Kutorginata Temporal range: | |
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Kutorgina cingulata, a kutorginid from the lower Cambrian of Canada. Seen looking onto the ventral valve (left), dorsal valve (right), and the side (top) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Brachiopoda |
Class: | † Kutorginata Williams et al., 1996 |
Order: | † Kutorginida Kuhn, 1949 |
Subgroups | |
See text. |
Kutorginates (Kutorginata) are an extinct class of early rhynchonelliform ("articulate") brachiopods. The class contains only a single order, Kutorginida (kutorginides). [2] Kutorginides were among the earliest rhynchonelliforms, restricted to the lower-middle part of the Cambrian Period ("Atdabanian" [stage 3] to "Mayan" [late Miaolingian]). [3]
Despite this short span of time, kutorginides were still a major order of Cambrian rhynchonelliforms during the lower Cambrian. [1] Kutorginide diversity was highest up to the "Toyonian", though they began to decline in the mid-Cambrian even as other brachiopod orders (particularly orthides and acrotretides) diversified. A similar pattern of diversity loss is seen in obollelides, naukatides, and chileides, three other early rhynchonelliform orders contemporary with kutorginides. [3]
Kutorginides typically have a ventribiconvex shell (both valves convex, the ventral valve moreso) and a strophic (straight) hinge line. Based on fossils of Nisusia, the shell’s internal (secondary) layer appears to have a microstructure of calcite fibers. [2]
Internally, the valves are simpler than most other rhynchonelliforms. Though kutorginides are technically articulate, the hinge is not braced by teeth and sockets, but rather a system of thin ridges and deep furrows along the hinge line. Likewise, cardinalia and dental plates are absent, with only a few subtle muscle scars in their place. The only other apparent structures are the mantle canals, which are pinnate in form (radiating, apart from the midline canals). [2]
The rear of the shell has a roughly kite- to diamond-shaped profile. The pseudodeltidium is voluminous, covering most of the delthyrium (ventral indentation) and taking up a large portion of the ventral valve from the rear. Conversely, the chilidium is rather low and undeveloped, leaving a large exposed notothyrium (dorsal indentation). A small pedicle foramen lies at the apex of the pseudodeltidium. [2]
Kutorginides also have another much larger and more enigmatic opening at the middle of the hinge line. This opening corresponds to the space encompassed by the notothyrium and the exposed portion of the delthyrium. [2] The soft-tissue relevance of this opening has been a subject of debate, and recent evidence has argued for a more nuanced interpretation with variation within the class. [1]
Several kutorginides are preserved in lagerstätten, elaborating on the structure and location of the pedicle relative to the two posterior openings. Kutorgina chengjiangensis, from the Chengjiang Lagerstätte of China, has a pedicle in the form of a thick annulated stalk. Despite its thickness, the pedicle is joined to the shell at the pseudodeltidium, strongly suggesting that it was originally derived from within the small pedicle foramen. The large posterior opening probably helped support strong diductor muscles in this interpretation. [1]
Several fossils of Nisusia sulcata, from the Marjum Limestone of Utah, tell a different story. [1] [4] Each fossil includes a distinctive silicified tube within the large posterior opening. This tube has long been interpreted as a coprolite, suggesting that kutorginides had a complete gut terminating at a gap in the rear of the shell. [5] [2] This condition is otherwise unknown in articulate brachiopods, and is more akin to inarticulates in the subphylum Craniiformea. [2] However, modern brachiopods have very small fecal pellets, so the silicified tube may be better interpreted as an adult pedicle. The smaller foramen at the tip of the pseudodeltidium may be a remnant of a larval pedicle, later rendered redundant by a second pedicle developing at the hinge in adulthood. In this scenario, the pedicle of other rhynchonelliforms would be homologous to the larval pedicle of Nisusia sulcata. [1] [4]
From The Treatise on Invertebrate Paleontology (Part H, Revised), unless stated otherwise: [2]
Some species of Kutorgina have a concavo-convex shell, with the smaller brachial (dorsal) valve dished in and the larger pedicle (ventral) valve broadly arched. The brachial valve has a rather prominent interarea at the back which is curved over by the prominent beak at the back of the pedicle valve.
It includes the species Kutorgina elanicaMalakhovskaya, 2013 and K. chengjiangensisZhang et al. 2007, among many others. K. chengjiangensis preserves soft anatomy, including a pedicle, lophophore, and gut.
NisusiaWalcott, 1905 (Walcott, 1889) is known from the Miaolingian-age Burgess Shale (~ 508 million years ago). It is a senior synonym to Orthisina alberta Walcott, 1889.
The pedicle of Nisusia emerges from between its valves, as displayed by silicified material of N. sulcata, though it still has an opening at the apex of the pedicle valve. [1]
Craniata is a class of brachiopods originating in the Cambrian period and still extant today. It is the only class within the subphylum Craniiformea, one of three major subphyla of brachiopods alongside linguliforms and rhynchonelliforms. Craniata is divided into three orders: the extinct Craniopsida and Trimerellida, and the living Craniida, which provides most information on their biology. Living members of the class have shells which are composed of calcite, though some extinct forms my have aragonite shells. The shells are inarticulate and are usually rounded in outline. There is no pedicle; the rear edge of the body cavity is a smooth and flat wall perforated by the anus. This class of brachiopods has an unsupported lophophore with only a single row of tentacles. In the absence of a pedicle, the shell is usually attached directly to a hard substrate. Many craniiforms are encrusting animals which attach directly to the shell of another animal, usually another brachiopod. The plicae from the host brachiopod will then appear within the shell of the craniiform.
The Obolellata are a class of Rhynchonelliform brachiopods with two orders, Obolellida and Naukatida. They are essentially restricted to the lower-middle Cambrian.
The Craniidae are a family of brachiopods, the only surviving members of the subphylum Craniiformea. They are the only members of the order Craniida, the monotypic suborder Craniidina, and the superfamily Cranioidea; consequently, the latter two taxa are at present redundant and rarely used.There are three living genera within Craniidae: Neoancistrocrania, Novocrania, and Valdiviathyris. As adults, craniids either live freely on the ocean floor or, more commonly, cement themselves onto a hard object with all or part of the ventral valve.
Strophomenida is an extinct order of articulate brachiopods which lived from the lower Ordovician period to the mid Carboniferous period. Strophomenida is part of the extinct class Strophomenata, and was the largest known order of brachiopods, encompassing over 400 genera. Some of the largest and heaviest known brachiopod species belong to this class. Strophomenids were among the most diverse and abundant brachiopods during the Ordovician, but their diversity was strongly impacted at the Late Ordovician mass extinction. Survivors rediversified into new morphologies in the Silurian, only to be impacted once again at the Late Devonian mass extinction. However, they still survived till the mid Carboniferous.
Brachiopods, phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection. Two major categories are traditionally recognized, articulate and inarticulate brachiopods. The word "articulate" is used to describe the tooth-and-groove structures of the valve-hinge which is present in the articulate group, and absent from the inarticulate group. This is the leading diagnostic skeletal feature, by which the two main groups can be readily distinguished as fossils. Articulate brachiopods have toothed hinges and simple, vertically-oriented opening and closing muscles. Conversely, inarticulate brachiopods have weak, untoothed hinges and a more complex system of vertical and oblique (diagonal) muscles used to keep the two valves aligned. In many brachiopods, a stalk-like pedicle projects from an opening near the hinge of one of the valves, known as the pedicle or ventral valve. The pedicle, when present, keeps the animal anchored to the seabed but clear of sediment which would obstruct the opening.
The origin of the brachiopods is uncertain; they either arose from reduction of a multi-plated tubular organism, or from the folding of a slug-like organism with a protective shell on either end. Since their Cambrian origin, the phylum rose to a Palaeozoic dominance, but dwindled during the Mesozoic.
Acrotretides (Acrotretida) are an extinct order of linguliform brachiopods in the class Lingulata. Acrotretida contains 8 families within the sole superfamily Acrotretoidea. They lived from the Lower Cambrian to the Middle Devonian, rapidly diversifying during the middle Cambrian. In the upper Cambrian, linguliforms reached the apex of their diversity: acrotretides and their relatives the lingulides together comprised nearly 70% of brachiopod genera at this time. Though acrotretides continued to diversify during the Ordovician, their proportional dominance declined, as rhynchonelliforms took on a larger role in brachiopod faunas.
Rhynchonelliformea is a major subphylum and clade of brachiopods. It is roughly equivalent to the former class Articulata, which was used previously in brachiopod taxonomy up until the 1990s. These so-called articulated brachiopods have many anatomical differences relative to "inarticulate" brachiopods of the subphyla Linguliformea and Craniformea. Articulates have hard calcium carbonate shells with tongue-and-groove hinge articulations and separate sets of simple opening and closing muscles.
The Rhynchonellata is a class of Lower Cambrian to Recent articulate brachiopods that combines orders from within the Rhynchonelliformea with well developed pedicle attachment. Shell forms vary from those with wide hinge lines to beaked forms with virtually no hinge line and from generally smooth to strongly plicate. Most all are biconvex. Lophophores vary and include both looped and spiraled forms. Although morphologically distinct, included orders follow a consistent phylogenetic sequence.
Mickwitziids are a Cambrian group of shelly fossils with originally phosphatic valves, belonging to the Brachiopod stem group, and exemplified by the genus Mickwitzia – the other genera are Heliomedusa and Setatella. The family Mickwitziidae is conceivably paraphyletic with respect to certain crown-group brachiopods.
Athyris is a brachiopod genus with a subequally biconvex shell that is generally wider than long and a range that extends from the Silurian into the Triassic. Athyris is the type genus for the Athyrididae, which belongs to the articulate order Athyridida. R.C. Moore (1952) gives a shorter range, from the Mid Devonian to the Lower Mississippian.
Gigantoproductus giganteus is an extinct species of brachiopods in the family Monticuliferidae, known only from its fossil remains. It was a marine invertebrate found on the seabed in shallow seas. It evolved during the Carboniferous period and it is believed to be the largest brachiopod that has ever existed.
Paterinata is an extinct class of linguliform brachiopods which lived from the lower Cambrian ("Tommotian") to the Upper Ordovician (Hirnantian). It contains the single order Paterinida and the subfamily Paterinoidea. Despite being some of the earliest brachiopods to appear in the fossil record, paterinides stayed as a relatively subdued and low-diversity group even as other brachiopods diversified later in the Cambrian and Ordovician. Paterinides are notable for their high degree of convergent evolution with rhynchonelliform (articulate) brachiopods, which have a similar set of muscles and hinge-adjacent structures.
Trimerellida is an extinct order of craniate brachiopods, containing the sole superfamily Trimerelloidea and the families Adensuidae, Trimerellidae, and Ussuniidae. Trimerellidae was a widespread family of warm-water brachiopods ranging from the Middle Ordovician to the late Silurian (Ludlow). Adensuidae and Ussuniidae are monogeneric families restricted to the Ordovician of Kazakhstan. Most individuals were free-living, though some clustered into large congregations similar to modern oyster reefs.
Argyrotheca is a genus of very small to minute lampshells. All species share a large pedicel opening, one ridge on the inside of the pedunculate valve, pits in a diamond pattern on the inside of both valves, and without radial ridges that end in tubercles. It occurs in depths between 6 and 1300 m. It is known since the latest Cretaceous.
Craniopsidae is an extinct family of craniiform brachiopods which lived from the mid-Cambrian to the Lower Carboniferous (Tournaisian). It is the only family in the monotypic superfamily Craniopsoidea and the monotypic order Craniopsida. If one includes the ambiguous Cambrian genus Discinopsis, craniopsids were the first craniiforms to appear, and may be ancestral to craniids and trimerellides. An even earlier Cambrian genus, Heliomedusa, has sometimes been identified as a craniopsid. More recently, Heliomedusa has been considered a stem-group brachiopod related to Mickwitzia.
Strophomenata is an extinct class of brachiopods in the subphylum Rhynchonelliformea.
Siphonotretida is an extinct order of linguliform brachiopods in the class Lingulata. The order is equivalent to the sole superfamily Siphonotretoidea, itself containing the sole family Siphonotretidae. Siphonotretoids were originally named as a superfamily of Acrotretida, before being raised to their own order.
The orthotetides (Orthotetida) are an extinct order of brachiopods in the class Strophomenata. Though not particularly diverse or abundant relative to strophomenides (Strophomenida) or productides (Productida), orthotetides were nevertheless the longest-lasting order of strophomenates, surviving from the Middle Ordovician (“Llanvirn”) up until the Late Permian. Externally, many orthotetides are difficult to distinguish from strophomenides. Most fundamental differences between the two orders are internal: orthotetides have more elaborate cardinal processes and a greater diversity of shell microstructure.
Chileata Wiliams et al., 1996 is a class of rhynchonelliform brachiopods, characterised, among others, by the presence of a perforation in the umbonal region of the ventral valve, usually (partly) covered by a colleplax. They are known from the Cambrian to the Lower Permian.