Cyclostomi Temporal range: Lochkovian - Recent | |
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Sea lamprey from Sweden | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Infraphylum: | Agnatha |
Superclass: | Cyclostomi Duméril, 1806 |
Classes | |
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Cyclostomi, often referred to as Cyclostomata /sɪkloʊˈstɒmətə/ , is a group of vertebrates that comprises the living jawless fishes: the lampreys and hagfishes. Both groups have jawless mouths with horny epidermal structures that function as teeth called ceratodontes, and branchial arches that are internally positioned instead of external as in the related jawed fishes. [1] The name Cyclostomi means "round mouths". [2] [3] [4] It was named by Joan Crockford-Beattie. [5]
This taxon is often included in the paraphyletic superclass Agnatha, which also includes several groups of extinct armored fishes called ostracoderms. Most fossil agnathans, such as galeaspids, thelodonts, and osteostracans, are more closely related to vertebrates with jaws (called gnathostomes) than to cyclostomes. [6] [7]
Biologists historically disagreed on whether cyclostomes are a clade. The "vertebrate hypothesis" held that lampreys are more closely related to gnathostomes than they are to the hagfish. The "cyclostome hypothesis", on the other hand, holds that lampreys and hagfishes are more closely related, making cyclostomi monophyletic. [8] [9]
Most studies based on anatomy have supported the vertebrate hypothesis, [10] while most molecular phylogenies have supported the cyclostome hypothesis. [2] [8] [11] [12]
There are exceptions in both cases, however. Similarities in the cartilage and muscles of the tongue apparatus also provide evidence of sister-group relationship between lampreys and hagfishes. [13] And at least one molecular phylogeny has supported the vertebrate hypothesis. [14] The embryonic development of hagfishes was once held to be drastically different from that of lampreys and gnathostomes, but recent evidence suggests that it is more similar than previously thought, which may remove an obstacle to the cyclostome hypothesis. [15]
Several groups of Paleozoic jawless fish have been suggested to be more closely related to cyclostomes than to jawed fish, including conodonts and anaspids. The presence of mineralised elements in these jawless fish, like the oral conodont elements and the armoured body covering of anaspids and scutes on other species like Lasanius suggests that mineralised tissues were present in the last common ancestor of all vertebrates, but were secondarily lost in hagfish and lampreys. [16]
Both hagfishes and lampreys have a single gonad, but for different reasons. In hagfishes the left gonad degenerates during their ontogeny and only the right gonad develops, whereas in lampreys the left and right gonads fuse into one. There are no gonoducts present. [17] [18]
Hagfishes have direct development, but lamprey go through a larval stage followed by metamorphosis into a juvenile form (or adult form in the non-parasitic species). Lamprey larvae live in freshwater and are called ammocoetes, and are the only vertebrates with an endostyle, an organ used for filter feeding that is otherwise found only in tunicates and lancelets. During metamorphosis the lamprey endostyle develops into the thyroid gland. [19]
The cyclostomi evolved oxygen transport hemoglobins independently from the jawed vertebrates. [20]
Hagfishes and lampreys lack a thymus, spleen, myelin and sympathetic chain ganglia. [21] [22] [23] Neither species has internal eye muscles and hagfishes also lack external eye muscles. [24] Both groups have only a single olfactory organ with a single nostril. The nasal duct ends blindly in a pouch in lampreys but opens into the pharynx in hagfishes. The branchial basket (reduced in hagfishes) is attached to the cranium. [25]
The common ancestor of both cyclostomes and gnathostomes went through a genome duplication before their split, and while a second genome duplicatio occurred in the stem-gnathostomes, the stem-cyclostomes experienced an independent genome triplication. [26]
The mouth apparatus in hagfishes and adult lampreys has some similarities, but differ from one another. Lampreys have tooth plates on the top of a tongue-like piston cartilage, and the hagfish have a fixed cartilaginous plate on the floor of its mouth with groves that allows tooth plates to slide backwards and forwards over it like a conveyor belt, and are everted as they move over the edge of the plate. Hagfishes also have a keratinous palatine tooth hanging from the roof of the mouth. [27] [28]
Unlike jawed vertebrates, which have three semicircular canals in each inner ear, lampreys have only two and hagfishes just one. The semicircular canal of hagfishes contains both stereocilia and a second class of hair cells, apparently a derived trait, whereas lampreys and other vertebrates have stereocilia only. Because the inner ear of hagfishes has two forms of sensory ampullae, their single semicircular canal is assumed to be a result of two semicircular canals that have merged into just one.
The hagfish blood is isotonic with seawater, while lampreys appears to use the same gill-based mechanisms of osmoregulation as marine teleosts. Yet the same mechanisms are apparent in the mitochondria-rich cells in the gill epithelia of hagfishes, but never develops the ability to regulate the blood's salinity, even if they are capable of regulating the ionic concentration of Ca and Mg ions. It has been suggested that the hagfish ancestors evolved from an anadromous or freshwater species that has since adapted to saltwater over a very long time, resulting in higher electrolyte levels in its blood. [29]
The lamprey intestine has a typhlosole that increases the inner surface like the spiral valve does in some jawed vertebrates. The spiral valve in the latter develops by twisting the whole gut, while the lamprey typhlosole is confined to the mucous membrane of the intestines. The mucous membranes of hagfishes have a primitive typhlosole in the form of permanent zigzag ridges. This trait could be a primitive one, since it is also found in some sea squirts such as Ciona . [30] The intestinal epithelia of lampreys also have ciliated cells, which have not been detected in hagfishes. Because ciliated intestines are also found in Chondrostei, lungfishes and the early stages of some teleosts, it is considered a primitive condition that has been lost in hagfishes. [31]
After Miyashita et al. 2019. [32]
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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.
Vertebrates are deuterostomal animals with bony or cartilaginous axial endoskeleton — known as the vertebral column, spine or backbone — around and along the spinal cord, including all fish, amphibians, reptiles, birds and mammals. The vertebrates consist of all the taxa within the subphylum Vertebrata and represent the overwhelming majority of the phylum Chordata, with currently about 69,963 species described.
Hagfish, of the class Myxini and order Myxiniformes, are eel-shaped jawless fish. Hagfish are the only known living animals that have a skull but no vertebral column, although they do have rudimentary vertebrae. Hagfish are marine predators and scavengers who can defend themselves against other larger predators by releasing copious amounts of slime from mucous glands in their skin.
Agnatha is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both living (cyclostomes) and extinct species. Among recent animals, cyclostomes are sister to all vertebrates with jaws, known as gnathostomes.
Gnathostomata are the jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all living vertebrates, including humans. In addition to opposing jaws, living gnathostomes have true teeth, paired appendages, the elastomeric protein of elastin, and a horizontal semicircular canal of the inner ear, along with physiological and cellular anatomical characters such as the myelin sheaths of neurons, and an adaptive immune system that has the discrete lymphoid organs of spleen and thymus, and uses V(D)J recombination to create antigen recognition sites, rather than using genetic recombination in the variable lymphocyte receptor gene.
Fish anatomy is the study of the form or morphology of fish. It can be contrasted with fish physiology, which is the study of how the component parts of fish function together in the living fish. In practice, fish anatomy and fish physiology complement each other, the former dealing with the structure of a fish, its organs or component parts and how they are put together, such as might be observed on the dissecting table or under the microscope, and the latter dealing with how those components function together in living fish.
A craniate is a member of the Craniata, a proposed clade of chordate animals with a skull of hard bone or cartilage. Living representatives are the Myxini (hagfishes), Hyperoartia, and the much more numerous Gnathostomata. Formerly distinct from vertebrates by excluding hagfish, molecular and anatomical research in the 21st century has led to the reinclusion of hagfish as vertebrates, making living craniates synonymous with living vertebrates.
Ostracoderms are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic and thus does not correspond to one evolutionary lineage. However, the term is still used as an informal way of loosely grouping together the armored jawless fishes.
The class Osteostraci is an extinct taxon of bony-armored jawless fish, termed "ostracoderms", that lived in what is now North America, Europe and Russia from the Middle Silurian to Late Devonian.
Pteraspidomorphi is an extinct class of early jawless fish. They have long been regarded as closely related or even ancestral to jawed vertebrates, but the few characteristics they share with the latter are now considered as basal traits for all vertebrates.
Cyclostome is a biological term used in a few different senses:
Marine vertebrates are vertebrates that live in marine environments. These are the marine fish and the marine tetrapods. Vertebrates are a subphylum of chordates that have a vertebral column (backbone). The vertebral column provides the central support structure for an internal skeleton. The internal skeleton gives shape, support, and protection to the body and can provide a means of anchoring fins or limbs to the body. The vertebral column also serves to house and protect the spinal cord that lies within the column.
Metaspriggina is a genus of chordate initially known from two specimens in the Middle Cambrian Burgess Shale and 44 specimens found in 2012 at the Marble Canyon bed in Kootenay National Park.
Lampreys are a group of jawless fish comprising the order Petromyzontiformes. The adult lamprey is characterized by a toothed, funnel-like sucking mouth. The common name "lamprey" is probably derived from Latin lampetra, which may mean "stone licker", though the etymology is uncertain. Lamprey is sometimes seen for the plural form.
Most bony fishes have two sets of jaws made mainly of bone. The primary oral jaws open and close the mouth, and a second set of pharyngeal jaws are positioned at the back of the throat. The oral jaws are used to capture and manipulate prey by biting and crushing. The pharyngeal jaws, so-called because they are positioned within the pharynx, are used to further process the food and move it from the mouth to the stomach.
The evolution of fish began about 530 million years ago during the Cambrian explosion. It was during this time that the early chordates developed the skull and the vertebral column, leading to the first craniates and vertebrates. The first fish lineages belong to the Agnatha, or jawless fish. Early examples include Haikouichthys. During the late Cambrian, eel-like jawless fish called the conodonts, and small mostly armoured fish known as ostracoderms, first appeared. Most jawless fish are now extinct; but the extant lampreys may approximate ancient pre-jawed fish. Lampreys belong to the Cyclostomata, which includes the extant hagfish, and this group may have split early on from other agnathans.
Priscomyzon riniensis is an extinct lamprey that lived some 360 million years ago during the Famennian in a marine or estuarine environment in South Africa. This small agnathan is anatomically similar to the Mazon Creek lampreys, but is some 35 million years older. Its key developments included the first known large oral disc, circumoral teeth and a branchial basket.
Romundina is a small, heavily armored extinct genus of acanthothoracid placoderms which lived in shallow marine environments in the early Devonian (Lochkovian). The name Romundina honors Canadian geologist and paleontologist Dr. Rómundur (Raymond) Thorsteinsson of Calgary, Alberta, Canada. Romundina are believed to have lived on Earth between 400 and 419 million years ago. The closest known relative to Romundina is the acanthothoracid Radotina. The type and only described species is R. stellina.
Myxinikela is an extinct genus of stem-hagfish known from the Late Carboniferous of Illinois, USA. It is the earliest definitive hagfish known from fossil remains, and one of only two alongside the Cretaceous crown-group hagfish Tethymyxine.
Fishes are a paraphyletic group and for this reason, the class Pisces seen in older reference works is no longer used in formal taxonomy. Traditional classification divides fish into three extant classes, and with extinct forms sometimes classified within those groups, sometimes as their own classes: