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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
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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
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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
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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
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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.
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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)
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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
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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).
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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,
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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
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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
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Do TT, Ho J-S. 1983b. Clavellopsis nodula sp. nov. (Copepoda: Lernaeopodidae) parasitic on
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the gills of Mugil cephalus Linnaeus (Pisces: Teleostei), with proposition of a new genus
Dermoergasilus. Hydrobiologia 89:247–252.
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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.
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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.
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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.
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Biol Lab. 37:275–303.
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Didemnum molle (Herdman), in Okinawa, Japan. Proc Biol Soc Wash. 122:414–425.
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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.]