Last updated

Temporal range: Carboniferous–Recent [1]
Notodoris minor.jpg
Notodoris minor
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Subclass: Heterobranchia
Informal group: Opisthobranchia
Milne-Edwards, 1848
Included groups

clade Cephalaspidea
clade Thecosomata
clade Gymnosomata
clade Aplysiomorpha
group Acochlidiacea
clade Sacoglossa
group Cylindrobullida
clade Umbraculida
clade Nudipleura


Opisthobranchs ( /əˈpɪsθəˌbræŋks,-θ-/ [2] ) is now an informal name for a large and diverse group of specialized complex gastropods which used to be united in the subclass Opisthobranchia. That taxon is no longer considered to represent a monophyletic grouping. [3]

Euopisthobranchia is a taxon containing a revised collection of opisthobranchs, and that taxon is considered monophyletic. Euopisthobranchia does not include some "traditional" opisthobranchs such as the Sacoglossa and the Acochlidiacea. [4] The subclass Heterobranchia now contains all the species which used to be assigned to Opisthobranchia, plus all the species in the Pulmonata.

The subclass Opisthobranchia included species in the order Cephalaspidea (bubble shells and headshield slugs), the sacoglossans, anaspidean sea hares, pelagic sea angels, sea butterflies, and many families of the Nudibranchia. [3]

Opisthobranch means "gills behind" (and to the right) of the heart. In contrast, Prosobranch means gills in front (of the heart). Opisthobranchs are characterized by two pairs of tentacles and a single gill behind and to the right of the heart. Opisthobranchia are known from as early as the Carboniferous. [5]

Bullina lineata Bullina lineata 1.jpg
Bullina lineata


Under the 1931 classification system drawn up by Johannes Thiele, the class Gastropoda was divided into three subclasses; Prosobranchia, Pulmonata and Opisthobranchia. The latter two were later combined into a single order. [6]

The current classification of the gastropods is more nuanced, with the following subclasses: [7]

As a non-monophyletic taxon

Aplysia californica, a sea hare. Sea-hare-morro-bay (431920585).jpg
Aplysia californica , a sea hare.

It was speculated as far back as 1985 that the Opisthobranchia were paraphyletic, based on morphological evidence, and had given rise to the Pulmonata, a group also of subclass rank. [8]

Because the Pulmonata are a sibling group to an opisthobranch taxon, some authors argued that the Opisthobranchia are therefore not a monophyletic group and can no longer be accepted as a valid taxon. The opisthobranchs are now included within the subclass Heterobranchia, although many manuals and websites still use the old classification.

A phylogenetic study published in November 2004, [9] gave new definitions of the seven main lineages of the Opisthobranchia.

However, in 2005, a study of rRNA gene sequences could not resolve monophyly versus paraphyly of the Opisthobranchia [10] Subsequent taxonomic classification in 2005 overturned the Opisthobranchia as a valid clade, reclassifying it as an informal group within the Heterobranchia. [3] Accordingly, articles no longer use the term Opisthobranchia, replacing it with Heterobranchia to emphasise a different concept. [4]

By 2011, the "Opisthobranchia" were declared artificial and obsolete and were replaced by new phylogenetic hypotheses. [11] The concept was abandoned. It now includes including "Lower Heterobranchia", Acteonimorpha, Ringipleura, Umbraculida, Cephalaspidea, Runcinida, Aplysiida, Pteropoda and Sacoglossa. [12]

Linnean taxonomy

Order Opisthobranchia Milne-Edwards, 1848 – sea slugs


The reduction or loss of the shell, the elaboration of the head, foot or mantle, and the acquisition of chemical defences are evolutionary trends shared by most opisthobranch taxa. [13]

Opisthobranchs have undergone detorsion, an evolutionary reversal of the half revolution torsion of their immediate ancestors. As a result of this detorsion, the visceral ganglia no longer overlap and are described as euthyneurous (as opposed to streptoneurous, the more common condition among gastropods, in which these ganglia form a distinct twist within the animal's body).

There is no marked distinction between head and mantle. The tentacles, situated close to the mouth, are used for orientation. Behind them are the rhinophores, olfactory organs which often have complex forms. The middle part of the foot is the sole, used for locomotion. The sides of the foot have evolved into parapodia, fleshy winglike outgrowths. In several suborders, such as the Thecosomata and Gymnosomata, these parapodia are used to move in a swimming motion.

Their eyes are simple pit-cup eyes with a lens and cornea capable of detecting light and the passage of shadows but not of producing a coherent image. [14]


Opisthobranchia represents a morphologically diverse group of gastropods occupying a great variety of ecological niches. Opisthobranchs have a global distribution, but are restricted almost exclusively to marine habitats with the only exception being few freshwater acochlidians. [13]


Principally soft-bodied marine creatures with a reduced or absent shell and no operculum, opisthobranchs use other methods for protection. Due to a combination of outstanding camouflage and aggressive toxicity they have few predators. However, some use warning colouration. Animals that do predate opisthobranchs include other opisthobranchs and toxin-resistant predators like sea spiders.

Opisthobranchs secrete irritants such as strong acids or accumulate toxins from their food. Aeolidioidea pirate the stinging cells from their cnidarian prey and use them for their own defense. [15] [16]


Oxynoe olivacea Oolivacea.Mgiangrasso.jpg
Oxynoe olivacea

Opisthobranchs may be herbivores, detritivores or carnivores. Being slow, the carnivores hunt sedentary prey. They may eat bryozoans, Cnidaria, or sponges, absorbing the sponge toxin for defensive purposes. Opisthobranchs may maintain the zooxanthellae of their coral prey and use their metabolic products for themselves. Some herbivorous slugs do the same with the chloroplasts of the algae they eat. [17]


Pleurobranchus mamillatus Pleurobranchus mamillatus.jpg
Pleurobranchus mamillatus

Like most lifeforms, they use chemical cues for much of their life cycle. The planktonic larvae float until a pheromone alerts them to a suitable settling site, sometimes delaying metamorphosis until favourable chemicals, such as prey pheromones, are detected. Some mating opisthobranchs release chemicals to attract conspecifics.


Winged snail Clione limacina, a sea angel. Sea angel.jpg
Winged snail Clione limacina , a sea angel.

Opisthobranchs are hermaphrodites and have complex reproductive strategies, typically involving reciprocal sperm transfer and storage until the eggs are ready for fertilisation. [18] Eggs are commonly laid in ribbons of varying structure. The egg ribbons are usually unique to each species and in some cases are the only means of differentiating them. [19]

See also

Related Research Articles

Nudibranch Order of gastropods

Nudibranchs are a group of soft-bodied, marine gastropod molluscs which shed their shells after their larval stage. 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," or "sea rabbit." Currently, about 3,000 valid species of nudibranchs are known.

Sea slug Group of marine invertebrates with varying levels of resemblance to terrestrial slugs

Sea slug is a common name for some marine invertebrates with varying levels of resemblance to terrestrial slugs. Most creatures known as sea slugs are actually gastropods, i.e. they are sea snails that over evolutionary time have either completely lost their shells, or have seemingly lost their shells due to having a greatly reduced or internal shell. The name "sea slug" is most often applied to nudibranchs, as well as to a paraphyletic set of other marine gastropods without obvious shells.

Prosobranchia Historic group of molluscs

Prosobranchia was a large taxonomic subclass of sea snails, land snails and freshwater snails. This taxon of gastropods dates back to the 1920s. It has however been proven to be polyphyletic. Generally speaking in biology taxonomy is required to reflect phylogeny, in other words the classification of a group must reflect its evolutionary descent, as far as that is known, so the taxon Prosobranchia is no longer considered suitable to be used.

Orthogastropoda was a major taxonomic grouping of snails and slugs, an extremely large subclass within the huge class Gastropoda according to the older taxonomy of the Gastropoda.

Heterobranchia Clade of gastropods

Heterobranchia, the heterobranchs, is a taxonomic clade of snails and slugs, which includes marine, aquatic and terrestrial gastropod mollusks.

Cephalaspidea Order of gastropods

The clade Cephalaspidea, also known as the headshield slugs and bubble snails, is a major taxon of sea slugs and bubble snails, marine gastropod mollusks within the larger clade Euopisthobranchia. Bubble shells is another common name for these families of marine gastropods, some of which have thin bubble-like shells. This clade contains more than 600 species.

Acteonoidea Superfamily of gastropods

Acteonoidea is a superfamily of sea snails, or bubble snails, marine gastropod mollusks.

Pulmonata Informal group of gastropods

Pulmonata, or "pulmonates", is an informal group of snails and slugs characterized by the ability to breathe air, by virtue of having a pallial lung instead of a gill, or gills. The group includes many land and freshwater families, and several marine families.

Sacoglossa Clade of gastropods

Sacoglossa, commonly known as the sacoglossans or the "sap-sucking sea slugs", are a clade of small sea slugs and sea snails, marine gastropod mollusks that belong to the clade Heterobranchia. Sacoglossans live by ingesting the cellular contents of algae, hence the adjective "sap-sucking".

Limapontiidae Family of gastropods

Limapontiidae is a taxonomic family of small to minute sacoglossan sea slugs. These are marine opisthobranch gastropod mollusks.

Notaspidea Group of molluscs

Notaspidea, also known as the sidegill slugs, is an artificial grouping of sea slugs which is now split into two unrelated groups, the Umbraculida and the Pleurobranchomorpha.

The taxonomy of the Gastropoda, as revised by Winston Ponder and David R. Lindberg in 1997, is an older taxonomy of the class Gastropoda, the class of molluscs consisting of all snails and slugs. The full name of the work in which this taxonomy was published is Towards a phylogeny of gastropod molluscs: an analysis using morphological characters.

Apogastropoda Group of molluscs

Apogastropoda was previously used as a major taxonomic grouping of sea snails, marine gastropod molluscs. This infraclass mostly consisted of marine limpets and operculate snails. At least 20,000 species were considered to exist within the two clades that were included, Heterobranchia and Caenogastropoda.

Euthyneura Clade of molluscs

Euthyneura is a taxonomic infraclass of snails and slugs, which includes species from freshwater, marine, aquatic and terrestrial gastropod mollusks in the clade Heterobranchia.

Lower Heterobranchia Group of molluscs

Lower Heterobranchia, also known as the Allogastropoda, is a group of rather specialized, highly evolved sea slugs and sea snails, within the subclass Heterobranchia.

Acochlidiacea Order of molluscs

Acochlidiacea, common name acochlidians, are a taxonomic clade of very unusual sea snails and sea and freshwater slugs, aquatic gastropod mollusks within the large clade Heterobranchia. Acochlidia is a variant spelling.

This overview lists proposed changes in the taxonomy of gastropods at the family level and above since 2005, when the taxonomy of the Gastropoda by Bouchet & Rocroi (2005) was published. In other words, these are recent updates in the way various groups of snails and slugs are classified.

Aiteng is a genus of two species of sea slugs, marine gastropod molluscs and one terrestrial species. Aiteng is the only genus in the family Aitengidae. The generic name Aiteng is derived from the name of a black puppet Ai Theng, which is one of the shadow play puppets in southern Thailand.

Euopisthobranchia is a taxonomic clade of snails and slugs in the clade Heterobranchia within the clade Euthyneura.

The oesophageal pouches are a pair of pouches connected to the oesophagus of all molluscs, and represent a synapomorphy of the phylum.


This article incorporates CC-BY-2.0 text from reference [13]

  1. Jensen, K. R. (1997). "Evolution of the Sacoglossa (Mollusca, Opisthobranchia) and the ecological associations with their food plants". Evolutionary Ecology. 11 (3): 301–335. doi:10.1023/A:1018468420368.
  2. "Opisthobranch". Oxford Dictionaries UK Dictionary. Oxford University Press . Retrieved 21 January 2016.
  3. 1 2 3 Bouchet, Philippe; Rocroi, Jean-Pierre; Frýda, Jiri; Hausdorf, Bernard; Ponder, Winston; Valdés, Ángel & Warén, Anders (2005). "Classification and nomenclator of gastropod families". Malacologia . Hackenheim, Germany: ConchBooks. 47 (1–2): 1–397. ISBN   3-925919-72-4. ISSN   0076-2997.
  4. 1 2 Jörger, K. M.; Stöger, I.; Kano, Y.; Fukuda, H.; Knebelsberger, T.; Schrödl, M. (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.
  5. (in Czech) Pek I., Vašíček Z., Roček Z., Hajn. V. & Mikuláš R.: Základy zoopaleontologie. – Olomouc, 1996. 264 pp., ISBN   80-7067-599-3.
  6. Knudsen, B.; Kohn, B.; Nahir, B.; Mcfadden, S.; Moroz, L. (February 2006). "Complete DNA sequence of the mitochondrial genome of the sea-slug, Aplysia californica: conservation of the gene order in Euthyneura". Molecular Phylogenetics and Evolution. 38 (2): 459–469. doi:10.1016/j.ympev.2005.08.017. ISSN   1055-7903. PMID   16230032.
  7. Gofas, S. (2014). Gastropoda. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=101 on 2015-02-12
  8. Haszprunar G. (1985). "The Heterobranchia – a new concept of the phylogeny of the higher Gastropoda". Zeitschrift für zoologische Systematik und Evolutionsforschung. 23 (1): 15–37.
  9. Cristina Grandea; Josè Templadoa; J. Lucas Cerverab; Rafael Zardoya (2004). "Phylogenetic relationships among Opisthobranchia (Mollusca: Gastropoda) based on mitochondrial cox 1, trnV, and rrnL genes". Molecular Phylogenetics and Evolution. 33 (2): 378–388. doi:10.1016/j.ympev.2004.06.008. PMID   15336672.
  10. Verena Vonnemann; Michael Schrödl; Annette Klussmann-Kolb; Heike Wägele (2005). "Reconstruction of the phylogeny of the Opisthobranchia (Mollusca: Gastropoda) by means of 18s and 28s rRNA gene sequences". Journal of Molluscan Studies . 71 (2): 113–125. doi: 10.1093/mollus/eyi014 .
  11. Schrödl M; Jörger KM; Klussmann-Kolb A; Wilson NG. "Bye bye "Opisthobranchia"! A review on the contribution of mesopsammic sea slugs to euthyneuran systematics" (PDF). Thalassas. 27 (2): 101–112.
  12. MolluscaBase eds. (2020). MolluscaBase. Opisthobranchia. Accessed through: World Register of Marine Species at: http://marinespecies.org/aphia.php?p=taxdetails&id=382226 on 2020-12-23
  13. 1 2 3 Kristof, Alen; Klussmann-Kolb, Annette (2010). "Neuromuscular development of Aeolidiella stephanieae Valdez, 2005 (Mollusca, Gastropoda, Nudibranchia)". Frontiers in Zoology. 7 (1): 5. doi:10.1186/1742-9994-7-5. PMC   2822759 . PMID   20205753.
  14. Hughes, H. P. I. (1970). "A light and electron microscope study of some opisthobranch eyes". Zeitschrift für Zellforschung und Mikroskopische Anatomie. 106: 79–98. doi:10.1007/BF01027719.
  15. Gosliner, Terrence (1987) Nudibranchs of Southern Africa p. 7 ISBN   0-930118-13-8
  16. Heike Wägele; Annette Klussmann-Kolb (2005). "Opisthobranchia (Mollusca, Gastropoda) – more than just slimy slugs. Shell reduction and its implications on defence and foraging". Frontiers in Zoology. 2 (3): 3. doi:10.1186/1742-9994-2-3. PMC   554092 . PMID   15715915.
  17. Gosliner, Terrence (1987) Nudibranchs of Southern Africa p. 9 ISBN   0-930118-13-8
  18. Debelius, Helmut (2001) Nudibranchs and Sea Snails Indo-Pacific Field Guide p. 7 Ikan, Frankfurt
  19. Gosliner, Terrence (1987) Nudibranchs of Southern Africa p. 11 ISBN   0-930118-13-8

Further reading