Opisthobranchia

Opisthobranchia Temporal range: Carboniferous–Recent[1] PreЄ Є O S D C P T J K Pg N
Notodoris minor
Scientific classification
Kingdom:	Animalia
Phylum:	Mollusca
Class:	Gastropoda
(unranked):	clade Heterobranchia informal group "Opisthobranchia"
Included groups
clade Cephalaspidea clade Thecosomata clade Gymnosomata clade Aplysiomorpha group Acochlidiacea clade Sacoglossa group Cylindrobullida clade Umbraculida clade Nudipleura

'Opisthobranchs ( /ɵˈpɪsθəbræŋks/) are a large and diverse group of specialized complex marine gastropods previously united under Opisthobranchia within the Heterobranchia, but no longer considered to represent a monophyletic grouping.^[2] Euopisthobrancha is a collection of opisthobranchs that is monophyletic, but this group leaves out some "traditional" opisthobranchs.^[3]

The Opisthobranchia includes a number of species in the order Cephalaspidea (bubble shells and headshield slugs), the sacoglossans, the anaspidean sea hares, the pelagic sea angels, the sea butterflies, and many families of Nudibranchia.^[2]

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.^[4]

Bullina lineata

Taxonomy

Under the old classification system by Johannes Thiele in 1931 Gastropoda were divided into the Prosobranchia, Pulmonata and Opisthobranchia. The later two were later combined into a single order.^[5]

Relation to pulmonates

Aplysia californica, a sea hare.

It is speculated that the Opisthobranchia may be paraphyletic,^[6] and may have given rise to the Pulmonata, although this is still disputed.

The Pulmonata may be a sibling group to an opisthobranch taxon. The Opisthobranchia are therefore not a monophyletic group and can no longer be accepted as a valid taxon. They are now included within the subclass Orthogastropoda. One can still encounter the old classification in many manuals and on most websites.

A new study of rRNA gene sequences, published in 2005, could not resolve monophyly versus paraphyly of the Opisthobranchia ^[7]

Linnean taxonomy

Order Opisthobranchia Milne-Edwards, 1848 - sea slugs

• Suborder Cephalaspidea P. Fischer, 1883 - headshield slugs and bubble shells
• Suborder Sacoglossa von Ihering, 1876 - sap-sucking slugs and bivalved gastropods
• Suborder Aplysiomorpha P. Fischer, 1883 - sea hares
• Suborder Notaspidea P. Fischer, 1883 - sidegill slugs
• Suborder Thecosomata Blainville, 1824 sea butterflies with shells
• Suborder Gymnosomata Blainville, 1824 - sea angels, no shells
• Suborder Nudibranchia Blainville, 1814 - nudibranchs
  • Infraorder Anthobranchia Férussac, 1819
  • Infraorder Cladobranchia Willan & Morton, 1984

A phylogenetic study published in November 2004,^[8] gave new definitions of the seven main lineages of the Opisthobranchia. However, the grouping is no longer considered to represent a valid clade, and thus appears as an informal group within the Heterobranchia in the taxonomy of the Gastropoda (Bouchet & Rocroi, 2005).^[2] Accordingly, recent articles do not use the term.^[3]

Description

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.^[9]

Opisthobranchs have undergone detorsion, an evolutionary reversal of the half revolution torsion of their immediate ancestors.

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.

They have pit-cup eyes^[ambiguous] with a lens and cornea.^[10]

Ecology

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.^[9]

Defence

Opisthobranchs are principally soft-bodied marine creatures with a reduced or absent shell and no operculum and utilize other methods for protection. Due to a combination of outstanding camouflage and aggressive toxicity they have few predators. However some utilize 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 defence.^[11][12]

Diet

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.^[13]

Communication

Pleurobranchus mamillatus

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

Reproduction

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.^[14] 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.^[15]

References

This article incorporates CC-BY-2.0 text from reference.^[9]

1. ^ Jensen, K. R. (1997). "Evolution of the Sacoglossa (Mollusca, Opisthobranchia) and the ecological associations with their food plants". Evolutionary Ecology 11: 301–335. doi:10.1023/A:1018468420368.  edit
2. ^ ^{a b c} Bouchet P., Rocroi J.-P., Frýda J., Hausdorf B., Ponder W., Valdés Á. & Warén A. (2005). "Classification and nomenclator of gastropod families". Malacologia: International Journal of Malacology (Hackenheim, Germany: ConchBooks) 47 (1-2): 1–397. ISBN 3925919724. ISSN 0076-2997. http://www.archive.org/details/malacologia47122005inst. 
3. ^ ^{a b} 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.
4. ^ (in Czech) Pek I., Vašíček Z., Roček Z., Hajn. V. & Mikuláš R.: Základy zoopaleontologie. - Olomouc, 1996. 264 pp., ISBN 80-7067-599-3.
5. ^ Knudsen, B.; Kohn, B.; Nahir, B.; Mcfadden, S.; Moroz, L. (Feb 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.  edit
6. ^ (Haszprunar, 1985)
7. ^ Verena Vonnemann, Michael Schrödl, Annette Klussmann-Kolb and 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. 
8. ^ Cristina Grandea, Josè Templadoa, J. Lucas Cerverab and 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. 
9. ^ ^{a b c} Kristof A. & Klussmann-Kolb A. (22 January 2010). "Neuromuscular development of Aeolidiella stephanieae Valdéz, 2005 (Mollusca, Gastropoda, Nudibranchia)". Frontiers in Zoology 7: 5. doi:10.1186/1742-9994-7-5.
10. ^ 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.  edit
11. ^ Gosliner, Terrence (1987) Nudibranchs of Southern Africa p.7 ISBN 0-930118-13-8
12. ^ Heike Wägele, Annette Klussmann-Kolb (2005). "Opisthobranchia (Mollusca, Gastropoda) – more than just slimy slugs. Shell reduction and its implications on defence and foraging". PubMed Central - Frontiers in Zoology 2 (3): 3. doi:10.1186/1742-9994-2-3. PMC 554092. PMID 15715915. http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pmcentrez&artid=554092. 
13. ^ Gosliner, Terrence (1987) Nudibranchs of Southern Africa p.9 ISBN 0-930118-13-8
14. ^ Debelius, Helmut (2001) Nudibranchs and Sea Snails Indo-Pacific Field Guide p.7 Ikan, Frankfurt
15. ^ Gosliner, Terrence (1987) Nudibranchs of Southern Africa p.11 ISBN 0-930118-13-8

Further reading

• Haszprunar G. (1985). "The Heterobranchia – a new concept of the phylogeny of the higher Gastropoda". Z. F. Zool. Systematik u. Evolutionsforschung Bd. 23 (1): 15–37. 
• Bieler R. (1990). "Haszprunar's "clado-evolutionary" classification of the Gastropoda—a critique". Malacologia 31(2): 371–380, 2 tabs. [28 May; G, Haszprunar's response published in Malacologia, 1990, 32(1): 195–202].
• Bieler R. (1992). "Gastropod phylogeny and systematics". Annual Review of Ecology and Systematics 23: 311–338. doi:10.1146/annurev.es.23.110192.001523. 
• Verena Vonnemann, Michael Schrödl, Annette Klussmann-Kolb and 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. 

External links

• The Sea Slug Forum, a resource from the Australian Museum.
• Sea slugs of Hawaii
• Born to be Wild: Sea Slugs, retrieved 30.4.2008
• Various opisthobranch species from Indonesia, Philippines and Thailand
• Nudi Pixel Nudi Pixel is a web-based identification tool for opisthobranchs worldwide