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Amblyseius bellatulus Tseng (Acari: Phytoseiidae): neotype designation with first description of a male

Liao, Jhih-Rong1; Ho, Chyi-Chen2 and Ko, Chiun-Cheng3

1Department of Entomology, National Taiwan University, Taipei City, 10617 Taiwan
2Taiwan Acari Research Laboratory, Taichung City, 41345 Taiwan
3✉ Department of Entomology, National Taiwan University, Taipei City, 10617 Taiwan

2017 - Volume: 57 Issue: 2 pages: 323-335

DOI: 10.1051/acarologia/20164157
ZooBank LSID: 05A3D8B4-BA50-421D-A2B1-DBAD5E4C7F2D
Date received: 2016-07-17, Date accepted: 2016-09-19, Date published: 2017-01-05, Edited by: Tixier, Marie-Stéphane

(CC BY-NC-ND 2.0) © 2017 Liao, Jhih-Rong; Ho, Chyi-Chen and Ko, Chiun-Cheng

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Keywords
Phytoseiids; Taiwan; Amblyseius bellatulus; neotype
Abstract
Yi-Hsiung Tseng provided fundamental information of phytoseiid fauna in Taiwan. However, according to an exclusive interview, all specimens included in Tseng’s collections were lost after his retirement; therefore, more than 20 holotypes are now lost. To ensure stability in nomenclature, designation of neotypes is thus necessary. In this study, we re-described a known species, Amblyseius bellatulus Tseng 1983, by collecting phytoseiid mites from the original type locality and the entirely of Taiwan. Because the morphology of the collected specimens is highly similar to that of specimens used for describing the species, we designated a neotype and redescribed female specimens, as well as male specimens, which have not been described before.


Introduction

Phytoseiid mites have received considerable attention because of their biocontrol potential. Some species can feed on phytophagous mites and small pests (McMurtry et al., 2013). So far, more than 2,700 species included in 91 genera and 3 subfamilies have been recorded worldwide (Chant and McMurtry, 2007; Demite et al., 2015). Fifty-three species are recorded from Taiwan (Ehara, 1970; Lo, 1970; Tseng, 1972; 1973; 1975; 1976; Chang and Tseng, 1978; Tseng, 1983; Ho and Lo, 1989; Ho et al., 2003). Yi-Hsiung Tseng provided the most complete survey of phytoseiid mites in this country from the 1970s to the early 1980s; he recorded 47 species including 20 newly described endemic ones. Collections of Tseng, including type specimens, were deposited at the Tainan Branch of the Bureau of Commodity Inspection and Quarantine (BCIQ) (Tseng, 1983). However, the Tainan Branch of BCIQ no longer possesses Tseng's collections and type specimens. The plant and animal quarantine work was transferred from Bureau of Commodity Inspection and Quarantine to the newly established Bureau of Animal and Plant Health Inspection and Quarantine (BAPHIQ) since 1998 and BCIQ was renamed to Bureau of Standards, Metrology and Inspection (BSMI). Y. H. Tseng retired from BSMI in 2008. The profile column of BAPHIQ Quarterly, a journal issued by BAPHIQ, had interviewed Yi-Hsiung Tseng in 2009 due to his great contribution to plant quarantine and talked about the mite collection of him. Tseng mentioned that he had discarded the whole mite collection of him after his retirement (BAPHIQ, 2009). The present work aims to retrieve specimens collected by Tseng to reconstitute his collection. It is thus the first step of further taxonomical studies.

Some species of Amblyseius are considered as generalist predators, they can feed on wide range preys, and can sometimes feed on pollens (McMurtry et al., 2013). In Taiwan, Ho and Chen (2001) reported that A. bellatulus and A. maai Tseng (=A. tamatavensis Bloommers) fed on melon thrips from cucumbers in the field. Field observation in Taiwan also reveals other that Amblyseius species (e.g., A. eharai Amitai & Swirski, A. herbicolus (Chant)) feed on spider mites and whiteflies. We consider Amblyseius species play an important role in controlling phytophagous mites and small insects. Correct identification and type references are thus essential.

Naming a neotype to ascertain its identity and noting the conditions required by the International Code of Zoological Nomenclature (ICZN) for such designation are necessary. Some examples already exist in the Taxonomy history of the family Phytoseiidae. Duso and Fontana (2001) reported for instance well-known controversy regarding the identity of Phytoseius plumifer (Canestrini & Fanzago) resulting from the loss of type materials. After the original description, many Acarologists subsequently provided controversial descriptions for this species on the basis of their specimens. Duso and Fontana (2001) investigated the type locality and designated a neotype for this species to solve the problem. By contrast, Thor (1930) described Lasioseius (Lasioseius) magnanalis Thor from Svalbard but did not designate any type materials. Chant (1959) moved the species "magnanalis" into family Phytoseiidae but he considered this species dubious because of the vague original description. Kolodochka and Gwiazdowicz (2014) discussed the fate of Thor's specimens as all specimens were lost after he died. They designated a neotype Neoseiulus magnanalis (Thor) to clarify its identity.

To collect the species described by Y. H. Tseng, we conducted a survey in entire Taiwan and surrounding islands. Concurrently, we also tried to collect phytoseiid mites in type localities and type habitat plants. However, we faced some difficulties because of typing errors in the names of the localities of collection, the old vague phonetic English transcription of the localities, and the use of general descriptive terms for plants (e.g., weeds, grasses). In addition, landscape greatly changed in the past 30 years. Nonetheless, we have found specimens that are nearly consistent with description of A. bellatulus by Tseng (1983). Tseng (1983) provided a critical description and illustration of A. bellatullus, but there is some lacking information regarding the characters and measurements that are currently used in phytoseiid species descriptions. We thus re-described Amblyseius bellatulus adding characteristics that were not described in Tseng (1983) and we designated the specimens collected as neotype. These specimens were compared to the elements provided in Tseng (1983). In addition, a description of the male (unknown until now) is provided.

Materials and methods

The specimens examined in this study were collected from various plants. Specimens were mounted in Hoyer's medium; also poor condition specimens were soaked with water, bleached by high concentration (50%) H2O2 until the cuticle color change back to normal condition, rinsed in ethanol (75%) (Yeh et al., 2008), and remounted in Hoyer's medium. Specimens examined under an optical microscope (Olympus BX51), and measured using stage-calibrated ocular micrometers and ImageJ 1.47 (Schneider et al., 2012). All measurements provided are in micrometers, neotype measurements are shown in bold type followed by their mean and range (in parentheses). The general terminology used for morphological descriptions in this study follows that of Chant and McMurtry (2007), while for the dorsal and ventral chaetotaxy we followed Rowell et al. (1978) and Chant and Yoshida-Shaul (1991, 1992); for adenotaxy and poroidotaxy terminology we followed Beard (2001).

The neotype and voucher specimens were deposited in the following institutions: ESALQ-USP (Escola Superior de Agricultura Luiz de Queiroz, Piracicaba, State of São Paulo, Brazil) (three females), NCHU (Department of Entomology, National Chung Hsing University, Taichung, Taiwan) (eleven females and one male), NMNS (National Museum of Natural Science, Taichung, Taiwan) (eight females), NTU (Department of Entomology, National Taiwan University, Taipei, Taiwan) (twenty-five females and two males), NPUST (National Pingtung University of Science and Technology, Pintung County, Taiwan) (fifteen females), TARI (Taiwan Agricultural Research Institute, Taichung City, Taiwan) (twenty-eight females), TARL (Taiwan Acari Research Laboratory, Taichung City, Taiwan) (ten females and one male). If necessary, the locality names were translated using the Geographic Name Information System, Department of Land Administration, Ministry of the Interior (Taiwan) ( http://gn.moi.gov.tw/geonames/Translation/Translation.aspx ). The distribution map was prepared using the SimpleMappr program (Shorthouse, 2010), based on the label data of examined material in this study.

Taxonomy
Family Phytoseiidae Berlese

Subfamily Amblyseiinae Muma

Tribe Amblyseiini Muma

Subtribe Amblyseiina Muma

Genus Amblyseius Berlese

Amblyseius bellatulus Tseng, 1983 (Figures 1-4)

Amblyseius (Amblyseius) bellatulus Tseng, 1983: 38.

Neotype Designation

Amblyseius bellatulus was described by Y. H. Tseng in 1983 from Mingchien (now Mingjian Township), Nantou Hsien on Morus alba. As mentioned in the introduction the mite collection of Y. H. Tseng does not longer exist (BAPHIQ, 2009), according to Article 75.3.4 of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 1999), we here designate adult females we collected as neotype of A. bellatulus to clarify its taxonomic status and provide a complete description. Additionally, as male specimens were not known, we provide male description for the first time.

Description: Female (n= 10). Dorsum (Figure 1A) — Idiosomal setal pattern: 10A:9B/JV-3:ZV. Dorsal shield lightly sclerotized, smooth, with lateral reticulation, 382 384 (353 – 449) long, 202 212 (192 – 250) wide at level of s4, 228 228 (212 – 276) wide at level of S4; with seven pairs of solenostomes (gd1, gd2, gd4, gd5, gd6, gd8, gd9), ten pairs of poroids (id1, id1a, id2, id4, is1, idl1, idl3, idl4, idm5, idm6); length of dorsal setae: j1 23 28 (23 – 32), j3 34 40 (34 – 46), j4 5 7 (5 – 13), j5 4 5 (4 – 7), j6 6 7 (4 – 8), J2 4 7 (4 – 9), J5 5 6 (4 – 9), z2 11 10 (8 – 11), z4 11 11 (8 – 13), z5 5 6 (4 – 8), Z1 8 8 (7 – 10), Z4 52 52 (47 – 57), Z5 119 136 (117 – 158), s4 46 49 (43 – 61), S2 10 11 (9 – 13), S4 5 8 (5 – 11), S5 6 7 (6 – 9), r3 21 19 (17 – 21), R1 13 10 (8 – 13). All setae smooth, except Z4 and Z5 slightly serrate.

Venter (Figure 1B) — Sternal shield smooth, posterior margin almost straight, wider than long, 66 69 (62 – 73) long, 87 94 (84 – 114) wide at st3 level, with three pairs of setae st1 25 27 (24 – 29), st2 24 24 (21 – 26), st3 23 23 (17 – 26) and two pairs of poroids (pst1, pst2). Metasternal platelets tear-shaped, with a pair metasternal setae, st4 22 19 (14 – 22), and one pair of poroids (pst3). Genital shield smooth, st5 23 22 (16 – 26), 77 78 (73 – 96) wide. Distances between st1-st1 59 58 (53 – 73), st2- st2 63 63 (59 – 75), st3- st3 78 74 (67 – 80), st1- st3 63 64 (61 – 74), st5- st5 59 64 (58 – 73). Ventrianal shield smooth, almost pentagonal, with slightly waist at JV2 level; 121 131 (121 – 151) long, 100 98 (90 – 117) wide at level of ZV2 and 86 90 (84 – 103) wide at level of anus; with three pairs of pre-anal setae, solenostome gv3 crescentic, JV1 13 15 (11 – 18), JV2 10 13 (10 – 18), ZV2 9 11 (9 – 14). Setae JV4 11 10 (7 – 11), JV5 45 51 (45 – 61), ZV1 15 16 (12 – 19), ZV3 9 10 (9 – 11) on interscutal membrane. All ventral setae smooth. Two metapodal plates 18 20 (18 – 23) long, 4 6 (4 – 7) wide, 12 15 (12 – 17) long, 2 2 (1 – 3) wide.

Peritreme (Figure 1A-B) — Peritreme extending beyond to j1, peritrematic shield smooth, lightly sclerotized, with one pair of solenostome (gd3).

Spermatheca (Figure 1D) — Calyx of spermatheca funnel-shaped and atrium well distinct with neck, embolus not visible, with a thin major duct.

Chelicera (Figure 1C) — Movable digit 31 32 (30 – 36) long, with three teeth; fixed digit 27 29 (26 – 34), with 10 teeth, pilus dentilis.

Figure 1. Amblyseius bellatulus Female: A – Dorsal shield; B – Ventral idiosoma; C – Chelicera; D – Spermatheca.

Legs (Figure 2) — Coxal formula 2, 2, 2, 1. Chaetotaxy (femur to basitarsus): leg I, 2-3/1-2/2-2, 2-2/1-2/1-2, 2-2/1-2/1-2, 1-1/1-1; leg II, 2-3/2-2/0-1, 2-2/0-2/0-1, 1-1/1-2/1-1, 1-1/1-1; leg III, 1-2/1-1/0-1, 1-2/1-2/0-1, 1-2/1-1/1-1, 1-1/1-1; leg IV, 1-2/1-1/0-1, 1-2/1-2/0-1, 1-1/1-2/0-1, 1-1/1-1. Macrosetae: Sge II (pd2) 27 29 (26 – 33), Sge III (ad2) 30 31 (27 – 37), Sge IV (ad2) 50 50 (40 – 59), Sti IV (ad) 45 46 (40 – 51), St IV (pd) 73 73 (66 – 84).

Figure 2. Amblyseius bellatulus Female: Legs (A – leg I dorso-anterior view; B – leg II dorso-anterior view; C – leg III dorsal view; D – leg IV dorso-anterior view).

Male (n= 2). Dorsum (Figure 3A) — Idiosomal setal pattern: 10A:9B/JV-3, 4:ZV-1, 3. Dorsal shield lightly sclerotized, smooth, with lateral reticulation, 264 (248 – 280) long, 178 (161 – 195) wide at level of s4, 138 (127 – 149) wide at level of S4; with seven pairs of solenostomes (gd1, gd2, gd4, gd5, gd6, gd8, gd9), ten pairs of poroids (id1, id1a, id2, id4, is1, idl1, idl3, idl4, idm5, idm6); length of dorsal setae: j1 20 (20 – 2), j3 28 (28 – 29), j4 5.4 (5 – 6), j5 5 (4 – 5), j6 5 (4 – 6), J2 5 (5 – 5), J5 5 (4 – 6), z2 10 (10 – 11), z4 9 (8 – 10), z5 4 (4 – 4), Z1 9 (8 – 9), Z4 35 (35 – 35), Z5 81 (77 – 85), s4 34 (34 – 34), S2 9 (9 – 9), S4 7 (7 – 7), S5 6 (6 – 6), r3 15 (15 – 16), R1 9 (7 – 11). All setae smooth, except Z4 and Z5 slightly serrate.

Venter (Figure 3B) — Sternogenital shield smooth, with slightly lateral reticulation, posterior margin almost straight, longer than wide 116 (110 – 123) long, 64 (55 – 73) wide at level st2, with five pairs of setae st1 18 (16 – 19), st2 18 (16 – 20), st3 16 (13 – 19), st4 14 (11 – 17), st5 14 (13 – 14) and three pairs of poroids (pst1, pst2, pst3); distance between st1-st1 42 (40 – 43), st2-st2 49 (46 – 52), st3-st3 50 (47 – 54), st4-st4 38 (36 – 40), st5-st5 30 (27 – 33), st1-st5 102 (97 – 106). Ventrianal shield subtriangular, with slightly reticulation, 108 (103 – 114) long, 137 (129 – 144) wide at level of anterior corner and 61 (61 – 62) wide at level of anus, fused with peritrematic shield cingulum; with three pairs of pre-anal setae, solenostome gv3 crescentic, JV1 9 (9 – 9), JV2 10 (10 – 10), ZV2 8 (8 – 8); JV5 20 (17 – 22) on interscutal membrane. All ventral setae smooth.

Peritreme (Figure 3A-B) — Peritreme extending to seta j1 level; peritrematic shield lightly sclerotized, with one pair of solenostome (gd3).

Chelicera (Figure 3C-D) — Movable digit 21 (21 – 22) long, with one tooth; fixed digit 22 (20 – 24), with 8 teeth, pilus dentilis. Spermatodactyl U-shaped, shaft 34 (30 – 37) long, heel rounded, foot 18 (17 – 19) long, with rounded toe and lateral thorn-like projection.

Figure 3. Amblyseius bellatulus Male: A – Dorsal shield; B – Ventral idiosoma; C – Chelicera; D – Ventral view of spermatodactyl.

Legs (Figure 4) — Coxal formula 2, 2, 2, 1. Chaetotaxy (femur to basitarsus): leg I, 2-3/1-2/2-2, 2-2/1-2/1-2, 2-2/1-2/1-2, 1-1/1-1; leg II, 2-3/2-2/0-1, 2-2/0-2/0-1, 1-1/1-2/1-1, 1-1/1-1; leg III, 1-2/1-1/0-1, 1-2/1-2/0-1, 1-2/1-1/1-1, 1-1/1-1; leg IV, 1-2/1-1/0-1, 1-2/1-2/0-1, 1-1/1-2/0-1, 1-1/1-1. Macrosetae: Sge II (pd2) 20 (19 – 20), Sge III (ad2) 20 (20 – 20) Sge IV (ad2) 29 (28 – 30), Sti IV (ad) 26 (25 – 26), St IV (pd) 56 (56 – 56).

Figure 4. Amblyseius bellatulus Male: Legs (A – leg I dorso-anterior view; B – leg II dorso-posterior view; C – leg III dorso-posterior view; D – leg IV dorso-posterior view).

Material examined — Neotype female: TARI, Wufeng District, Taichung, from Toona sinensis, 17.xii.2001, C. C. Ho (NTU).

Additional material examined — Hualien County, one female from rice leaf sheath, 2.xi.1977, K. C. Lo (TARL); Wanfeng Village, Wufeng District, Taichung City, one female from Polygonum perfoliatum, 3.ii.1978, C. C. Ho (TARI); Wanfeng Village, Wufeng District, Taichung City, one female from Bothriochloa ischaemum, 15.ii.1978, C. C. Ho (TARI); Wanfeng Village, Wufeng District, Taichung City, one female from mushroom, 15.xii.1981, K. C. Lo (TARI); TARI, Wufeng District, Taichung City, two females from Zea mays, 18.x.1985, C. C. Ho (TARL); TARI, Wufeng District, Taichung City, 17 females 1 male from Ageratum conyzoides, 22.xi.1985, C. C. Ho (TARI); NPIA, two females from Ageratum conyzoide, 22.xi.1985, C. C. Ho (TARL); Erlin Township, Changhua County, one female from Vigna unguiculata, 14.xii.1986, S. M. Yu (NPUST); Mingjian Township, Nantou County, two females from Ampelopsis brevipedunculata, 24.xii.1985, C. C. Ho (NTU); TARI, Wufeng District, Taichung City, one female from Zea mays, 4.vii.1987, C. C. Ho (TARI); Baihe District, Tainan City, one female from Sambucus formosana, 23.vi.1988, S. M. Yu (NMNS); TARI, Wufeng District, Taichung City, one female from Ageratum houstonianum, 12.x.1988, S. M. Yu (NMNS); TARI, Wufeng District, Taichung City, two females from Alternanthera sessilis, 17.x.1988, S. M. Yu (NPUST); TARI, Wufeng District, Taichung City, one female from Eclipta prostrata, 17.x.1988, S. M. Yu (NMNS); TARI, Wufeng District, Taichung City, nine females from Polygonum perfoliatum, 17.x.1988, S. M. Yu (NCHU); TARI, Wufeng District, Taichung City, two females (MZLQ-7644, 7645) from Arachis hypogaea, 17.x.1988, S. M. Yu (ESALQ-USP); TARI, Wufeng District, Taichung City, one female (MZLQ-7643) from unknown plant, 18.x.1988, S. M. Yu (ESALQ-USP); Caotun Township, Nantou County, three females from Psidium guajava, 27.x.1988, S. M. Yu (NTU); TARI, Wufeng District, Taichung City, two females from Ipomoea acuminata, 27.x.1988, S. M. Yu (NTU); TARI, Wufeng District, Taichung City, three females from Zingiber officinale, 2.xi.1988, S. M. Yu (TARL); TARI, Wufeng District, Taichung City, one female from Monorfiae choartiae, 2.xi.1988, S. M. Yu (TARL); Caotun Township, Nantou County, 11 females from Psidium guajava, 2.xi.1988, S. M. Yu (NPUST); TARI, Wufeng District, Taichung City, two females from Ageratum houstonianum, 16.xi.1988, S. M. Yu (NTU); TARI, Wufeng District, Taichung City, two females from Ipomoea acuminata, 24.xi.1988, S. M. Yu (NTU); TARI, Wufeng District, Taichung City, one female from Rorippa indica, 30.xi.1988, S. M. Yu (TARL); TARI, Wufeng District, Taichung City, two females one male from Oryza sativa, 2.xii.1988, S. M. Yu (NCHU); TARI, Wufeng District, Taichung City, one female from Ziziphus mauritiana, 7.xii.1988, C. C. Ho (TARI); Sanxiantai, Chenggong Township, Taitung County, one male from Calophyllum inophyllum, 4.iv.1989, C. C. Ho (TARL); TARI, Wufeng District, Taichung City, one female from Solanum melongena 25.vi.1991, C. C. Ho (NMNS); TARI, Wufeng District, Taichung City, two females from Solanum melongena 4.vi.1991, C. C. Ho (TARI); TARI, Wufeng District, Taichung City, one female from Solanum melongena 11.vi.1991, C. C. Ho (NMNS); TARI, Wufeng District, Taichung City, three females rear from unknown plant, 3.vii.1991, C. C. Ho (TARI); TARI, Wufeng District, Taichung City, one female from Solanum melongena 4.vii.1991, C. C. Ho (NPUST); TARI, Wufeng District, Taichung City, three females from Solanum melongena 6.vii.1991, C. C. Ho (NMNS); TARI, Wufeng District, Taichung City, nine females two males from Solanum melongena, 16.vii.1991, C. C. Ho (NTU); TARI, Wufeng District, Taichung City, one female from Solanum melongena, 23.vii.1991, C. C. Ho (NTU); TARI, Wufeng District, Taichung City, one female rear from unknown plant, 6.xi.1993, C. C. Ho (NTU); Lucao Township, Chiayi County, one female from Lagenaria siceraria soil, 14.viii.2001, C. C. Ho (NTU); Dayuan District, Taoyuan City, one female from Allium fistulosum soil, 25.x.2001, C. C. Ho (NTU).

Remarks — Tseng (1983) described this species by using a single female specimen collected from Mingchien (now Mingjian Township), Nantou Hsien, on Morus alba. We collected two female specimens from the type locality on Ampelopsis brevipedunculata, but they were in poor condition. Consequently, the neotype specimen was selected from among the entire collection depending on the slide condition. The neotype locality is Wufeng District, Taichung, which is not far from the type locality (Figure 5).

Figure 5. The distribution map of Amblyseius bellatulus from Taiwan.

The females of this species are unique among the known Amblyseius species from Taiwan because of the following characters: dorsal shield with lateral reticulation, ventrianal shield pentagonal with slightly lateral concaved, calyx of spermatheca funnel-shaped. A comparison of different body measurements among the holotype, neotype, and remaining specimens of A. bellatulus is provided in Table 1. Most specimens were similar to the neotype, with little morphological differences. We observed only one specimen with a larger body size and longer idiosomal setae (e.g. dorsal shield 449 long, 250 wide). Tseng (1983) reported the chaetotaxy of genu I-IV: 2-2/1-2/1-2, 1-2/1-2/0-1, 1-2/1-2/0-1, 1-2/1-2/0-1". We observed the chaetotaxy of leg I-IV and found that the chaetotaxy of genu I-IV differs from the original description: 2-2/1-2/1-2, 2-2/0-2/0-1, 1-2/1-2/0-1, 1-2/1-2/0-1. With regard to adenotaxy and poroidotaxy, Tseng (1983) reported "at least five pairs of pores on the dorsal shield". On the basis of his illustration, we observed three pairs of solenostomes (gd6, gd8, gd9). In the present study, we observed seven pairs of solenostomes (gd1, gd2, gd4, gd5, gd6, gd8, gd9), and ten pairs of poroids (id1, id1a, id2, id4, is1, idl1, idl3, idl4, idm5, idm6). The differences between the reported and observed adenotaxy and poroidotaxy may be due to the quality of microscopes.

Table 1. Comparisons on measurements1 (μm) of Amblyseius bellatulus between (1) holotype (in Tseng 1983), (2) presently studied neotype and (3) ten females including neotype

By comparing the collection records of this A. bellatulus, we observed that this species was mostly collected from central Taiwan; however, one specimen was collected from northern Taiwan and another from eastern Taiwan. Therefore, this species may be more widely distributed than is evident from the collection. We also observed A. bellatulus occurred on various plants, although these mites mostly inhabit weeds, they also inhabit some economically valuable plants (e.g., corn plants, eggplants, and guava). Ho and Chen (2001) reported that A. bellatulus could prey on melon thrips (Thrips palmi) in the field. Therefore, we consider A. bellatulus need further experiments for confirmation of the biocontrol potential.

Acknowledgements

We would like to thank H. T. Shih (TARI, Taiwan) providing equipment and useful advises for illustrations. We thank Y. N. Lee (NTU, Taiwan) for digital illustration, A. K. Dubey (FRI, India), M. J. Yang (NCYU, Taiwan), M. Y. Tsai (TFRI, Taiwan), J. F. Hsieh, M. J. Chiu, Y. Hsiao (NTU, Taiwan), S. F. Lin (NCHU, Taiwan) for their valuable suggestions. We are also thankful to Dr. G. J. de Moraes (ESALQ-USP, Brazil) and Dr. G. A. Evans (USDA, USA) for their encouragement to the first author. Thanks are extended to anonymous reviewers for help in improving the manuscript, to M. Andrews and Y. Sanyang for English editing of the draft. The study is supported by a grant (MOST105-2621-B-002-002-MY3) from the Ministry of Science and Technology, Taiwan.

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Photos: Varroa destructor (Varroidae) on a honey bee. Low temperature -Scanning electron microscopy photograph. Material collected and fixed for LT-SEM: J. Pettis, USDA-ARS-Bee Lab. and R. Ochoa, USDA-ARS-SEL; LT-SEM photo: E. Erbe and C. Pooley, USDA-ARS-CEMU © USDA; Petrobia harti (Tetranychidae) © INRA - Alain Migeon; Dermanyssus gallinae (Dermanysssidae) © Ecole Nationale Vétérinaire de Lyon - Lise Roy; Adult Ixodes sp. (Ixodidae) engorging on an Atlantic puffin Fratercula arctica © Andy Darrington; Panonychus ulmi (Tetranychidae): hatching of a winter egg © Université de Lausanne - Centre de microscopie électronique - Cazelles
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