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Research Article
New subgenera and a new species of the genus Raphignathus Dugès (Prostigmata, Raphignathidae), with taxonomic notes on the genus Neoraphignathus Smiley & Moser
expand article infoEid Muhammad Khan‡, Muhammad Kamran‡, Jawwad Hassan Mirza‡, Fahad Jaber Alatawi‡
‡ King Saud University, Riyadh, Saudi Arabia

Corresponding author: Fahad Jaber Alatawi ( falatawi@ksu.edu.sa )

Academic editor: Xiao-Feng Xue
© 2023 Eid Muhammad Khan, Muhammad Kamran, Jawwad Hassan Mirza, Fahad Jaber Alatawi.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Khan EM, Kamran M, Mirza JH, Alatawi FJ (2023) New subgenera and a new species of the genus Raphignathus Dugès (Prostigmata, Raphignathidae), with taxonomic notes on the genus Neoraphignathus Smiley & Moser. ZooKeys 1176: 165-180. https://doi.org/10.3897/zookeys.1176.106224
ZooBank: urn:lsid:zoobank.org:pub:91E50336-528C-43DE-8266-42D12D0C7C1C
Open Access

Abstract

Four new subgenera in the genus Raphignathus Dugès are hereby proposed: Raphignathus (Raphignathus), subgen. nov., Raphignathus (Monoraphignathus), subgen. nov., Raphignathus (Diraphignathus), subgen. nov., and Raphignathus (Triraphignathus), subgen. nov. These subgenera are diagnosed by the number of setae on the interscutal membrane of females. A new species, R. (D.) neohecmatanaensis sp. nov., is described and illustrated based on females collected from Ziziphus spina-christi Mill. (Rhamnaceae). The taxonomic status of the monotypic genus Neoraphignathus Smiley & Moser and three species (R. evidus, R. hsiufui, and R. johnstoni) are discussed. A key to world species of the family Raphignathidae is given.

Key words

Acari, new combinations, predatory mite, Raphignathoidea, Saudi Arabia

Introduction

Members of the family Raphignathidae Kramer (Prostigmata, Raphignathoidea) are active predators feeding on small arthropods (Meyer and Ueckermann 1989). They are mostly found in humus soil under dense bushes, leaf litter, lichens, and mosses and on a wide range of plants (Fan and Zhang 2005). Raphignathids have cervical peritremes and contiguous coxae and are divided into two genera: Raphignathus Dugès (76 species) and Neoraphignathus Smiley & Moser (one species, N. howei Smiley & Moser) (Smiley and Moser 1968; Khanjani et al. 2013). These genera are mainly differentiated by the presence and absence of dorsal shields, respectively. Raphignathus species have a worldwide distribution, whereas Neoraphignathus is only known from Louisiana, USA (Beron 2020).

The genus Raphignathus (type species: R. ruberrimus Dugés) was diagnosed as having three, or sometimes four, dorsal shields (Atyeo et al. 1961). While discussing the chaetotaxy of the superfamily Raphignathoidea, Atyeo et al. (1963) provided a general description of the genus Raphignathus. They considered the number of setae on prodorsal shields and on interscutal membrane as important taxonomic characteristics. In addition, the number of setae on the interscutal membrane was also used to differentiate among the species in recently published diagnostic keys (Nasrollahi et al. 2018; Pishehvar and Khanjani 2021). The validity of some Raphignathus species has been questionable due to ontogenetic development, and more females need to be collected and observed (Dönel and Doğan 2011). Raphignathus broomicus Podder was considered a species inquirenda due to uncertain and doubtful characters (Doğan and Erman 2019). Until now, most of published work on the family Raphignathidae has been regional and includes identification keys for China, India, Iran, and Turkey (Fan and Yin 2000; Doğan and Erman 2019; Pishehvar and Khanjani 2021).

In the present study, four new subgenera of Raphignathus are erected based on the prominent and consistent morphological character (number of setae on interscutal membrane). A new species, R. (D.) neohecmatanaensis sp. nov. is described and illustrated based on females. Some taxonomic notes on the monotypic genus, Neoraphignathus Smiley & Moser, and the identity of three species (R. evidus, R. hsiufui, and R. johnstoni) are concisely discussed. A diagnostic key to the world species is also provided.

Material and method

All published taxonomic literature on the family Raphignathidae was critically reviewed to confirm the validity of the species, subgeneric divisions and to prepare a diagnostic key of world species. The new raphignathoid species was collected by shaking foliage of Ziziphus spina-christi Mill. (Rhamnaceae) over a white sheet of paper; mite specimens were preserved in small vials containing 70% ethanol. The specimens were permanently mounted on glass slides in Hoyer’s medium and identified under a phase-contrast microscope (BX51, Olympus, Tokyo, Japan). All measurements of the holotype specimen are given in micrometers (μm), followed by those of paratypes in the parenthesis. The terminology and abbreviations used in the description of the new species follow those of Kethley (1990) and Grandjean (1939, 1944, 1946). The holotypes and paratypes were deposited at the King Saud Museum of Arthropods (KSMA, Acarology section), Department of Plant Protection, College of Food and Agriculture Sciences, King Saud University, Riyadh, Saudi Arabia.

Results

Four new subgenera of the genus Raphignathus are proposed: Raphignathus (Raphignathus), subgen. nov., Raphignathus (Monoraphignathus), subgen. nov., Raphignathus (Diraphignathus), subgen. nov., and Raphignathus (Triraphignathus), subgen. nov., on the basis of the number of setae on the interscutal membrane, a prominent and consistent morphological character. A new species, R. (D.) neohecmatanaensis sp. nov., is described and illustrated based on adult females. Furthermore, taxonomic notes on the status of the monotypic genus, Neoraphignathus Smiley & Moser, and the identity of three species (R. evidus, R. hsiufui and R. johnstoni) are discussed. A diagnostic key to the world species is also presented.

Raphignathidae Kramer, 1877

Raphignathidae Kramer, 1877: 215

Type genus

Raphignathus Dugès, 1834: 53

Diagnosis

(based on Krantz and Walter (2009) and Fan and Zhang (2005)). Peritremes linear, not imbedded in dorsal surface of stylophore; paired peritremes running laterally from base of stylophore to make short loops in collar membrane between gnathosoma and podosoma; coxae II and III contiguous; stigmata opening at base of chelicerae.

Taxonomic divisions of the genus Raphignathus

The presence or absence of prodorsal shields are diagnostic for the differentiation of the two existing raphignathid genera. Atyeo (1963) discussed in detail the chaetotaxy of the superfamily Raphignathoidea while describing seven Raphignathus species. This author also provided comprehensive diagnosis of Raphignathus and stated that the number of setae on the shields and the interscutal membrane is a consistent and important diagnostic character. The number of setae on the interscutal membrane is considered to be a strong character and has been used in identification keys to distinguish species (Nasrollahi et al. 2018; Pishehvar and Khanjani 2021). Through our extensive study of the literature of all 76 Raphignathus species, we find that the number of setae on the interscutal membrane can be used to erect subgenera.

In the current study, we categorize species of the genus Raphignathus into four new subgenera based on the number of setae on the interscutal membrane. These four subgenera are Raphignathus (Raphignathus), subgen. nov. without setae (11 spp.), Raphignathus (Monoraphignathus), subgen. nov. with one seta (14 spp.), Raphignathus (Diraphignathus), subgen. nov. with two setae (33 spp.), and Raphignathus (Triraphignathus), subgen. nov. with three setae (10 spp.).

Raphignathus (Raphignathus) , subgen. nov.

https://zoobank.org/4A81EA35-DC89-4971-BD24-0459A27A8621

Type species

Raphignathus ruberrimus Dugès, 1834: 53.

Diagnosis

Interscutal membrane without setae.

Etymology

The subgeneric epithet refers to the nominotypical subgenus.

Raphignathus (Monoraphignathus) , subgen. nov.

https://zoobank.org/D332A8E8-0390-4D39-95D4-A8C6E7E0B8D5

Type species

Raphignathus bathursti Meyer & Ryke, 1960: 229.

Diagnosis

Interscutal membrane with one pair of setae.

Etymology

The subgeneric epithet refers to the one pair of setae on interscutal membrane.

Raphignathus (Diraphignathus) , subgen. nov.

https://zoobank.org/0D7C8B12-10E8-4019-A2FA-591934FD9B17

Type species

Raphignathus gracilis (Rack, 1962): 281.

Diagnosis

Interscutal membrane with two pairs of setae.

Etymology

The sub-generic epithet refers to the two pairs of setae on interscutal membrane.

The species included in this new subgenus are widely distributed over the world.

Raphignathus (Triraphignathus) , subgen. nov.

https://zoobank.org/80C632B9-6521-47BB-A6E2-3EA44EE761DD

Type species

Raphignathus domesticus Shiba, 1969: 157.

Diagnosis

Interscutal membrane with three pairs of setae.

Etymology

The subgeneric epithet refers to the interscutal membrane with three pairs of setae.

Notes on the taxonomic status of the genus Neoraphignathus

To date, the family Raphignathidae has included two genera, Raphignathus and Neoraphignathus, which have been differentiated based on the presence or absence of shields on the dorsum. The monotypic genus, Neoraphignathus (type species: N. howei Smiley & Moser) was erected in 1968, based on a single female holotype specimen without detailed description and illustration. Based on observations and the collection of the immature specimens of the genus Raphignathus, prodorsal shields are weakly developed or absent in immatures. Atyeo et al. (1961) has reported that dorsal shields are sometime feebly developed. Since its first description, the type species, N. howei, has not been redescribed, nor have new Neoraphignathus species been described. We suggest that the type specimen of N. howei be re-examined and also that more specimens be collected from the type locality to confirm the absence of a dorsal shield to confirm the validity of Neoraphignathus.

Notes on the validity of Raphignathus evidus, R. hsiufui, and R. johnstoni

The taxonomic identity of Raphignathus evidus Fan, R. hsiufui Fan, and R. johnstoni Womersley are doubtful. These species were originally described based on single specimens, minor differential characteristics (i.e. number of dorsal setae on the lateral prodorsal shield; all three species have two pairs of setae on lateral shields), and small opisthosomal shields. In contrast, all other Raphignathus species have three pairs of setae on the lateral prodorsal shields along with the pores (ia). The immature stages of Raphignathus gradually develop the prodorsal shields, striation patterns, and leg setae (Fan and Yin 2000). For instance, the immature stages of R. giselae, R. lenis, and R. caspicus each have two setae on the lateral prodorsal shields (three setae in adult) and small lateral prodorsal shields with weakly developed striations. Moreover, we also observed the immatures from more than 10 populations of Raphignathus and found reduced size of weakly sclerotized lateral prodorsal shields and setae set on the edges of shields. Based on this evidence, R. evidus, R. hsiufui, and R. johnstoni should be revised and more specimens collected to confirm their validity.

New species

Raphignathus (Diraphignathus) neohecmatanaensis sp. nov.

https://zoobank.org/12E13CEE-478F-4A3F-A47E-B3C7F34C73FA
Figs 1, 2, 3, 4, 5–8

Diagnosis

Female: endopodal shield absent between coxae I–IV; two small shields present posterolateral to median prodorsal shield; palp femora with two setae; femora 6–5–3–2; genua 5(+κ)-5(+κ)–4–4; tibiae 5(+φρ)–5(+φρ)–4(+φρ)–4 (+φρ); tarsi 21(1ω) –20(1ω)–15–14.

Description

Female (n = 4). Idiosoma oval, length of body (including gnathosoma) 533 (525–545); width of body 345 (338–353).

Dorsum (Fig. 1). Propodosoma with one medial and two lateral shields each containing three setae; medial sclerite with setae vi, si and c1; paired ovoid lateral shields each with an eye, one cupule (ia) and sci, sce and seta c2 seta; opisthosomal setal pairs d1, e1 and the pair of cupuli (im); posterior opisthosomal shield large, rectangular, bearing four pairs of setae (f1, h1–3) and one pair of the cupule (ip); all dorsal shields finely punctate; dorsal body setae setiform, smooth and acute; pseudanal setae ps1 dorsally located. Lengths of dorsal setae: vi 25 (23–27), ve 28 (26–30), sci 27 (26–29), sce 28 (26–29), c1 23 (17–19), c2 25 (19–23), d1 21 (20–23), e1 22 (21–23), f1 21 (19–22), h1 24 (23–25), h2 25 (23–25), h3 22 (21–24), ps1 23 (21–25); distances between dorsal 195 setae: vi–vi 27 (29–31), sci–sci 123 (118–128), vi– sci 70 (66–72), sce–c2 68 (65–72), c1–c1 30 (28–32), d1–d1 99 (92–101), c1–d1 53 (49–55), d1–e1 22 (21–24), f1–f1 88 (83–90), e1–f1 65 (61–68), h1–h1 45 (42–48), h1–h2 43 (41–46), h2–h2 77 (72–80), h3–h3 101 (98–106).

Figure 1. 

Raphignathus (Diraphignathus) neohecmatanaensis sp. nov. (female), dorsum. Scale bar: 100 µm.

Venter (Fig. 2). Venter entirely striated, without punctations; coxisternal shields absent (Fig. 2); ventral setae lengths: 1a 42 (41–44), 1b 38 (35–39), 1c 34 (33–36), 2b 36 (30–34), 2c 32 (30–34), 3a 24 (23–26), 3b 18 (17–20), 3c 36 (34–38), 4a 22 (23–27), 4c 38 (35–40); two pairs of aggenital setae (ag1–2) with one cupule (ih) on each side of the genital shield; anal opening and genital shields separate; genital shield prominent with a few punctations, bearing three pairs of genital setae (g1–3); anal opening terminal, with three pairs of setae (ps1–3), ps1 dorsal ps3 and ps2 ventral; ventral setal lengths: ag1 29 (27–32); ag2 27 (25–28); g1 31 (28–33); g2 25 (23–28); g3 20 (19–22); ps2 22 (21–23); ps3 21 (20–22). Distances between ventral setae: 1a–1a 55 (53–58), 3a–3a 115 (97–109), 4a–4a 70 (68–73), ag2–ag2 42 (41–43), g1–g1 38 (35–40), g2–g2 52 (48–55), g3–g3 75 (74–75), 2b–2c 30 (25–29), 1a–3a 50 (49–53), 3a–4a 72 (68–75), 4a–ag1 85 (82–88), ag1–ag2 65 (63–68), ag2–g1 63 (60–65), g1–g2 15 (14–17), g2–g3 22 (21–25), ag–g1 42 (40–45), g3–ps3 16 (15–18), ps2– ps3 16 (14–18).

Figure 2. 

Raphignathus (Diraphignathus) neohecmatanaensis sp. nov. (female), venter. Scale bar: 100 µm.

Gnathosoma (Figs 3, 4). Ventral infracapitular with two pairs of very long setae (m and n), m 40 (39–42), n 52 (49–54) and two pairs of pilose adoral setae (or1–2), or1 23 (21–24), or2 20 (19–22) (Fig. 3); stylophore conical and striated; palp chaetotaxy (femur-tarsus) as follows: 3–2–4+1 claw 4+1ω+4 eupathidia (ζ) (Fig. 4).

Figure 3. 

Raphignathus (Diraphignathus) neohecmatanaensis sp. nov. (female), gnathosoma. Scale bar: 50 µm.

Figure 4. 

Raphignathus (Diraphignathus) neohecmatanaensis sp. nov. (female), palp. Scale bar: 20 µm.

Legs (Figs 5–8). Length of legs I–IV (without coxae): 340 (328–348); 270 (276–286); 325 (317–328); 375 (367–384), respectively. Chaetotaxy on legs I–IV (solenidia in parentheses and not included in setal counts): coxa 2–2–2–1; trochanter 1–1–2–1; femora 6–5–3–2; genua 5(+κ)-5(+κ)–4–4; tibiae 5(+φρ)–5(+φρ)–4(+φρ)–4(+φρ); tarsi 21(1ω) –20(1ω)–15–14.

Figures 5–8. 

Raphignathus (Diraphignathus) neohecmatanaensis sp. nov. (female), Legs I–IV. Scale bar: 20 µm.

Male and immature stages

Unknown.

Type materials

Holotype female and three paratype females, Faifa, Jazan, 24°30.412'N, 39°36.578'E, 8 Oct., 2020, collected from Ziziphus spina-christi Mill. (Rhamnaceae) by Eid M. Khan, Jawwad H. Mirza & Hafiz S. Mushtaq.

Etymology

The specific epithet is in reference to the similarity of the new species to R. (D.) hecmatanaensis; neo = new.

Remarks

Raphignathus (D.) neohecmatanaensis sp. nov., belongs to the subgenus Diraphignathus subgen. nov. The new species resembles to R. (D.) hecmatanaensis Khanjani & Ueckermann in having two pairs of setae (d1 and e1) on the interscutal membrane, two setae on the palp femora, and two small plates present posterolateral to the median prodorsal shield. However, the new species differs from R. (D.) hecmatanaensis in the absence of an endopodal shield (vs present), femur IV with two setae (vs three), and leg tarsus I with one solenidion (vs two solenidia).

Key to genera, subgenera, and all known species of the family Raphignathidae

Five species are not included in the key. Raphignathus longimanus (Koch), R. impressus (Koch), R. hispidus (Dugès), and R. deserticula (Trägårdh) because their descriptions are incomplete, and R. lanuginosus Atyeo is excluded, as it was described on the male.

1 Dorsum with well-developed shields, one medial, one pair of lateral shields on prodorsum and one hysterosomal shield Genus Raphignathus [2]
– Dorsum without shields Genus Neoraphignathus , N. howei Smiley & Moser
2 Interscutal membrane without setae, hysterosomal shield with six pairs of setae Raphignathus (Raphignathus) subgen. nov. [3]
– Interscutal membrane with ≥1 setae 13
3 Opisthosoma or opisthosomal shield reticulated 4
– Opisthosomal shield smooth or otherwise, not reticulated 6
4 Prodorsal shields reticulated R. (R.) crustus Fan & Zhang, New Zealand
– Prodorsal shields smooth or punctate 5
5 Tibia I with 5 +2φ, Tibia III with 5+1φ, dorsal body setae comparatively long (54–74) R. (R.) kurdistaniensis Nasrollahi, Khanjani & Mirfakhraei, Iran
– Tibia I with 5 +1φ, Tibia III with 4+1φ, dorsal body setae comparatively short (24–36) R. (R.) darjeelingensis Gupta, India
6 Opisthosoma without distinct shield; setae d1, e1, and f1 very minute, 1/3–1/2 length of setae v1 R. (R.) guajavae Gupta, India
– Opisthosoma with distinct opisthosomal shield; setae d1, e1, and f1 at least 2/3 length of setae v1 7
7 Genu I and II each with a large leaf-like solenidion R. (R.) pycnonotus (Gupta & Paul), India
– Genu I and II each with a small, slender solenidion 8
8 Setae c1 short, reaching 1/3 length of interscutal membrane, far behind the anterior margin of opisthosomal shield; tarsus IV with large solenidion R. (R.) inornata (Gupta & Paul), India
– Setae c1 crossing interscutal membrane and anterior margin of opisthosomal shield; tarsus IV with small solenidion 9
9 Tibia I with one solenidion 10
– Tibia I with two solenidia 12
10 Genu IV with three setae R. (R.) hirtellus Athias-Henriot, Algeria
– Genu IV with four setae 11
11 Ratios d1–d1/c1–c1 = 3.00, c2–c2/d1–d1 = 3, d1–d1/e1–e1 = 0.58, e1–e1/f1–f1 = 0.94–1.00 R. (R.) neocardinalis Atyeo, The Bahamas
– Ratios d1–d1/c1–c1 = 6.00, c2–c2/d1–d1 = 1.22, d1–d1/e1–e1 = 1.38, e1–e1/f1–f1 = 0.65 R. (R.) conspicuus (Berlese), Colombia
12 Dorsal setae comparatively long; setae c1, and d1 crossing bases of d1 and e1, respectively R. (R.) khorramabadensis Bagheri, Jafari & Paktinat, Iran
– Dorsal setae comparatively short; setae c1, and d1 far behind bases of d1 and e1, respectively R. (R.) cardinalis (Ewing), USA
13 Interscutal membrane with one pair of setae Raphignathus (Monoraphignathus) subgen. nov. [14]
– Interscutal membrane with more than one pair of setae 27
14 Palp femur with two setae; femur I with three setae R. (M.) arabicus Gomaa & Hassan, Egypt
– Palp femur with three setae; femur I with five or six setae 15
15 Femur IV with two or three setae 16
– Femur IV with four setae 21
16 Femur IV with two setae 17
– Femur IV with three setae 19
17 Genu II with five setae including micro setae R. (M.) costatus Chaudhri, Akbar & Rasool, Pakistan
– Genu II with six setae including microsetae 18
18 Setae e1 reaching to bases of h1; dorsal body setae with spinules along entire length R. (M.) zhaoi Hu, Jing & Liang, China
– Setae e1 reaching half distance to bases of h1 (or distance e1–h1), dorsal body setae with spinules along entire length R. (M.) kuznetzovi Dogan & Ayyildiz, Turkey
19 Setae c2 crossing bases of d1, Setae c1 extending to bases of e1, setae e1 extending to bases of h1 R. (M.) ueckermanni Koç & Kara, Turkey
– Setae c2 crossing setae c1 far behind to the bases of e1, setae e1 far behind to the bases of h1 20
20 Dorsal body setae ensiform, setae c1 far behind bases of d1, d1–d1 distance almost five times more than c1–c1 distance R. (M.) ensipilosus Meyer & Ueckermann, South Africa
– Dorsal body setae setiform, setae c1 far behind bases of d1, d1–d1 distance almost equal to c1–c1 R. (M.) cometes Atyeo, Bahama-Islands
21 Small shields absent posterolateral to median prodorsal shield 22
– Small shields present posterolateral to median prodorsal shield 25
22 Genital plates/covers with four pairs of setae R. (M.) koseiensis Dönel & Doğan, Turkey
– Genital plates/covers with three pairs of setae 23
23 Femur I and II each with five setae R. (M.) solimani Hassan & Gomaa, Egypt
– Femur I and II each with six setae 24
24 Dorsal setae comparatively long; most setae cross base of next consecutive setae R. (M.) kelkitensis Dönel & Doğan, Turkey
– Dorsal setae comparatively short; most setae far behind base of next consecutive setae R. (M.) fani Doğan & Ayyildiz, Turkey
25 Dorsal setae comparatively long; most setae reach or cross base of next consecutive setae R. (M.) bathursti Meyer & Ryke, South Africa
– Dorsal setae comparatively short; most setae far behind base of next consecutive setae 26
26 Trochanter III with three setae R. (M.) afyonensis Akyol & Koç, Turkey
– Trochanter III with two setae R. (M.) collegiatus Atyeo, Baker, & Crossley, USA
27 Interscutal membrane with two pairs of setae R. (Diraphignathus) subgen. nov. [28]
– Interscutal membrane with three or four pairs of setae R. (Triraphignathus) subgen. nov. [60]
28 Medial prodorsal shield with two pairs of setae 29
– Medial prodorsal shield with three pairs of setae 30
29 Setae c1 present, setae vi absents; presence of plates behind the anteromedian plate; femur IV with 2 setae R. (D.) evansi Zaher & Gomaa, Egypt
– Setae c1 absent, setae vi present; dorsum without а pair of small plates behind anteromedian plate; femur IV with 3 setae R. (D.) ehari Zaher & Gomaa, Egypt
30 Genital plates/covers with four pairs of setae 31
– Genital plates/covers with three pairs of setae 32
31 Two small shields posterior to median prodorsal shield absent; endopodal shields absent; setae f1 behind the anterior margin of opisthosomal shield R. (D.) saboorii Ghorbani & Bagheri, Iran
– Two small shields posterior to median prodorsal shield present; endopodal shields present; setae f1 on the anterior margin of opisthosomal shield R. (D.) karabagiensis Akyol & Koç, Turkey
32 Palp femur with two setae 33
– Palp femur with three setae 44
33 Two small shields posterior to median prodorsal shield present 34
– Two small shields posterior to median prodorsal shield absent 38
34 Endopodal shield present; femur IV with three setae 35
– Endopodal shield absent; femur IV with two setae 36
35 Tarsus I with two solenidia R. (D.) hecmatanaensis Khanjani & Ueckermann, Iran
– Tarsus I with one solenidion R. (D.) arcus Akyol, Turkey
36 Femur IV with two setae; tarsus I with one solenidion R. (D.) neohecmatanaensis sp. nov. Alatawi & Kamran, Saudi Arabia
– Femur IV with three setae; tarsus I with two solenidia 37
37 Lateral prodorsal shield with one pair of pob; tibiae III 5(+1φp) tarsi 18(+1ω+1ω2) R. (D.) seraji Pishehvar & Khanjani, Iran
– Lateral prodorsal shield without pob; tibiae III 5 tarsi 19(+1ω+1ω2) R. (D.) rakhshandehi Pishehvar & Khanjani, Iran
38 Dorsal setae distally forked or tricarinate R. (D.) furcisetosus Meyer & Ueckermann, South Africa
– Dorsal setae simple, not distally forked or tricarinate 39
39 Femur IV with two setae R. (D.) erzincanica Doğan, Turkey
– Femur IV with three setae 40
40 Opisthosomal shield reduced; interscutal membrane more longer than opisthosomal shield 41
– Opisthosomal shield equally long or longer than interscutal membrane 43
41 Dorsal setae stout, serrate and blunt-tipped R. (D.) membranus Fan & Yin, China
– Dorsal setae simple, distally pointed 42
42 Interscutal membrane four times longer than much reduced opisthosomal shield; f1 on anterior margin of opisthosomal shield R. (D.) vahiti Doğan, Turkey
– Interscutal membrane slightly longer than opisthosomal shield; f1 behind anterior margin of opisthosomal shield R. (D.) giselae Meyer & Ueckermann, Zimbabwe
43 Median prodorsal shield anteriorly extending to peritremes and wider anteriorly near setae sci as compared to posterior half; setae f1 on anterior margin of opisthosomal shield R. (D.) gracilis (Rack), Germany
– Median prodorsal shield anteriorly far behind peritremes and almost equally wide anteriorly near setae sci and at posterior half; setae f1 just behind anterior margin of opisthosomal shield R. (D.) bakeri Zaher & Gomaa, Egypt
44 Small shields posterior to median prodorsal shield absent 49
– Two small shields posterior to median prodorsal shield present 45
45 Coxae II with one seta R. (D.) atyeoi Meyer & Ueckermann, South Africa
– Coxae II with two setae 46
46 Femur IV with two setae 47
– Femur IV with three setae 48
47 Coxae III and IV with endopodal shields; setae f1 on posterior margin of interscutal membrane; distance f1–f1 < d1–d1 R. (D.) summersi Robaux, USA
– Coxae III and IV without endopodal shields; setae f1 far behind posterior margin of interscutal membrane; distance f1–f1 > d1–d1 R. (D.) aciculatus Fan, China
48 Tarsus I–IV 19+ ω, 15+1ω, 13, 12 R. (D.) africanus Meyer & Ueckermann, South Africa
– Tarsus I–IV 21+1 ω, 21+1ω, 15, 14 R. (D.) hatamii Khanjani & Pishehvar, Iran
49 Coxa II with one seta R. (D.) rarus Kuznetsov, USSR
– Coxa II with two setae 50
50 Coxae III and IV without endopodal shields 51
– Coxae III and IV with endopodal shields 53
51 c1–f1/f1–f1 = 0.70; c1–f1 < f1–f1; space between setae f1–f1 twice as wide as between setae d1–d1 R. (D.) atomatus Fan & Zhang, New Zealand
– c1–f1/f1–f1 = 1.50–1.87; c1–f1 1.5–2.0 times more than f1–f1; distance f1–f1 ≤ d1–d1 52
52 Dorsal setae barbed; setae c1, d1, and e1 reach or cross bases of next consecutive setae, distances d1–f1 ≤ f1–f1; setae f1 near anterior margin of opisthosomal shield R. (D.) satoi Shiba, Malay Peninsula
– Dorsal setae simple; setae c1, d1, and e1 not reaching base of next consecutive seta; distances d1–f1 1.31 times more as f1–f1, f1 behind anterior margin of opisthosomal shield R. (D.) kamiensis Meyer & Ueckermann, South Africa
53 Femur I and II with five and four setae, respectively R. (D.) hexeris Chaudhri, Akbar & Rasool, Pakistan
– Femur I and II with six and five setae, respectively 54
54 Opisthosomal shield 2–4 times wider than interscutal membrane 55
– Interscutal membrane as wide as or more wider than opisthosomal shield 56
55 Setae c1 far behind posterior margin of prodorsal shield; tibia III with five setae excluding solenidion; opisthosomal shield four times wider than interscutal membrane R. (D.) neogracilis Robaux, USA
– Setae c1 on the posterior margin of prodorsal shield; tibia III with four setae excluding solenidion; opisthosomal shield twice as wide as interscutal membrane R. (D.) scutatus Kuznetsov, USSR
56 Femur IV with two setae 57
– Femur IV with three setae 59
57 Dorsal shields without striations, tarsus I with 22 setae R. (D.) tumidus Kuznetsov, USSR
– Dorsal shields with fine, sparse puncta and faint striae; tarsi I with 21 setae 58
58 Tarsi III–IV with 14 and 13 setae, respectively; femur II with five setae R. (D.) caspicus Doustaresharaf and Kazemi, Colombia
– Tarsi III–IV with 15 and 14 setae, respectively; femur II with six setae R. (D.) tamaricis Poudineh, Ramroodi & Bagheri, Iran
59 Setae f1–f1 ≤ c1–c1 and d1–d1 R. (D.) giresuniensis Doğan, Turkey
– Setae f1 twice as widely spaced as c1–c1 R. (D.) orientalis Fan & Li, China
60 Medial prodorsal shield with two pairs of setae R. (T.) lenis Barillo, Uzbekistan
– Medial prodorsal shield with ≥3 pairs of setae 61
61 Genital shield with four pairs of setae 62
– Genital shield with three pairs of setae 63
62 Two small shields present posterolateral to prodorsal shield; endopodal shields near coxae III and IV absent R. (T.) sceptrum Chaudhri, Akbar & Rasool, Pakistan
– Small shields absent posterolateral to prodorsal shield; endopodal shields near coxae III and IV present R. (T.) quadrigeminus Dönel & Doğan, Turkey
63 Palp femur with three setae R. (T.) aethiopicus (Meyer & Ryke), South Africa
– Palp femur with two setae 64
64 Femur IV with two setae R. (T.) karrooi Meyer & Ueckermann, South Africa
– Femur IV with three setae 65
65 Femur I with five setae, femur II with four setae R. (T.) domesticus Shiba, Japan
– Femur I with six setae, femur II with five setae 66
66 Two small shields present posterolateral to prodorsal shield 67
– Small shields absent posterolateral to prodorsal shield 68
67 Tibiae III with four setae, solenidia absent R. (T.) hamooniensis Poudineh, Ramroodi & Bagheri, Iran
– Tibiae III with five setae with one solenidion R. (T.) larestanensis Bagheri, Akrami & Majidi, Iran
68 Genu II with four tactile setae; endopodal shields near coxae I–II present R. (T.) emirdagiensis Akyol & Koç, Turkey
– Genu II with five tactile setae; endopodal shields near coxae I–II absent R. (T.) ozkani Doğan, Turkey

Discussion

The taxonomic classification of predatory mites of the genus Raphignathus are revised, and for the first time, the genus is divided into four subgenera by considering the morphologically valid, persistent, and prominent characters (Atyeo 1963). The use of subgenera supports the identification of raphignathoid species and will help to avoid designation of new species based on variable characters. Raphignathus evidus, R. hsiufui, and R. johnstoni are considered doubtfully valid. They were described based on size of lateral prodorsal shields and number of setae, but these in these characteristics they resemble immature stages (Fan and Yin 2000). The monotypic genus Neoraphignathus was erected based on only the single type specimen with a restricted geographical region and its description is brief. We suspect it might have been described based on the immature stage of a Raphignathus species, and we suggest that the type species be revised and more specimens collected to confirm the validity of the species and genus.

Acknowledgements

The authors extend their appreciation and special thanks to Edward Ueckermann (North-West University, South Africa), Carlos Holger Wenzel Flechtmann (Universidade de São Paulo, Piracicaba, São Paulo, Brazil), Zhi-Qiang Zhang (Landcare Research, Auckland, New Zealand), and Daochao Jin (Guizhou University, Guiyang, China) for providing useful literature.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The authors would like to extend their sincere appreciation to the researchers supporting project number (RSPD2023R807), King Saud University, Riyadh, Saudi Arabia.

Author contributions

Conceptualization: MK, FJJA. Data curation: JHHM, EMK. Funding acquisition: FJJA. Investigation: EMK, MK, FJJA. Methodology: JHHM, EMK. Resources: FJJA. Supervision: FJJA. Visualization: JHHM, EMK. Writing - original draft: JHHM, EMK. Writing - review and editing: FJJA, MK.

Author ORCIDs

Eid Muhammad Khan https://orcid.org/0000-0001-5008-2524

Muhammad Kamran http://orcid.org/0000-0001-6084-203X

Jawwad Hassan Mirza https://orcid.org/0000-0002-1726-4331

Fahad Jaber Alatawi https://orcid.org/0000-0002-6824-2650

Data availability

All of the data that support the findings of this study are available in the main text.

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