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Nautiloid

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Nautiloids
Temporal range: 495–0 Ma Late Cambrian – Recent
Nautilus-JB-01.jpg
Nautilus pompilius
Cameroceras trentonese.jpg
the endocerid Cameroceras, shown feeding on the tarphycerid Aphetoceras, while a quartet of ellesmerocerid Cyclostomiceras swim by.
Scientific classification e
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Subclass: Nautiloidea
Agassiz, 1847
Major subdivisions

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.

In a broad sense, "nautiloid" refers to a major cephalopod subclass or collection of subclasses (Nautiloidea sensu lato). Nautiloids are typically considered one of three main groups of cephalopods, along with the extinct ammonoids (ammonites) and living coleoids (such as squid, octopus, and kin). While ammonoids and coleoids are monophyletic clades with exclusive ancestor-descendant relationships, this is not the case for nautiloids. Instead, nautiloids are a paraphyletic grade of various early-diverging cephalopod lineages, including the ancestors of ammonoids and coleoids. Some authors prefer a narrower definition of Nautiloidea (Nautiloidea sensu stricto), as a singular subclass including only those cephalopods which are close to living nautiluses.

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Cephalopod

Cephalopod

A cephalopod is any member of the molluscan class Cephalopoda such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles modified from the primitive molluscan foot. Fishers sometimes call cephalopods "inkfish", referring to their common ability to squirt ink. The study of cephalopods is a branch of malacology known as teuthology.

Mollusca

Mollusca

Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks. Around 85,000 extant species of molluscs are recognized. The number of fossil species is estimated between 60,000 and 100,000 additional species. The proportion of undescribed species is very high. Many taxa remain poorly studied.

Nautilus (genus)

Nautilus (genus)

Nautilus is a genus of cephalopods in the family Nautilidae. Species in this genus differ significantly in terms of morphology from those placed in the sister taxon Allonautilus. The oldest fossils of the genus are known from the Late Eocene Hoko River Formation, in Washington State and from Late-Eocene to Early Oligocene sediments in Kazakhstan. The oldest fossils of the modern species Nautilus pompilius are from Early Pleistocene sediments off the coast of Luzon in the Philippines.

Allonautilus

Allonautilus

The genus Allonautilus contains two species of nautiluses, which have a significantly different morphology from those placed in the sister taxon Nautilus. Allonautilus is now thought to be a descendant of Nautilus, rendering the latter genus paraphyletic.

Fossil

Fossil

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

Nautilus

Nautilus

The nautilus is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina.

Ammonoidea

Ammonoidea

Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids than they are to shelled nautiloids such as the living Nautilus species. The earliest ammonites appeared during the Devonian, with the last species vanishing during the Cretaceous–Paleogene extinction event.

Coleoidea

Coleoidea

Subclass Coleoidea, or Dibranchiata, is the grouping of cephalopods containing all the various taxa popularly thought of as "soft-bodied" or "shell-less". Unlike its extant sister group, Nautiloidea, whose members have a rigid outer shell for protection, the coleoids have at most an internal cuttlebone, gladius, or shell that is used for buoyancy or support. Some species have lost their cuttlebone altogether, while in some it has been replaced by a chitinous support structure. A unique trait of the group is the ability to edit their own RNA.

Octopus

Octopus

An octopus is a soft-bodied, eight-limbed mollusc of the order Octopoda. The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like other cephalopods, an octopus is bilaterally symmetric with two eyes and a beaked mouth at the center point of the eight limbs. The soft body can radically alter its shape, enabling octopuses to squeeze through small gaps. They trail their eight appendages behind them as they swim. The siphon is used both for respiration and for locomotion, by expelling a jet of water. Octopuses have a complex nervous system and excellent sight, and are among the most intelligent and behaviourally diverse of all invertebrates.

Monophyly

Monophyly

In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade — that is, a grouping of taxa which meets these criteria:the grouping contains its own most recent common ancestor, i.e. excludes non-descendants of that common ancestor the grouping contains all the descendants of that common ancestor, without exception

Clade

Clade

In biological phylogenetics, a clade, also known as a monophyletic group or natural group, is a grouping of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. In the taxonomical literature, sometimes the Latin form cladus is used rather than the English form.

Evolutionary grade

Evolutionary grade

A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit.

Taxonomic relationships

Nautiloids are among the group of animals known as cephalopods, an advanced class of mollusks which also includes ammonoids, belemnites and modern coleoids such as octopus and squid. Other mollusks include gastropods, scaphopods and bivalves.

Traditionally, the most common classification of the cephalopods has been a four-fold division (by Bather, 1888), into the orthoceratoids, nautiloids, ammonoids, and coleoids. This article is about nautiloids in that broad sense, sometimes called Nautiloidea sensu lato.

Cladistically speaking, nautiloids are a paraphyletic assemblage united by shared primitive (plesiomorphic) features not found in derived cephalopods. In other words, they are a grade group that is thought to have given rise to orthoceratoids, ammonoids and coleoids, and are defined by the exclusion of those descendent groups. Both ammonoids and coleoids have traditionally been assumed to have descended from bactritids, which in turn arose from straight-shelled orthoceratoids. The ammonoids appeared early in the Devonian period (some 400 million years ago) and became abundant in the Mesozoic era, before their extinction at the end of the Cretaceous.

Some workers apply the name Nautiloidea to a more exclusive group, called Nautiloidea sensu stricto. This taxon consists only of those orders that are clearly related to the modern nautilus to the exclusion of other modern cephalopods. In this restricted defition, membership is somewhat variable between authors, but it usually includes Tarphycerida, Oncocerida, and Nautilida.

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Mollusca

Mollusca

Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks. Around 85,000 extant species of molluscs are recognized. The number of fossil species is estimated between 60,000 and 100,000 additional species. The proportion of undescribed species is very high. Many taxa remain poorly studied.

Ammonoidea

Ammonoidea

Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids than they are to shelled nautiloids such as the living Nautilus species. The earliest ammonites appeared during the Devonian, with the last species vanishing during the Cretaceous–Paleogene extinction event.

Belemnoidea

Belemnoidea

Belemnoids are an extinct group of marine cephalopod, very similar in many ways to the modern squid and closely related to the modern cuttlefish. Like them, the belemnoids possessed an ink sac, but, unlike the squid, they possessed ten arms of roughly equal length, and no tentacles. The name "belemnoid" comes from the Greek word βέλεμνον, belemnon meaning "a dart or arrow" and the Greek word είδος, eidos meaning "form".

Coleoidea

Coleoidea

Subclass Coleoidea, or Dibranchiata, is the grouping of cephalopods containing all the various taxa popularly thought of as "soft-bodied" or "shell-less". Unlike its extant sister group, Nautiloidea, whose members have a rigid outer shell for protection, the coleoids have at most an internal cuttlebone, gladius, or shell that is used for buoyancy or support. Some species have lost their cuttlebone altogether, while in some it has been replaced by a chitinous support structure. A unique trait of the group is the ability to edit their own RNA.

Gastropoda

Gastropoda

The gastropods, commonly known as slugs and snails, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda.

Bivalvia

Bivalvia

Bivalvia, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class of marine and freshwater molluscs that have laterally compressed bodies enclosed by a shell consisting of two hinged parts. As a group, bivalves have no head and they lack some usual molluscan organs, like the radula and the odontophore. The class includes the clams, oysters, cockles, mussels, scallops, and numerous other families that live in saltwater, as well as a number of families that live in freshwater. The majority are filter feeders. The gills have evolved into ctenidia, specialised organs for feeding and breathing. Most bivalves bury themselves in sediment, where they are relatively safe from predation. Others lie on the sea floor or attach themselves to rocks or other hard surfaces. Some bivalves, such as the scallops and file shells, can swim. The shipworms bore into wood, clay, or stone and live inside these substances.

Orthoceratoidea

Orthoceratoidea

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.

Cladistics

Cladistics

Cladistics is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived characteristics (synapomorphies) that are not present in more distant groups and ancestors. However, from an empirical perspective, common ancestors are inferences based on a cladistic hypothesis of relationships of taxa whose character states can be observed. Theoretically, a last common ancestor and all its descendants constitute a (minimal) clade. Importantly, all descendants stay in their overarching ancestral clade. For example, if the terms worms or fishes were used within a strict cladistic framework, these terms would include humans. Many of these terms are normally used paraphyletically, outside of cladistics, e.g. as a 'grade', which are fruitless to precisely delineate, especially when including extinct species. Radiation results in the generation of new subclades by bifurcation, but in practice sexual hybridization may blur very closely related groupings.

Evolutionary grade

Evolutionary grade

A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit.

Bactritida

Bactritida

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 Pragian stage before 409 million years ago, and persisted until 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.

Devonian

Devonian

The Devonian is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, 419.2 million years ago (Mya), to the beginning of the Carboniferous, 358.9 Mya. It is named after Devon, England, where rocks from this period were first studied.

Mesozoic

Mesozoic

The Mesozoic Era is the second-to-last era of Earth's geological history, lasting from about 252 to 66 million years ago, comprising the Triassic, Jurassic and Cretaceous Periods. It is characterized by the dominance of archosaurian reptiles, like the dinosaurs; an abundance of conifers and ferns; a hot greenhouse climate; and the tectonic break-up of Pangaea. The Mesozoic is the middle of the three eras since complex life evolved: the Paleozoic, the Mesozoic, and the Cenozoic.

Shell

A cross-section of a Nautilus pompilius shell, showing the large body chamber, shrinking camerae, concave septa, and septal necks (partial siphuncle supports)
A cross-section of a Nautilus pompilius shell, showing the large body chamber, shrinking camerae, concave septa, and septal necks (partial siphuncle supports)

All nautiloids have a large external shell, divided into a narrowing chambered region (the phragmocone) and a broad, open body chamber occupied by the animal in life. The outer wall of the shell, also known as the conch, defines its overall shape and texture. The chambers (camerae) of the phragmocone are separated from each other by thin curved walls (septa), which formed during growth spurts of the animal. During a growth spurt, the rear of the mantle secretes a new septum, adding another chamber to the series of shell chambers. At the same time, shell material is added around the shell opening (aperture), enlarging the body chamber and providing more room for the growing animal. Sutures (or suture lines) appear where each septum contacts the wall of the outer shell. In life, they are visible as a series of narrow wavy lines on the outer surface of the shell. Like their underlying septa, the sutures of the nautiloids are simple in shape, being either straight or slightly curved. This is different from the "zigzag" sutures of the goniatites and the highly complex sutures of the ammonites.

The septa are perforated by the siphuncle, a fleshy tube which runs through each of the internal chambers of the shell. Surrounding the fleshy tube of the siphuncle are structures made of aragonite (a polymorph of calcium carbonate – which during fossilisation is often recrystallized to calcite, a more stable form of calcium carbonate [CaCO3]): septal necks and connecting rings. Some of the earlier nautiloids deposited calcium carbonate in the empty chambers (called cameral deposits) or within the siphuncle (endosiphuncular deposits), a process which may have been connected with controlling buoyancy. The nature of the siphuncle and its position within the shell are important in classifying nautiloids and can help distinguish them from ammonoids. The siphuncle is on the shell periphery in most ammonoids whereas it runs through the center of the chambers in some nautiloids, including living nautiluses.[1]

The subclass Nautiloidea, in its broader definition, is distinguished from other cephalopods by two main characteristics: the septa are smoothly concave in the forward direction, producing external sutures which are generally simple and smooth. The siphuncle is supported by septal necks which point to the rear (i.e. retrosiphonate) throughout the ontogeny of the animal.

Modern nautiluses have deeply coiled shells which are involute, meaning that the larger and more recent whorls overlap and obscure older whorls. The shells of fossil nautiloids may be either straight (i.e., orthoconic as in Orthoceras and Rayonnoceras), curved (as in Cyrtoceras) coiled (as in Cenoceras), or rarely a helical coil (as in Lorieroceras). Some species' shells—especially in the late Paleozoic and early Mesozoic—are ornamented with spines and ribs, but most have a smooth shell. The shells are formed of aragonite, although the cameral deposits may consist of primary calcite.[2] The coloration of the shell of the modern nautilus is quite prominent, and, although somewhat rarely, the shell coloration has been known to be preserved in fossil nautiloids. They often show color patterns only on the dorsal side, suggesting that the living animals swam horizontally.

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Phragmocone

Phragmocone

The phragmocone is the chambered portion of the shell of a cephalopod. It is divided by septa into camerae.

Camera (cephalopod)

Camera (cephalopod)

Camerae are the spaces or chambers enclosed between two adjacent septa in the phragmocone of a nautiloid or ammonoid cephalopod molluscus. These can be seen in cross-sections of a nautilus shell and in the polished cross-sections of ammonites. In life these chambers are filled with gas, mediated by the siphuncle, and used to control buoyancy.

Mantle (mollusc)

Mantle (mollusc)

The mantle is a significant part of the anatomy of molluscs: it is the dorsal body wall which covers the visceral mass and usually protrudes in the form of flaps well beyond the visceral mass itself.

Aperture (mollusc)

Aperture (mollusc)

The aperture is an opening in certain kinds of mollusc shells: it is the main opening of the shell, where the head-foot part of the body of the animal emerges for locomotion, feeding, etc.

Aragonite

Aragonite

Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, CaCO3. It is formed by biological and physical processes, including precipitation from marine and freshwater environments.

Buoyancy

Buoyancy

Buoyancy, or upthrust, is an upward force exerted by a fluid that opposes the weight of a partially or fully immersed object. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus the pressure at the bottom of a column of fluid is greater than at the top of the column. Similarly, the pressure at the bottom of an object submerged in a fluid is greater than at the top of the object. The pressure difference results in a net upward force on the object. The magnitude of the force is proportional to the pressure difference, and is equivalent to the weight of the fluid that would otherwise occupy the submerged volume of the object, i.e. the displaced fluid.

Ontogeny

Ontogeny

Ontogeny is the origination and development of an organism, usually from the time of fertilization of the egg to adult. The term can also be used to refer to the study of the entirety of an organism's lifespan.

Orthocone

Orthocone

An orthocone is an unusually long straight shell of a nautiloid cephalopod. During the 18th and 19th centuries, all shells of this type were named Orthoceras, creating a wastebasket taxon, but it is now known that many groups of nautiloids developed or retained this type of shell.

Orthoceras

Orthoceras

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.

Cyrtoceras

Cyrtoceras

Cyrtoceras is an extinct genus of oncoceridan nautiloids that lived from the middle Ordovician to the middle Devonian, in Africa, Europe, North America and South America.

Cenoceras

Cenoceras

Cenoceras is an extinct genus within the cephalopod mollusc family Nautilidae, which in turn makes up part of the superfamily Nautilaceae. This genus has been described by Hyatt in 1884. The type species is Cenoceras intermedium (Sowerby).

Lorieroceras

Lorieroceras

Lorieroceras lorieri is an extinct oncocerid nautilitoidean cephalopod belonging to the family Nothoceratidae. Fossils are found in Lower Devonian marine strata of France. Its shell is unusual among nautiloids in that it is a loosely coiled, turban-shaped helix that is sinistrally coiled.

Modern nautiloids

Much of what is known about the extinct nautiloids is based on what we know about modern nautiluses, such as the chambered nautilus, which is found in the southwest Pacific Ocean from Samoa to the Philippines, and in the Indian Ocean off the coast of Australia. It is not usually found in waters less than 100 meters (328 feet) deep and may be found as far down as 500 to 700 meters (1,600 to 2,300 feet).

Nautili are free swimming animals that possess a head with two simple lens-free eyes and arms (or tentacles). They have a smooth shell over a large body chamber, which is divided into subchambers filled with an inert gas (similar to the composition of atmospheric air, but with more nitrogen and less oxygen) making the animal neutrally buoyant in the water. As many as 90 tentacles are arranged in two circles around the mouth. The animal is predatory, and has jaws which are horny and beak-like, allowing it to feed on crustaceans.

Empty nautilus shells may drift a considerable distance and have been reported from Japan, India and Africa. Undoubtedy the same applies to the shells of fossil nautiloids, the gas inside the shell keeping it buoyant for some time after the animal's death, allowing the empty shell to be carried some distance from where the animal lived before finally sinking to the seafloor.

Nautili propel themselves by jet propulsion, expelling water from an elongated funnel called the hyponome, which can be pointed in different directions to control their movement. Unlike the belemnites and other cephalopods, modern nautili do not have an ink sac, and there is no evidence to suggest that the extinct forms possessed one either. Furthermore, unlike the extinct ammonoids, the modern nautilus lacks an aptychus, a biomineralized plate which is proposed to act as an operculum which closes the shell to protect the body. However, aptychus-like plates are known from some extinct nautiloids, and they may be homologous to the fleshy hood of a modern nautilus.[3]

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Nautilus

Nautilus

The nautilus is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina.

Nautilus belauensis

Nautilus belauensis

The Palau nautilus, Nautilus belauensis, is mainly found in Palau in the Western Carolines as its name suggests. It can be found on fore reef slopes in depths ranging from 95m to 504m but typically prefers to remain within 150-300m where water temperatures range between 16.6 to 9.4ºC. N. belauensis are highly mobile epibenthic scavenging herbalist and opportunistic predators that rely mostly on scent detection. They are active both diurnally and nocturnally within the preferred depth range although most shallow-water-incursions are usually nocturnal events that coincide with greatly diminished fish activities.

Chambered nautilus

Chambered nautilus

The chambered nautilus, also called the pearly nautilus, is the best-known species of nautilus. The shell, when cut away, reveals a lining of lustrous nacre and displays a nearly perfect equiangular spiral, although it is not a golden spiral. The shell exhibits countershading, being light on the bottom and dark on top. This is to help avoid predators, because when seen from above, it blends in with the darkness of the sea, and when seen from below, it blends in with the light coming from above.

Indian Ocean

Indian Ocean

The Indian Ocean is the third-largest of the world's five oceanic divisions, covering 70,560,000 km2 (27,240,000 sq mi) or ~19.8% of the water on Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia to the east. To the south it is bounded by the Southern Ocean or Antarctica, depending on the definition in use. Along its core, the Indian Ocean has some large marginal or regional seas such as the Arabian Sea, Laccadive Sea, Bay of Bengal, and Andaman Sea.

Australia

Australia

Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands. Australia is the largest country by area in Oceania and the world's sixth-largest country. Australia is the oldest, flattest, and driest inhabited continent, with the least fertile soils. It is a megadiverse country, and its size gives it a wide variety of landscapes and climates, with deserts in the centre, tropical rainforests in the north-east, and mountain ranges in the south-east.

Nitrogen

Nitrogen

Nitrogen is the chemical element with the symbol N and atomic number 7. Nitrogen is a nonmetal and the lightest member of group 15 of the periodic table, often called the pnictogens. It is a common element in the universe, estimated at seventh in total abundance in the Milky Way and the Solar System. At standard temperature and pressure, two atoms of the element bond to form N2, a colorless and odorless diatomic gas. N2 forms about 78% of Earth's atmosphere, making it the most abundant uncombined element. Nitrogen occurs in all organisms, primarily in amino acids (and thus proteins), in the nucleic acids (DNA and RNA) and in the energy transfer molecule adenosine triphosphate. The human body contains about 3% nitrogen by mass, the fourth most abundant element in the body after oxygen, carbon, and hydrogen. The nitrogen cycle describes the movement of the element from the air, into the biosphere and organic compounds, then back into the atmosphere.

Crustacean

Crustacean

Crustaceans belong to the subphylum Crustacea,, and form a large, diverse group of arthropods including decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can be treated as a subphylum under the clade Mandibulata. It is now well accepted that the hexapods emerged deep in the Crustacean group, with the completed group referred to as Pancrustacea. Some crustaceans are more closely related to insects and the other hexapods than they are to certain other crustaceans.

Japan

Japan

Japan is an island country in East Asia. It is situated in the northwest Pacific Ocean and is bordered on the west by the Sea of Japan, extending from the Sea of Okhotsk in the north toward the East China Sea, Philippine Sea, and Taiwan in the south. Japan is a part of the Ring of Fire, and spans an archipelago of 14,125 islands, with the five main islands being Hokkaido, Honshu, Shikoku, Kyushu, and Okinawa. Tokyo is the nation's capital and largest city, followed by Yokohama, Osaka, Nagoya, Sapporo, Fukuoka, Kobe, and Kyoto.

India

India

India, officially the Republic of India, is a country in South Asia. It is the seventh-largest country by area and the second-most populous country. Bounded by the Indian Ocean on the south, the Arabian Sea on the southwest, and the Bay of Bengal on the southeast, it shares land borders with Pakistan to the west; China, Nepal, and Bhutan to the north; and Bangladesh and Myanmar to the east. In the Indian Ocean, India is in the vicinity of Sri Lanka and the Maldives; its Andaman and Nicobar Islands share a maritime border with Thailand, Myanmar, and Indonesia.

Africa

Africa

Africa is the world's second-largest and second-most populous continent, after Asia in both aspects. At about 30.3 million km2 including adjacent islands, it covers 20% of Earth's land area and 6% of its total surface area. With 1.4 billion people as of 2021, it accounts for about 18% of the world's human population. Africa's population is the youngest amongst all the continents; the median age in 2012 was 19.7, when the worldwide median age was 30.4. Despite a wide range of natural resources, Africa is the least wealthy continent per capita and second-least wealthy by total wealth, behind Oceania. Scholars have attributed this to different factors including geography, climate, tribalism, colonialism, the Cold War, neocolonialism, lack of democracy, and corruption. Despite this low concentration of wealth, recent economic expansion and the large and young population make Africa an important economic market in the broader global context.

Fossil

Fossil

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

Belemnitida

Belemnitida

Belemnitida is an extinct order of squid-like cephalopods that existed from the Late Triassic to Late Cretaceous. Unlike squid, belemnites had an internal skeleton that made up the cone. The parts are, from the arms-most to the tip: the tongue-shaped pro-ostracum, the conical phragmocone, and the pointy guard. The calcitic guard is the most common belemnite remain. Belemnites, in life, are thought to have had 10 hooked arms and a pair of fins on the guard. The chitinous hooks were usually no bigger than 5 mm (0.20 in), though a belemnite could have had between 100 and 800 hooks in total, using them to stab and hold onto prey.

Fossil record

Fossil nautiloid Trilacinoceras from the Ordovician of China.
Fossil nautiloid Trilacinoceras from the Ordovician of China.
Fossil orthoconic nautiloid from the Ordovician of Kentucky; an internal mold showing siphuncle and half-filled camerae, both encrusted.
Fossil orthoconic nautiloid from the Ordovician of Kentucky; an internal mold showing siphuncle and half-filled camerae, both encrusted.
Cross-section of an Orthoceras nautiloid from the Siluro-Devonian of Erfoud, Morocco.
Cross-section of an Orthoceras nautiloid from the Siluro-Devonian of Erfoud, Morocco.

Nautiloids are often found as fossils in early Palaeozoic rocks (less so in more recent strata).

The rocks of the Ordovician period in the Baltic coast and parts of the United States contain a variety of nautiloid fossils, and specimens such as Discitoceras and Rayonnoceras may be found in the limestones of the Carboniferous period in Ireland. The marine rocks of the Jurassic period in Britain often yield specimens of Cenoceras, and nautiloids such as Eutrephoceras are also found in the Pierre Shale formation of the Cretaceous period in the north-central United States.

Specimens of the Ordovician nautiloid Endoceras have been recorded measuring up to 5.7 meters (19 feet) in shell length, and there is a description of a specimen estimated to have reached 9.1 meters (30 feet), although that specimen is reported as destroyed.[4] These large nautiloids would have been formidable predators of other marine animals at the time they lived.

In some localities, such as Scandinavia and Morocco, the fossils of orthoconic nautiloids accumulated in such large numbers that they form limestones composed of nonspecific assemblages known as cephalopod beds, cephalopod limestones, nautiloid limestones, or Orthoceras limestones in the geological literature. Although the term Orthoceras now only refers to a Baltic coast Ordovician genus, in prior times it was employed as a general name given to all straight-shelled nautiloids that lived from the Ordovician to the Triassic periods (but were most common in the early Paleozoic era).

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China

China

China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and borders fourteen countries by land, the most of any country in the world, tied with Russia. With an area of approximately 9.6 million square kilometres (3,700,000 sq mi), it is the world's third largest country by total land area. The country consists of 22 provinces, five autonomous regions, four municipalities, and two special administrative regions. The national capital is Beijing, and the most populous city and largest financial center is Shanghai.

Kentucky

Kentucky

Kentucky, officially the Commonwealth of Kentucky, is a state in the Southeastern region of the United States and one of the states of the Upper South. It borders Illinois, Indiana, and Ohio to the north, West Virginia to the northeast, Virginia to the east, Tennessee to the south, and Missouri to the west. Its northern border is defined by the Ohio River. Its capital is Frankfort, and its two largest cities are Louisville and Lexington. Its population was approximately 4.5 million in 2020.

Fossil

Fossil

A fossil is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in amber, hair, petrified wood and DNA remnants. The totality of fossils is known as the fossil record.

Baltic Sea

Baltic Sea

The Baltic Sea is an arm of the Atlantic Ocean that is enclosed by Denmark, Estonia, Finland, Germany, Latvia, Lithuania, Poland, Russia, Sweden and the North and Central European Plain.

Discitoceras

Discitoceras

Discitoceras is an extinct genus of nautiloids from the Lower Carboniferous.

Limestone

Limestone

Limestone is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of CaCO3. Limestone forms when these minerals precipitate out of water containing dissolved calcium. This can take place through both biological and nonbiological processes, though biological processes, such as the accumulation of corals and shells in the sea, have likely been more important for the last 540 million years. Limestone often contains fossils which provide scientists with information on ancient environments and on the evolution of life.

Carboniferous

Carboniferous

The Carboniferous is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period 358.9 million years ago (Mya), to the beginning of the Permian Period, 298.9 million years ago. The name Carboniferous means "coal-bearing", from the Latin carbō ("coal") and ferō, and refers to the many coal beds formed globally during that time.

Jurassic

Jurassic

The Jurassic is a geologic period and stratigraphic system that spanned from the end of the Triassic Period 201.4 million years ago (Mya) to the beginning of the Cretaceous Period, approximately 145 Mya. The Jurassic constitutes the middle period of the Mesozoic Era and is named after the Jura Mountains, where limestone strata from the period were first identified.

Eutrephoceras

Eutrephoceras

Eutrephoceras is an extinct genus of nautilus from the Late Jurassic to the Miocene. They are characterized by a highly rounded involute shell with slightly sinuous suture patterns.

Cretaceous

Cretaceous

The Cretaceous is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of the entire Phanerozoic. The name is derived from the Latin creta, "chalk", which is abundant in the latter half of the period. It is usually abbreviated K, for its German translation Kreide.

Endoceras

Endoceras

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 485 to 419 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.

Morocco

Morocco

Morocco, officially the Kingdom of Morocco, is a country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to the east, and the disputed territory of Western Sahara to the south. Mauritania lies to the south of Western Sahara. Morocco also claims the Spanish exclaves of Ceuta, Melilla and Peñón de Vélez de la Gomera, and several small Spanish-controlled islands off its coast. It spans an area of 446,300 km2 (172,300 sq mi) or 710,850 km2 (274,460 sq mi), with a population of roughly 37 million. Its official and predominant religion is Islam, and the official languages are Arabic and Berber; the Moroccan dialect of Arabic and French are also widely spoken. Moroccan identity and culture is a mix of Arab, Berber, and European cultures. Its capital is Rabat, while its largest city is Casablanca.

Evolutionary history

Nautiloids are first known from the late Cambrian Fengshan Formation of northeastern China, where they seem to have been quite diverse (at the time this was a warm shallow sea rich in marine life). However, although four orders have been proposed from the 131 species named, there is no certainty that all of these are valid, and indeed it is likely that these taxa are seriously oversplit.

Life reconstruction of Cyrtoceras sp
Life reconstruction of Cyrtoceras sp
Gyronaedyceras eryx, an oncocerid from the Middle Devonian of Wisconsin
Gyronaedyceras eryx, an oncocerid from the Middle Devonian of Wisconsin
Acleistoceras whitfieldi, an oncocerid from the Middle Devonian of Wisconsin
Acleistoceras whitfieldi, an oncocerid from the Middle Devonian of Wisconsin
Trocholites, an tarphycerid from the Upper Ordovician of Wisconsin
Trocholites, an tarphycerid from the Upper Ordovician of Wisconsin

Most of these early forms died out, but a single family, the Ellesmeroceratidae, survived to the early Ordovician, where it ultimately gave rise to all subsequent cephalopods. In the Early and Middle Ordovician the nautiloids underwent an evolutionary radiation.[5][6] Some eight new orders appeared at this time, covering a great diversity of shell types and structure, and ecological lifestyles.

Nautiloids remained at the height of their range of adaptations and variety of forms throughout the Ordovician, Silurian, and Devonian periods, with various straight, curved and coiled shell forms coexisting at the same time. Several of the early orders became extinct over that interval, but others rose to prominence.

Nautiloids began to decline in the Devonian, perhaps due to competition with their descendants and relatives the Ammonoids and Coleoids, with only the Nautilida holding their own (and indeed increasing in diversity). Their shells became increasingly tightly coiled, while both numbers and variety of non-nautilid species continued to decrease throughout the Carboniferous and Permian.

The massive extinctions at the end of the Permian were less damaging to nautiloids than to other taxa and a few groups survived into the early Mesozoic, including pseudorthocerids, bactritids, nautilids and possibly orthocerids. The last straight-shelled forms were long thought to have disappeared at the end of the Triassic, but a possible orthocerid has been found in Cretaceous rocks. Apart from this exception, only a single nautiloid suborder, the Nautilina, continued throughout the Mesozoic, where they co-existed quite happily with their more specialised ammonoid cousins. Most of these forms differed only slightly from the modern nautilus. They had a brief resurgence in the early Tertiary (perhaps filling the niches vacated by the ammonoids in the end Cretaceous extinction), and maintained a worldwide distribution up until the middle of the Cenozoic Era. With the global cooling of the Miocene and Pliocene, their geographic distribution shrank and these hardy and long-lived animals declined in diversity again. Today there are only six living species, all belonging to two genera, Nautilus (the pearly nautilus), and Allonautilus.

Timeline of orders

CenozoicMesozoic EraPaleozoic EraQuaternaryNeogenePaleogeneCretaceousJurassicTriassicPermianCarboniferousDevonianSilurianOrdovicianCambrianNautilidaDiscosoridaOncoceridaTarphyceridaEndoceridaActinoceridaEllesmeroceridaPlectronoceridaCenozoicMesozoic EraPaleozoic EraQuaternaryNeogenePaleogeneCretaceousJurassicTriassicPermianCarboniferousDevonianSilurianOrdovicianCambrian


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China

China

China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and borders fourteen countries by land, the most of any country in the world, tied with Russia. With an area of approximately 9.6 million square kilometres (3,700,000 sq mi), it is the world's third largest country by total land area. The country consists of 22 provinces, five autonomous regions, four municipalities, and two special administrative regions. The national capital is Beijing, and the most populous city and largest financial center is Shanghai.

Ellesmeroceratidae

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 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).

Ordovician

Ordovician

The Ordovician is a geologic period and system, the second of six periods of the Paleozoic Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period 485.4 million years ago (Mya) to the start of the Silurian Period 443.8 Mya.

Devonian

Devonian

The Devonian is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, 419.2 million years ago (Mya), to the beginning of the Carboniferous, 358.9 Mya. It is named after Devon, England, where rocks from this period were first studied.

Nautilida

Nautilida

The Nautilida constitute a large and diverse order of generally coiled nautiloid cephalopods that began in the mid Paleozoic and continues to the present with a single family, the Nautilidae which includes two genera, Nautilus and Allonautilus, with six species. All told, between 22 and 34 families and 165 to 184 genera have been recognised, making this the largest order of the subclass Nautiloidea.

Carboniferous

Carboniferous

The Carboniferous is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period 358.9 million years ago (Mya), to the beginning of the Permian Period, 298.9 million years ago. The name Carboniferous means "coal-bearing", from the Latin carbō ("coal") and ferō, and refers to the many coal beds formed globally during that time.

Permian

Permian

The Permian is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period 298.9 million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleozoic Era; the following Triassic Period belongs to the Mesozoic Era. The concept of the Permian was introduced in 1841 by geologist Sir Roderick Murchison, who named it after the region of Perm in Russia.

Mesozoic

Mesozoic

The Mesozoic Era is the second-to-last era of Earth's geological history, lasting from about 252 to 66 million years ago, comprising the Triassic, Jurassic and Cretaceous Periods. It is characterized by the dominance of archosaurian reptiles, like the dinosaurs; an abundance of conifers and ferns; a hot greenhouse climate; and the tectonic break-up of Pangaea. The Mesozoic is the middle of the three eras since complex life evolved: the Paleozoic, the Mesozoic, and the Cenozoic.

Bactritida

Bactritida

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 Pragian stage before 409 million years ago, and persisted until 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.

Orthocerida

Orthocerida

Orthocerida is an order of extinct Orthoceratoid cephalopods also known as the Michelinocerida 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. They were most common however from the Ordovician to the Devonian.

Cretaceous

Cretaceous

The Cretaceous is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of the entire Phanerozoic. The name is derived from the Latin creta, "chalk", which is abundant in the latter half of the period. It is usually abbreviated K, for its German translation Kreide.

Nautilina

Nautilina

The Nautilina is the last suborder of the Nautilida and the only nautiloids living since the end of the Triassic. The Nautilina, proposed by Shimanskiy, is basically the Nautilaceae of Kummel, 1964, defined by Furnish and Glenister, but differs in omitting two families, the Paracenoceratidae and Pseudonautilidae which instead are placed in the Liroceratina.

Classification

Older classification systems

A consensus on nautiloid classification has traditionally been elusive and subject to change, as different workers emphasize different fundamental traits when reconstructing evolutionary events. The largest and most widely cited publication on nautiloid taxonomy is the Treatise on Invertebrate Paleontology Part K by Teichert et al. 1964, though new information has rendered this volume outdated and in need of revision. Treatise Part K was based on previous classification schemes by Flower & Kummel (1950) and the Russian Osnovy Paleontologii Vol. 5 (1962) textbook.

Other comprehensive taxonomic schemes have been devised by Wade (1988), Teichert (1988), and Shevyrev (2006). Wade (1988) divided the subclass Nautiloidea (sensu lato) into 6 superorders, incorporating orders that are phylogenetically related. They are:

Three of these superorders were established for orders of uncertain placement: Endocerida, Actinocerida, and Discosorida. The other three unite related orders which share a common ancestor and form a branch of the nautiloid taxonomic tree: Plectronoceratoidea, which consists mostly of small Cambrian forms that include the ancestors of subsequent stocks; Orthoceratoidea, which unites different primarily orthoconic orders (including the ancestors for Bacritida and Ammonoidea); and Nautilitoidea, which includes the first coiled cephalopods, Tarphycerida, as well as Nautilida, which includes the recent Nautilus. Another order, Bactritida, which is derived from Orthocerida, is sometimes included with Nautiloidea, sometimes with Ammonoidea, and sometimes placed in a subclass of its own, Bactritoidea.

Recently some workers in the field have come to recognize Dissidocerida as a distinct order, along with Pseudorthocerida, both previously included in Orthocerida as subtaxa.

Early cladistic efforts

Cladistic approaches are rare in nautiloid systematics. Many nautiloid orders (not to mention the group as a whole) are not monophyletic clades, but rather paraphyletic grades. This means that they include some descendant taxa while excluding others. For example, the paraphyletic order Orthocerida includes numerous orthocerids stretching through the Paleozoic, but it excludes coleoids, despite coleoids having a well-established ancestry among the orthocerids. Interpretations by Engeser (1996-1998) suggests that nautiloids, and indeed cephalopods in general, should be split into two main clades: Palcephalopoda (including all the nautiloids except Orthocerida and Ascocerida) and Neocephalopoda (the rest of the cephalopods). Palcephalopoda is meant to correspond to groups which are closer to living nautilus, while Neocephalopoda is meant to correspond to groups closer to living coleoids. One issue which this scheme is the necessity of establishing a firm ancestry for nautilus, to contextualize which cephalopods are closer to which of the two living end members. On the basis of morphological traits, Nautilida is most similar to coiled early nautiloids such as the Tarphycerida and Oncocerida. However, these orders diverged from coleoid ancestors in the early Ordovician at the latest, while genetic divergence estimates suggest that Nautilida diverged in the Silurian or Devonian.[7]

A more recent phylogenetic study by Lindgren et al. (2004), which supports the monophyly of cephalopods, does not bear on the Palcephalopod/Neocephalopod question, since the only cephalopods included were Nautilus and coleoids.[8]

Recent revisions

For an in-process revision of Treatise Part K, King & Evans (2019) reclassified nautiloids sensu lato into five subclasses. Major groups were primarily defined by variation in their muscle attachment types. Other traits referenced during this reclassification include protoconch morphology, connecting ring structure, and the extent of cameral and endosiphuncular deposits. While most previous studies referred to subclasses with the suffix '-oidea', these authors instead opted for the suffix '-ia', to prevent confusion between group levels. For example, Nautiloidea sensu stricto was renamed to Nautilia, to differentiate it from the informal broader definition of "nautiloid". In addition, they used the unsimplified names for orders, with the suffix '-atida' rather than the common simplified form, '-ida'.[9]

Nautiloid relationships and body types according to Pohle et al. (2022)
Nautiloid relationships and body types according to Pohle et al. (2022)

Traditional nautiloid classification schemes emphasize certain character traits over others, potentially involving personal bias as to which traits are worth emphasizing according to different authors. This issue may be resolved by sampling all morphological traits equally through bayesian phylogenetic inference. The first cephalopod-focused paper to use this technique was published by Pohle et al. (2022). They recovered several previously hypothesized groups, though many orders were determined to be paraphyletic. The study was focused on early cephalopod diversification in the Late Cambrian and Ordovician, and did not discuss in detail the origin of post-Ordovician groups. The following is a simplified version of their cladogram, showing early cephalopod relationships to the order level (although various isolated families also originated during this diversification event):[7]

Cephalopoda

Plectronocerida

Yanhecerida

Ellesmerocerida (paraphyletic to Endoceratoidea, Multiceratoidea, and Orthoceratoidea)

Endoceratoidea

Endocerida

Bisonocerida

Multiceratoidea

Cyrtocerinida

Tarphycerida (possibly paraphyletic to Nautilida)

Oncocerida (paraphyletic to Ascocerida and Discosorida)

Ascocerida

Discosorida

Orthoceratoidea

Riocerida (possibly paraphyletic to later orthoceratoids)

Dissidocerida (paraphyletic to later orthoceratoids)

Lituitida

Actinocerida

Pseudorthocerida

Orthocerida (paraphyletic to Bactritida, Ammonoidea, and Coleoidea)

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Plectronoceratoidea

Plectronoceratoidea

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.

Ellesmerocerida

Ellesmerocerida

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.

Endocerida

Endocerida

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.

Orthoceratoidea

Orthoceratoidea

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.

Orthocerida

Orthocerida

Orthocerida is an order of extinct Orthoceratoid cephalopods also known as the Michelinocerida 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. They were most common however from the Ordovician to the Devonian.

Ascocerida

Ascocerida

The Ascocerida are comparatively small, bizarre Orthoceratoids known only from Ordovician and Silurian sediments in Europe and North America, uniquely characterized by a deciduous conch consisting of a longiconic juvenile portion and an inflated breviconic adult portion that separate sometime in maturity.

Nautilitoidea

Nautilitoidea

The Nautilitoidea is a superorder within the subclass Nautiloidea, comprising the phylogenetically related Nautilida, Oncocerida, and Tarphycerida.

Oncocerida

Oncocerida

The Oncocerida comprise a diverse group of generally small nautiloid cephalopods known from the Middle Ordovician to the Mississippian, in which the connecting rings are thin and siphuncle segments are variably expanded. At present the order consists of some 16 families, a few of which, such as the Oncoceratidae, Brevicoceratidae, and Acleistoceratidae contain a fair number of genera each while others like the Trimeroceratidae and Archiacoceratidae are represented by only two or three.

Nautilida

Nautilida

The Nautilida constitute a large and diverse order of generally coiled nautiloid cephalopods that began in the mid Paleozoic and continues to the present with a single family, the Nautilidae which includes two genera, Nautilus and Allonautilus, with six species. All told, between 22 and 34 families and 165 to 184 genera have been recognised, making this the largest order of the subclass Nautiloidea.

Actinocerida

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 deposits formed within contain a system of radial canals and a narrow space along the inner side of the connecting ring known as a paraspatium. Septal necks are generally short and cyrtochoanitic, some being recumbent, some hook shaped. Most grew to lengths of about 60 to 90 cm but some, like the Huroniidae of the Silurian grew significantly larger.

Discosorida

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 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.

Bactritida

Bactritida

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 Pragian stage before 409 million years ago, and persisted until 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.

Gallery

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Nautilus

Nautilus

The nautilus is a pelagic marine mollusc of the cephalopod family Nautilidae. The nautilus is the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina.

Allonautilus

Allonautilus

The genus Allonautilus contains two species of nautiluses, which have a significantly different morphology from those placed in the sister taxon Nautilus. Allonautilus is now thought to be a descendant of Nautilus, rendering the latter genus paraphyletic.

Endoceras

Endoceras

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 485 to 419 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.

Alaskoceras

Alaskoceras

Alaskoceras is a genus of lower Ordovician coiled nautiloid cephalopods; the shell moderately expanded, ribbed, with a divergent living chamber; whorl section more broadly rounded ventrally than dorsally; siphuncle marginal at maturity, septal necks short, almost achoanitic; connecting rings thick, layered.

Goldringia

Goldringia

Goldringia is an extinct nautilid of the Rutoceratidae family that lived during the Middle Devonian. It is known from New York, Ohio, and Indiana in the United States.

Devonian

Devonian

The Devonian is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, 419.2 million years ago (Mya), to the beginning of the Carboniferous, 358.9 Mya. It is named after Devon, England, where rocks from this period were first studied.

Lorieroceras

Lorieroceras

Lorieroceras lorieri is an extinct oncocerid nautilitoidean cephalopod belonging to the family Nothoceratidae. Fossils are found in Lower Devonian marine strata of France. Its shell is unusual among nautiloids in that it is a loosely coiled, turban-shaped helix that is sinistrally coiled.

Mesozoic

Mesozoic

The Mesozoic Era is the second-to-last era of Earth's geological history, lasting from about 252 to 66 million years ago, comprising the Triassic, Jurassic and Cretaceous Periods. It is characterized by the dominance of archosaurian reptiles, like the dinosaurs; an abundance of conifers and ferns; a hot greenhouse climate; and the tectonic break-up of Pangaea. The Mesozoic is the middle of the three eras since complex life evolved: the Paleozoic, the Mesozoic, and the Cenozoic.

Cenoceras

Cenoceras

Cenoceras is an extinct genus within the cephalopod mollusc family Nautilidae, which in turn makes up part of the superfamily Nautilaceae. This genus has been described by Hyatt in 1884. The type species is Cenoceras intermedium (Sowerby).

Eocene

Eocene

The Eocene Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name Eocene comes from the Ancient Greek ἠώς and καινός and refers to the "dawn" of modern ('new') fauna that appeared during the epoch.

Hoko River Formation

Hoko River Formation

The Hoko River Formation is a Late Eocene marine sedimentary geologic formation. The formation is exposed in outcrops along the Strait of Juan de Fuca on the Olympic Peninsula in Washington state, USA. It is known for containing numerous fossils of crabs. It overlies the older Lyre Formation and underlies the younger Makah Formation.

Cooperoceras

Cooperoceras

Cooperoceras is a genus of Tainoceratid nautiloid cephalopod molluscs within the superfamily Tainocerataceae, characterized by and evolute shell with an open, perforate, umbilicus, sinuous ribs at maturity, and recurved hollow spines along the ventro-lateral shoulders. The flanks and venter are flattened, the flanks converge on the dorsum, the venter has a shallow median groove. The suture is with rounded ventral and lateral lobes. The siphuncle is small, tubular, and subcentral.

Source: "Nautiloid", Wikipedia, Wikimedia Foundation, (2023, February 3rd), https://en.wikipedia.org/wiki/Nautiloid.

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References
  1. ^ "The Cephalopoda". The University of California Museum of Paleontology. Retrieved 2014-05-11.
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