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Cyclostomi, often referred to as Cyclostomata , is a group of
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
that comprises the living
jawless fish Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct (conodonts and ostracoderms) species. Among recent animals, cyclostomes ...
es: the
lamprey Lampreys (sometimes inaccurately called lamprey eels) are an ancient extant lineage of jawless fish of the order Petromyzontiformes , placed in the superclass Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like s ...
s and
hagfish Hagfish, of the class Myxini (also known as Hyperotreti) and order Myxiniformes , are eel-shaped, slime-producing marine fish (occasionally called slime eels). They are the only known living animals that have a skull but no vertebral column, a ...
es. Both groups have jawless mouths with horny epidermal structures that function as teeth called ceratodontes, and
branchial arch Branchial arches, or gill arches, are a series of bony "loops" present in fish, which support the gills. As gills are the primitive condition of vertebrates, all vertebrate embryos develop pharyngeal arches, though the eventual fate of these arc ...
es that are internally positioned instead of external as in the related
jawed fishes Gnathostomata (; from Greek: (') "jaw" + (') "mouth") 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 ...
. The name Cyclostomi means "round mouths". It was named by
Joan Crockford-Beattie Joan Marion Crockford-Beattie (January 1919 – 2015) was an Australian geologist and palaeontologist who specialised in Permian bryozoan faunas. Early life Joan Marion Crockford was born in January 1919 at Hunters Hill, New South Wales. ...
.


Possible external relationships

This
taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
is often included in the
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
superclass
Agnatha Agnatha (, Ancient Greek 'without jaws') is an infraphylum of jawless fish in the phylum Chordata, subphylum Vertebrata, consisting of both present (cyclostomes) and extinct ( conodonts and ostracoderms) species. Among recent animals, cyclosto ...
, which also includes several groups of extinct armored fishes called
ostracoderm Ostracoderms () are the armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes) (may also be polyphyletic if anaspids are closer to cyclostomes) an ...
s. Most fossil agnathans, such as
galeaspids Galeaspida (from Latin, 'Helmet shields') is an extinct taxon of jawless marine and freshwater fish. The name is derived from ''galea'', the Latin word for ''helmet'', and refers to their massive bone shield on the head. Galeaspida lived in shallo ...
, thelodonts, and osteostracans, are more closely related to vertebrates with jaws (called
gnathostome Gnathostomata (; from Greek: (') "jaw" + (') "mouth") 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 ...
s) than to cyclostomes. Cyclostomes seem to have split off before the evolution of dentine and bone, which are present in many fossil agnathans, including
conodonts Conodonts (Greek ''kōnos'', "cone", + ''odont'', "tooth") are an extinct group of agnathan (jawless) vertebrates resembling eels, classified in the class Conodonta. For many years, they were known only from their tooth-like oral elements, which ...
. Biologists disagree on whether cyclostomes are a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
. The "vertebrate hypothesis" holds that lampreys are more closely related to
gnathostomes Gnathostomata (; from Greek: (') "jaw" + (') "mouth") 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 ...
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 In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. Most studies based on
anatomy Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having its ...
have supported the vertebrate hypothesis, while most molecular phylogenies have supported the cyclostome hypothesis. 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. And at least one molecular phylogeny has supported the vertebrate hypothesis. 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. There is at present no consensus on the correct topology.


Internal differences and similarities

Both hagfishes and lampreys have a single gonad, but for different reasons. In hagfishes the left gonad degenerates during their
ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
and only the right gonad develops, whereas in lampreys the left and right gonads fuse into one. There are no gonoducts present. Hagfishes have direct development, but lamprey go through a larval stage followed by
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
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 The endostyle is an anatomical feature found in invertebrate chordates and larval lampreys. It is an organ which assists chordates in filter-feeding. It is found in adult urochordates and cephalochordates, as well as in the larvae of the vertebra ...
, an organ used for filter feeding that is otherwise found only in
tunicate A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata (). It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one time ca ...
s and
lancelet The lancelets ( or ), also known as amphioxi (singular: amphioxus ), consist of some 30 to 35 species of "fish-like" benthic filter feeding chordates in the order Amphioxiformes. They are the modern representatives of the subphylum Cephalochorda ...
s. During metamorphosis the lamprey endostyle develops into the
thyroid The thyroid, or thyroid gland, is an endocrine gland in vertebrates. In humans it is in the neck and consists of two connected lobes. The lower two thirds of the lobes are connected by a thin band of tissue called the thyroid isthmus. The thy ...
gland. The cyclostomi evolved oxygen transport hemoglobins independently from the jawed vertebrates. Hagfishes and lampreys lack a
thymus The thymus is a specialized primary lymphoid organ of the immune system. Within the thymus, thymus cell lymphocytes or ''T cells'' mature. T cells are critical to the adaptive immune system, where the body adapts to specific foreign invaders. ...
,
spleen The spleen is an organ found in almost all vertebrates. Similar in structure to a large lymph node, it acts primarily as a blood filter. The word spleen comes .
,
myelin Myelin is a lipid-rich material that surrounds nerve cell axons (the nervous system's "wires") to insulate them and increase the rate at which electrical impulses (called action potentials) are passed along the axon. The myelinated axon can be ...
and sympathetic chain ganglia. Neither species has internal eye muscles and hagfishes also lack external eye muscles. 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. 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 cartilagous 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. Unlike jawed vertebrates, which have three
semicircular canal In mathematics (and more specifically geometry), a semicircle is a one-dimensional locus of points that forms half of a circle. The full arc of a semicircle always measures 180° (equivalently, radians, or a half-turn). It has only one line of ...
s in each inner ear, lampreys have only two and hagfishes just one. The semicircular canal of hagfishes contains both
stereocilia Stereocilia (or stereovilli or villi) are non-motile apical cell modifications. They are distinct from cilia and microvilli, but are closely related to microvilli. They form single "finger-like" projections that may be branched, with normal cel ...
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
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...
s. 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. The lamprey intestine has a
typhlosole {{Unreferenced, date=December 2009 A typhlosole is an internal fold of the intestine or intestine inner wall. Typhlosoles occur in bivalve mollusks, lampreys and some annelids and echinoderms. In earthworms, it is a dorsal flap of the intestine ...
that increases the inner surface like the
spiral valve A spiral valve or scroll valve is the corkscrew-shaped lower portion of the intestine of some sharks, Acipenseriformes (sturgeon and paddlefish), rays, skates, bichirs, Lepisosteiformes (gars), and lungfishes. A modification of the ileum, the s ...
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 A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It is ...
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 squirt Ascidiacea, commonly known as the ascidians, tunicates (in part), and sea squirts (in part), is a polyphyletic class in the subphylum Tunicata of sac-like marine invertebrate filter feeders. Ascidians are characterized by a tough outer "tunic" m ...
s such as ''
Ciona ''Ciona'' is a genus of tunicate, sea squirts in the family Cionidae. The body of ''Ciona'' is bag-like and covered by a tunica (biology), tunic, which is a secretion of the Squamous epithelium, epidermal cells. The body is attached at a permanen ...
''. The intestinal epiphelia of lampreys also have ciliated cells, which have not been detected in hagfishes. Because ciliated intestines are also found in
Chondrostei Chondrostei is a group of non-neopterygian ray-finned fish, while the term originally referred to a paraphyletic group of all non-neopterygian ray-finned fish, it was redefined by Patterson in 1982 to be a clade comprising the Acipenseriformes (w ...
, lungfishes and the early stages of some teleosts, it is considered a primitive condition that has been lost in hagfishes.


Phylogeny

After Miyashita et al. 2019.


References

*
Related text and image resources
* Nelson, Joseph S. (2006). ''
Fishes of the World ''Fishes of the World'' by the American ichthyologist Joseph S. Nelson (1937–2011) is a standard reference for fish systematics. Now in its fifth edition (2016), the work is a comprehensive overview of the diversity and classification of the ...
''. John Wiley & Sons, Inc. {{Authority control Agnatha Superclasses (biology) Taxa named by André Marie Constant Duméril