Pteropoda
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Pteropoda (
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often con ...
pteropods, from the
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
meaning "wing-foot") are specialized free-swimming
pelagic The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...
sea snail Sea snails are slow-moving marine (ocean), marine gastropod Mollusca, molluscs, usually with visible external shells, such as whelk or abalone. They share the Taxonomic classification, taxonomic class Gastropoda with slugs, which are distinguishe ...
s and
sea slug Sea slug is a common name for some Marine biology, marine invertebrates with varying levels of resemblance to terrestrial Slug, slugs. Most creatures known as sea slugs are gastropods, i.e. they are Sea snail, sea snails (marine gastropod moll ...
s, marine
opisthobranch Opisthobranchs () is a now 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. Euopistho ...
gastropods Gastropods (), commonly known as slugs and snails, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and from the land. Ther ...
. Most live in the top 10 m of the ocean and are less than 1 cm long. The
monophyly In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent comm ...
of Pteropoda is the subject of a lengthy debate; they have even been considered as
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
with respect to
cephalopods A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
. Current consensus, guided by molecular studies, leans towards interpreting the group as
monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...
. Pteropoda encompasses the two
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
s
Thecosomata The Thecosomata (collective/plural: ''thecosomes'', meaning "case/shell-body"), or sea butterflies, are a taxonomic suborder of small, pelagic, free-swimming sea snails known as holoplanktonic opisthobranch gastropod mollusks, in the order Pt ...
, the
sea butterflies The Thecosomata (collective/plural: ''thecosomes'', meaning "case/shell-body"), or sea butterflies, are a taxonomic suborder of small, pelagic, free-swimming sea snails known as holoplanktonic opisthobranch gastropod mollusks, in the order ...
, and
Gymnosomata Sea angels (clade Gymnosomata) are a large group of small free-swimming sea slugs, classified into six separate families. They are pelagic opisthobranchs in the clade Gymnosomata within the larger mollusc clade Heterobranchia. Sea angels were pre ...
, the sea angels. The Thecosomata ( "case-body") have a shell, while the Gymnosomata ("naked body") do not. The two clades may or may not be
sister taxa In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
; if not, their similarity (in that they are both pelagic, small, and transparent, and both groups swim using wing-like flaps (
parapodia In invertebrates, the term parapodium ( Gr. ''para'', beyond or beside + ''podia'', feet; : parapodia) refers to lateral outgrowths or protrusions from the body. Parapodia are predominantly found in annelids, where they are paired, unjointed late ...
) which protrude from their bodies) may reflect convergent
adaptation 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 their particular lifestyle.


Taxonomy

The group Pteropoda was established by
Cuvier Jean Léopold Nicolas Frédéric, baron Cuvier (23 August 1769 – 13 May 1832), known as Georges Cuvier (; ), was a French natural history, naturalist and zoology, zoologist, sometimes referred to as the "founding father of paleontology". Cuv ...
as "'" in 1804. François Péron and
Charles Alexandre Lesueur Charles Alexandre Lesueur (; 1 January 1778 in Le Havre – 12 December 1846 in Le Havre) was a French Natural history, naturalist, artist, and explorer. He was a prolific natural-history collector, gathering many type specimens in Australia ...
thought the group to be larger, and so they also included the
opisthobranch Opisthobranchs () is a now 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. Euopistho ...
taxa ('' Phyllirhoë'' and ''
Glaucus In Greek mythology, Glaucus (; ) was a Greek prophetic sea-god, born mortal and turned immortal upon eating a magical herb. It was believed that he came to the rescue of sailors and fishermen in storms, having earlier earned a living from the ...
''), the heteropoda taxa ('' Carinaria'' and '' Firola''), and even the
Ctenophora Ctenophora (; : ctenophore ) is a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and they are ...
('' Callianira''). In 1810, these authors divided the whole group in two separate groups: Those with a shell and those without a shell. In 1824, H.M.D. de Blainville named these two groups Gymnosomata and Thecosomata and named the combining
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 ...
Aporobranchia instead of Pteropoda. He rejected the additional
genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...
, except ''Phyllirhoë'' which he upgraded to a third group that he called Psilosomata. Only much later was Phyllirhoë classified within the order Nudibranchia. Other attempts were made to describe the Pteropoda: J.E. Gray divided the Pteropoda into Dactylobranchia (with just the genus '' Cavolinia'') and Pterobranchia (including all the other genera).
Cuvier Jean Léopold Nicolas Frédéric, baron Cuvier (23 August 1769 – 13 May 1832), known as Georges Cuvier (; ), was a French natural history, naturalist and zoology, zoologist, sometimes referred to as the "founding father of paleontology". Cuv ...
(and his followers) did not accept the classification by de Blainville; they preferred the original classification as described in '.
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(1829) followed the Cuvierian classification but tried to include the character of having a distinct head or not. The German naturalist L. Oken went one step further and, for the sake of symmetry, wanted each order to contain four families and each family to contain four genera. P.A. 1829, divided the Pteropoda according to the size of their fins: "'" (including only Pneumonoderma) and "'" (including all the others). W.B. Clark (1829) treated the Pteropoda as a family and emended the spelling to Pteropodidae (a name now re-used for a family of fruit bats) Finally, all these attempts were abandoned and, as more and more species were described as a result of several scientific expeditions, the classification of the Pteropoda into Thecosomata and Gymnosomata was generally adopted. The relationship between these two clades is not unequivocally established, but it seems that they are sister taxa.


Evolutionary history

Pteropods are estimated to have originated during the Early Cretaceous, around 133 million years ago, with the diversification into the major lineages occurring during the mid-late Cretaceous. The oldest known fossil pteropod is a member of Limacinidae from the early-middle
Campanian The Campanian is the fifth of six ages of the Late Cretaceous epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campa ...
deposits of the
San Juan Islands The San Juan Islands is an archipelago in the Pacific Northwest of the United States between the U.S. state of Washington and Vancouver Island, British Columbia, Canada. The San Juan Islands are part of Washington state, and form the core of ...
.


Phylogeny


Threats


Vulnerability to ocean acidification

A study was conducted on the West Coast of the United States to see
ocean acidification Ocean acidification is the ongoing decrease in the pH of the Earth's ocean. Between 1950 and 2020, the average pH of the ocean surface fell from approximately 8.15 to 8.05. Carbon dioxide emissions from human activities are the primary cause of ...
's effects on pteropods. '' Limacina helicina'' was used to test the sensitivity to decreasing pH. This species of pteropod is potentially vulnerable to the corrosive waters associated with ocean acidification due to their
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 ...
shell. The shell of a pteropod was immersed in ocean water with the projected pH level that the water will reach by the year 2100. After a month and a half in the water, the shell had almost completely dissolved.


Distribution

Pteropods are found in all major oceans, usually below the ocean surface and in all levels of latitude. Pteropods can be found lower than 10 meters, but in less amounts in terms of biomass, however, pteropod distribution is more spread out deeper based on findings. This can be explained as Pteropods from tropical areas become more common in deeper areas. They are not found commonly in the deep sea, in fact, few live lower than 500 meters below sea level.
Continental shelves A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an island ...
, areas containing many opportunities for nutrients, and productivity are locations in which Pteropods are usually populous, according to patterns in data. Springtime is a peak season for Pteropoda, as they reach higher populations, though data shows that Pteropoda south of the
equator The equator is the circle of latitude that divides Earth into the Northern Hemisphere, Northern and Southern Hemisphere, Southern Hemispheres of Earth, hemispheres. It is an imaginary line located at 0 degrees latitude, about in circumferen ...
are less abundant seasonally. In addition, current data suggests that 93% of the world's pteropods are part of the
Thecosomata The Thecosomata (collective/plural: ''thecosomes'', meaning "case/shell-body"), or sea butterflies, are a taxonomic suborder of small, pelagic, free-swimming sea snails known as holoplanktonic opisthobranch gastropod mollusks, in the order Pt ...
suborder, while the 7% are
Gymnosomata Sea angels (clade Gymnosomata) are a large group of small free-swimming sea slugs, classified into six separate families. They are pelagic opisthobranchs in the clade Gymnosomata within the larger mollusc clade Heterobranchia. Sea angels were pre ...
.


Footnotes


References


External links

* — by German planktologist
Carl Chun Carl Chun or Karl Friedrich Gustav Chun (1 October 1852 – 11 April 1914) was a German marine biologist who worked as a professor at the Universities of Königsberg (1883), Breslau (1891) and Leipzig (1898). He was a pioneer of German oceanogr ...
; beautifully illustrated; free download * — archived documentary website with film clips & photos * * * {{Authority control Euopisthobranchia Obsolete gastropod taxa