New Genus and New Subgenera of Camerobiid Mites (Acari: Prostigmata: Camerobiidae) with a Key to World Species of the Genus Neophyllobius

Abstract

Simple Summary

The present study erects a new genus, Monobius Alatawi and Kamran, where all the leg tarsi in females have one midventral seta. Moreover, the genus Tillandsobius Bolland is synonymized with the genus Tycherobius Bolland and the genus Neophyllobius Berlese is categorized in two new subgeneric divisions. For the first time, a key to all known species of the genus Neophyllobius is provided. The ambiguities in the ventral idiosoma setal notation are highlighted and discussed.

Abstract

A new genus, Monobius Alatawi and Kamran, is hereby proposed for the two already described species, viz; M. electrus (Żmudziński) and M. meyerae (Bolland). In addition, the monospecific genus Tillandsobius Bolland is synonymized with the genus Tycherobius Bolland due to variations in the setae number of tibiae I–IV. Further, the genus Neophyllobius Berlese is categorized in two new subgeneric divisions as Neophyllobius Berlese and Monophyllobius Mirza. The number and position of the midventral setae on tarsi I–IV are considered as strong diagnostic generic and subgeneric diagnostic characters. The present study also includes the key to all known species of the genus Neophyllobius. The morphological characters of ten poorly described Neophyllobius species were studied in detail through published literature. The ambiguities in the ventral idiosoma setal notation are highlighted and discussed. It is concluded that two intercoxal setae 3a–4a are always present on small platelets, paired aggenital setae (ag) are present anteriorly and paired genital setae (g) present posteriorly on genital shield. In addition, five records of new species for Saudi Arabia are reported along with re–descriptions of three species.

1. Introduction

The family Camerobiidae Southcott (Acari: Prostigmata) is the second largest family in the superfamily Raphignathoidea. It consists of more than 170 species in seven genera that can be differentiated in two groups based on the position of the solenidion on tarsi I–II. The three genera, viz; Neophyllobius Berlese [1], Tillandsobius Bolland [2] and Tycherobius Bolland [2] have a solenidion present on the basal half of the tarsi I–II, while the four genera, Acamerobia Fan and Walter [3], Camerobia Southcott [4], Bisetalobius du Toit, Theron and Ueckermann [5] and Decaphyllobius Bolland [2] have a solenidion present on distal half of tarsi I–II.

The camerobiid mites, also known as stilt–legged mites, are non-potential predators feeding on different phytophagous pest mites and crawlers of scale insects [2]. Although they are widely distributed in both, temperate and tropical zones, their biology is not yet studied [2,3]. In the field, camerobiid mites are present in low numbers as reported for all the described species, globally [2,3,4,5]. There are four active developmental stages, viz. larva, protonymph, deutonymph and adult), which can be found on ground cover grasses, on plant foliage and under the bark of the trees [2,3,4].

Neophyllobius is the largest genus of the family Camerobiidae, having 139 species to date [6,7,8,9]. Berlese [1] erected the genus Neophyllobius with the type species N. elegans and designated it to the family Tetranychidae. Later, this genus was transferred to different families, including Stigmaeidae [10], Raphignathidae [11] and Calligonellidae [12,13], mainly due to the misinterpretations of mouthparts and stylophore. Southcott [4] identified the uniqueness of the genus, erected the new family Neophyllobiidae for Neophyllobius (without a camerostome and mouthparts anterior) and proposed a new family Camerobiidae for the genus Camerobia (with a camerostome and mouthparts inferior). Gerson [14] declared these morphologies as “misinterpretations” and synonymized Neophyllobiidae with the Camerobiidae and gave detailed diagnoses of the two genera included.

Based on the number of setae on tibiae I–IV and the position of two midventral setae on leg tarsi I–II, Bolland [2] erected two genera, Tycherobius and Tillandsobius, in the family Camerobiidae. The type species of the two genera (T. lombardinii and Ti. floridensis, respectively) were transferred from the genus Neophyllobius [2]. The genera Tycherobius and Tillandsobius currently include 25 and 1 species, respectively [7,15]. The systematics of the genus Neophyllobius was intensively studied by Bolland [16], where 50 new species were proposed, and 35 species were redescribed. Up to now, some regional keys of the genus have been published from Iran [17,18,19,20], Turkey [4,21,22] and Mexico [6]. Recently, Nasrollahi et al. [23] and Fan and Walter [3] provided the morphological characters of 25 species of Tycherobius and the camerobiid genera, respectively.

The ventral idiosoma chaetotaxy is mostly fixed and has less taxonomical importance [24]. However, the species of Neophyllobius have been distinguished based on differences in the lengths of coxal setae [16]. In this genus, the setal notation on the ventral idiosoma has been inconsistent in the literature. There are four different kinds of descriptions/illustrations present based on absence of intercoxal setae (3a or 4a), the absence of aggenital setae (ag) and the presence of one or two pairs of genital setae (g or g1–2). Kethley [25] and Fan and Walter [3] made some efforts towards highlighting this confusion. However, to date, the situation remains ambiguous.

The global camerobiid mite fauna, including that of Saudi Arabia (SA), require special consideration. Previously, two camerobiid genera (Neophyllobius and Decaphyllobius) and six species (N. muscantribii Bolland, N. fissus De Leon, N. hispanicus Bolland, N. gonzali Zaher and Gomaa, N. communis Gerson, and D. gersoni Bolland) have been reported from SA [26,27]. In the present research work, a new genus; Monobius Alatawi and Kamran gen. nov. is proposed. The monospecific genus Tillandsobius is synonymized with Tycherobius, raising the number of species to 26 in the latter genus. The comparative morphological characters of the three genera (Monobius, Neophyllobius and Tycherobius) are provided. The genus Neophyllobius is divided into two new subgenera, Neophyllobius Berlese and Monophyllobius Mirza. The inconsistencies in the ventral idiosoma setal notation are discussed. The morphological characters of a few poorly described species of the genus Neophyllobius are added. The present research provides the key to the world species of the genus Neophyllobius. Also, five new records from Saudi Arabia including three species redescriptions were given.

2. Materials and Methods

The camerobiid mites were collected using two different methods: (a) shaking plant foliage on a sheet of white paper, picking the freely moving mites by a camel hair brush, and storing in 1.5 mL vials filled with 70% ethanol, (b) scooping out the soil debris and leaf litter under trees and shrubs and storing it in a labelled plastic bag. The soil debris and leaf litter samples were processed through a Berlese funnel for at least eight hours where the specimens were collected in a water filled plastic bowl placed underneath the funnel. The collected camerobiid mites from both methods were permanently slide mounted using the Hoyer’s medium under a stereomicroscope (Olympus^®, SZX10, Tokyo Japan). The slide mounted specimens were identified under the phase contrast microscope (Olympus^®, BX51, Tokyo, Japan). Different body parts of mites for re–descriptions and illustrations, were pictured with the Auto–montage Software System (SYNCROSCOPY^®, Cambridge, UK) attached to a phase contrast microscope (Leica^®, DM2500, Wetzlar, Germany). Final processing of drawings was done in Adobe Illustrator (Adobe Systems Incorporated, San Jose, CA, USA). The terminology used in this study follows that of Grandjean [28], Bolland [2] and Kethley [25]. All the measurements of morphological characters in the redescriptions are provided as ranges in micrometres. Only the published species descriptions and illustrations were used in the present study to compare morphological differences and variations. The collected specimens from Saudi Arabia were deposited at King Saud University Museum of Arthropods (KSMA, Acarology section), Department of Plant Protection, College of Food and Agriculture Sciences, King Saud University, Riyadh, Saudi Arabia.

3. Results

3.1. Family Camerobiidae Southcott, 1957

Camerobiidae Southcott, 1957: 311 [4].

Neophyllobiidae Southcott, 1957: 311, synonymized by Gerson, 1972: 507 [14].

Type genus: Camerobia Southcott, 1957: 311.

Diagnosis: Idiosoma ovoid or nearly circular, dorsoventrally compressed; some or all legs longer than idiosoma; free leg segments annulated, femora and tibiae longer than other leg segments; genua short often with a whip–like long setae, gnathosoma jointed to idiosoma in an inferior position; palpi weak, short, without tibial claw; peritremes in one or several loops.

3.2. Synonymy of the Genus Tillandsobius

Considering the genera of the family Camerobiidae up to now, three genera i.e., Neophyllobius, Tillandsobius and Tycherobius are differentiated based on the number of setae on tibiae I–IV and the difference in position of the midventral setae on tarsi I–II [2]. As reported in different published literature [3,23,29], we found that the number of setae on tibiae I–IV were variable and not suitable for generic differentiation (Table 1). For instance, the species T. rhytis [29] has an almost similar tibial chaetotaxy to that for the type species of the genus Tillandsobius, Ti. floridensis i.e., 8–7–7–7 vs. 8–7–6–6 (excluding solenidion) [2,23,29,30]. The type species of the genus Tycherobius, T. lombardinii (McGregor), was redescribed by Bolland [2] with tibial chaetotaxy as 9–8–7–7. However, the recent publications reported tibiae I–IV with 8–7–6–6, which is exactly similar to the tibiae I–IV setal count for Ti. floridensis [23,29]. This further represents the variation in this morphological character.

Table 1.

Morphological characters (excluding solenidion) of the three genera (after Fan & Walter, 2011 [3]; Uluçay et al., 2016 [15]; Nasrollahi et al., 2019 [23]).

		Monobius gen. nov. (2 Species)	Tycherobius (26 Species)	Neophyllobius
				Neophyllobius (114)	Monophyllobius (15)
Coxa I		2–3	2–3	2–3	3
Coxa II		1	1	1–2	1
Coxa III		2	2	1–2	2
Coxa IV		2	1–2	1–2	2
Femur I		4	3–4	3–5	3–4
Femur II		3	3	2–4	2–3
Femur III		2–3	1–4	1–3	1–2
Femur IV		2	1–3	1–3	1–2
Genu I		1	1	1–2	1–2
Genu II		1	1	1–2	1–2
Genu III		1	1	1	1
Genu IV		1	1	1	1
Tibiae I		9	8–9	8–10	8–10
Tibiae II		8	7–8	7–9	7–9
Tibiae III		8	6–8	7–9	7–9
Tibiae IV		7	6–7	6–8	6–8
Tarsus I		9	7 or 9 or 10	7–11	10–11
Tarsus II		9	7–10	6/8–11	9–10
Tarsus III		7	7	6–8	7–8
Tarsus IV		7	7	7–8	7
Midventral setae on tarsi I–IV	Number	1–1–1–1	2–2–1–1	2–2–2–2	2–2–2(1)–1
	Position	–	not in a longitudinal line, variously spaced	in a longitudinal line

Additionally, there are two Neophyllobius species, N. fani Doğan and Ayyildiz [31] and N. succineus Bolland and Magowski [32], which have almost the same number of setae on tibiae I–IV as in Ti. floridensis (McGregor) [2] (8–7–7–6 vs. 8–7–6–6). Furthermore, there are three Neophyllobius species, N. podocarpi Bolland [16], N. nemoralis Kuznetsov and Livshits [33] and N. parthenocissi Bolland [16], which have the same number of setae on tibiae I–IV as in most of species of the genus Tycherobius (9–8–7–7 vs. 9–8–7–7) (Table 1) [15].

These three genera were also differentiated based on the position of midventral setae on tarsi I–II. The genus Neophyllobius has one or two midventral setae on tarsi I–IV, if two setae are present, they are in a longitudinal line. While the genera Tillandsobius and Tycherobius always have two midventral setae on tarsi I–II, consistently not in a longitudinal line and variously spaced (Table 1). In this aspect, the genera Tillandsobius (one species) and Tycherobius (25 species) are closely related.

Based on the evidence provided above, the number of setae on tibiae I–IV does not represent a strong morphological character to differentiate the three genera Neophyllobius, Tillandsobius and Tycherobius. However, the number and position of the mid–ventral setae on tarsi I–II, remain a constant and persistent generic diagnostic character. Therefore, the monospecific genus Tillandsobius is hereby synonymized with the genus Tycherobius. In addition, we propose a new genus, Monobius Alatawi and Kamran gen. nov., for the two species (one midventral seta on all leg tarsi along with a proximal solenidion) namely, M. meyerae (Bolland) and M. electrus (Żmudziński), described originally in the genus Neophyllobius. The diagnoses of the genera Neophyllobius and Tycherobius are modified and provided below. The morphological characters of these three genera, including the new genus, are summarized in Table 1.

3.3. New Genus Monobius Alatawi and Kamran

urn:lsid:zoobank.org:act:3B429EFF-A148-46A5-8BE4-32127A03E707

Type species: Neophyllobius electrus Żmudziński, 2020:3 [9].

Diagnosis: Leg tarsi I–II with one mid-ventral seta present on distal half and a proximal solenidion, tarsi I–IV with 9–9–7–7 tactile setae.

Remarks: The new genus Monobius, is morphologically closer to the genera Neophyllobius and Tycherobius (based on proximal solenidion on leg tarsi I–II) and distinct from Acamerobia, Camerobia, Bisetolobius and Decaphyllobius (based on the distal solenidion on leg tarsi I–II). It can be further distinguished from Neophyllobius and Tycherobius due to the presence of one midventral seta distally on leg tarsi I–II vs. two midventral setae on leg tarsi I–II in later two genera.

Etymology: The generic epithet is derived from the diagnostic character of one midventral seta on all leg tarsi (mono = one)

The new genus Monobius includes two species M. meyerae [16] and M. electrus [9]. Both species were originally described in the genus Neophyllobius.

3.3.1. Monobius meyerae (Bolland) comb. nov.

Neophyllobius meyerae Bolland, 1991:63 [16].

Remarks: The species M. meyerae (Bolland) is referred to the new genus Monobius due to presence of one midventral seta on all leg tarsi. The species can be differentiated from the second species of the genus M. electrus (Żmudziński) based on number of setae on femur III (2 vs. 3), number of dorsal body setae (15 vs. 14) and state of pdx setae (present vs. absent).

Distribution: South Africa

3.3.2. Monobius electrus (Żmudziński) comb. nov.

Neophyllobius electrus Żmudziński, 2020:3 [9].

Remarks: The species M. electrus (Żmudziński) is referred to the new genus Monobius due to presence of one midventral seta on all leg tarsi.

Distribution: Fossil preserved in Baltic Amber, Poland

3.4. Genus Tycherobius Bolland

Tycherobius Bolland, 1986: 205 [2].

Tillandsobius Bolland, 1986: 205; synonym nov.

Diagnosis: Two midventral setae on tarsi I–II, not present in a longitudinal line,

Remarks: The monospecific genus Tillandsobius was erected by Bolland [2] and it was distinguished from Tycherobius only by difference in number of setae on tibiae I–IV. As mentioned earlier, tibial setal counts are variable among the species of the genus Tycherobius and cannot be considered as a generic diagnostic character (Table 1).

Tycherobius floridensis (Bolland) comb. nov.

Tillandsobius floridensis Bolland, 1986:205 [2].

Neophyllobius floridensis McGregor, 1950:61 [10].

Remarks: The species T. floridensis resembles all 25 species of the genus based on two midventral setae on leg tarsi I–II not in a longitudinal line and leg tarsi III–IV, each, always with one midventral seta. It is closely related to the species T. rhytis based on tibiae I–II with 8–7 setae. However, it differs from later due to differences in number setae on tibiae III–IV (6–6 vs 7–7), tarsi I–IV (10–10–7–7 vs. 7–7–8–8), femur II–III (3–2 vs. 4–3) and differences in length of dorsal body setae (less than half the distance to the setae next in line vs. reaching to the base of setae next in line).

Distribution: Florida, USA

3.5. Genus Neophyllobius Berlese

Neophyllobius Berlese, 1886:19 [1].

Type species: Neophyllobius elegans Berlese, 1886:19 [1].

Diagnosis: Two midventral setae on tarsi I–II, present in a longitudinal line.

3.6. Subgenera in the Genus Neophyllobius

Among the species of the genus Neophyllobius, 114 species have two midventral setae on all leg tarsi. In contrast, 14 species have one midventral seta on leg IV and seven species with no such information available [6,8,9,16]. In the present study, the genus Neophyllobius is categorized in two new subgenera; Neophyllobius Berlese and Monophyllobius Mirza, based on the number of midventral setae on leg tarsi III–IV.

3.6.1. New subgenus Neophyllobius Berlese

urn:lsid:zoobank.org:act:D34B5F98-1F09-45C8-8C64-62CA683C154E

Type species: Neophyllobius elegans Berlese, 1886:19 [1].

Diagnosis: Leg tarsi III–IV always with two midventral setae

Number of species included: 114

3.6.2. New subgenus Monophyllobius Mirza

urn:lsid:zoobank.org:act:8648D2D3-7F38-442A-8C15-680FB8D560A7

Type species: Neophyllobius texanus McGregor, 1950:66 [10].

Diagnosis: Leg tarsus III often and tarsus IV always with one midventral seta

Etymology: The subgeneric epithet refers to the presence of one midventral seta on leg tarsi III and IV (mono = one)

Number of species included: 14

3.7. Redescriptions

The present research reported five new records of camerobiid mites from Saudi Arabia, viz; N. combreticola, N. lorestanicus, N. denizliensis, T. emadi and C. southcotii. The species N. combreticola along with two previously reported species, viz; N. muscantribii and N. fissus are redescribed in detail. In addition, the species N. lorestanicus and N. denizliensis were previously misidentified as N. communis and N. hispanicus, respectively.

3.7.1. Neophyllobius combreticola Bolland

Neophyllobius combreticola Bolland, 1991:196 [16]; Beyzavi et al., 2013:393 [34].

Redescription (Figure 1, Figure 2, Figure 3, Figure 4, Figure 5, Figure 6, Figure 7, Figure 8 and Figure 9)

Figure 1.

Neophyllobius combreticola. Female. Gnathosoma. Scale bar: 20 µm.

Figure 2.

Neophyllobius combreticola. Female. Palp. Scale bar: 10 µm.

Figure 3.

Neophyllobius combreticola. Female. Dorsum. Scale bar: 100 µm.

Figure 4.

Neophyllobius combreticola. Female. Venter. Scale bar: 100 µm.

Figure 5.

Neophyllobius combreticola. Female. Genital and anal region. Scale bar: 20 µm.

Figure 6.

Neophyllobius combreticola. Female. Genu I. Scale bar: 50 µm.

Figure 7.

Neophyllobius combreticola. Female. Genu II. Scale bar: 50 µm.

Figure 8.

Neophyllobius combreticola. Female. Genu III. Scale bar: 50 µm.

Figure 9.

Neophyllobius combreticola. Female. Genu IV. Scale bar: 50 µm.

Female (n = 3)

Gnathosoma (Figure 1 and Figure 2): 59–64 long, subcapitulum (Figure 1) with a subcapitular seta m 20–23 and two pairs of adoral setae Or1 7–9 and Or2 8–9, these three pairs simple and slender, chelicerae 24–28 long, palp five–segmented (Figure 2) with the following chaetotaxy: trochanter without setae; femora with two serrated setae, d 9–11 and l’ 34–38, genua with one long, slender, simple dorsal seta d 35–38 (Figure 2), tibiae with three tactile setae (l’, l” and d) and one sword-like seta, tarsus with two eupathidia (acmζ and sulζ), two simple setae (ba and va) and one small solenidion (ω) (Figure 2).

Dorsum (Figure 3): 369–378 long (excluding gnathosoma), integument transversely striated between all dorso–central setae, with 15 pairs of finely serrated setae set on small tubercles, all dorso–central setae longer than distance to the setae next in-line, two pairs of eyes positioned between setae sci and sce. Length of setae: vi 60–63, ve 65–71, sci 64–67, sce 66–70, pdx 75–82, c1 85–89, c2 64–71, d1 80–86, d2 60–65, e1 78–84, e2 59–65, f1 76–81, f2 45–49, h1 40–47, h2 34–38. Distances between setae: pdx–pdx 24–26, c1–c1 16–19, d1–d1 14–17, e1–e1 10–13, f1–f1 11–13, h1–h1 8–10, pdx–c1 20–24, c1–d1 63–67, d1–e1 54–61, e1–f1 50–55, f1–h1 70–73, pdx–d1 84–90, c1–e1 119–127, d1–f1 110–120, e1–h1 120–125.

Venter (Figure 4 and Figure 5): Ventral idiosoma, striated longitudinally between coxae I–IV, coxal setae slender serrate, intercoxal setae 1a present on the coxa I, coxa I grouped with coxa II and coxa III with IV but not completely fused (Figure 4). Length of setae: 1b 22–26, 1c 50–54, 2c 42–45, 3b 32–36, 3c 42–44, 4b 15–19, 4c 23–25, intercoxal setae length: 1a 26–28, 3a 33–38, 4a 15–17, one pair of aggenital setae (ag) present, genito-anal valves with a pair of genital setae (g) and three pairs of anal setae ps1, ps2 and ps3 (Figure 5).

Legs (Figure 6, Figure 7, Figure 8 and Figure 9): Slender and long, lengths (excluding coxae and including ambulacra): leg I 568–572, leg II 500–510, leg III 510–518, leg IV 570–576. Leg segment chaetotaxy (solenidia in parenthesis) as follows: coxae 3–1–2–2, leg segment chaetotaxy (solenidia in parenthesis) as follows: coxae: 3–1–2–2, trochanters 1–1–1–1, femur 4–3–2–2, genu 1(κ)–1(κ)–1–1, tibiae 9(φ)–8(φ)–8(φ)–7(φ), tarsi 10(ω)–10(ω)–8–8. All tarsi with ambulacrum bearing a pair of claws and an empodium with two rows of tenent hairs, all genu setae long and whip like, dorsal seta on genu I 290–300 (Figure 6), genu II 300–305 (Figure 7), genu III 336–342 (Figure 8), genu IV 358–360 (Figure 7). All leg tarsi with two in-line midventral setae and tarsi I–II with a basal solenidion.

Remarks: The species Neophyllobius combreticola is a new species record for the camerobiid mite fauna of Saudi Arabia. It belongs to the Neophyllobius subgenus nov. based on presence of two in-line midventral setae on tarsi I–IV. The specimens are almost similar to the original description [16] except all dorsal body setae are longer (5–10 µm) in the current collection. Furthermore, Alatawi and Kamran [27] reported N. hispanicus, which was a misidentification of this redescribed species.

Material Examined: Two females, unidentified plant, Al–Bashyer, Asir, SA, 19°15.884′ N, 42°05.261′ E, 29 October 2019, coll. M. Kamran and H. M. S. Mushtaq; one female, Acacia spp., Al–Baha, SA, 20°12.653′ N, 41°37.970′ E, 27 October 2019, coll. M. Kamran and H. M. S. Mushtaq.

Previous Distribution: Iran, South Africa

3.7.2. Neophyllobius fissus de Leon

Neophyllobius fissus de Leon, 1967:31 [35]; Bolland, 1991:212 [16].

Redescription (Figure 10, Figure 11, Figure 12, Figure 13, Figure 14, Figure 15, Figure 16 and Figure 17)

Figure 10.

Neophyllobius fissus. Female. Gnathosoma. Scale bar: 20 µm.

Figure 11.

Neophyllobius fissus. Female. Dorsum. Scale bar: 100 µm.

Figure 12.

Neophyllobius fissus. Female. Venter. Scale bar: 100 µm.

Figure 13.

Neophyllobius fissus. Female. Genital and anal region. Scale bar: 20 µm.

Figure 14.

Neophyllobius fissus. Female. Genu I. Scale bar: 50 µm.

Figure 15.

Neophyllobius fissus. Female. Genu II. Scale bar: 50 µm.

Figure 16.

Neophyllobius fissus. Female. Genu III. Scale bar: 50 µm.

Figure 17.

Neophyllobius fissus. Female. Genu IV. Scale bar: 50 µm.

Female (n = 3)

Gnathosoma (Figure 10): 51–55 long, subcapitulum with a subcapitular seta m 24–25 long and two pairs of adoral setae Or1 7 and Or2 8, these three pairs simple and slender, chelicerae 21 long, palp five-segmented with the following chaetotaxy: trochanter without setae; femora with two serrated setae, d 22–23 and l′ 31–34, genua with one long, slender, simple dorsal seta d 34–35, tibiae with three tactile setae (l′, l” and d) and one sword–like seta, tarsus with two eupathidia (acmζ and sulζ), two simple setae (ba and va) and one small solenidion (ω).

Dorsum (Figure 11): 260–271 long (excluding gnathosoma), integument transversely striated between all dorso-central setae, with 15 pairs of finely serrated setae set on small tubercles, all dorso-central setae longer than distance to the setae next in-line, two pairs of eyes positioned between setae sci and sce. Length of setae: vi 49–51, ve 46–48, sci 40–44, sce 36–40, pdx 40–45, c1 52–54, c2 15–19, d1 59–63, d2 41–45, e1 78–84, e2 46–48, f1 63–67, f2 45–47, h1 28–32, h2 35–39. Distances between setae: pdx–pdx 14–16, c1–c1 11–13, d1–d1 9–10, e1–e1 6–7, f1–f1 6–8, h1–h1 8–10, pdx–c1 20–22, c1–d1 55–57, d1–e1 40–44, e1–f1 44–48, f1–h1 38–41, pdx–d1 78–83, c1–e1 99–103, d1–f1 85–90, e1–h1 83–85.

Venter (Figure 12 and Figure 13): Ventral idiosoma, striated longitudinally between coxae I–IV, coxal setae slender serrate, intercoxal setae 1a present on the coxa I, coxa I grouped with coxa II and coxa III with IV but not completely fused. Length of setae: 1b 40–43, 1c 73–75, 2c 57–62, 3b 30–33, 3c 63–67, 4b 10–11 (curved dorsally), 4c 43–44, intercoxal setae length: 1a 32–35, 3a 34–46, 4a 25–27 (Figure 12), one pair of aggenital setae (ag) 14–15, genito-anal valves with a pair of genital setae (g) 10–13 and three pairs of anal setae ps1, ps2 and ps3 (Figure 13).

Leg (Figure 14, Figure 15, Figure 16 and Figure 17): Slender and long, lengths (excluding coxae and including ambulacra): leg I 536–540, leg II 450–460, leg III 485–493, leg IV 529–534. Leg segment chaetotaxy (solenidia in parenthesis) as follows: coxae 3–1–2–2, leg segment chaetotaxy (solenidia in parenthesis) as follows: coxae: 3–1–2–2, trochanters 1–1–1–1, femur 4–3–2–2, genu 1(κ)–1(κ)–1–1, tibiae 9(φ)–8(φ)–8(φ)–7(φ), tarsi 10(ω)–10(ω)–8–8. All tarsi with ambulacrum bearing a pair of claws and an empodium with two rows of tenent hairs, all genu setae long and whiplike, dorsal seta on genu I 218–220 (Figure 14), genu II 233–237 (Figure 15), genu III 313–318 (Figure 16), genu IV 377–382 (Figure 17). All leg tarsi with two in-line midventral setae and tarsi I–II with a basal solenidion.

Remarks: The species Neophyllobius fissus was previously reported from Saudi Arabia [27]. It belongs to the Neophyllobius subgenus nov. based on the presence of two in-line midventral setae on tarsi I–IV. The specimens are almost similar to what was described in the original description [16] except some variations in the length of dorsal body setae.

Material Examined: Two females, date palm (Arecaceae), Al–Qatif, SA, 26°34.36.7′ N, 49°59.07.1′ E, 25 January 2014 and 11 December 2013, coll. Kamal Alsahwan; one female, date palm (Arecaceae), Al–Imam University, Riyadh, SA, 24°48.785′ N, 46°42.142′ E, 19 March 2011, coll. Kamal Alsahwan.

Previous Distribution: Trinidad

3.7.3. Neophyllobius muscantribii Bolland

Neophyllobius muscantribii Bolland, 1991:73 [16].

Neophyllobius eragrostidis Bolland, 1991:73 [16]; Eragrostis (plant), syn. by du Toit, Theron and Ueckermann, 1998 [5].

Redescription (Figure 18, Figure 19, Figure 20, Figure 21, Figure 22, Figure 23, Figure 24 and Figure 25)

Figure 18.

Neophyllobius muscantribii. Female. Gnathosoma. Scale bar: 20 µm.

Figure 19.

Neophyllobius muscantribii. Female. Dorsum. Scale bar: 100 µm.

Figure 20.

Neophyllobius muscantribii. Female. Venter. Scale bar: 100 µm.

Figure 21.

Neophyllobius muscantribii. Female. Genital and anal region. Scale bar: 20 µm.

Figure 22.

Neophyllobius muscantribii. Female. Genu I. Scale bar: 50 µm.

Figure 23.

Neophyllobius muscantribii. Female. Genu II. Scale bar: 50 µm.

Figure 24.

Neophyllobius muscantribii. Female. Genu III. Scale bar: 50 µm.

Figure 25.

Neophyllobius muscantribii. Female. Genu IV. Scale bar: 50 µm.

Female (n = 3)

Gnathosoma (Figure 18): 54–60 long, subcapitulum with a subcapitular seta m 12–17 and two pairs of adoral setae Or1 6–7 and Or2 7–8, these three pairs simple and slender, chelicerae 14–16 long, palp five-segmented with the following chaetotaxy: trochanter without setae; femora with two serrated setae, d 8–11 and l’ 23–25, genua with one long, slender, simple dorsal seta d 20–22, tibiae with three tactile setae (l’, l” and d) and one sword-like seta, tarsus with two eupathidia (acmζ and sulζ), two simple setae (ba and va) and one small solenidion (ω).

Dorsum (Figure 19): 355–361 long (excluding gnathosoma), integument transversely striated between all dorso-central setae, with 15 pairs of finely serrated setae, broadly lanceolate and set on small tubercles, almost all dorso-central setae shorter than distance to the setae next in-line, two pairs of eyes positioned between setae sci and sce. Length of setae: vi 50–53, ve 35–38, sci 33–37, sce 37–41, pdx 30–35, c1 28–30, c2 56–58, d1 33–37, d2 36–41, e1 50–56, e2 35–40, f1 55–57, f2 32–34, h1 27–28, h2 24–27. Distances between setae: pdx–pdx 15–18, c1–c1 20–21, d1–d1 18–20, e1–e1 15–20, f1–f1 12–16, h1–h1 14–15, pdx–c1 43–48, c1–d1 53–58, d1–e1 58–60, e1–f1 47–50, f1–h1 56–58, pdx–d1 100–105, c1–e1 118–119, d1–f1 110–111, e1–h1 100–108.

Venter (Figure 20 and Figure 21): Ventral idiosoma, striated longitudinally between coxae I–IV, coxal setae slender serrate, intercoxal setae 1a present on the coxa I, simple, hair like (Figure 20). Length of setae: 1b 23–24, 1c 50–53, 2c 38–40, 3b 15–17, 3c 40–42, 4b 10–12, 4c 21–22, intercoxal setae length: 1a 15–17, 3a 25–27, 4a 12–14, one pair of aggenital setae (ag) 14–15 present, genito–anal valves with a pair of genital setae (g) 7–9 and three pairs of anal setae (ps1, ps2 and ps3) (Figure 21).

Leg (Figure 22, Figure 23, Figure 24 and Figure 25): Slender and long, lengths (excluding coxae and including ambulacra): leg I 450–452, leg II 388–390, leg III 410–415, leg IV 460–466. Leg segment chaetotaxy (solenidia in parenthesis) as follows: coxae 3–1–2–2, leg segment chaetotaxy (solenidia in parenthesis) as follows: coxae: 3–1–2–2, trochanters 1–1–1–1, femur 4–3–2–2, genu 1(κ)–1(κ)–1–1, tibiae 9(φ)–8(φ)–8(φ)–7(φ), tarsi 10(ω)–10(ω)–8–8. All tarsi with ambulacrum bearing a pair of claws and an empodium with two rows of tenent hairs, dorsal seta on genu I (38–41) twice as long as the segment (Figure 22), seta on genu II (26–28) equal to the segment length (Figure 23), genu III seta (63–65) long, not reaching the first row of setae on tibia III (Figure 24), genu IV seta (118–120) also, not reaching the first row of setae on tibia IV (Figure 25). All leg tarsi with two in-line midventral setae and tarsi I–II with a basal solenidion.

Remarks: The species Neophyllobius muscantribii was previously reported from Saudi Arabia [27]. It belongs to the Neophyllobius subgenus nov. based on presence of two in-line midventral setae on tarsi I–IV. The specimens are almost similar to the brief original description [16].

Material Examined: 1 female, Grasses (Poaceae), Asir, SA, 19°15.884′ N, 42°05.261′ E, 29 October 2019, coll. M. Kamran and H. M. S. Mushtaq, 2 females, Kavi plant and Sider like, Al–Baha, SA, 19°59.807′ N, 41°25.715′ E, 25 April 2013, coll. Kamal Alsahwan.

Previous Distribution: South Africa

3.8. New Records

3.8.1. Neophyllobius lorestanicus Khanjani, Hoseini, Yazdanpanah and Masoudian

Neophyllobius lorestanicus Khanjani, Hoseini, Yazdanpanah and Masoudian, 2014: 441 [36].

Remarks: The species N. lorestanicus is a new record for the camerobiid mite fauna of Saudi Arabia and no distinct morphological differences were found between Saudi Arabian specimens of N. lorestanicus and the original description. It belongs to the Neophyllobius subgenus nov. based on presence of two in-line midventral setae on tarsi I–IV.

Material Examined: Two females, date palm (Arecaceae), Irqa, Riyadh, SA, 24°41.130′ N, 46°34.550′ E, 11 December 2013, coll. Kamal Alsahwan; one female, Acacia spp, (Fabaceae). Al–Ahsa, SA, 25°55.371′ N, 48°59.037′ E, 22 October 2020, coll. J. H. Mirza, N. A. Elgoni, H. M. Sajid and M. W. Khan.

Previous distribution: Iran

3.8.2. Neophyllobius denizliensis Akyol

Neophyllobius denizliensis Akyol, 2020:88 [8].

Remarks: The species N. denizliensis is a new record for the camerobiid mite fauna of Saudi Arabia. It was previously misidentified as N. hispanicus [27]. It belongs to the Neophyllobius subgenus nov. based on presence of two in-line midventral setae on tarsi I–IV. No morphological differences were found between Saudi Arabian specimens and original description of the N. denizliensis.

Material Examined: One female, Tamarix spp., (Tamaricaceae), Tabuk, SA, 28°30.653′ N, 36°28.168′ E, September 29, 2020, coll. J. H. Mirza, H. M. S. Mushtaq and E. M. Khan; one female, Phoenix dactylifera (Arecaceae), Irqa, Riyadh, SA, 13 December 2020, coll. Kamal Alsahwan; one female, grasses (Poaceae). Al–Imam University, Riyadh, SA, 24°48.785′ N, 46°42.142′ E, 7 April 2016, coll. M. Kamran.

Previous distribution: Turkey

3.8.3. Genus Tycherobius Bolland

Tycherobius Bolland, 1986:205 [16].

Type species: Neophyllobius lombardinii Summers and Schlinger, 1955 [12].

Tycherobius emadi Khanjani, Hajizadeh, Ostovan and Asali Fayaz

Tycherobius emadi Khanjani, Hajizadeh, Ostovan and Asali Fayaz, 2013:134 [37]; Hoseini and Khanjani, 2013:212 [38].

Remarks: The species Tycherobius emadi is first time reported from Saudi Arabia and is morphologically similar to the original description [37].

Material Examined: One female, soil, and leaf debris, Tabuk, SA, 26°58.271′ N, 49°40.220′ E, 19 October 2019, coll. M. Kamran and H. M. S. Mushtaq.

Previous distribution: Iran

3.8.4. Genus Camerobia Southcott

Camerobia Southcott, 1957: 306 [4].

Type species: Camerobia australis Southcott, 1957 [4]

Camerobia southcotti Gerson

Camerobia southcotti Gerson, 1972: 502 [14].

Remarks: The species Camerobia southcotti is a new record for the camerobiid mite fauna in Saudi Arabia and is morphologically similar to the original description [14].

Material Examined: Three females, unidentified wild host plant, Jeddah, SA, 26°58.271′ N, 49°40.220′ E, 25 April 2016, coll. M. Kamran and J. H. Mirza.

Previous distribution: Israel

3.9. Additional Notes

The following 10 species are not included in the key either due to lack of available literature or incomplete descriptions/illustrations. The available but scarce information from different literatures about these species are summarized in the Table 2.

Table 2.

Morphological characters of Neophyllobius species with incomplete information.

Species		vanderwieli	ornatus		mexicanus	summersi	saxatilis	elegans	guajavae	hyderabadensis	Species 1	Species 2
Published Year		1926	1940		1950	1950	1938	1886	2002	1980	2006	2005
Country		Netherland	Australia		Mexico	California	Ireland	Italy	India	India	Yemen	Hungary
Habitat/Host plant		Nest of Talpa europaea.	Apiomorpha galls on Eucalyptus sp.	–	Zanthoxylon sp.	Saltgrass	Lichen covered rocks	–	Psidium guajava	Caryota urens	Malaise Trap	D–Vac Sample
Body	Length	–	250	250	–	–	320	250	240	No Literature/Description Available
	Width	–	175	175	–	–	210	220	178
Number of dorsal setae	mc	6	6 (?)	–	15?	6	6	6	6
	l	9	9 (?)	–		9	9	9	9
Number of setae on leg segments	coxae	3–1–2–2	3–1–?–?	1a+2–1–?–?	–		3–1–2–2	3–1–2–2	–
	trochanter	1–1–1–1	1–1–1–1	1–1–1–1	–	1–1–1–1	1–1–1–1	1–1–1–1	–
	femora	4–3–2–2	4–2?–2?–1?	4–2?–2?–1?	–	4–––	3?–1?–2–2	4–3–2–2	–
	genua	1–1–1–1	1–1–1–1	1–1–1–1	–	1–1–1–1	1–1–1–1	1–1–1–1	–
	tibiae	9–8–8–7	?–8–8–7	?–8–8–7	–	–	?–8–8–7	9–8–8–7	–
	tarsi	10–10–8–8	2–2–2–2	2–2–2–2	–	2–––	?–10–8–8	10–10–8–8	–
Number of setae on palp	trochanter	0	–	0	–	–	0	0	–
	femora	2	2	2	–	3	2	2	–
	genua	1	2	2	–	–	1	1	–
	tibiae	3+1 sword like	2?+1 sword like	2+1 claw	–	–	3+1 sword like	?+?	–
	tarsi	2+2 eup	2+2 eup	2+2 eup	–	–	2+2 eup	?+2 eup	–
Reference		[16]	[16]	[3]	[10,16]	[10,16]	[16]	[16]	[41]	[7]	[48]	[49]

“?” = the original author was not sure of the information and provided no specific details. “–” = unavailable.

• Neophyllobius elegans Berlese

Neophyllobius elegans Berlese, 1886:19 [1]; 1900:288 [39]; Canestrini, 1889:457 [40]; Bolland, 1991:212 [16].

Bolland [16] reported that the type specimen was not available for study. Most of the characters provided by Bolland [16] were based on research papers of Berlese and Canestrini and those are also provided in the Table 2

• Neophyllobius guajavae Chatterjee and Gupta

Neophyllobius guajavae Chatterjee and Gupta, in Gupta, 2002:38 [41].

The description and illustrations provided are very poor and didn’t help to place the species in the diagnostic key provided in present work. Although, the original authors have compared the species with N. natalensis.

Based on the present work, the species N. guajavae belongs to the new subgenus Monophyllobius based on tarsi III–IV, each, with one midventral seta (as only illustrated) and closely resembling N. variegata. The dorso-central setae in both the species are short where c1 is 1/2 and 1/3 the length of c1–d1 in N. guajavae and N. variegata, respectively.

• Neophyllobius hyderabadensis Indra, Rao and Thakur

Neophyllobius hyderabadensis Indra, Rao and Thakur, 1980:121 [42].

The original published description and illustration were not found. The first author contacted Mr. Mahran Zeity who published a new camerobiid species from India, but was not able to get any information about N. hyderabadensis (personal communication). Hence, it was not possible to provide any conclusive remarks on this species.

• Neophyllobius mexicanus McGregor

Neophyllobius mexicanus McGregor, 1950:49 [10]; de Leon, 1958:181 [35]; Bolland, 1991:218 [16].

The original description and illustration provided by McGregor [10] were not enough to compare with other species of the genus. de Leon [35] and Bolland [16] were unable to retrieve the type specimen hence they did not provide any details on species description. McGregor [10] and de Leon [35] mentioned that the dorso-central setae are shorter than the distance between the setae next in line and that genual setae are longer than twice the length of respective genua.

• Neophyllobius ornatus Womersley

Neophyllobius ornatus Womersley, 1940:248 [43]; Bolland, 1991:219 [16]; Fan and Walter, 2011:7 [3].

The status of this species is unresolved. The type specimen is lost as reported by Bolland [16] and Fan and Walter [3]. The original description and illustration provided by Womersley [43] were poor and both the previous references have tried to guess the leg chaetotaxy. Fan and Walter [3] mentioned two midventral setae on all leg tarsi. Bolland [16] suggested that this species was close to N. fissus, N. aegyptium, N. niloticus due to the femur IV with one seta.

Based on available information and classification of species in the present work, N. ornatus could belong to group of eight species (N. fissus, N. aegyptium, N. niloticus, N. lalbaghensis, N. womersleyi, N. bamiensis, N. punctulatus, N. ferrugineus) where all leg tarsi have two mid-ventral setae and femur I–IV with 4–3–2–1 setae.

• Neophyllobius saxatilis Halbert

Neophyllobius saxatilis Halbert, 1923:384 [44]; van Eyndhoven, 1938:25 [45]; Bolland, 1991:214 [16].

van Eyndhoven [45] reported this species with few morphological characters and illustrations. Bolland [16] also provided short description and illustration from a co-type specimen as the type slide was in bad condition.

In the present study, the N. saxatlilis could be placed among 48 species of the new subgenus Neophyllobius having all leg tarsi with two in-line midventral setae, femur I–IV with 4–3–2–2 setae, palp tarsus having two setae and two eupathidia, one solenidion on leg tarsi I–IV, dorsal setae c1 reaching the bases of setae d1, setae d1 not reaching the bases of setae f1.

• Neophyllobius summersi McGregor

Neophyllobius summersi McGregor, 1950:67 [10]; Bolland, 1991:218 [16].

McGregor [10] first time described this species with a unique character of palpfemur with three setae. All other Neophyllobius species, to date, do not contain such palp chaetotaxy. Bolland [16] added some other characters and confirmed this unique morphological feature. Although, Zaher and Gomaa [46] considered N. mangiferus to be close to N. summersi and differed from the latter on the basis of length of dorso-central setae and genu I–IV setae. However, it was not possible to include N. summersi, in the present diagnostic key, even with its unique character of palpfemur.

• Neophyllobius vanderwieli Oudemans

Neophyllobius vanderwieli Oudemans, 1926:121 [47]; Bolland, 1991:219 [16].

Bolland [16] mentioned, description based on a male specimen, that there are two solenidion distally on tibia I. We could consider this species close to three other species (N. kamalii, N. karabagiensis and N. sycomorus) which share the same character.

• Neophyllobius sp.

This unidentified species was reported from Yemen by Ueckermann et al. [48]. The authors provided no information (morphological description or illustrations) except the material examined.

• Neophyllobius sp.

This unidentified species was mentioned by Ripka et al. [49], which was collected during a survey of Hungarian mite fauna. The authors suspected this species was near to N. dichantii, but no morphological data was provided

3.10. Ventral idiosoma chaetotaxy

Bolland [2,16] provided a detailed family description and comprehensively reviewed the genus Neophyllobius, with 50 new species and 35 species redescriptions. A detailed genus diagnosis was given including; the ventral idiosoma with three pairs of ventral setae, two pairs of genital setae and three pairs of anal setae. In the remarks, the author stated that pregenital setae were never found in any other species of Neophyllobius, described until that time, in contrast to what was illustrated by Kuznetsov and Livshits [33]; presence of a paired pregenital setae.

It is difficult to discern the ventral idiosomal chaetotaxy from the genus diagnosis provided by Bolland [16] as that huge taxonomic review of the genus lacks ventral morphology of all 85 described and illustrated species. The presence of three pairs of ventral setae, as stated by the author, is confusing because the setae 1a is found on the coxa I in all species of Neophyllobius and it was counted along with coxal setal count. The similar concept is evident in the genus diagnosis provided by Bolland [16] and all the species of Neophyllobius described to date. If the three pairs of ventral setae include the coxal seta 1a, then it supports the absence of aggenital seta (ag) in adult females as reported by Bolland [16]. However, if the three pairs of ventral setae do not include coxal seta 1a, then the total number of setae on the ventral idiosoma according to Bolland [16], add up to five pairs (paired three ventral and two genital setae). This has not been reported in any species of the seven camerobiid genera.

Adding to this confusion, four different ventral idiosoma descriptions and illustrations represented by 55 Neophyllobius, one each of Tycherobius (T. dazkiriensis), Bisetolobius (B. varius) species are available till date. For the ease of understanding, we here consider them as four different ventral idiosoma chaetotaxy (VIC) descriptions. All these cases have a total of four pairs of setae excluding coxal and anal setae. The species representing these cases are also presented in the Table 3. The VIC #1 is represented by 16 species where paired 3a, 4a, ag and g setae are present. The VIC #2 is represented by 10 species where paired 3a is absent. The VIC #3 is represented by 14 species of Neophyllobius and a species, T. dazkiriensis, where paired setae 4a is absent. The VIC #4 includes 13 species of Neophyllobius and a species B. varius, where paired aggenital setae (ag) is absent. It is noteworthy that, in the cases 2–4, where either one of the paired setae was absent, two pairs of genital setae were described and illustrated (Table 3). In addition, the discrepancies were also found between the description and illustration of four species for 3a–4a–ag–g setae including N. bamiensis (0–1–1–2 vs. 1–1–1–1, respectively), N. lorioi (genitalia with two setae vs. no illustration, respectively), N. saberi (0–0–2–2 vs. 0–1–1–2, respectively) and N. zolfigolii (0–1–1–2 vs. 1–1–0–2, respectively).

Table 3.

Ventral idiosoma setal notation described and illustrated in 55 species of Neophyllobius.

Species	Ventral Idiosomal Setae				Species	Ventral Idiosomal Setae
	3a	4a	ag	g		3a	4a	ag	g
abiegnus	1	1	1	1	lamimani	1	1	1	1
afyonensis	1	1	1	1	lorestanicus	0	1	1	2
asalii	0	1	1	2	mamaneae	1	1	1	1
askalensis	1	1	0	2	mangiferus	1	1	0	2
astragalusi	1	0	1	2	mitrae	0	1	1	2
ayvalikensis	1	0	1	2	olurensis	1	1	0	2
ayyildizi	1	0	1	2	orhani	1	1	0	2
bamiensis	0	1	1	2	ostovani	0	1	1	2
bisetalis	1	1	1	1	parisianus	1	0	1	2
bolvadinensis	1	0	1	2	parthenocissi	1	1	1	1
camelli	1	0	1	2	persiaensis	1	0	1	2
cibyci	1	1	1	1	pistaciae	1	0	1	2
communis	1	1	0	2	podocarpi	1	1	0	2
consobrinus	1	1	1	1	populus	1	1	0	2
crinitus	1	0	1	2	punctulatus	1	1	0	2
demirsoyi	1	0	1	2	quercus	1	1	1	1
denizliensis	1	1	1	1	saberi	0	0	2	2
dogani	0	1	1	2	saxatilis	1	1	–	–
edwardi	0	1	1	2	seemani	0	1	1	2
euonymi	1	0	1	2	sturmerwoodi	1	1	1	1
fani	1	1	0	2	sultanensis	1	1	1	1
ferrugineus	1	1	0	2	sycomorus	1	1	0	2
foliosetosus	1	1	0	2	tepoztalensis	1	1	1	1
gonzali	1	1	0	2	tescalicola	1	1	1	1
izmirensis	1	0	1	2	womersleyi	1	1	1	1
karabagiensis	1	1	1	1	yunusi	1	0	1	2
lachishensis	1	0	1	2	zolfigolii	0	1	1	2
lalbaghensis	1	1	1	1

N. bamiensis: illustration depicts presence of paired 3a, 4a, ag and g setae. N. saberi: illustration depicts presence of paired 4a, ag and g1–2 setae N. zolfigolii: illustration depicts presence of paired 3a, 4a and g1–2.

In the genus Neophyllobius, few descriptions are available for immature stages, of which most are poor, which makes it further difficult to understand the setal ontology. Paredes–León et al. [6] studied the idiosomal and leg setal ontogeny for the species N. cibyci. It was reported that on the ventral idiosoma, the intercoxal seta 3a is present in larval stages where the setae 4a appear in protonymphs and aggenital setae (ag) is only present in adult females with one pair of genital (g) setae. The redefinition of the family Camerobiidae provided by Fan and Walter [3], with notes on idiosomal chaetotaxy, also stated setae 4a, ag and g present in females.

In its support, the species of closely related genus of Neophyllobius, the Tycherobius (other than the exception mentioned above) have similar situation as that presented in VIC#1. The females of the other two genera, Camerobia (Type genus; six species) and Acamerobia (one species) also follow the similar ventral idiosoma setal notation.

In the present and previous studies from Saudi Arabia [26,27], eight species from four genera, viz; Camerobia, Decaphyllobius, Neophyllobius and Tycherobius have been reported. As a case study, the ventral idiosoma of all these species are presented (Figure 4, Figure 12, Figure 20, Figure 26,Figure 27,Figure 28,Figure 29,Figure 30a,b). Consistent in females of these species, was the presence of four pairs of ventral setae, excluding 1a, and three pairs of anal setae. The longitudinal striations were present from the level of coxa I till coxa IV. The first pair of ventral setae, present on small platelets, appears in between the coxal setae 3b and 3c, where longitudinal striations curve to become transverse. The second pair of ventral setae, also present on small platelets, appears after that distinct transverse striation pattern, more or less at the level of coxal seta 4c. The position of third pair of setae was variable. Although always anterior to the genital shield, this setal pair was found on or off the anterior margin of genital shield (Figure 4, Figure 12, Figure 20, Figure 26,Figure 27,Figure 28,Figure 29,Figure 30a,b). The fourth pair of setae was always found on the flaps of genital shield.

Figure 26.

Camerobia southcotti. Female. Venter.

Figure 27.

Decaphyllobius gersoni. Female. Venter.

Figure 28.

Neophyllobius lorestanicus Female. Venter.

Figure 29.

Neophyllobius denizliensis. Female. Venter.

Figure 30.

(a,b) Tycherobius emadi. Female. Venter.

Based on the understanding from all the published literature on the family Camerobiidae and after carefully observing the relative position of ventral setae in Saudi Arabian camerobiid species, we reach the conclusion that four pairs of setae were present on the ventral idiosoma, excluding setae 1a and three pairs of anal setae (ps1–3); which were two pairs of intercoxal setae 3a–4a always present on small platelets, a pair of aggenital seta (ag), present on ventral integument and on or off the anterior margin of genital shield and a pair of genital setae (g) present on sides of genital opening. We support this notation based on the evidence discussed above and recommend future works to follow it.

3.11. Key to World Species of the Genus Neophyllobius (Modified after Bolland 1991)

• 1 Leg tarsi III–IV always with two midventral setae………………new subgenus Neophyllobius……15

• 1‘Leg tarsus III with one or two and tarsus IV always with one midventral seta ………………new subgenus Monophyllobius.……2

• 2 Tarsi III–IV with one midventral ……………………………………………N. variegata Fan and Walter

• 2‘Tarsus III with two and tarsus IV with one midventral setae…………………………………………3

• 3 Femur I–III with 4–3–2 setae ……………………………………………………………………………..4

• 3‘ Femur I–III with 3–2–1 setae ……………………………………………………………………………14

• 4 Femur IV with one seta ………………………………………………………………………N. orhani Doğan and Ayyildiz

• 4‘ Femur IV with two setae ………………………………………………………………………………5

• 5 Dorsal striations typically hooked between c1–d1 and d1–e1 …………………………………………….N. interruptus Bolland

• 5‘ Dorsal striations not hooked between c1–d1 and d1–e1………………………………………………6

• 6 Dorsal seta d1 the longest setae ……………………………………………………………………N. dichantii Bolland

• 6‘ Dorsal seta d1 not the longest setae ………………………………………………………………………………7

• 7 Dorsal setae, both e1 and f1, the longest setae …………………………………………………………………8

• 7‘ Either of the dorsal setae e1 or f1 the longest setae ………………….………………………………………………9

• 8 Tarsus II with 10(ω) setae ………………………………………………………………………N. yunusi Bolland *

• 8‘ Tarsus II with 9(ω) setae …………………………………………………………N. fani

• 9 Dorsal setae e1 the longest setae ………………………………………………………………10

• 9‘ Dorsal setae f1 the longest setae ………………………………………………………………11

• 10 Genu I–II with 1–1 tactile setae, setae pdx 10 µm long …….…………………………………………….N. panici Bolland

• 10‘ Genu I–II with 2–2 tactile setae, setae pdx 57 µm long….……………………………………………………N. mamaneae Bolland and Swift

• 11 Genu IV setae two times the genu length …………………………………………………N. texanus McGregor

• 11‘ Genu IV setae more than two times the genu length ………………………….………………………………………………………………12

• 12 Dorsal setae d1 longer than setae h1 ………………………….………………………………………………………………N. muscantribii

• 12‘ Dorsal setae d1shorter than setae h1 ………………………….………………………………………………………………………………...13

• 13 Five pairs of dorso–central setae, pdx absent, second seta on femur II is the shortest …………………………………………… N. quinquepilis Bolland

• 13‘ Six pairs of dorso–central setae, pdx present, second seta on femur II is the longest …………………………………………………N. graminicola Bolland

• 14 Dorsal setae d1 set on strong tubercles and much longer than e1 and f1, setae h1 based close to f1 …………………………………N. bialagorensis Bolland

• 14‘ Dorsal setae d1 not on strong tubercles and equal to e1 and f1, setae h1 based far from f1 …………………………………………N. vandebundi Bolland

• 15 Coxae II with two setae……………………………………….…………………………………………………………………..............……16

• 15‘ Coxae II with one seta ……………………………………………………………………………….....................……………………………...17

• 16 Coxae III–IV each with two setae; genu I–II setae long, reaching half the length of respective tibiae ………………………N. bisetalis Bolland and Swift +

• 16‘ Coxae III–IV each with one setae; genu I–II setae short, less than half the length of respective tibiae ……………………………N. spatulus De Leon +

• 17 Femur I with 5 setae …………………………………………………………………………………………………N. gonzali Zaher and Gomaa

• 17‘ Femur I with 3 or 4 setae………………………………………………………………………………………………………………................. 18

• 18 Femur I with 3 setae ……..….….. ………………………………………………………………………………N. crinitus du Toit, Theron and Ueckermann

• 18‘ Femur I with 4 setae …………………………………………………………………………………………………………....................……… 19

• 19 Femur II with 4 setae ……………………………………………………………………………………………………………….................... …20

• 19‘ Femur II with 3 setae …………………………………………………………………………………….....................................................…21

• 20 All dorsal body setae reaching base of setae in line, dorso–central setae d1 reaching base of e1 ………………………………N. quadrisetosus De Leon

• 20‘ All dorsal body setae very long, extending beyond the base of setae next in line, dorso–central setae d1 reaching base of h1 …… N. sultanensis Akyol and Koç

• 21 Femur III with 3 setae ………………………………………………………………………………………………………………..............…22

• 21‘ Femur III with 2 setae ………………………………………………………………………………………………………………..............…33

• 22 Femur IV with 1 or 2 setae ………………………………………………………………………………………………………………..............23

• 22‘ Femur IV with 3 setae ………………………………………………………………………………………………………………..............…24

• 23 Femur IV with 1 seta ……………………………………………………………………………………………………………N. foliosetosus Fan

• 23‘ Femur IV with 2 setae ………………………………………………………………………………………………………………..............…25

• 24 Lateral setae vi long, at least two times of h2, two most proximal setae on femur III on one level ……………………………………………………….... N. ueckermanni Bolland

• 24‘ Lateral setae vi normal, shorter than two times of h2, two most proximal setae on femur III not on one level …………………………………………. N. sanctaeluciae Bolland

• 25 Dorso–central setae longer than interval to setae next behind …..................... ………………………………26

• 25‘ Dorso–central setae just reach or shorter than interval to setae next behind .. …………………………………………29

• 26 Dorsal setae d1 longest setae …………………………….………………………N. trisetosus Bolland

• 26‘ Dorsal setae e1 longest setae ………………………………………………………………………………………………………………..............…27

• 27 Dorsal setae d1 longer than c1, coxal setae different in length ………………. 28

• 27‘ Dorsal setae d1 as long as c1, coxal setae equal in length … N. montanus Bolland

• 28 Genu II and III setae as long as genu, palps small ………. N. capparidis Bolland

• 28‘ Genu II and III setae longer than genu, palps thicker …… N. graminum Bolland

• 29 Dorso–central setae reaching to setae next in line …….. N. glaesus Zmudzinski

• 29‘ Dorso–central setae shorter than the distance between setae next in line....... 30

• 30 Some dorso–central setae shorter than interval to setae next behind ……….. 31

• 30‘ All dorso–central setae shorter than interval to setae next behind ………………………………………………………….... N. bequartiodendri Bolland

• 31 Genu IV seta five times longer than genu and longer than half the length of tibia IV ………………………………………………………………. N. mkuzensis du Toit et al.

• 31‘ Genu IV seta twice as long as genu and less than half the length of tibia IV.. 32

• 32 Dorsal setae c1 and d1 shorter than interval to setae next behind, third and fourth seta on femur I equal in length, distal seta on palpfemur at least two times longer than the proximal seta, coxa I setae nearly equal in length ………… N. gigantorum Bolland

• 32‘ Dorsal setae c1 shorter and d1 longer than interval to setae next behind, third seta on femur I much shorter than fourth seta, distal seta on palpfemur not two times longer than the proximal seta, coxa I setae much different in length N. hypoleanae Bolland

• 33 Femur IV with 2 setae …………………………………………………… 34

• 33‘ Femur IV with 1 seta ………………………………………………….............. 122

• 34 Palptarsus with 1 eupathidion ………………………………………………….. 35

• 34‘ Palptarsus with 2 eupathidia ……………………………………………………. 37

• 35 Palptarsus with 1 seta ………………………….… N. euonymi Bolland and Ripka

• 35‘ Palptarsus with 2 setae …………………………………………………………… 36

• 36 Dorso–central setae d1, e1, f1 equal in length ………………. N. plumifer Bolland

• 36‘ Dorso–central setae d1 the longest dorsal setae ……N. demirsoyi Akyol and Koç

• 37 Palptarsus with 3 setae …………………............................................................... 38

• 37‘ Palptarsus with 2 setae …………………………………………………………… 42

• 38 Genu I–II, each with 1 solenidion ……………………………………………….. 39

• 38‘ Genu without solenidia ………………….…. α N. edwardi Khanjani and Hoseini

• 39 Tibiae II with 9 tactile setae …………………………… N. zolfigolii Khanjani et al.

• 39‘ Tibiae II with 8 tactile setae …………………………………………….………. 40

• 40 Dorso–central setae c1 and d1 equal in length .. N. dogani Khanjani and Hoseini

• 41‘ Dorso–central seta d1 longer than c1 ………........................................................ 41

• 41 Tarsi I–II with 9–8 tactile setae ……………….. N. seemani Khanjani and Hoseini

• 41‘ Tarsi I–II with 10–9 tactile setae ………………………… N. mitrae Khanjani et al.

• 42 Band of coarse striae interrupted and hooked between setae c1 and d1 …………………………………..................................................... N. natalensis Meyer and Ryke

• 42‘ Striae neither interrupted nor hooked between setae c1 and d1 .…………….. 43

• 43 Two solenidia on distal end of tibia I, one solenidion on the distal end of tibiae II–IV …………………………………………………………………………………………….. 44

• 43‘ One solenidion on distal end of tibiae I–IV ……………………………………. 46

• 44 Tarsus II with 9 tactile setae …………………………. N. kamalii Khanjani et al. *

• 44‘ Tarsus II with 10 tactile setae ……………………………………………………. 45

• 45 In males, genu I seta less than fifth the length of tibiae I, coxae I–IV without polygonal dimples …………………………………………. N. karabagiensis Akyol and Koç *

• 45‘ In males, genu I seta less than third the length of tibiae I, coxae I–IV with polygonal dimples …………………………………………… N. sycomorus Zaher and Gomaa β

• 46 Dorsal setae c1 long, passes at least bases of e1 ……………………………….. 47

• 46‘ Dorsal setae c1 just reaching or shorter than the distance to bases of e1 ......... 58

• 47 Dorsal setae e1 longer than c1 ……………………………………………………. 48

• 47‘ Dorsal setae e1 as long as or shorter than c1 …..……………………………….. 52

• 48 Dorsal setae c1 longer than d1 …………… N. nemoralis Kuznetsov and Livshits

• 48‘ Dorsal setae c1 shorter than d1 …………………………………………………... 49

• 49 Setae h1 longer than h2 …………………………………………………………… 51

• 49‘ Setae h1 equal to or shorter than setae h2 ………………………………………. 50

• 50 Tarsus II with 9 tactile setae, dorso–central setae c1, d1, e1 very long > 200 µm in length …………………………………………… N. astragalusi Khanjani and Ueckermann

• 50‘ Tarsus II with 10 tactile setae, dorso–central setae c1, d1, e1 < 200 µm in length ……………………………………………………………….. N. platanobius Bolland

• 51 Tarsus II with 10 setae, tibiae III with 7 setae, setae e1 the longest…………………………………………………………………………… N. podocarpi Bolland

• 51‘ Tarsus II with 9 setae, tibiae III with 8 setae, setae d1 and e1 almost same in length ………………………………………………………………………... N. izmirensis Akyol

• 52 Setae e1 as long as setae c1 ………………………………. N. parthenocissi Bolland

• 52‘ Setae e1 shorter than setae c1…………………………………………………….. 53

• 53 Dorsal setae very small, distal seta on femur I not reaching femur–genu boarder …………………………………………………………………………………………. 57

• 53‘ Dorsal setae thicker, distal seta on femur I easily reaching femur–genu boarder ………………............................................................................................................ 54

• 54 Dorsal setae d1 and e1 unequal in length ………………………………………. 55

• 54‘ Dorsal setae d1 equal in length to setae e1 …………………. N. femoralis Bolland

• 55 Tarsus II with 10 tactile setae ……………………... N. turcicus Koç and Ayyildiz

• 55‘ Tarsus II with 9 tactile setae …………………………………………………….. 56

• 56 Dorso–central setae pdx, unpaired, single, 75 µm long ……………………………………………….….. N. ostovani Khanjani and Hoseini

• 56‘ Dorso–central setae pdx, paired, 88 µm long…………………………………………………N. asalii Khanjani and Ueckermann

• 57 Dorso–central setae c1 not reaching the base of setae f1 … N. tenuipilis Bolland

• 57‘ Dorso–central setae c1 long, reaching the base of setae f1 .………………………………………………………… N. afyonensis Akyol and Koç

• 58 Setae d1 reach or pass bases of setae f1 …………………………………………. 59

• 58‘ Setae d1 do not reach at all to bases of setae f1 …………………………………. 75

• 59 Setae e1 do not reach setae h2 ……………………………………………………. 60

• 59‘ Setae e1 reach or extend beyond the setae h2 ………………………………….. 70

• 60 Setae e1 reach bases of h1 ……………………………............................................. 61

• 60‘ Setae e1 does not reach the bases of h1 ……………............................................. 63

• 61 Genu I–II setae long, extending beyond half the length of respective tibiae ………………………………………………………….. N. populus Akyol and Koç

• 61‘ Genu I–II setae short, reaching less than half the length of respective tibiae .. 62

• 62 Dorsal setae c1 longer than pdx ………………. N. mangiferus Zaher and Gomaa

• 62‘ Dorsal setae pdx and c1 equal in length ………….……… N. theobromae Bolland

• 63 Setae f1 pass easily the bases of setae h1 ………….……………………………. 64

• 63‘ Setae f1 just reach the bases of setae h1 …………………. N. marginatus De Leon

• 64 Setae c1 easily reach bases of setae d1 ………………………………………….. 66

• 64‘ Setae c1 and pdx do not reach bases of setae d1 ……………………………….. 65

• 65 Leg tarsi I–III with 10–10–8 tactile setae, genu IV seta not reaching tibial border ………………………………………………………………….. N. longulus De Leon

• 65‘ Leg tarsi I–III with 9–9–7 tactile setae, genu IV seta long, reaching tibial border …………………………………………. N. persiaensis Khanjani and Ueckermann

• 66 Setae pdx reach bases of setae d1 ………………………………………………… 68

• 66‘ Setae pdx do not reach bases of setae d1 ………………………………………… 67

• 67 Second seta on the femur II the longest, palptarsus without solenidion …………………........................................................................ N. hispanicus Bolland

• 67‘ First and second setae on femur II equal in length, palptarsus with one solenidion ……………………………………………………………………….. N. denizlyensis Akyol

• 68 Genu I–III setae not whip like, not extending beyond half the length of corresponding tibiae ……………………………………………………………………………….. 69

• 68‘ Genu I–III setae whip like, extending till corresponding tarsi border …………………………………………………….. N. bolvadinensis Akyol and Koç

• 69 Setae d1 the longest dorsal body setae ……..………………… N. deleoni Bolland

• 69‘ Setae c2 the longest dorsal body setae.. N. helichrysi du Toit, Theron and Ueckermann

• 70 Most distal seta on femur I longer than the third one ………………………….71

• 70‘ Most distal seta on femur I shorter than the third one …… N. communis Gerson

• 71 Setae c1 and pdx shorter than f1 …………………………………………………. 72

• 71‘ Setae c1 longer than f1 ……………………………………………………………. 73

• 72 Tarsus II with 9 tactile setae ………………………. N. lorestanicus Khanjani et al.

• 72‘ Tarsus II with 10 tactile setae …………………………….. N. lachishensis Bolland

• 73 Dorso–central seta c1, d1 and e1 not equal in length ………………………….. 74

• 73‘ Dorso–central setae c1, d1 and e1 equal in length ………………………………………………… N. pistaciae Bolland and Mehrnejad

• 74 Tarsi I–II with 10 setae each ………………………………..N. levanticola Bolland

• 74‘ Tarsi I–II with 9 setae each …………………………….. N. saberi Ahaniazad et al.

• 75 Third seta on femur I shorter than the fourth seta …………………………….. 76

• 75‘ Third seta on femur I longer than the fourth seta …………………………….. 83

• 76 Most distal seta on femur I not reaching the genu border, pdx reaching the marginal side of the dorsum ……………………………………………………………………… 77

• 76‘ Most distal seta on femur I passing the genu border, pdx not reaching the marginal side of the dorsum ……………………………………………………………………… 78

• 77 All genu setae shorter than the length of respective leg ………………………………...…………………………N. quercus Uluçay and Koç

• 77‘ All genu setae very long, extending beyond the length of respective leg ……………………………………………………………….. N. lamimani McGregor

• 78 Most distal seta on femur I about 3–4 times longer than the third seta .......... 79

• 78‘ Most distal seta on femur I shorter than 3 times the third seta ……………… 82

• 79 Distal seta on femur II shorter than 1/3 of the length of the proximal one …………………………………………………………………….. N. coxalis Bolland

• 79‘ Distal seta on femur II longer than 1/3 of the length of the proximal one ….. 80

• 80 Dorso–central setae pdx and c1 equal in length, shorter than d1, e1 and f1….. 81

• 80‘ Setae c1 the longest dorso–central setae …….. N. olurensis Doğan and Ayyildiz

• 81 Setae c1 reach half to the distance c1–d1, third seta on tibia I close to seta one and two, setae e1 longer than f1 …………………………………………. N. arenarius Bolland

• 81‘ Setae c1 reach till bases of d1, third seta on tibia I far away from seta one and two, setae e1 as long as f1 ………………………………………………. N. oregonensis Bolland

• 82 Setae c1 do not reach the bases of d1, the distance e1–f1 much shorter than normal ………………………………………………………………………. N. danthoniae Bolland

• 82‘ Setae c1 reach the bases of d1, setae e1 not based close to setae f1 ……………………………………………………………….. N. coloradensis Bolland

• 83 Distal femur I seta passes genu border ….............................................................. 84

• 83‘ Distal femur I seta just reaches or not reaching the genu border at all ……… 90

• 84 Third seta of femur I as long as or longer than the fourth seta ……………… 88

• 84‘ Third seta of femur I equal to or more than half the length of the fourth ……85

• 85 Setae d1 longer than e2 ……………………………………………………………. 86

• 85‘ Setae d1 shorter than or as long as e2 …………………………………………… 87

• 86 Second seta on femur I shorter than the third one, femur I setae more lanceolate ………………………………………………………………… N. parisianus Bolland

• 86‘ Second seta on femur I longer than the third one, femur I setae more whip like …………………………………………………………………N. burrellis McGregor

• 87 Setae c1 do not reach bases of d1 …………………………. N. ponderosus Bolland

• 87‘ Setae c1 just reach or pass bases of d1 …………………… N. dickansoni Bolland

• 88 Genu I–III setae equal to the length of segment ……………………………..…….………………………………………………N. succineus Bolland and Magowski

• 88‘ Genu I–III setae longer than the length of segment …………………………. 89

• 89 Genu III setae reaching or just passing first row of tibia setae, distal setae on femur III reach genu border ………………………………………… N. sturmerwoodi Bolland

• 89‘ Genu III setae longer than tarsus border, distal setae on femur III does not reach genu border ……………………………………………………………………N. aesculi Bolland

• 90 None of the dorso–central setae reach bases of the setae next in line, all dorsal and femoral setae short, small and equal in length, genu setae passing tarsus end …………………………………………………………………. N. tamarindi Bolland

• 90‘ Some or all dorso–central setae reach bases of the setae next in line …………91

• 91 Some of the dorso–central setae reach or cross the bases of setae next in line (i.e., e1 or e1 and f1) ….………………………………………………………………………… 92

• 91‘ Nearly all dorsal or dorso–central setae reach the bases of setae next in line . 95

• 92 Setae e1 and f1 only longest dorso–central setae ……………………………… 93

• 92‘ Only setae e1 the longest dorso–central setae ..………………………………... 94

• 93 Tarsi II with 10(+ω) setae, genu IV setae not reaching base of corresponding tarsi ………………………………………………………………………… N. conocarpi Bolland

• 93‘ Tarsi II with 9(+ω) setae, genu IV setae reaching base of corresponding tarsi ………………………………………………………….............N. ayvalikensis Akyol

• 94 Femur setae very short, femur I setae 3 and 4 nearly at one level, coxal setae 1c twice the length of 1b, genus I seta passes tibiae and genu II and IV seta passing tarsi ends ………………………………………………………..…. N. lorioi Smiley and Moser

• 94‘ Femur setae longer, coxal seta 1c nearly as long as 1b, none of the genu setae reaching tibia end ……………………………………………………………. N. acaciae Bolland

• 95 Femur I–IV strongly serrulate at their proximal posterior margins …………………………………………………………………………………N. lobatus De Leon

• 95‘ Femur I–IV not strongly serrulate at their bases ……………………………… 96

• 96 Femur I setae 3 and 4 not based on one or nearly one level ………………….. 97

• 96‘ Femur I setae 3 and 4 based on one or nearly one level ……...........................113

• 97 Genu I–III and femur I–IV setae strongly lanceolate ………………………….. 98

• 97‘ Genu I–III setae not lanceolate …………………………………………………... 99

• 98 Distal setae on femur IV not passing genu border, coxal seta 1c longer than 1b…………………………………………………………………. N. ceratoniae Bolland

• 98‘ Distal setae on femur IV passing genu border, coxal seta 1c shorter than 1b…………………………………………………………………N. lanceolatus Bolland

• 99 Genu I setae not reaching second row of tibia setae ………………………... 100

• 99‘ Genu I setae reaching or longer than second row of tibia setae …………….. 104

• 100 Genu II setae not reaching second row of tibia setae …………………………. 101

• 100‘ Genu II setae reaching second row of tibia setae ……….. N. binisetosus Bolland

• 101 Genu III setae not whip like and plumed at the top ……….. N. atriplicis Bolland

• 101‘ Genu III setae whip like …………………………………………………………. 102

• 102 Genu II setae not reaching the first row of tibiae setae, dorso–central setae d1, e1 and f1 much longer than other dorsal setae, coxal setae 1c much shorter than 1b …… …………………………………………………………………….N. ambulans Meyer

• 102‘ Genu II setae reaching at least the first row of tibiae setae …......................... 103

• 103 Dorso–central setae d1 and e1 much longer than other dorsal setae ……… …………………………………………………. N. cavumarboris Meyer and Ryke

• 103‘ Dorso–central setae c1 longer than other dorsal setae …………………………………………….. N. camelli Khanjani and Ueckermann

• 104 Genu II setae not whip like ….…………………………………………………. 105

• 104‘ Genu II setae whip like ………………………………………………………….. 107

• 105 Proximal setae on femur IV reaching the base of distal setae ......................... 106

• 105‘ Proximal setae on femur IV not reaching the base of distal setae………………………………………………………………N. armeniaca Bolland

• 106 Genu II seta long whiplike, crossing first row of setae on respective tibiae ……………………………………………… N. abiegnus Khaustov and Abramov

• 106‘ Genu II seta short, not reaching first row of setae on respective tibiae …………………………………………………..… N. ayyildizi Koç and Madanlar

• 107 Genu I and II setae reach till second row of tibia setae ……………………… 108

• 107‘ Genu I and II setae are passing tibial border …………………………………. 109

• 108 Dorsal setae small, not pointed, body small, coxa 1 seta 1c:1b = 6:18 …………………………………………………………………… N. iramus De Leon

• 108‘ Dorsal setae more broad and occupied with very strong stings, coxa I seta 1c:1b = 6:17 ………………………………………………………………………… N. equalis De Leon

• 109 Dorsal setae specially spinosed ………………………………………………………………………… N. setosus Bolland

• 109‘ Dorsal setae not specially spinosed ………………………………………………………………………… 110

• 110 First seta on femur II not reaching the third, proximal femur I seta not the longest ……………………………………………………………................................................. 111

• 110‘ First seta on the femur II the longest femur seta which passes easily the third, proximal femur I seta the longest …………………………………………… N. pruni Bolland

• 111 Setae e1 longest among the dorso–central setae……………………………………

• ……………………………………………………N. askalensis Doğan and Ayyildız

• 111‘ Setae e1 not the longest among the dorso–central setae …………………….. 112

• 112 Proximal seta on femur I equal in length with the second, distal seta on femur IV long, passes genu border, first seta on femur I reaches the bases of the fourth ……................................................................................................. N. viticola Bolland

• 112‘ Proximal seta on femur I shorter than the second, distal seta on femur IV short, not passing genu border, first seta on femur I does not reach the fourth seta at all ………………………………………………………………………… N. combreticola

• 113 Dorsal setae broad, setae d1 strong, genu II seta as long as genu, genu I and II setae not reaching first row of tibiae setae ………………………………………………… 114

• 113‘ Dorsal setae smaller, genu I and II setae at least reaching first row of tibiae setae ……………………………………………………………………………………... 115

• 114 Genual setae distinctly spinose …………………………….. N. curtipilis De Leon

• 114‘ Genual setae linear and faintly spinose …………………….. N. spatulus De Leon

• 115 Genu I setae reach till first row of tibiae setae, palp femur swollen, palp genu short ………………………………………………………………………… N. sierrae McGregor

• 115‘ Genu I setae longer, palp femur longer ……………………………………… 116

• 116 Two eupathidia on the palp tarsus based very different in level, dorsal setae small with many spicules ……………………………………………….. N. americanus Banks

• 116‘ Two eupathidia on the palp tarsus on similar level, dorsal setae broader with less spicules …………………………………………………………………………………... 117

• 117 Genu III–IV setae are passing tarsus end ……………………………………… 118

• 117‘ Genu III–IV setae not reaching beyond end of respective legs ……………… 120

• 118 Dorso–central setae pdx and c1 grouped on a small finely striated platelet ……………………………………………………. N. tescalicola Parades–Leon et al.

• 118‘ Dorso–central setae pdx and c1 not grouped on a platelet …………………… 119

• 119 Lengths of dorsal setae c1 and d1 are the same as the distance between setae c1–d1 and d1–e1 respectively ………………………………………………. N. farrieri De Leon

• 119‘ Dorsal setae c1 and d1 are distinctly longer than the distance between setae c1–d1 and d1–e1 respectively ……………………………………… N. cibyci Parades–Leon et al.

• 120 Setae sce shorter sci ……………………………………………………… 121

• 120‘ Setae sce equal in length to sci …………………………ΩN. consobrinus De Leon

• 121 Lateral setae vi, ve and sci subequal in length ……………ΩN. inequalis De Leon

• 121‘ Lateral setae sci distinctly longer than vi and ve ………… ΩN. piniphilus Bolland

• 122 Setae e1 passes the bases of h1 ………….……. N. aegyptium Soliman and Zaher

• 122‘ Setae e1 do not reach the bases of h1 ……………………………………… 123

• 123 Genu I seta short not reaching second row of setae on corresponding tibiae ……………………………………… 124

• 123‘ Genu I seta long, extend beyond corresponding tarsus base ……………… 126

• Tarsi I–II with 8 setae each ……………………………………N. ferrugineus Fan

• 124‘ Tarsi I–II with 10 setae each ……………………………………… 125

• 125 Coxal seta 1c twice as long as coxal seta 1b …. N. lalbaghensis Zeity and Gowda

• 125‘ Coxal seta 1c 1.5 times as long as coxal seta 1b … N. womersleyi Fan and Walter

• 126 Distance e1–f1 two times longer as distance d1–e1, setae f1 the longest ………………………………………………………………… N. niloticus Bolland

• 126‘ Distance e1–f1 sub–equal to d1–e1 ……………………………………………. 127

• 127 Palp femur with both setae short, not reaching end of palp genu; long setae of genu I–III not reaching end of respected legs …………………. N. bamiensis Khanjani et al.

• 127‘ Palp femur with one of two setae reaches end of tarsus ……………………. 128

• 128 Dorsal setae d1 reaching base of e1, coxal setae 1c longer than 2c in length ………………………………………………………………………………... N. fissus

• 128‘ Dorsal setae d1 extend beyond the bases of e1, coxal setae 1c almost equal to 2c in length ……………………………………………………………………N. punctulatus Fan

• * = species described based on male holotype.

• + = two species, viz; N. bisetalis and N. spatulus with two setae on coxae II as mentioned by Bolland and Swift [50] and Bolland [16], respectively.

• α = The character of genu I–IV without solenidion is mentioned in the original description [38].

• β = The male specimens were not reported at the time of original description for N. sycomorus. However, Bolland [16] provided very few morphological characters of N. sycomorus with the illustrations of male and female.

• Ω = These three species have minute differences among them. Bolland and Swift [50] have also questioned the close similarity of N. consobrinus and N. inequalis suggesting later could be a deutonymph of former. However, these species are placed in the diagnostic key based on available information. but types of each species require re–examination.

Author Contributions

Conceptualization, J.H.M. and M.K.; methodology, J.H.M. and M.K.; validation, M.K., F.J.A.; formal analysis, J.H.M., F.J.A. and M.K.; investigation, J.H.M. and M.K.; data curation, J.H.M. and M.K.; writing—original draft preparation, J.H.M.; writing—review and editing, M.K. and F.J.A.; supervision, F.J.A.; funding acquisition, F.J.A. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by Deanship of Scientific Research at King Saud University, through the research project RG–1437–043.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

This published work and the nomenclatural acts it contains have been registered in ZooBank, the online reg-istration system for the ICZN (International Code of Zoological Nomenclature).

Conflicts of Interest

The authors declare no conflict of interest.