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Ammonoids are extinct, (typically) coiled-shelled comprising the subclass Ammonoidea. They are more closely related to living , squid, and cuttlefish (which comprise the clade Coleoidea) than they are to (family Nautilidae). (2015). 9789401796323, Springer. ISBN 9789401796323 The earliest ammonoids appeared during the Emsian stage of the Early Devonian (410.62 million years ago), with the last species vanishing during or soon after the Cretaceous–Paleogene extinction event (66 million years ago). They are often called ammonites, which is most frequently used for members of the order Ammonitida, the only remaining group of ammonoids from the Jurassic up until their extinction.
Ammonoids exhibited considerable diversity over their evolutionary history, with over 10,000 species having been described. Ammonoids are excellent index fossils, and they have been frequently used to link rock layers in which a particular species or genus is found to specific geologic time periods. Their fossil shells usually take the form of , although some helically spiraled and nonspiraled forms (known as ) have been found, primarily during the Cretaceous period.
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 sheep' horns. Pliny the Elder ( 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. NH 37.40.167 Often, the name of an ammonite genus ends in - ceras, which is from ancient Greek κέρας () meaning "horn".
In some classifications, these are left as suborders, included in only three orders: Goniatitida, Ceratitida and Ammonitida. The classification of ammonoids is based in part on the ornamentation and structure of the septa comprising their shells' gas chambers.
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 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.
Ammonoid Septum 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. The septal curvature in nautiloids and ammonoids also differ in that the septa curves towards the opening in nautiloids, and away from the opening in ammоnoids. Phylum Mollusca Class Cephalopoda 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 (or ventral) saddle, when present, lies directly on the lower midline of the shell. As a result, it is often called the median saddle. On suture diagrams the median saddle is supplied with an arrow which points towards the aperture. The median saddle is edged by fairly small external (or ventral) lobes. The earliest ammonoids lacked a median saddle and instead had a single midline ventral lobe, which in later forms is split into two or more components.
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. The lateral saddle and lobe are usually larger than the ventral saddle and lobe. 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. In many cases the distinction between the lateral and umbilical regions are unclear; new umbilical features can develop from subdivisions of other umbilical features, or from subdivisions of lateral features. Lobes and saddles which are so far towards the center of the whorl that they are covered up by succeeding whorls are labelled internal (or dorsal) lobes and saddles.
Three major types of suture patterns are found in the Ammonoidea:
Only recently has sexual variation in the shells of ammonites 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, because the dimorphic sizes are so consistently found together, they are more likely an example of sexual dimorphism within the same species.
Whorl width in the body chamber of many groups of ammonites, as expressed by the width:diameter ratio, is another sign of dimorphism. This character has been used to separate "male" (Largiventer conch "L") from "female" (Leviventer conch "l"). (1999). 9781461371939 ISBN 9781461371939
Major shell forms include:
Ammonites vary greatly in the ornamentation (surface relief) of their shells. Some may be smooth and relatively featureless, except for growth lines, resembling that of the modern Nautilus. In others, various patterns of spiral ridges, ribs, nodes, or spines are presented. This type of complex ornamentation of the shell is especially evident in the later ammonites of the Cretaceous.
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 symmetric coiling. Upon closer inspection, though, the shell proves to be a three-dimensional network of connected "U" shapes. Nipponites occurs in rocks of the upper part of the Cretaceous in Japan and the United States.
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 symmetric 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.
Although ammonites do occur in exceptional lagerstatten such as the Solnhofen Limestone, their soft-part record is surprisingly sparse. Beyond a tentative ink sac and possible digestive organs, no soft parts were known until 2021. In this year an isolated specimen showing some of the internal soft anatomy including organs was described. When neutron imaging was used on a fossil found in 1998, part of the musculature became visible and showed they were able to retract themselves into the shell for protection, and that the retractor muscles and hyponome that work together to enable jet propulsion in nautilus worked independently in ammonites. 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 reproductive organs show possible traces of spermatophores, which would support the hypothesis that the microconchs were males. They likely bore a radula and Cephalopod beak, and marginal siphuncle. They operated by direct development with sexual reproduction, were carnivorous, and had a crop for food storage. They are unlikely to have dwelt in fresh or brackish water. Many ammonites were likely , so adaptations associated with this lifestyle like sieves probably occurred. ammonites Garniericeras (left) and Kachpurites (right)|320x320px]], which had a pair of tentacles tipped with hooks, largely unique among ammonites]]A 2021 study reported specimens of the Scaphitidae ammonite genera Rhaeboceras and Hoploscaphites'' with mineralised hooks, which were likely present on the ends of a pair of enlarged tentacles. However, these mineralised hooks appear to be present only in scaphitids and were not typical of ammonites as a whole. The number of arms has been subject considerable speculation, with different artists either opting for a nautilus-like restoration with many arms, or a more squid-like restoration with much fewer arms, with a 1996 study suggesting that they probably had 10 arms like modern squid, cuttlefish and octopuses, but that nothing could be said for certain. Paleontologist Mark Witton has stated that "The basic details of ammonite life appearance are far from clear . . . While we can be certain that a squid-like organism lived in the last chamber of their shells . . . little else can be said with certainty about their appearance. ... Despite being creatures which occur so commonly as fossils that it seems like we should know everything about them, ammonites are creatures fraught with uncertainty for artists and palaeontologists alike. Until new data comes to light, all life reconstructions of ammonites should be taken as extremely tentative, almost speculative renditions of their actual appearance."
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 benthic zone life is found. In general, they appear to have inhabited the upper of the water column. 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 aptychus 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 Cephalopod ink, much like modern cephalopods; ink is occasionally preserved in fossil specimens.
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 and even reproduced there.
During the Permian, the Capitanian mass extinction event severely reduced the diversity of Goniatitida and Prolecanitida, while the Ceratitida, which originated during the Middle Permian, likely from the Daraelitidae, was largely unaffected and radiated in the Late Permian, becoming the dominant group of ammonoids in this period represented by two groups, the Araxoceratidae and Xenodiscidae. The end-Permian mass extinction again reduced ammonoids to the verge of extinction, though both main ceratitd lineages lineages survived, though the xenodiscids were more successful and ancestral to all later ammonoids.
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 , and would have been heavily affected. , exemplified by modern , 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. Some reports suggest that a few ammonite species may have persisted into the very early Danian stage of the Paleocene, before going extinct.
Others believed ammonites, which they referred to as "salagrana" were composed of Coprolite, and could be used to ward off witches.
Ammonites from the Gandaki River in Nepal are known as , and are believed by to be a concrete manifestation of Vishnu.
The Ammonite order, developed by George Dance the Younger, is an Classical order featuring ammonite-shaped that has seen a handful of uses in British architecture. (2025). 9780199674992, Oxford University Press. ISBN 9780199674992
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