Anaspid
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Anaspida ("shieldless ones") is an extinct group of
jawless fish Agnatha (; ) or jawless fish is a paraphyletic infraphylum of animals in the subphylum Vertebrata of the phylum Chordata, characterized by the lack of jaws. The group consists of both extant taxon, living (Cyclostomi, cyclostomes such as hagfish ...
that existed from the early
Silurian The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...
period to the late
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
period. They were classically regarded as the ancestors of
lampreys Lampreys (sometimes inaccurately called lamprey eels) are a group of Agnatha, jawless fish comprising the order (biology), order Petromyzontiformes , sole order in the Class (biology), class Petromyzontida. The adult lamprey is characterize ...
, but it is denied in recent
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analysis, although some analysis show these group would be at least related. Anaspids were small marine fish that lacked a heavy bony shield and paired fins, but were distinctively hypocercal.


Anatomy

Compared to many other
ostracoderm Ostracodermi () or ostracoderms is an informal group of vertebrate animals that include all armored jawless fish of the Paleozoic Era. The term does not often appear in classifications today because it is paraphyletic (excluding jawed fishes and ...
s, such as the
Heterostraci Heterostraci (Ancient Greek, ἕτερος+ὄστρακον "those
ith The Ith () is a ridge in Germany's Central Uplands which is up to 439 m high. It lies about 40 km southwest of Hanover and, at 22 kilometers, is the longest line of crags in North Germany. Geography Location The Ith is i ...
a different shell" i is pl. of -us is an extinct subclass (biology), subclass of Pteraspidomorphi, pteraspidomorph, ostracoderm, jawless vertebrate that lived prima ...
and
Osteostraci The class Osteostraci (meaning "bony shells") 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. Anatomically speaking, ...
, anaspids did not possess a bony shield or armor, hence their name. The anaspid head and body are instead covered in an array of small, weakly mineralized scales, with a row of massive scutes running down the back, and, at least confirmed among the birkeniids, the body was covered in rows of tile-like scales made of aspidine, an acellular bony tissue. Anaspids all had prominent, laterally placed eyes with no
sclerotic ring The scleral ring or sclerotic ring is a hardened ring of plates, often derived from bone, that is found in the eyes of many animals in several groups of vertebrates. Some species of mammals, amphibians, and crocodilians lack scleral rings. The rin ...
, with the
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...
s opened as a row of holes along either side of the animal, typically numbering anywhere from 6-15 pairs. The major
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
for the anaspids is the large, tri-radiate spine behind the series of the gill openings.


Taxonomy

Now that ''
Jamoytius ''Jamoytius kerwoodi'' is an extinct species of primitive, eel-like jawless fish known from the Patrick Burn Formation in Scotland, dating to the Llandovery epoch of the Early Silurian period. Long thought of as a "basal anaspid," ''J. kerwood ...
'' and its close cohorts, i.e., '' Euphanerops'', have been moved to
Jamoytiiformes ''Jamoytius kerwoodi'' is an extinct species of primitive, eel-like jawless fish known from the Patrick Burn Formation in Scotland, dating to the Llandovery epoch of the Early Silurian period. Long thought of as a "basal anaspid," ''J. kerwoodi' ...
,Sansom, Robert S., et al. "Taphonomy and affinity of an enigmatic Silurian vertebrate, Jamoytius kerwoodi White." Palaeontology 53.6 (2010): 1393-1409.
Class Class, Classes, or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used d ...
Anaspida now consists of two
orders Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * H ...
, the
monogeneric In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...
Lasaniida, which contains the genus ''
Lasanius ''Lasanius'' is a genus of basal jawless fish from the Early Silurian, around 443.8 million years ago, known from fossils found near Lesmahagow, Scotland. Specimens range from 13.3 to 74.5 mm in length. ''Lasianus'' has a pair of eyes, and a ...
'' and represents a basal anaspid group,Blom, Henning. "New birkeniid anaspid from the Lower Devonian of Scotland and its phylogenetic implications." Palaeontology 55.3 (2012): 641-652. and Birkeniida, which contains all other recognized anaspid taxa.Blom, Henning, T. Märss, and C. G. Miller. "Silurian and earliest Devonian birkeniid anaspids from the Northern Hemisphere." Earth and Environmental Science Transactions of the Royal Society of Edinburgh 92.03 (2001): 263-323. Birkeniida is further divided into several families, including Birkeniidae, Pterygolepididae, Rhyncholepididae and Pharyngolepididae, which contain those taxa known from whole body fossils (in addition to several taxa known only from scales) and the family Septentrioniidae, whose subtaxa are known exclusively from scales. Two recently described genera, '' Kerreralepis'' and '' Cowielepis'',Blom, Henning. "A new anaspid fish from the Middle Silurian Cowie Harbour fish bed of Stonehaven, Scotland." Journal of Vertebrate Paleontology 28.3 (2008): 594-600. are considered to be Birkeniida ''
incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
''. Some recent studies have suggested that anaspids are stem-
cyclostomes Cyclostome is a biological term (from the Greek for "round mouth") used in a few different senses: * for the taxon Cyclostomi, which comprises the extant jawless fishes: the hagfish (Myxini) and the lampreys (Petromyzontidae). This was thought for ...
, more closely related to
hagfish Hagfish, of the Class (biology), class Myxini (also known as Hyperotreti) and Order (biology), order Myxiniformes , are eel-shaped Agnatha, jawless fish (occasionally called slime eels). Hagfish are the only known living Animal, animals that h ...
and
lampreys Lampreys (sometimes inaccurately called lamprey eels) are a group of Agnatha, jawless fish comprising the order (biology), order Petromyzontiformes , sole order in the Class (biology), class Petromyzontida. The adult lamprey is characterize ...
than to
jawed fish Gnathostomata (; from Ancient Greek: (') 'jaw' + (') 'mouth') are jawed vertebrates. Gnathostome diversity comprises roughly 60,000 species, which accounts for 99% of all extant taxon, extant vertebrates, including all living bony fishes (bot ...
. A newer taxonomy based on the work of Mikko's Phylogeny Archive, Nelson, Grande and Wilson 2016 and van der Laan 2018. * Class †Anaspida Janvier 1996 non Williston 1917 ** Order † Endeiolepidiformes Berg 1940 *** Family † Endeiolepididae Stensiö 1939 corrig. **** Genus †'' Endeiolepis'' Stensiö 1939 ** Order † Birkeniiformes Berg 1940 *** Genus †'' Cowielepis'' Blom 2008 *** Genus †'' Hoburgilepis'' Blom, Märss & Miller 2002 *** Genus †'' Kerreralepis'' Blom 2012 *** Genus †'' Maurylepis'' Blom, Märss & Miller 2002 *** Genus †'' Rytidolepis'' Pander 1856 *** Genus †'' Schidiosteus'' Pander 1856 *** Genus †'' Silmalepis'' Blom, Märss & Miller 2002 *** Genus †'' Vesikulepis'' Blom, Märss & Miller 2002 *** Family † Pharyngolepididae Kiær 1924 corrig. **** Genus †'' Pharyngolepis'' Kiaer 1911 *** Family † Pterygolepididae Obručhev 1964 corrig. **** Genus †'' Pterygolepis'' Cossmann 1920 'Pterolepis'' Kiaer 1911 non Rambur 1838 non De Candolle ex Miquel 1840; ''Pterolepidops'' Fowler 1947 *** Family † Rhyncholepididae Kiær 1924 corrig. **** Genus †''
Rhyncholepis ''Rhyncholepis'' was an extinct genus of anaspid from the Late Silurian. Fossils of species ''R. butriangula'' and ''R. parvula'' have been found in Ringerike, Norway, and Oesel, Estonia. The genus Genus (; : genera ) is a taxonomic rank ab ...
'' Kiær 1911 non Miquel 1843 non Nuttall 1841 *** Family † Tahulalepididae Blom, Märss & Miller 2002 **** Genus †'' Tahulalepis'' Blom, Märss & Miller 2002 **** Genus †'' Trimpleylepis'' Miller, Märss & Blom 2004 *** Family † Lasaniidae Goodrich 1909 **** Genus †''
Lasanius ''Lasanius'' is a genus of basal jawless fish from the Early Silurian, around 443.8 million years ago, known from fossils found near Lesmahagow, Scotland. Specimens range from 13.3 to 74.5 mm in length. ''Lasianus'' has a pair of eyes, and a ...
'' Traquair 1898 *** Family † Ramsaasalepididae Blom, Märss & Miller 2003 **** Genus †'' Ramsaasalepis'' Blom, Märss & Miller 2003 *** Family † Birkeniidae Traquair 1899 **** Genus ?†'' Vilkitskilepis'' Märss 2002 **** Genus †'' Ctenopleuron'' Matthew 1907 **** Genus †'' Saarolepis'' Robertson 1945 'Anaspis'' Robertson 1941 non Geoffroy 1762 non Thomson 1893**** Genus †'' Birkenia'' Traquair 1898 *** Family † Septentrioniidae Blom, Märss & Miller 2002 **** Genus †'' Liivilepis'' Blom, Märss & Miller 2002 **** Genus †'' Manbrookia'' Blom, Märss & Miller 2002 **** Genus †'' Ruhnulepis'' Blom, Märss & Miller 2002 **** Genus †'' Spokoinolepis'' Blom, Märss & Miller 2002 **** Genus †'' Septentrionia'' Blom, Märss & Miller 2002


Notes


External links

* * {{Taxonbar, from=Q132837 Prehistoric fish classes Paleozoic jawless fish Llandovery first appearances Early Devonian extinctions