| Proetida Temporal range: | |
|---|---|
 | |
| Aulacopleura konincki , Kosov u Berouna, Czech Republic | |
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | † Trilobita |
| Order: | † Proetida Fortey & Owens, 1975 |
| Superfamilies | |
| |
Proetida is an order of trilobite that lived from the Ordovician to the Permian. It was the last surviving order of trilobite, dying out in the Permian-Triassic extinction event. [1]
These typically small trilobites resemble those of the order Ptychopariida, from which the new order Proetida was separated in 1975 by Fortey and Owens. Like the order Phacopida, the proetids have exoskeletons that sometime have pits or small tubercles, especially on the glabella (middle portion of the head). Because of their resemblance to the Ptychopariida in some features, the proetids are included in the subclass Librostoma.
Unlike the trilobites of the phacopid suborder Phacopina, whose eyes are schizochroal, the proetids have the more common holochroal eyes. These eyes are characterized by close packing of biconvex lenses beneath a single corneal layer that covers all of the lenses. Each lens is generally hexagonal in outline and in direct contact with the others. They range in number from one to more than 15,000 per eye. Eyes are usually large, and because the individual lenses are hard to make out, they look smooth and sometimes bead-like.
The thorax of proetids was made up of anywhere between 8–22 segments, but most commonly 10. Many also extend the backcorners of the headshield into so-called genal spines. These two features can aid in distinguishing proetids from some phacopid trilobites in the suborder Phacopina, to which they can be very similar.
Opinions about the composition of and the affinities within the proetids, and to other trilobites, have been very divergent over time. In 2011 it was suggested to retain in Proetida only the families Proetidae and Tropidocoryphidae. The remainder of the families should be combined in a new proposed order, Aulacopleurida (Adrian, 2011), that would consist of the families Aulacopleuridae, Brachymetopidae, Dimeropygidae, Rorringtoniidae, Scharyiidae, Bathyuridae, Telephinidae, Holotrachelidae and Hystricuridae (considered Proetida before), combined with the Ptychopariid families Alokistocaridae, Crepicephalidae, Ehmaniellidae, Marjumiidae, Solenopleuridae and Tricrepicephalidae.
The reasoning for this proposed split is based on differences in early larval stages. While the remaining Proetida taxa have globular larvae very unlike the adult form, the Aulacopleurids have adultlike larvae with paired spines. [2] Others observe that globular non-adult larvae also occur in some taxa within the proposed order Aulacopleurida. More recently phylogenetic analysis of both larval and adult characters suggests the proetids as earlier understood probably are monophyletic. Two larval characters are unique to all Proetida; the first is that the eye develops on the side of the headshield, not at the front, and the second is a forwardly tapering glabella that is distanced from the rim of the headshield.
The analysis identifies the taxa Asaphida, Olenina and Phacopida (including the Holotrachelidae) as sister groups. The earliest branch in Proetida is the family Hystricuridae. This is followed by a branch that consists of the families Dimeropygidae and Toernquistiidae. At the third node, the superfamily Aulacopleuroidea (consisting of Aulacopleuridae and Brachymetopidae) split off. The fourth branch is the family Scharyiidae. The fifth branch consists of the families Roringtoniidae and Tropidocoryphidae. The sixth node combines a restricted Bathyuridae split off from Bathyurella with the family Proetidae (including Phillipsiidae, which, according to Lamsdell, has been demoted to the subfamily Phillipsiinae). [3]
The following superfamilies, families and genera are recognized: [4]
Family Aulacopleuridae
Family Brachymetopidae
Family Rorringtoniidae
Family Bathyuridae
Family Dimeropygidae (including Celmidae)
Family Holotrachelidae
Family Hystricuridae
Family Raymondinidae? (including Glaphuridae) order placement considered uncertain by Adrain, 2011 [5]
Family Telephinidae
Family Toernquistiidae
Family Phillipsiidae
Family Proetidae
Family Tropidocoryphidae
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 trilobites 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.
Redlichiida is an order of trilobites, a group of extinct marine arthropods. Species assigned to the order Redlichiida are among the first trilobites to appear in the fossil record, about halfway during the Lower Cambrian. Due to the difficulty to relate sediments in different areas, there remains some discussion, but among the earliest are Fallotaspis, and Lemdadella, both belonging to this order. The first representatives of the orders Corynexochida and Ptychopariida also appear very early on and may prove to be even earlier than any redlichiid species. In terms of anatomical comparison, the earliest redlichiid species are probably ancestral to all other trilobite orders and share many primitive characters. The last redlichiid trilobites died out before the end of the Middle Cambrian.
Phacopida ("lens-face") is an order of trilobites that lived from the Late Cambrian to the Late Devonian. It is made up of a morphologically diverse assemblage of taxa in three related suborders.
Ptychopariida is a large, heterogeneous order of trilobite containing some of the most primitive species known. The earliest species occurred in the second half of the Lower Cambrian, and the last species did not survive the Ordovician–Silurian extinction event.
The Phacopina comprise a suborder of the trilobite order Phacopida. Species belonging to the Phacopina lived from the Lower Ordovician (Tremadocian) through the end of the Upper Devonian (Famennian). The one unique feature that distinguishes Phacopina from all other trilobites are the very large, separately set lenses without a common cornea of the compound eye.
Phacops is a genus of trilobites in the order Phacopida, family Phacopidae, that lived in Europe, northwestern Africa, North and South America and China from the Late Ordovician until the very end of the Devonian, with a broader time range described from the Late Ordovician. It was a rounded animal, with a globose head and large eyes, and probably fed on detritus. Phacops is often found rolled up ("volvation"), a biological defense mechanism that is widespread among smaller trilobites but further perfected in this genus.
Olenellina is a suborder of the order Redlichiida of trilobites that occurs about halfway during the Lower Cambrian, at the start of the stage called the Atdabanian. Olenellina are arguably the earliest trilobites in the fossil record as members of Redlichiina, although Ptychopariida and Eodiscina follow soon after. The suborder died out when the Lower Cambrian passed into the Middle Cambrian, at the end of the stage called Toyonian. A feature uniting the Olenellina is the lack of rupture lines in the headshield, which in other trilobites assist the periodic moulting, associated with arthropod growth. Some derived trilobites have lost facial sutures again, but all of these are blind, while all Olenellina have eyes.
Paciphacops is a genus of trilobites within order Phacopida, suborder Phacopina. This genus is found primarily in the United States and Australia and is easily mistaken for the genera Phacops and Kainops, which are also popular among collectors. One major difference between Paciphacops and Phacops is that the central raised area of the headshield extends beyond the headshield's anterior margin. A major difference between Paciphacops and Kainops is the greater number of eye facets in Kainops. The skin of the visual surface in Paciphacops is thickened and bulged compared to the edge of each lens.
Ductina is a genus of extinct, small to average sized, eyeless phacopid trilobite, that lived during the Devonian.
Chotecops is a genus of trilobites from the order Phacopida, suborder Phacopina, family Phacopidae. It was initially erected as a subgenus of Phacops but some later authors thought it distinctive enough to raise its status. Species assigned to this genus occur between the Emsian and the Famennian. Chotecops is the most abundant trilobite in the Hunsrück Slate and due to the excellent preservation, often soft tissue such as antennae and legs have been preserved as a thin sheet of pyrite.
Acastoidea is a superfamily of trilobites from the order Phacopida, suborder Phacopina. This superfamily is divided into two families, Acastidae and Calmoniidae. This superfamily is distinguishable from the Phacopidae in that eyes are closer to the glabella and that the glabella has lobes, unlike the genera in Phacopidae. Acastoidea was first proposed by T.J.A. Reijers in 1972.
Proetidae is a family of proetid trilobites. The first species appeared in the Upper Ordovician, and the last genera survived until the Middle Permian. However, if the closely related family Phillipsiidae is actually a subfamily of Proetidae, then the proetids of Proetidae survive until the end of the Permian, where the last perish during the Permian–Triassic extinction event.
Phillipsiidae is a family of proetid trilobites, the various genera of which comprise some of the last of the trilobites, with a range that extended from the Kinderhookian epoch of the Lower Mississippian, to the end of Changhsingian age at Permian-Triassic extinction event in the latest Permian period.
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.
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. 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.
Viaphacops is a genus of trilobites in the order Phacopida, family Phacopidae, that lived during the Middle Devonian, and is known from North and South America, Asia.
Librostoma is a subclass of trilobites defined by having a natant hypostome, which is a hypostome that is free from the anterior doublure and aligned with the anterior of the glabella, this is unlike a conterminant hypostome, which is attached to the exoskeleton.
Gerastos is a genus of proetid trilobite in the family Proetidae that lived between the Pragian and Eifelian of the Lower-Middle Devonian, spanning approximately 21 million years.
Anisopyge is an extinct genus of trilobite belonging to the order Proetida and family Phillipsiidae. Specimens have been found in Permian beds in North and Central America.
Proetoidea is a superfamily of trilobites in the order Proetida.
