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The Spanish slug (''Arion vulgaris'', but also widely known as ''Arion lusitanicus'' owing to a misidentification) is an 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 ...
, a
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
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 ...
mollusk 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 e ...
in the family
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 slug ...
, the roundback slugs. Other vernacular names are Lusitanian slug, Iberian slug, and killer slug. It is a large, conspicuous slug, which has spread across much of Europe since the 1950s and now reached North America. It may attain high densities and be a serious horticultural and agricultural pest. The life cycle is annual, with adults appearing in summer and dying off before winter.


Confusion over nomenclature

The Spanish slug was identified as ''
Arion lusitanicus ''Arion lusitanicus'', also known by its common name Portuguese slug, is a species of air-breathing land slug, a terrestrial pulmonate gastropod mollusk in the family Arionidae. Distribution ''Arion lusitanicus'' is endemic to forests of the ...
'' when it was first reported as an invading species in France in 1956, and hence it is sometimes called the Lusitanian slug (e.g.). This was a case of misidentification. In slugs, it is often impossible to find external characters that distinguish closely related species using external features, as colouration can be quite variable, and the rather plastic anatomy makes diagnostic anatomical features difficult to establish. The current consensus is that the true ''Arion lusitanicus'' is a species of the western part of the Iberian Peninsula. Examination of slugs from the
Serra da Arrábida Serra (Latin for "saw") may refer to: People * Serra (footballer) (born 1961), Portuguese footballer * Serra (surname) * Serra (given name) Cities, towns, municipalities Brazil *Serra, Espírito Santo, a city in the Greater Vitória area *Ampar ...
mountains in
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 ...
from where it was originally described by
Jules François Mabille Jules François Mabille (Tours, 5 December 1831 − 18 January 1904) was a French malacologist, biologist and zoologist who in many trips around the world discovered and studied many species of mollusc. In 1882−83 Mabille participated in the Fr ...
in 1868 showed that the true ''A. lusitanicus'' differed from the invader in its internal anatomy, the shape of the
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 ...
and the number of
chromosomes A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
. The misidentification was first recognised in 1997, and more widely publicised in an atlas of British molluscs. ''Arion vulgaris'' was proposed as a substitute name based on a drawing of the genitalia in an 1855 work by
Alfred Moquin-Tandon Christian Horace Benedict Alfred Moquin-Tandon (7 May 1804 – 15 April 1863) was a French naturalist and doctor. Moquin-Tandon was professor of zoology at Marseille from 1829 until 1833, when he was appointed professor of botany and director of ...
. However, it is debatable whether the name applies to this drawing, so one temporary solution was to use the name ''Arion lusitanicus'' auct. non Mabille (i.e. "as used by authors other than Mabille"). Nevertheless, ''A. vulgaris'' has increasingly been used since, and this is the proposal that has been formally submitted to the
International Commission on Zoological Nomenclature The International Commission on Zoological Nomenclature (ICZN) is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 26 commissioners from 20 countries. Orga ...
. Still today, we are confronted with the invader being reported as having originally been introduced with vegetables from Spain. These reports are usually based on outdated information published in pre-1999 literature. The common name "Spanish slug" was further based on the unsubstantiated assumption that the species would not only live in Portugal, but also in Spain. ''Arion vulgaris'' seems to be rare in Spain. Another name sometimes applied is the "Iberian slug".


Distribution

The native distribution of ''Arion vulgaris'' is not exactly known. Genetic evidence (the higher incidence of rare
alleles An allele (, ; ; modern formation from Greek ἄλλος ''állos'', "other") is a variation of the same sequence of nucleotides at the same place on a long DNA molecule, as described in leading textbooks on genetics and evolution. ::"The chro ...
) suggests an origin in France or Spain, contrary to earlier genetic analyses that did not adequately sample these regions. In Britain the slug was first recorded in 1954,Kerney M. (1999). ''Atlas of the land and freshwater molluscs of Britain and Ireland''. pp. 1-264. Colchester. (Harley). which is not an indication of it being native there. It is presumed that the specimen illustrated in Moquin-Tandon's original 1855 description was from France.


Non-indigenous distribution

The non-indigenous distribution of ''Arion vulgaris'' includes almost the whole of Western and Central Europe and extends to various parts of Southern, Northern and Eastern Europe. It is now also recorded from Canada (2009) and Mexico. A 2017 report from the Asian part of Turkey was not based on dissection and is liable to be ''A. ater'' s.l. Reports of "''A. lusitanicus''" from Madeira may be copied from an 1895 report, in which case they need confirmation; similarly, a 1975 report from the Azores could be of the true ''A. lusitanicus''. Claims of its presence in Algeria also appear unsupported. Chronological overview of expansion of ''Arion vulgaris'' in Europe: *
Great Britain Great Britain is an island in the North Atlantic Ocean off the northwest coast of continental Europe. With an area of , it is the largest of the British Isles, the largest European island and the ninth-largest island in the world. It is ...
– since 1954, later spread to Lincolnshire and Norfolk in 2012 *
France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...
– (expansion from native range) since 1955 *
Switzerland ). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...
– since at least 1956 *
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– since 1965 *
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– since 1966 *
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– since 1969 *
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– since 1971 *
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– since 1973 *
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– since 1975 *
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– since 1982–83 *
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– since 1982–83 *
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 ...
– since 1984 *
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– since 1985 *
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– likely since 1987, certainly by 1993 *
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– since 1988 *
Netherlands ) , anthem = ( en, "William of Nassau") , image_map = , map_caption = , subdivision_type = Sovereign state , subdivision_name = Kingdom of the Netherlands , established_title = Before independence , established_date = Spanish Netherl ...
– since 1989 *
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 ...
– (possibly native; only known from north-east) since 1990 *
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– since 1990; Valovirta I. (2001)
''Tehokkaan leviämisen mestari''
. accessed 27 September 2009.
abundant in south *
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– since 1991 *
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– since 1992 *
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– since 1996 *
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– since 2002 *
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– since 2002 *
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– since 2003 * Macedonia – since at least 2003 *
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– since 2007, subsequently spread across almost whole country *
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– since 2008 *
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– since 2009 *
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– since 2009 (greenhouses), since 2019 (outdoors) (2018 record not confirmed anatomically) *
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– since 2012Păpureanu A.-M., Reise H. & Varga A. (2014). "First records of the invasive slug ''Arion lusitanicus'' auct. non Mabille (Gastropoda: Pulmonata: Arionidae) in Romania". ''
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'' 13: 6-11
PDF
/ref> *
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– since 2020 (2018 record not confirmed anatomically) This species has not yet become established in the USA, but it is considered to represent a potentially serious threat as a pest, an
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
which could negatively affect agriculture, natural ecosystems, human health or commerce. Therefore, it has been suggested that this species be given top national
quarantine A quarantine is a restriction on the movement of people, animals and goods which is intended to prevent the spread of disease or pests. It is often used in connection to disease and illness, preventing the movement of those who may have been ...
significance in the USA. British authorities were also concerned as of 2014 that it may become a major pest.


Description

The size of the adult slug is 60–140 mm. The colour ranges from yellow to black, but is most commonly brick-red, dirty orange, or brown. The
tentacles 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 ...
are darker. Adult ''A. vulgaris'' may not differ in external appearance from ''
Arion rufus The red slug (''Arion rufus''), also known as the large red slug, However, juveniles of other large ''Arion'' species such as '' Arion flagellus'' also have longitudinal bands.
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 ...
: The atrium is small. The adjacent part of the oviduct is dilated and muscular, with the same diameter as the atrium and containing a longitudinal ligula. This distinguishes ''A. vulgaris'' from ''Arion ater'' s.l., in which the oviduct is thinner and the atrium is larger and contains the ligula. ''Arion flagellus'' also has a ligula in the dilated part of the oviduct, but the ligula does not reach as far towards the atrium as in ''A. vulgaris''. The spermatheca is spherical, its diameter twice that of the oviduct.


Ecology

The habitat of ''Arion vulgaris'' includes all agricultural ecosystems, as well as natural environments such as river and lake margins, forest edges, forests in valleys or meadows. In Switzerland it has been found up to 1700 m altitude. It is a serious agricultural and horticultural pest in large parts of Europe, eating a cosmopolitan range of growing plant parts as well as decaying vegetation. Opportunistically it eats carrion, including squashed conspecifics. It is active mostly during the night and in wet weather during the day. Densities can reach 50 individuals per m2 or locally even higher. Whilst a slug can crawl several metres within a night, long-distance dispersal is believed to be on vegetables, on horticultural seedlings, and on plant debris disposed of as waste. The species has an annual life cycle with mating starting in July and eggs first laid some weeks later in late summer. Clutches are laid on the soil surface or in crevices up to 10 cm underground, with an average clutch size of about 70 eggs; an adult slug typically lays about 400 eggs in its lifetime. The eggs hatch from autumn to spring. Neither eggs nor slugs can survive temperatures below ―3 °C, so overwinter survival depends on hiding under shelters. Adults normally die off in autumn before winter frosts.


As an invasive species

''Arion vulgaris'' is considered among the 100 worst
alien species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived there ...
in Europe in DAISIE European Invasive Alien Species Gateway,DAISIE European Invasive Alien Species Gateway
''Arion vulgaris''
Last updated 30 October 2006, accessed 27 September 2009.
and this is the only land gastropod among them. ''Arion vulgaris'' is the worst slug pest in Europe and it has an important economic, ecological, and social impact. The local name of the slug in the regions it has invaded is typically a translation of "Spanish slug". In recent years, as its dominance has increased, it has been nicknamed "killer slug", perhaps due to its tendency to eat dead or weaker individuals of the species, although its destructive impact on gardens may seem just as appropriate a reason for the name. Like other pulmonate snails and slugs, it 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 ...
and this species has the capacity to self-fertilize, so that one single slug can start an infestation. Long-distance transport of produce and garden plants has been assumed to be a common means of its rapid dispersal. Besides causing economic damage, the arrival of ''A. vulgaris'' has often been associated with the disappearance within a few years of the similarly sized congener ''Arion ater'' s.l., at least in
synanthropic A synanthrope (from the Greek σύν ''syn'', "together with" + ἄνθρωπος ''anthropos'', "man") is a member of a species of wild animal or plant that lives near, and benefits from, an association with human beings and the somewhat artific ...
habitats. While the two species coexist they may mate with each other and produce fertile offspring. Hybrids are often identifiable by their intermediate genital anatomy. It has been proposed that in Scandinavia crossing of ''A. vulgaris'' and the indigenous ''A. ater ater'' might have produced a more frost-resistant variety. However, genetic investigations have not shown that
introgression Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Intr ...
of ''A. ater'' genes into ''A. vulgaris'' persist for long once the native species has disappeared. In the Swiss Alps, '' Arion ater rufus'' persists only at high elevations, and hybrids with ''A. vulgaris'' occur in a contact zone along the altitudinal gradient. Given the densities that ''A. vulgaris'' can attain, other ecological effects of its invasion on the native flora and fauna are to be expected. For instance, the seeds that it consumes are less likely to survive to germinate than with other gastropods with which it was compared. In Sweden complete defoliation of native shrubs in natural woodland has been reported.


References


External links


''Arion vulgaris''
at
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''Arion vulgaris''
images at
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* Slug controls (on Wikibooks) {{DEFAULTSORT:Spanish Slug Arion (gastropod) Gastropods described in 1855 Articles containing video clips Fauna of Spain