Cameroceras Temporal range: | |
---|---|
Partial internal mold of C. inaequabile, Upper Ordovician of northern Kentucky | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Mollusca |
Class: | Cephalopoda |
Subclass: | Nautiloidea |
Order: | † Endocerida |
Family: | † Endoceratidae |
Genus: | † Cameroceras Conrad, 1842 |
Type species | |
†Cameroceras trentonense Conrad, 1842 | |
Species | |
See text. |
Cameroceras ("chambered horn") is an extinct genus of endocerid cephalopod which lived in equatorial oceans during the entire Ordovician period. Like other endocerids, it was an orthocone, meaning that its shell was fairly straight and pointed. It was particularly abundant and widespread in the Late Ordovician, inhabiting the shallow tropical seas in and around Laurentia, Baltica and Siberia (equivalent to modern North America, Europe, and Asia). [1]
Cameroceras exhibited a broad range of sizes, and some species were fairly large by extinct cephalopod standards. One species, C. turrisoides from the Boda Limestone of Sweden, [2] is estimated to have shell around 2 metres (6.6 ft) in length, [3] while that of C. rowenaense was about 70 centimetres (2.3 ft). [1] Some books and older scientific papers previously treated Cameroceras as the absolute largest nautiloid-grade cephalopod, with a shell length reaching 5.7 metres (19 ft) [4] [5] or even 9.14 metres (30.0 ft). [6] [4] [1] More recent studies have indicated that the largest orthocone fossils do not belong to the genus Cameroceras, but rather Endoceras giganteum. Moreover, the maximum length estimate is based on a highly doubtful field observation. [5] Cameroceras and Endoceras are indistinguishable in most anatomical aspects, only differing in their shell texture. [7]
Cameroceras is a cephalopod, the same group of molluscs that includes the octopuses, squids and cuttlefish. The only portion of the animal to fossilize is the shell (formally known as the conch). Like other orthoconic nautiloids, Cameroceras had a narrow conical shell with smooth, simple sutures dividing a series of septa (internal chambers). In cross-section, the shell may be perfectly circular or slightly depressed (elliptical, wider than tall). [7]
The position of the siphuncle varies, but in most species it runs close to the lower edge of the shell. In some species the siphuncle is voluminous, reaching a diameter equal to half of the shell diameter. The siphuncle is filled with stacked funnel-shaped concretions known as endocones, which have a simple conical form in Cameroceras. [7] A thin endosiphuncular tube passes through the apex of each endocone, in the lower part of the siphuncle. The surface of the siphuncle is supported by septal necks which are holochoanitic, meaning that they fully sheath the siphuncle and extend all the way between each septum. The main difference between Cameroceras and Endoceras is that Cameroceras lacks annulations (thin concentric rings) on the outer surface of the shell. [7]
From comparison with living cephalopods, particularly the shelled nautilus, some inferences about the biology of Cameroceras can be made. The head of the animal would have been soft muscular tissue situated at the opening of the shell, with the mantle (sheath-like body wall) lying within the shell for protection. Tentacles would have grown from the base of the head, and these tentacles would have been used to seize and manipulate prey. At the base of these tentacles within the buccal mass (analogous to the mouth) a hard keratinous beak would have bitten into the bodies of its prey, and is assumed to have been strong enough to breach the prey's exoskeleton or shell. Modern cephalopods beaks contain a radula, or 'toothed' tongue, which is used to rasp out soft tissue from within the prey's shell.
Cameroceras has historically been utilized as a "wastebasket taxon" in which species of large orthoconic endocerids such as Endoceras, Vaginoceras , and Meniscoceras were originally placed. This poses difficulty for describing Cameroceras as a distinct genus. The type species Cameroceras trentonense was named by Conrad in 1842, based on fossils from the Trenton Limestone of western New York state. [8] The original specimen of C. trentonense is apparently lost, which complicates comparisons to other endocerids. [6]
Hall, who named and described Endoceras annulatum in 1847, recognized C. trentonense as a valid combination, but used Endoceras for other specimens of large endocerids from the Trenton Limestone. Sardeson (1925/1930) suggested that Cameroceras and Endoceras are potentially different growth stages of the same genus, [9] [10] though other authors have doubted this perspective. [6]
For many historical studies, Cameroceras was considered to take precedence over Endoceras whenever the two refer to the same species, according to the principle of priority. [11] [9] [10] [1] Cameroceras' vague early descriptions have led other authors to prefer Endoceras or other better-described genera when the nomenclature is in question. [6] Recent studies generally accept both Cameroceras and Endoceras as valid genera, even some species are in an unstable state between the two. [7] [2] [3] [12]
Fossils assigned to Cameroceras have been found in North America, Asia, and Europe throughout the Ordovician, though most species occur in the Katian stage of the Late Ordovician. Reports of Cameroceras fossils from the Wenlock epoch of the Silurian are based on Rossicoceras hudsonicum , [13] an Ontarian endocerid species sometimes placed within Cameroceras. [7] Species which are currently referred to Cameroceras rather than to other endocerids include:
Species | Author(s) | Year | Temporal range | Notes & description |
---|---|---|---|---|
Cameroceras akpatokense [2] | (Foerste & Cox) | 1936 | Upper Ordovician (late Katian) | From Akpatok Island in Nunavut. Originally known as Endoceras akpatokense. [2] |
Cameroceras alternatum [14] | Flower | 1968 | Upper Ordovician ("Mohawkian") | A very large species (diameter ~ 16.5 cm) based on a single fossil from the Black River Group of Quebec. [14] |
Cameroceras coxi [2] | (Foerste & Cox) | 1936 | Upper Ordovician (late Katian) | From Akpatok Island in Nunavut. Originally known as Endoceras coxi. [2] |
Cameroceras curvatum | Ruedemann | 1906 | Lower Ordovician | From Vermont. |
Cameroceras hasta [2] | (Eichwald) | 1857 | Upper Ordovician (late Katian) | A widespread European species. Previously known as Endoceras hasta, Endoceras megastoma, or Rossicoceras pirguense. [2] |
Cameroceras hennepini [11] | Clarke | 1897 | Upper Ordovician ("Shermanian") | A large species (diameter ~ 10 cm, length ~ 4 ft) from the Galena Limestone of Minnesota. [9] |
Cameroceras huzzohense [15] | Ulrich & Foerste | 1930 | Lower Ordovician | A common small species (diameter ~ 2.7 cm) from the Gasconade Formation of Missouri. [15] |
Cameroceras inaequabile [16] | (Miller) | 1882 | Upper Ordovician ("Richmondian") | A fairly uncommon but widespread American species found throughout "Richmondian" strata in Ohio, [17] [18] Kentucky, Indiana, [17] and Illinois. [16] Originally known as Endoceras inaequabile. [16] |
Cameroceras inopinatum [19] | Stauffer | 1937 | Lower Ordovician | A tiny species (diameter ~ 1 cm) from the Shakopee Dolomite of Minnesota. [19] |
Cameroceras motsognir [12] | Kröger & Aubrechtová | 2019 | Upper Ordovician (late Sandian – Katian?) | A small species (diameter ~ 4.3 cm) with a slightly curved shell. From the Kullsberg Limestone Formation of Sweden. [12] |
Cameroceras regulus [2] | (Eichwald) | 1860 | Upper Ordovician (late Katian) | A medium-sized European species (diameter ~ 7.2 cm) found in Estonia and Sweden. [2] Originally known as Endoceras regulus. |
Cameroceras rowenaense [1] | Frey | 1995 | Upper Ordovician ("Maysvillian") | A medium-sized species (diameter ~ 8 cm, length > 70 cm) from the Leipers Limestone of Kentucky. [1] |
Cameroceras stillwaterense [20] | Stauffer | 1937 | Lower Ordovician | A small species (diameter ~ 1.5 cm, length > 10 cm) from the Shakopee Dolomite of Minnesota. [20] |
Cameroceras styliforme | Grabau | 1922 | Lower Ordovician | From Hubei, China. |
Cameroceras trentonense (type species) | Conrad | 1842 | Upper Ordovician (late Katian) | The type species, from the Trenton Limestone of New York and possibly the Lexington Limestone of Kentucky. [1] |
Cameroceras turrisoides [2] | Kröger | 2013 | Upper Ordovician (mid-late Katian) | A very large species (maximum diameter ~ 17 cm, total length ~ 2 m) [3] from the Boda Limestone of Sweden and the Bardahessaigh Formation of Ireland. [2] |
Cameroceras vertebrale | (Eichwald) | 1860 | Middle Ordovician | |
Cameroceras windriverense [2] | (Miller) | 1932 | Upper Ordovician (late Katian) | From the Lander Sandstone. Originally known as Endoceras windriverense. [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.
Orthoceras is a genus of extinct nautiloid cephalopod restricted to Middle Ordovician-aged marine limestones of the Baltic States and Sweden. This genus is sometimes called Orthoceratites. Note it is sometimes misspelled as Orthocera, Orthocerus or Orthoceros.
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.
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.
The Bactritida are a small order of more or less straight-shelled (orthoconic) cephalopods that first appeared during the Emsian stage of the Devonian period with questionable origins in the Pragian stage before 409 million years ago, and persisted until the Carnian pluvial event in the upper middle Carnian stage of the Triassic period. They are considered ancestors of the ammonoids, as well as of the coleoids.
Aulacocerida is an order of primitive coleoid cephalopods, possibly derived from michelinoceraitids (Orthocerida) early in the Devonian, which in turn gave rise to the Belemnites.
Endoceras is an extinct genus of large, straight shelled cephalopods that gives its name to the Nautiloid order Endocerida. The genus lived during the middle and upper Ordovician 470 to 443 million years ago. The cross section in the mature portion is slightly wider than high, but is narrower laterally in the young. Sutures are straight and transverse. Endoceras has a large siphuncle, located close to the ventral margin, composed of concave segments, especially in the young but which may be tubular in the adult stage. Endocones are simple, subcircular in cross section, and penetrated by a narrow tube which may contain diaphragms reminiscent of the Ellesmerocerid ancestor.
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.
Mcqueenoceras is an extinct genus of early endocerid, a nautiloid from the Floian epoch of the late early Ordovician period. It was similar in overall form to Clitendoceras, from which it may have been derived. Mcqueenoceras, like Clitendoceras, has ventral siphuncle but the endocones are thicker on the ventral side and thinner on the dorsal. Also the sutures in Mcqueenoceras retreat rearward, forming lobes as they cross the venter. The type species is Mcqueenoceras jeffersonense, named by E.O. Ulrich and A.F. Foerste in 1935, and it is known from Missouri and New York. In 1956, Rousseau H. Flower named two other species, M. cariniferum and M. ventrale, both known from Maryland.
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).
Bactroceras is a genus of orthoceratoid cephalopods that lived during the early Middle Ordovician, from about 472—464 mya, existing for approximately 8 million years.
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.
Endoceratidae is a family of large to very large straight shelled nautiloid cephalopods belonging to the order Endocerida that lived during the Middle and Late Ordovician. They include the largest known Paleozoic invertebrates, represented by Endoceras and Cameroceras.
The cephalopods have a long geological history, with the first nautiloids found in late Cambrian strata.
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.
Proterovaginoceras is a medium to large sized endocerid from the Early and Middle Ordovician included in the family Endoceratidae.
Cartersoceras is a genus of nautiloid cephalopods placed in the Orthocerid family Baltoceratidae, established by Rousseau Flower (1964), for species formally included in part in Murrayoceras, and in part in "Sacotoceras". This group is typical of Murrayoceras in all features except that the siphuncle segments are convex in outline. It is known from the Middle Ordovician of the eastern United States.
Murrayoceras is a nautilid cephalopod included in the orthocerid family Baltoceratidae, widespread in the Middle Ordovician of North America, characterized by a depressed orthoconic shell with a subtriangular cross section and flattened venter and a proportionally large ventral siphuncle, 0.15 to 0.3 the dorso-ventral shell diameter. Septa are close spaced with sutures forming broad lobes on the upper flanks and ventral surface.
Protactinocerida is a proposed order of Late Cambrian nautiloid cephalopods. Their fossils have only been found in the Late Cambrian of North China, a diversity pattern similar to other early cephalopod orders.
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.