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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) than they are to shelled nautiloids such as the living Nautilus species. The earliest ammonites appeared during the Devonian, with the last species vanishing during the Cretaceous–Paleogene extinction event.

Ammonites are excellent index fossils, and linking the rock layer in which a particular species or genus is found to specific geologic time periods is often possible. Their fossil shells usually take the form of planispirals, although some helically spiraled and nonspiraled forms (known as heteromorphs) have been found.

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 rams' horns. Often, the name of an ammonite genus ends in -ceras, which is from κέρας (kéras) meaning "horn".

Appearence[]

Ammonites (subclass Ammonoidea) can be distinguished by their septa, the dividing walls that separate the chambers in the phragmocone, by the nature of their sutures where the septa join the outer shell wall, and in general by their siphuncles.

The smallest ammonoid was Maximites from the Upper Carboniferous. Adult specimens reached only 10 mm (0.39 in) in shell diameter. Few of the ammonites occurring in the lower and middle part of the Jurassic period reached a size exceeding 23 cm (9.1 in) 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 cm (1.74 ft) in diameter, and Parapuzosia seppenradensis of the Cretaceous period of Germany, which is one of the largest-known ammonites, sometimes reaching 2 m (6.6 ft) in diameter. The largest-documented North American ammonite is Parapuzosia bradyi from the Cretaceous, with specimens measuring 137 cm (4.5 ft) in diameter.

Septa[]

Ammonoid septa characteristically have bulges and indentations and are to varying degrees convex when seen 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[]

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 ("peaks" that point towards the aperture) and lobes ("valleys" which point away from the aperture). The suture line has four main regions. The external or ventral region refers to sutures along the lower (outer) edge of the shell, where the left and right suture lines meet. The external saddle lies directly on the lower midline of the shell and is edged by external lobes. On suture diagrams the external saddle is supplied with an arrow which typically points towards the aperture. The lateral region involves the first saddle and lobe pair past the external region as the suture line extends up the side of the shell. Additional lobes developing towards the inner edge of a whorl are labelled umbilical lobes, which increase in number through ammonoid evolution as well as an individual ammonoid's development. Lobes and saddles which are so far towards the center of the whorl that they are covered up by succeeding whorls are labelled internal lobes and saddles. 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 is a narrow tubular structure that runs along the shell's outer rim, known as the venter, connecting the chambers of the phragmocone to the body or living chamber. This distinguishes them from living nautiloides (Nautilus and Allonautilus) and typical Nautilida, in which the siphuncle runs through the center of each chamber. However the very earliest nautiloids from the Late Cambrian and Ordovician typically had ventral siphuncles like ammonites, although often proportionally larger and more internally structured. The word "siphuncle" comes from the New Latin siphunculus, meaning "little siphon".

Life history[]

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, because their fossils are often found in rocks 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 larvae in its buccal cavity, indicating at least this kind of ammonite fed on plankton. 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.

Many ammonite shells have been found with round holes once interpreted as a result of limpets attaching themselves to the shells. However, the triangular formation of the holes, their size and shape, and their presence on both sides of the shells, corresponding to the upper and lower jaws, is more likely evidence of the bite of a medium-sized mosasaur preying upon ammonites.

Some ammonites appear to have lived in cold seeps and even reproduced there.

Evolutionary history[]

Goniatites, which were a dominant component of Early and Middle Permian faunas, became rare in the Late Permian, and no goniatite is thought to have crossed into the Triassic.

Ceratitida originated during the Middle Permian, likely from the Daraelitidae, and radiated in the Late Permian. In the aftermath of the Permian–Triassic extinction event, Ceratitids represent the dominant group of Triassic ammonites.

Ammonites were devastated by the end-Triassic extinction, with only a handful of genera belonging to the family Psiloceratidae of the suborder Phylloceratina surviving and becoming ancestral to all later Jurassic and Cretaceous ammonites. Ammonites explosively diversified during the Early Jurassic, with the orders Psiloceratina, Ammonitina, Lytoceratina, Haploceratina, Perisphinctina and Ancyloceratina all appearing during the Jurassic.

Heteromorph ammonites (ammonites with open or non-spiral coiling) of the order Ancyloceratina became common during the Cretaceous period.

At least 57 species of ammonites, which were widespread and belonged to six superfamilies, were extant during the last 500,000 years of the Cretaceous, indicating that ammonites remained highly diverse until the very end of their existence. All ammonites were wiped out during or shortly after the K-Pg extinction event, caused by the Chicxulub impact. It has been suggested that ocean acidification generated by the impact played a key role in their extinction, as the larvae of ammonites were likely small and planktonic, and would have been heavily affected. Nautiloids, exemplified by modern nautiluses, are conversely thought to have had a reproductive strategy in which eggs were laid in smaller batches many times during the lifespan, and on the sea floor well away from any direct effects of such a bolide strike, and thus survived. Many ammonite species were filter feeders, so they might have been particularly susceptible to marine faunal turnovers and climatic change.

Species-based characters[]

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