| Tolkienia Temporal range: | |
|---|---|
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | † Trilobita |
| Order: | † Phacopida |
| Family: | † Acastidae |
| Genus: | † Tolkienia Lieberman & Kloc 1997 |
| Species | |
| |
Tolkienia is an extinct genus of trilobite in the family Acastidae. It consists of three species split from Comura based on morphological differences. The genus was present during the Emsian stage of the Early Devonian epoch around 400 Ma. [1]
The front edge of the glabellar lobe is round when viewed from the top. There are very long furrows in the pygidium. The flaps in these furrows bulge outwards to the edges of the pygidium and have a pointed tip. The rearmost furrow is teardrop-shaped and does not reach as far to the rear as the other flaps. The pygidium becomes narrow to the rear of the sixth axial ring, and the tops of each segment are rounded. On the outside of the exoskeleton there is a prosopon made of small rounded projections. [2]
Tolkienia was first described in 1997 by Bruce Lieberman and Gerald Kloc. The paleontologists named the genus for fantasy writer J. R. R. Tolkien, author of The Lord of the Rings. The genus was split from Comura because the morphology of its pygidium differed from the diagnostic description of Comura. However, no specimens of Comura are intact enough to be used for a phylogenetic analysis in comparison to the new species of Tolkienia, [2] so its taxonomic status remains uncertain. [3] T. granulispina, formerly Comura granulispina, was chosen as the type species for the genus because of the relative completeness of its fossils compared to the other species. [2]
In addition to the three accepted species, several other specimens have been doubtfully placed within Tolkienia. These include a partial fossil from Michigan previously placed in Greenops and a species proposed in 2003 by Martin Basse named †Tolkienia wiltzense. [4]
Fossils of the genus have been extracted from the site of Neuerburg 1 in Germany, [5] the Armorican Massif in France, and the Cantabrian Mountains in Spain. [4] These trilobites were capable of fast movement, and would have lived on the surface of the benthic zone and preyed on small fauna. [5]
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.
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.
Dalmanites is a genus of trilobite in the order Phacopida. They lived from the Late Ordovician to Middle Devonian.
Greenops is a mid-sized Devonian trilobite of the order Phacopida, subfamily Asteropyginae. They are mainly reported from the mid-Devonian Hamilton Group of upstate New York and southwestern Ontario. A similar-looking trilobite from Morocco is often mis-labelled Greenops. Greenops had schizocroidal eyes, large genal spines and short, sharp spines at the tip of each segment of the pygidium ("tail"). Greenops lived in warm, fairly deep water. In the Hamilton Group of New York, they are found with Eldredgeops, Dipleura and Bellacartwrightia, a trilobite that resembles Greenops but has much larger pygidial spines. In Ontario, they are found in the Widder Formation, which outcrops at Arkona, where they are, by far, the dominant trilobite.
Walliserops is a genus of spinose phacopid trilobite, of the family Acastidae, found in Lower to Middle Devonian age rocks from the Anti-Atlas Mountains of Morocco. All species of Walliserops possess a three-pronged "trident" that protrudes from the glabella. Walliserops is most closely related to the genus Comura.
Dikelocephalus is a genus of very large trilobites of up to 50 cm (20 in) long, that lived during the last 3 million years of the Cambrian (Sunwaptan). Their fossils are commonly found as disarticulated sclerites, in the upper Mississippi Valley and in Canada (Alberta). The exoskeleton is rounded anteriorly, with the thorax and sides of the tailshield slightly tapering to about 2⁄3× of the width across the base of the spines at the back of the headshield. At the side corners of the pygidium there may be triangular or hooked spines, pointing backwards, while between the spines the posterior margin is at a 30-75° angle with the lateral margin, gently convex or nearly straight. If pygidial spines are lacking, the margin is gradually rounded. The thorax has 12 segments.
Meteoraspis is an extinct genus of ptychopariid trilobites of the family Tricrepicephalidae. The various species lived from 501 to 497 million years ago during the Dresbachian faunal stage of the late Cambrian Period. Fossils of Meteoraspis are characteristic of Late Cambrian strata in North America, though they are found in Late Cambrian strata elsewhere in the world, such as M. nevensis from Victoria Land, Antarctica.
Erbenochile is a genus of spinose phacopid trilobite, of the family Acastidae, found in Lower to Middle Devonian age rocks from Algeria and Morocco. Originally described from an isolated pygidium, the first complete articulated specimen of E. erbeni revealed the presence of extraordinarily tall eyes:
"Straight-sided towers of lenses... with [up to] 18 lenses in a vertical file"
Dipleura is a genus of trilobites in the order Phacopida. It was described by Green in 1832, and the type species is Dipleura dekayi. The type locality was in the Hamilton Group in New York.
Peachella is an extinct genus of trilobites, fossil marine arthropods, with species of average size. It lived during the Toyonian stage, 516 to 513 million years ago, in what is today the southwestern United States. It can easily be distinguished from other trilobites by its club-like genal spines.
Odontochile is a genus of trilobites in the order Phacopida, family Dalmanitidae.
Lochmanolenellus is an extinct genus of trilobites, fossil marine arthropods, with one small species, L. mexicana. It lived during the Botomian stage (Olenellus-zone), 522–513 million years ago, in the South-West of the former continent of Laurentia, in what are today Mexico, and the South-Western United States.
Glyptagnostus reticulatus is a species of agnostid trilobite belonging to the genus Glyptagnostus. It existed during the Paibian Age of the Cambrian. It has a cosmopolitan distribution and is an important index fossil in biostratigraphy. It was characterized by an unusual net-like pattern of furrows on both the cephalon and the pygidium.
Jinghediscus Xiang & Zhang, 1985 is a genus of Eodiscinid trilobite belonging to the family Weymouthiidae Kobayashi T. (1943), Order Agnostida It lived during the lower Middle Cambrian, with remains found in China (Xinjiang) and Australia (Queensland).
Coltraneia is a genus of trilobite, that lived during the upper Emsian and lower Eifelian, and has been found in Algeria, France, Germany, Morocco and Spain.
Minicryphaeus is a genus of trilobite, that lived during the Pragian in what is today Morocco (Anti-Atlas).
Psychopyge is a genus of trilobite, that lived during the upper Emsian and has been found in Germany and Morocco. It is characterized by the swordlike extension from the front of the head.
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.
Hollardops is a genus of trilobite in the order Phacopida that lived during the Devonian. Their fossils are found in the upper Emsian of western Europe and in the lower Emsian to lowermost Eifelian of North Africa. The type species, Asteropyge mesocristata, was described from Algeria by Le Maître in 1952. The genus Hollardops was erected by Morzadec in 1997. In the same year, Lieberman & Kloc erected Modellops and Philipsmithiana but those genera are regarded as subjective synonyms of Hollardops. The 10-segmented thoracic condition of Hollardops is a rare feature among acastid trilobites that almost always have 11 thorax segments. Van Viersen & Kloc (2022) revisited Hollardops and described a number of new species from the Devonian of Morocco. They also regarded Pennarbedops Bignon & Crônier, 2013 as a synonym of Hollardops. Van Viersen & Kloc construed Hollardops as a scavenger or predator with well-developed eyes, that used its shovel-like cephalon to plough the top layers of the sediment in search of food. Hollardops had small pits horizontally along the fringe of the exoskeleton; these are believed to have housed setae that allowed the trilobite to closely monitor its surroundings.
Bruce Smith Lieberman is an American paleontologist.