Monoplacophora

{{Short description|Class of molluscs}} {{Automatic taxobox | image = Neopilina.jpg | image_caption = The holotype of ''Neopilina galatheae'' at the Zoological Museum, Copenhagen | fossil_range = {{fossil range|538.8|0|Early Cambrian – Recent|refs=<ref name="ReferenceA"/><ref name="Lindberg2009"/>}} | taxon = Monoplacophora | authority = Odhner, 1940 | subdivision_ranks = | subdivision = }} thumb|Shell of Monoplacophora '''Monoplacophora''' {{IPAc-en|ˌ|m|ɒ|n|oʊ|p|l|ə|ˈ|k|ɒ|f|ər|ə}}, meaning "bearing one plate", is a paraphyletic class of molluscs with a cap-like shell, inhabiting deep sea environments. Extant representatives were not recognized as such until 1952; previously they were known only from the fossil record, and were thought to have become extinct 375 million years ago.<ref name="Lindberg2009" />

Although the shell of many monoplacophorans is limpet-like in shape, they are not gastropods, nor do they have any close relation to gastropods.<ref name="ReferenceA">{{cite journal |author1=Taviani, M. |author2=Sabelli, B. |author3=Candini, F. |year=1990 |title=A fossil Cenozoic monoplacophoran |journal=Lethaia |volume=23 |issue=2 |pages=213–216|doi=10.1111/j.1502-3931.1990.tb01361.x |bibcode=1990Letha..23..213T }}</ref>

== Definition == Discussion about monoplacophorans is made difficult by the slippery definition of the taxon; some authors take it to refer to all non-gastropod molluscs with a single shell, or all single-shelled molluscs with serially repeated units; whereas other workers restrict the definition to cap-shaped forms, excluding spiral and other shapes of shell.<ref name="Lindberg2009">{{cite journal |last1=Lindberg |first1=D.R. |year=2009 |title=Monoplacophorans and the origin and relationships of mollusks |journal=Evolution: Education and Outreach |volume=2 |issue=2 |pages=191–203 |doi=10.1007/s12052-009-0125-4 |s2cid=26108547 |url=https://cloudfront.escholarship.org/dist/prd/content/qt342063m8/qt342063m8.pdf|doi-access=free }}</ref> The inclusion of the gastropod-like Bellerophontoidea within the group is also contentious.<ref name="Gubanov2001">{{cite journal |last1=Gubanov |first1=A.P. |last2=Peel |first2=J.S. |year=2001 |title=Latest helcionelloid molluscs from the Lower Ordovician of Kazakhstan |journal=Palaeontology |volume=44 |issue=4 |pages=681–694 |doi=10.1111/1475-4983.00198|bibcode=2001Palgy..44..681G |doi-access=free }}</ref>

One attempt to resolve this confusion was to separate out the predominantly coiled helcionelloids from the traditional, cap-like tergomyans, this latter group containing extant Tryblidiids.<ref name="Gubanov2001" />

==Taxonomy== {{Hatnote|For a taxonomy of living monoplacophoran species, see Tryblidiida.}} Taxonomy of Monoplacophora per Bouchet, et al. (2017):<ref>Bouchet, Philippe & Jean-Pierre Rocroi, et al. (2017) Revised Classification, Nomenclator and Typification of Gastropod and Monoplacophoran Families. ''Malacologia'', 2017, 61(1–2): 1–526. {{doi|10.4002/040.061.0201}}</ref>

Class '''Monoplacophora''' *† Subclass Cyrtolitiones **† Order Sinuitopsida ***† Superfamily Cyrtolitoidea S. A. Miller, 1889 ****† Family Cyrtolitidae S. A. Miller, 1889 ****† Family Carcassonnellidae Horný, 1997 ***† Superfamily Cyclocyrtonelloidea Horný, 1962 ****† Family Cyclocyrtonellidae Horný, 1962 (= Yochelsoniidae Horný, 1962 (inv.)) ****† Family Multifariitidae Bjaly, 1973 ****† Family Sinuellidae Starobogatov & Moskalev, 1987 ****† Family Sinuitinidae Starobogatov & Moskalev, 1987 *† Subclass Cyrtonelliones **† Order Cyrtonellida ***† Superfamily Cyrtonelloidea Knight & Yochelson, 1958 ****† Family Cyrtonellidae Knight & Yochelson, 1958 (= Cyrtonellopsinae Horný, 1965) *† Subclass Eomonoplacophora **(Unassigned to Order) ***† Superfamily Maikhanelloidea Missarzhevsky, 1989 ****† Family Maikhanellidae Missarzhevsky, 1989 (= Purellidae Vassiljeva, 1990) *Subclass Tergomya (= Pilinea) **† Order Kirengellida (= Romaniellida) ***† Superfamily Archaeophialoidea Knight & Yochelson, 1958 ****† Family Archaeophialidae Knight & Yochelson, 1958 ****† Family Peelipilinidae Horný, 2006 ****† Family Pygmaeoconidae Horný, 2006 ***† Superfamily Kirengelloidea Starobogatov, 1970 ****† Family Kirengellidae Starobogatov, 1970 ****† Family Romaniellidae Rozov, 1975 ****† Family Nyuellidae Starobogatov & Moskalev, 1987 ***† Superfamily Hypseloconoidea Knight, 1952 ****† Family Hypseloconidae Knight, 1952 **Order Tryblidiida ***Superfamily Tryblidioidea Pilsbry, 1899 ****† Family Tryblidiidae Pilsbry, 1899 ****† Family Proplinidae Knight & Yochelson, 1958 ****† Family Drahomiridae Knight & Yochelson, 1958 ****† Family Bipulvinidae Starobogatov, 1970 ***Superfamily Neopilinoidea Knight & Yochelson, 1958 ****Family Neopilinidae Knight & Yochelson, 1958 *****Subfamily Neopilininae Knight & Yochelson, 1958 (= Vemidae Moskalev, Starobogatov & Filatova, 1983; = Laevipilinidae Moskalev, Starobogatov & Filatova, 1983; = Monoplacophoridae Moskalev, Starobogatov & Filatova, 1983) *****Subfamily Veleropilininae Starobogatov & Moskalev, 1987 (= Rokopellidae Starobogatov & Moskalev, 1987; = Micropilinidae Haszprunar & Schaefer, 1997)

==Anatomy and physiology== [[File:Monoplacophora.svg|thumb|left|upright=1.8|Internal anatomy of ''Micropilina''. The head region is on the left by the mouth.]] Monoplacophorans are univalved (though not gastropodal), limpet-shaped, and are untorted. They have a pseudometamerism of bilaterally symmetrical repeated organs and muscles. The extant members of the class live only in the deep ocean (the abyssal zone, the continental shelf, and the continental slope) at depths below {{convert|180|m|ft}}. Cambrian forms predominately lived in shallow seas, whereas later Paleozoic forms are more commonly found in deeper waters with soft, muddy sea floors.<ref name="Lindberg2009"/>

Although superficially resembling limpets when viewed dorsally, monoplacophorans are not anatomically similar to gastropods. Some similarities are shared with the chitons, such as having segmented anatomy (organs arranged in series). There are eight pairs of dorso-ventral muscles (shell muscles). The nervous system is relatively simple, with no true ganglion present.

The repeated organs include from three to six pairs of "gills" (actually ctenidia) located in a curved line along each side of the foot (though the number is not always considered definitive of a given species), and as many as six "kidneys" (actually nephridia). The tip or point of their low shells points forward rather than towards the back. The shell ranges from 3 mm to 37 mm in diameter depending on species. Like in chitons, the head is poorly defined, and there are no eyes. The mouth is located within the animal's undeveloped head in front of its single large foot and contains a radula, a defining characteristic of the mollusca. Tentacles are situated behind the mouth. They also have a cone-shaped stomach with a single crystalline style though no gastric shield. The intestines are long and make between four and six loops before reaching the posteriorly-positioned anus. Monoplacophorans also have oesophageal pouches.

The sexes are separate with any given animal having two pairs of either ovaries or testes connected to either the third or fourth pair of kidneys. One genus, ''Micropilina'', has apparently been recorded as brooding young in the distal oviduct and pallial groove, releasing the young when approximately 300 micrometers in diameter.<ref>{{cite book |last1=Counts |first1=Clement L. |year=2006 |chapter=Chapter 17: Monoplacophora |editor1-last=Sturm |editor1-first=C.F. |editor2-last=Pearce |editor2-first=T.A. |editor3-last=Valdes |editor3-first=Angel |title=The Mollusks: A guide to their study, collection, and preservation |pages=212–214 |publisher=Universal |isbn=9781581129304 |chapter-url=https://books.google.com/books?id=-NbmHx93s8gC&q=%22monoplacophora%22&pg=PA212 |via=Google Books}}</ref>

== Phylogenetic position == In 2006 a molecular study on ''Laevipilina antarctica'' suggested that extant Monoplacophora and Polyplacophora form a well-supported clade with the researched ''Neopilina'' closest to the chitons.<ref name="Giribet2006">{{cite journal |last1=Giribet |first1=G. |last2=Okusu |first2=A. |last3=Lindgren |first3=A.R. |last4=Huff |first4=S.W. |last5=Schrödl |first5=M. |last6=Nishiguchi |first6=M.K. |title=Evidence for a clade composed of molluscs with serially repeated structures: Monoplacophorans are related to chitons |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=103 |issue=20 |pages=7723–7728 |date=May 2006 |pmid=16675549 |pmc=1472512 |doi=10.1073/pnas.0602578103 |bibcode=2006PNAS..103.7723G|doi-access=free }}</ref> The two classes in this new clade, with the proposed name Serialia, all show a variable number of serially repeated gills and eight sets of dorsoventral pedal retractor muscles.

This study contradicts the fossil evidence, which suggests that the Monoplacophora are the sister group to the remainder of the conchiferans,<ref>{{cite journal |author=Scheltema, A.H. |title=''Aplacophora'' as progenetic ''Aculiferans'' and the ''Coelomate'' origin of mollusks as the sister taxon of ''Sipuncula'' |journal=The Biological Bulletin |volume=184 |issue=1 |pages=57–78 |date=1 February 1993 |doi=10.2307/1542380 |pmid=29300613 |jstor=1542380 |url=http://www.biolbull.org/cgi/content/abstract/184/1/57?ijkey=d45e9547a760e1cb9921f81243d05ae05773204c&keytype2=tf_ipsecsh |access-date=12 November 2007 |archive-url=https://web.archive.org/web/20110616214155/http://www.biolbull.org/cgi/content/abstract/184/1/57?ijkey=d45e9547a760e1cb9921f81243d05ae05773204c&keytype2=tf_ipsecsh |archive-date=16 June 2011 |url-status=dead }}</ref><ref>{{cite journal |author=Haszprunar, G. |year=2000 |title=[no title cited] |journal=Am. Malacol. Bull |volume=15 |pages=115–130}}</ref><ref>{{cite book |author1=Salvini-Plawen, L.V. |author2=Steiner, G. |year=1996 |title=Origin and Evolutionary Radiation of the Mollusca |editor=Taylor, J.D. |publisher=Oxford University Press |place=Oxford, UK |pages=29–51}}</ref> and that the cephalopods (squids, octopuses, and relatives) arose from within the monoplacophoran lineage.<ref name=mollusca12-2>{{cite book |chapter=Main features of cephalopod evolution |series=The Mollusca |volume=12 |title=Palaeontology and Neontology of Cephalopods |editor1=Clarke, M.R. |editor2=Trueman, E.R. |isbn=0-12-751412-0 |year=1988 |publisher=Academic Press |location=Orlando, FL}}</ref> However, some authors dispute this view and do not necessarily see modern Monoplacophora as related to their presumed fossil ancestors.<ref name=Budd2000>{{cite journal |last1=Budd |first1=G.E. |last2=Jensen |first2=S. |year=2000 |title=A critical re-appraisal of the fossil record of the bilaterian phyla |volume=75 |issue=2 |pages=253–95 |journal=Biological Reviews of the Cambridge Philosophical Society |doi=10.1111/j.1469-185X.1999.tb00046.x |pmid=10881389|s2cid=39772232 }}</ref>

The concept of Serialia is supported by other molecular studies.<ref>{{Cite journal |last1=Wilson |first1=N. |last2=Rouse |first2=G. |last3=Giribet |first3=G. |year=2010 |title=Assessing the molluscan hypothesis Serialia (Monoplacophora+Polyplacophora) using novel molecular data. |journal=Molecular Phylogenetics & Evolution |volume=54 |issue=1 |pages=187–193 |doi=10.1016/j.ympev.2009.07.028 |pmid=19647088}}</ref>

The fossil record does indicate that the ancestral mollusc was monoplacophoran-like and that the Polyplacophora arose from within the Monoplacophora – not the other way around.<ref name=Runnegar1974>{{cite journal |first1=B. |last1=Runnegar |first2=J. Jr. |last2=Pojeta |date=October 1974 |title=Molluscan phylogeny: The paleontological viewpoint |journal=Science |volume=186 |issue=4161 |pages=311–317 |jstor=1739764 |pmid=17839855 |doi=10.1126/science.186.4161.311 |bibcode=1974Sci...186..311R|s2cid=46429653 }}</ref> This could be reconciled if a secondary loss of shells caused a monoplacophoran body form to re-appear secondarily, which is plausible: At the very least, modern monoplacophorans are not closely related to vent-dwelling representatives from the Silurian.<ref>{{cite journal |last1=Little |first1=C. |last2=Vrijenhoek |first2=R.C. |title=Are hydrothermal vent animals living fossils? |journal=Trends in Ecology & Evolution |volume=18 |issue=11 |pages=582–588 |year=2003 |doi=10.1016/j.tree.2003.08.009}}</ref>

Cambrian monoplacophoran ''Knightoconus antarcticus'' is hypothesised to be an ancestor to the cephalopods.

== Fossil species == {{Expand section|date=August 2009}} Living families: * Tryblidiida ** Laevipilinidae ** Micropilinidae ** Monoplacophoridae ** Neopilinidae

Extinct families: * Tryblidiida ** † Tryblidiidae von Zittel, 1899 * † Palaeacmaeidae (uncertain, as the Cambrian type species is a cnidarian. It is maintained here as a receptacle for the Paleozoic genus ''Parmophorella'') ** † ''Palaeacmaea'' Hall & Whitfield, 1872 ** † ''Parmophorella'' Matthew, 1886 ** † ''Knightoconus'' *** † ''Knightoconus antarcticus'' Many Cambrian-Devonian species have been described as "monoplacophorans", but the only fossil members of the crown group date to the Pleistocene.<ref name="ReferenceA"/>

The Taxonomy of the Gastropoda (Bouchet & Rocroi, 2005){{refn|name="Bouchet"|{{cite book |editor1=Bouchet, P. |editor2=Rocroi, J.-P. |year=2005b |title=''Classification and nomenclator of gastropod families'' |title-link=Taxonomy of the Gastropoda (Bouchet & Rocroi, 2005) |publisher=ConchBooks |place=Hackenheim, Germany |isbn=3-925919-72-4}} {{cite journal |title=(online data ref) |journal = Malacologia: International Journal of Malacology|year = 2005|issue = 1–2|url=https://www.vliz.be/Vmdcdata/imis2/ref.php?refid=78278|last1 = Bouchet|first1 = P.|last2 = Rocroi|first2 = J. -P}} reprinted from Bouchet & Rocroi (2005a)<ref name=BouchetRocroi2005/>}}<ref name=BouchetRocroi2005>{{cite journal |author1=Frýda, J. |author2=Hausdorf, B. |author3=Ponder, W. |author4=Valdés, Á. |author5=Warén, A. |editor1=Bouchet, P. |editor2=Rocroi, J.-P. |title=Classification and nomenclator of gastropod families |year=2005a |journal=Malacologia: International Journal of Malacology |volume=47 |issue=1–2 |publisher=Institute of Malacology |place=Ann Arbor |issn=0076-2997 |id=e- |url=https://www.biodiversitylibrary.org/item/81069#page/5/mode/1up}}</ref> also contains Paleozoic molluscs of uncertain systematic position. It is not known whether these were gastropods or monoplacophorans.

== References == {{Reflist|25em}}

* {{cite journal |author=Lemche, Henning |year=1957 |title=A new living deep-sea mollusc of the Cambro-Devonian class Monoplacophora |journal=Nature |volume=179 |issue=4556 |pages=413–416 |doi=10.1038/179413a0 |bibcode=1957Natur.179..413L |s2cid=4173823 |place=London, UK}} * {{cite book |author=Lemche, Henning |editor=Jenkins, Marie |year=1972 |title=The Curious Mollusks |place=New York, NY}}

== Further reading == * {{cite journal |author1=Horný, Radvan |year=1963 |title=On the systematic position of cyrtonelloids (Mollusca) |journal=Časopsis národního Muzea, oddil přírodovědný |volume=132 |issue=2 |pages=90–94 |place=Prague, CZ}} * {{cite journal |author=Rozov, S.N. |year=1975 |title=A new order of the Monoplacophora |journal=Paleontological Journal |volume=9 |pages=39–43 |place=Washington, DC}} * {{cite journal |author1=Schrödl, Michael |author2=Linse, Katrin |author3=Schwabe, Enrico |name-list-style=amp |title=Review on the distribution and biology of Antarctic Monoplacophora, with first abyssal record of Laevipilina antarctica |journal=Polar Biology |date=August 2006 |volume=29 |issue=9 |pages=721–727 |doi=10.1007/s00300-006-0132-7|bibcode=2006PoBio..29..721S |s2cid=23753587 }}

== External links == {{Commons category|Monoplacophora}} * {{cite web |url=http://bio.classes.ucsc.edu/bio136/molluscs/monoplacophora/monoplacophora.html |archive-date=2007-11-20 |df=dmy-all |archive-url=https://web.archive.org/web/20071120214523/http://bio.classes.ucsc.edu/bio136/molluscs/monoplacophora/monoplacophora.html |title=monoplacophora |series=Bio 136: Anatomy |publisher=University of California – Santa Cruz (UCSC)}} * {{cite web |url=https://www.discoverlife.org/mp/20q?guide=Mollusca_Monoplacophora_species |website=DiscoverLife |title=Taxonomy of Monoplacophora}} * {{cite web |url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Monoplacophora.html |publisher=University of Michigan |department=Animal Diversity Web |title=Monoplacophora}} * {{cite web |url=http://depts.washington.edu/natmap/mollusks/6monofile.html |archive-date=2007-07-15 |df=dmy-all |archive-url=https://web.archive.org/web/20070715220601/http://depts.washington.edu/natmap/mollusks/6monofile.html |website=Nature Mapping Program |publisher=University of Washington |place=Seattle, WA |title=Monoplacophora}} * {{cite web |url=http://palaeos.com/metazoa/mollusca/tryblidiida/tryblidiida.html |website=Palaeos.com |title=''Tryblidiida (Monoplacophora)''}} Includes pictures and thorough discussion of cladistic issues. * {{cite web |url=http://www.zmuc.dk/inverweb/Galathea/Pdf_filer/Volume_03/Plates_41-56-galathea-vol.03-pp_009-072.pdf |archive-date=2011-06-16 |df=dmy-all |archive-url=https://web.archive.org/web/20110616155103/http://www.zmuc.dk/inverweb/Galathea/Pdf_filer/Volume_03/Plates_41-56-galathea-vol.03-pp_009-072.pdf |title=images of Monoplacophora}}

Category:Monoplacophora Category:Extant Cambrian first appearances Category:Conchifera Category:Taxa named by Nils Hjalmar Odhner