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Home , Chanodichthys, Corallanidae, Neocaridina, Neocaridina davidi, Palaemon paucidens, Shimane Prefecture

Research 2018 Vol.47: 73–88 ©Carcinological Society of . doi: 10.18353/crustacea.47.0_73 Tachaea chinensis (: Corallanidae), an ectoparasite of freshwater and prawns, from western Japan, with a list of its known collection localities and hosts

Kazuya Nagasawa, Tadashi Imai, Hidetoshi Saito

Abstract.̶ The corallanid isopod Tachaea chinensis Thielemann, 1910 is a parasite attached on the carapace of freshwater shrimps and prawns. The isopod is reported from paucidens De Haan, 1884 () at two localities in () and two localities in Shimane Prefecture (western ), and from sp. () at one locality in Shimane Prefecture, Japan. This represents the first record of T. chinensis from Shikoku. The is briefly described and illustrated based on non-ovigerous females. Prevalence of T. chinensis in populations of P. paucidens and Neocaridina sp. ranged from 10.7–57.1%, and the intensity of infestation was almost constantly one, which indicates its intra-specific competition on the host carapace. Palaemon paucidens is considered as a preferred host of T. chinensis in Honshu and Shikoku, Japan. Based on the literature published between 1910 and 2018 including this paper, T. chinensis has been reported from 14 nominal and some unidentified species of shrimps and prawns belonging to three atyid genera (Caridina, Neocaridina, and Paratya), two palaemonid genera (Macrobrachium and Palaemon), and one penaeid genus (Penaeus), ranging from temperate (Honshu, Shikoku, and Kyushu in Japan, and central-eastern China) through subtropical (Okinawa in Japan, and Hong Kong in southeastern China) to tropical (Hainan in southeastern China, Vietnam, Thailand, and Malaysia) regions in East Asia.

Key words: parasite, prevalence, geographical distribution, host utilization

■ Introduction and western Japan from 2009 to 2017 (e.g., Imai, 2012; Imai & Oonuki, 2013, 2014, 2017; Tachaea chinensis Thielemann, 1910 is a Imai et al., 2015, 2016, 2017, 2018), we col- parasite found attached on the carapace of lected T. chinensis parasitic on Palaemon pau- freshwater shrimps and prawns in East Asia cidens De Haan, 1884 (Palaemonidae) and (e.g., Thielemann, 1910; Tattersall, 1921; Ko- Neocaridina sp. (Atyidae) at four localities in mai, 1927; Nierstrasz, 1931; Shen, 1936; Iwa- Shikoku and western Honshu. This paper brief- sa, 1947; Karim & Fernando, 1963; Shiino, ly describes T. chinensis and reports on these 1965; Matsumoto, 1973; Delaney, 1989; Yu & collections with a new record of the species Li, 1998; Takeda et al., 2000; Saito, 2011, see from Shikoku and its occurrence on freshwater Table 2). There is as yet scant information on shrimps at the four localities. A list of the the biology of T. chinensis because the previ- known collection localities and hosts of T. chi- ous papers of the species mostly dealt with its nensis is also given based on the literature pub- and distributional records. During a lished between 1910 and 2018 and this paper. faunal study of freshwater shrimps in central

Received: 2 May 2018. Accepted: 14 Aug 2018. Published online: 28 Sept 2018. 73 KAZUYA NAGASAWA, TADASHI IMAI, HIDETOSHI SAITO

■ Materials and Methods species were recognized in this study. Howev- er, these two species are tentatively reported Shrimp sampling herein as sp. 1 and sp. 2 because the recent tax- Freshwater shrimps were collected using a onomy of shrimps of the genus from Japan is D-shaped hand net (mesh size, 2.5 mm; frame highly confused due to the invasion of alien width, 33 cm; handle length, 60 cm) in Kagawa congeneric species to the country (e.g., Nishi- Prefecture (Shikoku) on 26 October 2014 and no, 2017). Since the propodus of their third pe- Shimane Prefecture (western Honshu) on 22 reiopod is similar in both sexes (cf. Mitsugi et October 2017. The collection localities in al., 2017), both species are not a recently intro- Kagawa Prefecture were a pond in Tsuda Park duced species, (Bouvier, (34°17′37″N, 134°13′58″E, locality 29 in Fig. 1904) and will be reported in a separate article. 2), Tsuda, Sanuki, and the Sakamoto River Isopod-infested shrimps were later transported (34°12′36″N, 134°25′31″E, locality 30 in Fig. to the laboratory at Hiroshima University, Hi- 2), Sakamoto, Higashi-Kagawa, and the collec- gashi-Hiroshima, where isopods were carefully tion localities in Shimane Prefecture were removed using forceps and preserved in 70% Myôji Reservoir (35°27′50″N, 132°48′51″E, ethanol. Isopods were measured for body locality 23 in Fig. 2), Takudani, , and Ko length (from the anterior margin of the cepha- Reservoir (35°26′21″N, 132°48′43″E, locality lon to the posterior margin of the pleotelson), 24 in Fig. 2), Hirata, Izumo. At each locality, sexed (based on the presence or absence of an submerged vegetation and root masses were appendix masculina arising from the proximal scooped five times to quantitatively collect margin of the endpod of pleopod 2, cf. Del- shrimps. At the two localities in Kagawa Pre- aney, 1989, fig. 34F), and identified. Isopods fecture, shrimps were identified and counted on from Myôji Reservoir were examined for mor- site, and only isopod-infested individuals were phological characters using an Olympus taken and fixed in 10% formalin, while at the SZX10 stereo microscope and an Olympus two localities in Shimane Prefecture, shrimps BX51 light microscope, and drawings were were identified, separated into two groups by made with the aid of a drawing tube fitted on the presence or absence of isopod, and fixed in each of these microscopes. Voucher specimens 10% formalin individually in small plastic of T. chinensis have been deposited in the bags. During the field samplings, we carefully Crustacea (Cr) collection of the National Mu- scooped the shrimps and took firstly the infest- seum of Nature and Science, Tsukuba, Ibaraki ed ones: we found no detached isopod in the Prefecture, Japan (NSMT-Cr 25830 [n＝10] nets. from P. paucidens at a pond in Tsuda Park; NSMT-Cr 25831 [n＝10] from P. paucidens at Laboratory examination Myôji Reservoir). The remaining specimens of Shrimps from the four localities were T. chinensis are retained by the senior author brought to the laboratory at the National Re- (K. N.) for a future morphological study of the search Institute of Fisheries and Environment species from various localities in Japan and of Inland Sea, Takamatsu, where their on-site will be deposited in the same collection. Preva- identification was confirmed and they were lence and intensity of infestation are those de- measured for body length (from the orbital fined by Bush et al. (1997). The scientific edge of the carapace to the posterior margin of names of Japanese shrimps and prawns men- the telson) in mm. In particular, shrimps of the tioned in this paper follow Toyota & Seki atyid genus Neocaridina were carefully ob- (2014). served for morphological features, and two

74 Crustacean Research 47 Crustacean Research 47 TACHAEA CHINENSIS FROM JAPANESE SHRIMPS

■ Results Body slightly elongated, with many chro- matophores scattered on dorsal surface. Cepha- Description of T. chinensis lon small, nearly triangular, anteriorly with The following description of T. chinensis rounded rostrum. Eyes laterally separated. An- (Fig. 1A, Fig. 2) is based on 15 non-ovigerous tennula short, with 8–9 articles (2 peduncle ar- females (4.0–7.5 mm long) from Myôji Reser- ticles and 6–7 flagellum articles); peduncle voir. basal article slightly enlarged, article 2 cylin- drical; flagellum article 1 about as long as pe- duncle article 2. Antenna long, reaching to pe- reonite 4, with 18–19 articles (5 peduncle articles and 13–14 flagellum articles); peduncle articles 4 and 5 subequal. Pereonites 1–4 slightly larger than pereonites 5–7; posterolateral corners of pereonites 4–7 increasing in size posteriorly; pleonite 4 with lateral portions produced posteriorly and ex-

Fig. 2. Tachaea chinensis, non-ovigerous female, 6.4 mm Fig. 1. Tachaea chinensis, non-ovigerous female, 7.1 mm long, NSMT-Cr 25831, from Palaemon paucidens in Myôji long, NSMT-Cr 25831, from Palaemon paucidens in Myôji Reservoir, Izumo, Shimane Prefecture, western Japan. Reservoir, Izumo, Shimane Prefecture, western Japan, ethanol- A, cephalon and anterior pereonites (right antenna is not preserved specimen. A, body, dorsal view; B, individual illustrated), dorsal view; B, pleotelson and left uropod (right attached on the lateral side of the carapace of P. paucidens. uropod is not illustrated), dorsal view. Scale bars: 0.5 mm in A Scale bars: 2 mm in A; 3 mm in B. and B.

Crustacean Research 47 Crustacean Research 47 75 KAZUYA NAGASAWA, TADASHI IMAI, HIDETOSHI SAITO

Table 1. Occurrence of Tachaea chinenis on freshwater shrimps at four localities in western Japan

Body length Percent Intensity Locality Prefecture Collection date Shrimp species (mm) prevalence (mean)

Pond in Tsuda Park Kagawa 26 October 2014 Palaemon paucidens 16.7–24.4 48.5 (16/33*) 1 (1.0) Paratya improvisa 10.6–17.8 0 (0/4) 0 Neocaridina sp. 1** n.m*** 0 (0/37) 0 Sakamoto River Kagawa 26 October 2014 Palaemon paucidens 17.7–31.7 10.7 (6/56) 1 (1.0) Myôji Reservoir Shimane 22 October 2017 Palaemon paucidens 19.9–28.1 37.9 (11/29) 1–3 (1.4) Neocaridina sp. 1 and sp. 2** 10.0–15.6 0 (0/6) 0 Ko Reservoir Shimane 22 October 2017 Palaemon paucidens 18.7–22.4 57.1 (4/7) 1 (1.0) Neocaridina sp. 1 8.3–22.3 12.5 (1/8) 1 (1.0)

* No. infested/examined. ** See the Materials and Methods section for the identication of these species. *** Not measured. tending to pleotelson. Pleotelson semi-triangu- spectively. lar, with 8–9 (mostly 8) robust setae and longer There was no particular relationship between plumose setae on posterior margin. Exopods of occurrence of T. chinensis and shrimp body uropods fusiform; endopods wider than exo- size: the infested and uninfested individuals of pods; both rami with robust setae and plumose P. paucidens from Shimane Prefecture mea- setae on lateral and posterior margins. Pereo- sured 19.9–26.6 (mean, 22.9, n＝15) and 18.7– pods 1–3 short, prehensile; pereopods 4–7 lon- 28.1 (23.4, n＝21) mm long, respectively. ger than pereopods 1–3, ambulatory. Exopods A total of 42 specimens of T. chinensis was of pleopods 1–5 with marginal plumose setae. collected in this study: 16 and six specimens from a pond in Tsuda Park and the Sakamoto Occurrence of T. chinensis in shrimp popula- River, Kagawa Prefecture; 15 and five speci- tions mens from Myôji and Ko reservoirs, Shimane Four species of freshwater shrimps were col- Prefecture, respectively. These specimens were lected in this study: Palaemon paucidens, Neo- all non-ovigerous females. Their body length caridina sp. 1, Neocaridina sp. 2, and Paratya was similar between localities, measuring 4.5– improvisa Kemp, 1917. There was a difference 7.6 (mean, 6.3, n＝16), 4.8–7.1 (6.1, n＝6), in occurrence of T. chinensis between these 4.0–7.5 (6.3, n＝15), and 4.5–6.5 (5.6, n＝5) shrimp species (Table 1). Palaemon paucidens mm from the above four localities, respective- was infested by T. chinensis at four localities in ly. Kagawa and Shimane prefectures, and the prevalence of the isopod infestation ranged Mode of attachment by T. chinensis from 10.7–57.1%. Neocaridina sp. 1 harbored Tachaea chinensis was found firmly attached T. chinensis at one locality in Shimane Prefec- on the lateral side of the host carapace with its ture with prevalence being 12.5%. No infesta- cephalon posteriorly oriented (Fig. 1B). tion was found on Paratya improvisa, although only four individuals of the species were exam- ■ Discussion ined. The intensity of infestation by T. chinensis Morphology and identification of T. chinensis was almost constantly one (Table 1). Two and The specimens of isopod collected in this three isopods were found on three (8.1%) and study are nearly identical with the morphology one (2.7%) of the 37 infested P. paucidens, re- of T. chinensis from China (Thielemann, 1910;

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Shen, 1936; Wei, 1991; Cheng et al., 2011). body length of the species. There are several differences between Chinese In Japan, the generic name of the isopod has and Japanese specimens. The number of pleon- been often misspelled as “Tachea” (Komai, ites was reported as six in Chinese specimens 1927; Iwasa, 1947; Shiino, 1965; Kamita, (Shen, 1936) but it is five in our specimens. 1970; Matsumoto, 1973; Ueki et al., 1988; The antenna flagellum is composed of 13–14 Nishino, 1993; Nunomura, 1995; Sato & Kato, articles in our specimens but was reported to 1996; Narita, 2010; Takeda et al., 2000; consist of 11–12 (Shen, 1936; Cheng et al., Hayashi, 2014; Kawaguchi & Shimizu, 2013; 2011) or 16 articles (Wei, 1991) in Chinese Kimura et al., 2014; Anonymous, 2014; Shiba- specimens. Our specimens are more or less in ta et al., 2014; Mori et al., 2017; Saito et al., agreement with the descriptions of T. chinensis 2017). This is because Komai (1927) mistak- by Komai (1927), Iwasa (1947), Shiino (1965), enly reported the generic name as “Tachea,” and Aoyagi (2014) from Japan, although no de- which has been used by most of the subsequent tailed description of the species yet has been Japanese scientists. made using material from this country. Like in Chinese specimens, the antenna flagellum Occurrence of T. chinensis in shrimp popula- showed wide variations in number of articles tions (13–20 [Iwasa, 1947], ca. 16 [Shiino, 1965; In the present study, T. chinensis was collect- Aoyagi, 2014], 13–14 [this paper]) in Japanese ed in Kagawa Prefecture (Shikoku) and Shi- specimens. In this study, we did not collect any mane Prefecture (western Honshu), western Ja- ovigerous female or male of T. chinensis and pan. The collection in Kagawa Prefecture describe the species using the non-ovigerous represents the first record of the species from females. It is thus important to collect and de- Shikoku as well as its new prefectural record scribe fully grown adult specimens of both (see Table 2). sexes for a better understanding of the mor- During a period from 2009 to 2017, the sec- phology and taxonomic status of Japanese T. ond author (T. I.) of this paper conducted chinensis. The sequence data of the species freshwater shrimp and prawn samplings at 486 from P. paucidens in Osaka, Japan, have been localities (280 in rivers and 206 in lakes, registered in GenBank (Hata et al., 2017). ponds, and reservoirs) in central and western Body length of T. chinensis vary between in- Japan (see the literature in the Introduction dividuals reported from Japan (6–9 mm long section) and collected T. chinensis only at the [Tattersall, 1921]; 7–12 mm long [Komai, four localities reported herein, which indicates 1927]; ca. 10 mm long [Iwasa, 1947; Shiino, that this isopod parasite has a wide geographi- 1965]; 5.3–8.4 mm long [Delaney, 1989]; 4.2– cal distribution range in Japan (see below) but 8.4 mm long [Takeda et al., 2000]; 4–9 mm occurs in much fewer water bodies than do its long [Aoyagi, 2014]; 4.0–7.6 mm long [from hosts. Based on field samplings of freshwater the four localities in this paper]), China (7– in Shimane Prefecture, Hayashi (2011) 12 mm [Thielemann, 1910]; 3–4.5 mm long similarly stated that T. chinensis was collected [Tattersall, 1921]; 4.2 mm long [Shen, 1936]; only at some of the sites where freshwater 5.3 mm long [Delaney, 1989]; 9 mm long [Wei, shrimps occurred. Such distributional patterns 1991]; 5–7.5 mm long [Wang & Lin, 2008]; of the species are probably determined by vari- 5–10.3 mm long [Cheng et al., 2011]), and ous environmental and biological factors, and Malaysia (2.9–4.5 mm long [Delaney, 1989]). research is necessary to explain those patterns. These differences may have been influenced by The infested shrimps usually harbored one T. sampling seasons and/or regional variations in chinensis and only a few shrimps were parasit-

Crustacean Research 47 Crustacean Research 47 77 KAZUYA NAGASAWA, TADASHI IMAI, HIDETOSHI SAITO ized by two or three isopods (Table 1). This 1860, Caridina leucosticta Stimpson, 1860, implies that an intra-specific competition is Caridina typus H. Milne Edwards, 1837, and present between individuals of T. chinensis on Macrobrachium japonicum (De Haan, 1849)) the host carapace. Since detached individuals and two unidentified species (Palaemon sp. of T. chinensis have been observed to swim in and Neocaridina sp. 1) (Table 2). Palaemon the water (Shen, 1936; Takahashi, 2015), those paucidens was frequently infested by T. chi- individuals may seek uninfested shrimps after nensis in Shikoku and western Honshu in this they lose out in the competition. study (Table 1) and has also been often report- The specimens of T. chinensis collected in ed as the host of the isopod in Honshu (Table this study were all non-ovigerous females. A 2), which indicates that P. paucidens is a pre- similar female predominance was recorded by ferred host of T. chinensis in Honshu and Shi- Delaney (1989: 51) for the species from East koku. Nonetheless, it may be probable that T. Asia including Japan: eight of the 10 sexed chinensis differently utilizes shrimps and specimens examined were females (2.9– prawns on Okinawa-jima Island, one of the 8.4 mm long) and all of the six Japanese speci- (Aoyagi, 2014) because the mens were females (5.3–8.4 mm long). How- freshwater shrimp and prawn fauna of these is- ever, the number of individuals examined was lands differs from that of the Japanese main is- not numerous in our and Delaney’s (1989) lands (Shokita, 1975). studies (n＝42 and 10, respectively): we col- In the present study, P. paucidens and Neo- lected the specimens only two times (October caridina sp. 1 were found to be infested by T. 2014 and 2017) in this study. Quantitative sur- chinensis (Table 1). It is interesting to note that veys on a consecutive monthly basis, including Neocaridina sp. 1 was not infested at two of research on changes in body size (growth), sex the three localities where this shrimp species ratio, and maturity of T. chinensis, are needed occurred. This may have been influenced by to understand the observed female predomi- the difference in abundance of P. paucidens, a nance and the life cycle of the species. preferred host of T. chinensis, between locali- ties. At the two localities (a pond in Tsuda Park Hosts of T. chinensis and Myôji Reservoir), P. paucidens was abun- Based on the past literature and this paper, 14 dant (based on the number of individuals col- nominal and some unidentified species of lected) and T. chinensis could have easily shrimps and prawns belonging to three atyid found its preferred host, while at another local- genera (Caridina, Neocaridina, and Paratya), ity (Ko Reservoir), P. paucidens was much less two palaemonid genera (Macrobrachium and abundant and Neocaridina sp. 1 was probably Palaemon), and one penaeid genus (Penaeus) utilized as an additional host by T. chinensis. have been reported as hosts of T. chinensis (Ta- Further, the infested individual of Neocaridina ble 2). These species become infested in the sp. 1 was the third largest (18.0 mm long) wild, but the white leg shrimp, Penaeus vanna- among eight individuals of the species collect- mei Boone, 1931 is exceptional, being com- ed and about as large as those infested individ- mercially farmed in Tianjin, China (Wang & uals of P. paucidens (19.9–22.4 mm long, n＝ Lin, 2008). 4). Tachaea chinensis has been well studied in Tachaea chinensis was sometimes found on Japan, where it is known to parasitize seven freshwater fishes, such as the nominal species (Palaemon paucidens, Chanodichthys erythropterus (Basilewsky, Paratya improvisa, Paratya compressa (De 1855) (as brevicauda Günther, 1868 Haan, 1849), Caridina grandirostris Stimpson, [: ]) in China (Shen,

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1936), but those fishes can be regarded as acci- Geographical distribution of T. chinensis dental hosts of the isopod. Tachaea chinensis was originally described by Thielemann (1910) based on specimens Mode of attachment and pathological impact from a market in Shanghai, China. Subsequent- of T. chinensis ly, the species was reported from eastern China Tachaea chinensis was attached on the host (Tattersall, 1921; Shen, 1936; Delaney, 1989; carapace with its cephalon posteriorly oriented Bruce, 1990; Wei, 1991; Xu & Wang, 2006; (Fig. 1B). A similar observation was made by Qin, 2008; Wang & Lin, 2008; Yu, 2009; Aoyagi (2014). The reason of the posterior ori- Cheng et al., 2011; Li et al., 2018), Japan (e.g., entation of the species is unknown. Tattersall, 1921; Komai, 1927; Iwasa, 1947; A mass mortality occurred in a population of Delaney, 1989; Takeda et al., 2000, see Table 2 P. paucidens that had been captured in irriga- for the other papers), Vietnam (Dang et al., tion ponds and then reared in a concrete tank in 1979; Dang, 1980; Tran et al., 2011; Ngo et al., , western Japan (Ueki et 2013; Ngo & Ngo, 2014), Thailand (Delaney, al., 1988). Tachaea chinensis was found to par- 1989), and Malaysia (Karim & Fernando, asitize the diseased shrimps, from which Vibrio 1963; Delaney, 1989) (Figs. 3–4). No record of sp., a pathogenic bacterium, was isolated. Lat- T. chinensis is available from Korea, where er, all individuals of P. paucidens co-reared Tachaea sp. from Jindo Island has been report- with T. chinensis died by experimentally add- ed as “a new species” (Ahn et al., 2016). In ad- ing Vibrio sp. to the rearing water, and it has dition to T. chinensis and T. sp., three more been suggested that shrimp’s skin scars caused species of the genus have been reported from by the isopod serve as a portal of entry for the East Asia: Tachaea crassipes Schioedte and bacterium (Ueki et al., 1988). In this study, no Meinert, 1879 from Singapore (see Delaney, detailed observation was made on the mode of 1989), Tachaea lacustris Weber, 1892 from In- attachment by T. chinensis or scars caused by donesia (see Delaney, 1989; Bowman & Boto- such attachment. Various studies, such as histo- saneanu, 1992), and Tachaea tonlesapensis pathological observations on the scars and ex- Nunomura, 2006 from Cambodia (Nunomura, perimental infestation of shrimps with T. chi- 2006). nensis, are necessary to assess the impact of In Japan, T. chinensis has been found from the isopod on shrimps. Further, no bacterial more than 30 localities in 15 prefectures (Ao- disease has so far been reported from wild mori, Ibaraki, , Chiba, Kanagawa, Fukui, freshwater shrimp populations in Japan, but it Shiga, Mie, Kyoto, Osaka, Shimane, Okayama is important to note that T. chinensis has the [Honshu], Kagawa [Shikoku], Fukuoka [Kyu- potential to contribute to bacterial infections in shu], and Okinawa) (Table 2, Fig. 3). the wild. To summarize its geographical distribution, T. Tachaea chinensis was found infesting chinensis widely occurs in East Asia, ranging Penaeus vannamei cultured in Tianjin, China from temperate (Honshu, Shikoku, and Kyushu (Wang & Lin, 2008). Each infested shrimp har- in Japan, and central-eastern China) through bored two to five individuals of T. chinensis on subtropical (Okinawa in Japan, and Hong the carapace and/or abdomen. Infested shrimps Kong in southeastern China) to tropical (Hai- swam slowly and then died. These observa- nan in southeastern China, Vietnam, Thailand, tions indicate that the isopod is also an impor- and Malaysia) regions. tant parasite in the shrimp culture. Currently, Saito (2018) reported that P. pau- cidens caught in Lake Biwa, central Japan, is transported live to fishing bait shops in western

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Table 2. Known collection localities and shrimp/prawn hosts of Tachaea chinensis. A pelagic record of T. chinensis from the East China Sea (Xu & Wang, 2006) is not included herein

Prefecture/ Number in Shrimp/prawn Country Locality Reference Province Figs. 2–3 Family Species Japan Aomori Lake Ushirogata 1 ̶* NR** Kimura et al. (2014) Lake Myôjinnuma 2 ̶ NR Kimura et al. (2014) Ibaraki Lake Kasumigaura 3 Palaemonidae Palaemon paucidens Tattersall (1921) (as Leander paucidens) Tokyo ̶ ̶ NR Delaney (1989) Reservoir in Mizumoto Park 4 Palaemonidae Palaemon paucidens Takahashi (2015) (as Paraemon paucidens) Ôba River 5 ̶ NR Anonymous (2014) Inside moat of the Imperial Palace 6 Atyidae Paratya improvisa Takeda et al. (2000) (as P. compressa improvisa) 6 Palaemonidae Palaemon paucidens Takeda et al. (2000) Ponds in Meiji Jingu Shrine 7 Palaemonidae Palaemon paucidens Kawaguchi & Shimizu (2013) Chiba Obitsu River 8 Palaemonidae Palaemon paucidens Narita (2010) Kanagawa Pool in Yokohama 9 Palaemonidae Palaemon sp. Delaney (1989) Fukui Lake Kitagata 10 Palaemonidae Palaemon paucidens Nakachi et al. (2010) 10 Atyidae Paratya compressa Nakachi et al. (2010) Shiga Lake Biwa 11 Palaemonidae Palaemon paucidens Shiga-Kenritsu Biwako- Bunkakan (1980), Nishino (1993), Ota (2015) 11 Atyidae atyids Ota (2015) 11 ̶ NR Shibata et al. (2014) Mie Mouth of a small stream near Toba 12 ̶ NR Delaney (1989) Kyoto Ogura Pond 13 Palaemonidae Palaemon paucidens Tattersall (1921) (as Leander paucidens) Osaka Matoba Pond 14 ̶ NR Iwasaki & Morimoto (2012) Unknown 15 Palaemonidae Palaemon paucidens Hata et al. (2017) Shimane Kumi River 16 ̶ shrimps Kamita (1970), Sato & Kato (1996) 17 ̶ shrimps Nunomura (2000), Yamauchi (2002) Tamatsukuri River 18 ̶ NR Sato & Kato (1996) Pond in Koshin-in Temple, 19 ̶ NR Sato & Kato (1996) Nishikawatsu, Matsue 20 ̶ NR Sato & Kato (1996) Two reservoirs at Kosakai, Izumo 21 Atyidae Paratya compressa Hayashi (2011) (as P. c. compressa) Four reservoirs at Hirata, Izumo 22 Atyidae Paratya compressa Hayashi (2014) (as P. c. compressa) Myôji Reservoir 23 Palaemonidae Palaemon paucidens This paper Ko Resevoir 24 Atyidae Neocaridina sp. 1*** This paper Kumogi-sichijo-kozasa 25 ̶ NR Sato & Kato (1996) 26 ̶ NR Sato & Kato (1996) Okayama Okayama Prefectural Nature 27 ̶ NR Mori et al. (2017) Conservation Center Reservoir in southern region 28 Palaemonidae Palaemon paucidens Ueki et al. (1988) Kagawa Pond in Tsuda Park 29 Palaemonidae Palaemon paucidens This paper Sakamoto River 30 Palaemonidae Palaemon paucidens This paper Fukuoka Rivers and a reservoir in Hakata 31 ̶ NR Anonymous (2013) Okinawa Hajiô River 32 Atyidae Caridina grandirostris Aoyagi (2014) 32 Atyidae Caridina leucosticta Aoyagi (2014) 32 Atyidae Caridina typus Aoyagi (2014) 32 Palaemonidae Macrobrachium japonicum Aoyagi (2014) ̶ ̶ freshwater shrimps Komai (1927) ̶ Lakes and ponds in various localities Palaemonidae Palaemon paucidens Iwasa (1947) ̶ ̶ Palaemonidae Palaemon paucidens Shiino (1965) ̶ ̶ Palaemonidae Palaemon paucidens Toyota & Seki (2014: 239) ̶ ̶ Atyidae Paratya improvisa Toyota & Seki (2014: 240)

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Table 2. Continued

Prefecture/ Number in Shrimp/prawn Country Locality Reference Province Figs. 2–3 Family Species China Liaoning Rice field in Panjin 33 Palaemonidae Palaemon sinensis Li et al. (2018) (as Palaemonetes sinensis) Hebei Pei-tai-ho 34 ̶ NR Shen (1936, as Hopei province) 34 Atyidae Neocaridina davidi Yu (2009) (as Neocardinia heteropoda heteropoda) 34 Palaemonidae Palaemon sinensis Yu (2009) (as Palaemonetes sinensis) Beijing Pei-hai, Yü-chuan-shan, Tsin-hwa yuan 35 Atyidae Neocaridina sp. Shen (1936, as Peiping, (as Cardinia denticulata)**** Hopei province) 35 Palaemonidae Palaemon sinensis Shen (1936, as Peiping, (as Palaemonetes sinensis) Hopei province) ̶ 35 ̶ NR Delaney (1989, as Peking) Tianjin shrimp farm 36 Penaeidae Penaeus vannamei Wang & Lin (2008) Henan Xinxiang Yellow River Wetland Bird 37 Palaemonidae Macrobrachium spp., Cheng et al. (2011) National Nature Reserve, food markets Palaemon sinensis (as Palaemonetes sinensis), Palaemon modestus (as Exopalaemon modestus) 37 Atyidae Neocaridina davidi Cheng et al. (2011) (as Neocaridina heteropoda) Jiangsu Lake Taihu 38 Atyidae Cardinia gracilipes (as Tattersall (1921, as Tai Hu) Cardinia nilotica gracilipes) 38 Palaemonidae Palaemon sinensis Tattersall (1921, as Tai Hu) (as Palaemonetes sinensis) 38 ̶ NR Qin (2008) Shanghai Market (type locality) 39 ̶ NR Thielemann (1910) Zhejiang Jiaxing 40 ̶ NR Shen (1936, as Kashing) Hangzhou 40 Palaemonidae Macrobrachium nipponense Wei (1991) Zhoushan 40 ̶ NR Cited from Li (2003) Fujian Haiteng 41 ̶ NR Delaney (1989, as Fukien) ̶ 41 ̶ NR Cited from Li (2003) Hong Kong ̶ 42 ̶ NR Delaney (1989) Mai Po 42 Palaemonidae Macrobrachium sp. Bruce (1990) Deep Bay 42 ̶ NR Cited from Ong Che & Morton (1992) Hainan ̶ 43 ̶ NR Shen (1936)

Vietnam ̶ Mountain and plain areas ̶ NR Dang et al. (1979) Hoa Binh Northen plain, north-central plain 44 ̶ NR Dang (1980) Ninh Binh Karst area 45 ̶ NR Tran et al. (2011) Mekong Lower Mekong River 46 ̶ NR Ngo et al. (2013) Delta 46 ̶ NR Ngo & Ngo (2014)

Thailand ̶ Huai Nam Puat***** ̶ NR Delaney (1989) Ubon Ubon Ubon 47 ̶ NR Delaney (1989, as Ratchathani Ratchathani Province)

Malaysia Johor Gunung Pulai 48 Palaemonidae Macrobrachium malayanum Karim & Fernando (1963, (as M. geron) as Johore, Gunong Pulai) 48 Palaemonidae Macrobrachium sp. Delaney (1989, as Johore, Gunong Pulai)

* No information. ** Not reported. *** See the Materials and Methods section for the identification of this species. **** Due to the current taxonomic confusion in Neocaridina from Far East Asia (Nishino, 2017), this shrimp is herein regarded as an unidentified species. ***** “Huai Nam” means a brook in Thai, and due to the limited information on the reported locality, it is impossibe to find it in maps.

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Fig. 3. A map of the Japanese Archipelago, showing the collection localities of Tachaea chinensis based on the previous and present studies. Information on individual localities (1–32) is given in Table 2. Prefectural boundaries are indicated by dotted lines. and central Japan. In this country, some of in Tianjin, China (Wang & Lin, 2008). There freshwater shrimps used as fishing bait are dis- are records of T. chinensis associated with oys- carded to rivers by amateur fishermen after ters Crassostrea gigas (Thunberg, 1793) in their fishing (Saito et al., 2011) and P. pauci- Hong Kong, China (Delaney, 1989; see also dens is infested by T. chinensis in Lake Biwa Ong Che & Morton, 1992: 226). Moreover, T. (Shiga-Kenritsu Biwako-Bunkakan, 1980; chinensis was collected from the East China Nishino, 1993; Ota, 2015; Saito et al., 2017 [as Sea off China (Xu & Wang, 2006; cf. Huang & “Shirasa ebi”]). It is thus likely that T. chinensis Zheng, 2012). These collections suggest that T. has spread its geographical distribution in Ja- chinensis is not a genuine freshwater but eury- pan by live individuals of P. paucidens trans- haline species, and it is desirable to examine ported from Lake Biwa. the salinity tolerance of the species. Lake Kitagata is one of the localities where T. Salinity tolerance of T. chinensis chinensis was collected (Nakachi et al., 2010, In Japan, the specimens of T. chinensis were see Table 2 and Fig. 3). This lake is connected collected from fresh waters in the previous and with the by a short river and is present studies (see Table 2 for the literature), composed of freshwater and brackish-water re- except for the individuals collected from gions. Thus, it may be possible to clarify a sa- “brackish water” at the mouth of a small stream linity-related distributional pattern of T. chi- in (Delaney, 1989). Infestation nensis in the lake by examining its occurrence of T. chinensis on the white leg shrimp farmed at various sites differing in salinity. in brackish-water ponds has also been reported

82 Crustacean Research 47 Crustacean Research 47 TACHAEA CHINENSIS FROM JAPANESE SHRIMPS

ture in Honshu, through Shikoku and Kyushu, southward to Okinawa, the southernmost pre- fecture in Japan (Fig. 2). In addition, seasonal changes in occurrence of T. chinensis were only preliminarily studied in Japan (Takahashi, 2015). The species occurs in China, Vietnam, Thailand, and Malaysia as well, and it would be important to compare its life history and seasonal dynamics between temperate (Japa- nese main islands, and central-eastern China), subtropical (Okinawa in Japan, and Hong Kong in southeastern China) and tropical (Hainan in southeastern China, Vietnam, Thai- land, and Malaysia) regions.

■ Acknowledgements

We thank Tran Thi Thuy Ha (Research Insti- tute for Aquaculture No. 1, Tu Son), Masato Nitta (Kobe University, Kobe), Masubon Thongnam (Kasetsart University, Bangkok), Hironori Komatsu (National Science Museum of Nature and Science, Tsukuba), Masakazu Hayashi (Hoshizaki Green Foundation, Izumo), and Nobuhiro Saito (Suidosha, Kawasaki) for their assistance with the literature. We are also grateful to two anonymous reviewers who pro- vided constructive comments on the manu- script of the paper. This study was partially Fig. 4. A map of East Asia, showing the collection localities supported by JSPS KAKENHI Grant Number of Tachaea chinensis based on the previous and present 15K06932. studies. Information on individual localities (33–48) is given in Table 2. Country boundaries are indicated by dotted lines. See Fig. 3 and Table 2 for detailed collection localities (1–32) in ■ Literature Cited the Japanese Archipelago. Ahn, D.-H., Lee, C.-W., Yang, H.-M., Song, J.-H., Kwon, J.-I., Ji, S.-J., Park, M.-H., & Min, Future work G.-S., 2016. Freshwater invertebrates of Jin- In the present study, samplings were made do Island in Korea. Systematics, only in October 2014 and 2017 in Kagawa and Evolution and Diversity, Special Issue, 9: Shimane prefectures, but it is necessary to con- 37–44. duct more samplings for clarifying seasonal Anonymous, 2013. [General Report of the Envi- and regional variations in prevalence and in- ronmental Monitoring Survey in 2012 for tensity of T. chinensis in different regions of the Integration and Transference of Kyushu Japan because the species is widely distributed, University]. 232 pp. Kyushu University, Fu- kuoka. [In Japanese] ranging from Aomori, the northernmost prefec-

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of the systematic study of Isopoda in China. 424–0886, Japan. Marine Sciences, 1998(4): 28–29. [In Chi- (TI) National Research Institute of Fisheries nese with English title] and Environment of Inland Sea, Japan Fisheries Research and Education Agency, Addresses 234 Yashima-Higashi, Takamatsu, Kagawa (KN and HS) Graduate School of Biosphere 761–0111, Japan. Science, Hiroshima University, 1–4–4 Kagamiyama, Higashi-Hiroshima, Hiroshima 739–8528, Japan. E-mail address of corresponding author (KN’s present address) Aquaparasitology (KN) [email protected] Laboratory, 365–61 Kusanagi, Shizuoka

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