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The taxonomy of commonly fossilized invertebrates combines both traditional and modern paleozoological terminology. This article compiles various invertebrate taxa in the fossil record, ranging from protists to arthropods. This includes groups that are significant in paleontological contexts, abundant in the fossil record, or have a high proportion of extinct species. Special notations are explained below:
The paleobiologic systematics that follow are not intended to be comprehensive, rather, they are designed to encompass invertebrates that (a) are popularly collected as fossils and (b) extinct. As a result, some groups of invertebrates are not listed. [1]
If an invertebrate animal is mentioned below using its common (vernacular) name, it is an extant (living) taxon, but if it is cited by its scientific genus, then it is typically an extinct invertebrate known only from the fossil record. [2]
Invertebrate clades that are important fossils (e.g. ostracods, frequently used as index fossils), and clades that are very abundant as fossils (e.g. crinoids, easily found in crinoidal limestone), [3] are highlighted with a bracketed exclamation mark [ ! ].
Eukaryotes are cellular organisms bearing a central, organized nucleus with DNA.
Opisthokonts; the animal-related kingdoms. These include proto-spongal choanoflagellates; proto-fungal microsporidians; and true fungi; true animals.
Metazoans are multicellular "true" animals (multicellular creatures that capture and ingest their organic food).
Parazoans are typically sessile, basal non-eumetazoans. They are the most primitive animals, comprising simple, colonial, attached, bottom-dwelling marine invertebrates.
Cone-shaped archaeocyathids/archeocyathids; cup-shaped archaeocyathans/archeocyathans; reef-building pleosponges; calcareous "ancient-cups".
Includes fossil genera such as Archaeocyathus , Cambrocyathus , Atikonia , Tumuliolynthus , Kotuyicyathus , Metaldetes , Ajacicyathus and Paranacyathus.
Archaeocyatha is sometimes classified as a class of Porifera below.
Quintessential true sponges; marine, colonial, pore-bearing animals; organized collar-flagellates; poriferans - today mostly siliceous – half of all documented species of Porifera are fossils and extinct. [4]
Porifera may eventually be broken up into separate phyla:
Eumetazoans; true metazoans (typically mobile, multicellular animals).
Eumetazoa contains most of the living and deceased species of recorded life, including most invertebrates (extinct and extant), as well as all vertebrate animals.
Radiates; non-bilaterian eumetazoans.
Lophotrochozoan bilaterians, such as flatworms, ribbon worms, lophophorates, and molluscs.
Bryozoans – half of all documented species of Bryozoa are fossils and extinct. [5]
Lampshells, brachiopods or "brachs," (not to be confused with the hard-shelled marine mollusks below) – 99% of all documented species of Brachiopoda are now extinct.
Segmented worms such as earthworms and leeches.
Molluscs or mollusks, not to be confused with the hard-shelled marine brachiopods above.
Ecdysozoans, such as nematodes, horsehair worms, and molting bilaterians/panarthropods
Panarthropodic water bears.
Panarthropodic velvet worms,
Arthropods; jointed legged creatures with an exoskeleton.
Second-mouthed bilaterians called deuterostomians, such as chordates and echinoderms.
Echinoderms – 72% of all documented species of Echinodermata are fossils and extinct. [7]
Hemichordates such as extant acorn worms – Less than half of the documented species of Hemichordata are fossils and extinct.
Both invertebrate and vertebrate chordates; are animals possessing a notochord.
Bryozoa are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about 0.5 millimetres long, they have a special feeding structure called a lophophore, a "crown" of tentacles used for filter feeding. Most marine bryozoans live in tropical waters, but a few are found in oceanic trenches and polar waters. The bryozoans are classified as the marine bryozoans (Stenolaemata), freshwater bryozoans (Phylactolaemata), and mostly-marine bryozoans (Gymnolaemata), a few members of which prefer brackish water. 5,869 living species are known. Originally all of the crown group Bryozoa were colonial, but as an adaptation to a mesopsammal life or to deep-sea habitats, secondarily solitary forms have since evolved. Solitary species has been described in four genera; Aethozooides, Aethozoon, Franzenella and Monobryozoon). The latter having a statocyst-like organ with a supposed excretory function.
Cnidaria is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in fresh water and marine environments, including jellyfish, hydroids, sea anemones, corals and some of the smallest marine parasites. Their distinguishing features are a decentralized nervous system distributed throughout a gelatinous body and the presence of cnidocytes or cnidoblasts, specialized cells with ejectable flagella used mainly for envenomation and capturing prey. Their bodies consist of mesoglea, a non-living, jelly-like substance, sandwiched between two layers of epithelium that are mostly one cell thick. Cnidarians are also some of the only animals that can reproduce both sexually and asexually.
Invertebrates is an umbrella term describing animals that neither develop nor retain a vertebral column, which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordate subphylum Vertebrata, i.e. vertebrates. Well-known phyla of invertebrates include arthropods, mollusks, annelids, echinoderms, flatworms, cnidarians, and sponges.
An echinoderm is any animal of the phylum Echinodermata, which includes starfish, brittle stars, sea urchins, sand dollars and sea cucumbers, as well as the sessile sea lilies or "stone lilies". While bilaterally symmetrical as larvae, as adults echinoderms are recognisable by their usually five-pointed radial symmetry, and are found on the sea bed at every ocean depth from the intertidal zone to the abyssal zone. The phylum contains about 7,600 living species, making it the second-largest group of deuterostomes after the chordates, as well as the largest marine-only phylum. The first definitive echinoderms appeared near the start of the Cambrian.
Archaeocyatha is a taxon of extinct, sessile, reef-building marine sponges that lived in warm tropical and subtropical waters during the Cambrian Period. It is believed that the centre of the Archaeocyatha origin is now located in East Siberia, where they are first known from the beginning of the Tommotian Age of the Cambrian, 525 million years ago (mya). In other regions of the world, they appeared much later, during the Atdabanian, and quickly diversified into over a hundred families.
Sponges or sea sponges are members of the metazoan phylum Porifera, a basal animal clade and a sister taxon of the diploblasts. They are sessile filter feeders that are bound to the seabed, and are one of the most ancient members of macrobenthos, with many historical species being important reef-building organisms.
Parazoa are a taxon with sub-kingdom category that is located at the base of the phylogenetic tree of the animal kingdom in opposition to the sub-kingdom Eumetazoa; they group together the most primitive forms, characterized by not having proper tissues or that, in any case, these tissues are only partially differentiated. They generally group a single phylum, Porifera, which lack muscles, nerves and internal organs, which in many cases resembles a cell colony rather than a multicellular organism itself. All other animals are eumetazoans, which do have differentiated tissues.
Eumetazoa, also known as diploblasts, Epitheliozoa or Histozoa, are a proposed basal animal clade as a sister group of Porifera (sponges). The basal eumetazoan clades are the Ctenophora and the ParaHoxozoa. Placozoa is now also seen as a eumetazoan in the ParaHoxozoa. The competing hypothesis is the Myriazoa clade.
The calcareoussponges are members of the animal phylum Porifera, the cellular sponges. They are characterized by spicules made of calcium carbonate, in the form of high-magnesium calcite or aragonite. While the spicules in most species are triradiate, some species may possess two- or four-pointed spicules. Unlike other sponges, calcareans lack microscleres, tiny spicules which reinforce the flesh. In addition, their spicules develop from the outside-in, mineralizing within a hollow organic sheath.
Stenolaemata are a class of exclusively marine bryozoans. Stenolaemates originated and diversified in the Ordovician, and more than 600 species are still alive today. All extant (living) species are in the order Cyclostomatida, the third-largest order of living bryozoans.
An endoskeleton is a structural frame (skeleton) on the inside of an animal, overlaid by soft tissues and usually composed of mineralized tissue. Endoskeletons serve as structural support against gravity and mechanical loads, and provide anchoring attachment sites for skeletal muscles to transmit force and allow movements and locomotion.
Demosponges (Demospongiae) are the most diverse class in the phylum Porifera. They include greater than 90% of all species of sponges with nearly 8,800 species worldwide. They are sponges with a soft body that covers a hard, often massive skeleton made of calcium carbonate, either aragonite or calcite. They are predominantly leuconoid in structure. Their "skeletons" are made of spicules consisting of fibers of the protein spongin, the mineral silica, or both. Where spicules of silica are present, they have a different shape from those in the otherwise similar glass sponges. Some species, in particular from the Antarctic, obtain the silica for spicule building from the ingestion of siliceous diatoms.
The Treatise on Invertebrate Paleontology published by the Geological Society of America and the University of Kansas Press, is a definitive multi-authored work of some 50 volumes, written by more than 300 paleontologists, and covering every phylum, class, order, family, and genus of fossil and extant invertebrate animals. The prehistoric invertebrates are described as to their taxonomy, morphology, paleoecology, stratigraphic and paleogeographic range. However, taxa with no fossil record whatsoever have just a very brief listing.
Eleutherozoa is a subphylum of echinoderms. They are mobile animals with the mouth directed towards the substrate. They usually have a madreporite, tube feet, and moveable spines of some sort. It includes all living echinoderms except for crinoids. The monophyly of Eleutherozoa has been proven sufficiently well to be considered "uncontroversial."
The Chancelloriids are an extinct family of superficially sponge-like animals common in sediments from the Early Cambrian to the early Late Cambrian. Many of these fossils consists only of spines and other fragments, and it is not certain that they belong to the same type of organism. Other specimens appear to be more complete and to represent sessile, radially symmetrical hollow bag-like organisms with a soft skin armored with star-shaped calcareous sclerites from which radiate sharp spines.
Marine invertebrates are the invertebrates that live in marine habitats. Invertebrate is a blanket term that includes all animals apart from the vertebrate members of the chordate phylum. Invertebrates lack a vertebral column, and some have evolved a shell or a hard exoskeleton. As on land and in the air, marine invertebrates have a large variety of body plans, and have been categorised into over 30 phyla. They make up most of the macroscopic life in the oceans.
Brachiopods, phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the front can be opened for feeding or closed for protection. Two major categories are traditionally recognized, articulate and inarticulate brachiopods. The word "articulate" is used to describe the tooth-and-groove structures of the valve-hinge which is present in the articulate group, and absent from the inarticulate group. This is the leading diagnostic skeletal feature, by which the two main groups can be readily distinguished as fossils. Articulate brachiopods have toothed hinges and simple, vertically oriented opening and closing muscles. Conversely, inarticulate brachiopods have weak, untoothed hinges and a more complex system of vertical and oblique (diagonal) muscles used to keep the two valves aligned. In many brachiopods, a stalk-like pedicle projects from an opening near the hinge of one of the valves, known as the pedicle or ventral valve. The pedicle, when present, keeps the animal anchored to the seabed but clear of sediment which would obstruct the opening.
Rhynchonelliformea is a major subphylum and clade of brachiopods. It is roughly equivalent to the former class Articulata, which was used previously in brachiopod taxonomy up until the 1990s. These so-called articulated brachiopods have many anatomical differences relative to "inarticulate" brachiopods of the subphyla Linguliformea and Craniformea. Articulates have hard calcium carbonate shells with tongue-and-groove hinge articulations and separate sets of simple opening and closing muscles.
Fossils of many types of water-dwelling animals from the Devonian period are found in deposits in the U.S. state of Michigan. Among the more commonly occurring specimens are bryozoans, corals, crinoids, and brachiopods. Also found, but not so commonly, are armored fish called placoderms, snails, sharks, stromatolites, trilobites and blastoids.
The biological systematics and taxonomy of invertebrates as proposed by Richard C. Brusca and Gary J. Brusca in 2003 is a system of classification of invertebrates, as a way to classify animals without backbones.