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

cephalopod A cephalopod is any member of the molluscan class Cephalopoda (Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head ...

s that first appeared during the Emsian stage of the

Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...

period (407 million years ago) with questionable origins in

Pragian The Pragian is one of three faunal stages in the Early Devonian Epoch. It lasted from 410.8 ± 2.8 million years ago to 407.6 ± 2.8 million years ago. It was preceded by the Lochkovian Stage and followed by the Emsian Stage. The most important La ...

stage before 409 million years ago, and persisted until

Carnian pluvial event The Carnian pluvial episode (CPE), often called the Carnian pluvial event, was an interval of major change in global climate synchronous with significant changes in Earth's biota both in the sea and on land. It occurred during the latter part of ...

in the upper middle Carnian stage of the Triassic period (231 million years ago). They are considered ancestors of the

ammonoid Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttlefish) ...

s, as well as of the

coleoids Subclass Coleoidea, or Dibranchiata, is the grouping of cephalopods containing all the various taxa popularly thought of as "soft-bodied" or "shell-less" (i.e., octopuses, squid and cuttlefish). Unlike its extant sister group, Nautiloidea, whose ...

(

octopus An octopus ( : octopuses or octopodes, see below for variants) 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, cuttle ...

squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...

cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. 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 of ...

, 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 cyrtochonic 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 cyrtochonic 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 a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttle ...

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 Orthocerida is an order of extinct Orthoceratoid cephalopods also known as the Michelinocerida that lived from the Early Ordovician () possibly to the Late Triassic (). A fossil found in the Caucasus suggests they may even have survived until ...

, nautiloid cephalopods that first appeared in the Ordovician. '' Bactroceras'' (''Eobactrites''), an early middle-Ordovician cephalopod, is considered as a true bactritid by some paleontologists, because of its spherical apex and ventral

siphuncle The siphuncle is a strand of tissue passing longitudinally through the shell of a cephalopod mollusk. Only cephalopods with chambered shells have siphuncles, such as the extinct ammonites and belemnites, and the living nautiluses, cuttlefish, and ...

. 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 Ordovician

orthocerid Orthocerida is an order of extinct Orthoceratoid cephalopods also known as the Michelinocerida that lived from the Early Ordovician () possibly to the Late Triassic (). A fossil found in the Caucasus suggests they may even have survived until ...

s, 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

orthocone An orthocone is an unusually long straight shell of a nautiloid cephalopod.; During the 18th and 19th centuries, all shells of this type were named ''Orthoceras'', creating a wastebasket taxon, but it is now known that many groups of nautiloids d ...

s 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