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Conulariida are an extinct group of medusozoan
cnidarians Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
known from fossils spanning from the latest
Ediacaran The Ediacaran ( ) is a geological period of the Neoproterozoic geologic era, Era that spans 96 million years from the end of the Cryogenian Period at 635 Million years ago, Mya to the beginning of the Cambrian Period at 538.8 Mya. It is the last ...
up until the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
. They are almost exclusively known from their hard external structures (alternatively referred to as a theca, periderm or test), which were pyramidal in shape and made up of numerous lamellae (thin layers). They are thought to have been sessile animals that grew with the narrower tip anchored to the seafloor, with the wider end bearing an array of
tentacles 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 ...
used to ensnare prey.


Structure

The conulariids are
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 preserved as shell-like structures made up of rows of calcium phosphate rods, resembling an ice-cream cone with fourfold symmetry, usually four prominently-grooved corners. New rods were added as the organism grew in length; the rod-based growth falsely gives the fossils a segmented appearance. Exceptional soft-part preservation has revealed that soft tentacles protruded from the wider end of the cone, and a holdfast from the pointed end attached the organisms to hard substrate. The prevailing reconstruction of the organism has it look superficially like a
sea anemone Sea anemones ( ) are a group of predation, predatory marine invertebrates constituting the order (biology), order Actiniaria. Because of their colourful appearance, they are named after the ''Anemone'', a terrestrial flowering plant. Sea anemone ...
sitting inside an angular, hard cone held perpendicular to the substrate. Conulariid shell is composed of francolite with carbonate ion concentration 8.1 wt%. The lattice parameters of conulariid apatite are a = 9.315(7) Å, c = 6.888(3) Å. The fine structure of their shell comprises multiple lamellae of alternately organic-rich and organic-poor layers.


Fossil record

With the inclusion of the possible
Ediacaran The Ediacaran ( ) is a geological period of the Neoproterozoic geologic era, Era that spans 96 million years from the end of the Cryogenian Period at 635 Million years ago, Mya to the beginning of the Cambrian Period at 538.8 Mya. It is the last ...
conulariid '' Vendoconularia'', which may or may not be a conulariid at all, and the definite late Ediacaran conulariid '' Paraconularia ediacara'', the Conulata fossil record begins with undeniable specimens in the Upper Ediacaran and extends without significant break through numerous major mass extinctions. The Conulariids finally disappear from the fossil record during the
Late Triassic The Late Triassic is the third and final epoch (geology), epoch of the Triassic geologic time scale, Period in the geologic time scale, spanning the time between annum, Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch a ...
, by which time they were very rare, with only 8 documented occurrences across the entire Triassic. Their extinction may have been due to the rise of
durophagous Durophagy is the eating behavior of animals that consume hard-shelled or exoskeleton-bearing organisms, such as corals, shelled mollusks, or crabs. It is mostly used to describe fish, but is also used when describing reptiles, including fossil t ...
organisms as part of the
Mesozoic marine revolution The Mesozoic marine revolution (MMR) refers to the increase in shell-crushing (durophagy, durophagous) and drilling, boring predation in benthic organisms throughout the Mesozoic era (251 Mya (unit), Mya to 66 Mya), along with bulldozing and sed ...
. In North America, conulariids are generally more common in rocks of
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 ...
and
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 ...
age.


Lifestyle

Conulariids were
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
animals that were sessile and attached to a substrate at the base of the theca, older individuals may have become recumbent (tipped over). They are generally proposed to have been predators, using tentacles to ensnare prey.


Phylogeny

About 20 genera and 150 species are known, but except for local occurrences, Conulariids are relatively uncommon. The conulariids were originally thought to be
anthozoa Anthozoa is one of the three subphyla of Cnidaria, along with Medusozoa and Endocnidozoa. It includes Sessility (motility), sessile marine invertebrates and invertebrates of brackish water, such as sea anemones, Scleractinia, stony corals, soft c ...
n cnidarians. However, the lack of septa or other features diagnostic of anthozoans led researchers to abandon this hypothesis. Ivantsov and Fedonkin (2002) posit that the conulariids were ancestrally tri-radially symmetrical, as typified with '' Vendoconularia'', typical of the structure seen in Vendozooans. Conulariids are, however, technically a part of the
Ediacaran biota The Ediacaran (; formerly Vendian) biota is a taxonomic period classification that consists of all life forms that were present on Earth during the Ediacaran Period (). These were enigmatic tubular and frond-shaped, mostly sessile, organis ...
as their fossil record starts at latest parts of that period. It is now also thought that the conulate trilobozoans derived their fourfold symmetry from a sixfold symmetry, as seen in ''Vendoconularia''. This in turn, is thought to be originally derived from an ancestral disk-like
trilobozoa Trilobozoa, from Ancient Greek τρεῖς (''treîs''), meaning "three", λοβός (''lobós''), meaning "lobe", and ζῷον (''zôion''), meaning "animal", is a phylum of extinct, sessile animals that were originally classified into the Cnid ...
n three-fold symmetry. Conulariids have generally been thought to be of
Cnidaria Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
n affinity, occupying a position near the base of the Cnidarian family tree. However, since the 2010s, authors consider conulariids to be members of the subclade
Medusozoa Medusozoa is a clade in the phylum Cnidaria, and is often considered a subphylum. It includes the classes Hydrozoa, Scyphozoa, Staurozoa and Box jellyfish, Cubozoa, and possibly the parasitic Polypodium (animal), Polypodiozoa. Medusozoans are dis ...
, though their exact placement within the clade is uncertain.


Pearls

Conulariids produced
pearl A pearl is a hard, glistening object produced within the soft tissue (specifically the mantle (mollusc), mantle) of a living Exoskeleton, shelled mollusk or another animal, such as fossil conulariids. Just like the shell of a mollusk, a pear ...
s within their shells, similar to the way
molluscs Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
such as oysters, other
bivalve Bivalvia () or bivalves, in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of aquatic animal, aquatic molluscs (marine and freshwater) that have laterally compressed soft bodies enclosed b ...
s, and some
gastropod Gastropods (), commonly known as slugs and snails, belong to a large Taxonomy (biology), taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and fro ...
s do today. These pearls give a clue as to the internal anatomy of the conulariid animal. But due to their calcium phosphate composition, their crystal structure, and their extreme age, these pearls tend to be rather unattractive for use in or as decorative objects.


List of genera

*'' Aciconularia'' *'' Adesmoconularia'' *'' Anaconularia'' *'' Archaeoconularia'' *'' Australoconularia'' *'' Barbigodithreca'' *'' Calloconularia'' *'' Circonularia'' *'' Climacoconus'' *'' Conchopeltis'' *'' Conomedusites''? *'' Conulariella'' *'' Conularia'' *'' Conularina'' *'' Conulariopsis'' *'' Ctenoconularia'' *'' Diconularia'' *'' Eoconularia'' *'' Exoconularia'' *'' Flectoconularia'' *'' Garraconularia'' *'' Glyptoconularia'' *'' Gondaconularia'' *'' Hexangulaconularia'' *'' Holoconularia'' *'' Mabianoconullus'' *'' Mesoconularia'' *'' Metaconularia'' *'' Neoconularia'' *'' Notoconularia'' *'' Palaenigma'' *'' Paraconularia'' *'' Pseudoconularia'' *'' Quadrosiphogonuchites'' *'' Reticulaconularia'' *'' Tasmanoconularia'' *'' Vendoconularia''


References


Bibliography

* * * *


External links

* *http://www.uga.edu/strata/cincy/fauna/conulariida/Conularia.html {{Taxonbar, from=Q2567954 Staurozoa Prehistoric cnidarians Early Triassic extinctions Cambrian genus extinctions Ediacaran first appearances