Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Spiralia
Cladus: Lophotrochozoa
Phylum: Mollusca
Classis: Cephalopoda
Subclassis: Nautiloidea
Ordo: †Orthocerida
Familiae: Brachycycloceratidae - Clinoceratidae - Dawsonoceratidae - Geisonoceratidae - Lamellorthoceratidae - Offleyoceratidae - Paraphragmitidae - Sactorthoceratidae - Sphooceratidae - Troedssonellidae
Orthocerida, also known as the Michelinocerida, is an order of extinct orthoceratoid cephalopods that lived from the Early Ordovician (490 million years ago) possibly to the Late Triassic (240 million years ago).[1] A fossil found in the Caucasus suggests they may even have survived until the Early Cretaceous (150 million years ago),[2] and the Eocene fossil Antarcticeras is sometimes considered a descendant of the orthocerids although this is disputed.[3][4] They were most common however from the Ordovician to the Devonian.
Shell form
The shell is usually long, and may be straight ("orthoconic") or gently curved. In life, these animals may have been similar to the modern squid, except for their long shell. The internal structure of the shell consists of concavo-convex chambers linked by a centrally-placed tube called a siphuncle. There is a tendency for the chambers to develop cameral deposits, which were used as ballast to balance the long gas-filled shell. Depending on the family, the siphuncle has orthochoanitic (short and straight) or cyrtochoanitic (outwardly curved) septal necks, which protrude from the septa. The shell surface may be (depending on the species or genus) smooth, transversely ribbed, or ornamented by a network of fine lirae. Fossils are common and have been found on many continents, including the Americas, Africa, Europe, and Asia.
Ecology
Orthocerids may have swum near the sea bed with their buoyant shell resting horizontally in the water, floated more passively among plankton, or rested on the sea floor. Like modern cephalopods they would have used jet-propulsion for locomotion. Their long, bulky shell and relatively weak muscle attachments however make it unlikely that they were as agile as ammonoids or modern cephalopods. They most likely fed on trilobites and small arthropods.
Taxonomy
Orthocerid taxonomy is based primarily on morphological characters found in the shell, principally in the nature of the siphuncle. Parsing these relationships out however can become complicated due to similar characteristics through convergent evolution, making certain taxa appear to have a much longer stratigraphic range than in actuality.[5]
Well preserved embryonic shells of the family Pseudorthoceratidae from the Mississippian (Lower Carboniferous) Imo Formation of Arkansas revealed significant diversity in the shells' morphology, as well as indicating that Pseudorthoceratidae may be in need of revision.[6]
Evolutionary history
Orthocerids likely arose from the Baltoceratidae, a family of the Ellesmerocerida. However, the phylogeny of Orthocerida is heavily debated as it lacks clearly defined characters. Some important characteristics include the diameter and thickness of a calciosiphonate connecting ring and shape of orthocone siphuncles.[7][8] Orthocerida may thus be a polyphyletic group, having arisen as several lineages from early Ordovician cephalopods.[9] Some workers have split off the Pseudorthocerida and Dissidocerida as separate orders, the latter on the grounds that it arose from a different baltoceratid ancestor. The pseudorthocerids are thought to be distinct because their protoconch and septal necks significantly different than orthocerids.[10]
Orthocerids flourished in the Paleozoic Era, giving rise to multiple orders, most notably ascocerids. The spherical protoconch, or first chamber, of some orthocerids suggests they were ancestors to the Bactritida, small orthoconic forms that gave rise to both the ammonoids and coleoids. Fossilized radulas from orthocerids also suggest a closer affinity with modern coleoids than with Nautilus.
There is some dispute as to when the orthocerids became extinct. Although they are said to have survived into the Triassic Period, the two genera that date from that period may actually be pseudorthocerids. In that case, the last orthocerids may date only to the Permian. However, the discovery in the Caucasus of a possible orthocerid from the Early Cretaceous suggests that they may have endured much longer as a ghost lineage.[2] Even a potential Cenozoic descendant of orthocerids is known in Antarcticeras, an enigmatic cephalopod from the Eocene of Antarctica with an internal shell akin to that of coleoids but with an anatomy closely resembling that of orthocerids, suggesting that it may represent an orthocerid offshoot that diverged as its own subclass during the Paleozoic and survived up to the Eocene, convergently evolving a coleoid-esque internal shell.[3] However, other studies recover it as a primitive oegopsid squid.[4]
References
"Orthocerida". Atlas of Ordovician Life. 2013-12-03. Retrieved 2023-10-10.
"An Early Cretaceous orthocerid cephalopod from north-western Caucasus | The Palaeontological Association". www.palass.org. Retrieved 2023-10-10.
Doguzhaeva, Larisa A.; Bengtson, Stefan; Reguero, Marcelo A.; Mörs, Thomas (2017-03-01). "An Eocene orthocone from Antarctica shows convergent evolution of internally shelled cephalopods". PLOS ONE. 12 (3): e0172169. doi:10.1371/journal.pone.0172169. ISSN 1932-6203. PMC 5332165. PMID 28248970.
Fuchs, Dirk; Keupp, Helmut; Klug, Christian (2020-01-02). "A critical review of Antarcticeras Doguzhaeva, 2017 – teuthid affinities can explain the poorly mineralized phragmocone". Historical Biology. 32 (1): 49–54. doi:10.1080/08912963.2018.1467905. ISSN 0891-2963.
Kröger, B.; Mapes, H. (2005). "Revision of some common carboniferous genera of North american orthocerid nautiloids" (PDF). Journal of Paleontology. 5 (79): 954–963.
KröGer, B.; Mapes, R. H. (May 2004). "Lower Carboniferous (Chesterian) Embryonic Orthoceratid Nautiloids". Journal of Paleontology. 78 (3): 560–573. doi:10.1666/0022-3360(2004)078<0560:LCCEON>2.0.CO;2. ISSN 0022-3360.
Mutvei, Harry (2016-04-02). "Siphuncular structures in Calciosiphonate nautiloid orders Actinocerida, Orthocerida and Barrandeocerida (Cephalopoda)". GFF. 138 (2): 295–305. doi:10.1080/11035897.2015.1123768. ISSN 1103-5897.
Kröger, Björn (March 2007). "Early growth‐stages and classification of orthoceridan Cephalopods of the Darriwillian (Middle Ordovician) of Baltoscandia". Lethaia. 39 (2): 129–139. doi:10.1080/00241160600623749. ISSN 0024-1164.
Kröger, Björn (December 2008). "A New Genus of Middle Tremadocian Orthoceratoids and the Early Ordovician Origin of Orthoceratoid Cephalopods". Acta Palaeontologica Polonica. 53 (4): 745–749. doi:10.4202/app.2008.0417. ISSN 0567-7920.
Kröger, Björn (1 February 2006). "Revision of annulated orthoceridan cephalopods of the Baltoscandic Ordovician" (PDF). Fossil Record. 1 (9).
Doguzhaeva, Larisa (1994). "An Early Cretaceous orthocerid cephalopod from North-western Caucasus". Palaeontology. 37 (4): 889–899.
Sweet, Walter C., (1964), Nautiloidea—Orthocerida, in Treatise on Invertebrate Paleontology. Part K. Mollusca 3. (Geological Society of America, and University of Kansas Press, New York, New York and Lawrence, Kansas)
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