Zootaxa 4378 (4): 563–572 ISSN 1175-5326 (print edition)

http://www.mapress.com/j/zt/
Copyright © 2018 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.4378.4.7
http://zoobank.org/urn:lsid:zoobank.org:pub:0AF08ABE-D5F6-4743-9C01-8747E493E7AB

A new species of spider mite, Oligonychus neocastaneae sp. nov.

(Acari: Tetranychidae), from Japan

TEA ARABULI1, 2 & TETSUO GOTOH1, 3

1
Laboratory of Applied Entomology and Zoology, Faculty of Agriculture, Ibaraki University, Ami, Ibaraki 300-0393, Japan
2
Institute of Zoology, Ilia State University, Kakutsa Cholokashvili Ave 3/5, Tbilisi 0162, Georgia
3
Corresponding author. E-mail: tetsuo.gotoh.acari@vc.ibaraki.ac.jp
ORCID: T. Gotoh http://orcid.org/0000-0001-9108-7065

Abstract

A new species Oligonychus neocastaneae sp. nov. is described and illustrated from Castanea crenata Sieb. et Zucc. (Fa-
gaceae). The new species closely resembles Oligonychus castaneae Ehara & Gotoh, 2007, which inhabits the same host
plant, Castanea crenata, but mainly differs by the aedeagus in having a longer distal portion which forms a small sigmoid
and acuminate tip, instead of having a shorter distal portion which ends in a truncate tip. Several differences were also
observed between the two species in the leg setal counts of tarsus II in the male and female; the number of tactile setae
and solenidia (in parentheses) was 11(1) for the female and male in the new species, but in O. castaneae this count is 12(1)
for the female and 12(2) for the male. A maximum likelihood tree based on the cytochrome c oxidase subunit I (COI) gene
of mitochondrial DNA (mtDNA) showed that O. neocastaneae sp. nov. was clearly separated from O. castaneae and other
related species. A key to all species of the genus Oligonychus known in Japan is given.

Key words: Japan, spider mite, Oligonychus, Castanea crenata, Quercus acutissima, Quercus serrata

Introduction

Spider mites are considered as one of the most important phytophagous pests that devastate agricultural crops,
ornamental plants and fruit trees in greenhouses and fields. Migeon & Dorkeld (2006–2017) listed 1305 valid
species in the world. To date there are 206 known species of Oligonychus (Migeon & Dorkeld 2006–2017), and 18
species have been recorded in Japan (Ehara & Gotoh 2009), which is the second largest genus in Japan: the first is
the genus Eotetranychus consisting of 23 species. In the course of our studies on the relationship between
intraspecific variation of the cytochrome c oxidase subunit I (COI) gene of mitochondrial DNA (mtDNA) and
endosymbiotic bacteria infecting Oligonychus castaneae Ehara & Gotoh, 2007, an unknown species that
morphologically closely resembles to O. castaneae was found, but the morphology of its aedeagus and number of
tactile setae of tarsus II in both sexes differ from O. castaneae. After minute morphological investigation, we
describe it as a new species. DNA sequences of the COI gene have been commonly used to identify and separate
species. Here, we examined whether this new species could be distinguished from O. castaneae and other related
species by COI sequences. We also provide a key to the Japanese species of the genus Oligonychus.

Materials and methods

The collected mites were preserved in 70% ethyl alcohol and later mounted in Hoyer’s medium on glass slides for
examination. The specimens were examined using the Olympus® BX 53 differential interference contrast
microscope and illustrated with the Olympus® DP72 digital camera. Measurements were performed using the
imaging software Sensiv Measure® ver. 2.6.0. All measurements are given in micrometers (µm) and correspond to
the holotype followed by minimum and maximum values from paratypes and other specimens. The aedeagal

Accepted by O. Seeman: 18 Dec. 2017; published: 12 Feb. 2018 563

morphological parameters of the new species were compared with O. castaneae based on the parameters provided
by Beard (2008). Setal nomenclature and abbreviations on the body are according to Lindquist (1985). Leg setal
counts are given in the order: coxa, trochanter, femur, genu, tibia and tarsus. Leg tactile and eupathidial setal
numbers are provided first followed by solenidia in parentheses. The holotype and some of the paratypes of the
new species are deposited in the collections of the National Museum of Nature and Science, Tsukuba (NSMT); the
remainder of the paratypes and other specimens are placed in the Laboratory of Applied Entomology and Zoology,
Ibaraki University (AEZ-IU) under the serial voucher specimen numbers.
Molecular analyses were performed for two strains of Oligonychus neocastaneae sp. nov. collected from
Saitama (voucher specimen no. 346) and Tochigi (voucher specimen no. 349; see ‘Other specimens’ section in
detail), Japan. Mites randomly selected from each strain were used for molecular analyses. Total DNA was
extracted from the whole body of each female individual by using PrepMan Ultra Reagent (Applied Biosystems,
Foster City, CA). Primers C1-J-1718 (Simon et al. 1994; 5'-GGAGGATTTGGAAATTGATTAGTTCC-3') and
COI REVA (Gotoh et al. 2009; 5'-GATAAAACGTAATGAAAATGAGCTAC-3') were used for the polymerase
chain reaction (PCR). PCR amplification was performed with the following profile: 3 min at 94°C, followed by 35
cycles of 1 min at 94°C, 1 min at 45°C and 1.5 min at 72°C. An additional 10 min at 72°C was allowed for last
strand elongation. The resultant DNA solutions were purified by using MinElute PCR Purification Kit (Qiagen,
Valencia, CA) and sequenced directly. Sequencing was carried out in both directions using the amplifying primers
with a BigDye Terminator Cycle Sequencing Kit v.3.1 (Applied Biosystems) and on an ABI 3130xl automated
sequencer.
All sequence data obtained were deposited in DDBJ/EMBL/GenBank International Nucleotide Sequence
Databases under the accession number LC341206 (Saitama strain) and LC341207 (Tochigi strain). Sequences
obtained were aligned using CLUSTALW in MEGA7 software (Kumar et al. 2016). For the maximum-likelihood
(ML) analysis, we used the best-fit model (GTR+G model) chosen by MEGA7. ML tree was constructed with
MEGA7. Sequences of the COI gene of mtDNA for Panonychus and Oligonychus species, except for O.
neocastaneae sp. nov., were obtained from previously published data (Matsuda et al. 2012). The robustness of the
branches was tested by bootstrap analysis with 100 replications.

Taxonomy

Family Tetranychidae Donnadieu, 1875

Subfamily Tetranychinae Berlese, 1913

Tribe Tetranychini Reck, 1950

Genus Oligonychus Berlese, 1886

Oligonychus Berlese, 1886: 24; Pritchard & Baker, 1955: 270; Wainstein, 1960: 203; Tuttle & Baker, 1968: 116; Meyer, 1974:
248; Mitrofanov, 1977: 1801-1802.
Type-species: Heteronychus brevipodus Targioni Tozzetti, 1878.

Oligonychus neocastaneae Arabuli & Gotoh sp. nov.

[Japanese name: Nise-kuri-no-tsumehadani]
(Figs. 1–18)

Description. Female (n = 10). Body greenish brown. Length of body including gnathosoma 485–520; gnathosoma
94–107 long; width of body 321–350.
Dorsum. Dorsal body setae long and sparsely setose; lengths of setae: v2 75–86; sc1 110–118; sc2 95–111; c1
99–113; c2 94–108; c3 94–107; d1 98–112; d2 109–117; e1 101–109; e2 106–113; f1 94–99; f2 83–93; h2 50–54.
Dorsal setae not set on tubercles, length of setae longer than distance between their bases; second pair of prodorsal
setae (sc1) longer than first (v2) and third (sc2) pairs. Medial hysterosomal striation with V-shaped striae between

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d1-d1 and e1-e1, small area of longitudinal striae between e1-e1, and transverse striae posterior to e1-e1 (Fig. 1);
lobes on dorsal striae square shaped with rounded corners, broader than tall (Fig. 2).
Gnathosoma. Peritreme straight, distal end enlarged forming a simple bulb sometimes posteriorly directed
(Fig. 6). Palptarsus spinneret about as long as wide: 4.1–4.8 long and 3.6–4.3 wide; dorsal sensillum (solenidion)
slender 5.1–5.7 long; eupathidia ul’ζ and ul”ζ subequal in length 6.1–6.8 long (Fig. 7).
Venter. Genital flap with transverse striae; pregenital striae longitudinal (Fig. 8).

FIGURES 1–2. Oligonychus neocastaneae sp. nov. 1. Female (dorsum), 2. Lobes on dorsal striation.

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FIGURES 3–8. Oligonychus neocastaneae sp. nov., female. 3. Leg I, 4. Leg II 5. Empodium I, 6. Distal part of the peritremes,
7. Distal segments of palpus, 8. Genital flap and anterogenital area.

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FIGURES 9–13. Oligonychus neocastaneae sp. nov., male. 9. Tarsus and tibia I, 10. Tarsus and tibia II, 11. Empodium I, 12.
Distal part of the peritremes, 13. Distal segments of palpus.

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FIGURES 14–18. Oligonychus neocastaneae sp. nov. 15–18. Aedeagi, variations (15. holotype), 14a–d. Measured parameters
for aedeagi comparisons (based on Beard (2008)).

Legs. Leg I 238–255, leg II 195–220, leg III 218–235, leg IV 229–long (from trochanter to tarsus). Lengths of
segments of leg I as follows: femur 77–87, genu 39–46, tibia 40–45, tarsus 72–79 long. Leg setal count as follows
(Figs. 3, 4):
I 2 – 1 – 8 – 5 – 7 + (1) – 11 + (1) + 2 duplexes
II 2 – 1 – 6 – 5 – 5 – 11 + (1) + 1 duplex
III 1 – 1 – 2 – 2 – 5 – 8 + (1)
IV 1 – 1 – 1 – 2 – 5 – 8 + (1)
Tarsus I distal duplex solenidion longer than proximal solenidion: 70–75 and 49–55, respectively. Tactile setae
either of distal and proximal duplexes subequal in length: 14–17 and 17–19, respectively. Tarsus II solenidion of
duplex setae 58–69 long and tactile seta 15–18 (Figs. 3, 4). Tarsus I with 3 tactile setae and 1 solenidion proximal
to proximal set of duplex setae (Fig. 3); tarsus II with 3 tactile setae and 1 solenidion proximal to duplex setae (Fig.
4).
Empodia I–IV uncinate, long, clawlike with 4–5 pairs of proximoventral hairs; empodial claw as long as
proximoventral hairs (Fig. 5).
Male (n = 11). Body greenish brown with pale reddish tint at the tip of prodorsum. Holotype—length of body
including gnathosoma 446 (428–451); gnathosoma 82 (78–94) long; width of body 230 (225–240).
Dorsum. Dorsal body setae long, far longer than distance between bases of consecutive setae, length of setae:
v2 68 (64–76); sc1 97 (84–97); sc2 78 (74–83); c1 81 (76–88); c2 86 (74–86); c3 92 (85–93); d1 84 (76–88); d2 91
(75–91); e1 85 (75–86); e2 90 (82–94); f1 69 (65–70); f2 58 (50–58); h2 34 (29–35).
Gnathosoma. Peritreme as in female (Fig. 12). Palpus with spinneret about as long as wide: 3.7 (3.7–4.1) long
and 3.4 (3.2–3.5) wide; solenidion obviously spindle-shaped 4.1 (4.0–4.3) long; eupathidia ul’ζ and ul”ζ subequal
in length 5.1 (5.0–5.3) long (Fig. 13).
Legs. Leg I 231 (221–235); leg II 185 (184–199); leg III 182 (180–195); leg IV 195 (195–209) long (from
trochanter to tarsus). Length of first leg segments as follows: femur 65 (64–79); genu 41 (38–44); tibia 35 (33–37);
tarsus 72 (71–79) long. Leg setal count as follows (Figs. 9, 10):
I 2 – 1 – 8 – 5 – 7 + (4) – 11 + (3) + 2 duplexes
II 2 – 1 – 6 – 5 – 5 – 11 + (1) + 1 duplex
III 1 – 1 – 2 – 2 – 5 – 8 + (1)
IV 1 – 1 – 1 – 2 – 5 – 8 + (1)
Tarsus I distal duplex solenidion longer than proximal solenidion: 64 (61–65) and 47 (44–52), respectively.
Tactile setae of distal and proximal duplexes subequal in length: 16 (14–16) and 17 (16–18), respectively (Fig. 9).
Tarsus II solenidion of duplex setae 41 (40–46) long, tactile 16 (15–17) (Fig. 10). Tarsus I with 3 tactile setae and 3
solenidia proximal to proximal set of duplex setae (Fig. 9); tarsus II with 3 tactile setae and 1 solenidion proximal
to duplex setae (Fig. 10).

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 Empodia I with 3 pairs of proximoventral hairs, thick, claw-like, the proximoventral spurs much shorter than
dorsal claw and each with 2 fine dorsal hairs (Fig. 11). Empodia II –IV uncinate, long, claw-like with 3 pairs of
proximoventral hairs; empodial claw as long as proximoventral hair (Fig. 11).
Aedeagus. Shaft of aedeagus narrowing, bent ventrad, distally less than at right angle and form a very small
sigmoid, acuminate tip; sigmoid part tapers (Figs. 14–18). Measurements of aedeagus (Fig. 14): a 6.3 (5.8–6.8); b
12.4 (10.5–12.5); c 3.6 (3.4–4.1); d 31.3 (26.7–32.1). Aedeagus measurements (n=6) of O. castaneae (Figs. 19–23;
voucher specimen no. 696) collected from Kanegasaki (39°16’N 141°05’), Isawa Prov., Iwate Pref., 19-VIII-2013
(Y. Kitashima leg.): a 4.3–4.9; b 9.1–11.4; c 3.4–4; d 33.2–35.5.
Type series. Holotype: male (NSMT-Ac 14218), Kanegasaki (39°13’N 141°05’E), Isawa Prov., Iwate Pref., 5-
VII-2009 (N. Nishizawa leg.), on Castanea crenata Sieb. & Zucc. (Fagaceae). Paratypes: 23 males and 27 females
(voucher specimen no. 327) including 4 males (NSMT-Ac 14219-14222) and 5 females (NSMT-Ac 14223-14227)
deposited to NSMT, data same as for holotype.
Other specimens. Remainder of the paratypes and additional specimens are retained in AEZ-IU under the
serial voucher specimen numbers. 4 males and 8 females (voucher specimen no. 325), Tomi (36°21’N 138°20’E),
Nagano Pref., 20-VII-2009 (N. Nishizawa leg.), on Castanea crenata; 5 males and 8 females (voucher specimen
no. 326), Kawachi-Nagano (34°24’N 135°32’E), Ohsaka Pref., 21-VII-2009 (A. Suwa leg.), on Quercus acutissima
Carruth. (Fagaceae); 3 males and 8 females (voucher specimen no. 346), Kumagaya (36°10’N 139°24’E), Saitama
Pref., 16-VIII-2009 (N. Nishizawa leg.), on Quercus serrata Murray (Fagaceae); 4 males and 7 females (voucher
specimen no. 349), Fujioka (36°17’N 139°37’E), Shimotsuga Prov., Tochigi Pref., 16-VIII-2009 (N. Nishizawa
leg.), on Q. acutissima.
Diagnosis. The female of the new species is closely related to O. castanea. Both infest the upper side of leaves
of fagaceous host plants and are brownish green in colour. However, the main morphological character that
separates these two species is the shape and size of the aedeagus. That of the new species has a longer distal portion
that forms a very small sigmoid shape and has an acuminate tip. In contrast, the aedeagus of O. castaneae is bent at
a right angle to the shaft axis and it has a straight, truncate tip bent anteriorly. In addition, the new species is
distinguished from O. castaneae by having 11 tactile setae and one solenidion on tarsus II of the female, instead of
12 tactile setae and one solenidion; and 11 tactile setae and one solenidion on tarsus II of the male, instead of 12
tactile setae and 2 solenidia. The number of tactile setae proximal to the duplex setae on tarsus I is sometimes
useful for distinguishing species. However, females of O. castanea from Ami (the type locality) are characterized
by having tarsus I with 4 tactile setae and 1 solenidion proximal to proximal duplex setae, but tarsus I of female
specimens from other localities have 3 or 4 tactile setae and 1 solenidion proximal to proximal duplex setae. So, the
number of proximal tactile setae on the tarsus I is not useful for distinguishing these species.

FIGURES 19–23. Oligonychus castaneae. 20–23. Aedeagi variations, 19a–d. Measured parameters for aedeagi comparisons
(based on Beard (2008)).

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 This new species resembles to O. newcomeri (McGregor, 1950); however, aedeagus of O. neocastaneae is
distinct from that of O. newcomeri in being widened at the bend with the upper surface straight (Pritchard & Baker
1955).
The new species also closely resembles to O. castaneae, O. gotohi Ehara, 1999, O. amiensis Ehara, 1999, and
O. coffeae (Nietner, 1861) (Ehara & Gotoh 2007). However, the new species can be easily separated from the other
species by using the morphology of aedeagus, because aedeagi of the latter three species have closely similar shape
with O. castaneae. There are other two conspicuous differences between the new species and these four species:
(1) there are four or more proximoventral hairs in the male leg I of the latter four species, but only three in the new
species; and (2) the proximoventral hairs in the male leg I are free, but there are fused (at least partially) in the new
species. The shape of aedeagus of the new species is also closely related to that of O. clavatus (Ehara, 1959), but
the latter differs from the new species in having 6 tactile setae on tibia I instead of 7 in the new species.
Molecular analyses. After alignment, the COI fragment had 714 nucleotide sites, of which 180 were
parsimony informative sites and contained no insertions or deletions. In the COI tree, the taxonomic status of each
species was robustly supported because all eight Oligonychus species analyzed in the current study were
independently clustered (Fig. 24, bootstrap values = 100), meaning that O. neocastaneae sp. nov. was a clearly
separate species from O. castaneae.
Etymology. This species inhabits the same host plant than O. castaneae to which it is morphologically closely
related. As it was discovered after O. castaneae we designated it by the species name neocastaneae.

FIGURES 24. Maximum likelihood (ML) tree based on the COI gene (714 bp) of mtDNA using GTR+G model. Bootstrap
values based on 100 replications are indicated at the nodes. Only bootstrap values >50% are shown. Each operational
taxonomic unit is indicated by accession number and species.

Key to the Japanese species of the genus Oligonychus (mainly based on female characters)

1. Tibia I with 6–7 tactile setae; aedeagus curved ventrad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

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- Tibia I with 9 tactile setae; aedeagus curved dorsad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
2. Tibia I with 6 tactile setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Tibia I with 7 tactile setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3. Dorsal idiosomal setae set on tubercles, longer than intervals between their bases . . . . . . . . . . . . . . . . . . . . . . . . . . pustulosus
- Dorsal idiosomal setae not set on tubercles, mostly shorter than intervals between their bases . . . . . . . . . . . . . . . . . . . clavatus
4. Dorsocentral opisthosomal setae (c1, d1, e1, f1) shorter than distances between their bases . . . . . . . . . . . . . . . . . . . . . . . . . . 5
- Dorsocentral opisthosomal setae about as long as, or longer than distances between their bases . . . . . . . . . . . . . . . . . . . . . . . 6
5. Dorsal idiosomal setae similar in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . karamatus
- Setae v2, sc1 and c3 longer than the other dorsal idiosomal setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hondoensis
6. Dorsocentral opisthosomal setae noticeably longer than longitudinal distance to the bases of consecutive setae . . . . . . . . . . 7
- Dorsocentral opisthosomal setae approximately as long as longitudinal distance to the bases of consecutive setae . . tsudomei
7. Setae f2 and f1subequal in length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
- Seta f2 much shorter than f1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ilicis
8. Palpus with spinneret about 1.5–2.0 times as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
- Palpus with spinneret about as long as wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
9. Tarsus I with only 4 tactile setae proximal to proximal duplex setae; on Camellia spp. . . . . . . . . . . . . . . . . . . . . . . . . camelliae
- Tarsus I with more than 4 tactile setae proximal to proximal duplex setae; on coniferous trees . . . . . . . . . . . . . . . . . . . . . . . 10
10. Tarsus I and II each with 2 tactile seta ventrad of duplex setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . perditus
- Tarsus I and II each with 1 tactile seta ventrad of duplex setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ununguis
11. Tarsus IV with 7 tactile setae and 1 solenidion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gotohi
- Tarsus IV with 8 tactile setae and 1 solenidion; on evergreen plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12. Tarsus II with 3 tactile setae and 1 solenidion proximal to duplex setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
- Tarsus II with 4 tactile setae and 1 solenidion proximal to duplex setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
13. Peritreme hooked distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . coffeae
- Peritreme enlarged, but not hooked distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . amiensis
14. Aedeagus bent at less than a right angle with a very small sigmoid, acuminate tip . . . . . . . . . . . . . . . . neocastaneae sp. nov.
- Aedeagus bent at right angle to shaft axis ending in a truncate tip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . castaneae
15. Aedeagus with obvious terminal knob . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
- Aedeagus without terminal knob . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . orthius
16. Female peritreme hooked distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
- Female peritreme straight distally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . modestus
17. Terminal knob of aedeagus with posterior projection very small . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
- Terminal knob of aedeagus with posterior projection very long, slender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . biharensis
18. Terminal knob of aedeagus forming strong angle with shaft axis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . uruma
- Terminal knob of aedeagus tiny, forming a weak angle with shaft axis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rubicundus

Acknowledgments

We are greatly indebted to N. Nishizawa, A. Suwa, Y. Kitashima, T. Matsuda, R. Suzuki, N. Sakamoto and M. W.
A. A. M. Negm for providing specimens and their kind help with this study. We also wish to express our sincere
gratitude to Dr. P. Auger for critical reading of the draft of this manuscript and many invaluable comments, and Dr.
O. D. Seeman for his kind suggestions on setal nomenclature.

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