| Ellesmerocerida Temporal range: | |
|---|---|
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| Life positions of Ellesmerocerida [3] | |
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Mollusca |
| Class: | Cephalopoda |
| Subclass: | Nautiloidea |
| Superorder: | † Plectronoceratoidea |
| Order: | † Ellesmerocerida Flower, 1950 |
| Subgroups | |
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. [4]
The Ellesmerocerida are characterized by shells that are typically small, some even tiny, with close-spaced septa and relatively large ventral siphuncles. In some genera (e.g. Paleoceras ), the septa are uniformly spaced. [5] Shells of ellesmerocerids are typically smooth and compressed and vary in form. They may be breviconic (short) or longiconic (elongate), straight (orthoconic) or curved (cyrtoconic). Cyrtoconic forms are usually endogastric, with longitudinally convex ventral margins. The apeces of straight forms typically have an endogastric curvature. [6] Some may have grown to as much as 15 cm.
Siphuncle segments are tubular or concave. Septal necks are short. Connecting rings which may appear layered are thick and typically wedge shaped with their maximum width at or near where they join the previous septum. The siphuncle interior is commonly crossed by irregular partitions, known as diaphragms, but are otherwise free of internal deposits [5] [6]
As soft parts are not prone to fossilization, little can be surmised as to their soft part anatomy. Preserved muscle attachment scars indicate that they may have had segmented muscles reminiscent of primitive monoplacophoran molluscs. [1] As for arms or tentacles, little can be said except that eight or ten, retained in modern coleoids, seems to be Plesiomorphic in origin.
Rousseau Flower defined the Ellesmerocerida as containing all archaic, ancestral cephalopods and established three suborders within: the Plectronoceratina, Ellesmeroceratina, and Cyrtocerinina. [5] Furnish and Glenister, in the Treatise on Invertebrate Paleontology, Part K, [7] essentially followed suit with minor differences at the family level. Mary Wade (1988) [8] included the Ellesmerocerida in the superorder Plectonoceratoidea, which she defined as containing the Plectronocerida, Ellesmerocerida and two orders introduced by Chen and Teichert in 1983, [9] the Cambrian Yanhecerida and Protactinocerida. The Plectronocerida, also Cambrian, includes forms once included in the suborder Plectronoceratina, now elevated in rank.
The Ellesmerocerida have been revised to include only primitive nautiloid cephalopods with thick connecting rings and siphuncle segments that are concave in outline. [10] [11] Accordingly, the order includes the Ellesmeroceratidae, Protocycloceratidae, Cyclostomiceratidae, Bassleroceratidae, Eothinoceratidae, Bathmoceratidae, and Cyrtocerinidae. The Ellesmeroceratidae, Protocycloceratidae, Cyclostomiceratidae, Bassleroceratidae are found in Flower's basic Ellesmeroceratina. The Eothinoceratidae, Bathmoceratidae, and Cyrtocerinidae are combined in the Cyrtocerinina. The Schideleroceratidae, Apocrinoceratidae, Baltoceratidae and certain members of the Protocycloceratidae, all which have thin tubular or expanded siphuncles, are now excluded. The Apocrinoceratidae, once included, is now assigned to the Discosorida [12]
The Ellesmerocerida are derived from the Plectronocerida, [5] [8] having first appeared early in the Trempealeauan Stage of the Late Cambrian and quickly diversifying into four families, only one of which, the Ellesmeroceratidae, continued by means of the genera Ectenolites and Clarkoceras into the Gasconadian in the Lower Ordovician. The other three, Acaroceratidae, Huaiheceratidae, and Xiaoshanoceratidae having perished in the extinction event that occurred late in the Trempealeauan, before the end of the Cambrian. [9]
The dominant family of Ordovician Ellesmerocerida is the Ellesmeroceriatidae which are distinguished from the generally similar Protocycloceratidae by the presence of broad lateral lobes in the suture. [11] Sutures in the Protocycloceratidae are straight and transverse. Both contain forms that are annunlate (transversally ribbed) as well as smooth. Other Ordovician families are the broad and beviconic Cyclostomiceratidae and exogastric Bassleroceratidae, rounding out the Ellesmeroceratina. [5] Three families, the Bathmoceratidae, Cyrtocerinidae, and Eothinoceratidae differ from the others in that their connecting rings are greatly thickened inwardly as annular lobes. The Ellesmerocerida gave rise to the Endocerida through Pachendoceras and to the Tarphycerida and Oncocerida through Bassleroceras and is the source for the Orthocerida, Pseudorthocerida, Actinocerida, and Discosorida.
The Ellesmerocerida mostly died out by the end of the early Ordovician (Arenigian), although some stragglers survived until the end of the Ordovician. [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 speciose, 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). Only a handful of rare 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.
Discosorida are an order of cephalopods that lived from the beginning of the Middle Ordovician, through the Silurian, and into the Devonian. Discosorids are unique in the structure and formation of the siphuncle, the tube that runs through and connects the camerae (chambers) in cephalopods, which unlike those in other orders is zoned longitudinally along the segments rather than laterally. Siphuncle structure indicated that the Discosorida evolved directly from the Plectronoceratida rather than through the more developed Ellesmerocerida, as did the other orders. Finally and most diagnostic, discosorids developed a reinforcing, grommet-like structure in the septal opening of the siphuncle known as the bullette, formed by a thickening of the connecting ring as it draped around the folded back septal neck.
Plectronocerida is a primitive order from which subsequent cephalopod orders are ultimately derived.
Clarkoceras is a genus of breviconic ellesmerocerid cephalopods, one of only two genera known to have crossed from the Late Cambrian, Trempealeauan, into the Early Ordovician, Gasconadian. ; the other being Ectenolites.
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.
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).
The Reudemannoceratidae are the ancestral and most primitive of the Discosorida, an order of cephalopods from the early Paleozoic. The Reudemannoceratidae produced generally medium-sized endogastric and almost straight shells with the siphuncle slightly ventral from the center.
Ulrichoceras is recognised as the basal cyrtogomphoceratid genus, which is the source for the rest of the Cyrtogomphoceratidae as well as for the Westonoceratidae. The Cyrtogomphoceratidae are endogastric, the Westonoceratidae, exogastric.
Baltoceratidae is an extinct family of orthoconic cephalopods belonging to the subclass Nautiloidea endemic to what would be Asia, Australia, Europe, North America, and South America during the Ordovician living from about 480–460 mya, existing for approximately 20 million years.
Basslerocerida is an order of nautiloid cephalopods from the Ordovician comprising exogastric longiconic cyrtocones, that is no longer in common use.
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.
Protcycloceratidae is an extinct family of slender, commonly annulate, members of the cephalopod order Ellesmerocerida that lived during the Early Ordovician.
The cephalopods have a long geological history, with the first nautiloids found in late Cambrian strata, and purported stem-group representatives present in the earliest Cambrian lagerstätten.
Bassleroceras is an elongate upwardly curved, exogastric, genus with the venter on the under side more sharply rounded than the dorsum on the upper. The siphuncle is ventral, composed of thick-walled tubular segments in which connection rings thicken in towardly as in both the Ellesmerocerida and primitive Tarphycerida.
Shangtungendoceras is a conical plectronoceratoid cephalopod from the Upper Cambrian of north-eastern China, described by Sun (1937) as a primitive endoceroid. Because of its apparently poor preservation its ordinal and familial position is uncertain. It could be included in either the Plectronocerida, family Plectronoceratidae, or the Ellesmerocerida, family Ellesmeroceratidae. On the other hand, Teichert (1964) included Shungtangendoceras in the Ellesmoerocatidae.
Hectoceras is a genus in the nautiloid cephalopod order Discosorida from the Upper Ordovician of Australia (Tasmania), known from a few isolated siphuncle specimens.
Ellesmeroceras is the type genus for the Ellesmeroceratidae, a family of primitive nautiloid cephalopods, that is characterized by its small, generally compressed, gradually expanded, orthoconic shell, found in Lower Ordovician marine sediments. The septa are close spaced and the siphuncle is ventral, about 0.2 the diameter of the shell. Septal necks are typically orthochoanitic but may slant inwardly (loxochoanitic) or reach halfway to the previous septum (hemichoanitic). Connecting rings are thick. As common for the Ellesmerocerida, Ellesmeroceras has diaphragms within the siphuncle tube.
Apocrinoceratidae constitutes a family of Middle Ordovician nautiloid cephalopods characterized by straight or slightly curved, transversely ribbed shells having siphuncles composed of expanded segments, short recurved septal necks, and thick connecting rings. Derivation is from the Protocycloceratidae, a family of ellesmerocerids, which differ in having straight or concave siphuncle segments, but are otherwise similar in form.
Eburoceras is an early nautiloid cephalopod from the Upper Yenchau and Wanwankau, Upper Cambrian Trempealeauan of China, assigned to the Ellesmeroceratidae.
Notable fossil cephalopods (listed by first occurrence) | |||||||||
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| Cenozoic |
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| Mesozoic |
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| Palaeozoic |
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| Misidentified |
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