Septa (biology)
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Septa (singular septum) are thin walls or partitions between the internal chambers (
camera A camera is an instrument used to capture and store images and videos, either digitally via an electronic image sensor, or chemically via a light-sensitive material such as photographic film. As a pivotal technology in the fields of photograp ...
e) of the
shell Shell may refer to: Architecture and design * Shell (structure), a thin structure ** Concrete shell, a thin shell of concrete, usually with no interior columns or exterior buttresses Science Biology * Seashell, a hard outer layer of a marine ani ...
of a
cephalopod A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
, namely
nautiloid Nautiloids are a group of cephalopods (Mollusca) which originated in the Late Cambrian and are represented today by the living ''Nautilus'' and ''Allonautilus''. Fossil nautiloids are diverse and species rich, with over 2,500 recorded species. Th ...
s or
ammonoid Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...
s. As the creature grows, its body moves forward in the shell to a new living chamber, secreting septa behind it. This adds new chambers to the shell, which can be clearly seen in cross-sections of the shell of the living
nautilus A nautilus (; ) is any of the various species within the cephalopod family Nautilidae. This is the sole extant family of the superfamily Nautilaceae and the suborder Nautilina. It comprises nine living species in two genera, the type genus, ty ...
, or in ammonoid and nautiloid
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 preserve ...
s. The septa are attached to the inside wall of the shell, thus dividing the
phragmocone The phragmocone is the chambered portion of the shell of a cephalopod. It is divided by septa into camerae. In most nautiloids and ammonoids, the phragmocone is a long, straight, curved, or coiled structure, in which the camerae are linked by ...
into camerae. Where the septum meets the shell a suture line forms; in some ammonoids these lines became extremely complex and elaborate, providing strength without the necessity of added weight. Elaborate sutures allowed for thinner shells, and hence less time needed for shell growth and less time spent in the vulnerable juvenile stage. The nature and structure of the septa, as with the camerae, and
siphuncle The siphuncle is a strand of biological tissue, tissue passing longitudinally through the mollusc shell, shell of a cephalopod mollusc. Only cephalopods with chambered shells have siphuncles, such as the extinct ammonites and belemnites, and the li ...
, and the presence or absence of deposits, are important in classification of nautiloids. In some nautiloids, such as the
Orthoceratidae Orthoceratidae, from Ancient Greek ὀρθός (''orthós''), meaning "straight", and κέρας (''kéras''), meaning "horn", is an extinct family of actively mobile carnivorous cephalopods, subclass Nautiloidea, that lived in what would be Nort ...
, the septa tend to be widely spaced, resulting in large, long camarae. In others such as the
Ellesmerocerida The Ellesmerocerida is an extinct order of primitive cephalopods belonging to the subclass Nautiloidea with a widespread distribution that lived during the Late Cambrian and Ordovician. Morphology The Ellesmerocerida are characterized by shell ...
,
Oncocerida The Oncocerida comprise a diverse group of generally small nautiloid cephalopods known from the Middle Ordovician to the Mississippian (early Carboniferous; one possible member is known from the Early Permian), in which the connecting rings are t ...
and
Discosorida Discosorida are an order of cephalopods that lived from the beginning of the Middle Ordovician, through the Silurian, and into the Devonian. Discosorids are unique in the structure and formation of the siphuncle, the tube that runs through and co ...
the septa are crowded closely together. In some straight-shelled forms like ''
Actinoceras ''Actinoceras'' is the principal and root genus of the Actinoceratidae, a major family in the Actinocerida, that lived during the Middle and Late Ordovician. It is an extinct genus of nautiloid cephalopod that thrived in the warm waters of the Un ...
'',
calcium carbonate Calcium carbonate is a chemical compound with the chemical formula . It is a common substance found in Rock (geology), rocks as the minerals calcite and aragonite, most notably in chalk and limestone, eggshells, gastropod shells, shellfish skel ...
deposits extend from the camera (mural deposits) to the septa (episeptal deposits). It is possible to calculate the strength of cephalopod septa on the basis of their thickness and curvature, and from this the shell's implosion depth can be estimated. This has in turn been used to estimate maximum depth ranges for many living and extinct cephalopod groups, on the assumption that these animals would not normally venture deeper than two-thirds of their shell's implosion depth. Ordered by increasing depth, these estimated maximum depth ranges are:
Discosorida Discosorida are an order of cephalopods that lived from the beginning of the Middle Ordovician, through the Silurian, and into the Devonian. Discosorids are unique in the structure and formation of the siphuncle, the tube that runs through and co ...
(<100 m);
Oncocerida The Oncocerida comprise a diverse group of generally small nautiloid cephalopods known from the Middle Ordovician to the Mississippian (early Carboniferous; one possible member is known from the Early Permian), in which the connecting rings are t ...
and
Tarphycerida The Tarphycerida were the first of the coiled cephalopods, found in marine sediments from the Lower Ordovician (middle and upper Canad) to the Middle Devonian. Some, such as '' Aphetoceras'' and '' Estonioceras'', are loosely coiled and gyroconi ...
(<150 m);
Actinoceroidea The Actinocerida are an order of generally straight, medium to large cephalopods that lived during the early and middle Paleozoic, distinguished by a siphuncle composed of expanded segments that extend into the adjacent chambers, in which deposit ...
(50–150 m);
Ellesmerocerida The Ellesmerocerida is an extinct order of primitive cephalopods belonging to the subclass Nautiloidea with a widespread distribution that lived during the Late Cambrian and Ordovician. Morphology The Ellesmerocerida are characterized by shell ...
(50–200 m);
Belemnitida Belemnitida (or belemnites) is an extinct order (biology), order of squid-like cephalopods that existed from the Late Triassic to Late Cretaceous (And possibly the Eocene). Unlike squid, belemnites had an internal skeleton that made up the cone ...
(50–200 m, exceptionally to 350 m);
Bactritoidea The Bactritida are a small order of more or less straight-shelled (orthoconic) cephalopods that first appeared during the Emsian stage of the Devonian period (407 million years ago) with questionable origins in the Pragian stage before 409 mill ...
( 400 m);
Endoceroidea Endocerida, from Ancient Greek ἔνδον (''éndon''), meaning "inside", and κέρας (''kéras''), meaning "horn", is an extinct nautiloid order, a group of cephalopods from the Lower Paleozoic with cone-like deposits in their siphuncle. End ...
(100–450 m);
Orthocerida Orthocerida, from Ancient Greek ὀρθός (''orthós''), meaning "straight", and κέρας (''kéras''), meaning "horn", also known as the Michelinocerida, is an order of extinct Orthoceratoidea, orthoceratoid cephalopods that lived from the ...
(150–500 m);
Nautilida The Nautilida constitute a large and diverse order of generally coiled nautiloid cephalopods that began in the mid Paleozoic and continues to the present with a single family, the Nautilidae which includes two genera, ''Nautilus'' and ''Allonauti ...
(200–600 m);
Aulacocerida Aulacocerida is an order of primitive coleoid cephalopods, possibly derived from michelinoceraitids (Orthocerida) early in the Devonian, which in turn gave rise to the Belemnites. Aulococerids are characterized by a straight, slim, conical she ...
(200–900 m); and
Sepiida Cuttlefish, or cuttles, are marine molluscs of the suborder Sepiina. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control ...
(200–1000 m).Westermann, G.E.G. (1973). Strength of concave septa and depth limits of fossil cephalopods. ''Lethaia'' 6(4): 383–403.


References

{{DEFAULTSORT:Septum (Marine Biology) Cephalopod zootomy Mollusc shells