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The Kerry slug or Kerry spotted slug (''Geomalacus maculosus'') is a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
of terrestrial, pulmonate, gastropod mollusc. It is a medium-to-large sized, air-breathing land slug in the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
of roundback slugs,
Arionidae Arionidae, common name the "roundback slugs" or "round back slugs" are a taxonomic family of air-breathing land slugs, terrestrial pulmonate gastropod mollusks in the superfamily Arionoidea. Distribution The distribution of this family of slu ...
. Adult Kerry slugs generally measure in length; they are dark-grey or brown with yellowish spots. The internal anatomy of the slug has some unusual features and some characteristic differences from the genus ''Arion'', also part of Arionidae. The Kerry slug was described in 1843—later than many other relatively large land gastropods present in Ireland and Great Britain—an indication of its restricted distribution and secretive habits. Although the distribution of this slug species includes south-western
Ireland Ireland ( ; ga, Éire ; Ulster-Scots: ) is an island in the North Atlantic Ocean, in north-western Europe. It is separated from Great Britain to its east by the North Channel, the Irish Sea, and St George's Channel. Ireland is the s ...
—including
County Kerry County Kerry ( gle, Contae Chiarraí) is a county in Ireland. It is located in the South-West Region and forms part of the province of Munster. It is named after the Ciarraige who lived in part of the present county. The population of the co ...
—the species is more widespread in north-western Spain and central-to-northern Portugal. Given that the slug has thus far been recorded exclusively at locations in Ireland and north-western
Iberia The Iberian Peninsula (), ** * Aragonese language, Aragonese and Occitan language, Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a pe ...
, it can be said to tentatively possess a
Lusitanian distribution In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but considerably separated from each other geographically. The causes are varied and might demonstrate either the expansion or contraction of a s ...
. The species appears to require environments that have high humidity, warm summer temperatures and acidic soils with no calcium carbonate. The slug is mostly nocturnal or crepuscular but in Ireland it is active on overcast days. It feeds on lichens, liverworts, mosses and
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
, which grow on boulders and tree trunks. The Kerry slug is protected by conservation laws in the three countries in which it occurs. It is now known to be less dependent on sensitive, wild habitats than when these laws were introduced. Attempts have been made to establish breeding populations in captivity to ensure the survival of this slug species but these have been only partly successful.


Taxonomy and etymology

The Kerry slug is a gastropod, a
class Class or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used differently ...
of molluscs that includes all snails and slugs, including terrestrial, freshwater and marine species. The Kerry slug, a member of the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
Panpulmonata, is terrestrial; it breathes air with a lung. It is in the clade
Stylommatophora Stylommatophora is an orderPhilippe Bouchet, Jean-Pierre Rocroi, Bernhard Hausdorf, Andrzej Kaim, Yasunori Kano, Alexander Nützel, Pavel Parkhaev, Michael Schrödl and Ellen E. Strong. 2017. Revised Classification, Nomenclator and Typification ...
, members of which have two sets of retractable
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work main ...
s, the upper pair of which have eyes on their tips. Its
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
is
Arionidae Arionidae, common name the "roundback slugs" or "round back slugs" are a taxonomic family of air-breathing land slugs, terrestrial pulmonate gastropod mollusks in the superfamily Arionoidea. Distribution The distribution of this family of slu ...
, the round-backed slugs. The Kerry slug has no
keel The keel is the bottom-most longitudinal structural element on a vessel. On some sailboats, it may have a hydrodynamic and counterbalancing purpose, as well. As the laying down of the keel is the initial step in the construction of a ship, in Br ...
on its back, unlike the slugs in the families
Limacidae Limacidae, also known by their common name the keelback slugs, are a taxonomic family of medium-sized to very large, air-breathing land slugs, terrestrial pulmonate gastropod molluscs in the superfamily Limacoidea. Distribution The distributio ...
and Milacidae. Many of its anatomical features are shared with species in the genus '' Arion'', which is a more species-rich and widely distributed group of slugs within Arionidae. The Kerry slug is placed in the genus ''
Geomalacus ''Geomalacus'' is a genus of large air-breathing land slugs, terrestrial pulmonate gastropod mollusks in the family Arionidae, the roundback slugs. Etymology The Ancient Greek word () means the Earth. The Greek word () means mollusc. Dis ...
'', which means "earth mollusc". The Kerry slug's scientific name is ''Geomalacus maculosus'', where ''maculosus'' means "spotted" from the Latin word ''macula'', a spot. The English-language
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contrast ...
is derived from County Kerry in the south-west of Ireland, where the
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
s that were used for the formal scientific description were collected. In 1842, a Dublin-based naturalist William Andrews (1802–1880) sent specimens he had found at
Caragh Lake Caragh Lake ( ga, Loch Cárthaí), also Lough Caragh, is a lake in the Reeks District in County Kerry, Ireland. The lake was formed by the damming of the Caragh River. Caragh Lake railway station was on the Great Southern and Western Railwa ...
in County Kerry to the Irish biologist George James Allman. The next year, Allman exhibited them at the Dublin Natural History Society and published a formal description of the new species and genus in the London literary magazine ''The Athenaeum''. The full scientific name, including the taxonomic authority, is ''Geomalacus maculosus'' Allman, 1843. The synonyms are other binomial names that were given over time to this taxon by authors who were unaware that the specimens they were describing belonged to a species already described by Allman. The species' binomial name is sometimes written as ''Geomalacus (Geomalacus) maculosus'' because the genus ''Geomalacus'' contains two subgenera; the nominate subgenus (subgenus of the same name) ''Geomalacus'' and a second subgenus ''Arrudia'' Pollonera, 1890. The subgenus ''Geomalacus'' contains only one species, the Kerry slug; three species comprise ''Arrudia''. The Kerry slug has been included in
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies since 2001.


Description

The body length of adult Kerry slugs is . These slugs are difficult to measure accurately because of their unusual
startle response In animals, including humans, the startle response is a largely unconscious defensive response to sudden or threatening stimuli, such as sudden noise or sharp movement, and is associated with negative affect.Rammirez-Moreno, David. "A computation ...
. Kerry slugs can also elongate themselves within crevices up to . Official measurements of this species vary; Kerney et al. (1983) give a range of measurements of . The body of a fixed (preserved) adult specimen was long with a mantle length of . The body of the Kerry slug is glossy and is covered on both sides with about 25 longitudinal rows of polygonal granulations. The slugs have two
colour morph In biology, polymorphism is the occurrence of two or more clearly different morphs or forms, also referred to as alternative ''phenotypes'', in the population of a species. To be classified as such, morphs must occupy the same habitat at the s ...
s, brown and black. In Ireland the black morph occurs in open habitats and the brown morph occurs in woodland; this correlates with the colours of the surroundings, suggesting camouflage. Experiments indicate the dark colouration is induced by exposure to light as the slug develops. There is also variation in banding; on each side of the body there can be two bands: one band just below the summit of the back and the other band further down the side of the body. When these bands are present they usually extend the whole length of the body and are overspread by numerous, ovoid yellow spots that are distributed approximately in five longitudinal zones. Behind the animal's head is the shield-shaped outer surface of the
mantle A mantle is a piece of clothing, a type of cloak. Several other meanings are derived from that. Mantle may refer to: *Mantle (clothing), a cloak-like garment worn mainly by women as fashionable outerwear **Mantle (vesture), an Eastern Orthodox ve ...
, which is about a third of the length of the body when the slug is actively crawling and thus extended; when the slug is stationary and contracted, the shield is about half the length of the body. The front of the shield is rounded and its rear is bluntly pointed. The surface texture of this area resembles the underside of undyed leather; it is spotted with pale, buff or light-coloured spots that are similar to those on the body but are more uniformly distributed. The foot fringe, a band of tissue around the edge of the foot, is not distinctly separated; it is very pale and somewhat expanded and has indistinct lines on it. The sole of the foot is pale grey-yellow and is divided into three indistinct bands; the mid-area is somewhat darker and more transparent than the side bands. There is a
caudal mucous pit The caudal mucous pit, or caudal mucous horn, is an anatomical structure on the tail end of the foot of various land snails and slugs, terrestrial animal, terrestrial pulmonate gastropod mollusc, mollusks. The function of this pit is the resor ...
situated between the foot and the body on the upper surface of the tip of the tail. The pit, which collects extra mucus, is not conspicuous, triangular and opens transversely. The mucous pit often carries a transparent, yellowish ball of mucus. The Kerry slug's upper tentacles are smoky-black or grey, short and thick with oval ends, and have eye spots at their tips. The
genital pore A sex organ (or reproductive organ) is any part of an animal or plant that is involved in sexual reproduction. The reproductive organs together constitute the reproductive system. In animals, the testis in the male, and the ovary in the female, a ...
or opening lies behind and below the right eye tentacle.Scharff, R. F. (1891)
''The slugs of Ireland''
. The Scientific Transactions of the Royal Dublin Society, volume IV., series II. Dublin, Royal Dublin Society; London, Williams & Norgate. 513–563. Cited pages
551
–556.
The lower tentacles are pale-grey and translucent. The skin mucus is usually pale yellow and varies in viscosity. The locomotory mucus is tenacious and usually colourless but is sometimes yellow because of mixing with body slime.


Internal anatomy


Shell

Within the mantle, most land slugs have the remnants of what was in the evolutionary past a larger, external shell. Usually this remnant is either a small, thin, shell-like plate or a collection of
calcareous Calcareous () is an adjective meaning "mostly or partly composed of calcium carbonate", in other words, containing lime or being chalky. The term is used in a wide variety of scientific disciplines. In zoology ''Calcareous'' is used as an ad ...
(chalky) granules. The Kerry slug has an internal shell or shell plate that resembles those found in land slugs of the genus '' Limax''; it is ovoid, solid and chalky with a transparent
conchiolin Conchiolins (sometimes referred to as conchins) are complex proteins which are secreted by a mollusc's outer epithelium (the mantle). These proteins are part of a matrix of organic macromolecules, mainly proteins and polysaccharides, that asse ...
(horny) base. The shell plate is usually convex above and concave beneath and has some indistinct, concentric lines of growth. According to Godwin-Austen, the exterior of the shell plate is covered with a thin, transparent protein layer called the periostracum and with the nucleus—the first part to form—situated near the front. In young Kerry slugs the shell is very thin and convex, abruptly cut off behind, and with an extremely thin layer that projects in front and contains minute granules. Authors have differed in their depictions of the Kerry slug's shell plate but they are consistent in showing it as a solid plate.


Various organ systems

The
circulatory The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, t ...
and excretory systems of the Kerry slug are closely related; the heart is surrounded by the triangular kidney, which has a lamellate (layered) structure and two ureters. In this species, the ventricle of the heart is directed towards, and is very close to, the anal and respiratory openings. The ventricle of the heart is further away and further back than it is in species of the related genus '' Arion'', the type-genus of the family Arionidae. The gland above the foot, the
suprapedal gland The suprapedal gland or mucous pedal gland is an anatomical feature found in some snails and slugs. It is a gland located inside the front end of the foot of gastropods.Barker G. M. (2001) "Gastropods on Land: Phylogeny, Diversity and Adaptive Mo ...
, is deeply imbedded in the tissues and reaches far back. The cephalic (head) gland known as
Semper's organ The Semper's organ is an anatomical structure, a gland located in the head of some land snails, pulmonate gastropod mollusks.Lane J. N. (1964) "Semper's Organ, a Cephalic Gland in Certain Gastropods". ''Quarterly Journal of Microscopical Science ...
is well developed and shows as two strong, flattened lobes. The salivary and digestive glands are the same as those found in ''Arion'' species but the vestigial osphradium (kidney-like structure) within the mantle chamber is more distinct than it is in ''Arion'' species.


Muscles

In the Kerry slug, the cephalic retractors (muscles for pulling in the head) are very similar to those in ''Arion'' species. The right and left tentacular muscles, which pull in all four of the tentacles, divide early for the upper and lower tentacles but only the muscles of the ommatophores—the two upper tentacles, which have eye spots—are darkly pigmented. The right and left muscles that pull in the eye-spot tentacles are attached at the base to the back edge of the mantle on the right and left respectively. The pharyngeal (throat) retractor muscle is furcate (split) where it attaches to the back of the buccal bulb (mouth bulb); its other end is anchored on the right side of the body, just behind the site of attachment of the right tentacular muscle.


Reproductive system

The Kerry slug is a hermaphrodite, as are all pulmonates. Various authors have depicted its
reproductive system The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are ...
: Godwin-Austen (1882), Godwin-Austen, H. H. (1882). ''Land and freshwater mollusca of India, including South Arabia, Baluchistan, Afghanistan, Kashmir, Nepal, Burma, Pegu, Tenasserim, Malaya Peninsula, Ceylon and other islands of the Indian Ocean; Supplementary to Masers Theobald and Hanley's Conchologica Indica.'' ''Plates to Volume I.'' Taylor and Francis, London
Plate XII
, figure 5.
Sharff (1891), Simroth (1891, 1894), Taylor (1907), Germain (1930), Quick (1960) and Platts & Speight (1988). Platts & Speight considered the depiction by Godwin-Austen (1882) to be the most accurate of those by earlier authors; others depicted the atrium too short. The ovotestis—a combination of ovary and testis—is small, compact and darkly pigmented. The hermaphroditic duct, where sperm is stored, is long and convoluted, and ends in a small, spherical, seminal vesicle. The albumen gland, which produces albumen for the eggs, is elongated and shaped like a tongue. The ovispermatoduct, along which both eggs and sperm pass, is greatly twisted. This turns into the free oviduct after the vas deferens carrying the sperm branches off. The free oviduct is long and consistently thin. It opens into the atrium near the genital pore, where the muscular atrium is greatly but irregularly enlarged and connected by muscle fibres to the oviduct. The vas deferens is long, complexly twisted, and rolled in a bundle. The bursa copulatrix for digesting spermatophore and sperm—earlier literature refers to this as the spermatheca—is globular and has a short
bursa duct ( grc-gre, Προῦσα, Proûsa, Latin: Prusa, ota, بورسه, Arabic:بورصة) is a city in northwestern Turkey and the administrative center of Bursa Province. The fourth-most populous city in Turkey and second-most populous in the ...
. There is a long retractor muscle from the bursa duct, its other end is anchored near the tail of the slug at the midline. The vas deferens and the bursa duct open nearly together into the far extremity of the atrium, the duct into which both the male and the female systems open and which connects to the outside via the genital pore. A special feature of the genus ''Geomalacus'', is the extremely elongated atrium. The elongated portion of the atrium further from the genital pore than the insertion of the oviduct is termed the atrial diverticulum. In ''Geomalacus'', the penis and its penial retractor muscle have been lost. The atrial diverticulum has been proposed to be the functional equivalent, homoplasy) of a penis, acting as a copulatory organ. Pilsbry H. A. (1898). "Phylogeny of the genera of Arionidae". '' Proceedings of the Malacological Society of London'' 3
94
-104.
It is presumed that the bursa retractor muscle retracts the atrial diverticulum. In ''Geomalacus maculosus'', the atrial diverticulum is longer than the bursa duct; this situation is reversed in ''
Geomalacus anguiformis ''Geomalacus anguiformis'' is a species of air-breathing land slug, a terrestrial pulmonate gastropod mollusc in the family Arionidae, the round back slugs. Distribution This species occurs in south Portugal (Serra de Monchique) and central Sp ...
''. Godwin-Austen noted that the part of the atrium just inside the genital pore—he called this region the "vagina"—has "a curious arrangement" of flattened folds. The central part, situated close to the genital pore, has a pointed end. He compared this to the calcareous darts in other genera; on the preceding pages he had described such structures in the Asian slug genus Anadenus).


Apparatus for feeding


=Radula

= The
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
, which is located inside the mouth, is a feeding structure that is unique to molluscs. Typically, it is a small, strong, ribbon-like structure that bears numerous complex rows of tiny teeth across it. In the Kerry slug, the radula is long and wide, and has 240 slightly curved, transverse rows of denticles; tiny teeth. Each row of teeth is composed of one median tooth and 10 lateral and marginal teeth on each side. The median teeth are small, have one cusp and are slightly shouldered. The lateral teeth have two cusps. The admedian (next to the middle) teeth are larger than the median row and the mesocone—an extra protrusion in the middle of the tooth—is well developed. The only difference between the lateral and marginal series is that the ectocone (extra little side protrusion) present on the admedian teeth recedes in position and slightly diminishes in size in the succeeding teeth up to about the 20th row on the radula. In the marginal series, however, the ectocone gradually grows in size and importance as the margin is approached while the mesocone becomes almost correspondingly diminished. The outermost teeth show a more embryonic character.


=Jaw

= The jaw of the Kerry slug is about from side to side and is distinctly arched from front to rear, crescent-shaped and very wide with broad and slightly rounded ends. The jaw is solid, dark-brown and has about 10 broad flat ribs in the middle part of the jaw. These ribs are absent or scarcely discernible on the side areas. Where the ribs meet the upper edge, they sometimes form crenulations (a scalloped effect) and may also produce the same effect on the lower edge of the jaw. In other individuals, the ribs extend across the jaw, making both the upper and the cutting edges of the jaw clearly toothed in outline. In the Kerry slug, as in all species within the family Arionidae, the alimentary canal of the digestive system forms two loops."Family summary for Arionidae"
. AnimalBase, last change 12-06-2009, accessed 4 August 2010.


Distribution

The Kerry slug has a discontinuous or disjunct distribution; it is found only in
Ireland Ireland ( ; ga, Éire ; Ulster-Scots: ) is an island in the North Atlantic Ocean, in north-western Europe. It is separated from Great Britain to its east by the North Channel, the Irish Sea, and St George's Channel. Ireland is the s ...
—mostly the south-western corner— in north-western
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = '' Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , ...
, and central-to-northern
Portugal Portugal, officially the Portuguese Republic ( pt, República Portuguesa, links=yes ), is a country whose mainland is located on the Iberian Peninsula of Southwestern Europe, and whose territory also includes the Atlantic archipelagos of th ...
. It was once reported as occurring in France but this has not been confirmed and that record is considered suspect. Similar distribution patterns have been observed in other species of animals and plants. This particular disjunct distribution in Iberia and Ireland with no intermediate localities is known as a "
Lusitanian distribution In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but considerably separated from each other geographically. The causes are varied and might demonstrate either the expansion or contraction of a s ...
". There has been speculation that ''G. maculosus'' was introduced to Ireland from Iberia by prehistoric humans; a similar introduction appears to have happened with the
Eurasian pygmy shrew The Eurasian pygmy shrew (''Sorex minutus''), often known simply as the pygmy shrew, is a widespread shrew of the northern Palearctic. Description It measures about in length, not including the tail, and has an average weight of .The hair on t ...
. In support of such an origin or of a more recent human-mediated introduction, the genetic diversity of the Kerry slug in Ireland was found to be greatly reduced compared with that of the Iberian populations.


Ireland

Within Ireland, the Kerry slug is known to occur in areas with sandstone geology in West Cork and
County Kerry County Kerry ( gle, Contae Chiarraí) is a county in Ireland. It is located in the South-West Region and forms part of the province of Munster. It is named after the Ciarraige who lived in part of the present county. The population of the co ...
, an area of around . In 2010, a previously unknown population was recorded further north in County Galway.


Protected sites

A significant proportion of the Kerry slug's range in Ireland is protected by being included in Special Areas of Conservation (SACs). In response to European environmental legislation, Ireland has designated seven SACs with the slug named as a "selection feature": Glengarriff Harbour and Woodland; Caha Mountains; Sheep's Head;
Killarney National Park Killarney National Park ( ga, Páirc Náisiúnta Chill Airne), near the town of Killarney, County Kerry, was the first national park in Ireland, created when the Muckross Estate was donated to the Irish Free State in 1932. The park has sinc ...
, MacGillycuddy's Reeks and Caragh River Catchment; Lough Yganavan and Lough Nambrackdarrig; Cloonee and Inchiquin Loughs, Uragh Wood and
Blackwater River (Kerry) The Blackwater River or Kerry Blackwater is a river in County Kerry, southwestern Ireland. It is not to be confused with the much longer Munster Blackwater, which also rises in County Kerry. Geography The Kerry Blackwater has a drainage basin ...
. In addition, St. Gobnet's Wood SAC (which was designated in relation to other selection criteria) was expanded in 2008 to protect Cascade Wood, a small area of woodland which is inhabited by the slug.Ketch, Catherine (2012)
Kerry Slug researcher visits Baile Bhúirne and Beara
''
The Corkman ''The Corkman'' is a weekly Irish newspaper. It is part of the Corkman Group and based in Mallow. It is owned by the Independent News & Media group. The paper is primarily a North Cork newspaper, and also publishes a separate edition for M ...
''
The species has also been recorded at other SACs where it is not a selection feature, for example in Derryclogher Bog, County Cork.Threat Response Plan
, Ireland: National Parks & Wildlife Service. Retrieved 24 June 2012


Iberia: Spain and Portugal

Despite its first discovery at Caragh Lake and its English common name of Kerry slug, Ireland is at the periphery of this slug species' distribution; in terms of genetic diversity the distribution is centred on the north-western parts of the
Iberian peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, def ...
. The Kerry slug has been known in northern Spain since 1868 and in northern Portugal since 1873.


Portugal

The southernmost locality where this species is found is the mountain range Serra da Estrela in Portugal. It is also found in the provinces
Beira Alta Beira Alta may refer to: * Beira Alta Province, a province in the north of Portugal * Beira Alta (region) Beira Alta was one of the 13 regions of continental Portugal identified by geographer Amorim Girão, in a study published between 1927 and 19 ...
,
Douro Litoral Douro Litoral () is a historical province of Portugal. It is centered on the city of Porto, now the capital of the Norte Region. Other important cities in the province are Vila Nova de Gaia, Matosinhos, Maia, Póvoa de Varzim, and the historical ...
, Minho,
Trás-os-Montes e Alto Douro Trás-os-Montes () is a geographical, historical and cultural region of Portugal. Portuguese for "behind the mountains", Trás-os-Montes is located northeast of the country in an upland area, landlocked by the Douro and Tâmega rivers to south ...
and in the Peneda-Gerês National Park.


Spain

In Spain, the distribution of this species includes coastal locations in Galicia and extends through the Cantabrian Mountains as far east as Mount Ganekogorta in the Basque Country. These localities fall within the boundaries of various autonomous communities: Galicia, Asturias, Cantabria,
Castile and León Castile and León ( es, Castilla y León ; ast-leo, Castiella y Llión ; gl, Castela e León ) is an autonomous community in northwestern Spain. It was created in 1983, eight years after the end of the Francoist regime, by the merging of th ...
(provinces of León, Palencia and Zamora), and the Basque Country provinces of
Biscay Biscay (; eu, Bizkaia ; es, Vizcaya ) is a province of Spain and a historical territory of the Basque Country, heir of the ancient Lordship of Biscay, lying on the south shore of the eponymous bay. The capital and largest city is Bilbao. ...
and
Álava Álava ( in Spanish) or Araba (), officially Araba/Álava, is a Provinces of Spain, province of Spain and a historical territory of the Basque Country (autonomous community), Basque Country, heir of the ancient Basque señoríos#Lords of Álav ...
. There have been unconfirmed findings of this slug from
Navarra Navarre (; es, Navarra ; eu, Nafarroa ), officially the Chartered Community of Navarre ( es, Comunidad Foral de Navarra, links=no ; eu, Nafarroako Foru Komunitatea, links=no ), is a foral autonomous community and province in northern Spa ...
.


=Protected sites

= Natura 2000 sites for this species in Spain include 48 localities (listed below, grouped by region). As at 2017, some of these sites have yet to be designated as Special Areas of Conservation: *Asturias :: Muniellos; PongaAmieva; Redes; *Cantabria :: Camesa river; Liebana (Special Area of Conservation; Liébana ( Special Protection Area); "Upper valleys of the Nansa and Saja and
Alto Campoo Alto Campoo is a ski resort in the Cantabrian Mountains of northern Spain. The resort is located in the Cantabrian comarca of Campoo. The source of the river Ebro is near the resort in the town of Fontibre. Resort With of marked pistes, it i ...
"); *Castile and León :: Hoces de Vegacervera; Lake Sanabria and its vicinities; Montes Aquilanos (
Site of Community Importance A Site of Community Importance (SCI) is defined in the European Commission Habitats Directive (92/43/EEC) as a site which, in the biogeographical region or regions to which it belongs, contributes significantly to the maintenance or restoration ...
); Montes Aquilanos y Sierra de Teleno (SPA); :: Natural Park of Fuentes Carrionas and Fuente Cobre-Montaña Palentina (SAC);
Sierra de la Cabrera The Sierra de la Cabrera is a mountain range in northern Spain. The landscape shows evidence of past glaciation. Ecological importance The Sierra de la Cabrera gives its name to a Site of Community Importance (ES4190110), where species of inter ...
(SCI partially overlapping with a SPA of the same name). *Galicia ::A Marronda;
Anllóns river The Anllóns is a Galician river that begins in the Xalo Mountains at an elevation of above sea level and flows into the Atlantic via the Ria of Corme and Laxe. Along its course, it crosses the municipalities of Cabana de Bergantiños, Carball ...
; Baixa Limia;
Baixa Limia - Serra do Xurés The Baixa ''(Downtown)'' (), also known as the Baixa Pombalina (''Pombaline Downtown'') is a neighborhood in the historic center of Lisbon, Portugal. It consists of the grid of streets north of the Praça do Comércio, roughly between the Cais do ...
; Baixo Miño; Bidueiral de Montederramo; Carballido, a yew wood in
A Fonsagrada A Fonsagrada is a town and municipality in the province of Lugo in the autonomous community of Galicia in northwest Spain which is 25 miles east-north-east of Lugo by road. Its population in 2004 was 5,007. A Fonsagrada is situated above sea ...
;
Carnota Carnota is a municipality of northwestern Spain in the province of A Coruña, autonomous community of Galicia. It belongs to the comarca of Muros. It has an area of 66.4 km2, a population of 5,285 (2004 estimate) and a population density ...
 – Monte Pindo;
Cíes Islands The Cíes Islands ( gl, Illas Cíes, link=no; es, Islas Cíes, link=no) are an archipelago off the coast of Pontevedra in Galicia, Spain, in the mouth of the Ria de Vigo. They were declared a Nature Reserve in 1980 and are included in the Atla ...
; Costa Ártabra; Costa da Morte—''Costa da Morte'' and ''Costa da Morte (Northern)''; Cruzul-Agüeira; Encoro de Abegondo-
Cecebre Cambre is a municipality in the Province of A Coruña, in the autonomous community of Galicia in northwestern Spain. It is located 12 km from the capital city of A Coruña and ten minutes away from the city's airport, Alvedro. According to ...
;
Eo river The Eo is a river, long, in northwestern Spain. Its estuary forms the boundary between the regions of Galicia and Asturias. The river is known for its salmon fishing. In the western Cantabrian mountains the river forms the axis of one of Asturi ...
is included among the Galician sites although the estuary forms the boundary with Asturias; Costa de Ferrolterra-
Valdoviño Valdoviño is a municipality in the province of A Coruña in the autonomous community of Galicia in northwestern Spain. It is located in the comarca of Ferrol. Valdoviño has a population of 6,926 inhabitants (INE, 2011). Geography The gr ...
;
Fragas do Eume The Fragas do Eume is a natural park situated in north-west Spain. ''Fraga'' is a Galician word for "natural woodland", (old-growth forest) and the park is an example of a temperate rainforest in which oak (''Quercus robur'' and ''Quercus pyren ...
; Macizo Central, Ourense (province); Monte Aloia; Monte Maior; Negueira;
Pena Trevinca Peña Trevinca ( gl, Pena Trevinca), also known as Trevinca, is a mountain in northern Spain. It is located at the confluence of the Montes de León and the Macizo Galaico on the boundary between the autonomous communities of Galicia and Castile ...
; Pena Veidosa; Serra do Candán; Serra do Cando; Serra do Xistral; Sil river canyon; Sobreirais do Arnego;
Tambre Tambre is a '' comune'' (municipality) in the Province of Belluno in the Italian region Veneto, located about north of Venice and about east of Belluno. Tambre borders the following municipalities: Aviano Aviano ( fur, Davian; cim, Pleif) is ...
- two areas, the river and its estuary; Támega river; Ulla-Deza river system *More than one region ::Ancares - This district is divided between Galicia and
Castile and León Castile and León ( es, Castilla y León ; ast-leo, Castiella y Llión ; gl, Castela e León ) is an autonomous community in northwestern Spain. It was created in 1983, eight years after the end of the Francoist regime, by the merging of th ...
. Sierra de los Ancares is a mountain range that forms the boundary between the two autonomous communities and gives its name to a Natura 2000 site in the province of León. On the Galician side of the sierra are two relevant sites—''Ancares'' (protected under the Birds Directive) and ''Ancares-Courel'' (protected under the Habitats Directive). :: Picos de Europa - This mountain range is divided between three autonomous communities. The three sites listed are ''Picos de Europa'', ''Picos de Europa (Asturias)'', and ''Picos de Europa en Castilla y León'', all of which include protected areas in the
Picos de Europa National Park The Picos de Europa National Park ( es, Parque Nacional de Picos de Europa) is a National Park in the Picos de Europa mountain range, in northern Spain. It is within the boundaries of three autonomous communities, Asturias, Cantabria and Casti ...
and in a regional park in Castile and Leon that is also called Picos de Europa.


Behaviour

The Kerry slug is primarily nocturnal. During daylight hours, the slug usually hides in crevices of rocks and under loose bark on trees. In Iberia, juvenile Kerry slugs become active during twilight and adults become active at night, especially on rainy or very humid nights. Because Ireland is much further north and has a considerably cooler, wetter and more humid climate, the Kerry slug is sometimes active there in the daytime if the weather is humid and overcast. The species has in unusual defensive behaviour; whereas most land slugs retract the head and contract the body but stay firmly attached to the substrate when they are attacked or threatened, the Kerry slug retracts its head, lets go of the substrate and rolls itself into a ball-like shape. This is behaviour is unique among species in Arionidae and among slugs in Ireland.


Ecology


Habitat

It was once thought that ''Geomalacus maculosus'' lives only in wild habitats. 9 pp. In the Iberian Peninsula, it occurs on tree trunks in oak (''Quercus'') and chestnut (''Castanea'') forest but it is easiest to find in synanthropic habitats such as rocky walls in oak or chestnut orchards, in ruins, near houses, churches and cemeteries. In Ireland, it also occurs in upland conifer plantations and areas of clear-fell. The Kerry slug is not considered an agricultural pest, unlike some other slugs in the family Arionidae. In Ireland, the Kerry slug occurs in woodland with oak trees, oligotrophic open moorland, blanket bogs and lake shores, especially if boulders covered with lichens and mosses are present in these habitats. Although there was a geographical association with sandstone areas, the new locality in Galway is on granite. In Iberia it usually occurs in granite mountains, and on slates, quartzite, schists, gneiss and serpentine. The best predictor of its occurrence is high rainfall and high summer temperatures.


Feeding

The food of ''Geomalacus maculosus'' includes lichens, liverworts, mosses,
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
(''
Fistulina hepatica ''Fistulina hepatica'' (beefsteak fungus, also known as beefsteak polypore, poor man’s steak, ox tongue, or tongue mushroom) is an unusual bracket fungus classified in the Agaricales, that is commonly seen in Britain, but can be found in Nort ...
'') and bacteria that grow on boulders and on tree trunks. In captivity, the Kerry slug has been fed on porridge, bread, dandelion leaves, lichen ''
Cladonia fimbriata ''Cladonia fimbriata'' or the trumpet cup lichen is a species cup lichen belonging to the family Cladoniaceae. As of July 2021, its conservation status has not been estimated by the IUCN. In Iceland, its conservation status is denoted as data ...
'', carrot, cabbage, cucumber and lettuce. It can be carnivorous in captivity; there are records of it consuming the snail ''
Vitrina pellucida ''Vitrina pellucida'' is species of small land snail, a terrestrial pulmonate gastropod mollusk in the family Vitrinidae, the glass snails. Description This species is a 'semi-slug' with a flattened, globular shell. The animal is pale grey w ...
''.


Life cycle

The Kerry slug mates in head-to-head position with partners' genital openings facing each other. The sexual organs, called atria—singular:atrium—are funnel-shaped with fluted edges after mating. As in ''Arion'', sperm is transferred in a
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophore ...
. In the wild, eggs are laid between July and October, and from February to October in captivity.
Self-fertilisation Autogamy, or self-fertilization, refers to the fusion of two gametes that come from one individual. Autogamy is predominantly observed in the form of self-pollination, a reproductive mechanism employed by many flowering plants. However, species ...
is also possible in this species. Eggs are laid in clusters of 18 to 30, and are held together by a film of mucus. The egg masses are about . The eggs are very large compared with the size of the animal. The largest eggs are more elongate, being ; the smallest are more ovoid and are . All are semi-translucent, milky-white or opalescent when fresh, although some of the larger and more elongate eggs have a semi-transparent area at the smaller end. The opalescent lustre disappears in a few days and the eggs turn yellowish and later brown or black. The young appear to hatch in six to eight weeks, at this stage the spots on the body are barely present. The lateral bands are distinct and black, and are more conspicuous than they are in mature slugs of this species. In juveniles the shield shows lyre-shaped markings, as is the case in slugs of the genus ''Arion''. These lyre-shaped markings become indistinct as the slugs grow larger. The Kerry slug probably overwinters in the sexually immature stage. The bodies of preserved juvenile specimens are up to long with a mantle length of . Juveniles reach maturity in two years, at a length about . In the wild, the Kerry slug can live for up to seven years but in captivity, the lifespan rarely exceeds three years. In numerous localities in Spain, very few individuals of the species were observed at any one time. Until 2014, the natural enemies of ''Geomalacus maculosus'' were not known. The Kerry slug's predators include larvae of the third
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...
of the fly ''
Tetanocera elata ''Tetanocera elata'' is a species of fly in the family Sciomyzidae. It is found in the Palearctic Larvae of ''T. elata'' are known obligate feeders on slugs both as parasitoids and predators.Lloyd Vernon Knutson and Jean-Claude Vala, 2011''Biol ...
''.


Parasites

''G. maculosus'' suffers
parasitism Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson ha ...
by nematodes. Carnaghi et al 2017 find '' Phasmarhabditis californica'' parasitising ''G. maculosus'' in Ireland however they also find the slug is immune to a similar nematode, '' P. hermaphrodita''.


Threats

The most serious threat to the Kerry slug is probably the modification of habitat, which reduces its lichen and moss food sources. This can lead to the local disappearance of the species, which was documented in Spain. Other threats include intensification of land use, land reclamation, use of pesticides, overgrazing by sheep, removal of shrubs, tourism, general development pressures, planting of conifer plantations, the spread of invasive plants such as '' Rhododendron ponticum'' and
habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological process ...
(see also Moorkens 2006). Other potential dangers to the species are
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
and
air pollution Air pollution is the contamination of air due to the presence of substances in the atmosphere that are harmful to the health of humans and other living beings, or cause damage to the climate or to materials. There are many different type ...
, which negatively affect the lichens eaten by the Kerry slug. Climate change will probably affect the Iberian populations more acutely because the
climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...
there is already hot and dry relative to that of Ireland, which is generally cool and damp.


Conservation measures


International protection

Because of its perceived rarity and its restricted distribution, the Kerry slug is protected under the Convention on the Conservation of European Wildlife and Natural Habitats (Bern Convention), EIS Bern Invertebrates Project. This decision was backed by studies of its distribution and ecology in Ireland, which concluded that evidence of a decline in Iberia and uncertainty over its status in Ireland tended to support its inclusion in the convention. Since 2006, ''Geomalacus maculosus'' has been considered a
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. ...
species in the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biolo ...
; between 1994 and 2006, however, the slug was rated as vulnerable. ''Geomalacus maculosus'' is also protected by the European Union's Habitats Directive and has been listed as an Annex II and Annex IV species since 1992. The principal mechanisms used by the Directive to protect habitats and species are the creation of Special Areas of Conservation (SACs) and the protection of species independently of their habitats by other means. Seven SACs have been designated for this species in Ireland and 49 SCIs in Spain. Threats to the Kerry slug will probably be greatest in areas not specifically protected as SACs. The Habitats Directive protects the Kerry slug outside the SACs by Article 12 (1), which obliges European Union member states to: * establish 'a system of strict protection' for listed species * prohibit deliberate capture or killing * prohibit 'deliberate disturbance ... particularly during the period of breeding, rearing, hibernation and migration' * prohibit 'deliberate destruction or taking of eggs from the wild' * prohibit the deliberate or non-deliberate 'deterioration or destruction of breeding sites or resting places'.


Protection in Iberia

Conservation status reports from Portugal and Spain were not yet available in August 2009. Its conservation status in Spain for the IUCN criteria is vulnerable.


Protection in Ireland

In 1988, Platts and Speight noted that only three of the Irish sites where the slug occurred were protected;
Glengariff Forest Glengarriff Forest is an area of woodland near Glengarriff, West Cork, Ireland. Most of the woodland is a nature reserve in public ownership which is sometimes referred to as Glengarriff "forest park" or "state forest". Glengarriff Forest is ...
, West Cork; Uragh Wood Nature Reserve, South Kerry; and
Killarney National Park Killarney National Park ( ga, Páirc Náisiúnta Chill Airne), near the town of Killarney, County Kerry, was the first national park in Ireland, created when the Muckross Estate was donated to the Irish Free State in 1932. The park has sinc ...
, North Kerry. They concluded that the species could not be adequately safeguarded with only three sites and supported its inclusion in the Bern list, to which the Irish government is a signatory. The Habitats Directive was
transposed In linear algebra, the transpose of a matrix is an operator which flips a matrix over its diagonal; that is, it switches the row and column indices of the matrix by producing another matrix, often denoted by (among other notations). The tr ...
into
Irish law Law of Ireland or Irish law may refer to: * Early Irish law (Brehon law) of Medieval Ireland * Alternative law in Ireland prior to 1921 * Law of the Republic of Ireland The law of Ireland consists of constitutional, statute, and common law. ...
by: * The EC (Natural Habitats) Regulations 1997. This was the principal legislation transposing the Habitats Directive and upgraded the protection of the Kerry slug's habitat by the designation of Special Areas of Conservation. * Adapting existing legislation. The Kerry slug has been protected since 1990 under the Irish Wildlife Act of 1976; it was added to the list of protected species by
Statutory Instrument In many countries, a statutory instrument is a form of delegated legislation. United Kingdom Statutory instruments are the principal form of delegated or secondary legislation in the United Kingdom. National government Statutory instrumen ...
112/1990, and was the only gastropod so protected. 15 pp., cited page: p. 12
mentioned here on pg 20 and 22
/ref> The Wildlife Act does not protect the slug from indirect damage but only from wilful direct damage such as collecting. The Irish National Parks and Wildlife Service published a Species Action Plan for the Kerry slug in January 2008. Efforts were made to protect the slug from indirect damage, for instance from commercial forestry. Following a legal challenge to Ireland's transposition and implementation of the Habitats Directive, however, the Action Plan was superseded in May 2010 by a Threat Response Plan that addressed problems that arose when the
European Court of Justice The European Court of Justice (ECJ, french: Cour de Justice européenne), formally just the Court of Justice, is the supreme court of the European Union in matters of European Union law. As a part of the Court of Justice of the European U ...
held that Ireland was not protecting the Kerry slug with the strictness the directive required for a species listed in annex 4.


Monitoring

In a report to the
European Commission The European Commission (EC) is the executive of the European Union (EU). It operates as a cabinet government, with 27 members of the Commission (informally known as "Commissioners") headed by a President. It includes an administrative body ...
covering 1988–2007, the conservation status of the species in Ireland was declared "favourable (FV)" in all evaluated criteria; range, population, habitat and future prospects. The validity of this assessment, however, was put into question by the European Court of Justice ruling that held that Ireland was not monitoring the slug properly. The need to improve monitoring was discussed by the NPWS Threat Response Plan of 2010, which recognised that population statistics were still deficient, particularly outside the SACs. As the Threat Response Plan noted, species monitoring is a process in which distribution and status of the subject are evaluated systematically over time. Under this definition, no monitoring of the Kerry Slug had been undertaken in Ireland as of May 2010. The Kerry Slug Survey of Ireland, a collaboration between the National Parks and Wildlife Service and the Applied Ecology Unit at the National University of Ireland, Galway, researched a "suitable monitoring protocol" for the species.Kerry Slug Survey of Ireland (Official Website)
Retrieved 3 July 2010.
The Kerry Slug Survey's investigations resulted in the publication of a guide to the population dynamics of the Kerry slug, which was published as part of the ''Irish Wildlife Manual'' series in 2011.Mc Donnell, R.J. and Gormley M.J. (2011)
Distribution and population dynamics of the Kerry Slug
''Irish Wildlife Manual'' No 54, National Parks and Wildlife Service, Department of Arts, Heritage and the Gaeltacht, Dublin


Captive breeding

Since 1990, the Kerry slug has been successfully bred in captivity. The Wildfowl & Wetlands Trust, a British conservation organisation, operates a captive breeding programme in terraria at its "Endangered Species Breeding Unit". The project is located at the Martin Mere Wetland Centre in
Lancashire Lancashire ( , ; abbreviated Lancs) is the name of a historic county, ceremonial county, and non-metropolitan county in North West England. The boundaries of these three areas differ significantly. The non-metropolitan county of Lancas ...
, England. During the 1990s, slugs from the breeding programme were given out to a number of zoos and individuals to set up their own breeding programmes but very few of those breeding groups survived.


References

This article incorporates public domain text from Taylor (1907).


Further reading

* Allman, G. J. (1844). "On a new genus of terrestrial gastropod". ''Report of the British Association for the Advancement of Science'
1843
77. * Allman, G. J. (1846). "Description of a new genus of pulmonary gastropods". '' Annals and Magazine of Natural History'' 17: 297–299, plate 9. * * Oldham, C. (1942).
Notes on ''Geomaculus maculosus''
. ''Proceedings of the Malacological Society of London'' 25. * Heynemann, D. F. (1873). "On the French species of the genus ''Geomalacus''". ''Annals and Magazine of Natural History'', pages 271–275. * Heynemann, D. F. (1869). "Zur Kenntniss von ''Geomalacus''". ''Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft'', pages 165–168. * Heynemann, D. F. (1871). "''Geomalacus maculosus''". ''Nachrichtsblatt der Deutschen Malakozoologischen Gesellschaft'' 3(1)
126
* Heynemann, D. F. (1873). "Ueber ''Geomalacus''". ''Malakozoologische Blätter'
xxi25
��36
table 1
fig. 1–6. * Moorkens, E. A. (2006). "Irish non-marine molluscs – an evaluation of species threat status". ''Bulletin of the Irish Biogeographical Society'' 30: 348–371. . * * *


External links


''Geomalacus maculosus''
at Animalbase taxonomy, short description, distribution, biology, status (threats), images
Bridges & Species: Post-Glacial Colonisation

A photograph of a live individual

Mollusc Ireland
{{Good article Arionidae Gastropods described in 1843 Molluscs of Europe Taxa named by George Allman (natural historian) Taxonomy articles created by Polbot