Ellesmeroceratida
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The Ellesmerocerida is an extinct order of primitive
cephalopods A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
belonging to the subclass
Nautiloidea Nautiloids are a group of 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. Th ...
with a widespread distribution that lived during the Late
Cambrian The Cambrian ( ) is the first geological period of the Paleozoic Era, and the Phanerozoic Eon. The Cambrian lasted 51.95 million years from the end of the preceding Ediacaran period 538.8 Ma (million years ago) to the beginning of the Ordov ...
and
Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
.


Morphology

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. 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. 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 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. As for arms or tentacles, little can be said except that eight or ten, retained in modern coleoids, seems to be
Plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, an ...
in origin.


Taxonomy

Rousseau Flower Rousseau Hayner Flower (1913–1988) was an extremely prolific 20th century paleontologist, known for his eccentric personality. Career Although trained as an entomologist, and a specialist in dragonflies and orthopterans, Flower began study ...
defined the Ellesmerocerida as containing all archaic, ancestral cephalopods and established three suborders within: the Plectronoceratina, Ellesmeroceratina, and Cyrtocerinina. Furnish and Glenister, in the ''Treatise on Invertebrate Paleontology'', Part K,Furnish & Glenister. 1964, Ellesmerocerida, in the ''
Treatise on Invertebrate Paleontology The ''Treatise on Invertebrate Paleontology,'' published from 1953–2007 by the Geological Society of America and the University of Kansas, then 2009–present by the University of Kansas Paleontological Institute, is a definitive multi-authore ...
'', Part-K (Nautiloidea; Geological Society of America and University of Kansas Press; pp K160–K188; figs.
essentially followed suit with minor differences at the family level.
Mary Wade Mary Wade (17 December 1775 – 17 December 1859) was a British teenager and convict who was transported to Australia when she was 13 years old. She was the youngest convict aboard , part of the Second Fleet. Her family grew to include five g ...
(1988)Wade 1988. Nautiloids and their descendants:cephalopod classification in 1986; in Memoir 44, New Mexico Bureau of Mines and Mineral Resources, 1988. 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, 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. Accordingly, the order includes the
Ellesmeroceratidae 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 d ...
,
Protocycloceratidae Protcycloceratidae is an extinct family of slender, commonly annulate, members of the cephalopod order Ellesmerocerida that lived during the Early Ordovician. Protocycloceratidae, named by Kobayashi (1935), are characterized by transversely ribbe ...
,
Cyclostomiceratidae Cyclostomiceratidae, from Ancient Greek κύκλος (''kúklos''), meaning "circle", στόμα (''stóma''), meaning "mouth", and κέρας (''kéras''), meaning "horn", is an extinct family of Early Ordovician The Ordovician ( ) is a geol ...
,
Bassleroceratidae 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 typical ...
,
Eothinoceratidae Eothinoceratidae is a family of Lower Ordovician Nautiloidea, nautiloid cephalopods included in the Cyrtocerinida. The family was originally established for the genus ''Eothinoceras''. Morphology The Eothinoceratidae as typified by ''Eothinocera ...
,
Bathmoceratidae ''Bathmoceras'' is a primitive cephalopod genus from the Middle and Upper Ordovician. It is a member of the order Cyrtocerinida and is the only genus in the family Bathmoceratidae. Derivation At one time ''Bathmoceras'' was thought to have pos ...
, and
Cyrtocerinidae Cyrtocerinidae is a family of nautiloid cephalopods in the Order Cyrtocerinida Cyrtocerinida is an order of Ordovician nautiloid cephalopods. The order includes the families Cyrtocerinidae and Eothinoceratidae, as well as the genera '' Bathmo ...
. 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 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 conn ...
, once included, is now assigned to the
Discosorida 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 co ...
Apocrinoceratidae, Pdb
/ref>


Evolution and phylogeny

The Ellesmerocerida are derived from the
Plectronocerida Plectronocerida is a primitive order from which subsequent cephalopod orders are ultimately derived.Curt Teichert, 1988. Main Features of Cephalopod Evolution. The Mollusca Vol. 12 Paleontology and Neontology of Cephalopds; Academic Pres Inc. Oc ...
, having first appeared early in the Trempealeauan Stage of the Late Cambrian and quickly diversifying into four families, only one of which, the
Ellesmeroceratidae 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 d ...
, continued by means of the genera ''
Ectenolites ''Ectenolites'' is a genus of small, slender, cylindrical Ellesmeroceratids that resemble ''Ellesmeroceras'' but are smaller and proportionally narrower. Septa, as typical for ellesmerocerids, are close spaced with shallow lobes on either flank. ...
'' and ''
Clarkoceras ''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. (Flower 1964, Teichert 1988); the other being ''Ect ...
'' into the Gasconadian in the Lower Ordovician. The other three,
Acaroceratidae Acaroceratidae is family of Upper Cambrian ( Trempealeauan) nautiloid cephalopods included in the Ellesmerocerida that contains two known genera, ''Acaroceras'' and ''Weishanuceras'', both found in northern China. Acaroceratidae represent some ...
, Huaiheceratidae, and Xiaoshanoceratidae having perished in the extinction event that occurred late in the Trempealeauan, before the end of the Cambrian. 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. 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. 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 Endocerida, from Ancient Greek ἔνδον (''éndon''), meaning "inside", and κέρας (''kéras''), meaning "horn", is an extinct nautiloid order, a group of cephalopods from the Lower Paleozoic with cone-like deposits in their siphuncle. End ...
through '' Pachendoceras'' and to the
Tarphycerida The Tarphycerida were the first of the coiled cephalopods, found in marine sediments from the Lower Ordovician (middle and upper Canad) to the Middle Devonian. Some, such as '' Aphetoceras'' and '' Estonioceras'', are loosely coiled and gyroconi ...
and
Oncocerida The Oncocerida comprise a diverse group of generally small nautiloid cephalopods known from the Middle Ordovician to the Mississippian (early Carboniferous; one possible member is known from the Early Permian), in which the connecting rings are t ...
through ''
Bassleroceras ''Bassleroceras'' is an elongate upwardly curved, exogastric, genus of mollusks 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 co ...
'' and is the source for the
Orthocerida Orthocerida, from Ancient Greek ὀρθός (''orthós''), meaning "straight", and κέρας (''kéras''), meaning "horn", also known as the Michelinocerida, is an order of extinct Orthoceratoidea, orthoceratoid cephalopods that lived from the ...
,
Pseudorthocerida Pseudorthocerida is an order of generally straight longiconic (consisting of longicones - long, thin conic shells) orthoceratoids with a subcentral to marginal cyrtochoanitic siphuncle composed of variably expanded segments which may contain in ...
,
Actinocerida The Actinocerida are an order of generally straight, medium to large cephalopods that lived during the early and middle Paleozoic, distinguished by a siphuncle composed of expanded segments that extend into the adjacent chambers, in which depo ...
, and
Discosorida 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 co ...
. The Ellesmerocerida mostly died out by the end of the early Ordovician (Arenigian), although some stragglers survived until the end of the Ordovician.


References

{{Taxonbar, from=Q3723510 Ordovician molluscs Nautiloids Prehistoric cephalopod orders Cambrian first appearances Late Ordovician extinctions Taxa named by Rousseau H. Flower Paraphyletic groups