informal group Sigmurethra
clade limacoid clade
Limax maximus (literally, "great slug"), known by the common names great grey slug and leopard slug, is a species of slug in the family Limacidae, the keeled slugs. It is among the largest keeled slugs, Limax cinereoniger being the largest.
Limax maximus is the type species of the genus Limax. The adult slug measures 10-20 cm (4-8 in) in length and is generally a light greyish or grey-brown with darker spots and blotches, although the coloration and exact patterning of the body of this slug species is quite variable.
This species has a very unusual and distinctive mating method, where the pair of slugs use a thick thread of mucus to hang suspended in the air from a tree branch or other structure.
Although native to Europe, this species has been accidentally introduced to many other parts of the world.
- 1 Description
- 2 Distribution
- 3 Behavior
- 4 Ecology
- 5 Images
- 6 References
- 7 Notes
- 8 External links
- 9 Further reading
The greater part of the body is rounded, but there is a short keel on its tail, with about 48 longitudinal rows of elongate, detached tubercles. The body color is pale-grey, ash-colored, brownish or sometimes yellowish-white. The body is longitudinally streaked or spotted with black. The pattern of spotting is variable. The shield is always black-spotted. The sole of the foot is a uniform ash or yellowish-ash color. The foot-fringe is pale, with a row of minute submarginal blackish tubercles.
The shield is oblong, about one-third of the total length of the animal. The shield is rounded in front, angular behind, and forming an angle of about 80 degrees when in motion, usually of a similar tint to the body, but boldly marbled or maculate with black, somewhat concentrically and interruptedly ridged around a sub-posterior nucleus.
The pneumostome is just posterior to the midpoint of the mantle, as it is in all Limacidae.
Although color varieties have no actual taxonomic significance, a large number of color varieties have been described, prominent among them being the varieties serpentinus, vulgaris, cellarius (typical), johnstoni, maculatus, ferrussaci, obscurus, fasciatus and rufescens, of Alfred Moquin-Tandon, and cornaliee, of Pini.
The shell of Limax maximus is reduced and internal, under the shield. The occurrence of this internal shell was known to Pliny the Elder; the shell was used by the ancient physicians for the sake of its carbonate of lime
The calcitic shell is situated beneath the hinder part of the shield, and is perceptible through the skin. The color of the shell is whitish. The shape of the shell is oblong-oval and thin, slightly convex above, and correspondingly concave beneath, with a membranous margin. The apex or nucleus is at the posterior margin but inclined towards the left side, and forming the apophysis by which the shell is organically attached to the animal. The length of the shell is 13 mm (1/2 inch) and the width of the shell is 7 mm (1/4 inch). Shells of different Limacidae species are undiagnostic: in other words, they are not helpful for identification purpose.
Digestive system: The formula of the radula is: 62-73/ × 138-157. The intestine has six convolutions and is without a caecum. Of the six convolutions of the intestine, four are imbedded in the liver, and two hang freely in the body cavity.
The nervous system is composed of the typical ganglia. The pedal ganglia are placed beneath the radula sac and joined together by an anterior and a posterior commissure. The abdominal ganglion lies a little to the right of the median line. The visceral ganglia occupy the angle between the lingual sheath and the oesophagus and the buccal ganglia are widely separated but joined together by a commissure nearly as thick as the ganglia themselves.
Reproductive system: The hermaphrodite gland (HG) is elongated and large, and is connected with spermoviduct (SD) by means of the hermaphrodite duct (HD) which takes its course through a portion of the albumen gland (AG). The spermoviduct is thick and well convoluted, and separates further down into a vas deferens or sperm-duct (VD) and an oviduct (OV). The former opens into the upper end of a very long penis (P), to which a strong retractor muscle (PRM) is attached. The lower portion of the penis unites with that of the oviduct at the genital orifice, so that there is no vestibule. The receptaculum seminis (RS) opens into the lower end of the penis near the junction of the two ducts.
The internal shells of the different species of Limacidae are not recognizable to the species level. Therefore, the fossil distribution of Limax maximus (and other Limacidae species) is unknown. Unidentified calcitic shells of Limacidae are known from European Tertiary and Quaternary deposits.
Within Europe, this species occurs in many areas, including:
- Great Britain - Limax maximus was first described England in Animalium Angliae Tres Tractatus … Alter de Cochleis tum Terrestribus tum Fluviatilibus by Martin Lister in 1678. The presence of Limax maximus in England had been noted twelve years earlier in Christopher Merret's Pinax Rerum Naturalium Britannicarum.
- Benelux: Belgium, Netherlands, Luxembourg
The non-indigenous distribution of Limax maximus includes:
- Central Europe: Czech Republic, Poland, Slovakia, Hungary, Germany, Austria
- Northern Europe - Scandinavia: Norway, Sweden (possibly the type locality)
- Eastern Europe: Latvia, Lithuania, Belarus, Ukraine
Its introduction into the United States was first announced by George Washington Tryon in 1867, when it was discovered in cellars in Philadelphia. Within a few years its presence was noticed at Newport, Rhode Island, Brooklyn, and Pittsburgh, and it is now common in many areas, including:
- East Coast statesSouthern Ontario Canada
- South Central United States
- West Coast states, including California since 1896, and Montana
It can be found in some parts of Canada. In 2012 it was found in south-central Ontario, Nova Scotia, and western Newfoundland. Other non-indigenous distribution areas include Madeira, southern Africa, Australia, including Tasmania, New Zealand, Chile, and southern Brazil.
Limax maximus is nocturnal, feeding at night. It is not very active or prolific. When alarmed, or at rest, this slug merely draws its head within the shield, but does not otherwise contract its body. When irritated, it is said to expand its shield.
The homing faculty is strongly developed in this species, which, after its nocturnal rambles or foraging expeditions, usually returns to the particular crevice or chink in which it has established itself.
Limax maximus is capable of associative learning, specifically classical conditioning, because it is capable of aversion learning and other types of learning. It can also detect deficiencies in a nutritionally incomplete diet if the essential amino acid methionine is experimentally removed from its food.
The slug is almost always found near human habitation — usually in lawns, gardens, cellars or in other damp areas.
This species is not gregarious. It frequents gardens, damp and shady hedgerows and woods, hiding during the day beneath stones, under fallen trees, or other obscure and damp places. It does however exhibit a decided preference for the vicinity of human habitations, and readily takes up its abode in damp cellars or outbuildings.
In Ireland, this predilection for human dwellings is not exhibited, and the species is restricted to woods and other similar places. It may even be met with almost within high-water mark on the seashore.
Limax maximus is omnivorous. It is a detrivore, cleaning up dead plants and fungi, and a carnivore known to pursue other slugs at a top speed of 6 inches per minute. It also eats young crops faster than they can grow and so is listed as a major agricultural pest by state departments of agriculture in the US from Florida to Oregon.
The eggs of this slug are deposited in a cluster, slightly attached to each other. Eggs are transparent, elastic and slightly yellowish in color. The size of the egg is 6×4.5 mm. They hatch in about a month.
The tiny slugs which emerge from the eggs need at least two years to reach sexual maturity.
The lifespan of Limax maximus is 2.5–3 years.
The mating habits of Limax maximus are considered unusual among slugs: the hermaphrodite slugs court, usually for hours, by circling and licking each other. After this, the slugs will climb into a tree or other high area and then, entwined together, lower themselves on a thick string of mucus, evert their white translucent mating organs (penises) from their gonopores (openings on the right side of the head), entwine these organs, and exchange sperm. Both participants will later lay hundreds of eggs.
A commonly seen practice among many slugs is apophallation, when one or both of the slugs chews off the other's penis. The penis of these species is curled like a corkscrew and often becomes entangled in their mate's genitalia in the process of exchanging sperm. When all else fails, apophallation allows the slugs to separate themselves. Once its penis has been removed, a slug is still able to participate in mating subsequently, but only using the female parts of its reproductive system.
Parasites of Limax maximus include the nematode Agfa flexilis, which lives in its salivary glands,[unreliable source?] the nematode Angiostoma limacis, which lives in its rectum,[unreliable source?] and Angiostrongylus costaricensis.
A meningitis-causing nematode, Angiostrongylus cantonensis, which normally infests the lungs of rats, has a larval stage which can only live in molluscs, including slugs. This nematode was once known to be a problem only in tropical areas, but it has since spread to other regions. Live slugs that are accidentally eaten with improperly cleaned vegetables, such as lettuce, slugs which have been improperly cooked, can act as vectors for the parasite.
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- Taylor, J. W. (7 November) 1902. part 8, pages 1-52. Monograph of the land and freshwater Mollusca of the British Isles. Testacellidae. Limacidae. Arionidae. Taylor Brothers, Leeds. Introduction page XV., pages 34-52.
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- Pini. 1876. Bull. Soc. Mal. Ital. ii., 83. (There is described Limax cornaliae Pini)
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- Florida IFAS.
- UC Davis IPM.
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- Sanjaya N. Senanayake, Don S. Pryor, John Walker & Pam Konecny 2003. First report of human angiostrongyliasis acquired in Sydney. The Medical Journal of Australia, 179 (8): 430-31.
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|Wikimedia Commons has media related to Limax maximus.|
- Images of Mating
- Giant Garden Slug, Aliens Among Us. Virtual Exhibit of the Virtual Museum of Canada.
- Gelperin A (March 1974). "Olfactory Basis of Homing Behavior in the Giant Garden Slug, Limax maximus". Proceedings of the National Academy of Sciences of the United States of America 71 (3): 966–70. PMC 388138. PMID 16592149.
- Wieland SJ, Gelperin A (September 1983). "Dopamine elicits feeding motor program in Limax maximus". The Journal of Neuroscience 3 (9): 1735–45. PMID 6886743.