Limacina helicina

Last updated

Limacina helicina
LimacinaHelicinaNOAA.jpg
A live individual of Limacina helicina
Limacina helicina 7.jpg
A live individual of Limacina helicina
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Clade: Euopisthobranchia
Order: Pteropoda
Family: Limacinidae
Genus: Limacina
Species:
L. helicina
Binomial name
Limacina helicina
(Phipps, 1774) [1]
Synonyms [2]
  • Argonauta arctica O. Fabricius, 1780 [3]
  • Clio helicinaPhipps, 1774 (original combination)
  • Limacina helicialis Lamarck, 1819 [4]
  • Spiratella helicina(Phipps, 1774)
  • Spiratella limacina Blainville, 1824 [5]
  • Limacina helicina var. typica Meisenheimer, 1906 [6]
  • Spiratella planospiraRehder, 1942

Limacina helicina is a species of small swimming planktonic sea snail, in the family Limacinidae. It belongs to the group commonly known as sea butterflies (Thecosomata). [7] [8]

Contents

Limacina helicina is a keystone species of mesozooplankton in Arctic pelagic ecosystems. [9] [10] [11]

The first written record of this species was by Friderich Martens from Spitsbergen in 1675. [12] [13] Limacina helicina was also observed during a 1773 expedition to the Arctic led by Constantine John Phipps on the ships HMS Racehorse and on HMS Carcass and the species was described one year later, in 1774. [1]

Limacina helicina is the type species of the genus Limacina. [14]

In contrast to the traditional view, it was shown in 2010 that the distribution of this species is not bipolar; Arctic and Antarctic individuals belong to two genetically distinct species: Limacina helicina in the Arctic, and Limacina antarctica in the Antarctic. [11] [15]

Subspecies

Limacina helicina has been recognised as a species complex comprising two sub-species and at least five forms. [15] In addition, the taxonomic category “forma” has been applied to designate at least three morphotypes of Limacina helicina helicina (acuta, helicina and pacifica) and two morphotypes of Limacina helicina antarctica (antarctica and rangi). [15] It is also known as Limacina helicina rangii (d'Orbigny, 1835). [16] These forms typically have different geographical ranges, but it remains unclear as to whether forms represent morphological responses to different environmental conditions or are indeed taxonomically distinct, and if the latter, their level of taxonomic separation. [15]

However, at the species level the geographical distribution is considered to be bipolar, as it occurs in both the Arctic and Antarctic oceans. [15] Remigio and Hebert (2003) [19] provided initial evidence for the genetic separation of Limacina helicina helicina and Limacina helicina antarctica. [15] Hunt et al. (2010) [15] have quantified genetic distance within these taxa. Hunt 2010 found a 33.56% difference in cytochrome c oxidase subunit I (COI) gene sequences between the "Limacina helicina" which were collected from the Arctic and the Antarctic oceans. [15] This degree of separation is sufficient for ordinal level taxonomic separation in other organisms, and provides strong evidence for the Arctic and Antarctic populations of Limacina helicina differing at least at the species level. [15] Subspecies Limacina helicina antarctica Woodward, 1854 can be considered as a separate species Limacina antarctica Woodward, 1854. [16] A conservative divergence time estimate of 31 Ma (95% HPD interval 12–53 Ma) for Arctic and Antarctic taxa, indicates that they have undergone rapid independent evolution since the establishment of cold water provinces in the early Oligocene. [15] Also there is different structure of the shell between Limacina helicina and Limacina antarctica. [9]

Distribution

The type locality of Limacina helicina is "Arctic seas". [1] Limacina helicina is the only thecosome pteropod in Arctic waters. [11]

The distribution of Limacina helicina is arctic and subarctic (subpolar–polar) [20] especially in the Arctic Ocean and countries include:

Description

Drawing of part of Limacina helicina from ventral view shows parapodia, anterior lobe of the parapodia, lower part of the foot and foldings on the ventral side of the foot. Limacina helicina 5.png
Drawing of part of Limacina helicina from ventral view shows parapodia, anterior lobe of the parapodia, lower part of the foot and foldings on the ventral side of the foot.
Detailed drawing of the same ventral view. Limacina helicina 6.png
Detailed drawing of the same ventral view.

Limacina helicina has wing-like parapodia which evolved from the original gastropod foot (as is the case in all other pteropods). [11]

In this species, the color of the soft parts is dark purple or violet, with paler pellucid (translucent) parapodia. [24]

Dorsal view. Limacina helicina 2 color.png
Dorsal view.
Left lateral view. Limacina helicina 3.png
Left lateral view.
Frontal view. Limacina helicina 4.png
Frontal view.

The shell is sinistral, subglobose, subdiscoidal, [25] hyaline and very thin. The spire is depressed [25] [26] but it can be considered rather high in comparison of other Limacina species. [21] The shell has 5-6 transversally striated whorls. [21] [26] The suture is distinct. [21] The last whorl is large and with very obscure keel next to its umbilicus. [21] [25] The shell has a wide umbilicus. [21] The aperture is higher than it is wide. [21]

The width of the shell is 5–10 mm [10] or up to 13 mm. [13] The height of the shell is up to 6 mm (when maximum width was 8 mm). [21]

Apical view. Limacina helicina shell 2.png
Apical view.
Apertural view. Limacina helicina shell.png
Apertural view.
Umbilical view. Limacina helicina shell 3.png
Umbilical view.
Drawing of operculum of Limacina helicina. Limacina helicina operculum.png
Drawing of operculum of Limacina helicina.

Adult specimens in the genus Limacina have usually lost the operculum. [21]

The radula consist of 10 rows. [24] Each row consist of one central tooth and two lateral teeth. [24] The Digestive system also includes an esophagus, gizzard sac and gut. [13]

Drawing of the whole radula. Magnified 60x. Limacina helicina radula.png
Drawing of the whole radula. Magnified 60×.
Detail of radula showing 3 rows of teeth. Magnified 190x. Limacina helicina radula 2.png
Detail of radula showing 3 rows of teeth. Magnified 190×.

Ecology

Habitat

Live Limacina helicina. The white rectangle shows the shell opening, the area where Comeau et al. (2009) studied the effect of ocean acidification. Limacina helicina.png
Live Limacina helicina. The white rectangle shows the shell opening, the area where Comeau et al. (2009) studied the effect of ocean acidification.

Pteropods are strict pelagic mollusks that are highly adapted to life in the open ocean. [11] They are actively swimming in the water. Limacina helicina is a holoplanktonic species. Habitat of Limacina helicina is upper epipelagic and glacial. [16] It lives in temperatures from -0.4 °C to +4.0 °C or rarely up to 7 °C. [27]

Vertical distribution is affected by the size and also by other factors. [28] Limacina helicina of the size from 0.2 to 0.4 mm lives mainly in depths from 0 m to 50 m. [28] Larger pteropods lives from 0 m to 150 m. [28] For example, Gilmer & Harbison (1991) [13] have found larger specimen of Limacina helicina to occur mainly in depths 5–25 m with abundance up to 2.5 adults in m3. [13] They do not occur much in upper 4 m probably because of turbulence. [13]

Already Constantine John Phipps mentioned its "innumerable quantities" in arctic seas in 1774. [1] Limacina helicina is a major component of the polar zooplankton. [15] It can comprise >50% of total zooplankton abundance (number of individuals per unit volume). [15]

Species of the clade Thecosomata produce a fragile external calcium carbonate shell, which could serve as a ballast enabling large vertical migrations and as a protection against predators. [11] The aragonitic composition of the shell makes it very sensitive to dissolution. [11] Aragonite is a metastable form of calcium carbonate and it is more soluble in seawater than calcite. [10] Because of its highly soluble [11] aragonite shell and polar distribution, Limacina helicina may be one of the first organisms affected by ocean acidification, and it is therefore a key indicator species of this process. [15] As a key indicator of the acidification process, and a major component of polar ecosystems, Limacina helicina has become a focus for acidification research. [15] Based on laboratory experiments, they are able to precipitate calcium carbonate at low aragonite saturation state. [11] Limacina helicina seems to be relatively more resilient to elevated concentration of carbon dioxide (CO2) than other aragonitic organisms such as corals. [11] Laboratory experiments results support the current concern for the future of Arctic pteropods, as the production of their shell appears to be very sensitive to decreased pH. [10] [11] A decline of pteropod populations would likely cause dramatic changes to various pelagic ecosystems. [10] [11] However, in response to acute abiotic stress, Limacina helicina shows high transcriptomic plasticity, [29] suggesting this species may have a limited capacity to acclimate to the effects of ocean acidification and ocean warming. Shelled pteropods also play a geochemical role in carbon cycle in the oceans, as they contribute to the export of calcium carbonate and can represent a major component of the carbon transport to the deep ocean. [10] [11] [15]

Researchers found 24-53% individuals of Limacina helicina with shells damaged by dissolution off the U.S. West Coast in 2011. [30]

Feeding habits

They produce large mucus webs to filter-feed on phytoplankton but also small zooplankton. [11] They eat the web with the captured prey and then re-reproduce a web net. [26] The web is large and spherical [31] and it is difficult to see during the day because of diffuse reflection. [13] Webs are easier to see at night. [13] Limacina helicina is easily disturbed (like all other Thecosomata); when disturbed, it retracts into its shell and destroys its web. [13] Gilmer & Harbison (1991) [13] have assumed, that Limacina helicina feeds while motionless (without actively swimming). [13] Its web enables them neutral buoyancy or allows them slow sinking only. [31]

Limacina helicina plays a significant ecological role as a phytoplankton grazer. [15] Limacina helicina is an obligate ciliary feeder. [28] Gilmer & Harbison (1991) [13] hypothesized that Limacina helicina are "web trappers", who are also chemically attracting their motile prey. [13]

Major parts of the food of Limacina helicina include tintinnid (Tintinnida), small crustaceans - copepods (Copepoda) and juvenile specimen of its own species (cannibalism). [13] Danish zoologist Johan Erik Vesti Boas reported diatoms (Bacillariophyceae), dinoflagellates (Dinoflagellata) and tintinnids in the digestive system of Limacina helicina in 1888 already. [32] Diatoms and dinoflagellates appear to pass the digestive system of adults largely intact. [8] [13] Fecal pellets of Limacina helicina contains small cells, dinoflagellates and diatoms as main largely intact food items and also few small fragments of tintinnids, Limacina and copepods. [13] All experiments performed on Limacina helicina in the laboratory were done on starved specimens, because they do not feed in unnatural conditions. [13] Ocean acidification and ocean warming may cause Limacina helicina to increase its metabolic rate and use more storage compounds, especially during the Arctic winter when food is limited. This may result in a decline in reproductive success and survival. [33]

Gilmer & Harbison (1991) [13] also suggested that smaller specimens may be herbivores feeding preferentially on phytoplankton and protozoans and that larger specimens became omnivores. [13]

Life cycle

Limacina helicina is a protandric hermaphrodite. [8] [28] Males are smaller, at sizes of 4–5 mm [8] and then they change to females, which are larger than 5 mm. [8] Sperm is transferred by spermatophores during copulation. [34] They lay eggs in ribbon-like clusters [8] mainly in summer, but also a little in winter. [28]

The size of the veliger larvae is about 0.15 mm. [26] When animals reached 0.7 mm in size, gonads have been detected in them. [28] Fully mature individuals are 0.8 mm in size. [28]

The life cycle of Limacina helicina lasts about 1 year [8] or 1.5–2 years. [28]

Predators

Limacina helicina plays an important role in the marine food web as a major dietary component for predators such as large zooplankton, herring Clupea sp., [22] chum salmon Oncorhynchus keta, [22] pink salmon Oncorhynchus gorbuscha, [22] rorquals, [22] [27] Phoca hispida [27] and other seals [27] and birds. [11]

The pteropod Clione limacina feeds almost entirely on the genus Limacina: on Limacina helicina and on Limacina retroversa . [34] [35] [36] Also the pteropod Paedoclione doliiformis feeds on those two species only, but solely on juveniles with shells smaller than 1 mm. [34]

Locomotion

A snapshot of the flows and forces at work in Torkel Weis-Fogh's clap and fling mechanism for insect flight. Movements of Limacina's wing-like parapodia create vortices in the water flow, which generate lift. Clap and Fling 6- fling 3.svg
A snapshot of the flows and forces at work in Torkel Weis-Fogh's clap and fling mechanism for insect flight. Movements of Limacina's wing-like parapodia create vortices in the water flow, which generate lift.

Limacina helicina possesses a pair of flexible, wing-like appendages called parapodia which it beats in a complex 3D stroke pattern which resembles the wing kinematics of flying insects. By doing so, the animal effectively flies through the water. The sea butterfly uses a high angle of attack of approximately 45-50 degrees to generate lift, and it beats its wings 4 to 10 times per second. It propels itself using a version of the clap and fling mechanism described by Torkel Weis-Fogh in small insects such as thrips. [37] [38] Another aspect of locomotion by Limacina helicina is the extreme, forward-back pitching (called hyper-pitching) which it experiences during each half-stroke of its wings. The animal rocks forward and backward by up to 60 degrees during each half-stroke. No other species is known to experience such extreme hyper-pitching during normal locomotion. [38] [39]

See also

Related Research Articles

<span class="mw-page-title-main">Sea butterfly</span> Suborder of molluscs

The Thecosomata, or sea butterflies, are a taxonomic suborder of small, pelagic, free-swimming sea snails known as holoplanktonic opisthobranch gastropod mollusks, in the order Pteropoda. Most pteropods have some form of calcified shell, although it is often very light, even translucent.

<span class="mw-page-title-main">Sea angel</span> Clade of gastropods

Sea angels are a large group of small free-swimming sea slugs, not to be confused with Cnidarians, classified into six different families. They are pelagic opisthobranchs in the clade Gymnosomata within the larger mollusc clade Heterobranchia. Sea angels were previously referred to as a type of pteropod.

<i>Limacina</i> Genus of gastropods

Limacina is a genus of swimming predatory sea snails commonly known as sea butterflies in the family Limacinidae. This genus contains some of the world's most abundant gastropod species.

<i>Clione</i> Genus of gastropods

Clione is a genus of small, floating sea slugs, pelagic marine gastropod mollusks in the family Clionidae, the sea angels.

Paedoclione doliiformis is a species of sea angel, a small floating sea slug, a pelagic marine gastropod mollusk in the family Clionidae.

<span class="mw-page-title-main">Pteropoda</span> Order of molluscs

Pteropoda 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 monophyly of Pteropoda is the subject of a lengthy debate; they have even been considered as paraphyletic with respect to cephalopods. Current consensus, guided by molecular studies, leans towards interpreting the group as monophyletic.

<i>Clione limacina</i> Species of gastropod

Clione limacina, known as the naked sea butterfly, sea angel, and common clione, is a sea angel found from the surface to greater than 500 m (1,600 ft) depth. It lives in the Arctic Ocean and cold regions of the North Atlantic Ocean. It was first described by Friderich Martens in 1676 and became the first gymnosomatous "pteropod" to be described.

<i>Diplulmaris antarctica</i> Species of jellyfish

Diplulmaris antarctica is a species of Antarctic jellyfish in the family Ulmaridae.

<i>Limacina rangii</i> Species of gastropod

Limacina rangii is a species of swimming sea snail in the family Limacinidae, which belong to the group commonly known as sea butterflies (Thecosomata).

Clione antarctica is a species of "sea angel", a sea slug, a pelagic marine gastropod mollusk in the family Clionidae, the "sea angels".

Limacina retroversa is a distinct species of swimming planktonic gastropods, belonging to a group of predatory sea snails known as sea butterflies (Thecosomata). The name Limacina retroversa describes the unique morphology of this sea snail, including its slug-like body and coiled, backwards-turning shell. They are typically found in the epipelagic zone of cold, polar waters, but can be found worldwide, in any ocean. L. retroversa are currently under threat, as their numbers are decreasing due to rising global carbon levels and other human-caused climate threats.

<i>Clio</i> (gastropod) Genus of gastropods

The genus Clio is a taxonomic group of small floating sea snails, pelagic marine opisthobranch gastropod mollusks, the sole genus belonging to the family Cliidae

<span class="mw-page-title-main">Wildlife of Antarctica</span> Antarctic wildlife

The wildlife of Antarctica are extremophiles, having adapted to the dryness, low temperatures, and high exposure common in Antarctica. The extreme weather of the interior contrasts to the relatively mild conditions on the Antarctic Peninsula and the subantarctic islands, which have warmer temperatures and more liquid water. Much of the ocean around the mainland is covered by sea ice. The oceans themselves are a more stable environment for life, both in the water column and on the seabed.

<span class="mw-page-title-main">Ocean acidification in the Arctic Ocean</span>

The Arctic Ocean covers an area of 14,056,000 square kilometers, and supports a diverse and important socioeconomic food web of organisms, despite its average water temperature being 32 degrees Fahrenheit. Over the last three decades, the Arctic Ocean has experienced drastic changes due to climate change. One of the changes is in the acidity levels of the ocean, which have been consistently increasing at twice the rate of the Pacific and Atlantic oceans. Arctic Ocean acidification is a result of feedback from climate system mechanisms, and is having negative impacts on Arctic Ocean ecosystems and the organisms that live within them.

<span class="mw-page-title-main">Human impact on marine life</span>

Human activities affect marine life and marine habitats through overfishing, habitat loss, the introduction of invasive species, ocean pollution, ocean acidification and ocean warming. These impact marine ecosystems and food webs and may result in consequences as yet unrecognised for the biodiversity and continuation of marine life forms.

Cavolinia inflexa is a species of small pteropod.

Clione okhotensis is a species of sea angel, a pelagic marine gastropod in the family Clionidae.

Clione elegantissima is a species of sea angel in the family Clionidae.

Callianira antarctica is a species of ctenophore that physically resembles Mertensia ovum, but lacks the oil sacs. Just like other ctenophores, over 95% of its body mass and composition is water.

References

This article incorporates CC-BY-2.5 text from the reference [11] [15] and public domain text from the reference [25]

  1. 1 2 3 4 Phipps C. J. (1774). A Voyage towards the North Pole undertaken by His Majesty's Command 1773. J. Nourse, London, viii + 253 pp. Page 195.
  2. ↑ "Limacina helicina (Phipps, 1774)". CLEMAM, accessed 29 January 2011.
  3. ↑ (in Latin) Fabricius O. (1780). Fauna Groenlandica systematice sistens animalis Groenlandiae occidentalis hactenus indagata. Hafniae et Lipsiae. J. G. Rothe pp. XVI + 452 + 1 pl. Page 386-388.
  4. ↑ (in French) Lamarck J.-B. (1819). Histoire naturelle des animaux sans vertèbres. Paris. Volume 6. page 291.
  5. ↑ (in French) de Blainville H. M. D. (1824). In: Cuvier F. (ed.). (1804-1845). Dictionnaire des Sciences Naturelles dans lequel on traite méthodiquement des différens êtres de la nature. Levrault, Strasbourg & Paris, and Le Normant, Paris, 32: page 284.
  6. 1 2 (in German) Meisenheimer J. (1906). Die Pteropoden der deutschen Sud-polar Expedition 1901-1903. In: Deutsche Sudpolar-Expedition 1901-1903. 9 (Zool.), 1(2): 92-152, pl. 5-7. page 96-98. Plate 5, figure 1 a-b, 3.
  7. ↑ Bouchet P. & Gofas S. (2011). Limacina helicina (Phipps, 1774). Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=140223 on 2011-02-07
  8. 1 2 3 4 5 6 7 Gannefors, C.; Böer, M.; Kattner, G.; Graeve, M.; Eiane, K.; Gulliksen, B. R.; Hop, H.; Falk-Petersen, S. (2005). "The Arctic sea butterfly Limacina helicina: Lipids and life strategy". Marine Biology. 147 (1): 169–177. Bibcode:2005MarBi.147..169G. doi:10.1007/s00227-004-1544-y. hdl: 11250/174238 . S2CID   55386624.
  9. 1 2 Sato-Okoshi, W.; Okoshi, K.; Sasaki, H.; Akiha, F. (2010). "Shell structure of two polar pelagic molluscs, Arctic Limacina helicina and Antarctic Limacina helicina antarctica forma antarctica". Polar Biology. 33 (11): 1577–1583. Bibcode:2010PoBio..33.1577S. doi:10.1007/s00300-010-0849-1. S2CID   24975779.
  10. 1 2 3 4 5 6 7 8 Comeau, S.; Gorsky, G.; Jeffree, R.; Teyssié, J. -L.; Gattuso, J. -P. (2009). "Impact of ocean acidification on a key Arctic pelagic mollusc (Limacina helicina)". Biogeosciences. 6 (9): 1877–1882. Bibcode:2009BGeo....6.1877C. doi: 10.5194/bg-6-1877-2009 . hdl: 10453/14721 .
  11. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Comeau, S.; Jeffree, R.; Teyssié, J. L.; Gattuso, J. P. (2010). Stepanova, Anna (ed.). "Response of the Arctic Pteropod Limacina helicina to Projected Future Environmental Conditions". PLOS ONE. 5 (6): e11362. Bibcode:2010PLoSO...511362C. doi: 10.1371/journal.pone.0011362 . PMC   2894046 . PMID   20613868.
  12. ↑ (in German) Martens F. (1675). Spitzbergische oder Groenlandische Reise Beschreibung gethan im Jahr 1671. Schultzen, Hamburg. page 110
  13. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Gilmer, R. W.; Harbison, G. R. (1991). "Diet of Limacina helicina (Gastropoda: Thecosomata) in Arctic waters in midsummer" (PDF). Marine Ecology Progress Series. 77: 125–134. Bibcode:1991MEPS...77..125G. doi: 10.3354/meps077125 .
  14. ↑ Janssen A. W. (2007). "Holoplanktonic Mollusca (Gastropoda: Pterotracheoidea, Janthinoidea, Thecosomata and Gymnosomata) from the Pliocene of Pangasinan (Luzon, Philippines)". Scripta Geologica 137 http://www.scriptageologica.nl/07/nr135/a02 Archived 2011-07-24 at the Wayback Machine
  15. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Hunt, B.; Strugnell, J.; Bednarsek, N.; Linse, K.; Nelson, R. J.; Pakhomov, E.; Seibel, B.; Steinke, D.; Würzberg, L. (2010). Finkel, Zoe (ed.). "Poles Apart: The "Bipolar" Pteropod Species Limacina helicina is Genetically Distinct Between the Arctic and Antarctic Oceans". PLOS ONE. 5 (3): e9835. Bibcode:2010PLoSO...5.9835H. doi: 10.1371/journal.pone.0009835 . PMC   2847597 . PMID   20360985.
  16. 1 2 3 4 5 6 7 Bouchet, P.; Gofas, S. (2011). Limacina helicina (Phipps, 1774). Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=140223 on 2011-01-29
  17. ↑ Spoel S. van der (1967). "Euthecosomata, a group with remarkable developmental stages (Gastropoda, Pteropoda)". J. Noorduyn and Zn, Gorinchem, 375 pp.
  18. ↑ Dall W. H. (1871). "Descriptions of sixty new forms of mollusks from the west coast of North America and the north Pacific Ocean, with notes on others already described". American Journal of Conchology 7(2): 93-160, plates 13-16. Pages 138-139.
  19. ↑ Remigio, E.; Hebert, P. D. (2003). "Testing the utility of partial COI sequences for phylogenetic estimates of gastropod relationships". Molecular Phylogenetics and Evolution. 29 (3): 641–647. doi:10.1016/S1055-7903(03)00140-4. PMID   14615199.
  20. ↑ Orr, J. C.; Fabry, V. J.; Aumont, O.; Bopp, L.; Doney, S. C.; Feely, R. A.; Gnanadesikan, A.; Gruber, N.; Ishida, A.; Joos, F.; Key, R. M.; Lindsay, K.; Maier-Reimer, E.; Matear, R.; Monfray, P.; Mouchet, A.; Najjar, R. G.; Plattner, G. K.; Rodgers, K. B.; Sabine, C. L.; Sarmiento, J. L.; Schlitzer, R.; Slater, R. D.; Totterdell, I. J.; Weirig, M. F.; Yamanaka, Y.; Yool, A. (2005). "Anthropogenic ocean acidification over the twenty-first century and its impact on calcifying organisms" (PDF). Nature. 437 (7059): 681–686. Bibcode:2005Natur.437..681O. doi:10.1038/nature04095. PMID   16193043. S2CID   4306199. PDF Archived 2008-06-25 at the Wayback Machine . (author's draft).
  21. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 Spoel S. van der (1972). "Pteropoda Thecosomata". Zooplankton, Conseil International Pour L’Exploration de la Mer, sheet 140-142.
  22. 1 2 3 4 5 Wing B. (not dated). "Auke Bay Laboratory (ABL). Ocean Carrying Capacity Program. Unusual Observations of Fish and Invertebrates From the Gulf of Alaska, 2004-05". accessed 29 January 2011.
  23. ↑ Hermans C. O. & Satterlie R. A. (1992). "Fast-Strike Feeding Behaviour in a Pteropod Mollusk, Clione limacina Phipps". The Biological Bulletin, Marine Biological Laboratory, 182: 1-7.
  24. 1 2 3 (in Latin and Norwegian) Sars G. O. (1878). Bidrag til kundskaben om norges arktiske fauna. I. Mollusca regonis arcticae Norwegiae. Oversigt over de i norges arktiske region forekommende bløddyr. Christiania, Brøgger. pages 328-329, Plate 29, figure 1. Plate 16, figure 17.
  25. 1 2 3 4 Tryon G. W. (1873). American marine conchology: or, Descriptions of the shells of the Atlantic coast of the United States from Maine to Florida. Philadelphia. page 121-122. plate 14, figure 253.
  26. 1 2 3 4 Hopcroft R. (created 31 January 2009). "Sea Butterfly: Limacina helicina (Phipps, 1774)". Arctic Ocean biodiversity, accessed 30 January 2011.
  27. 1 2 3 4 "Limacina helicina helicina helicina". Marine Species Identification Portal, accessed 8 February 2011.
  28. 1 2 3 4 5 6 7 8 9 Kobayashi, H. A. (1974). "Growth cycle and related vertical distribution of the thecosomatous pteropod Spiratella (?Limacina?) helicina in the central Arctic Ocean". Marine Biology. 26 (4): 295–301. Bibcode:1974MarBi..26..295K. doi:10.1007/BF00391513. S2CID   84668586.
  29. ↑ Johnson, K. M.; Hofmann, G. E. (2020). "Combined stress of ocean acidification and warming influence survival and drives differential gene expression patterns in the Antarctic pteropod". Conservation Physiology. 8 (1): coaa013. doi:10.1093/conphys/coaa013. PMC   7098371 . PMID   32257214.
  30. ↑ Bednaršek, N.; Feely, R. A.; Reum, J. C. P.; Peterson, B.; Menkel, J.; Alin, S. R.; Hales, B. (2014). "Limacina helicina shell dissolution as an indicator of declining habitat suitability owing to ocean acidification in the California Current Ecosystem". Proceedings of the Royal Society B . 281 (1785): 1785. doi:10.1098/rspb.2014.0123. PMC   4024287 . PMID   24789895.
  31. 1 2 Gilmer, R. W.; Harbison, G. R. (1986). "Morphology and field behavior of pteropod molluscs: Feeding methods in the families Cavoliniidae, Limacinidae and Peraclididae (Gastropoda: Thecosomata)". Marine Biology. 91 (1): 47–57. Bibcode:1986MarBi..91...47G. doi:10.1007/BF00397570. S2CID   84378241.
  32. ↑ (in Danish) Boas J. E. V. (1888). "Spolia Atlantica. Bidrag til Pteropodernes. Morfologi og Systematik samt til Kundskaben om deres geografiski Udbredelse". Det Kongelige Danske videnskabernes selskabs skrifter. København, serie 6, number 4: 1-231. Page 33, 40-43. Plate 5, figure 69-70.
  33. ↑ Lischka, Silke; Riebesell, Ulf (2017). [Lischka, S., Riebesell, U., 2016. Metabolic response of Arctic pteropods to ocean acidification and warming during the polar night/twilight phase in Kongsfjord (Spitsbergen). Polar Biology.. doi:10.1007/s00300-016-2044-5 "Metabolic response of Arctic pteropods to ocean acidification and warming during the polar night/twilight phase in Kongsfjord (Spitsbergen)"]. Polar Biology. 40 (6): 1211–1227. Bibcode:2017PoBio..40.1211L. doi:10.1007/s00300-016-2044-5. S2CID   25305403.{{cite journal}}: Check |url= value (help)
  34. 1 2 3 Lalli C. M. & Gilmer R. W. (1989). Pelagic Snails. The biology of holoplanktonic gastropod molluscs. Stanford University Press: Stanford, California. page 185, 188.
  35. ↑ Böer, M.; Gannefors, C.; Kattner, G.; Graeve, M.; Hop, H.; Falk-Petersen, S. (2005). "The Arctic pteropod Clione limacina: Seasonal lipid dynamics and life-strategy". Marine Biology. 147 (3): 707–717. Bibcode:2005MarBi.147..707B. doi:10.1007/s00227-005-1607-8. hdl: 11250/174234 . S2CID   54050600.
  36. ↑ Kallevik, I.H.F. (2013). Alternative prey choice in the pteropod Clione limacina (Gastropoda) studied by DNA-based methods. Archived 2016-03-04 at the Wayback Machine Biology Field of study - Arctic Marine Ecology and Resource Biology. Bio-3950 (60 ECT). The University Center in Svalbard.
  37. ↑ Weisberger, Mindy (19 February 2016). "Swim Like a Butterfly? Sea Snail 'Flies' Through Water". Scientific American . Retrieved 20 February 2016.
  38. 1 2 Murphy, D., Adhikari, D., Webster, D., and Yen, J. (2016). "Underwater flight by the planktonic sea butterfly". Journal of Experimental Biology. 219 (4): 535–543. doi: 10.1242/jeb.129205 . PMID   26889002.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  39. ↑ Adhikari, D., Webster, D., and Yen, J. (2016). "Portable tomographic PIV measurements of swimming shelled Antarctic pteropods". Experiments in Fluids. 57 (12): 180–197. Bibcode:2016ExFl...57..180A. doi:10.1007/s00348-016-2269-7. S2CID   125624301.{{cite journal}}: CS1 maint: multiple names: authors list (link)

Further reading

This page is based on this Wikipedia article
Text is available under the CC BY-SA 4.0 license; additional terms may apply.
Images, videos and audio are available under their respective licenses.