Belemnitida


Belemnitida (or the belemnite) is an extinct order of squid-like cephalopods that existed from the Late Triassic to Late Cretaceous. Unlike squid, belemnites had an internal skeleton that made up the cone. The parts are, from the arms-most to the tip: the tongue-shaped pro-ostracum, the conical phragmocone, and the pointy guard. The calcitic guard is the most common belemnite remain. Belemnites, in life, are thought to have had 10 hooked arms and a pair of fins on the guard. The chitinous hooks were usually no bigger than 5 mm (0.20 in), though a belemnite could have had between 100 and 800 hooks in total, using them to stab and hold onto prey.

Belemnites were an important food source for many Mesozoic marine creatures, both the adults and the planktonic juveniles, and likely played an important role in restructuring marine ecosystems after the Triassic–Jurassic extinction event. They may have laid between 100 and 1,000 eggs. Some species may have been adapted to speed and swam in the turbulent open ocean, whereas others resided in the calmer littoral zone (nearshore) and fed off the seafloor. The largest belemnite known, Megateuthis elliptica, had guards of 60 to 70 cm (24 to 28 in).

Belemnites were coleoids, a group that includes squid and octopuses, and are often grouped into the superorder Belemnoidea, though the higher classification of cephalopods is volatile and there is no clear consensus how belemnites are related to modern coleoids. Guards can give information on the climate, habitat, and the carbon cycle of the ancient waters they inhabited. Guards have been found since antiquity and have become part of folklore.

The belemnite cone is composed of three parts. Going from arms to tip, these are the tongue-shaped pro-ostracum; the conical, chambered phragmocone; and the spear-shaped guard at the very tip.[2][3] The guard attached to the phragmocone in a socket called the alveolus.[3][4] The cone, in life, would have been encased in muscle and connective tissue. They had calcite guards,[5] and aragonite pro-ostraca and phragmocones,[2] though a few belemnites also had aragonite guards,[6] and the alveolar side of the guards of belemnitellids may have also been of aragonite.[4] The pro-ostracum probably supported the soft parts of the belemnite, similar to the gladius of squid, and completely surrounded the phragmocone.[3][7]

The phragmocone was divided by septa into chambers, much like the shells of cuttlefish and nautiluses.[7] The chambered phragmocone was probably the center of buoyancy, and so was positioned directly above the center of mass for stability purposes. In regards to buoyancy, belemnites may have behaved much like modern ram's horn squid, having the chambers of the phragmocone flooded and slowly releasing more seawater via the siphuncle tube as the animal increases in size and weight over its lifetime to maintain neutral buoyancy.[2] At the tip of the phragmocone beneath the guard is a tiny, cup-like protoconch, the remains of the embryonic shell.[1][7]

The dense guard probably served to counterbalance the weight of the soft parts in the mantle cavity near the arms on the opposite end of the animal, analogous to the camera of nautiloids. This would have allowed the animal to move horizontally through the water.[2][3] The guard may have also served to cut through waves while swimming at the surface, though modern cephalopods generally stay completely submerged. Though unlikely, it is possible fossilization increased the perceived density of the guard, and it may have been up to 20% more porous in life. Fins may have been attached to the guard, or the guard may have lent support for large fins. Including arms, guards could have accounted for one fifth to one third of the total length of a belemnite.[2]


Cone diagram:
   r pro-ostracum,
   a alveolus,
   p phragmocone,
   g guard
Preserved soft body elements of the Late Jurassic Acanthoteuthis (above) and the Early Jurassic Passaloteuthis (below)
Reconstruction of a typical belemnite
The Jurassic Youngibelus reconstructed with hectocotyli
Cephalopod embryonic shells. A) Ammonoidea (B) Sepiida (C) Belemnoidea (D) Spirulida (E) Orthocerida (F) Nautilida (G) Oncocerida (H) Pseudorthocerida (I) Oegopsina (J) Bactritida. IC indicates the initial chamber.
MRI of a deformed Late Cretaceous Gonioteuthis guard
Map of the world in the Late Triassic
Opalized Peratobelus guard from the Early Cretaceous
Hibolites from the early Cretaceous at the State Museum of Natural History Stuttgart
Belemnites with Early Jurassic Dactylioceras ammonites
The shark Hybodus with belemnite guards in its stomach, State Museum of Natural History Stuttgart
A large aggregation of belemnite guards