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Tryblidiida is a taxon of
monoplacophora Monoplacophora , meaning "bearing one plate", is a polyphyletic superclass 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 ...
n
mollusca Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
ns containing the only extant representatives: 37 species are still alive today, inhabiting the ocean at depths of between .


History of discoveries

The first captured living monoplacophoran was '' Veleropilina zografi'' in 1896, but at that time it was described as if it were an archaeogastropod, a
true limpet The Patellogastropoda, common name true limpets and historically called the Docoglossa, are members of a major phylogenetic group of marine gastropods, treated by experts either as a clade or as a taxonomic order. The clade Patellogastropoda ...
, mainly because of its patelliform (limpet-like) shell. This species was finally revealed to be monoplacophoran 87 years later, in 1983. In April 1952, a living specimen was collected from deep depths in the
Middle America Trench The Middle America Trench is a major subduction zone, an oceanic trench in the eastern Pacific Ocean off the southwestern coast of Middle America, stretching from central Mexico to Costa Rica. The trench is 1,700 miles (2,750 km) long an ...
off
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
's Pacific coast. In 1957 that species was described and named ''
Neopilina galatheae ''Neopilina galatheae'' is a species of monoplacophoran, a superficially limpet-like marine mollusc. Its name means new '' Pilina'' (after ''Pilina'', an extinct monoplacophoran species that lived about 400 million years ago). It lives in depths ...
'' by its discoverer, Danish biologist Henning Mourier Lemche (1904–1977). An expert in the field has called this discovery "one of the greatest sensations in the wentiethcentury."Schwabe, above, at p. 205. As of 2008, there were 31 living species known, discovered in waters from 200 meters in depth to
hadal The hadal zone, also known as the hadopelagic zone, is the deepest region of the ocean, lying within oceanic trenches. The hadal zone ranges from around below sea level, and exists in long, narrow, topographic V-shaped depressions. The cumula ...
depths, or more than 6,000 meters in the deepest ocean trenches. The first specimen photographed alive was '' Vema hyalina'', at a depth of 400 meters off
Catalina Island, California Santa Catalina Island ( xgf, Pimuu'nga or ; es, Isla Santa Catalina) is a rocky island off the coast of Southern California in the Gulf of Santa Catalina. The island name is often shortened to Catalina Island or just Catalina. The island is ...
, in 1977. Scientists believe that the taxon Monoplacophora is probably
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...
and have proposed including all the living members in the
order 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 d ...
Tryblidiida. In 1989, fossils in Italy from the middle
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
were described which appear to be identical with the living species '' Micropilina minuta''.


Anatomy

Little is known about monoplacophorans. They have a single, flat, rounded bilateral shell that is often thin and fragile; it ranges in size from 3 to 30 millimetres (in recent species). The
apex The apex is the highest point of something. The word may also refer to: Arts and media Fictional entities * Apex (comics), a teenaged super villainess in the Marvel Universe * Ape-X, a super-intelligent ape in the Squadron Supreme universe *Apex ...
of the shell is at the anterior end. The fossil shells exhibit a series of muscular attachment scars on the inner side, suggesting metamerism; indeed, with living Monoplacophora to study, it can be seen that their body segments exhibit a serial repetition of kidneys, gills and reproductive structure. This used to be interpreted as a true segmentation, which suggested a " missing link" between
mollusks Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
and
annelids The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecolo ...
. More recent studies have shown that the repetition of these organs is secondary. Monoplacophorans move on a rounded foot. Their reduced head lacks eyes or tentacles. The mantle cavity forms a horseshoe-shaped groove running around the muscular foot, in a similar fashion to that of the
chiton Chitons () are marine molluscs of varying size in the class Polyplacophora (), formerly known as Amphineura. About 940 extant and 430 fossil species are recognized. They are also sometimes known as gumboots or sea cradles or coat-of-mail s ...
s, and contains five or six gills on either side. The mouth opens on the underside between the ends of the groove, while the
anus The anus (Latin, 'ring' or 'circle') is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, the residual semi-solid waste that remains after food digestion, which, d ...
opens into the hindmost part. Like chitons, monoplacophorans possess a sensory subradular organ, as well as a rasping
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
. A fold of ciliated tissue surrounds the mouth to the front and sides, while a smaller fold, bearing a number of tentacles, lies just behind it. The stomach contains a
style Style is a manner of doing or presenting things and may refer to: * Architectural style, the features that make a building or structure historically identifiable * Design, the process of creating something * Fashion, a prevailing mode of clothing ...
, projecting from a diverticulum, or "style sac". The mouth has a chevron-shaped lip in front of it, and bears tentacles behind it, which have various shapes and layouts in different species. The heart is divided into two equal halves, each with its own auricle, ventricle and
aorta The aorta ( ) is the main and largest artery in the human body, originating from the left ventricle of the heart and extending down to the abdomen, where it splits into two smaller arteries (the common iliac arteries). The aorta distributes ...
. The left and right aorta fuse shortly after leaving the heart, and supply blood to the
open circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
. There are six pairs of
nephridia The nephridium (plural ''nephridia'') is an invertebrate organ, found in pairs and performing a function similar to the vertebrate kidneys (which originated from the chordate nephridia). Nephridia remove metabolic wastes from an animal's body. Neph ...
l excretory organs, which empty into the mantle cavity. The nervous system has small
ganglia A ganglion is a group of neuron cell bodies in the peripheral nervous system. In the somatic nervous system this includes dorsal root ganglia and trigeminal ganglia among a few others. In the autonomic nervous system there are both sympatheti ...
around the
oesophagus The esophagus (American English) or oesophagus (British English; both ), non-technically known also as the food pipe or gullet, is an organ in vertebrates through which food passes, aided by peristaltic contractions, from the pharynx to the ...
from which two pairs of main nerve cords run through the body; one pair supplying the foot, and the other the visceral organs. As in the chitons, these main nerve cords are connected by a series of lateral nerves, giving the layout of the nervous system an appearance somewhat like a ladder. There are two pairs of
gonad A gonad, sex gland, or reproductive gland is a mixed gland that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, produces sper ...
s, which release
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
s into the water through one of the pairs of nephridia. The sexes are separate, and fertilisation is
external External may refer to: * External (mathematics), a concept in abstract algebra * Externality In economics, an externality or external cost is an indirect cost or benefit to an uninvolved third party that arises as an effect of another party' ...
.


Ecology


Habitat

Monoplacophora are a geographically widespread component of the
benthos Benthos (), also known as benthon, is the community of organisms that live on, in, or near the bottom of a sea, river, lake, or stream, also known as the benthic zone.Laevipilinidae ''Laevipilina'' is a genus of monoplacophoran Monoplacophora , meaning "bearing one plate", is a polyphyletic superclass of molluscs with a cap-like shell inhabiting deep sea environments . Extant representatives were not recognized as such un ...
** Genus ''
Laevipilina ''Laevipilina'' is a genus of monoplacophoran molluscs. They are very small, mostly deepwater animals which have a superficially limpet-like shell.Schwabe, Enrico (2008)A summary of reports of abyssal and hadal Monoplacophora and Polyplacophora ...
'' J. H. McLean, 1979 *** ''
Laevipilina antarctica ''Laevipilina antarctica'' is a species of monoplacophoran, a superficially limpet-like marine mollusk. It is found in the Weddell Sea and the Lazarev Sea of Antarctica.Schwabe, Enrico (2008)A summary of reports of abyssal and hadal Monoplacoph ...
'' Warén & Hain, 1992 *** ''
Laevipilina cachuchensis ''Laevipilina cachuchensis'' is a species of monoplacophoran, a superficially limpet-like marine mollusk. Two live specimens were collected off the coast of Asturias Asturias (, ; ast, Asturies ), officially the Principality of Asturias ...
'' Urgorri, García-Alvarez & Luque, 2005 *** ''
Laevipilina hyalina ''Laevipilina hyalina'' is a species of monoplacophoran, a superficially limpet-like marine mollusk. It is found off the coast of southern California Southern California (commonly shortened to SoCal) is a geographic and cultural region that ...
'' J. H. McLean, 1979 *** ''
Laevipilina rolani ''Laevipilina rolani'' is a species of monoplacophoran, a superficially limpet-like marine mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollu ...
'' Warén & Bouchet, 1990 *** ''
Laevipilina theresae ''Laevipilina theresae'' is a species of monoplacophoran, a superficially limpet-like marine mollusk. It is known from a live specimen collected in the Eastern Weddell Sea of Antarctica Antarctica () is Earth's southernmost and least-p ...
'' Schrödl, 2006 * Family Micropilinidae ** Genus '' Micropilina'' Warén, 1989 *** '' Micropilina arntzi'' Warén & Hain, 1992 *** '' Micropilina minuta'' Warén, 1989 *** '' Micropilina rakiura'' Marshall, 1998 *** '' Micropilina reingi'' Marshall, 2006 *** '' Micropilina tangaroa'' Marshall, 1992 *** '' Micropilina wareni'' Marshall, 2006 * Family Monoplacophoridae ** Genus '' Monoplacophorus'' Moskalev, Starobogatov & Filatova, 1983 *** '' Monoplacophorus zenkevitchi'' Moskalev, Starobogatov & Filatova, 1983 * Family
Neopilinidae Tryblidiida is a taxon of monoplacophoran molluscans containing the only extant representatives: 37 species are still alive today, inhabiting the ocean at depths of between . History of discoveries The first captured living monoplacophoran wa ...
** Genus '' Adenopilina'' Starobogatov & Moskalev, 1987 *** '' Adenopilina adenensis'' (Tebble, 1967) ** Genus ''
Neopilina ''Neopilina'' is a highly derived genus of modern monoplacophoran.Organisms, Genes and Evolution: Evolutionary Theory at the Crossroads ; Proceedings of the 7th International Senckenberg Conference. By Dieter Stefan Peters, Michael Weingarten. Co ...
'' H. Lemche, 1957 *** ''
Neopilina bruuni ''Neopilina bruuni'' is a species of monoplacophoran, a superficially limpet-like marine mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks () ...
'' Menzies, 1968 *** ''
Neopilina galatheae ''Neopilina galatheae'' is a species of monoplacophoran, a superficially limpet-like marine mollusc. Its name means new '' Pilina'' (after ''Pilina'', an extinct monoplacophoran species that lived about 400 million years ago). It lives in depths ...
'' Lemche, 1957 *** ''
Neopilina rebainsi ''Neopilina rebainsi'' is a species of monoplacophoran, a superficially limpet-like marine mollusc. The holotype was collected southeast of the Falkland Islands in the Southern Atlantic Ocean.Schwabe, Enrico (2008)A summary of reports of abyssal ...
'' Moskalev, Starobogatov & Filatova, 1983 *** ''
Neopilina starobogatovi ''Neopilina starobogatovi'' is a species of monoplacophoran, a superficially limpet-like marine mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or moll ...
'' Ivanov & Moskalev, 2007 ** Genus '' Rokopella'' Starobogatov & Moskalev, 1987 *** '' Rokopella brummeri'' Goud & Gittenberger, 1993 *** '' Rokopella capulus'' Marshall, 2006 *** '' Rokopella euglypta'' (Dautzenberg & Fischer, 1897) *** '' Rokopella goesi'' (Warén, 1988) *** '' Rokopella oligotropha'' (Rokop, 1972) *** '' Rokopella segonzaci'' Warén & Bouchet, 2001 ** Genus '' Veleropilina'' Starobogatov & Moskalev, 1987 *** '' Veleropilina brummeri'' (Goud & Gittenberger, 1993) *** '' Veleropilina capulus'' (B. A. Marshall, 2006) *** '' Veleropilina euglypta'' (Dautzenberg & H. Fischer, 1897) *** '' Veleropilina goesi'' (Warén, 1988) *** '' Veleropilina oligotropha'' (Rokop, 1972) *** '' Veleropilina reticulata'' (Seguenza, 1876) *** '' Veleropilina segonzaci'' (Warén & Bouchet, 2001) *** '' Veleropilina seisuimaruae'' Kano, S. Kimura, T. Kimura & Warén, 2012 *** '' Veleropilina veleronis'' (Menzies & Layton, 1963) *** '' Veleropilina zografi'' (Dautzenberg & H. Fischer, 1896) ** Genus '' Vema'' (Clarke & Menzies, 1959) *** '' Vema bacescui'' (Menzies, 1968) *** '' Vema ewingi'' (Clarke & Menzies, 1959) *** '' Vema levinae'' Warén, 1996 *** '' Vema occidua'' B. A. Marshall, 2006


References

{{Taxonbar, from=Q874793 Monoplacophora Mollusc orders