Ammonite
Encyclopedia
Ammonite, as a zoological or paleontological term, refers to any member of the Ammonoidea an extinct subclass within the Molluscan class
Cephalopoda which are more closely related to living coleoids (i.e. octopus
es, squid
, and cuttlefish
) than they are to shelled nautiloids such as living species of Nautilus
.
Ammonites are excellent index fossils, and it is often possible to link the rock layer in which they are found to specific geological time periods
. Their fossil
shells usually take the form of planispirals, although there were some helically-spiraled and non-spiraled forms (known as heteromorphs).
The name ammonite, from which the scientific term is derived, was inspired by the spiral shape of their fossilized shells, which somewhat resemble tightly coiled rams' horns. Pliny the Elder
(d. 79 AD. near Pompeii) called fossils of these animals ammonis cornua ("horns of Ammon") because the Egyptian god Ammon (Amun
) was typically depicted wearing ram's horns. Often the name of an ammonite genus ends in -ceras, which is Greek
(κέρας) for "horn".
and Allonautilus
) and typical Nautilida
. However the very earliest nautiloids from the Late Cambrian and Ordovician typically had ventral siphuncles, although often proportionally larger than those in ammonites and more internally structured.
nautiloids, the ammonoid cephalopods first appeared in the Devonian
(circa 400 million years ago) and became extinct at the close of the Cretaceous
(65.5 Ma) along with the dinosaur
s. The classification of ammonoids is based in part on the ornamentation
and structure of the septa
comprising their shells' gas chambers; by these and other characteristics we can divide subclass Ammonoidea into three orders and eight known suborders. While nearly all nautiloids show gently curving sutures, the ammonoid suture line (the intersection of the septum with the outer shell) is variably folded, forming saddles (or peaks) and lobes (or valleys).
The Ammonoidea can be divided into eight orders, listed here starting with the most primitive and going to the more derived.
In some classifications these are left as suborders, included in only three orders: Goniatitida, Ceratitida
, and Ammonitida
.
(Part L, 1957) divides the Ammonoidea, regarded simply as an order, into eight suborders, the Anarcestina, Clymeniina, Goniatitina, and Prolecanitina from the Paleozoic; the Ceratitina from the Triassic; and the Ammonitina, Lytoceratina, and Phylloceratina from the Jurassic and Cretaceous. In subsequent taxonomies these are sometimes regarded as orders within the subclass Ammonoidea.
Because ammonites and their close relatives are extinct, little is known about their way of life. Their soft body parts are very rarely preserved in any detail. Nonetheless, much has been worked out by examining ammonoid shells and by using models of these shells in water tanks.
Many ammonoids probably lived in the open water of ancient seas, rather than at the sea bottom. This is suggested by the fact that their fossils are often found in rocks that were laid down under conditions where no bottom-dwelling life is found. Many of them (such as Oxynoticeras) are thought to have been good swimmers with flattened, discus-shaped, streamlined shells, although some ammonoids were less effective swimmers and were likely to have been slow-swimming bottom-dwellers. Synchrotron
analysis of an aptychophoran ammonite revealed remains of isopod and mollusc larva in its buccal cavity, indicating that at least this kind of ammonite fed on plankton. Fossilized ammonoids have been found showing tooth marks from such attacks. They may have avoided predation by squirting ink
, much like modern cephalopods; ink is occasionally preserved in fossil specimens.
The soft body of the creature occupied the largest segments of the shell at the end of the coil. The smaller earlier segments were walled off and the animal could maintain its buoyancy by filling them with gas. Thus the smaller sections of the coil would have floated above the larger sections.
The chambered part of the ammonite shell is called a phragmocone
. The phragmocone contains a series of progressively larger chambers, called camerae
(sing. camera) that are divided by thin walls called septa
(sing. septum). Only the last and largest chamber, the body chamber
, was occupied by the living animal at any given moment. As it grew, it added newer and larger chambers to the open end of the coil.
A thin living tube called a siphuncle
passed through the septa, extending from the ammonite's body into the empty shell chambers. Through a hyperosmotic active transport process, the ammonite emptied water out of these shell chambers. This enabled it to control the buoyancy
of the shell and thereby rise or descend in the water column.
A primary difference between ammonites and nautiloids is that the siphuncle of ammonites (excepting Clymeniina) runs along the ventral periphery of the septa and camerae (i.e., the inner surface of the outer axis of the shell), while the siphuncle of nautiloids runs more or less through the center of the septa and camerae.
of the animal, the shell of the male being slightly smaller and wider than that of the female. This sexual dimorphism
is thought to be an explanation for the variation in size of certain ammonite shells of the same species, the larger shell (called a macroconch) being female, and the smaller shell (called a microconch) being male. This is thought to be because the female required a larger body size for egg production. A good example of this sexual variation is found in Bifericeras from the early part of the Jurassic
period of Europe
.
It is only in relatively recent years that the sexual variation in the shells of ammonites has been recognized. The macroconch and microconch of one species were often previously mistaken for two closely related but different species occurring in the same rocks. However, these "pairs" were so consistently found together that it became apparent that they were in fact sexual forms of the same species.
s). Still other species' shells are coiled helically, superficially like that of a large gastropod (as in and ). Some species' shells are even initially uncoiled, then partially coiled, and finally straight at maturity (as in ). These partially uncoiled and totally uncoiled forms began to diversify mainly during the early part of the Cretaceous and are known as heteromorphs.
Perhaps the most extreme and bizarre looking example of a heteromorph is Nipponites
, which appears to be a tangle of irregular whorls lacking any obvious symmetrical coiling. However, upon closer inspection the shell proves to be a three-dimensional network of connected "U" shapes. occurs in rocks of the upper part of the Cretaceous
in Japan
and the USA.
Ammonites vary greatly in the ornamentation (surface relief) of their shells. Some may be smooth and relatively featureless, except for growth lines, and resemble that of the modern Nautilus. In others various patterns of spiral ridges and ribs or even spines are shown. This type of ornamentation of the shell is especially evident in the later ammonites of the Cretaceous.
Some ammonites have been found in association with a single horny plate or a pair of calcitic plates. In the past it was assumed that these plates served to close the opening of the shell in much the same way as an operculum
, however more recently it has been postulated that they were instead a jaw apparatus.
The plates are collectively termed the aptychus
or aptychi in the case of a pair of plates, and anaptychus in the case of a single plate. The paired aptychi were symmetrical to one another and equal in size and appearance.
Anaptychi are relatively rare as fossils. They are found representing ammonites from the Devonian period through those of the Cretaceous period.
Calcified aptychi only occur in ammonites from the Mesozoic
era. They are almost always found detached from the shell, and are only very rarely preserved in place. Still, sufficient numbers have been found closing the apertures of fossil ammonite shells as to leave no doubt as to their identity as part of the anatomy of an ammonite.
Large numbers of detached aptychi occur in certain beds of rock (such as those from the Mesozoic in the Alps
). These rocks are usually accumulated at great depths. The modern Nautilus lacks any calcitic plate for closing its shell, and only one extinct nautiloid
genus is known to have borne anything similar. Nautilus does, however, have a leathery head shield (the hood) which it uses to cover the opening when it retreats inside.
There are many forms of aptychus, varying in shape and the sculpture of the inner and outer surfaces, but because they are so rarely found in position within the shell of the ammonite it is often unclear to which species of ammonite one kind of aptychus belongs. A number of aptychi have been given their own genus and even species names independent of their unknown owners' genus and species, pending future discovery of verified occurrences within ammonite shells.
, their soft part record is surprisingly bleak. Beyond a tentative ink sac and possible digestive organs, no soft parts are known at all. They likely bore a radula and beak, a marginal siphuncle, and ten arms. They operated by direct development with sexual reproduction, were carnivorous and had a crop
for food storage. It is unlikely that any ammonoids dwelt in fresh or brackish water.
Few of the ammonites occurring in the lower and middle part of the Jurassic period reach a size exceeding 23 centimetres (9 inches) in diameter. Much larger forms are found in the later rocks of the upper part of the Jurassic and the lower part of the Cretaceous, such as Titanites from the Portland Stone of Jurassic of southern England, which is often 53 centimetres (2 feet) in diameter, and Parapuzosia seppenradensis
of the Cretaceous period of Germany, which is one of the largest known ammonites, sometimes reaching 2 metres (6.5 feet) in diameter. The largest documented North America
n ammonite is Parapuzosia bradyi from the Cretaceous with specimens measuring 137 centimetres (4.5 feet) in diameter, although a new 2.3-metre (7.5-foot) British Columbian specimen, if authentic, would appear to trump even the European champion.
Starting from the mid-Devonian, ammonoids were extremely abundant, especially as ammonites during the Mesozoic
era. Many genera
evolved and ran their course quickly, becoming extinct in a few million years. Due to their rapid evolution
and widespread distribution, ammonoids are used by geologist
s and paleontologists for biostratigraphy
. They are excellent index fossils, and it is often possible to link the rock layer in which they are found to specific geological time periods.
Due to their free-swimming
and/or free-floating
habits, ammonites often happened to live directly above seafloor waters so poor in oxygen as to prevent the establishment of animal life on the seafloor. When upon death the ammonites fell to this seafloor and were gradually buried in accumulating sediment, bacterial decomposition of these corpses often tipped the delicate balance of local redox
conditions sufficiently to lower the local solubility of minerals dissolved in the seawater, notably phosphates and carbonates. The resulting spontaneous concentric precipitation of minerals around a fossil is called a concretion
and is responsible for the outstanding preservation of many ammonite fossils.
When ammonites are found in clay
s their original mother-of-pearl
coating is often preserved. This type of preservation is found in ammonites such as Hoplites from the Cretaceous Gault clay
of Folkestone in Kent, England.
The Cretaceous Pierre Shale formation of the United States and Canada
is well known for the abundant ammonite fauna it yields, including Baculites, Placenticeras, Scaphites
, Hoploscaphites, and Jeletzkytes, as well as many uncoiled forms. Many of these also have much or all of the original shell, as well as the complete body chamber, still intact. Many Pierre Shale ammonites, and indeed many ammonites throughout earth history, are found inside concretion
s.
Other fossils, such as many found in Madagascar
and Alberta
(Canada
), display iridescence
. These iridescent ammonites are often of gem quality (ammolite
) when polished. In no case would this iridescence have been visible during the animal's life; additional shell layers covered it.
The majority of ammonoid specimens, especially those of the Paleozoic
era, are preserved only as internal molds; that it to say, the outer shell (composed of aragonite
) has been lost during the fossilization process. It is only in these internal-mold specimens that the suture lines can be observed; in life the sutures would have been hidden by the outer shell.
The ammonoids as a group continued through several major extinction event
s, although it appears that often only a few species survived. Each time, however, this handful of species diversified into a multitude of forms. Ammonite fossils became less abundant during the latter part of the Mesozoic
, with none surviving into the Cenozoic
era. The last surviving lineages disappeared, along with the dinosaurs, 65 million years ago in the Cretaceous-Tertiary extinction event
. The reason why no ammonites survived the extinction event
at the end of the Cretaceous, whereas some nautiloid cousins survived, might be due to differences in ontogeny
. If their extinction was due to a bolide strike, plankton
around the globe could have been severely diminished, thereby dooming ammonite reproduction during its planktonic stage.
s, has been attributed to a bolide impact, marking the end of the Cretaceous Period. Regardless of what effect an impact may have had, many of these groups, including ammonoids, were already in serious decline. Previously ammonoid cephalopods barely survived several earlier major extinction event
s, often with only a few species surviving from which a multitude of forms diversified.
Eight or so species from only two families made it almost to the end of the Cretaceous, the order having gone through a more or less steady decline since the middle of the period. Six other families made it well into the upper Maastrichtian
(uppermost stage of the Cretaceous) but were extinct well before the end. All told, 11 families entered the Maastrichtian, a decline from the 19 families known from the Cenomanian
in the middle of the Cretaceous.
One reason given for their demise is that Cretaceous ammonites, being closely related to coleoids, had a similar reproductive strategy in which a huge number of eggs is laid in a single batch at the end of the life span. These, along with juvenile ammonites, are thought to have been part of the plankton
at the surface of the ocean where they were killed off by the effects of an impact. Nautiloid
s, exemplified by modern nautilus
es, are thought on the other hand to have had a reproductive strategy in which eggs were laid in smaller batches many times during the life span and on the sea floor well away from any direct effects of such a bolide strike, and thus survived.
, a myth referenced in Sir Walter Scott
's Marmion, and Saint Patrick
, and were held to have healing or oracular
powers. Traders would occasionally carve the head of a snake onto the empty, wide end of the ammonite fossil, and then sell them to the public. In other cases the snake's head would be simply painted on. Ammonites from the Gandaki river in Nepal
are known as saligrams, and are believed by Hindu
s to be a concrete manifestation of God or Vishnu
.
(or sometimes even order Ammonitida
).
Class (biology)
In biological classification, class is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, order, family, genus, and species, with class fitting between phylum and order...
Cephalopoda which are more closely related to living coleoids (i.e. octopus
Octopus
The octopus is a cephalopod mollusc of the order Octopoda. Octopuses have two eyes and four pairs of arms, and like other cephalopods they are bilaterally symmetric. An octopus has a hard beak, with its mouth at the center point of the arms...
es, squid
Squid
Squid are cephalopods of the order Teuthida, which comprises around 300 species. Like all other cephalopods, squid have a distinct head, bilateral symmetry, a mantle, and arms. Squid, like cuttlefish, have eight arms arranged in pairs and two, usually longer, tentacles...
, and cuttlefish
Cuttlefish
Cuttlefish are marine animals of the order Sepiida. They belong to the class Cephalopoda . Despite their name, cuttlefish are not fish but molluscs....
) than they are to shelled nautiloids such as living species of Nautilus
Nautilus
Nautilus is the common name of marine creatures of cephalopod family Nautilidae, the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in two genera, the type of which is the genus Nautilus...
.
Ammonites are excellent index fossils, and it is often possible to link the rock layer in which they are found to specific geological time periods
Geologic time scale
The geologic time scale provides a system of chronologic measurement relating stratigraphy to time that is used by geologists, paleontologists and other earth scientists to describe the timing and relationships between events that have occurred during the history of the Earth...
. Their fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...
shells usually take the form of planispirals, although there were some helically-spiraled and non-spiraled forms (known as heteromorphs).
The name ammonite, from which the scientific term is derived, was inspired by the spiral shape of their fossilized shells, which somewhat resemble tightly coiled rams' horns. Pliny the Elder
Pliny the Elder
Gaius Plinius Secundus , better known as Pliny the Elder, was a Roman author, naturalist, and natural philosopher, as well as naval and army commander of the early Roman Empire, and personal friend of the emperor Vespasian...
(d. 79 AD. near Pompeii) called fossils of these animals ammonis cornua ("horns of Ammon") because the Egyptian god Ammon (Amun
Amun
Amun, reconstructed Egyptian Yamānu , was a god in Egyptian mythology who in the form of Amun-Ra became the focus of the most complex system of theology in Ancient Egypt...
) was typically depicted wearing ram's horns. Often the name of an ammonite genus ends in -ceras, which is Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...
(κέρας) for "horn".
Diagnostic characters
Ammonites (subclass Ammonoidea) can be distinguishes by their septa, the dividing walls that separate the chambers in the phragmocone, by the nature of their sutures where the septa joint the outer shell wall, and in general by their siphuncles.Septa
Ammonoid septa characteristically have bulges and indentations and are to varying degrees convex from the front, distinguishing them from nautiloid septa which are typically simple concave dish-shaped structures. The topology of the septa, especially around the rim, results in the various suture patterns found.Suture patterns
Three major types of suture patterns are found in the Ammonoidea:- Goniatitic - numerous undivided lobes and saddles; typically 8 lobes around the conch. This pattern is characteristic of the Paleozoic ammonoids.
- Ceratitic - lobes have subdivided tips, giving them a saw-toothed appearance, and rounded undivided saddles. This suture pattern is characteristic of Triassic ammonoids and appears again in the Cretaceous "pseudoceratites".
- Ammonitic - lobes and saddles are much subdivided (fluted); subdivisions are usually rounded instead of saw-toothed. Ammonoids of this type are the most important species from a biostratigraphical point of view. This suture type is characteristic of Jurassic and Cretaceous ammonoids but extends back all the way to the Permian.
Siphuncle
The siphuncle in most ammonoids by far is a narrow tubular structure that runs along the outer rim, known as the venter, connecting the chambers of the phragmocone to the body or living chamber. This distinguishes them from living nautiloides (NautilusNautilus
Nautilus is the common name of marine creatures of cephalopod family Nautilidae, the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in two genera, the type of which is the genus Nautilus...
and Allonautilus
Allonautilus
The genus Allonautilus contains two species of nautiluses, which differ significantly in terms of morphology from those placed in the sister taxon Nautilus. Allonautilus is now thought to be a descendant of Nautilus and the latter paraphyletic.-External links:*...
) and typical 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...
. However the very earliest nautiloids from the Late Cambrian and Ordovician typically had ventral siphuncles, although often proportionally larger than those in ammonites and more internally structured.
Classification of the Ammonoidea
Originating from within the bactritoidBactritida
The Bactritida form a small order of more or less straight-shelled cephalopods that first appeared during the Emsian Stage of the Devonian Period and persisted until the Carnian Stage of the Triassic Period...
nautiloids, the ammonoid cephalopods first appeared in the Devonian
Devonian
The Devonian is a geologic period and system of the Paleozoic Era spanning from the end of the Silurian Period, about 416.0 ± 2.8 Mya , to the beginning of the Carboniferous Period, about 359.2 ± 2.5 Mya...
(circa 400 million years ago) and became extinct at the close of the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
(65.5 Ma) along with the dinosaur
Dinosaur
Dinosaurs are a diverse group of animals of the clade and superorder Dinosauria. They were the dominant terrestrial vertebrates for over 160 million years, from the late Triassic period until the end of the Cretaceous , when the Cretaceous–Paleogene extinction event led to the extinction of...
s. The classification of ammonoids is based in part on the ornamentation
Biological ornament
A biological ornament is a structure of an animal that appears to serve a decorative function rather than an ostensible, utilitarian function. Ornaments are used in displays to attract mates. An animal may shake, lengthen, or spread out its ornament in order to get the attention of the opposite...
and structure of the septa
Septa (biology)
Septa are thin walls or partitions between the internal chambers of the shell of a cephalopod, namely nautiloids or ammonoids....
comprising their shells' gas chambers; by these and other characteristics we can divide subclass Ammonoidea into three orders and eight known suborders. While nearly all nautiloids show gently curving sutures, the ammonoid suture line (the intersection of the septum with the outer shell) is variably folded, forming saddles (or peaks) and lobes (or valleys).
Orders and suborders
- Anarcestida, Devonian
- ClymeniidaClymeniidaThe Clymeniida constitute a unique order of ammonoid cephalods from the Upper Devonian characterized by having an unusual dorsal siphuncle. They measured about 4 cm in diameter and are restricted to Europe, North Africa, and possibly Australia....
, Upper Devonian - Goniatitida, Middle Devonian - Upper Permian
- ProlecanitidaProlecanitidaThe Prolecanitida are extinct ammonoid cephalopods with discoidal to thinly lenticular shells with goniatitic or ceratitic sutures and which retained the simple retrochoanitic siphuncle with backward extending septal necks. As typical for ammonoids the siphuncle is along the ventral margin...
, Upper Devonian - Upper Triassic - CeratitidaCeratitidaThe Ceratitida is an order that contains almost all ammonoid cephalopod genera from the Triassic as well as ancestral forms from the Upper Permian, the exception being the phylloceratids which gave rise to the great diversity of post Triassic ammonites....
, Permian - Triassic - Phylloceratida, Triassic - Cretaceous
- Lytoceratida, Jurassic - Cretaceous
- AmmonitidaAmmonitidaThe Ammonitida is an order of more highly evolved ammonoid cephalopods from the Jurassic and Cretaceous time periods, commonly with intricate ammonitic sutures....
, Lower Jurassic - Upper Cretaceous
In some classifications these are left as suborders, included in only three orders: Goniatitida, Ceratitida
Ceratitida
The Ceratitida is an order that contains almost all ammonoid cephalopod genera from the Triassic as well as ancestral forms from the Upper Permian, the exception being the phylloceratids which gave rise to the great diversity of post Triassic ammonites....
, and Ammonitida
Ammonitida
The Ammonitida is an order of more highly evolved ammonoid cephalopods from the Jurassic and Cretaceous time periods, commonly with intricate ammonitic sutures....
.
Taxonomy of the Treatise
The Treatise on Invertebrate PaleontologyTreatise on Invertebrate Paleontology
The Treatise on Invertebrate Paleontology 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 covering every phylum, class, order, family, and genus of fossil and...
(Part L, 1957) divides the Ammonoidea, regarded simply as an order, into eight suborders, the Anarcestina, Clymeniina, Goniatitina, and Prolecanitina from the Paleozoic; the Ceratitina from the Triassic; and the Ammonitina, Lytoceratina, and Phylloceratina from the Jurassic and Cretaceous. In subsequent taxonomies these are sometimes regarded as orders within the subclass Ammonoidea.
Life
Many ammonoids probably lived in the open water of ancient seas, rather than at the sea bottom. This is suggested by the fact that their fossils are often found in rocks that were laid down under conditions where no bottom-dwelling life is found. Many of them (such as Oxynoticeras) are thought to have been good swimmers with flattened, discus-shaped, streamlined shells, although some ammonoids were less effective swimmers and were likely to have been slow-swimming bottom-dwellers. Synchrotron
Synchrotron
A synchrotron is a particular type of cyclic particle accelerator in which the magnetic field and the electric field are carefully synchronised with the travelling particle beam. The proton synchrotron was originally conceived by Sir Marcus Oliphant...
analysis of an aptychophoran ammonite revealed remains of isopod and mollusc larva in its buccal cavity, indicating that at least this kind of ammonite fed on plankton. Fossilized ammonoids have been found showing tooth marks from such attacks. They may have avoided predation by squirting ink
Cephalopod ink
Cephalopod ink is a dark pigment released into water by most species of cephalopod, usually as an escape mechanism. All cephalopods, with the exception of the Nautilidae and the species of octopus belonging to the suborder Cirrina, are able to release ink....
, much like modern cephalopods; ink is occasionally preserved in fossil specimens.
The soft body of the creature occupied the largest segments of the shell at the end of the coil. The smaller earlier segments were walled off and the animal could maintain its buoyancy by filling them with gas. Thus the smaller sections of the coil would have floated above the larger sections.
Basic shell anatomy
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 siphuncle which determines buoyancy by means of gas...
. The phragmocone contains a series of progressively larger chambers, called camerae
Camerae
Camerae are the spaces or chambers enclosed between two adjacent septa in the phragmocone of a nautiloid or ammonoid cephalopod. These can be seen in cross-sections of a nautilus shell and in the polished cross-sections of ammonites...
(sing. camera) that are divided by thin walls called septa
Septa (biology)
Septa are thin walls or partitions between the internal chambers of the shell of a cephalopod, namely nautiloids or ammonoids....
(sing. septum). Only the last and largest chamber, the body chamber
Body chamber
The body chamber, also called the living chamber, is the outermost or last chamber in the shell of a nautiloid or ammonoid cephalopod. The body of the animal occupies the living chamber, apart from the siphuncle which extends through the rest of septa to provide buoyancy....
, was occupied by the living animal at any given moment. As it grew, it added newer and larger chambers to the open end of the coil.
A thin living tube called a siphuncle
Siphuncle
The siphuncle is a strand of tissue passing longitudinally through the shell of a cephalopod mollusk. Only cephalopods with chambered shells have siphuncles, such as the extinct ammonites and belemnites, and the living nautiluses, cuttlefish, and Spirula...
passed through the septa, extending from the ammonite's body into the empty shell chambers. Through a hyperosmotic active transport process, the ammonite emptied water out of these shell chambers. This enabled it to control the buoyancy
Buoyancy
In physics, buoyancy is a force exerted by a fluid that opposes an object's weight. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus a column of fluid, or an object submerged in the fluid, experiences greater pressure at the bottom of the...
of the shell and thereby rise or descend in the water column.
A primary difference between ammonites and nautiloids is that the siphuncle of ammonites (excepting Clymeniina) runs along the ventral periphery of the septa and camerae (i.e., the inner surface of the outer axis of the shell), while the siphuncle of nautiloids runs more or less through the center of the septa and camerae.
Sexual dimorphism
One feature found in shells of the modern Nautilus is the variation in the shape and size of the shell according to the sexSex
In biology, sex is a process of combining and mixing genetic traits, often resulting in the specialization of organisms into a male or female variety . Sexual reproduction involves combining specialized cells to form offspring that inherit traits from both parents...
of the animal, the shell of the male being slightly smaller and wider than that of the female. This sexual dimorphism
Sexual dimorphism
Sexual dimorphism is a phenotypic difference between males and females of the same species. Examples of such differences include differences in morphology, ornamentation, and behavior.-Examples:-Ornamentation / coloration:...
is thought to be an explanation for the variation in size of certain ammonite shells of the same species, the larger shell (called a macroconch) being female, and the smaller shell (called a microconch) being male. This is thought to be because the female required a larger body size for egg production. A good example of this sexual variation is found in Bifericeras from the early part of the Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...
period of Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...
.
It is only in relatively recent years that the sexual variation in the shells of ammonites has been recognized. The macroconch and microconch of one species were often previously mistaken for two closely related but different species occurring in the same rocks. However, these "pairs" were so consistently found together that it became apparent that they were in fact sexual forms of the same species.
Variations in shape
The majority of ammonite species feature a shell that is a planispiral flat coil, but other species feature a shell that is nearly straight (as in baculiteBaculite
Baculites is an extinct cephalopod genus with a nearly straight shell that is included in the heteromorph ammonites...
s). Still other species' shells are coiled helically, superficially like that of a large gastropod (as in and ). Some species' shells are even initially uncoiled, then partially coiled, and finally straight at maturity (as in ). These partially uncoiled and totally uncoiled forms began to diversify mainly during the early part of the Cretaceous and are known as heteromorphs.
Perhaps the most extreme and bizarre looking example of a heteromorph is Nipponites
Nipponites
Nipponites is an extinct genus of heteromorph ammonites. The species of Nipponites are famous for the way their shells form "ox-bow" bends, resulting in some of the most bizarre shapes ever seen among ammonites.The ecology of Nipponites, as with many other nostoceratids, is subject to much...
, which appears to be a tangle of irregular whorls lacking any obvious symmetrical coiling. However, upon closer inspection the shell proves to be a three-dimensional network of connected "U" shapes. occurs in rocks of the upper part of the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...
in Japan
Japan
Japan is an island nation in East Asia. Located in the Pacific Ocean, it lies to the east of the Sea of Japan, China, North Korea, South Korea and Russia, stretching from the Sea of Okhotsk in the north to the East China Sea and Taiwan in the south...
and the USA.
Ammonites vary greatly in the ornamentation (surface relief) of their shells. Some may be smooth and relatively featureless, except for growth lines, and resemble that of the modern Nautilus. In others various patterns of spiral ridges and ribs or even spines are shown. This type of ornamentation of the shell is especially evident in the later ammonites of the Cretaceous.
Aptychus
Operculum (gastropod)
The operculum, meaning little lid, is a corneous or calcareous anatomical structure which exists in many groups of sea snails and freshwater snails, and also in a few groups of land snails...
, however more recently it has been postulated that they were instead a jaw apparatus.
The plates are collectively termed the aptychus
Aptychus
An aptychus is a type of marine fossil, a hard anatomical structure like a curved shelly plate, which was part of the body of an ammonite. Paired aptychi have, on rare occasions, been found at or within the aperture of ammonite shells....
or aptychi in the case of a pair of plates, and anaptychus in the case of a single plate. The paired aptychi were symmetrical to one another and equal in size and appearance.
Anaptychi are relatively rare as fossils. They are found representing ammonites from the Devonian period through those of the Cretaceous period.
Calcified aptychi only occur in ammonites from the Mesozoic
Mesozoic
The Mesozoic era is an interval of geological time from about 250 million years ago to about 65 million years ago. It is often referred to as the age of reptiles because reptiles, namely dinosaurs, were the dominant terrestrial and marine vertebrates of the time...
era. They are almost always found detached from the shell, and are only very rarely preserved in place. Still, sufficient numbers have been found closing the apertures of fossil ammonite shells as to leave no doubt as to their identity as part of the anatomy of an ammonite.
Large numbers of detached aptychi occur in certain beds of rock (such as those from the Mesozoic in the Alps
Alps
The Alps is one of the great mountain range systems of Europe, stretching from Austria and Slovenia in the east through Italy, Switzerland, Liechtenstein and Germany to France in the west....
). These rocks are usually accumulated at great depths. The modern Nautilus lacks any calcitic plate for closing its shell, and only one extinct nautiloid
Nautiloid
Nautiloids are a large and diverse group of marine cephalopods belonging to the subclass Nautiloidea that began in the Late Cambrian and are represented today by the living Nautilus. Nautiloids flourished during the early Paleozoic era, where they constituted the main predatory animals, and...
genus is known to have borne anything similar. Nautilus does, however, have a leathery head shield (the hood) which it uses to cover the opening when it retreats inside.
There are many forms of aptychus, varying in shape and the sculpture of the inner and outer surfaces, but because they are so rarely found in position within the shell of the ammonite it is often unclear to which species of ammonite one kind of aptychus belongs. A number of aptychi have been given their own genus and even species names independent of their unknown owners' genus and species, pending future discovery of verified occurrences within ammonite shells.
Soft-part anatomy
Although ammonites do occur in exceptional lagerstatten such as the Solnhofen limestoneSolnhofen limestone
The Solnhofen Plattenkalk is a Jurassic Konservat-Lagerstätte that preserves a rare assemblage of fossilized organisms, including highly detailed imprints of soft bodied organisms such as sea jellies...
, their soft part record is surprisingly bleak. Beyond a tentative ink sac and possible digestive organs, no soft parts are known at all. They likely bore a radula and beak, a marginal siphuncle, and ten arms. They operated by direct development with sexual reproduction, were carnivorous and had a crop
Crop
Crop may refer to:* Crop, a plant grown and harvested for agricultural use* Crop , part of the alimentary tract of some animals* Crop , a modified whip used in horseback riding or disciplining humans...
for food storage. It is unlikely that any ammonoids dwelt in fresh or brackish water.
Size
Parapuzosia seppenradensis
Parapuzosia seppenradensis is the largest known species of ammonite. It lived during the Late Cretaceous period. A specimen found in Germany in 1895 measures 1.95 m in diameter, although the living chamber is incomplete. It is estimated that if complete, this specimen would have had a diameter of...
of the Cretaceous period of Germany, which is one of the largest known ammonites, sometimes reaching 2 metres (6.5 feet) in diameter. The largest documented North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...
n ammonite is Parapuzosia bradyi from the Cretaceous with specimens measuring 137 centimetres (4.5 feet) in diameter, although a new 2.3-metre (7.5-foot) British Columbian specimen, if authentic, would appear to trump even the European champion.
Distribution
Mesozoic
The Mesozoic era is an interval of geological time from about 250 million years ago to about 65 million years ago. It is often referred to as the age of reptiles because reptiles, namely dinosaurs, were the dominant terrestrial and marine vertebrates of the time...
era. Many genera
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
evolved and ran their course quickly, becoming extinct in a few million years. Due to their rapid evolution
Evolution
Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...
and widespread distribution, ammonoids are used by geologist
Geologist
A geologist is a scientist who studies the solid and liquid matter that constitutes the Earth as well as the processes and history that has shaped it. Geologists usually engage in studying geology. Geologists, studying more of an applied science than a theoretical one, must approach Geology using...
s and paleontologists for biostratigraphy
Biostratigraphy
Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them. Usually the aim is correlation, demonstrating that a particular horizon in one geological section represents the same period...
. They are excellent index fossils, and it is often possible to link the rock layer in which they are found to specific geological time periods.
Due to their free-swimming
Nekton
Nekton refers to the aggregate of actively swimming aquatic organisms in a body of water able to move independently of water currents....
and/or free-floating
Plankton
Plankton are any drifting organisms that inhabit the pelagic zone of oceans, seas, or bodies of fresh water. That is, plankton are defined by their ecological niche rather than phylogenetic or taxonomic classification...
habits, ammonites often happened to live directly above seafloor waters so poor in oxygen as to prevent the establishment of animal life on the seafloor. When upon death the ammonites fell to this seafloor and were gradually buried in accumulating sediment, bacterial decomposition of these corpses often tipped the delicate balance of local redox
Redox
Redox reactions describe all chemical reactions in which atoms have their oxidation state changed....
conditions sufficiently to lower the local solubility of minerals dissolved in the seawater, notably phosphates and carbonates. The resulting spontaneous concentric precipitation of minerals around a fossil is called a concretion
Concretion
A concretion is a volume of sedimentary rock in which a mineral cement fills the porosity . Concretions are often ovoid or spherical in shape, although irregular shapes also occur. The word 'concretion' is derived from the Latin con meaning 'together' and crescere meaning 'to grow'...
and is responsible for the outstanding preservation of many ammonite fossils.
When ammonites are found in clay
Clay
Clay is a general term including many combinations of one or more clay minerals with traces of metal oxides and organic matter. Geologic clay deposits are mostly composed of phyllosilicate minerals containing variable amounts of water trapped in the mineral structure.- Formation :Clay minerals...
s their original mother-of-pearl
Nacre
Nacre , also known as mother of pearl, is an organic-inorganic composite material produced by some mollusks as an inner shell layer; it is also what makes up pearls. It is very strong, resilient, and iridescent....
coating is often preserved. This type of preservation is found in ammonites such as Hoplites from the Cretaceous Gault clay
Gault Clay
Gault is a clay formation of stiff blue clay deposited in a calm, fairly deep water marine environment during the Lower Cretaceous Period...
of Folkestone in Kent, England.
The Cretaceous Pierre Shale formation of the United States and Canada
Canada
Canada is a North American country consisting of ten provinces and three territories. Located in the northern part of the continent, it extends from the Atlantic Ocean in the east to the Pacific Ocean in the west, and northward into the Arctic Ocean...
is well known for the abundant ammonite fauna it yields, including Baculites, Placenticeras, Scaphites
Scaphites
Scaphites is a genus of extinct cephalopod belonging to the family of heteromorph ammonites . They were a widespread genus that thrived during the Cretaceous period....
, Hoploscaphites, and Jeletzkytes, as well as many uncoiled forms. Many of these also have much or all of the original shell, as well as the complete body chamber, still intact. Many Pierre Shale ammonites, and indeed many ammonites throughout earth history, are found inside concretion
Concretion
A concretion is a volume of sedimentary rock in which a mineral cement fills the porosity . Concretions are often ovoid or spherical in shape, although irregular shapes also occur. The word 'concretion' is derived from the Latin con meaning 'together' and crescere meaning 'to grow'...
s.
Madagascar
The Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...
and Alberta
Alberta
Alberta is a province of Canada. It had an estimated population of 3.7 million in 2010 making it the most populous of Canada's three prairie provinces...
(Canada
Canada
Canada is a North American country consisting of ten provinces and three territories. Located in the northern part of the continent, it extends from the Atlantic Ocean in the east to the Pacific Ocean in the west, and northward into the Arctic Ocean...
), display iridescence
Iridescence
Iridescence is generally known as the property of certain surfaces which appear to change color as the angle of view or the angle of illumination changes...
. These iridescent ammonites are often of gem quality (ammolite
Ammolite
Ammolite is a rare and valuable opal-like organic gemstone found primarily along the eastern slopes of the Rocky Mountains of North America. It is made of the fossilized shells of ammonites, which in turn are composed primarily of aragonite, the same mineral that makes up nacreous pearls...
) when polished. In no case would this iridescence have been visible during the animal's life; additional shell layers covered it.
The majority of ammonoid specimens, especially those of the Paleozoic
Paleozoic
The Paleozoic era is the earliest of three geologic eras of the Phanerozoic eon, spanning from roughly...
era, are preserved only as internal molds; that it to say, the outer shell (composed of aragonite
Aragonite
Aragonite is a carbonate mineral, one of the two common, naturally occurring, crystal forms of calcium carbonate, CaCO3...
) has been lost during the fossilization process. It is only in these internal-mold specimens that the suture lines can be observed; in life the sutures would have been hidden by the outer shell.
The ammonoids as a group continued through several major extinction event
Extinction event
An extinction event is a sharp decrease in the diversity and abundance of macroscopic life. They occur when the rate of extinction increases with respect to the rate of speciation...
s, although it appears that often only a few species survived. Each time, however, this handful of species diversified into a multitude of forms. Ammonite fossils became less abundant during the latter part of the Mesozoic
Mesozoic
The Mesozoic era is an interval of geological time from about 250 million years ago to about 65 million years ago. It is often referred to as the age of reptiles because reptiles, namely dinosaurs, were the dominant terrestrial and marine vertebrates of the time...
, with none surviving into the Cenozoic
Cenozoic
The Cenozoic era is the current and most recent of the three Phanerozoic geological eras and covers the period from 65.5 mya to the present. The era began in the wake of the Cretaceous–Tertiary extinction event at the end of the Cretaceous that saw the demise of the last non-avian dinosaurs and...
era. The last surviving lineages disappeared, along with the dinosaurs, 65 million years ago in the Cretaceous-Tertiary extinction event
Cretaceous-Tertiary extinction event
The Cretaceous–Paleogene extinction event, formerly named and still commonly referred to as the Cretaceous-Tertiary extinction event, occurred approximately 65.5 million years ago at the end of the Maastrichtian age of the Cretaceous period. It was a large-scale mass extinction of animal and plant...
. The reason why no ammonites survived the extinction event
Extinction event
An extinction event is a sharp decrease in the diversity and abundance of macroscopic life. They occur when the rate of extinction increases with respect to the rate of speciation...
at the end of the Cretaceous, whereas some nautiloid cousins survived, might be due to differences in ontogeny
Ontogeny
Ontogeny is the origin and the development of an organism – for example: from the fertilized egg to mature form. It covers in essence, the study of an organism's lifespan...
. If their extinction was due to a bolide strike, plankton
Plankton
Plankton are any drifting organisms that inhabit the pelagic zone of oceans, seas, or bodies of fresh water. That is, plankton are defined by their ecological niche rather than phylogenetic or taxonomic classification...
around the globe could have been severely diminished, thereby dooming ammonite reproduction during its planktonic stage.
Extinction
The extinction of the ammonites along with other marine animals and of course, non-avian dinosaurDinosaur
Dinosaurs are a diverse group of animals of the clade and superorder Dinosauria. They were the dominant terrestrial vertebrates for over 160 million years, from the late Triassic period until the end of the Cretaceous , when the Cretaceous–Paleogene extinction event led to the extinction of...
s, has been attributed to a bolide impact, marking the end of the Cretaceous Period. Regardless of what effect an impact may have had, many of these groups, including ammonoids, were already in serious decline. Previously ammonoid cephalopods barely survived several earlier major extinction event
Extinction event
An extinction event is a sharp decrease in the diversity and abundance of macroscopic life. They occur when the rate of extinction increases with respect to the rate of speciation...
s, often with only a few species surviving from which a multitude of forms diversified.
Eight or so species from only two families made it almost to the end of the Cretaceous, the order having gone through a more or less steady decline since the middle of the period. Six other families made it well into the upper Maastrichtian
Maastrichtian
The Maastrichtian is, in the ICS' geologic timescale, the latest age or upper stage of the Late Cretaceous epoch or Upper Cretaceous series, the Cretaceous period or system, and of the Mesozoic era or erathem. It spanned from 70.6 ± 0.6 Ma to 65.5 ± 0.3 Ma...
(uppermost stage of the Cretaceous) but were extinct well before the end. All told, 11 families entered the Maastrichtian, a decline from the 19 families known from the Cenomanian
Cenomanian
The Cenomanian is, in the ICS' geological timescale the oldest or earliest age of the Late Cretaceous epoch or the lowest stage of the Upper Cretaceous series. An age is a unit of geochronology: it is a unit of time; the stage is a unit in the stratigraphic column deposited during the corresponding...
in the middle of the Cretaceous.
One reason given for their demise is that Cretaceous ammonites, being closely related to coleoids, had a similar reproductive strategy in which a huge number of eggs is laid in a single batch at the end of the life span. These, along with juvenile ammonites, are thought to have been part of the plankton
Plankton
Plankton are any drifting organisms that inhabit the pelagic zone of oceans, seas, or bodies of fresh water. That is, plankton are defined by their ecological niche rather than phylogenetic or taxonomic classification...
at the surface of the ocean where they were killed off by the effects of an impact. Nautiloid
Nautiloid
Nautiloids are a large and diverse group of marine cephalopods belonging to the subclass Nautiloidea that began in the Late Cambrian and are represented today by the living Nautilus. Nautiloids flourished during the early Paleozoic era, where they constituted the main predatory animals, and...
s, exemplified by modern nautilus
Nautilus
Nautilus is the common name of marine creatures of cephalopod family Nautilidae, the sole extant family of the superfamily Nautilaceae and of its smaller but near equal suborder, Nautilina. It comprises six living species in two genera, the type of which is the genus Nautilus...
es, are thought on the other hand to have had a reproductive strategy in which eggs were laid in smaller batches many times during the life span and on the sea floor well away from any direct effects of such a bolide strike, and thus survived.
Mythology
In medieval Europe, fossilised ammonites were thought to be petrified coiled snakes, and were called "snakestones" or, more commonly in medieval England, "serpentstones". They were considered to be evidence for the actions of saints such as Hilda of WhitbyHilda of Whitby
Hilda of Whitby or Hild of Whitby was a Christian saint and the founding abbess of the monastery at Whitby, which was chosen as the venue for the Synod of Whitby...
, a myth referenced in Sir Walter Scott
Walter Scott
Sir Walter Scott, 1st Baronet was a Scottish historical novelist, playwright, and poet, popular throughout much of the world during his time....
's Marmion, and Saint Patrick
Saint Patrick
Saint Patrick was a Romano-Briton and Christian missionary, who is the most generally recognized patron saint of Ireland or the Apostle of Ireland, although Brigid of Kildare and Colmcille are also formally patron saints....
, and were held to have healing or oracular
Divination
Divination is the attempt to gain insight into a question or situation by way of an occultic standardized process or ritual...
powers. Traders would occasionally carve the head of a snake onto the empty, wide end of the ammonite fossil, and then sell them to the public. In other cases the snake's head would be simply painted on. Ammonites from the Gandaki river in Nepal
Nepal
Nepal , officially the Federal Democratic Republic of Nepal, is a landlocked sovereign state located in South Asia. It is located in the Himalayas and bordered to the north by the People's Republic of China, and to the south, east, and west by the Republic of India...
are known as saligrams, and are believed by Hindu
Hindu
Hindu refers to an identity associated with the philosophical, religious and cultural systems that are indigenous to the Indian subcontinent. As used in the Constitution of India, the word "Hindu" is also attributed to all persons professing any Indian religion...
s to be a concrete manifestation of God or Vishnu
Vishnu
Vishnu is the Supreme god in the Vaishnavite tradition of Hinduism. Smarta followers of Adi Shankara, among others, venerate Vishnu as one of the five primary forms of God....
.
Terminological note
The words ammonite and ammonoid are both used quite loosely in common parlance to refer to any member of subclass Ammonoidea. However, in stricter usage the term ammonite is reserved for members of suborder AmmonitinaAmmonitina
The Ammonitina comprises a diverse suborder of ammonoid cephalopods that lived during the Jurassic and Cretaceous periods of the Mesozoic Era. They are excellent index fossils, and it is often possible to link the rock layer in which they are found to specific geological time periods.The shells of...
(or sometimes even order Ammonitida
Ammonitida
The Ammonitida is an order of more highly evolved ammonoid cephalopods from the Jurassic and Cretaceous time periods, commonly with intricate ammonitic sutures....
).
See also
- AmmoliteAmmoliteAmmolite is a rare and valuable opal-like organic gemstone found primarily along the eastern slopes of the Rocky Mountains of North America. It is made of the fossilized shells of ammonites, which in turn are composed primarily of aragonite, the same mineral that makes up nacreous pearls...
- a gemstoneGemstoneA gemstone or gem is a piece of mineral, which, in cut and polished form, is used to make jewelry or other adornments...
formed from fossil ammonite shells. - BelemnoideaBelemnoideaBelemnoids 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...
- ColeoideaColeoideaSubclass Coleoidea, or Dibranchiata, is the grouping of cephalopods containing all the primarily soft-bodied creatures. Unlike its sister group Nautiloidea, whose members have a rigid outer shell for protection, the coleoids have at most an internal bone or shell that is used for buoyancy or support...
- Fossils and the geological timescale
- Nautiloidea
External links
- Descriptions and pictures of ammonite fossils
- goniat.org, a palaezoic ammonoid database
- paleozoic.org: gallery of ammonite photographs
- photos of ammonites at Lyme Regis, UK
- TaxonConcept's data on cretaceous ammonites
- The ammonites of Peacehaven - photos of giant cretaceous ammonites in Southern England
- tonmo.com: The octopus news magazine online, Cephalopod fossil articles.