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Cladus: Eukaryota
Supergroup: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Spiralia
Cladus: Lophotrochozoa
Phylum: Mollusca
Classis: Cephalopoda
Subclassis: †Ammonoidea
Ordo: Goniatitida
Subordines: Goniatitina - Tornoceratatina


Goniatitida Hyatt, 1884

Goniatites are extinct ammonoids, shelled cephalopods related to squid, octopus, and belemnites, that form the order Goniatitida. The Gonatitida originated from within the more primitive anarcestine ammonoids in the Middle Devonian some 390 million years ago. Surviving the Late Devonian extinction, goniatitids flourished during the Carboniferous and Permian only to become extinct at the end of the Permian some 251.4 million years ago. They were survived by their cousins the ceratite ammonoids, indirect descendants of the Anarcestida.


The goniatites all possessed an external shell, which is divided internally into chambers. The animal lived in the largest of the external chambers, and the internal chambers would have been filled with gas, making the animal buoyant in the water. The general structure of the goniatites would have been similar to that of their relatives the ammonites, being a free swimming animal possessing a head with two well developed eyes and arms (or tentacles).
A polished Goniatite fossil

Goniatite shells are small to medium in size, almost always less than 15 centimeters (6 inches) in diameter and often smaller than 5 centimeters (2 inches) in diameter. The shell is always planispirally coiled, unlike those of Mesozoic ammonites in which some are trochoidal and even aberrant (called heteromorphs). Goniatitid shells vary in form from thinly discoidal to broadly globular and may be smooth or distinctly ornamented. Their shape suggests many were poor swimmers

The thin walls between the internal chambers of the shell are called the septa, and as the goniatite grew it would move its body forward in the shell secreting septa behind it, thereby adding new chambers to the shell. The sutures (or suture lines) are visible as a series of narrow, wavy lines on the surface of the shell. The sutures appear where each septa contacts the wall of the outer shell.

The typical goniatitid has a suture with smooth saddles and lobes, which gives the name "goniatitic" to this particular suture pattern. In some the sutures has a distinctive "zigzag" pattern Not all goniatitid ammonoides have goniatitic sutures. In some the sutures are ceratitic, in others, even ammonitic. Nor are goniatitic sutures limited to the Goniatidia. The sutures of nautiloids are by comparison somewhat simpler, being either straight or slightly curved, whereas later ammonoids showed suture patterns of increasing complexity. One explanation for this increasing extravagancy in suture pattern is that it leads to a higher strength of the shell.


Ecologically, goniatites were limited to environments of normal-marine salinity—as appears to be the case for all cephalopods throughout their history. Goniatites are much more abundant and speciose in rocks understood to represent cratonic (also called epicontinental or inland) sea sediments than they are in rocks understood to represent open ocean sediments. Within these inland seas, goniatites' greatest abundance and diversity appears to have been achieved in offshore deep ramp and basinal environments rather than in nearshore environments. Known nearshore (e.g., lagoonal) occurrences have generally been ascribed to wash-in of shells from offshore waters.

Due to lack of strong evidence for any particular life mode (e.g., nektonic, planktonic, demersal, planktivorous, piscivorous), it remains unclear what resources goniatites were capitalizing on in these offshore environments. Only a few goniatites' full trophic apparatuses have ever been described, and reports of stomach contents in these creatures' fossils remain questionable at best. However, goniatites clearly lacked the calcified jaw apparatuses developed later in ammonoid history by the ammonites; this has been cited as evidence against a durophagous (shell-crushing) diet for goniatites.


Goniatites are found in North America, Europe, North Africa and Australasia. However, they seem to occur mostly in areas which at the time would have been tropical to subtropical. Almost any fossil-bearing limestone or shale from inland seas of the late Paleozoic tropics or subtropics is likely to yield some goniatites. In the USA, such rocks are found from Maine, New York, and Virginia and in every state west to Nebraska and south to Texas and Alabama; as well as in parts of almost every western state (with North Dakota, Oregon, Washington, and Hawai'i as the exceptions).

Notable goniatite occurrences include the following: Certain limestones in western part of the Republic of Ireland are packed with beautifully preserved goniatite fossils. They are also found in marine bands of the Carboniferous coal measures in Europe, and in marine rocks of the Pennsylvanian period in Arkansas. Large numbers of goniatites occur in rocks from the Devonian period of Morocco, and they are important zone, or index fossils used in dating the rocks of that period.


* Miller, Furnish, and Schindewolf (1964); Paleozoic Ammonoidea, in the Treatise on Invertebrate Paleontology, Part L, Ammonoidea; Geological Society of America and Univ of Kansas Press (1964)
* Saunders et al. (1999) Evolution of Complexity in Paleozoic Ammonoid Sutures, Supplementary Material. (http//www.sciencemag.org/feature/data/1043099.dtl)

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