Temporal range: Eocene–Recent
Revised division (2013)
Amphipoda is an order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods range in size from 1 to 340 millimetres (0.039 to 13 in) and are mostly detritivores or scavengers. There are more than 9,500 amphipod species so far described. They are mostly marine animals, but are found in almost all aquatic environments. Some 1900 species live in freshwater, and the order also includes terrestrial animals and sandhoppers such as Talitrus saltator.
Etymology and names
The name Amphipoda comes, via the New Latin amphipoda, from the Greek roots ἀμφί ("different") and πούς ("foot"), in reference to two kinds of legs that amphipods possess. This contrasts with the related Isopoda, which have a single kind of thoracic leg. Particularly among anglers, amphipods are known as freshwater shrimp, scuds or sideswimmers.
The thorax and abdomen are usually quite distinct and bear different kinds of legs; they are typically laterally compressed, and there is no carapace. The thorax bears eight pairs of uniramous appendages, the first of which are used as accessory mouthparts; the next four pairs are directed forwards, and the last three pairs are directed backwards. Gills are present on the thoracic segments, and there is an open circulatory system with a heart, using haemocyanin to carry oxygen in the haemolymph to the tissues. The uptake and excretion of salts is controlled by special glands on the antennae.
The abdomen is divided into two parts: the pleosome which bears swimming legs; and the urosome, which comprises a telson and three pairs of uropods which do not form a tail fan as they do in animals such as true shrimp.
Amphipods are typically less than 10 millimetres (0.39 in) long, but the largest recorded living amphipods were 28 centimetres (11 in) long, and were photographed at a depth of 5,300 metres (17,400 ft) in the Pacific Ocean. Samples from the Atlantic Ocean with a reconstructed length of 34 centimetres (13 in) have been assigned to the same species, Alicella gigantea. The smallest known amphipods are less than 1 millimetre (0.04 in) long. The size of amphipods is limited by the availability of dissolved oxygen, such that the amphipods in Lake Titicaca at an altitude of 3,800 metres (12,500 ft) can only grow up to 22 millimetres (0.87 in), compared to lengths of 90 millimetres (3.5 in) in Lake Baikal at 455 metres (1,500 ft).
Reproduction and life cycle
Mature females bear a marsupium, or brood pouch, which holds her eggs while they are fertilised, and until the young are ready to hatch. As a female ages, she produces more eggs in each brood. Mortality is around 25%–50% for the eggs. There are no larval stages; the eggs hatch directly into a juvenile form, and sexual maturity is generally reached after 6 moults. Some species have been known to eat their own exuviae after moulting.
Diversity and classification
Over 9,500 species of amphipods are currently recognized. Traditionally they were placed in the four suborders Gammaridea (which contained the majority of taxa, including all the freshwater and terrestrial species), Caprellidea, Hyperiidea, and Ingolfiellidea (the latter with only 40 species).
The classification of the Amphipoda is however being rearranged to better reflect their phylogeny, the relationships within the suborder Gammaridea having suffered from the most confusion. A new classification has been developed in the works of Lowry & Myers, where a new large suborder Senticaudata was split off from the Gammaridea in 2013. That taxon, which also encompasses the previous Caprellidea, now comprises over half of the known amphipod species.
The classification given below, from the rank of suborder down to superfamily, however still represents the traditional division as given in Martin & Davis (2001), except that superfamilies are recognised here[according to whom?] within the Gammaridea.
Amphipods are thought to have originated in the Lower Carboniferous. Despite the group's age, however, the fossil record of the order Amphipoda is meagre, comprising specimens of 12 species dating back only as far as the Upper Eocene, where they have been found in Baltic amber. The species Rosagammarus minichiellus, known from the Late Triassic (Norian) Luning Formation of Nevada, was originally described as an amphipod; however, the holotype specimen was subsequently reinterpreted as the right half of a decapod tail of uncertain taxonomic affinity.
Amphipods are found in almost all aquatic environments, from fresh water to water with twice the salinity of sea water. They are almost always an important component of aquatic ecosystems. Most species in the suborder Gammaridea are epibenthic, although they are often collected in plankton samples. Members of the Hyperiidea are all planktonic and marine. Many are symbionts of gelatinous animals, including salps, medusae, siphonophores, colonial radiolarians and ctenophores, and most hyperiids are associated with gelatinous animals during some part of their life cycle. Some 1,900 species, or 20% of the total amphipod diversity, live in fresh water or other non-marine waters. Notably rich endemic amphipod faunas are found in the ancient lakes Baikal and waters of the Caspian Sea basin.
The landhoppers of the family Talitridae (which also includes semi-terrestrial and marine animals) are terrestrial, living in damp environments such as leaf litter. Landhoppers have a wide distribution in areas that were formerly part of Gondwanaland, but have colonised parts of Europe and North America in recent times.
Around 750 species in 160 genera and 30 families are troglobitic, and are found in almost all suitable habitats, but with their centres of diversity in the Mediterranean Basin, southeastern North America and the Caribbean.
Compared to other crustacean groups, such as the Isopoda, Rhizocephala or Copepoda, relatively few amphipods are parasitic on other animals. The most notable example of parasitic amphipods are the whale lice (family Cyamidae); unlike other amphipods, these are dorso-ventrally flattened, and have large, strong claws, with which they attach themselves to baleen whales. They are the only parasitic crustaceans which cannot swim during any part of their life cycle.
Most amphipods are detritivores or scavengers, with some being grazers of algae, omnivores or predators on small insects and crustaceans. Food is grasped with the front two pairs of legs which are armed with large claws. More inactive species have been observed to exhibit compensatory feeding to minimize predation risk in comparison to more mobile species. The incidence of cannibalism and intraguild predation is relatively high in some species, although adults may decrease cannibalistic behaviour directed at juveniles when they are likely to encounter their own offspring.
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