Anaspida
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Anaspida ("without shield") is an extinct group of primitive jawless vertebrates that lived primarily during the
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 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 shortest period of the Paleoz ...
period, and became extinct soon after the start of the
Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wh ...
. They were classically regarded as the ancestors of
lampreys 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 su ...
. Anaspids were small marine
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 ...
ns that lacked a heavy bony shield and paired fins, but have a striking highly hypocercal tail. They first appeared in the early
Silurian The Silurian ( ) is a geologic period and system spanning 24.6 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 shortest period of the Paleoz ...
, and flourished until the early Devonian, when they disappear from the fossil record.


Anatomy

Compared to other prehistoric
agnathan 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, cyclostome ...
groups, such as the
Heterostraci Heterostraci ( Ancient Greek, ἕτερος+ὄστρακον "those itha different shell" i is pl. of -us is an extinct subclass of pteraspidomorph jawless vertebrate that lived primarily in marine and estuary environments. Heterostraci ...
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, t ...
, 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 Birkeniiformes (Birkeniida or Birkeniids) is an extinct order of jawless fish belonging to the class Anaspida. Subtaxa A newer taxonomy based on the work of Mikko's Phylogeny Archive, Nelson, Grande and Wilson 2016 and van der Laan 2018. * Ge ...
, 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 Sclerotic rings are rings of bone found in the eyes of many animals in several groups of vertebrates, except for mammals and crocodilians. They can be made up of single bones or multiple segments and take their name from the sclera. They are bel ...
, with the
gill A gill () is a respiratory organ that many 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 respiration on land provided they ar ...
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 character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...
for the anaspids is the large, tri-radiate spine behind the series of the gill openings.


Taxonomy

Now that ''
Jamoytius ''Jamoytius kerwoodi'' was a species of primitive, eel-like jawless fish that lived in the Llandovery epoch of the Early Silurian period. Long thought of as a "basal anaspid," ''J. kerwoodi'' is now recognized as the best-known member of the H ...
'' and its close cohorts, i.e., '' Euphanerops'', have been moved to
Jamoytiiformes Jamoytiiformes is an extinct order of prehistoric jawless fish in the superclass Anaspidomorphi Anaspidomorphi (anaspidomorphs) is an extinct superclass of jawless fish. According to the newer taxonomy based on the work of Nelson, Grande and W ...
,Sansom, Robert S., et al. "Taphonomy and affinity of an enigmatic Silurian vertebrate, Jamoytius kerwoodi White." Palaeontology 53.6 (2010): 1393-1409.
Class Class 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 differently ...
Anaspida now consists of two
orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
, 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 "unispec ...
Lasaniida, which contains the genus '' Lasanius'' 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 Birkeniiformes (Birkeniida or Birkeniids) is an extinct order of jawless fish belonging to the class Anaspida. Subtaxa A newer taxonomy based on the work of Mikko's Phylogeny Archive, Nelson, Grande and Wilson 2016 and van der Laan 2018. * Ge ...
, 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 ''Cowielepis'' is an extinct genus of jawless fish in the class Anaspida. It is from the middle Silurian Cowie Harbour fish bed of Stonehaven, Scotland. Habits It was possibly a nectonic filter, of active locomotion, due to its physiognomy. ...
'',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 ''problematica'' is a term used for a 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, uncertain ...
''. 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 Birkeniiformes (Birkeniida or Birkeniids) is an extinct order of jawless fish belonging to the class Anaspida. Subtaxa A newer taxonomy based on the work of Mikko's Phylogeny Archive, Nelson, Grande and Wilson 2016 and van der Laan 2018. * Ge ...
Berg 1940 *** Genus †''
Cowielepis ''Cowielepis'' is an extinct genus of jawless fish in the class Anaspida. It is from the middle Silurian Cowie Harbour fish bed of Stonehaven, Scotland. Habits It was possibly a nectonic filter, of active locomotion, due to its physiognomy. ...
'' 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 ''Pharyngolepis'' is an extinct genus of primitive jawless fish that lived in the Silurian period of what is now Norway. ''Pharyngolepis'' had well-developed anal and caudal fins, but no paired or dorsal fins that would have helped stabilise i ...
Kiær 1924 corrig. **** Genus †'' Pharyngolepis'' Kiaer 1911 *** Family †
Pterygolepididae ''Pterygolepis'' is an extinct genus of jawless fish belonging to the order Birkeniiformes. File:Pterygolepis nitidus Tuebingen.jpg, Fossil of ''Pterygolepis nitidus'', an extinct fish. Took the picture at Museum of Paleontology, Tübingen F ...
Obručhev 1964 corrig. **** Genus †''
Pterygolepis ''Pterygolepis'' is an extinct genus of jawless fish belonging to the order Birkeniiformes Birkeniiformes (Birkeniida or Birkeniids) is an extinct order of jawless fish belonging to the class Anaspida. Subtaxa A newer taxonomy based on th ...
'' Cossmann 1920 'Pterolepis'' Kiaer 1911 non Rambur 1838 non De Candolle ex Miquel 1840; ''Pterolepidops'' Fowler 1947 *** Family †
Rhyncholepididae ''Rhyncholepis'' is an extinct genus of anaspida from the Late Silurian. It was 26 cm long. Fossils have been found in southern Norway Norway, officially the Kingdom of Norway, is a Nordic country in Northern Europe, the mainla ...
Kiær 1924 corrig. **** Genus †''
Rhyncholepis ''Rhyncholepis'' is an extinct genus of anaspida from the Late Silurian. It was 26 cm long. Fossils have been found in southern Norway Norway, officially the Kingdom of Norway, is a Nordic countries, Nordic country in Northern E ...
'' Kiær 1911 non Miquel 1843 non Nuttall 1841 *** Family †
Tahulalepididae Tahulalepididae is an extinct family of jawless fish belonging to the order Birkeniiformes. References External links * Prehistoric jawless fish families Birkeniiformes {{paleo-jawless-fish-stub ...
Blom, Märss & Miller 2002 **** Genus †'' Tahulalepis'' Blom, Märss & Miller 2002 **** Genus †'' Trimpleylepis'' Miller, Märss & Blom 2004 *** Family †
Lasaniidae ''Lasanius'' is a genus of basal anaspida from the Early Silurian. Its fossils are around 10 mm to 150 mm in length. Fossils have been found in Early Silurian-aged strata in Ayrshire, Muirkirk, Scotland Scotland (, ) is a co ...
Goodrich 1909 **** Genus †'' Lasanius'' Traquair 1898 *** Family †
Ramsaasalepididae ''Ramsaasalepis'' is an extinct genus of jawless fish. Its the type and only genus of the family Ramsaasalepididae, and contains the single species ''Ramsaasalepis porosa''. References External links * Birkeniiformes genera Fossil taxa ...
Blom, Märss & Miller 2003 **** Genus †''
Ramsaasalepis ''Ramsaasalepis'' is an extinct genus of jawless fish. Its the type and only genus of the family Ramsaasalepididae, and contains the single species ''Ramsaasalepis porosa''. References External links * Birkeniiformes genera Fossil taxa ...
'' Blom, Märss & Miller 2003 *** Family †
Birkeniidae Birkeniidae is an extinct family of jawless fish belonging to the order Birkeniiformes. They were defined in 1899 by Ramsay H. Traquair, and were a successful family among the many jawless fish in the Silurian seas. Morphology Birkeniidae had ...
Traquair 1899 **** Genus ?†'' Vilkitskilepis'' Märss 2002 **** Genus †''
Ctenopleuron ''Ctenopleuron'' is an extinct genus of jawless fish in the class Anaspida Anaspida ("without shield") is an extinct group of primitive jawless vertebrates that lived primarily during the Silurian period, and became extinct soon after the s ...
'' Matthew 1907 **** Genus †''
Saarolepis ''Saarolepis'' is an extinct genus of jawless fish belonging to the family Birkeniidae. References External links * Birkeniiformes genera Fossil taxa described in 1945 {{paleo-jawless-fish-stub ...
'' Robertson 1945 'Anaspis'' Robertson 1941 non Geoffroy 1762 non Thomson 1893**** Genus †''
Birkenia ''Birkenia'' is a genus of extinct anaspid fish from Middle Silurian strata of Northern Europe, and Middle Silurian to possibly Earliest Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 mil ...
'' Traquair 1898 *** Family †
Septentrioniidae Septentrioniidae is an extinct family of jawless fish belonging to the order Birkeniiformes. References External links * Prehistoric jawless fish families Birkeniiformes {{paleo-jawless-fish-stub ...
Blom, Märss & Miller 2002 **** Genus †''
Liivilepis ''Liivilepis'' is an extinct genus of jawless fish belonging to the family Septentrioniidae Septentrioniidae is an extinct family of jawless fish belonging to the order Birkeniiformes. References External links * Prehistoric jaw ...
'' 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