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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 appropriate s ...
of terrestrial,
pulmonate Pulmonata or pulmonates, is an informal group (previously an order, and before that a subclass) of snails and slugs characterized by the ability to breathe air, by virtue of having a pallial lung instead of a gill, or gills. The group includ ...
,
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
. It is a medium-to-large sized, air-breathing land
slug Slug, or land slug, is a common name for any apparently shell-less terrestrial gastropod mollusc. The word ''slug'' is also often used as part of the common name of any gastropod mollusc that has no shell, a very reduced shell, or only a smal ...
in the
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of roundback slugs, Arionidae. 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 dialect, Ulster-Scots: ) is an island in the Atlantic Ocean, North Atlantic Ocean, in Northwestern Europe, north-western Europe. It is separated from Great Britain to its east by the North Channel (Grea ...
—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 and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, defi ...
, 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 Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
or
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
but in Ireland it is active on overcast days. It feeds on
lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.liverwort The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of g ...
s,
moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es 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 from ...
, 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 The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
, 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 differentl ...
of
molluscs Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant taxon, extant species of molluscs are recognized. The number of fossil sp ...
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 d ...
Panpulmonata Panpulmonata is a taxonomic clade of snails and slugs in the clade Heterobranchia within the clade Euthyneura. Panpulmonata was established as a new taxon by Jörger et al. in October 2010. The older name "Pulmonata" referred to a group of gast ...
, 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 Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
is Arionidae, 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 distr ...
and
Milacidae Milacidae is a family of air-breathing, keeled, land slugs. These are shell-less terrestrial gastropod mollusks in the superfamily Parmacelloidea. This family has no subfamilies (according to the taxonomy of the Gastropoda by Bouchet & Rocro ...
. Many of its anatomical features are shared with species in the genus ''
Arion Arion (; grc-gre, Ἀρίων; fl. c. 700 BC) was a kitharode in ancient Greece, a Dionysiac poet credited with inventing the dithyramb. The islanders of Lesbos claimed him as their native son, but Arion found a patron in Periander, tyrant ...
'', which is a more species-rich and widely distributed group of slugs within Arionidae. The Kerry slug is placed in the genus '' Geomalacus'', which means "earth mollusc". The Kerry slug's
scientific name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
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 wiktionary:en:specimen, 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 a ...
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 George James Allman FRS FRSE (181224 November 1898) was an Irish ecologist, botanist and zoologist who served as Emeritus Professor of Natural History at Edinburgh University in Scotland. Life Allman was born in Cork, Ireland, the son of Jam ...
. 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 In biology, taxonomy () is the scientific study of naming, defining ( circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given ...
, is ''Geomalacus maculosus'' Allman, 1843. The
synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
s 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 Nomination is part of the process of selecting a candidate for either election to a public office, or the bestowing of an honor or award. A collection of nominees narrowed from the full list of candidates is a short list. Political office In the ...
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 (psychology), affect.Rammirez-Moreno, D ...
. 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, 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 pulmonate gastropod mollusks. The function of this pit is the resorption of mucus when the gast ...
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 The Royal Dublin Society (RDS) ( ga, Cumann Ríoga Bhaile Átha Cliath) is an Irish philanthropic organisation and members club which was founded as the 'Dublin Society' on 25 June 1731 with the aim to see Ireland thrive culturally and economi ...
; 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 Shell may refer to: Architecture and design * Shell (structure), a thin structure ** Concrete shell, a thin shell of concrete, usually with no interior columns or exterior buttresses ** Thin-shell structure Science Biology * Seashell, a hard ou ...
. 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 adje ...
(chalky) granules. The Kerry slug has an internal shell or shell plate that resembles those found in land slugs of the genus ''
Limax ''Limax'' is a genus of air-breathing land slugs in the terrestrial pulmonate gastropod mollusk family Limacidae. The generic name ''Limax'' literally means "slug". Some species, such as the leopard slug (''L. maximus'') and the tawny garden ...
''; 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 assem ...
(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 The periostracum ( ) is a thin, organic coating (or "skin") that is the outermost layer of the shell of many shelled animals, including molluscs and brachiopods. Among molluscs, it is primarily seen in snails and clams, i.e. in gastropods and ...
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, tha ...
and
excretory system The excretory system is a passive biological system that removes excess, unnecessary materials from the body fluids of an organism, so as to help maintain internal chemical homeostasis and prevent damage to the body. The dual function of excreto ...
s of the Kerry slug are closely related; the heart is surrounded by the triangular kidney, which has a lamellate (layered) structure and two
ureter The ureters are tubes made of smooth muscle that propel urine from the kidneys to the urinary bladder. In a human adult, the ureters are usually long and around in diameter. The ureter is lined by urothelial cells, a type of transitional epit ...
s. In this species, the ventricle of the heart is directed towards, and is very close to, the
anal Anal may refer to: Related to the anus *Related to the anus of animals: ** Anal fin, in fish anatomy ** Anal vein, in insect anatomy ** Anal scale, in reptile anatomy *Related to the human anus: ** Anal sex, a type of sexual activity involvin ...
and
respiratory The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies grea ...
openings. The ventricle of the heart is further away and further back than it is in species of the related genus ''
Arion Arion (; grc-gre, Ἀρίων; fl. c. 700 BC) was a kitharode in ancient Greece, a Dionysiac poet credited with inventing the dithyramb. The islanders of Lesbos claimed him as their native son, but Arion found a patron in Periander, tyrant ...
'', the type-genus of the family Arionidae. The gland above the foot, the suprapedal gland, is deeply imbedded in the tissues and reaches far back. The cephalic (head) gland known as Semper's organ is well developed and shows as two strong, flattened lobes. The
salivary The salivary glands in mammals are exocrine glands that produce saliva through a system of ducts. Humans have three paired major salivary glands (parotid, submandibular, and sublingual), as well as hundreds of minor salivary glands. Salivary glan ...
and digestive glands are the same as those found in ''Arion'' species but the vestigial
osphradium The osphradium is a pigmented chemosensory epithelium patch in the mantle cavity present in six of the eight extant classes of molluscs (it is absent in the scaphopoda and monoplacophora; among cephalopoda, only the nautilus has what appears to be ...
(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 In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...
, 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 als ...
: 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 An ovotestis is a gonad with both testicular and ovarian aspects. In humans, ovotestes are an infrequent anatomical variation associated with gonadal dysgenesis. The only mammals where ovotestes are not symptomatic of an intersex variation are mole ...
—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 The vas deferens or ductus deferens is part of the male reproductive system of many vertebrates. The ducts transport sperm from the epididymis to the ejaculatory ducts in anticipation of ejaculation. The vas deferens is a partially coiled tube ...
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. 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 Homoplasy, in biology and phylogenetics, is the term used to describe a Phenotypic trait, feature that has been gained or lost independently in separate lineages over the course of evolution. This is different from Homology (biology), homology, w ...
) 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 The ''Journal of Molluscan Studies'' is the peer-reviewed scientific journal of the Malacological Society of London, covering research in malacology.
'' 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''. 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 Darts or dart-throwing is a competitive sport in which two or more players bare-handedly throw small projectile point, sharp-pointed projectile, missiles known as dart (missile), darts at a round shooting target, target known as a #Dartboard, dar ...
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 The human digestive system consists of the gastrointestinal tract plus the accessory organs of digestion (the tongue, salivary glands, pancreas, liver, and gallbladder). Digestion involves the breakdown of food into smaller and smaller compone ...
forms two loops."Family summary for Arionidae"
.
AnimalBase AnimalBase is a project brought to life in 2004 and is maintained by the University of Göttingen, Germany. The goal of the AnimalBase project is to digitize early zoological literature, provide copyright-free open access to zoological works, and pr ...
, last change 12-06-2009, accessed 4 August 2010.


Distribution

The Kerry slug has a discontinuous or
disjunct 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 ...
; it is found only in
Ireland Ireland ( ; ga, Éire ; Ulster Scots dialect, Ulster-Scots: ) is an island in the Atlantic Ocean, North Atlantic Ocean, in Northwestern Europe, north-western Europe. It is separated from Great Britain to its east by the North Channel (Grea ...
—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") , i ...
, 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 ...
. 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 West Cork ( ga, Iarthar Chorcaí) is a tourist region and municipal district in County Cork, Ireland. As a municipal district, West Cork falls within the administrative area of Cork County Council, and includes the towns of Bantry, Castletownbe ...
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 "Righteousness and Justice" , anthem = () , image_map = Island of Ireland location map Galway.svg , map_caption = Location in Ireland , area_footnotes = , area_total_km2 = ...
.


Protected sites

A significant proportion of the Kerry slug's range in Ireland is protected by being included in
Special Areas of Conservation A Special Area of Conservation (SAC) is defined in the European Union's Habitats Directive (92/43/EEC), also known as the ''Directive on the Conservation of Natural Habitats and of Wild Fauna and Flora''. They are to protect the 220 habitats and ap ...
(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 The Caha Mountains (''An Cheacha'' in Irish) are a range of low sandstone mountains situated on the Beara peninsula in south-west County Cork, in Ireland. The highest peak is Hungry Hill, tall. Other notable peaks include Knocknagree, Sugarloaf ...
;
Sheep's Head Sheep's Head, also known as Muntervary ( ga, Rinn Mhuintir Bháire), is the headland at the end of the Sheep's Head peninsula situated between Bantry Bay and Dunmanus Bay in County Cork, Ireland. The peninsula is popular with walkers, 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 since ...
,
MacGillycuddy's Reeks , photo=MacGuillycuddy's Reeks.jpg , photo_caption= , country=Ireland , country1= , location = County Kerry , region = Munster , region_type = Provinces of Ireland , parent= , border= , length_km=19 , length_orientation=East–West ...
and Caragh River Catchment;
Lough Yganavan and Lough Nambrackdarrig Lough Yganavan and Lough Nambrackdarrig ( Irish: ''Loch Gaineamháin agus Loch na mBreac Dearg'') are two lakes of ecological importance in County Kerry, Ireland. In 1988, Lough Nambrackdarrig and the southern half of Lough Yganavan were design ...
;
Cloonee and Inchiquin Loughs, Uragh Wood Cloonee and Inchiquin Loughs, Uragh Wood is the name of a Natura 2000 site in a valley in Tuosist, County Kerry, Ireland. Habitats include lowland oligotrophic lakes (loughs) and oceanic oak woodland. Uragh Wood has been protected as a nature ...
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, defi ...
. 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 Serra da Estrela () is the highest mountain range in Continental Portugal. Together with the Serra da Lousã it is the westernmost constituent range of the Sistema Central and also one of the highest in the system. It includes mainland Portugal's ...
in Portugal. It is also found in the provinces Beira Alta,
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 language, Portuguese for "behind the mountains", Trás-os-Montes is located northeast of the country in an highland, upland area, landlocked by the Douro ...
and in the
Peneda-Gerês National Park The Peneda-Gerês National Park ( pt, Parque Nacional da Peneda-Gerês, ), also known simply as Gerês, is a national park located in northern Portugal. Created in May 1971, it is the oldest protected area and the only national park in Portugal. ...
.


Spain

In Spain, the distribution of this species includes coastal locations in Galicia and extends through the
Cantabrian Mountains , etymology=Named after the Cantabri , photo=Cordillera Cantábrica vista desde el Castro Valnera.jpg , photo_caption=Cantabrian Mountains parallel to the Cantabrian Sea seen from Castro Valnera in an east-west direction. In the background, th ...
as far east as Mount Ganekogorta in the Basque Country. These localities fall within the boundaries of various
autonomous communities eu, autonomia erkidegoa ca, comunitat autònoma gl, comunidade autónoma oc, comunautat autonòma an, comunidat autonoma ast, comunidá autónoma , alt_name = , map = , category = Autonomous administra ...
: Galicia,
Asturias Asturias (, ; ast, Asturies ), officially the Principality of Asturias ( es, Principado de Asturias; ast, Principáu d'Asturies; Galician-Asturian: ''Principao d'Asturias''), is an autonomous communities of Spain, autonomous community in nor ...
,
Cantabria Cantabria (, also , , Cantabrian: ) is an autonomous community in northern Spain with Santander as its capital city. It is called a ''comunidad histórica'', a historic community, in its current Statute of Autonomy. It is bordered on the east ...
,
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 the ...
(provinces of León,
Palencia Palencia () is a city of Spain located in the autonomous community of Castile and León. It is the capital and most populated municipality of the province of Palencia. Located in the Northwest of the Iberian Peninsula, in the northern half o ...
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. B ...
and
Álava Álava ( in Spanish) or Araba (), officially Araba/Álava, is a province of Spain and a historical territory of the Basque Country, heir of the ancient Lordship of Álava, former medieval Catholic bishopric and now Latin titular see. Its ca ...
. 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 Spain, ...
.


=Protected sites

=
Natura 2000 Natura 2000 is a network of nature protection areas in the territory of the European Union. It is made up of Special Areas of Conservation and Special Protection Areas designated under the Habitats Directive and the Birds Directive, respectively ...
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; Ponga
Amieva Amieva is a municipality in the autonomous community of Asturias, Spain. It is also the name of one of the municipality's subdivisions (''parroquias'', or parish). Amieva municipality is sparsely inhabited, having a resident population of 868 ( ...
; Redes; *Cantabria :: Camesa river; Liebana (Special Area of Conservation;
Liébana Liébana is a ''comarca'' of Cantabria (Spain). It covers 575 square kilometres and is located in the far southwest of Cantabria, bordering Asturias, León and Palencia. It is made up of the municipalities of: Cabezón de Liébana, Camaleño, Cil ...
(
Special Protection Area A Special Protection Area (SPA) is a designation under the European Union Directive on the Conservation of Wild Birds. Under the Directive, Member States of the European Union (EU) have a duty to safeguard the habitats of migratory birds and cert ...
); "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 is ...
"); *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 at ...
); 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; 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 d ...
; Baixo Miño; Bidueiral de Montederramo; Carballido, a yew wood in A Fonsagrada;
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 of ...
 – 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 (; es, Costa de la Muerte; "Death Coast") is part of the Galician coast. The Costa da Morte extends from the villages of Muros and Malpica. The Costa da Morte received its name because there have been so many shipwrecks along it ...
—''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 Ferrol is a coastal ''comarca'' in the northwest of the province of A Coruña, Galicia, Spain. It is also known as Ferrolterra. The area is 613.4 km2, and the overall population of this ''comarca'' was 161,154 at the 2011 Census; the latest offic ...
-
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 gra ...
;
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) Ourense (in Spanish, ''Orense'') is a Spanish province, in the southeastern part of the autonomous community of Galicia. It is bordered by the provinces of Pontevedra to the west, Lugo to the north, León and Zamora, (which both belong to Cas ...
;
Monte Aloia Monte Aloia is a summit in the mountains of Galicia, Spain, which was declared a natural park on 4 December 1978. The park covers an area of 746 hectares and is located within the municipality of Tui, a town on the River Miño. Topography and ...
; Monte Maior;
Negueira Negueira is one of seven parishes (administrative divisions) in the municipality of Grandas de Salime, within the province and autonomous community of Asturias, in northern Spain. The population is 35 (INE INE, Ine or ine may refer to: Instit ...
; Pena Trevinca; 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, Barcis, Budoia, Caneva, Chies d' ...
- 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 the ...
. 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 The Picos de Europa ("Peaks of Europe", also the Picos) are a mountain range extending for about , forming part of the Cantabrian Mountains in northern Spain. The range is situated in the Autonomous Communities of Asturias, Cantabria and Castil ...
- 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 Castil ...
and in a regional park in Castile and Leon that is also called Picos de Europa.


Behaviour

The Kerry slug is primarily
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
. 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 An oligotroph is an organism that can live in an environment that offers very low levels of nutrients. They may be contrasted with copiotrophs, which prefer nutritionally rich environments. Oligotrophs are characterized by slow growth, low rates of ...
open moorland,
blanket bog Blanket bog or blanket mire, also known as featherbed bog, is an area of peatland, forming where there is a climate of high rainfall and a low level of evapotranspiration, allowing peat to develop not only in wet hollows but over large expanses o ...
s and lake shores, especially if boulders covered with
lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.moss Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es 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,
liverwort The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of g ...
s, 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 from ...
(''
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 Nor ...
'') 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'', 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 molluscs, terrestrial pulmonate gastropod mollusk in the family Vitrinidae, the glass snails. Description This species is a 'semi-slug' with a flattened, globular shell. The ...
''.


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. Spermatophores ...
. In the wild,
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
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 of ...
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 ass ...
of the fly '' Tetanocera elata''.


Parasites

''G. maculosus'' suffers
parasitism Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
by
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s. 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 Land reclamation, usually known as reclamation, and also known as land fill (not to be confused with a waste landfill), is the process of creating new land from oceans, seas, riverbeds or lake beds. The land reclaimed is known as reclamati ...
, use of pesticides, overgrazing by sheep, removal of shrubs, tourism, general development pressures, planting of
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
plantations, the spread of invasive plants such as ''
Rhododendron ponticum ''Rhododendron ponticum'', called common rhododendron or pontic rhododendron, is a species of ''Rhododendron'' native to the Iberian Peninsula in southwest Europe and the Caucasus region in northern West Asia. Description ''R. ponticum'' is a d ...
'' 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 processes ...
(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 E ...
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 types ...
, 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 meteorologic ...
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 The Bern Convention on the Conservation of European Wildlife and Natural Habitats, also known as the Bern Convention (or Berne Convention), is a binding international legal instrument in the field of Nature Conservation, it covers the natural h ...
(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. T ...
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 biol ...
; between 1994 and 2006, however, the slug was rated as vulnerable. ''Geomalacus maculosus'' is also protected by the European Union's
Habitats Directive The Habitats Directive (more formally known as Council Directive 92/43/EEC on the Conservation of natural habitats and of wild fauna and flora) is a directive adopted by the European Community in 1992 as a response to the Berne Convention. The E ...
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 A Special Area of Conservation (SAC) is defined in the European Union's Habitats Directive (92/43/EEC), also known as the ''Directive on the Conservation of Natural Habitats and of Wild Fauna and Flora''. They are to protect the 220 habitats and ap ...
(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, 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 since ...
, 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 * Law of Northern Ireland The law of Northern Ireland is the legal system ...
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 instrument ...
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 Un ...
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
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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
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, 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,
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, Dublin


Captive breeding

Since 1990, the Kerry slug has been successfully bred in captivity. The
Wildfowl & Wetlands Trust The Wildfowl & Wetlands Trust (WWT) is an international wildfowl and wetland conservation charity in the United Kingdom. Its patron is Charles III, and its president is Kate Humble. History The WWT was founded in 1946 by the ornithologist a ...
, a British conservation organisation, operates a captive breeding programme in
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at its "Endangered Species Breeding Unit". The project is located at the
Martin Mere Wetland Centre WWT Martin Mere is a wetland nature reserve and wildfowl collection managed by the Wildfowl and Wetlands Trust at Tarlscough, Burscough, Lancashire, England, on the West Lancashire Coastal Plain, from Ormskirk and from Southport (Merseyside) ...
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 Lancashi ...
, 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
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'
1843
77. * Allman, G. J. (1846). "Description of a new genus of pulmonary gastropods". ''
Annals and Magazine of Natural History The ''Journal of Natural History'' is a scientific journal published by Taylor & Francis focusing on entomology and zoology. The journal was established in 1841 under the name ''Annals and Magazine of Natural History'' (''Ann. Mag. Nat. Hist.'') a ...
'' 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
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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