J. Acarol. Soc. Jpn., 13 (1): 1-30. 2004  The Acarological Society of Japan 1

[REVIEW] Checklist and Keys to Japanese (: Gamasina: )

1* 2 Shôzô EHARA and Hiroshi AMANO

1Hamasaka 2–15–7, Tottori 680–0001, Japan 2Laboratory of Applied Entomology and Zoology, Faculty of Horticulture, Chiba University, 648 Matsudo, Chiba 271–8510, Japan (Received 5 February 2004; Accepted 16 March 2004)

ABSTRACT

Forty-nine Japanese species of Amblyseiinae, a subfamily of Phytoseiidae, are assigned to 10 genera, namely: Hughes, De Leon, Muma, Berlese, Wainstein, Muma, Okiseius Ehara, Amblyseiulella Muma, Paraphy- toseius Swirski and Shechter, and Indoseiulus Ehara. An annotated list of these species and keys to the genera and species are given, with redescription of Neoseiulus repletus (Wu and Li), recorded from Japan for the first time. urumanus (Ehara) differs from P. orientalis (Narayanan, Kaur and Ghai) in that leg IV is provided with five instead of four macrosetae. Key words: Amblyseiinae, Euseius, Neoseiulus, Paraphytoseius, Typhlodromalus, Typhlodromips

INTRODUCTION

Forty-seven Japanese species belonging to the subfamily Amblyseiinae Muma were previously referred to five genera, Amblyseius Berlese, Okiseius Ehara, Amblyseiulella Muma, Paraphytoseius Swirski and Shechter, and Indoseiulus Ehara (Ehara et al., 1994, 2000; Ehara and Amano, 1998, 2002). In the present paper 49 Japanese ambliseiine are revised and listed with annotations, in accordance with the current classification of the subfamily, in which more genera based on narrower concepts are placed (Beard, 2001; Chant and McMurtry, 2003a). Such a classification was founded generally by Floridan acarolo- gists about forty years ago (Muma, 1961; Muma and Denmark, 1968; Muma et al., 1971), but it has not always been accepted by many taxonomists for a long time. However, its gradual acceptance in their later papers is largely owing to the striking increase of the number of described species of phytoseiids. In this paper the diagnostic characters including leg setation are mentioned about female mites unless otherwise stated. One species newly found from Japan is redescribed and illustrated. The tribes and species groups are omitted below. The setal notations follow those of Rowell et al. (1978). In order to avoid the confusion in spermathecal terminology, the term calyx is used here instead of the cervix. The term cervix has been

*Corresponding author: e-mail: [email protected] 2 Shôzô EHARA and Hiroshi AMANO referred to the calyx by many acarologists, but it was also used often for the connection (neck) between the calyx and the atrium (cf. Beard, 2001). All the measurements are given in micrometers.

Key to the Genera of Japanese Amblyseiinae (Females) 1. Peritrematic shields not fused anteriorly with dorsal shield ...... Indoseiulus Ehara Peritrematic shields fused anteriorly with dorsal shield...... 2 2. Either or both setae J2 and S4 absent...... 3 Both setae J2 and S4 present...... 6 3. Seta S4 present; seta J2 absent ...... Proprioseiopsis Muma Seta S4 and J2 absent ...... 4 4. Dorsal shield without lateral incision at level of seta s4 ...... Okiseius Ehara Dorsal shield with lateral incision at level of seta s4 ...... 5 5. Seta S2 present ...... Amblyseiulella Muma Seta S2 absent...... Paraphytoseius Swirski and Shechter 6. Chelicerae tiny; 2 or all 3 pairs of preanal setae more or less in a transverse line on anterior part of preanal region...... Euseius Wainstein Chelicerae not tiny; 2 or all 3 pairs of preanal setae not in a transverse line on anterior part of preanal region, with exceptions ...... 7 7. Macrosetae present on leg IV, absent on legs I–III (may be scarcely discernible on leg III in N. haimatus) ...... Neoseiulus Hughes Macrosetae present on legs I–IV (sometimes absent on legs I and II)...... 8 8. Posterior margin of sternal shield trilobate ...... Typhlodromalus Muma Posterior margin of sternal shield nearly straight or concave ...... 9 9. Seta Z5 much longer than distance between bases of Z5; tarsus I with an erect macroseta near base...... Amblyseius Berlese Seta Z5 shorter than, or approximately as long as distance between bases of Z5 (except for T. ochii and T. indocalami); tarsus I without an erect proximal seta ...... Typhlodromips De Leon

Genus Neoseiulus Hughes, 1948

Neoseiulus Hughes, 1948: 141; Muma and Denmark 1968: 235; Muma et al. 1971: 100; Beard 2001: 79; Chant and McMurtry 2003a: 15. [Type species: Neoseiulus barkeri Hughes, 1948, by original designation] Amblyseius (Neoseiulus): Ehara and Amano 1998 (in part): 28.

The genus Neoseiulus can be distinguished from Typhlodromips by the absence of macrosetae on legs I–III. However, genu III may bear a scarcely discernible macroseta in N. haimatus. Leg IV of this genus is usually provided with 1 macroseta on the basitarsus, and may bear 1 macroseta each on the genu and tibia. The posterior margin of the sternal shield is nearly straight or concave in Japanese species. Japanese Amblyseiinae 3

Key to Species of the Genus Neoseiulus in Japan (Females) 1. Setae j4 to j6 and J2 longer than distances between their bases ..... womersleyi (Schicha) Setae j4 to j6 and J2 shorter than distances between their bases...... 2 2. Atrium of spermatheca very large, bifurcate at junction with major duct ...... 3 Atrium of spermatheca not bifurcate at junction with major duct ...... 4 3. Genu IV with macroseta...... makuwa (Ehara) Genu IV without macroseta...... barkeri Hughes 4. All 3 pairs of preanal setae and pores on anterior one-third to two-fifths of preanal region of ventrianal shield...... 5 At least 1 pair of preanal setae and pores not on anterior one-third to two-fifths of preanal region of ventrianal shield...... 6 5. Seta Z5 about 20 µm long ...... hinoki (Ehara) Seta Z5 about 60 µm long ...... repletus (Wu and Li) 6. Leg IV with 3 macrosetae ...... 7 Leg IV with 1 macroseta ...... 9 7. Dorsal shield slender; ventrianal shield oval...... inabanus (Ehara) Dorsal shield not slender; ventrianal shield approximately pentagonal...... 8 8. Seta Z5 approximately as long as Z4 ...... yanoi (Ehara) Seta Z5 noticeably longer than Z4 ...... haimatus (Ehara) 9. Seta Z4 approximately as long as S4 ...... bicaudus (Wainstein) Seta Z4 noticeably longer than S4...... 10 10. Seta Z4 approximately as long as distance to base of S5...... californicus (McGregor) Seta Z4 much shorter than distance to base of S5...... koyamanus (Ehara and Yokogawa)

1. Neoseiulus californicus (McGregor, 1954)

Typhlodromus californicus McGregor, 1954: 89, figs. 1–4. [Type locality: Whittier, California, U.S.A.; type habitat: lemon] Typhlodromus chilenensis Dosse, 1958: 55, figs. 11–15. [Type loc.: Chile; type hab.: Eichhornia crassipes L.] Synonymy by Athias-Henriot (1977) Amblyseius chilenensis: González and Schuster 1962: 10, fig. 3; Ehara 1964: 383, figs. 9–16. Cydnodromus californicus: Athias-Henriot 1977: 62, figs. 10, 13, 14, 18, 27–29. Neoseiulus californicus: Moraes et al. 1986: 73; Chant and McMurtry 2003a: 21. Amblyseius californicus: Ehara and Amano 1993: 10, fig. 5. Amblyseius (Amblyseius) californicus: Ehara et al. 1994: 126. Amblyseius (Neoseiulus) californicus: Ehara and Amano 1998: 33.

Basitarsus IV with distinct macroseta; tibia and genu IV with macroseta practically absent to scarcely discernible. Spermathecal calyx considerably variable in shape. In Japan this was found preying on the cyclamen mite, Phytonemus pallidus (Banks), on Boehmeria nivea Gaud. in Meguro, Tokyo (Ehara, 1964). At present it is well known as being an effective predator of spider mites (McMurtry, 1982) and is used as one of the biotic pesticides for them in the world. 4 Shôzô EHARA and Hiroshi AMANO

New collection records. Two ♀, Yahatahama, Ehime Pref., 29–VII–2002 (R. Ohnishi), on citrus. Distribution. Japan (Honshu; Shikoku, new record); Europe, Algeria, U.S.A., Central and South America.

2. Neoseiulus haimatus (Ehara, 1967) (Fig. 7)

Amblyseius (Amblyseius) haimatus Ehara, 1967b: 214, figs. 8–12; Ehara 1972: 149, figs. 49–53; Ehara et al. 1994: 126. [Type loc.: Mt. Muine, Hokkaido; type hab.: Pinus pumila Rgl.] Neoseiulus haimatus: Moraes et al. 1986: 83; Denmark and Edland 2002: 214, fig. 3W, map 8. Amblyseius haimatus: Kolodochka 1989: 13, fig. 1. Amblyseius (Neoseiulus) haimatus: Ehara and Amano 1998: 37, fig. 23.

Genu III without or with macroseta scarcely discernible. In the original description of this species the spermathea was illustrated in comparison with that of one female of “Amblyseius reticulatus (Oudemans, 1930a),” which was borrowed from Dr. W. Karg, Germany. Recently, spermatheca of “A. reticulatus” was shown erroneously as that of A. haimatus in Ehara and Amano (1998, fig. 28). In addition, Dr. Karg lately informed Ehara that the “A. reticulatus” specimen had been misidentified by him and ought to have been assigned to other species (personal communication to Ehara). Distribution. Japan (Hokkaido, Honshu); Europe.

3. Neoseiulus koyamanus (Ehara and Yokogawa, 1977)

Amblyseius (Amblyseius) koyamanus Ehara and Yokogawa, 1977: 50, figs. 1–8; Zhu and Chen 1983b: 181, fig. 1; Ehara 1985: 118, figs. 13, 14; Ryu and Lee 1992: 24, figs. 10–15; Ehara et al. 1994: 125. [Type loc.: Koyama, Tottori City, Honshu; type hab.: a graminaceous plant] Neoseiulus koyamanus: Moraes et al. 1986: 85; Chant and McMurtry 2003a: 23. Amblyseius (Neoseiulus) koyamanus: Ehara and Amano 1998: 35.

Distribution. Japan (Honshu, Kyushu); Korea, China.

4. Neoseiulus yanoi (Ehara, 1972)

Amblyseius (Amblyseius) yanoi Ehara, 1972: 151, figs. 54–62; Ehara et al. 1994: 125. [Type loc.: Mt. Ontake, Kiso, Honshu; type hab.: Abies veitchii Lindl. at 2350m above the sea level] Neoseiulus yanoi: Moraes et al. 1986: 100; Chant and McMurtry 2003a: 23. Amblyseius yanoi: Kolodochka 1989: 16, fig. 2. Amblyseius (Neoseiulus) yanoi: Ehara and Amano 1998: 37.

Distribution. Japan (Honshu); Ukraine. Japanese Amblyseiinae 5

5. Neoseiulus bicaudus (Wainstein, 1962)

Amblyseius bicaudus Wainstein, 1962b: 237, figs. 40, 41; Livschitz and Kuznetzov 1972: 26, fig. 33; Kolodochka 1978: 34, fig. 12. [Type loc.: Kargalink, Alma-Ata, Kazakhstan; type hab.: grass] Amblyseius (Amblyseius) hirotae Ehara, 1985: 119, figs. 15–24; Ehara et al. 1994: 126. [Type loc.: Tottori Sand Dune, Tottori Pref., Honshu; type hab.: Ischaemum anthephoroides (Steud.) Miq.] New synonymy. Neoseiulus bicaudus: Moraes et al. 1986: 72; Chant and McMurtry 2003a: 23. Neoseiulus hirotae: Moraes et al. 1986: 83; Chant and McMurtry 2003a: 23. Amblyseius (Neoseiulus) hirotae: Ehara and Amano 1998: 35.

The spermathecal calyx of Amblyseius hirotae is individually rather variable in shape; in some specimens the calyx appears to have just the same shape as that of A. bicaudus (Livschitz and Kuznetzov, 1972; Kolodochka, 1978). Therefore, A. hirotae Ehara, 1985 is recognized as a junior synonym of A. bicaudus Wainstein, 1962. Distribution. Japan (Honshu); Central Asia, Caucasus, Ukraine, Moldavia, France.

6. Neoseiulus inabanus (Ehara, 1972)

Amblyseius (Amblyseius) inabanus Ehara, 1972: 146, figs. 36–41; Ehara and Yokogawa 1977: 52, fig. 9; Ehara et al. 1994: 126. [Type loc.: Koyama, Tottori City, Honshu; type hab.: Phragmites communis Trin.] Neoseiulus inabanus: Moraes et al. 1986: 84; Chant and McMurtry 2003a: 27. Amblyseius (Neoseiulus) inabanus: Ehara and Amano 1998: 33, figs. 18, 37.

Distribution. Japan (Honshu).

7. Neoseiulus barkeri Hughes, 1948

Neoseiulus barkeri Hughes, 1948: 141, figs. 200–206; Hughes 1976: 343, figs. 391–394; Moraes et al. 1986: 70. [Type loc.: London docks, England; type hab.: germinating barley] Typhlodromus (Neoseiulus) barkeri: Nesbitt 1951: 35, pl. 12, figs. 31, 33. Typhlodromus (Amblyseius) barkeri: Hughes 1961: 222, figs. 300–303. Amblyseius barkeri: Athias-Henriot 1961: 440, figs. 124–126; Athias-Henriot 1966: 215, figs. 24, 32, 129. Amblyseius (Amblyseius) barkeri: Ehara 1972: 146, figs. 42–48; Ehara et al. 1994: 124. Amblyseius (Neoseiulus) barkeri: Ehara and Amano 1998: 34, figs. 27, 44.

New collection records. Four ♀, Aki, Kochi Pref., 11–X–2000 (T. Hirose), on eggplant. Distribution. Japan (Hokkaido, Honshu; Shikoku, new record; Kyushu); Korea, China, Israel, Turkey, Georgia, U.K., Europe, Africa.

8. Neoseiulus makuwa (Ehara, 1972)

Amblyseius (Amblyseius) makuwa Ehara, 1972: 154, figs. 70–74; Ehara and Hamaoka 1980: 6, fig. 8; Ryu 1993: 109, figs. 77–81; Ehara et al. 1994: 124. [Type loc.: Kita-Usa, Usa, Oita Pref., Kyushu; type hab.: Cucumis melo L. var. makuwa Makino] Amblyseius makuwa: Chen et al. 1984: 335, fig. 14 (32); Ehara and Amano 1993: 6, fig. 3; Wu et al. 1997b: 99, fig. 69. 6 Shôzô EHARA and Hiroshi AMANO

Neoseiulus makuwa: Moraes et al. 1986: 87; Chant and McMurtry 2003a: 37. Amblyseius (Neoseiulus) makuwa: Ehara and Amano 1998: 37; Ehara 2002b: 127, fig. 10.

New collection record. One ♀, Higashitsuno, Kochi Pref., 19–VI–2002 (K. Komi), on strawberry. Distribution. Japan (Honshu; Shikoku, new record; Kyushu); Korea, China, Taiwan, Indonesia (Sumatra).

9. Neoseiulus womersleyi (Schicha, 1975)

Amblyseius womersleyi Schicha, 1975: 101, figs. 1–9; Ehara and Amano 1993: 8, fig. 4 [Type loc.: Rydalmere, New South Wales, Australia; type hab.: strawberry] Typhlodromus longispinosus (nec Evans, 1952): Ehara 1958: 55, figs. 4–6. Amblyseius longispinosus (nec Evans, 1952): Ehara 1961: 95, fig. 5. Neoseiulus longispinosus: Moraes et al. 1986: 85 (in part). Amblyseius (Amblyseius) womersleyi: Ehara et al. 1994: 123. Amblyseius (Neoseiulus) womersleyi: Ehara and Amano 1998: 30. Neoseiulus womersleyi: Beard 2001: 84, figs. 6a–d; Chant and McMurtry 2003a: 37.

Distribution. Japan (Hokkaido, Kunashiri, Honshu, Shikoku, Kyushu, Amami-oshima I., Okinawa I., Ishigaki I.); Korea, Sakhalin, China, Taiwan, Australia, New Zealand.

10. Neoseiulus hinoki (Ehara, 1972), n. comb.

Amblyseius (Amblyseius) hinoki Ehara, 1972: 165, figs. 111–115; Ehara et al. 1994: 126. [Type loc.: Mt. Ontake, Kiso, Honshu; type hab.: Chamaecyparis obtusa (Sieb. et Zucc.) Endl. at 1550m above the sea level] Typhlodromalus hinoki: Moraes et al. 1986: 129. Amblyseius (Neoseiulus) hinoki: Ehara and Amano 1998: 31.

Sternal shield with posterior margin nearly straight. Leg IV with macroseta on tibia and basitarsus. Distribution. Japan (Honshu).

11. Neoseiulus repletus (Wu and Li, 1985), n. comb. (Japanese name: Hoonoki-kaburidani) (Figs. 1–6)

Amblyseius (Amblyseius) repletus Wu and Li, 1985: 268, figs. 1–5. [Type loc.: Shen Nong Jia, Hubei Province, China; type hab.: Rhus sp., Vitis adstricta Hance, or Prunus salicina Lindl. (Holotype and 8 paratype specimens taken from the 3 plant species.)]

Female. Dorsal shield (Fig. 1) heavily sclerotized, reticulate, with at least 6 pairs of solenostomes. Setae on dorsal shield: Z5 stout, serrate; remaining setae much shorter, smooth, seta Z4 slightly longer than S4. Setae r3 and R1 similar to most setae on dorsal shield. Peritreme Japanese Amblyseiinae 7

Figs. 1, 2. Neoseiulus repletus (♀). 1, dorsum of idiosoma; 2, posterior ventral surface.

Figs. 3–6. Neoseiulus repletus (♀). 3, sternal shield; 4, spermatheca; 5, chelicera; 6, genu, tibia and basitarsus of leg IV. A, atrium; C, calyx; MA, major duct; MI, minor duct. extending anterior to seta j1; posterior extension of peritrematic shield with termination more or less truncate (Fig. 2). Sternal shield wider than long, with posterior margin concave (Fig. 3). Ventrianal shield nearly pentagonal, slightly longer than wide, approximately as wide as genital shield, with lateral margins slightly concave (Fig. 2); 3 pairs of preanal setae on anterior two fifths of preanal region, ZV2 and JV2 in a transverse line with pair of crescentic 8 Shôzô EHARA and Hiroshi AMANO pores. Two pairs of slender metapodal platelets. Spermatheca with calyx bowl-shaped, with atrium conspicuous (Fig. 4). Fixed digit of chelicera with 7 teeth, the movable digit bidentate (Fig. 5). Chaetotaxic formula: genu II 2–2/0, 2/0–1; genu III 1–2/1, 2/0–1. Leg IV with mac- roseta on basitarsus, without macrosetae on tibia and genu (Fig. 6). Measurements (mean, n=4): length of idiosoma 372, width of idiosoma 283; length of dorsal shield 346, width of dorsal shield 228; lengths of setae (mean±SE, n=8): j1 16.3±0.5, j3 17.6±0.2, j4 10.5±0.3, j5 9.9±0.3, j6 10.7±0.3, J2 13.1±0.4, J5 9.8±0.1, z2 12.7±0.2, z4 13.4±0.2, z5 10.6±0.1, Z1 12.5±0.4, Z4 18.7±0.8, Z5 60.5±0.5, s4 15.3±0.3, S2 13.8±0.3, S4 13.3±0.3, S5 13.4±0.3, r3 14.8±0.3, R1 12.2±0.3, JV5 22.7±0.5; macroseta on basitarsus IV 27.1±0.2. Male. Not known. New collection records. Three ♀, Experimental Forest of Nagoya Univ., Inabu-cho, Kita- shitara-gun, Aichi Pref., 10–X–2002 (E. Shibata), on Magnolia hypoleuca Sieb. et Zucc.; 1 ♀, Expt. Forest of Kyoto Univ., Miyama-cho, Kita-kuwada-gun, Kyoto Pref., 2–X–2002 (A. Takafuji), on the above plant species. Distribution. Japan (Honshu, new record); China. Remarks. Neoseiulus repletus is close to N. hinoki (Ehara) but distinct in having Z5 (61 µm) much longer than that (21 µm) in hinoki.

Genus Typhlodromips De Leon, 1965

Typhlodromips De Leon, 1965: 23; Muma et al. 1971: 78; Moraes et al. 1986: 135. [Type species: Typhlodromus simplissimus De Leon, 1959, by subsequent designation, De Leon (1965)]

The genus Typhlodromips differs from Neoseiulus in that the macrosetae are usually present on genua II and III, and may also be present on genu I and tibia III. Leg IV has 3 macrosetae: genu, tibia, basitarsus. The posterior margin of the sternal shield is nearly straight or concave.

Key to Species of the Genus Typhlodromips in Japan (Females) 1. Genu II with 8 setae ...... paraki (Ehara) Genu II with 7 setae ...... 2 2. Seta Z5 much longer than distance between bases of Z5...... 3 Seta Z5 shorter than, or approximately as long as distance between bases of Z5 ...... 4 3. Seta S2 noticeably longer than Z1...... ochii (Ehara and Yokogawa) Seta S2 as long as Z1...... indocalami (Zhu and Chen) 4. Ventrianal shield with 3 pairs of preanal setae and pair of pores on anterior third of preanal region...... ezoensis (Ehara) Ventrianal shield with 3 pairs of preanal setae and pair of pores not on anterior third of preanal region...... 5 5. Ventrianal shield with lateral margins strongly concave, widest at level of anus...... cantonensis (Schicha) Ventrianal shield with lateral margins slightly concave to nearly straight...... 6 6. Spermatheca with calyx cup or bowl-shaped...... 7 Japanese Amblyseiinae 9

Spermatheca with calyx fundibular or saccular ...... 10 7. Seta Z4 approximately as long as S4 ...... neoparaki (Ehara) Seta Z4 much longer than S4 ...... 8 8. Seta Z5 slightly longer than Z4...... rademacheri (Dosse) Seta Z5 at least twice as long as Z4 ...... 9 9. Macroseta on basitarsus IV much longer than that on tibia IV...... tsugawai (Ehara) Macroseta on basitarsus IV approximately as long as that on tibia IV ...... ishikawai (Ehara) 10. Dorsal shield wholly reticulate...... 11 Dorsal shield nearly smooth, or reticulate only along lateral margins...... 12 11. Seta Z5 approximately as long as distance between bases of Z5...... alpicola (Ehara) Seta Z5 noticeably shorter than distance between bases of Z5 ...... ainu (Ehara) 12. Ventrianal pores just behind the posterior preanal setae (JV2) ...... morii (Ehara) Ventrianal pores between the posterior preanal setae (JV2) ...... 13 13. Z4 longer than distance between its base and that of Z5 ...... oguroi (Ehara) Z4 shorter than distance between its base and that of Z5...... okinawanus (Ehara)

12. Typhlodromips paraki (Ehara, 1967), n. comb. (Fig. 8)

Amblyseius (Amblyseius) paraki Ehara, 1967b: 216, figs. 16–20; Ehara and Yokogawa 1977: 52, figs. 10–16; Ehara et al. 1994: 126. [Type loc.: Sapporo, Hokkaido; type hab.: apple] Neoseiulus paraki: Moraes et al. 1986: 92; Chant and McMurtry 2003a: 23. Amblyseius (Neoseiulus) paraki: Ehara and Amano 1998: 35, fig. 35.

Genu II with 8 setae, without macroseta; genu III with distinctive macroseta. Genu I without macroseta. Distribution. Japan (Hokkaido, Honshu).

13. Typhlodromips ochii (Ehara and Yokogawa, 1977), n. comb.

Amblyseius (Amblyseius) ochii Ehara and Yokogawa, 1977: 54, figs. 17–24; Ehara et al. 1994: 124; Ehara and Amano 1998: 41. [Type loc.: Kyushozan, Tottori City, Honshu; type hab.: bamboo] Amblyseius ochii: Chen et al. 1984: 333, fig. 14 (30); Wu et al. 1997b: 94, fig. 63. Neoseiulus ochii: Moraes et al. 1986: 91.

Leg III with distinctive macroseta each on genu and tibia; basitarsus III with macroseta scarcely discernible or practically absent. Genua I and II without macrosetae. Distribution. Japan (Honshu); China.

14. Typhlodromips indocalami (Zhu and Chen, 1983), n. comb.

Amblyseius indocalami Zhu and Chen, 1983a: 385, figs. 7–12; Wu et al. 1997b: 61, fig. 32. [Type loc.: Mt. Wuyi, Jiangxi Prov., China; type hab.: Indocalamus tessellatus (Munro.) Keng. F.] 10 Shôzô EHARA and Hiroshi AMANO

Amblyseius (Amblyseius) indocalami: Ehara et al. 1994: 130, figs. 18–26; Ehara and Amano 1998: 40, fig. 31.

Genua I–III and tibia III each with macroseta. Distribution. Japan (Honshu); China.

15. Typhlodromips neoparaki (Ehara, 1972), n. comb. (Fig. 9)

Amblyseius (Amblyseius) neoparaki Ehara, 1972: 153, figs. 63–69; Ehara et al. 1994: 124. [Type loc.: Shiga Heights, Nagano Pref., Honshu; type hab.: a bamboo, Sasa kurilensis (Rupr.) Makino et Shibata] Neoseiulus neoparaki: Moraes et al. 1986: 90; Chant and McMurtry 2003a: 23. Amblyseius (Neoseiulus) neoparaki: Ehara and Amano 1998: 36.

Leg III with macroseta each on genu and tibia; genua I and II without macrosetae. Distribution. Japan (Honshu).

16. Typhlodromips rademacheri (Dosse, 1958)

Amblyseius rademacheri Dosse, 1958: 44, figs. 1–5; Ehara 1959: 288, figs. 6–11; Ehara 1961: 96; Kolodochka 1978: 22, fig. 3; Ryu and Ehara 1992: 727, figs. 15–23. [Type loc.: Stuttgart-Hohenheim, Germany; type hab.: apple] Amblyseius (Amblyseius) rademacheri: Ehara 1966: 23; Zhu and Chen 1983b: 182, fig. 2; Ehara et al. 1994: 130. Amblyseius (Typhlodromips) rademacheri: Karg 1971: 185, figs. 129e, 132h. Typhlodromips rademacheri: Moraes et al. 1986: 145. Amblyseius (Neoseiulus) rademacheri: Ehara and Amano 1998: 31.

Fig. 7. Neoseiulus haimatus, spermatheca (Ehara, 1967b). Fig. 8. Typhlodromips paraki (♀), genu III. Fig. 9. T. neoparaki (♀), genu and tibia III. Fig. 10. Typhlodromalus japonicus (♀), sternal shield. Japanese Amblyseiinae 11

Genua I–III each with macroseta; tibia III with macroseta. New collection record. One ♀, Ayakami, Kagawa Pref., 9–VII–2001 (E. Matsumoto), on Pueraria lobata (Willd.) Ohwi. Distribution. Japan (Hokkaido; Shikoku, new record; Honshu); Korea, China, Siberia, Caucasus.

17. Typhlodromips tsugawai (Ehara, 1959)

Amblyseius tsugawai Ehara, 1959: 290, figs. 12, 13; Ehara 1961: 95, fig. 6; Ehara 1964: 386, figs. 24–27; Chen et al. 1984: 341, fig. 14 (40); Ehara and Amano 1993: 14, fig. 7; Wu et al. 1997b: 64, fig. 35; Ryu and Kim 1998: 101, figs. 1–6. [Type loc.: Kuroishi, Aomori Pref., Honshu; type hab.: apple] Amblyseius (Amblyseius) tsugawai: Ehara 1966: 23; Ehara 1967b: 221, figs. 35–39; Ehara 1972: 157, fig. 79; Ehara et al. 1994: 130; Ehara and Amano 1998: 41, fig. 38. Typhlodromips tsugawai: Moraes et al. 1986: 151.

Genua I–III each with macroseta; tibia III with macroseta. Distribution. Japan (Hokkaido, Honshu, Shikoku); Korea, China.

18. Typhlodromips ishikawai (Ehara, 1972)

Amblyseius (Amblyseius) ishikawai Ehara, 1972: 158, figs. 83–91; Ehara et al. 1994: 130. [Type loc.: Mt. Ontake, Kiso, Honshu; type hab.: Alnus maximowiczii Callier at 2570m above the sea level] Typhlodromips ishikawai: Moraes et al. 1986: 141. Amblyseius (Neoseiulus) ishikawai: Ehara and Amano 1998: 31.

Genua II and III and tibia III each with macroseta; genu I without macroseta. Distribution. Japan (Honshu).

19. Typhlodromips alpicola (Ehara, 1982)

Amblyseius (Amblyseius) alpicola Ehara, 1982: 40, figs. 1–9; Ehara et al. 1994: 125. [Type loc.: Shiga Heights, Nagano Pref., Honshu; type hab.: Acer ukurunduense Trautv. et Mey.] Typhlodromips alpicola: Moraes et al. 1986: 136. Amblyseius (Neoseiulus) alpicola: Ehara and Amano 1998: 36.

Genua II and III and tibia III each with macroseta; genu I without macroseta. Distribution. Japan (Honshu).

20. Typhlodromips ainu (Ehara, 1967)

Amblyseius (Amblyseius) ainu Ehara, 1967b: 218, figs. 21–27; Ehara 1972: 156, fig. 77; Ehara et al. 1994: 125. [Type loc.: Mombetsu, Hidaka Prov., Hokkaido; type hab.: Cirsium kamtschaticum Ledeb.] Amblyseius ainu: Chen et al. 1984: 337, fig. 14 (35); Wu et al. 1997b: 85, fig. 55. Typhlodromips ainu: Moraes et al. 1986: 135. Amblyseius (Neoseiulus) ainu: Ehara and Amano 1998: 32, fig. 25. 12 Shôzô EHARA and Hiroshi AMANO

Genua I–III and tibia III each with macroseta. Distribution. Japan (Hokkaido, Honshu); China.

21. Typhlodromips morii (Ehara, 1967), n. comb.

Amblyseius (Amblyseius) morii Ehara, 1967b: 219, figs. 28–34; Ehara 1972: 156, fig. 78; Ehara et al. 1994: 123. [Type loc.: Toyotomi, Sarobetsu wasteland, Hokkaido; type hab.: Salix hultenii Floderus var. angustifolia Kitamura] Amblyseius morii: Moraes et al. 1986: 22. Amblyseius (Neoseiulus) morii: Ehara and Amano 1998: 32.

Genua II and III and tibia III each with macroseta; genu I without macroseta. Distribution. Japan (Hokkaido, Honshu).

22. Typhlodromips oguroi (Ehara, 1964)

Amblyseius oguroi Ehara, 1964: 384, figs. 17–23; Wu et al. 1997b: 90, fig. 60. [Type loc.: Sendai, Miyagi Pref., Honshu; type hab.: Mallotus japonicus (Thunb.) Muell. Arg. Amblyseius (Amblyseius) oguroi: Ehara 1966: 21; Ryu and Lee 1992: 23, figs. 1–9; Ehara et al. 1994: 124. Typhlodromips oguroi: Moraes et al. 1986: 144. Amblyseius (Neoseiulus) oguroi: Ehara and Amano 1998: 32.

Macrosetae on legs I–III: genua I–III, tibiae II and III, and basitarsus II; basitarsus III with macroseta practically absent. New collection records. Eight ♀, Monobe, Kochi Pref., 17–IX–2002 (K. Komi), on Mallotus japonicus; 2 ♀, Takamori, Kumamoto Pref., 9–V–1983 (S. Ehara), on Boehmeria spicata Thunb. Distribution. Japan (Hokkaido, Honshu; Shikoku, new record; Kyushu, new record); Korea, China.

23. Typhlodromips okinawanus (Ehara, 1967), n. comb.

Amblyseius (Amblyseius) okinawanus Ehara, 1967a: 72, figs. 17–24; Ehara and Lee 1971: 64, fig. 8; Tseng 1976: 115, figs. 55–58; Ehara and Hamaoka 1980: 6, figs. 9–11; Ryu and Lee 1992: 27, figs. 22–30; Ehara et al. 1994: 124. [Type loc.: Tomigusuku, Okinawa I.; type hab.: Verbena officinalis L.] Amblyseius okinawanus: Chen et al. 1984: 342, fig. 14 (41); McMurtry and Moraes 1985; 80; Wu et al. 1997b: 89, fig. 59. Neoseiulus okinawanus: Moraes et al. 1986: 91. Amblyseius (Neoseiulus) okinawanus: Ehara and Amano 1998: 37; Ehara 2002b: 127, figs. 11, 12.

Leg III: genu and tibia each with macroseta; basitarsus with macroseta practically absent. Genua I and II without macrosetae. New collection record. One ♀, Kamojima, Tokushima Pref., 21–X–2001 (Y. Shimada), on Japanese Amblyseiinae 13

Rhynchosia acuminatifolia Makino. Distribution. Japan (Honshu; Shikoku, new record; Kyushu, Amami-oshima I., Okinawa I.); Korea, China, Taiwan, Thailand, Sumatra, Papua New Guinea, Russia.

24. Typhlodromips ezoensis (Ehara, 1967), n. comb.

Amblyseius (Amblyseius) ezoensis Ehara, 1967b: 223, figs. 45–49; Ehara 1972: 167, figs. 116, 117; Ehara et al. 1994: 127. [Type loc.: Mt. Moiwa, Sapporo, Hokkaido; type hab.: Heracleum dulce Fisch.] Thphlodromalus ezoensis: Moraes et al. 1986: 129. Amblyseius ezoensis: Wu et al. 1997b: 70, fig. 41. Amblyseius (Neoseiulus) ezoensis: Ehara and Amano 1998: 31.

Genua I–III and tibia III each with macroseta. Distribution. Japan (Hokkaido, Honshu); China.

25. Typhlodromips cantonensis (Schicha, 1982), n. comb.

Amblyseius cantonensis Schicha, 1982: 48, figs. 10–15. [Type loc.: Canton, China; type hab.: citrus] Amblyseius (Amblyseius) newsami (nec Evans, 1953): Ehara and Lee 1971: 66, figs. 13–20; Ehara and Bhandhufalck 1977: 62; Tseng 1983: 44, figs. 30–35. Amblyseius newsami (nec Evans, 1953): Chen et al. 1984: 330, figs. 14 (26); Wu et al. 1997b: 74, fig. 45. Amblyseius (Neoseiulus) cantonensis: Ehara and Amano 2002: 322, figs. 1, 2.

Macrosetae on legs I–III: genua I–III, tibiae II and III, and basitarsi II and III. Distribution. Japan (Ishigaki I., Taketomi I.); China, Hainan I., Taiwan, Thailand.

Genus Typhlodromalus Muma, 1961

Amblyseius (Typhlodromalus) Muma, 1961: 288. [Type species: Amblyseius (Typhlodromalus) peregrinus Muma, 1961, by original designation] Typhlodromalus: De Leon 1966: 87; Muma et al. 1971: 86; Moraes et al. 1986: 128. Amblyseius (Neoseiulus): Ehara and Amano 1998 (in part): 28.

The genus Typhlodromalus is distinguished from Neoseiulus and Typhlodromips by having the sternal shield with trilobate posterior margin. The arrangement of the macrosetae on legs I–III is similar to that of Typhlodromips. Leg IV is provided with 3 macrosetae: genu, tibia, basitarsus. One species belonging to Typhlodromalus is known from Japan.

26. Typhlodromalus japonicus (Ehara, 1958) (Fig. 10)

Typhlodromus japonicus Ehara, 1958: 56, figs. 7–9. [Type loc.: Koshino-o, Nishimera, Miyazaki Pref., Kyushu; type hab.: Mallotus japonicus (Thunb.) Muell. Arg.] Amblyseius japonicus: Ehara 1961: 95, fig. 3; Schicha 1980: 28, figs. 48–54; Schicha 1987: 94, pl. 47. Amblyseius (Amblyseius) japonicus: Ehara 1966: 21; Ehara et al. 1994: 125. 14 Shôzô EHARA and Hiroshi AMANO

Typhlodromalus japonicus: Moraes et al. 1986: 130. Amblyseius (Neoseiulus) japonicus: Ehara and Amano 1998: 33.

Dorsal shield reticulate except for a narrow smooth area connecting setae j5, z5 and j6. Spermatheca with tubular, coiled calyx. Chaetotaxic formula: genu II 2–2/0, 2/0–1, genu III 1–2/0, 2/0–1. Genu and tibia of leg III with macroseta; genua I and II without macrosetae. Measurements of holotype (♀): length of idiosoma 332, width of idiosoma 277, length of dorsal shield 358, width of dorsal shield 244; lengths of setae: j1 16.2, j3 20.5, j4 9.7, j5 11.5, j6 13.2, J2 16.2, J5 11.5, z2 16.0, z4 17.0, z5 11.5, Z1 15.0, Z4 21.3, Z5 76.4, s4 19.8, S2 19.0, S4 15.8, S5 13.8, r3 16.1, R1 14.8, JV5 24.5; macrosetae on leg IV: genu 39.3, tibia 29.0, basitarsus 39.1. A Chinese mite from bamboo was identified with Amblyseius japonicus by Chen et al. (1984), but it does not belong to japonicus because of the straight posterior margin of the sternal shield and the different lengths of the macrosetae on leg IV. Distribution. Japan (Kyushu).

Genus Amblyseius Berlese, 1914

Amblyseius Berlese, 1914: 143. [Type species: Zercon obtusus Koch, 1839, by original designation]

The genus Amblyseius is distinctive in having seta Z5 stout, whiplike, much longer than distance between bases of Z5, and tarsus I with an erect macroseta near base (Fig. 14). Setae Z4 and s4 are also stout, remarkably longer than remaining body setae. The posterior margin of the sternal shield is nearly straight, rarely protruded. The macrosetae are present on legs I–IV; leg IV always bears 3 macrosetae: genu, tibia, basitarsus.

Key to Species of the Genus Amblyseius in Japan (Females) 1. Ventrianal shield vase-shaped, with distinct waist...... 2 Ventrianal shield triangular or pentagonal, with a slight waist at most...... 3 2. Sternal shield with posterior margin slightly convex to trilobate ...... eharai Amitai and Swirski Sternal shield with posterior margin nearly straight or concave ...... kokufuensis Ehara and Kato 3. Ventrianal shield wider than long...... ishizuchiensis Ehara Ventrianal shield longer than, or about as long as wide ...... 4 4. Macroseta on tibia IV shorter than that on basitarsus IV...... obtuserellus Wainstein and Begljarov Macroseta on tibia IV longer than, or approximately as long as that on basitarsus IV....5 5. Ventrianal pores just behind posterior pair of preanal setae (JV2) ...... kaguya Ehara Ventrianal pores not as above...... 6 6. Ventrianal shield approximately as long as wide ...... neofirmus Ehara and Okada Ventrianal shield longer than wide ...... 7 7. Spermatheca with calyx constricted in middle...... firmus Ehara Japanese Amblyseiinae 15

Spermatheca with calyx not constricted...... 8 8. Spermatheca with atrium incorporated into proximal portion of calyx ...... tamatavensis Blommers Spermatheca with atrium directly proximal to calyx ...... 9 9. Spermatheca with calyx very slender, atrium distinct...... orientalis Ehara Spermatheca with calyx thick, atrium indistinct ...... shiganus Ehara

27. Amblyseius eharai Amitai and Swirski, 1981 (Figs. 11–14)

Amblyseius eharai Amitai and Swirski, 1981: 60, figs. 1–3, 6–8, 12, 13; McMurtry and Moraes 1984: 35, figs. 19, 23; Wu and Lan 1989: 450; Ehara and Amano 1993: 16, fig. 8. [Type loc.: Tai Po Hui, New Territories, Hong Kong; type hab.: Euphoria longana Lam.] Amblyseius largoensis (nec Muma, 1955): Ehara 1959: 293, figs. 17, 18; Ehara 1961: 96, fig. 8. Amblyseius deleoni (nec Muma and Denmark, 1971): Ehara 1977: 34, figs. 1, 2. Amblyseius (Amblyseius) eharai: Ehara et al. 1994: 127, figs. 1–7; Ehara and Amano 1998: 38, figs. 5–9, 16, 29, 42; Ehara and Amano 2002: 322; Ehara 2002a: 32, fig. 6.

This species is distinctive in having the sternal shield with posterior margin more or less convex. Genua I-III, tibia III and basitarsus III each with macroseta. Distribution. Japan (Honshu, Shikoku, Kyushu, Amami-oshima I., Okinawa I., Ishigaki I., Taketomi I.); Korea, China, Taiwan, Malaysia.

28. Amblyseius kokufuensis Ehara and Kato, 1994 (Figs. 15, 16)

Amblyseius (Amblyseius) kokufuensis Ehara and Kato in Ehara et al. 1994: 128, figs. 8–15; Ryu 1995: 80, figs. 16–

Figs. 11–14. Amblyseius eharai (♀). 11, variation of sternal shield (posterior margin); 12, 13, spermatheca; 14, tarsus I, showing erect macroseta (E). Figs. 15, 16. A. kokufuensis, spermatheca. 11–13, 15, 16, Ehara et al. (1994). 16 Shôzô EHARA and Hiroshi AMANO

25; Ehara and Amano 1998: 38, fig. 1. [Type loc.: Kamikihara, Kokufu, Tottori Pref., Honshu, 14–X–1987; type hab.: Phyllostachys sp.]

The spermathecal calyx of this species is tubular, gradually widening from base to distal extremity, whilst in eharai it is narrower and flaring only in distal half. Genua I–III, tibia III and basitarsus III each with macroseta; basitarsus II with macroseta present or absent. Distribution. Japan (Honshu); Korea.

29. Amblyseius ishizuchiensis Ehara, 1972

Amblyseius (Amblyseius) ishizuchiensis Ehara, 1972: 162, figs. 97–102; Ehara et al. 1994: 127; Ehara and Amano 1998: 40, figs. 10, 24. [Type loc.: Mt. Kamegamori, Shikoku; type hab.: litter] Amblyseius ishizuchiensis: Denmark and Muma 1989: 35, figs. 158–162.

Genua I–III with macroseta; tibiae I and III with macroseta. Distribution. Japan (Shikoku).

30. Amblyseius obtuserellus Wainstein and Begljarov, 1971

Amblyseius obtuserellus Wainstein and Begljarov, 1971: 1806, fig. 3; Denmark and Muma 1989: 124, figs. 660– 664. [Type loc.: Khasan, Primorsky Territory, Russia; type hab.: Potentilla anserina L.] Amblyseius (Amblyseius) obtuserellus: Ehara and Yokogawa 1977: 54, figs. 25–31; Wu 1984: 224, figs. 12, 13; Ryu and Lee 1992: 27, figs. 16–21; Ehara et al. 1994: 132; Ehara and Amano 1998: 40.

The spermatheca of this species is distinctive in that the calyx is tubular, gradually narrowing distally, and with granulate surface. Genua I–III and tibia III with macroseta. New collection records. Two ♀, Chiran, Kagoshima Pref., 18–V–2000 (K. Matsuhira), on tea. Distribution. Japan (Honshu; Kyushu, new record); Korea, China, Siberia.

31. Amblyseius kaguya Ehara, 1966

Amblyseius (Amblyseius) kaguya Ehara, 1966: 12, figs. 15–19; Ehara 1972: 160, fig. 80; Wu 1987: 266, figs. 27–32; Ehara et al. 1994: 132; Ehara and Amano 1998: 40. [Type loc.: Kochi, Kochi Pref., Shikoku; type hab.: bamboo] Amblyseius kaguya: Denmark and Muma 1989: 40, figs. 191–195.

Genua I–III and tibia III with macroseta. Distribution. Japan (Shikoku); China.

32. Amblyseius neofirmus Ehara and Okada, 1994

Amblyseius (Amblyseius) neofirmus Ehara and Okada in Ehara et al. 1994: 133, figs. 31–38; Ryu 1995: 83, figs. 26– 31; Ehara and Amano 1998: 40. [Type loc.: Koyama, Tottori City, Honshu; type hab.: Pleioblastus chino (Franchet et Savatier) Makino var. viridis (Makino) S. Suzuki] Japanese Amblyseiinae 17

Genua I–III and tibia III with macroseta. Distribution. Japan (Honshu); Korea.

33. Amblyseius firmus Ehara, 1967

Amblyseius (Amblyseius) firmus Ehara, 1967b: 222, figs. 40–44; Ehara et al. 1994: 132; Ehara and Amano 1998: 40. [Type loc.: Mombetsu, Hidaka Prov., Hokkaido; type hab.: Magnolia kobus DC. var. borealis Sarg.] Amblyseius firmus: Denmark and Muma 1989: 43, figs. 207–211.

Genua I–III and tibia III with macroseta. New collection records. Three ♀ & 1 ♂, Niimi, Okayama Pref., 31–VIII–2001 (Y. Shimada), on Saxifraga fortunei Hook. f. var. incisolobata (Engl. et Irmsch) Nakai. Distribution. Japan (Hokkaido; Honshu, new record); China.

34. Amblyseius tamatavensis Blommers, 1974

Amblyseius (Amblyseius) tamatavensis Blommers, 1974: 144, figs. 6–12; Ehara 2002a: 33, figs. 7–14; Ehara and Amano 2002: 322, fig. 3. [Type loc.: Ivoloina, near Tamatave, Madagascar; type hab.: Citrus (Papeda) hystrix De Candolle] Amblyseius (Amblyseius) maai Tseng, 1976: 123, figs. 78–83. [Type loc.: Tainan Hsien, Taiwan; type hab.: pineapple] Synonymy by Denmark and Muma (1989). Amblyseius tamatavensis: Schicha 1981: 40; Schicha 1987: 54, pl. 16; Denmark and Muma 1989: 13, figs. 37–43.

Genua I–III and tibia III with macroseta. Distribution. Japan (Ishigaki I.); Taiwan, Malaysia, Singapore, Java, Philippines, Papua New Guinea, Australia, New Caledonia, New Hebrides, Fiji, Samoa, Madagascar.

35. Amblyseius orientalis Ehara, 1959

Amblyseius orientalis Ehara, 1959: 291, figs. 14–16; Ehara 1961: 96, fig. 7; Ehara 1962: 53, figs. 1–5; Chen et al. 1984: 345, fig. 14 (44); Denmark and Muma 1989: 42, figs. 201–206; Ehara and Amano 1993: 18, fig. 9; Wu et al. 1997b: 49, fig. 20. [Type loc.: Sapporo, Hokkaido; type hab.: Quercus crispula Blume] Amblyseius (Amblyseius) orientalis: Ehara 1966: 23; Ehara and Yokogawa 1977: 56, fig. 32; Ryu 1993: 103, figs. 45–52; Ehara et al. 1994: 133, figs. 27–30; Ehara and Amano 1998: 41, figs. 11, 17, 30, 43.

Genua I–III and tibia III with macroseta. Distribution. Japan (Hokkaido, Honshu, Shikoku), Korea, China, Siberia.

36. Amblyseius shiganus Ehara, 1972

Amblyseius (Amblyseius) shiganus Ehara, 1972: 160, figs. 92–96; Ehara et al. 1994: 133; Ehara and Amano 1998: 41. [Type loc.: Shiga Heights, Nagano Pref., Honshu; type hab.: a bamboo, Sasa kurilensis (Rupr.) Makino et Shibata] Amblyseius shiganus: Denmark and Muma 1989: 47, figs. 232–236. 18 Shôzô EHARA and Hiroshi AMANO

Genua I–III and tibiae I and III with macroseta. Distribution. Japan (Honshu).

Genus Euseius Wainstein, 1962

Amblyseius (Amblyseius) section Euseius Wainstein, 1962a: 15. [Type species: Seiulus finlandicus Oudemans, 1915, by original designation] Amblyseius (Euseius): De Leon 1965: 125; Ehara and Amano 1998: 42. Euseius: De Leon 1966: 86; Muma et al. 1971: 92; McMurtry 1983: 257; Moraes et al. 1986: 35.

The genus Euseius is distinctive in that all 3 pairs or 2 pairs of preanal setae are arranged in a transverse line on the anterior part of the preanal region, and the chelicerae are tiny, with fixed digit dentate only on distal portion. The sternal shield is provided with trilobate posterior margin. The ventrianal shield is nearly oval or vase-shaped, and widest at the level of the anus. The macrosetae are usually present on the genu, tibia and basitarsus of leg IV, and often also on the genu and tibia of leg III.

Key to Species of the Genus Euseius in Japan (Females) 1. Three pairs of preanal setae nearly in a transverse line; peritreme very short, not extending beyond level of seta z4 ...... finlandicus (Oudemans) Two mesial pairs of preanal setae nearly in a transverse line; peritreme extending between setae j3 and z2...... 2 2. Seta j1 slightly longer than j3...... sojaensis (Ehara) Seta j1 at least twice the length of j3...... ovalis (Evans)

37. Euseius finlandicus (Oudemans, 1915)

Seiulus finlandicus Oudemans, 1915: 183. [Type loc.: Åbo, Turun Porin Laani, Finland; type hab.: Salix caprea L.] Typhlodromus finlandicus: Nesbitt 1951: 25, pl. 3, pl. 9 (fig. 5), pl. 10 (fig. 12), pl. 11 (fig. 19); Ehara 1958: 53, figs. 1–3; Ehara 1961: 95, fig. 4. Amblyseius finlandicus: Athias-Henriot 1958: 34; Chant and Hansell 1971: 706, figs. 1–4; Chen et al. 1984: 328, fig. 14 (23). Typhlodromus (Amblyseius) finlandicus: Chant 1959: 67, figs. 94, 95. Typhlodromus (Typhlodromus) finlandicus: Westerboer and Bernhard 1963: 592, figs. 365–376. Amblyseius (Amblyseius) finlandicus: Ehara 1966: 24; Ehara et al. 1994: 123. Euseius finlandicus: Karg 1971: 178, fig. 125c; Moraes et al. 1986: 41; Wu et al. 1997b: 120, fig. 88. Amblyseius (Euseius) finlandicus: Ehara and Amano 1998: 42.

Genu and tibia of leg III with macroseta often inconspicuous. Distribution. Japan (Hokkaido, Honshu); Korea, China, Java, India, Iran, Kazakhstan, Siberia, Caucasus, Turkey, U.K., Europe, Africa, North America, Central and South America. This species has been recorded from various parts of the world, as enumerated above. However, it is possible that incorrect identifications are included among early records of the Japanese Amblyseiinae 19 species, as mentioned partly in Moraes et al. (1986).

38. Euseius sojaensis (Ehara, 1964)

Amblyseius sojaensis Ehara, 1964: 381, figs. 5–8; Ehara and Amano 1993: 12, fig. 6. [Type loc.: Sôja, Gumma Pref., Honshu; type hab.: mulberry] Euseius sojaensis: Moraes et al. 1986: 54. Amblyseius (Amblyseius) sojaensis: Ehara 1966: 24; Ehara 1972: 168, figs. 118–121; Ehara et al. 1994: 123. Amblyseius (Euseius) sojaensis: Ehara and Amano 1998: 42, fig. 12.

Genu and tibia of leg III with macroseta; basitarsus III with macroseta often inconspicuous. Distribution. Japan (Honshu, Shikoku, Kyushu).

39. Euseius ovalis (Evans, 1953)

Typhlodromus ovalis Evans, 1953: 458, figs. 5, 6. [Type loc.: Kuala Lumpur, Malaysia; type hab.: rubber plant] Amblyseius (Amblyseius) ovalis: Ehara 1966: 24; Ehara 1967a: 74, figs. 25–30; Ehara 1970: 58; Ehara et al. 1994: 123. Amblyseius ovalis: Corpuz and Rimando 1966: 122, fig. 5; Schicha 1977: 127, figs. 28–34; Chen et al. 1884: 327, fig. 14 (22); Schicha and Corpuz-Raros 1992: 41, pl. 18. Euseius ovalis: Gupta 1978: 335; McMurtry and Moraes 1985: 85; Wu et al. 1997b: 117, fig. 85. Amblyseius (Euseius) ovalis: Ehara and Amano 1998: 43, fig. 36; Ehara and Amano 2002: 323.

Genu and tibia of leg III with macroseta. New collection record. One ♀, Naze, Amami-oshima I., Kagoshima Pref., 13–III–1999 (T. Yamaguchi), on mango. Distribution. Japan (Amami-oshima I., new record; Okinawa I., Ishigaki I.); China, Taiwan, Philippines, Malaysia, India, Mauritius, Mexico, Hawaii, Fiji, Cook Islands, Papua New Guinea, Australia, New Zealand.

Genus Proprioseiopsis Muma, 1961

Proprioseiopsis Muma, 1961: 277. [Type species: Typhlodromus (Amblyseius) terrestris Chant, 1959, by original designation]

The genus Proprioseiopsis is readily distinguished from its allied genera by the absence of seta J2 and the presence of seta S2. The ventrianal shield is as long as, or longer than wide. Leg IV always has 3 macrosetae: genu, tibia, basitarsus.

Key to Species of the Genus Proprioseiopsis in Japan (Females) 1. Seta z4 extending to base of seta s4 ...... scurra (Wainstein and Begljarov) Seta z4 not extending to base of s4 ...... nemotoi (Ehara and Amano) 20 Shôzô EHARA and Hiroshi AMANO

40. Proprioseiopsis scurra (Wainstein and Begljarov, 1971)

Amblyseius scurra Wainstein and Begljarov, 1971: 1803, fig. 1. [Type loc.: Kamen Rýbolov, Primorsky Territory, Russia; type hab.: herb] Amblyseius (Amblyseius) scurra: Ehara 1972: 162, figs. 103–110. Amblyseius scurra: Begljarov 1981: 7, fig. 85. Proprioseiopsis scurra: Moraes et al. 1986: 123. Amblyseius (Proprioseiopsis) scurra: Ehara et al. 1994: 135; Ehara and Amano 1998: 43.

Genua II and III with macroseta. Distribution. Japan (Hokkaido); Siberia.

41. Proprioseiopsis nemotoi (Ehara and Amano, 1998), n. comb.

Amblyseius (Proprioseiopsis) nemotoi Ehara and Amano, 1998: 43, figs. 46–54; Ehara and Amano: 2002, 323. [Type loc.: Kuki, Saitama Pref., Honshu; type hab.: pear]

Genua II and III with macroseta. New collection records. Two ♀, Aki, Kochi Pref., 11–X–2000 (T. Hirose), on eggplant. Distribution. Japan (Honshu; Shikoku, new record; Hahajima I., Amami-oshima I.).

Genus Okiseius Ehara, 1967

Okiseius Ehara, 1967a: 77; Chant and McMurtry 2003b: 199. [Type species: Okiseius subtropicus Ehara, 1967a, by original designation] Okiseius (Okiseius): Kolodochka and Denmark 1996: 233. Okiseius (Kampimodromellus): Kolodochka and Denmark 1996: 241. Amblyseius (Okiseius): Ehara and Amano 1998: 45. Amblyseius (Kampimodromellus): Ehara and Amano 1998: 45.

The genus Okiseius is distinctive in lacking setae S4 and J2, and in having setae S2 and S5. Seta R1 of female is present on the dorsal shield in some species and on the interscutal membrane in the other species. Dorsal shield with or without lateral incision posterior to seta R1. Legs I–III without macrosetae. According to Chant and McMurtry (2003b) 17 species in the world are assigned to this genus.

Key to Species of the Genus Okiseius in Japan (Females) 1. Dorsal shield with seta R1 and a posterolateral incision...... subtropicus Ehara Dorsal shield without seta R1 and a posterolateral incision; seta R1 on interscutal membrane ...... maritimus (Ehara)

42. Okiseius subtropicus Ehara, 1967

Okiseius subtropicus Ehara, 1967a: 77, figs. 36–39; Ehara and Hamaoka 1980: 6, fig. 12; Wu and Qian 1983: 76, Japanese Amblyseiinae 21

figs. 5, 6; Ehara et al. 1994: 136, figs. 39–41; Wu et al. 1997a: 142, figs. 1–7; Walter 1999: 90, fig. 14; Chant and McMurtry 2003b: 199, figs. 34–37. [Type loc.: Itoman, Okinawa Island; type hab.: Hibiscus tiliaceus L.] Platyseiella (Noeledius) subtropicus: Tseng 1976: 102, figs. 33–37. Amblyseius () subtropicus: Ueckermann and Loots 1985: 195. Okiseius (Okiseius) subtropicus: Kolodochka and Denmark 1996: 235, figs. 8–19; Ehara and Amano 2002: 323. Amblyseius (Okiseius) subtropicus: Ehara and Amano 1998, fig. 55.

Leg IV with macroseta on basitarsus and telotarsus. Okiseius subtropicus was reported as being an active predator of the citrus rust mite, Phyllocoptruta oleivorus (Ashmead) on citrus in China (Yang, 1986), and the other eriophyid mite, Aculops pelekassi (Keifer) on citrus in Japan (Ehara et al., 1994). Distribution. Japan (Honshu, Okinawa I., Ishigaki I.); China, Taiwan, Australia.

43. Okiseius maritimus (Ehara, 1967)

Amblyseius (Kampimodromus) maritimus Ehara, 1967b: 224, figs. 50–57; Ehara 1972: 168; Zhu and Chen 1983b: 183; Ehara et al. 1994: 135. [Type loc.: Hamakoshimizu, Abashiri Prov., Hokkaido; type hab.: Rosa rugosa Thunb.] Okiseius maritimus: Moraes et al. 1986: 102; Wu et al. 1997a: 145, figs. 13–16; Chant and McMurtry 2003b: 20. Okiseius (Kampimodromellus) maritimus: Kolodochka and Denmark 1996: 241, figs. 54–63. Amblyseius (Kampimodromellus) maritimus: Ehara and Amano 1998: 45, fig. 56.

Leg IV with 4 macrosetae: genu, tibia, basitarsus, telotarsus. Distribution. Japan (Hokkaido, Honshu); China.

Genus Amblyseiulella Muma, 1961

Amblyseiulella Muma, 1961: 276. [Type species: Typhlodromus heveae Oudemans, 1930b, by original designation]

The genus Amblyseiulella is characterized by having the dorsal shield with a lateral incision near seta s4, setae S4, S5 and J2 absent, and seta S2 present. Leg IV with 4 knobbed to spatulate macrosetae: genu, tibia, basitarsus, telotarsus.

Key to Species of the Genus Amblyseiulella in Japan (Females)

1. Spermatheca with calyx V-shaped, shorter than wide...... amanoi Ehara Spermatheca with calyx saccular, much longer than wide...... yaeyamana Ehara and Amano

44. Amblyseiulella amanoi Ehara, 1994

Amblyseiulella amanoi Ehara in Ehara et al. 1994: 137, figs. 42–48; Ehara and Amano 1998: 46, figs. 20, 33, 46, 57. [Type loc.: Ienoko, Togane, Chiba Pref., Honshu; type hab.: pear] 22 Shôzô EHARA and Hiroshi AMANO

Genu II with macroseta barely discernible; genua I and II without macrosetae. Distribution. Japan (Honshu).

45. Amblyseiulella yaeyamana Ehara and Amano, 2002

Amblyseiulella yaeyamana Ehara and Amano, 2002: 323, figs. 4–11. [Type loc.: Higashi-yashiki, Taketomi Island; type hab.: Hibiscus tiliaceus L.]

Genua I and II with distinctive macroseta; genu III with macroseta practically absent. Distribution. Japan (Taketomi I.).

Genus Paraphytoseius Swirski and Shechter, 1961

Paraphytoseius Swirski and Shechter, 1961: 113. [Type species: Typhlodromus (Amblyseius) orientalis Narayanan, Kaur and Ghai, 1960 (=Paraphytoseius multidentatus Swirski and Shechter, 1961, by Chant and McMurtry, 2003b)]

The genus Paraphytoseius is distinctive in having the dorsal shield with a lateral incision near seta s4, and in lacking setae J2, S2 and S4. Seta S5 present or absent. Leg IV with 4 or 5 spatulate to knobbed macrosetae: (femur), genu, tibia, basitarsus, and telotarsus.

Key to Species of the Genus Paraphytoseius in Japan (Females) 1. Seta S5 present; leg IV with 4 spatulate to knobbed macrosetae...... cracentis (Corpuz and Rimando) Seta S5 absent; leg IV with 8 spatulate to knobbed setae including 4 or 5 macrosetae...2 2. Leg IV with 4 macrosetae: genu, tibia, basitarsus, telotarsus ...... orientalis (Narayanan, Kaur and Ghai) Leg IV with 5 macrosetae: femur, genu, tibia, basitarsus, telotarsus....urumanus (Ehara)

46. Paraphytoseius cracentis (Corpuz and Rimando, 1966)

Ptenoseius cracentis Corpuz and Rimando, 1966: 115, fig. 1. [Type loc.: Gamu, Isabela, the Philippines; type hab.: Achyranthes aspera L.] Amblyseius (Paraphytoseius) multidentatus (nec Swirski and Shechter, 1961): Ehara and Lee 1971: 69, figs. 26–31; Ehara and Bhandhufalck 1977: 79, figs. 131–133. Paraphytoseius cracentis: Schicha and Corpuz-Raros 1985: 69, figs. 10–18; Ho and Lo 1989: 94, fig. 3; Wu et al. 1997b: 133, fig. 99; Ehara et al. 2000: 114, figs. 1–8; Chant and McMurtry 2003b: 220, figs. 99–103.

Genu II with macroseta; genua I and III without macrosetae. Distribution. Japan (Honshu, Kyushu); China, Taiwan, Philippines, Thailand, Singapore, New Caledonia, Papua New Guinea. Japanese Amblyseiinae 23

47. Paraphytoseius orientalis (Narayanan, Kaur and Ghai, 1960) (Japanese name: Uruma-kaburidani-modoki)

Typhlodromus (Amblyseius) orientalis Narayanan, Kaur and Ghai, 1960: 394, fig. 6 (2), a–g. [Type loc.: Chembur, Bombay, India; type hab.: Ipomea and cotton (syntypes)] Paraphytoseius multidentatus Swirski and Shechter, 1961: 114, figs. 7, 26–28; Schicha and Corpuz-Raros 1985: 67, figs. 1–9; Ho and Lo 1989: 91, fig. 1; Wu et al. 1997b: 134, fig. 100. [Type loc.: Tai Po (N. T.), Hong Kong; type hab.: Bambusa sp. Synonymy by Chant and McMurtry (2003b). Paraphytoseius multidentatus (in part): Ehara and Amano 1998: 47, figs. 34, 58; Ehara et al. 2000: 116, figs. 9–16. Paraphytoseius orientalis: Chant and McMurtry 2003b: 216.

Genua I and II with macroseta; genu III without macroseta. Leg IV with 4 spatulate to knobbed, hyaline-tipped macrosetae: genu, tibia, basitarsus, telotarsus (Ehara et al. 2000). Leg IV also with 1 spatulate non-macroseta each on femur, genu, tibia, and basitarsus. The measurements and illustrations of Kyushu specimens were shown in Ehara et al. (2000). Distribution. Japan (Honshu, Kyushu); China, Taiwan, Philippines, India, Pakistan, Madagascar, Africa, Costa Rica, New Caledonia.

48. Paraphytoseius urumanus (Ehara, 1967) (Figs. 17–23)

Amblyseius (Paraphytoseius) urumanus Ehara, 1967a: 76, figs. 31–35. [Type loc.: Tomigusuku, Okinawa Island; type hab.: Ficus erecta Thunb.] Paraphytoseius multidentatus (nec Swirski and Shechter, 1961): Ehara et al. 1994: 139; Ehara and Amano 2002: 323. Paraphytoseius multidentatus (in part): Ehara and Amano 1998: 47, figs. 34, 58; Ehara et al. 2000: 116, figs. 9–16.

Female. Spermatheca disc-shaped (Figs. 17, 18). Fixed digit of chelicera with about 8 teeth; movable digit bidentate (Fig. 19). Macrosetae on legs as figured (Figs. 20–23); genu III without macroseta. Setal lengths (specimens from Ishigaki I.; mean±S.E., n=10): j1 34.8±0.3, j3 75.9±0.9, j4 4.1±0.2, j5 4.5±0.2, j6 5.5±0.2, J5 3.5±0.1, z2 9.8±0.3, z4 10.3±0.3, z5 4.7±0.1, Z1 6.0±0.1, Z4 64.3±0.7, Z5 87.2±1.5, s4 116.9±1.8, r3 44.0±0.4, R1 31.4±0.5, JV5 66.3±1.4; macrosetae on legs: genu I 6.7±0.2, genu II 10.8±0.3, femur IV 25.0±0.6, genu IV 29.4±0.1, tibia IV 37.4±0.4 basitarsus IV 47.9±0.3, telotarsus IV 36.3± 0.4. Paraphytoseius urumanus is readily recognized by having leg IV with 5 knobbed to spatulate, hyaline-tipped macrosetae: femur, genu, tibia, basitarsus, telotarsus. Leg IV also with 1 knobbed to spatulate non-macroseta each on genu, tibia, and basitarsus. P. urumanus is similar to P. santurcensis De Leon, 1965, described from Puerto Rico in having 5 macrosetae on leg IV, but differs in having much longer main idiosomal setae, j3, s4, Z4, Z5 and JV5. Amblyseius (P.) urumanus was synonymized with multidentatus by Matthysse and Denmark (1981), but Chant and McMurtry (2003b) rejected this synonymy on account of seta R1 of urumanus longer and serrate. However, length of seta R1 (female) was described 24 Shôzô EHARA and Hiroshi AMANO

Figs. 17–23. Paraphytoseius urumanus (♀). 17, 18, spermatheca; 19, chelicera; 20, genu II; 21, genu III; 22, femur, genu and tibia of leg IV; 23, tarsus of leg IV. as 20 to 33 µm in the original description of multidentatus, while it was described as 35 µm in the type series of urumanus. In the original description of orientalis the length of seta R1 was not given. According to our measurement, length of seta R1 of female urumanus is 26 to 33 µm (mean±S.E., 31.4±0.5 µm) in Ishigaki specimens and 37 µm in the holotype (Okinawa I.), whilst in females of orientalis (=multidentatus) from Kyushu this seta measures 22 to 28 µm (26.0±0.7 µm) (Ehara et al., 2000). Specimens examined. In addition to the type series, the following specimens were studied: Ishigaki I.: 5 ♀, Momosato, 11–XI–2000 (H. Amano), on Hibiscus tiliaceus L.; 3 ♀, Hirakubo, 11–XI–2000 (H. A.), on Mallotus japonicus (Thunb.) Muell. Arg.; 1 ♀, Ishisuku, 11–XI–2000 (H. A.), on M. japonicus; 18 ♀, Kabira, 13–XI–2000 (H. A.), on H. tiliaceus. Taketomi I.: 6 ♀, Higashi-yashiki, 12–XI–2000 (H. A.), on H. tiliaceus. Iriomote I.: 1 ♀, Uehara, Taketomi, 24–VIII–1984 (collector unknown), on Pueraria montana (Lour.) Merr. Distribution. Japan (Okinawa I., Ishigaki I., Taketomi I., Iriomote I.)

Genus Indoseiulus Ehara, 1982

Indoseius Ghai and Menon, 1969: 347 (nec Indoseius Evans, 1955: 107). Amblyseius (Indoseiulus) Ehara, 1982: 42; McMurtry and Moraes 1984: 29. Indoseiulus: Moraes et al. 1986: 59; Denmark and Kolodochka 1993: 249; Ehara et al. 1994: 139; Ehara and Amano 1998: 48. [Type species: Indoseius ricini Ghai and Menon, 1969, by original designation by Ghai and Menon (1969)]

The genus Indoseiulus is unique in having the peritrematic shields of both sexes not fused Japanese Amblyseiinae 25 anteriorly with dorsal shield. Seta S4 absent; setae J2, Z1, S2 and S5 present. Ventrianal shield poorly sclerotized. Leg IV with 3 macrosetae: genu, tibia, basitarsus.

49. Indoseiulus liturivorus (Ehara, 1982)

Amblyseius (Indoseiulus) liturivirus Ehara, 1982: 43, figs. 10–18; McMurtry and Moraes 1984: 29; Ehara 1985: 120, figs. 25, 26. [Type loc.: Kishigawa, Wakayama Pref., Honshu; type hab.: soy bean (in greenhouse)] Amblyseius (Amblyseius) liturivorus: Tseng 1983: 54, figs. 73–79. Indoseiulus liturivorus: Moraes et al. 1986: 60; Denmark and Kolodochka 1993: 253, figs. 13–17; Ehara et al. 1994: 139; Ehara and Amano 1998: 48, figs. 59–61.

Genua I and III with macroseta; genu II without macroseta. Tibia III with macroseta. Distribution. Japan (Honshu, Shikoku); China, Taiwan.

ACKNOWLEDGEMENTS

We are very grateful to Dr. A. Takafuji (Kyoto University) and Mr. E. Shibata (Nagoya University) for placing specimens of Neoseiulus repletus at our disposal. Our appreciation extends to Messrs. T. Hirose, K. Komi (Kochi Pref.), K. Matsuhira, T. Yamaguchi (Kagoshima Pref.), E. Matsumoto (Kagawa Pref.) and R. Ohnishi (Ehime Pref.), and Miss Y. Shimada (Chiba Pref.) for furnishing some of the present materials. This study was supported in part by Grant-in-Aid from the Ministry of Education, Culture, Sports, Science and Technology of Japan to H. A. (Nos. 11660043 and 14360026).

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摘要 日本産ムチカブリダニ亜科（カブリダニ科）の再検討：属と種のチェックリスト及び検索表 江原 昭三（鳥取市浜坂 2–15–7）・天野 洋（千葉大学園芸学部） 30 Shôzô EHARA and Hiroshi AMANO

カブリダニ科に属する Amblyseiinae（ムチカブリダニ亜科，新称）の日本産種の分類学的 再検討を行い，チェックリストと検索表を掲げた．49 種を取り扱い，これらを Neoseiulus （11），Typhlodromips（14），Typhlodromalus（1），Amblyseius（10），Euseius（3），Proprioseiopsis （2），Okiseius（2），Amblyseiulella（2），Paraphytoseius（3），Indoseiulus（1）の 10 属に分類 した（括弧内は種数）．日本未記録の Neoseiulus repletus（Wu and Li）（ホオノキカブリダニ， 新称）を再記載した．なお，酷似する 2 種，Paraphytoseius urumanus（Ehara）（ウルマカブリ ダニ）と P. orientalis（Narayanan, Kaur and Ghai）（ウルマカブリダニモドキ，新称）は，第 IV 脚の巨大毛の数によって容易に識別できることを明らかにした．