Librostoma Temporal range: | |
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Asaphus lepidurus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | † Trilobita |
Subclass: | † Librostoma Fortey, 1990 [1] |
Orders | |
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Librostoma is a subclass of trilobites defined by having a natant hypostome, which is a hypostome (mineralized trilobite mouthpart) that is free from the anterior doublure and aligned with the anterior of the glabella, [4] this is unlike a conterminant hypostome, which is attached to the exoskeleton. [5]
They are a large and long lasting group, with some of the candidates for the first trilobites being librostomes in the order ptychopariida; and all post-Devonian [lower-alpha 2] trilobites being in the order proetida, who are librostomes.
Some librostomes may not even be within the group, with Whittington (2003) placing some genera (Hemibarrandia, Lakaspis, Peraspis and Symphysurina) of the asaphid family Nileidae in the order Corynexochida. [6]
Librostomes make up a large component of the Trilobita. with Traditionally, four orders are placed in the librostoma; they are the: Asaphida, Harpetida, Proetida, and Ptychopariida; however, two primitive Phacopids (Pharostomina and Bavarilla) have natant hypostomes, indicating that the Phacopids are also librostomes. [2] In more recent publications, the asaphid superfamily Trinucleioidea is placed in its own order, Trinucleida, [3] [7] however, even today some studies recover them as asaphids. [8]
The first librostomes were Ellipsocephaloid Ptychopariida in the family Bigotinidae from the early cambrian. [9] This group was once thought to be in the order redlichiida, but are now considered primitive Ptychopariida.
During the Middle Cambrian, the Ptychopariida became the dominant trilobites. Around that time, the Asaphida and Trinucleida would first appear.
The first Harpetida would appear in the late cambrian with the primitive genera Baikadamaspis and Entomaspis of the family Entomaspididae.
The first Proetida appeared during the cambrian-ordovician transition. Since the earliest Phacopida (which are most likely librostomes) appeared during the early ordovician, it is assumed that a late cambrian sister taxon was present. [2]
During the Ordovician, the Ptychopariida declined. After the decline of the Ptychopariida, the order Asaphida rose to prominence, becoming the dominant order of trilobites by the end of the mid-ordovician.
The End-Ordovician extinction event would drastically reduce trilobite diversity, with librostomes included. The already endangered Ptychopariida would go extinct. All trilobites with planktonic or Nektonic life stages would go extinct, [10] which included the swimming Proetida family Telephinidae, and all Asaphids & Trinucleids except for the raphiophorid Raphiophorus, which would go extinct shortly after in the Silurian.
Librostomes occupied many niches, sizes, and ecologies. Both some of the largest (Ordovician asaphids Isotelus rex, and Hungioides ) and smallest Trilobites (Ordovician ptychopariid Acanthopleurella stipulae). [11] Many trilobites in the group had some ability to stay above the sea floor, such as asaphids in the superfamilies Cyclopygoidea and Remopleuroidea, [12] [13] proetids in the family Telephinidae, and perhaps some Cheirurid phacopids such as Deiphon and Crotalocephalus.
The Asaphida, Ptychopariida (and Phacopida if included) are some of the most morphologically and cladistically diverse orders of Trilobites, while the Proetida and Harpetida had a more modest morphological diversity, the former (sometimes referred to as 'garden variety trilobites') [14] containing the most diverse family of trilobites in terms of genera, and the latter being more modest in both terms of diversity.
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 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.
Nektaspida is an extinct order of non-mineralised artiopodan arthropods. They are known from the lower-Cambrian to the upper Silurian. Originally classified as trilobites, which they superficially resemble, they are now placed as close relatives as members of the Trilobitomorpha within Artiopoda. The order is divided into three major families; Emucarididae, Liwiidae, and Naraoiidae.
Harpetida is one of the eleven orders of the extinct arthropod class Trilobita. The first harpetid trilobites appear in the Upper Cambrian, and the last species die out at the end of the Devonian period.
Asaphida is a large, morphologically diverse order of trilobites found in marine strata dated from the Middle Cambrian until their extinction during the Silurian. Asaphida contains six superfamilies, but no suborders. Asaphids comprise some 20% of described fossil trilobites.
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.
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.
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.
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.
Redlichiina is a suborder of the order Redlichiida of Trilobites. The suborder contains three superfamilies: Emuelloidea, Redlichioidea and Paradoxidoidea. These trilobites are some of the oldest trilobites known. They originated at the beginning of the Cambrian Period and disappeared at the end of the middle Cambrian.
Aulacopleura is a genus of proetid trilobite that lived from the Middle Ordovician to the Middle Devonian. Some authors may classify this group as subgenus Otarion (Aulacopleura). The cephalon is semicircular or semielliptical, with border and preglabellar field. The glabella is short, with or without defined eye ridges connecting it with eyes of variable size. Spines at the rear outer corners of the cephalon are present, typically reaching back to the 2nd to 4th thorax segment. The 'palate' is not connected to the dorsal shield of the cephalon. The cephalon is pitted, or has small tubercles. The thorax has up to 22 segments. The pleural ends are usually rounded. The pygidium is small (micropygous), with an even margin. A. koninckii had a modern type of compound eye.
Asaphidae is a family of asaphid trilobites. Although the first genera originate in Upper Cambrian marine strata, the family becomes the most widely distributed and most species-rich trilobite family during the Ordovician. 754 species assigned to 146 genera are included in Asaphidae.
The hypostome is the hard mouthpart of trilobites found on the ventral side of the cephalon (head). The hypostome can be classified into three types based on whether they are permanently attached to the rostrum or not and whether they are aligned to the anterior dorsal tip of the glabella.
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.
Typhlokorynetes plana is a species of small, button-shaped asaphid trilobites of the family Raphiophoridae that lived during the Early Tremadocian of Vermont, United States.
The Saint-Chinian Formation is a geological formation composed of shales with limestone inclusions, dating from the Lower Ordovician (Tremadocian).
Trinucleioidea is a superfamily of trilobites. Traditionally placed within the Asaphida, it is now sometimes considered its own order, Trinucleida.