Cyclestheriidae
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Clam shrimp are a group of bivalved branchiopod crustaceans that resemble the unrelated bivalved molluscs. They are extant and also known from the fossil record, from at least the Devonian period and perhaps before. They were originally classified in the former
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
Conchostraca, which later proved to be
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
and was subsumed into the superorder Diplostraca. Clam shrimp now make up three of the seven orders in Diplostraca, Cyclestherida, Laevicaudata, and Spinicaudata, in addition to the fossil family Leaiidae.


Characteristics

Both valves of the shell are held together by a strong closing
muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...
. The animals react to danger by contracting the muscle, so that the valves close tightly and the crustacean, as if dead, lies motionlessly at the bottom of the pool. In most species the head is dorsoventrally compressed. The sessile compound eyes are close together and located on the forehead; in the genus ''Cyclestheria'' they are truly fused. In front of them is a simple naupliar eye. The first pair of antennae is reduced and unsegmented. The second pair of antennae, however, is long and biramous. Both branches are covered with numerous bristles. The crustaceans swim primarily by swooping the antennae. In the common genus ''Lynceus'', which can open its spherical valves wide, the thoracic legs move in an oar-like manner along with the antennae. The number of segments constituting the thorax varies from 10 to 32, and the number of legs varies accordingly. They are similar in structure to the legs of
tadpole shrimp 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 ...
, and similarly, their size decreases from front to back. In females, the outer lobes of several middle legs are modified into long, upward-bending threadlike outgrowths, used to hold the eggs on the dorsal side of the body under the shell. However, the main functions of the thoracic legs are respiration and carrying food forward to the mouth. The gills are basically the outer lobes of all thoracic legs that are closest to the base of the leg. The legs are in constant movement, and the water between the valves of the
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 ...
is quickly renewed. The body ends in a large
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
ised telson, which is either laterally compressed and bears a pair of large hooks, or dorsoventrally compressed, with short hooks.


Reproduction and development


Reproduction

Clam shrimp have different reproductive strategies. For example, within the family
Limnadiidae Limnadiidae is a family of crustaceans in the order Spinicaudata that live in seasonal wetlands, inland saline pools and lakes. They are found on all the world's continents except Antarctica, and are distinguished from other families in the same ...
are found
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
(male-female), hermaphroditic (only hermaphrodites), and androdioecious (male-hermaphrodite) species.


Life cycle

The eggs are surrounded by a tough shell and can withstand drying out, freezing and other hostile conditions. In some species these eggs can hatch after as long as 7 years. When the egg arrives in a suitable pool, a larva hatches out at the nauplius stage (the nauplius stage is absent in Cyclestherida).How the cladoceran heterogonic life cycle evolved--insights from gamogenetic reproduction and direct development in Cyclestherida
/ref> Clam shrimp nauplii are distinguished by very small front antennae. At the second stage ( metanauplius), the larva develops the small shell. They develop very quickly. For instance, ''Cyzicus'' reaches sexual maturity in 19 days after hatching.


Taxonomy

Extant clam shrimp belong to three orders, divided into five families; some notable genera and
prehistoric Prehistory, also known as pre-literary history, is the period of human history between the use of the first stone tools by hominins 3.3 million years ago and the beginning of recorded history with the invention of writing systems. The use of ...
taxa are also listed: Spinicaudata Linder, 1945 * Cyzicidae Stebbing, 1910 ** '' Caenestheria'' ** '' Caenestheriella'' ** '' Cyzicus'' ** '' Eocyzicus'' * Eosestheriidae Zhang & Chen, 1976 ( fossil) ** '' Bairdestheria'' ** '' Menucoestheria'' Gallego & Covacevich, 1998 * Euestheridae Defretin, 1965 ( fossil) ** '' Laxitextella'' Kozur, 1982 * Leptestheriidae Daday, 1923 ** ''
Eoleptestheria ''Eoleptestheria ticinensis'' is a species of clam shrimp in the family Leptestheriidae. Although up to eight species have formerly been recognised in the genus ''Eoleptestheria'', they are more usually all considered part of one species with a c ...
'' ** '' Leptestheria'' ** '' Leptestheriella'' ** '' Maghrebestheria'' ** '' Sewellestheria'' *
Limnadiidae Limnadiidae is a family of crustaceans in the order Spinicaudata that live in seasonal wetlands, inland saline pools and lakes. They are found on all the world's continents except Antarctica, and are distinguished from other families in the same ...
Baird, 1849 (including Imnadiidae) ** ''
Eulimnadia ''Eulimnadia'' is a genus of branchiopods in the family Limnadiidae. There are about 13 described species in ''Eulimnadia''. Species * ''Eulimnadia agassizii'' Packard, 1874 * ''Eulimnadia antillarum'' (Baird, 1852) * ''Eulimnadia antlei'' Macki ...
'' ** ''
Imnadia ''Imnadia'' is a genus of conchostracans found only in Europe. It has occasionally been placed in a monotypic family, "Imnadiidae", but is more usually placed in the Limnadiidae. It contains the following species: *'' Imnadia cristata'' MarinĨe ...
'' ** '' Limnadia'' ** '' Limnadiopsis'' ** '' Limnadiopsidum'' ** '' Metalimnadia'' * Palaeolimnadiidae Tasch, 1956 ( fossil) ** '' Krasiestheria'' Olempska, 2004 (tentatively placed here) Cyclestherida Sars, 1899 * Cyclestheriidae Sars, 1899 ** '' Cyclestheria'' ** '' Paracyclestheria'' Laevicaudata Linder, 1945 * Lynceidae Baird, 1845 ** '' Lynceiopsis'' ** '' Lynceus'' ** '' Paralimnetis''


Geological history

Modern clam shrimp have little significance to humans. However, extinct species of these crustaceans are often studied by geologists. In
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...
deposits, generally poor in fossils, the well-preserved clam shrimp shells are found quite often. They help identify the age of the corresponding
strata In geology and related fields, a stratum ( : strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as ei ...
. During the past geological periods clam shrimp were apparently more numerous and diverse than they are now. 300 extinct species are known, and half as many living species. The oldest clam shrimp, such as ''
Asmussia murchisoniana ''Asmussia murchisoniana'' was a Devonian crustacean of the class Branchiopoda. Like other clam shrimp Clam shrimp are a group of bivalved branchiopod crustaceans that resemble the unrelated bivalved molluscs. They are extant and also known ...
'', were found in
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
deposits. Many extinct species, mostly Triassic specimens, once lived in marine environments, where no extant clam shrimp inhabit today.


References


External links




Introduction to the Branchiopoda
* * * {{Taxonbar, from=Q500917 Branchiopoda Extant Devonian first appearances Arthropod common names