Difference between revisions of "Nudibranch" - New World Encyclopedia

Latest revision as of 17:15, 14 October 2021

Nudibranch

Spanish shawl, Flabellina iodinea

Scientific classification

Kingdom:	Animalia
Phylum:	Mollusca
Class:	Gastropoda
Subclass:	Orthogastropoda
Superorder:	Heterobranchia
Order:	Opisthobranchia
Suborder:	Nudibranchia

Infraorders

Anthobranchia
Cladobranchia

See text for superfamilies

A nudibranch is any of the soft-bodied, shell-less marine gastropods comprising the mollusk taxon (order or suborder) Nudibranchia. Found on the bottom of oceans throughout the world, from reefs and sandy shallows to seabeds a mile deep, there are more than 3,000 known species of nudibranchs. The word "nudibranch," which comes from the Latin nudus for "naked," and the Greek brankhia for gills, describes their feathery gills and horns that many have on their dorsal sides (NGS 2008). Nudibranchs sometimes are called sea slugs, although this term technically applies to a more inclusive taxon of marine gastropod mollusks, Heterobranchia (which includes, but is not exclusive to, the nudibranchs).

Nudibranchs are known for their often extraordinary colors, striking forms, and intricate patterns. Such brilliant hues and fascinating shapes have added greatly to the wonder of nature. However, nudibranchs also provide important ecological functions, playing an important role in marine food chains, both as predator (corals, sponges, anemones, barnacles, fish) and prey (fish, sea spiders, turtles, sea stars)—the latter role despite protective mechanisms ranging from poisons, to camouflage, to stinging cells.

Overview and description

Hermissenda crassicornis in a tide pool in Moss Beach, California

Nudibranchs are members of the gastropod class (Gastropoda) of mollusks. As gastropods, their body plan involves a torsion or twisting during larval development whereby the visceral mass twists 180 degrees in relation to the head. Gastropoda (meaning "stomach-foot") is typified by a large, ventral, muscular foot for locomotion, and a distinct head that has eyes and sensory tentacles.

The taxon that nudibranchs comprise, Nudibranchia, generally is placed either at the suborder level, under the order Opisthobranchia of the superorder Heterobranchia and subclass Orthogastropoda, or at the order level under the subclass Opisthobranchia (ITIS 2004). While often casually called "sea slugs," there are numerous other kinds of sea slugs belonging to several taxonomic groups that are not very closely related to nudibranchs. A fair number of these other sea slugs are colorful and thus are even more easily confused with nudibranchs.

The body forms of nudibranchs vary enormously, but because they are opisthobranchs, unlike most other gastropods they are bilaterally symmetrical because they have undergone secondary detorsion. The adult form is without a shell or operculum (a bony or horny plate covering the opening of the shell, when the body is withdrawn). Often oblong in shape, some are thick and other flattened, and some are short and others long (NGS 2008). They vary in adult size from six millimeters (0.25 inches) to 31 centimeters (12 inches) in length (NGS 2008). Nudibranchs have cephalic (head) tentacles and oral tentacles, which are sensitive to touch, taste, and smell. Two club-shaped rhinophores (highly sensitive head-mounted sensory appendages or tentacles) detect odors.

The name nudibranch is appropriate, since the dorids (infraclass Anthobranchia) breathe through a branchial plume of bushy extremities on their back, rather than using gills. By contrast, on the back of the aeolids in infraclass Cladobranchia, there are brightly colored sets of tentacles called cerata.

Nudibranchs occur in marine environments worldwide, in all the world's oceans, and can be found depths nearly a mile down, but are most abundant, and reach their greatest size and variation, in shallow, tropical waters (NGS 2008).

Protective mechanisms: Colors, camouflage, and stinging cells

Hypselodoris bullocki
Sipadan, Malaysia

While they lack shells, nudibranchs have developed other defense mechanisms: poisons, stinging cells, and camouflage.

Some nudibranchs have toxic secretions, mostly derived from the poisons of their prey but some have poisons of their own making (Holland 2008). For example, some species consume toxic sponges and alter and store the poisonous compounds and secrete them from skin cells or glands as needed (Holland 2008).

Nembrotha chamberlaini
Sipadan, Malaysia

Some utilize stinging cells, ingested from anemones, fire corals, and hydroids, and employ these along their own extremities (Holland 2008). Nudibranchs that feed on hydroids can store the hydroid's nematocysts (stinging cells) in the dorsal body wall, the cerata (Frick 2003). The nematocysts can move through the alimentary tract without harming the nudibranch. Once further into the organism, the cells are brought to specific placements on the creature's hind body via intestinal protuberances. It is not yet clear how the nudibranches can protect themselves from the hydroids and their nematocysts, but special cells with large vacuoles probably play an important role.

The vivid color of various nudibranchs helps to warn predators of such defenses. Among the nudibranchs can be found the most colorful creatures on earth. The intense and bright coloring, such as especially seen on Chromodorids, warns that they are distasteful or poisonous (aposematic coloration).

Camouflage also is used. The anatomy and color of nudibranchs may resemble the texture and color of the surrounding plants, sponges, and substrate, allowing them to be difficult to discern. Some may mimic toxic nudibranches.

The tough-skin and abrasive quality of nudibranchs also may be a detriment (Holland 2008). Another way of protection is the release of a sour liquid from the skin. Once the specimen is physically irritated or touched by another creature, it will release the slime automatically.

Life habits

Acanthodoris lutea laying eggs

Feeding and predation

Nudibranchs are carnivorous, feeding on sponges, hydroids, bryozoans, coral, barnacles, eggs, tunicates, anemones, or small fish. Some eat other nudibranches, including members of their own species. Some graze on algae. Some nudibranchs derive nutrition from ingested photosynthetic algae (Holland 2008).

Despite their protective mechanisms, nudibranchs have a number of predators, including certain species of fish, sea spiders, turtles, crabs, and sea stars, as well as some people (such as Chileans and islanders from off Alaska and Russia) who consume them after removing the toxic organs (Holland 2008).

Reproduction

Nudibranch eggs in Moss Beach, California

Nudibranchs are simultaneous hermaphroditic, and thus have a set of reproductive organs for both genders. They can rarely fertilize themselves, but can mate with any other mature members of their species (NGS 2008).

Nudibranchs typically deposit their eggs within a gelatinous spiral (Klussmann-Kolb 2001). Some masses (coils, ribbons, or tangled clumbs) may reach up to two million at a time (Holland 2008).

Taxonomy

"Nudibranchia," from Ernst Haeckel's Artforms of Nature, 1904.

The taxonomy of the Nudibranchia is still under investigation and is subject to frequent revision. Many taxonomists in the past treated the Nudibranchia as an order, based on the authoritative work of Johannes Thiele (1931), who built on the concepts of Henri Milne-Edwards (1848). Newer insights derived from morphological data and gene-sequence research have supported the basic taxonomic divisions. On the basis of investigation of 18S rDNA sequence data, there is strong evidence for support of the monophyly of the Nudibranchia and its two major groups, the Anthobranchia/Doridoidea and Cladobranchia.

Competing taxonomies remain, with Nudibranchia variously considered an order or a suborder, and diverse classes and subclasses recognized; for example, Opisthobranchia may variously be considered an order or a subclass. Ponder and Lindberg (1997) recognize two subclasses within Gastropoda (Orthogastropoda and Eogastropoda), with Nudibranchia placed within the subclass Orthogastropoda. The Integrated Taxonomic Information System recognizes Nudibranchia as an order in the subclass Opisthobranchia (ITIS 2004).

A proposed classification of Nudibranchia, recognizing the subclass Orthogastropoda, Nudibranchia as a suborder, and two major infraorders (Anthobranchia/Doridoidea and Cladobranchia) is as follows:

Class Gastropoda (Cuvier, 1797)

Subclass Orthogastropoda (Ponder & Lindberg, 1996) (earlier Prosobranchia, Opisthobranchia)

Superorder Heterobranchia J.E. Gray, 1840

Order Opisthobranchia Milne-Edwards, 1848

Suborder Nudibranchia Blainville, 1814 (nudibranchs)

Infraorder Anthobranchia Férussac, 1819 (dorids)

Superfamily Doridoidea Rafinesque, 1815

Superfamily Doridoxoidea Bergh, 1900

Superfamily Onchidoridoidea Alder & Hancock, 1845

Superfamily Polyceroidea Alder & Hancock, 1845

Infraorder Cladobranchia Willan & Morton, 1984 (aeolids)

Superfamily Dendronotoidea Allman, 1845

Superfamily Arminoidea Rafinesque, 1814

Superfamily Metarminoidea Odhner in Franc, 1968

Superfamily Aeolidioidea J.E. Gray, 1827

The dorids (infraorder Anthobranchia) have the following characteristics: The branchial plume forms a cluster on the posterior part of the neck, around the eyes. Fringes on the mantle do not contain any intestines.

The aeolids (infraorder Cladobranchia) have the following characteristics: Instead of the branchial plume, they have cerata. They lack a mantle. Only species of the Cladobranchia are reported to house zooxanthellae.

Schrödl et al. (2001) have offered other revisions in the taxonomy of the Nudibranchia. Here they are divided into two major clades:

Anthobranchia (= Bathydoridoidea + Doridoidea)
Dexiarchia nom. nov. (= Doridoxoidea + Dendronotoidea + Aeolidoidea + “Arminoidea”).

Images

Red nudibranch
Hurghada, Egypt
Pyjamas nudibranch
Hurghada, Egypt
Godiva quadricolor
laying eggs, Swan River, Western Australia
Collingwood's chromodoris
Papua New Guinea
Chromodoris lochi
Papua New Guinea
Two clown nudibranchs
New Zealand
Notodoris minor on the ribbon reefs off Cooktown, Queensland, Australia
Chromodoris willani
Lembeh Straits, Indonesia
Nembrotha kubaryana
Lembeh Straits, Indonesia
Chromodoris annae
Lembeh Straits, Indonesia
Pteraeolidia ianthina
Lembeh Straits, Indonesia
Clown nudibranch
Triopha catalinae, Northern California
Acanthodoris lutea
Northern California

References
ISBN links support NWE through referral fees

Frick, K. 2003. Predator suites and flabellinid nudibranch nematocyst complements in the Gulf of Maine Proceedings of the 22nd Annual Scientific Diving Symposium, American Academy of Underwater Sciences. Greenville, North Carolina. Retrieved October 14, 2021.
Integrated Taxonomic Information System (ITIS). 2004. Nudibranchia Blainville, 1814 ITIS Taxonomic Serial No.: 78156. Retrieved October 14, 2021.
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. Retrieved October 14, 2021.
National Geographic Society (NGS). 2008. Nudibranch National Geographic Society. Retrieved October 14, 2021.
Ponder, W. F., and D. R. Lindberg. 1997. “Towards a phylogeny of gastropod molluscs: An analysis using morphological characters.” Zoological Journal of the Linnean Society 119: 83-2651.
Schrödl, M., H. Wägele, and R. C. Willan. 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. Retrieved October 14, 2021.
Wägele, H., and R. C. Willan. 2000. Phylogeny of the Nudibranchia. Zoological Journal of the Linnean Society 1(1): 83–181.

Credits

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+{{Copyedited}}{{Images OK}}{{Approved}}
 {{Taxobox
 | name = Nudibranch
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 See text for superfamilies
 }}
+A '''nudibranch''' is any of the soft-bodied, shell-less [[marine (ocean)|marine]] [[gastropod]]s comprising the [[mollusk]] taxon (order or suborder) '''Nudibranchia'''. Found on the bottom of [[ocean]]s throughout the world, from reefs and sandy shallows to seabeds a mile deep, there are more than 3,000 known [[species]] of nudibranchs. The word "nudibranch," which comes from the [[Latin]] ''nudus'' for "naked," and the [[Greek language|Greek]] ''brankhia'' for [[gill]]s, describes their feathery gills and horns that many have on their dorsal sides (NGS 2008). Nudibranchs sometimes are called '''sea slugs''', although this term technically applies to a more inclusive taxon of marine gastropod mollusks, Heterobranchia (which includes, but is not exclusive to, the nudibranchs).
-A '''nudibranch''' is any of the soft-bodied, shell-less [[marine (ocean)|marine]] [[gastropod]]s comprising the [[mollusk]] taxon (order or suborder) '''Nudibranchia'''. Found on the bottom of oceans throughout the world, from reefs and sandy shallows to seabeds a mile deep, there are more than 3,000 known species of nudibranchs (Holland 2008). The word "nudibranch," which comes from the [[Latin]] ''nudus'' for "naked," and the [[Greek language|Greek]] ''brankhia'' for [[gill]]s, describes their feathery gills and horns that many have on their dorsal sides (NGS 2008). Nudibranchs sometimes are called '''sea slugs''', although this term technically applies to a more inclusive taxon of marine gastropod mollusks, Heterobranchia, which includes the nudibranchs.
+{{toc}}
+Nudibranchs are known for their often extraordinary [[color]]s, striking forms, and intricate patterns. Such brilliant hues and fascinating shapes have added greatly to the wonder of nature. However, nudibranchs also provide important ecological functions, playing an important role in marine [[food chain]]s, both as predator ([[coral]]s, [[sponge]]s, [[anemone]]s, [[barnacle]]s, [[fish]]) and prey (fish, [[sea spider]]s, [[turtle]]s, [[sea star]]s)&mdash;the latter role despite protective mechanisms ranging from poisons, to [[camouflage]], to stinging cells.
-Nudibranchs are known for their often extraordinary colors, striking forms, and intricate patterns. Such brilliant hues and fascinating shapes have added greatly to the wonder of nature. However, nudibranchs also provide important ecological functions, playing an important role in marine [[food chain]]s, both as predator ([[coral]]s, [[sponge]]s, [[anemone]]s, [[barnacle]]s, [[fish]]) and prey (fish, [[sea spider]]s, [[turtle]]s, [[sea star]]s)&mdash;the latter role despite protective mechanisms ranging from poisons, to [[camouflage]], to stinging cells.
 ==Overview and description==
+[[Image:Nudi from tidepool.jpg|thumb|right|350px|''[[Hermissenda crassicornis]]'' in a tide pool in [[Moss Beach, California]]]]
+Nudibranchs are members of the [[gastropod]] class (Gastropoda) of mollusks. As gastropods, their body plan involves a ''torsion'' or twisting during larval development whereby the visceral mass twists 180 degrees in relation to the head. Gastropoda (meaning "stomach-foot") is typified by a large, ventral, muscular foot for locomotion, and a distinct head that has eyes and sensory tentacles.
-Nudibranchs are members of the [[gastropod]] class (Gastropoda) of mollusks. As gastropods, their body plan involves a ''torsion'' or twisting during larval development whereby the visceral mass twists 180 degrees in relation to the head, bringing the mantle cavity to the anterior of the animal. Gastropoda (meaning "stomach-foot") is typified by a large, ventral, muscular foot for locomotion, and a distinct head that has eyes and sensory tentacles.
+The [[taxon]] that nudibranchs comprise, ''Nudibranchia'', generally is placed either at the suborder level, under the order Opisthobranchia of the superorder Heterobranchia and subclass Orthogastropoda, or at the order level under the subclass Opisthobranchia (ITIS 2004). While often casually called "sea [[slug]]s," there are numerous other kinds of sea slugs belonging to several [[taxonomic]] groups that are not very closely related to nudibranchs. A fair number of these other sea slugs are colorful and thus are even more easily confused with nudibranchs.
-The taxon that nudibranchs comprise, '''Nudibranchia''', generally is placed either at the suborder level, under the
+The body forms of nudibranchs vary enormously, but because they are opisthobranchs, unlike most other gastropods they are [[bilateral symmetry|bilaterally symmetrical]] because they have undergone secondary [[torsion (gastropod)|detorsion]]. The adult form is without a shell or [[operculum (gastropod)|operculum]] (a bony or horny plate covering the opening of the shell, when the body is withdrawn). Often oblong in shape, some are thick and other flattened, and some are short and others long (NGS 2008). They vary in adult size from six millimeters (0.25 inches) to 31 centimeters (12 inches) in length (NGS 2008). Nudibranchs have cephalic (head) tentacles and oral tentacles, which are sensitive to touch, taste, and smell. Two club-shaped [[rhinophore]]s (highly sensitive head-mounted sensory appendages or tentacles) detect odors.
-order Opisthobranchia of the superorder Heterobranchia and subclass Orthogastropoda, or at the order level under the subclass Opisthobranchia (ITIS 2004). While often casually called "sea [[slug]]s," there are numerous other kinds of sea slugs belonging to several [[taxonomic]] groups that are not very closely related to nudibranchs. A fair number of these other sea slugs are colorful and thus are even more easily confused with nudibranchs.
-The body forms of nudibranchs vary enormously, but because they are opisthobranchs, unlike most other gastropods they are [[bilateral symmetry|bilaterally symmetrical]] because they have undergone secondary [[torsion (gastropod)|detorsion]].
+The name nudibranch is appropriate, since the dorids (infraclass [[Anthobranchia]]) breathe through a branchial plume of bushy extremities on their back, rather than using gills. By contrast, on the back of the aeolids in infraclass [[Cladobranchia]], there are brightly colored sets of tentacles called [[cerata]].
-They lack a [[mantle (mollusc)|mantle]] cavity.
+Nudibranchs occur in marine environments worldwide, in all the world's oceans, and can be found depths nearly a mile down, but are most abundant, and reach their greatest size and variation, in shallow, tropical waters (NGS 2008).
-They vary in adult size from 20 to 600 [[millimetre|mm]].
+===Protective mechanisms: Colors, camouflage, and stinging cells===
+[[Image:Hypselodoris bullocki.JPG|thumb|right|400px|''[[Hypselodoris bullocki]]''<br/>Sipadan, [[Malaysia]]]]
+While they lack shells, nudibranchs have developed other defense mechanisms: poisons, stinging cells, and camouflage.
-The adult form is without a shell or [[operculum (gastropod)|operculum]] (a bony or horny plate covering the opening of the shell, when the body is withdrawn).
+Some nudibranchs have toxic secretions, mostly derived from the poisons of their prey but some have poisons of their own making (Holland 2008). For example, some species consume toxic sponges and alter and store the poisonous compounds and secrete them from skin cells or glands as needed (Holland 2008).
-The name nudibranch is appropriate, since the dorids (infraclass [[Anthobranchia]]) breathe through a branchial plume of bushy extremities on their back, rather than using gills. By contrast, on the back of the aeolids in infraclass [[Cladobranchia]] there are brightly colored sets of tentacles called [[cerata]].
+[[Image:Nebrotha Chamberlaini.jpg|thumb|right|400px|''[[Nembrotha chamberlaini]]''<br/>Sipadan, Malaysia]]
+Some utilize stinging cells, ingested from [[anemone]]s, fire corals, and [[hydroid]]s, and employ these along their own extremities (Holland 2008). Nudibranchs that feed on hydroids can store the hydroid's [[nematocyst]]s (stinging cells) in the [[dorsum (biology)|dorsal]] body wall, the [[cerata]] (Frick 2003). The nematocysts can move through the [[alimentary tract]] without harming the nudibranch. Once further into the organism, the cells are brought to specific placements on the creature's hind body via intestinal protuberances. It is not yet clear how the nudibranches can protect themselves from the hydroids and their nematocysts, but special cells with large [[vacuole]]s probably play an important role.
-Nudibranchs have cephalic (head) tentacles, which are sensitive to touch, taste, and smell. Club-shaped [[rhinophore]]s detect odors.
+The vivid [[color]] of various nudibranchs helps to warn predators of such defenses. Among the nudibranchs can be found the most colorful creatures on earth. The intense and bright coloring, such as especially seen on [[Chromodoris quadricolor|Chromodorids]], warns that they are distasteful or poisonous ([[Aposematism|aposematic coloration]]).
+[[Camouflage]] also is used. The anatomy and color of nudibranchs may resemble the texture and color of the surrounding plants, sponges, and substrate, allowing them to be difficult to discern. Some may mimic toxic nudibranches.
+The tough-skin and abrasive quality of nudibranchs also may be a detriment (Holland 2008). Another way of protection is the release of a sour liquid from the skin. Once the specimen is physically irritated or touched by another creature, it will release the slime automatically.
-==Distribution==
+==Life habits==
-Nudibranchs occur worldwide.
+[[Image:Editing Image-Acanthodoris lutea laying eggs 2.jpg|thumb|400px|right|''[[Acanthodoris lutea]]'' laying eggs]]
+===Feeding and predation===
+Nudibranchs are [[carnivore|carnivorous]], feeding on [[sea sponge|sponge]]s, [[hydroid]]s, [[bryozoa]]ns, [[coral]], [[barnacle]]s, [[egg]]s, [[tunicate]]s, [[anemone]]s, or small [[fish]]. Some eat other nudibranches, including members of their own species. Some graze on [[algae]]. Some nudibranchs derive nutrition from ingested photosynthetic algae (Holland 2008).
-==Habitat==
+Despite their protective mechanisms, nudibranchs have a number of predators, including certain species of fish, sea spiders, [[turtle]]s, [[crab]]s, and sea stars, as well as some [[human|people]] (such as Chileans and islanders from off Alaska and Russia) who consume them after removing the toxic organs (Holland 2008).
-Nudibranchs live at virtually all depths of salt water, but reach their greatest size and variation in warm, shallow waters.
-[[Image:Nudi from tidepool.jpg|thumb|right|200px|''[[Hermissenda crassicornis]]'' in a tide pool in [[Moss Beach, California]]]]
-==Life habits==
 ===Reproduction===
-[[Image:Eggs of nudibranch.JPG|thumb|Nudibranch eggs in Moss Beach, California]]
+[[Image:Eggs of nudibranch.JPG|thumb|400px|Nudibranch eggs in Moss Beach, California]]
-[[Image:Editing Image-Acanthodoris lutea laying eggs 2.jpg|thumb|200px|left|''[[Acanthodoris lutea]]'' laying eggs]]
+Nudibranchs are simultaneous [[hermaphrodite|hermaphroditic]], and thus have a set of reproductive organs for both genders. They can rarely fertilize themselves, but can mate with any other mature members of their species (NGS 2008).
-Nudibranchs are [[hermaphrodite|hermaphroditic]], and thus have a set of reproductive organs for both genders, but they can rarely fertilize themselves.
-Nudibranchs typically deposit their eggs within a gelatinous spiral. <ref name="Reproductive systems ">{{cite journal | author=Klussmann-Kolb A| title= The Reproductive Systems of the Nudibranchia (Gastropoda, Opisthobranchia): Comparative Histology and Ultrastructure of the Nidamental Glands with Aspects of Functional Morphology| journal=Zoologischer Anzeiger| year=2001| volume=240| issue=2| url=http://www.ingentaconnect.com/content/urban/351/2001/00000240/00000002/art00011| pages=119–136| doi= 10.1078/0044-5231-00011}}</ref>
+Nudibranchs typically deposit their eggs within a gelatinous spiral (Klussmann-Kolb 2001). Some masses (coils, ribbons, or tangled clumbs) may reach up to two million at a time (Holland 2008).
-===Feeding===
+==Taxonomy==
-Nudibranchs are [[carnivore|carnivorous]]. Some feed on [[sea sponge|sponge]]s, others on [[hydroid]]s, others on [[bryozoa]]ns, and some are [[cannibal]]s, eating other sea slugs, or, on some occasions, members of their own species. There is also a group that feeds on [[tunicate]]s and [[barnacle]]s.
+[[Image:Haeckel Nudibranchia.jpg|300px|thumb|"Nudibranchia," from [[Ernst Haeckel]]'s ''Artforms of Nature'', 1904.]]
-They even sometimes feed on anemones.
+The [[taxonomy]] of the Nudibranchia is still under investigation and is subject to frequent revision. Many taxonomists in the past treated the Nudibranchia as an order, based on the authoritative work of [[Johannes Thiele]] (1931), who built on the concepts of [[Henri Milne-Edwards]] (1848). Newer insights derived from morphological data and gene-sequence research have supported the basic taxonomic divisions. On the basis of investigation of 18S rDNA sequence data, there is strong evidence for support of the [[monophyly]] of the Nudibranchia and its two major groups, the Anthobranchia/Doridoidea and Cladobranchia.
-==Colors and camouflage==
+Competing taxonomies remain, with Nudibranchia variously considered an order or a suborder, and diverse classes and subclasses recognized; for example, Opisthobranchia may variously be considered an order or a subclass. Ponder and Lindberg (1997) recognize two subclasses within Gastropoda (Orthogastropoda and Eogastropoda), with Nudibranchia placed within the subclass ''Orthogastropoda''. The Integrated Taxonomic Information System recognizes Nudibranchia as an order in the subclass Opisthobranchia (ITIS 2004).
-Among this group can be found the most colorful creatures on earth. In the course of evolution, sea slugs have lost their shell because they have developed other defense mechanisms. Their anatomy may resemble the texture and color of the surrounding plants, allowing them to [[camouflage]] (''cryptic behavior''). Others, as seen especially well on [[Chromodoris quadricolor|Chromodorids]], have an intense and bright coloring, which warns that they are distasteful or poisonous ([[Aposematism|aposematic coloration]]).
-Nudibranchs that feed on hydroids can store the hydroid's [[nematocyst]]s (stinging cells) in the [[dorsum (biology)|dorsal]] body wall, the [[cerata]].<ref>{{cite journal |author=Frick, K |title=Predator Suites and Flabellinid Nudibranch Nematocyst Complements in the Gulf of Maine. |journal=In: SF Norton (ed).  Diving for Science...2003. |volume=Proceedings of the American Academy of Underwater Sciences |issue=22nd Annual Scientific Diving Symposium |date=2003 |url=http://archive.rubicon-foundation.org/4744 |accessdate=2008-07-03 }}</ref> The nematocysts wander through the [[alimentary tract]] without harming the nudibranch. Once further into the organ, the cells are brought to the specific placements on the creature's hind body via intestinal protuberances. Nudibranches can protect themselves from the hydroids and their nematocysts. It is not yet clear how, but special cells with large [[vacuole]]s probably play an important role. They can also take in plants' chloroplasts (plant cell organelles used for photosynthesis) and use them to make food for themselves.
+A proposed classification of Nudibranchia, recognizing the subclass Orthogastropoda, Nudibranchia as a suborder, and two major infraorders (Anthobranchia/Doridoidea and Cladobranchia) is as follows:
-Another way of protection is the release of a sour liquid from the skin. Once the specimen is physically irritated or touched by another creature, it will release the slime automatically.
+'''Class Gastropoda''' ''([[Georges Cuvier|Cuvier]], 1797)''<br />
+:'''Subclass [[Orthogastropoda]]* ''' (Ponder & Lindberg, 1996) (earlier [[Prosobranchia]]*, [[Opisthobranchia]]*)<br />
+::Superorder Heterobranchia [[John Edward Gray|J.E. Gray]]*, 1840
+:::Order Opisthobranchia [[Alphonse Milne-Edwards|Milne-Edwards]]*, 1848
+::::Suborder Nudibranchia Blainville, 1814 (nudibranchs)
+:::::Infraorder Anthobranchia Férussac, 1819 ('''dorids''')
+::::::Superfamily Doridoidea [[Constantine Samuel Rafinesque|Rafinesque]]*, 1815
+::::::Superfamily Doridoxoidea Bergh, 1900
+::::::Superfamily Onchidoridoidea Alder & Hancock, 1845
+::::::Superfamily Polyceroidea Alder & Hancock, 1845
+:::::Infraorder Cladobranchia Willan & Morton, 1984 ('''aeolids''')
+::::::Superfamily Dendronotoidea Allman, 1845
+::::::Superfamily Arminoidea [[Constantine Samuel Rafinesque|Rafinesque]]*, 1814
+::::::Superfamily Metarminoidea Odhner in Franc, 1968
+::::::Superfamily Aeolidioidea [[John Edward Gray|J.E. Gray]]*, 1827
-==Taxonomy==
+The dorids (infraorder Anthobranchia) have the following characteristics: The branchial plume forms a cluster on the posterior part of the neck, around the eyes. Fringes on the mantle do not contain any intestines.
-[[Image:Haeckel Nudibranchia.jpg|250px|thumb|"Nudibranchia", from [[Ernst Haeckel]]'s ''Artforms of Nature'', 1904.]]
-The [[taxonomy]] of the Nudibranchia is still under investigation. Many taxonomists in the past treated the Nudibranchia as an order, based on the authoritative work of [[Johannes Thiele]] (1931), who built on the concepts of [[Henri Milne-Edwards]] (1848). Newer insights derived from morphological data and gene-sequence research, have confirmed these ideas. On the basis of investigation of 18S rDNA sequence data, there is strong evidence for support of the [[monophyly]] of the Nudibranchia and its two major groups, the Anthobranchia/Doridoidea and Cladobranchia.
-*Infraorder [[Anthobranchia]]  [[André Étienne d'Audebert de Férussac|Férussac]], 1819 ('''dorids''')
+The aeolids (infraorder Cladobranchia) have the following characteristics: Instead of the branchial plume, they have [[cerata]]. They lack a mantle. Only species of the Cladobranchia are reported to house [[zooxanthella]]e.
-**Superfamily [[Doridoidea]] [[Constantine Samuel Rafinesque|Rafinesque]], 1815
-**Superfamily [[Doridoxoidea]] Bergh, 1900
-**Superfamily [[Onchidoridoidea]] Alder & Hancock, 1845
-**Superfamily [[Polyceroidea]] Alder & Hancock, 1845
-*Infraorder [[Cladobranchia]] Willan & Morton, 1984 ('''aeolids''')
-**Superfamily [[Aeolidioidea]] [[John Edward Gray|J. E. Gray]], 1827
-**Superfamily [[Arminoidea]] [[Constantine Samuel Rafinesque|Rafinesque]], 1814
-**Superfamily [[Dendronotoidea]] Allman, 1845
-**Superfamily [[Metarminoidea]] Odhner in  Franc, 1968
-A study published in May 2001, has again revised the taxonomy of the Nudibranchia <ref name="A new taxonomic redescription of the Doridoxidae">{{cite journal | author=Schrödl M.; Wägele H.2 Willan R.C.| title=Taxonomic Redescription of the Doridoxidae(Gastropoda: Opisthobranchia), an Enigmatic Family of Deep Water Nudibranchs, with Discussion of Basal Nudibranch Phylogeny| journal=Zoologischer Anzeiger,| year=2001| volume=240| issue=1| page=83-97| url=http://www.ingentaconnect.com/content/urban/351/2001/00000240/00000001/art00008 | pages=83 | doi=10.1078/0044-5231-00008}}</ref>. They are thus divided into two major clades:
+Schrödl et al. (2001) have offered other revisions in the taxonomy of the Nudibranchia. Here they are divided into two major clades:
 * Anthobranchia (= Bathydoridoidea + Doridoidea)
 * Dexiarchia nom. nov. (= Doridoxoidea + Dendronotoidea + Aeolidoidea + “Arminoidea”).
-The dorids (infraorder Anthobranchia) have the following characteristics: the branchial plume forms a cluster on the posterior part of the neck, around the eyes. Fringes on the mantle do not contain any intestines.
-The aeolids (infraorder Cladobranchia) have the following characteristics: Instead of the branchial plume, they have [[cerata]]. They lack a mantle. Only species of the Cladobranchia are reported to house [[zooxanthella]]e.
 ==Images==
 <gallery>
-Image:Red nudibranch from Hurghada.jpg|[[Red nudibranch]]<br>[[Hurghada]], [[Egypt]]
+Image:Red nudibranch from Hurghada.jpg|[[Red nudibranch]]<br/>[[Hurghada]], [[Egypt]]
-Image:Pyjamas nudibranch from Hurghada.jpg|[[Pyjamas nudibranch]]<br>[[Hurghada]], [[Egypt]]
+Image:Pyjamas nudibranch from Hurghada.jpg|[[Pyjamas nudibranch]]<br/>[[Hurghada]], [[Egypt]]
-Image:Godiva Quadicolor Wes Australia.jpg|''[[Godiva quadricolor]]''<br>laying eggs, [[Swan River (Western Australia)|Swan River]], [[Western Australia]]
+Image:Godiva Quadicolor Wes Australia.jpg|''[[Godiva quadricolor]]''<br/>laying eggs, [[Swan River (Western Australia)|Swan River]], [[Western Australia]]
-Image:Hypselodoris bullocki.JPG|''[[Hypselodoris bullocki]]''<br>[[Sipadan]], [[Malaysia]]
+File:Collingwood's Chromodoris.JPG|[[Collingwood's chromodoris]]<br/>[[Papua New Guinea]]
-Image:Nebrotha Chamberlaini.jpg|''[[Nembrotha chamberlaini]]''<br>[[Sipadan]], [[Malaysia]]
+Image:Chromodoris_lochi.jpg|''[[Chromodoris lochi]]''<br/>[[Papua New Guinea]]
-Image:Collingwood%27s_Chromodoris.JPG|[[Collingwood's chromodoris]]<br>[[Papua New Guinea]]
+Image:Two_Clown_Nudibranchs.JPG|Two [[clown nudibranch]]s<br/>[[New Zealand]]
-Image:Chromodoris_lochi.jpg|''[[Chromodoris lochi]]''<br>[[Papua New Guinea]]
-Image:Two_Clown_Nudibranchs.JPG|Two [[clown nudibranch]]s<br>[[New Zealand]]
 Image:Notodoris_minor.jpg|''[[Notodoris minor]]'' on the ribbon reefs off [[Cooktown, Queensland]], [[Australia]]
-Image:Chromodoris willani Lembeh.JPG|''[[Chromodoris willani]]''<br>Lembeh Straits, [[Indonesia]]
+Image:Chromodoris willani Lembeh.JPG|''[[Chromodoris willani]]''<br/>Lembeh Straits, [[Indonesia]]
-Image:Nembrotha kubaryana.JPG|''[[Nembrotha kubaryana]]''<br>Lembeh Straits, [[Indonesia]]
+Image:Nembrotha kubaryana.JPG|''[[Nembrotha kubaryana]]''<br/>Lembeh Straits, [[Indonesia]]
-Image:Chromodoris annae.JPG|''[[Chromodoris annae]]''<br>Lembeh Straits, [[Indonesia]]
+Image:Chromodoris annae.JPG|''[[Chromodoris annae]]''<br/>Lembeh Straits, [[Indonesia]]
-Image:Pteraeolidia ianthina1.JPG|''[[Pteraeolidia ianthina]]''<br>Lembeh Straits, [[Indonesia]]
+Image:Pteraeolidia ianthina1.JPG|''[[Pteraeolidia ianthina]]''<br/>Lembeh Straits, [[Indonesia]]
-Image:Nudibranch in tidepool.jpg|Clown nudibranch<br>''[[Triopha catalinae]]'', Northern [[California]]
+Image:Nudibranch in tidepool.jpg|Clown nudibranch<br/>''[[Triopha catalinae]]'', Northern [[California]]
-Image:Acanthodoris lutea in california tide pools 2.jpg|''[[Acanthodoris lutea]]''<br>Northern [[California]]
+Image:Acanthodoris lutea in california tide pools 2.jpg|''[[Acanthodoris lutea]]''<br/>Northern [[California]]
 </gallery>
-== Footnotes ==
-{{reflist}}
 ==References==
-* {{cite journal | author = H. Wägele and R. C. Willan | title = Phylogeny of the Nudibranchia | journal = Zoological Journal of the Linnean Society | date = September 2000 | volume = 1 | issue = 1 | pages = 83–181}}
+* Frick, K. 2003. [https://web.archive.org/web/20110106062453/http://archive.rubicon-foundation.org/4744 Predator suites and flabellinid nudibranch nematocyst complements in the Gulf of Maine] ''Proceedings of the 22nd Annual Scientific Diving Symposium, American Academy of Underwater Sciences. Greenville, North Carolina''. Retrieved October 14, 2021.
+* Integrated Taxonomic Information System (ITIS). 2004. [http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=78156 Nudibranchia Blainville, 1814] ''ITIS Taxonomic Serial No.: 78156''. Retrieved October 14, 2021.
-National Geographic Society. 2008. [http://animals.nationalgeographic.com/animals/printable/nudibranch.html Nudibranch (Nudibranchia)].
+* Klussmann-Kolb, A. 2001. [https://www.sciencedirect.com/science/article/abs/pii/S004452310470011X 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. Retrieved October 14, 2021.
+* National Geographic Society (NGS). 2008. [https://www.nationalgeographic.com/animals/invertebrates/facts/nudibranchs-1 Nudibranch] ''National Geographic Society''. Retrieved October 14, 2021.
-Holland, J. S. 2008. [http://ngm.nationalgeographic.com/2008/06/nudibranchs/holland-text Living color: Toxic nudibranchs&mdash;soft, seagoing slugs&mdash;produce a brilliant defense]. ''National Geographic'' June 2008. Retrieved July 23, 2008.
+* Ponder, W. F., and D. R. Lindberg. 1997. “Towards a phylogeny of gastropod molluscs: An analysis using morphological characters.” ''Zoological Journal of the Linnean Society'' 119: 83-2651.
+* Schrödl, M., H. Wägele, and R. C. Willan. 2001. [https://www.sciencedirect.com/science/article/abs/pii/S0044523104700091 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. Retrieved October 14, 2021.
+* Wägele, H., and R. C. Willan. 2000. Phylogeny of the Nudibranchia. ''Zoological Journal of the Linnean Society'' 1(1): 83–181.
-* Integrated Taxonomic Information System (ITIS). 2004. [http://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=78156 Nudibranchia  Blainville, 1814] ''ITIS Taxonomic Serial No.: 78156''. Retrieved July 23, 2008.
-==External links==
-* [http://www.nudibranch.com.au/ Lots of images and information on Australian nudibranchs]
-* [http://scubadiving.naroomaonline.com Nudibranchs in their natural environment, Scuba Diving - Narooma NSW]
-* [http://www.seaslugforum.net/ Sea Slug Forum: many images and much valuable information worldwide]
-* [http://www.nudipixel.net/ Nudi Pixel: Online resource for nudibranchs and sea slugs identification using photographs]
-* [http://www.sergeyphoto.com/underwater/nudibranchs.html Nudibranch gallery]
-* [http://www.seaslug.com/ Bibliography and portal to opisthobranch, nudibranch & seaslug information]
-* [http://www.waterworxbali.com/nudibranchs.shtml Nudibranch photo gallery from Bali]
-* [http://www.scottishnudibranchs.co.uk/index.html Scottish Nudibranchs: Online resource for identification of species found in Scottish waters]
-* [http://www.gmanews.tv/video/21917/Born-to-be-Wild-Sea-Slugs Born to be Wild: Sea Slugs, 04/30/2008]
-* http://ngm.nationalgeographic.com/2008/06/nudibranchs/doubilet-photography?email=focus_20080523
 [[Category:Life sciences]]