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Nudibranch

January 31, 2023

Nudibranchs (/ˈnjuːdɪbræŋk/[2]) are a group of soft-bodied marine gastropod molluscs that shed their shells after their larval stage.[3] They are noted for their often extraordinary colours and striking forms, and they have been given colourful nicknames to match, such as "clown", "marigold", "splendid", "dancer", "dragon", and[4] "sea rabbit".[5] Currently, about 3,000 valid species of nudibranchs are known.[6]

Nudibranch
Berghia coerulescens
Chromodoris lochi pair in Puerto Galera, the Philippines
Scientific classification
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Subclass: Heterobranchia
Infraclass: Euthyneura
Subterclass: Ringipleura
Superorder: Nudipleura
Order: Nudibranchia
Cuvier, 1817
Clades

See text for superfamilies

Diversity[1]
about 3000 species

The word "nudibranch" comes from the Latin nudus "naked" and the Ancient Greek βράγχια (bránkhia) "gills".

Nudibranchs are often casually called sea slugs, as they are a family of opistobranchs (sea slugs), within the phylum Mollusca (molluscs), but many sea slugs belong to several taxonomic groups that are not closely related to nudibranchs. A number of these other sea slugs, such as the photosynthetic Sacoglossa and the colourful Aglajidae, are often confused with nudibranchs.

Distribution and habitat

 
Janolus sp. in Anilao, the Philippines
 
Kalinga ornata from Anilao Pier dive site, depth 5 m

Nudibranchs occur in seas worldwide, ranging from the Arctic, through temperate and tropical regions, to the Southern Ocean around Antarctica.[6][7][8] They are almost entirely restricted to salt water, although a few species are known to inhabit lower salinities in brackish water.[9]

Nudibranchs live at virtually all depths, from the intertidal zone to depths well over 700 m (2,300 ft).[7] The greatest diversity of nudibranchs is seen in warm, shallow reefs, although one nudibranch species was discovered at a depth near 2,500 m (8,200 ft).[10]

Nudibranchs are benthic animals, found crawling over the substrate.[7] The only exceptions to this are the neustonic Glaucus nudibranchs, which float upside down just under the ocean's surface; the pelagic nudibranchs Cephalopyge trematoides, which swim in the water column;[11][12] and Phylliroe bucephalum.[13]

Anatomical description

 
Berghia stephanieae nudibranch body: Note the oral tentacles (ot), foot tentacles (ft), eye (e), rhinophores (r), and cerata (c). This species has cnidosacs (cn) at the cerata tips. Scale bar is 100 μm.

The body forms of nudibranchs vary a great deal, but because they are opisthobranchs, unlike most other gastropods, they are apparently bilaterally symmetrical externally (but not internally) because they have undergone secondary detorsion. In all nudibranchs, the male and female sexual openings are on the right side of the body, reflecting their asymmetrical origins. They lack a mantle cavity. Some species have venomous appendages (cerata) on their sides, which deter predators. Many also have a simple gut and a mouth with a radula.[14]

The eyes in nudibranchs are simple and able to discern little more than light and dark.[15] The eyes are set into the body, are about a quarter of a millimeter in diameter, and consist of a lens and five photoreceptors.[16]

Nudibranchs vary in adult size from 4 to 600 mm (0.16 to 23.62 in).

The adult form is without a shell or operculum (in shelled gastropods, the operculum is a bony or horny plate that can cover the opening of the shell when the body is withdrawn). In most species, there is a swimming veliger larva with a coiled shell, but the shell is shed at metamorphosis when the larva transforms into the adult form. Some species have direct development, and the shell is shed before the animal emerges from the egg mass.[14]

 
The Glaucus atlanticus is an example of a nudibranch that has its cerata positioned like wings instead of on its back.

The name nudibranch is appropriate, since the dorids (infraclass Anthobranchia) breathe through a "naked gill" shaped into branchial plumes in a rosette on their backs.[17] By contrast, on the back of the aeolids in the clade Cladobranchia, brightly coloured sets of protruding organs called cerata are present.

Nudibranchs have cephalic (head) tentacles, which are sensitive to touch, taste, and smell. Club-shaped rhinophores detect odors.

Defence mechanisms

 
Nudibranchs (Nembrotha kubaryana) eating Clavelina tunicate colonies

In the course of their evolution, nudibranchs have lost their shells, while developing alternative defence mechanisms. Some species evolved an external anatomy with textures and colours that mimicked surrounding sessile invertebrate animals (often their prey sponges or soft corals) to avoid predators (camouflage). Other nudibranchs, as seen especially well on Chromodoris quadricolor, have an intensely bright and contrasting colour pattern that makes them especially conspicuous in their surroundings. Nudibranch molluscs are the most commonly cited examples of aposematism in marine ecosystems, but the evidence for this has been contested,[18] mostly because few examples of mimicry are seen among species, many species are nocturnal or cryptic, and bright colours at the red end of the spectrum are rapidly attenuated as a function of water depth. For example, the Spanish dancer nudibranch (genus Hexabranchus), among the largest of tropical marine slugs, potently chemically defended, and brilliantly red and white, is nocturnal and has no known mimics.[19] Other studies of nudibranch molluscs have concluded they are aposematically coloured, for example, the slugs of the family Phylidiidae from Indo-Pacific coral reefs.[20]

Nudibranchs that feed on hydrozoids can store the hydrozoids' nematocysts (stinging cells) in the dorsal body wall, the cerata.[21] These stolen nematocysts, called kleptocnidae, wander through the alimentary tract without harming the nudibranch. Once further into the organ, the cells are assimilated by intestinal protuberances and brought to specific placements on the creature's hind body. Nudibranchs can protect themselves from the hydrozoids and their nematocysts; the specific mechanism is yet unknown, but special cells with large vacuoles probably play an important role. Similarly, some nudibranchs can also take in plant cells (symbiotic algae from soft corals) and reuse these to make food for themselves. The related group of sacoglossan sea slugs feed on algae and retain just the chloroplasts for their own photosynthetic use, a process known as kleptoplasty.

Nudibranchs use a variety of chemical defences to aid in protection,[22] but it is not necessary for the strategy to be lethal to be effective; in fact, good arguments exist that chemical defences should evolve to be distasteful rather than toxic.[23] Some sponge-eating nudibranchs concentrate the chemical defences from their prey sponge in their bodies, rendering themselves distasteful to predators.[19][24] One method of chemical defense used by nudibranchs are secondary metabolites, which play an important role in mediating relationships among marine communities.[25] The evidence that suggests the chemical compounds used by dorid nudibranchs do in fact come from dietary sponges lies in the similarities between the metabolites of prey and nudibranchs, respectively. Furthermore, nudibranchs contain a mixture of sponge chemicals when they are in the presence of multiple food sources, as well as change defence chemicals with a concurrent change in diet.[26] This, however, is not the only way for nudibranchs to develop chemical defences. Certain Antarctic marine species defense mechanisms are believed to be controlled by biological factors like predation and competition, and selective pressures.[27] Certain species are able to produce their own chemicals de novo without dietary influence. Evidence for the different methods of chemical production comes with the characteristic uniformity of chemical composition across drastically different environments and geographic locations found throughout de novo production species compared to the wide variety of dietary and environmentally dependent chemical composition in sequestering species.[28]

Another method of protection is the release of the ugdon acid from the skin.[29] Once the specimen is physically irritated or touched by another creature, it will release the mucus automatically, eating the animal from the inside out.

Apparent production of sound

In 1884, Philip Henry Gosse reported observations by "Professor Grant" (possibly Robert Edmond Grant) that two species of nudibranchs emit sounds that are audible to humans.[30]

Two very elegant species of Sea-slug, viz., Eolis punctata [i.e. Facelina annulicornis], and Tritonia arborescens [i.e. Dendronotus frondosus], certainly produce audible sounds. Professor Grant, who first observed the interesting fact in some specimens of the latter which he was keeping in an aquarium, says of the sounds, that 'they resemble very much the clink of a steel wire on the side of the jar, one stroke only been given at a time, and repeated at intervals of a minute or two; when placed in a large basin of water the sound is much obscured, and is like that of a watch, one stroke being repeated, as before, at intervals. The sound is longest and most often repeated when the Tritonia are lively and moving about, and is not heard when they are cold and without any motion; in the dark I have not observed any light emitted at the time of the stroke; no globule of air escapes to the surface of the water, nor is any ripple produced on the surface at the instant of the stroke; the sound, when in a glass vessel, is mellow and distinct.' The Professor has kept these Tritonia alive in his room for a month, and during the whole period of their confinement they have continued to produce the sounds with very little diminution of their original intensity. In a small apartment they are audible at the distance of twelve feet. The sounds obviously proceed from the mouth of the animal; and at the instant of the stroke, we observe the lips suddenly separate, as if to allow the water to rush into a small vacuum formed within. As these animals are hermaphrodites, requiring mutual impregnation, the sounds may possibly be a means of communication between them, or, if they are of an electric nature, they may be the means of defending from foreign enemies one of the most delicate, defenceless, and beautiful Gasteropods that inhabit the deep.

Lifecycle

 
Mating behavior in Nembrotha purpureolineata
 
Acanthodoris lutea laying eggs

Nudibranchs are hermaphroditic, thus having a set of reproductive organs for both sexes, but they cannot fertilize themselves.[31] Mating usually takes a few minutes, and involves a dance-like courtship. Nudibranchs typically deposit their eggs within a gelatinous spiral,[32] which is often described as looking like a ribbon. The number of eggs varies; it can be as few as just 1 or 2 eggs (Vayssierea felis) or as many as an estimated 25 million (Aplysia fasciata{not a nudibranch}). The eggs contain toxins from sea sponges as a means of deterring predators.[33] After hatching, the infants look almost identical to their adult counterparts, albeit smaller. Infants may also have fewer cerata. The lifespan of nudibranchs can range from a few weeks to a year, depending on the species.

Feeding and ecological role

 
Pteraeolidia ianthina has adapted cerata to house symbiotic zooxanthellae obtained from its diet, which continue to photosynthesize and provide energy to the nudibranch.

All known nudibranchs are carnivorous.[31] Some feed on sponges, others on hydroids (e.g. Cuthona),[34] others on bryozoans (phanerobranchs such as Tambja, Limacia, Plocamopherus and Triopha),[35] and some eat other sea slugs or their eggs (e.g. Favorinus)[36] or, on some occasions, are cannibals and prey on members of their own species. Other groups feed on tunicates (e.g. Nembrotha, Goniodoris),[37] other nudibranchs (Roboastra, which are descended from tunicate-feeding species),[37] barnacles (e.g. Onchidoris bilamellata),[38] and anemones (e.g. the Aeolidiidae and other Cladobranchia).[35]

The surface-dwelling nudibranch, Glaucus atlanticus, is a specialist predator of siphonophores, such as the Portuguese man o' war. This predatory mollusc sucks air into its stomach to keep it afloat, and using its muscular foot, it clings to the surface film. If it finds a small victim, Glaucus simply envelops it with its capacious mouth, but if the prey is a larger siphonophore, the mollusc nibbles off its fishing tentacles, the ones carrying the most potent nematocysts. Like some others of its kind, Glaucus does not digest the nematocysts; instead, it uses them to defend itself by passing them from its gut to the surface of its skin.[39]

Taxonomy

 
Dorids (Chromodoris willani shown) breathe with the branchial plume, which projects from around their anus.
 
Aeolids (Hermissenda crassicornis pictured) have many cerata over their back which are used for defense and respiration.
Nudibranchs are frequently differentiated as either dorid or aeolid.

Nudibranchs are commonly divided into two main kinds, dorid and aeolid (also spelled eolid) nudibranchs:[40][41]

  • Dorids (clade Anthobranchia, Doridacea, or Doridoidea) are recognised by having an intact digestive gland and the feather-like branchial (gill) plume, which forms a cluster on the posterior part of the body, around the anus. Fringes on the mantle do not contain any intestines.[citation needed] Additionally, dorid nudibranchs commonly have distinct pockets, bumps, and/or mantle dermal formations, which are distortions on their skin, used to store bioactive defense chemicals.[42]
  • Aeolids (clade Cladobranchia) have cerata (spread across the back) instead of the branchial plume. The cerata function in place of gills and facilitate gas exchange through the epidermis. Additionally, aeolids possess a branched digestive gland, which may extend into the cerate and often has tips that contain cnidosacs (stinging cells absorbed from prey species and then used by the nudibranch).[43] They lack a mantle.[citation needed] Some are hosts to zooxanthellae.

The exact systematics of nudibranchs are a topic of recent revision. Traditionally, nudibranchs have been treated as the order Nudibranchia, located in the gastropod mollusc subclass Opisthobranchia (the marine slugs: which consisted of nudibranchs, sidegill slugs, bubble snails, algae sap-sucking sea slugs, and sea hares).[40] Since 2005,[44] pleurobranchs (which had previously been grouped among sidegill slugs) have been placed alongside nudibranchs in the clade Nudipleura (recognising them as more closely related to each other than to other opisthobranchs).[45] Since 2010, Opisthobranchia has been recognised as not a valid clade (it is paraphyletic) and instead Nudipleura has been placed as the first offshoot of Euthyneura (which is the dominant clade of gastropods).[46]


Traditional hierarchy

This classification was based on the work of Johannes Thiele (1931),[47] who built on the concepts of Henri Milne-Edwards (1848).[48]

 
1852 watercolor of a Nudibranch by Jacques Burkhardt

Order Nudibranchia:

Early revisions

Newer insights derived from morphological data and gene-sequence research seemed to confirm those ideas. On the basis of investigation of 18S rDNA sequence data, strong evidence supports the monophyly of the Nudibranchia and its two major groups, the Anthobranchia/Doridoidea and Cladobranchia.[49] A study published in May 2001, again revised the taxonomy of the Nudibranchia.[50] They were thus divided into two major clades:

However, according to the taxonomy by Bouchet & Rocroi (2005), currently the most up-to-date system of classifying the gastropods, the Nudibranchia are a subclade within the clade of the Nudipleura. The Nudibranchia are then divided into two clades:

Gallery

This gallery shows some of the great variability in the color and form of nudibranchs, and nudibranch egg ribbons.

See also

References

  1. ^ Wägele, H.; Klussmann-Kolb, A. (2005). "Opisthobranchia (Mollusca, Gastropoda) more than just slimy slugs. Shell reduction and its implications on defence and foraging". Frontiers in Zoology. 2 (1): 1–18. doi:10.1186/1742-9994-2-3. PMC 554092. PMID 15715915.
  2. ^ Longman Pronunciation Dictionary (2nd edition), ISBN 0-582-36467-1
  3. ^ Thompson, T. E. (2009). "Feeding in nudibranch larvae" (PDF). Journal of the Marine Biological Association of the United Kingdom. 38 (2): 239–248. doi:10.1017/S0025315400006044. S2CID 86275359.
  4. ^ Turnbull, John (Spring 2016). "The Nudibranch – Creature Feature". Nature New South Wales. 60 (3): 16–17.
  5. ^ Bronson, Wilfrid. Water People, 1935
  6. ^ a b Ocean Portal (2017). A Collage of Nudibranch Colors. Smithsonian National Museum of Natural History. Retrieved 17 April 2018.
  7. ^ a b c Nudibranchs 2013-08-14 at the Wayback Machine, Fishermen Scuba.
  8. ^ Ekimova, I.; T. Korshunova; D. Schepetov; T. Neretina; N. Sanamyan; A. Martynov (2015). "Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus (Mollusca, Gastropoda), with descriptions of three new species". Zoological Journal of the Linnean Society. 173 (4): e0192177. doi:10.1111/zoj.12214.
  9. ^ Korshunova, T.; K. Lundin; K. Malmberg; B. Picton; A. Martynov (2018). "First true brackish-water nudibranch mollusc provides new insights for phylogeny and biogeography and reveals paedomorphosis-driven evolution". PLOS ONE. 13 (1): e0192177. Bibcode:2018PLoSO..1392177K. doi:10.1371/journal.pone.0192177. PMC 5851531. PMID 29538398.
  10. ^ . Monterey Bay Aquarium Research Institute. Archived from the original on 9 October 2013. Retrieved 16 October 2013.
  11. ^ Steinberg, J. E. (1956). "The pelagic nudibranch, Cephalopyge trematoides (Chun, 1889), in New South Wales with a note on other species in this genus". Proceedings of the Linnean Society of New South Wales. 81: 184–192.
  12. ^ G.M. Mapstone & M.N. Arai, Siphonophora (Cnidaria, Hydrozoa) of Canadian Pacific Waters, p.33. "The best documented predators of pelagic cnidarians from the phylum Mollusca are the neustonic nudibranchs and snails [...and] the pelagic nudibranch [...]"
  13. ^ Gosliner TM, Valdes A Behrens DW 2015 Nudibranch and Sea Slug Identification Indo-Pacific New World Publications Jacksonville Florida USA
  14. ^ a b Thompson, T. E. 1976. Biology of opisthobranch molluscs, vol. 1, 207 pp., 21 pls. Ray Society, no. 151.
  15. ^ "Nudibranchs – National Geographic Magazine". Ngm.nationalgeographic.com. 2013-04-25. Retrieved 2013-07-04.
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  17. ^ Dayrat, B. (2005). "Advantages of naming species under the PhyloCode: An example of how a new species of Discodorididae (Mollusca, Gastropoda, Euthyneura, Nudibranchia, Doridina) may be named" (PDF). Marine Biology Research. 1 (3): 216–232. doi:10.1080/17451000510019141. S2CID 53557429. Retrieved 2009-06-14.
  18. ^ Edmunds, M. (1991). "Does warning colouration occur in nudibranchs?". Malacologia. 32: 241–255.
  19. ^ a b Pawlik, JR; et al. (1988). "Defensive chemicals of the Spanish Dancer nudibranch, Hexabranchus sanguineus, and its egg ribbons: Macrolides derived from a sponge diet". Journal of Experimental Marine Biology and Ecology. 119 (2): 99–109. doi:10.1016/0022-0981(88)90225-0.
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  21. ^ Frick, K (2003). . In: SF Norton (Ed). Diving for Science...2003. Proceedings of the American Academy of Underwater Sciences (22nd Annual Scientific Diving Symposium). Archived from the original on January 29, 2009. Retrieved 2008-07-03.{{cite journal}}: CS1 maint: unfit URL (link)
  22. ^ Karuso, P. (1987). "Chemical Ecology of the Nudibranchs". In Scheuer, PJ (ed.). Bioorganic Marine Chemistry. Vol. 1. Springer-Verlag. pp. 31–60. doi:10.1007/978-3-642-72726-9_2. ISBN 978-3-642-72728-3., a comprehensive review of the chemical ecology of the nudibranchs
  23. ^ Pawlik, JR (2012). Fattorusso, E.; et al. (eds.). Antipredatory defensive roles of natural products from marine invertebrates. Handbook of Marine Natural Products. NY: Springer Science. pp. 677–710.
  24. ^ Gosliner, T. M. (1987). Nudibranchs of Southern Africa. ISBN 978-0-930118-13-6.
  25. ^ Avila, C; Iken, K; Fontana, A; Cimino, G (2000-09-05). "Chemical ecology of the Antarctic nudibranch Bathydoris hodgsoni Eliot, 1907: defensive role and origin of its natural products". Journal of Experimental Marine Biology and Ecology. 252 (1): 27–44. doi:10.1016/S0022-0981(00)00227-6. ISSN 0022-0981. PMID 10962063.
  26. ^ Faulkner, D. J.; Ghiselin, M. T. (1983). "Chemical defence and evolutionary ecology of dorid nudibranchs and some other opisthobranch gastropods". Marine Ecology Progress Series. 13: 295–301. Bibcode:1983MEPS...13..295F. doi:10.3354/meps013295.
  27. ^ Avila, C; Iken, K; Fontana, A; Cimino, G (2000-09-05). "Chemical ecology of the Antarctic nudibranch Bathydoris hodgsoni Eliot, 1907: defensive role and origin of its natural products". Journal of Experimental Marine Biology and Ecology. 252 (1): 27–44. doi:10.1016/S0022-0981(00)00227-6. ISSN 0022-0981. PMID 10962063.
  28. ^ Barsby, T.; Linington, R. G.; Andersen, R. J. (2002). "De Novo terpenoid biosynthesis by the dendronotid nudibranch Melibe leonina". Chemoecology. 12 (4): 199–202. doi:10.1007/PL00012669. S2CID 35384332.
  29. ^ Edmunds, M. (1968). "Acid secretion in some species of Doridacea (Mollusca, Nudibranchia)". Proceedings of the Malacological Society of London. 38 (2): 121–133. Archived from the original on 2013-04-15.
  30. ^ P.H. Gosse, Evenings at the Microscope, 1884 edition,[1] p57
  31. ^ a b "Nudibranch". Aquaticcommunity.com. Retrieved 2013-07-04.
  32. ^ Klussmann-Kolb A (2001). "The Reproductive Systems of the Nudibranchia (Gastropoda, Opisthobranchia): Comparative Histology and Ultrastructure of the Nidamental Glands with Aspects of Functional Morphology". Zoologischer Anzeiger. 240 (2): 119–136. doi:10.1078/0044-5231-00011.
  33. ^ "Diving with Nudibranchs". Dive the World.
  34. ^ NC Folino (1997). (PDF). American Malacological Bulletin. Archived from the original (PDF) on 2012-06-25. Retrieved 2013-03-07.
  35. ^ a b Domínguez, M.; Troncoso, J. S.; García, F. J. (2008). "The family Aeolidiidae Gray, 1827 (Gastropoda Opisthobranchia) from Brazil, with a description of a new species belonging to the genus Berghia Trinchese, 1877". Zoological Journal of the Linnean Society. 153 (2): 349–368. doi:10.1111/j.1096-3642.2008.00390.x.
  36. ^ Rudman, W.B. (1999-03-19). "Favorinus tsuruganus Baba & Abe, 1964. [In] Sea Slug Forum. Australian Museum".
  37. ^ a b Valdés, Á. (2004). "Phylogeography and phyloecology of dorid nudibranchs (Mollusca, Gastropoda)". Biological Journal of the Linnean Society. 83 (4): 551–559. doi:10.1111/j.1095-8312.2004.00413.x.
  38. ^ Barnes, H.; Powell, H. T. (1954). "Onchidoris fusca (Müller); A Predator of Barnacles". Journal of Animal Ecology. 23 (2): 361–363. doi:10.2307/1986. JSTOR 1986.
  39. ^ Piper, Ross (2007), Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals, Greenwood Press.
  40. ^ a b Hans Bertsch, Nudibranchs: Marine slugs with verve. "Navanax inermis[..] is the bane of all nudibranchs, because it is one of the few known predators on this group of slugs. [...] Dorids mainly eat sponges, bryozoans, and tunicates, whereas aeolids principally eat cnidarians."
  41. ^ "Facts About Nudibranchs". Marinelife.about.com. 2011-11-10. Retrieved 2013-07-04.
  42. ^ Dean, Lewis J.; Prinsep, Michèle R. (2017). "The chemistry and chemical ecology of nudibranchs". Natural Product Reports. 34 (12): 1359–1390. doi:10.1039/C7NP00041C. ISSN 0265-0568. PMID 29135002.
  43. ^ Cella, Kristen; Carmona, Leila; Ekimova, Irina; Chichvarkhin, Anton; Schepetov, Dimitry; Gosliner, Terrence M. (2016-12-15). "A Radical Solution: The Phylogeny of the Nudibranch Family Fionidae". PLOS ONE. 11 (12): e0167800. Bibcode:2016PLoSO..1167800C. doi:10.1371/journal.pone.0167800. ISSN 1932-6203. PMC 5158052. PMID 27977703.
  44. ^ Taxonomy of the Gastropoda, Bouchet & Rocroi, 2005
  45. ^ Poppe, Guido T.; Tagaro, Sheila P. (February 23, 2006). (PDF). Visaya. Archived from the original (PDF) on September 27, 2007.
  46. ^ Jörger, K. M.; Stöger, I.; Kano, Y.; et al. (2010). "On the origin of Acochlidia and other enigmatic euthyneuran gastropods, with implications for the systematics of Heterobranchia". BMC Evolutionary Biology. 10: 323. doi:10.1186/1471-2148-10-323. PMC 3087543. PMID 20973994. At the basis of the Euthyneura the Nudipleura split off
  47. ^ Thiele, J. (1931). Handbuch der systematischen Weichtierkunde, II. Verlag von Gustav Fischer, Jena, Germany.
  48. ^ Milne-Edwards H (1848). Note sur la classification naturelle chez Mollusques Gasteropodes. Annales des Sciences Naturelles, series 3, 9: 102-112.
  49. ^ Wägele H. & Willan R. C. (September 2000). "Phylogeny of the Nudibranchia". Zoological Journal of the Linnean Society. 130 (1): 83–181. doi:10.1111/j.1096-3642.2000.tb02196.x.
  50. ^ Schrödl M., Wägele H. & Willan R. C. (2001). "Taxonomic Redescription of the Doridoxidae (Gastropoda: Opisthobranchia), an Enigmatic Family of Deep Water Nudibranchs, with Discussion of Basal Nudibranch Phylogeny". Zoologischer Anzeiger. 240 (1): 83–97. doi:10.1078/0044-5231-00008.

Further reading

  • Thompson, T. E. (1976). Biology of opisthobranch molluscs Vol. 1. 207 pp., 21 pls. Ray Society, no. 151.
  • Thompson, T. E., & G. H. Brown (1984). Biology of opisthobranch molluscs Vol. 2. 229 pp., 41 pls. Ray Society, no. 156.
  • McDonald, Gary R. (7 July 2021). Institute of Marine Sciences. Bibliographia Nudibranchia, 3rd online Edition. A listing, by Author, of publications on nudibranchs.
  • McDonald, Gary R. (7 July 2021). Institute of Marine Sciences. Nudibranch Systematic Index, 3rd online Edition. An index of names given to nudibranchs and their subsequent use, referenced to Bibliographia Nudibranchia.
  • McDonald, Gary R. & J. W. Nybakken. (November 5, 2014). List of the Worldwide Food Habits of Nudibranchs
  • Coleman, Neville (2008). Nudibranchs Encyclopedia: Catalogue of Asia/Indo-Pacific Sea Slugs. Neville Coleman's Underwater Geographic. ISBN 0-947325-41-7

External links

 
Wikispecies has information related to Nudibranchia.
 
Wikimedia Commons has media related to Nudibranchia.
  • Sea Slug Forum by William B. Rudman
  • Nudibranchs of the British Isles
  • OPK Opistobranquis – Iberian and Mediterranean Opisthobranchs
  • Mediterranean slug site (actually a misnomer – Worldwide coverage
  • The Slug Site, Michael D. Miller 2002–2014
  • The Okinawa Slug Site
  • Nudibranch Photos by Mick Tait
  • Nudi Pixel: Online resource for nudibranchs and sea slugs identification using photographs 2011-05-27 at the Wayback Machine
  • Various nudibranch species from Indonesia, Philippines and Thailand
  • Scottish Nudibranchs: Online resource for identification of species found in Scottish waters
  • National Geographic Nudibranch Photo Gallery
  • Sea Slugs of Hawaii
  • Slug City – Molluscs. Brain & Behavior, from the University of Illinois at Urbana-Champaign

Videos

  • Attack of the Sea Slugs at YouTube
  • The Lynx Nudibranch: HD clip of Phidiana lynceus carefully consuming a hydroid Myrionema amboinense.
  • Slug City – Molluscs. Brain & Behavior: many videos of nudibranchs from the University of Illinois at Urbana-Champaign

January 31, 2023
nudibranch, group, soft, bodied, marine, gastropod, molluscs, that, shed, their, shells, after, their, larval, stage, they, noted, their, often, extraordinary, colours, striking, forms, they, have, been, given, colourful, nicknames, match, such, clown, marigol. Nudibranchs ˈ nj uː d ɪ b r ae ŋ k 2 are a group of soft bodied marine gastropod molluscs that shed their shells after their larval stage 3 They are noted for their often extraordinary colours and striking forms and they have been given colourful nicknames to match such as clown marigold splendid dancer dragon and 4 sea rabbit 5 Currently about 3 000 valid species of nudibranchs are known 6 NudibranchBerghia coerulescensChromodoris lochi pair in Puerto Galera the PhilippinesScientific classificationKingdom AnimaliaPhylum MolluscaClass GastropodaSubclass HeterobranchiaInfraclass EuthyneuraSubterclass RingipleuraSuperorder NudipleuraOrder NudibranchiaCuvier 1817CladesEuctenidiacea DexiarchiaSee text for superfamiliesDiversity 1 about 3000 speciesThe word nudibranch comes from the Latin nudus naked and the Ancient Greek bragxia brankhia gills Nudibranchs are often casually called sea slugs as they are a family of opistobranchs sea slugs within the phylum Mollusca molluscs but many sea slugs belong to several taxonomic groups that are not closely related to nudibranchs A number of these other sea slugs such as the photosynthetic Sacoglossa and the colourful Aglajidae are often confused with nudibranchs Contents 1 Distribution and habitat 2 Anatomical description 3 Defence mechanisms 3 1 Apparent production of sound 4 Lifecycle 5 Feeding and ecological role 6 Taxonomy 6 1 Traditional hierarchy 6 2 Early revisions 7 Gallery 8 See also 9 References 10 Further reading 11 External links 11 1 VideosDistribution and habitat Edit Janolus sp in Anilao the Philippines Kalinga ornata from Anilao Pier dive site depth 5 m Nudibranchs occur in seas worldwide ranging from the Arctic through temperate and tropical regions to the Southern Ocean around Antarctica 6 7 8 They are almost entirely restricted to salt water although a few species are known to inhabit lower salinities in brackish water 9 Nudibranchs live at virtually all depths from the intertidal zone to depths well over 700 m 2 300 ft 7 The greatest diversity of nudibranchs is seen in warm shallow reefs although one nudibranch species was discovered at a depth near 2 500 m 8 200 ft 10 Nudibranchs are benthic animals found crawling over the substrate 7 The only exceptions to this are the neustonic Glaucus nudibranchs which float upside down just under the ocean s surface the pelagic nudibranchs Cephalopyge trematoides which swim in the water column 11 12 and Phylliroe bucephalum 13 Anatomical description Edit Berghia stephanieae nudibranch body Note the oral tentacles ot foot tentacles ft eye e rhinophores r and cerata c This species has cnidosacs cn at the cerata tips Scale bar is 100 mm The body forms of nudibranchs vary a great deal but because they are opisthobranchs unlike most other gastropods they are apparently bilaterally symmetrical externally but not internally because they have undergone secondary detorsion In all nudibranchs the male and female sexual openings are on the right side of the body reflecting their asymmetrical origins They lack a mantle cavity Some species have venomous appendages cerata on their sides which deter predators Many also have a simple gut and a mouth with a radula 14 The eyes in nudibranchs are simple and able to discern little more than light and dark 15 The eyes are set into the body are about a quarter of a millimeter in diameter and consist of a lens and five photoreceptors 16 Nudibranchs vary in adult size from 4 to 600 mm 0 16 to 23 62 in The adult form is without a shell or operculum in shelled gastropods the operculum is a bony or horny plate that can cover the opening of the shell when the body is withdrawn In most species there is a swimming veliger larva with a coiled shell but the shell is shed at metamorphosis when the larva transforms into the adult form Some species have direct development and the shell is shed before the animal emerges from the egg mass 14 The Glaucus atlanticus is an example of a nudibranch that has its cerata positioned like wings instead of on its back The name nudibranch is appropriate since the dorids infraclass Anthobranchia breathe through a naked gill shaped into branchial plumes in a rosette on their backs 17 By contrast on the back of the aeolids in the clade Cladobranchia brightly coloured sets of protruding organs called cerata are present Nudibranchs have cephalic head tentacles which are sensitive to touch taste and smell Club shaped rhinophores detect odors Defence mechanisms Edit Nudibranchs Nembrotha kubaryana eating Clavelina tunicate colonies In the course of their evolution nudibranchs have lost their shells while developing alternative defence mechanisms Some species evolved an external anatomy with textures and colours that mimicked surrounding sessile invertebrate animals often their prey sponges or soft corals to avoid predators camouflage Other nudibranchs as seen especially well on Chromodoris quadricolor have an intensely bright and contrasting colour pattern that makes them especially conspicuous in their surroundings Nudibranch molluscs are the most commonly cited examples of aposematism in marine ecosystems but the evidence for this has been contested 18 mostly because few examples of mimicry are seen among species many species are nocturnal or cryptic and bright colours at the red end of the spectrum are rapidly attenuated as a function of water depth For example the Spanish dancer nudibranch genus Hexabranchus among the largest of tropical marine slugs potently chemically defended and brilliantly red and white is nocturnal and has no known mimics 19 Other studies of nudibranch molluscs have concluded they are aposematically coloured for example the slugs of the family Phylidiidae from Indo Pacific coral reefs 20 Nudibranchs that feed on hydrozoids can store the hydrozoids nematocysts stinging cells in the dorsal body wall the cerata 21 These stolen nematocysts called kleptocnidae wander through the alimentary tract without harming the nudibranch Once further into the organ the cells are assimilated by intestinal protuberances and brought to specific placements on the creature s hind body Nudibranchs can protect themselves from the hydrozoids and their nematocysts the specific mechanism is yet unknown but special cells with large vacuoles probably play an important role Similarly some nudibranchs can also take in plant cells symbiotic algae from soft corals and reuse these to make food for themselves The related group of sacoglossan sea slugs feed on algae and retain just the chloroplasts for their own photosynthetic use a process known as kleptoplasty Nudibranchs use a variety of chemical defences to aid in protection 22 but it is not necessary for the strategy to be lethal to be effective in fact good arguments exist that chemical defences should evolve to be distasteful rather than toxic 23 Some sponge eating nudibranchs concentrate the chemical defences from their prey sponge in their bodies rendering themselves distasteful to predators 19 24 One method of chemical defense used by nudibranchs are secondary metabolites which play an important role in mediating relationships among marine communities 25 The evidence that suggests the chemical compounds used by dorid nudibranchs do in fact come from dietary sponges lies in the similarities between the metabolites of prey and nudibranchs respectively Furthermore nudibranchs contain a mixture of sponge chemicals when they are in the presence of multiple food sources as well as change defence chemicals with a concurrent change in diet 26 This however is not the only way for nudibranchs to develop chemical defences Certain Antarctic marine species defense mechanisms are believed to be controlled by biological factors like predation and competition and selective pressures 27 Certain species are able to produce their own chemicals de novo without dietary influence Evidence for the different methods of chemical production comes with the characteristic uniformity of chemical composition across drastically different environments and geographic locations found throughout de novo production species compared to the wide variety of dietary and environmentally dependent chemical composition in sequestering species 28 Another method of protection is the release of the ugdon acid from the skin 29 Once the specimen is physically irritated or touched by another creature it will release the mucus automatically eating the animal from the inside out Apparent production of sound Edit In 1884 Philip Henry Gosse reported observations by Professor Grant possibly Robert Edmond Grant that two species of nudibranchs emit sounds that are audible to humans 30 Two very elegant species of Sea slug viz Eolis punctata i e Facelina annulicornis and Tritonia arborescens i e Dendronotus frondosus certainly produce audible sounds Professor Grant who first observed the interesting fact in some specimens of the latter which he was keeping in an aquarium says of the sounds that they resemble very much the clink of a steel wire on the side of the jar one stroke only been given at a time and repeated at intervals of a minute or two when placed in a large basin of water the sound is much obscured and is like that of a watch one stroke being repeated as before at intervals The sound is longest and most often repeated when the Tritonia are lively and moving about and is not heard when they are cold and without any motion in the dark I have not observed any light emitted at the time of the stroke no globule of air escapes to the surface of the water nor is any ripple produced on the surface at the instant of the stroke the sound when in a glass vessel is mellow and distinct The Professor has kept these Tritonia alive in his room for a month and during the whole period of their confinement they have continued to produce the sounds with very little diminution of their original intensity In a small apartment they are audible at the distance of twelve feet The sounds obviously proceed from the mouth of the animal and at the instant of the stroke we observe the lips suddenly separate as if to allow the water to rush into a small vacuum formed within As these animals are hermaphrodites requiring mutual impregnation the sounds may possibly be a means of communication between them or if they are of an electric nature they may be the means of defending from foreign enemies one of the most delicate defenceless and beautiful Gasteropods that inhabit the deep Lifecycle EditThis section needs expansion You can help by adding to it January 2018 Mating behavior in Nembrotha purpureolineata Acanthodoris lutea laying eggs Nudibranchs are hermaphroditic thus having a set of reproductive organs for both sexes but they cannot fertilize themselves 31 Mating usually takes a few minutes and involves a dance like courtship Nudibranchs typically deposit their eggs within a gelatinous spiral 32 which is often described as looking like a ribbon The number of eggs varies it can be as few as just 1 or 2 eggs Vayssierea felis or as many as an estimated 25 million Aplysia fasciata not a nudibranch The eggs contain toxins from sea sponges as a means of deterring predators 33 After hatching the infants look almost identical to their adult counterparts albeit smaller Infants may also have fewer cerata The lifespan of nudibranchs can range from a few weeks to a year depending on the species Feeding and ecological role Edit Pteraeolidia ianthina has adapted cerata to house symbiotic zooxanthellae obtained from its diet which continue to photosynthesize and provide energy to the nudibranch All known nudibranchs are carnivorous 31 Some feed on sponges others on hydroids e g Cuthona 34 others on bryozoans phanerobranchs such as Tambja Limacia Plocamopherus and Triopha 35 and some eat other sea slugs or their eggs e g Favorinus 36 or on some occasions are cannibals and prey on members of their own species Other groups feed on tunicates e g Nembrotha Goniodoris 37 other nudibranchs Roboastra which are descended from tunicate feeding species 37 barnacles e g Onchidoris bilamellata 38 and anemones e g the Aeolidiidae and other Cladobranchia 35 The surface dwelling nudibranch Glaucus atlanticus is a specialist predator of siphonophores such as the Portuguese man o war This predatory mollusc sucks air into its stomach to keep it afloat and using its muscular foot it clings to the surface film If it finds a small victim Glaucus simply envelops it with its capacious mouth but if the prey is a larger siphonophore the mollusc nibbles off its fishing tentacles the ones carrying the most potent nematocysts Like some others of its kind Glaucus does not digest the nematocysts instead it uses them to defend itself by passing them from its gut to the surface of its skin 39 Taxonomy Edit Dorids Chromodoris willani shown breathe with the branchial plume which projects from around their anus Aeolids Hermissenda crassicornis pictured have many cerata over their back which are used for defense and respiration Nudibranchs are frequently differentiated as either dorid or aeolid Nudibranchs are commonly divided into two main kinds dorid and aeolid also spelled eolid nudibranchs 40 41 Dorids clade Anthobranchia Doridacea or Doridoidea are recognised by having an intact digestive gland and the feather like branchial gill plume which forms a cluster on the posterior part of the body around the anus Fringes on the mantle do not contain any intestines citation needed Additionally dorid nudibranchs commonly have distinct pockets bumps and or mantle dermal formations which are distortions on their skin used to store bioactive defense chemicals 42 Aeolids clade Cladobranchia have cerata spread across the back instead of the branchial plume The cerata function in place of gills and facilitate gas exchange through the epidermis Additionally aeolids possess a branched digestive gland which may extend into the cerate and often has tips that contain cnidosacs stinging cells absorbed from prey species and then used by the nudibranch 43 They lack a mantle citation needed Some are hosts to zooxanthellae The exact systematics of nudibranchs are a topic of recent revision Traditionally nudibranchs have been treated as the order Nudibranchia located in the gastropod mollusc subclass Opisthobranchia the marine slugs which consisted of nudibranchs sidegill slugs bubble snails algae sap sucking sea slugs and sea hares 40 Since 2005 44 pleurobranchs which had previously been grouped among sidegill slugs have been placed alongside nudibranchs in the clade Nudipleura recognising them as more closely related to each other than to other opisthobranchs 45 Since 2010 Opisthobranchia has been recognised as not a valid clade it is paraphyletic and instead Nudipleura has been placed as the first offshoot of Euthyneura which is the dominant clade of gastropods 46 Traditional hierarchy Edit This classification was based on the work of Johannes Thiele 1931 47 who built on the concepts of Henri Milne Edwards 1848 48 1852 watercolor of a Nudibranch by Jacques Burkhardt Order Nudibranchia Infraorder Anthobranchia Ferussac 1819 dorids Superfamily Doridoidea Rafinesque 1815 Superfamily Doridoxoidea Bergh 1900 Superfamily Onchidoridoidea Alder amp Hancock 1845 Superfamily Polyceroidea Alder amp Hancock 1845 Infraorder Cladobranchia Willan amp Morton 1984 aeolids Superfamily Aeolidioidea J E Gray 1827 Superfamily Arminoidea Rafinesque 1814 Superfamily Dendronotoidea Allman 1845 Superfamily Metarminoidea Odhner in Franc 1968Early revisions Edit Newer insights derived from morphological data and gene sequence research seemed to confirm those ideas On the basis of investigation of 18S rDNA sequence data strong evidence supports the monophyly of the Nudibranchia and its two major groups the Anthobranchia Doridoidea and Cladobranchia 49 A study published in May 2001 again revised the taxonomy of the Nudibranchia 50 They were thus divided into two major clades Anthobranchia Bathydoridoidea Doridoidea Dexiarchia nom nov Doridoxoidea Dendronotoidea Aeolidoidea Arminoidea However according to the taxonomy by Bouchet amp Rocroi 2005 currently the most up to date system of classifying the gastropods the Nudibranchia are a subclade within the clade of the Nudipleura The Nudibranchia are then divided into two clades Euctenidiacea Holohepatica Gnathodoridacea contains only Bathydorididae Doridacea Doridoidea Phyllidioidea Onchidoridoidea Polyceroidea Phanerobranchiata Non Suctoria Dexiarchia Actenidiacea Pseudoeuctenidiacea Doridoxida Cladobranchia Cladohepatica Euarminida Dendronotida Aeolidida Unassigned Cladobranchia previously Metarminoidea Charcotiidae Dironidae Goniaeolididae Heroidae Proctonotidae Madrellidae Pinufiidae EmbletoniidaeGallery EditThis gallery shows some of the great variability in the color and form of nudibranchs and nudibranch egg ribbons Tritoniopsis elegans Nembrotha cristata in Bunaken National Park Clown nudibranch Triopha catalinae Northern California Chromodoris annae from Lembeh Straits Indonesia Spanish dancer taken at night Red Sea Nembrotha chamberlaini from Verde Island the Philippines Glossodoris atromarginata Chromodoris dianae from Verde Island the Philippines A pair of Nembrotha milleri mating at Verde Island the Philippines Regal sea goddess Felimare picta in the Gray s Reef National Marine Sanctuary Savannah Georgia Flabellina affinis at La Herradura Mediterranean Sea Spain Dorid nudibranch egg ribbon in Moss Beach California Nudibranch egg ribbon at Shaab Mahmoud Red Sea Egypt Nudibranch egg ribbon at Malahi Red Sea Egypt Goniobranchus kuniei off the coast of Papua New Guinea NudibranchSee also EditSymposia and workshops on opisthobranchsReferences Edit Wagele H Klussmann Kolb A 2005 Opisthobranchia Mollusca Gastropoda more than just slimy slugs Shell reduction and its implications on defence and foraging Frontiers in Zoology 2 1 1 18 doi 10 1186 1742 9994 2 3 PMC 554092 PMID 15715915 Longman Pronunciation Dictionary 2nd edition ISBN 0 582 36467 1 Thompson T E 2009 Feeding in nudibranch larvae PDF Journal of the Marine Biological Association of the United Kingdom 38 2 239 248 doi 10 1017 S0025315400006044 S2CID 86275359 Turnbull John Spring 2016 The Nudibranch Creature Feature Nature New South Wales 60 3 16 17 Bronson Wilfrid Water People 1935 a b Ocean Portal 2017 A Collage of Nudibranch Colors Smithsonian National Museum of Natural History Retrieved 17 April 2018 a b c Nudibranchs Archived 2013 08 14 at the Wayback Machine Fishermen Scuba Ekimova I T Korshunova D Schepetov T Neretina N Sanamyan A Martynov 2015 Integrative systematics of northern and Arctic nudibranchs of the genus Dendronotus Mollusca Gastropoda with descriptions of three new species Zoological Journal of the Linnean Society 173 4 e0192177 doi 10 1111 zoj 12214 Korshunova T K Lundin K Malmberg B Picton A Martynov 2018 First true brackish water nudibranch mollusc provides new insights for phylogeny and biogeography and reveals paedomorphosis driven evolution PLOS ONE 13 1 e0192177 Bibcode 2018PLoSO 1392177K doi 10 1371 journal pone 0192177 PMC 5851531 PMID 29538398 Discoveries of deep sea biomass and biodiversity using an ROV Monterey Bay Aquarium Research Institute Archived from the original on 9 October 2013 Retrieved 16 October 2013 Steinberg J E 1956 The pelagic nudibranch Cephalopyge trematoides Chun 1889 in New South Wales with a note on other species in this genus Proceedings of the Linnean Society of New South Wales 81 184 192 G M Mapstone amp M N Arai Siphonophora Cnidaria Hydrozoa of Canadian Pacific Waters p 33 The best documented predators of pelagic cnidarians from the phylum Mollusca are the neustonic nudibranchs and snails and the pelagic nudibranch Gosliner TM Valdes A Behrens DW 2015 Nudibranch and Sea Slug Identification Indo Pacific New World Publications Jacksonville Florida USA a b Thompson T E 1976 Biology of opisthobranch molluscs vol 1 207 pp 21 pls Ray Society no 151 Nudibranchs National Geographic Magazine Ngm nationalgeographic com 2013 04 25 Retrieved 2013 07 04 CHASE RONALD June 1 1974 The Electrophysiology of Photoreceptors in the Nudibranch Mollusc Tritonia Diomedia Journal of Experimental Biology 60 3 707 19 doi 10 1242 jeb 60 3 707 PMID 4847278 Dayrat B 2005 Advantages of naming species under the PhyloCode An example of how a new species of Discodorididae Mollusca Gastropoda Euthyneura Nudibranchia Doridina may be named PDF Marine Biology Research 1 3 216 232 doi 10 1080 17451000510019141 S2CID 53557429 Retrieved 2009 06 14 Edmunds M 1991 Does warning colouration occur in nudibranchs Malacologia 32 241 255 a b Pawlik JR et al 1988 Defensive chemicals of the Spanish Dancer nudibranch Hexabranchus sanguineus and its egg ribbons Macrolides derived from a sponge diet Journal of Experimental Marine Biology and Ecology 119 2 99 109 doi 10 1016 0022 0981 88 90225 0 Ritson Williams R Paul VJ 2007 Marine benthic invertebrates use multimodal cues for defence against reef fish Marine Ecology Progress Series 340 29 39 Bibcode 2007MEPS 340 29R doi 10 3354 meps340029 Frick K 2003 Predator Suites and Flabellinid Nudibranch Nematocyst Complements in the Gulf of Maine In SF Norton Ed Diving for Science 2003 Proceedings of the American Academy of Underwater Sciences 22nd Annual Scientific Diving Symposium Archived from the original on January 29 2009 Retrieved 2008 07 03 a href Template Cite journal html title Template Cite journal cite journal a CS1 maint unfit URL link Karuso P 1987 Chemical Ecology of the Nudibranchs In Scheuer PJ ed Bioorganic Marine Chemistry Vol 1 Springer Verlag pp 31 60 doi 10 1007 978 3 642 72726 9 2 ISBN 978 3 642 72728 3 a comprehensive review of the chemical ecology of the nudibranchs Pawlik JR 2012 Fattorusso E et al eds Antipredatory defensive roles of natural products from marine invertebrates Handbook of Marine Natural Products NY Springer Science pp 677 710 Gosliner T M 1987 Nudibranchs of Southern Africa ISBN 978 0 930118 13 6 Avila C Iken K Fontana A Cimino G 2000 09 05 Chemical ecology of the Antarctic nudibranch Bathydoris hodgsoni Eliot 1907 defensive role and origin of its natural products Journal of Experimental Marine Biology and Ecology 252 1 27 44 doi 10 1016 S0022 0981 00 00227 6 ISSN 0022 0981 PMID 10962063 Faulkner D J Ghiselin M T 1983 Chemical defence and evolutionary ecology of dorid nudibranchs and some other opisthobranch gastropods Marine Ecology Progress Series 13 295 301 Bibcode 1983MEPS 13 295F doi 10 3354 meps013295 Avila C Iken K Fontana A Cimino G 2000 09 05 Chemical ecology of the Antarctic nudibranch Bathydoris hodgsoni Eliot 1907 defensive role and origin of its natural products Journal of Experimental Marine Biology and Ecology 252 1 27 44 doi 10 1016 S0022 0981 00 00227 6 ISSN 0022 0981 PMID 10962063 Barsby T Linington R G Andersen R J 2002 De Novo terpenoid biosynthesis by the dendronotid nudibranch Melibe leonina Chemoecology 12 4 199 202 doi 10 1007 PL00012669 S2CID 35384332 Edmunds M 1968 Acid secretion in some species of Doridacea Mollusca Nudibranchia Proceedings of the Malacological Society of London 38 2 121 133 Archived from the original on 2013 04 15 P H Gosse Evenings at the Microscope 1884 edition 1 p57 a b Nudibranch Aquaticcommunity com Retrieved 2013 07 04 Klussmann Kolb A 2001 The Reproductive Systems of the Nudibranchia Gastropoda Opisthobranchia Comparative Histology and Ultrastructure of the Nidamental Glands with Aspects of Functional Morphology Zoologischer Anzeiger 240 2 119 136 doi 10 1078 0044 5231 00011 Diving with Nudibranchs Dive the World NC Folino 1997 The role of prey mobility in the population ecology of the nudibranch Cuthona nana Gastropoda Opisthobranchia PDF American Malacological Bulletin Archived from the original PDF on 2012 06 25 Retrieved 2013 03 07 a b Dominguez M Troncoso J S Garcia F J 2008 The family Aeolidiidae Gray 1827 Gastropoda Opisthobranchia from Brazil with a description of a new species belonging to the genus Berghia Trinchese 1877 Zoological Journal of the Linnean Society 153 2 349 368 doi 10 1111 j 1096 3642 2008 00390 x Rudman W B 1999 03 19 Favorinus tsuruganus Baba amp Abe 1964 In Sea Slug Forum Australian Museum a b Valdes A 2004 Phylogeography and phyloecology of dorid nudibranchs Mollusca Gastropoda Biological Journal of the Linnean Society 83 4 551 559 doi 10 1111 j 1095 8312 2004 00413 x Barnes H Powell H T 1954 Onchidoris fusca Muller A Predator of Barnacles Journal of Animal Ecology 23 2 361 363 doi 10 2307 1986 JSTOR 1986 Piper Ross 2007 Extraordinary Animals An Encyclopedia of Curious and Unusual Animals Greenwood Press a b Hans Bertsch Nudibranchs Marine slugs with verve Navanax inermis is the bane of all nudibranchs because it is one of the few known predators on this group of slugs Dorids mainly eat sponges bryozoans and tunicates whereas aeolids principally eat cnidarians Facts About Nudibranchs Marinelife about com 2011 11 10 Retrieved 2013 07 04 Dean Lewis J Prinsep Michele R 2017 The chemistry and chemical ecology of nudibranchs Natural Product Reports 34 12 1359 1390 doi 10 1039 C7NP00041C ISSN 0265 0568 PMID 29135002 Cella Kristen Carmona Leila Ekimova Irina Chichvarkhin Anton Schepetov Dimitry Gosliner Terrence M 2016 12 15 A Radical Solution The Phylogeny of the Nudibranch Family Fionidae PLOS ONE 11 12 e0167800 Bibcode 2016PLoSO 1167800C doi 10 1371 journal pone 0167800 ISSN 1932 6203 PMC 5158052 PMID 27977703 Taxonomy of the Gastropoda Bouchet amp Rocroi 2005 Poppe Guido T Tagaro Sheila P February 23 2006 The New Classification of Gastropods according to Bouchet amp Rocroi 2005 PDF Visaya Archived from the original PDF on September 27 2007 Jorger K M Stoger I Kano Y et al 2010 On the origin of Acochlidia and other enigmatic euthyneuran gastropods with implications for the systematics of Heterobranchia BMC Evolutionary Biology 10 323 doi 10 1186 1471 2148 10 323 PMC 3087543 PMID 20973994 At the basis of the Euthyneura the Nudipleura split off Thiele J 1931 Handbuch der systematischen Weichtierkunde II Verlag von Gustav Fischer Jena Germany Milne Edwards H 1848 Note sur la classification naturelle chez Mollusques Gasteropodes Annales des Sciences Naturelles series 3 9 102 112 Wagele H amp Willan R C September 2000 Phylogeny of the Nudibranchia Zoological Journal of the Linnean Society 130 1 83 181 doi 10 1111 j 1096 3642 2000 tb02196 x Schrodl M Wagele H amp Willan R C 2001 Taxonomic Redescription of the Doridoxidae Gastropoda Opisthobranchia an Enigmatic Family of Deep Water Nudibranchs with Discussion of Basal Nudibranch Phylogeny Zoologischer Anzeiger 240 1 83 97 doi 10 1078 0044 5231 00008 Further reading EditThompson T E 1976 Biology of opisthobranch molluscs Vol 1 207 pp 21 pls Ray Society no 151 Thompson T E amp G H Brown 1984 Biology of opisthobranch molluscs Vol 2 229 pp 41 pls Ray Society no 156 McDonald Gary R 7 July 2021 Institute of Marine Sciences Bibliographia Nudibranchia 3rd online Edition A listing by Author of publications on nudibranchs McDonald Gary R 7 July 2021 Institute of Marine Sciences Nudibranch Systematic Index 3rd online Edition An index of names given to nudibranchs and their subsequent use referenced to Bibliographia Nudibranchia McDonald Gary R amp J W Nybakken November 5 2014 List of the Worldwide Food Habits of Nudibranchs Coleman Neville 2008 Nudibranchs Encyclopedia Catalogue of Asia Indo Pacific Sea Slugs Neville Coleman s Underwater Geographic ISBN 0 947325 41 7External links Edit Wikispecies has information related to Nudibranchia Wikimedia Commons has media related to Nudibranchia Sea Slug Forum by William B Rudman Nudibranchs of the British Isles OPK Opistobranquis Iberian and Mediterranean Opisthobranchs Mediterranean slug site actually a misnomer Worldwide coverage The Slug Site Michael D Miller 2002 2014 The Okinawa Slug Site Images information and identification of Nudibranchs Nudibranch Photos by Mick Tait Nudibranchs in their natural environment Scuba Diving Narooma NSW offline 26 Nov 2014 Nudi Pixel Online resource for nudibranchs and sea slugs identification using photographs Archived 2011 05 27 at the Wayback Machine Various nudibranch species from Indonesia Philippines and Thailand Nudibranch gallery Sergey Parinov offline 26 Nov 2014 Opisthobranch Newsletter Bibliography and portal to opisthobranch nudibranch amp seaslug information Scottish Nudibranchs Online resource for identification of species found in Scottish waters National Geographic Nudibranch Photo Gallery Sea Slugs of Hawaii Slug City Molluscs Brain amp Behavior from the University of Illinois at Urbana ChampaignVideos Edit Attack of the Sea Slugs at YouTube The Lynx Nudibranch HD clip of Phidiana lynceus carefully consuming a hydroid Myrionema amboinense Slug City Molluscs Brain amp Behavior many videos of nudibranchs from the University of Illinois at Urbana Champaign Retrieved from https en wikipedia org w index php title Nudibranch amp oldid 1136167854, wikipedia, wiki, book, books, library,

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