Cepaea nemoralis is able to respond to painful and non-painful stimuli. During the night, C. nemoralis feels more pain. Calcium channels are involved in the regulation of neuronal functions in mollusks in a manner like vertebrates. Similar intermediary messenger systems also exist between Cepaea and rodents.
Cepaea nemoralis predominantly moves in an upwind direction. They first randomly move in any direction before following the upwind stream. The decision to move upwind is made when the odor of favored foods is detected. Interactions between C. nemoralis exist. As the number of individuals within a colony increases, there is a decline in juvenile growth rates and birth rates. Limited resources is not the explanation because even in areas of normal competition, there is still growth rate and birth rate declines. This indicates that there is either a chemical or behavioral type of communication within the species that is responsible for the declination.
Communication Channels: visual ; tactile ; chemical
Perception Channels: visual ; infrared/heat ; tactile ; chemical ; magnetic
Cepaea nemoralis is listed as "least concern" in the Czech Republic. It is not specially listed by any other countries or organizations.
US Federal List: no special status
CITES: no special status
State of Michigan List: no special status
IUCN Red List of Threatened Species: no special status
In Europe, from 30-80 eggs (2.3-3.0 mm in diameter) are laid and hatch in 15-20 days. High temperatures and low humidity reduce the snail's activity and therefore inhibits growth rate. Cepaea nemoralis can form the peristome lip, which indicates it is an adult, in one active season. Juveniles may take to three years to develop into adults.
The color variation in the shells of Cepaea nemoralis is determined genetically by allelic series. Yellow shell alleles are recessive to pink shell alleles, and both yellow and pink shell alleles are recessive to brown shell alleles. The unbanded shell allele is dominant to banded.
There are no known adverse affects of Cepaea nemoralis on humans.
Cepaea nemoralis display a significant amount of polymorphism in their shells. This polymorphism has been the subject of many evolutionary studies in Europe.
Positive Impacts: research and education
Phorid flies are parasitic on C. nemoralis.
Ecosystem Impact: biodegradation
Commensal/Parasitic Species:
Cepaea nemoralis prefers to eat dead plant material rather than fresh. They also prefer to eat herbs rather than grasses. Adult C. nemoralis show greater selectivity in their eating habits than the juveniles, although they eat some grasses while juveniles do not. They eat Poterium sanguisorba and Leontodon hispidus. Lotus corniculatus and Urtica diocia are examples of rarely consumed greens. Cepaea nemoralis avoid vetch, shrub and grass in their diet. Remains of ants, beetles, spiders, mites, springtails and aphids are found in the diet of C. nemoralis, but these are probably the remains that were eaten along with greens and herbs.
In a colony that is as dense as 5 adults per square meter, the mean annual biomass consumed is 1.03 g per square meter. A large C. nemoralis can eat 125 mg of food per week, meaning a daily consumption rate of 59.5 mg of food per 1 g of dry tissue weight of the snail. Cepaea nemoralis relies on large lumps of food and its hydrolytic enzymes in its gut for nutrition. Vitamins A and B and some sterols are required components of C. nemoralis nutrition. Adults require sitosterol, a plant sterol.
Animal Foods: insects
Plant Foods: leaves; wood, bark, or stems; fruit; flowers
Primary Diet: herbivore (Folivore , Frugivore , Lignivore)
The northern distribution boundary of Cepaea nemoralis is in Scotland and southern Scandinavia. The range extends south to the Iberian Peninsula and Croatia in the south. Capaea nemoralis is found in the western and eastern coasts of both Ireland and the UK, Belgium, and France. The eastward distribution extends to the northwestern areas of Poland. This species was introduced in southeastern Poland, where it currently thrives. Cepaea nemoralis was introduced into North America during the nineteenth century and is currently found in Virginia, New York, Ontario, and Massachusetts.
Biogeographic Regions: nearctic (Introduced ); palearctic (Native )
Cepaea nemoralis is found in habitats ranging from hedgerows to downland turf and from beech woods to sand dunes near the sea. This species is also found throughout grasses and herbs. A relatively small amount is found in Marram grass. Unbanded and yellow C. nemoralis are mostly found in open habitats. Colonies with a green background have a high proportion of yellow C. nemoralis. Yellow C. nemoralis can also be found in shaded areas and banded shells of this species are found in areas of hedgerows and mixed rough herbage. In areas where the type of land is discontinuous, branded C. nemoralis are found. In Southeastern Poland, where C. nemoralis has been introduced, the species is found in urban environments where it inhabits gardens, orchards, cemeteries, hedgerows and other vegetation made up of herbs. In the hot and dry months of the summer, C. nemoralis is also found in tall plants and plants with large leaves or stems. Two of these plants include goldenrod and Centaurea. During hibernation C. nemoralis are found underground on land and underwater where they can survive for 2-3 weeks.
Habitat Regions: temperate ; terrestrial
Terrestrial Biomes: savanna or grassland
Other Habitat Features: urban
Cepaea nemoralis may live up to six years in the wild, but this is uncommon because of predation. In captivity, they may live up to 10 years. The average lifespan of C. nemoralis is 2.3 years. The survival rates of the young greatly differs and can range anywhere from 0.3 to 0.7. However, in most cases, the rate is closer to 0.3. It depends on the region and the temperature.
Range lifespan
Status: wild: 6 (high) years.
Average lifespan
Status: wild: 2.3 years.
Range lifespan
Status: captivity: 10 (high) years.
Average lifespan
Status: wild: 2.3 years.
Cepaea nemoralis has a yellow, pink, or brown shell. The shell contains as many as five dark bands (each 360 degree revolution constitutes one band). The shells are made up of different layers. The outer layer (periostracum) is made up of conchiolin and the layer directly layer is much thicker and is composed of calcium carbonate. Calcium is an important component of the shell and it is an important factor that determines the shell strength. Calcium concentrations vary from 319 to 359 mg/g. The shell strength, measured in Newtons required to break it, varies from 35 to 63 N. Cepaea nemoralis shell thickness varies from 0.17 to 0.21 mm. The dry weight of the shell varies from 0.43 to 0.72 g. The shell volume measures anywhere from 2230 to 3012 cubic mm.
Range mass: 0.72 (high) g.
Range basal metabolic rate: 0.014 to 0.251 cm3.O2/g/hr.
Average basal metabolic rate: 0.134 cm3.O2/g/hr.
Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry ; polymorphic
Sexual Dimorphism: sexes alike
Birds, mice, and rats are the most significant predators of Cepaea nemoralis. In Europe, the most common predator of C. nemoralis is song thrushes. Predators feed on snails by cracking the shells on nearby hard objects or with the use of their teeth.
Other predators of C. nemoralis include the rook, the brown rat, hedgehogs, moles, field mice, Sorex, rabbits, and maggots. Phorid flies are parasitic and consume C. nemoralis.
Anti-predator adaptations of C. nemoralis include the complexity of color and bands on shell varying with the complexity of the landscape, known as background matching. Brown shelled C. nemoralis are found in landward parts which are generally complex landscapes. Unbanded shells are found in the most pitted areas. The greatest complexity in the variation of shell colors and whether or not the shells are un/banded correlates with the amount of enclosed spaces such as pits that serve as amphitheaters for other organisms. Song Thrushes, a main predator of C. nemoralis, preys on shells that are distinguishable from the environment.
Another anti-predator adaptation of C. nemoralis is shell thickening. Thickening of the shell prevents the ability of a predator to crush the shell. It also increases the "handling time" of the snail. Increasing handling time makes C. nemoralis less energetically rewarding prey. Crushing resistance positively correlates with calcium concentration.
Known Predators:
Anti-predator Adaptations: cryptic
Cepaea nemoralis is hermaphroditic and mates more than once. A mating partner for C. nemoralis is random, according to color, size and banding patterns. The snails can store sperm for long periods of time. Each individual C. nemoralis may produce offspring from several matings in each brood. The average number of mates per brood is two. Cross fertilization is obligatory.
The offspring of each C. nemoralis is divided into a number of broods that is produced over a period of months or even years. This makes it unlikely that the fates between different broods will mimic each other. Each brood consists of genetic contributions from one female parent and two male parents. This reduces the dependence of single parents for survival within a whole brood unit. This system of multiple mating and sperm storage protects organisms that are minimally mobile from complete fatality within a gene pool.
Courtship of C. nemoralis is elaborate. A calcareous dart is jabbed into a potential partner before mating begins. In order to prevent accidental mating between C. nemoralis and a closely related species, Cepaea hortensis, the two species have different darts.
Mating System: polygynandrous (promiscuous)
The breeding interval of Cepaea nemoralis runs from April through October. The relative seasons are spring, summer and the beginning of the fall. The number of offspring per brood is around 23. Eggs are laid simultaneously in a dug nest in soil. The snail's foot is used to create a cavity in soil for laying eggs. The laying of the eggs can take up to three days and when complete, its foot is used again to cover the nest.
The average number of hatched young per year is 33. The eggs of one brood may survive or die as a unit; however, once they hatch, the individual's survival is not correlated to the rest of the brood. High temperatures and low humidity reduce the snail's activity and therefore inhibits growth rate and egg production.
Breeding interval: This species breeds 1 to 2 times yearly.
Breeding season: Breeding season is from April to October.
Average number of offspring: 23.
Range age at sexual or reproductive maturity (female): 1 to 2 weeks.
Range age at sexual or reproductive maturity (male): 1 to 2 weeks.
Key Reproductive Features: seasonal breeding ; sequential hermaphrodite; sexual ; induced ovulation ; fertilization (Internal ); oviparous ; sperm-storing ; delayed fertilization
Cepaea nemoralis produces eggs at the same time and are laid in nest they dig and cover up with soil. Beyond hatching, there is no parental care.
Parental Investment: male parental care ; female parental care ; pre-fertilization
The grove snail, brown-lipped snail or lemon snail (Cepaea nemoralis) is a species of air-breathing land snail, a terrestrial pulmonate gastropod mollusc.[3]
It is one of the commonest large species of land snail in Europe, and has been introduced to North America.
Cepaea nemoralis is the type species of the genus Cepaea.[5] It is used as a model organism in ecological genetics, including in citizen science projects.[6][7]
Cepaea nemoralis is among the largest and, because of its bright colouration, one of the best-known snails in Western Europe.[6] The colour of the shell is highly variable; it ranges from brown, through pink, to yellow or even whitish, with or without one to five dark-brown bands.[8] Names for many colour variants were coined in the nineteenth century but this system has been replaced by an independent scoring of shell colour and the presence/absence and fusion of individual bands numbered 1 to 5.[9]
The thickened and slightly out-turned apertural lip of adults is usually dark brown, but can be white in some regions.[10] The umbilicus is closed in adults but narrowly open in juveniles.[8] The shell surface is semi-glossy. An adult shell consists of 4½–5½ whorls, with a width of 18–25 mm and a height of 12–22 mm.[8]
Cepaea nemoralis is closely related to Cepaea hortensis. They share much the same habitat and exhibit a similar range of shell colours and banding patterns. Cepaea nemoralis tends to grow larger, but usually the species can most easily be recognised by the colour of the lip of adult shells. In a high proportion of regions, C. nemoralis consistently has a dark-brown lip to its shell, whilst C. hortensis has a white lip.[10]
In areas where lip colour is variable, dissection is necessary. A cross-section of the love dart of C. nemoralis shows a cross with simple blades, whereas that of C. hortensis has bifurcated blades. The mucus gland has 3 or fewer branches in C. nemoralis, but 4 or more in C. hortensis.[12]
Two superficially similar species Caucasotachea vindobonensis and Macularia sylvatica both have a lip that is brown near the columella becoming pale towards the suture, and they have fine growth ridges on the shell whereas in both Cepaea species it is smooth. Also, M. sylvatica is distinct in having a small blunt tooth in its aperture, whilst the lowest brown band on the shell of C. vindobonensis lies noticeably closer to the columella than in Cepaea.[13]
Cepaea nemoralis is highly polymorphic in shell colour and banding. The background colour of the shell various along a continuum from brown through pink to yellow and sometimes almost white.[14] Additionally the shells can be with or without dark bands. The bands vary in intensity of colour, in width and in number, from zero to five. The genetics underlying this variation is extensively understood and is shared with C. hortensis.[10]
The polymorphism has also been intensely studied for its evolution and ecology. For instance, in stable habitats shells tend to be darker in woodland than in open habitats.[15] The explanation might be camouflage or climatic selection: paler, more reflective colours in sunny environments reduce water loss and overheating. Climatic selection can also explain why yellow shells are commoner in the south.[6]
Another question is why the variation persists, usually even within a locality. Researchers have variously argued that the cause is random genetic drift and founder effects, different selection pressures in different areas with mixing by migration, and balanced polymorphism. Balanced polymorphism could arise when a predator like the song thrush develops a 'search image' for the commonest morph, so that the rarer morphs are less likely to be predated. Natural selection would then favour a diversity of colours and patterns as an antipredator adaptation. Most probably, the polymorphism has multiple causes.[10][16]
A diversity of forms
Yellow unbanded
Yellow with several bands
Yellow mid-banded
Reddish-brown unbanded
The native distribution of C. nemoralis is from Western and Northern Europe to Central Europe, but it has been spreading eastwards especially over the last few decades.[17][18] Thus it is known from most of the Iberian Peninsula,[19] France, Great Britain, Ireland, Belgium, the Netherlands, Germany, Denmark, southern Sweden and Norway, Switzerland, Austria, the Czech Republic,[20][21] Hungary, Bosnia, Croatia, and the northern half of Italy.[8][22] In Central and Eastern Europe it has spread particularly along the Baltic coast (e.g. in Poland,[18] Latvia, Estonia, southern Finland,[23] the east coast of Sweden)[24] but also now elsewhere in Poland,[18] and in Ukraine, Belarus and Russia.[25] In Slovakia it was still known only from a single garden centre by 2020.[26]
At the northern edge of its range,C. nemoralis is rare and scattered in northern Scotland, where it has been introduced.[8] It is not found in the Hebrides, Orkney or Shetland.[8] It seems to have been affected by air pollution and soil acidification in some parts of England.[8]
Starting in 1857, there have been multiple introductions to North America, where it now occurs widely in Canada (from British Columbia to Newfoundland; in Ontario it is a pest of vineyards) and in the north-eastern part of the US, with further occurrences further south (e.g. California, Colorado, Texas and South Carolina).[27][28][29]
Both in America and Eastern Europe it is known that some introductions have been deliberate.[28][25]
The range of C. hortensis mostly overlaps that of C. nemoralis but extends further north and less far south.[12]
This is a very common and widespread species in Western Europe, occupying a wide range of habitats from coastal dunes to woodlands with full canopy cover, including gardens and abandoned land.[6] In Eastern Europe, where it is a new arrival, it has turned up particularly in urban areas and other disturbed habitats.[18] It can be found up to an altitude of 1600 m in the Alps,[30] 1800 m in the Pyrenees, 900 m in Wales, 600 m in Scotland.[8] Density of adults is often of the order of 2 per square metre.[31]
This species feeds mainly on dead or senescent plants.[6][8] It prefers broad-leaved plants over grasses.[32] Although mostly not a pest of crops,[8] it can be a nuisance in vineyards because it is inadvertently picked with the grapes.[33]
Like all pulmonate land snails, it is hermaphrodite, and this species must mate to produce fertile eggs.[6] Mating tends to be concentrated in late spring and early summer, though it can continue through the autumn.[6] The snails may store the sperm they receive from their partner for some time, and individual broods can have mixed paternity.[6] In Britain it lays clutches of 30–50 (in France 40–80) oval eggs between June and August (in France May–October, in W France until November).[8] The size of the egg is 3.1 × 2.6 mm[34] or egg diameter can be 2.3–3.0 mm.[8] Juveniles hatch after 15–20 days.[8]
This snail is comparatively slow-growing, taking 1 to 3 years to develop from an egg to a breeding adult.[35] The life-span for this species is up to seven or eight years, with the annual survival rate of adults about 50% (= 3% over five years, older adults suffer higher mortalities).[8] In winter, the snails may hibernate, but can become active again during warm spells.[6]
Cepaea nemoralis is known experimentally to be a host for Angiostrongylus vasorum.[36] Predators of Cepaea nemoralis include the song thrush (Turdus philomelos).
This article includes public domain text from the reference[8] and CC-BY-2.5 text from the reference[6]
The grove snail, brown-lipped snail or lemon snail (Cepaea nemoralis) is a species of air-breathing land snail, a terrestrial pulmonate gastropod mollusc.
It is one of the commonest large species of land snail in Europe, and has been introduced to North America.
Subspecies Cepaea nemoralis etrusca (Rossmässler, 1835) Cepaea nemoralis nemoralis (Linnaeus, 1758)Cepaea nemoralis is the type species of the genus Cepaea. It is used as a model organism in ecological genetics, including in citizen science projects.
