Ergasilus Argulus
Ergasilus Argulus
Ergasilus Argulus
Copepod
Ergasilus spp.
I. Morphology
Ergasilus spp. are primarily parasites of freshwater hosts but are common on several marine
fishes. Ergasilidae show primitive morphological characteristics reminiscent of free-living
copepods, with few but effective adaptations for parasitism. Their antennules are sensory, but
the antennae have become modified into powerful organs of prehension. Ergasilus spp. females
usually are found clinging by their antennae to one of the fish’s gill filaments. Each antenna ends
in a sharp claw. The third segment and claw are opposable with the second (subchelate). Rather
than depending on muscle and heavy sclerotization of antennae, the antennal tips may be fused
or locked so that a gill filament is completely encircled (E. amplectens, E. tenax). When removed
from their position on a gill, most Ergasilus spp. can swim reasonably well; their pereiopods
retain the flat copepod form, with setae and hairs well adapted for swimming. Their first legs,
however, show adaptation for their feeding habit. These appendages are supplied with heavy,
bladelike spines; in some species the second and third endopodal segments are fused,
presumably lending greater rigidity to the leg. Such modifications increase the animal’s ability to
rasp off mucus and tissue from the gill to which it is clinging. The first legs dislodge epithelial and
underlying cells in this manner and sweep them forward to the mouth.
Ergasilus spp. have three naupliar and five copepodid stages, all free-living. Adult males are
planktonic as well, and females are fertilized before attaching to a fish host. Only adult females
have been found as parasites. In one species even females are planktonic as adults (E.
chautauquaensis, which may be the only nonparasitic species in the genus), although females of
several other species are sometimes encountered plankton in. The life cycle of Ergasilus spp. is
very temperature dependent. Parasite reproduction starts in spring (eggs start to hatch at
around 8°C) and continues into late autumn. At relatively low temperatures (12-15°C) it can take
approximately 10 weeks for the parasite to develop from an egg to adult. However, this is
reduced to 22 days in warmer conditions. Infection levels increase throughout the year.
V. Treatment
Chemical treatments cannot be used to remove the parasite and its free-living stages.
The only way to eradicate it would be to de-stock, drain and lime the fishery. This is
often very costly, undesirable and impractical.
References:
Roberts, L. S., Schmidt, G. D., Janovy, J. J., & Nadler, S. (2013). Gerald D. Schmidt & Larry S. Roberts
Foundations of Parasitology. Boston: McGraw-Hill.
Thatcher, V. E. (1984). Ergasilus Pitalicus, New Species (Copepoda: Poecilostomatoida: Ergasilidae), a Gill
Parasite of a Cichlid Fish from the Pacific Coast of Colombia. Journal of Crustacean Biology, 4(3),
495–501. doi: 10.2307/1548045
4
Branchuria
Argulus spp.
Ventral view of Argulus viridis, female. Note suctorial proboscis, modification of maxillules into sucking discs, and
lateral expansion of carapace into alae. Source: Drawing by William Ober and Claire Garrison after M. F. Martin,
“On the morphology and classification of Argulus (Crustacea),” in Proc. Zool. Soc. London 1932:771–806.
I. Morphology
They are dorsoventrally flattened, reminiscent of caligid copepods with which they are
sometimes confused and can adhere closely to the host’s surface. Some species are moderately
large, up to 12 mm or so. Argulus spp. can swim well as adults; females must leave their hosts to
deposit eggs on the substrate. Many Argulus spp. are not host specific and so have been
recorded from many fish species. Their carapace expands laterally to form respiratory alae. The
parasites have two pairs of antennae. Homologies of the remaining head appendages have been
disputed, but best evidence suggests that the only appendages in the suctorial proboscis, or
mouth tube, are mandibles. 50 The large, prominent sucking discs are modified maxillules.
Immediately posterior to the maxillular discs are large maxillae, apparently used to maintain the
animal’s position on its host and to clean other appendages. Argulus spp. have four pairs of
thoracic swimming legs of typically crustacean biramous form. Exopods of the first two pairs
often bear an odd, recurved process, or flabellum, thought by some to indicate affinities with
the subclass Branchiopoda.
While the branchiuran life cycle averages 30 to 60 days, the actual duration depends on the
parasite species and the water temperature. All life stages of both sexes are parasitic. Argulus
spp adults can survive for several days off the fish host. After a male/female pair mates, the
adult female detaches from the fish host to lay eggs onto hard surfaces and vegetation in the
environment. After she releases her eggs, the female returns to the fish host. The time required
for Argulus spp eggs to hatch will vary, depending upon the species and temperature. Argulus
japonicus eggs hatch in 10 days at 35°C but require 61 days at 15°C. At 23°C the eggs of a closely
related species, A. foliaceus, hatch in 17 days, whereas at 20°C they hatch in 30 days. If laid in
the fall, eggs are capable of overwintering (surviving) until the following spring. In many species,
the first stage larvae (known as the “metanauplius”) must find a parasitic host within 2–3 days
of hatching or they will die. Once attached to the fish host, juveniles undergo a series of molts
(11 molts or 12 “stages” in A. foliaceus) until they reach sexual maturity, roughly 30–40 days
after hatching. Juveniles can overwinter within the fish’s mucus.
V. Treatment
There are currently no FDA-approved drugs for the treatment of Argulus spp.
Prolonged immersion of an organophosphate pesticide, such as trichlorfon (Dylox® 80,
Bayer), which acts by disrupting the nervous system, has been an effective treatment
when dosed at 0.25–0.50 mg/L active ingredient, once a week for 4 treatments.
Diflubenzuron (Dimilin®, Chemtura) is another pesticide, which acts by interfering with
the parasite’s molting (“shedding”) of the outer skin (“exoskeleton”) during growth and
development. Diflubenzuron effectively kills both adult and larval stages, but it is a
restricted-use pesticide, and applicators must follow label instructions.
Other compounds with a similar mechanism of action as diflubenzuron, such as
lufenuron (Program®, Novartis Animal Health) at 0.13 mg/L, have also been used with
success.
Potassium permanganate (10 mg/L for 30 minutes, or 1.3 mg/L applied twice over 3
days) has been shown to be effective in killing the juvenile and adult stages of
freshwater species in several studies.
References:
Hoffman, G. L. 1977. Argulus, a branchiuran parasite of freshwater fishes. US Fish and Wildlife Services
Fish Disease Leaflet 49: 1–9.
Hoffman, G. L. 1999. Parasites of North American freshwater fishes. Cornell University Press, Ithaca, NY.
Noga, E. J. 2010. Fish Disease: Diagnosis and Treatment, 2nd ed. Wiley-Blackwell, Ames, Iowa.
Roberts, L. S., Schmidt, G. D., Janovy, J. J., & Nadler, S. (2013). Gerald D. Schmidt & Larry S. Roberts
Foundations of Parasitology. Boston: McGraw-Hill.
Stoskopf, M. K. 1993. Fish medicine. W.B. Saunders Company, Philadelphia, PA.