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Heterobranchia, the ''heterobranchs'' (meaning "different-gilled snails"), is a taxonomic
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
of
snail A snail is, in loose terms, a shelled gastropod. The name is most often applied to land snails, terrestrial pulmonate gastropod molluscs. However, the common name ''snail'' is also used for most of the members of the molluscan class ...
s and
slugs Slug, or land slug, is a common name for any apparently shell-less terrestrial molluscs, terrestrial gastropod mollusc. The word ''slug'' is also often used as part of the common name of any gastropod mollusc that has no shell, a very reduced ...
, which includes
marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military ...
, aquatic and terrestrial
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is e ...
s. Heterobranchia is one of the main clades of gastropods. Currently Heterobranchia comprises three informal groups: the lower heterobranchs, the opisthobranchs and the pulmonates.Bouchet P. & Rocroi J.-P. (Ed.); Frýda J., Hausdorf B., Ponder W., Valdes A. & Warén A. 2005. ''Classification and nomenclator of gastropod families''. Malacologia: International Journal of Malacology, 47(1-2). ConchBooks: Hackenheim, Germany. . . 397 pp. http://www.vliz.be/Vmdcdata/imis2/ref.php?refid=78278


Diversity

The three subdivisions of this large clade are quite diverse: * The Lower Heterobranchia includes shelled marine and freshwater species. *
Opisthobranchia Opisthobranchs () 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. Euopi ...
are almost all marine species, some shelled and some not. The internal organs of the opisthobranchs have undergone detorsion (unwinding of the viscera that were twisted during torsion). * The
Pulmonata Pulmonata or pulmonates, is an informal group (previously an Order (biology), order, and before that a Class (biology), subclass) of snails and slugs characterized by the ability to breathe air, by virtue of having a Respiratory system of gastro ...
includes the majority of land snails and slugs, many freshwater snails, and a small number of marine species. The mantle cavity of the Pulmonata is modified into an air-breathing organ. They are also characterized by detorsion and a symmetrically-arranged nervous system. The pulmonates almost always lack an operculum and are
hermaphroditic In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have s ...
.


Taxonomy


Older taxonomy

The families currently included in Heterobranchia have historically been placed in many different parts of the taxonomic class of gastropods. Earlier authors (such as J.E. Gray, 1840) considered Heterobranchia to consist of only marine gastropods, and conceptualized it as a borderline category, intermediate between the
Opisthobranchia Opisthobranchs () 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. Euopi ...
&
Pulmonata Pulmonata or pulmonates, is an informal group (previously an Order (biology), order, and before that a Class (biology), subclass) of snails and slugs characterized by the ability to breathe air, by virtue of having a Respiratory system of gastro ...
, and all the other gastropods. The (sometimes recognized) category Heterostropha within the Heterobranchia, which includes such families as Architectonicidae, the sundial or staircase snails, is primarily characterized by a shell which has a heterostrophic
protoconch A protoconch (meaning first or earliest or original shell) is an embryonic or larval shell which occurs in some classes of molluscs, e.g., the initial chamber of an ammonite or the larval shell of a gastropod. In older texts it is also called " ...
, in other words the apical whorls are coiled in the opposite plane to the adult whorls. The classification of this group was revised by Ponder & Warén in 1988. According to the older taxonomy of the Gastropoda (Ponder & Lindberg, 1997) the Heterobranchia were ranked as a superorder.


2005 taxonomy

Heterobranchia is currently one of the main clades of gastropods. For a detailed taxonomy, see Taxonomy of the Gastropoda (Bouchet & Rocroi, 2005)#Clade Heterobranchia.


2010 taxonomy

Jörger et al. (2010) have redefined major groups within the Heterobranchia: they created the new clades Euopisthobranchia and Panpulmonata. A
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
showing phylogenic relations of Heterobranchia as proposed by Jörger et al. (2010):


References


Further reading

* Dinapoli A. (2009). ''Phylogeny and Evolution of the Heterobranchia (Mollusca, Gastropoda)''. Thesis, Frankfurt am Main, 176 pp
PDF
* {{Taxonbar, from=Q133143 Gastropod taxonomy Protostome unranked clades Taxa named by John Edward Gray