This art icle was downloaded by: [ Hiroshim a Universit y] , [ I . Madinabeit ia] On: 19 March 2013, At : 00: 40 Publisher: Taylor & Francis I nform a Lt d Regist ered in England and Wales Regist ered Num ber: 1072954 Regist ered office: Mort im er House, 37- 41 Mort im er St reet , London W1T 3JH, UK Journal of Natural History Publicat ion det ails, including inst ruct ions f or aut hors and subscript ion inf ormat ion: ht t p: / / www. t andf online. com/ loi/ t nah20 Contribution of Ju-shey Ho to the systematics of symbiotic copepods of Japan K. Nagasawa a , D. Tang a , D. Uyeno a b & I. Madinabeit ia a a Graduat e School of Biosphere Science, Hiroshima Universit y, 1-4-4 Kagamiyama, Higashi-Hiroshima, Hiroshima, 739-8528, Japan b Facult y of Science, Universit y of t he Ryukyus, 1 Senbaru, Nishihara, Okinawa, 903-0213, Japan Version of record f irst published: 11 Feb 2013. To cite this article: K. Nagasawa , D. Tang , D. Uyeno & I. Madinabeit ia (2013): Cont ribut ion of Jushey Ho t o t he syst emat ics of symbiot ic copepods of Japan, Journal of Nat ural Hist ory, 47: 5-12, 517-527 To link to this article: ht t p: / / dx. doi. org/ 10. 1080/ 00222933. 2012. 742586 PLEASE SCROLL DOWN FOR ARTI CLE Full t erm s and condit ions of use: ht t p: / / www.t andfonline.com / page/ t erm s- andcondit ions This art icle m ay be used for research, t eaching, and privat e st udy purposes. Any subst ant ial or syst em at ic reproduct ion, redist ribut ion, reselling, loan, sub- licensing, syst em at ic supply, or dist ribut ion in any form t o anyone is expressly forbidden. The publisher does not give any warrant y express or im plied or m ake any represent at ion t hat t he cont ent s will be com plet e or accurat e or up t o dat e. The accuracy of any inst ruct ions, form ulae, and drug doses should be independent ly verified wit h prim ary sources. The publisher shall not be liable for any loss, act ions, claim s, proceedings, dem and, or cost s or dam ages what soever or howsoever caused arising direct ly or indirect ly in connect ion wit h or arising out of t he use of t his m at erial. Journal of Natural History, 2013 Vol. 47, Nos. 5–12, 517–527, http://dx.doi.org/10.1080/00222933.2012.742586 Contribution of Ju-shey Ho to the systematics of symbiotic copepods of Japan Kazuya Nagasawaa* , Danny Tanga , Daisuke Uyenoa,b and Ione Madinabeitiaa Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 a Graduate School of Biosphere Science, Hiroshima University, 1-4-4 Kagamiyama, Higashi-Hiroshima, Hiroshima 739-8528, Japan; b Faculty of Science, University of the Ryukyus, 1 Senbaru, Nishihara, Okinawa 903-0213, Japan (Received 9 October 2011; final version received 18 October 2012; first published online 11 February 2013) This note reviews the work done by Ju-shey Ho, currently Professor Emeritus at California State University, Long Beach, CA, USA, on the systematics of symbiotic copepods from aquatic animals in Japanese waters. Since 1980, he has reported 110 species of symbiotic copepods from Japanese fish and marine invertebrates, including those representing one new family, seven new genera and 41 new species, and has greatly contributed to clarifying the symbiotic copepod fauna of Japan. Research using symbiotic copepods as bioindicators of the phylogeny and evolution of host animals was conducted by him for the first time in Japan. He also made significant contributions to the taxonomy and biology of caligid copepods, a group that poses a serious threat to the aquaculture industry, found on farmed fish in Japan. Keywords: Ju-shey Ho; invertebrates; fish; biology; fauna; Japan Introduction When the Eleventh International Conference on Copepoda was held in Mérida, Mexico, in July 2011, a special symposium was organized in honour of Dr Ju-shey Ho, who is currently Professor Emeritus in Biological Sciences at California State University, Long Beach (CSULB), CA, USA. He is recognized as a world-renowned scientist working on the biology, especially taxonomy and systematics, of symbiotic copepods. He began working on symbiotic copepods in Taiwan in 1960 and, as of October 2011, he has produced 257 publications (three books and 254 articles) on symbiotic copepods collected from around the world. In the course of his 52-year research career, Ju-shey has been very interested in the symbiotic copepod fauna of Japan, because Japan has a very rich marine fauna affected by the four surrounding seas (i.e. North Pacific Ocean, East China Sea, Sea of Japan and Sea of Okhotsk) each with different oceanographic features. Moreover, studies on symbiotic copepods in Japanese waters were restricted to particular regions, such as the Seto Inland Sea and the North Pacific Ocean. The impetus to study the symbiotic copepods of Japan was firmly planted in Ju-shey’s mind when he met Dr Sueo M. Shiino for the first time in Washington, D.C. in the summer of 1964. Dr Shiino was a professor at the Prefectural University of Mie in Tsu, Japan, and greatly contributed to the systematics of Crustacea, especially Copepoda. When *Corresponding author. Email: ornatus@hiroshima-u.ac.jp © 2013 Taylor & Francis Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 518 K. Nagasawa et al. Dr Shiino was returning to Japan from Rome after attending the First International Conference of Parasitology, he stopped in Washington, D.C. to visit Dr Thomas E. Bowman at the Smithsonian Institution. At that time, Ju-shey was working on his PhD with Dr Arthur G. Humes at Boston University. Ju-shey took a Greyhound bus south to Washington, D.C. It was a long ride, but he did not mind because his mind was entirely filled with the excitement of meeting Dr Shiino. Inspired by his meeting with Dr Shiino, Ju-shey used his first sabbatical leave from CSULB in August 1978 to make the first of many research visits to Japan. This note, therefore, reviews the work done by Ju-shey from 1978 to the present on the symbiotic copepods of Japan. Throughout the text, the phrase “symbiotic copepods” is used rather than “parasitic copepods”, because the former contains a wider meaning, i.e. copepods parasitic on or in and those associated with their hosts. Research by Ju-shey Ho on the symbiotic copepods of Japan Ju-shey’s research on the symbiotic copepods of Japan is too vast to describe in detail so several highlights are discussed below. Explaining the reason why he visited Japan in August 1978, he wrote “Japan is one of the few countries in the world where the marine copepod parasite fauna is relatively well known, with nearly 300 species being recorded from both fishes and invertebrates. However, an analysis of their geographical distribution indicates that a disproportionately large number of them, 251 species (about 84%), were recorded from the warm-temperate waters of the Pacific coast south of Cape Inubo and only 28 species (about 9%) are known from the Sea of Japan. In order to correct this artificial imbalance in the distribution of copepod parasites, I spent four months (from August to November) in 1978 to collect the copepod parasites of marine animals in the cold-temperate waters of the Sea of Japan” (Ho 1980). Ju-shey stayed at two marine biological stations facing the Sea of Japan: the Sado Marine Biological Station of Niigata University in Tassha, Sado Island, Niigata Prefecture, and the Noto Marine Biological Station of Kanazawa University in Noto-Ogi, Ishikawa Prefecture. During his 4-month stay at these stations, he dissected 109 species of marine fish and 86 species of marine invertebrates (Ho 1980). Even though the number of individuals examined for each host species is unknown, there is a record that 267 and 539 specimens, respectively, for two species of embiotocid fish, Ditrema temmincki and Neoditrema ransonneti, were dissected (Ho 1983). His examination of 195 marine species resulted in a huge collection of symbiotic copepods from 50 and 35 species of fish and invertebrates, respectively (Ho 1980). More surprisingly, using this material collected within only 4 months in 1978 in Japan, he managed to publish as many as 15 papers from 1980 to 1995 dealing with a total of 48 species of symbiotic copepods from the Sea of Japan, including those representing two new genera and 19 new species (Ho 1980, 1981, 1982, 1983, 1984, 1986, 1988, 1991; Ho et al. 1981; Ho and Honma 1983; Ho and Do 1985; Dojiri and Ho 1987, 1988; Ho and Kim 1995a, 1995b). Furthermore, in a book written in Japanese on the parasites of aquatic animals, he wrote a scientific essay about his experience in 1978 and his findings on the symbiotic copepods from Japanese bivalves (Ho 2005). Deep-sea fish are one of the host animals targeted by Ju-shey for symbiotic copepods. In the summer of 1984, he stayed at the Laboratory of Marine Zoology, Faculty of Science, Kochi University in Kochi, where he dissected rattails and other Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 Journal of Natural History 519 deep-sea fish accessioned in the laboratory collection. Based on the copepod specimens obtained from these dissections, he described four (two new, one known, and one unidentified) species of Lophoura (Sphyriidae) (Ho and Kim 1989), six new species of Clavella (Lernaeopodidae) (Ho 1993) and four (two new and two known) species and one new genus of Chondracanthidae (Ho 1994). Ju-shey also stayed as an invited scientist at the National Research Institute of Far Seas Fisheries in Shimizu (currently Shizuoka) from May to August 2000. He examined copepod samples collected by one of us (KN) from marine fish in Japanese waters. Using this material, he reported a variety of symbiotic copepods from 2000 to 2007, e.g. six species of copepods from offshore pelagic fishes such as marlin, yellowfin tuna, wahoo and shortfin mako (Ho and Nagasawa 2001a), chalimus larvae of Caligus sp. (Caligidae) from a mid-water sternoptychid fish (Ho and Nagasawa 2001b), caligid copepods from farmed marine fish (Ho and Nagasawa 2001c; Ho et al. 2001b) and a new species of Sarcotretes (Pennellidae) from a nomeid fish found in the stomachs of short-finned pilot whales (Ho et al. 2007). He also found, based on the stomach contents of juvenile pleuronectids, that they occasionally served as cleaner fish by feeding on copepods infecting other fishes (Ho 2001a). Ju-shey Ho has written a total of 41 publications so far on the symbiotic copepods of Japan, consisting of 38 peer-reviewed papers (nine as a single author, 20 as a senior author and nine as a coauthor), one review (as a coauthor), one book chapter (as a contributor) and one scientific article (as a single author). These publications were produced in a 30-year timespan from 1980 to 2009. The total number of pages printed in them is 696 (an average of 17.0 pages per publication), which is almost equivalent to the volume of two ordinary books. Considering that he is based in California, USA, we can say without reservation that his work on the Japanese symbiotic copepods is massive. In addition to his copepod work, he has written a paper on the monogenean parasites (which includes the establishment of one new species) of marine fish from Japanese waters (Ho and Perkins 1980). Number of species of the symbiotic copepods reported by Ju-shey Ju-shey’s taxonomic descriptions and illustrations of symbiotic copepods are of high standard. He has always attempted to make the best description for each species he found, including those from Japan. Between 1980 and 2009, Ju-shey reported, as a single author or a co-author, a total of 110 named species and three unidentified species of symbiotic copepods from Japan (Table 1). He established one new family (Umazuracolidae Ho, Ohtsuka and Nakadachi, 2006), seven new genera (Coelotrophus Ho, Katsumi and Honma, 1981; Humesulus Ho, 1982; Dermoergasilus Ho and Do 1982; Naricolax Ho, Do and Kasahara, 1983; Chelonichondria Ho, 1994; Umazuracola Ho, Ohtsuka and Nakadachi, 2006; Lobomolgus Ho and Kim, 2009) and 41 new species. The copepods reported by him belong to three genera in three families of the order Harpacticoida, one genus in one family of the order Cyclopoida, 33 genera in 14 families of the order Poecilostomatoida and 26 genera in 10 families of the order Siphonostomatoida. Among those 28 families, eight are well represented: he reported 16 species each for Chondracanthidae and Lernaeopodidae, 11 species (including two unidentified species) for Caligidae, eight species each for Bomolochidae, Lichomolgidae and Taeniacanthidae, six species for Ergasilidae and five species for Pennellidae. 520 K. Nagasawa et al. Table 1. Symbiotic copepods reported by Ju-shey Ho from Japan between 1980 and 2009. Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 Species Order Harpacticoida Sars, 1903 Family Canuellidae Lang, 1943 Sunaristes japonicus Ho, 1986∗ Family Porcellidiidae Boek, 1865 Porcellidium paguri Ho, 1986∗ Family Tisbidae Stebbing, 1910 Tisbe japonica Ho, 1982∗ Order Cyclopoida Burmeister, 1834 Family Notodelphyidae Dana, 1853 Bonnierilla mollia Ho, 1984∗ Doroixys capillosus Ho and Kim, 2009∗ Doropygus brevipes Ho, 1984∗ Doropygus pinguis (Ooishi, 1962) Order Poecilostomatoida Thorell, 1859 Family Anthessiidae Humes, 1986 Panaietis haliotis Yamaguti, 1936 Panaietis yamagutii Izawa, 1976 Family Bomolochidae Sumpf, 1871 Bomolochus bellones Burmeister, 1835 Bomolochus decapteri Yamaguti, 1936 Naricolax atypicus Ho, Do and Kasahara, 1983†∗ Nothobomolochus lateolabracis (Yamaguti and Yamasu, 1959) Nothobomolochus thambus Ho, Do and Kasahara, 1983∗ Nothobomolochus triceros (Bassett-Smith, 1898) Orbitacolax hapalogenyos (Yamaguti and Yamasu, 1959) Pumiliopes squamosus Cressey and Boyle, 1973 Family Chondracanthidae Milne Edwards, 1840 Acanthochondria inimici Yamaguti, 1939 Acanthochondria macrocephala Gusev, 1951 Acanthochondria priacanthi Shiino, 1964 Acanthochondria sixteni Wilson, 1922 Acanthochondria uranoscopi Ho and Kim, 1995∗ Chelonichondria okamurai Ho, 1994†∗ Chondracanthodes deflexus Wilson, 1932 Chondracanthus irregularis Fraser, 1920 Chondracanthus neali Leigh-Sharpe, 1930 Chondracanthus parvus Ho, Kim and Nagasawa, 2005∗ Chondracanthus solidus (Gusev, 1951) Chondracanthus yanezi Atria, 1980 Chondracanthus zei Delaroche, 1811 Diocus sadoensis (Shiino, 1960) Jusheyhoea ryukyuensis Ho, 1994∗ Pseudacanthocanthopsis bicornutus (Shiino, 1960) Reference Ho (1986, 1988) Ho (1986) Ho (1982) Ho (1984) Ho and Kim (2009) Ho (1984) Ho (1984) Ho (1981) Ho (1981) Ho et al. (1983) Ho (1983) Ho et al. (1983) Ho et al. (1983) Ho et al. (1983) Ho et al. (1983) Ho et al. (1983) Ho et al. (1983) Dojiri and Ho (1988) Ho and Kim (1995b) Ho and Kim (1995b, 1996) Dojiri and Ho (1988); Ho and Kim (1995b) Ho and Kim (1995b) Ho (1994) Ho (1994) Ho and Kim (1996) Ho (1994) Ho et al. (2005) Ho and Kim (1995a) Ho and Kim (1996) Ho (1991) Ho and Kim (1995a) Ho (1994) Ho and Kim (1995a) (Continued) Journal of Natural History 521 Table 1. (Continued). Species Family Ergasilidae von Nordmann, 1832 Dermoergasilus amplectens (Dogiel and Akhmerov, 1952)† Ergasilus bani Ohtsuka, Ho and Nagasawa, 2004∗ Ergasilus genuinus (Kokubo, 1914) Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 Ergasilus hypomesi Yamaguti, 1936 Nipergasilus bora (Yamaguti, 1939) Thersitina gasterostei (Pagenstrecher, 1861) Family Lamippidae Joliet, 1882 Enalcyonium digitigerum Ho, 1984∗ Family Lernaeosoleidae Yamaguti, 1963 Bobkabata kabatabobbus Hogans and Benz, 1990 Family Lichomolgidae Kossmann, 1877 Humesulus lobatus Ho, 1982†∗ Lichomolgus bidentipes Ho, 1980∗ Lichomolgus sadoensis Ho, 1980∗ Lobomolgus okinawaensis Ho and Kim, 2009†∗ Modiolicola gracilicaudus Avdeev, 1977 Parapanaietis tegulae Hoshina and Sugiura, 1953 Synstellicola longicauda Ho, 1982∗ Synstellicola similis Ho, 1982∗ Family Myicolidae Yamaguti, 1936 Pseudomyicola spinosus (Raffaele and Monticelli, 1885) Family Oncaeidae Giesbrecht, 1892 Oncaea venusta Philippi, 1843 Family Philoblennidae Izawa, 1976 Philoblenna tumida Ho, 1981∗ Family Rhynchomolgidae Humes and Stock, 1972 Doridilicola isoawamochi Ho, 1981∗ Family Synapticolidae Humes and Boxshall, 1996 Scambicornus affinis Ho, 1982∗ Family Taeniacanthidae Wilson, 1911 Anchistrotos kojimensis Do and Ho, 1983∗ Cirracanthus inimici (Yamaguti and Yamasu, 1954) Taeniacanthus balistae (Claus, 1864) Taeniacanthus platycephali (Yamaguti, 1939) Taeniacanthus rotundiceps (Shiino, 1957) Taeniacanthus sebastichthydis Yamaguti, 1939 Reference Ho and Do (1982) Ohtsuka et al. (2004a) Ohtsuka et al. (2004a) Ohtsuka et al. (2004a) Ho and Do (1982) Ohtsuka et al. (2004b, c) Ho (1984) Ho et al. (2005) Ho (1982) Ho (1980) Ho (1980) Ho and Kim (2009) Ho (1980) Ho (1981) Ho (1982) Ho (1982) Ho (1980); Do et al. (1984) Ho (1984) Ho (1981) Ho (1981) Ho (1982) Do and Ho (1983a) Dojiri and Ho (1987) Dojiri and Ho (1987) Dojiri and Ho (1987) Dojiri and Ho (1987) Dojiri and Ho (1987) (Continued) 522 K. Nagasawa et al. Table 1. (Continued). Species Taeniacanthus similis Dojiri and Cressey, 1987 Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 Taeniacanthus yamagutii (Shiino, 1957) Family Umazuracolidae Ho, Ohtsuka and Nakadachi, 2006‡ Umazuracola elongatus Ho, Ohtsuka and Nakadachi, 2006†∗ Undetermined family Coelotrophus nudus Ho, Katsumi and Honma, 1981†∗ Order Siphonostomatoida Burmeister, 1835 Family Artotrogidae Brady, 1880 Pteropontius decorus Ho, 1984∗ Family Asterocheridae Giesbrecht, 1899 Asterocheres aesthetes Ho, 1984∗ Collocheres inaequalis Ho, 1982∗ Family Caligidae Burmeister, 1835 Caligus coryphaenae Steenstrup and Lütken, 1861 Caligus lalandei Barnard, 1948 Caligus sclerotinosus Roubal, Armitage and Rohde, 1983 Caligus tanago Yamaguti, 1939 Caligus sp. Gloiopotes huttoni (Thompson, 1889) Gloiopotes hygomianus Steenstrup and Lütken, 1861 Lepeophtheirus elegans Gusev, 1951 Lepeophtheirus longiventralis Yü and Wu, 1932 Lepeophtheirus salmonis (Krøyer, 1837) Lepeophtheirus sp. Family Dichelesthiidae Milne Edwards, 1840 Anthosoma crassum (Abildgaard, 1794) Family Hatschekiidae Kabata, 1979 Hatschekia conifera Yamaguti, 1939 Family Lernaeopodidae Milne Edwards, 1840 Alella ditrematis (Yamaguti, 1936) Alella macrotrachellus (Brian, 1906) Clavella collaris Ho, 1993∗ Clavella diversia Ho, 1993∗ Clavella gadomi Ho, 1993∗ Clavella longicauda Ho, 1993∗ Clavella okamurai Ho, 1993∗ Clavella sokodara Ho, 1993∗ Reference Dojiri and Ho (1987) Dojiri and Ho (1987) Ho et al. (2006) Ho et al. (2006) Ho et al. (1981) Ho (1984) Ho (1984) Ho (1982) Ho and Nagasawa (2001a) Ho et al. (2001b) Ho et al. (2004) Ho (1983) Ho and Nagasawa (2001b) Ho and Kim (1996); Ho and Nagasawa (2001a) Ho and Nagasawa (2001a) Ho and Kim (1996) Ho et al. (2004) Ho and Nagasawa (2001c) Ho et al. (2001a) Ho and Kim (1996); Ho and Nagasawa (2001a) Ho and Kim (1996) Ho (1983) Ho (1983) Ho (1993) Ho (1993) Ho (1993) Ho (1993) Ho (1993) Ho (1993) (Continued) Journal of Natural History 523 Table 1. (Continued). Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 Species Reference Clavellisa dorosomatis Yamaguti, 1939 Clavellopsis nodula Do and Ho, 1983∗ Naobranchia occidentalis Wilson, 1915 Parabrachiella amphipacifica Ho, 1982 [as Neobrachiella amphipacifica Ho, 1982] Parabrachiella brevicapita (Ho and Do, 1984) [as Neobrachiella brevicapita (Ho and Do, 1984)∗ ] Parabarchiella seriolae (Yamaguti and Yamasu, 1960) [as Eobrachiella elegans f. seriolae (Yamaguti and Yamasu, 1960)] Parabrachiella trichiuri (Yamaguti, 1939) [as Neobrachiella trichiuri (Yamaguti, 1939)] Pseudocharopinus markewitschi (Gusev, 1951) Family Lernanthropidae Kabata, 1979 Lernanthropinus sauridae Do, 1985 Lernanthropsis mugilii (Shishido, 1898) Lernanthropus atrox Heller, 1865 Lernanthropus chrysophrys Shishido, 1898 Lernanthropus cornutus Kirtisinghe, 1937 Mitrax heteropodus (Yü, 1933) Family Pandaridae Milne Edwards, 1840 Echthrogaleus denticulatus Smith, 1874 Pandarus satyrus Dana, 1849 Family Pennellidae Burmeister, 1835 Haemobaphes diceraus Wilson, 1917 Haemobaphes pannosus Kabata, 1979 Lernaeolophus aceratus Ho and Honma, 1983∗ Pennella filosa (Linnaeus, 1758) Sarcotretes longirostris Ho, Nagasawa and Kim, 2007∗ Family Sphyriidae Wilson, 1919 Lophoura cardusa (Leigh-Sharpe, 1934) Lophoura tetraloba Ho and Kim, 1989∗ Lophoura ventricula Ho and Kim, 1989∗ Lophoura sp. Taxa marked with ‡, † and respectively. ∗ Do and Ho (1983c) Do and Ho (1983b) Ho and Kim (1996) Ho et al. (2005) Ho and Do (1984) Ho and Do (1984) Ho and Do (1984) Ho and Kim (1996) Ho and Do (1985) Ho and Do (1985) Ho and Do (1985) Ho and Do (1985) Ho and Do (1985) Ho and Do (1985) Ho and Kim (1996) Ho and Nagasawa (2001a) Ho and Kim (1996) Ho and Kim (1996) Ho and Honma (1983) Ho and Nagasawa (2001a) Ho et al. (2007) Ho and Kim (1989) Ho and Kim (1989) Ho and Kim (1989) Ho and Kim (1989) were described as a new family, a new genus and a new species, Significance of research by Ju-shey Ho When Ju-shey visited Japan in 1978, “nearly 300 species” (Ho 1980) of symbiotic copepods were recorded from this country. Now, more than 400 species are known to occur there. This increase in the number of species of symbiotic copepods is largely the result of his continued research after 1978. Before his work, the symbiotic copepod fauna of Japan was intensively studied by two scientists, Dr S.M. Shiino and Dr S. Yamaguti, who published their papers from 1932 to 1965 and from 1936 to 1974, Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 524 K. Nagasawa et al. respectively. Ju-shey is the successor to these two scientists. Through his work, knowledge of the symbiotic copepod fauna from the Sea of Japan has been dramatically improved. Ju-shey described many copepods associated with marine invertebrates from Japan. The invertebrates he examined consisted of a wide range of taxa, including sponges (Porifera), hydroids and corals (Cnidaria), peanut worms (Sipuncula), bivalves and gastropods (Mollusca), hermit crabs (Arthropoda), sea stars, feather stars and sea cucumbers (Echinodermata) and sea squirts (Tunicata) (Ho 1980, 1981, 1982, 1984, 1986, 1988; Ho et al. 1981; Do et al. 1984; Ho and Kim 2009). In Japan, only a few studies had been conducted on the symbiotic copepods of marine invertebrates before his research. Therefore, his work made Japanese scientists aware of the diversity of symbiotic copepods. He stated that “it is hoped that my report of these associated copepods will stimulate and encourage more Japanese marine biologists to become interested in working on them, not only on the systematics, but also on all aspects of their biology” (Ho 1984). Research using symbiotic copepods as bio-indicators of the phylogeny and evolution of host animals was conducted by Ju-shey for the first time in Japan. As previously mentioned, he dissected numerous embiotocid fish from Japanese waters and, by comparing their symbiotic copepod fauna with that of North American embiotocids, he inferred the phylogeny and biogeography of these fish occurring in the Far East (Ho 1983). He also assessed the phylogenetic relationships of the genera of psychrolutid fish based on symbiotic copepods collected from fish kept in the Marine Zoology of Hokkaido University in Hakodate (Ho et al. 2005). Caligid copepods pose a serious threat to the aquaculture industry in Asia, and their exact identification is essential to facilitate prevention and control (Ho 2000a). He reported Caligus lalandei parasitic on farmed and wild amberjacks in Japan, which represented the first record from East Asia (Ho et al. 2001b). He identified and redescribed two species of caligids, Caligus sclerotinosus and Lepeophtheirus longiventralis, as new country records, from marine fish cultured in Japan (Ho et al. 2004). He also investigated and explained why Lepeophtheirus salmonis, a highly pathogenic parasite of salmonids cultured in Europe and North America, does not cause serious problems in coho salmon farms in Japan (Ho and Nagasawa 2001c). Furthermore, he wrote a scientific article in Japanese about the caligid copepods as pathogens of farmed fish (Ho 2000b). Ju-shey Ho’s supportive family Why has Ju-shey been so active and productive? Needless to say, it is a result of his enthusiasm for research on symbiotic copepods, but there is an unwritten fact in his publications, that is, his life has been supported in various ways by his family, especially his wife, Pao-Hsi Ho. When he received the Taiwanese Science and Engineering Achievement Award, which is called “the Taiwanese Nobel Prize” in Taiwan, in November 2001, he noted in a speech given at the ceremony party that “I would not have been awarded without her assistance. Although she has always been saying that she could help him much more if she would become a biologist, I am completely satisfied to see that she has been skilfully handling complex domestic things and been devoted to the development and growth of our two children” (Nagasawa and Ohtsuka 2002). Journal of Natural History 525 Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 Conclusion It is obvious that Ju-shey has greatly contributed to our knowledge of the symbiotic copepods of Japan. However, this research constitutes only a small part of his entire work. He has described numerous species of symbiotic copepods from many other countries and regions of the world, including Asia (Taiwan, Korea, China, Philippines, Palau, Malaysia, Thailand, India, Pakistan, Iraq and Kuwait), Europe (Portugal, Spain and the UK), North America (USA and Mexico), South America (Venezuela, Ecuador, Peru, Chile and Brazil), Africa (South Africa, Mauritania and Senegambia) and Antarctica. The number of his publications on the Japanese symbiotic copepods constitutes only 16% (41/257) of his entire publication list. Since 1996, he has actively studied the symbiotic copepods of Taiwan, which is his motherland, in collaboration with Dr Ching-Long Lin, a professor at the National Chiayi University in Chiayi, Taiwan. At the Seventh International Conference on Copepoda held in Curitiba, Brazil in July 1999, as the president of the World Association of Copepodologists, Ju-shey stated in his “Maxilliped Lecture” that symbiotic copepods are estimated to comprise more than one-third of known copepods and we need more knowledge on this group (Ho 2001). Fortunately, Ju-shey and a younger generation of copepodologists from all over the world are steadfastly contributing to this field. References Do TT, Ho J-S. 1983a. Anchistrotos kojimensis sp. nov. (Copepoda: Taeniacanthidae) parasitic on Acanthogobius flavimanus (Pisces: Teleostei) in Kojima Bay, Japan. Fish Pathol. 18:1–5. Do TT, Ho J-S. 1983b. Clavellopsis nodula sp. nov. (Copepoda: Lernaeopodidae) parasitic on sea bream, Mylio macrocephalus (Basilewsky) (Pisces: Sparidae) in Japan. Fish Pathol. 18:31–36. Do TT, Ho J-S. 1983c. Redescription of Clavellisa dorosomatis Yamaguti, 1939, a lernaeopodid copepod parasitic on gizzard shad, Konosirus punctatus (Temminck & Schlegel) (Pisces: Dorosomatidae) in Japan. Fish Pathol. 18:41–43. Do TT, Kajihara T, Ho J-S. 1984. The life history of Pseudomyicola spinosus (Raffaele & Monticelli, 1885) from the blue mussel, Mytilus edulis galloprovincialis in Tokyo Bay, Japan, with notes on the production of atypical male. Bull Ocean Res Inst Univ Tokyo. 17:i–ii, 1–65. Dojiri M, Ho J-S. 1987. Copepods of the Taeniacanthidae (Poecilostomatoida) parasitic on fishes of Japan. Rep Sado Mar Biol Stat Niigata Univ. 17:32–42. Dojiri M, Ho J-S. 1988 Two species of Acanthochondria (Copepoda: Poecilostomatoida) parasitic on fishes of Japan. Rep Sado Mar Biol Stat Niigata Univ. 18:47–56. Ho J-S. 1980. Origin and dispersal of Mytilus edulis in Japan deduced from its present status of copepod parasitism. Publ Seto Mar Biol Lab. 25:293–313. Ho J-S. 1981. Parasitic Copepoda of gastropods from the Sea of Japan. Annu Rep Sado Mar Biol Stat Niigata Univ. 11:23–41. Ho J-S. 1982. Copepoda associated with echinoderms of the Sea of Japan. Annu Rep Sado Mar Biol Stat Niigata Univ. 12:33–61. Ho J-S. 1983. Copepod parasites of Japanese surfperches: their inference on the phylogeny and biogeography of Embiotocidae in the Far East. Ann Rep Sado Mar Biol Stat Niigata Univ. 13:31–62. Ho J-S. 1984. Copepoda associated with sponges, cnidarians, and tunicates of the Sea of Japan. Rep Sado Mar Biol Stat Niigata Univ. 14:23–61. Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 526 K. Nagasawa et al. Ho J-S. 1986. Harpacticoid copepods of the genera Sunaristes and Porcellidium associated with hermit crabs in Japan. Rep Sado Mar Biol Stat Niigata Univ. 16:21–38. Ho J-S. 1988. Cladistics of Sunaristes, a genus of harpacticoid copepods associated with hermit crabs. Hydrobiologia 167/168:555–560. Ho J-S. 1991. Redescription of Chondracanthus zei Delaroche (Copepoda, Poecilostomatoida) parasitic on Zeus faber L. in the Sea of Japan, with a preliminary review of the genus. Rep Sado Mar Biol Stat Niigata Univ. 21:49–79. Ho J-S. 1993. New species of Clavella (Copepoda: Lernaeopodidae) parasitic on Japanese rattails (Pisces: Macrouridae). Publ Seto Mar Biol Lab. 36:107–118. Ho J-S. 1994. Chondracanthid copepods (Poecilostomatoida) parasitic on Japanese deep-sea fishes, with a key to the genera of the Chondracanthidae. J Nat Hist. 28:505–517. Ho J-S. 2000a. The major problem of cage aquaculture in Asia relating to sea lice. In: Liao IC, Lin CK, editors. Cage aquaculture in Asia: Proceedings of the First International Symposium on Cage Aquaculture in Asia. Manila: Asian Fish Soc and Bangkok: World Aquacult Soc Southeast Asian Chap. p. 13–19. Ho J-S. 2000b. Sea lice (Copepoda, Caligidae): the potential pests to cage aquaculture in Asia. Aquabiology 22:442–447. [In Japanese with English abstract.] Ho J-S. 2001. Why do symbiotic copepods matter? Hydrobiologia 453/454:1–7. Ho J-S. 2005. Inferring the origin of blue mussels established in Japanese waters from the parasites. In: Nagasawa K. editor. Introduction to Copepodology – World of Tiny Giants in Water. Hadano (Japan): Tokai University Press. p. 272–280, 307–308. [In Japanese.] Ho J-S, Do TT. 1982. Two species of Ergasilidae (Copepoda: Poecilostomatoida) parasitic on the gills of Mugil cephalus Linnaeus (Pisces: Teleostei), with proposition of a new genus Dermoergasilus. Hydrobiologia 89:247–252. Ho J-S, Do TT. 1984. Three species of Lernaeopodidae (Copepoda) parasitic on fishes of Japan, with proposition of a new genus and discussion of Charopinus Yamaguti, 1963. Publ Seto Mar Biol Lab. 29:333–358. Ho J-S, Do TT. 1985. Copepods of the family Lernanthropidae parasitic on Japanese marine fishes, with a phylogenetic analysis of the lernanthropid genera. Rep Sado Mar Biol Stat Niigata Univ. 15:31–76. Ho J-S, Do TT, Kasahara S. 1983. Copepods of the family Bomolochidae parasitic on fishes of Kojima Bay, Okayama Prefecture. J Fac Applied Biol Sci Hiroshima Univ. 22:1–41. Ho J-S, Gómez S, Ogawa K, Aritaki M. 2004. Two species of parasitic copepods (Caligidae) new to Japan. Syst Parasitol. 57:19–34. Ho J-S, Honma Y. 1983. Lernaeolophus aceratus, a new species of copepod parasitic on rainbowfish from the Sea of Japan, with notes on food and feeding. J Crust Biol. 3:321–328. Ho J-S, Katsumi F, Honma Y. 1981. Coelotrophus nudus gen. et sp. nov., an endoparasitic copepod causing sterility in a sipunculan Phascolosoma scolops (Selenka and De Man) from Sado Island, Japan. Parasitology 82:481–488. Ho J-S, Kim I-H. 1989. Lophoura (Copepoda: Sphyriidae) parasitic on the rattails (Pisces: Macrouridae) in the Pacific, with note on Sphyrion lumpi from the Sea of Japan. Publ Seto Mar Biol Lab. 34:37–54. Ho J-S, Kim I-H. 1995a. Chondratacanthid copepods (Poecilostomatoida) parasitic on fishes of the Sea of Japan. Rep Sado Mar Biol Stat Niigata Univ. 25:31–44. Ho J-S, Kim I-H. 1995b. Acanthochondria (Copepoda: Chondracanthidae) parasitic on fishes of Sado Island in the Sea of Japan, with a preliminary review of the genus. Rep Sado Mar Biol Stat Niigata Univ. 25:45–67. Ho J-S, Kim I-H. 1996. Copepods parasitic on fishes of western North Pacific. Publ Seto Mar Biol Lab. 37:275–303. Ho J-S, Kim I-H. 2009. Two species of Copepoda parasitic in the algal-bearing ascidian, Didemnum molle (Herdman), in Okinawa, Japan. Proc Biol Soc Wash. 122:414–425. Downloaded by [Hiroshima University], [I. Madinabeitia] at 00:40 19 March 2013 Journal of Natural History 527 Ho J-S, Kim I-H, Nagasawa K. 2005. Copepod parasites of the flatheads (Pisces, Psychrolutidae) and their implication on the phylogenetic relationships of psychrolutid genera. Zool Sci. 22:411–425. Ho J-S, Nagasawa K. 2001a. New records of parasitic Copepoda from the offshore pelagic fishes of Japan. Bull Nat Res Inst Far Seas Fish. 38:1–5. Ho J-S, Nagasawa K. 2001b. Implication of the occurrence of chalimus larvae (Copepoda, Caligidae) on the larvae of Maurolicus muelleri (Pisces, Sternoptychidae) in the Sea of Japan. Bull Nat Res Inst Far Seas Fish. 38:7–11. Ho J-S, Nagasawa K. 2001c. Why infestation by Lepeophtheirus salmonis (Copepoda: Caligidae) is not a problem in the coho salmon farming industry in Japan. J Crust Biol. 21:954–960. Ho J-S, Nagasawa K, Takatsu T. 2001a. The juvenile cresthead flounder (Pleuronectes schrenki): an occasional cleaner occurring in Lake Notoro, Hokkaido. Bull Fish Sci Hokkaido Univ. 52:1–3. Ho J-S, Nagasawa K, Kim I-H, Ogawa K. 2001b. Occurrence of Caligus lalandei Barnard, 1948 (Copepoda, Siphonostomatoida) on amberjacks (Seriola spp.) in the western North Pacific. Zool Sci. 18:423–431. Ho J-S, Nagasawa K, Kim I-H. 2007. Sarcotretes longirostris n. sp. (Copepoda: Pennellidae) parasitic on bluefin driftfish (Psenes pellucidus) from the stomachs of short-finned pilot whales off Japan. J Crust Biol. 27:116–120. Ho J-S, Ohtsuka S, Nakadachi N. 2006. A new family of poecilostomatoid copepods (Umazuracolidae) based on specimens parasitic on the black scraper (Thamnaconus modestus) in Japan. Zool Sci. 23:483–496. Ho J-S, Perkins PS. 1980. Monogenea from fishes of the Sea of Japan. Part I. Order Monopisthocotylea. Ann Rep Sado Mar Biol Stat Niigata Univ. 10:1–10. Nagasawa K, Ohtsuka S. 2002. Great award to the former WAC president, Professor Ju-shey Ho. Monoculus 43:5–6. Ohtsuka S, Ho JS, Nagasawa K. 2004a. Ergasilid copepods (Poecilostomatoida) in plankton samples from Hokkaido, Japan, with reconsideration of the taxonomic status of Limnoncaea Kokubo, 1914. J Nat Hist. 38:471–498. Ohtsuka S, Ho J-S, Nagasawa K, Morozinska-Godol J, Piasecki W. 2004b. The identity of Limnoncaea Kokubo, 1914 (Copepoda: Poecilostomatoida) from Hokkaido, Japan, with the relegation of Diergasilus Do, 1981 to a junior synonym of Thersitina Norman, 1905. Syst Parasitol. 57:35–44. Ohtsuka S, Nagasawa K, Ho J-S, Grygier M. 2004c. The true nature of the enigmatic copepod genus Limnoncaea from plankton in Japan and the importance of parasito-planktology. Bull Plankton Soc Jpn. 51:13–24. [In Japanese with English abstract.]