Cyclopygidae Temporal range: | |
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Cyclopygid cephalon, probably Symphysops stevaninae, 24mm, lateral view | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | † Trilobita |
Order: | † Asaphida |
Superfamily: | † Cyclopygoidea |
Family: | † Cyclopygidae Raymond, 1925 |
Subfamilies | |
And see text for genera |
Cyclopygidae is a family of asaphid trilobites from the Ordovician. Cyclopygids had an extratropical distribution, and there is evidence that they lived in darker parts of the water column (around 175m deep). Cyclopygids are characterized by enlarged eyes, with a wide angle of view, both horizontal and vertical, reminiscent of the eyes of dragonflies. These typically touch the glabella directly on the side. Cyclopygids all lack genal spines, but Symphysops carries a forward directed frontal spine on the glabella. It is presumed that at least the members of the genus Pricyclopyge swam upside down and had bioluminescent organs on the third thorax segment. Cyclopygids had between 7 and 5 thorax segments, a wide and stout axis, and short side lobes (or pleurae).
The genera of Cyclopygidae are grouped into the following subfamilies:
The extinction ending the Ordovician was one of the most radical for life to have experienced, and the trilobites were heavily affected. Those with pelagic or deep water benthic life styles (such as species in Olenidae and Agnostida) died out. Also those trilobites having planktonic larvae became extinct, and these include most of the superfamilies in the order Asaphida, save for Trinucleoidea. A reduction in diversity already occurred before this major extinction, but many families persisted into the Hirnantian, and it is possible that they would quickly have been restored to their former diversity. The crisis that started the Silurian must have exceptionally severe, and was associated with low oxygen levels in the oceans after an ice age. [1]
Cyclopygids have particularly large eyes with a wide angle view, also vertically, that occupy most of the free cheeks, and the fixed cheeks absent or reduced to a very narrow strip at the sides of the glabella, and a zone between the both eyes. In the earliest cyclopygids ( Prospectatrix ) the eyes are less enlarged, [2] but in some later taxa, eyes are so big they have even fused. The most backward lobe of the glabella (the occipital ring) cannot be identified, except in the Ellipsotaphrinae subfamily. Further furrows crossing the glabella may be absent or are reduced to pairs of slight depressions. Genal spines are lacking. Cyclopygids have between 5 and 7 thorax segments. The pleurae become successively wider further back, making the thorax widest across the last segment. [1]
In pelagic trilobites, such as the species in the proetid family Telephinidae, and in Cyclopygidae, as in many extant pelagic crustaceans, the eyes are particularly large and have very wide angles of view, both horizontal and vertical. This is in stark contrast to contemporary benthic trilobites, that may have an extensive horizontal angle of view, but always have a limited vertical angle of view. [3]
In a few species of the genera Cyclopyge, Microparia, Ellipsotaphrus, Pricyclopyge and in Symphysops the eyes are merged in front of the head creating a visor. This development improves the sensitivity of the eye for objects that move relative to the eye, which might have been particularly useful under low-light conditions and when rapidly moving. The extant hyperiid amphipod Cystisoma also has such fused eyes. Monocular trilobites are always younger than closely related species with normal paired eyes, and is an example of a trend that occurred several times in parallel. Only in Pricyclopyge binodosa several stages in this development can be seen as a consecutive series of subspecies collected from successive zones in the late Arenig to the Llanvirn. Although the distance between the eyes varies within any one population of the earlier subspecies, the eyes only touch and merge in P. binodosa synophthalma. [1] [4]
Cyclopygids are absent from shallow water strata, such as alluvial and calcareous deposits. They are not found together with well-sighted benthic trilobite species or corals. They do occur with blind or nearly blind benthic trilobites, a typical adaptation to a lightless environment, and oceanic free-floating graptolites. Hence, cyclopygids are considered to have been confined to deeper water, swimming at the lower limit of the photic zone (or mesopelagic), [1] but still high above the benthic species they were deposited with. This is also evidenced by the presumed present of bioluminescent organs on the third thorax segment of Pricyclopyge, which also occur on the functional underside of extant mesopelagic species. This is why it is assumed Pricyclopyge may have swum upside down. [3] Very large, convex eyes and a narrow zone of thoracic pleurae are typical for all Cyclopygidae, and are indications of a pelagic lifestyle. The stout exoskeleton is consistent with rapid swimming and it is likely cyclopygids actively hunted zooplankton.
Agnostida are an order of extinct arthropods which have classically been seen as a group of highly modified trilobites, though some recent research has doubted this placement. Regardless, they appear to be close relatives as part of the Artiopoda. They are present in the Lower Cambrian fossil record along with trilobites from the Redlichiida, Corynexochida, and Ptychopariida orders, and were highly diverse throughout the Cambrian. Agnostidan diversity severely declined during the Cambrian-Ordovician transition, and the last agnostidans went extinct in the Late Ordovician.
Trilobites are extinct marine arthropods that form the class Trilobita. Trilobites form one of the earliest known groups of arthropods. 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 before slipping into a long decline, when, during the Devonian, all trilobite orders except the Proetida died out. The last extant trilobites finally disappeared in the mass extinction at the end of the Permian about 251.9 million years ago. Trilobites were among the most successful of all early animals, existing in oceans for almost 270 million years, with over 22,000 species having been described.
Phacopidae is a family of phacopid trilobites that ranges from the Lower Ordovician to the Upper Devonian, with representatives in all paleocontinents.
The Paradoxididae are a family of trilobites, a group of extinct marine arthropods. They occurred during the late Lower Cambrian (Toyonian) and disappeared at the end of the Middle Cambrian. Representatives of this family have been found in the paleocontinents of Avalonia, Baltica, and Gondwana, now Canada, USA, England, Wales, Morocco, Spain, Czech Republic, Poland, Russia, Mongolia, and Turkey. Species in this family can typically grow large to very large, are relatively flat, have an inverted egg-shaped outline, opisthoparian sutures, a glabella that in early genera has parallel sides and expands forward in later representatives, and approaches or reaches the frontal border. All species have an almost semicircular headshield with long backward-directed genal spines. The articulate middle part of the body consists of 15 to 21 segments ending in sickle-shaped spines that to the back curve increasingly further backwards. The tailshield is small.
Acadagnostus is a genus of trilobite from the Middle Cambrian, with 7 species currently recognized. The type species A. acadicus has the widest distribution known from any peronopsid and has been found in North America, Greenland, England, Western Europe, Eastern Europe, Central Asia, the Altai Mountains, the Siberian shield, China, and Australia.
Biceratops is an extinct genus of olenelloid redlichiid trilobites, of average size, with the largest specimen 8 centimetres or 3.1 inches long, not including the huge pleural spines of the 3rd segment of the thorax. It lived during the Toyonian stage, in what is today the South-Western United States. Biceratops can easily be distinguished from other members of Biceratopsidae by the absence of genal spines, in combination with effaced features of the raised axial area of the head shield, that is bordering the two horn-like projections that carry the eyes. Biceratops nevadensis is the only known species in this genus.
Conocoryphe is a genus of primarily eyeless trilobites belonging to the family Conocoryphidae. They lived during the Middle Cambrian period, about 505 million years ago. These arthropods lived on the sea bottom (epifaunal) and lived off dead particulate organic matter.
Eodiscina is trilobite suborder. The Eodiscina first developed near the end of the Lower Cambrian period and became extinct at the end of the Middle Cambrian. Species are tiny to small, and have a thorax of two or three segments. Eodiscina includes six families classified under one superfamily, Eodiscoidea.
Ogygiocarella Brongniart, 1822, is a genus of asaphid trilobites. It occurred during the Middle Ordovician.
Cyclopyge is a genus of small to average size trilobites that lived during the Ordovician. Like all members of the family Cyclopigidae, it has very large convex eyes, that cover most of the free cheeks, and in some species touch each other. The eyes almost touch the large glabella. The occipital ring has merged with the rest of the glabella. The glabella does not extend into a frontal thorn. The cephalon lacks genal spines. The 6 thorax segments have short pleurae. The pygidium is rather large, and often rather effaced. These are features that also occur in other Cyclopygidae, and are indications of a pelagic lifestyle.
Emmrichops is a genus of average size trilobite, assigned to the Cyclopygidae family, that lived during the Middle Ordovician (Llanvirn), and have been found in what are today the Czech Republic and in Wales. Like other cyclopygids, it has huge eyes, that occupy almost the entire free cheeks, six thorax segments and a wide tail shield. Like the other cyclopygids, Emmrichops probably lived hunting plankton in the water column. Only one species, E. planicephala, has been described sofar.
Symphysops is a genus of trilobites of average size, belonging to the Cyclopygidae family. It had a cosmopolitan distribution and lived from the Middle to the Upper Ordovician. It has been found in Canada, China, the Czech Republic (Bohemia), Iran, Ireland, Kazakhstan, Poland, Morocco, Spain, Scotland and Wales. The name Symphysops refers to the fused eyes, common to the species of this genus. Some (sub)species of the cyclopygid genera Cyclopyge and Pricyclopyge share this character, but Symphysops uniquely combines the merged eye with a frontal thorn on the head and the "lower eyelid".
Prospectatrix is a genus of trilobites of average size, that lived in the Lower Ordovician and is probably ancestral to the other genera of the Cyclopygidae family. Its eyes are only moderately enlarged and it has six or seven thorax segments.
Sagavia is a genus of trilobites that lived during the Middle and Upper Ordovician in what are now Northwest and Southeast China, North Kazakhstan and Wales. It is a typical cyclopygid that can be distinguished by its large but separate eyes, elongated glabella, five thorax segments and a pygidium with clearly defined axis and border.
Pricyclopyge is a genus of trilobites assigned to the family Cyclopygidae that occurs throughout the Ordovician. Pricyclopyge had an extratropical distribution, and there is evidence that it lived in darker parts of the water column. Pricyclopyge has huge eyes, an inverted pear-shaped glabella, six thorax segments, with on the 3rd two small discs. Pricyclopyge is known from what are today China, the Czech Republic, France, and the United Kingdom.
Carolinites is a genus of trilobite, assigned to the Telephinidae family, that occurs during the Lower and Middle Ordovician. Carolinites had a pantropical distribution, and there is evidence that it lived in upper parts of the water column. The free cheeks of Carolinites are largely covered by its huge eyes, except for the attachment of large genal spines that extend downward, backward and lateral and gradually curving further backward. The glabella is slightly bulbous, the occipital ring is well defined, but further transglabellar furrows are lacking. The thorax has 10 segments. The axis of the pygidium is highly vaulted, with a curved spine emerging almost perpendicular to the midline and ending parallel to it and a node on each of the other three segments. Carolinites is known from what are today Australia (Tasmania), Canada (Alberta), China, France, Spitsbergen, and the United States (Utah).
Raphiophoridae is a family of small to average-sized trilobites that first occurred at the start of the Ordovician and became extinct at the end of the Middle Silurian.
Pliomera is a genus of trilobites that lived during the Middle Ordovician on the paleocontinent Baltica, now Norway, Sweden, Estonia and the Russian Federation, and in Argentina. It can be recognized for its pentagonal glabella widest between the frontal corners, with an inverted V-shaped occipital ring. In front of the occipital furrow that crosses the entire glabella, two pairs of dead-ending furrows create three side lobes left and right. The front of the glabella also has three dead-ending furrows, a very short one on the midline and left and right a longer one, directed inward and slightly backward. The eyes are small and are not connected to the glabella by an eye ridge. The thorax and pygidium are very regularly divided into up to 23 rather narrow segments, without a furrow within each of the pleurae. The pleurae are clearly wider than the axis. The pygidium ends in downward pointing toothlike spines.
The Paradoxidoidea Hawle & Corda 1847, are a superfamily of trilobites, a group of extinct marine arthropods. They occurred during the late Lower Cambrian (Toyonian) and disappeared at the end of the Middle Cambrian.
Telephinidae is a family of pelagic trilobites with large wide-angle eyes, occupying most of the free cheeks, downward directed facial spines and 9-10 thorax segments. The family is known during the entire Ordovician and occurred in deep water around the globe.