Cyclida
Encyclopedia
Cyclida is an order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 of fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

 arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

s that lived from the Carboniferous
Carboniferous
The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...

 to the Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to  Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...

. Their classification is uncertain, but they are generally treated as a group of maxillopod
Maxillopoda
Maxillopoda is a diverse class of crustaceans including barnacles, copepods and a number of related animals. It does not appear to be a monophyletic group, and no single character unites all the members.-Description:...

 crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...

s.

Description

Cycloids have a "striking" resemblance to crab
Crab
True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...

s, and are thought to have inhabited a similar ecological niche
Ecological niche
In ecology, a niche is a term describing the relational position of a species or population in its ecosystem to each other; e.g. a dolphin could potentially be in another ecological niche from one that travels in a different pod if the members of these pods utilize significantly different food...

, and to have been driven to extinction when crabs became widespread and diverse. The largest members are over 6 centimetres (2.4 in) across the carapace
Carapace
A carapace is a dorsal 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 tortoises, the underside is called the plastron.-Crustaceans:In crustaceans, the...

. Their gill
Gill
A gill is a respiratory organ found in many aquatic organisms that extracts dissolved oxygen from water, afterward excreting carbon dioxide. The gills of some species such as hermit crabs have adapted to allow respiration on land provided they are kept moist...

s are often preserved in three dimensions, and do not resemble those of other crustaceans. Cycloid taxa differ in the number of walking legs, in the form of the mouthparts and in other significant ways.

Affinities

There is considerable debate about the placement of cycloids within the Arthropod
Arthropod
An arthropod is an invertebrate animal having an exoskeleton , a segmented body, and jointed appendages. Arthropods are members of the phylum Arthropoda , and include the insects, arachnids, crustaceans, and others...

a. While they are generally considered to be crustacean
Crustacean
Crustaceans form a very large group of arthropods, usually treated as a subphylum, which includes such familiar animals as crabs, lobsters, crayfish, shrimp, krill and barnacles. The 50,000 described species range in size from Stygotantulus stocki at , to the Japanese spider crab with a leg span...

s of some kind, doubts have been expressed about the homology
Homology (biology)
Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...

 of cycloids' respiratory structures with those of other crustaceans, and parallels drawn instead with chelicerates.

The first description of a cycloid was in the 1836 treatise Illustrations of the Geology of Yorkshire by John Phillips
John Phillips (geologist)
John Phillips FRS was an English geologist.- Life and work :Philips was born at Marden in Wiltshire...

, where Phillips described "Agnostus ? radialis" among the trilobite
Trilobite
Trilobites are a well-known fossil group of extinct marine arthropods that form the class Trilobita. The first appearance of trilobites in the fossil record defines the base of the Atdabanian stage of the Early Cambrian period , and they flourished throughout the lower Paleozoic era before...

s, with the text "ribs radiating, with acute puncta; abdomen mucronate". In 1838, Hermann von Meyer
Christian Erich Hermann von Meyer
Christian Erich Hermann von Meyer was a German palaeontologist.He was born at Frankfurt am Main.In 1832 von Meyer issued a work entitled Palaeologica, and in course of time he published a series of memoirs on various fossil organic remains: molluscs, crustaceans, fishes and higher vertebrata,...

 described a species of trilobite, albeit in the genus Limulus, and later transferred it to a new genus, Halicyne, recognising that it was something different. In 1841, Laurent-Guillaume de Koninck
Laurent-Guillaume de Koninck
Laurent-Guillaume de Koninck was a Belgian palaeontologist and chemist, born at Leuven.He studied medicine in the university of his native town, and in 1831 he became assistant in the chemical schools. He pursued the study of chemistry in Paris, Berlin and Gießen, and was subsequently engaged in...

 transferred Phillips' species to a new genus, Cyclus, away from the trilobites, although he later described a second species of Cyclus which was later recognised as the hypostome
Hypostome
A hypostome , is a calcified harpoon-like structure near the mouth area of certain parasitic arthropods including ticks and mites , that allows them to anchor themselves firmly in place on a host mammal while sucking blood...

 of a trilobite. Cycloids were later considered to be members of the Xiphosura
Xiphosura
Xiphosura is an order of marine chelicerates which includes a large number of extinct lineages and only four recent species in the family Limulidae, which include the horseshoe crabs...

, true crabs
Crab
True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...

, and branchiura
Branchiura
The family Argulidae contains the carp lice or fish lice – a group of parasitic crustaceans of uncertain position within the Maxillopoda. Although they are thought to be primitive forms, they have no fossil record...

ns.

In an unpublished dissertation, Neil D. L. Clark proposed in 1989 that cycloids were copepod
Copepod
Copepods are a group of small crustaceans found in the sea and nearly every freshwater habitat. Some species are planktonic , some are benthic , and some continental species may live in limno-terrestrial habitats and other wet terrestrial places, such as swamps, under leaf fall in wet forests,...

s. In 1997, Frederick Schram
Frederick Schram
Frederick Robert Schram is an American palaeontologist and carcinologist. He received his B.S. in biology from Loyola University Chicago in 1965, and a Ph.D...

 and his co-authors classified them as the sister group to copepods, within the Maxillopoda
Maxillopoda
Maxillopoda is a diverse class of crustaceans including barnacles, copepods and a number of related animals. It does not appear to be a monophyletic group, and no single character unites all the members.-Description:...

, and in 2008, Jerzy Dzik placed them as an order within the maxillopod suborder Branchiura
Branchiura
The family Argulidae contains the carp lice or fish lice – a group of parasitic crustaceans of uncertain position within the Maxillopoda. Although they are thought to be primitive forms, they have no fossil record...

, which previously contained only the modern fish lice.

Taxa and stratigraphy

Cycloids are known from deposits ranging from Carboniferous
Carboniferous
The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...

 (Cyclus spp.) to Maastrichtian
Maastrichtian
The Maastrichtian is, in the ICS' geologic timescale, the latest age or upper stage of the Late Cretaceous epoch or Upper Cretaceous series, the Cretaceous period or system, and of the Mesozoic era or erathem. It spanned from 70.6 ± 0.6 Ma to 65.5 ± 0.3 Ma...

 (Maastrichtocaris rostrata). They are one of only three groups of "generally Palaeozoic" arthropods to survive the Permian–Triassic extinction event, the others being the freshwater-living Euthycarcinoida and the marine Thylacocephala
Thylacocephala
The Thylacocephala are a unique group of extinct arthropods, with possible crustacean affinities. As a class they have a short research history, having been erected in the early 1980s. They typically possess a large, laterally flattened carapace that encompasses the entire body...

.

The order Cyclida contains 15 genera. In approximate age order, they are:
  • Cyclus contains a number of species found in Carboniferous
    Carboniferous
    The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...

     deposits.
  • Schramine comprises three species formerly included in Halicyne of mid to late Carboniferous age.
  • Americlus contains several species from the Carboniferous, including A. americanus (formerly Cyclus americanus), a well known species from Mazon Creek.
  • Apionicon apioides is found in Pennsylvanian
    Pennsylvanian
    The Pennsylvanian is, in the ICS geologic timescale, the younger of two subperiods of the Carboniferous Period. It lasted from roughly . As with most other geochronologic units, the rock beds that define the Pennsylvanian are well identified, but the exact date of the start and end are uncertain...

     shale
    Shale
    Shale is a fine-grained, clastic sedimentary rock composed of mud that is a mix of flakes of clay minerals and tiny fragments of other minerals, especially quartz and calcite. The ratio of clay to other minerals is variable. Shale is characterized by breaks along thin laminae or parallel layering...

    s in Illinois
    Illinois
    Illinois is the fifth-most populous state of the United States of America, and is often noted for being a microcosm of the entire country. With Chicago in the northeast, small industrial cities and great agricultural productivity in central and northern Illinois, and natural resources like coal,...

    , including Mazon Creek.
  • Hemitrochiscus paradoxus is found in Permian
    Permian
    The PermianThe term "Permian" was introduced into geology in 1841 by Sir Sir R. I. Murchison, president of the Geological Society of London, who identified typical strata in extensive Russian explorations undertaken with Edouard de Verneuil; Murchison asserted in 1841 that he named his "Permian...

     dolomite
    Dolomite
    Dolomite is a carbonate mineral composed of calcium magnesium carbonate CaMg2. The term is also used to describe the sedimentary carbonate rock dolostone....

     near Wünschendorf/Elster
    Wünschendorf/Elster
    Wünschendorf/Elster is a municipality in the district of Greiz, in Thuringia, Germany....

     (Germany
    Germany
    Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...

    ).
  • Oonocarcinus insignis and Paraprosopon reussi are found in the Permian limestone
    Limestone
    Limestone is a sedimentary rock composed largely of the minerals calcite and aragonite, which are different crystal forms of calcium carbonate . Many limestones are composed from skeletal fragments of marine organisms such as coral or foraminifera....

    s of the Sosio valley on Sicily
    Sicily
    Sicily is a region of Italy, and is the largest island in the Mediterranean Sea. Along with the surrounding minor islands, it constitutes an autonomous region of Italy, the Regione Autonoma Siciliana Sicily has a rich and unique culture, especially with regard to the arts, music, literature,...

     (Italy
    Italy
    Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

    ).
  • Halicyne contains several species from the Triassic.
  • Carcinaspides pustulosus is found in limestone rocks in the Isar
    Isar
    The Isar is a river in Tyrol, Austria and Bavaria, Germany. Its source is in the Karwendel range of the Alps in Tyrol; it enters Germany near Mittenwald, and flows through Bad Tölz, Munich, and Landshut before reaching the Danube near Deggendorf. At 295 km in length, it is the fourth largest river...

     valley of Bavaria
    Bavaria
    Bavaria, formally the Free State of Bavaria is a state of Germany, located in the southeast of Germany. With an area of , it is the largest state by area, forming almost 20% of the total land area of Germany...

     (Germany
    Germany
    Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...

    ) and is probably of Mid
    Middle Triassic
    In the geologic timescale, the Middle Triassic is the second of three epochs of the Triassic period or the middle of three series in which the Triassic system is divided. It spans the time between 245 ± 1.5 Ma and 228 ± 2 Ma...

     to Late Triassic
    Late Triassic
    The Late Triassic is in the geologic timescale the third and final of three epochs of the Triassic period. The corresponding series is known as the Upper Triassic. In the past it was sometimes called the Keuper, after a German lithostratigraphic group that has a roughly corresponding age...

     age.
  • Opolanka decorosa is found in Late Triassic
    Late Triassic
    The Late Triassic is in the geologic timescale the third and final of three epochs of the Triassic period. The corresponding series is known as the Upper Triassic. In the past it was sometimes called the Keuper, after a German lithostratigraphic group that has a roughly corresponding age...

     rocks at Krasiejów
    Krasiejów
    Krasiejów is a village in the administrative district of Gmina Ozimek, within Opole County, Opole Voivodeship, in south-western Poland...

     in Poland
    Poland
    Poland , officially the Republic of Poland , is a country in Central Europe bordered by Germany to the west; the Czech Republic and Slovakia to the south; Ukraine, Belarus and Lithuania to the east; and the Baltic Sea and Kaliningrad Oblast, a Russian exclave, to the north...

    .
  • Cyclocarcinoides serratus and Mesoprosopon triasinum are found in the grey Norian
    Norian
    The Norian is a division of the Triassic geological period. It has the rank of an age or stage . The Norian lasted from 216.5 ± 2.0 to 203.6 ± 1.5 million years ago. It was preceded by the Carnian and succeeded by the Rhaetian.-Stratigraphic definitions:The Norian was named after the Noric Alps in...

     (late Triassic) limestones of Bad Ischl
    Bad Ischl
    Bad Ischl is a spa town in Austria. It lies in the southern part of Upper Austria, at the Traun River in the centre of the Salzkammergut region. The town consists of the Katastralgemeinden Ahorn, Bad Ischl, Haiden, Jainzen, Kaltenbach, Lauffen, Lindau, Pfandl, Perneck, Reiterndorf and Rettenbach...

     in the Austria
    Austria
    Austria , officially the Republic of Austria , is a landlocked country of roughly 8.4 million people in Central Europe. It is bordered by the Czech Republic and Germany to the north, Slovakia and Hungary to the east, Slovenia and Italy to the south, and Switzerland and Liechtenstein to the...

    n Salzkammergut
    Salzkammergut
    The Salzkammergut is a resort area located in Austria. It stretches from City of Salzburg to the Dachstein mountain range, spanning the federal states of Upper Austria , Salzburg , and Styria . The main river of the region is the Traun, a tributary of the Danube...

    .
  • Juracyclus posidoniae is found in the Jurassic
    Jurassic
    The Jurassic is a geologic period and system that extends from about Mya to  Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...

      near Tübingen
    Tübingen
    Tübingen is a traditional university town in central Baden-Württemberg, Germany. It is situated south of the state capital, Stuttgart, on a ridge between the Neckar and Ammer rivers.-Geography:...

    , southern Germany
    Germany
    Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...

    .
  • Alsasuacaris nostradamus was found in late Albian
    Albian
    The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous epoch/series. Its approximate time range is 112.0 ± 1.0 Ma to 99.6 ± 0.9 Ma...

     to early Cenomanian
    Cenomanian
    The Cenomanian is, in the ICS' geological timescale the oldest or earliest age of the Late Cretaceous epoch or the lowest stage of the Upper Cretaceous series. An age is a unit of geochronology: it is a unit of time; the stage is a unit in the stratigraphic column deposited during the corresponding...

     (Cretaceous
    Cretaceous
    The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

    ) rocks near Alsasua (Navarre
    Navarre
    Navarre , officially the Chartered Community of Navarre is an autonomous community in northern Spain, bordering the Basque Country, La Rioja, and Aragon in Spain and Aquitaine in France...

    , Spain
    Spain
    Spain , officially the Kingdom of Spain languages]] under the European Charter for Regional or Minority Languages. In each of these, Spain's official name is as follows:;;;;;;), is a country and member state of the European Union located in southwestern Europe on the Iberian Peninsula...

    ).
  • Maastrichtocaris rostrata is found in Maastrichtian
    Maastrichtian
    The Maastrichtian is, in the ICS' geologic timescale, the latest age or upper stage of the Late Cretaceous epoch or Upper Cretaceous series, the Cretaceous period or system, and of the Mesozoic era or erathem. It spanned from 70.6 ± 0.6 Ma to 65.5 ± 0.3 Ma...

     (late Cretaceous) rocks from that period's type locality
    Type locality (geology)
    Type locality , also called type area or type locale, is the where a particular rock type, stratigraphic unit, fossil or mineral species is first identified....

     in the southern Netherlands
    Netherlands
    The Netherlands is a constituent country of the Kingdom of the Netherlands, located mainly in North-West Europe and with several islands in the Caribbean. Mainland Netherlands borders the North Sea to the north and west, Belgium to the south, and Germany to the east, and shares maritime borders...

    .


Stagmacaris quenstedti, a species from the late Kimmeridgian
Kimmeridgian
In the geologic timescale, the Kimmeridgian is an age or stage in the Late or Upper Jurassic epoch or series. It spans the time between 155.7 ± 4 Ma and 150.8 ± 4 Ma . The Kimmeridgian follows the Oxfordian and precedes the Tithonian....

 of southern Germany
Germany
Germany , officially the Federal Republic of Germany , is a federal parliamentary republic in Europe. The country consists of 16 states while the capital and largest city is Berlin. Germany covers an area of 357,021 km2 and has a largely temperate seasonal climate...

, has been reinterpreted as part of the abdomen of a hermit crab
Hermit crab
Hermit crabs are decapod crustaceans of the superfamily Paguroidea. Most of the 1100 species possess an asymmetrical abdomen which is concealed in an empty gastropod shell that is carried around by the hermit crab.-Description:...

, rather than a cycloid.
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