The siphonal canal is an anatomical feature of the shells of certain groups of

sea snails Sea snail is a common name for slow-moving marine gastropod molluscs, usually with visible external shells, such as whelk or abalone. They share the taxonomic class Gastropoda with slugs, which are distinguished from snails primarily by the ...

within the clade

Neogastropoda Neogastropoda is an order of sea snails, both freshwater and marine gastropod molluscs. Description The available fossil record of Neogastropoda is relatively complete, and supports a widely accepted evolutionary scenario of an Early Creta ...

. Some sea marine

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 ...

s have a soft tubular

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

extension of the mantle called a

siphon A siphon (from grc, σίφων, síphōn, "pipe, tube", also spelled nonetymologically 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 a ...

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 respiratory organ that many 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 respiration on land provided they are ...

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 chemorecept ...

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 A stimulus is something that causes a physiological response. It may refer to: *Stimulation **Stimulus (physiology), something external that influences an activity **Stimulus (psychology), a concept in behaviorism and perception *Stimulus (economi ...

, 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 food and energy requirements derive from animal tissues (mainly muscle, fat and other sof ...

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, an aperture is a hole or an opening through which light travels. More specifically, the aperture and focal length of an optical system determine the cone angle of a bundle of rays that come to a focus in the image plane. An opt ...

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 about 1,600 living species, the Muricidae represent almost 10% of the Neogas ...

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 gastropod mollusk in the family Muricidae, the rock snails or murex snails. The shell of this Indo-Pacific species has a very long siph ...

. Some gastropods have a simplified

siphonal notch A siphonal notch is a feature of the shell anatomy in some sea snails, marine gastropod mollusks. In these particular groups of sea snails the animal has a soft tubular anterior extension of the mantle called a siphon through which water is dra ...

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 clade, 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 clade is the most diverse and ecologically successfu ...

(14 instances of siphon evolution) and

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

(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. The Ordovician spans 41.6 million years from the end of the Cambrian Period million years ago (Mya) to the start ...

, late

Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ' ...

, and

Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceo ...

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 in ...

23 times, and was subsequently lost 17 times. Secondary loss was associated with miniaturization, the adoption of a

sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...

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 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...

and

Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...

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