Conulariida Temporal range: Late Ediacaran-late Triassic, | |
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Conulariid from the Mississippian (c. 360 to 325 mya) of Indiana; scale in mm. | |
Two Conularia gratiosa specimens from the Salem Limestone aged to the Middle Mississippian. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Cnidaria |
Order: | † Conulatae |
Clade: | † Conulariida Miller and Gurley, 1896 |
Genera | |
See text. |
Conulariida are an extinct group of medusozoan cnidarians known from fossils spanning from the latest Ediacaran up until the Late Triassic. [1] [2] [3] 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.
The conulariids are fossils 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. [4] 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 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. [5] The fine structure of their shell comprises multiple lamellae of alternately organic-rich and organic-poor layers. [6]
With the inclusion of the possible Ediacaran conulariid Vendoconularia , which may or may not be a conulariid at all, [7] and the definite late Ediacaran conulariid Paraconularia ediacara , [2] 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, 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 organisms as part of the Mesozoic marine revolution. [8]
In North America, conulariids are generally more common in rocks of Ordovician and Carboniferous age.
Conulariids were benthic animals that were sessile and attached to a substrate at the base of the theca, older individuals may have become recumbent (tipped over). [9] They are generally proposed to have been predators, using tentacles to ensnare prey. [10]
About 20 genera and 150 species are known, [11] but except for local occurrences, Conulariids are relatively uncommon.
The conulariids were originally thought to be anthozoan 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. [12] Conulariids are, however, technically a part of the Ediacaran biota as their fossil record starts at latest parts of that period. [2]
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 trilobozoan three-fold symmetry.[ citation needed ]
Conulariids have generally been thought to be of Cnidarian 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, though their exact placement within the clade is uncertain. [3]
Conulariids produced pearls within their shells, similar to the way molluscs such as oysters, other pelecypods, and some gastropods 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. [13]
Pteridinium is an erniettomorph found in a number of Precambrian deposits worldwide. It is a member of the Ediacaran biota.
Dickinsonia is a genus of extinct organism, most likely an animal, that lived during the late Ediacaran period in what is now Australia, China, Russia, and Ukraine. It is one of the best known members of the Ediacaran biota. The individual Dickinsonia typically resembles a bilaterally symmetrical ribbed oval. Its affinities are presently unknown; its mode of growth has been considered consistent with a stem-group bilaterian affinity, though various other affinities have been proposed. It lived during the late Ediacaran. The discovery of cholesterol molecules in fossils of Dickinsonia lends support to the idea that Dickinsonia was an animal, though these results have been questioned.
Ausia fenestrata is a curious Ediacaran period fossil represented by only one specimen 5 cm long from the Nama Group, a Vendian to Cambrian group of stratigraphic sequences deposited in the Nama foreland basin in central and southern Namibia. It has similarity to Burykhia from Ediacaran (Vendian) siliciclastic sediments exposed on the Syuzma River of Arkhangelsk Oblast, northwest Russia. This fossil is of the form of an elongate bag-like sandstone cast tapering to a cone on one end. The surface of the fossil is covered with oval depressions ("windows") regularly spaced over the surface in the manner of concentric/parallel rows. The taxonomic identity of Ausia is unresolved.
A trace fossil, also known as an ichnofossil, is a fossil record of biological activity by lifeforms but not the preserved remains of the organism itself. Trace fossils contrast with body fossils, which are the fossilized remains of parts of organisms' bodies, usually altered by later chemical activity or by mineralization. The study of such trace fossils is ichnology - the work of ichnologists.
Kimberella is an extinct genus of bilaterian known only from rocks of the Ediacaran period. The slug-like organism fed by scratching the microbial surface on which it dwelt in a manner similar to the gastropods, although its affinity with this group is contentious.
Yorgia waggoneri is a discoid Ediacaran organism. It has a low, segmented body consisting of a short wide "head", no appendages, and a long body region, reaching a maximum length of 25 cm (9.8 in). It is classified within the extinct animal phylum Proarticulata.
Parvancorina is a genus of shield-shaped bilaterally symmetrical fossil animal that lived in the late Ediacaran seafloor. It has some superficial similarities with the Cambrian trilobite-like arthropods.
Trilobozoa is a phylum of extinct, sessile animals that were originally classified into the Cnidaria. The basic body plan of trilobozoans is often a triradial or radial sphere-shaped form with lobes radiating from its centre. Fossils of trilobozoans are restricted to marine strata of the Late Ediacaran period.
The Ediacaranbiota 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, organisms. Trace fossils of these organisms have been found worldwide, and represent the earliest known complex multicellular organisms. The term "Ediacara biota" has received criticism from some scientists due to its alleged inconsistency, arbitrary exclusion of certain fossils, and inability to be precisely defined.
Albumares brunsae is a tri-radially symmetrical fossil animal that lived in the late Ediacaran seafloor. It is a member of the extinct group Trilobozoa.
Anfesta stankovskii is a tri-radially symmetrical fossil animal that lived in the late Ediacaran (Vendian) seafloor. It is a member of the extinct group Trilobozoa.
Rugoconites is a genus of Ediacaran biota found as fossils in the form of a circular or oval-like impression preserved in high relief, six or more centimeters in diameter. The fossils are surrounded by frills that have been interpreted as sets of tentacles. The bifurcating radial ribs, spreading from a central dome, serve to distinguish this genus from the sponge Palaeophragmodictya, and may represent the channels of the gastrovascular system. Fossils of Rugoconites have been interpreted as early sponges, although this is countered by Sepkoski et al. (2002), who interpreted the organism as a free-swimming jellyfish-like cnidarian; similar to Ovatoscutum. However, the fossil is consistently preserved as a neat circular form and its general morphology does not vary, therefore a benthic and perhaps slow-moving or sessile lifestyle is more likely. Ivantstov & Fedonkin (2002), suggest that Rugoconites may possess tri-radial symmetry and be a member of the Trilobozoa.
The small shelly fauna, small shelly fossils (SSF), or early skeletal fossils (ESF) are mineralized fossils, many only a few millimetres long, with a nearly continuous record from the latest stages of the Ediacaran to the end of the Early Cambrian Period. They are very diverse, and there is no formal definition of "small shelly fauna" or "small shelly fossils". Almost all are from earlier rocks than more familiar fossils such as trilobites. Since most SSFs were preserved by being covered quickly with phosphate and this method of preservation is mainly limited to the late Ediacaran and early Cambrian periods, the animals that made them may actually have arisen earlier and persisted after this time span.
Vendoconularia is a genus of Ediacaran organism consisting of a hexagonal cone, which is thought to have housed a tentaculate organism. Three longitudinal bands are interspersed between the six sides of the cone. The discovery of vendoconulariids in Proterozoic strata of Russia confirmed a 1987 prediction that conulariids constituted part of Ediacaran biota.
Sphenothallus is a problematic extinct genus lately attributed to the conulariids. It was widespread in shallow marine environments during the Paleozoic.
Ventogyrus is an Ediacaran fossil found in the White Sea-Arkhangelsk region of Russia. It was first discovered in the Teska member of the Ust'-Pinega formation, in a thick lens of sandstone, originally sand dumped by storm waves that cut a deep channel through the shallow sea bottom where the organisms lived. Many individuals were preserved on top of each other, often torn or in distorted positions. As a result, it was originally thought to have had a "boat shaped" form and to have lived anchored in the sea floor. However, a nearby site discovered later by Mikhail Fedonkin yielded separate specimens which were beautifully preserved in an upright position and showed the internal anatomy.
The petalonamids (Petalonamae) are an extinct group of archaic animals typical of the Ediacaran biota, also called frondomorphs, dating from approximately 635 million years ago to 516 million years ago. They are benthic and motionless animals, that have the shape of leaves, fronds (frondomorphic), feathers or spindles and were initially considered algae, octocorals or sea pens. It is now believed that there are no living descendants of the group, which shares a probable relation to the Ediacaran animals known as Vendozoans.
Archaeaspinus fedonkini is an extinct proarticulatan organism from the Late Precambrian (Ediacaran) period.
Staurinidia is a genus of Ediacaran soft-bodied organism from the deposits of the Ust'-Pinega formation. It is a monotypic genus, containing only the single species Staurinidia crucicula. The genus was first described in 1985 by Russian palaeontologist Mikhail A. Fedonkin. S. crucicula's four-fold symmetry is present as a result of four canals radiating from the middle of a small cavity in the middle of the body. Other forms with four way symmetry, mainly Medusoid forms, from the Ediacaran comprise an essential chunk of the Ediacaran diversity of symmetry; their organisations are similar to, though smaller than, those of a modern-day Scyphozoan Cnidarian.