Reviews in Aquaculture (2014) 6, 147–161 doi: 10.1111/raq.

12035

The invasive potential of parasitic monogenoids

(platyhelminthes) via the aquarium fish trade: an appraisal
with special reference to India
Amit Tripathi
Department of Zoology, Rajiv Gandhi University, Itanagar, Arunachal Pradesh, India

Correspondence
Amit Tripathi, Department of Zoology,
Rajiv Gandhi University, Itanagar 791 112,
Arunachal Pradesh, India.
Email: tripathi_amit02@yahoo.co.in
Received 2 February 2013; accepted 9 April
2013.
Abstract
Aquaculture, the rearing of aquatic organisms under controlled or semi-con-
trolled conditions to produce organisms, mainly for food, ornamental and sport-
ing purposes, is a fastest-growing animal-food-producing sector in the world. The
human-mediated introduction of exotic fish has played an important role in the
success story of aquaculture. Nonetheless, many exotic fish have been implicated
in the loss of native fish biodiversity, largely through the transmission of parasites
and diseases amongst others. Whilst the transmission of parasites following the
introduction of exotic food fish is well documented, transfer of those carried by
exotic ornamental fish has received little attention. This article provides the first
summary review of the global translocation of the platyhelminthes class Monoge-
noidea via the aquarium fish trade to draw attention to the growing parasitologi-
cal risk factors associated with this form of commerce. By examining the invasive
characteristics of both aquarium fish and monogenoids, I review how the monog-
enoids fit into the different stages that an introduced species goes through when
invading the destination environment. For this, I use a theoretical framework,
synthesising published reports on the aquarium fish trade with current knowledge
on monogenoids to model invasion success. Next, I provide examples of invasive
monogenoids on exotic Indian aquarium fish and briefly discuss the vulnerability
of India to colonisation by imported aquarium fish and their parasitic monoge-
noids. I conclude that the aquarium fish trade is a perfect gateway for worldwide
translocation of monogenoids. As this trade continues to increase and intensify,
global translocation of monogenoids will expand even further.
Key words: aquarium fish trade, exotic, India, invasive potential, Monogenoidea, parasite.

The human-mediated introduction of nonnative (also

Introduction
Aquaculture is the rearing of aquatic organisms under
controlled or semi-controlled conditions to produce
organisms, mainly for food, ornamental and sporting
purposes (modified from Landau 1992). Although the
practice of aquaculture dates back at least 2000 years in
China and the Roman Empire (Balon 1995; Dunham
et al. 2001), its transformation into an important and
highly productive agricultural activity is rather recent. In
fact, aquaculture has witnessed a phenomenal growth in
the last 50 years and today stands out as the fastest-
growing animal-food-producing sector in the world,
with an average annual growth rate of 6.6% (FAO
2010).
known as alien or exotic) fish has played a very important
role in the success story of aquaculture (FAO 2005–2012a;
Gozlan et al. 2010). Therefore, about 17% of the world’s
finfish is contributed by introduced species alone; the pro-
duction of African tilapia and European turbot is signifi-
cantly higher in Asia than in most areas of Africa and
Europe, respectively; introduced salmonids in Chile con-
tribute about 20% of the world’s farmed salmon; about half
of the global production of American channel catfish origi-
nates in China and several other countries. Nonetheless, as
more and more data are generated, many exotic fish have
been implicated in the loss of native fish biodiversity, either
through direct effects, for example, competition for food
and space, predation or hybridisation (Courtenay & Moyle

© 2013 Wiley Publishing Asia Pty Ltd 147

A. Tripathi
1992; Fuller et al. 1999; Sala et al. 2000; Britton et al. 2010;
Gozlan et al. 2010), or indirect effects, for example, trans-
mission of diseases and pathogens (Kennedy 1975a; Arthur
2005; FAO 2005–2012b). In the latter instance, introduced
fish and their accompanying pathogens/diseases together
may impact native fisheries synergistically and thus more
drastically (see Fevre et al. 2006). This explains why the
introduction and spread of nonnative fish are considered
the greatest threat to fish biodiversity conservation, second
only to habitat destruction (Cambray & Pister 2002; Lepri-
eur et al. 2009), followed by chemical pollution, hybridisa-
tion and over-harvesting (Allan & Flecker 1993; Saunders
et al. 2002).
Whilst the transmission of diseases and pathogens fol-
lowing the introduction of exotic food fish holds the atten-
tion of the general public and scientists alike, those carried
by exotic ornamental fish have been poorly documented
(see Andrews 1990; Hedrick 1996; Whittington & Chong
2007; Copp et al. 2010). Strangely, this is the situation
when imported aquarium fish are known to be either
infected with or susceptible to all major groups of micro-
bial and parasitic pathogens, for example viruses (Adair &
Ferguson 1981; Hedrick & McDowell 1995; Humphrey
1995; Hegde et al. 2003), bacteria (Dixon & Contreras
1991; Humphrey & Ashburner 1993; Humphrey 1995),
fungi (Miyazaki & Egusa 1972; Lilley & Roberts 1997),
protozoan and metazoan parasites including copepods,
monogenoids, trematodes, cestodes, nematodes and
acanthocephalans (Humphrey 1995; Moravec et al. 1999;
Evans & Lester 2001; Kim et al. 2002; Thilakaratne et al.
2003). In particular, the translocation of monogenoidean
parasites via the aquarium trade and their subsequent
establishment and epidemic mortalities in wild native fish
populations have never been systematically studied and
documented.
In this article, I provide a first summary review of the
global translocation of the platyhelminthes class Mono-
genoidea via the aquarium fish trade. By examining the
invasive characteristics of both aquarium fish and monoge-
noids, I review how the monogenoids fit into the different
stages that an introduced species goes through when invad-
ing the destination environment. For this, I use a theoreti-
cal framework, synthesising published reports on the
aquarium trade with current knowledge on monogenoids
to model invasion success. Next, I provide examples of
invasive monogenoids on exotic Indian aquarium fish and
briefly discuss the vulnerability of India to colonisation by
imported ornamental fish and their parasitic monogenoids.
The objective of this study was to draw attention of all
persons, including invasion biologists, involved with the
international transport and regulation thereof of aquarium
fish to the growing parasitological risk factors associated
with this form of commerce so that immediate economic
148
benefits should not result in future risks of disease translo-
cation.
Monogenoidean parasites: pathogens of concern
Monogenoidea (adj. monogenoidean) is a class of the Phy-
lum Platyhelminthes. The member species of Monogenoi-
dea are mainly ectoparasitic on gills and/or external
surfaces of freshwater and marine fish (Bychowsky 1957); a
few species become endoparasitic by inhabiting the palleal
cavity of cephalopods, and the urinary bladder and rectum
of amphibians and reptiles (Euzet & Combes 1998 and
other references mentioned therein). None infect birds, but
one (Oculotrema hippopotami Stunkard, 1924) is known to
infect the eye of a hippopotamus, a mammal (Stunkard
1924).
These worms feed on blood (Williams & Jones 1994;
Anshary et al. 2002; Hayward et al. 2007) and/or the epi-
thelial cells and mucus (Buchmann & Bresciani 2006) of gill
tissues, leading to their fusion and hyperplasia, respiratory
epithelium displacement, and ruptured pillar cells (Martins
& Romero 1996). This causes direct losses due to mortality,
usually to younger fish and those in intensive culture or
aquaria (Thoney & Hargis 1991; Jones 2001). Damage is
frequently indirect when their haptor armed with sclero-
tised structures degrade the mucous layer, rendering the
host fish susceptible to secondary pathogens such as bacte-
ria and fungi (Grimes et al. 1985; Harris et al. 1998; Mou-
ton et al. 2001). They also block respiratory surfaces
mechanically by their huge numbers, leading to a decrease
in gill surface area and a consequent decrease in efficiency
of respiration, often causing the death of host fish by
asphyxia (Kritsky et al. 1994; Molnar 1994). In the natural
environment, however, they usually cause few problems
unless the host is continually being re-infected, leading to
the accumulation of massive numbers of worms (Yamam-
oto et al. 1984). The economic effects of their infestation
include a decrease in and/or rejection of otherwise edible
fish products (Thoney & Hargis 1991) leading to subse-
quent loss of interest in the aquaculture industries (Jones
2001; Marcogliese et al. 2001).
Aquarium fish trade: a perfect gateway for
parasitic monogenoids
The aquarium fish sector, which began as small export fish-
eries in parts of the Indo-Pacific region during the early
20th century, is now a widespread and global component
of international trade, fisheries, aquaculture and develop-
ment (FAO 2005–2012c). Over 1.5 billion fish comprising
more than 4000 freshwater and 1400 marine species are
traded worldwide each year (Whittington & Chong 2007)
generating in excess of US $ 6 billion annually (NABARD
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd

2007) with a growth rate of 14% annually since 1985 (FAO
2005–2012c). The majority of aquarium fish traded (>90%)
are of freshwater origin and farm-raised (Dawes 1998; Cato
& Brown 2003); however, 90% of marine aquarium fish are
wild-caught (Andrews 1990). The USA is the largest impor-
ter of ornamental fish, with over 10% of households pos-
sessing home aquaria (Chapman 2000), followed by
Western Europe, Japan, Taiwan and Australia (Walton
1994); however, no less than two-thirds of total –worldwide
exports originate from Asia (FAO 2005–2012c). The top
exporting country (with the percentage contributions to
global trade) is Singapore (19.8%) followed by the Czech
Republic (7.8%), Japan (7.4%), Malaysia (7.3%), Indonesia
(5.3%), Israel (4.3%), Thailand (3.9%), Sri Lanka (2.9%)
and India (0.008%) (NABARD 2007). However, it should
be noted that the ornamental fish trade data can easily be
misleading because it is often impossible to know the coun-
try of origin of fish due to transhipment and relabelling.
Although trade in aquarium fish and associated aquar-
ium products have a huge potential to contribute to the
socio-economic growth of the nations concerned (FAO
2005–2012c; Del Rio Rodriguez 2006), the beneficial impli-
cations are not always straightforward. In fact, the growth
in the aquarium trade is paradoxically leading to the col-
lapse of fishery stocks worldwide for three reasons: first,
with the increasing aquarium trade, ornamental fish are
over-harvested, threatening the sustainability of fisheries
(Cato & Brown 2003; Lunn & Moreau 2004); second, with
the increasing aquarium trade, more nonnative aquarium
fish are translocated worldwide, leading to the demise of
native fish fauna (Pimentel 2002); and third, the uncon-
trolled transfer of aquarium fish facilitates the interconti-
nental translocation of many parasites and diseases
(Ferguson et al. 1994; Robertson & Austin 1994; Hedrick &
McDowell 1995; Humphrey 1995; Evans & Lester 2001;
Kim et al. 2002) which may have a distressing impact on
the native fish of the importing country (Goodwin 2002;
Thilakaratne et al. 2003; Gozlan et al. 2005; Go et al.
2006).
There are many sporadic examples of the translocation
of monogenoidean parasites together with ornamental
fish. For example, Gyrodactylus cyprini Diarova, 1964
(Gyrodactylidae) was introduced into the United States
with the common carp Cyprinus carpio (Cypriniformes:
Cyprinidae), from Europe (Bauer & Hoffman 1976).
Dactylogyrus ostraviensis Rehulka 1988 (Dactyologyridae)
was translocated to Czechoslovakia with the rosy barb
Barbus conchonius (now Puntius conchonius) (Cyprinifor-
mes: Cyprinidae) imported from India (Rehulka 1988). A
plethora of dactylogyrid species was introduced into the
United States with the goldfish Carassius auratus (Cyprin-
iformes: Cyprinidae), from endemic regions of Eurasia
(Bauer & Hoffman 1976; Kritsky & Heckmann 2002).
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd
Invasive monogenoids via aquarium trade
Thaparocleidus caecus (Mizelle & Kritsky 1969) Lim 1996;
(Dactylogyridae) was introduced into the United States
(Mizelle & Kritsky 1969) and Peninsular Malaysia (Lim
1996) along with the striped catfish Pangasius sutchi (now
Pangasianodon hypophthalmus) (Siluriformes: Pangasii-
dae), from Thailand. Gussevia asota Kritsky, Thatcher and
Boeger, 1989 (Dactylogyridae) and Gyrodactylus bullataru-
dis Turnbull, 1956 (Gyrodactylidae) were translocated to
Korea along with the oscar Astronotus ocellatus (Percifor-
mes: Cichlidae), and the platy Xiphophorus maculatus
(Cyprinodontiformes: Poeciliidae), respectively, from
South-East Asia (Kim et al. 2002). Heteropriapulus
heterotylus (Jogunoori et al. 2004) Kritsky 2007 (Dactylo-
gyridae), Diaphorocleidus armillatus Jogunoori et al.
2004 (Dactylogyridae) and Urocleidoides vaginoclaustrum
Jogunoori et al. 2004 (Dactylogyridae) were translocated
to India along with Hypostomus sp. (Siluriformes:
Loricariidae), the black tetra Gymnocorymbus ternetzi
(Characiformes: Characidae), and the green swordtail
Xiphophorus helleri (Cyprinodontiformes: Poeciliidae),
respectively, from the neotropics (Jogunoori et al. 2004;
Kritsky 2007). A wide range of worms, including Dactylo-
gyrus spp. (Dactylogyridae) and Gyrodactylus spp. (Gyro-
dactylidae), entered Australia via the aquarium fish trade
over the period 1995–2004 (Humphrey 1995; Whittington
& Chong 2007). Dactylogyrus coartatus Rehulkova & Gel-
nar, 2006; D. macrocolpius Rehulkova & Gelnar 2006 and
D. melanopteri Rehulkova & Gelnar, 2006 (Dactylogyri-
dae) were translocated to Czech Republic along with the
bala sharkminnow Balantiocheilos melanopterus (Cyprini-
formes: Cyprinidae), from Thailand (Rehulkova & Gelnar
2006). Sciadicleithrum iphthimum Kritsky, Thatcher, and
Boeger, 1989 (Dactylogyridae) was introduced into India
along with the freshwater angelfish Pterophyllum scalare
(Perciformes: Cichlidae), from South America (Tripathi
et al. 2010). Two Onchocleidus spp. (Ancyrocephalidae),
one Onchocleidus dispar Muller, 1936 (sensu Wheeler and
Beverley-Burton 1989) and the other probably Onchoclei-
dus similis Muller 1936, were introduced to Norway along
with the pumkinseed Lepomis gibbosus (Perciformes:
Centrarchidae), most likely from Czech Republic (Sterud
& Jorgensen 2006). Lepomis gibbosus, a native to North
America, has also been responsible for the introduction
of Actinocleidus recurvatus Mizelle and Donahue, 1944
and A. oculatus Mueller, 1934 (Dactylogyridae), and Uro-
cleidus similis Mueller, 1936 and U. dispar Mueller, 1936
(Dactylogyridae) to Italy (Galli et al. 2003).
Unfortunately, due to the absence of any organised
studies of most of the above cases, transfer of exotic mo-
nogenoids from ornamental fish to native fish and their
subsequent mortalities in the wild has not yet been shown
(as far as I know). Nonetheless, we know from experience
with global introductions of food fish that imported exo-
149
A. Tripathi
tic monogenoidean parasites may pose a devastating
impact on the importing aquaculture industry, threaten-
ing human economies and health. For example, the intro-
duction of Nitzschia sturionis Abildgaard, 1794
(Capsalidae) from the Caspian Sea into the Aral Sea along
with sturgeon Huso huso (Acipenseriformes: Acipenseri-
dae) led to the total collapse of the endemic Aral Sea
sturgeon Acipenser nudiventris (Acipenseriformes: Acipen-
seridae) (Dogiel & Lutta 1937). Norway suffered huge
losses of wild salmon populations because of the translo-
cation of Gyrodactylus salaris Malmberg, 1957 (Gyrodac-
tylidae) along with Atlantic salmon smolts Salmo salar
(Salmoniformes: Salmonidae) into their natural environ-
ment from Sweden (Johnsen & Jensen 1991); the resulting
economic losses ran in excess of US $500 million (King
& Cable 2007). The Japanese aquaculture industry was
seriously affected when Neobenedenia girellae (Hargis,
1955) Yamaguti, 1963 (Capsalidae) was introduced there
along with imported amberjack fry Seriola dumerili (Perc-
iformes: Carangidae) from China (Ogawa et al. 1995;
Gaughan 2001). Similarly, Neoheterobothrium hirame Og-
awa, 1999 (Diclidophoridae), believed to have invaded
Asian waters from North America via commercial impor-
tation of either the summer flounder Paralichthys denta-
tus (Pleuronectiformes: Paralichthyidae) (Hayward et al.
2001a), or the southern flounder Paralichthys lethostigma
(Pleuronectiformes: Paralichthyidae) (Tsutsumi 2004), has
led to a marked decline in the commercial catch of the
olive flounder Paralichthys olivaceus (Pleuronectiformes:
Paralichthyidae), in Japan (Anshary et al. 2002). Unfortu-
nately, this parasite has not only become ubiquitous
throughout most Japanese waters but has expanded its
range into Korean waters (Mushiake 2001). Pseudodacty-
logyrus anguillae (Yin & Sproston, 1948) Gusev, 1965 and
P. bini (Kikuchi, 1929) Gusev, 1965 (Pseudodactylogyri-
dae) were introduced from Asia to North America and
Europe with the importation of the Japanese eel
Anguilla japonica (Anguilliformes: Anguillidae); both spe-
cies then infected the native North American eel
Anguilla rostrata and the native European eel A. Anguilla
(Anguilliformes: Anguillidae), causing mass mortality
(Hayward et al. 2001a,b). Similarly, Dactylogyrus vastator
Nybelin, 1924 (Dactylogyridae), introduced into Europe
with the grass carp Ctenopharyngodon idella (Cyprinifor-
mes: Cyprinidae) from Asia, caused losses in European
carp cultures (Schaperclaus 1990).
The aquarium trade and monogenoids: assessing
the invasive potential
Understanding the different stages that an introduced
species goes by in order to successfully invade an ecosys-
tem could be of great practical importance for predicting
150
and preventing future invasions (Garcia-Berthou 2007).
Host–parasitic invasions are complex processes dependent
upon the probability of moving through the following
chain of five sequential phases: (i) occurrence of infec-
tion; (ii) translocation to a new region; (iii) release to the
wild; (iv) establishment; and (v) the ecological and/or
socio-economic impacts (modified from Williamson
1996; Kolar & Lodge 2001). It is to be emphasised here
that the invasive success of a parasite, including monoge-
noids, should essentially be seen as a function of the
combined invasive success of both the parasite and the
host it has been co-introduced with.
Occurrence
Occurrence, the probability that a parasitic infection will
take place, is the first and most important step in the inva-
sion process. As each species of fish is typically infected
with at least one species of Monogenoidea (Whittington
1998; Williams & Jones 1994), the probability of occurrence
of parasitic monogenoids is greater than that for any other
pathogens associated with the importation of ornamental
fish.
The taxonomic diversity of a species is said to be one of
the best indicators of invasion success (Carlton & Geller
1993; Levine & D’Antonio 2003; Shannon et al. 2005).
Thus, the huge taxonomic diversity of both monogenoids
(Poulin 2002) and imported aquarium fish (Blanc 2001)
further optimise the occurrence of monogenoids. The esti-
mated number of described species of Monogenoidea is
already ‘on the order’ of 10 000 (Delane C. Kritsky, pers.
comm., 2011), whilst that of traded aquarium fish is about
5400 plus (Whittington & Chong 2007).
Translocation
The monogenoids have a high probability of transloca-
tion together with their host fish. This appears to be a
function of two key factors: host specificity and quaran-
tine procedures. Host specificity, defined as the ‘restric-
tion of a parasite to certain host species’ for maximising
its reproductive success (Kennedy 1975b), is an extremely
important parameter of invasion success (Shea & Chesson
2002). Monogenoids are the most host-specific of all fish
parasites (Whittington et al. 2000) and thus score high
by this criterion. The fact that the monogenoids display a
high level of co-evolution via co-speciation by virtue of
their high host specificity (Noble et al. 1989; Kearn 1994)
leads us to believe these worms remain in exceptionally
close contact with their hosts and consequently will be
co-transported together with them. In other words, the
high host specificity of monogenoids ensures that they
are not lost or ‘left behind’ but remain on their hosts
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd

during the translocation process. The growing list of mo-
nogenoids that are being translocated around the world
by the aquarium trade (reviewed above) is an eloquent
testimony of this theory.
Either many countries do not have proper quarantine
procedures for the import of aquarium fish, or their imple-
mentation is inadequate (see Thomas et al. 2009; Secretar-
iat of the Convention on Biological Diversity 2010); nor are
there sufficient data for risk assessment of their monoge-
noids. Even when a quarantine procedure has been per-
formed, fish in quarantine can still be harbouring
monogenoids for two reasons. First, the morpho-taxo-
nomic detection of monogenoids, especially freshwater
varieties, is an extremely difficult task owing to their small
sizes. Second, host fish infected with monogenoids often
show few or no histopathological signs (Paperna 1996)
except for pale gills, excess mucus production, skin erosion
and focal haemorrhagic lesions (Tripathi 1957; Kabata
1985). Thus, the mortalities in wild Atlantic salmon popu-
lations in Norway were long attributed to industrial pollu-
tion when they were actually caused by the monogenoid
G. salaris (Johnsen & Jensen 1986).
Release
Traditionally, aquarium fish are not believed to come into
contact with farmed or wild fish (Davenport 2000), and
hence, it is assumed that their parasites, including monoge-
noids, will not have a chance to infect the native fish. How-
ever, there is increasing evidence that ornamental fish are
frequently released, accidentally or otherwise, into the nat-
ural waters of importing countries, where they establish
permanent self-reproducing populations. For example, of
about 100 species of ornamental fish that have been intro-
duced into North America via the aquarium trade, up to 40
have established populations (Crossman & Cudmore 1999;
Fuller et al. 1999). Similarly, at least 22 species of ornamen-
tal fish that were imported into Australia have now estab-
lished breeding populations in the wild (Whittington &
Chong 2007). In the same way, at least 18 species of
imported ornamental fish have established permanent self-
reproducing populations in Indian waters, especially in
peninsular India (see section ‘Indian perspective’ below).
In fact, there is growing evidence that many instances of
ornamental fish into the wild are due to deliberate human
abandonment of these fish (Courtenay & Taylor 1986;
Copp et al. 2005). For example, aquarium fish species are
known to be routinely released into natural water bodies by
retailers and private owners, because either their large sizes,
aggressiveness or high reproductive rates prove inconve-
nient to them (Padilla & Williams 2004); and by the general
public for spiritual gains (see section ‘Indian perspective’
below).
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd
Invasive monogenoids via aquarium trade
Establishment
The factors often cited as associated with the establishment
success of invading species are propagule pressure, biologi-
cal attributes and global invasion history. The propagule
pressure, that is, the number of individuals introduced and
the number of times they are introduced, enhances the
establishment success of ornamental fish (Marchetti et al.
2004; Lockwood et al. 2005; Gertzen et al. 2008). Thus, the
most popular fish sold in the aquarium trade are also the
most likely to become introduced and established in the
wild (Duggan et al. 2006). Because the success of the orna-
mental fish industry depends heavily on the export and
import of fish species (FAO 2005–2012c), the ever-growing
and profitable trade in aquarium fish can only be expected
to introduce more fish in even greater volumes (i.e. high
propagule pressure) in the future, which, in turn, will lead
to higher rates of successful establishment of aquarium fish
and their monogenoids in the wild.
The establishment success is further optimised using
two key features of the biology of monogenoidean para-
sites. First, the direct life cycle (i.e. intermediate hosts are
absent) of monogenoids necessarily means that more
than one suitable host species (a definitive and an inter-
mediate host) is not necessary for the establishment of
parasite in the new locality. For example, it might be
much more unlikely for a digenean parasite to establish
when a specific snail and a definitive host are required to
complete the life cycle. Second, monogenoids have short
life spans with high reproductive potential/fecundity
(Bychowsky 1957; Lester 1972), which, coupled with their
hermaphroditic nature, allows them to reproduce rapidly
and directly reinfect their hosts successfully (Bauer 1991).
Hence, Tautog Tautoga onitis (Perciformes: Labridae)
developed massive infestations of an unknown species of
Gyrodactylus, which is not a normal parasite of tautog,
within a month after being placed in aquaria with several
other types of fish at the New York Aquarium (Thoney
& Hargis 1991).
Information available is not sufficient to judge the role of
the past history of successful invasion of monogenoids.
Nonetheless, there are hints that some species of both fish
and monogenoids can establish exotic populations more
readily than others and hence should be dealt with caution.
For example, all 18 exotic Indian ornamental species of fish
(reviewed below) have established themselves in at least
one other country (see Froese & Pauly 2012). Similarly,
some monogenoids, for example, Gyrodactylus anguillae
Ergens, 1960 (Gyrodactylidae), Pseudodactylogyrus anguil-
lae and P. bini (Pseudodactylogyridae), who were translo-
cated with the shipment of angullid eels for aquaculture,
have established their population worldwide (see Gelnar
et al. 1996; Hayward et al. 2001b,c).
151
A. Tripathi
Whether exotic monogenoids remain on their intro-
duced fish or switch to native ones is a debating point. The
establishment of exotic monogenoids on the native host
fish via host switching is often considered less likely than
for other parasitic groups due to the generally high host
specificity of monogenoids, combined with the phyloge-
netic dissimilarity of native and exotic fish (Ernst & Dove
1998). However, there are many known species of Monoge-
noidea, which show an amazingly low host specificity and
thus are potential candidates for infecting native hosts. For
instance, Dactylogyrus anchoratus (Dujardin, 1845) Wagen-
er, 1857 and D. extensus Mueller and Van Cleave, 1932
(Dactylogyridae) have been reported from 19 and seven
genera of fish, respectively (see Gibson et al. 1996). In fact,
D. anchoratus, which was imported to Iran along with
genetically improved specimens of Cyprinus carpio from
Romania and Hungary, has now switched over to Bar-
bus sharpeyi, an important native host species in the
Khuzestan Province of the Tigris system of Iran (Shamsi
et al. 2009). Transfer from introduced African Cichlids to
native Mexican cichlids has also been shown for Cichlidogy-
rus longicornis Paperna and Thurston, 1969, C. sclerosus
Paperna and Thurston, 1969, C. tilapiae Paperna, 1960
(Dactylogyridae) and Enterogyrus malmbergi Bilong Bilong,
1988 (Dactylogyridae) (Jimenez-Garcia et al. 2001). D. vas-
tator, which was introduced into Europe with the grass carp
Ctenopharyngodon idella from Asia, has established itself on
native populations of European cyprinid fish (Schaperclaus
1990). King and Cable (2007) have categorically shown, at
least under aquarium conditions, the establishment and
reproduction of Gyrodactylus turnbulli Harris, 1986 (Gyro-
dactylidae) on a range of poeciliids and even more distantly
related fish. In fact, Gyrodactylus spp., considered to be
amongst the most invasive fish parasites (Kennedy 1994),
have a habit of recurrent ‘host switching’ to promote speci-
ation (Harris et al. 2004). As Gyrodactylus spp. have a
viviparous mode of reproduction, with an exponential
growth rate and no specialised transmission stage (oncomi-
racidia) (Scott & Anderson 1984), they are likely to be
amongst the most successful invaders of an exotic host
(Torchin et al. 2003), infecting new hosts and environ-
ments following introduction.
Besides, monogenoids frequently exhibit low host speci-
ficity in aquaria (Thoney & Hargis 1991), where they may
infect any available species of fish (Bakke et al. 1991, 1992).
Consequently, the very nature of ‘aquaria’ (where numerous
ornamental species of fish from multiple source countries
co-habit) provides monogenoids an ideal opportunity to
infect other, including native, ornamental species of fish in
the importing country. For instance, Neobenedenia melleni
(MacCallum, 1927) Yamaguti 1963 (Capsalidae) has been
found infesting as many as 21 families of fish at the New
York Aquarium, USA (Jahn & Kuhn 1932; Nigrelli 1937).
152
Ecological and/or socio-economic impact
The actual damage caused by invasive monogenoids intro-
duced through the aquarium trade in any one area is open
to question, as experimental studies are unavailable. Part of
the problem, of course, is that it is difficult to detect the
mortalities in the wild. The scientific community still needs
to devise a blueprint measuring the impact of an intro-
duced parasite species. Nonetheless, once exotic monoge-
noids introduced via the aquarium trade become
successfully established in the wild, we do not have any rea-
son to doubt they can destroy native fish just the way mo-
nogenoids from food fish have done, as reviewed above.
Hoffman (1970) noted: ‘the potential already exists: the risk
of disease should not be ignored just because a particular
disease cannot be detected’.
Indian perspective
The Indian aquarium fish sector is a small but vivacious
segment with an export value of approximately US$ 1.17
million (Silas et al. 2011). There are over 470 recorded fish
species in India that are either exported or have the poten-
tial to be exported as aquarium fish (Froese & Pauly 2012).
In contrast to global trends, 90% of the export trade in
India is based on wild collection (Sundaresan 2011). Unfor-
tunately, concise details on the introduction of exotic
Indian ornamental fish and their subsequent release into
the wild are mostly unavailable (Singh & Lakra 2011). Nev-
ertheless, an exhaustive review of the literature shows that
more than 170 species of ornamental fish have been intro-
duced into India via the aquarium trade, of which at least
18 now have permanent self-reproducing populations in
natural waters, especially in peninsular India, representing
six families (see Table 1). The invasive success of ornamen-
tal fish in India (and probably elsewhere too) is evidenced
by the fact that against 18 exotic ornamental fish, only nine
exotic food fish (i.e. having little or no aquarium value)
have been able to establish populations in Indian waters
(see Table 2).
In India, there have been several studies, which docu-
mented invasive aquarium fish species as a direct source of
biodiversity erosion (e.g. Sehgal 1989; Shetty et al. 1989;
Jhingran 1991; Sreenivasan 1996; Singh & Mishra 2001;
Singh & Das 2006; Singh & Lakra 2006, 2011; Raghavan
et al. 2008a,b; Knight & Rema Devi 2009; Krishnakumar
et al. 2009; Knight 2010). However, the parasitic fauna of
exotic Indian ornamental fish are poorly characterised rela-
tive to those in other countries, and for monogenoids there
have been no all-inclusive and ‘aquarium fish–targeted’
surveys to date. To the best of my knowledge, 13 species of
Monogenoidea have been described/recorded from exotic
Indian ornamental fish (see Table 3). Assuming that each
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd
 Invasive monogenoids via aquarium trade

Table 1 Exotic ornamental fish species intro-

duced and reproductively established in Indian Family Species Origin Year Distribution
waters (after many contributions in Rema Devi Cyprinidae Cyprinus carpio Thailand, 1939 Throughout India
1987; Shetty et al. 1989; Biju Kumar 2000; carpio Sri Lanka
Kharat et al. 2003; Chandrasekhar 2004; Carassius auratus Japan Unknown Peninsular India
Global Invasive Species Database 2005; Daniels auratus
2006; Singh & Das 2006; Raghavan et al. Tinca tinca UK 1870 Throughout India
2008a,b; Knight & Rema Devi 2009; Krish- Carassius carassius UK 1870 Throughout India
nakumar et al. 2009; Knight 2010; Singh & Barbonymus gonionotus Indonesia 1972 East India
Lakra 2011; Froese & Pauly 2012) Cichlidae Oreochromis Thailand, 1952 Throughout India
mossambicus Sri Lanka
Cichlasoma trimaculatum Unknown Unknown Peninsular India
Poeciliidae Gambusia affinis Japan Unknown Throughout India
Gambusia holbrooki Italy 1928 Throughout India
Xiphophorus hellerii Unknown Unknown Peninsular India
Xiphophorus maculates Unknown Unknown Peninsular India
Poecilia reticulata South America 1908 Peninsular India
Osphronemidae Osphronemus Mauritius, 1800–1849 Peninsular India
goramy Indonesia
Trichopodus trichopterus Unknown Unknown Peninsular India
Loricariidae Pterygoplichthys Unknown Unknown Peninsular India
multiradiatus
Pterygoplichthys Unknown Unknown North India
disjunctivus
Pterygoplichthys pardalis Unknown Unknown North India
Characidae Pygocentrus natterei Unknown Unknown Peninsular India

Table 2 Exotic food fish species introduced

and reproductively established in Indian waters Family Species Origin Year Distribution
(after many contributions in Jhingran 1991; Salmonidae Salmo trutta fario UK 1863 Upland waters
Singh & Mishra 2001; Singh 2004; Global Inva- of North and
sive Species Database 2005; Singh & Lakra South India
2006; Lakra et al. 2008; Singh & Lakra 2011; Oncorhynchus Sri Lanka, U.K., New 1906 Upland waters of
Froese & Pauly 2012) mykiss Zealand, Germany North India
Salmo trutta trutta UK 1900 Upland waters of
North India
Salvelinus fontinalis Canada 1960 Upland waters of
North India
Cyprinidae Ctenopharyngodon idella Hong Kong 1959 Throughout India
Hypophthalmichthys Hong Kong, Japan 1959 North India
molitrix
Clariidae Clarias gariepinus Thailand 1993 Throughout India
Cichlidae Oreochromis mossambicus Thailand, Sri Lanka 1952 Throughout India
Oreochromis niloticus Thailand 1990 Throughout India

fish species in the world harbours a minimum of one spe-
cies of Monogenoidea (Whittington 1998), and all the 170
plus exotic Indian ornamental host fish were included, the
total number of monogenoidean species from India known
at present reflects only a tiny proportion of those actually
in the country.
The hypothesis that introduced species of Monogenoidea
might cause a serious parasitological problem in India can
be appreciated against the background of the large number
of documented examples of exotic species other than
monogenoids already impacting the country. In the 1990s,
an exotic virus species (WSSV; White spot syndrome
virus), believed to have entered India from the People’s
Republic of China, cost the Indian shrimp industry millions
of dollars in just one decade (Vijayan & Rao 2009). During
1992–1995, Aphanomyces invadans, an exotic fungus of
freshwater and estuarine fish, believed to have entered India
from the neighbouring state of Bangladesh through their
common river system, caused an estimated loss of 42.5 mil-
lion US dollars (Lilley et al. 2002).

Reviews in Aquaculture (2014) 6, 147–161

© 2013 Wiley Publishing Asia Pty Ltd 153
A. Tripathi

Table 3 Invasive monogenoids from exotic Indian ornamental fish

Species Host fish Reference

Diplozoon nipponicum Goto, 1891 Cyprinus carpio carpio Fotedar and Parveen (1987)
Gyrodactylus medius Cyprinus carpio carpio Devaraj et al. (1977)
Kathariner, 1895
Carassius auratus Devaraj et al. (1977) and
Chanda et al. (2011)
Dactylogyrus sp. Carassius auratus Chanda et al. (2011)
Pellucidhaptor kritskyia Singh, Rastogi Carassius sp. Singh et al. (2003)
and Rastogi, 2003
Heteropriapulus heterotylus (Jogunoori, Kritsky and Hypostomus sp. Kritsky (2007)
Venkatanarasaiah, 2004) Kritsky, 2007
Diaphorocleidus armillatus Jogunoori, Kritsky and Gymnocormbus ternetzi Jogunoori et al. (2004)
Venkatanarasaiah, 2004
Urocleidoides vaginoclaustrum Jogunoori, Kritsky Xiphophorus helleri Jogunoori et al. (2004)
and Venkatanarasaiah, 2004
Sciadicleithrum iphthimum Kritsky, Thatcher Pterophyllum scalare Tripathi et al. (2010)
and Boeger, 1989
Gussevia spiralocirra Kohn and Paperna, 1964 Pterophyllum scalare Personal observation;
unpublished data
Silurodescoides exotica Rastogi, Mishra, Rastogi, Corydoras melanistius Rastogi et al. (2008)
Sharma and Singh, 2008
Silurodiscoides vistulensis Siwak (1932) Bychowsky Pangasianodon hypophthalmus Rastogi et al. (2008)
and Nagibina, 1957
Thaparocleidus siamensis (Lim, 1990) Lim, 1996 Pangasianodon hypophthalmus Personal observation;
unpublished data
Thaparocleidus caecus (Mizelle & Kritsky, Pangasianodon hypophthalmus Personal observation;
1969) Lim, 1996 unpublished data

India appears to be particularly vulnerable to colonisa-
tion by imported ornamental fish (and their parasitic mo-
nogenoids) due to the following reasons:
1 The country, and its north-east and western ghat regions,
in particular, is bestowed with a model combination of
climatic conditions ideally conducive for the growth,
maturation and breeding of escapee exotic ornamental
fish, especially those originating from parts of South
America and South-East Asia having similar climates.
2 The aquarium trade in India imports ornamental species
of fish with a huge taxonomic structure, both in terms of
the total number of species and their taxonomic diversity
(see Froese & Pauly 2012). As the taxonomic diversity of
a species is an important indicator of invasion success
(reviewed above), the vast taxonomic structure of
imported ornamental species of fish represents a serious
invasive potential for the aquatic ecosystems of India.
3 In Indian mythology, fish are considered to be an incar-
nation of Lord Vishnu and hence a sanctified animal. In
many places, followers of Hinduism regularly ‘release’ or
‘liberate’ live fish back into the river water to earn per-
sonal spiritual gains. Because it is both easier and time-
saving to purchase fish from aquarium markets, rather
than going to a river spot and catching a fish with a myr-
iad array of fishing gears, devotees (end-users) purchase
the fish directly from aquarium markets and release
them into the wild. The exact numbers of purchased-
and-released fish are not available, but no fewer than
thousands of both native and exotic ornamental fish spe-
cies are purposely and regularly released from aquarium
shops into inland waters of the country every year.
4 India has one of the largest networks of rivers in the
world; the mainland is drained by 15 major (drain-
age basin area > 20 000 km2), 45 medium (2000–
20 000 km2) and over 120 minor (<2000 km2) rivers,
along with their tributaries and distributaries (Rao
1975). Such a vast network of rivers maximises the dis-
persal options for both native and nonnative escapee fish
to ‘trickle’ and colonise new habitats across state bound-
aries (see Hendry et al. 2004 for discussion).
5 In India, the state of knowledge of fish hygiene is not
advanced nor controls enforced (Anonymous 1998, 1999
cited in Kamat et al. 2002). This is a serious concern
because polluted conditions are known to increase the
infestation levels of monogenoids possibly by compro-
mising the immunity of host fish (Skinner 1982; Khan &
Kiceniuk 1988; Koskivaara et al. 1992; Bagge & Valtonen
1996).
The presence of 18 exotic ornamental fish species in
Indian freshwaters identifies a major threat to be addressed

Reviews in Aquaculture (2014) 6, 147–161

154 © 2013 Wiley Publishing Asia Pty Ltd

by researchers. A comparison of their monogenoids both
from India and exporting countries may help quantifying
their parasitological impact(s) on the regional ecology and
economy of India.
Conclusions and recommendations
The aquarium fish trade is a perfect gateway for the world-
wide translocations of monogenoidean parasites, and the
situation is no different in India. As this trade continues to
increase and intensify, global translocation of monogenoids
will expand even further. Although we lack substantial and
quantifiable data to confirm actual risks consequences aris-
ing out of these introductions to date, given the historical
instances of the harmful consequences of exotic monoge-
noids from imported food fish, potential risks are always
associated with the aquarium fish trade. To make matters
even more complicated, many countries have either no leg-
islation or proper quarantine procedures for the import of
ornamental fish nor are there enough data for risk assess-
ment of monogenoids. The fact that we have no precise
answer yet to a question as simple as to how many exotic
monogenoids we have worldwide shows an alarming
knowledge gap in this field.
To redress the situation, I recommend following five key
steps be taken:
• Key Step 1. Develop an international database or a basic
taxonomic monograph, something similar to that of Ya-
maguti’s (1963) classical manual, with a list of species of
Monogenoidea on a wide range of ornamental species of
fish from different parts of the world. This is absolutely
crucial for formulating or upgrading vital procedures for
introduction, quarantine and certification.
• Key Step 2. Initiate international collaborations for con-
certed and sustained parasitological sampling efforts
from the source and destination environments of intro-
duced aquarium fish. This will help validating the inva-
sive potential of monogenoids.
• Key Step 3. Formulate a strong risk assessment frame-
work to prevent the import of ornamental fish infested
with monogenoids. However, a major problem exists in
obtaining not-so-easily accessible background informa-
tion, for example, biodiversity, general development and
life cycle, and a history of successful establishment, if
any, of both imported ornamental species of fish and
associated monogenoid(s); ecosystem of both exporting
and importing countries; and cultural and legal perspec-
tives of importing country. The valuable discussions on
risk assessment for aquarium fish trade are available in
the literature (see AQIS 1999; BNZ 2005; Rixon et al.
2005; Whittington & Chong 2007; Roelofs 2008), and
the same can be refined and improved to ‘fit’ into the
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd
Invasive monogenoids via aquarium trade
remedial investigations on monogenoids introduced by
aquarium fish trade.
• Key Step 4. Develop molecular tools, which can detect
the presence of monogenoids from the gill mucus of host
fish using nonlethal sampling methods (see for example,
Mozhdeganlou et al. 2011; Ek-Huchim et al. 2012). As
freshwater monogenoids are tiny and can be easily over-
looked, especially when their incidence and intensity is
on a lower side, such tools may help in putative diagno-
sis of the worms in question.
• Key step 5. Prepare a comprehensive active/targeted sur-
veillance system as a part of quarantine strategy to gather
and assess information on fish health status with respect
to specific monogenoids. The design of the system
should incorporate importer facilities, breeding facilities
and higher-risk water bodies, for example, those situated
close to ornamental fish culture sites. This will help
maximising the likelihood of detecting exotic monoge-
noids.
Acknowledgements
This study was supported by research grants from Univer-
sity Grant Commission, New Delhi, India [39- 603/2010
(SR)], and Department of Science and Technology, New
Delhi, India [SR/SO/AS-56/2011].
References
Adair BM, Ferguson HW (1981) Isolation of infectious pancre-
atic necrosis (IPN) virus from non-salmonid fish. Journal of
Fish Diseases 4: 69–76.
Allan JD, Flecker AS (1993) Biodiversity conservation in running
waters. BioScience 43: 32–43.
Andrews C (1990) The ornamental fish trade and fish conserva-
tion. Journal of Fish Biology 37: 53–59.
Anonymous (1998) EU import ban affects 40% frozen fish as 30
countries remain on waiting list, Marine Products Export
Development Authority Cochin. Newsletter 42–43.
Anonymous (1999) Value added shrimp export is under threat,
Marine Products Export Development Authority, Cochin,
India. Newsletter 30–32.
Anshary H, Yamamoto E, Miyanaga T, Ogawa K (2002) Infec-
tion dynamics of the monogenean Neoheterobothrium hirame
infecting Japanese flounder in the western Sea of Japan. Fish
Pathology 37: 131–140.
AQIS (1999) Import Risk Analysis on Live Ornamental Fish. Aus-
tralian Quarantine and Inspection Service, Commonwealth of
Australia, Canberra, 172 pp. (Available at: http://www.daff.
gov.au/__data/assets/pdf_file/0018/16362/finalornamental.pdf)
Arthur JR (2005) A historical overviews of pathogen introduc-
tions and their trans-boundary spread in Asia. In: Subasinghe
RP, Arthur JR (eds) Preparedness and Response to Aquatic
155
A. Tripathi
Animal Heath Emergencies in Asia, pp. 21–39. FAO Fisheries
Proceedings, No 4. FAO, Rome.
Bagge A, Valtonen ET (1996) Experimental study on the influ-
ence of paper and pulp mill effluent on the gill parasite
communities of roach (Rutilus rutilus). Parasitology 112:
499–508.
Bakke TA, Jansen PA, Hansen LP (1991) Experimental transmis-
sion of Gyrodactylus salaris Malmberg, 1957 (Platyhelminthes,
Monogenea) between the Atlantic salmon (Salmo salar) and
the European eel (Anquilla anquilla). Canadian Journal of
Zoology 69: 733–737.
Bakke TA, Harris PD, Jansen PA, Hansen LP (1992) Host speci-
ficity and dispersal strategy in gyrodactylid monogeneans,
with particular reference to Gyrodactylus salaris Malmberg
(Platyhelminthes, Monogenea). Diseases of Aquatic Organisms
13: 63–74.
Balon EK (1995) Origin and domestication of the wild carp,
Cyprinus carpio: from Roman gourmets to the swimming
flowers. Aquaculture 129: 3–48.
Bauer ON (1991) Spread of parasites and diseases of aquatic
organism by acclimatization: a short review. Journal of Fish
Biology 39: 679–686.
Bauer ON, Hoffman GL (1976) Helminth range extension by
translocation of fish. In: Page LA (ed.) Wildlife Diseases, pp.
163–172. Plenum Publication, New York.
Biju Kumar A (2000) Exotic fishes and freshwater fish diversity.
Zoos’ Print Journal 15: 363–367.
Blanc G (2001) Introduction of pathogens in European aquatic
ecosystems: attempt of evaluation and realities. In: Uriarte A,
Basurco B (eds) Environmental Impact Assessment of Mediter-
ranean Aquaculture Farms, pp. 37–56. CIHEAM-IAMZ, Zar-
agoza, Spain.
BNZ (2005) Import Risk Analysis: Ornamental Fish. Biosecurity
New Zealand, Ministry of Agriculture and Forestry, Welling-
ton, 270 pp. Available at: http://www.biosecurity.govt.nz/files/
regs/imports/risk/ornamental-fish-ira.pdf).
Britton JR, Davies GD, Harrod C (2010) Trophic interactions
and consequent impacts of the invasive fish Pseudorasbora
parva in a native aquatic foodweb: a field investigation in the
UK. Biological Invasions 12: 1533–1542.
Buchmann K, Bresciani J (2006) Monogenea (Phylum Platyhel-
minthes). In: Woo PTK (ed.) Fish Diseases and Disorders, Vol.
1. Protozoan and Metazoan Infections, pp. 297–344. CAB
International, Wallingford.
Bychowsky BE (1957) Monogenetic Trematodes, Their Systematic
And Phylogeny. Akademii Nauk SSR, Moscow-Leningrad, pp.
509. [In Russian; English translation edited by Hargis WJ, Jr
(1961), American Institute of Biological Sciences, Washing-
ton, DC. pp. 627.]
Cambray JA, Pister EP (2002) The role of scientists in creating
public awareness for the conservation of fish species: Afri-
can and American case studies. In: Collares-Pereira MJM,
Coelho M, Cowx I (eds) Conservation of Freshwater Fishes:
Options for the Future, pp. 414–423. Blackwell Science,
Oxford.
156
Carlton JT, Geller KB (1993) Ecological roulette: the global
transport of nonindigenous marine organisms. Science 261:
78–82.
Cato JC, Brown CL (2003) Marine Ornamental Species: Collec-
tion, Culture, and Conservation. Iowa State Press, Blackwell
Publishers, Ames.
Chanda M, Paul M, Maity J, Dash G, Gupta SC, Patra BC (2011)
Ornamental fish Goldfish, Carassius auratus and related para-
sites in three districts of West Bengal, India. Chronicles of
Young Scientists 2: 51–54.
Chandrasekhar SVA (2004) Fish fauna of Hyderabad and its
environs. Zoos’ Print Journal 19: 1530–1533.
Chapman FA (2000) Ornamental fish culture, freshwater. In:
Stickney RR (ed.) Encyclopaedia of Aquaculture, pp. 602–610.
John Wiley & Sons, New York.
Copp GH, Wesley KJ, Vilizzi L (2005) Pathways of ornamental
and aquarium fish introductions into urban ponds of Epping
Forest (London, England): the human vector. Journal of
Applied Ichthyology 21: 263–274.
Copp GH, Vilizzi L, Gozlan RE (2010) The demography of
introduction pathways, propagule pressure and occurrences
of nonnative freshwater fish in England. Aquatic Conservation
– Marine and Freshwater Ecosystems 20: 595–601.
Courtenay WR Jr, Moyle PB (1992) Crimes against biodiversity:
the lasting legacy of fish introductions. Transactions of the
North American Wildlife and Natural Resources Conference 57:
365–372.
Courtenay WR Jr, Taylor JN (1986) Strategies for reducing risks
from introductions of aquatic organisms: a philosophical per-
spective. Fisheries 11: 30–33.
Crossman EJ, Cudmore BC (1999) Summary of North American
fish introductions through the aquarium/horticulture trade.
In: Claudi R, Leach JH (eds) Nonindigenous Freshwater
Organisms: Vectors, Biology, and Impacts, pp. 129–134. Lewis
Publishers, Boca Raton, FL.
Daniels RJR (2006) Introduced fishes: a potential threat to the
native freshwater fishes of peninsular India. Journal of the
Bombay Natural History Society 103: 346–348.
Davenport K (2000) Querying assumptions of risk in the orna-
mental fish trade. In: Rodgers CJ (ed) Risk Analysis in Aquatic
Animal Health, pp. 117–124. Office International des Epizoo-
ties, Paris.
Dawes J (1998) International experiences in ornamental marine
species management. Part I: perspectives. Proceedings of the
Management strategies for the Marine Ornate Species of the
Gulf of California, La Paz, Baja California Sur, Mexico, 18–19
November 1998. Taken from the official website of the Orna-
mental Fish International organisation at http://ornamental--
fish-int.org/.
Del Rio Rodriguez R (2006) Ornamental fish: economic, eco-
logic and health implications of the trade. Jaina Bolet
ın Infor-
mativo 16: 6–27.
Devaraj KV, Subramanya MN, Manissery JK (1977) New record
of the monogenetic trematode Gyrodactylus medius (Kathari-
ner) on cyprinid fishes in Karnataka. Current Research 6: 105.
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd

Dixon B, Contreras B (1991) Isolation of Edwardsiella tarda
(Ewing and McWhorter) from the freshwater tropical petfish
Meytynnis schreitmeuelleri (Ahl) and Trichogaster trichopterus
(Pallas). Journal of Aquariculture and Aquatic Sciences 6: 31–
32.
Dogiel VA, Lutta A (1937) Mortality among spiny sturgeon of
the Aral Sea in 1936. Rybn Khoz 12: 26–27.
Duggan IC, Rixon CAM, MacIsaac HJ (2006) Popularity and
propagule pressure: determinants of introductions and estab-
lishment of aquarium fish. Biological Invasions 8: 377–382.
Dunham RA, Majumdar K, Hallerman E, Mair G, Hulata G, Liu
Z et al. (2001) Review of the status of aquaculture genetics.
In: Subasinghe RP, Bueno P, Phillips MJ, Hough C, McGlad-
dery SE, Arthur JR (eds) Proceedings of the Conference on
Aquaculture in the Third Millennium, Bangkok, 20–25 February
2000, pp. 129–157. NACA, FAO, Bangkok, Rome.
Ek-Huchim JP, Jimenez-Garcia I, Perez-Vega JA, Rodriguez-
Canul R (2012) Non-lethal detection of DNA from Cichlido-
gyrus spp. (Monogenea, Ancyrocephalinae) in the gill mucus
of the Nile tilapia Oreochromis niloticus. Diseases of Aquatic
Organisms 98: 155–162.
Ernst I, Dove AM (1998) Concurrent invaders – four exotic spe-
cies of Monogenea now established on exotic freshwater fishes
in Australia. International Journal for Parasitology 28: 1755–
1764.
Euzet L, Combes C (1998) The selection of habitats among the
monogenea. International Journal for Parasitology 28: 1645–
1652.
Evans BB, Lester RJG (2001) Parasites of ornamental fish
imported into Australia. Bulletin of the European Association
of Fish Pathologists 21: 51–55.
FAO (2005–2012a). Fisheries and Aquaculture topics. Introduc-
tion of species. Topics Fact Sheets. Text by Devin Bartley. In:
FAO Fisheries and Aquaculture Department [online]. Rome.
Updated 27 May 2005. [Cited 2 January 2012]. http://www.
fao.org/fishery/topic/13532/en
FAO (2005–2012b). Fisheries and Aquaculture topics. Introduc-
tion of species. Topics Fact Sheets. Text by Devin Bartley. In:
FAO Fisheries and Aquaculture Department [online]. Rome.
Updated 27 May 2005. [Cited 2 January 2012]. http://www.
fao.org/fishery/topic/13599/en–>
FAO (2005–2012c). Fisheries and Aquaculture topics. Ornamen-
tal fish. Topics Fact Sheets. Text by Devin Bartley. In: FAO
Fisheries and Aquaculture Department [online]. Rome.
Updated 27 May 2005. [Cited 2 January 2012]. http://www.
fao.org/fishery/topic/13611/en
FAO (2010) The State of Fisheries and Aquaculture. Food and
Agriculture Organization (FAO), Rome.
Ferguson HW, Morales JA, Ostland VE (1994) Streptococcosis in
aquarium fish. Diseases of Aquatic Organisms 19: 1–6.
Fevre EM, Bronsvoort BM, Hamilton KA, Cleaveland S (2006)
Animal movements and the spread of infectious diseases.
Trends in Microbiology 14: 125–131.
Fotedar DN, Parveen QN (1987) On two species of trematode
genus Diplozoon Nordmann 1832, from fishes of Kashmir and
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd
Invasive monogenoids via aquarium trade
notes on some species of the genus. Indian Journal of Helmin-
thology 39: 66–76.
Froese R, Pauly D (2012) FishBase. World Wide Web electronic
publication. www.fishbase.org, version (01/2012).
Fuller PL, Nico LG, Williams JD (1999) Non-Indigenous Fishes
Introduced into Inland Waters of the United States. American
Fisheries Society Special Publication, 27, Maryland, USA, 613
pp.
Galli P, Stefani F, Benzoni F, Crosa G, Zullini A (2003) New
records of alien monogeneans from Lepomis gibbosus and Silu-
rus glanis in Italy. Parassitologia 45: 147–149.
Garcia-Berthou E (2007) The characteristics of invasive fishes:
what has been learned so far? Journal of Fish Biology 71: 33–
55.
Gaughan DJ (2001) Disease-translocation across geographic
boundaries must be recognized as a risk even in the absence
of disease identification: the case with Australian Sardinops.
Review of Fishery Biology and Fisheries 11: 113–123.
Gelnar M, Scholz T, Matejusova I, Konecny R (1996) Occur-
rence of Pseudodactylogyrus anguillae (Yin & Sproston, 1948)
and P. bini (Kikuchi, 1929), parasites of eel, Anguilla anguilla
L., in Austria (Monogeneana: Dactylogyridae). Annalen des
Naturhistorischen Museums in Wien Serie B 98: 1–4.
Gertzen E, Familiar O, Leung B (2008) Quantifying invasion
pathways: fish introductions from the aquarium trade.
Canadian Journal of Fisheries and Aquatic Sciences 65: 1265–
1273.
Gibson DI, Timofeeva TA, Gerasev PI (1996) A catalogue of the
nominal species of the monogenean genus Dactylogyrus Die-
sing, 1850 and their host genera. Systematic Parasitology 35:
3–48.
Global Invasive Species Database (2005) Available from: http://
www.issg.org/database/species/search.asp?sts=sss&st=sss&fr=
1&Image1.x=27&Image1.y=9&sn=&rn=india&hci=-1&ei=
162&lang=EN [Accessed 2 November 2011].
Go J, Lancaster M, Deece K, Dhungyel O, Whittington R (2006)
The molecular epidemiology of iridovirus in Murray cod
(Maccullochella peelii peelii) and dwarf gourami (Colisa lalia)
from distant biogeographical regions suggests a link between
trade in ornamental fish and emerging iridoviral diseases.
Molecular and Cellular Probes 20: 212–222.
Goodwin AE (2002) First report of spring viraemia of carp virus
(SVCV) in North America. Journal of Aquatic Animal Health
14: 161–164.
Gozlan RE, St-Hilaire S, Feist SW, Martin P, Kent ML (2005) Bio-
diversity: disease threat to European fish. Nature 435: 1046.
Gozlan RE, Britton JR, Cowx I, Copp GH (2010) Current
knowledge on non-native freshwater fish introductions. Jour-
nal of Fish Biology 76: 751–786.
Grimes DJ, Gruber SH, May EB (1985) Experimental infection
of lemon sharks, Negaprion brevirostris (Poey), with Vibrio
species. Journal of Fish Diseases 8: 173–180.
Harris PD, Soleng A, Bakke TA (1998) Killing of Gyrodactylus
salaries (Platyhelminthes, Monogenea) mediated by host
complement. Parasitology 2: 137–143.
157
A. Tripathi
Harris PD, Shinn AP, Cable J, Bakke TA (2004) Nominal species
of the genus Gyrodactylus von Nordmann 1832 (Monogenea:
Gyrodactylidae), with a list of principal host species. System-
atic Parasitology 59: 1–27.
Hayward CJ, Kim JH, Heo GJ (2001a) Spread of Neoheteroboth-
rium hirame (Monogenea), a serious pest of olive flounder
Paralichthys olivaceus, to Korea. Diseases of Aquatic Organisms
45: 209–213.
Hayward CJ, Iwashita M, Crane JS, Ogawa K (2001b) First
report of the invasive eel pest Pseudodactylogyrus bini in North
America and in wild American eels. Diseases of aquatic organ-
isms 44: 53–60.
Hayward CJ, Iwashita M, Ogawa K, Ernst I (2001c) Global
spread of the eel parasite Gyrodactylus anguillae (Monogenea).
Biological Invasions 3: 417–424.
Hayward CJ, Bott NJ, Naoki I, Iwashita M, Okihiro M, Nowak
BF (2007) Three species of parasites emerging on the gills of
mulloway, Argyrosomus japonicus (Temminck and Schlegel,
1843), cultured in Australia. Aquaculture 265: 27–40.
Hedrick RP (1996) Movements of pathogens with the interna-
tional trade of live fish: problems and solutions. Revue Scien-
tifique et Technique (International Office of Epizootics) 15:
523–531.
Hedrick RP, McDowell TS (1995) Properties of iridoviruses
from ornamental fish. Veterinary Research 26: 423–427.
Hegde A, Teh HC, Lam TJ, Sin YM (2003) Nodavirus infection
in freshwater ornamental fish, guppy, Poicelia reticulate-com-
parative characterization and pathogenicity studies. Archives
of Virology 148: 575–586.
Hendry AP, Bohlin T, Jonsson B, Berg OK (2004) To sea or not
to sea? Anadromy versus non-anadromy in Salmonids. In:
Hendry AP, Stearns SC (eds) Evolution Illuminated: Salmon
and Their Relatives, pp. 93–125. Oxford University Press,
Oxford.
Hoffman GL (1970) Intercontinental and transcontinental dis-
semination and transfaunation of fish parasites with emphasis
on whirling disease (Myxosoma cerebralis). In: Snieszko SF
(ed) Proceedings of the Diseases of Fish and Shellfish, pp. 69–
81. American Fisheries Society Special Publication No. 5.
Humphrey JD (1995) Australian Quaran tine Policies and Prac-
tices for Aquatic Animals and their Products: A Review for the
Scientific Working Party on Aquatic Animal Quarantine.
Bureau of Resource Sciences, Canberra 264 pp.
Humphrey JD, Ashburner LD (1993) Spread of the bacterial fish
pathogen Aeromonas salmonicida after importation of infected
goldfish, Carassius auratus, into Australia. Australian Veteri-
nary Journal 70: 453–454.
Jahn TL, Kuhn LR (1932) The life-history of Epibdella melleni,
Mac&hum, 1927, a monogenetic trematode parasitic on mar-
ine fishes. Biological Bulletin 62: 89–111.
Jhingran AG (1991) Performance of tilapia in Indian waters and
its possible impact on the native ichthyofauna. FAO Fisheries
Report No. 458, 1–281.
Jimenez-Garcia MI, Vidal-Martinez VM, Lopez-Jimenez S
(2001) Monogeneans in introduced and native cichlids in
158
Mexico: evidence for transfer. Journal of Parasitology 87: 907–
909.
Jogunoori W, Kritsky DC, Venkatanarasaiah V (2004) Neotropi-
cal Monogenoidea. 46. Three new species from the gills of
introduced aquarium fishes in India, the proposal of Heteroty-
lus n. g. and Diaphorocleidus n. g., and the reassignment of
some previously described species of Urocleidoides Mizelle &
Price, 1964 (Polyonchoinea: Dactylogyridae). Systematic Para-
sitology 58: 115–124.
Johnsen BO, Jensen AJ (1986) Infestations of Atlantic salmon,
Salmo salar, by Gyrodactylus salaris in Norwegian rivers. Jour-
nal of fish biology 29: 233–241.
Johnsen BO, Jensen AJ (1991) The Gyrodacytlus story in Nor-
way. Aquaculture 98: 289–302.
Jones SR (2001) The occurrence and mechanism of innate
immunity against parasites in fish. Developmental and Com-
parative Immunology 25: 841–852.
Kabata Z (1985) Parasites and Diseases of Fish Cultured in the
Tropics. Taylor and Francis, London.
Kamat AS, Bandekar JRM, Karani S, Jadhav R, Shashidhar A,
Kakatkar S et al. (2002) Microbiological quality of some
major fishery products exported from India. Proceedings of a
Final Research Coordination Meeting on Determination of
Human Pathogen Profiles in Food by Quality Assured Micro-
bial Assays, Mexico City, 22–26 July, 2002, pp. 51–61. Inter-
national Atomic Energy Agency, Vienna.
Kearn GC (1994) Evolutionary expansion of the Monogenea.
International Journal for Parasitology 24: 1227–1271.
Kennedy CR (1975a) The distribution of crustacean fish para-
sites in Britain in relation to the introduction and movement
of freshwater fish. Journal of the Institute of Fishery Manage-
ment 6: 36–41.
Kennedy CR (1975b) Ecological Animal Parasitology. Blackwell
Scientific Publications, Oxford.
Kennedy CR (1994) The ecology of introductions. In: Pike AW,
Lewis JW (eds) Parasitic Diseases of Fish, pp. 189–208. Samara
Publishers, Tresaith, Wales.
Khan RA, Kiceniuk JW (1988) Effect of petroleum aromatic
hydrocarbons on monogeneids parasitizing Atlantic cod,
Gadus morhua L. Bulletin of Environmental Contamination
and Toxicology 41: 94–100.
Kharat SS, Dahanukar N, Raut R, Mahabaleshwarkar M (2003)
Long term changes in the freshwater fish fauna in the North-
ern Western Ghats, Pune. Current Science 84: 816–820.
Kim JH, Hayward CJ, Joh SJ, Heo GJ (2002) Parasitic infections
in live freshwater tropical fishes imported to Korea. Diseases
of Aquatic Organisms 52: 169–173.
King TA, Cable J (2007) Experimental infections of the monoge-
nean Gyrodactylus turnbulli indicate that it is not a strict spe-
cialist. International Journal for Parasitology 37: 663–672.
Knight JDM (2010) Invasive ornamental fish: a potential threat
to aquatic biodiversity in peninsular India. Journal of Threa-
tened Taxa 2: 700–704.
Knight JD, Rema Devi K (2009) On a record of Amphilophus
trimaculatum (G€ unther) (Teleostei: Perciformes: Cichlidae) in
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd

the natural waters of Tamil Nadu, India. Journal of Bombay
Natural History Society 106: 143–144.
Kolar CS, Lodge DM (2001) Progress in invasion biology:
predicting invaders. Trends in Ecology and Evolution 16:
199–204.
Koskivaara M, Valtonen ET, Vuori KM (1992) Microhabitat dis-
tribution and coexistence of Dactylogyrus species (Monogene-
a) on the gills of roach. Parasitology 104: 273–281.
Krishnakumar K, Rajeev R, Prasad G, Bijukumar A, Mani S,
Pereira B et al. (2009) When pets become pests-exotic aquar-
ium fishes and biological invasions in Kerala, India. Current
Science 97: 474–476.
Kritsky DC (2007) Heteropriapulus nom. nov. (Monogenoidea:
Dactylogyridae) for Heterotylus Jogunoori, Kritsky & Venka-
tanarasaiah, 2004, a junior homonym of Heterotylus Kirsch in
Reitter, 1913 (Coleoptera: Curculionidae). Systematic Parasi-
tology 68: 233.
Kritsky DC, Heckmann R (2002) Species of Dactylogyrus
(Monogenoidea: Dactylogyridae) and Trichodina mutabilis
(Ciliata) Infesting Koi Carp, Cyprinus carpio, during mass
mortality at a Commercial Rearing Facility in Utah, U.S.A.
Comparative Parasitology 69: 217–218.
Kritsky DC, Vidal-Martinez VM, Rodriguez-Canul R (1994)
Neotropical Monogenoidea. 19 Dactylogyridae of Chichlidae
(Perciformes) from the Yucatan Peninsula, with descriptions
of three new species of Sciadecleithrum Kritsky, Thatcher, and
Boeger, 1989. Journal of Helminthology 61: 26–33.
Lakra WS, Singh AK, Ayyappan S (2008) Fish Introduction in
India: Status, Potential and Challenges. Narendra Publishers,
New Delhi.
Landau M (1992) Introduction on Aquaculture. Wiley, New
York.
Leprieur F, Brosse S, Garcia-Berthou E, Oberdorff T, Olden JD,
Townsend CR (2009) Scientific uncertainty and the assess-
ment of risks posed by non-native freshwater fishes. Fish and
Fisheries 10: 88–97.
Lester RJG (1972) Attachment of Gyrodactylus to Gasterosteus
and host response. Journal of Parasitology 58: 717–722.
Levine JM, D’Antonio CM (2003) Forecasting biological inva-
sions with increasing international trade. Conservation Biology
17: 322–326.
Lilley JH, Roberts RJ (1997) Pathogenicity and culture studies
comparing the Aphanomyces involved in epizootic ulcerative
syndrome (EUS) with other similar fungi. Journal of Fish
Diseases 20: 135–144.
Lilley JH, Callinan RB, Khan MH (2002) Social, economic and
biodiversity impact of epizootic ulcerative syndrome (EUS).
In: Arthur JR, Phillips MJ, Subasinghe RP, Reantaso MB,
MacRae IH (eds) Primary Aquatic Animal Health Care in
Rural, Small-Scale, Aquacultural Development, pp. 127–139.
FAO Fisheries Technical Paper No. 406. FAO, Rome.
Lim SLH (1996) Silurodiscoides Gussev, 1961 (Monogenea:
Ancyrocephalidae) from Pangasius sutchi Fowler, 1931 (Pang-
asiidae) cultured in Peninsular Malaysia. Raffles Bulletin of
Zoology 38: 55–63.
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd
Invasive monogenoids via aquarium trade
Lockwood JL, Cassey P, Blackburn T (2005) The role of propa-
gule pressure in explaining species invasions. Trends in Ecol-
ogy and Evolution 20: 223–228.
Lunn KE, Moreau MA (2004) Unmonitored trade in marine
ornamental fishes: the case of Indonesia’s Banggai cardinal
fish (Pterapogon kauderni). Coral Reefs 23: 344–351.
Marchetti MP, Moyle PB, Levine R (2004) Alien fishes in Cali-
fornia watersheds: Characteristics of successful and failed
invaders. Ecological Applications 14: 587–596.
Marcogliese DJ, Ball M, Lankester MW (2001) Potential impacts
of clearcutting on parasites of minnows in small boreal lakes.
Folia Parasitologica 48: 269–274.
Martins ML, Romero NG (1996) Efectos del parasitismo
sobre el tejido branquial en peces cultivados: estudioparasi-
tologico e histopatologico. Revista brasileira de Zoologia 13:
489–500.
Miyazaki T, Egusa S (1972) Studies on mycotic granulomatosis
in freshwater fish. I: mycotic granulomatosis in goldfish. Fish
Pathology 7: 15–25.
Mizelle J, Kritsky DC (1969) Studies on monogenetic trema-
todes. XXXIX. Exotic species of monopisthocotylea with the
proposal of Archidiplectallum gen. n. and Longihaptor gen. n.
American Midland Naturalist 81: 370–386.
Molnar K (1994) Effect of decreased water oxygen content on
common carp fry with Dactylogyrus vastator (Monogenea)
infection of varying severity. Diseases of Aquatic Organisms 20:
153–157.
Moravec F, Wolter J, Korting W (1999) Some nematodes and
acanthocephalans from exotic ornamental freshwater fishes
imported into Germany. Folia Parasitologica 46: 296–310.
Mouton A, Basson L, Impson D (2001) Health status of orna-
mental freshwater fishes imported to South Africa: a pilot
study. Aquarium Sciences and Conservation 3: 327–333.
Mozhdeganlou Z, Ebrahimzadeh Mousavi H, Shayan P, Soltani
M, Ebrahimzadeh E, Rostami M (2011) Detection of single
Dactylogyrus spp. in DNA extracted from infected gill tissue of
fishes using polymerase chain reaction. International Journal
of Veterinary Research 5: 77–80.
Mushiake K (2001) Epizootiology of Neoheterobothrium infec-
tion in wild Japanese flounder. Fish Pathology 36: 252–253.
NABARD (2007) Available from URL: http://www.nabard.org/
modelbankprojects/fish_ornamental_fish.asp (accessed November
2, 2010).
Nigrelli RF (1937) Further studies on the susceptibility and
acquired immunity of marine fishes to Epibdetla melleni, a
monogenetic trematode. Zoologica-New York 22: 185–191.
Noble ER, Noble GA, Schad GA, MacInnes AJ (1989) Parasitol-
ogy: The Biology of Animal Parasites. Lea and Febiger, Phila-
delphia, PA.
Ogawa K, Bondad-Reantaso M, Fukudome M, Wakabayashi H
(1995) Neobenedenia girellae (Hargis, 1955) Yamaguti, 1963
(Monogenea: Capsalidae) from cultured marine fish of Japan.
Journal of Parasitology 81: 223–227.
Padilla DK, Williams SL (2004) Beyond ballast water: aquarium
and ornamental trades as sources of invasive species in aquatic
159
A. Tripathi
ecosystem. Frontiers in Ecology and the Environment 2: 131–
138.
Paperna I (1996) Parasites, infections and diseases of fishes in
Africa – an update. Central Institute of Freshwater Aquacul-
ture Technical Paper of FAO, Rome, No. 31: 220.
Pimentel D (2002) Biological Invasions: Economic and Environ-
mental Costs of Alien Plant, Animal and Microbe Species. CRC
press, London.
Poulin R (2002) The evolution of monogenean diversity. Inter-
national Journal for Parasitology 32: 245–254.
Raghavan R, Prasad G, Anvar-Ali PH, Pereira B (2008a) Exotic
fish species in a global biodiversity hotspot: observations from
River Chalakudy, part of Western Ghats, Kerala, India. Biolog-
ical Invasions 10: 37–40.
Raghavan R, Prasad G, Anvar-Ali PH, Pereira B (2008b) Fish
fauna of Chalakudy River, part of Western Ghats biodiversity
hotspot, Kerala, India: patterns of distribution, threats and
conservation needs. Biodiversity Conservation 17: 3119–3131.
Rao KL (1975) India’s Water Wealth: Its Assessment, Uses and
Projections. Orient Longman, New Delhi.
Rastogi P, Mishra D, Rastogi R, Sharma V, Singh HS (2008) On
a New Species of Genus Silurodescoides (Achmerow, 1964)
Gussev, 1973 with Redescription, Copulation Biology and
Neuroanatomy of S. vistulensis (New Combination) from
Meerut (U.P.), India. Asian Journal of Experimental Sciences
22: 329–342.
Rehulka J (1988) Dactylogyrus ostraviensis n.sp. (Dactylogytri-
dae: Monogenea) from the gills of Barbus conchonius. System-
atic Parasitology 12: 77–80.
Rehulkova E, Gelnar M (2006) Three new species of Dactylogyrus
Diesing, 1850 (Monogenea: Dactylogyridae) from the gills of
the bala sharkminnow Balantiocheilos melanopterus (Cyprini-
dae) from Thailand. Systematic Parasitology 64: 215–223.
Rema Devi K (1987) A golden variation. Black Buck 3: 22–24.
Rixon CAM, Duggan IC, Bergeron NMN, Ricciardi A, MacIsaac
HJ (2005) Invasion risks posed by the aquarium trade and live
fish markets on the Laurentian Great Lakes. Biodiversity and
Conservation 14: 1365–1381.
Robertson PAW, Austin B (1994) Disease associated with cypri-
nids imported into the United Kingdom. International Sym-
posium on Aquatic Animal Health. University of California,
Davis, CA.
Roelofs A (2008) Ecological Risk Assessment of the Queensland
Marine Aquarium Fish Fishery. Department of Primary Indus-
tries and Fisheries, Brisbane, Qld.
Sala OE, Chapin FS, Armesto JJ, Berlow E, Bloomfield J, Dirzo R
et al. (2000) Global biodiversity scenarios for the year 2100.
Science 287: 1770–1774.
Saunders DL, Meeuwig JJ, Vincent CJ (2002) Freshwater pro-
tected areas: strategies for conservation. Conservation Biology
16: 30–41.
Schaperclaus W (1990) Fischkrankheiten, 5th edn. Akademie
Verlag, Berlin.
Scott ME, Anderson RM (1984) The population dynamics of
Gyrodactylus bullatarudis (Monogenea) within laboratory
160
populations of the fish host Poecilia reticulata. Parasitology 89:
159–194.
Secretariat of the Convention on Biological Diversity (2010)
Pets, Aquarium, and Terrarium Species: Best Practices for
Addressing Risks to Biodiversity. Secretariat of the Convention
on Biological Diversity, Montreal, QC.
Sehgal KL (1989) Present status of exotic coldwater fish species
in India. In: Joseph MM (ed.) Proceedings of the Exotic Aquatic
Species in India, pp. 41–47. Asian Fisheries Society (Indian
Branch, Mangalore) Special Publication No. 1.
Shamsi S, Jalali B, Aghazadeh MM (2009) Infection with Dacty-
logyrus spp. among introduced cyprinid fishes and their geo-
graphical distribution in Iran. Iranian Journal of Veterinary
Research 10: 70–74.
Shannon MW, Smith LD, Carlton JT, Pederson J (2005) Assess-
ing the risk of introducing exotic species via the live marine
species trade. Conservation Biology 19: 213–223.
Shea K, Chesson P (2002) Community ecology theory as a
framework for biological invasions. Trends in Ecology and
Evolution 17: 170–176.
Shetty HPC, Nandeesha MC, Jhingran A (1989) Impact of exotic
aquatic species in Indian waters. In: DeSilva SS (ed.) Proceed-
ings of Introduction of Exotic Aquatic Organisms in Asia, Dar-
win, Australia, 1988, pp. 45–55. Asian Fisheries Society Special
publication 3.
Silas EG, Gopalakrishnan A, Ramachandran A, Anna Mercy
TV, Kripan Sarkar, Pushpangadan KR et al. (2011) Guide-
lines for Green Certification of Freshwater Ornamental Fish.
The Marine Products Export Development Authority,
Kochi, India. xii + 106 p. Available at: http://www.mpeda.
com/tender/green.pdf
Singh AK (2004) Aquaculture diversification and species
enhancement: problems and perspectives. In: Verma A (ed.)
Zoology and Human Welfare, pp. 252–261. S.P. Mukherji Gov-
ernment College, University of Allahabad, Allahabad, India.
Singh AK, Das P (2006) Status of common carp (Cyprinus car-
pio-Linnaeus, 1758) in aquaculture and its environmental
impact. Aquaculture 7: 245–257.
Singh AK, Lakra WS (2006) Impact of alien fish species in India:
emerging scenario. Journal of Ecophysiology and Occupational
Health 6: 165–174.
Singh AK, Lakra WS (2011) Risk and benefit assessment of alien
fish species of the aquaculture and aquarium trade into India.
Reviews in Aquaculture 3: 3–18.
Singh AK, Mishra A (2001) Natural breeding of Thai magur in a
village pond-a case report. Aquaculture 2: 175–179.
Singh HS, Rastogi P, Rastogi R (2003) On a new species of Pellu-
cidhaptor Price and Mizelle, 1964 (Monogenea: Dactylogyri-
dae) with a note on its zoogeographical distribution. Uttar
Pradesh Journal of Zoology 23: 7–13.
Skinner RH (1982) The interrelation of water quality, gill para-
sites, and gill pathology of some fishes from south Biscayne
Bay, Florida. Fishery Bulletin 80: 269–280.
Sreenivasan A (1996) Why exotic species. Fishing Chimes 16: 9–
10.
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd

Sterud E, Jorgensen A (2006) Pumpkinseed Lepomis gibbosus
(Linnaeus, 1758) (Centrarchidae) and associated parasites
introduced to Norway. Aquatic Invasions 1: 278–280.
Stunkard HW (1924) A new trematode, Oculotrema hippopotami
n. g., n. sp. from the eye of the hippopotamus. Parasitology
16: 436–440.
Sundaresan J (2011) Ornamental fish industry. Indian Journal of
Marine Sciences 40: 9–10.
Thilakaratne ID, Rajapaksha G, Hewakopara A, Rajapakse RP,
Faizal AC (2003) Parasitic infections in freshwater ornamental
fish in Sri Lanka. Diseases of Aquatic Organisms 54: 157–162.
Thomas VG, Vasarhelyi C, Niimi AJ (2009) Legislation and the
capacity for rapid-response management of nonindigenous
species of fish in contiguous waters of Canada and the USA.
Aquatic Conservation: Marine and Freshwater Ecosystems 19:
354–364.
Thoney DA, Hargis WHJ (1991) Monogenea (Platyhelminthes)
as hazards for fish in confinement. Annual Review of Fish
Diseases 1: 133–153.
Torchin ME, Lafferty KD, Dobson AP, McKenzie VJ (2003)
Introduced species and their missing parasites. Nature 421:
628–630.
Tripathi YR (1957) Monogenetic trematodes from fishes of
India. Indian Journal of Helminthology 9: 1–149.
Tripathi A, Agrawal N, Pandey KC (2010) Monogenoidea on
Exotic Indian Freshwater Fishes. 1. A new geographical record
of Sciadicleithrum iphthimum Kritsky, Thatcher & Boeger,
1989 (Dactylogyridae: Ancyrocephalinae) with the first
description of its egg. Comparative Parasitology 77: 83–86.
Tsutsumi N (2004) The origin and spreads of the monogenean
parasite Neoheterobothrium hirame infecting wild olive floun-
der. PhD Thesis, University of Tokyo, Tokyo.
Reviews in Aquaculture (2014) 6, 147–161
© 2013 Wiley Publishing Asia Pty Ltd
Invasive monogenoids via aquarium trade
Vijayan KK, Rao GS (2009) Invasive Alien Species: animals
including fishes and their pests. Souvenir: International Day
for Biological Diversity, Ministry of Environment and Forests.
Government of India, New Delhi, pp. 26–38. Available from
URL: http://eprints.cmfri.org.in/7145/1/vijayan_souvenir.pdf.
Walton A (1994) Marketability of a cyanide detection kit for use
with the ornamental marine fish trade. Department of Biolog-
ical Science Report submitted to the Industrial Liaisons
Office, Simon Fraser University, pp. 112.
Whittington ID (1998) Diversity ‘down under’: monogeneans in
the Antipodes (Australia) with a prediction of monogenean
biodiversity world-wide. International Journal for Parasitology
28: 1481–1493.
Whittington RJ, Chong R (2007) Global trade in ornamental fish
from an Australian perspective: the case for revised import
risk analysis and management strategies. Preventive Veterinary
Medicine 81: 92–116.
Whittington ID, Cribb BW, Hamwood TE, Halliday J (2000)
Host-specificity of monogenean (platyhelminth) parasites: a
role for anterior adhesive areas? International Journal for Para-
sitology 30: 305–320.
Williams H, Jones A (1994) Parasitic Worms of Fish. Taylor and
Francis, London.
Williamson M (1996) Biological Invasions, Population and Com-
munity Biology Series. Vol. 15. Chapman and Hall, London.
Yamaguti S (1963) Systema Helminthum. IV. Monogenea and
Aspidocotylea. Interscience Publication, New York.
Yamamoto K, Takagi S, Matsuoka M (1984) Mass mortality of
the Japanese anchovy (Engraulis japonica) caused by a gill
monogenean Pseudanthocotyloides sp. (Mazocraeidae) in the
Sea of Iyo (‘Iyo-Nada’), Eime Prefecture. Fish Pathology 9:
119–123.
161