Lernaeocera branchialis | |
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The gills of a whiting infested by two blood-sucking Lernaeocera branchialis | |
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Subphylum: | Crustacea |
Class: | Copepoda |
Order: | Siphonostomatoida |
Family: | Pennellidae |
Genus: | Lernaeocera |
Species: | L. branchialis |
Binomial name | |
Lernaeocera branchialis | |
Synonyms [1] | |
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Lernaeocera branchialis, sometimes called cod worm, is a parasite of marine fish, found mainly in the North Atlantic. [2] It is a marine copepod which starts life as a small pelagic crustacean larva. It is among the largest of copepods, ranging in size from 2 to 3 millimetres (3⁄32 to 1⁄8 inch) when it matures as a copepodid larva to more than 40 mm (1+1⁄2 in) as a sessile adult.
Lernaeocera branchialis is ectoparasitic, 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 and some of which are sessile. It goes through two parasitic stages, one where it parasitizes as a secondary host a flounder or lumpsucker, and another stage where it parasitizes as a primary host a cod or other fishes of the cod family (gadoids). It is a pathogen that negatively impacts the commercial fishing and mariculture of cod-like fish.
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 stages and the adult copepod, each separated by a moult". [3]
The cycle begins with the females laying eggs which hatch into a nauplius, the usual early larval stage of crustaceans. [4] 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. [3]
The next stage is finding a secondary or intermediate host, a demersal fish like a flounder or lumpfish 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. [4]
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 setae, detach from the flounder or lumpfish and again swim freely as pelagic organisms. [4] [5]
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. With her fertilised eggs, she looks for a cod or a fish belonging to the same family as cod, such as a haddock or whiting. [4]
When a suitable definite host is located, females enter the gill 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. [4] Females now measure about 20 mm long, but can grow up to 50 mm. [6] The oral end of the female copepod penetrates the body of the cod until it enters the rear bulb of the host's heart. 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. 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. [4] [5]
It is not known how L. branchialis searches for its fish hosts, but it probably uses chemoreceptors and mechanoreceptors, and follows physical clues in the water column such as those provided by haloclines and thermoclines. [3]
The most serious parasitic crustaceans among fish in general are sea lice. [7] 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 gonads. 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. [7] These can have commercial impacts on wild fisheries, making cod-like fishes more expensive to market. [7] [8] Gadoids, particularly cod, are emerging marine aquaculture species in some North Atlantic countries. L. branchialis present potential problems for their successful mariculture. [9]
Copepods are a group of small crustaceans found in nearly every freshwater and saltwater habitat. Some species are planktonic, some are benthic, a number of species have parasitic phases, and some continental species may live in limnoterrestrial habitats and other wet terrestrial places, such as swamps, under leaf fall in wet forests, bogs, springs, ephemeral ponds, puddles, damp moss, or water-filled recesses of plants (phytotelmata) such as bromeliads and pitcher plants. Many live underground in marine and freshwater caves, sinkholes, or stream beds. Copepods are sometimes used as biodiversity indicators.
The haddock 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 it is an important species for fisheries, especially in northern Europe, where it is marketed fresh, frozen and smoked; smoked varieties include the Finnan haddie and the Arbroath smokie.
Tantulocarida is a highly specialised group of parasitic crustaceans that consists of about 33 species, treated as a class in superclass Multicrustacea. They are typically ectoparasites that infest copepods, isopods, tanaids, amphipods and ostracods.
The family Argulidae, whose members are commonly known as carp lice or fish lice, are parasitic crustaceans in the class Ichthyostraca. It is the only family in the monotypic subclass Branchiura and the order Arguloida, although a second family, Dipteropeltidae, has been proposed. Although they are thought to be primitive forms, they have no fossil record.
Sea lice are copepods of the family Caligidae within the order Siphonostomatoida. They are marine ectoparasites that feed on the mucus, epidermal tissue, and blood of host fish. The roughly 559 species in 37 genera include around 162 Lepeophtheirus and 268 Caligus species.
Monstrilloida is an order of copepods with a cosmopolitan distribution in the world's oceans. The order contains a single family, Monstrillidae. The name of the first ever described genus Monstrilla is derived from latin, meaning "tiny monster", because the lack of usual diagnostic features of copepods puzzled early taxonomists.
Lernaea is a genus of copepod crustaceans whose members are commonly called anchor worms and are parasitic on freshwater fishes.
The salmon louse is a species of copepod in the genus Lepeophtheirus. It is a sea louse, a parasite living mostly on salmon, particularly on Pacific and Atlantic salmon and sea trout, but is also sometimes found on the three-spined stickleback. It feeds on the mucus, skin and blood of the fish. Once detached, they can be blown by wind across the surface of the sea, like plankton. When they encounter a suitable marine fish host, they adhere themselves to the skin, fins, or gills of the fish, and feed on the mucus or skin. Sea lice only affect fish and are not harmful to humans.
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 larvae of crustaceans often bear little resemblance to the adult, and there are still cases where it is not known what larvae will grow into what adults. This is especially true of crustaceans which live as benthic adults, more-so than where the larvae are planktonic, and thereby easily caught.
Nicothoë astaci or the 'lobster louse' is an ectoparasitic copepod that parasitises the gills of the European lobster species Homarus gammarus. The lobster louse was first reported in 1826 by Audoin & Milne-Edwards. N. astaci has been found on lobsters inhabiting locations including Scotland, Lundy Island in the Bristol Channel and as far south as France and Portugal. The louse possesses a narrow suctorial mouthpart to feed on host haemolymph. Internally, In its adult form, Nicothoe is barely mobile and most likely remains in the same position for most of its life. The parasite occurs in groups, particularly near the base of the gills, and study has gone into its effects on the lobsters, which are considerably important, commercially. Not much is known about its life cycle, since there are significant gaps in knowledge of certain stages of its growth.
Pennella is a genus of large copepods which are common parasites of large pelagic fishes. They begin their life cycle as a series of free-swimming planktonic larvae. The females metamorphose into a parasitic stage when they attach to a host and enter into its skin. The males are free swimming. Due to their large size and mesoparasitic life history there have been a number of studies of Pennella, the members of which are among the largest of the parasitic Copepoda. All species are found as adults buried into the flesh of marine bony fish, except for a single species, Pennella balaenopterae which can be found in the muscles and blubber of cetaceans and occasionally other marine mammals, and is the largest species of copepod.
Pennella balaenopterae is a large ectoparasitic copepod specialising in parasitising marine mammals. It is the largest member of the genus Pennella, the other species of which are parasites of larger marine fish.
Lepeophtheirus pectoralis is a species of parasitic copepod from the northeast Atlantic Ocean, and the type species of the genus Lepeophtheirus. It is a parasite of flatfish, with the European flounder, the plaice, and the dab as the most frequent hosts. It feeds on the mucus, skin, and blood of the fish, with egg-producing females infecting the pectoral and pelvic fins of the host, while immature individuals and males are found on the rest of the body.
Tigriopus brevicornis is a coastal marine copepod. They are a dominant member of shallow supra tidal rock pools along the North Western European coastline. A broad range of studies have been carried out on this species, including: its ecology, physiology, phylogeography, metapopulation genetics, development and reproductive behaviour. T. brevicornis has also recently been used in ecotoxicology studies and has been trialled as a live feed for larvae in several aquaculture-based studies for the past 30 years.
Peniculisa is a genus of marine parasitic copepods in the family Pennellidae.
Pennella exocoeti is a large ectoparasitic copepod, a specialist parasite of flying fish. The adult female copepod clings to the fish's gills or skin and feeds on its body fluids.
Lernaeenicus sprattae is a species of copepod in the family Pennellidae. It is a parasite of the European sprat and certain other fish and is sometimes known as the sprat eye-maggot.
Acanthochondria cornuta is a species of parasitic copepod from the northeast Atlantic Ocean, and the type species of the genus Acanthochondria. It infects the gills of several species of flatfish, particularly the European flounder. Copepodids and immature females infect the holobranch of the host, while adult females prefer the pseudobranch and the internal wall, suggesting they migrate upstream in the gills of the host as they mature.
Contracaecum is genus of parasitic nematodes from the family Anisakidae. These nematodes are parasites of warm-blooded, fish eating animals, i.e. mammals and birds, as sexually mature adults. The eggs and the successive stages of their larvae use invertebrates and increasing size classes of fishes as intermediate hosts. It is the only genus in the family Anisakidae which can infect terrestrial, marine and freshwater animals.