The calcichordate hypothesis holds that each separate lineage of chordate (Cephalochordates, Urochordates, Craniates) evolved from its own lineage of mitrate, and thus the echinoderms and the chordates are sister groups, with the hemichordates as an out-group. [1]
It was formulated by British Museum paleontologist Richard Jefferies. The hypothesis has been disproven as of 2019; exceptional preservation of soft tissues in the single appendage of the stylophorans Thoralicystis and Hanusia revealed clear traces consistent with a water vascular system—an ambulacral canal with tube feet—covered by movable plates, where the calcichordate hypothesis would require the anatomy be a tail containing a notochord protected by fixed plates. The enlarged area at the base of the appendage, which in the calcichordate hypothesis would contain muscles to move the tail, contains an extension of the body cavity. The findings also disprove the hypothesis that the group were stalked echinoderms. Based on the internal anatomy, "calcichordates" (stylophorans) are revealed as echinoderms with a single, somewhat starfish-like arm that gathered food with tube feet and transferred it to a mouth at the base of the arm. The food-gathering arm may also have been used for locomotion, as in starfish. The fossils from the Bou Izargane Lagerstätte from the Lower Ordivician of Morocco were unknown to Jeffries; they had not been discovered when he formulated the hypothesis in the early 1960s based on the examination of a group of mitrate fossils, and were still unknown when he wrote subsequent papers in 1981 and 1997 defending the hypothesis. Stylophorans are classed as echinoderms based on their possession of at least two shared and unique features (apomorphies) of the phylum; stereom plates and a water vascular system. Because they show no sign of radial symmetry, the position of stylophorans within Echinodermata remains unresolved as of 2019. [2]
The carpoids Cornuta and Mitrata are grouped together in a clade called Calcichordata. Cornutes and mitrates are viewed as sister groups, and mitrates represent stem group chordates. The mitrates (and thus the chordates) are all Dexiothetes, dexiothetism being a synapomorphy for the clade. [3] [4]
In the evolution of the chordates, the ancestors of the chordates underwent a profound remodeling of their bauplan, becoming dexiothetetic. All chordates share a common ancestor which lost its echinoderm stereom calcite skeleton. However, later revisions of the theory had each separate lineage losing its calcitic skeleton independently, [3] as it evolved from its own mitrate ancestor, making the chordates a paraphyletic group[ citation needed ].
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The central part of the Calcichordate Theory lies in the interpretation of the phylogeny of the two groups of stylophorans, which are termed calcichordates in the theory. Mitrates (and the rest of the calcichordates that evolved from them) are dexiothetic as a synapomorphy, having evolved from a cornute. Mitrates are thought to have formed their tail from the proximal part of the cornute tail, with the distal part atomised, [4] and evolving new appendages. The left hand side in this scheme would be cognate with the Pterobranch left-hand side, with the right hand side a novel feature. This would explain the bizarre embryology of Amphioxus, a basal cephalochordate widely held to be the prime example of a chordate bauplan.
The appendage of the carpoids is regarded as a tail, with the central canal probably containing a notochord. [5] The large orifice seen is most likely the mouth, with many of the slits along the side assumed to be gill slits. While the Cornuta were interpreted as lying with the flat side ventrally, Jefferies suggested that in Mitrata the flat side was dorsal and the convex side ventral, while the tail was curved underneath to provide forward thrust; many mitrates are preserved with the tail underneath.
The calcichordate theory is not widely accepted as a viable theory on the origins of the chordates. [4] Many cite its overall unparsimonius nature as unnecessary, however there are many specific points that can be raised. All in all, the carpoids are much closer to echinoderms than to chordates.
Stereom calcite is considered to be a synapomorphy of the echinoderms, and there is no evidence for it ever having evolved in any other lineage, and there is no crown group echinoderm that seems to have lost it secondarily[ citation needed ]. The theory implies either that it must have been lost once or three times, which is considered to be very unlikely.
Chordates are known to exist from the Cambrian with Pikaia , which is around the same time that carpoids are found, although the mitrates may not be stem group.
Genetic evidence has shown that Hemichordata is the sister group to Echinodermata. [6]
A chordate is an animal of the phylum Chordata. All chordates possess five synapomorphies, or primary characteristics, at some point during their larval or adulthood stages that distinguish them from all other taxa. These five synapomorphies include a notochord, dorsal hollow nerve cord, 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 structure and movement. Chordates are also bilaterally symmetric, have a coelom, possess a circulatory system, and exhibit metameric segmentation.
Pikaia gracilens is an extinct, primitive chordate animal known from the Middle Cambrian Burgess Shale of British Columbia. Sixteen specimens are known from the Greater Phyllopod bed, where they comprised 0.03% of the community. It resembled the lancelet and perhaps swam much like an eel.
A cephalochordate is an animal in the chordate subphylum, Cephalochordata. They are commonly called lancelets. Cephalochordates possess 5 synapomorphies, or primary characteristics, that all chordates have at some point during their larval or adulthood stages. These 5 synapomorphies include a notochord, dorsal hollow nerve cord, endostyle, pharyngeal slits, and a post-anal tail. The fine structure of the cephalochordate notochord is best known for the Bahamas lancelet, Asymmetron lucayanum. Cephalochordates are represented in modern oceans by the Amphioxiformes and are commonly found in warm temperate and tropical seas worldwide. With the presence of a notochord, adult amphioxus are able to swim and tolerate the tides of coastal environments, but they are most likely to be found within the sediment of these communities.
Balanoglossus is an ocean-dwelling acorn worm (Enteropneusta) genus of zoological importance because, being a Hemichordate, it is an "evolutionary link" between invertebrates and vertebrates. Balanoglossus is a deuterostome, and resembles the Ascidians or sea squirts, in that it possesses branchial openings, or "gill slits". It has a notochord in the upper part of the body and has no nerve chord. It does have a stomochord, however, which is a gut chord within the collar. Their heads may be as small as per 2.5 mm (1/10 in) or as large as 5 mm (1/5 in).
The dorsal nerve cord is a unique feature to chordates, and it is mainly found in the Vertebrata chordate subphylum. The dorsal nerve cord is only one embryonic feature unique to all chordates, among the other four chordate features-- a notochord, a post-anal tail, an endostyle, and pharyngeal slits. The dorsal hollow nerve cord is a hollow cord dorsal to the notochord. It is formed from a part of the ectoderm that rolls, forming the hollow tube. This is important, as it distinguishes chordates from other animal phyla, such as Annelids and Arthropods, which have solid, ventral tubes. The process by which this is performed is called invagination. The cells essentially convolute into the body cavity, arranging themselves on the dorsal plane above the notochord, as mentioned above. The evolutionary explanation to this adaptation from a solid cord to hollow tube is unknown. In vertebrates, the dorsal nerve cord is modified into the central nervous system, which comprises the brain and spinal cord.
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.
Eleutherozoa is a proposed 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, and some have Tiedemann's bodies on the ring canal. All living echinoderms except Crinozoa and Blastozoa belong here.
Walter Garstang FLS FZS, a Fellow of Lincoln College, Oxford and Professor of Zoology at the University of Leeds, was one of the first to study the functional biology of marine invertebrate larvae. His best known works on marine larvae were his poems published as Larval Forms and Other Zoological Verses, especially The Ballad of the Veliger. They describe the form and function of several marine larvae as well as illustrating some controversies in evolutionary biology of the time.
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.
Deuterostomia are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some examples of deuterostomes include vertebrates, sea stars, and crinoids.
Homalozoa is an obsolete extinct subphylum of Paleozoic era echinoderms, prehistoric marine invertebrates. They are also referred to as carpoids.
The Upper and Lower Fezouata Formations of Morocco are Burgess shale-type deposits dating to the Early Ordovician, filling an important preservational window between the common Cambrian Lagerstätten and the Late Ordovician Soom Shale. Found fossilized fauna were numerous organisms previously thought to have died out after the mid-Cambrian.
In evolutionary developmental biology, inversion refers to the hypothesis that during the course of animal evolution, the structures along the dorsoventral (DV) axis have taken on an orientation opposite that of the ancestral form.
Dexiothetism refers to a reorganisation of a clade's bauplan, with right becoming ventral and left becoming dorsal. The organism would then recruit a new left hand side.
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.
Yanjiahella biscarpa is an extinct species of Early Cambrian deuterostome which may represent the earliest stem group echinoderms.
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