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Sea butterflies, scientific name Thecosomata (thecosomes, "case / shell-body"), are a taxonomic
suborder Order ( la, ordo) 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 ...
of small
pelagic The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or w ...
swimming sea snails. They are holoplanktonic
opisthobranch Opisthobranchs () is now an informal name for a large and diverse group of specialized complex gastropods which used to be united in the subclass Opisthobranchia. That taxon is no longer considered to represent a monophyletic grouping. Euopis ...
gastropod
mollusk 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 es ...
s. Most Thecosomata have some form of calcified
shell Shell may refer to: Architecture and design * Shell (structure), a thin structure ** Concrete shell, a thin shell of concrete, usually with no interior columns or exterior buttresses ** Thin-shell structure Science Biology * Seashell, a hard o ...
, although it is often very light and / or transparent. The sea butterflies include some of the world's most abundant gastropod species, and because of their large numbers are an essential part of the food chain, and a significant contributor to the
oceanic carbon cycle The oceanic carbon cycle (or marine carbon cycle) is composed of processes that exchange carbon between various pools within the ocean as well as between the atmosphere, Earth interior, and the seafloor. The carbon cycle is a result of many inte ...
. The sea butterflies are included in the
Pteropoda Pteropoda (common name pteropods, from the Greek meaning "wing-foot") are specialized free-swimming pelagic sea snails and sea slugs, marine opisthobranch gastropods. Most live in the top 10 m of the ocean and are less than 1 cm long. The mono ...
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 ...
, and are also included in the informal group
Opisthobranchia Opisthobranchs () is now an informal name for a large and diverse group of specialized complex gastropods which used to be united in the subclass Opisthobranchia. That taxon is no longer considered to represent a monophyletic grouping. Euopisth ...
.


Morphology

Sea butterflies float and swim freely in the water, and are carried along with the currents. This has led to a number of adaptations in their bodies. The shell and 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 ...
have disappeared in several families. Their gastropodal foot has taken the form of two wing-like lobes, or ''
parapodia In invertebrates, the term parapodium ( Gr. ''para'', beyond or beside + ''podia'', feet; plural: parapodia) refers to lateral outgrowths or protrusions from the body. Parapodia are predominantly found in annelids, where they are paired, unjointed ...
'', which propel the animal through the sea by slow flapping movements. left, Unidentified thecosome Most Thecosomata have some form of calcified
shell Shell may refer to: Architecture and design * Shell (structure), a thin structure ** Concrete shell, a thin shell of concrete, usually with no interior columns or exterior buttresses ** Thin-shell structure Science Biology * Seashell, a hard o ...
, although often very light. They are rather difficult to see, since their shell (if present) is mostly transparent, very fragile, and usually tiny: Less than 1 cm in length. Although their shell may be so fine as to be transparent, it is nevertheless
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an a ...
, and an important part of the ocean calcite cycle. Their shells are
bilaterally symmetric Symmetry in biology refers to the symmetry observed in organisms, including plants, animals, fungi, and bacteria. External symmetry can be easily seen by just looking at an organism. For example, take the face of a human being which has a pla ...
and can vary widely in shape: coiled, needle-like, triangular, or globular. The shell is present in all stages of the Cavolinioidea (euthecosomata) life cycle. In the Cymbulioidea (pseudothecosomata), adult Peraclididae also bear shells; the Cymbuliidae shed their larval shells and develop a cartilaginous ''pseudoconch'' in adulthood; only the Desmopteridaen lack any rigid covering as adults.


Behaviour and distribution

Sea butterflies range from the tropics to the poles. They are " holoplanktonic": That is, they spend their whole lives floating among the
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a cr ...
, rather than just being planktonic during their
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. Th ...
l stage. As such, the thecosomata are the most common (in terms of diversity, species richness, and abundance) in the top of the ocean, and become rarer the deeper one samples. Sometimes, they swarm in large numbers and can be found washed up in
flotsam In maritime law, flotsam'','' jetsam'','' lagan'','' and derelict are specific kinds of shipwreck. The words have specific nautical meanings, with legal consequences in the law of admiralty and marine salvage. A shipwreck is defined as the rema ...
, especially along the coast of
eastern Australia The eastern states of Australia are the states adjoining the east continental coastline of Australia. These are the mainland states of Victoria, New South Wales and Queensland, and the island state of Tasmania. The Australian Capital Terr ...
.


Diurnal vertical migration

Thecosomata beat their wing-like parapodia to "fly" through the water. When descending to deeper water, they hold their wings up. They
migrate vertically Diel vertical migration (DVM), also known as diurnal vertical migration, is a pattern of movement used by some organisms, such as copepods, living in the ocean and in lakes. The word ''diel'' comes from the Latin ''dies'' day, and means a 24-ho ...
from day to night, so the community structure changes on a 24 hour cycle; during the day many organisms take refuge at water depths in excess of 100 m.


Feeding

Little is known about the behaviour of sea butterflies, but they are known to have a peculiar way of feeding. They are generally herbivorous, mostly passive
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a cr ...
feeders, just floating along with the currents,
ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
-side up, although some may become active feeders at times. They catch planktonic food by entangling it in a
mucous Mucus ( ) is a slippery aqueous secretion produced by, and covering, mucous membranes. It is typically produced from cells found in mucous glands, although it may also originate from mixed glands, which contain both serous and mucous cells. It ...
web that can be up to 5 cm wide – many times larger than themselves. If disturbed, they abandon the net and flap slowly away. Every day, they migrate vertically in the water column, following their planktonic prey. At night they graze at the ocean surface and return to deeper water in the morning.


Fossil record

This is, geologically speaking, a rather young group, having evolved from the Late
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...
in the
Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configu ...
Era. The group is represented in the fossil record from shells of those groups within the clade that mineralized. These carbonate shells are a major contributor to the
oceanic carbon cycle The oceanic carbon cycle (or marine carbon cycle) is composed of processes that exchange carbon between various pools within the ocean as well as between the atmosphere, Earth interior, and the seafloor. The carbon cycle is a result of many inte ...
, making up as much as 12% of global carbonate flux. However the low stability of their aragonitic shells means that few end up being preserved in sediments as fossils, mostly being deposited in shallow waters of tropical seas.


Importance in the food chain

These creatures, which range from
lentil The lentil (''Lens culinaris'' or ''Lens esculenta'') is an edible legume. It is an annual plant known for its lens-shaped seeds. It is about tall, and the seeds grow in pods, usually with two seeds in each. As a food crop, the largest pro ...
- to orange-sized, are eaten by various marine species, including a wide variety of fish that are, in turn, consumed by
penguin Penguins ( order Sphenisciformes , family Spheniscidae ) are a group of aquatic flightless birds. They live almost exclusively in the Southern Hemisphere: only one species, the Galápagos penguin, is found north of the Equator. Highly adapt ...
s and polar bears. The sea butterflies form the sole food source of their relatives, the
Gymnosomata Sea angels (clade Gymnosomata) are a large group of small free-swimming sea slugs, not to be confused with Cnidarians (Jellyfish and other similar creatures), classified into six different families. They are pelagic opisthobranchs in the clade Gy ...
. They are also consumed by sea birds, whales, and commercially important fish. However, if sea butterflies are consumed in large quantities fish can get "black gut", which makes them unsellable.


Taxonomy

Along with its sister group, the sea angels (
Gymnosomata Sea angels (clade Gymnosomata) are a large group of small free-swimming sea slugs, not to be confused with Cnidarians (Jellyfish and other similar creatures), classified into six different families. They are pelagic opisthobranchs in the clade Gy ...
), the sea butterflies (Thecosomata) are included 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 ...
Pteropoda Pteropoda (common name pteropods, from the Greek meaning "wing-foot") are specialized free-swimming pelagic sea snails and sea slugs, marine opisthobranch gastropods. Most live in the top 10 m of the ocean and are less than 1 cm long. The mono ...
. The validity of the pteropod order is not universally accepted; it fell out of favour for a number of years, but recent molecular evidence suggests that the taxon should be revived. Although most Thecosomata have some form of calcified
shell Shell may refer to: Architecture and design * Shell (structure), a thin structure ** Concrete shell, a thin shell of concrete, usually with no interior columns or exterior buttresses ** Thin-shell structure Science Biology * Seashell, a hard o ...
, mature Gymnosomata have none.


Ponder & Lindberg

Order Thecosomata de Blainville, 1824 * Infraorder
Euthecosomata ''Euthecosomata'' is a taxonomic unit used to classify sea snails. It is a suborder of the order Pteropoda. MolluscaBase eds. (2021). MolluscaBase. Euthecosomata. Accessed through: World Register of Marine Species at: http://www.marinespecies.org ...
** Superfamily
Limacinoidea The superfamily Limacinoidea is a taxonomic group of small floating sea snails, pelagic marine opisthobranch gastropod mollusks.Bouchet, P. (2012). Limacinoidea. Accessed through: World Register of Marine Species at http://www.marinespecies.o ...
*** Family
Limacinidae The Limacinidae are a family of small sea snails, pteropods, pelagic marine gastropod mollusks in the clade Thecosomata (sea butterflies).Bouchet, P. (2012). Limacinidae. Accessed through: World Register of Marine Species at http://www.marine ...
de Blainville, 1823 ** Superfamily
Cavolinioidea The superfamily Cavolinioidea is the most speciose group of sea butterflies. They belong to the suborder Euthecosomata. MolluscaBase eds. (2021). MolluscaBase. Cavolinioidea Gray, 1850 (1815). Accessed through: World Register of Marine Species at ...
*** Family
Cavoliniidae The family Cavoliniidae is a taxonomic group of small floating sea snails, pelagic marine opisthobranch gastropod mollusks.Gofas, S. (2011). Cavoliniidae. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphi ...
H. and A. Adams, 1854 *** Family Clioidae *** Family
Creseidae Creseidae is a family of gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwat ...
*** Family Cuvierinidae *** Family
Praecuvierinidae The Praecuvierinidae are a family of extinct, small, floating sea snails, pelagic marine opisthobranch gastropod mollusks in the superfamily Cavolinioidea. Genera * Genus '' Praecuvierina'' Janssen, 2005 ** '' Praecuvierina lura'' (Hodgki ...
* Infraorder
Pseudothecosomata Cymbulioidea is a taxonomic superfamily of pelagic " sea butterflies", one group of swimming sea snails. They are holoplanktonic opisthobranch gastropod molluscs in the clade Thecosomata. Anatomy Some groups within this superfamily possess a sh ...
** Superfamily Peraclidoidea *** Family Peraclidae Tesch, 1913 ** Superfamily Cymbulioidea *** Family
Cymbuliidae Cymbuliidae is a family of pelagic sea snails or "sea butterflies", marine gastropod mollusks in the superfamily Cymbulioidea. Description Instead of an external calcareous shell, they possess a pseudoconch, consisting of conchioline, a cart ...
Gray Grey (more common in British English) or gray (more common in American English) is an intermediate color between black and white. It is a neutral or achromatic color, meaning literally that it is "without color", because it can be composed ...
, 1840
*** Family
Desmopteridae Desmopteridae is a family of pelagic sea snails or "sea butterflies", marine gastropod mollusks in the superfamily Cymbulioidea. This family has no subfamilies (according to the taxonomy of the Gastropoda by Bouchet & Rocroi, 2005). '' Desmopt ...
Dall Dall may refer to: People * Anders Bendssøn Dall (died 1607), Danish Lutheran bishop * Bobby Dall (born 1963), American musician * Caroline Healey Dall (1822–1912), American feminist writer *Clarrie Dall (1887–1953), Australian footballer * C ...
, 1921


Bouchet & Rocroi

In the new taxonomy of Bouchet & Rocroi (2005) Thecosomata is treated differently : Clade Thecosomata : *Superfamily Cavolinioidea Gray, 1850 ( = Euthecosomata) **Family Cavoliniidae Gray, 1850 (1815) ***Subfamily Cavoliinae Gray, 1850 (1815) (formerly Hyalaeidae Rafinesque, 1815 ) ***Subfamily Clioinae Jeffreys, 1869 (formerly Cleodoridae Gray, 1840 - ''nomen oblitum'') ***Subfamily Cuvierininae van der Spoel, 1967 (formerly : Cuvieriidae Gray, 1840 (nom. inv.); Tripteridae Gray, 1850 ) ***Subfamily Creseinae Curry, 1982 **Family Limacinidae Gray, 1840 (formerly : Spirialidae Chenu, 1859 ; Spiratellidae Dall, 1921 ) ** † Family Sphaerocinidae A. Janssen & Maxwell, 1995 *Superfamily Cymbulioidea Gray, 1840 ( = Pseudothecosomata) **Family Cymbuliidae Gray, 1840 ***Subfamily Cymbuliinae Gray, 1840 ***Subfamily Glebinae van der Spoel, 1976 **Family Desmopteridae Chun, 1889 **Family Peraclidae Tesch, 1913 (formerly Procymbuliidae Tesch, 1913 Bouchet & Rocroi (2005) move the family Limacinidae into the superfamily Cavolinioidea, making redundant the superfamily Limacinoidea erected for it in Ponder & Lindberg's taxonomy. The families Creseidae and Cuvierinidae are demoted to subfamilies of Cavoliniidae (Creseinae and Cuvierininae). The infraorder Pseudothecosomata becomes the superfamily Cymbulioidea. The family Peraclididae is included in the superfamily Cymbulioidea as the family Peraclidae, making the superfamily Peraclidoidea redundant.


See also

*
Ocean acidification Ocean acidification is the reduction in the pH value of the Earth’s ocean. Between 1751 and 2021, the average pH value of the ocean surface has decreased from approximately 8.25 to 8.14. The root cause of ocean acidification is carbon dioxi ...
* ''
Clione antarctica ''Clione antarctica'' is a species of "sea angel", a sea slug, a pelagic marine gastropod mollusk in the family Clionidae, the "sea angels". Distribution The distribution of ''Clione antarctica'' is within the Southern Hemisphere, in the ...
''


Footnotes


References


Sources

* * * * * {{taxonbar, from=Q772011 Euopisthobranchia Extant Thanetian first appearances