Aquaculture Magazine November/December 2003 1

OTHER ECTOPARASITES INFESTING FISH;

COPEPODS, BRANCHIURANS, ISOPODS,
MITES AND BIVALVES
RICHARD HECKMANN
I
remember one of my first fishing trips to Yellowstone
Lake, Yellowstone National Park seeing white
threadlike organisms attached to the cutthroat trout
we had caught. My fishing friend told me the fish
had lice. I thought lice existed on land creatures
not fish. The fish did not appear to be damaged by
the ectoparasite. Three years ago, a research colleague
brought some small cyprinid fishes back from Brazil that
had isopods attached. In both cases the parasites were visible
without magnification. Copepods, isopods and acarine mites,
ectoparasites of fish, belong to the Phylum Arthropoda, the
jointed appendage group with a chitinous exoskeleton. The
other major group included in this article are the bivalves,
Phylum Mollusca.
The Phylum Arthropoda contains most of the known
animals in the world and many are very abundant as
individuals. They are highly adapted to their habitat and
many are successful as symbiotic organisms living at the
expense of their hosts.
There are three parasitic groups for fish in the Class
Crustacea part of the Phylum Arthropoda: the Branchiura
(fish lice), parasitic copepods (Entomostraca), and parasitic
isopods. Males of the parasitic Entomostraca disappear after
copulation in the preadult stages, so identification is usually
based on females only. Both sexes of the Branchiura and
Isopoda are parasitic and have been described. The copepods
in general are small to microscopic with both free living and
parasitic forms. There are numerous examples of
ectoparasites on fish for the true copepods and Branchiura.
Certain copepod species of the genera Argulus, Lernaea,
Ergasilus and salmon lice have been considered serious pests
for fish culture and have become increasingly important in
recent years. Control of dangerous crustaceans has been
difficult. One unique observation is that parasitic copepods
are not found in acidic seepage waters, and Ergasilus sieboldi
is highly sensitive to hydrogen sulfide.
With the exception of salmon lice, parasitic crustaceans,
when present in small numbers, usually cause only minor
harm to their hosts. In the case of heavy infections severe
damage to skin, muscle, and gill tissues can occur. Secondary
infections (other parasites, bacteria, fungi, and viruses),
anemia, emaciation, mortality, and sometimes severe
epizootics have been reported. Parasitic crustaceans
sometimes undergo marked modifications in external and
internal morphology, including partial or complete fusion
of body segments, reduction and change of function of
appendages, and reduction or disappearance of body cavity
organs. Most of these changes enhance the parasitic mode
of life. Sensory organs and the reproductive system are
usually well-developed. Sexes are usually separate for this
parasitic group (dioecious), and sexual dimorphism often is
presentespecially dwarfism among males.
For the bivalve parasites, the mollusks have soft
unsegmented bodies with a larval stage (glochidia) that can
infest fish. Before the use of plastic for buttons, freshwater
bivalves represented the main source for this product. The
Upper Mississippi was the main area for this industry until
overfishing depleted the number of hosts (fish) for the
glochidia. The glochidia attach to the gill filaments of the
host and depend on the fish for their nutrient needs. Many
of the freshwater bivalves require fish to complete their life
cycle. The larvae became encysted in the epithelium of the
gill filament of the host and when initially observed resemble
metacercariae cysts of trematode.
Examples of these ectoparasites will be listed and
summarized starting with the copepods.
Salmincola is considered a copepod or fish louse
belonging to the family Lernacopodidae. This copepod is
highly modified for its parasitic mode of life and is a major
problem to salmonids in certain areas of North America.
Recently the author received specimens of Salmincola
infecting rainbow trout (Oncorhynchus clarki) in Colorado,
which have had a devastating effect on the fish. Records
show that during the past 40 years this ectoparasite native to
the western United States has spread via fish shipments to
as far east as New Jersey. It has become a major problem in
commercial salmon operations. It has a typical life cycle
with the characteristic nauplius and copepodid stages.
Salmincola attaches to the gill filaments, opercula, mouth
cavity and the tongue. Like other ergasilids, Salmincola
feeds on blood and epithelium. Heavy infections may result
in respiratory problems due to parasite location and,
secondary infections, similar to what happens for the
copepod, Ergasilus.
Aquaculture Magazine November/December 2003 2
For Salmincola there are at least 17 species described to
date. For the female, the cephalothorax is short, stout, and
inclining to the body axis. The trunk is short and stout, often
flattened dorsoventrally.
An example of one of the 17 species is Salmincola
edwardsii which has been reported from the gill filaments,
gill arches and fins of brook trout (Salvelinus fontinalis) and
other salmonids. Reports indicate that the parasite can cause
disease and losses in all stages of cultured fish. Severe
infestation brings about blood loss and the proliferation of
branchial mucous membrane, whereby respiration is
obstructed. The fish become sluggish and discolored. The
reproduction of this parasite takes place primarily during
the hot summer season. Older fish suffer the most severe
attack. The intensity of infestation has been found to be not
less than 500 per fish.
The measures recommended for the prevention of
copepod infestations including Salmincola which have had
some success include:
a) separating fish of different age groups;
b) arranging intense sources of electric light on the pond
and capturing the larval parasites attracted by positive
phototaxis;
c) biological control by introducing cleaning-fish such
as the genus Notropis which consume the larvae of the
copepods through a process of cleaning symbiosis.
It is well known that the salmon louse, Lepeophtheirus
salmonis, is parasitic on marine salmon, Salmo salar,
Oncorhynchus and sea trout, Oncorhynchus trutta. With
the increase in farming both atlantic (Salmo) and pacific
salmon (Oncorhynchus) there is renewed interest in
alleviating the infestation by lice. Salmon lice occur on wild
marine and farmed salmon and have become a serious
problem to the salmon industry. There are other genera and
species of fish lice that can infest the organism. It was
common knowledge in the northern European countries that
the lice will leave the fish in fresh water and survive only a
few days in brackish or fresh water. Salmon are also host to
another caligid, Caligus elongatus, a copepod it shares with
many other teleosts such as tilapia.
From a modest start at the end of the 1960s, the marine
salmon aquaculture program has grown immensely in several
countries such as Chile, Norway, England and Canada. The
major country producing salmon is Norway. Scotland,
Iceland and the Faroes also have many salmon farms. More
and more coastal countries with suitable water have become
interested in salmon production.
Lepeophtheirus salmonis has become a serious pest in
marine aquaculture. Feeding on the fishs skin, mucus and
blood, these lice can cause small haemorrhages and sores,
and may erode the skin and expose the underlying tissue to
secondary infection. It has been reported that 400-500 lice
on a single cultured salmon is not exceptional, and that close
to 2000 lice have been recorded from a single host. Lice on
wild salmon can be the source of the infestation in farmed
marine salmon.
Sea lice such as Lepeophtheirus salmonis and Caligus
elongatus have the potential to damage, weaken and kill
salmon. These ectoparasites may transmit microbial
pathogens, and farm infestations may have impact upon wild
salmonids. Infestations between wild and farmed fish shows
a usually higher prevalence and abundance for the latter,
indicating enhanced transmission of sea lice in farm
conditions. Farmed or cultured salmon are usually isolated
and packed in smaller areas, which enhances parasite
transmission.
Aquaculture Magazine November/December 2003 3
For management and control of the parasites, the choice
of a control method depends upon its efficacy, stress to the
fish, environmental effects, cost, hazard to staff, marketing
implications and ease of application. Each method of
control will vary in adaptability to different farm conditions.
Alternative methods need to be planed to enable control
under different farming conditions and prevent development
of resistance by lice, if pharmaceuticals are used.
Adult females of L. salmonis infesting wild salmon are
mainly attached at the sides of, and behind, the dorsal and
adipose fins where, being dark brown, they do not show
well against the dark color of the fish. Single adult females
may attach anywhere on the fishs body, but few are found
on top of the head, where they seem to aggregate in farmed
salmon. For L. salmonis on cultured salmon, preference is
the summer season, even though the lice can reproduce
during the entire year. Enormous quantities of nauplii must
be produced since these larvae can be a food source for
other animals. The nauplii develop into copepodids.
For those readers primarily interested in the ectoparasitic
crustacea of sea water reared salmon, Johnson, Kent and
Margolis wrote an excellent review article for Aquaculture
Magazine Vol 23 (2). There has been increased interest
pertaining to these parasites with the accelerated farming
of Atlantic and Pacific salmon.
Currently there is considerable interest in the problem
of sea lice control for farm-raised salmon. In an issue of
Fish Farming News (vol. 9. May/June. 2001) a summary
of a conference for pest management for sea lice infesting
trout was held at the University of Aberdeen (England). At
the conference three main sessions were held; (a) the science
base of sea lice control, (b) epidemiology and modeling,
and (c) management systems. In an email to all members
of the Fish Health section, American Fisheries Society a
news release from Canada June 29, 2001 cited a main
problem of sea lice in the Brought Archipelalgo near
Vancouver Island , Canada. It was noted that salmon
farmers watch very closely for sea lice on fish.
CALIGUS
Besides the species that infest salmon, another example
of the copepod Caligus is C. epidemicus, which has been
reported infesting fingerling tilapia. It is a common parasite
of low-salinity water in the coastal and estuarine zones of
the Western Pacific. It has been reported from both wild
and cultured fishes. Recently it was reported on the cultured
tiger shrimp (Penaeus monodon Fabricius) from Chantaburi
Province, Thailand, which is most unusual, not only for this
species but for the entire family of sea lice.
In Taiwan C. epidemicus was first noticed in March 1990
on the Mozambique tilapia (Oreochromis mossambicus)
being reared in the salt-water ponds at the Tainan branch of
the Fisheries Research Institute. The infestation was so
severe that many fish died from the infestation. With the
increased interest in production of tilapia, fish farmers need
to be aware of this ectoparasite.
Ergasilus
Another common ectoparasitic crustacean on fish is the
Argulus, branchiuran parasite of fish. Note highly
modified maxillae for attachment (arrows) and other
parasitic modifications. (View 1)
Argulus, branchiuran parasite of fish. Note highly
modified maxillae for attachment (arrows) and other
parasitic modifications. (View 2)
genus Ergasilus. This organism has evolved into a very
effective parasite modified for its mode of life. The body
attaches to a fish host with prominent clawlike second
antenae. The cephalothorax or body is highly modified and
the body is narrowed posteriorly. Females (1.5 to 2.5 mm)
are larger than males (1.0 mm). The highly visible egg sacs
for the female are long and often cigar-shaped. There are
many described species in the world that infest fish.
Ergasilus attaches to the gill filaments, where it feeds on
blood and epithelium. Heavy infestations can result in
mechanical damage, petechial hemorrhaging, impaired
respiration, epithelial hyperplasia, anemia and retarded
Aquaculture Magazine November/December 2003 4
growth. When the parasite attach, the piscine host becomes
more susceptible to secondary infections. Adult parasites
occasionally infest gill rakers or other external locations.
For the life cycle, which is similar to other copepods,
mature eggs are released from egg sacs. After the eggs are
released, embryonic development takes place and a free-
swimming nauplius hatches from each egg. Larvae pass
through four copepodid stages, each accompanied by
molting. Copulation occurs when an individual is free-
swimming, after which the male dies. The female copepod
enters the gill cavity, where she is retained by gillrakers,
creeps to the gills, and
attaches with highly
modified appendages.
Ergasilosis is of
c o n s i d e r a b l e
importance in fishery
m a n a g e m e n t ,
especially for fish
inhabiting lakes.
Among ergasilids,
Ergasilus sieboldii.
lives as a parasite on the gills of various freshwater fish, and
has become common for fishes in central Europe. Its wide
geographical distribution and the high incidence of its
infestation have caused quite a serious problem for those
managing lake fisheries. The practical consequences of
heavy attacks have been fairly enormous in the fishery
economy of lakes and probably surpass the losses known
from many of the other fish parasites. Both small fish and
large fish may be severly impacted by the copepod.
A common observation for the severity of attack on
different freshwater fish, by adult and copulated females,
can be attributed to the differences in the species-
characteristic movements (speed of swimming) and shape
of fishes. More swift movements and greater swimming
activity do not favor the attachment of parasites.
Lernaea Anchor worm, Thread worm
The freshwater fish tanks at my university annually offer
examples of fish infested with this copepod. Characteristics
of this Lernaeidid copepod include the long filamentous
nature of the female, which averages 5-22mm in length with
trailing egg sacs attached. Modifications include the head
which is a rounded knob inserted into the musculature of its
host with one or two pair of anchors to hold it in position. It
has a typical life cycle with eggs released by the female
hatching in one to three days with subsequent nauplia larvae.
The nauplius metamorphoses into first or second copepodid
stage in 4 to 16 days. No further development occurs unless
it attaches to a host. Larvae pass through five successive
copepodid stages before attachment; copulation occurs
during the fourth coepodid stage and the male disappears
similar to the Ergasilus species. In the temperate zone, the
female overwinters as a larva or juvenile adult attached to a
fish. The copepodid stages will feed on fish mucus.
Following infestation, damage to the fish host includes;
hemorrhagic and ulcerated lesions, anemia, retardation of
growth, loss of weight, and loss of equilibrium. The potential
for secondary infections by other parasites, including
Saprolegnia, Trichodina Ichthyobodo; and bacteria, and
viruses has been reported. Massive mortalities have occurred
and fish kills due to heavy Lernaea infections have been
reported in lakes and streams. Many species of Lernaea
have been reported in the USA (see references).
Clinical Signs: Lernaea
The first reactions to the irritation from a parasite attack
by Lernaea are aberrant swimming, flashy, scouring or
gliding movements against the bottom or on submerged
objects (similar to
those fish infested
with Costia,
T r i c h o d i n a ,
Ichthophthirius and
other ectoparasites
(See articles on
parasitic protozoa).
The infestation is
accompanied by
marked emaciation
and loss of weight.
This parasitic copepod is readily visible without the use
of magnification. Anchor worm is a general term for species
of highly modified copepods that possess anchor-like
processes for securing themselves to the host. Anchor worms
may be introduced into an aquarium from wild or pond-raised
fish (parasitic reservoir). Goldfish, koi, or wild native fish
are most commonly affected. Lernaea and related genera
infest freshwater fish. Lernaea is most likely to be seen in
F
or management and control of the
parasites, the choice of a control
method depends upon its efficacy,
stress to the fish, environmental effects, cost,
hazard to staff, marketing implications and
ease of application.
Aquaculture Magazine November/December 2003 5
summer, when copepod reproduction is at a peak.
Treatment: Management of Lernaea infestations.
For lernaeid infestations in aquarium fish, individual
parasites can be removed with forceps. Wounds should be
watched closely for secondary infections. Be sure to isolate
infected fish from the other stock. Organophosphate is
usually effective; prolonged immersion treatment should be
repeated every 7 days for 28 days. Remember only a limited
number of pharmaceuticals are approved for aquaculture.
Among the pathogenic crustaceans, members of the family
Argulidae represent some of the most widespread and
dangerous ectoparasites of freshwater and marine fish. These
organisms damage the fish directly by extracting blood and
vital tissue fluids from the host with their modified mouth
parts. Secondarily, they can initiate access for other disease
organisms (virus, bacteria, etc) by breaking down the first
line of defense for the fish, the integument. The genus
Argulus constitutes the greatest number of known species of
branchiuran fish parasites and the widest distribution through
out the world. Not all species of Argulus have been
described.
Species of Argulus are characterized by a depressed,
ovoid and broad body. The head is fused with the thoracic
segment forming a cephalothorax covered by a dorsal, broad,
flat shield (carapace) with posterolateral lobes. Two pair of
maxillae are present, the first maxillae in the genus Argulus
is highly modified to form cup-like suckers with strongly
reinforced walls.
Argulus, commonly called fish-lice, crustacean
ectoparasites of fish, are large (4-20 mm long) ectoparasites,
which move freely over the body (and sometimes gills) of
fishes. Thus they can readily be observed without
magnification. They puncture the hosts skin, inject a
cytolytic toxin through the oral sting, and feed on blood.
Feeding sites often become hemorrhagic and ulcerated, and
provide access to secondary infections by other parasites,
fungi, bacteria, or viruses. Heavy infestations can cause
emaciation, loss of equilibrium, and on occasion high
mortality for both cultured and free-ranging fishes.
Argulus is an ectoparasite from the subclass Branchiura,
Phylum Arthropoda belonging to a taxonomic group separate
from other copepods. They are considered modified
copepods, and are parasites of both freshwater and marine
fish. Argulids have long been recognized as pests of cultured
trout in Europe and carp in China. Recently, they have been
considered a major problem for the sea-caged salmon
industry. Argulus has a world-wide distribution for fishes.
After attaching to fish, the argulids grow and metamorphose
several times and become sexually mature in 30 to 35 days.
Since their first mention in the literature, the biology of
argulids and their pathological effects have attracted the
attention of biologists and fishermen. With the development
of intensive fish culture in many countries, the problem of
controlling this group of dangerous ectoparasites has again
been brought to the attention of researchers.
There are two structures used directly for feeding by these
organisms, the preoral spine and the mouthtube. The preoral
spine punctures the host integument and injects the fish with
a toxic secretion that can cause an inflammatory, hemorrhagic
response. The highly adapted maxillules are the primary
organs for attachment to the host and they look like a toilet
plunger and provide the easiest means of identifying the
genus since they are the most noticeable structures on the
animals ventral surface.
Argulus (fish louse), is uncommon in freshwater aquarium
fish but may occur if wild or pond-raised fish are introduced
into the tank. Argulus is especially common on goldfish
and koi and on many wild freshwater fish.
Fish can display flashing or violent erratic swimming,
similar to other crustacean parasites, or other behavioral
abnormalities because of the irritation caused by the parasite.
Fish are damaged by the repeated piercing of the skin by the
stylet or preoral spine. Also hooks and spines are on the
appendages, which may cause mechanical damage. One or
two parasites usually cause no clinical signs in large fish,
but Argulus has a high reproductive rate thus increasing the
parasite infestation.
Cyprinus carpio (common carp) is a favored host for fish
lice; thus Argulus are sometimes called carp-lice. Argulus
has been reported in stickleback, salmonids, and white
crappie. Where common sites for infection are around the
gills, operculum, mouth, back, and pelvic and pectoral fins.
Treatment
As with other crustacean parasites, organophosphates are
usually an effective treatment. Review regulations relating
to the use of this chemical for fish culture. The time needed
to complete the life cycle varies for Argulus; therefore, it is
Aquaculture Magazine November/December 2003 6
useful to rid tanks of egg contamination by using disinfectant
or by allowing the tanks to dry thoroughly preferably outside
exposed to the sun for several days. Otherwise, multiple
chemical treatment may be needed. Individual parasites can
be removed from fish by using forceps, but this is labor
intensive and does not eliminate parasites in the environment.
Isopods
This is a commonly overlooked group of fish parasites.
There are many examples of highly modified isopods
parasitic to marine fish such as the flabelliferan group. Many
have yet to be recorded and described. Isopods are members
of the Phylum Arthropoda, organisms with jointed
appendages and usually a flattened three-part body. There
has been limited research conducted on this group.
Parasitic isopods, with approximately 450 described
species, are fairly common crustacean ectoparasites of wild
tropical marine fish. They are less common in cold marine
waters and rarely found on freshwater fish. The samples
recently brought to our lab were from freshwater fish in
Brazil. They are rare for cultured fish, although some
infestations have caused problems in sea-caged salmonids
in Australia. The life cycle is simple. Most are parasitic as
both juveniles and adults.
There are two main groups of isopods with parasitic
species; Flabellifera and Gnathiidea. The Flabellifera have
a typical isopod shape and are up to 6 cm in length. Nearly
all the major fish groups are represented as hosts to this
group. For some only the larvae are parasitic living on the
skin or gills of fish. Most of the adults are nonfeeding, live
in mud tubes, and produce infective larvae.
Due to their large size, single isopods, readily seen on
the fish, can cause considerable damage with their biting
and sucking mouthparts including pressure necrosis of gill
tissue limiting respiratory surface and growth retardation.
Heavy infestations of parasitic juveniles have the potential
to kill small fingerlings when they attach. There are no
published studies of treatments for parasitic isopods.
Individuals can also be removed from fish by using forceps.
Isopods are also found on coastal freshwater fishes or on
brackish water fishes inhabiting freshwater. Most are
associated with marine, brackish water, or euryhaline fishes.
Freshwater fish isopods have been imported to North
America on aquarium fishes. Thus, shipments of aquarium
fish with parasites should be isolated from the other stock.
Cymothoids parasitize freshwater fishes in the rivers and
lakes of South America, Africa, and Asia. Many of these
cymothoids are exclusively freshwater parasites. Some
species burrow into the flesh of their hosts. Two examples
of parasitic isopods are Nerocila and Lironeca (Livoneca)
Nerocilia belongs to the family Cymothoidae, cymothoid
parasites. It has a body two to three times longer than wide
and a head with wide anterior margins. Trunk and tail possess
numerous pointed projections. Nerocilia infests a variety
of marine and brackish water fishes and is also found on
coastal freshwater or euryhaline fishes. Species often show
little or no host specificity attaching to any fish in the area.
Most occur on the bodies and fins of their hosts; while one
species infests the gill chamber.
The second representative, Lironeca (Livoneca) is also
part of the family Cymothoidea. It is described by a body
weakly vaulted, neither flat nor highly inflated. The head
not surrounded by shoulders or extensions of the body.
This isopod is limited to the New World, with described
species on the U.S. Atlantic and Pacific coasts, and one in
the Caribbean. Lironeca occurs has been described in the
gill chamber of both marine and brackish water fishes.
There has been additional interest in describing and
studying isopods from marine fish in Pakistan. In the
December issue (1999) of Proceedings of Parasitology an
article by Shireen described two new species of isopods,
Cymothoa eremita and Joryma engraulidas from marine
fishes off the Karachi coast in Pakistan. This has been a
pattern for other issues of the Proceedings of Parasitology
describing new species of this parasite.
There are many parts of the world where isopods are not
that well known. Considerable research has been done in
India due to the interest in culturing fish. The importance of
parasitic isopods has become critical due to the interest in
prawn and fishes as a food source. Samples of isopods
attached to the gills of freshwater fishes from Brazil were
brought to my laboratory for identification.
Acarina: Mites of the Phylum Arthropoda
Very little is known about mite infestation in fish. There
have been a few records listed in journals. The habitat and
behavior of fishes are not conducive to mite infestation.
Recently samples of mites were brought to our lab infesting
mosquitoes not fish. It is the larval stage of the acarine mites
that infests fish and other animals. The larvae are usually
encapsulated with a layer of collagenous connective tissue.
Most researches consider mites to be accidental parasites.
Very few records exist for infestations of North American
fishes.
Several genera of mites have been reported from the skin,
gills and esophagus of fishes in Europe and North America.
Fishes can become heavily infested and damaged. In North
America the mites observed in fishes have not been classified.
They have been observed in the esophagus, gills, intestine
and air bladder.
The six-legged larvae of several genera of mites have
also been reported from the skin, gills, and esophagus of
fishes in Europe. As indicated above, the larvae are usually
encapsulated. Even though most authors consider them
accidental parasites, if they are present in large numbers in
the water the fishes can become heavily infected and
damaged.
Phylum Mollusca
Glochidia, the larval stage of freshwater bivalves, can be
a major problem for fishes occupying the same habitat as
the molluscan parasite. The symbiotic relationship was
observed many years ago (1921) and before the use of
plastics, the glochidia became a major problem for the button
industry. The larvae, with exceptions, of most of the
freshwater bivalves (clams) must go through a parasitic stage
on the gills or fins of fishes. The invasion of larval bivalves
Aquaculture Magazine November/December 2003 7
has multiple effects on the host; a) masks respiratory surfaces
on the gill, b) opens the host to secondary infection and
invasion, c)gains sustenance from the host, d) decreases
the growth and development of the freshwater clam.
References
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Argulus
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thesis, Brigham Young University, Provo, Utah. 24 pages.
Boxshall, G.A. and D. Defaye. 1993. Pathogens of Wild and Farmed
Fish: Sea Lice. Ellis Horwood Limited Publisher. England. 378
pages.
Bristow, G..A. & Berland, B. 1991. A report on some metazoan parasites
of wild marine salmon (Salmo salar L.) from the west
coast of Norway with comments on their interactions with
farmed salmon. Aquaculture 98: 311-318.
Buttner, J.K. 1980. Occurrence of Argulus (Crustacea: Branchiura) on
the white crappie (Pomoxis annularis) and free
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the Illinois State Academy of Science. 72:5-7.
Gardner, J. and R.A. Heckmann. 1999. New techniques for observing
whole amounts of parasites by SEM. The American
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Gresty, K.A., G.A. Boxshall, and K. Nagasawa. 1993. The fine structure
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Heckmann, R.A. 1996. Protozoan parasites of fish, Parts I & II.
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Hoffman, G.L. 1999. Parasites of North American Freshwater Fishes.
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Kabata, Z. 1970. Diseases of Fishes. I. Crustacea as Enemies of Fishes.
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Kabata, Z. 1974. Mouth and mode of feeding of Caligidae (Copepoda)
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About The Author
Richard D. Heckmann Ph.D.
Department of Zoology
Brighan Young University