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 9,900 amphipod species so far described. They are mostly marine animals, but are found in almost all aquatic environments. Some 1,900 species live in fresh water, and the order also includes terrestrial animals and sandhoppers such as Talitrus saltator.
Temporal range: Eocene–Recent
Revised division (2013)
Etymology and namesEdit
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 (0.4 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 cycleEdit
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 classificationEdit
Over 9,950 species of amphipods are currently recognised. 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.
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, often acting as mesograzers. 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 Lake 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.
In populations found in Benthic ecosystems, amphipods play an essential role in controlling brown algae growth. The mesograzer behaviour of amphipods greatly contributes to the suppression of brown algal dominance in the absence of amphipod predators. Amphipods display a strong preference for brown algae in Benthic ecosystems, but due to removal of mesograzers by predators such as fish, brown algae is able to dominate these communities over green and red algae species.
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 of small insects and crustaceans. Food is grasped with the front two pairs of legs, which are armed with large claws. More immobile species of amphipods eat higher quantities of less nutritious food rather than actively seeking more nutritious food. This is a type of compensatory feeding. This behaviour may have evolved to minimise predation risk when searching for other foods. Ampithoe longimana, for example, is more sedentary than other species and have been observed to remain on host plants longer. In fact, when presented with both high- and low-nutrition food options, the sedentary species Ampithoe longimana does not distinguish between the two options. Other amphipod species, such as Gammarus mucronatus and Elasmopus levis, which have superior predator avoidance and are more mobile, are better able to pursue different food sources. In species without the compensatory feeding ability, survivorship, fertility, and growth can be strongly negatively affected in the absence of high-quality food. Compensatory feeding may also explain the year-round presence of A. longimana in certain waters. Because algal presence changes throughout the year in certain communities, the evolution of flexible feeding techniques such as compensatory feeding may have been beneficial to survival.
Ampithoe longimana has been observed to avoid certain compounds when foraging for food. In response to this avoidance, species of seaweed such as Dictyopteris membranacea or Dictyopteris hoytii have evolved to produce C11 sulfur compounds and C-9 oxo-acids in their bodies as defense mechanisms that specifically deter amphipods instead of deterrence to consumption by other predators.
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. In addition to age, the gender seems to affect cannibalistic behaviour as males cannibalised newly moulted females less than males.
They have, rarely, been identified as feeding on humans; in Melbourne in 2017 a boy who stood in the sea for about half an hour had severe bleeding from wounds on his legs that did not coagulate easily. This was found to have been caused by "sea fleas" identified as lysianassid amphipods, possibly in a feeding group. Their bites are not venomous and do not cause lasting damage.
- Lowry J, ed. (2014). "Amphipoda". World Amphipoda database. World Register of Marine Species. Retrieved 2014-05-23.
- "Amphipoda". Integrated Taxonomic Information System.
- "Amphipoda". Oxford English Dictionary (3rd ed.). Oxford University Press. September 2005. (Subscription or UK public library membership required.)
- Wade, Sam; Corbin, Tracy; McDowell, Linda-Marie (2004). "Class Crustacea". Critter Catalogue. A guide to the aquatic invertebrates of South Australian inland waters (PDF). Waterwatch South Australia. ISBN 1-876562-67-6. Archived from the original (PDF) on 2009-10-17.
- Chan, Brian. "Freshwater shrimp (scuds, sideswimmers) – Class: Crustacea, Order: Amphipoda". Fly Fishers' Republic. Archived from the original on 23 March 2010. Retrieved April 7, 2010.
- "Order Amphipoda". Guide to the marine zooplankton of south eastern Australia. Tasmanian Aquaculture & Fisheries Institute. 2008. Archived from the original on 2008-07-20.
- Holsinger, John R. "What are amphipods?". Old Dominion University. Archived from the original on July 20, 2011. Retrieved April 7, 2010.
- Barnard, J. Laurens; Bowers, Darl E.; Haderlie, Eugene C. (1980). "Amphipoda: The Amphipods and Allies". In Morris, Robert H.; Morris, Robert Hugh; Abbott, Donald Putnam; Haderlie, Eugene Clinton (eds.). Intertidal Invertebrates of California. Stanford University Press. pp. 559–566. ISBN 0-8047-1045-7.
- Barnard, J. Laurens; Ingram, Camilla L. (1986). "The supergiant amphipod Alicella gigantea Chevreux from the North Pacific Gyre". Journal of Crustacean Biology. 6 (4): 825–839. doi:10.2307/1548395. JSTOR 1548395.
- Wolff, T. (1969). "The fauna of Rennell and Bellona, Solomon Islands". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 255 (800): 321–343. doi:10.1098/rstb.1969.0014. JSTOR 2416857.
- Peck, L. S.; Chapelle, G. (2003). "Reduced oxygen at high altitude limits maximum size". Proceedings of the Royal Society B. 270: S166–S167. doi:10.1098/rsbl.2003.0054. PMC 1809933. PMID 14667371.
- Introduction World Amphipoda Database (read November 2016)
- Vonk, R.; Schram, F. R. (2003). "Ingolfiellidea (Crustacea, Malacostraca, Amphipoda): a phylogenetic and biogeographic analysis". Contributions to Zoology. 72 (1): 39–72.
- Lowry, J. K.; Myers, A. A. (2013). "A phylogeny and classification of the Senticaudata subord. nov. (Crustacea: Amphipoda)" (PDF). Zootaxa. 3610 (1): 1–80. doi:10.11646/zootaxa.3610.1.1.
- Horton T (2013). Lowry J (ed.). "Senticaudata". World Amphipoda database. World Register of Marine Species. Retrieved October 1, 2013.
- Martin, Joel W.; Davis, George E. (2001). An Updated Classification of the Recent Crustacea (PDF). Natural History Museum of Los Angeles County. p. 132.
- Lowry, J.K.; Myers, A.A. (2017). "A Phylogeny and Classification of the Amphipoda with the establishment of the new order Ingolfiellida (Crustacea: Peracarida)". Zootaxa. Magnolia Press. 4265 (1): 001–089. doi:10.11646/zootaxa.4265.1.1.
- Bousfield, E. L.; Poinar, G. O., Jr. (1994). "A new terrestrial amphipod from tertiary amber deposits of Chiapas province, Southern Mexico". Historical Biology. 7 (2): 105–114. doi:10.1080/10292389409380448.
- The species Rosagammarus minichiellus from the considerably older Late Triassic Luning Formation of Nevada was originally described as an amphipod, but subsequently reinterpreted as the right half of a decapod tail (Starr, Hegna & McMenamin 2015, The Geological Society of America North-Central Section 49th Annual Meeting )
- Lowry, J. K.; Springthorpe, R. T. "Introduction". Amphipoda: Families. Australian Museum. Retrieved April 5, 2010.
- Duffy, J. E.; Hay, Mark E. (2000). "Strong impacts of grazing amphipods on the organization of a benthic community". Ecological Monographs. 70 (2): 237–263. CiteSeerX 10.1.1.473.4746. doi:10.1890/0012-9615(2000)070[0237:SIOGAO]2.0.CO;2.
- Harbison, G. R.; Biggs, D. C.; Madin, L. P. (1977). "The associations of Amphipoda Hyperiidea with gelatinous zooplankton. II. Associations with Cnidaria, Cteuophora and Radiolaria". Deep-Sea Research. 24 (5): 465–488. doi:10.1016/0146-6291(77)90484-2.
- Väinölä, R.; Witt, J. D. S.; Grabowski, M.; Bradbury, J. H.; Jazdzewski, K.; Sket, B. (2008). "Global diversity of amphipods (Amphipoda, Crustacea) in freshwater" (PDF). Hydrobiologia. 595 (1): 241–255. doi:10.1007/s10750-007-9020-6.
- Minor, M. A.; Robertson, A. W. (March 5, 2010). "Amphipoda". Guide to New Zealand Soil Invertebrates. Massey University. Archived from the original on 10 May 2010. Retrieved April 7, 2010.
- Hobbs, Horton H., III (2003). "Crustacea". In Gunn, John (ed.). Encyclopedia of Caves and Karst Science (PDF). Routledge. pp. 254–257. ISBN 978-1-57958-399-6.
- Goater, Tim (May 4, 1996). "Parasitic Amphipoda". Interactive Parasitology. Vancouver Island University. Archived from the original on July 14, 2010. Retrieved April 7, 2010.
- Cruz-Rivera, Edwin; Hay, Mark E. (2000). "Can quantity replace quality? Food choice, compensatory feeding, and fitness of marine mesograzers". Ecology. 81: 201–219. doi:10.1890/0012-9658(2000)081[0201:CQRQFC]2.0.CO;2.
- Cruz-Rivera, Edwin; Hay, Mark E. (2001). "Macroalgal traits and the feeding and fitness of an herbivorous amphipod: the roles of selectivity, mixing, and compensation". Marine Ecology Progress Series. 218: 249–266. doi:10.3354/meps218249.
- Schnitzler, Iris; Pohnert, Georg; Hay, Mark; Boland, Wilhelm (2001). "Chemical defense of brown algae (Dictyopteris spp.) against the herbivorous amphipod Ampithoe longimana". Oecologia. 126 (4): 515–521. doi:10.1007/s004420000546.
- Dick, Jaimie T. A. (1995). "The cannibalistic behaviour of two Gammarus species (Crustacea: Amphipoda)". Journal of Zoology. 236 (4): 697–706. doi:10.1111/j.1469-7998.1995.tb02740.x.
- Lewis, Susan E.; Dick, Jaimie T. A.; Lagerstrom, Erin K.; Clarke, Hazel C. (2010). "Avoidance of filial cannibalism in the amphipod Gammarus pulex". Ethology. 116 (2): 138–146. doi:10.1111/j.1439-0310.2009.01726.x.
- "Museum identifies 'Flesh Eating Creatures'". Bayside City Council, Victoria, Australia. 7 August 2017. Retrieved 8 August 2017.