''Lernaeocera branchialis'', sometimes called cod worm, is a

parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...

of marine fish, found mainly in the

North Atlantic The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the "Old World" of Africa, Europe and ...

. It is a marine

copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...

which starts life as a small

pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or wa ...

crustacean larva. It is among the largest of copepods, ranging in size from when it matures as a copepodid larva to more than as a sessile adult. ''Lernaeocera branchialis'' is

ectoparasitic Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has c ...

, which means it is a parasite that lives primarily on the surface of its hosts. It has many life stages, some of which are

motile Motility is the ability of an organism to move independently, using metabolic energy. Definitions Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...

and some of which are

sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...

. It goes through two parasitic stages, one where it parasitizes as a secondary host a

flounder Flounders are a group of flatfish species. They are demersal fish, found at the bottom of oceans around the world; some species will also enter estuaries. Taxonomy The name "flounder" is used for several only distantly related species, thou ...

lumpsucker The Cyclopteridae are a family of marine fishes, commonly known as lumpsuckers or lumpfish, in the order Scorpaeniformes. They are found in the cold waters of the Arctic, North Atlantic, and North Pacific oceans. The greatest number of species ar ...

, and another stage where it parasitizes as a primary host a

cod Cod is the common name for the demersal fish genus '' Gadus'', belonging to the family Gadidae. Cod is also used as part of the common name for a number of other fish species, and one species that belongs to genus ''Gadus'' is commonly not call ...

or other fishes of the cod family (

gadoid The Gadidae are a family of marine fish, included in the order Gadiformes, known as the cods, codfishes, or true cods. It contains several commercially important fishes, including the cod, haddock, whiting, and pollock. Most gadid species a ...

s). It is a

pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...

that negatively impacts the

commercial fishing Commercial fishing is the activity of catching fish and other seafood for commercial profit, mostly from wild fisheries. It provides a large quantity of food to many countries around the world, but those who practice it as an industry must often ...

and

mariculture Mariculture or marine farming is a specialized branch of aquaculture (which includes freshwater aquaculture) involving the cultivation of marine organisms for food and other animal products, in enclosed sections of the open ocean ( offshore mari ...

of cod-like fish.

Life stages

The life-cycle of a cod worm involves a complex progression of life stages, including two successive hosts. It comprises "two free-swimming nauplius stages, one infective copepodid stage, four

chalimus Crustaceans may pass through a number of larval and immature stages between hatching from their eggs and reaching their adult form. Each of the stages is separated by a moult, in which the hard exoskeleton is shed to allow the animal to grow. The ...

stages and the adult copepod, each separated by a

moult In biology, moulting (British English), or molting (American English), also known as sloughing, shedding, or in many invertebrates, ecdysis, is the manner in which an animal routinely casts off a part of its body (often, but not always, an outer ...

". The cycle begins with the females laying eggs which hatch into a nauplius, the usual early

larval A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The l ...

stage of crustaceans. This nauplius I moults about 10 minutes after hatching to produce nauplius II, and 48 hours later, nauplius II moults to a copepodid stage. At this point the copepodid is pelagic and free-swimming with an average length of about 0.5 mm. The next stage is finding a secondary or

intermediate host In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include a ...

, a

demersal fish Demersal fish, also known as groundfish, live and feed on or near the bottom of seas or lakes (the demersal zone).Walrond Carl . "Coastal fish - Fish of the open sea floor"Te Ara - the Encyclopedia of New Zealand. Updated 2 March 2009 They occ ...

like a

lumpfish The Cyclopteridae are a family of marine fishes, commonly known as lumpsuckers or lumpfish, in the order Scorpaeniformes. They are found in the cold waters of the Arctic, North Atlantic, and North Pacific oceans. The greatest number of species ar ...

which is often stationary and therefore easy to catch. The copepodid has only a day to find such a fish and attach itself to the fish's gills. When they locate such a fish, they capture it with grasping hooks at the front of their body. They penetrate the fish with a thin filament which they use to suck its blood. The nourished cod worms then progress via four moults from the naupliar stage to the mature chalimus stage. At this point the males transfer sperm to the females. Both sexes develop swimming

seta In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...

e, detach from the flounder or lumpfish and again swim freely as pelagic organisms. The female cod worm still resembles a copepod and is 2 to 3 mm long. Females undergo another pelagic quest, searching this time for a definitive or

primary host In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include a ...

. With her fertilised eggs, she looks for a

or a fish belonging to the same family as cod, such as a

haddock The haddock (''Melanogrammus aeglefinus'') is a saltwater ray-finned fish from the family Gadidae, the true cods. It is the only species in the monotypic genus ''Melanogrammus''. It is found in the North Atlantic Ocean and associated seas where ...

or whiting. When a suitable definite host is located, females enter the

gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...

chamber. There, while attached to a gill, the female develops a plump, sinusoidal, worm-like body, with a coiled mass of egg strings at the posterior end. Females now measure about 20 mm long, but can grow up to 50 mm. The oral end of the female copepod penetrates the body of the cod until it enters the rear bulb of the host's

heart The heart is a muscular organ in most animals. This organ pumps blood through the blood vessels of the circulatory system. The pumped blood carries oxygen and nutrients to the body, while carrying metabolic waste such as carbon dioxide t ...

. There, firmly rooted in the cod's circulatory system, the front part of the parasite develops in the shape of antlers or branches on a tree, reaching into the main

artery An artery (plural arteries) () is a blood vessel in humans and most animals that takes blood away from the heart to one or more parts of the body (tissues, lungs, brain etc.). Most arteries carry oxygenated blood; the two exceptions are the pul ...

. In this way, while safely tucked beneath the cod's gill cover, the female's deeply embedded oral end can feed on blood while eggs develop and are released into the water column from the posterior end.

Behaviour

It is not known how ''L. branchialis'' searches for its fish hosts, but it probably uses

chemoreceptor A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance (endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorecept ...

s and

mechanoreceptor A mechanoreceptor, also called mechanoceptor, is a sensory receptor that responds to mechanical pressure or distortion. Mechanoreceptors are innervated by sensory neurons that convert mechanical pressure into electrical signals that, in animals, ...

s, and follows physical clues in the

water column A water column is a conceptual column of water from the surface of a sea, river or lake to the bottom sediment.Munson, B.H., Axler, R., Hagley C., Host G., Merrick G., Richards C. (2004).Glossary. ''Water on the Web''. University of Minnesota-D ...

such as those provided by

halocline In oceanography, a halocline (from Greek ''hals'', ''halos'' 'salt' and ''klinein'' 'to slope') is a cline, a subtype of chemocline caused by a strong, vertical salinity gradient within a body of water. Because salinity (in concert with temp ...

s and

thermocline A thermocline (also known as the thermal layer or the metalimnion in lakes) is a thin but distinct layer in a large body of fluid (e.g. water, as in an ocean or lake; or air, e.g. an atmosphere) in which temperature changes more drastically with ...

Effects on fisheries

The most serious parasitic crustaceans among fish in general are

sea lice Sea lice (singular: sea louse) are copepods (small crustaceans) of the family Caligidae within the order Siphonostomatoida. They are marine ectoparasites (external parasites) that feed on the mucus, epidermal tissue, and blood of host fish. Th ...

. However, ''L. branchialis'' is probably the most serious parasitic crustacean among cod. Infestation reduces the efficiency with which food can be utilised, delaying the development of the

gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...

s. Up to 30% loss in weight can occur, with increases in mortality because of open lesions with loss of blood, and possibly occlusion of vessels or aorta. These can have commercial impacts on

wild fisheries A wild fishery is a natural body of water with a sizeable free-ranging fish or other aquatic animal (crustaceans and molluscs) population that can be harvested for its commercial value. Wild fisheries can be marine (saltwater) or lacustrine/ri ...

, making cod-like fishes more expensive to market.

Gadoid The Gadidae are a family of marine fish, included in the order Gadiformes, known as the cods, codfishes, or true cods. It contains several commercially important fishes, including the cod, haddock, whiting, and pollock. Most gadid species a ...

s, particularly cod, are emerging marine aquaculture species in some North Atlantic countries. ''L. branchialis'' present potential problems for their successful

.Fisheries Research Services (2005
Final report of the Aquaculture Health Joint Working Group sub-group on disease risks and interactions between farmed salmonids and emerging marine aquaculture species
Page 29. Scotland.

Life stages

Behaviour

Effects on fisheries

References

Further reading