cephalopod
A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...
s (
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 esti ...
) which originated in the
Late Cambrian
Late may refer to:
* LATE, an acronym which could stand for:
** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia
** Local-authority trading enterprise, a New Zealand business law
** Local average treatment effect, ...
and are represented today by the living ''
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.
It comprises six living species in ...
'' and ''
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 t ...
''.
Fossil
A fossil (from Classical Latin , ) 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 ...
nautiloids are diverse and speciose, with over 2,500 recorded species. They flourished during the early
Paleozoic
The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon.
The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838
by combining the Greek words ''palaiós'' (, "old") and ' ...
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
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 (i.e., octopuses, squid and cuttlefish) ...
squid
True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...
,
octopus
An octopus ( : octopuses or octopodes, see below for variants) 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, cuttle ...
, and kin). While ammonoids and coleoids are
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
clades
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, t ...
with exclusive ancestor-descendant relationships, this is not the case for nautiloids. Instead, nautiloids are a
paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
grade
Grade most commonly refers to:
* Grade (education), a measurement of a student's performance
* Grade, the number of the year a student has reached in a given educational stage
* Grade (slope), the steepness of a slope
Grade or grading may also ref ...
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.
Taxonomic relationships
Nautiloids are among the group of animals known as cephalopods, an advanced class of
mollusks
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 esti ...
which also includes
ammonoids
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 (i.e., octopuses, squid and cuttlefish) ...
, belemnites and modern coleoids such as octopus and squid. Other mollusks include
gastropods
The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda ().
This class comprises snails and slugs from saltwater, from freshwater, and from land. The ...
, scaphopods and bivalves.
Traditionally, the most common classification of the cephalopods has been a four-fold division (by Bather, 1888), into the orthoceratoids, nautiloids,
ammonoid
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 (i.e., octopuses, squid and cuttlefish) ...
s, and
coleoid
Subclass Coleoidea,
or Dibranchiata, is the grouping of cephalopods containing all the various taxa popularly thought of as "soft-bodied" or "shell-less" (i.e., octopuses, squid and cuttlefish). Unlike its extant sister group, Nautiloidea, whose ...
s. This article is about nautiloids in that broad sense, sometimes called Nautiloidea ''sensu lato''.
Cladistically
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 char ...
speaking, nautiloids are a
paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
assemblage united by shared primitive ( plesiomorphic) features not found in derived cephalopods. In other words, they are a
grade
Grade most commonly refers to:
* Grade (education), a measurement of a student's performance
* Grade, the number of the year a student has reached in a given educational stage
* Grade (slope), the steepness of a slope
Grade or grading may also ref ...
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-shelledorthoceratoids. The ammonoids appeared early in the
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
period (some 400 million years ago) and became abundant in the
Mesozoic
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
era
An era is a span of time defined for the purposes of chronology or historiography, as in the regnal eras in the history of a given monarchy, a calendar era used for a given calendar, or the geological eras defined for the history of Earth.
Comp ...
, 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.
Shell
All nautiloids have a large external shell, divided into a narrowing chambered region (the
phragmocone
The phragmocone is the chambered portion of the shell of a cephalopod. It is divided by septa into camerae.
In most nautiloids and ammonoids, the phragmocone is a long, straight, curved, or coiled structure, in which the camarae are linked by a ...
) and a broad, open
body chamber
The body whorl is part of the morphology of the shell in those gastropod mollusks that possess a coiled shell. The term is also sometimes used in a similar way to describe the shell of a cephalopod mollusk.
In gastropods
In gastropods, the ...
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 (
camera
A camera is an Optics, optical instrument that can capture an image. Most cameras can capture 2D images, with some more advanced models being able to capture 3D images. At a basic level, most cameras consist of sealed boxes (the camera body), ...
e) of the phragmocone are separated from each other by thin curved walls (
septa
The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five c ...
), 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
In optics, an aperture is a hole or an opening through which light travels. More specifically, the aperture and focal length of an optical system determine the cone angle of a bundle of rays that come to a focus in the image plane.
An opt ...
), 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
Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including prec ...
(a polymorph of calcium carbonate – which during fossilisation is often recrystallized to calcite, a more stable form of calcium carbonate 3">aCO3: 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
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 p ...
. 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.
The subclass Nautiloidea, in its broader definition, is distinguished from other cephalopods by two main characteristics: the
septa
The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people in five c ...
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
Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
of the animal.
Modern nautiluses have deeply coiled shells which are involute, meaning that the larger and more recent
whorls
A whorl ( or ) is an individual circle, oval, volution or equivalent in a whorled pattern, which consists of a spiral or multiple concentric objects (including circles, ovals and arcs).
Whorls in nature
File:Photograph and axial plane floral ...
overlap and obscure older whorls. The shells of fossil nautiloids may be either straight (i.e.,
orthoconic
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 ...
as in ''
Orthoceras
''Orthoceras'' ("straight horn") 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 misspelle ...
'' 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. 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
Dorsal (from Latin ''dorsum'' ‘back’) may refer to:
* Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism
* Dorsal, positioned on top of an aircraft's fuselage
* Dorsal c ...
side, suggesting that the living animals swam horizontally.
Modern nautiloids
Much of what is known about the extinct nautiloids is based on what we know about modern
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.
It comprises six living species in ...
Pacific Ocean
The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the continen ...
from
Samoa
Samoa, officially the Independent State of Samoa; sm, Sāmoa, and until 1997 known as Western Samoa, is a Polynesian island country consisting of two main islands (Savai'i and Upolu); two smaller, inhabited islands (Manono Island, Manono an ...
to the
Philippines
The Philippines (; fil, Pilipinas, links=no), officially the Republic of the Philippines ( fil, Republika ng Pilipinas, links=no),
* bik, Republika kan Filipinas
* ceb, Republika sa Pilipinas
* cbk, República de Filipinas
* hil, Republ ...
, and in the
Indian Ocean
The Indian Ocean is the third-largest of the world's five oceanic divisions, covering 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 th ...
off the coast of
Australia
Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
. It is not usually found in waters less than 100 meters (328 feet) deep and may be found as far down as .
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
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 se ...
and less
oxygen
Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as wel ...
) making the animal neutrally buoyant in the water. As many as 90
tentacles
In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work main ...
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
crustacean
Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
s.
Empty nautilus shells may drift a considerable distance and have been reported from
Japan
Japan ( ja, 日本, or , and formally , ''Nihonkoku'') 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, while extending from the Sea of Okhotsk in the north ...
,
India
India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
and
Africa
Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
. Undoubtedy the same applies to the shells of
fossil
A fossil (from Classical Latin , ) 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 ...
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
A siphon is an anatomical structure which is part of the body of aquatic molluscs in three classes: Gastropoda, Bivalvia and Cephalopoda (members of these classes include saltwater and freshwater snails, clams, octopus, squid and relatives).
Si ...
, 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
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 (i.e., octopuses, squid and cuttlefish) ...
, the modern nautilus lacks an
aptychus
An aptychus is a type of marine fossil. It is a hard anatomical structure, a sort of curved shelly plate, now understood to be part of the body of an ammonite. Paired aptychi have, on rare occasions, been found at or within the aperture of ammo ...
, 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.
Fossil record
Nautiloids are often found as
fossil
A fossil (from Classical Latin , ) 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 ...
s in early Palaeozoic rocks (less so in more recent strata).
The rocks of the
Ordovician
The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
period in the
Baltic coast
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.
The sea stretches from 53°N to 66°N latitude and from 10 ...
and parts of the United States contain a variety of nautiloid fossils, and specimens such as '' Discitoceras'' and ''Rayonnoceras'' may be found in the
limestone
Limestone ( calcium carbonate ) 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 . Limestone forms whe ...
s of the
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
period in
Ireland
Ireland ( ; ga, Éire ; Ulster Scots dialect, Ulster-Scots: ) is an island in the Atlantic Ocean, North Atlantic Ocean, in Northwestern Europe, north-western Europe. It is separated from Great Britain to its east by the North Channel (Grea ...
. The marine rocks of the
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
period in
Britain
Britain most often refers to:
* The United Kingdom, a sovereign state in Europe comprising the island of Great Britain, the north-eastern part of the island of Ireland and many smaller islands
* Great Britain, the largest island in the United King ...
often yield specimens of ''Cenoceras'', and nautiloids such as ''
Eutrephoceras
''Eutrephoceras'' is an extinct genus of nautilus from the Late Jurassic to the Miocene (around 161 to 5 million years ago). They are characterized by a highly rounded involute shell with slightly sinuous suture patterns.
Description
''Eutrep ...
'' are also found in the Pierre Shale formation of the
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 th ...
period in the north-central United States.
Specimens of the
Ordovician
The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
nautiloid ''
Endoceras
''Endoceras'' (Ancient Greek for "inner horn") is an extinct genus of large, straight shelled cephalopods from the Middle and Upper Ordovician that gives its name to the Nautiloid order Endocerida. The cross section in the mature portion is slig ...
'' have been recorded measuring up to in shell length, and there is a description of a specimen estimated to have reached , although that specimen is reported as destroyed. These large nautiloids would have been formidable predators of other marine animals at the time they lived.
In some localities, such as
Scandinavia
Scandinavia; Sámi languages: /. ( ) is a subregion#Europe, subregion in Northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. In English usage, ''Scandinavia'' most commonly refers to Denmark, ...
and
Morocco
Morocco (),, ) officially the Kingdom of Morocco, is the westernmost 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
fossil
A fossil (from Classical Latin , ) 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 ...
s of
orthoconic
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 ...
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
''Orthoceras'' ("straight horn") 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 misspelle ...
'' now only refers to a
Baltic coast
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.
The sea stretches from 53°N to 66°N latitude and from 10 ...
Ordovician
The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
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
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
periods (but were most common in the early
Paleozoic
The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon.
The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838
by combining the Greek words ''palaiós'' (, "old") and ' ...
era).
Evolutionary history
Nautiloids are first known from the late Cambrian Fengshan Formation of northeastern
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 ...
, 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
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
named, there is no certainty that all of these are valid, and indeed it is likely that these taxa are seriously oversplit.
Most of these early forms died out, but a single family, 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 do ...
, survived to the early
Ordovician
The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
, where it ultimately gave rise to all subsequent cephalopods. In the Early and Middle Ordovician the nautiloids underwent an evolutionary radiation. 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
The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
, and
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
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
Coleoid
Subclass Coleoidea,
or Dibranchiata, is the grouping of cephalopods containing all the various taxa popularly thought of as "soft-bodied" or "shell-less" (i.e., octopuses, squid and cuttlefish). Unlike its extant sister group, Nautiloidea, whose ...
s, with only the
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 ''Allonauti ...
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
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
and
Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
.
The massive extinctions at the end of the Permian were less damaging to nautiloids than to other
taxa
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
and a few groups survived into the early
Mesozoic
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
, including pseudorthocerids, bactritids, nautilids and possibly orthocerids. The last straight-shelled forms were long thought to have disappeared at the end of the
Triassic
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
, but a possible orthocerid has been found in
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 th ...
rocks. Apart from this exception, only a single nautiloid suborder, the
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 ...
, continued throughout the
Mesozoic
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
, 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
Tertiary ( ) is a widely used but obsolete term for the geologic period from 66 million to 2.6 million years ago.
The period began with the demise of the non-avian dinosaurs in the Cretaceous–Paleogene extinction event, at the start ...
(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
The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
Era. With the global cooling of the
Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
and
Pliocene
The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Nautilus
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.
It comprises six living species in ...
'' (the pearly nautilus), and ''
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 t ...
Cambrian
The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
from: -488.3 till: -443.7 color:ordovician text:
Ordovician
The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
from: -443.7 till: -416 color:silurian text:
Silurian
The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
from: -416 till: -359.2 color:devonian text:
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
from: -359.2 till: -299 color:carboniferous text:
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
from: -299 till: -251 color:permian text:
Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
from: -251 till: -199.6 color:triassic text:
Triassic
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
from: -199.6 till: -145.5 color:jurassic text:
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
from: -145.5 till: -65.5 color:cretaceous text:
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 th ...
from: -65.5 till: -23.03 color:paleogene text:
Paleogene
The Paleogene ( ; British English, also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period, geologic period and system that spans 43 million years from the end of the Cretaceous Period million yea ...
Paleozoic Era
The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon.
The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838
by combining the Greek words ''palaiós'' (, "old") and ' ...
from: -251 till: -65.5 color:mesozoic text:
Mesozoic Era
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising ...
from: -65.5 till: 0 color:cenozoic text:
Cenozoic
The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
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.
O ...
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.
Morphology
The Ellesmerocerida are characterized by shells that ...
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 deposit ...
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 Silu ...
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 gyroconic; ...
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 ...
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 an ...
color:mesozoic bar:NAM8 from:-419.2 till:0 text:
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 ''Allonauti ...
Cambrian
The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million ...
from: -488.3 till: -443.7 color:ordovician text:
Ordovician
The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...
from: -443.7 till: -416 color:silurian text:
Silurian
The Silurian ( ) is a geologic period and system spanning 24.6 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the shortest period of the Paleozo ...
from: -416 till: -359.2 color:devonian text:
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
from: -359.2 till: -299 color:carboniferous text:
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
from: -299 till: -251 color:permian text:
Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
from: -251 till: -199.6 color:triassic text:
Triassic
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
from: -199.6 till: -145.5 color:jurassic text:
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
from: -145.5 till: -65.5 color:cretaceous text:
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 th ...
from: -65.5 till: -23.03 color:paleogene text:
Paleogene
The Paleogene ( ; British English, also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period, geologic period and system that spans 43 million years from the end of the Cretaceous Period million yea ...
Paleozoic Era
The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon.
The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838
by combining the Greek words ''palaiós'' (, "old") and ' ...
from: -251 till: -65.5 color:mesozoic text:
Mesozoic Era
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising ...
from: -65.5 till: 0 color:cenozoic text:
Cenozoic
The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...
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
The ''Treatise on Invertebrate Paleontology'' (or ''TIP'') published by the Geological Society of America and the University of Kansas Press, is a definitive multi-authored work of some 50 volumes, written by more than 300 paleontologists, and co ...
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:
*†
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 ...
= †
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.
O ...
, †
Protactinocerida
Protactinocerida is a proposed order of Late Cambrian nautiloid Cephalopod, cephalopods. Their fossils have only been found in the Late Cambrian (late Jiangshanian to early Cambrian Stage 10, Stage 10) of North China Craton, North China, a diversi ...
, †
Yanhecerida
Yanhecerida is a small order of Late Cambrian (mid- Stage 10) nautiloid cephalopods. They were similar to the more diverse Plectronocerida and Ellesmerocerida, with short shells, closely-spaced septa, and diaphragms (partitions) within the sip ...
, and †
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.
Morphology
The Ellesmerocerida are characterized by shells that ...
.
*†
Endoceratoidea
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 ...
= †
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 Silu ...
*†
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. Orthocerato ...
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 inflate ...
, and †
Pseudorthocerida
Pseudorthocerida is an order of generally straight longiconic orthoceratoids with a subcentral to marginal cyrtochoanitic siphuncle composed of variably expanded segments which may contain internal deposits that may develop into a continuous p ...
(the Orthoceratoidea of Kröger 2007)
* Nautilitoidea = †
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 gyroconic; ...
, †
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 ...
, and
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 ''Allonauti ...
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 deposit ...
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 an ...
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
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 (i.e., octopuses, squid and cuttle ...
, and sometimes placed in a subclass of its own,
Bactritoidea
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 (407 million years ago) with questionable origins in Pragian stage before 409 million ...
.
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
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 char ...
approaches are rare in nautiloid systematics. Many nautiloid orders (not to mention the group as a whole) are not
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
clades
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, t ...
, but rather
paraphyletic
In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
grades
Grade most commonly refers to:
* Grade (education), a measurement of a student's performance
* Grade, the number of the year a student has reached in a given educational stage
* Grade (slope), the steepness of a slope
Grade or grading may also r ...
. 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.
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.
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'.
* Subclass † Plectronoceratia (formerly Plectronoceratoidea)
** Order † Plectronoceratida
** Order † Yanheceratida
** Order † Protactinoceratida
* Subclass † Multiceratia (formerly Multiceratoidea)
** Order † Ellesmeroceratida
** Order †
Cyrtocerinida
Cyrtocerinida is an order of Ordovician nautiloid cephalopods. The order includes the families Cyrtocerinidae and Eothinoceratidae, as well as the genera '' Bathmoceras'' and '' Rummoceras''.
Cyrtocerinids can be characterized by a broad si ...
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 an ...
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 ''Allonauti ...
Pseudorthoceratida
Pseudorthocerida is an order of generally straight longiconic orthoceratoids with a subcentral to marginal cyrtochoanitic siphuncle composed of variably expanded segments which may contain internal deposits that may develop into a continuous pa ...
Endoceratida
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 Silu ...
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
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
, showing early cephalopod relationships to the order level (although various isolated
families
Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
also originated during this diversification event):
=Gallery=
File:Nautilus pompilius 3.jpg, The
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.
It comprises six living species in ...
, one of only two surviving nautiloid genera
File:Allonautilus scrobiculatus.jpg, ''
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 t ...
'', the other surviving nautliod genus
File:Endoceras_sp.png, ''
Endoceras
''Endoceras'' (Ancient Greek for "inner horn") is an extinct genus of large, straight shelled cephalopods from the Middle and Upper Ordovician that gives its name to the Nautiloid order Endocerida. The cross section in the mature portion is slig ...
'' was one of the largest nautiloids to have ever lived
File:Alaskoceras sewardi.jpg, ''
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, sept ...
'' was a member of 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 gyroconic; ...
, the first cephalopods to evolve coiled shells
File:Plectronoceras.png, ''
Plectronoceras
''Plectronoceras'' is the earliest known shelled cephalopod, dating to the Late Cambrian.Goldringia'' was a Rutoceratid nautiloid from the middle
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
of the
United States
The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...
File:Lorieroceras lorieri.jpg, '' Lorieroceras'' was a bizarre oncocerid nautiloid from the Devonian that possessed a helically coiled shell
File:Cenoceras NT.jpg, A reconstruction of the
Mesozoic
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
nautiloid '' Cenoceras''
File:Hoko River Nautilids.jpg, Two nautiloids from the
Eocene
The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
Nautilus cookanum
''Nautilus cookanum'' is an extinction, extinct species of nautilus. It lived during the Eocene Epoch (geology), epoch. ''N. cookanum'' placed within the genus ''Nautilus'', together with extant species based on their shared shell characters.Ward ...
'' and '' Aturia alabamensis''
File:Diorama of a Permian seafloor - coiled cephalopod, sponges, brachiopods (43887749560).jpg, ''
Cooperoceras
''Cooperoceras'' is a genus of Tainoceratid nautiloid cephalopod molluscs within the superfamily Tainocerataceae
The Tainoceratoidea are a superfamily in the cephalopod order Nautilida characterized by straight to loosely coiled shells, genera ...
Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
that possessed hollow recurved spines running along its shell
See also
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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 (i.e., octopuses, squid and cuttle ...
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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 o ...