Around 1,300 species of freshwater crabs are distributed throughout the

tropics The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...

and

subtropics The subtropical zones or subtropics are geographical and climate zones immediately to the north and south of the tropics. Geographically part of the temperate zones of both hemispheres, they cover the middle latitudes from to approximately ...

, divided among eight

families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...

. They show direct development and maternal care of a small number of offspring, in contrast to marine crabs, which release thousands of planktonic

larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...

. This limits the dispersal abilities of freshwater crabs, so they tend to be

endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

to small areas. As a result, a large proportion are threatened with extinction.

Systematics

More than 1,300 described species of freshwater crabs are known, out of a total of 6,700 species of crabs across all environments. The total number of species of freshwater crabs, including

undescribed species In taxonomy, an undescribed taxon is a taxon (for example, a species) that has been discovered, but not yet formally described and named. The various Nomenclature Codes specify the requirements for a new taxon to be validly described and named. U ...

, is thought to be up to 65% higher, potentially up to 2,155 species, although most of the additional species are currently unknown to science. They belong to eight families, each with a limited distribution, although various crabs from other families are also able to tolerate freshwater conditions (

euryhaline Euryhaline organisms are able to adapt to a wide range of salinities. An example of a euryhaline fish is the short-finned molly, '' Poecilia sphenops'', which can live in fresh water, brackish water, or salt water. The green crab ('' Carcinus m ...

) or are secondarily adapted to fresh water. The phylogenetic relationships between these families is still a matter of debate, so how many times the freshwater lifestyle has evolved among the true crabs is unknown. The eight families are: ;Superfamily Trichodactyloidea * Trichodactylidae (

Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...

and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

) ;Superfamily Potamoidea: * Potamidae ( Mediterranean Basin and

Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...

) * Potamonautidae (

Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...

, including

Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...

) * Deckeniidae (

East Africa East Africa, also known as Eastern Africa or the East of Africa, is a region at the eastern edge of the Africa, African continent, distinguished by its unique geographical, historical, and cultural landscape. Defined in varying scopes, the regi ...

and

Seychelles Seychelles (, ; ), officially the Republic of Seychelles (; Seychellois Creole: ), is an island country and archipelagic state consisting of 155 islands (as per the Constitution) in the Indian Ocean. Its capital and largest city, Victoria, ...

) – also treated as part of Potamonautidae * Platythelphusidae (East Africa) – also treated as part of Potamonautidae ;Superfamily Gecarcinucoidea *

Gecarcinucidae The Gecarcinucidae are a family of true freshwater crabs. They are found throughout South Asia, Southeast Asia and New Guinea, with a single genus found in Australia. Taxonomy The family Parathelphusidae is now demoted to the rank of subfamily ...

(Asia) * Parathelphusidae (Asia and

Australasia Australasia is a subregion of Oceania, comprising Australia, New Zealand (overlapping with Polynesia), and sometimes including New Guinea and surrounding islands (overlapping with Melanesia). The term is used in a number of different context ...

) – nowadays treated as a junior synonym of Gecarcinucidae ;Superfamily Pseudothelphusoidea: *

Pseudothelphusidae Pseudothelphusidae is a family of freshwater crabs found chiefly in mountain streams in the Neotropics. They are believed to have originated in the Greater Antilles and then crossed to Central America via a Pliocene land bridge. Some species of t ...

(Central America and South America) The

fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

of freshwater organisms is typically poor, so few fossils of freshwater crabs have been found. The oldest is '' Tanzanonautes tuerkayi'', from the

Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...

, and the evolution of freshwater crabs is likely to postdate the break-up of the supercontinent

Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...

. Members of the family Aeglidae and '' Clibanarius fonticola'' are also restricted to fresh water, but these "crab-like" crustaceans are members of the infraorder

Anomura Anomura (sometimes Anomala) is a group of decapod crustaceans, including hermit crabs and others. Although the names of many anomurans include the word ''crab'', all true crabs are in the sister group to the Anomura, the Brachyura (the two groups ...

(true crabs are Brachyura).

Description and lifecycle

The external morphology of freshwater crabs varies very little, so the form of the

gonopod Gonopods are specialized appendages of various arthropods used in reproduction or egg-laying. In males, they facilitate the transfer of sperm from male to female during mating, and thus are a type of intromittent organ. In crustaceans and millipe ...

(first abdominal appendage, modified for insemination) is of critical importance for

classification Classification is the activity of assigning objects to some pre-existing classes or categories. This is distinct from the task of establishing the classes themselves (for example through cluster analysis). Examples include diagnostic tests, identif ...

Development Development or developing may refer to: Arts *Development (music), the process by which thematic material is reshaped * Photographic development *Filmmaking, development phase, including finance and budgeting * Development hell, when a proje ...

of freshwater crabs is characteristically direct, where the eggs hatch as juveniles, with the larval stages passing within the egg. The broods comprise only a few hundred eggs (compared to hundreds of thousands for marine crabs), each of which is quite large, at a diameter around . The colonisation of fresh water has required crabs to alter their water balance; freshwater crabs can reabsorb salt from their

urine Urine is a liquid by-product of metabolism in humans and many other animals. In placental mammals, urine flows from the Kidney (vertebrates), kidneys through the ureters to the urinary bladder and exits the urethra through the penile meatus (mal ...

, and have various

adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...

s to reduce the loss of water. In addition to their

gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, 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 r ...

s, freshwater crabs have a "pseudolung" in their gill chamber that allows them to breathe in air. These developments have preadapted freshwater crabs for terrestrial living, although freshwater crabs need to return to water periodically to excrete

ammonia Ammonia is an inorganic chemical compound of nitrogen and hydrogen with the chemical formula, formula . A Binary compounds of hydrogen, stable binary hydride and the simplest pnictogen hydride, ammonia is a colourless gas with a distinctive pu ...

Ecology and conservation

Freshwater crabs are found throughout the tropical and subtropical regions of the world. They live in a wide range of water bodies, from fast-flowing rivers to

swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...

s, as well as in tree boles or

cave Caves or caverns are natural voids under the Earth's Planetary surface, surface. Caves often form by the weathering of rock and often extend deep underground. Exogene caves are smaller openings that extend a relatively short distance undergrou ...

s. They are primarily

nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...

, emerging to feed at night; most are

omnivore An omnivore () is an animal that regularly consumes significant quantities of both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize t ...

s, although a small number are specialist predators, such as '' Platythelphusa armata'' from

Lake Tanganyika Lake Tanganyika ( ; ) is an African Great Lakes, African Great Lake. It is the world's List of lakes by volume, second-largest freshwater lake by volume and the List of lakes by depth, second deepest, in both cases after Lake Baikal in Siberia. ...

, which feeds almost entirely on

snail A snail is a shelled gastropod. The name is most often applied to land snails, terrestrial molluscs, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class Gas ...

s. Some species provide important food sources for various

vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...

s. A number of freshwater crabs (for example species from the genus ''

Nanhaipotamon ''Nanhaipotamon'' is a genus of freshwater crabs, in the subfamily Potamiscinae, found in southern China and Taiwan. As of 2018, 18 species have been described. The genus is named after the South China Sea (; literally: 'South Sea'), for it occur ...

'') are secondary hosts of flukes in the genus '' Paragonimus'', which causes

paragonimiasis Paragonimiasis is a food-borne parasitic disease caused by several species of lung flukes belonging to genus '' Paragonimus''. Infection is acquired by eating crustaceans such as crabs and crayfishes which host the infective forms called metacerc ...

in humans. The majority of species are narrow endemics, occurring in only a small geographical area. This is at least partly attributable to their poor dispersal abilities and low

fecundity Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the capability to produc ...

, and to

habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological proces ...

caused by the world's human population. In

West Africa West Africa, also known as Western Africa, is the westernmost region of Africa. The United Nations geoscheme for Africa#Western Africa, United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Gha ...

, species that live in

savanna A savanna or savannah is a mixed woodland-grassland (i.e. grassy woodland) biome and ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach th ...

s have wider ranges than species from the

rainforest Rainforests are forests characterized by a closed and continuous tree Canopy (biology), canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforests can be generally classified as tropi ...

; in

, species from the mountains have restricted distributions, while lowland species are more widespread. Every species of freshwater crab described so far has been assessed by the

International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the stat ...

; of the species for which data are available, 32% are threatened with extinction. For instance, all but one of

Sri Lanka Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka, also known historically as Ceylon, is an island country in South Asia. It lies in the Indian Ocean, southwest of the Bay of Bengal, separated from the Indian subcontinent, ...

's 50 freshwater crab species are endemic to that country, and more than half are

critically endangered An IUCN Red List critically endangered (CR or sometimes CE) species is one that has been categorized by the International Union for Conservation of Nature as facing an extremely high risk of extinction in the wild. As of December 2023, of t ...

References

External links

*{{cite web , url=http://faculty.nmu.edu/ncumberl/Neil/MainFWC-website/FWCBiology.html , title=Freshwater Crab Biology , author1=Neil Cumberlidge , author2=Sadie K. Reed , date=April 4, 2009 , publisher=

Northern Michigan University Northern Michigan University (Northern Michigan, Northern or NMU) is a public university in Marquette, Michigan, United States. It was established in 1899 by the Michigan Legislature as Northern State Normal School. In 1963, the state designa ...

, url-status=dead , archive-url=https://web.archive.org/web/20110720014222/http://faculty.nmu.edu/ncumberl/Neil/MainFWC-website/FWCBiology.html , archive-date=July 20, 2011 Crabs Freshwater crustaceans