Goniatids, informally goniatites, are
ammonoid
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) ...
cephalopods that form the order Goniatitida, derived from the more primitive
Agoniatitida
Agoniatitida, also known as the Anarcestida, is the ancestral order within the cephalopod subclass Ammonoidea originating from bactritoid nautiloids, that lived in what would become Africa, Asia, Australia, Europe, and North America during t ...
during the Middle
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
some 390 million years ago (around
Eifelian
The Eifelian is the first of two faunal stages in the Middle Devonian Epoch. It lasted from 393.3 ± 1.2 million years ago to 387.7 ± 0.8 million years ago. It was preceded by the Emsian Stage and followed by the Givetian Stage.
North American s ...
stage). Goniatites (goniatitids) survived the
Late Devonian extinction
The Late Devonian extinction consisted of several extinction events in the Late Devonian Epoch, which collectively represent one of the five largest mass extinction events in the history of life on Earth. The term primarily refers to a major exti ...
to flourish during the
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
and
Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
only to become
extinct at the end of the Permian some 139 million years later.
Morphology
All goniatites possessed an external shell, which is divided internally into chambers filled with gas giving it
buoyancy
Buoyancy (), or upthrust, is an upward force exerted by a fluid that opposes the weight of a partially or fully immersed object. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus the p ...
during the life of the animal. An open chamber at the front of the shell provided living space for the goniatitid animal, with access to open water through a ventral siphuncle.
The general morphology and habit of goniatites was probably similar to that of their later relatives the
ammonites
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) ...
, being free swimming and possessing a head with two well developed eyes and arms (or
tentacle
In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work main ...
s).
Goniatite shells are small to medium in size, almost always less than in diameter and often smaller than in diameter. The shell is always planispirally coiled, unlike those of
Mesozoic
The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...
ammonites
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) ...
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 have 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.
Ecology
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 sediments that represent
epicontinental
An inland sea (also known as an epeiric sea or an epicontinental sea) is a continental body of water which is very large and is either completely surrounded by dry land or connected to an ocean by a river, strait, or "arm of the sea". An inland se ...
seas than they are in those that represent the open ocean. Within these inland seas, goniatites' greatest abundance and diversity appears to have been achieved in deeper offshore and
basinal environments rather than in nearshore environments. Known nearshore (e.g.,
lagoon
A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'') a ...
al) 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.,
benthic
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
nekton
Nekton or necton (from the ) refers to the actively swimming aquatic organisms in a body of water. The term was proposed by German biologist Ernst Haeckel to differentiate between the active swimmers in a body of water, and the passive organisms ...
ic,
plankton
Plankton are the diverse collection of organisms found in Hydrosphere, water (or atmosphere, air) that are unable to propel themselves against a Ocean current, current (or wind). The individual organisms constituting plankton are called plankt ...
ic), it remains unclear what resources goniatites were capitalizing on in these offshore environments. Only a few goniatites' full
trophic
Trophic, from Ancient Greek τροφικός (''trophikos'') "pertaining to food or nourishment", may refer to:
* Trophic cascade
* Trophic coherence
* Trophic egg
* Trophic function
* Trophic hormone
* Trophic level index
* Trophic level
* Trop ...
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 in later
ammonites
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) ...
; this has been cited as evidence against their having a
durophagous
Durophagy is the eating behavior of animals that consume hard-shelled or exoskeleton bearing organisms, such as corals, shelled mollusks, or crabs. It is mostly used to describe fish, but is also used when describing reptiles, including fossil tu ...
(shell-crushing) diet.
Distribution
Goniatites are found in
North America
North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
,
Europe
Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
,
North Africa
North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
and
Australasia
Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecologica ...
where they are used as
index fossil
Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock Stratum, strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictiona ...
s in age determination and stratigraphic correlation. 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 the exception of North Dakota, Oregon, Washington, and Hawaii).
Notable goniatite occurrences are found in certain
limestone
Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
s in western part of Ireland (particularly
Slieve Anierin
Sliabh an Iarainn (Irish for "iron mountain"), anglicized Slieve Anierin, is a mountain in County Leitrim, Ireland. It rises to and lies east of Lough Allen and northeast of Drumshanbo. Its present form evolved from the southwestward movement ...
) which are packed with beautifully preserved goniatite
fossil
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s. They are also found in marine bands of the
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
coal measures in Europe and in marine rocks of the
Pennsylvanian period in Arkansas. Large numbers of goniatites also occur in rocks from the
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
period of Morocco.
References
*Miller, Furnish, and Schindewolf (1964); Paleozoic Ammonoidea, in the
Treatise on Invertebrate Paleontology
The ''Treatise on Invertebrate Paleontology'' (or ''TIP'') 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 co ...
, 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)
{{Taxonbar, from=Q2116552
*
Middle Devonian first appearances
Lopingian extinctions
Fossil taxa described in 1884