Craniidae
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The Craniidae are a
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
of
brachiopods Brachiopods (), phylum Brachiopoda, are a phylum of animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, while the fron ...
, the only surviving members of the subphylum
Craniiformea Craniata is a class of brachiopods originating in the Cambrian period and still extant today. It is the only class within the subphylum Craniiformea, one of three major subphyla of brachiopods alongside linguliforms and rhynchonelliforms. Cran ...
. They are the only members of the
order 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 ...
Craniida, the
monotypic 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 "unisp ...
suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...
Craniidina, and the
superfamily SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
Cranioidea; consequently, the latter two
taxa In biology, a taxon (back-formation from ''taxonomy''; : 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 name and ...
are at present redundant and rarely used. There are three living genera within Craniidae: ''
Neoancistrocrania ''Neoancistrocrania'' is a monotypic genus of brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in ...
'', ''
Novocrania ''Novocrania'' is a genus of brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. ...
'', and ''
Valdiviathyris ''Valdiviathyris'' is a genus of craniate brachiopods that has changed little since the Silurian, from when fossils are known. The extant species ''V. quenstedti'' is known from the late Eocene. It was initially known only from the holotype col ...
''. As adults, craniids either live freely on the ocean floor or, more commonly, cement themselves onto a hard object with all or part of the
ventral Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
valve. Like the burrowing lingulids, Craniids are inarticulate brachiopods. There are no outgrowths forming a hinge between both valves, nor is there any support for the
lophophore The lophophore () is a characteristic feeding organ possessed by four major groups of animals: the Brachiopoda, Bryozoa, Hyolitha, and Phoronida, which collectively constitute the protostome group Lophophorata.apatite Apatite is a group of phosphate minerals, usually hydroxyapatite, fluorapatite and chlorapatite, with high concentrations of Hydroxide, OHβˆ’, Fluoride, Fβˆ’ and Chloride, Clβˆ’ ion, respectively, in the crystal. The formula of the admixture of ...
and organic material, craniids have shells composed mainly of
calcium carbonate Calcium carbonate is a chemical compound with the chemical formula . It is a common substance found in Rock (geology), rocks as the minerals calcite and aragonite, most notably in chalk and limestone, eggshells, gastropod shells, shellfish skel ...
. No craniids are known to bear a
pedicle Pedicle or pedicel may refer to: Human anatomy *Pedicle of vertebral arch, the segment between the transverse process and the vertebral body, and is often used as a radiographic marker and entry point in vertebroplasty and kyphoplasty procedures ...
at any development stage.


Evolution


Extinct craniids

Most craniid genera are
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
, known only from
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...
s like other craniiforms. Craniids first appeared in the later part of the
Tremadocian The Tremadocian is the lowest stage of Ordovician. Together with the later Floian Stage it forms the Lower Ordovician Epoch. The Tremadocian lasted from to million years ago. The base of the Tremadocian is defined as the First appearance datum ...
, the first stage of the
Lower Ordovician The Early Ordovician is the first epoch of the Ordovician period, corresponding to the Lower Ordovician series of the Ordovician system. It began after the Age 10 of the Furongian epoch of the Cambrian and lasted from to million years ago, unt ...
. In the Lower Ordovician, they were mostly restricted to peri-
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
n
terrane In geology, a terrane (; in full, a tectonostratigraphic terrane) is a crust fragment formed on a tectonic plate (or broken off from it) and accreted or " sutured" to crust lying on another plate. The crustal block or fragment preserves its d ...
s (modern central Europe) in the South Polar region. By the Middle Ordovician, they had spread northwards to
Baltica Baltica is a paleocontinent that formed in the Paleoproterozoic and now constitutes northwestern Eurasia, or Europe north of the Trans-European Suture Zone and west of the Ural Mountains. The thick core of Baltica, the East European Craton, i ...
. In the Late Ordovician, their range expanded eastwards to
Avalonia Avalonia was a microcontinent in the Paleozoic era. Crustal fragments of this former microcontinent are terranes in parts of the eastern coast of North America: Atlantic Canada, and parts of the East Coast of the United States, East Coast of the ...
before crossing the
Iapetus Ocean The Iapetus Ocean (; ) existed in the late Neoproterozoic and early Paleozoic eras of the geologic timescale (between 600 and 400 million years ago). It was in the southern hemisphere, between the paleocontinents of Laurentia, Baltica and Avalon ...
to
Laurentia Laurentia or the North American craton is a large continental craton that forms the Geology of North America, ancient geological core of North America. Many times in its past, Laurentia has been a separate continent, as it is now in the form of ...
. Craniid diversity and abundance was respectable but still fairly low during the Late Ordovician, and even lower through the rest of the
Paleozoic The Paleozoic ( , , ; or Palaeozoic) Era is the first of three Era (geology), geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma a ...
. The craniid fossil record is patchy, with fossils unknown from the
Upper Carboniferous Upper may refer to: * Shoe upper or ''vamp'', the part of a shoe on the top of the foot * Stimulant, drugs which induce temporary improvements in either mental or physical function or both * ''Upper'', the original film title for the 2013 found f ...
,
Upper Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the sixth and last period o ...
, and the entire
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
. They reappear in the Oxfordian stage of the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 Β± 1.0 to 143.1 Β± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
and briefly regain their Ordovician level of diversity in the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
.


Living craniids

Craniids are remarkable for their slow rate of
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
. Approximately 11 species of this 480-million-year-old lineage still
survive Survival or survivorship, the act of surviving, is the propensity of something to continue existing, particularly when this is done despite conditions that might kill or destroy it. The concept can be applied to humans and other living things ...
today, with minimal differences relative to their fossil counterparts. One species, '' Valdiviathyris quenstedti,'' has remained essentially unchanged for the last 35 million years or so. Although some evolution would have taken place in the meantime, this was essentially
silent mutation Silent mutations, also called synonymous or samesense mutations, are mutations in DNA that do not have an observable effect on the organism's phenotype. The phrase ''silent mutation'' is often used interchangeably with the phrase '' synonymous mut ...
s and marginal
adaptations In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
to cooler
habitat In ecology, habitat refers to the array of resources, biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species' habitat can be seen as the physical manifestation of its ...
. Present-day ''Valdiviathyris'' are indistinguishable from fossils of the Late
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
, and the
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
cannot even be divided into
chronospecies A chronospecies is a species derived from a sequential development pattern that involves continual and uniform changes from an extinct ancestral form on an evolutionary scale. The sequence of alterations eventually produces a population that is p ...
. ''V. quenstedti'' can be considered a
living fossil A living fossil is a Deprecation, deprecated term for an extant taxon that phenotypically resembles related species known only from the fossil record. To be considered a living fossil, the fossil species must be old relative to the time of or ...
and one of the oldest and most long-lived
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
known to science. ''Valdiviathyris'' and ''Neoancistrocrania'' have occasionally been separated into their own family, Valdiviathyrididae, though few authors follow this suggestion. ''Craniscus'' has sometimes been cited as a fourth living craniid genus,'''' based on "''Craniscus japonica''", a putative species from waters off Japan. Genetic evidence has clarified this misconception, revealing that "''Craniscus japonica''" actually represents a misattributed species of ''Neoancistrocrania''.


List of genera

From the ''
Treatise on Invertebrate Paleontology The ''Treatise on Invertebrate Paleontology,'' published from 1953–2007 by the Geological Society of America and the University of Kansas, then 2009–present by the University of Kansas Paleontological Institute, is a definitive multi-authore ...
Part H, Revised'' (unless stated otherwise): * †'' Acanthocrania'' Williams, 1943
Caradoc Caradoc Vreichvras (; Modern , ) was a semi-legendary ancestor to the kings of Gwent. He may have lived during the 5th or 6th century. He is remembered in the Matter of Britain as a Knight of the Round Table, under the names King Carados and C ...
") – Lower Carboniferous] * †''Ancistrocrania'' Dall, 1877 Synonym (taxonomy), = ''Cranopsis'' [Upper Cretaceous (Coniacian) – Paleocene (Danian?)] * †''Celidocrania'' Liu, Xhy, & Xue, 1985 iddle Ordovician(possible synonym of ''Acanthocrania'') * † ''Crania'' Retzius, 1781 pper Cretaceous (Campanian – Maastrichtian">Campanian.html" ;"title="pper Cretaceous (Campanian">pper Cretaceous (Campanian – Maastrichtian)] * †''Craniscus'' Dall, 1871 [Upper Jurassic ( Oxfordian)] *†'' Conocrania'' Smirnova, 1996 ower Cretaceous (Berriasian – Valanginian">Berriasian.html" ;"title="ower Cretaceous (Berriasian">ower Cretaceous (Berriasian – Valanginian)] *†''Danocrania'' Rozenkrantz, 1964 [Upper Cretaceous (Maastrichtian) – Paleocene (Danian?)] *†''Deliella'' Halamski, 2004 [Lower Devonian (
Emsian The Emsian is one of three faunal stages in the Early Devonian Epoch. It lasted from 410.62 Β±1.95 million years ago to 393.47 Β±0.99 million years ago. It was preceded by the Pragian Stage and followed by the Eifelian Stage. It is named after ...
) – Middle Devonian (
Givetian The Givetian is one of two faunal stages in the Middle Devonian Period. It lasted from million years ago to million years ago. It was preceded by the Eifelian Stage and followed by the Frasnian Stage. It is named after the town of Givet in ...
)] *†'' Isocrania'' JΓ€kel, 1902 pper Cretaceous (Campanian) – Paleocene (Danian?)*†'' Lepidocrania'' Cooper and Grant, 1974 nowiki/>Lower Permian">Lower_Permian.html" ;"title="nowiki/>Lower Permian">nowiki/>Lower Permian*†''Mesocrania'' Smirnova, 1997 [Upper Jurassic (Oxfordian) – Lower Cretaceous (Berriasian)] *†''Nematocrania'' Grant, 1976 [Lower Permian (Artinskian)] *''
Neoancistrocrania ''Neoancistrocrania'' is a monotypic genus of brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in ...
'' Laurin, 1992 *''
Novocrania ''Novocrania'' is a genus of brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. ...
'' Lee and Bruton, 2001 = ''Neocrania'' *†'' Orthisocrania'' Rowell, 1963 rdovician*†'' Petrocrania'' Raymond, 1911 – Lower Carboniferous">ower Ordovician (Tremadocian) – Lower Carboniferous*†'' Philhedra'' Koken, 1889 rdovician*†'' Pseudocrania'' McCoy, 1851 ower Ordovician ("Arenig")*''
Valdiviathyris ''Valdiviathyris'' is a genus of craniate brachiopods that has changed little since the Silurian, from when fossils are known. The extant species ''V. quenstedti'' is known from the late Eocene. It was initially known only from the holotype col ...
'' Helmcke, 1940


Gallery

File:EncrustedStroph.JPG, '' Petrocrania'' (small rounded shells) encrusting an Ordovician strophomenide brachiopod File:Estonian Museum of Natural History Specimen No 193665 photo (g32 g32-3-2 jpg).jpg, '' Pseudocrania'' ''sp''., from the Ordovician of
Estonia Estonia, officially the Republic of Estonia, is a country in Northern Europe. It is bordered to the north by the Gulf of Finland across from Finland, to the west by the Baltic Sea across from Sweden, to the south by Latvia, and to the east by Ru ...
File:Isocrania costata Sowerby 1823.jpg, '' Isocrania costata'', from the Upper Cretaceous of
The Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
File:Novocrania anomala 001.png, Illustration of '' Novocrania anomala'', a living craniid native to the
North Atlantic The Atlantic Ocean is the second largest of the world's five oceanic divisions, with an area of about . It covers approximately 17% of Earth's surface and about 24% of its water surface area. During the Age of Discovery, it was known for ...


References

{{Authority control Brachiopod families Extant Ordovician first appearances Craniata