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This article is about the group of sea snails. For other uses, see Conus (disambiguation).
Not to be confused with Telescopium (gastropod).

Cone snail
A group of shells of various species of cone snails
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Subclass: Caenogastropoda
Order: Neogastropoda
Superfamily: Conoidea
Family: Conidae
Fleming, 1822[1]
Subfamilies and genera

See text

Synonyms
  • Californiconinae Tucker & Tenorio, 2009
  • Conilithidae Tucker & Tenorio, 2009
  • Profundiconinae Limpalaër & Monnier, 2018· accepted, alternate representation
  • Puncticulinae Tucker & Tenorio, 2009
  • Taranteconidae Tucker & Tenorio, 2009

Cone snails, or cones, are highly venomous sea snails that constitute the family Conidae.[2] Conidae is a taxonomic family (previously subfamily) of predatory marine gastropod molluscs in the superfamily Conoidea.

The 2014 classification of the superfamily Conoidea groups only cone snails in the family Conidae. Some previous classifications grouped the cone snails in a subfamily, Coninae. As of March 2015 Conidae contained over 800 recognized species, varying widely in size from lengths of 1.3 cm to 21.6 cm. Working in 18th-century Europe, Carl Linnaeus knew of only 30 species that are still considered valid.

Fossils of cone snails have been found from the Eocene to the Holocene epochs.[3] Cone snail species have shells that are roughly conical in shape. Many species have colorful patterning on the shell surface.[4] Cone snails are almost exclusively tropical in distribution.

All cone snails are venomous and capable of stinging. Cone snails use a modified radula tooth and a venom gland to attack and paralyze their prey before engulfing it. The tooth, which is likened to a dart or a harpoon, is barbed and can be extended some distance out from the head of the snail at the end of the proboscis.

Cone snail venoms are mainly peptide-based, and contain many different toxins that vary in their effects. The sting of several larger species of cone snails can be serious, and even fatal to humans. Cone snail venom also shows promise for medical use.[5][6]

Distribution and habitat

[edit]

Species in the family Conidae are found in the tropical and subtropical seas of the world, in four biogeographic regions, including: the Indo-Pacific (with 60% of all species), the Tropical Eastern Pacific, the western Tropical Atlantic, and the eastern Tropical Atlantic, plus 10 species in the warm temperate Agulhas bioregion on the southern coast of South Africa. Fewer than one percent of fossil species have been found in more than one of the above regions.[7]

Cone snails are typically found in warm tropical seas and oceans worldwide. Cone snails reach their greatest diversity in the Western Indo-Pacific region. While the majority of cone snails are found in warm tropical waters, some species have adapted to temperate/semi-tropical environments and are endemic to areas such as the Cape coast of South Africa,[8][9] the Mediterranean,[10] or the cool subtropical waters of southern California (Californiconus californicus).[11]

They live on a variety of substrates, from the intertidal zone and deeper areas, to sand, rocks or coral reefs.

Paleontology

[edit]

The oldest known fossil of Conidae is from the lower Eocene, about 55 million years ago. Analysis of nucleotide sequences indicate that all living species of Conidae belong to one of two clades that diverged about 33 million years ago. One clade includes most of the species in the eastern Pacific and western Atlantic regions, which were connected by the Central American Seaway until the emergence of the Isthmus of Panama less than three million years ago. The other clade includes most of the species in the eastern Atlantic and Indo-Pacific regions, which were connected by the Neo-Tethys Sea until 21 to 24 million years ago.[7]

Shell

[edit]

Cone snails have a large variety of shell colors and patterns, with local varieties and color forms of the same species often occurring. This variety in color and pattern has led to the creation of a large number of known synonyms and probable synonyms, making it difficult to give an exact taxonomic assignment for many snails in this genus. As of 2009, more than 3,200 different species names have been assigned, with an average of 16 new species names introduced each year.[12]

The shells of cone snails vary in size and are conical in shape. The shell is whorled in the form of an inverted cone, with the anterior end being narrower. The protruding parts of the top of the whorls, that form the spire, are in the shape of another more flattened cone. The aperture is elongated and narrow with the sharp operculum being very small. The outer lip is simple, thin, and sharp, without a callus, and has a notched tip at the upper part. The columella is straight.

The larger species of cone snails can grow up to 23 cm (9.1 in) in length. The shells of cone snails are often brightly colored with a variety of patterns. Some species color patterns may be partially or completely hidden under an opaque layer of periostracum. In other species, the topmost shell layer is a thin periostracum, a transparent yellowish or brownish membrane.

Physiology and behavior

[edit]

The snails within this family are sophisticated predatory animals.[13] They hunt and immobilize prey using a modified radular tooth along with a venom gland containing neurotoxins; the tooth is launched out of the snail's mouth in a harpoon-like action.

Cone snails are carnivorous. Their prey consists of marine worms, small fish, molluscs, and other cone snails. Cone snails are slow-moving, and use their venomous harpoon to disable faster-moving prey.

The osphradium in cone snails is more specialized than in other groups of gastropods. It is through this sensory modality that cone snails are able to sense their prey. The cone snails immobilize their prey using a modified, dartlike, barbed radular tooth, made of chitin, along with a venom gland containing neurotoxins.

Molecular phylogeny research has shown that preying on fish has evolved at least twice independently in cone snails. Some species appear to have also evolved prey mimicry, where they release chemicals that resemble the sex pheromones certain ragworms release during their short breeding season. The researchers hypothesize that these chemicals cause the prey to be more easily harpooned, but are still uncertain as to exactly how this occurs in the wild.[14]

Harpoon

[edit]
An individual (Conus pennaceus) attacking one of a cluster of three snails of the species Cymatium nicobaricum, in Hawaii

Cone snails use a harpoon-like structure called a radula tooth for predation. Radula teeth are modified teeth, primarily made of chitin and formed inside the mouth of the snail, in a structure known as the toxoglossan radula. Each specialized cone snail tooth is stored in the radula sac, except for the tooth that is in current use.[15]

The radula tooth is hollow and barbed, and is attached to the tip of the radula in the radular sac, inside the snail's throat. When the snail detects a prey animal nearby, it extends a long flexible tube called a proboscis towards the prey. The radula tooth is loaded with venom from the venom bulb and, still attached to the radula, is fired from the proboscis into the prey by a powerful muscular contraction. The venom can paralyze smaller fish almost instantly. The snail then retracts the radula, drawing the subdued prey into the mouth. After the prey has been digested, the cone snail will regurgitate any indigestible material, such as spines and scales, along with the harpoon. There is always a radular tooth in the radular sac. A tooth may also be used in self-defense when the snail feels threatened.[16][17]

The harpoon attack of the species Conus catus has been found to be one of the fastest complete movements recorded in animals, with a maximum speed of 90 km/h (56 mph), an acceleration of 400,000 m/s², and a deceleration of 700,000 m/s². The speed of other animals such as the peacock mantis shrimp and the trap-jaw ant was measured at the free end of a fixed appendage, while the speed of the harpoon was measured from its base and traveling inside the proboscis.[18]

The reason for this speed relies in hydrostatic pressure by the fluid inside the proboscis which propels the harpoon inside until it is almost completely out. A sphincter acts as a valve to keep fluid in the proximal half and in the distal half a constriction of ephitelial tissue together with a thicker harpoon base helps to build up hydrostatic pressure when the sphincter opens. The deceleration may help release the venom from the harpoon.[18]

Venom

[edit]
Cone snail venom apparatus

There are approximately 30 records of humans killed by cone snails. Human victims suffer little pain, because the venom contains an analgesic component. Some species reportedly can kill a human in under five minutes, thus the name "cigarette snail" as supposedly one only has time to smoke a cigarette before dying. Cone snails can sting through a wetsuit with their harpoon-like radular tooth, which resembles a transparent needle.[19]

Normally, cone snails (and many species in the superfamily Conoidea) use their venom to immobilize prey before engulfing it. The venom consists of a mixture of peptides, called conopeptides. The venom is typically made up of 10 to 30 amino acids, but in some species as many as 60. The venom of each cone snail species may contain as many as 200 pharmacologically active components. It is estimated that more than 50,000 conopeptides can be found, because every species of cone snail is thought to produce its own specific venom.

Cone-snail venom has come to interest biotechnologists and pharmacists because of its potential medicinal properties. Production of synthetic conopeptides has started, using solid-phase peptide synthesis.

A component of the venom of Conus magus, ω-conotoxin, is now marketed as the analgesic ziconotide, which is used as a last resort in chronic and severe pain. Conopeptides are also being looked at as anti-epileptic agents and to help stop nerve-cell death after a stroke or head injury. Conopeptides also have potential in helping against spasms due to spinal cord injuries, and may be helpful in diagnosing and treating small cell carcinomas in the lung.

The biotechnology surrounding cone snails and their venom has promise for medical breakthroughs; with more than 50,000 conopeptides to study, the possibilities are numerous.[20]

Reproduction

[edit]

Most cone snails appear to reproduce sexually, with separate sexes and internal fertilization. Varying numbers of eggs in egg capsules are laid in substrate by cone snails. Hatchlings are of two types, the veligers (larvae that swim freely) and veliconcha (baby snail).[21]

Relevance to humans

[edit]

Because all cone snails are venomous and capable of stinging humans, live ones should be handled with great care or preferably not at all.

Dangers

[edit]
A live textile cone (Conus textile), one of several species whose venom can cause serious harm to a human

Cone snails are prized for their brightly colored and patterned shells,[22] which may tempt people to pick them up. This is risky, as the snail often fires its harpoon in self defense when disturbed. The harpoons of some of the larger species of cone snail can penetrate gloves or wetsuits.

The sting of many of the smallest cone species may be no worse than a bee or hornet sting,[23] but the sting of a few of the larger tropical fish-eating species, such as Conus geographus (geography cone), Conus tulipa and Conus striatus, can be fatal. Other dangerous species are Conus pennaceus, Conus textile, Conus aulicus, Conus magus and Conus marmoreus.[24] According to Goldfrank's Toxicologic Emergencies, about 27 human deaths can be confidently attributed to cone snail envenomation, though the actual number is almost certainly much higher; some three dozen people are estimated to have died from geography cone envenomation alone.[25]

Most of the cone snails that hunt worms are not a risk to humans, with the exception of larger species. One of the fish-eating species, the geography cone is also known colloquially as the "cigarette snail", a gallows humor exaggeration implying that, when stung by this creature, the victim will have only enough time to smoke a cigarette before dying.[16][26]

Symptoms of a more serious cone snail sting include severe, localized pain, swelling, numbness and tingling, and vomiting. Symptoms can start immediately or can be delayed for days. Severe cases involve muscle paralysis, changes in vision and respiratory failure that can lead to death. If stung, one should seek medical attention as soon as possible.[27]

Medical use

[edit]

The appeal of conotoxins for creating pharmaceutical drugs is the precision and speed with which the chemicals act; many of the compounds target only a particular class of receptor. This means that they can reliably and quickly produce a particular effect on the body's systems without side effects; for example, almost instantly reducing heart rate or turning off the signaling of a single class of nerve, such as pain receptors.

Ziconotide, a powerful atypical painkiller, was initially isolated from the venom of the magician cone snail, Conus magus.[28] It was approved by the U.S. Food and Drug Administration in December 2004 under the name Prialt. Other drugs based on cone snail venom targeting Alzheimer's disease, Parkinson's disease, depression, and epilepsy are in clinical or preclinical trials.[29][30]

Many peptides produced by the cone snails show prospects for being potent pharmaceuticals, such as AVC1, isolated from the Australian species, the Queen Victoria cone, Conus victoriae, and have been highly effective in treating postsurgical and neuropathic pain, even accelerating recovery from nerve injury.

Geography and tulip cone snails are known to secrete a type of insulin that paralyzes nearby fish by causing hypoglycaemic shock. They are the only two non-human animal species known to use insulin as a weapon.[31] Cone snail insulin is capable of binding to human insulin receptors and researchers are studying its use as a potent fast-acting therapeutic insulin.[32]

Shell collecting

[edit]

The intricate color patterns of cone snails have made them one of the most popular species for shell collectors.[33][34]

Conus gloriamaris, also known as "Glory of the Seas", one of the most famous and sought-after seashells in past centuries, with only a few specimens in private collections. The rarity of this species' shells led to high market prices for the objects, until the habitat of this cone snail was discovered, which decreased prices dramatically.[35]

As jewelry

[edit]

Naturally occurring, beach-worn cone shell tops can function as beads without any further modification. In Hawaii, these natural beads were traditionally collected from the beach drift to make puka shell jewelry. Since it is difficult to obtain enough naturally occurring cone snail tops, almost all modern puka shell jewelry uses cheaper imitations, cut from thin shells of other species of mollusk, or made of plastic.

Species

[edit]
Main article: List of Conus species

Until 2009 all species within the family Conidae were placed in one genus, Conus. Testing of the molecular phylogeny of the Conidae was first conducted by Christopher Meyer and Alan Kohn,[36] and has continued, particularly with the advent of nuclear DNA testing.

In 2009, J.K. Tucker and M.J. Tenorio proposed a classification system consisting of three distinct families and 82 genera for living species of cone snails. This classification is based on shell morphology, radular differences, anatomy, physiology, and cladistics, with comparisons to molecular (DNA) studies.[37] Published accounts of Conidae that use these new genera include J.K. Tucker & M.J. Tenorio (2009), and Bouchet et al. (2011).[38] Tucker and Tenorio's proposed classification system for the cone shells and other clades of Conoidean gastropods is shown in Tucker & Tenorio cone snail taxonomy 2009.

Some experts, however, still prefer to use the traditional classification. For example, in the November 2011 version of the World Register of Marine Species, all species within the family Conidae were placed in the genus Conus. The binomial names of species in the 82 genera of living cone snails listed in Tucker & Tenorio 2009 were recognized by the World Register of Marine Species as "alternative representations".[39] Debate within the scientific community regarding this issue has continued, and additional molecular phylogeny studies are being carried out in an attempt to clarify the issue.[37][40][41][42][43][44][45][46][47][48]

In 2015, in the Journal of Molluscan Studies, Puillandre, Duda, Meyer, Olivera & Bouchet presented a new classification for the old genus Conus. Using 329 species, the authors carried out molecular phylogenetic analyses. The results suggested that the authors should place all cone snails in a single family, Conidae, containing four genera: Conus, Conasprella, Profundiconus and Californiconus. The authors group 85% of all known cone snail species under Conus. They recognize 57 subgenera within Conus, and 11 subgenera within the genus Conasprella.[2]

Current taxonomy

[edit]

In the Journal of Molluscan Studies, in 2014, Puillandre, Duda, Meyer, Olivera & Bouchet presented a new classification for the old genus Conus. Using 329 species, the authors carried out molecular phylogenetic analyses. The results suggested that the authors should place all living cone snails in a single family, Conidae, containing the following genera:

The authors grouped 85% of all known cone snail species under Conus. They recognized 57 subgenera within Conus, and 11 subgenera within the genus Conasprella.[2]

History of the taxonomy

[edit]

Prior to 1993, the family Conidae contained only Conus species. In 1993 significant taxonomic changes were proposed by Taylor, et al.,:[49] the family Conidae was redefined as several subfamilies. The subfamilies included many subfamilies that had previously been classified in the family Turridae, and the Conus species were moved to the subfamily Coninae.

In further taxonomic changes that took place in 2009 and 2011, based upon molecular phylogeny (see below), the subfamilies that were previously in the family Turridae were elevated to the status of families in their own right. This left the family Conidae once again containing only those species that were traditionally placed in that family: the cone snail species.

1993, Taylor et al., Bouchet & Rocroi

[edit]

According to Taylor, et al. (1993),[49] and the taxonomy of the Gastropoda by Bouchet & Rocroi, 2005,[50] this family consisted of seven subfamilies.

  • Coninae Fleming, 1822 — synonyms: Conulinae Rafinesque, 1815 (inv.); Textiliinae da Motta, 1995 (n.a.)
  • Clathurellinae H. Adams & A. Adams, 1858 — synonyms: Defranciinae Gray, 1853 (inv.); Borsoniinae A. Bellardi, 1875; Pseudotominae A. Bellardi, 1888; Diptychomitrinae L. Bellardi, 1888; Mitrolumnidae Sacco, 1904; Mitromorphinae Casey, 1904; Lorinae Thiele, 1925
  • Conorbiinae de Gregorio, 1880—synonym: Cryptoconinae Cossmann, 1896
  • Mangeliinae P. Fischer, 1883—synonym: Cytharinae Thiele, 1929
  • Oenopotinae Bogdanov, 1987—synonym: Lorinae Thiele, 1925 sensu Thiele
  • Raphitominae A. Bellardi, 1875—synonyms: Daphnellinae Casey, 1904; Taraninae Casey, 1904; Thatcheriidae Powell, 1942; Pleurotomellinae F. Nordsieck, 1968; Andoniinae Vera-Pelaez, 2002
  • † Siphopsinae Le Renard, 1995

2009, Tucker & Tenorio

[edit]

In 2009 John K. Tucker and Manuel J. Tenorio proposed a classification system for the cone shells and their allies (which resorb their inner walls during growth) was based upon a cladistical analysis of anatomical characters including the radular tooth, the morphology (i.e., shell characters), as well as an analysis of prior molecular phylogeny studies, all of which were used to construct phylogenetic trees.[51] In their phylogeny, Tucker and Tenorio noted the close relationship of the cone species within the various clades, corresponding to their proposed families and genera; this also corresponded to the results of prior molecular studies by Puillandre et al. and others.[52][53][54][55][56][57][58] This 2009 proposed classification system also outlined the taxonomy for the other clades of Conoidean gastropods (that do not resorb their inner walls), also based upon morphological, anatomical, and molecular studies, and removes the turrid snails (which are a distinct large and diverse group) from the cone snails, and creates a number of new families.[51] Tucker and Tenorio's proposed classification system for the cone shells and their allies (and the other clades of Conoidean gastropods ) is shown in Tucker & Tenorio cone snail taxonomy 2009.

2011, Bouchet et al.

[edit]

In 2011 Bouchet et al. proposed a new classification in which several subfamilies were raised to the rank of family:[59]

The classification by Bouchet et al. (2011)[59] was based on mitochondrial DNA and nuclear DNA testing, and built on the prior work by J.K. Tucker & M.J. Tenorio (2009), but did not include fossil taxa.[51][59]

Molecular phylogeny, particularly with the advent of nuclear DNA testing in addition to the mDNA testing (testing in the Conidae initially began by Christopher Meyer and Alan Kohn[60]), is continuing on the Conidae.[52][53][54][55][56][57][58][61][62][63][64][65][66][67]

2009, 2011, list of genera from Tucker & Tenorio, and Bouchet et al.

[edit]

This is a list of what were recognized extant genera within Conidae as per J.K. Tucker & M.J. Tenorio (2009), and Bouchet et al. (2011):[51][59] However, all these genera have become synonyms of subgenera within the genus Conus as per the revision of the taxonomy of the Conidae in 2015[2]

  • Afonsoconus Tucker & Tenorio, 2013: synonym of Conus (Afonsoconus) Tucker & Tenorio, 2013 represented as Conus Linnaeus, 1758
  • Africonus Petuch, 1975: synonym of Conus (Lautoconus) Monterosato, 1923 represented as Conus Linnaeus, 1758
  • Arubaconus Petuch, 2013: synonym of Conus (Ductoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Asprella Schaufuss, 1869: synonym of Conus (Asprella) Schaufuss, 1869 represented as Conus Linnaeus, 1758
  • Atlanticonus Petuch & Sargent, 2012: synonym of Conus (Atlanticonus) Petuch & Sargent, 2012 represented as Conus Linnaeus, 1758
  • Attenuiconus Petuch, 2013: synonym of Conus (Attenuiconus) Petuch, 2013 represented as Conus Linnaeus, 1758
  • Austroconus Tucker & Tenorio, 2009 synonym of Conus (Austroconus) Tucker & Tenorio, 2009 represented as Conus Linnaeus, 1758
  • Bathyconus Tucker & Tenorio, 2009: synonym of Conasprella (Fusiconus) Thiele, 1929, represented as Conasprella Thiele, 1929
  • Bermudaconus Petuch, 2013: synonym of Conus (Bermudaconus) Petuch, 2013 represented as Conus Linnaeus, 1758
  • Boucheticonus Tucker & Tenorio, 2013: synonym of Conasprella (Boucheticonus) Tucker & Tenorio, 2013 represented as Conasprella Thiele, 1929
  • Brasiliconus Petuch, 2013: synonym of Conus (Brasiliconus) Petuch, 2013 represented as Conus Linnaeus, 1758
  • Calamiconus Tucker & Tenorio, 2009: synonym of Conus (Lividoconus) Wils, 1970 represented as Conus Linnaeus, 1758
  • Calibanus da Motta, 1991: synonym of Conus (Calibanus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Cariboconus Petuch, 2003: synonym of Conus (Dauciconus) Cotton, 1945 represented as Conus Linnaeus, 1758
  • Californiconus Tucker & Tenorio, 2009
  • Chelyconus Mörch, 1852: synonym of Conus (Chelyconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Cleobula Iredale, 1930: synonym of Dendroconus Swainson, 1840
  • Coltroconus Petuch, 2013: synonym of Conasprella (Coltroconus) Petuch, 2013 represented as Conasprella Thiele, 1929
  • Conasprella Thiele, 1929: accepted name
  • Conasprelloides Tucker & Tenorio, 2009: synonym of Conus (Dauciconus) Cotton, 1945 represented as Conus Linnaeus, 1758
  • † Conilithes Swainson, 1840
  • Continuconus Tucker & Tenorio, 2013
  • Conus Linnaeus, 1758: accepted name
  • Cornutoconus Suzuki, 1972: synonym of Taranteconus Azuma, 1972
  • Coronaxis Swainson, 1840: synonym of Conus (Conus) Linnaeus, 1758 represented as Conus Linnaeus, 1758
  • Cucullus Röding, 1798: synonym of Conus (Conus) Linnaeus, 1758 represented as Conus Linnaeus, 1758
  • Cylinder Montfort, 1810: synonym of Conus (Cylinder) Montfort, 1810 represented as Conus Linnaeus, 1758
  • Cylindrella Swainson, 1840: synonym of Asprella Schaufuss, 1869synonym of Conus (Asprella) Schaufuss, 1869 represented as Conus Linnaeus, 1758
  • Cylindrus Batsch, 1789: synonym of Cylinder Montfort, 1810synonym of Conus (Cylinder) Montfort, 1810 represented as Conus Linnaeus, 1758
  • Dalliconus Tucker & Tenorio, 2009: synonym of Conasprella (Dalliconus) Tucker & Tenorio, 2009 synonym of Conasprella Thiele, 1929
  • Darioconus Iredale, 1930: synonym of Conus (Darioconus) Iredale, 1930 represented as Conus Linnaeus, 1758
  • Dauciconus Cotton, 1945: synonym of Conus (Dauciconus) Cotton, 1945 represented as Conus Linnaeus, 1758
  • Dendroconus Swainson, 1840: synonym of Conus (Dendroconus) Swainson, 1840 represented as Conus Linnaeus, 1758
  • Ductoconus da Motta, 1991: synonym of Conus (Ductoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Duodenticonus Tucker & Tenorio, 2013: synonym of Conasprella (Conasprella) Thiele, 1929 represented as Conasprella Thiele, 1929
  • Dyraspis Iredale, 1949: synonym of Conus (Virroconus) Iredale, 1930 represented as Conus Linnaeus, 1758
  • Elisaconus Tucker & Tenorio, 2013: synonym of Conus (Elisaconus) Tucker & Tenorio, 2013 represented as Conus Linnaeus, 1758
  • Embrikena Iredale, 1937: synonym of Conus (Embrikena) Iredale, 1937 represented as Conus Linnaeus, 1758
  • Endemoconus Iredale, 1931: synonym of Conasprella (Endemoconus) Iredale, 1931 represented as Conasprella Thiele, 1929
  • Eremiconus Tucker & Tenorio, 2009: synonym of Conus (Eremiconus) Tucker & Tenorio, 2009 represented as Conus Linnaeus, 1758
  • Erythroconus da Motta, 1991: synonym of Conus (Darioconus) Iredale, 1930 represented as Conus Linnaeus, 1758
  • Eugeniconus da Motta, 1991: synonym of Conus (Eugeniconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Floraconus Iredale, 1930: synonym of Conus (Floraconus) Iredale, 1930 represented as Conus Linnaeus, 1758
  • Fraterconus Tucker & Tenorio, 2013: synonym of Conus (Fraterconus) Tucker & Tenorio, 2013 represented as Conus Linnaeus, 1758
  • Fulgiconus da Motta, 1991: synonym of Conus (Phasmoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Fumiconus da Motta, 1991: synonym of Conasprella (Fusiconus) da Motta, 1991 represented as Conasprella Thiele, 1929
  • Fusiconus da Motta, 1991: synonym of Conasprella (Fusiconus) da Motta, 1991 represented as Conasprella Thiele, 1929
  • Gastridium Modeer, 1793: synonym of Conus (Gastridium) Modeer, 1793 represented as Conus Linnaeus, 1758
  • Genuanoconus Tucker & Tenorio, 2009: synonym of Conus (Kalloconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Gladioconus Tucker & Tenorio, 2009: synonym of Conus (Monteiroconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Globiconus Tucker & Tenorio, 2009: synonym of Conasprella (Ximeniconus) Emerson & Old, 1962 represented as Conasprella Thiele, 1929
  • Gradiconus da Motta, 1991: synonym of Conus (Dauciconus) Cotton, 1945 represented as Conus Linnaeus, 1758
  • Graphiconus da Motta, 1991: synonym of Conus (Phasmoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Harmoniconus da Motta, 1991: synonym of Conus (Harmoniconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Hermes Montfort, 1810: synonym of Conus (Hermes) Montfort, 1810 represented as Conus Linnaeus, 1758
  • Heroconus da Motta, 1991: synonym of Conus (Pionoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Isoconus Tucker & Tenorio, 2013: synonym of Conus (Splinoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Jaspidiconus Petuch, 2004: synonym of Conasprella (Ximeniconus) Emerson & Old, 1962 represented as Conasprella Thiele, 1929
  • Kalloconus da Motta, 1991: synonym of Conus (Kalloconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Kellyconus Petuch, 2013: synonym of Conus (Kellyconus) Petuch, 2013 represented as Conus Linnaeus, 1758
  • Kenyonia Brazier, 1896: genus incertae sedis
  • Kermasprella Powell, 1958: synonym of Conasprella (Endemoconus) Iredale, 1931 represented as Conasprella Thiele, 1929
  • Ketyconus da Motta, 1991: synonym of Conus (Floraconus) Iredale, 1930 represented as Conus Linnaeus, 1758
  • Kioconus da Motta, 1991: synonym of Conus (Splinoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Klemaeconus Tucker & Tenorio, 2013: synonym of Conus (Klemaeconus) Tucker & Tenorio, 2013 represented as Conus Linnaeus, 1758
  • Kohniconus Tucker & Tenorio, 2009: synonym of Conasprella (Kohniconus) Tucker & Tenorio, 2009 represented as Conasprella Thiele, 1929
  • Kurodaconus Shikama & Habe, 1968: synonym of Conus (Turriconus) Shikama & Habe, 1968 represented as Conus Linnaeus, 1758
  • Lamniconus da Motta, 1991: synonym of Conus (Lamniconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Lautoconus Monterosato, 1923: synonym of Conus (Lautoconus) Monterosato, 1923 represented as Conus Linnaeus, 1758
  • Leporiconus Iredale, 1930: synonym of Conus (Leporiconus) Iredale, 1930 represented as Conus Linnaeus, 1758
  • Leptoconus Swainson, 1840: synonym of Conus (Leptoconus) Swainson, 1840 represented as Conus Linnaeus, 1758
  • Lilliconus Raybaudi Massilia, 1994: synonym of Conasprella (Lilliconus) G. Raybaudi Massilia, 1994 represented as Conasprella Thiele, 1929
  • Lindaconus Petuch, 2002: synonym of Conus (Lindaconus) Petuch, 2002 represented as Conus Linnaeus, 1758
  • Lithoconus Mörch, 1852: synonym of Conus (Lithoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Lividoconus Wils, 1970: synonym of Conus (Lividoconus) Wils, 1970 represented as Conus Linnaeus, 1758
  • Lizaconus da Motta, 1991synonym of Profundiconus Kuroda, 1956
  • Magelliconus da Motta, 1991: synonym of Conus (Dauciconus) Cotton, 1945 represented as Conus Linnaeus, 1758
  • Malagasyconus Monnier & Tenorio, 2015
  • Mamiconus Cotton & Godfrey, 1932: synonym of Endemoconus Iredale, 1931synonym of Conasprella (Endemoconus) Iredale, 1931 represented as Conasprella Thiele, 1929
  • Miliariconus Tucker & Tenorio, 2009: synonym of Conus (Virroconus) Iredale, 1930 represented as Conus Linnaeus, 1758
  • Mitraconus Tucker & Tenorio, 2013: synonym of Conus (Turriconus) Shikama & Habe, 1968 represented as Conus Linnaeus, 1758
  • Monteiroconus da Motta, 1991: synonym of Conus (Monteiroconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Nataliconus Tucker & Tenorio, 2009: synonym of Conus (Leptoconus) Swainson, 1840 represented as Conus Linnaeus, 1758
  • Nimboconus Tucker & Tenorio, 2013: synonym of Conus (Phasmoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Nitidoconus Tucker & Tenorio, 2013: synonym of Conus (Splinoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Ongoconus da Motta, 1991: synonym of Conus (Splinoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Papyriconus Tucker & Tenorio, 2013: synonym of Conus (Papyriconus) Tucker & Tenorio, 2013 represented as Conus Linnaeus, 1758
  • Parviconus Cotton & Godfrey, 1932: synonym of Conasprella (Parviconus) Cotton & Godfrey, 1932 represented as Conasprella Thiele, 1929
  • Perplexiconus Tucker & Tenorio, 2009: synonym of Conasprella (Ximeniconus) Emerson & Old, 1962 represented as Conasprella Thiele, 1929
  • Phasmoconus Mörch, 1852: synonym of Conus (Phasmoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Pionoconus Mörch, 1852: synonym of Conus (Pionoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Plicaustraconus Moolenbeek, 2008: synonym of Conus (Plicaustraconus) Moolenbeek, 2008 represented as Conus Linnaeus, 1758
  • Poremskiconus Petuch, 2013: synonym of Conus (Dauciconus) Cotton, 1945 represented as Conus Linnaeus, 1758
  • Profundiconus Kuroda, 1956: accepted name
  • Protoconus da Motta, 1991: synonym of Tenorioconus Petuch & Drolshagen, 2011
  • Protostrioconus Tucker & Tenorio, 2009: synonym of Conus (Gastridium) Modeer, 1793 represented as Conus Linnaeus, 1758
  • Pseudoconorbis Tucker & Tenorio, 2009: synonym of Conasprella (Pseudoconorbis) Tucker & Tenorio, 2009, represented as Conasprella Thiele, 1929
  • Pseudohermes Tucker & Tenorio, 2013: synonym of Conus (Virgiconus) Cotton, 1945 represented as Conus Linnaeus, 1758
  • Pseudolilliconus Tucker & Tenorio, 2009: synonym of Conus (Pseudolilliconus) Tucker & Tenorio, 2009 represented as Conus Linnaeus, 1758
  • Pseudonoduloconus Tucker & Tenorio, 2009: synonym of Conus (Pseudonoduloconus) Tucker & Tenorio, 2009 represented as Conus Linnaeus, 1758
  • Pseudopterygia Tucker & Tenorio, 2013: synonym of Conus (Pseudopterygia) Tucker & Tenorio, 2013 represented as Conus Linnaeus, 1758
  • Puncticulis Swainson, 1840: synonym of Conus (Puncticulis) Swainson, 1840 represented as Conus Linnaeus, 1758
  • Purpuriconus da Motta, 1991: synonym of Conus (Dauciconus) Cotton, 1945 represented as Conus Linnaeus, 1758
  • Pygmaeconus Puillandre & Tenorio, 2017
  • Pyruconus Olsson, 1967: synonym of Conus (Pyruconus) Olsson, 1967 represented as Conus Linnaeus, 1758
  • Quasiconus Tucker & Tenorio, 2009: synonym of Conus (Quasiconus) Tucker & Tenorio, 2009 represented as Conus Linnaeus, 1758
  • Regiconus Iredale, 1930: synonym of Conus (Darioconus) Iredale, 1930 represented as Conus Linnaeus, 1758
  • Rhizoconus Mörch, 1852: synonym of Conus (Rhizoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Rhombiconus Tucker & Tenorio, 2009: synonym of Conus (Stephanoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Rhombus Montfort, 1810: synonym of Rhombiconus Tucker & Tenorio, 2009, synonym of Conus (Stephanoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Rolaniconus Tucker & Tenorio, 2009: synonym of Conus (Strategoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Rollus Montfort, 1810 :synonym of Conus (Gastridium) Modeer, 1793 represented as Conus Linnaeus, 1758
  • Rubroconus Tucker & Tenorio, 2013: synonym of Conus (Rubroconus) Tucker & Tenorio, 2013 represented as Conus Linnaeus, 1758
  • Sandericonus Petuch, 2013: synonym of Conus (Sandericonus) Petuch, 2013 represented as Conus Linnaeus, 1758
  • Sciteconus da Motta, 1991: synonym of Conus (Sciteconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Seminoleconus Petuch, 2003: synonym of Conus (Stephanoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Socioconus da Motta, 1991: synonym of Conus (Pionoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Splinoconus da Motta, 1991: synonym of Conus (Splinoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Spuriconus Petuch, 2003: synonym of Conus (Lindaconus) Petuch, 2002 represented as Conus Linnaeus, 1758
  • Stellaconus Tucker & Tenorio, 2009: synonym of Conus (Splinoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Stephanoconus Mörch, 1852: synonym of Conus (Stephanoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Strategoconus da Motta, 1991: synonym of Conus (Strategoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Strioconus Thiele, 1929: synonym of Pionoconus Mörch, 1852, synonym of Conus (Pionoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Sulciconus Bielz, 1869: synonym of Asprella Schaufuss, 1869, synonym of Conus (Asprella) Schaufuss, 1869 represented as Conus Linnaeus, 1758
  • Taranteconus Azuma, 1972: synonym of Conus (Stephanoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Tenorioconus Petuch & Drolshagen, 2011: synonym of Conus (Stephanoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Tesselliconus da Motta, 1991: synonym of Conus (Tesselliconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Textilia Swainson, 1840: synonym of Conus (Textilia) Swainson, 1840 represented Conus Linnaeus, 1758
  • Thalassiconus Tucker & Tenorio, 2013: synonym of Calibanus da Motta, 1991, synonym of Conus (Calibanus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Theliconus Swainson, 1840: synonym of Hermes Montfort, 1810, synonym of Conus (Hermes) Montfort, 1810 represented as Conus Linnaeus, 1758
  • Thoraconus da Motta, 1991: synonym of Fulgiconus da Motta, 1991, synonym of Conus (Phasmoconus) Mörch, 1852 represented as Conus Linnaeus, 1758
  • Trovaoconus Tucker & Tenorio, 2009, synonym of Conus (Kalloconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Tuckericonus Petuch, 2013: synonym of Conus (Dauciconus) Cotton, 1945 represented as Conus Linnaeus, 1758
  • Tuliparia Swainson, 1840: synonym of Gastridium Modeer, 1793, synonym of Conus (Gastridium) Modeer, 1793 represented as Conus Linnaeus, 1758
  • Turriconus Shikama & Habe, 1968, synonym of Conus (Turriconus) Shikama & Habe, 1968 represented as Conus Linnaeus, 1758
  • Utriculus Schumacher, 1817: synonym of Gastridium Modeer, 1793, synonym of Conus (Gastridium) Modeer, 1793 represented as Conus Linnaeus, 1758
  • Varioconus da Motta, 1991: synonym of Conus (Lautoconus) Monterosato, 1923 represented as Conus Linnaeus, 1758
  • Viminiconus Tucker & Tenorio, 2009: synonym of Conasprella (Fusiconus) da Motta, 1991 represented as Conasprella Thiele, 1929
  • Virgiconus Cotton, 1945: synonym of Conus (Virgiconus) Cotton, 1945 represented as Conus Linnaeus, 1758
  • Virroconus Iredale, 1930: synonym of Conus (Virroconus) Iredale, 1930 represented as Conus Linnaeus, 1758
  • Vituliconus da Motta, 1991: synonym of Conus (Strategoconus) da Motta, 1991 represented as Conus Linnaeus, 1758
  • Ximeniconus Emerson & Old, 1962: synonym of Conasprella (Ximeniconus) Emerson & Old, 1962 represented as Conasprella Thiele, 1929
  • Yeddoconus Tucker & Tenorio, 2009: synonym of Conasprella (Endemoconus) Iredale, 1931 represented as Conasprella Thiele, 1929

1993 to 2011 list of genera

[edit]

Following Taylor et al., from 1993 to 2011, the family Conidae was defined as including not only the cone snails, but also a large number of other genera which are commonly known as "turrids". However, as a result of molecular phylogeny studies in 2011, many of those genera were moved back to the Turridae, or were placed in new "turrid" families within the superfamily Conoidea. The following list of genera that used to be included in Conidae is retained as a historical reference:


See also

[edit]
  • ConoServer, a database of cone snail toxins, known as conopeptides.[68] These toxins are of importance to medical research.
  • Conotoxin

References

[edit]

Further reading

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Cone snail

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Cone snails are predatory marine gastropods belonging to the family Conidae, renowned for their distinctive cone-shaped shells and sophisticated venom delivery systems. These snails, primarily in the Conus, comprise approximately 850 extant that inhabit tropical and subtropical ocean waters worldwide, often in shallow coastal environments such as coral reefs, sandy flats, and seagrass beds, though some dwell at depths up to several hundred meters. As active hunters, cone snails employ a extendable proboscis to deploy a harpoon-like radular tooth that injects a complex venom cocktail known as conotoxins, which are disulfide-rich peptides targeting channels, receptors, and transporters to paralyze prey. Their diet varies by species and evolutionary lineage: vermivorous types primarily consume polychaete worms, molluscivorous ones prey on other gastropods, and piscivorous species hunt , with the latter often possessing the most potent venoms. This specialized predation has driven remarkable venom diversity, with each species producing hundreds of unique conotoxins tailored to their ecological niche. While most encounters with humans are harmless, stings from certain piscivorous cone snails can cause severe , including and, in rare cases, , due to the neurotoxic effects of their . Conversely, the pharmacological potential of conotoxins has led to significant biomedical applications; for instance, , derived from the venom of , is an FDA-approved analgesic for . Conservation efforts highlight threats from habitat loss and shell collection, underscoring the need to protect this biodiverse group.

Description and anatomy

Shell

The shells of cone snails exhibit an elongated, conical shape, typically forming an inverted with a narrow anterior end, a long and slender , and a short siphonal . These dimensions allow for efficient retraction of the soft body while maintaining a streamlined profile. Shell lengths vary widely among , generally ranging from 10 to 230 mm. Surface features of cone snail shells include axial ribs, nodules, and spiral ridges that contribute to structural reinforcement and visual distinction. Intricate color patterns, often featuring bands, spots, or mottling in , , and , provide against sandy or substrates and assist in identification. The shell's material composition consists primarily of in the polymorph, arranged in a crossed-lamellar microstructure of rod-shaped that enhances toughness and fracture resistance. An outer periostracum layer, composed of organic proteins, covers the shell and offers initial against environmental abrasion. These shells serve adaptive roles in predator protection, with their robust, layered structure deterring crushing by larger marine animals such as octopuses or . Additionally, the shell provides , anchoring muscles that enable the projection of the during feeding. Variations in shell morphology occur across , with some displaying smooth surfaces and others featuring pronounced sculpturing like ribs or tubercles. Color morphs are more vibrant and diverse in tropical , aiding in reefs, while temperate tend toward subdued patterns better suited to cooler, sediment-rich environments. Shell morphology also plays a crucial role in taxonomic , as differences in , ornamentation, and patterning help delineate boundaries.

Radula and harpoon

In cone snails, the has undergone significant modification compared to the typical gastropod structure, which consists of a flexible bearing multiple rows of chitinous arranged in transverse series for scraping or cutting food. Instead, the Conus is reduced to a single per row, forming a specialized, hollow, -like structure that serves as the primary tool for prey capture. This single , known as the radular , is barbed to anchor into the prey and features a that allows for upon deployment. The tooth exhibits a distinct tripartite morphology consisting of a bulbous base, a narrow shaft, and a bladed tip equipped with backward-facing barbs. The bulbous base attaches to the for loading and propulsion, the hollow shaft provides structural support and serves as a conduit for delivery, and the bladed tip facilitates penetration and securement of the prey. This design enables the to function like a , piercing tissues to inject directly. The integrates with the apparatus by drawing fluid from the venom bulb through its hollow core during projection. Projection of the occurs via a hydrostatic mechanism, where pressurized fluid from the venom bulb builds within the extended , engaging a cellular at the base until release. Upon prey contact, the disengages, propelling the forward at peak velocities averaging 19.3 m/s and exceeding 25 m/s in fish-hunting species, with average peak accelerations exceeding 280,000 m/s² (approximately 28,600 g) and maximal accelerations over 400,000 m/s² (about 40,800 g). This rapid deployment represents an evolutionary adaptation for overcoming the escape responses of mobile prey like . Evolutionarily, the Conus radula's specialization as a disposable, single-use reflects adaptations from ancestral worm-hunting forms, with piscivorous lineages independently evolving enhanced barbs and speed for tethering larger prey. Each is expended after use—often regurgitated shortly thereafter—and the radular sac continuously produces replacements, regenerating a new within days to maintain readiness for subsequent hunts. This disposable nature contrasts sharply with the multi-tooth, reusable radulae of other gastropods, underscoring the Conidae's shift toward venom-mediated predation.

Venom apparatus

The venom apparatus of cone snails comprises an unpaired venom gland, a venom duct, and a muscular . The venom gland is a long, highly convoluted structure extending from the posterior end of the animal, lined with glandular responsible for synthesis, and connected to the via the duct. The duct facilitates the transport of components, while the muscular bulb at the proximal end of the gland serves for storage and generates hydrostatic pressure to propel the venom during injection. Cone snail venom consists of a complex mixture of 50 to 200 conotoxins per , primarily small disulfide-rich peptides known as conopeptides that target specific neuronal components. These include alpha-conotoxins, which antagonize nicotinic receptors; omega-conotoxins, which block N-type voltage-gated calcium channels; and mu-conotoxins, which inhibit voltage-gated sodium channels, among other families. Conotoxins are produced through the synthesis of precursor proteins in the glandular of the , where they undergo post-translational processing, including disulfide bond formation and proteolytic cleavage. Genetic diversity arises from hypermutation in conotoxin-encoding genes, enabling rapid and species-specific variations in sequences and structures. Venom composition exhibits significant variability across cone snail species, tailored to their dietary preferences: approximately 70% of species are vermivores with venoms optimized for immobilizing polychaete worms, 15% are molluscivores targeting other gastropods, and 15% are piscivores adapted for capturing . This prey-specific ensures efficient strategies distinct from those in other clades. The evolutionary refinement of conotoxins over millions of years has positioned them as valuable models for , owing to their precise targeting of channels and receptors honed by for prey capture.

Habitat and ecology

Distribution

Cone snails are predominantly distributed in tropical and subtropical marine waters worldwide, exhibiting their highest species diversity in the region, where the majority of the over 850 described occur. This region accounts for approximately 60% of known cone snail diversity, spanning vast coral reef systems from the in the west to in the east. In contrast, species richness is considerably lower in the Atlantic Ocean and eastern Pacific, with around 98 species documented in the eastern Atlantic and even fewer in the western Atlantic and eastern Pacific regions. These gastropods occupy a wide bathymetric range, from shallow intertidal zones to depths exceeding 700 meters, though most species prefer shallow waters less than 50 meters deep. Deep-water species, particularly those in genera such as Profundiconus, extend this range further, with records from collections reaching up to 1,260 meters in areas like New Caledonia's . Endemism is a prominent feature of cone snail distribution, with numerous confined to specific islands or archipelagos due to biogeographic isolation. For instance, the Islands host 55 , 52 of which are endemic (noting one recent as of 2025), highlighting hotspots of localized diversity in the eastern Atlantic. Similarly, many Indo-Pacific islands support unique assemblages, contributing to the overall pattern of restricted ranges observed in about 37.5% of with areas of occupancy under 100 km². Limited larval dispersal plays a key role in shaping these distribution patterns, as most cone snails produce planktotrophic larvae that, despite enabling some oceanic transport, often result in constrained by oceanographic barriers and developmental constraints. This contributes to low migration rates across major biogeographic divides, such as those separating the from the Atlantic and eastern Pacific.

Preferred environments

Cone snails, genus Conus, predominantly occupy tropical and subtropical marine habitats, with a strong preference for coral reefs where they achieve the highest species diversity and abundance. They also inhabit sandy or muddy bottoms, beds, and fringes, often in shallow coastal waters from the to depths of around 50 meters. These environments provide the structural complexity and prey availability essential for their predatory lifestyle. Within these habitats, cone snails exploit diverse microhabitats, such as burrowing partially or fully into loose sediment on flats or floors to remain concealed during daylight hours, or attaching to rocks, rubble, and sponges for stability. Many species are nocturnal, emerging at night to while retreating to protective crevices, algal mats, or under overhangs during the day to avoid predators and in intertidal areas. For instance, species like frequently burrow in near reefs, enhancing their capabilities. Cone snails exhibit environmental tolerances suited to stable tropical conditions, typically thriving in seawater salinities of 30–35 ppt and temperatures ranging from 20–30°C, though some can endure broader fluctuations such as salinities from 5–40 ppt under optimal temperatures around 20°C. Certain deep-water or sediment-dwelling show adaptations to lower oxygen levels, such as in hypoxic environments or deeper slopes. These tolerances reflect their reliance on consistent and Atlantic coastal ecosystems. Adaptations to these environments include intricate shell coloration and patterning that provide camouflage against sandy, rubbly, or coral backgrounds, allowing the snails to blend seamlessly with their surroundings for predatory ambushes. Burrowing behaviors, facilitated by a muscular foot, enable them to create temporary shelters in sediment while maintaining sensory awareness through extended siphons. Such traits are particularly evident in reef-associated species, where shell patterns mimic surrounding algae or rubble. Habitat change poses acute risks to cone snails, particularly those reliant on reefs, where bleaching events driven by rising sea temperatures and degrade structural habitats and reduce microhabitat availability. For example, widespread mortality has led to population declines in reef-dependent species across the , disrupting their preferred niches.

Feeding behavior

Cone snails are classified into three primary dietary guilds based on their prey preferences: piscivorous species that primarily hunt fish, molluscivorous species that target other mollusks such as other snails, and vermivorous species that feed on polychaete worms and similar invertebrates. This specialization influences their venom composition and hunting efficiency, with piscivorous species generally exhibiting the most potent and rapid-acting toxins to subdue fast-moving fish. For instance, species like Conus geographus and Conus textile are piscivorous and rely on fish as their main prey. Hunting strategies among cone snails emphasize stealth and precision, often involving ambush predation where the snail remains camouflaged on the substrate until prey approaches within striking distance. Piscivorous frequently employ a lightning strike technique, rapidly everting the —up to 20 times their body length in milliseconds—to impale the prey with a harpoon-like radular . Alternatively, some utilize a "net-hunting" or "cabling" strategy, extending the in a lasso-like manner to ensnare and tether before , as observed in like Conus catus. Vermivorous and molluscivorous tend toward slower, more deliberate approaches, such as probing sediments or luring prey with protrusible structures mimicking food or mates. The process is initiated by the strike, which injects a complex mixture of through the apparatus, leading to near-instantaneous —often within 2-5 seconds for prey—by disrupting neuromuscular transmission and channels. Following immobilization, external are sometimes released to begin liquefying the prey's tissues externally, facilitating consumption. The entire prey is then engulfed whole via the highly extensible and mouth, with ongoing effects preventing any post-capture resistance; indigestible remnants, such as shells or exoskeletons, are expelled hours later. Behavioral variations in feeding reflect ecological adaptations, with many species exhibiting nocturnal to exploit crepuscular prey activity, though some temperate piscivores like Conus californicus hunt diurnally in response to availability. Active involves siphon-mediated prey detection and pursuit over short distances, contrasting with passive tactics where snails position themselves in high-traffic prey areas and wait. These patterns can shift ontogenetically, as seen in , where juveniles are strictly vermivorous with cautious probing behaviors before transitioning to piscivory in adulthood via bolder strikes.

Evolutionary history

Paleontology

The fossil record of cone snails (genus Conus) dates back to the early Eocene epoch, approximately 55 million years ago, when primitive Conus-like forms first appeared in shallow marine deposits across what is now . These earliest known s, including such as Eoconus edwardsi (Hampshire Basin) and E. deperditus (Paris Basin), have been recovered from Lower Eocene sediments in the of and the Hampshire Basin of , indicating an origin in temperate to subtropical shelf environments during a period of global warming following the Paleocene-Eocene Thermal Maximum. Fossil diversity expanded significantly through the , with over 1,000 described extinct species documented to date, though many may represent synonyms due to similarities in shell morphology. Diversity peaked during the epoch (23–5 million years ago), a time of rapid and geographic radiation coinciding with the expansion of tropical shallow-water habitats and the closure of the Tethys Sea, which facilitated dispersal into the and Atlantic regions. Key fossil assemblages from this period include those from coral reef deposits in the , where well-preserved shells reveal insights into coloration patterns and ecological roles, and chalk and limestone formations in , such as those in and the Mediterranean Basin. Morphological evolution in the fossil record reflects adaptations to predatory lifestyles, with shell coiling transitioning from loosely coiled, more globose forms in Eocene ancestors to the tightly coiled, elongated conical shapes characteristic of later , enhancing mobility and in environments. Concurrently, the underwent specialization, evolving from a multi- configuration typical of ancestral vermivorous neogastropods to the single, hypodermic harpoon-like seen in modern molluscivorous and piscivorous Conus, enabling precise delivery for prey capture. These changes are evident in comparative analyses of radular remnants and shell microstructures from and sites. Significant extinctions occurred during the Pleistocene epoch (2.6 million–11,700 years ago), driven by repeated glacio-eustatic sea-level fluctuations that contracted shallow tropical habitats, particularly affecting island-endemic and reef-associated species in the western Atlantic and . Fossil evidence from deposits shows a decline in and average body size among surviving lineages, with approximately 70% of regional Conus diversity lost in the western Atlantic due to and reduced nutrient availability.

Phylogenetic relationships

Cone snails, belonging to the family Conidae, are placed within the superfamily Conoidea of the order , formerly grouped under the toxoglossan gastropods. This positioning reflects their shared characteristics with other venomous marine gastropods, including a harpoon-like for prey . Within Conoidea, molecular phylogenies indicate that Conidae is monophyletic and forms a distinct lineage, with some analyses suggesting a close relationship to (auger snails) as part of a broader diversification of toxiferous families. The family underwent significant diversification approximately 50 million years ago during the Eocene, coinciding with the radiation of and the evolution of complex venom systems. Large-scale molecular studies, incorporating (e.g., COI ) and nuclear markers (e.g., 28S rRNA and ), reveal a rapid radiation of Conidae primarily in the region, the ancestral cradle of the group. This phylogeny, based on over , identifies four major highly divergent clades (Clades A–D), with limited subsequent dispersal to other oceans, underscoring the Indo-Pacific's role in driving through ecological opportunities like diverse prey availability. Phylogenetic analyses further delineate key clades by dietary specialization, separating piscivorous (fish-hunting) lineages from non-piscivorous ones (e.g., vermivorous or molluscivorous). Piscivory has arisen independently at least three times, in distinct subclades such as those including and , highlighting in venom composition and hunting strategies. Non-piscivorous clades, often basal, retain ancestral worm- or mollusk-hunting behaviors, with transitions to fish predation linked to innovations in conotoxin diversity. The of Conus sensu lato has been challenged by these molecular data, which demonstrate within the traditional genus, prompting a 2014 reclassification that recognizes multiple genera across the four clades to better reflect evolutionary relationships. This shift emphasizes the polyphyletic nature of shell-based taxonomy and advocates for integrating genetic evidence in Conidae systematics.

Life history

Reproduction

Cone snails are gonochoristic, possessing separate sexes with distinct s and s. occurs during copulation, in which the mounts the using its foot and inserts a ribbon-like verge—analogous to a —into the 's mantle cavity opening to transfer . is facilitated by chemical cues, as individuals secrete pheromones to attract potential partners; the then approaches the , and copulation may last several hours depending on the . Following fertilization, females deposit eggs in gelatinous capsules attached to hard substrates such as rocks or , often in clusters to protect against environmental hazards. Each capsule typically contains dozens to hundreds of eggs, with species like producing about 40 eggs per capsule in masses of multiple capsules, while others may yield up to 1,000 eggs per capsule across 20–50 capsules per mass. Within these capsules, embryos develop either into free-swimming veliger larvae, which hatch and enter the , or directly into juvenile snails via intracapsular , depending on the species' life history strategy. Cone snails exhibit no after egg deposition; females abandon the capsules, relying on high —often thousands of eggs per reproductive event—to offset high rates of predation and mortality in early life stages. is minimal across the , with males and females generally similar in shell shape, color, and size, though some show slight differences in maximum adult size, such as females growing marginally larger in certain vermivorous taxa.

Development and growth

Cone snails display two primary larval developmental modes: planktotrophic veligers, which are free-swimming larvae that actively feed on to fuel their growth, and lecithotrophic larvae, which are non-feeding and depend on internal yolk reserves for energy during their brief pelagic phase. Most species in the Conus employ planktotrophic development, allowing larvae to sustain longer planktonic periods, whereas lecithotrophic development occurs in select species, such as those endemic to the archipelago, where larvae derive nourishment from an egg sac without external feeding. In planktotrophic species, veliger larvae can disperse for 10 to 50 days—or up to several weeks to months in natural conditions—via currents, facilitating the wide geographic distribution observed across tropical and subtropical marine environments. This pelagic phase ends with settlement onto appropriate substrates, such as sandy or coralline bottoms, triggering ; during this process, the velum (a larval organ) regresses, the operculum forms, and the protoconch transitions into the adult teleoconch shell, marking the onset of benthic life. Following , juvenile cone snails exhibit steady shell growth, typically at rates of 1 to 2 mm per month in shallow-water like Conus pennaceus, though variability exists across taxa due to environmental factors such as and availability. is generally attained within 1 to 3 years, with smaller reaching reproductive size sooner (e.g., 6 to 12 months in Conus geographus) compared to larger forms. Lifespans range from 5 to 20 years, influenced by ; deep-water often display slower growth rates and extended longevity owing to lower metabolic demands in cooler, stable environments.

Taxonomy and classification

Current taxonomy

The family Conidae comprises over 1,000 extant species of cone snails, classified into 8 accepted genera and numerous subgenera following updates to the 2015 taxonomic revision. This revision, building on a comprehensive molecular phylogeny, restructured the traditional broad genus Conus to reflect evolutionary relationships more accurately. The current system, as endorsed by the (WoRMS) as of 2025, recognizes the family Conidae with 8 genera: Californiconus, Conasprella, Conus, Kenyonia, Lilliconus, Profundiconus, Pseudolilliconus, and Pygmaeconus, along with many subgenera. For instance, Conus includes well-known subgroups like the textile cones (Conus textile and relatives), while Conasprella, Profundiconus, and others represent distinct lineages adapted to varied habitats, such as deep-water environments in the latter. This framework has evolved from the 803 valid species recognized in 2015, with subsequent additions and reclassifications bringing the total over 1,000. The classification criteria integrate molecular data from mitochondrial genes, including subunit I (COI), 16S rRNA, and 12S rRNA, analyzed across 329 species, with supporting morphological evidence from shell morphology and radular dentition. These approaches revealed four major divergent clades initially, but subsequent studies have justified further separations into additional genera. Post-2015 updates have included descriptions of new deep-water species and taxonomic adjustments, such as the 2023 review of New Caledonian fauna that described one new species (Conus samadiae) but preserved the overall structure while noting ongoing refinements. As of November 2025, the system continues to evolve, with WoRMS reflecting the latest accepted genera.

Historical developments

The genus Conus was established by in his (10th edition) in , initially encompassing all known cone snails within a single , with approximately 30 species described based on shell characteristics from global collections. This Linnaean framework treated the diverse morphologies of cone snails as variations within one , reflecting the limited understanding of their biology at the time. Throughout the 19th and early 20th centuries, taxonomists expanded the by introducing subgenera primarily based on shell , , and coloration, as anatomical details like the apparatus were not yet central to . A key contribution came from William J. Clench in 1942, who proposed subgenera such as Dauciconus and Jaspidiconus for western Atlantic , emphasizing regional shell variations to organize the growing number of described taxa. By the , around 500 were recognized within Conus, highlighting the genus's remarkable diversity but also the challenges in delineating boundaries solely on morphological grounds. In 1993, John D. Taylor and colleagues provided the first detailed anatomical classification of the superfamily Conoidea, analyzing structures and feeding mechanisms across families, which suggested the of Conus by revealing deep divergences unsupported by traditional shell-based groupings. This work marked a shift toward integrating internal , laying groundwork for questioning the of the genus. Building on emerging molecular data, John K. Tucker and Manuel J. Tenorio proposed a major revision in 2009, elevating many subgenera to full genera and recognizing 82 genera within three families for over 600 living cone snail species, based on combined morphological and preliminary genetic evidence. In 2011, Philippe Bouchet and coauthors refined this framework in a new operational classification of the Conoidea, incorporating molecular phylogenies to validate 82 taxa and underscore the need for further splits in Conus sensu lato, paving the way for contemporary .

Diversity and genera

Cone snails, belonging to the family Conidae, exhibit remarkable diversity, with over 1,000 valid worldwide. This is classified into 8 principal genera—Californiconus, Conasprella, Conus, Kenyonia, Lilliconus, Profundiconus, Pseudolilliconus, and Pygmaeconus—encompassing numerous subgenera, based on molecular phylogenetic analyses and updates to the 2015 framework. The genus Conus dominates with over 700 , primarily inhabiting shallow tropical waters, while Conasprella includes around 160 specialized as worm-hunters. Deep-sea adapted groups, such as Profundiconus with approximately 30 , occupy colder, abyssal environments. Diversity is highest in the Indo-West Pacific region, which hosts over 60% of all known species, exceeding 600 taxa. Regional hotspots include the , with more than 150 species recorded, and , supporting around 150 species, reflecting the family's concentration in and subtropical habitats. Species delineation in cone snails faces significant challenges due to cryptic —morphologically similar but genetically distinct forms—and evidence of hybridization, complicating traditional . Molecular studies suggest that up to 20% of the actual diversity remains undescribed, particularly in understudied deep-water and remote populations. Recent assessments confirm one (Conus lugubris, 2025 IUCN update). Conservation assessments by the IUCN indicate that approximately 10-15% of evaluated cone snail species are threatened or near-threatened, primarily due to their rarity, limited distributions, and pressures from habitat loss and collection.

Human relevance

Risks to humans

Cone snails pose risks to humans primarily through accidental envenomation via their harpoon-like radular tooth, which can penetrate skin during handling of live specimens, often by shell collectors or divers. The venom, composed of conotoxins, includes peptides such as δ-conotoxins that block or modulate voltage-gated sodium channels, leading to intense localized pain by disrupting nerve signaling. Envenomations are rare globally, with fewer than 200 documented cases historically, most occurring among individuals handling the snails in tropical marine environments. Symptoms of a cone snail sting typically begin with sharp, burning pain at the site, followed by swelling, numbness, and tingling that may radiate proximally. In severe cases, particularly from piscivorous species like Conus geographus (known as the "cigarette snail" due to the time victims have to smoke one last cigarette before death), systemic effects can include muscle , , and , progressing within hours if untreated. Approximately 30 to 36 fatalities have been recorded worldwide, predominantly before 2000 and attributed to C. geographus, with no confirmed deaths reported in recent years owing to increased awareness and prompt medical intervention. There is no specific antivenom available for cone snail envenomations, so first aid focuses on symptom management and rapid transport to a medical facility. Immediate measures include immobilizing the affected limb to slow venom spread, immersing the sting site in hot water (as tolerable, up to 45–50°C) for 30–90 minutes to denature heat-labile toxins and alleviate pain, and monitoring for respiratory distress. Supportive care in a hospital setting, such as mechanical ventilation if needed, is critical for severe cases.

Biomedical applications

Cone snail venoms contain conotoxins, a diverse array of disulfide-rich peptides with high specificity for channels and receptors, making them valuable for biomedical research and . The most prominent example is (Prialt), an ω-conotoxin MVIIA derived from , approved by the FDA in 2004 for intrathecal treatment of severe chronic pain in patients unresponsive to other therapies. By selectively antagonizing N-type voltage-gated calcium channels (Cav2.2), ziconotide inhibits glutamate and release in the , providing non-opioid analgesia without respiratory depression or addiction risk. Ongoing research explores conotoxins for additional therapeutic applications, particularly in . Mu-conotoxins, such as μ-conotoxin CnIIIC, block voltage-gated sodium channels (Nav1.4), offering potential for treating muscle disorders like by reducing hyperexcitability without affecting cardiac or neuronal channels. Alpha-conotoxins, including α-conotoxin Vc1.1, target nicotinic receptors (nAChRs) and are under investigation for , where nAChR dysregulation contributes to seizures, as well as and inflammatory conditions. Several conotoxin analogs, such as contulakin-G and χ-conotoxin MrIA, have advanced to phase I/II clinical trials for pain and , though challenges like delivery and persist; as of , at least five compounds remain in early-stage development. Drug development from conotoxins involves solid-phase to produce these complex, 10-40 sequences with multiple bonds, followed by oxidative folding to achieve native . A key hurdle is their poor pharmacokinetic profile, including rapid enzymatic degradation and limited oral ; this is addressed through backbone cyclization, which links the N- and C-termini via amide bonds, enhancing serum stability by up to 100-fold while preserving activity, as demonstrated in cyclic analogs of α-conotoxin RgIA and Vc1.1. Beyond , conotoxins show promise in other areas. Chi-conotoxins, like χ-conotoxin MrIA, inhibit norepinephrine transporters and have icidal potential by disrupting neurotransmitter systems, with recombinant forms exhibiting lethality against agricultural pests in preclinical assays. Certain conotoxins, such as those targeting voltage-gated potassium channels, display anti-cancer effects by inducing in tumor cells, with studies on venom showing against ovarian and cell lines via modulation. Biotech firms have driven this field, with historical examples like Cognetix patenting over 100 conotoxins.

Collecting and trade

Cone snails have long been prized by collectors for their ornate shells, with interest dating back to the 17th and 18th centuries when they featured prominently in European due to their aesthetic appeal and exotic origins. During this period, a frenzy of shell collecting swept through Europe, fueled by trade routes such as those of the , which supplied rare specimens from tropical waters. Collecting peaked in the 1970s and 1980s, particularly in the , where the shell industry became a major export earner, with cone shells among the highly sought-after items traded internationally. Today, the trade in cone snail shells persists primarily for ornamental purposes, with specimens sold to collectors and jewelers; rare species can fetch prices ranging from $10 to over $500 per shell, depending on size, condition, and scarcity. While exact global volumes are difficult to quantify, the ornamental shell trade, including cones, involves tens of thousands of specimens annually, sourced mainly from regions. Collection methods remain labor-intensive, typically involving hand-gathering by free-diving or snorkeling in shallow tropical waters, often using tongs or nets to safely extract live snails without direct contact, as their venomous can pose risks to handlers. Efforts to establish for cone snails have been limited and largely unsuccessful outside settings, due to challenges in replicating their complex dietary and environmental needs, with most attempts focused on extraction rather than commercial shell production. Overharvesting has significantly impacted endemic species, such as Conus gloriamaris (the glory-of-the-sea cone), once among the rarest shells known, with only a few specimens documented until the 1960s; intensified collecting in the subsequently flooded the market but depleted local populations. Habitat loss from coastal development, pollution, and destructive fishing exacerbates these pressures, particularly in biodiversity hotspots like the . In October 2025, the confirmed the extinction of Conus lugubris, an endemic species from , underscoring the severity of these threats. According to the , as of 2013 approximately 6.5% of assessed cone snail species (out of 632) are threatened with extinction globally, with higher rates—up to 45% as of 2016—in isolated regions like Cape Verde, where endemism amplifies vulnerability. Regulatory measures aim to curb these threats, though cone snails are not currently listed under appendices. In , collection of marine snails, including cones, is strictly regulated under state fisheries laws, with bans on taking live specimens in many areas to protect native and prevent risks. The , a major sourcing hub, has implemented fishing quotas and marine protected areas to limit cone snail harvesting, with ongoing enforcement as of 2025 to sustain populations amid trade demands.

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