J. Acarol. Soc. Jpn., 25(S1): 19-25. March 25, 2016 © The Acarological Society of Japan http://www.acarology-japan.org/ 19

Histiostomatids on common Dermaptera in Japan

Kazumi TAGAMI* University of Tsukuba, Institute of Health and Sports Sciences, 1-1-1 Tennodai, Tsukuba, Ibaraki 305-8574, Japan

ABSTRACT

In 2013, phoresy of Histiostoma mahunkai Fain, 1974, on the Japanese common , Anisolabella (Gonolabis) marginalis (Dohrn, 1864), was recorded in Ibaraki Prefecture, central Japan. To investigate the extent of this phenomenon, I studied A. marginalis and Anisolabis maritima (Bonelli, 1832) collected in Kochi (Kochi Pref.), Matsue (Shimane Pref.), Yonago (Tottori Pref.), Echizen (Fukui Pref.), Shizuoka (Shizuoka Pref.), Misato and Iwatsuki (Saitama Pref.), and Mito (Ibaraki Pref.). Phoretic histiostomatids were isolated, reared, and identified as H. mahunkai and H. piscium. Key words: phoresy, earwig, , Histiostoma, Astigmata

INTRODUCTION

Some Dermaptera carry numerous deutonymphs of histiostomatids in an association known as phoresy. The best known association is Histiostoma polypori (Oudemans), 1914 (: Histiostomatidae), on auricularia L., 1758 (Dermaptera: ), a herbivorous, ground-dwelling, cosmopolitan earwig (Oudemans, 1914; Behura, 1957; Chmielewski, 1984, 2009, 2010). Histiostoma feroniarum (Dufour, 1839) is also found on this host (Rebolledo and Arroyo, 1996). Other combinations are also known: Labidura riparia (Labiduridae) and Histiostoma sp. (Strandberg and Tucker, 1974); L. riparia and H. camphori Eraky, 1999 (Negm and Alatawi, 2011); Titanolabis colossea () and H. humiditatis (Vitzthum, 1927), H. australiense Mahunka, 1975, H. feroniarum (Dufour, 1839), and H. titanolabi Tagami and Halliday, 2013 (Tagami and Halliday, 2013); and Anisolabella (Gonolabis) marginalis (Dohrn, 1864) (Anisolabididae, Anisolabididae) and H. mahunkai Fain, 1974 (Tagami, 2013). Phoretic associations first occurred at least 49 million years ago (Dunlop et al., 2012). Japan is especially interesting in this respect because it is geographically distant from the Dermapteran ancestral area in Africa or southern Gondwana (Popham, 2000; Jarvis et al., 2005). This paper reports a survey of the diversity of Histiostoma species phoretic on Japanese .

Tetsuo GOTOH and DeMar TAYLOR (eds.), Acarology XIV: Proceedings of the International Congress. Journal of Acarological Society of Japan 25 (Suppl. 1): 1-192. * Corresponding author: e-mail: [email protected]; [email protected] DOI: 10.2300/acari.25.Suppl_19 20 Kazumi TAGAMI

MATERIALS AND METHODS

My colleagues and I collected earwigs (Dermaptera) outside Kochi Castle Park, Kochi Pref. (collected 2011 09 30); in Matsue Castle Park, Shimane Pref. (2012 09 29); in a private garden in Echizen, Fukui Pref. (2012 10 10); at Hirota Shrine, Yonago, Tottori Pref. (2013. 10. 31); in Shizuoka Gokoku-Shrine Park, Shizuoka Pref. (2013 09 28); in Hinuma Natural Park, Mito, Ibaraki Pref. (2012 09 16); and in Misato Park (2012 04 29) and Iwatsuki Castle Park, Saitama Pref. (2012 04 15). All specimens were collected beneath rotten wood or leaf litter. I identified the species. The earwigs were placed in tubes with wet paper for transport to the laboratory. Each was then killed by decapitation and maintained on a moist 42.5-mm paper disc with powdered yeast in a screw-cap jar (60 ml, Nalgene, USA) to avoid drying. Two to three days later, feeding stages of were detected in the culture (Tagami and Kuwahara, 2005; Tagami, 2013). Collected specimens were cleared in Keifer’s clearing solution (Keifer, 1953) at 50℃, mounted on a 24-mm ×24-mm cover glass in gum-chloral medium, and covered with a 15- or 18-mm round cover glass. The two cover glasses were sealed with paint to retain moisture and mounted on a metal arm with an 18-mm central hole (Tagami and Kuwahara, 2005). All specimens will be deposited in the Collection of the Department of Zoology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba, Ibaraki 305-0005, Japan: Kochi 01 to 05 (NSMT accession numbers 14160 to 14164), Matsue 1-1 to 1-6 (14165 to 14170), Matsue 2-1 to 2-6 (14171 to 14176), Yonago 01 to 05 (14177 to 14181), Echizen 01 to 05 (14182 to 14186), Shizuoka 01 to 04 (14187 to 14190), Mito 01 to 06 (14191 to 14196), Misato 01 to 06 (14197 to 14202), and Iwatsuki 01 to 04 (14203 to 14206). Four to six deutonymph specimens from each earwig species were identified and measured. Characteristics of adult mites specimens were used as additional evidence for identification. The following features were measured: relative length of a structure to idiosomal length (RLI), total leg I length (L1), total leg II length (L2), tarsus I length (T1), tarsus II length (T2), 1a–1a distance

(D1 a), and 3a–3 a distance (D3 a). The attachment organ width (Wao, from the center of the median left sucker to the center of the right) was calculated (Kurosa, 1987).

RESULTS

The earwigs from Kochi, Mito, Matsue 1, Echizen, and Shizuoka were identified as A. marginalis, and those from Iwatsuki, Misato, Matsue 2, and Yonago as Anisolabis maritima (Bonelli, 1832) (Fig. 1). These earwigs are very common throughout Japan. From their measurements (Table 1), the histiostomatids were identified as H. mahunkai, recorded previously from soil in the Ivory Coast, Africa (Fain, 1974), and from the earwig A. marginalis from Tsuchiura, Ibaraki, Japan (Tagami, 2013); and as Histiostoma piscium Fain and Lambrechts, 1985, described from a swim-bladder collected from a dead iridescent shark (Pangasius sutchi) in an aquarium, and from a Japanese cockroach, Periplaneta fuliginosa (Tagami, 2007). Histiostoma mahunkai was isolated from A. marginalis collected in Matsue 1, Japanese histiostomatids phoretic on Dermaptera 21

Fig. 1. Localities of Dermaptera sampled for mites.

Echizen, and Shizuoka, and from A. maritima in Matsue 2. Histiostoma piscium was isolated from A. marginalis collected in Mito, and from A. maritima in Kochi, Yonago, Misato, and Iwatsuki.

DISCUSSION

The Dermapteran ancestor emerged in Africa at least 200 million years ago, and diversified into seven families, with 2000 species distributed worldwide. In addition to F. auricularia, only three other species of earwig (Anisolabella marginalis, Anisolabididae; and Titanolabis colossea and Labidura riparia, Labiduridae) are known as hosts of phoretic histiostomatid mites. Two very common anisolabiid earwigs, Anisolabella marginalis and Anisolabis maritima, are 22

Table 1. Measurements of idiosomal length (µm), and relative lengths (RLI) of leg 1 (L1), leg II (L2), tarsus I (T1), tarsus II (T2),

1 a–1 a distance (D1 a), 3 a–3 a distance (D3 a), and width of attachment organ (Wao). Locality and Idiosomal L (RLI) L (RLI) T (RLI) T (RLI) D (RLI) D (RLI) W (RLI) slide No. length (µm) 1 2 1 2 1 a 3 a ao Kochi 01 160 145.0 (90.6) 92.5 (57.8) 65.0 (40.6) 45.0 (28.1) 37.5 (23.4) 21.3 (13.3) 30.0 (18.8) Kochi 02 162.5 120.0 (73.8) 90.0 (55.4) 70.0 (43.1) 45.0 (27.7) 37.5 (23.1) 21.3 (13.1) 32.5 (20.0) Kochi 03 142.5 102.5 (71.9) 82.5 (57.9) 58.8 (41.3) 42.5 (29.8) 33.8 (23.7) 20.0 (14.0) 30.0 (21.1) Kochi 04 150 116.3 (77.5) 87.5 (58.3) 65.0 (43.3) 43.8 (29.2) 35.0 (23.3) 20.0 (13.3) 30.0 (20.0) Kochi 05 168.8 115.0 (68.1) 88.8 (52.6) 65.0 (38.5) 43.8 (25.9) 38.8 (23.0) 22.5 (13.3) 31.3 (18.5) 119.8±15.6 88.2±3.71 64.8±4.0 44.0±1.0 36.5±2.1 21.0±1.1 30.8±1.1 Mean±SD 156.8±10.5 (76.4±8.6) (56.4±2.4) (41.4±2.0) (28.2±1.5) (23.3±0.3) (13.4±0.4) (19.7±1.0) Matsue 1-1 158.8 135.0 (85.0) 100.0 (63.0) 75.0 (47.2) 50.0 (31.5) 40.0 (25.2) 23.8 (15.0) 35.0 (22.0) Matsue 1-2 172.5 155.0 (89.9) 110.0 (63.8) 82.5 (47.8) 57.5 (33.3) 42.5 (24.6) 26.3 (15.2) 37.5 (21.7) Matsue 1-3 162.5 137.5 (84.6) 97.5 (60.0) 75.0 (46.2) 52.5 (32.3) 40.0 (24.6) 23.8 (14.6) 35.0 (21.5) Matsue 1-4 160 130.0 (81.3) 97.5 (60.9) 72.5 (45.3) 50.0 (31.3) 38.8 (24.3) 22.5 (14.1) 35.0 (21.9)

Matsue 1-5 160 130.0 (81.3) 102.5 (64.1) 72.5 (45.3) 52.5 (32.8) 37.5 (23.4) 22.5 (14.1) 32.5 (20.3) Kazumi T Matsue 1-6 167.5 142.5 (85.1) 105.0 (62.7) 77.5 (46.3) 53.8 (32.1) 41.3 (24.7) 23.8 (14.2) 35.0 (20.9) 138.3±9.4 102.1±4.9 75.8±3.8 52.7±2.8 40.0±1.8 23.8±1.4 35.0±1.6 Mean±SD 163.6±5.4 (84.5±3.2) (62.4±1.6) (46.4±1.0) (32.2±0.8) (24.5±0.6) (14.5±0.5) (21.4±0.7) AGAMI Matsue 2-1 180 155.0 (86.1) 117.5 (65.3) 87.5 (48.6) 60.0 (33.3) 42.5 (23.6) 25.0 (13.9) 36.3 (20.2) Matsue 2-2 175 147.5 (84.3) 112.5 (64.3) 82.5 (47.1) 56.3 (32.2) 41.3 (23.6) 23.8 (13.6) 33.8 (19.3) Matsue 2-3 172 140.0 (81.4) 107.5 (62.5) 77.5 (45.1) 52.5 (30.5) 42.5 (24.7) 25.0 (14.5) 33.8 (19.7) Matsue 2-4 182 145.0 (79.7) 110.0 (60.4) 81.3 (44.7) 53.8 (29.6) 42.5 (23.4) 25.0 (13.7) 35.0 (19.2) Matsue 2-5 176.3 153.8 (87.2) 117.5 (66.6) 85.0 (48.2) 57.5 (32.6) 43.8 (24.8) 26.3 (14.9) 37.5 (21.3) Matsue 2-6 170 137.5 (80.9) 106.3 (62.5) 75.0 (44.1) 52.5 (30.9) 40.0 (23.5) 23.8 (14.0) 35.0 (20.6) 146.5±7.1 111.9±4.8 81.5±4.6 55.4±3.0 42.1±1.3 24.8±0.9 35.2±1.4 Mean±SD 175.9±4.6 (83.3±3.1) (63.6±2.2) (46.3±1.9) (31.5±1.4) (23.9±0.7) (14.1±0.5) (20.0±0.8) Yonago 01 156.3 115.0 (73.6) 92.5 (59.2) 65.0 (41.6) 45.0 (28.8) 38.8 (24.8) 22.5 (14.4) 30.0 (19.2) Yonago 02 157.5 107.5 (68.3) 85.0 (54.0) 60.0 (38.1) 40.0 (25.4) 40.0 (25.4) 23.8 (15.1) 30.0 (19.0) Yonago 03 165 116.3 (70.5) 92.5 (56.1) 65.0 (39.4) 47.5 (28.8) 41.3 (25.0) 23.8 (14.4) 30.0 (18.2) Yonago 04 170 122.5 (72.1) 93.8 (55.2) 68.8 (40.5) 47.5 (27.9) 40.0 (23.5) 22.5 (13.2) 30.0 (17.6) Yonago 05 170 122.5 (72.1) 97.5 (57.4) 57.5 (33.8) 47.5 (27.9) 41.3 (24.3) 22.5 (13.2) 31.3 (18.4) 116.8±6.2 92.3±4.5 63.3±4.5 45.5±3.3 40.3±1.1 23.0±0.7 30.3±0.6 Mean±SD 163.8±6.6 (71.3±2.0) (56.4±2.0) (38.7±3.0) (27.8±1.4) (24.6±0.7) (14.1±0.8) (18.5±0.6) Echizen 01 166.3 － 110.0 (66.1) － 56.3 (33.9) 43.8 (26.3) 22.5 (13.5) 36.3 (21.8) Echizen 02 162.5 140.0 (86.2) 105.0 (64.6) 77.5 (47.7) 53.8 (33.1) 42.5 (26.2) 22.5 (13.8) 32.5 (20.0) Echizen 03 162.5 136.3 (83.9) 105.0 (64.6) 77.5 (47.7) 50.0 (30.8) 41.3 (25.4) 23.8 (14.6) 32.5 (20.0) Echizen 04 175 145.0 (82.9) 112.5 (64.3) 75.0 (42.9) 57.5 (32.9) 45.0 (25.7) 25.0 (14.3) 33.8 (19.3) Echizen 05 155 127.5 (82.3) 97.5 (62.9) 70.0 (45.2) 48.8 (31.5) 37.5 (24.2) 21.3 (13.7) 32.5 (21.0) 137.2±7.4 106.0±5.8 75.0±3.5 53.3±3.8 42.0±2.9 23.0±1.4 33.5±1.7 Mean±SD 164.3±7.3 (83.8±1.7) (64.5±1.2) (45.9±2.3) (32.4±1.3) (25.6±0.8) (14.0±0.5) (20.4±1.0) Shizuoka 01 150 127.5 (85.0) 97.5 (65.0) 72.5 (48.3) 48.8 (32.5) 40.0 (26.7) 23.8 (15.9) 30.0 (20.0) Shizuoka 02 150 125.0 (83.3) 97.5 (65.0) 70.0 (46.7) 47.5 (31.7) 38.8 (25.9) 21.3 (14.2) 30.0 (20.0) Shizuoka 03 167.5 135.0 (80.6) 105.0 (62.7) 75.0 (44.8) 51.3 (30.6) 40.0 (23.9) 23.8 (14.2) 32.5 (19.4) Shizuoka 04 － 125.0 (－) 87.5 (－) 68.8 (－) 41.3 (－) － (－) － (－) － (－) 128.1±4.7 96.9±7.2 71.6±2.8 47.2±4.3 39.6±0.7 23.0±1.4 30.8±1.4 Mean±SD 155.8±10.1 (83.0±2.2) (64.2±1.3) (46.6±1.8) (31.6±1.0) (25.5±1.4) (14.8±1.0) (19.8±0.3) Mito 01 152.5 120.0 (78.7) 95.0 (62.3) 65.0 (42.6) 45.0 (29.5) 37.5 (24.6) 23.8 (15.6) 28.8 (18.9) Mito 02 158.8 121.3 (76.4) 96.3 (60.6) 70.0 (44.1) 46.3 (29.2) 38.8 (24.4) 23.8 (15.0) 31.3 (19.7)

Mito 03 145 116.3 (80.2) 88.8 (61.2) 65.0 (44.8) 45.0 (31.0) 36.3 (25.0) 22.5 (15.5) 30.0 (20.7) Japanese histiostomatidsphoreticonDermaptera Mito 04 143.8 107.5 (74.8) 82.5 (57.4) 60.0 (41.7) 40.0 (27.8) 37.5 (26.1) 22.5 (15.6) 28.8 (20.0) Mito 05 157.5 120.0 (76.2) 91.3 (58.0) 65.0 (41.3) 45.0 (28.6) 37.5 (23.8) 22.5 (14.3) 31.3 (19.9) Mito 06 157.5 125.0 (79.4) 95.0 (60.3) 72.5 (46.0) 48.8 (31.0) 38.8 (24.6) 23.8 (15.1) 32.5 (20.6) 118.4±6.0 91.5±5.2 66.3±4.4 45.0±2.9 37.7±0.9 23.2±0.7 30.5±1.5 Mean±SD 152.5±6.7 (77.6±2.1) (60.0±1.9) (43.4±1.9) (29.5±1.3) (24.8±0.8) (15.2±0.5) (20.0±0.7) Misato 01 155 118.8 (76.6) 95.0 (61.3) 66.3 (42.8) 47.5 (30.6) 38.8 (25.0) 23.8 (15.4) 31.3 (20.2) Misato 02 155 118.8 (76.6) 93.8 (60.5) 53.8 (34.7) 45.0 (29.0) 38.8 (25.0) 22.5 (14.5) 31.3 (20.2) Misato 03 157.5 120.0 (76.2) 93.8 (59.6) 66.3 (42.1) 46.3 (29.4) 37.5 (23.8) 23.8 (15.1) 32.5 (20.6) Misato 04 162.5 122.5 (75.4) 97.5 (60.0) 67.5 (41.5) 48.8 (30.0) 40.0 (24.6) 25.0 (15.4) 32.5 (20.0) Misato 05 165 125.0 (75.8) 97.5 (59.1) 70.0 (42.4) 47.5 (28.8) 41.3 (25.0) 25.0 (15.2) 32.5 (19.7) Misato 06 175 120.0 (68.6) 97.5 (55.7) 66.3 (37.9) 47.5 (27.1) 40.0 (22.9) 23.8 (13.6) 33.8 (19.3) 120.9±2.4 95.9±1.9 65.0±5.7 47.1±1.3 39.4±1.3 24.0±0.9 32.3±0.9 Mean±SD 161.7±7.7 (74.9±3.1) (59.4±1.9) (40.2±3.2) (29.2±1.2) (24.4±0.9) (14.9±0.7) (20.0±0.5) Iwatsuki 01 162.5 127.5 (78.5) 95.0 (58.5) 70.0 (43.1) 47.5 (29.2) 40.0 (24.6) 23.8 (14.6) 32.5 (20.0) Iwatsuki 02 153.8 120.0 (78.0) 95.0 (61.8) 66.3 (43.1) 47.5 (30.9) 38.8 (25.2) 22.5 (14.6) 30.0 (19.5) Iwatsuki 03 157.5 125.0 (79.4) 100.0 (63.5) 67.5 (42.9) 50.0 (31.7) 42.5 (27.0) 25.0 (15.9) 32.5 (20.6) Iwatsuki 04 162.5 95.0 (58.5) 85.0 (52.3) 67.5 (41.5) 42.5 (26.2) 40.0 (24.6) 22.5 (13.8) 33.8 (20.8) 116.9±14.9 93.8±6.3 67.8±1.6 46.9±3.1 40.3±1.6 23.5±1.2 32.2±1.6 Mean±SD 159.1±4.2 (73.6±10.1) (59.0±4.9) (42.7±0.7) (29.5±2.5) (25.4±1.1) (14.8±0.8) (20.2±0.6) H. feroniarum 1 162.5 － (－) 62.5 (38/5) 42.5 (26.2) 27.5 (16.9) 33.7 (20.7) 25.0 (15.4) － (－) H. feroniarum 2 150 83.5 (55.7) － (－) 40.0 (26.7) 27.5 (18.3) 31.3 (20.9) 23.7 (15.8) － (－) H. feroniarum 3 157.5 85.0 (54.0) 67.5 (42.9) 40.0 (25.4) 28.8 (18.3) 33.8 (21.5) 28.0 (17.8) － (－) H. feroniarum 4 140 77.5 (55.4) 60.0 (42.9) 42.5 (30.4) 25.0 (17.9) 28.8 (20.6) 23.5 (16.8) － (－) 82.0±4.0 63.3±3.8 41.3±1.4 27.2±1.6 31.9±2.4 25.1±2.1 － Mean±SD 152.5±9.8 (55.0±0.9) (41.4±2.5) (27.1±2.2) (17.9±0.7) (20.9±0.4) (16.4±1.1) (－) 23 Measurements of H. feroniarum 1-4 are from Tagami and Halliday (2013). 24 Kazumi TAGAMI

Table 2. Host-phoretic records of Dermaptera and mites of genus Histiostoma Host Phoretic Distribution records Anisolabididae H. mahunkai Fain, 1974 West-Central districts, Japan, Tagami, 2013 Anisolabella (Gonolabis) H. mahunkai Fain, 1974 Table 1 in this paper marginalis (Dohrn) H. piscium Fain & Lambrechts, 1985 Anisolabis maritima (Bonelli) H. mahunkai Fain, 1974 Table 1 in this paper H. piscium Fain & Lambrechts, 1985 Titanolabiinae H. feroniarum (Dufour), 1839 New South Wales, Australia, Tagami & Halliday, 2013 Titanolabis colossea (Dohrn) H. australiense Mahunka, 1975 H. humiditatis (Vitzthum), 1927 H. titanolabi Tagami & Halliday, 2013 Labiduridae Labidurinae H. camphori Eraky, 1999 Riyadh, Saudi Arabia, Negm & Labidura riparia (Pallas) Alatawi, 2011 Histiostoma sp. Florida, USA, Strandberg and Tucker, 1974 Forficulidae Forficulinae H. polypori (Oudemans), 1914 Arnheim, The Netherlands, L. Oudemans, 1900; Edinburgh, UK, Behura, 1950; Poland, Chmielewski, 1984; Pulawy, Poland, Chmielewski, 2009 and 2010 H. feroniarum (Dufour), 1839 Madrid, Spain, Rebolledo and Arroyo, 1996; Pulawy, Poland, Chmielewski, 2010 distributed throughout Japan; the former is distributed in eastern to southern Asia, and the latter is cosmopolitan. Their habitats are very similar, and both can be collected in semi-natural environments such as urban parks, long-maintained gardens in churches, shrines and temples, and abandoned spaces in downtown areas. Anisolabella marginalis, a known host of H. mahunkai, is a novel host for H. piscium. This finding suggests low specificity of phoretic selection in these mites. The former species was recorded from the swim-bladder of a dead fish in an aquarium in Antwerp (Fain and Lambrechts, 1985), and from a cockroach collected indoors in Tsukuba (Tagami, 2007). Although their adult stages are identifiable by their specific morphologies, their deutonymphs are very similar to each other. Only RLIs of leg I (80-85 in H. mahunkai vs 75-80 in H. piscium) and tarsus I (45-50 in H. mahunkai vs 40-45 in H. piscium) were diagnostic. The known phoretic histiostomatids on earwigs are summarized in Table 2. They may belong to a feroniarum group (Scheucher, 1957) because these deutonymphs have some common morphologies.

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