Mahdavi, Sayed Mosayeb ¹ ; Asadi, Mahdieh ² and Seeman, Owen D. ³

¹Research and Technology Institute of Plant Production, Shahid Bahonar University of Kerman, Kerman, Iran.
²✉ Research and Technology Institute of Plant Production, Shahid Bahonar University of Kerman, Kerman, Iran.
³Queensland Museum, South Brisbane, Queensland 4101, Australia.

2024 - Volume: 64 Issue: 1 pages: 63-75

ZooBank LSID: 09E530D8-1A37-4486-98E2-AA5B7EC34A25

Original research

Keywords

Abstract

Introduction

The Hystrichonychini are bryobiine spider mites with pad-like claws and empodia (Tetranychidae: Bryobiinae). Comprising 18 genera, the largest are Aplonobia Womersley, 1940, Neopetrobia Wainstein, 1956, and Paraplonobia Wainstein, 1960 (Migeon & Dorkeld 2022). Under the currently accepted classification (Smith Meyer 1974; Gutierrez 1985; Bolland et al. 1998), they differ from each other by somewhat minor differences: Neopetrobia has their f₁ setae displaced marginally, Aplonobia has dorsal setae set on strong tubercles, and Paraplonobia has smaller tubercles or none at all. As intermediate species exist, and the defining character states for the genera are subjective, the classification of species can be problematical, especially for Aplonobia and Paraplonobia.

Herein, we describe a new species of Aplonobia, classified into this genus after a reconsideration of genus definitions for Aplonobia and Paraplonobia. It is the first species of this genus recorded from the host plant Haloxylon ammodendron (C.A.Mey.) Bunge (Amaranthaceae).

Previously, within Iran, eight new species of Tetranychidae were described from Kerman (Khodayari et al. 2008; Mahdavi and Asadi, 2015; Mahdavi et al. 2013, 2018, 2021a,b; 2022a) and two species were recorded from H. ammodendron from Iran, namely Augeriflechtmannia haloxyloni Mahdavi, Latifi, Asadi & Auger, 2022 and Aegyptobia carmania Mahdavi & Asadi, 2023 (Mahdavi et al. 2022; Mahdavi & Asadi, 2023).

Material and Methods

Our genus re-evaluation was based on a literature review dealing with the genera Paraplonobia and Aplonobia, including Kamran et al. (2016), who provided a useful overview of morphological variability in Paraplonobia as currently defined.

For the description of the new species, leaves and sheaths of Haloxylon ammodendron (Amaranthaceae) were collected in bags and brought to the laboratory. The dipping-washing-filtering method was used to remove the mites (Boller, 1984) and then was filtered through a sieve (400 Mesh), and the mites retained on the sieve were washed with 70% ethanol into a petri dish, then collected individually under a stereomicroscope and mounted in Hoyer's medium. Mites were identified with an Olympus® BX51 phase-contrast compound microscope. All measurements are given in micrometers (μm) as ranges; the measurements of the holotype are given in parentheses after the range of measurements for the paratypes. The drawings were made using a drawing tube attached to the microscope with 1000x and 1600x magnifications. The terminology and setal notations used for the description follow that of Lindquist et al. (1985). Body size was measured between dorsal setae v₂–h₁ for length and between sc₂–sc₂ for width (Saito et al. 1999). Distances between setae were measured as the distance from the inside edge of one setal base to the other (i.e., the minimum distance between two setal bases). Leg setal numbers are written as the total number of setae, including eupathidia, followed by the additional number of solenidia in parentheses. Depositories are cited using the following abbreviations:

SBUK — Collection of the Acarology Laboratory, Shahid Bahonar University of Kerman, Kerman, Iran.

ACASI — carological Collection, Acarological Society of Iran (ASI), Department of Plant Protection, Faculty of Agriculture, University of Tehran, Karaj, Iran.

Results and discussion

Taxonomic status of the genera Aplonobia and Paraplonobia

Aplonobia and Paraplonobia have a confused taxonomic history that is intertwined with the subgenera Neopetrobia (Langella) Wainstein, 1960, Paraplonobia (Anaplonobia) Wainstein, 1960 and Paraplonobia (Brachynychus) Mitrofanov & Strunkova, 1971, as discussed by Khanjani et al. (2017). Adding to confusion around genus concepts, the current classification differs markedly from the original definitions of Wainstein (1960). Originally, Wainstein (1960) divided Aplonobia into three subgenera: Anaplonobia Wainstein, with empodia shortened and bearing just one group of tenent hairs (versus a long empodium with multiple groups of tenent hairs in the other two subgenera); Paraplonobia Wainstein (with unbranched peritremes and short dorsal setae set on small tubercles or no tubercles at all); and Aplonobia s.str. (with anastomosing peritremes and dorsal setae long and on large tubercles). Smith Meyer (1974) recognized each of Wainstein's subgenera as separate genera and provided new diagnoses, but instead emphasized the size of the tubercles (large in Aplonobia, small or absent in Anaplonobia and Paraplonobia) over the form of the peritremes (simple or anastomosing in Aplonobia, anastomosing in Anaplonobia, simple in Paraplonobia). The form of the empodia was discarded as diagnostic character, with importance placed instead on the combination of small tubercles and anastomosing peritremes. Later, Gutierrez (1985) demoted Brachynychus and Anaplonobia back to subgenera, but under Paraplonobia, thus further underlining the importance of dorsal tubercles over the form of the peritremes. That is, Aplonobia was defined by having strong tubercles and Paraplonobia, including Brachynychus and Anaplonobia as subgenera, was defined by having small tubercles or none at all.

At the time of Smith Meyer (1974), most described taxa seemed to fall reasonably neatly into their genera. However, new taxa have challenged the definitions of these genera, as the size of the tubercles is sometimes open to interpretation, as is apparent in the keys of Gutierrez (1985). In these keys, Aplonobia have ''some or all dorsal setae set on strong tubercles'' and Paraplonobia have ''dorsal setae not set on strong tubercles'' which, at the very least, asks for a judgment on what defines a strong tubercle and whether any of the dorsal setae – which may be on tubercles of different sizes – would be regarded as ''strong''. Not surprisingly, species of Paraplonobia with tubercles can be difficult to classify. We also note that the type species of Paraplonobia, Paraplonobia (Paraplonobia) echinopsili Wainstein, 1960, is one of these intermediate species as it has its dorsal setae set on small tubercles.

Due to this confusion, we propose to define Aplonobia and Paraplonobia based on having anastomosing peritremes (Aplonobia) or simple peritremes (Paraplonobia). The presence of anastomosing peritremes is much less open to interpretation: although some peritremes anastomose less in certain species, whether or not a peritreme is anastomosing is more obvious. We define simple peritremes as those that are a single linear line of chambers. Simple peritremes may bend but they are never divided into a multi-chambered terminal structure. In their most basic form, an anastomosing peritreme in Aplonobia divides just once, as in the circular structure of Aplonobia eurotiae comb. nov. (Mitrofanov & Strunkova, 1975) and the two lines of closely appressed chambers in Aplonobia (Brachynychus) cousiniae comb. nov. (Mitrofanov & Strunkova, 1971) (Çobanoğlu et al. 2021). In the new species, the peritreme divides just two or three times (Fig. 3b). Most species have peritremes that terminate in a complex, bulbous, multi-chambered structure.

Thus, Paraplonobia is narrowed to exclude Paraplonobia (Anaplonobia) and Paraplonobia (Brachynychus), which are moved to Aplonobia. As the boundary between Aplonobia and Paraplonobia (Anaplonobia) was vague (i.e. based on the strength of dorsal tubercles) we consider Anaplonobia a junior synonym of Aplonobia. We also excluded the possibility of retaining Anaplonobia as a subgenus of Aplonobia sensu Wainstein (1960). Under his original concept, Aplonobia (Anaplonobia) has empodia with just one group of tenent hairs while Aplonobia (Aplonobia) has empodia with at least two groups of tenent hairs. This feature possesses diagnostic value but we perceive a grade between species. In Wainstein (1960), the subgenus Anaplonobia was based on A. acharis and A. calame. The type species, A. calame, is unique in having what appears to be one pair of tenent hairs (Pritchard & Baker 1955). In contrast, A. acharis is little different from several other taxa, except that Wainstein (1960) interpreted its five pairs of tenent hairs as one group as they are clustered at their tips. While a monotypic subgenus for A. calame is an option, we prefer to avoid this as another problem is the lack of empodial drawings in many descriptions (e.g., Tuttle & Baker 1968; Baker & Tuttle 1972; Tuttle et al. 1976). Thus, without a full revisionary work based on type specimens, the original concept of Wainstein (1960) is untenable.

Genus Aplonobia Womersley, 1940

Aplonobia Womersley, 1940:252; Pritchard & Baker, 1955:58; Wainstein, 1960: 139; Tuttle & Baker, 1968:47.

Type species — Tetranycopsis histricina Berlese.

Paraplonobia (Anaplonobia) (Wainstein, 1960) syn. nov.

Genus diagnosis

Peritreme anastomosing distally; body oval; prodorsum without lobes and with three pairs of setae, opisthosoma with 10 pairs of setae; setae f₁ and c₃ in normal position; claws and empodium pad-like each with tenent hairs; tarsus of leg I with 2 sets ofduplex setae; coxal setal formula usually 3-3-1-1, rarely 4-3-2-2 (subgenus Brachynychus); dorsal setae with/without tubercles (Table 1), tubercles sometimes large.

Remarks

Aplonobia is distinguished from Paraplonobia by the form of the peritreme (anastomosing in Aplonobia vs. simple in Paraplonobia)

Genus Paraplonobia Wainstein, 1960

Aplonobia (Paraplonobia) Wainstein, 1960: 140.

Paraplonobia: Tuttle and Baker 1968: 48, Smith Meyer 1974: 119, Chaudhri et al. 1974: 28, Gutierrez 1985: 75, Bolland et al. 1998: 7.

Type species — Aplonobia (Paraplonobia) echinopsili Wainstein, 1960 by original designation.

Genus diagnosis

Peritremes simple; body oval; prodorsum without lobes and with three pairs of setae, opisthosoma with 10 pairs of setae; setae f₁ and c₃ in normal position; claws and empodium pad-like each with tenent hairs; tarsus of leg I with 2 sets ofduplex setae; coxal setal formula usually 3-3-1-1. Dorsal setae with/without tubercles.

Note: Only two species with small tubercles (P. (P.) echinopsili Wainstein, 1960 and P. (P.) penicillatus Chaudhri, Akbar & Rasool, 1974).

Remarks

As noted above, the peritremes of A. cousiniae and A. eurotiae are considered anastomosing because they branch. However, previous authors have considered these peritremes differently. Interpretations of the peritreme of A. cousiniae have varied, being multi-chambered in Mitrofanov & Strunkova (1971), anastomosing in Meyer (1974) and simple in Meyer (1987). Here, we consider the termination of the peritreme anastomosing because it divides into two lines of chambers. For A. eurotiae, the original description shows two series of chambers along the peritrematal stem that divide distally and form a circle. The specimens examined by Çobanoğlu et al. (2021) may need re-evaluation as no chambers are shown, but their peritreme does branch distally before reconnecting to form a circle. Thus, in both Mitrofanov & Strunkova (1975) and Çobanoğlu et al. (2021), A. eurotiae has an anastomosing peritreme.

Our re-evaluation of Aplonobia and Paraplonobia creates a homonym: Paraplonobia (Anaplonobia) ambrosiae (Tuttle, Baker & Abbatiello, 1976) [preoccupied by Aplonobia ambrosiae (Tuttle & Baker, 1968)] is renamed Aplonobia neoambrosiae nom. nov.

Key to the genera and subgenera of Paraplonobia/Aplonobia and near genera

The following key begins at couplet 12 of Bolland et al. (1998, p. 7) and includes Geogiobia, which was re-instated as a genus by Zhovnerchuk & Auger (2019).

1. Many coxal (11-9-9-9) and ventral setae, 1 species
...... Tauriobia Livshitz & Mitrofanov, 1967

— Coxal setae formula not exceeding 4-3-2-2
...... 2

2. Setae c₃ moved in lateral position, posterior to c₂ setae, forming an almost longitudinal arrangement with c₂ and e₂ setae, 8 species
...... Georgiobia Wainstein, 1960

— Setae c₃ in normal position
...... 3

3. Peritremes termination simple, 10 species
...... Paraplonobia Wainstein, 1960

— Peritreme termination anastomosing, 53 species
...... Aplonobia Womersley, 1940 – 4

4. Coxal setae formula not exceeding 3-3-1-1, 52 species
...... Aplonobia (Aplonobia) Wainstein, 1960

— Coxal setae formula 4-3-2-2, 1 species
...... Aplonobia (Brachynychus) Mitrofanov & Strunkova, 1971

Family Tetranychidae Donnadieu, 1875

Tribe Hystrichonychini Pritchard & Baker, 1955

Genus Aplonobia Womersley, 1940

Aplonobia Womersley, 1940:252; Pritchard & Baker, 1955:58; Wainstein, 1960: 139; Tuttle & Baker, 1968:47.

Type species — Tetranycopsis histricina Berlese.

Aplonobia aria sp. nov.

ZOOBANK: D55BEB8A-A80C-4D9D-AB87-0CCCAC6F081F

(Figs. 1–4)

Type material

Female (Holotype), 20 females, 1 deutonymph paratypes ex. Haloxylon ammodendron (Amaranthaceae), Kerman-Kerman Province, Iran, 30°14′N 57°13′E, altitude 1835 m; 03 June 2022, coll. S. M. Mahdavi.

Type deposition — Holotype, 19 females and juveniles deposited at SBUK, one female paratype deposited at ACASI.

Diagnosis

Dorsal body setae slender, tapering and pilose, longer than distances between consecutive setae and set on small tubercles; setal bases of e₁ and f₁ closer together than other dorsocentral setae; prodorsum medially with weak longitudinal striae bearing small lobes, longitudinal striae laterally; opisthosoma with transverse striae between dorsocentral setae and longitudinal to oblique striae between dorsolateral setae; stylophore with a small mediocephalic emargination; peritreme anastomosing, branching into 2–3 additional chambers; leg I shorter than body length; setation of legs I–IV lacking many setae: coxae 2-2-1-1; trochanters 0-0-1-1; femora 3-3-2-2; genua 4-4-2-2; tibiae 8+(1)-3-3-3; tarsi 6+2dupl.-7+1dupl.-7-7; empodia with two rows of four tenent hairs.

Description

Female — (n=20; measurement of holotype in parentheses; Figs. 1–4 except 3d). Length of body (excluding gnathosoma) 341–395 (367), v₂–h₁ 317–358 (323); width 265–301 (270), sc₂–sc₂ 169–190 (169).

Dorsum – (Fig. 1a). Prodorsum medially with weak longitudinal striae bearing small lobes, longitudinal striae laterally; opisthosoma with transverse striae between dorsocentral setae, longitudinal to oblique between dorsolateral setae and arched in caudal part; dorsal body setae slender, tapering and pilose (Fig. 1b), dorsocentral setae a little longer than distances between consecutive setae and e₂, f₂; dorsal setae set on small tubercles (Fig. 2), c₁–c₃ and h₁ tubercles without punctations, other opisthosomal setae surrounded by punctations; setal bases of e₁ and f₁ closer together than other dorsocentral setae; dorsal seta c₃ more or less in line with c₁ and c₂, not longitudinally aligned with c₂, d₂, e₂ and f₂. Lengths of setae: v₂ 40–45 (42), sc₁ 45–59 (58), sc₂ 39–47 (46), c₁ 52–65 (65), c₂ 48–55 (50), c₃ 37–48 (43), d₁ 50–64 (59), d₂ 48–69 (55), e₁ 52–64 (60), e₂ 60–77 (73), f₁ 70–80 (80), f₂ 57–68 (59), h₁ 41–49 (47). Distances between dorsal setae: v₂–v₂ 45–48 (45), sc₁–sc₁ 124–131 (124), sc₂–sc₂ 169–190 (169), c₁–c₁ 87–93 (88), c₂–c₂ 158–174 (158), c₃–c₃ 235–266 (235), d₁–d₁ 80–95 (83), d₂–d₂ 160–185 (160), e₁–e₁ 31–35 (31), e₂–e₂ 119–137 (123), f₁–f₁ 26–29 (29), f₂–f₂ 82–87 (84), h₁–h₁ 41–49 (41).

Venter – striae between setae (4a) transverse, anterior of agential setae (ag) and posterior of setae 3a longitudinal (Fig. 3a). Lengths of setae: 1a 34–36 (34), 3a 40–44 (42), 4a 53–55 (53), ag 70–75 (72), g₁ 99–101 (100), g₂ 100–102 (100), ps₁ 11–12 (11), ps₂ 11–12 (11), ps₂ 11–12 (11), h₂ 13–14 (13), h₃ 10–13 (12); distances between setae 1a–1a 34–37 (34), 3a–3a 70–86 (73), 4a–4a 52–55 (55), ag 13–14 (14), g₁ 7–9 (7), g₂ 25–28 (26), h₂–h₂ 11–15 (12), h₃–h₃ 22–24 (23). Coxal setae smooth, except 2c barbed.

Spermatheca – Elongate vesicle, bulb-shaped structure at distal end (Fig. 3c).

Gnathosoma – Peritreme anastomosing distally, branching into 2–3 additional chambers (Fig. 3b); stylophore with a small mediocephalic emargination; palpal setation typical, spinneret (suζ) about 3 times as long as broad (Fig. 4g).

Legs – (Figs. 4a–f). Setation of legs I–IV: coxae 2(1b, 1c)–2(2b, 2c)–1(3b)–1(4b); trochanters 0-0-1(v′)-1(v′); femora 3(d, bv″, l′)-3(d, bv″, l′)-2(d, ev′)-2(d, ev′); genua 4(d, v′, v″, l′)-4(d, v′, v″, l′)-2(d, l′)-2(d, l′); tibiae 9(db, v′, v″, l′, l″, l′1, l″1, v′1, φ)-3(d, v′, v″)-3(d, v′, v″)-3(d, v′, v″); tarsi 6(p′ζ, p″ζ, u′, u″, pv″, v′1+2dupl.(ft′, ω′, ft″, ω″))-7(p′ζ, p″ζ, u′, u″, pv″, v′1, ft′+1dupl.(ft″, ω″))-7(ft′, tc′, tc″, u′, u″, pv″, v′1)-7(ft′, tc′, tc″, u′, u″, pv″, v′1); tarsi II–IV without solenidia; coxal setae smooth, except 2c barbed; claws and empodia I–IV pad-like, empodia with two rows of four tenent hairs (Fig. 4f). Lengths of legs (from trochanter to tarsus): I 237–250 (248), II 174–190 (186), III 178–200 (193), IV 201–230 (218).

Deutonymph — (n=1; Fig. 3d). Length of body (excluding gnathosoma) 342, v₂–h₁ 292; width 250, sc₂–sc₂ 151.

Dorsum – Prodorsal striation pattern close to that of female; dorsal setae similar to female but narrower, only caudal part e, f and h with small tubercles. Lengths of setae: v₂ 35, sc₁ 31, sc₂ 31, c₁ 36, c₂ 36, c₃ 35, d₁ 33, d₂ 42–61, e₁ 49, e₂ 54, f₁ 51, f₂ 52, h₁ 41. Distances between dorsal setae: v₂–v₂ 40, sc₁–sc₁ 102, sc₂–sc₂ 151, c₁–c₁ 70, c₂–c₂ 133, c₃–c₃ 229, d₁–d₁ 71, d₂–d₂ 139, e₁–e₁ 23, e₂–e₂ 99, f₁–f₁ 23, f₂–f₂ 66, h₁–h₁ 28.

Venter – (Fig. 3d). Striation pattern similar to that of female; aggenital area with one pair each of aggenital setae (ag) and genital setae (g₁). Lengths of intercoxal setae: 1a 33, 3a 25, 4a 18, ag 18, g₁ 13, h₂ 10, h₃ 10; distances between intercoxal setae 1a–1a 32, 3a–3a 73, 4a–4a 59, ag–ag 14, h₂–h₂ 10, h₃–h₃ 20.

Gnathosoma – Palp and peritreme similar to female but stylophore completely rounded.

Legs – Setation of legs I–IV: coxae 2(1b, 1c)-2(2b, 2c)-1(3b)-1(4b); trochanters 0-0-0-0; femora 3(d, bv″, l′)-3(d, bv″, l′)-2(d, ev′)-2(d, ev′); genua 2(d, l′)-2(d, l′)-1(d)-1(d); tibiae 7(db, v′, v″, l′, l″, l′1, φ)-3(d, v′, v″)-3(d, v′, v″)-3(d, v′, v″); tarsi 5(p′ζ, p″ζ, u′, u″, pv″+2dupl.(ft′, ω′, ft″, ω″))-6(p′ζ, p″ζ, u′, u″, pv″, ft′+1dupl.(ft″, ω″))-6(ft′, tc′, tc″, u′, u″, pv″)-6(ft′, tc′, tc″, u′, u″, pv″). Setation for legs I–IV differs from adult female by seta (v′) absent on trochanters III-IV, setae (v′, v″) absent on genua I-II, seta (l′) absent on genua III-IV, setae (v′1, l″1) absent on tibiae I and seta (v′1) absent on tarsi I–IV. Lengths of legs (from trochanter to tarsus): I 178, II 136, III 132, IV 136.

Male — Unknown

Remarks

Aplonobia aria sp. nov. is close to Aplonobia calame Pritchard & Baker, 1955 reinst. comb. which shares slender pilose dorsal setae that are mostly longer than the distances between consecutive setae and set on small tubercles and a prodorsal region formed by small lobes. However, the new species differs from A. calame by leg setation tibia I [8+(1) in A. aria vs. 13+(1) in A. calame], leg setation tarsi III–IV (without solenidia in A. aria vs. with solenidia A. calame), the form of the peritreme (few anastomosing chambers in A. aria vs. many chambers, forming a club, in A. calame); the number of empodial hairs (four pairs tenent hairs vs. two hairs, not formed as tenent hairs); the anterior stylophore (with a small mediocephalic emargination in A. aria vs. rounded in A. calame); length of leg I as compared to body length (shorter than body length in A. aria vs. as long as in A. calame).

According to Kamran's key (i.e. couplet 28) in Kamran et al. (2016), Aplonobia aria sp. nov. is close to Aplonobia tshipensis (Smith Meyer, 1974) comb. nov. which was first described as Anaplonobia tshipensis Smith Meyer, 1974. The species differ by the form of the dorsal body setae (densely pilose in A. aria vs. weakly barbed in A. tshipensis); numerous differences in leg setation [A. tshipensis: femora 9-6-4-4; genua 5-5-4-4; tibiae 13(1)-8-6-7; tarsi 16(1)+2dupl.-13+1dupl.-12(1)-12(1)]; the anterior stylophore (small mediocephalic emargination in A. aria vs. deeply incised in A. tshipensis).

The leg chaetotaxy of Aplonobia aria sp. nov. also deserves comment as it has remarkably few leg setae, expressing fewer than all other species of Aplonobia and Paraplonobia. These losses are also apparent in the deutonymph, with numerous setae also absent in this life stage.

Etymology

This species is named after Aria, an old Persian ''satrapy'' – a system of government used in ancient Persia that was established by the Achaemenid Empire during the 6th century BC.

New record of Aplonobia tabukensis (Kamran & Alatawi) comb. nov. for the first time from Iran

Aplonobia tabukensis (Kamran & Alatawi, 2016), 44, Saudi Arabia ex. Haloxylon salicornicum. This species was collected for the first time from Haloxylon ammodendron (Amaranthaceae), Qalehganj, Kerman province, 27°31′N, 57°52′E, 20 Sep. 2018, coll. S. M. Mahdavi.

Depository: SBUK-No. Tet 32.

Notes — In Kamran et al. (2016) there is an inconsistency between their table, figures and text in terms of leg chaetotaxy as it seems all setae on tarsus I-II were not included in the table. Setation of legs I–IV of our samples completely match their drawing as follows: coxa 2-2-1-1; trochanter 1-1-1-1; femora 8-6-3-3; genua 4-5-3-3; tibiae 13(+1)-9-8-8; tarsi 14+2dupl.-11+1dupl.-11+1dupl.-11+1dupl., tarsi I with 12 tactile setae, two sets of duplex setae and two eupathidia; tarsi II with 9 tactile setae, one set of duplex setae and two eupathidia.

Acknowledgements

This work was funded by the National Elites Foundation, Research and Technology Institute of Plant Production, Shahid Bahonar University of Kerman, Kerman, Iran and Fars Elites Charity Association, whom we thank for their assistance. We are very grateful to Dr. Philippe Auger for his helpful comments.

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