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Heterobranchia composite 03.jpg
Four examples of Heterobranchia
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
J.E. Gray, 1840
Taxonomic subcategories

informal group  Lower Heterobranchia
clade Nudipleura
clade Euopisthobranchia
clade Panpulmonata


Heterobranchia, the heterobranchs (meaning "different-gilled snails"), is a taxonomic clade of snails and slugs, which includes marine, aquatic and terrestrial gastropod mollusks.

Taxonomy (biology) The science of identifying, describing, defining and naming groups of biological organisms

In biology, taxonomy is the science of defining and naming groups of biological organisms on the basis of shared characteristics. Organisms are grouped together into taxa and these groups are given a taxonomic rank; groups of a given rank can be aggregated to form a super-group of higher rank, thus creating a taxonomic hierarchy. The principal ranks in modern use are domain, kingdom, phylum, class, order, family, genus, and species. The Swedish botanist Carl Linnaeus is regarded as the founder of the current system of taxonomy, as he developed a system known as Linnaean taxonomy for categorizing organisms and binomial nomenclature for naming organisms.

Clade A group of organisms that consists of a common ancestor and all its lineal descendants

A clade, also known as monophyletic group, is a group of organisms that consists of a common ancestor and all its lineal descendants, and represents a single "branch" on the "tree of life".

Snail mollusc

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 Gastropoda that have a coiled shell that is large enough for the animal to retract completely into. When the word "snail" is used in this most general sense, it includes not just land snails but also numerous species of sea snails and freshwater snails. Gastropods that naturally lack a shell, or have only an internal shell, are mostly called slugs, and land snails that have only a very small shell are often called semi-slugs.

Heterobranchia is one of the main clades of gastropods. Currently Heterobranchia comprises three informal groups: the lower heterobranchs, the opisthobranchs and the pulmonates. [1]

Lower Heterobranchia taxon of molluscs

Lower Heterobranchia, also known as the Allogastropoda, is a group of rather specialized, highly evolved sea slugs and sea snails, within the clade Heterobranchia. Although the great majority of Lower Heterobranchs are indeed marine, a few have succeeded in making the transition to freshwater.

Opisthobranchia infraclass of molluscs

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.

Pulmonata order of molluscs

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.

A shell of the marine species Bulla quoyii , which is an opisthobranch. Bulla quoyii, underside.JPG
A shell of the marine species Bulla quoyii , which is an opisthobranch.


The three subdivisions of this large clade are quite diverse:

Gastropod shell part of the body of a gastropod or snail

The gastropod shell is part of the body of a gastropod or snail, a kind of mollusc. The shell is an exoskeleton, which protects from predators, mechanical damage, and dehydration, but also serves for muscle attachment and calcium storage. Some gastropods appear shell-less (slugs) but may have a remnant within the mantle, or the shell is reduced such that the body cannot be retracted within (semi-slug). Some snails also possess an operculum that seals the opening of the shell, known as the aperture, which provides further protection. The study of mollusc shells is known as conchology. The biological study of gastropods, and other molluscs in general, is malacology. Shell morphology terms vary by species group. An excellent source for terminology of the gastropod shell is "How to Know the Eastern Land Snails" by John B. Burch now freely available at the Hathi Trust Digital Library.

Torsion is a gastropod synapomorphy which occurs in all gastropods during larval development. Torsion is the rotation of the visceral mass, mantle, and shell 180˚ with respect to the head and foot of the gastropod. This rotation brings the mantle cavity and the anus to an anterior position above the head.

Operculum (gastropod) A hard structure which closes the aperture of a gastropod when the animal retreats into the shell

The operculum, meaning little lid, is a corneous or calcareous anatomical structure like a trapdoor which exists in many groups of sea snails and freshwater snails, and also in a few groups of land snails; the structure is found in some marine and freshwater gastropods, and in a minority of terrestrial gastropods, including the families Helicinidae, Cyclophoridae, Aciculidae, Maizaniidae, Pomatiidae, etc.


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 & Pulmonata, and all the other gastropods. [2]

John Edward Gray British zoologist and philatelist (1800–1875)

John Edward Gray, FRS was a British zoologist. He was the elder brother of zoologist George Robert Gray and son of the pharmacologist and botanist Samuel Frederick Gray (1766–1828). The standard author abbreviation J.E.Gray is used to indicate this person as the author when citing a botanical name. or zoological name.

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, 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. [3]

Heterostropha was a previously used taxonomic category, an order of sea snails, within the superorder Heterobranchia. In the most current gastropod taxonomy, that of Bouchet & Rocroi, this taxon is no longer in use.

Architectonicidae family of molluscs

Architectonicidae, common name the staircase shells or sundials, are a family of sea snails, marine gastropod mollusks in the informal group "Lower Heterobranchia" of the clade Heterobranchia.


A protoconch 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 "nucleus". The protoconch may sometimes consist of several whorls, but when this is the case, the whorls show no growth lines.

According to the older taxonomy of the Gastropoda (Ponder & Lindberg, 1997) the Heterobranchia were ranked as a superorder.

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.

2005 taxonomy

The graph of neighbor-joining phylogenetic tree shows that there is no clade-supporting pattern for the monophyly of Opisthobranchia (green) or of Pulmonata (yellow) based on datasets by Jorger et al. (2010). Heterobranchia tree.png
The graph of neighbor-joining phylogenetic tree shows that there is no clade-supporting pattern for the monophyly of Opisthobranchia (green) or of Pulmonata (yellow) based on datasets by Jörger et al. (2010).

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) [4] have redefined major groups within the Heterobranchia: they created the new clades Euopisthobranchia and Panpulmonata. [4]

A cladogram showing phylogenic relations of Heterobranchia as proposed by Jörger et al. (2010): [4]


Lower Heterobranchia (including Acteonoidea) - Lower Heterobranchia does not form a clade in the study by Jörger et al. (2010) [4]










Cephalaspidea s.s.















Related Research Articles

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.

Cephalaspidea suborder of molluscs

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 molluscs

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

Sea snail common name for snails that normally live in saltwater

Sea snail is a common name for snails that normally live in salt water, in other words marine gastropods. The taxonomic class Gastropoda also includes snails that live in other habitats, such as land snails and freshwater snails. Many species of sea snails are edible and exploited as food sources by humans.

Amphibolidae family of molluscs

Amphibolidae is a family of air-breathing snails with opercula, terrestrial pulmonate gastropod molluscs.

Amphiboloidea superfamily of molluscs

Amphiboloidea is a taxonomic superfamily of air-breathing land snails.

Euthyneura clade of molluscs

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

Neritimorpha subclass of molluscs

Neritimorpha is a taxonomic grouping, an unranked clade of snails, gastropod mollusks. This grouping includes land snails, sea snails, some deepwater limpets, and also freshwater snails. This clade used to be known as the superorder Neritopsina.

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.

Siphonarioidea superfamily of molluscs

Siphonarioidea is a taxonomic superfamily of air-breathing sea snails or false limpets, marine gastropod mollusks in the clade Panpulmonata.

Umbraculoidea superfamily of molluscs

Umbraculoidea is a superfamily of unusual false limpets with a thin soft patelliform shell, marine gastropod molluscs in the clade Umbraculida, within the clade Euopisthobranchia.

Runcinoidea is a taxonomic superfamily or a clade Runcinaecea of sea slugs, marine gastropod mollusks in the clade Euopisthobranchia.

Rhodopidae family of molluscs

Rhodopidae is a taxonomic family of sea snails, marine opisthobranch gastropod mollusks in the superfamily Murchisonelloidea.

Heteroneritidae is an extinct taxonomic family of sea snails, marine gastropod mollusks in the superfamilia Pyramidelloidea.

Hygrophila (gastropod) clade of molluscs

Hygrophila is a taxonomic clade of air-breathing freshwater snails, aquatic pulmonate gastropod mollusks within the clade Panpulmonata.

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.

Architectibranchia is a clade of marine snails, gastropod mollusks.


  1. 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. ISBN   3-925919-72-4. ISSN   0076-2997. 397 pp. http://www.vliz.be/Vmdcdata/imis2/ref.php?refid=78278
  2. 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. ISSN   0044-3808.
  3. Ponder, W. F.; Warén, A. (1988). "Classification of the Caenogastropoda and Heterostropha- A list of family-group names and higher taxa". Malacological review supplement. 4: 288–317.
  4. 1 2 3 4 5 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 Lock-green.svg. PMID   20973994.

Further reading