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''Lepidocaris rhyniensis'' is an extinct species 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, amphipods and mantis shrimp. The crustacean group ...
. It is the only species known from the order Lipostraca, and is the only abundant animal in the
Pragian The Pragian is one of three faunal stages in the Early Devonian Epoch. It lasted from 410.8 ± 2.8 million years ago to 407.6 ± 2.8 million years ago. It was preceded by the Lochkovian Stage and followed by the Emsian Stage. The most important La ...
-aged
Rhynie chert The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness (a Lagerstätte). It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is located ...
deposits. It resembles modern
Anostraca Anostraca is one of the four orders of crustaceans in the class Branchiopoda; its members are referred to as fairy shrimp. They live in vernal pools and hypersaline lakes across the world, and they have even been found in deserts, ice-covered m ...
, to which it is probably closely related, although its relationships to other orders remain unclear. The body is long, with 23 body segments and 19 pairs of
appendage An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including anten ...
s, but no
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 ...
. It occurred chiefly among
charophyte Charophyta () is a group of freshwater green algae, called charophytes (), sometimes treated as a phylum, division, yet also as a superdivision or an unranked clade. The terrestrial plants, the Embryophyta emerged within Charophyta, possibly fro ...
s, probably in alkaline temporary pools.


Biostratigraphy and taxonomy

All the known specimens of ''Lepidocaris rhyniensis'' have been excavated from the
Rhynie chert The Rhynie chert is a Lower Devonian sedimentary deposit exhibiting extraordinary fossil detail or completeness (a Lagerstätte). It is exposed near the village of Rhynie, Aberdeenshire, Scotland; a second unit, the Windyfield chert, is located ...
deposits in
Aberdeenshire Aberdeenshire ( sco, Aiberdeenshire; gd, Siorrachd Obar Dheathain) is one of the 32 Subdivisions of Scotland#council areas of Scotland, council areas of Scotland. It takes its name from the County of Aberdeen which has substantially differe ...
,
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to the ...
, which is a famous ', or site of exceptional preservation. ''Lepidocaris'' is the only abundant animal in the deposits, and is likely to be responsible for many of the frequent
coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is de ...
s found in the rocks. ''Lepidocaris'' was first described by D. J. Scourfield in a 1926 paper in the ''
Philosophical Transactions of the Royal Society B ''Philosophical Transactions of the Royal Society B: Biological Sciences'' is a biweekly peer-reviewed scientific journal published by the Royal Society. The editor-in-chief is John Pickett (Cardiff University). Overview Each issue covers a spe ...
''. Scourfield could not accommodate his new genus in the same
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 ...
as its closest relatives – the
Anostraca Anostraca is one of the four orders of crustaceans in the class Branchiopoda; its members are referred to as fairy shrimp. They live in vernal pools and hypersaline lakes across the world, and they have even been found in deserts, ice-covered m ...
– so he erected a new
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
and order for ''Lepidocaris'' alone: Lepidocarididae and Lipostraca, respectively. Until 2003, when '' Castracollis'' was described, ''Lepidocaris'' was the only crustacean known from the Rhynie chert.


Description

''Lepidocaris rhyniensis'' is a segmented animal with 23 body segments. The whole body measures around long. The head has two pairs of antennae, the second of which is used for swimming. As in Anostraca, there is no
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 ...
. There are eleven pairs of
appendages An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body. In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including anten ...
on the thorax and abdomen, of which the first three pairs are ''phyllopodia'', or leaf-like limbs, as seen in other branchiopods such as ''
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 ...
'', while the last eight pairs are similar to the swimming limbs of
copepod Copepods (; meaning "oar-feet") are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (inhabiting sea waters), some are benthos, benthic (living on the ocean floor) ...
s. The tail ends in a pair of caudal furcae.


Ecology

''Lepidocaris'' is one of the earliest preserved
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 ...
crustaceans. It is frequently found in association with the
charophyte Charophyta () is a group of freshwater green algae, called charophytes (), sometimes treated as a phylum, division, yet also as a superdivision or an unranked clade. The terrestrial plants, the Embryophyta emerged within Charophyta, possibly fro ...
'' Palaeonitella'' (
Characeae Characeae is a family of freshwater green algae in the order Charales, commonly known as stoneworts. They are also known as brittleworts or skunkweed, from the fragility of their lime-encrusted stems, and from the foul odor these produce when step ...
); if the ecology of ''Palaeonitella'' resembled that of its modern relatives, the water would have been
alkaline In chemistry, an alkali (; from ar, القلوي, al-qaly, lit=ashes of the saltwort) is a base (chemistry), basic, ionic compound, ionic salt (chemistry), salt of an alkali metal or an alkaline earth metal. An alkali can also be defined as ...
. Similarly, ''Lepidocaris'' is thought to have had a similar ecology to extant members of the
Anostraca Anostraca is one of the four orders of crustaceans in the class Branchiopoda; its members are referred to as fairy shrimp. They live in vernal pools and hypersaline lakes across the world, and they have even been found in deserts, ice-covered m ...
and the
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 ...
such as ''
Artemia ''Artemia'' is a genus of aquatic crustaceans also known as brine shrimp. It is the only genus in the family Artemiidae. The first historical record of the existence of ''Artemia'' dates back to the first half of the 10th century AD from Urmia La ...
'' and ''Triops'', inhabiting shallow, temporary pools.


Relationships

The phylogenetic position of ''Lepidocaris'' in relation to other orders of crustaceans is uncertain. In his original description of the species, Scourfield noted that ''Lepidocaris'' could not be accommodated in the existing order
Anostraca Anostraca is one of the four orders of crustaceans in the class Branchiopoda; its members are referred to as fairy shrimp. They live in vernal pools and hypersaline lakes across the world, and they have even been found in deserts, ice-covered m ...
, and even suggested that a position outside the
Branchiopoda Branchiopoda is a class of crustaceans. It comprises fairy shrimp, clam shrimp, Diplostraca (or Cladocera), Notostraca and the Devonian ''Lepidocaris''. They are mostly small, freshwater animals that feed on plankton and detritus. Description M ...
was not unthinkable. In 1986,
Frederick Schram Frederick Robert Schram (born August 11, 1943, in Chicago, Illinois, Chicago, Illinois) is an Americans, American palaeontologist and carcinologist. He received his Bachelor of Science, B.S. in biology from Loyola University Chicago in 1965, and ...
considered Lipostraca to be the
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
to Brachypoda, with the two orders together making up the
Cephalocarida The Cephalocarida are a class in the subphylum Crustacea comprising only 12 benthic species. They were discovered in 1955 by Howard L. Sanders, and are commonly referred to as horseshoe shrimp. They have been grouped together with the Remiped ...
. In 1997, Dieter Walossek considered ''Lepidocaris'' and '' Rehbachiella'' to be
stem-group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
anostracans, outside the extant Euanostraca. In 2001, Schram and Koenemann considered ''Lepidocaris'' and ''Rehbachiella'' to be stem-group lineages basal to the whole
Branchiopoda Branchiopoda is a class of crustaceans. It comprises fairy shrimp, clam shrimp, Diplostraca (or Cladocera), Notostraca and the Devonian ''Lepidocaris''. They are mostly small, freshwater animals that feed on plankton and detritus. Description M ...
.


References


Further reading

* {{Taxonbar, from=Q6527441 Branchiopoda genera Prehistoric crustacean genera Monotypic arthropod genera Devonian animals of Europe Fossil taxa described in 1926