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==Taxonomy and phylogeny== Squid are members of the class [[Cephalopod]]a, subclass [[Coleoidea]]. The squid [[Order (taxonomy)|orders]] [[Myopsida]] and [[Oegopsida]] are in the superorder [[Decapodiformes]] (from the [[Greek language|Greek]] for "ten-legged"). Two other orders of decapodiform cephalopods are also called squid, although they are [[Taxonomy (biology)|taxonomically]] distinct from squids and differ recognizably in their gross anatomical features. They are the [[bobtail squid]] of order [[Sepiolida]] and the [[ram's horn squid]] of the monotypic order [[Spirulida]]. The [[vampire squid]] (''[[Vampyroteuthis]] infernalis''), however, is more closely related to the octopus than to any squid.<ref name="Sanchez Setiamarga 2018">{{cite journal |last1=Sanchez |first1=Gustavo |last2=Setiamarga |first2=Davin H. E. |last3=Tuanapaya |first3=Surangkana |last4=Tongtherm |first4=Kittichai |last5=Winkelmann |first5=Inger E. |last6=Schmidbaur |first6=Hannah |last7=Umino |first7=Tetsuya |last8=Albertin |first8=Caroline |last9=Allcock |first9=Louise |last10=Perales-Raya |first10=Catalina |last11=Gleadall |first11=Ian |last12=Strugnell |first12=Jan M. |last13=Simakov |first13=Oleg |last14=Nabhitabhata |first14=Jaruwat |title=Genus-level phylogeny of cephalopods using molecular markers: current status and problematic areas |journal=[[PeerJ]] |volume=6 |date=February 2018 |doi=10.7717/peerj.4331 |pmid=29456885 |pmc=5813590 |page=e4331 |doi-access=free }}</ref> The [[cladogram]], not fully resolved, is based on Sanchez et al., 2018.<ref name="Sanchez Setiamarga 2018" /> Their [[molecular phylogeny]] used [[mitochondria]]l and [[nuclear DNA]] marker sequences; they comment that a robust [[phylogeny]] "has proven very challenging to obtain". If it is accepted that [[Sepiidae]] cuttlefish are a kind of squid, then the squids, excluding the vampire squid, form a [[clade]] as illustrated.<ref name="Sanchez Setiamarga 2018"/> Orders are shown in boldface; all the families not included in those orders are in the [[paraphyletic]] order "Oegopsida", except Sepiadariidae and Sepiidae that are in the polyphyletic order "Sepiida", {{clade |label1=[[Cephalopoda]] |1={{clade |1=''[[Nautilus]]'' [[File:Nautilus belauensis profile (white background).jpg|50px]] |label2=[[Coleoidea]] |2={{clade |label1=Octopuses and allies |1={{clade |1=[[Octopoda]] [[File:Octopus_vulgaris_Merculiano.jpg|50px]] |2=[[Vampyroteuthidae]] (vampire squid) [[File:Vampyroteuthis_infernalis.jpg|70px]] }} |label2='''Decapodiformes''' |2={{clade |1={{clade |1=[[Cranchiinae]] (glass squid A) [[File:Cranchiidae sp (cropped).jpg|50px]] |2={{clade |1=[[Cycloteuthidae]] |2={{clade |1=[[Psychroteuthidae]] (glacial squid) [[File:Psychroteuthis glacialis paralarva.jpg|70px]] |2={{clade |1={{clade |1={{clade |1={{clade |1={{clade |1=[[Onychoteuthidae]] (hooked squid) [[File:Onychoteuthis banksii2.jpg|70px]] |2=[[Taoniinae]] (glass squid B) [[File:Sandalops melancholicus.jpg|80px]] }} |2={{clade |1={{clade |1=[[Architeuthidae]] (giant squid) [[File:Architeuthis princeps image modified.PNG|80px]] |2={{clade |1=[[Lepidoteuthidae]] (Grimaldi scaled squid) [[File:Lepidoteuthis grimaldii 617 mm ML.jpg|60px]] |2=[[Octopoteuthidae]] (octopus squid) [[File:Taningia danae.gif|80px]] }} }} |2={{clade |1={{clade |1=[[Ancistrocheiridae]] (sharpear enope squid) [[File:Ancistrocheirus lesueurii.jpg|60px]] |2=[[Lycoteuthidae]] (firefly squid) [[File:Lycoteuthis lorigera male.jpg|70px]] }} |2=[[Pyroteuthidae]] (fire squid) [[File:Pyroteuthismargaritifera.jpg|60px]] }} }} |3=[[Bathyteuthidae]] [[File:Bathyteuthisabyssicola (top).jpg|70px]] }} |2={{clade |1={{clade |1={{clade |1=[[Ommastrephidae]] (flying squid) [[File:Todarodes pacificus (white background).jpg|50px]] |2={{clade |1=[[Pholidoteuthidae]] [[File:Pholidoteuthis massyae (cropped).jpg|70px]] |2=[[Gonatidae]] (armhook squid) [[File:Berryteuthis magister.jpg|50px]] }} }} |2=[[Chiroteuthidae]] (whip-lash squid{{efn|common name (whip-lash) is shared with [[Mastigoteuthidae]]}}) [[File:Chiroteuthid.png|70px]] }} |2={{clade |1={{clade |1={{clade |1='''[[Sepiolida]]''' (bobtail squid) [[File:Austrorossia mastigophora (rotated).jpg|60px]] |2=[[Sepiadariidae]] (pyjama and bottletail squid) [[File:Striped pyjama squid (white background).jpg|45px]] }} |2=[[Chtenopterygidae]] [[File:Chtenopteryx sicula 1 (rotated).jpg|60px]] }} |2={{clade |1=[[Thysanoteuthidae]] [[File:Thysanoteuthis rhombus (Merculiano).jpg|55px]] |2=[[Enoploteuthidae]] [[File:Mollusques méditeranéens (Abralia veranyi).jpg|70px]] }} }} }} }} |2={{clade |1=[[Brachioteuthidae]] [[File:Brachioteuthis riisei1.jpg|70px]] |2={{clade |1={{clade |1=[[Neoteuthidae]] |2=[[Histioteuthidae]] (cock-eyed squid) [[File:Histioteuthidae - Histioteuthis bonnellii.JPG|60px]] }} |2={{clade |1=[[Batoteuthidae]] (bush-club squid) |2={{clade |1=[[Mastigoteuthidae]] (whip-lash squid{{efn|common name (whip-lash) is shared with [[Chiroteuthidae]]}}) [[File:Mastigoteuthis agassizii1.jpg|70px]] |2={{clade |1=[[Joubiniteuthidae]] (Joubin's squid) [[File:Joubiniteuthis portieri.jpg|50px]] |2=[[Magnapinnidae]] (bigfin squid) [[File:Magnapinna talismani.jpg|50px]] }} }} }} }} }} }} |2={{clade |1='''[[Spirulida]]''' (ram's horn squid) [[File:Spirula spirula illustration.jpg|30px]] |2={{clade |1={{clade |1='''[[Myopsida]]''' (neritic squid) [[Loliginidae]] [[File:Loligo vulgaris1.jpg|70px]] |2=[[Sepiidae]] (cuttlefish) [[File:Sepia officinalis1.jpg|70px]] }} |2=[[Idiosepiidae]] (pygmy squid) [[File:Tropical Pygmy Squid (cropped).jpg|40px]] }} }} }} }} }} }} }} }} }} }} ===Evolution=== [[Crown group|Crown]] coleoids (the common ancestor of octopuses and squid) diverged in the late [[Paleozoic]] ([[Mississippian (geology)|Mississippian]]), according to fossils of ''[[Syllipsimopodi]]'', an early relative of vampire squids and octopuses.<ref>{{Cite journal |last1=Whalen |first1=Christopher D. |last2=Landman |first2=Neil H. |date=2022-03-08 |title=Fossil coleoid cephalopod from the Mississippian Bear Gulch Lagerstätte sheds light on early vampyropod evolution |journal=Nature Communications |language=en |volume=13 |issue=1 |page=1107 |doi=10.1038/s41467-022-28333-5 |pmid=35260548 |pmc=8904582 |bibcode=2022NatCo..13.1107W |issn=2041-1723}}</ref> True squid diverged during the Jurassic, but many squid families appeared in or after the [[Cretaceous]].<ref name="Tanner Fuchs Winkelmann Gilbert p=20162818">{{cite journal | last1=Tanner | first1=Alastair R. | last2=Fuchs | first2=Dirk | last3=Winkelmann | first3=Inger E. | last4=Gilbert | first4=M. Thomas P. | last5=Pankey | first5=M. Sabrina | last6=Ribeiro | first6=Ângela M. | last7=Kocot | first7=Kevin M. | last8=Halanych | first8=Kenneth M. | last9=Oakley | first9=Todd H. | last10=da Fonseca | first10=Rute R. | last11=Pisani | first11=Davide | last12=Vinther | first12=Jakob | title=Molecular clocks indicate turnover and diversification of modern coleoid cephalopods during the Mesozoic Marine Revolution | journal=Proceedings of the Royal Society B: Biological Sciences | volume=284 | issue=1850 | date=March 2017 | doi=10.1098/rspb.2016.2818 | pmid=28250188 | pmc=5360930 | page=20162818}}</ref> Both the coleoids and the teleost fish were involved in much adaptive radiation at this time, and the two modern groups resemble each other in size, ecology, habitat, morphology and behaviour, however some fish moved into fresh water while the coleoids remained in marine environments.<ref name=Ruppert/> The ancestral coleoid was probably nautiloid-like with a strait septate shell that became immersed in the mantle and was used for buoyancy control. Four lines diverged from this, Spirulida (with one living member), the [[cuttlefish]]es, the squids and the [[octopus]]es. Squid have differentiated from the ancestral [[mollusc]] such that the body plan has been condensed antero-posteriorly and extended dorso-ventrally. What may have been the [[Foot (mollusc)|foot]] of the ancestor is modified into a complex set of appendages around the mouth. The sense organs are highly developed and include advanced eyes similar to those of [[vertebrate]]s.<ref name=Ruppert>{{cite book |title=Invertebrate Zoology |edition=7th |last1=Ruppert |first1=Edward E. |last2=Fox |first2=Richard S. |last3=Barnes |first3=Robert D. |year=2004 |publisher=[[CEngage Learning]] |isbn=978-81-315-0104-7 |pages=343–367 |url=https://books.google.com/books?id=A3opAQAAMAAJ}}</ref> The ancestral shell has been lost, with only an internal [[gladius (cephalopod)|gladius]], or pen, remaining. The pen, made of a chitin-like material,<ref name=Ruppert/><ref name="ifuku">{{cite journal |last=Ifuku |first=S. |title=Chitin and chitosan nanofibers: preparation and chemical modifications |pmid=25393598 |pmc=6271128 |journal=Molecules |year=2014 |volume=19 |issue=11 |pages=18367–80 |doi=10.3390/molecules191118367 |doi-access=free }}</ref> is a feather-shaped internal structure that supports the squid's mantle and serves as a site for muscle attachment. The [[cuttlebone]] or sepion of the Sepiidae is [[calcareous]] and appears to have evolved afresh in the [[Tertiary]].<ref name="Bonnaud 2006">{{cite journal | last1=Bonnaud | first1=Laure | last2=Lu | first2=C. C. | last3=Boucher-Rodoni | first3=Renata | title=Morphological character evolution and molecular trees in sepiids (Mollusca: Cephalopoda): is the cuttlebone a robust phylogenetic marker? | journal=Biological Journal of the Linnean Society | volume=89 | issue=1 | date=2006 | doi=10.1111/j.1095-8312.2006.00664.x | pages=139–150| doi-access=free }}</ref> <gallery mode="packed"> File:Rhomboteuthis lehmani (cropped).jpg|Fossil ''[[Rhomboteuthis]]'' from the [[Callovian|Lower Callovian]] ({{circa| 164 [[Million years ago|Mya]]}}, middle Jurassic) of [[La Voulte-sur-Rhône (lagerstätte)|La Voulte-sur-Rhône]], France File:Plesioteuthis prisca 01.jpg|Fossil ''[[Plesioteuthis]]'' from the [[Tithonian]] ({{circa|150 Mya}}, upper Jurassic), [[Solnhofen limestone|Solnhofen]], Germany </gallery>
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