The siphonal canal is an anatomical feature of the shells of certain groups of sea snails within the clade Neogastropoda. Some sea marine

gastropod Gastropods (), commonly known as slugs and snails, belong to a large Taxonomy (biology), taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and fro ...

s have a soft tubular

anterior Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position pro ...

extension of the mantle called a

siphon A siphon (; also spelled syphon) is any of a wide variety of devices that involve the flow of liquids through tubes. In a narrower sense, the word refers particularly to a tube in an inverted "U" shape, which causes a liquid to flow upward, abo ...

through which water is drawn into the

mantle cavity The mantle (also known by the Latin word pallium meaning mantle, robe or cloak, adjective pallial) is a significant part of the anatomy of molluscs: it is the dorsal body wall which covers the visceral mass and usually protrudes in the form of ...

and over the

gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow r ...

and which serves as a

chemoreceptor A chemoreceptor, also known as chemosensor, is a specialized sensory receptor which transduces a chemical substance ( endogenous or induced) to generate a biological signal. This signal may be in the form of an action potential, if the chemorece ...

to locate food. Siphonal canals allow for active transport of water to sensory organs inside the shell. Organisms without siphonal canals in their shells rely on passive or diffuse transport or water into their shell. Those with siphonal canals have a direct inhalant stream of water that interacts with sensory organs to detect concentration and direction of a stimulus, such as food or mates. In certain groups of

carnivorous A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose nutrition and energy requirements are met by consumption of animal tissues (mainly mu ...

snails, where the siphon is particularly long, the structure of the shell has been modified in order to house and protect the soft structure of the siphon. Thus the siphonal canal is a semi-tubular extension of the

aperture In optics, the aperture of an optical system (including a system consisting of a single lens) is the hole or opening that primarily limits light propagated through the system. More specifically, the entrance pupil as the front side image o ...

of the shell through which the siphon is extended when the animal is active.

Morphology

The size and shape of the siphonal canal are important in taxonomic identification of mollusks. Sealed siphonal canals are a morphological feature of

Muricidae Muricidae is a large and varied taxonomic family of small to large predatory sea snails, marine gastropod mollusks, commonly known as murex snails or rock snails. With over 1,700 living species, the Muricidae represent almost 10% of the Neog ...

subfamilies Ocenebrinae and Typhinae, a feature not found in other subfamilies from this taxonomic group. One gastropod whose shell has an exceptionally long siphonal canal is the

Venus comb murex The Venus comb murex, scientific name ''Murex pecten'', is a species of large predatory sea snail, a marine (ocean), marine gastropod mollusk in the family (biology), family Muricidae, the rock snails or murex snails. The gastropod shell, shell o ...

. Some gastropods have a simplified siphonal notch at the edge of the aperture instead of a canal. For example, high intertidal gastropods often have a very short siphonal canal or a greatly reduced siphonal notch. There are close parallels between shell shape and presence of a siphonal canal. A study done by Vermeij found that shells that have a steep angle of their generating curve relative to their base axis (E) not do not have siphonal canals. Specifically, all shells examined with an E angle greater than 30% did not have siphonal canals, but shells with lower angles often did. A short siphonal canal is preferred by some species of obligate commensal bivalves. '' Curvemysella paula'' is specially adapted to live inside snail shells occupied by hermit crabs. The crescent shaped shell of ''C. paula'' facilitates entry into the narrow space inside snail shells where they attach to the columellae and siphonal canal via byssal threads. The position and location of attachment anteriorly near the final whorls of the shell is an evolutionary behavior of adult individuals which prevents removal by the host.

Evolution

Siphonal indentations have evolved multiple times in gastropods and are widespread among many clades. Euomphanilae gastropods in the genus ''Scalites'' developed siphons in the early Ordovician period (448-443 MYA); however, they are not observed in any other members of the clade. 22 of an estimated >23 instances of siphonal indentations evolved in Murchosinoniinae gastropods - the two major clades in this group are Vetigastropoda (1 siphonate group: ''Tylozone'') and Apogastropoda. Apodastropoda contains

Caenogastropoda Caenogastropoda is a taxonomic subclass of molluscs in the class Gastropoda. It is a large diverse group which are mostly sea snails and other marine gastropod mollusks, but also includes some freshwater snails and some land snails. The subcla ...

(14 instances of siphon evolution) and

Heterobranchia Heterobranchia, the heterobranchs (meaning "different gill"), is a taxonomic clade of snails and slugs, which includes marine, aquatic, and terrestrial gastropod molluscs. Heterobranchia is one of the main clades of gastropods. Currently Het ...

(3 instances of siphon evolution). The evolution of siphonal indentations and related parts are not associated with species diversification and thus are likely not key innovations. Diversification is observed in only 17% of siphonate clades and primarily occurred during the

Ordovician The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...

, late

Paleozoic The Paleozoic ( , , ; or Palaeozoic) Era is the first of three Era (geology), geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma a ...

, and

Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...

eras and was restricted to clades with few genera. Active predation only occurs in 1 siphonate clade (~4.5%) and is not considered a driver of evolution. Further, the evolution of the siphonal canal did not coincide with a change in feeding style as the basal ancestor was nonpredatory. One hypothesis is that the evolution of the siphonal canal was due to locomotion, because species without siphonal canals are largely sedentary, whereas siphonal families can move around the benthos. However, due to the widespread ecological contexts of gastropods with a siphonal canal, there is no consensus hypothesis or unified driver which can explain the evolution or loss of this structure. The siphonal canal arose independently in the

fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

23 times, and was subsequently lost 17 times. Secondary loss was associated with miniaturization, the adoption of a sessile lifestyle, and non-marine habits. A possible driver for these losses may have been species no longer relying on distance chemoreception, common in non-marine gastropods. During the

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...

and

Cenozoic The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...

periods, some species evolved long or closed siphonal canals. This was the earliest instance of long canal and closed canal evolution, which subsequently arose approximately 15 times. Closed canals in particular helped prevent predation by armoring the gastropod. More generally, the siphonal canal protects gastropods by allowing their shells to be fully closed, with their mantle protected, while still sensing cues from the environment.

Morphology

Evolution

See also

References

Further reading