Lepidurus Apus
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''Lepidurus apus'', commonly known as a tadpole shrimp, is a
notostraca The order Notostraca, containing the single family Triopsidae, is a group of crustaceans known as tadpole shrimp or shield shrimp. The two genera, ''Triops'' and '' Lepidurus'', are considered living fossils, with similar forms having existed sinc ...
n in the family Triopsidae, one of a lineage of shrimp-like
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s that have had a similar form since the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
period and are considered
living fossil A living fossil is an extant taxon that cosmetically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of origin of the extant clade. Living fossi ...
s. This species is
cosmopolitan Cosmopolitan may refer to: Food and drink * Cosmopolitan (cocktail), also known as a "Cosmo" History * Rootless cosmopolitan, a Soviet derogatory epithet during Joseph Stalin's anti-Semitic campaign of 1949–1953 Hotels and resorts * Cosmopoli ...
, inhabiting temporary freshwater ponds over much of the world, and the most widespread of the tadpole shrimps. Like other notostracans, ''L. apus'' has a broad
carapace A carapace is a Dorsum (biology), dorsal (upper) section of the exoskeleton or shell in a number of animal groups, including arthropods, such as crustaceans and arachnids, as well as vertebrates, such as turtles and tortoises. In turtles and tor ...
, long segmented
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
, and large numbers of paddle-like legs. It reproduces by a mixture of sexual reproduction and self-fertilisation of females.


Description

''Lepidurus apus'' grows to in length. Its long abdomen is divided into about 30 segment-like rings, with two long caudal rami or "tails" attached behind the last ring. Between the tails is a projection which distinguishes ''
Lepidurus ''Lepidurus'' is a genus of small crustaceans in the order Notostraca (tadpole shrimp). It is the larger of the two extant genera of the tadpole shrimps, the other being ''Triops''. They are commonly found in vernal pools and survive dry periods ...
'' from ''
Triops ''Triops'' is a genus of small crustaceans in the order Notostraca (tadpole shrimp). The long-lasting resting eggs of several species of ''Triops'' are commonly sold in kits as a pet. The animals hatch upon contact with fresh water. Most adult-st ...
'', the other notostracan
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
. Its carapace is flat with an average length of , and is attached only at the front, covering up to two thirds of the abdomen. The carapace is a mottled dark yellow/brown colour transitioning to a lighter edge, and bears a single pair of compound eyes. At the front of the abdomen are one or more (up to three) pairs of feelers. Under the body are 41–46 (average 44) pairs of paddle-like limbs used for swimming. Males are readily identifiable by the lack of ovisacs, and also have subtle differences in the carapace. Females and hermaphrodites look virtually identical, but hermaphrodites have testicular lobes amongst their ovarian lobes, which allows them to reproduce in isolation. ''L. apus'' is often referred to as a ‘living fossil’, virtually unchanged for over 300 million years. However, a recent study suggests that resemblance to fossil notostracans is probably a result of the "highly conserved general morphology in this group and of
homoplasy Homoplasy, in biology and phylogenetics, is the term used to describe a Phenotypic trait, feature that has been gained or lost independently in separate lineages over the course of evolution. This is different from Homology (biology), homology, w ...
". Recent species of ''Lepidurus'' are morphologically nearly identical to fossil records but may "represent very different evolutionary lineages".


Distribution and habitat


Global range

''Lepidurus apus'' is perhaps the most cosmopolitan of all the Notostraca, occurring widely around the world including, but not limited to,
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...
,
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
,
Iran Iran, officially the Islamic Republic of Iran, and also called Persia, is a country located in Western Asia. It is bordered by Iraq and Turkey to the west, by Azerbaijan and Armenia to the northwest, by the Caspian Sea and Turkmeni ...
,
Israel Israel (; he, יִשְׂרָאֵל, ; ar, إِسْرَائِيل, ), officially the State of Israel ( he, מְדִינַת יִשְׂרָאֵל, label=none, translit=Medīnat Yīsrāʾēl; ), is a country in Western Asia. It is situated ...
,
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
,
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
,
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...
,
Denmark ) , song = ( en, "King Christian stood by the lofty mast") , song_type = National and royal anthem , image_map = EU-Denmark.svg , map_caption = , subdivision_type = Sovereign state , subdivision_name = Danish Realm, Kingdom of Denmark ...
, and
Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
. ''Lepidurus apus'' is split into several geographic subspecies, such as ''L. apus viridis,'' present in parts of Australia, Tasmania and New Zealand.


Habitat preferences

''Lepidurus apus'' is found predominantly in temporary freshwater ponds, 10–100 cm deep, filled during autumn and winter, and drying out over spring and summer. It is less common in permanent water bodies such as swamps and ditches. ''Lepidurus apus viridis'', for example, is found throughout New Zealand in small ponds and ditches. Its life-cycle allows it to become dormant if the pond freezes over, is covered with snow, or dries out; it can persist in the dry sediment margin as a
cyst A cyst is a closed sac, having a distinct envelope and cell division, division compared with the nearby Biological tissue, tissue. Hence, it is a cluster of Cell (biology), cells that have grouped together to form a sac (like the manner in which ...
form, which can survive harsh conditions for many years until the pond reappears. ''L. apus'' prefers a pH between 6 and 7.8, and can tolerate relatively high concentrations of chemicals, such as levels of total
nitrate Nitrate is a polyatomic ion A polyatomic ion, also known as a molecular ion, is a covalent bonded set of two or more atoms, or of a metal complex, that can be considered to behave as a single unit and that has a net charge that is not zer ...
of 1 mg/L and
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phospho ...
of 0.1 mg/L, a level which would be detrimental to other aquatic species. Wetlands and temporary ponds worldwide are increasingly being converted to grasslands for agriculture, so the total land area available for ''L. apus'' is gradually diminishing. Some subspecies may become threatened in future, or may already be under threat, although our knowledge is limited. ''L. apus'' is well-adapted to variations of climate and location, disperses easily, and has highly resilient eggs, so appears to be less sensitive to human pressures across its wide geographical range.


Life cycle

''Lepidurus apus'' has an unusual life-cycle: it is able to produce microscopic cysts that can lie dormant for years at a time through extreme conditions, letting it survive in areas with vastly different climates such as Morocco and Denmark. These "resting eggs" are so drought resistant there is a record of hatching after being kept dry for 28 years. The species is hermaphroditic; no males are found in the New Zealand subspecies ''viridius'', but in Italy there are males that are non-functional. Different subspecies of ''Lepidurus apus'' have different methods of fertilisation, some by a male, some by hermaphroditic individuals. Cysts average 0.447 mm in diameter, and have been found at concentrations of 250 per 100 cm2. They are laid on gravel in the middle of ditches or ponds, to avoid (it is speculated) large animals such as sheep transporting the cysts onto land. The cysts can survive drought and sub-zero temperatures, and can even synthesise haemoglobin if there is a lack of oxygen. As the pond dries out in the summer, the cysts will lie dormant until immersed in water. Light is an important factor in hatching: experiments showed no cysts hatched in darkness, some hatched after 10 mins of bright light, and all hatched in continuous light. They hatched between 10 °C and 24 °C, though the optimum was 16 °C and 20 °C, and even then hatch rates did not exceed 60%. Hatching often happens after winter rainfall forms temporary ponds. Larvae feed and rapidly grow to maturity, in as little as 4 weeks in optimum conditions in the warmer summer and spring months. ''Lepidurus apus'' has been found globally in remote areas with no waterways to transport individuals. In dry conditions, the dust-like resting eggs are easily distributed by wind. ''Lepidurus apus'' eggs are also though to be distributed by water, people, wildlife, and migratory birds.


Diet

''Lepidurus apus'' is omnivorous, feeding on both plant matter, mostly floating detritus, and small aquatic invertebrates such as
Branchinecta ''Branchinecta'' is a genus of crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipod ...
and
Daphnia ''Daphnia'' is a genus of small planktonic crustaceans, in length. ''Daphnia'' are members of the order Anomopoda, and are one of the several small aquatic crustaceans commonly called water fleas because their saltatory swimming style resembl ...
. It swims along the bottom of ponds, stirring up the substrate as it forages. The genus
Lepidurus ''Lepidurus'' is a genus of small crustaceans in the order Notostraca (tadpole shrimp). It is the larger of the two extant genera of the tadpole shrimps, the other being ''Triops''. They are commonly found in vernal pools and survive dry periods ...
also feeds upon algae, myxozoa, bacteria and fung''i.''


Predators and parasites

Predators of ''Lepidurus'' include small wading birds such as
sandpiper Sandpipers are a large family, Scolopacidae, of waders. They include many species called sandpipers, as well as those called by names such as curlew and snipe. The majority of these species eat small invertebrates picked out of the mud or soil. ...
s or
stint A stint is one of several very small waders in the paraphyletic ''"Calidris"'' assemblage – often separated in ''Erolia'' – which in North America are known as peeps. They are scolopacid waders much similar in ecomorphology to the ...
, larger waterfowl like
duck Duck is the common name for numerous species of waterfowl in the family Anatidae. Ducks are generally smaller and shorter-necked than swans and geese, which are members of the same family. Divided among several subfamilies, they are a form t ...
s and
swan Swans are birds of the family (biology), family Anatidae within the genus ''Cygnus''. The swans' closest relatives include the goose, geese and ducks. Swans are grouped with the closely related geese in the subfamily Anserinae where they form t ...
s, and, in some ponds, fishes. Migrating birds feed from temporary pools, as well as itinerant birds already found in the area. The rapid abundance of ''Lepidurus apus'' and other small invertebrates in pools often results in an increase in bird numbers in the area. ''Nosema lepiduri'' is a microsporidian parasite found in water bodies less than 15 cm deep that internally parasitises ''Lepidurus'' with spores, in some cases killing the host. Infected ''Lepidurus'' have a milky white colouration on the legs and carapace due to internal infection.


Conservation status

The subspecies ''Lepidurus apus viridis'' has been classified by the New Zealand
Department of Conservation An environmental ministry is a national or subnational government agency politically responsible for the environment and/or natural resources. Various other names are commonly used to identify such agencies, such as Ministry of the Environment ...
as Nationally Endangered under the
New Zealand Threat Classification System The New Zealand Threat Classification System is used by the Department of Conservation to assess conservation priorities of species in New Zealand. The system was developed because the IUCN Red List, a similar conservation status system, had some ...
.


References


External links

* The tadpole shrimp was discussed on RadioNZ ''Critter of the Week''
1 April 2016
{{Taxonbar, from=Q1317340 Notostraca Taxa named by Carl Linnaeus Cosmopolitan arthropods Crustaceans described in 1758