Belemnites
Belemnites
Belemnites
Belemnites, familiar fossils in some rocks, are sometimes referred to as cigars or bullets. In fact they are extinct cephalopod molluscs related to the modern squid and cuttlefish, and their name is appropriately derived from the Greek word belemnos, meaning dart. The first true belemnites appeared in the Lower Jurassic, and the last died out with the ammonites at the end of the Cretaceous, although problematic belemnite-like forms have been found in Carboniferous and Eocene rocks (Eobelemnites and Buyanoteuthzs)
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1 he belemnite possessed a calcareous internal shell consisting of three parts: the cigar-shaped rostrum (or guard), which is the part most commonly found in fossil form, the phragmocone and the proostracum. The rostrum probably acted as a counterbalance for the main bulk of the living body, and its fossil is constructed of a series of massive, compact, radially-arranged calcite crystals. It is now thought that the rostrum of a living belemnite would have had a more open structure, growing concentrically at a probable rate of one major growth stage per year, suggesting a life span of 3-4 years for some species. Less commonly preserved than the rostrum is the phragmocone, a conical chambered shell, whose apical part fitted snugly into a matching cavity (the alveolus) in the anterior of the rostrum.
T h e pro-ostracum is the third, very rarely preserved, skeletal element, which consists of a thin calcareous shield that extended anteriorly from the dorsal surface of the phragmocone and covered the soft parts in life. The phragmocone is analogous to the chambered shell o: the nautiloids, and the proostracum represems a remnant of its body chamber. To some extent, the familiar calcareous cuttlebone of the cuttlefish Sepia resembles an opened out belemnite phragmocone and pro-ostracum. Although belemnites all look very similar to the untrained eye, they may be readily separated into taxonomic groups using the pattern of the grooves that adorn their rostra1 surfaces and by their different shapes, sizes and general form. T h e living belemnite probably possessed lateral fins, and lateral lines on the flanks of some rostra have been interpreted as the point of fin attachment. In addition to fins, the belemnite possessed a head with 10 arms, each equipped with a double row of small, but dangerous-looking, hooks. Ink sacs and surface colour patterns, similar to those of modern cephalopods, have also been found preserved within belemnite remains. Belemnites were probably voracious predators, living in the shallow shelf seas where prey, such as other mollusca and crustacea, abounded. No doubt they were able to move in search of food by gently pulsing their arms. When threatened, they probably moved backwards through a cloud of ink using the jet propulsion mechanism (water ejected through a fleshy tube - the hyponome) common to all modern
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Fig. 1. Reconstruction of the Middle Jurassic belemnite Megateuthts giganteus (young individual). The scale bar represents approximately 0.1 m.
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Fig. 2. Two views of the dimitobelid belemnite Dimitobelus diptychus from the Albian (mid-Cretaceous) of South Australia showing deeply incised alveolar grooves (top) and double lateral lines. The dimitobelids were restricted to the Indo-Pacific region in the Cretaceous. Lenrth = 81 mm.
cephalopods. This defensive system was not always effective, however, as fossil sharks and reptiles such as Ichthyosaurus have been found with stomach contents of hundreds of belemnite hooks - evidence perhaps that only the heads of these unfortunate cephalopods were consumed. M a n y present-day squids congregate in communal spawning grounds in the breeding season, an event often culminating in mass mortality. Similar events may have occurred during the Mesozoic, and possible belemnite spawning grounds are suggested by beds crowded with adult rostra. It is perhaps not suprising that such beds have been termed belemnite battlefields. Differences between sexes seen in the ammonites and the modern cephalopods, such as size, have also been identified in the belemnites. I n early Jurassic times the belemnites, like many ammonites, were fairly cosmopolitan; but by the mid-Cretaceous distinct, geographically separate belemnite groups can be identified in the north (Boreal Realm), the Mediterranean region (Tethyan Realm), and in the Indo-Pacific region (Austral Realm). Possible causes of such separation may have been climatic, such as differences in water temperature, and/or physical, such as differences in water depth. The concept of temperate to cool-temperate
waters for the Boreal and Austral Realms, with warmer Tethyan waters, has been developed from calculation of the proportion of the oxygen isotopes 0 l 6 and O* (a proportion that varies with water temperature) locked into the calcite of belemnite rostra from these regions, but recently the validity of the technique has been questioned. T h e r e is great potential for future belemnite research, and recent discoveries include rostra preserved with their soft parts (in the Posidonia Shale of the German Lower Jurassic). Perhaps this will help revive interest in a neglected group.