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Gymnosomes slowly beat their wing-like parapodia in a rowing motion to propel their "perfectly streamlined" bodies through the upper 20 m of the water column. Although usually slow-moving, beating their wings once or twice per second, they are capable of bursts of speed when they need to pursue their
349:
The wing comprises seven groups of muscles, anterior oblique muscles for dorsal and ventral sides, posterior oblique muscles for dorsal and ventral sides, the longitudinal and transverse wing retractors muscles, and dorso-ventral muscles. The first four groups form a continuous sheet where the dorsal
457:
The gymnosomes, like other shell-less opisthobranchs, discard their shells at metamorphosis, with the retractor muscles being severed and the shell lost. The group does not truly, therefore, lack a shell. Few larval shells have been described (and consequently an understanding of their fossil record
350:
muscles are perpendicular to the ventral muscles. Transverse and longitudinal retractors muscles retract the wing along the span and the chord. The dorso-ventral muscles control the thickness of the wing by changing the haemocoelic pressure inside the wing, this also ejects the buccal apparatus from
430:
stimulates the neuromuscular system to chase the prey, this notably increases the motion of the wings, minimizes the gap between wingtips at the end of each stroke, lessening the aspect ratio, and increases the deflection of the wings. Therefore, reducing the drag forces and overcoming stall. Also,
421:
Their hunting strategies are variable; some forms are ambush predators, sitting and waiting for their prey; whilst others actively pursue their prey; their metabolic rate is closely linked to that of their prey species. Even the size of the gymnosomes is correlated to the size of their prey, which
369:
By rowing their "wings" back and forth at 1–3 Hz, the sea angels swim at speeds up to 100 mm/s (0.22 mph). This is about twice as fast as their prey, the sea butterfly. Tracing the wingtip showed a ‘figure-of-eight’ trajectory and stroke plane orthogonal to the caudal-rostral axis,
261:
Recent molecular data suggest the
Gymnosomata form a sister group to the Thecosomata (other planktonic, weakly or nonmineralized gastropods), but this long-standing hypothesis has also had some recent detractors. Fossils of the group go back to the Middle Frasnian stage of the
295:, the foot of the gastropod has developed into wing-like flapping appendages (parapodia) and larval sea angels discard their embryonic shells a few days after hatching. Both adaptations suit their free-swimming oceanic lives. The adaptations also explain the common name
313:
The lack of a heavy shell for gymnosomes allows them to hover on the upper water surfaces for longer periods, along with a streamlined body that reduces the drag coefficient. Also, gymnosomes have a low aspect ratio which aids them to maneuver and accelerate quickly.
282:
These organisms have a wide geographic range, from polar regions, under sea ice, to equatorial (tropic) seas. From spring to autumn, sea angels live at a depth of 200 m in the Sea of
Okhotsk. In winter, they migrate to the coast of north Hokkaido with drift ice.
1223:
Janssen, A.W. (2003). "Notes on the systematics, morphology and biostratigraphy of fossil holoplanktonic
Mollusca, 13. Considerations on a subdivision of Thecosomata, with the emphasis on genus group classification of Limacinidae".
1102:
1079:
Szymik, B. G., & Satterlie, R. A. (2011). Changes in wingstroke kinematics associated with a change in swimming speed in a pteropod mollusk, Clione limacina. Journal of
Experimental Biology, 214(23),
453:
with internal fertilization. A fertilized animal later releases a gelatinous egg mass, and the eggs float freely until hatching. Their embryonic shells are lost within the first few days after hatching.
393:
causes deleterious lift-drag ratios. (2) High angle of attack typically ranges from 60 to 80 degrees. (3) The orthogonality between the surface of the wing and ascending trajectory of the sea angel.
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422:
they recognize by means of touch and grab using their sometimes-suckered buccal cones. A combination of hooks and a toothed radula are employed to scour the flesh from the thecosomes' shells.
1361:
1005:
SATTERLIE, R. A., LABARBERA, M., & SPENCER, A. N. (1985). Swimming in the
Pteropod Mollusc, Clione Umacina: I. Behaviour and Morphology. Journal of experimental biology, 116(1), 189-204.
1127:
993:
Karakas, F., Wingate, J., Blanco-Bercial, L., Maas, A. E., & Murphy, D. W. (2020). Swimming and sinking behavior of warm water pelagic snails. Frontiers in Marine
Science, 749.
756:
1014:
Murphy, D. W., Adhikari, D., Webster, D. R., & Yen, J. (2016). Underwater flight by the planktonic sea butterfly. Journal of
Experimental Biology, 219(4), 535-543.
1502:
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1541:
366:. Their lifestyles have coevolved with those of their prey, with their feeding strategy adapting to the morphology and consistency of the thecosome shell.
1066:
Borrell, B. J., Goldbogen, J. A., & Dudley, R. (2005). Aquatic wing flapping at low
Reynolds numbers: swimming kinematics of the Antarctic pteropod,
1243:
Peijnenburg, Katja T.C.A.; Janssen, Arie W.; Wall-Palmer, Deborah; Goetze, Erica; Maas, Amy E.; Todd, Jonathan A.; Marlétaz, Ferdinand (2020-09-24).
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the increment in the angle of attack along with low
Reynold number for sea angels resembles the fluid dynamic forces acting on insects such as
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878:
901:"Systematic position of the pelagic Thecosomata and Gymnosomata within Opisthobranchia (Mollusca, Gastropoda) - revival of the Pteropoda"
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is a polar species; those found in warmer waters are far smaller. Some species of sea angels feed exclusively on
708:
as the containing clade is traditional, but contested. Others would omit
Pteropoda and use its containing clade
1357:
https://web.archive.org/web/20040607043553/http://www.biol.sc.edu/~vogt/courses/neuro/neurobehavior.html#clione
250:
Sea angels are also sometimes known as "sea butterflies" but this is potentially misleading because the family
378:. It is not yet clear whether the sea angel uses its swimming appendages as 'rowing paddles' or as 'wings'.
1340:
748:. They have larger, broader parapodia, and most of that species retain a shell; they are commonly known as
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is just one of the families within this clade, and the term "sea butterfly" is also applied to the shelled
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1455:
564:
1564:
621:
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1424:
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726:, is superficially similar to sea angels, but are not closely related; some authorities include both
1245:"The origin and diversification of pteropods precede past perturbations in the Earth's carbon cycle"
1161:
953:"Metabolic temperature compensation and coevolution of locomotory performance in pteropod molluscs"
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1317:
Order Gymnosomata by L. Newman pages 985–989; Beesley, P.L., Ross, G.J.B. & Wells, A (eds) -
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likely accommodates drag-based force production for several reasons, (1) Low aspect ratio for
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prey, calling a separate suite of muscles into action to obtain the higher beat frequency.
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may reach extraordinary levels; up to 300 animals per cubic metre have been recorded.
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325:
319:
228:(Jellyfish and other similar creatures), classified into six different families. They are
199:
54:
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out of the water column, and carry it around for protection. Local population density of
337:
common to mollusks, and tentacles to grasp their prey, sometimes with suckers similar to
317:
Sea angels are gelatinous, mostly transparent, and very small, with the largest species (
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Reconstruction of the phylogeny of Opisthobranchia; Journal of Molluscan studies
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406:. Because of this secretion, predators will not eat the sea angel. A species of
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Gymnosomata are carnivorous, feeding only on their fellow pteropods, the
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takes advantage of this trait: The amphipod will seize an individual of
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1095:"Observations on the Gymnosomatous Pteropod Clione Limacina (Phipps)"
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1157:, with a Review of Development in the Notaspidea (Opisthobranchia)"
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Opisthobranchs with no shells are the gymnosomes (sea angels), the
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The group that used to be the family Thliptodontidae is treated as
370:
this pattern was observed in some aerial insects, small birds, and
1103:
Journal of the Marine Biological Association of the United Kingdom
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292:
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214:
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Journal of Zoological Systematics and Evolutionary Research
741:, whereas others list them as distinct orders within the
1024:
Murphy, D.; Adhikari, D.; Webster, D.; Yen, J. (2016).
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999:
1070:. Journal of Experimental Biology, 208(15), 2939-2949.
243:. Sea angels were previously referred to as a type of
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1026:"Underwater flight by the planktonic sea butterfly"
951:Seibel, B. A.; Dymowska, A.; Rosenthal, J. (2007).
449:Like many gastropods, sea angels are simultaneous
670: (19 species in 13 genera)
1250:Proceedings of the National Academy of Sciences
871:"Order Gymnosomata Blainville 1824 (sea angel)"
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333:; the angels have terminal mouths with the
299:and the scientific name of the order: From
235:in the clade Gymnosomata within the larger
220:) are a large group of small free-swimming
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657: (sea angels)
44:
31:
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1153:"Larval Development and Metamorphosis in
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899:Klussmann-Kolb, A.; Dinapoli, A. (2006).
862:
835:
826:, a subfamily of the family Clionidae.
7:
1557:20d78daa-722b-4943-a384-1767eb20dfc9
763:Gymnosomata is arranged as follows:
958:Integrative and Comparative Biology
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511:
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492:
479:
469:Cladogram of sea angels according
25:
1315:Mollusca - The Southern Synthesis
722:The other suborder of pteropods,
1355:Video of a sea angel in motion:
1334:
920:10.1111/j.1439-0469.2006.00351.x
757:taxonomy of Bouchet & Rocroi
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66:
38:Temporal range: Frasnian-present
1350:entries in the Sea Slug Forum:
1205:from the original on 2011-06-16
1133:from the original on 2017-09-22
1031:Journal of Experimental Biology
881:from the original on 2020-06-18
543:
278:Sea angels in Australian waters
381:However, cold water sea angel
1:
734:as separate branches of the
445:Reproduction and development
1622:
1151:Gibson, Glenys D. (2003).
601:Spongiobranchaea australis
224:, not to be confused with
1362:Mikko's Phylogeny Archive
1116:10.1017/S0025315400023687
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471:to Peijnenburg, Janssen,
374:such as hummingbirds and
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63:Scientific classification
61:
52:
43:
34:
1155:Pleurobranchaea maculata
1272:10.1073/pnas.1920918117
35:Gymnosomata (sea angel)
1093:Morton, J. E. (2009).
323:) reaching 5 cm.
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1565:Paleobiology Database
704:Note that the use of
622:Pneumoderma violaceum
277:
1343:at Wikimedia Commons
678:sea butterflies
458:is as yet unknown).
306:meaning "naked" and
1263:2020PNAS..11725609P
1257:(41): 25609–25617.
1162:Biological Bulletin
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586:
565:Pneumodermopsis spe
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27:Clade of gastropods
1226:Cainozoic Research
1045:10.1242/jeb.129205
972:10.1093/icb/icm089
416:C. antarctica
412:C. antarctica
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1606:Euopisthobranchia
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310:meaning "body".
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1416:Gymnosomata
1410:Wikispecies
1386:Gymnosomata
1341:Gymnosomata
848:nudibranchs
755:In the new
732:Gymnosomata
728:Thecosomata
724:Thecosomata
664:Thecosomata
498:Gymnosomata
427:C. limacina
364:Thecosomata
339:cephalopods
287:Description
256:Thecosomata
218:Gymnosomata
143:Gymnosomata
18:Gymnosomata
1209:2010-06-04
1137:2019-07-12
1080:3935-3947.
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858:References
844:sacoglossa
782:Cliopsidae
769:Clionoidea
434:Drosophila
404:pteroenone
226:Cnidarians
211:Sea angels
165:Cliopsidae
139:Suborder:
110:Gastropoda
1281:0027-8424
830:Footnotes
776:Clionidae
739:Pteropoda
706:Pteropoda
486:Pteropoda
297:sea angel
266:period.
252:Clionidae
222:sea slugs
158:Clionidae
149:Families
133:Pteropoda
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80:Eukaryota
1600:Category
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1346:list of
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1128:Archived
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1054:26889002
981:21669767
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462:Taxonomy
408:amphipod
358:Behavior
291:In this
245:pteropod
100:Mollusca
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90:Animalia
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1290:7568333
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1508:10364
1503:IRMNG
1495:49153
1469:7NFBD
1195:S2CID
1179:JSTOR
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301:Greek
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