The Bactritida are a small order of more or less straight-shelled (orthoconic)

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 ...

s that first appeared during the Emsian stage of 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 (407 million years ago) with questionable origins in the Pragian stage before 409 million years ago, and persisted until the Carnian pluvial event in the upper middle

Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227.3 ...

stage of the

Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...

period (231 million years ago). They are considered ancestors of the ammonoids, as well as of the coleoids (

octopus An octopus (: octopuses or octopodes) is a soft-bodied, eight-limbed mollusc of the order Octopoda (, ). The order consists of some 300 species and is grouped within the class Cephalopoda with squids, cuttlefish, and nautiloids. Like oth ...

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 ...

cuttlefish Cuttlefish, or cuttles, are Marine (ocean), marine Mollusca, molluscs of the order (biology), suborder Sepiina. They belong to the class (biology), class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique ...

, and the extinct belemnites). Bactritids are distinguished from the more primitive nautiloids by the small size and globular shape of the protoconch, the so-called embryonic shell. Nautiloids have relatively large embryonic shells, and living species lay a few large eggs. In contrast, bactritids and ammonoids produced large numbers of small eggs, each housing a small embryonic shell.

Classification and description

Bactritida (Erben 1964) are characterized by orthoconic to cyrtoconic shells that may be long or short with a narrow siphuncle invariably in contact with the ventral wall and sutures uniformly with V-shaped ventral lobes. Septal necks are orthochoanitic to cyrtochoanitic, the apical angle may be small or large, and the protoconch is globular to egg-shaped. The Bactritida comprise two families, the Bactritidae and the Parabactritidae.

Bactritidae

The BactritidaeErben H.K. 1964; Bactritoidea, in the Treatise on Invertebrate Paleontology Vol K Nautiloidea, K491 –, Teichert, C and Moore R.C. Eds, GSA and Univ Kansas Press. 1964) are characterized by long orthoconic to cyrtoconic shells with a small apical angle (less than 10°) and septal necks that are orthochoanitic. Chamber length is variable. The Bactritidae contain eight recognized genera. ''Bactrites'' has the longest range, from the Lower Devonian to the Upper Permian, and even possibly from the Silurian. The Bactritidae gave rise to the

Ammonoidea 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 ...

in the Early Devonian starting with an early ''Bactrites'' and going with increasingly tight curvature from ''Lobobactrites'' to ''Cyrtobactrites'', leading to the gyroconic '' Anetoceras'' of the Anarcestida (Ammonoidea).Klug, C. and Korn, D. 2004. The origin of ammonoid locomotion. Acta Palaeontologica Polonica 49 (2): 235–242

Parabactritidae

The Parabactritidae are characterized by orthoconic and breviconic shells with a large apical angle (greater than about 10°) and septal necks that are vary from orthochoanitic or suborthochoanitic to cyrtochoanitic. The Parabactritidae contain some five described genera and are thought to have given rise to the Belemnoidea (Coleoidea).

Derivation

The Bactritida have their origin in the Orthocerida, nautiloid cephalopods that first appeared in the

Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...

. '' Bactroceras'' (''Eobactrites''), an early middle-Ordovician cephalopod, is considered as a true bactritid by some paleontologists, because of its spherical

apex The apex is the highest point of something. The word may also refer to: Arts and media Fictional entities * Apex (comics) A-Bomb Abomination Absorbing Man Abraxas Abyss Abyss is the name of two characters appearing in Ameri ...

and

ventral Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...

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 ...

. This view is challenged by more recent research. The shell of ''Bactroceras'' has important differences from those of true bactritids. For instance, the first chamber of ''Bactroceras'' resembles that of other

orthocerids, such as ''Archigeisonoceras'' and ''Hedstroemoceras'': it is about 10 mm in diameter, and is short, forming a spherical cap. True bactritids more strongly resemble late Silurian and Devonian orthocerids, whose first chamber is only about 5 mm across. Moreover, a large stratigraphic gap of nearly 50 million years occurs between ''Bactroceras'' and the next orthocones with a spherical apex and ventral siphuncle.

Paleoecology and lifestyle

Bactritids, especially the Bactritidae, seem to have lived in a vertical orientation, head down, as probably did many of the Orthocerida such as ''

Michelinoceras ''Michelinoceras'' is the oldest known genus of the Michelinocerida, more commonly known as the Orthocerida, characterized by long, slender, nearly cylindrical orthocones with a circular cross section, long camerae, very long body chambers, and a ...

'' and '' Buttsoceras'', or these forms without extensive internal ballast may simply have been more flexible in their vertical orientation than the well-ballasted bottom-dwellers such as the actinocerids and endocerids. The ventral siphuncle of the Bactritida would have facilitated chamber dewatering in either horizontal or vertical orientation or in between. A central siphuncle would have made horizontal or near-horizontal dewatering more difficult, though. Bactritids, like cephalopods in general, were probably active predators with tentacles of some sort. As with other externally shelled forms, they were probably not active swimmers in the sense of fish or squid. Rather, they were likely stalkers and ambush predators hiding among the seaweeds, snatching prey found below.

References

{{Taxonbar, from=Q26704395 Prehistoric nautiloids Prehistoric cephalopod orders Paleozoic cephalopods Paleozoic cephalopods of Europe Transitional fossils Carboniferous cephalopods Devonian cephalopods Permian cephalopods Triassic cephalopods Early Devonian first appearances Carnian extinctions Paraphyletic groups Neocephalopoda