Ammonites are an extinct group of marine animals of the subclass Ammonoidea in the class Cephalopoda, phylum Mollusca. 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. Ammonites' closest living relative is probably not the modern Nautilus (which they outwardly resemble), but rather the subclass Coleoidea (octopus, squid, and cuttlefish). Their fossil shells usually take the form of planispirals, although there were some helically-spiraled and non-spiraled forms (known as "heteromorphs"). Their name came from their spiral shape as their fossilized shells somewhat resemble tightly-coiled rams' horns. Plinius the Elder (died 79 A.D. 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" (for instance, Pleuroceras).

Classification

seealso List of ammonites Originating from within the bactritoid nautiloids, the ammonoid cephalopods first appeared in the Late Silurian to Early Devonian (circa 400 million years ago) and became extinct at the close of the Cretaceous (65 m.y.a.) along with the dinosaurs. 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) was folded, forming saddles (or peaks) and lobes (or valleys).

Suture patterns

Three major types of suture patterns in Ammonoidea have been noted:

• 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.

Orders and suborders

The three orders and various suborders of Ammonoidea are herein listed from most primitive to more derived.

• Goniatitida (Devonian to Permian) -- have round saddles, pointed lobes
  • Anarcestina (Devonian only)
  • Clymeniina (upper Upper Devonian only)
  • Goniatitina (Devonian to Upper Permian) -- includes the true goniatites
• Ceratitida (Carboniferous to Triassic) -- have round saddles, serrated lobes
  • Prolecanitina (Upper Devonian to Upper Triassic)
  • Ceratitina (Permian to Triassic) -- includes the true ceratites
• Ammonitida (Permian to Cretaceous) -- have folded saddles and lobes, fractal patterns
  • Phylloceratina (Lower Triassic to Upper Cretaceous)
  • Ammonitina (Lower Jurassic to Upper Cretaceous) -- includes the true ammonites
  • Lytoceratina (Lower Jurassic to Upper Cretaceous)
  • Ancyloceratina (Upper Jurassic to Upper Cretaceous) -- the heteromorph ammonites

Life

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. Ammonites and their kin probably preyed on fishes, crustaceans and other small creatures; while they themselves were preyed upon by such marine reptiles as mosasaurs. Fossilized ammonoids have been found showing teeth marks from such attacks.

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. Many illustrations make the mistake of placing the larger end of the coil at the top for aesthetic reasons but this is factually incorrect.

Shell anatomy and diversity

Basic shell anatomy

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.

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 gender 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.

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 baculites). 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.

Aptychus

Like the modern nautilus, many ammonites were probably able to withdraw their body into the living chamber of the shell and developed either a single horny plate or a pair of calcitic plates with which they were able to close the opening of the shell. The opening of the shell is called the aperture. 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 aptychi were identical and equal in size.

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 and are normally found detached from the shell and are 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 intended purpose. (This long-standing and wide-spread interpretation of the function of the aptychus has long been disputed. The latest studies suggest that the anaptychus may have in fact formed part of a special jaw apparatus).

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 many aptychi belong. 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.

Size

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 American ammonite is Parapuzosia bradyi from the Cretaceous with specimens measuring 137 centimetres (4.5 feet) in diameter, although a new British Columbian specimen, if authentic, would appear to trump even the European champion.

Distribution

Starting from the late Silurian, 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 geologists 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 clays 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 concretions.

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 through fossilization. It is only in these internal-moldic specimens that the suture lines can be observed; in life the sutures would have been hidden by the outer shell.

The ammonoids survived several major extinction events, with often only a few species surviving. Each time, however, this handful would diversify 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 lines disappeared along with the dinosaurs 65 million years ago in the Cretaceous-Tertiary extinction event. That no ammonites survived the extinction event at the end of the Cretaceous, while some nautiloid cousins survived, might be due to differences in ontogeny. If their extinction was due to a meteor strike, plankton around the globe could have been severely diminished, thereby dooming ammonite reproduction during its planktonic stage.

Mythology

In medieval Europe, fossilised ammonites were thought to be petrified snakes, and were called "snakestones". They were taken to be evidence for the actions of saints such as St Hilda and St Patrick. Traders would occasionally carve the face of a snake into the empty, wide end of the ammonite fossil and sell them to the public. Ammonites from the Gandaki river in Nepal are known as saligrams, and are believed by Hindus to be a concrete manifestation of God or Vishnu.

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 Ammonitina (or sometimes even order Ammonitida).

See also

• Fossils and the geological timescale
• Belemnoidea
• Nautiloidea
• Coleoidea
• Ammolite - a gemstone formed from fossil ammonite shells.

References and further reading

• Neal L Larson, Steven D Jorgensen, Robert A Farrar and Peter L Larson. Ammonites and the other Cephalopods of the Pierre Seaway. Geoscience Press, 1997.
• Lehmann, Ulrich. The Ammonites: Their life and their world. Cambridge University Press, New York, 1981. Translated from German by Janine Lettau.
• Monks, Neale and Palmer, Phil. Ammonites. Natural History Museum, 2002.
• Walker, Cyril and Ward, David. Fossils. Dorling, Kindersley Limited, London, 2002.
• A Broad Brush History of the Cephalopoda by Dr. Neale Monks, from The Cephalopod Page.
• Ammonite maturity, pathology and old age By Dr. Neale Monks, from The Cephalopod Page. Essay about the life span of Ammonites.
• Cretaceous Fossils Taxonomic Index for Order Ammonoitida
• Deeply Buried Sediments Tell Story of Sudden Mass Extinction

External links

• Cephalopod Fossil Articles from TONMO.com
• Photos of Ammonites at Lyme Regis, UK
• Paleozoic.org: Gallery of ammonite photographs
• Descriptions and pictures of ammonite fossils