Johngarthia lagostoma
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''Johngarthia lagostoma'' is a species of
terrestrial crab A number of lineages of crabs have evolved to live predominantly on land. Examples of terrestrial crabs are found in the families Gecarcinidae and Gecarcinucidae, as well as in selected genera from other families, such as '' Sesarma'', althoug ...
that lives on Ascension Island and three other islands in the
South 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 an ...
. It grows to a carapace width of on Ascension Island, where it is the largest native land animal. It exists in two distinct colour morphs, one yellow and one purple, with few intermediates. The yellow morph dominates on Ascension Island, while the purple morph is more frequent on
Rocas Atoll The Rocas Atoll ( pt, Atol das Rocas ) is the only atoll in the South Atlantic Ocean. It belongs to the Brazilian State of Rio Grande do Norte. It is located approximately northeast of Natal and west of the Fernando de Noronha archipelago. The ...
. The species differs from other '' Johngarthia'' species by the form of the third maxilliped. ''Johngarthia lagostoma'' lives in burrows among vegetation, at altitudes of up to , emerging at night to feed on plant matter and occasionally on animals. From January to March there is an annual migration to the sea to release the
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
ic larvae. The species was first described (as ''Gecarcinus lagostoma'') by Henri Milne-Edwards in 1837 from material sent to him by the naturalists Jean René Constant Quoy and Joseph Paul Gaimard, collected by the French ship '' Astrolabe''.


Distribution and biogeography

''Johngarthia lagostoma'' is found on Ascension Island, Trindade Island,
Fernando de Noronha Fernando de Noronha () is an archipelago in the Atlantic Ocean, part of the State of Pernambuco, Brazil, and located off the Brazilian coast. It consists of 21 islands and islets, extending over an area of . Only the eponymous main island is in ...
and the
Rocas Atoll The Rocas Atoll ( pt, Atol das Rocas ) is the only atoll in the South Atlantic Ocean. It belongs to the Brazilian State of Rio Grande do Norte. It is located approximately northeast of Natal and west of the Fernando de Noronha archipelago. The ...
. On Ascension Island, ''J. lagostoma'' is restricted outside the breeding season to the slopes of Green Mountain, where there is sufficient moisture and vegetation, the rest of the island being too arid for the crab to survive. All the land above is suitable habitat for the crabs, as is much of the land above . They are occasionally found at lower altitudes, including the well-watered gardens of Georgetown, and the sooty tern breeding colony in the south-west of the island (known as the Wideawake Fairs). On Trindade Island, ''J. lagostoma'' is abundant wherever plants grow, including the highest parts of the island. The geographic distribution of ''J. lagostoma'' across a small number of islands in the southern
Atlantic Ocean 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 ...
is very unusual, and difficult to explain by planktonic dispersal. Its nearest relatives are the other species of '' Johngarthia'', two of which ('' J. malpilensis'' and '' J. planata'') inhabit islands in the
Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the contin ...
off
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
, and one ('' J. weileri'') is found on islands in the
Gulf of Guinea The Gulf of Guinea is the northeasternmost part of the tropical Atlantic Ocean from Cape Lopez in Gabon, north and west to Cape Palmas in Liberia. The intersection of the Equator and Prime Meridian (zero degrees latitude and longitude) is in ...
. Some authors have suggested the existence of former islands, now submerged, which could have acted as " stepping stones" for the colonisation of Ascension Island; the isolation of Ascension Island from any other land mass makes transportation of either larvae or adults difficult.


Description

Mature specimens of ''J. lagostoma'' are typically wide across the carapace on Ascension Island; individuals from the Rocas Atoll are somewhat smaller. In the family Gecarcinidae, species are normally separated by the form of the first pleopod (gonopod), which is used by males during mating, but there is no difference in the gonopod between ''J. lagostoma'' and '' J. planata''. Instead, ''J. lagostoma'' differs from other species in the genus by the form of the third maxilliped; it has a fissure which is a narrow slit, but which gapes open in other species. The third maxilliped is also larger, covering the
epistome Stomotheca is the term applied to the feeding apparatus in front of the mouth of harvestmen, and sometimes the related scorpions. Usually it consists of the ''epistome'' ( labrum), two pairs of ''coxapophyses'' (endites, maxillary lobes) and often a ...
and the antennules in ''J. lagostoma'' but not in other species. Within the species, two distinct colour morphs can be seen. The "yellow" morph has a bright yellow or orange
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
, with white patches on the underside of the walking legs and claws. The "purple" morph has a dark purple exoskeleton, but with the same white patches as on the yellow morph. A few intermediate individuals occur, which are predominantly yellow, but with purple patches on the carapace. On Ascension Island, the yellow morph is the more frequent one, while on the Rocas Atoll, the distribution is more equal. Darker individuals are thought to be better concealed from
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
s, but more prone to
heat stress Hyperthermia, also known simply as overheating, is a condition in which an individual's body temperature is elevated beyond normal due to failed thermoregulation. The person's body produces or absorbs more heat than it dissipates. When extreme ...
during long migrations; The Rocas Atoll is much smaller than Ascension Island, and the migration routes are therefore shorter, which may explain the difference in frequency of the two morphs.


Ecology and conservation

Before Ascension Island was colonised by Europeans in the 19th century, ''Johngarthia lagostoma'' was the only large land animal on the island. Since then, many species of mammal have been introduced to Ascension Island, and now compete with ''J. lagostoma''; they include mice, rats and rabbits. ''J. lagostoma'' is active at night and after rain, when it emerges from its burrows, which can be up to deep. In 1915, H. A. Baylis reported that it feeds on "decaying vegetation and perhaps a certain amount of excreta from sea-birds"; most of its diet is thought to be made up of plant matter, but can also include animals. In the Wideawake Fairs, ''J. lagostoma'' feeds on chicks and eggs of the sooty tern, and they have been proposed as a
limiting factor A limiting factor is a variable of a system that causes a noticeable change in output or another measure of a type of system. The limiting factor is in a pyramid shape of organisms going up from the producers to consumers and so on. A factor not l ...
in the breeding of
petrel Petrels are tube-nosed seabirds in the bird order Procellariiformes. Description The common name does not indicate relationship beyond that point, as "petrels" occur in three of the four families within that group (all except the albatross f ...
s on Ascension Island. In May, green sea turtles (''Chelonia mydas'') hatch, and ''J. lagostoma'' is one of several predators to target the emerging
hatchling In oviparous biology, a hatchling is a newly hatched fish, amphibian, reptile, or bird. A group of mammals called monotremes lay eggs, and their young are hatchlings as well. Fish Fish hatchlings generally do not receive parental care, similar to ...
s, alongside the
Ascension frigatebird The Ascension frigatebird (''Fregata aquila'') is a seabird of the frigatebird family Fregatidae which breeds on Boatswain Bird Island and Ascension Island in the tropical Atlantic Ocean. The Ascension frigatebird is a large lightly built se ...
(''Fregata aquila'') and
feral cat A feral cat or a stray cat is an unowned domestic cat (''Felis catus'') that lives outdoors and avoids human contact: it does not allow itself to be handled or touched, and usually remains hidden from humans. Feral cats may breed over dozens ...
s. The crabs find water on and under rocks, especially after condensation has begun to form on their cool surfaces at night. In the late nineteenth century, a
bounty Bounty or bounties commonly refers to: * Bounty (reward), an amount of money or other reward offered by an organization for a specific task done with a person or thing Bounty or bounties may also refer to: Geography * Bounty, Saskatchewan, a g ...
was levied on ''J. lagostoma'' in order to reduce its numbers. Hunters were paid 6 pence (alternatively paid in
rum Rum is a liquor made by fermenting and then distilling sugarcane molasses or sugarcane juice. The distillate, a clear liquid, is usually aged in oak barrels. Rum is produced in nearly every sugar-producing region of the world, such as the Ph ...
) for every hundred claws collected. In 1879, the total bag for the island comprised 66 cats, 4,013 mice, 7,683 rats and 80,414 land crabs; in the eight years to 1887, bounties were collected on more than 330,000 crabs. The current population is much smaller, and the low level of recruitment and lack of juveniles suggests that the species may become endangered.


Life cycle

Although juvenile and adult ''J. lagostoma'' are almost exclusively terrestrial, the larvae are marine and
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
ic. In order to release their offspring, the adults therefore have to migrate to the sea, as seen in other land crabs, such as the
Christmas Island red crab The Christmas Island red crab (''Gecarcoidea natalis'') is a species of land crab that is endemic to Christmas Island and Cocos (Keeling) Islands in the Indian Ocean. Although restricted to a relatively small area, an estimated 43.7 million a ...
, ''Gecarcoidea natalis''. In most members of the family
Gecarcinidae The Gecarcinidae, the land crabs, are a family of true crabs that are adapted for terrestrial existence. Similar to all other crabs, land crabs possess a series of gills. In addition, the part of the carapace covering the gills is inflated and e ...
, migration coincides with the
rainy season The rainy season is the time of year when most of a region's average annual rainfall occurs. Rainy Season may also refer to: * ''Rainy Season'' (short story), a 1989 short horror story by Stephen King * "Rainy Season", a 2018 song by Monni * '' ...
, which reduces the risk of desiccation; migration in ''J. lagostoma'' occurs from January until March, while the period of highest rainfall is from March to May. The crabs travel approximately per day, and mating can take place anywhere along the route. The proportion of males on the migration therefore decreases as the migration continues. The
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
are larger than in other gecarcinid species and consequently fewer in number; females with a carapace width of had a mean
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 natural capability to pr ...
of 72,000. Spawning occurs in the last quarter of the
lunar cycle Concerning the lunar month of ~29.53 days as viewed from Earth, the lunar phase or Moon phase is the shape of the Moon's directly sunlit portion, which can be expressed quantitatively using areas or angles, or described qualitatively using the t ...
, during
neap tide Tides are the rise and fall of sea levels caused by the combined effects of the gravitational forces exerted by the Moon (and to a much lesser extent, the Sun) and are also caused by the Earth and Moon orbiting one another. Tide tables c ...
s, on rocky shores.


Taxonomy and history

''Johngarthia lagostoma'' was first described by Henri Milne-Edwards in 1837, under the name ''Gecarcinus lagostoma''. The specific epithet ' means " hare-lip", from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
(', "hare") and (', "mouth"); Milne-Edwards also named the species in French ' ("hare-lipped land crab"). Milne-Edwards erroneously gave the type locality as "" (
Australasia Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecologi ...
); he had been given the material he used for the description by Jean René Constant Quoy and Joseph Paul Gaimard, who were working on the molluscs collected on the '' Astrolabe'' excursion, which had visited Australasia and stopped at Ascension Island in 1829. ''G. lagostoma'' was placed in the subgenus ''Johngarthia'' by Michael Türkay in 1970, who raised the subgenus to the
rank Rank is the relative position, value, worth, complexity, power, importance, authority, level, etc. of a person or object within a ranking, such as: Level or position in a hierarchical organization * Academic rank * Diplomatic rank * Hierarchy * ...
of
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
in 1987. Many subsequent voyages halted at Ascension Island, and many visitors commented on the land crabs they found there, including
William Dampier William Dampier (baptised 5 September 1651; died March 1715) was an English explorer, pirate, privateer, navigator, and naturalist who became the first Englishman to explore parts of what is today Australia, and the first person to circumnav ...
,
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
and Charles Wyville Thomson.
Bernard Stonehouse Bernard Stonehouse (1 May 1926 – 12 November 2014) was a British scientist who specialised in animal behaviour, polar research and popular science. In 1953 he received the Polar Medal. Early life and military service Stonehouse was born in Hu ...
reported on an expedition by the British Ornithologists' Union to Ascension Island, stating: Cited in Manning & Chace (1990).
We met land crabs, too, and were sadly disappointed. They were small, no larger than an ordinary edible crab, with bloated shells and moderately long pincers. Some were of beetroot purple, others yellow or orange, apparently two colour-phases of the same species. They were shy, freezing with pincers erect when alarmed, usually to be found within a short distance of their burrows and ready to scuttle down at the drop of a hat. In rainy weather they promenaded more freely, sometimes appearing at the side of road with arms waving like diminutive but aggressive hitch-hikers. Fringed mandibles hird maxillipedssuggest a permanently turned-down mouth, giving them a disgruntled, unhappy expression; they were disagreeable rather than sinister and fell far short of their reputation. The first land crab I met was sitting in a prickly pear bush, sadly munching one of the brilliant red fruits and dribbling juice. I could never take them seriously after that.
The closely related crabs from the islands of the
Gulf of Guinea The Gulf of Guinea is the northeasternmost part of the tropical Atlantic Ocean from Cape Lopez in Gabon, north and west to Cape Palmas in Liberia. The intersection of the Equator and Prime Meridian (zero degrees latitude and longitude) is in ...
were formerly included in ''Gecarcinus lagostoma'', but are now treated as a separate species, ''
Johngarthia weileri ''Johngarthia weileri'' is a species of land crab in the genus '' Johngarthia'' from the eastern Atlantic Ocean. Distribution ''Johngarthia weileri'' is found on the coast of Cameroon and on the islands in the Gulf of Guinea The Gulf of Guine ...
''.


References

{{Taxonbar, from=Q3182828 Grapsoidea Crustaceans of the Atlantic Ocean Fauna of Ascension Island Crustaceans described in 1837 Taxa named by Henri Milne-Edwards