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Slug

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For marine slugs, seeSea slug.For snails, seeSnail.For other uses, seeSlug (disambiguation).

Slug
Various species of British land slugs, including (from the top) the larger drawings: Arion ater, Kerry slug, Limax maximus and Limax flavus
Various species ofBritishland slugs, including (from the top) the larger drawings:Arion ater,Kerry slug,Limax maximusandLimax flavus
Arion sp., from Vancouver, BC
Arionsp., fromVancouver,BC
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Groups included
A slug on a wall inKanagawa,Japan.

Slug,orland slug,is acommon namefor any apparently shell-lessterrestrialgastropodmollusc.The wordslugis also often used as part of the common name of any gastropod mollusc that has no shell, a very reduced shell, or only a small internal shell, particularlysea slugsandsemi-slugs(this is in contrast to the common namesnail,which applies to gastropods that have a coiled shell large enough that they can fully retract their soft parts into it).

Varioustaxonomicfamilies of land slugs form part of several quite different evolutionary lineages, which also include snails. Thus, the various families of slugs are not closely related, despite the superficial similarity in overall body form. The shell-less condition has arisen many times independently as an example ofconvergent evolution,and thus the category "slug" ispolyphyletic.

Taxonomy

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Of the six orders ofPulmonata,two – theOnchidiaceaandSoleolifera– solely comprise slugs. A third group, theSigmurethra,contains various clades of snails, semi-slugs (i.e. snails whose shells are too small for them to retract fully into), and slugs.[1]The taxonomy of this group is in the process of being revised in light of DNA sequencing.[2]Research suggests that pulmonates are paraphyletic and basal to theopisthobranchs,which are a terminal branch of the tree.[3]The familyEllobiidaeare also polyphyletic.

Description

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Drawing of slug with labels for the foot (bottom side), the foot fringe that surrounds it, the mantle behind the head, the pneumostome for breathing, and the optical and sensory tentacles
The external anatomy of a slug

The external anatomy of a slug includes the following:

  • Tentacles:Like other pulmonate land gastropods, the majority of land slugs have two pairs of 'feelers' ortentacleson their head. The upper pair is light-sensing and has eyespots at the ends, while the lower pair provides thesense of smell.Both pairs are retractable instylommatophoranslugs, but contractile inveronicellidslugs.
  • Mantle:On top of the slug, behind the head, is the saddle-shapedmantle.In stylommatophoran slugs, on the right-hand side of the mantle is arespiratoryopening, thepneumostome,which is easier to see when open; also on the right side at the front are the genital opening and anus. Veronicellid slugs have a mantle covering the whole dorsal part of the body, they have no respiratory opening, and the anus opens posteriorly.
  • Tail:The part of a slug behind the mantle is called the 'tail'.
  • Keel:Some species of slugs, for exampleTandonia budapestensis,have a prominent ridge running over their back along the middle of the tail (sometimes along the whole tail, sometimes only the posterior part).
  • Foot:The bottom side of a slug, which is flat, is called the 'foot'. Like almost all gastropods, a slug moves by rhythmic waves ofmuscular contractionon the underside of its foot. It simultaneously secretes a layer ofmucusthat it travels on, which helps prevent damage to the foot tissues.[4]Around the edge of the foot in some slugs is a structure called the 'foot fringe'.
  • Vestigial shell:Most slugs retain a remnant of their shell, which is usually internalized. This organ generally serves as storage for calcium salts, often in conjunction with the digestive glands.[5]An internal shell is present in theLimacidae[6]and Parmacellidae.[7]AdultPhilomycidae,[6]Onchidiidae[8]andVeronicellidae[9]lack shells.

Physiology

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An activeAmbigolimaxslug inFremont, California

Slugs' bodies are made up mostly of water and, without a full-sized shell, their soft tissues are prone todesiccation.They must generate protective mucus to survive. Many species are most active following rainfall or during nighttime since there is increased moisture on the ground. In drier conditions, they hide in damp places such as undertree bark,fallenlogs,rocks and manmade structures, such asplanters,to help retain body moisture.[4]Like all other gastropods, they undergotorsion(a 180° twisting of the internal organs) during development. Internally, slug anatomy clearly shows the effects of this rotation—but externally, the bodies of slugs appear more or less symmetrical, except thepneumostome,which is on one side of the animal, normally the right-hand side.

Slugs produce two types ofmucus:one is thin and watery, and the other thick and sticky. Both kinds arehygroscopic.The thin mucus spreads from the foot's centre to its edges, whereas the thick mucus spreads from front to back. Slugs also produce thick mucus that coats the whole body of the animal.[4]The mucus secreted by the foot contains fibres that help prevent the slug from slipping down vertical surfaces.

The "slimetrail "a slug leaves behind has some secondary effects: other slugs coming across a slime trail can recognise the slime trail as produced by one of the same species, which is useful in finding a mate. Following a slime trail is also part of the hunting behaviour of some carnivorous slugs.[4]Body mucus provides some protection against predators, as it can make the slug hard to pick up and hold by a bird's beak, for example, or the mucus itself can be distasteful.[10]Some slugs can also produce very sticky mucus which can incapacitate predators and can trap them within the secretion.[11]Some species of slug, such asLimax maximus,secrete slime cords to suspend a pair during copulation.

Reproduction

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Derocerasjuvenile with eggs of another (larger) slug species, most likely Arionidae
Pair of mating slugs, suspended bymucousthread and pad on wall
A pair of mating slugs

Slugs arehermaphrodites,having both female and male reproductive organs.[12]Once a slug has located a mate, they encircle each other and sperm is exchanged through their protruded genitalia.

Apophallationhas been reported only in some species of banana slug (Ariolimax) and one species ofDeroceras.In the banana slugs, the penis sometimes becomes trapped inside the body of the partner. Apophallation allows the slugs to separate themselves by one or both of the slugs chewing off the other's or its own penis. Once the penis has been discarded, banana slugs are still able to mate using only the female parts of the reproductive system.[12][13][14]

In a temperate climate, slugs usually live one year outdoors. In greenhouses, many adult slugs may live for more than one year.[15]

Ecology

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Slugs play an important role in theecosystemby eating decaying plant material andfungi.[16]Most carnivorous slugs on occasion also eat dead specimens of their own kind.

Feeding habits

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Large slug nearManali, India

Most species of slugs are generalists, feeding on a broad spectrum of organic materials, including leaves from living plants,lichens,mushrooms,and evencarrion.[16][17]Some slugs arepredatorsand eat other slugs andsnails,orearthworms.[16][18]

Lehmanniasp. feeding on a small fruit inMexico City

Slugs can feed on a wide variety ofvegetablesandherbs,[19]including flowers such aspetunias,chrysanthemums,daisies,lobelia,lilies,dahlias,narcissus,gentians,primroses,tuberous begonias,hollyhocks,marigolds,and fruits such asstrawberries.[20]They also feed on carrots, peas, apples, and cabbage that are offered as a sole food source.[17]

Slugs from different families arefungivores.It is the case in thePhilomycidae(e. g.Philomycus carolinianusandPhylomicus flexuolaris) andAriolimacidae(Ariolimax californianus), which respectively feed on slime molds (myxomycetes) and mushrooms (basidiomycetes).[17]Species of mushroom producing fungi used as food source by slugs include milk-caps (Lactariusspp.), the oyster mushroom (Pleurotus ostreatus) and the penny bun (Boletus edulis). Other genera such asAgaricus,PleurocybellaandRussulaare also eaten by slugs. Slime molds used as food source by slugs includeStemonitis axiferaandSymphytocarpus flaccidus.[17]Some slugs are selective towards certain parts or developmental stages of the fungi they eat, though this is very variable. Depending on the species and other factors, slugs eat only fungi at specific stages of development. In other cases, whole mushrooms can be eaten, without any selection or bias towardsontogeneticstages.[17]

Predators

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Slugs are preyed upon by variousvertebratesandinvertebrates.The predation of slugs has been the subject of studies for at least a century. Because some species of slugs are consideredagricultural pests,research investments have been made to discover and investigate potential predators in order to establish biological control strategies.[21]

Vertebrates

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Slugs are preyed upon by virtually every major vertebrate group. With many examples amongreptiles,birds,mammals,amphibiansandfish,vertebrates can occasionally feed on, or be specialised predators of, slugs.[21]Fish that feed on slugs include the brown trout (Salmo trutta), which occasionally feeds onArion circumscriptus,anarionidslug.[21]Similarly, the shortjaw kokopu (Galaxias postvectis) includes slugs in its diet.[22]Amphibians such as frogs and toads have long been regarded as important predators of slugs. Among them are species in the genusBufo,RhinellaandCeratophrys.[21]

Reptiles that feed on slugs include mainlysnakesandlizards.[21]Somecolubridsnakes are known predators of slugs. Coastal populations of the garter snake,Thamnophis elegans,have a specialised diet consisting of slugs, such asAriolimax,while inland populations have a generalized diet.[23]One of itscongeners,the Northwestern garter snake (Thamnophis ordinoides), is not a specialized predator of slugs but occasionally feeds on them. The redbelly snake (Storeria occipitomaculata) and the brown snake (Storeria dekayi) feed mainly but not solely on slugs, while some species in the genusDipsas(e.g.Dipsas neuwiedi) and the common slug eater snake (Duberria lutrix), are exclusively slug eaters.[21][24]Several lizards include slugs in their diet. This is the case in the slowworm (Anguis fragilis), the bobtail lizard (Tiliqua rugosa), the she-oak skink (Cyclodomorphus casuarinae) and the common lizard (Zootoca vivipara).[21][25][26]

Birds that prey upon slugs includecommon blackbirds(Turdus merula),starlings(Sturnus vulgaris),rooks(Corvus frugilegus),jackdaws(Corvus monedula),owls,vulturesandducks.Studies on slug predation also citefieldfares(feeding onDeroceras reticulatum),redwings(feeding onLimaxandArion),thrushes(onLimaxandArion ater),red grouse(onDerocerasandArion hortensis),game birds,wrynecks(onLimax flavus),rock dovesandcharadriiformbirds as slug predators.[21]

Mammals that eat slugs includefoxes,badgersandhedgehogs.[27][28]

Invertebrates

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Beetles in the familyCarabidae,such asCarabus violaceusandPterostichus melanarius,are known to feed on slugs.[29][30]Ants are a common predator of slugs; some ant species are deterred by the slug's mucus coating, while others such asdriver antswill roll the slug in dirt to absorb its mucus.

Parasites and parasitoids

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Slugs areparasitisedby several organisms, includingacari[31][32]and a wide variety ofnematodes.[33][34]The slug mite,Riccardoella limacum,is known to parasitise several dozen species of molluscs, including many slugs, such asDeroceras reticulatum,Arianta arbustorum,Arion ater,Arion hortensis,Limax maximus,Tandonia budapestensis,Milax gagates,andTandonia sowerbyi.[31][32]R. limacumcan often be seen swarming about their host's body, and live in its respiratory cavity.

Several species of nematodes are known to parasitise slugs. The nematode wormsAgfa flexilisandAngiostoma limacisrespectively live in thesalivary glandsandrectumofLimax maximus.[35]Species of widely known medical importance pertaining to the genusAngiostrongylusare also parasites of slugs. BothAngiostrongylus costaricensisandAngiostrongylus cantonensis,ameningitis-causing nematode, have larval stages that can only live in molluscs, including slugs, such asLimax maximus.[33]

Insects such asdipteransare knownparasitoidsof molluscs. To complete their development, many dipterans use slugs as hosts during theirontogeny.Some species of blow-flies (Calliphoridae) in the genusMelindaare known parasitoids ofArionidae,LimacidaeandPhilomycidae.Flies in the familyPhoridae,specially those in the genusMegaselia,are parasitoids ofAgriolimacidae,including many species ofDeroceras.[36]House flies in the familyMuscidae,mainly those in the genusSarcophaga,are facultative parasitoids of Arionidae.[37]

Behavior

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A slug retracts its head and contracts its body after being touched
A brown and yellow spotted slug curled up into a tight ball so that its head is withdrawn completely, its mantle edge and tail are nearly touching, and none of its foot surface is exposed
The alarm response posture of theKerry slug,which is found only in this species

When attacked, slugs can contract their body, making themselves harder and more compact and more still and round. By doing this, they become firmly attached to thesubstrate.This, combined with the slippery mucus they produce, makes slugs more difficult for predators to grasp. The unpleasant taste of the mucus is also a deterrent.[10]Slugs can also incapacitate predators through the production of a highly sticky and elastic mucus which can trap predators in the secretion.[11]

Some species present different response behaviors when attacked, such as theKerry slug.In contrast to the general behavioral pattern, the Kerry slug retracts its head, lets go of the substrate, rolls up completely, and stays contracted in a ball-like shape.[38]This is a unique feature among all theArionidae,[39]and among most other slugs.[38]Some slugs can self-amputate (autotomy) a portion of their tail to help the slug escape from a predator.[40]Some slug specieshibernateunderground during the winter in temperate climates, but in other species, the adults die in the autumn.[20]

Intra- and inter-specificagonistic behavioris documented, but varies greatly among slug species. Slugs often resort to aggression, attacking bothconspecificsand individuals from other species when competing for resources. This aggressiveness is also influenced byseasonality,because the availability of resources such as shelter and food may be compromised due toclimaticconditions. Slugs are prone to attack during the summer, when the availability of resources is reduced. During winter, the aggressive responses are substituted by agregariousbehavior.[41]

Human relevance

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The great majority of slug species are harmless to humans and to their interests, but a small number of species are serious pests of agriculture and horticulture. They can destroy foliage faster than plants can grow, thus killing even fairly large plants. They also feed on fruits and vegetables prior to harvest, making holes in the crop, which can make individual items unsuitable to sell for aesthetic reasons, and can make the crop more vulnerable to rot and disease.[42]Excessive buildup of slugs within some wastewater treatment plants with inadequate screening have been found to cause process issues resulting in increased energy and chemical use.[43]

In a few rare cases, humans have developedAngiostrongylus cantonensis-inducedmeningitisfrom eating raw slugs.[44]Live slugs that are accidentally eaten with improperly cleaned vegetables (such aslettuce), or improperly cooked slugs (for use in recipes requiring larger slugs such asbanana slugs), can act as a vector for a parasitic infection in humans.[34][45]

Prevention

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Main article:Molluscicide

As control measures, baits are commonly used in both agriculture and the garden. In recent years,iron phosphatebaits have emerged and are preferred over the more toxicmetaldehyde,especially because domestic or wild animals may be exposed to the bait. The environmentally safer iron phosphate has been shown to be at least as effective as baits.[46]Methiocarb baits are no longer widely used. Parasitic nematodes (Phasmarhabditis hermaphrodita) are a commercially available biological control method that are effective against a wide range of common slug species. The nematodes are applied in water and actively seek out slugs in the soil and infect them, leading to the death of the slug. This control method is suitable for use in organic growing systems.

Other slug control methods are generally ineffective on a large scale, but can be somewhat useful in small gardens. These includebeer traps[de],[47][48]diatomaceous earth,[49]crushed eggshells,coffee grounds,and copper.[50]Salt kills slugs by causing water to leave the body owing toosmosis[51]but this is not used for agricultural control assoil salinityis detrimental to crops.[citation needed]Conservation tillageworsens slug infestations. Hammond et al. 1999 findmaize/cornandsoybeanin the US to be more severely affected under low till because this increases organic matter, thus providing food and shelter.[52]

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See also

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References

[edit]
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  2. ^White, T. R.; Conrad, M. M.; Tseng, R.; Balayan, S.; Golding, R.; de Frias Martins, A. M.; Dayrat, B. A. (2011)."Ten new complete mitochondrial genomes of pulmonates (Mollusca: Gastropoda) and their impact on phylogenetic relationships".BMC Evolutionary Biology.11(1): 295.Bibcode:2011BMCEE..11..295W.doi:10.1186/1471-2148-11-295.PMC3198971.PMID21985526.
  3. ^Kocot, Kevin (December 2013)."Phylogenomics supports Panpulmonata: Opisthobranch paraphyly and key evolutionary steps in a major radiation of gastropod molluscs".Molecular Phylogenetics and Evolution.69(2): 764–771 – via Science Direct.
  4. ^abcdDenny, M. W.; Gosline, J. M. (1980)."The physical properties of the pedal mucus of the terrestrial slug,Ariolimax columbianus"(PDF).Journal of Experimental Biology.88(1): 375–393.doi:10.1242/jeb.88.1.375.
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Further reading

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