Belemnoids are an extinct group of marine

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 ...

, very similar in many ways to the modern

squid A squid (: squid) is a mollusc with an elongated soft body, large eyes, eight cephalopod limb, arms, and two tentacles in the orders Myopsida, Oegopsida, and Bathyteuthida (though many other molluscs within the broader Neocoleoidea are also ...

. Like them, the belemnoids possessed an

ink sac An ink sac is an anatomical feature that is found in many cephalopod mollusks used to produce the defensive cephalopod ink. With the exception of nocturnal and very deep water cephalopods, all Coleoidea (squid, octopus and cuttlefish) which dwell ...

, but, unlike the squid, they possessed ten

arms Arms or ARMS may refer to: *Arm or arms, the upper limbs of the body Arm, Arms, or ARMS may also refer to: People * Ida A. T. Arms (1856–1931), American missionary-educator, temperance leader Coat of arms or weapons *Armaments or weapons **Fi ...

of roughly equal length, and no

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. The name "belemnoid" comes from the Greek word βέλεμνον, ''belemnon'' meaning "a dart or arrow" and the Greek word είδος, ''eidos'' meaning "form". Belemnoids include

belemnites Belemnites may refer to: *Belemnitida, an extinct order of cephalopods commonly known as "belemnites" *Belemnites (genus), ''Belemnites'' (genus), a belemnite genus from the Early Jurassic {{disambiguation ...

(which belong to the

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

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 ...

), aulacocerids (order

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 ...

), phragmoteuthids (order Phragmoteuthida), and diplobelids (order Diplobelida). Belemnoidea has been suggested to be paraphyletic by some authors.

Occurrence

Belemnoids were numerous during the

Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...

and

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...

periods, and their

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 are abundant in

Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...

marine rocks, often accompanying their cousins the

ammonite 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. The belemnoids become extinct at the end of the Cretaceous period along with the ammonites. The belemnoids' origin lies within the bactritoid nautiloids, which date from the

Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...

period; well-formed belemnoid guards can be found in rocks dating from the Mississippian (or Early

Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...

) onward through the Cretaceous. Other fossil

s include baculites,

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 and

goniatite Goniatids, informally goniatites, are ammonoid cephalopods that form the order Goniatitida, derived from the more primitive Agoniatitida during the Middle Devonian some 390 million years ago (around Eifelian stage). Goniatites (goniatitids) su ...

Anatomy

Belemnoids possessed a central rostrom/guard made of

aragonite Aragonite is a carbonate mineral and one of the three most common naturally occurring crystal forms of calcium carbonate (), the others being calcite and vaterite. It is formed by biological and physical processes, including precipitation fr ...

and with negative buoyancy. To the rear of the creature was a heavy

calcite Calcite is a Carbonate minerals, carbonate mineral and the most stable Polymorphism (materials science), polymorph of calcium carbonate (CaCO3). It is a very common mineral, particularly as a component of limestone. Calcite defines hardness 3 on ...

guard whose main role appears to have been to counterbalance the front (towards the head) of the organism; it positions the centre of mass below the centre of buoyancy, increasing the stability of the swimming organism. The guard would account for between a third and a fifth of the length of the complete organism, arms included. Like some modern squid, belemnoid arms carried a series of small hooks for grabbing

prey Predation is a biological interaction in which one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not ki ...

. Belemnoids were efficient

carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they ar ...

s that caught small

fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...

and other marine animals with their arms and ate them with their beak-like jaws. In turn, belemnites appear to have formed part of the diet of marine

reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...

s such as

Ichthyosaur Ichthyosauria is an order of large extinct marine reptiles sometimes referred to as "ichthyosaurs", although the term is also used for wider clades in which the order resides. Ichthyosaurians thrived during much of the Mesozoic era; based on fo ...

s, whose fossilized

stomach The stomach is a muscular, hollow organ in the upper gastrointestinal tract of Human, humans and many other animals, including several invertebrates. The Ancient Greek name for the stomach is ''gaster'' which is used as ''gastric'' in medical t ...

s frequently contain

phosphatic Phosphates are the naturally occurring form of the element phosphorus. In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphor ...

hooks from the arms of cephalopods.

Ecology

Belemnoids were effectively neutrally buoyant, and swam in near-shore to mid-shelf oceans. Their fins could be used to their advantage in all water speeds; in a gentle current they could be flapped for propulsion; in a stronger current they could be held erect to generate lift; and when swimming rapidly by jet propulsion they could be tucked in to the body for streamlining.

Preservation

Normally with fossil belemnoids only the back part of the shell (called the ''guard'' or ''rostrum'') is found. The guard is usually elongated and bullet-shaped (though in some subgroups the rostrum may only exist as a thin layer coating the phragmocone). The hollow region at the front of the guard is termed the ''alveolus'', and this houses a chambered conical-shaped part of the shell (called 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 ...

''). The phragmocone is usually only found with the better preserved specimens. Projecting forwards from one side of the phragmocone is the thin ''pro-ostracum''. While belemnoid phragmocones are homologous with the shells of other

cephalopods 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 ...

and are similarly composed of

, belemnoid guards are evolutionarily novel and are composed of

, thus tending to preserve well. Broken guards show a structure of radiating calcite fibers and may also display concentric

growth rings Dendrochronology (or tree-ring dating) is the scientific method of chronological dating, dating tree rings (also called growth rings) to the exact year they were formed in a tree. As well as dating them, this can give data for dendroclimatology, ...

. Diagenetic modifications of the shells are complex. Radiating calcitic crystals are thin, or very large, with a shape indicative of a strong alteration. In other samples, the aragonite - calcite boundary is not dependent on growth lines. In a given fossil sites, some specimens are calcite, others are aragonite. The guard, phragmocone and pro-ostracum were all internal to the living creature, forming a skeleton which was enclosed entirely by soft muscular tissue. The original living creature would have been larger than the fossilized shell, with a long streamlined body and prominent eyes. The guard would have been in place toward the rear of the creature, with the phragmocone behind the head and the pointed end of the guard facing backward. The guard of the belemnoid '' Megateuthis gigantea'', which is found in

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

and

Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...

, can measure up to in length, giving the living animal an estimated length of . Very exceptional belemnoid specimens have been found showing the preserved soft parts of the animal. Elsewhere in the fossil record, bullet-shaped belemnite guards are locally found in such profusion that such deposits are referred to semi-formally as "belemnite battlefields" (cf. "orthocone orgies"). It remains unclear whether these deposits represent post-mating mass death events, as are common among modern

and other

semelparous Semelparity and iteroparity are two contrasting reproductive strategies available to living organisms. A species is considered ''semelparous'' if it is characterized by a single reproduction, reproductive episode before death, and ''iteroparous ...

creatures.

Thunderstones

The name "

thunderbolt A thunderbolt or lightning bolt is a symbolic representation of lightning when accompanied by a loud thunderclap. In Indo-European mythology, the thunderbolt was identified with the 'Sky Father'; this association is also found in later Hel ...

" or "thunderstone" has also been traditionally applied to the fossilised rostra of belemnoids. The origin of these bullet-shaped stones was not understood, and thus a mythological explanation of stones created where lightning struck has arisen.

Uses

The

stable isotope Stable nuclides are Isotope, isotopes of a chemical element whose Nucleon, nucleons are in a configuration that does not permit them the surplus energy required to produce a radioactive emission. The Atomic nucleus, nuclei of such isotopes are no ...

composition of a belemnoid rostrum from the

Peedee Formation The Peedee Formation is a Formation (geology), geologic formation in North Carolina, North and South Carolina. A marine deposit representing an inner Neritic zone, neritic environment, named for exposures along the Great Peedee River, it preserve ...

(

, southeast USA) has long been used as a global standard (Peedee Belemnite, "PDB") against which other

isotope geochemistry Isotope geochemistry is an aspect of geology based upon the study of natural variations in the relative abundances of isotopes of various Chemical element, elements. Variations in isotopic abundance are measured by isotope-ratio mass spectrometry, ...

samples are measured, for both carbon isotopes and

oxygen isotopes There are three known stable isotopes of oxygen (8O): , , and . Radioactive isotopes ranging from to have also been characterized, all short-lived. The longest-lived radioisotope is with a half-life of , while the shortest-lived isotope is ...

. Some belemnoids (such as ''Belemnites'' of

) serve as

index fossil Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. "Biostratigraphy." ''Oxford Reference: Dictionary of Biology ...

s, particularly in the Cretaceous Chalk Formation of Europe, enabling

geologist A geologist is a scientist who studies the structure, composition, and History of Earth, history of Earth. Geologists incorporate techniques from physics, chemistry, biology, mathematics, and geography to perform research in the Field research, ...

s to date the age the

rocks In geology, rock (or stone) is any naturally occurring solid mass or aggregate of minerals or mineraloid matter. It is categorized by the minerals included, its chemical composition, and the way in which it is formed. Rocks form the Earth's ...

in which they are found.

Classification

Note: all families extinct *Clade Belemnoidea ** Basal and unresolved *** Genus '' Jeletzkya'' *** Genus '' Belemnotheutis'' ** Order

*** Family Aulacoceratidae *** Family Dictyoconitidae *** Family Hematitidae *** Family Palaeobelemnopseidae *** Family Xiphoteuthidae ** Order

*** Suborder

Belemnitina Belemnitina is a suborder of belemnites, an extinct group of cephalopods. They have been identified as the oldest of all the belemnites and are likely the stockgroup for them. They were extant from the early Jurassic to the early Cretaceous. Thei ...

**** Family Cylindroteuthidae **** Family Hastitidae **** Family Oxyteuthidae **** Family Passaloteuthidae **** Family Salpingoteuthidae *** Suborder Belemnopseina **** Family Belemnitellidae **** Family Belemnopseidae **** Family Dicoelitidae **** Family Dimitobelidae **** Family Duvaliidae *** Suborder Belemnotheutina **** Family Belemnotheutidae **** Family Chitinobelidae **** Family Sueviteuthidae ** Order Diplobelida *** Family Chondroteuthidae *** Family Diplobelidae ** Order Phragmoteuthida *** Family Phragmoteuthidae *** Family Rhiphaeoteuthidae

References

External links

TONMO.com Cephalopod Fossils articles and discussion forums
{{Taxonbar, from=Q14900832 Mesozoic cephalopods Paleozoic cephalopods Cephalopods by classification Cretaceous cephalopods Carboniferous cephalopods Devonian cephalopods Permian cephalopods Jurassic cephalopods Triassic cephalopods Devonian first appearances Maastrichtian extinctions