| Homalozoa | |
|---|---|
 | |
| Cothurnocystis , Paleozoic era. | |
Scientific classification  | |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Echinodermata |
| Subphylum: | † Homalozoa Whitehouse, 1941 |
| Groups included | |
| |
Homalozoa is an obsolete extinct subphylum of Paleozoic era echinoderms, prehistoric marine invertebrates. They are also referred to as carpoids. [1]
The Homalozoa lacked the typical pentamer body form of other echinoderms, but all were sessile animals. Instead all Homalozoans were markedly asymmetric, and were extremely variable in forms.
The body (theca) was covered with calcite plates with a number of openings. Their form is in some cases so unusual that it is unclear which openings are to be considered as mouth and anus. Many of them were stalked, similar to sea lilies (crinoids), but often their bodies were bent over, so that the mouth and anus projected forwards rather than upwards. Some forms, especially stylophorans, rested flat on the sea floor. [2]
In some forms the single ray (brachiole or aulacophore) possessed an ambulacral groove. [3]
It has been claimed that some forms possessed gills and gill slits. [4]
Homalozoans were traditionally considered to be stem-group echinoderms, [5] but had also been considered to lie in the stem lineage of the chordates (calcichordates). However, it is now generally accepted that homalozoans were echinoderms because their calcite skeleton was composed of the typical stereom crystalline structure. [6]
They include the unusual stylophorans (mitrates and cornutes), Homoiostelea (solutes), the Homostelea (cinctans), and the Ctenocystoidea (ctenoid-bearing homalozoans). [7] They have recently been recognised as a polyphyletic group. The stylophorans are now classified as a clade of the Crinozoa, whereas the other three are classified as clades of the Blastozoa. [7]
Unlike many other types of echinoderm, solute homalozoans lack radial symmetry (such as the five limbs of a starfish). [8] [9] Solutes are the sole order of the class Homoiostelea.
Solute fossils have an irregularly shaped flattened body covered in calcite plates, and are up to about 10 cm long. The body has two appendages, interpreted as a "feeding arm" at one end, bearing tube feet at its end, and a "stele" at the other, which may have been used by the animal to propel itself along the sea floor. [10]
A chordate is a deuterostomic animal belonging to the phylum Chordata. All chordates possess, at some point during their larval or adult stages, five distinctive physical characteristics (synapomorphies) that distinguish them from other taxa. These five synapomorphies are a notochord, a hollow dorsal nerve cord, an endostyle or thyroid, pharyngeal slits, and a post-anal tail. The name "chordate" comes from the first of these synapomorphies, the notochord, which plays a significant role in chordate body plan structuring and movements. Chordates are also bilaterally symmetric, have a coelom, possess a closed circulatory system, and exhibit metameric segmentation.
An echinoderm is any deuterostomal 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.
Machaeridia is an extinct group of armoured, segmented annelid worms, known from the Early Ordovician to Carboniferous. It consists of three distinct families: the plumulitids, turrilepadids and lepidocoleids.
Crinoids are marine invertebrates that make up the class Crinoidea. Crinoids that remain attached to the sea floor by a stalk in their adult form are commonly called sea lilies, while the unstalked forms, called feather stars or comatulids, are members of the largest crinoid order, Comatulida. Crinoids are echinoderms in the phylum Echinodermata, which also includes the starfish, brittle stars, sea urchins and sea cucumbers. They live in both shallow water and in depths as great as 9,000 meters (30,000 ft).
Bilateria is a large clade or infrakingdom of animals called bilaterians, characterized by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front and a rear end, as well as a left–right–symmetrical belly (ventral) and back (dorsal) surface. Nearly all bilaterians maintain a bilaterally symmetrical body as adults; the most notable exception is the echinoderms, which extend to pentaradial symmetry as adults, but are only bilaterally symmetrical as an embryo. Cephalization is also a characteristic feature among most bilaterians, where the special sense organs and central nerve ganglia become concentrated at the front/rostral end.
Palaeozoology, also spelled as Paleozoology, is the branch of paleontology, paleobiology, or zoology dealing with the recovery and identification of multicellular animal remains from geological contexts, and the use of these fossils in the reconstruction of prehistoric environments and ancient ecosystems.
Cladoselache is an extinct genus of shark-like chondrichthyan from the Late Devonian (Famennian) of North America. It was similar in body shape to modern lamnid sharks, but was not closely related to lamnids or to any other modern (selachian) shark. As an early chondrichthyan, it had yet to evolve traits of modern sharks such as accelerated tooth replacement, a loose jaw suspension, enameloid teeth, and possibly claspers.
Stromatoporoidea is an extinct clade of sea sponges common in the fossil record from the Middle Ordovician to the Late Devonian. They can be characterized by their densely layered calcite skeletons lacking spicules. Stromatoporoids were among the most abundant and important reef-builders of their time, living close together in flat biostromes or elevated bioherms on soft tropical carbonate platforms.
Cothurnocystis is a genus of small enigmatic echinoderms that lived during the Ordovician. Individual animals had a flat boot-shaped body and a thin rod-shaped appendage that may be a stem, or analogous to a foot or a tail. Fossils of Cothurnocystis species have been found in Nevada, Scotland, Czech Republic, France and Morocco.
The stylophorans are an extinct, possibly polyphyletic group allied to the Paleozoic Era echinoderms, comprising the prehistoric cornutes and mitrates. It is synonymous with the subphylum Calcichordata. Their unusual appearances have led to a variety of very different reconstructions of their anatomy, how they lived, and their relationships to other organisms.
Mitrates are an extinct group of stem group echinoderms, which may be closely related to the hemichordates. Along with the cornutes, they form one half of the Stylophora.
Deuterostomes are bilaterian animals of the superphylum Deuterostomia, typically characterized by their anus forming before the mouth during embryonic development. Deuterostomia is further divided into 4 phyla: Chordata, Echinodermata, Hemichordata, and the extinct Vetulicolia known from Cambrian fossils. The extinct clade Cambroernida is also thought to be a member of Deuterostomia.
Cystoidea is a class of extinct crinozoan echinoderms, termed cystoids, that lived attached to the sea floor by stalks. They existed during the Paleozoic Era, in the Middle Ordovician and Silurian Periods, until their extinction in the Devonian Period.
The Fezouata Formation or Fezouata Shale is a geological formation in Morocco which dates to the Early Ordovician. It was deposited in a marine environment, and is known for its exceptionally preserved fossils, filling an important preservational window beyond the earlier and more common Cambrian Burgess shale-type deposits. The fauna of this geological unit is often described as the Fezouata biota, and the particular strata within the formation which exhibit exceptional preservation are generally termed the Fezouata Lagerstätte.
The calcichordate hypothesis holds that each separate lineage of chordate evolved from its own lineage of mitrate, and thus the echinoderms and the chordates are sister groups, with the hemichordates as an out-group.
Stereom is a calcium carbonate material that makes up the internal skeletons found in all echinoderms, both living and fossilized forms. It is a sponge-like porous structure which, in a sea urchin may be 50% by volume living cells, and the rest being a matrix of calcite crystals. The size of openings in stereom varies in different species and in different places within the same organism. When an echinoderm becomes a fossil, microscopic examination is used to reveal the structure and such examination is often an important tool to classify the fossil as an echinoderm or related creature.
Cincta is an extinct class of echinoderms that lived only in the Middle Cambrian epoch. Homostelea is a junior synonym. The classification of cinctans is controversial, but they are probably part of the echinoderm stem group.
Soluta is an extinct class of echinoderms that lived from the Middle Cambrian to the Early Devonian. The class is also known by its junior synonym Homoiostelea. Soluta is one of the four "carpoid" classes, alongside Ctenocystoidea, Cincta, and Stylophora, which made up the obsolete subphylum Homalozoa. Solutes were asymmetric animals with a stereom skeleton and two appendages, an arm extending anteriorly and a posterior appendage called a homoiostele.
Ctenocystoidea is an extinct clade of echinoderms, which lived during the Cambrian and Ordovician periods. Unlike other echinoderms, ctenocystoids had bilateral symmetry, or were only very slightly asymmetrical. They are believed to be one of the earliest-diverging branches of echinoderms, with their bilateral symmetry a trait shared with other deuterostomes. Ctenocystoids were once classified in the taxon Homalozoa, also known as Carpoidea, alongside cinctans, solutes, and stylophorans. Homalozoa is now recognized as a polyphyletic group of echinoderms without radial symmetry. Ctenocystoids were geographically widespread during the Middle Cambrian, with one species surviving into the Late Ordovician.
Jaekelocarpus is a genus of mitrate stylophoran known from the Morrowan age of the Golf Course Formation (southern Oklahoma, US. It possessed two billaterally symmetrical complexes of four internal bars. The morphological similarity of these structures to the gill bars of cephalochordates and enteropneusts is considered a likely indicator of their homology.