Multiceratoidea Temporal range: Nautilida survives to present Possible descendant taxon | |
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Cyrtoceras , a member of the order Oncocerida | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Mollusca |
Class: | Cephalopoda |
Subclass: | Nautiloidea |
Superorder: | † Multiceratoidea Mutvei, 2013 |
Orders | |
Multiceratoidea is a major subclass or superorder of Paleozoic nautiloid cephalopods. [1] [2] Members of this group can be characterized by nautilosiphonate connecting rings, with an organic inner layer and outer layer of calcitic spherules and blades, similar to the modern nautilus. [1] [3] The earliest-diverging multiceratoids have oncomyarian muscle scars (with numerous small muscle attachments ringing the body chamber), [1] though several orders trend towards a ventromyarian condition (with muscle scar area concentrated at the bottom of the body chamber). [4] Multiceratoid shells are generally short and curled, with a relatively small aperture (opening). Cameral deposits are never found among the multiceratoids, though several orders are known to bear endosiphuncular deposits within their siphuncles. [1] [4]
When originally defined in 2013, Multiceratoidea included four nautiloid orders: Ellesmerocerida, Oncocerida, Discosorida, and Ascocerida. The order Tarphycerida was considered a potential member of the subclass, though their larger body chamber and specialized muscle attachments lent uncertainty to this idea. [1] A later analysis added the early-diverging order Cyrtocerinida, which was previously considered a suborder of Ellesmerocerida. Nautilida (the order containing the modern nautilus) was allied with multiceratoids in a broader group termed "Nautilosiphonata", defined by its namesake connecting ring structure. [3]
Solely on the basis of morphological traits, Tarphycerida and Nautilida appear to be well-nested within Multiceratoidea. [4] Multiceratoidea would be an equivalent term to Nautiloidea sensu stricto if nautilids are confirmed to be within the group, though molecular divergence timing disagrees with this assessment. [2] The recently-named order Bisonocerida shares traits with ellesmerocerids and endocerids, [5] and some studies have placed it as a member of Multiceratoidea. [4]
A 2022 bayesian phylogenetic analysis supports the existence of Multiceratoidea as a valid monophyletic clade (discounting the question of nautilus origins), with a membership that includes Tarphycerida and excludes most ellesmerocerids. This is because Ellesmerocerida is a paraphyletic group, an assemblage of early nautiloids ancestral to various later groups, more than just Multiceratoidea. The order Oncocerida was found to be paraphyletic and ancestral to ascocerids and discosorids, though it was an unambiguous member of Multiceratoidea regardless. Bisonocerids were excluded from Multiceratoidea and instead allied with Endocerida, similar to older perspectives on their relationships. [2]
Nautiloids are a group of marine cephalopods (Mollusca) which originated in the Late Cambrian and are represented today by the living Nautilus and Allonautilus. Fossil nautiloids are diverse and species rich, with over 2,500 recorded species. They flourished during the early Paleozoic era, when they constituted the main predatory animals. Early in their evolution, nautiloids developed an extraordinary diversity of shell shapes, including coiled morphologies and giant straight-shelled forms (orthocones). No orthoconic and only a handful of coiled species, the nautiluses, survive to the present day.
Endocerida is an extinct nautiloid order, a group of cephalopods from the Lower Paleozoic with cone-like deposits in their siphuncle. Endocerida was a diverse group of cephalopods that lived from the Early Ordovician possibly to the Late Silurian. Their shells were variable in form. Some were straight (orthoconic), others curved (cyrtoconic); some were long (longiconic), others short (breviconic). Some long-shelled forms like Endoceras attained shell lengths close to 6 metres (20 ft). The related Cameroceras is anecdotally reported to have reached lengths approaching 9 metres (30 ft), but these claims are problematic. The overwhelming majority of endocerids and nautiloids in general are much smaller, usually less than a meter long when fully grown.
Septa are thin walls or partitions between the internal chambers (camerae) of the shell of a cephalopod, namely nautiloids or ammonoids.
Orthocerida, also known as the Michelinocerida, is an order of extinct orthoceratoid cephalopods that lived from the Early Ordovician possibly to the Late Triassic. A fossil found in the Caucasus suggests they may even have survived until the Early Cretaceous, and the Eocene fossil Antarcticeras is sometimes considered a descendant of the orthocerids although this is disputed. They were most common however from the Ordovician to the Devonian.
Plectronocerida is a primitive order from which subsequent cephalopod orders are ultimately derived.
The Ellesmerocerida is an order of primitive cephalopods belonging to the subclass Nautiloidea with a widespread distribution that lived during the Late Cambrian and Ordovician.
The Tarphycerida were the first of the coiled cephalopods, found in marine sediments from the Lower Ordovician to the Middle Devonian. Some, such as Aphetoceras and Estonioceras, are loosely coiled and gyroconic; others, such as Campbelloceras, Tarphyceras, and Trocholites, are tightly coiled, but evolute with all whorls showing. The body chamber of tarphycerids is typically long and tubular, as much as half the length of the containing whorl in most, greater than in the Silurian Ophidioceratidae. The Tarphycerida evolved from the elongated, compressed, exogastric Bassleroceratidae, probably Bassleroceras, around the end of the Gasconadian through forms like Aphetoceras. Close coiling developed rather quickly, and both gyroconic and evolute forms are found in the early middle Canadian.
Lituitida is an order of orthoceratoid cephalopods. They correspond to the family Lituitidae of the Treatise, reranked as an order and combined with other orthoceratoids. They are considered to be more closely related to the Orthocerida than to the Ascocerida or Pseudorthocerida which are also included.
The Ellesmeroceratidae constitute a family within the cephalopod order Ellesmerocerida. They lived from the Upper Cambrian to the Lower Ordovician. They are characterized by straight and endogastric shells, often laterally compressed, so the dorso-ventral dimension is slightly greater than the lateral, with close spaced sutures having shallow lateral lobes and a generally large tubular ventro-marginal siphuncle with concave segments and irregularly spaced diaphragms. Connecting rings are thick and layered, externally straight but thickening inwardly with the maximum near the middle of the segment so as to leave concave depressions on internal siphuncle molds. Septal necks are typically orthochoanitic but vary in length from almost absent (achoanitic) to reaching halfway to the previous septum (hemichoanitic) and may even slope inwardly (loxochoanitic).
Basslerocerida is an order of nautiloid cephalopods from the Ordovician comprising exogastric longiconic cyrtocones, that is no longer in common use.
Orthoceratoidea is a major subclass of nautiloid cephalopods. Members of this subclass usually have orthoconic (straight) to slightly cyrtoconic (curved) shells, and central to subcentral siphuncles which may bear internal deposits. Orthoceratoids are also characterized by dorsomyarian muscle scars, extensive cameral deposits, and calciosiphonate connecting rings with a porous and calcitic inner layer.
The Bassleroceratidae is a family of gradually expanding, smooth ellesmerocerids with a slight to moderate exogastric curvature, subcircular to strongly compressed cross section, and ventral orthochaonitc siphuncle. The ventral side is typically more sharply rounded than the dorsal side and septa are close spaced. Connecting rings are thick and slightly expanded into the siphuncle, making the segments slightly concave; characteristic of the Ellesmerocerida.
Conostichoceras is a genus of exogastric, breviconic oncocerids included in the family Nothoceratidae, known from the Middle Devonian of central Europe and Upper Devonian of Australia. It probably lived on and swum above the sea floor.
Plectronoceratoidea is a superorder or subclass containing primitive nautiloids from the Late Cambrian and Early Ordovician. This group is best considered a paraphyletic grade of early cephalopods, as it contains the ancestors of subsequent post-Cambrian cephalopod orders.
The Nautilitoidea is a superorder within the subclass Nautiloidea, comprising the phylogenetically related Nautilida, Oncocerida, and Tarphycerida.
Cyrtocerinidae is a family of nautiloid cephalopods in the Order Cyrtocerinida, previously considered ellesmerocerids. Members of the family have slightly endogastric breviconic shells with ventral siphuncles that have connecting rings thickened as lobes that project straight into the interior.
Cyrtocerinida is an order of Ordovician nautiloid cephalopods. The order includes the families Cyrtocerinidae and Eothinoceratidae, as well as the genera Bathmoceras and Rummoceras.
Yanhecerida is a small order of Late Cambrian nautiloid cephalopods. They were similar to the more diverse Plectronocerida and Ellesmerocerida, with short shells, closely spaced septa, and diaphragms (partitions) within the siphuncle. Their most distinctive trait is the conical shape of the diaphragms, similar to the endocones which characterize the later nautiloid order Endocerida. While some authors have argued that Yanhecerida should be lumped into Ellesmerocerida, a phylogenetic analysis has maintained that Yanhecerida is a valid clade of early cephalopods closely related to a paraphyletic Ellesmerocerida.
Bisonocerida is an order of Ordovician to Silurian nautiloid cephalopods. Members of this order were originally placed in the order Endocerida, but later investigation argued that this broad usage of Endocerida was a polyphyletic assemblage encompassing two different groups of independent origin. Bisonocerida was differentiated from Endocerida in 2012 in order to resolve this issue.