setae Z4 70–72, s4 52
...... A. filixis Karg, 1970
— Setae Z4 ≥ 90, s4 > 75
...... 6
6. Setae j3 29, s4 98
...... A. nayaritensis De Leon, 1961
— Setae j3 > 40, s4 < 90
...... 8
7. Movable digit of chelicerae with one tooth, without transversely elongate platelet between genital and ventrianal shields; ventrianal shield distinctly wider at ZV1 than at anus level, Z4 and Z5 smooth
...... A. blosis Ahmad & Chaudhri, 1989
— Movable digit of chelicerae with two teeth, with a transversely elongate platelet between genital and ventrianal shields; ventrianal shield slightly wider at ZV1 than at anus level, Z4 and Z5 slightly serrate
...... A. excelsus Chaudhri, Akbar & Rasool, 1979
8. Setae Z4 distinctly longer than s4
...... 9
— Setae Z4 and s4 subequal in length
...... 10
9. Setae Z5 133, 4.5 times as long as j3 28
...... A. collinellus Athias-Henriot, 19661 Athias-Henriot, 1966
— Setae Z5 84, 1.8 times as long as j3 47
...... A. arbuti De Leon, 1961
10. Spermathecal calyx long, bell-shaped
...... A. similoides Buchelos & Pritchard, 1960
— Spermathecal calyx small, cup-, cone-, funnelor dish shaped
...... 11
11. Setae Z5 > 100
...... 12
— Setae Z5 < 90
...... 13
12. Setae j3 35, z4 29–33, movable digit of chelicerae without teeth
...... A. kochi (Koch, 1839) = replacement name of A. similis (Koch, 1839) by Chant and McMurtry 2004
— Setae j3 56, z4 46, movable digit of chelicerae with one tooth
...... A. sinuatus De Leon, 1961
13. Setae z4 23, half the length of j3 44, ventrianal shield more wider at anus level than at ZV1
...... A. marunumus Negm & Gotoh, 2020
— Setae z4 and j3 almost equal in length = 35, ventrianal shield more wider at ZV1 level than at anus level
...... A. yadongensis Wu, 1987
Amblyseius adhatodae Muma, 1967: 268; Moraes et al. 1989b: 79; Chant and McMurtry 2004: 203; Moraes et al. 2004: 12; Gupta 1985: 336; Döker et al. 2022: 866.
Two females, Tripidium bengalense (Retz) (Poaceae), Sargodha, near College of Agriculture, Punjab (32.134294° N, 72.681879° E), 16 Jan. 2016, coll. Muneeb Ur Rehman; one female, Chrysanthemum (Astreraceae), Mazar e Qaid, Karachi, 24.87379° N, 67.03901°° E, 11 Aug. 2023, coll. A. Hafeez and M. Zeeshan.
Brazil (Lawson-Balagbo et al. 2008), India and Pakistan (Muma 1967) and current study, Kenya (Moraes et al. 1989), Sri Lanka (Moraes et al. 2004), Maldives (Döker et al. 2022).
Dorsum — (Figure 5A). Dorsal shield smooth, 365–367 long, 221–224 wide at the level of s4, and 269–273 wide near S2, with seven pairs of solenostomes (gd1, gd2, gd4, gd5, gd6, gd8, gd9); poroids indistinct. Length of setae: j1 34–36, j3 46–48, j4 5–6, j5 5, j6 6, J2 6–7, J5 10–11, z2 7–8, z4 7–8, z5 5, Z1 8–9, Z4 128–132, Z5 248–268, s4 110–115, S2 11–12, S4 11–12, S5 10–11, r3 11, R1 10. All setae smooth, except for Z4, finely serrate. Peritreme extending to the level of j1.
Venter — (Figure 5B). Sternal shield smooth, slightly concave posteriorly, with three pairs of setae (st1-st3) and two pairs of poroids (iv1, iv2); distance st1–st3 61 (63), st2-st2 68 (66), st4 and poroid iv3 tylochorous; genital shield smooth, distance st5-st5 73–75; a transversely elongate platelet between genital and ventrianal shields. Ventrianal shield vase-shaped, smooth, 88–90 long, 62–64 wide at ZV2 level, and 72–74 wide at anus level; with three pairs of pre-anal setae (JV1, JV2, and ZV2) along with para-anal and post-anal setae, a pair of solenostomes (gv3), crescentic shape, posteromesad JV2; four pairs of setae (JV4, JV5, ZV1, and ZV3), four pairs of poroids (ivo and ivp), and two pairs of metapodal platelets on the membrane surrounding ventrianal shield, primary metapodal platelet 16–18, secondary 8–9 long. Setae JV5 63–65, smooth.
Chelicera — (Figure 6A, B). Movable cheliceral digit 31–33 long, with four teeth (three teeth in one specimen); fixed digit 34–35 long, with ten teeth and a pilus dentilis.
Spermatheca — (Figure 6C). Calyx fundibular tube, 25–27 long and 5 wide at apex, atrium distinct, bulbous, four times as wide as proximal section of calyx, calyx forming long tube, almost 4–5 times longer than the width of atrium.
Legs — (Figure 6D). Genua II-IV each with seven setae, macrosetae on genu I 42–44, genu II 34–36, and genu III 48–50, tibia III 34–36, tibia IV with six setae; leg IV with three macrosetae: SgeIV 123–126, StiIV 80–86, StIV 65–70. All macrosetae smooth with pointed tip.
Amblyseius adhatodae belongs to largoensis species subgroup of largoensis species group of the genus Amblyseius. This species subgroup includes 19 species worldwide so far. The morphological characters (shape of spermatheca, length and shape of dorsal setae and chelicerae, and chaetotaxy of body and legs) of specimens of A. adhatodae collected in the present study are very similar to that in the original description (Muma 1967) and subsequent/complementary descriptions by Moraes et al. (1989b) and Döker et al. (2022) except for some variations in length of setae Z4, Z5 and other dorsal setae. Döker et al. (2022) reported 11 teeth on fixed digit in A. adhatodae compared with ten teeth in the present study as well in holotype specimen described by Moraes et al. (1989b) (Table 1).
1. Spermathecal calyx comparatively long, forming a long tube and flared distally, neither widened basally nor saccular
...... 9
— Spermathecal calyx comparatively short, widened saccular or constricted basally forming short tube and flared/widened gradually toward distal end
...... 2
2. Spermathecal calyx almost saccular, not basally constricted
...... 3
— Spermathecal calyx constricted basally forming short tube and flared/widened gradually toward distal end
...... 6
3. Setae Z5 310–336, SgeIV 157
...... A. duplicisetus Moraes & McMurtry, 1988
— Setae Z5 < 270, SgeIV < 100–130
...... 4
4. Movable digit of chelicerae with two teeth
...... A. kokofuensis Ehara, Okada & Kato in Ehara et al. 1994
— Movable digit of chelicerae with four teeth
...... 5
5. Spermathecal calyx not deeply bifurcate, gradually widened toward distal end
...... A. conulus Karmakar, Bhowmik & Sherpa, 2017
— Spermathecal calyx deeply bifurcate, almost equally wide at base and distal end
...... A. sumatrensis Ehara, 2002
6. Sternal shield posteriorly straight
...... 7
—Sternal shield posteriorly with median protrusion
...... 8
7. Setae Z4 76–96, s4 82–96, Z5 240–296
...... A. fijiensis McMurtry & Moraes, 1984
— Setae Z4, Z5 and s4 comparatively long, Z4 = s4 128–133, Z5 76–96, s4 82–96, Z5 305–348
...... A. nambourensis Schicha, 1981
8. Setae StIV 86, Z4 91–125, spermathecal atrium two times as wide as proximal section of calyx
...... A. eharai Amitai & Swirski, 1981
— Setae StIV 38–42, Z4 59–67, spermathecal atrium more distinct, almost three times as wide as proximal section of calyx
...... A. herbiculoides McMurtry & Moraes, 1984
9. Setae Z5 110
...... A. talpa Khan & Ashfaq, 2008
— Setae Z5 > 200
...... 10
10. Setae Z4 59
...... A. keni Shicha, 1987
— Setae Z4 > 80
...... 11
11. Spermathecal calyx trumpet shaped
...... 13
— Spermathecal calyx tubular with parallel sides
...... 12
12. Setae Z5 355–362, Z4 158–167, s4 140–148
...... A. neolargoensis Van der Merwe, 1965
— Setae Z5 206, Z4 96, s4 94
...... A. largoensis (Muma, 1955)
13. Cheliceral fixed digit with six teeth
...... A. diverto Khan, Ashfaq & Hassan, 2010
— Cheliceral fixed digit with ≥ nine teeth
...... 14
14. Movable digit of chelicerae with two teeth
...... A. melicardus Khan & Ashfaq, 2005
— Movable digit of chelicerae with 3–4 teeth
...... 15
15. Spermathecal atrium almost two times as wide as proximal section of calyx, calyx forming short tube, the tube two times as long as the width of atrium, gradually widening toward distal end
...... A. herbicolus (Chant, 1959)
— Spermathecal atrium more bulbous, four times as wide as proximal section of calyx, calyx forming long tube, almost 3–5 times longer than the width of atrium
...... A. adhatodae Muma, 1967, A. impactus Chaudhri, 1968, A. fletcheri Schicha, 1981, A. phillipsi McMurtry & Schicha, 1987*
The morphological features of A. impactus Chaudhri, 1968, A. fletcheri Schicha, 1981, and A. phillipsi McMurtry & Schicha, 1987 are very similar to A. adhatodae (Table 1). So we could not separate these species in the diagnostic key from A. adhatodae. Further research is required to confirm if these species are either distinct species or junior synonyms of A. adhatodae.
Amblyseius isuki Chant & Hansell, 1971:714.
Amblyseius bajulus Chaudhri, Akbar & Rasool, 1979: 70. (Synonymy by Denmark and Muma 1989: 7)
Three females, grasses (Poaceae), Hadoot, Muzaffarabad (34.327106° N, 73.473219° E), 16 July 2023, three females, grasses, Kiran, Neelam Valley, A. J. Kashmir (34.6318277° N, 73.9191069° E), 18 July 2023, one female, grasses (Poaceae), Dam Gala, Mansehra, KPK (34.4062372° N, 73.3375557° E), 20 July 2023, coll. M. Kamran.
British Columbia, Canada (Chant and Hansell 1971); Kalam, Abotabad, Pakistan (Chaudhri et al. 1979).
Dorsum — (Figure 7A). Dorsal shield smooth, 400–410 long and 270–277 wide at the level of s4; with eight pairs of solenostomes (gd1, gd2, gd3, gd4, gd5, gd6, gd8, gd9). Setae length: j1 18–21, j3 29–35, j4 4, j5 4–5, j6 4–5, J2 6–7, J5 7–8, z2 = z4 4–5, z5 4–5, Z1 6–7, Z4 136–142, Z5 260–269, s4 82–89, S2 14–15, S4 9–10, S5 9–10, r3 11–12, R1 9–10. All setae smooth, except for Z4 and Z5, finely serrated near tips. Peritreme extending to the level of j1.
Venter — (Figure 7B). Sternal shield smooth, slightly concave slightly, with three pairs of setae (st1-st3) and two pairs of poroids (iv1, iv2); distance st1-st3 62–66, st2-st2 68–73, st4 and poroid iv3 tylochorous; genital shield smooth, distance st5-st5 69–75; a transversely elongate platelet between genital and ventrianal shields. Ventrianal shield almost pentagonal in shape, wider anteriorly at the level of JV1, with transverse striated pattern, 108–115 long, 103–109 wide at anterior corners, and 73 wide at anus level; with three pairs of pre-anal setae (JV1, JV2, and ZV2) along with para-anal and post-anal setae, a pair of solenostomes (gv3), round shape, posteriomesad JV2; four pairs of setae (JV4, JV5, ZV1, and ZV3), four pairs of poroids (ivo and ivp), and two pairs of metapodal platelets on the membrane surrounding ventrianal shield, primary metapodal platelet 20–22, secondary 12–13 long. Setae JV5 75–80 smooth.
Chelicerae — (Figure 8A). Movable cheliceral digit 28–32 long, with three teeth; fixed digit 30–35 long, with nine teeth and a pilus dentilis.
Spermatheca — (Figures 9A, B). Calyx dish-shaped, 13–14 wide, with a divided granulated stalk, 9 long; atrium distinct, not adjacent to the calyx; minor duct visible (Figures 9A, B).
Legs — (Figure 8B). Genua II-IV each with seven setae, macrosetae on genu I 35–38, genu II 39–42, genu III 60–63, and tibia III 44–47; tibia IV with six setae; leg IV with three macrosetae: SgeIV 98–107, StiIV 84–89, StIV 61–66. All macrosetae smooth with pointed tip.
Amblyseius bajulus, a species described from Pakistan considered as a junior synonym of A. obtusus based on the original descriptions and illustrations without observing its type specimens by Denmark and Muma (1989). While, Chant and McMurtry (2004) considered A. bajulus as uncertain species because of the brief description of spermatheca. The type specimens of A. bajulus were deposited at Acarology Laboratory, University of Agriculture, Faisalabad (AL, UAF). The senior author of the current study contacted AL, UAF and was informed that type specimens of A. bajulus had been lost. The senior author collected a few mite specimens of genus Amblyseius from the northern Pakistan near type locality of A. bajulus. The morphological characteristics (shape of spermatheca, length and shape of dorsal setae and chelicerae, and chaetotaxy of body and legs) of Amblyseius specimens collected in the present study are very similar to the original descriptions of both A. bajulus and A. isuki. Based on description and illustrations, we found that spermatheca of A. bajulus is similar to that of A. isuki where the spermathecal calyx stalk shorter than its width distally in both species (Figures 9C, D), as opposed to A. obtusus, which has a longer stalk of spermathecal calyx than its width distally (Figure 9E). As a result, A. bajulus would be considered as junior synonym of A. isuki, rather than A. obtusus.
The authors are thankful to E. M. Khan, H. M. S. Ali, N. A. Elgoni, and M.W. Shakoor (Acarology Research Laboratory, King Saud University) for their technical support. Thanks are also to the Researchers Supporting Project [RSPD2024R807], King Saud University, Riyadh, Saudi Arabia for supporting the present work.
No potential conflict of interest was reported by the author(s).
This research was funded by the Researchers Supporting Project [RSPD2024R807], King Saud University, Riyadh, Saudi Arabia.

