Villebrunaster Temporal range: Early Ordovician, ~ | |
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Fossil of V. thorali | |
Fossil holotype of V. fezouataensis | |
Scientific classification | |
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Family: | Fell, 1963 [1] |
Genus: | †Villebrunaster Spencer, 1951 |
Type species | |
†Villebrunaster thorali Spencer, 1951 | |
Other species | |
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Villebrunaster is an extinct genus of starfish-like animal belonging to Asterozoa that lived around 480 million years ago during Early Ordovician Period in modern-day southern France and Morocco. As of 2022, it contains two species, namely V. thorali and V. fezouataensis. [2] V. thorali was described in 1951 and V. fezouataensis was described in 2021. Villebrunaster represents one of the oldest members of asterozoans, and perhaps, according to a description in 2021, the earliest divergent stem-group (ancestral members) of Asterozoa. [3]
The first species was discovered and described by British palaeontologist William Kingdon Spencer in 1951. [4] The fragmentary specimens were collected from Saint-Chinian Formation in southern France. [5] Spencer recognised it as among "the earliest starfish known." [4] Another British palaeontologist Juliette Dean Shackleton identified new specimens as Ampullaster ubaghsi in 2005, [5] which were later reclassified as V. thorali. [6]
The second species, V.fezouataensis, was described by Aaron W. Hunter and Javier Ortega-Hernández at the University of Cambridge as Cantabrigiaster fezouataensis in 2021. [3] [7] The specimens, originally collected from Fezouata Shale Formation in Morocco, were reanalysed by American palaeontologist Daniel B. Blake and Frederick H.C.Hotchkiss who moved to taxonomic position the genus Villebrunaster in 2022. [2]
The body of Villebrunaster is that of a typical starfish having five radiating arms. Mouth is at the centre of the body. The mouth region is composed of three types of endoskeletons called ossicles, such as half-cylinders or the ambulacral, virgal that form skeleton between the ambulacral, and a pair of mouth plates that radiate into the mouth opening. [4] The arms are broad and evenly arranged to form pentagonal structure. [5]
The presence of virgal ossicles is a feature of Somasteodea. However, Villebrunaster lacks axially oriented ossicles along the lateral margins of the arms, which are found in somasteroids. This suggests that it is a primitive member of the group. [3] The radial water channel are large and run close to the ventral side of the body, while the transverse water channels small and difficult to recognise. The ossicle series are larger in V.fezouataensis than in V. thorali. [2]
Phylogenetic analysis indicate that Villebrunaster is oldest known and the earliest diverging group among Asterozoa, a group that includes starfish, brittle stars and basket stars. The absence of some ambulacral ossicles but presence of virgal ossicles show that the development and variation of ossicles are the important features in the evolution of later asterozoans. [3] [8] However, the genus does not necessarily represent the common ancestor of Asterozoa. [2]
An echinoderm is any member of the phylum Echinodermata. The adults are recognisable by their radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the sea lilies or "stone lilies". Adult echinoderms are found on the sea bed at every ocean depth, from the intertidal zone to the abyssal zone. The phylum contains about 7,000 living species, making it the second-largest grouping of deuterostomes, after the chordates. Echinoderms are the largest entirely marine phylum. The first definitive echinoderms appeared near the start of the Cambrian.
Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that are attached to the sea bottom by a stalk in their juvenile form are commonly called sea lilies, while the unstalked forms, called feather stars or comatulids, are members of the largest crinoid order, Comatulida. Crinoids are echinoderms in the phylum Echinodermata, which also includes the starfish, brittle stars, sea urchins and sea cucumbers. They live in both shallow water and in depths as great as 9,000 meters (30,000 ft).
Starfish or sea stars are star-shaped echinoderms belonging to the class Asteroidea. Common usage frequently finds these names being also applied to ophiuroids, which are correctly referred to as brittle stars or basket stars. Starfish are also known as asteroids due to being in the class Asteroidea. About 1,900 species of starfish live on the seabed in all the world's oceans, from warm, tropical zones to frigid, polar regions. They are found from the intertidal zone down to abyssal depths, at 6,000 m (20,000 ft) below the surface.
Sea cucumbers are echinoderms from the class Holothuroidea. They are marine animals with a leathery skin and an elongated body containing a single, branched gonad. They are found on the sea floor worldwide. The number of known holothurian species worldwide is about 1,786, with the greatest number being in the Asia-Pacific region. Many of these are gathered for human consumption and some species are cultivated in aquaculture systems. The harvested product is variously referred to as trepang, namako, bêche-de-mer, or balate. Sea cucumbers serve a useful role in the marine ecosystem as they help recycle nutrients, breaking down detritus and other organic matter, after which bacteria can continue the decomposition process.
Brittle stars, serpent stars, or ophiuroids are echinoderms in the class Ophiuroidea, closely related to starfish. They crawl across the sea floor using their flexible arms for locomotion. The ophiuroids generally have five long, slender, whip-like arms which may reach up to 60 cm (24 in) in length on the largest specimens.
Cothurnocystis is a genus of small enigmatic echinoderms that lived during the Ordovician. Individual animals had a flat boot-shaped body and a thin rod-shaped appendage that may be a stem, or analogous to a foot or a tail. Fossils of Cothurnocystis species have been found in Nevada, Scotland, Czech Republic, France and Morocco.
Articulata are a subclass or superorder within the class Crinoidea, including all living crinoid species. They are commonly known as sea lilies or feather stars. The Articulata are differentiated from the extinct subclasses by their lack of an anal plate in the adult stage and the presence of an entoneural system. Articulata first appeared in the fossil record during the Triassic period although other, now extinct crinoid groups, originated in the Ordovician.
The stylophorans are an extinct, possibly polyphyletic group allied to the Paleozoic Era echinoderms, comprising the prehistoric cornutes and mitrates. It is synonymous with the subphylum Calcichordata. Their unusual appearances have led to a variety of very different reconstructions of their anatomy, how they lived, and their relationships to other organisms.
Eiffelia is an extinct genus of sponges known from the Middle Cambrian Burgess Shale as well as several Early Cambrian small shelly fossil deposits. It is named after Eiffel Peak, which was itself named after the Eiffel Tower. It was first described in 1920 by Charles Doolittle Walcott. It belongs in the Hexactinellid stem group. 60 specimens of Eiffelia are known from the Greater Phyllopod bed, where they comprise 0.11% of the community.
Hamptonia is an extinct genus of sea sponge known from the Middle Cambrian Burgess Shale and the Lower Ordovician Fezouata formation. It was first described in 1920 by Charles Doolittle Walcott. 48 specimens of Hamptonia are known from the Greater Phyllopod bed, where they comprise < 0.1% of the community.
Pirania is an extinct genus of sea sponge known from the Middle Cambrian Burgess Shale and the Ordovician Fezouata formation. It is named after Mount St. Piran, a mountain situated in the Bow River Valley in Banff National Park, Alberta. It was first described in 1920 by Charles Doolittle Walcott. 198 specimens of Pirania are known from the Greater Phyllopod bed, where they comprise 0.38% of the community.
The Fezouata Formation or Fezouata Shale is a geological formation in Morocco which dates to the Early Ordovician. It was deposited in a marine environment, and is known for its exceptionally preserved fossils, filling an important preservational window beyond the earlier and more common Cambrian Burgess shale-type deposits.
The calcichordate hypothesis holds that each separate lineage of chordate evolved from its own lineage of mitrate, and thus the echinoderms and the chordates are sister groups, with the hemichordates as an out-group.
Thelxiope is a genus of Cambrian and Ordovician arthropod. Four named species are known, the type species T. palaeothalassia is known from the Burgess Shale, Canada T. holmani is from the Wheeler Shale of Utah, Thelxiope tangi from the Linyi Lagerstätte of Shandong, China, and T. spinosa, which is known from both the Linyi Lagerstätte and the Wheeler Shale. An indeterminate species is also known from the Ordovician (Floian) Fezouata Formation in Morocco. It is a member of Mollisoniida, alongside close relatives Mollisonia, Corcorania and Urokodia. They are suggested to be stem-chelicerates.
Furca is an extinct genus of marrellomorph arthropod known from the Sandbian stage of the Czech Republic, with a single currently described species, Furca bohemica. A tentative additional species, "Furca mauretanica": was proposed for specimens discovered in Morocco, but this species remains a nomen nudum until formally published, and probably belongs in a new separate genus.
Aegirocassis is an extinct genus of giant radiodont arthropod belonging to the family Hurdiidae that lived 480 million years ago during the early Ordovician in the Fezouata Formation of Morocco. It is known by a single species, Aegirocassis benmoulai. Van Roy initiated scientific study of the fossil, the earliest known of a "giant" filter-feeder discovered to date. Aegirocassis is considered to have evolved from early predatory radiodonts. This animal is characterized by its long, forward facing head sclerite, and the endites on its frontal appendages that bore copious amounts of baleen-like auxiliary spines. This animal evolving filter-feeding traits was most likely a result of the Great Ordovician Biodiversification Event, when environmental changes caused a diversification of plankton, which in turn allowed for the evolution of new suspension feeding lifeforms. Alongside the closely related Pseudoangustidontus, an unnamed hurdiid from Wales, the middle Ordovician dinocaridid Mieridduryn, and the Devonian hurdiid Schinderhannes this radiodont is one of the few known dinocaridids known from post-Cambrian rocks.
The Somasteroidea, or Stomasteroidea, is an extinct order of asterozoan echinoderms first defined in 1951 by W. K. Spencer. Their first appearance in the fossil record was in the Early Ordovician (Tremadocian) and they had become extinct by the Late Devonian (Famennian). They are similar to the asteroids in that their bodies are flattened dorsoventrally and they have five petaloid arms with broad bases. The ambulacral plates in somasteroids are simple and unspecialized, and the arms were thought to be not flexible and were unable to assist in feeding, but the oral mouth parts were more complex. According to the World Register of Marine Species, Stelleroidea is no longer considered valid and it has since been absorbed into Somasteroidea and Asterozoa.
Rathbunaster is a monospecific genus of sea stars in the family Asteriidae. The genus name was given by Walter Kenrick Fisher to honor the starfish biologist Richard Rathbun of the Smithsonian Institution. He originally ranged this genus under the family Pycnopididae, synonymous with Asteriidae.
Flexibilia is a superorder of specialized Paleozoic crinoids. They exhibited a conserved body plan and consistent suite of characteristics throughout their long history. Previously considered a subclass with unclear affinities, later investigation determined that flexibles are well-nested within Cladida, a broad group ancestral to living articulate crinoids. The Ordovician cladid Cupulocrinus acts as an intermediate form linking the generalized anatomy of other early cladids with the distinctive anatomy of flexibles, and several studies have considered it to be ancestral to the rest of the group. Although flexibles never reached the same abundance or diversity as many other crinoid groups, they remained a reliable component of crinoid faunas, particularly from the Silurian onwards. Flexible fossils are very rare in the Ordovician, but the Late Ordovician appears to have been an interval of rapid diversification for the group.
Mieridduryn is a genus of extinct dinocaridid arthropod that lived during the Middle Ordovician of what is now the United Kingdom. This animal was described in 2022 based on a singular fossil found in Castle Bank, a Burgess shale type lagerstätte located in the country of Wales. This animal's taxonomic affinities are somewhat unclear, but there are some hypotheses. One is that this animal represents a new grade of stem-euarthropods that evolved features similar to the Cambrian aged opabiniids. Another is that if the features seen in Mieridduryn are convergent, and not homologous, to those seen in radiodonts, then this animal represents a late surviving opabiniid.