Tarphycerida

Tarphycerida
Temporal range: Lower Ordovician–Devonian
Scientific classification
Kingdom: Animalia
Phylum: Mollusca
Class: Cephalopoda
Subclass: Nautiloidea
Order: Tarphycerida
Flower, 1950

The Tarphycerida were the first of the coiled cephalopods. They are found in marine sediments from the Lower Ordovician (middle and upper Canad) to the Middle Devonian. Some like Aphetoceras and Estonioceras are loosely coiled, gyroconic, others like Campbelloceras, Tarphyceras, and Trocholites are tightly coiled, but evolute with all whorls showing. The body chamber of tarphycerids is typically long and tubular (Furnish and Glenister 1964), as much as half the length of the containing whorl in most, greater than in the Silurian Ophidioceratidae.

The Tarphycerida evolved from the elongate, compressed, exogastric Bassleroceratidae, probably Bassleroceras, around the end of the Gasconadian through forms like Aphetoceras. Close coiling developed rather quickly and both gyroconic and evolute forms are found in the early middle Canadian.

Tarphycerids tend to uncoil in the late mature stage of their growth, indicating they settled into a benthic lifestyle as they became older. Younger, wholly coiled forms were probably more active, nekto-benthic, certainly more maneuverable

Composition and taxonomy

The Tarphycerida comprise three phylogenetically related groups of families. They are: the tarphyceratid group consisting of the Estonioceratidae, Tarphyceratidae, Trocholitidae, Lituitidae, and Ophidioceratidae; the barradeoceratid group, derived from Centrotarphyceras (Flower 1984), consisting of the Barrandeoceratidae, Bickmoritidae, Nephriticeratidae, and Uranoceratidae; and the plectoceratid group, derived from Campbelloceras (ibid), consisting of the Plectoceratidae, Lechritrochoceratidae, and Apsidoceratidae.

The tarphyceratids comprise the Tarphycerida of the Treatise on Invertebrate Paleontology Part K (Furnish and Glenister 1964) to which Flower added the ancestral Bassleroceratidae. The Estonioceratidae, Tarphyceratidae, and Trocholitidae are primitive forms characterized by siphuncles with thick-walled connecting rings. The Lituitidae and Ophidioceratidae are derived offshoots.

The barrandeoceratid and Plectoceratid families were once combined in the Barrandeocerida (Sweet, 1964), determined by Flower (1984) to be invalid due to having multiple ancestors in the Tarphyceriatidae and therefore abandoned. The common character of these forms are the thin-walled connecting rings in their siphuncles.

Some recent classifications (e.g. Teichert 1988) divide the Tarphycerida into to suborders, the Tarphycerina and Barrandeocerina, which were previously defined as separate orders.

Broader relationships

Tarphycerids are more closely related to the diverse Oncocerida, through the ancestral Bassleroceratidae in the Lower Ordovician, than to the other nautiloid orders. The Oncocerida in turn gave rise to the Nautilida which includes the recent Nautilus and Allonautilus. This puts the Tarphycerida in the broad group that includes the nautilids. It makes them also separate from the groups that include the Discosorida, Actinocerida, Endocerida, and Orthocerida, and from the superficially similar Ammonoidea .( Teichert and Moore 1964, Wade 1988)

Tarphycerid biology

The tarphycerid animal must have been rather elongate, like squid, although no close relationship is implied. Either that or it was able to retreat deep into its portable lair. As for what ever arms or tentacles they may have had, there is no indication. What may be surmised is that they went through two stages, first a younger more active and swimming stage with simple coiled shells, followed by a less active bottom dwelling stage with shells that diverge during which they mated and produced, probably, a single litter of young, like the modern but unrelated coleoids.

References