Deroceras invadens
Two individuals of Deroceras invadens from different sites on Sicily (identity checked by observing matings)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Subclass: Heterobranchia
Order: Stylommatophora
Family: Agriolimacidae
Genus: Deroceras
Species:
D. invadens
Binomial name
Deroceras invadens
Reise, Hutchinson, Schunack & Schlitt, 2011[2]

Deroceras invadens is a species of air-breathing land slug, a terrestrial pulmonate gastropod mollusc in the family Agriolimacidae. Until 2011, this widely distributed species was known as Deroceras panormitanum, and earlier as Deroceras caruanae or Agriolimax caruanae, but Reise et al. (2011)[2] showed that these names refer to a distinct species of similar external appearance known at that time only from Sicily and Malta. Consequently, although the more widespread species was already well known, it then had to be redescribed under the new name of D. invadens. Genetic evidence has indicated that D. invadens is native in southern Italy, including parts of Sicily, and possibly parts of central Italy. Elsewhere it has been introduced, predominantly within the last 100 years, but its spread has been constrained by cold winter temperatures.

Description

Adults are usually 20–35 mm long.[2] The skin and flesh are watery and fairly transparent.[3] The colour of the skin varies between light greyish-brown to almost black. Close inspection reveals fine dark spotting usually over the whole body; this shows up better in alcohol-preserved specimens.[2] Often, but not always, the respiratory pore is pale and unspotted.[3] The mucus is colourless.

D. invadens preserved in ethanol. Scale = 3 mm

Particularly in North America, some Deroceras laeve grow large enough that they closely resemble D. invadens.[3] A useful clue to distinguish them is the profile of the end of the tail when the animal has been disturbed or is preserved.[3][4] The tail of D. invadens usually slants vertically upward from the sole for a short distance, or even bends backwards. The tail of D. laeve slopes forward above the sole. Also, the tail of D. invadens is longer than the mantle, whereas it is the same length or shorter in D. laeve.[5]

Penis, twisted to reveal the ventral side

Nevertheless, dissection is required to distinguish D. invadens reliably from D. laeve and from various similar Deroceras species occurring in Europe, such as Deroceras sturanyi, the true Deroceras panormitanum, and Deroceras cecconii. Reise et al. (2011)[2] discuss and figure the most critical anatomical characters. In most populations of D. invadens, the proximal penis has two side pockets (the penial caecum and penial lobe); these have roughly equal widths and both have rounded, stout ends. A prominent appending penial gland with 3–7 branches attaches between the caecum and lobe on the dorsal side; these branches are less knobbly in outline than those of D. panormitanum. The penial retractor muscle attaches between the caecum and lobe on the ventral side. The intestinal caecum is either absent or represented merely by a widening of the rectum. Barker (1999)[6] and Sirgel (1973)[7] detail various other aspects of the anatomy.

Distribution

The distribution has been extensively reviewed by Hutchinson et al. (2014).[8] The native range is the southern and perhaps central parts of Italy[9] but the first certain record is from Britain in 1930.[10] Deroceras invadens now occurs in very many other parts of the world, although there is only one record from Asia.[11] The list below gives dates of the first reported findings (outdoors unless stated); oceanic Islands are considered separately at the end.

Distribution of Deroceras invadens. Based on Figure 4 of Hutchinson et al. 2014,[8] with subsequent published records from Israel and Montenegro added.[11][12] The circled i's indicate countries from which exported goods were found to contain this species. Otherwise each symbol indicates presence on a 1-degree grid of latitude and longitude

Europe

  • Italy – native to the south, including parts of Sicily[9]
  • Great Britain – 1930[10]
  • Denmark – 1937[13]
  • France – 1945[14]
  • Sweden – c. 1957 (greenhouses),[15] ≤1980 (outdoors)[16]
  • Ireland – 1959[17]
  • Finland – ≤1961 (greenhouses),[18] ≤2014 (outdoors)[19]
  • Norway – 1967 (greenhouses),[20] 1983 (outdoors)[8]
  • Belgium – 1968[21]
  • Netherlands – 1969[22]
  • Spain – 1974[23] (earlier record from Canaries)
  • Portugal – 1977[24] (earlier record from Azores)
  • Austria – 1977[25]
  • Germany – 1978[26]
  • Switzerland – 1982[27]
  • Czech Republic – 1996[28]
  • Luxembourg – 1997[8]
  • Andorra – ≤2000[29]
  • Poland – 2001[30]
  • Slovakia – 2003 (greenhouse),[31] 2018 (outdoors)[32]
  • Russia – 2009 (greenhouses, Tver Oblast)[33]
  • Greece – 2011[8][34]
  • Monaco – 2012[8]
  • San Marino – 2013[8]
  • Montenegro – 2014[12]
  • Liechtenstein – 2014 [9]
  • Hungary – 2019; a 1974 report was retracted by its author[35]

Earlier records of "D. panormitanum sensu lato" from south-eastern Europe (e.g. Bulgaria, parts of Greece) should be rechecked, given the taxonomic confusion prior to 2011; records from Romania and Lithuania are erroneous or unconfirmed.[8]

Africa

  • South Africa – 1963[36]
  • Egypt – 2005–7[37]
  • Kenya – 2012 (found on exports to USA)[8]

Asia and Australasia

  • Israel – 2013 (greenhouse)[11]
  • Australia – 1936[8]
  • New Zealand – 1974 (circumstantial evidence for much earlier date; slightly earlier record from Raoul Island)[6]

North America:

  • USA: California (1940),[38] Washington State and Oregon (2001),[8] Colorado (2004),[3] Utah (2006),[2] Washington D.C. (1998)[2]
  • Canada: Quebec (greenhouses 1966),[39] British Columbia (1974),[40] Alberta (2021),[41] Newfoundland (2012).[42] Records from Ontario require confirmation.[8]
  • Mexico – 1974[8]

Central and South America

  • Costa Rica – 2006[8]
  • Panamá – 2007 (found on exports to USA)[8]
  • Colombia – 1975[8]
  • Ecuador – 2012 (found on exports to USA in 2004)[8]
  • Peru – 2012 (found on exports to USA)[8]
  • Chile – ≤2003[43] (earlier record from Juan Fernández Islands)
  • Argentina – 2004[44]
  • Brazil – 1991[6]

Oceanic Islands

  • Faroe Islands (Denmark) – 1970[45]
  • Madeira (Portugal) – 1980[46]
  • Azores (Portugal) – 1957[15]
  • Canary Islands (Spain) – 1947[47]
  • Tristan da Cunha (UK) – 1982[48]
  • Raoul Island (New Zealand) – 1973[6]
  • Chatham Islands (New Zealand) – 1976[49]
  • Lord Howe Island (Australia) – 2000[50]
  • Norfolk Island (Australia) – 2013[51]
  • Marion Islands (South Africa) – 1972[52][53]
  • Juan Fernández Islands (Chile) – 1962[8]

Ecology

Deroceras invadens typically occurs in disturbed sites (e.g. the most widespread slug species in Manchester gardens[54]) and is often easiest to find under rubbish.[55] However, this species has also spread to natural habitats such as woodland and grassland (e.g. in Britain, Tenerife, South Africa and Australia).[8] It prefers areas of high humidity and cannot survive temperatures below –7 °C.[53][56] Its distribution appears restricted by low winter temperatures, which could explain its slow and only partial colonisation of central Europe and its recent range expansion in Sweden following climate amelioration.[8][57] It can be a significant pest in gardens, greenhouses, pasture, and arable fields.[8][56][58] In captivity, slugs eat their own body weight of lettuce in two or three days.[59]

In North Wales, most adults start to lay eggs in autumn and have died by early spring; the species can be found in any season, but it is most abundant in late spring.[60] In New Zealand pastureland, populations fall considerably in summer.[56] In agricultural fields in northern Italy, D. invadens was commonly trapped from November to May, but not at all in summer.[58]

Behaviour

The mating behaviour of this species has been particularly well studied, partly to provide taxonomic characters.[2][6][61][62] Mating starts when one individual closely follows the flattened tail of the other (precourtship). Typically after some minutes the leader turns back and protrudes its sarcobelum (a tapering finger-like part of the penis), as does its partner shortly afterwards. The partners often form a circle head to tail. Early courtship involves violent biting of the partner, lashing the sarcobelum against the partner, and bursts of tail wagging. This is gradually replaced by more gentle-looking stroking of the sarcobelum across the partner. During courtship, sperm accumulates in the penial caecum.[62] After about 90 minutes the partners orient themselves face-to-face, nibble the base of the partner's raised sarcobelum, and suddenly their penises simultaneously evert. The finger-like penial caecum curves round the back of the partner's sarcobelum, transferring sperm onto it. The penises remain everted for a variable period (typically some minutes) before the penial lobe and then the penial gland also evert. The latter transfers a secretion onto the partner.[62] This is directly followed by retraction of the penis (each partner thereby taking up the transferred sperm) and then separation.

Laboratory experiments suggest that D. invadens shows a diurnal rhythm of active locomotion and feeding at night, then inactivity in or below the litter layer during the day; the rhythm is entrained by light. This feeding cycle is accompanied by regular cytological changes in the organs of digestion.[59]

Deroceras invadens exhibited the highest crawling speed (4.9 millimetres (0.19 in)/s) amongst measurements from 28 species of terrestrial slug and snail.[63]

Deroceras invadens usually avoids areas treated with its parasite Phasmarhabditis hermaphrodita, a nematode which is used commercially to control slugs. However, the avoidance is replaced by attraction once the slug is infected, apparently an example of host manipulation. Also, another species of parasitic nematode, Phasmarhabditis neopapillosa, is attractive even to uninfected D. invadens.[64]

References

  1. ↑ Kappes, H. (2017). "Deroceras invadens". IUCN Red List of Threatened Species. 2017: e.T83478691A83478783. doi:10.2305/IUCN.UK.2017-3.RLTS.T83478691A83478783.en. Retrieved 12 November 2021.
  2. 1 2 3 4 5 6 7 8 9 Reise, H.; Hutchinson, J.M.C.; Schunack, S.; Schlitt, B. (2011). "Deroceras panormitanum and congeners from Malta and Sicily, with a redescription of the widespread pest slug as Deroceras invadens n. sp". Folia Malacologica. 19 (4): 201–233. doi:10.2478/v10125-011-0028-1. hdl:11858/00-001M-0000-002C-1000-1.
  3. 1 2 3 4 5 Reise, H.; Hutchinson, J.M.C.; Robinson, D.G. (2006). "Two introduced pest slugs: Tandonia budapestensis new to the Americas and Deroceras panormitatum new to the Eastern USA". The Veliger. 48 (2): 110–115.
  4. ↑ de Winter, A.G. (1988). "Remarks on the non-marine molluscan fauna of the Azores. 1-2". Basteria. 52: 105–109.
  5. ↑ Rowson, B.; Turner, J.; Anderson, R.; Symondson, W. (2014). Slugs of Britain and Ireland: identification, understanding and control. Telford: Field Studies Council. ISBN 978-1-908819-13-0.
  6. 1 2 3 4 5 Barker, G.M. (1999). Naturalised terrestrial Stylommatophora (Mollusca: Gastropoda). Fauna of New Zealand No. 38 (PDF). Manaaki Whenua Press, Canterbury, New Zealand. ISBN 978-0-478-09322-3.
  7. ↑ Sirgel, W. (1973). "Contributions to the morphology and histology of the genital system of the pulmonate Agriolimax caruanae". Annale van die Uniwersiteit van Stellenbosch. Reeks A. 48: 1–43.
  8. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Hutchinson, J.M.C.; Reise, H.; Robinson, D.G. (2014). "A biography of an invasive terrestrial slug: the spread, distribution and habitat of Deroceras invadens". NeoBiota. 23: 17–64. doi:10.3897/neobiota.23.7745.
  9. 1 2 3 Hutchinson, J.M.C.; Schlitt, B.; Kořínková, T.; Reise, H.; Barker, G.M. (2020). "Genetic evidence illuminates the origin and global spread of the slug Deroceras invadens". Journal of Molluscan Studies. 86 (4): 306–322. doi:10.1093/mollus/eyaa016.
  10. 1 2 Quick, H.E. (1960). "British slugs (Pulmonata: Testacellidae, Arionidae, Limacidae)". Bulletin of the British Museum (Natural History), Zoology. 6: 103–226. doi:10.5962/bhl.part.26845.
  11. 1 2 3 Mienis, H.K.; Mienis, D.; Vaisman, S. & Rittner, O. (2014). "Two exotic gastropods: Aegopinella nitidula and Deroceras invadens, recently discovered in Israel". Triton. 29: 21–25.
  12. 1 2 Soes, M. (2014). "The first record of Deroceras invadens (Gastropoda: Pulmonata: Agriolimacidae) for Montenegro". Ecologica Montenegrina. 1 (4): 189–190. doi:10.37828/em.2014.1.25.
  13. ↑ Lohmander, H. (1959). "Faunistiskt fältarbete i västra och norra Jylland 1954–1957. Landmolluskerna". Göteborgs Musei Årstryck. 1959: 33–104.
  14. ↑ Reygrobellet, D. (1963). "Une nouvelle espèce de limacidé, Deroceras meridionale n. sp". Bulletin de la Société zoologique de France. 88: 399–402.
  15. 1 2 Waldén, H.W. (1960). "Om ett par för Sverige nya anthropochora landmollusker, Limax valentianus Férussac och Deroceras caruanae (Pollonera) jämte några andra, kulturbunda arter". Göteborgs Kungliga Vetenskaps- och Vitterhets-Samhälles Handlingar, Sjätte Följden. Series B. 6 (8): 5–48.
  16. ↑ von Proschwitz, T. (2002). "Faunistikt nytt 2001—snäckor, sniglar och musslor". Göteborgs Naturhistoriska Museum Årstryck. 2002: 29–46.
  17. ↑ Makings, P. (1959). "Agriolimax caruanae Pollonera new to Ireland". Journal of Conchology. 24: 354–356.
  18. ↑ Brander, T.; Kantee, J. (1961). "Lounais-Hämeen nilviäiset, Mollusca". Lounais-Hämeen Luonto. 11: 70–72.
  19. ↑ Koivunen, A.; Malinen, P.; Ormio, H.; Terhivuo, J.; Valovirta, I. (2014). Suomen kotilot ja etanat: opas maanilviäisten maailmaan. Helsinki: Hyönteistarvike. ISBN 978-952-67544-6-8.
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  21. ↑ Van Goethem, J.L. (1974). "Sur la presence en Belgique de Deroceras caruanae (Pollonera, 1891) et de Deroceras agreste (Linnaeus, 1758) (Mollusca, Pulmonata, Limacidae)" (PDF). Bulletin de l'Institut Royal des Sciences Naturelles de Belgique. 50 (2): 1–21.
  22. ↑ Gittenberger, E.; Backhuys, W.; Ripken, Th.E.J. (1970). De landslakken van Nederland. Amsterdam: Koninklijke Nederlandse Natuurhistorische Vereniging.
  23. ↑ Castillejo, J. (1983). "Los pulmonados desnudos de Galicia, III. Estudio del género Deroceras Rafinesque, 1820 (Agriolimacidae, Gastropoda, Pulmonata)". Iberus. 3: 1–13.
  24. ↑ Seixas, M.M.P. (1978). "Descrição de uma espécie de Limacidae (Gasteropoda, Pulmonata), nova para a fauna portuguesa". Boletim da Sociedade Portuguesa de Ciências Naturais. 2nd Series. 18: 5–6.
  25. ↑ Reischütz, P.L. (1977). "Die Malakofauna des Waldviertels aus zoogeographischer Sicht". Jahres-Bericht des Bundesgymnasiums Horn. 99: 4–9.
  26. ↑ Falkner, G. (1979). "Ein Freilandvorkommen von Deroceras (D.) panormitanum (Lessona & Pollonera) [= D. caruanae (Pollonera)] in Deutschland". Mitteilungen der Zoologischen Gesellschaft Braunau. 3 (8/9): 239–242.
  27. ↑ Falkner, G. (1979). "Deroceras (D.) panormitanum (= D. caruanae) in der Schweiz". Mitteilungen der Zoologischen Gesellschaft Braunau. 4: 134–135.
  28. ↑ Horsák, M.; Dvořák, L. (2003). "First records of the introduced slug Deroceras panormitanum (Lessona et Pollonera, 1882) from the Czech Republic (Mollusca: Gastropoda: Agriolimacidae)". Folia Malacologica. 11 (1–2): 57–58. doi:10.12657/folmal.011.006.
  29. ↑ Bertran, A. (2000). "Les mollusques d'Andorre: atlas préliminaire de repartition". Documents Malacologiques. 1: 17–39.
  30. ↑ Wiktor, A. (2001). "Deroceras (Deroceras) panormitatum (Lessona et Pollonera) – a new introduced slug species in Poland (Gastropoda, Pulmonata, Agriolimacidae)". Folia Malacologica. 9: 155–157. doi:10.12657/folmal.009.018.
  31. ↑ Dvořák, L.; Čejka, T.; Horsák, M. (2003). "First record of Deroceras panormitanum (Gastropoda, Agriolimacidae) from Slovakia". Biologia. 58: 917–918.
  32. ↑ Čejka, T. (2018-02-22). "First outdoor records of the slug Deroceras invadens from the Slovak Republic". Malakologický bulletin. [scientist's blog]. Retrieved 22 September 2018.
  33. ↑ Schikov, E.V. (2016). "Adventive species of terrestrial malacofauna in the central portion of the Russian plain [In Russian]". Ruthenica. 26 (3–4): 153–164.
  34. ↑ Rowson, B.; Anderson, R.; Turner, J.A.; Symondson, W.O.C. (2014). "The slugs of Britain and Ireland: undetected and undescribed species increase a well-studied, economically important fauna by more than 20%". PLOS ONE. 9 (4): e91907. Bibcode:2014PLoSO...991907R. doi:10.1371/journal.pone.0091907. PMC 3989179. PMID 24740519.
  35. ↑ Turóci, Á.; Hutchinson, J.M.C.; Schlitt, B.; Reise, H.; Rapala, M.; Páll-Gergely, B. (2023). "Five new introduced terrestrial slugs in Hungary". BioInvasions Records. 12 (3): 711–729. doi:10.3391/bir.2023.12.3.08. S2CID 261543248.
  36. ↑ van Regteren Altena, C.O. (1966). "Notes on land slugs 11. Arionidae, Milacidae and Limacidae from South Africa (Mollusca, Gastropoda, Pulmonata)". Zoologische Mededelingen. 41: 269–298.
  37. ↑ Obuid-Allah, A.H.; Abdel-Tawab, H.S.; El-Bakary, Z.; Abd El-Wakeli, K.F.; El-Sanabany, A. (2008). "A survey and population dynamics of terrestrial slugs (Mollusca, Gastropoda) at Assuit Governate, Egypt". Egyptian Journal of Zoology. 51: 585–608.
  38. ↑ Pilsbry, H.A. (1948). Land Mollusca of North America (North of Mexico). Vol. II, Part 2. Acad. Nat. Sci. Philadelphia.
  39. ↑ Chichester, L.F.; Getz, L.L. (1969). "The zoogeography and ecology of arionid and limacid slugs introduced into north-eastern North America". Malacologia. 7: 313–346.
  40. ↑ Rollo C.D.; Wellington W.G. (1975). "Terrestrial slugs in the vicinity of Vancouver, British Columbia". Nautilus. 89: 107–115.
  41. ↑ Patuwatha Withanage, D.B.M.; Howe, D.K.; Richart, C.H.; Mc Donnell, R.J.; Denver, D.R.; Luong, L.T. (2023). "Pestiferous slugs and their associated nematodes in agricultural fields, greenhouses, and nurseries in Alberta, Canada". Journal of Helminthology. 97 (e41): e41. doi:10.1017/S0022149X23000226. PMID 37194264. S2CID 258717942.
  42. ↑ Forsyth, R. (2014). "First record of Deroceras invadens Reise, Hutchinson, Schunack & Schlitt, 2011 (Gastropoda: Pulmonata: Agriolimacidae) from the island of Newfoundland, Canada". Check List. 10 (1): 149–150. doi:10.15560/10.1.149.
  43. ↑ Letelier, S.; Vega, M.A.; Ramos, A.M.; Carreño, E. (1969). "Base de datos del Museo Nacional de Historia Natural: moluscos de Chile". Revista de Biología Tropical. 51 (Supplement 3): 33–137.
  44. ↑ Gutiérrez Gregoric, D.E.; Beltramino, A.A.; Vogler, R.E.; Cuezzo, M.G.; Núñez, V.; Gomes, S.R.; Virgillito, M.; Miquel, S.E. (2013). "First records of four exotic slugs in Argentina" (PDF). American Malacological Bulletin. 31 (2): 245–256. doi:10.4003/006.031.0204. S2CID 62892096.
  45. ↑ McMillan, N.F. (1972). "Agriolimax caruanae Pollonera and other non-marine Mollusca in Faroe". Journal of Conchology. 27: 419–421.
  46. ↑ Rähle, W. (1992). "Nacktschnecken (Arionidae, Milacidae, Agriolimacidae und Limacidae) von Madeira und Porto Santo (Mittelatlantische Inseln) (Gastropoda: Pulmonata)". Malakologische Abhandlungen aus dem Staatlichen Museum für Tierkunde Dresden. 16: 13–24.
  47. ↑ van Regteren Altena, C.O. (1950). "The Limacidae of the Canary Islands". Zoologische Verhandelingen. 11: 1–34.
  48. ↑ Preece, R.C. (2001). "Introduced land molluscs on the islands of the Tristan da Cunha-Gough Group (South Atlantic)". Journal of Conchology. 37: 253–259.
  49. ↑ Barker, G.M. (1979). "The introduced slugs of New Zealand (Gastropoda: Pulmonata)". New Zealand Journal of Zoology. 6 (3): 411–437. doi:10.1080/03014223.1979.10428382.
  50. ↑ Hyman, I.; Köhler, F. (2020). A field guide to the land snails of Lord Howe Island. Sydney: Australian Museum. ISBN 9780975047682.
  51. ↑ Colgan, D.J. (2017). "The invasive slug Deroceras invadens Reise, Hutchinson, Schunack and Schlitt, 2011 occurs on Norfolk Island". BioInvasions Records. 6: 9–12. doi:10.3391/bir.2017.6.1.02.
  52. ↑ Smith, V.R. (1992). "Terrestrial slug recorded from sub-Antarctic Marion Island". Journal of Molluscan Studies. 58: 80–81. doi:10.1093/mollus/58.1.80.
  53. 1 2 Lee, J.E.; Janion, C.; Marais, E.; Jansen van Vuuren, B.; Chown, S.L. (2009). "Physiological tolerances account for range limits and abundance structure in an invasive slug". Proceedings of the Royal Society B. 276 (1661): 1459–1468. doi:10.1098/rspb.2008.1240. PMC 2677229. PMID 19324817.
  54. ↑ North, M.C.; Bailey, S.E.R. (1989). "Distribution of Boettgerilla pallens in North-west England". In Henderson, I.F. (ed.). BCPC Monograph No. 41, Slugs and snails in world agriculture. pp. 327–329.
  55. ↑ Kerney, M. (1999). Atlas of the land and freshwater molluscs of Britain and Ireland. Harley Books: Great Horkesley, Colchester. ISBN 978-0-946589-48-7.
  56. 1 2 3 Barker, G.M. (2002). "Chapter 18: Gastropods as pests in New Zealand pastoral agriculture, with emphasis on Agriolimacidae, Arionidae and Milacidae". In Barker, G.M. (ed.). Molluscs as crop pests. CABI. pp. 361–423. ISBN 978-0-85199-320-1.
  57. ↑ von Proschwitz, T. (2010). "Faunistical news from the Göteborg Natural History Museum 2009—snails, slugs and mussels—with some notes on Pupilla pratensis (Clessin)—a land snail species new to Sweden". Göteborgs Naturhistoriska Museum Årstryck. 2010: 41–62.
  58. 1 2 Scaccini, D.; Panini, M.; Chiesa, O.; Aldini, R.N.; Tabaglio, V.; Mazzoni, E. (18 June 2020). "Slug monitoring and impacts on the ground beetle community in the frame of sustainable pest control in conventional and conservation agroecosystems". Insects. 11 (6): 380. doi:10.3390/insects11060380. PMC 7349495. PMID 32570919.
  59. 1 2 Morton, B. (1979). "The diurnal rhythm and the cycle of feeding and digestion in the slug Deroceras caruanae". Journal of Zoology. 187 (2): 135–152. doi:10.1111/j.1469-7998.1979.tb03938.x.
  60. ↑ Dirzo, R. (1980). "Experimental studies on slug-plant interactions. I. The acceptability of thirty plant species to the slug Agriolimax caruanae". Journal of Ecology. 68 (3): 981–998. doi:10.2307/2259470. JSTOR 2259470.
  61. ↑ Reise, H. (2006). "A review of mating behavior in slugs of the genus Deroceras (Pulmonata: Agriolimacidae)". American Malacological Bulletin. 23: 137–156. doi:10.4003/0740-2783-23.1.137. S2CID 29257202.
  62. 1 2 3 Benke, M.; Reise, H.; Montagne-Wajer, K.; Koene, J. (2010). "Cutaneous application of an accessory-gland secretion after sperm exchange in a terrestrial slug (Mollusca:Pulmonata)". Zoology. 113 (2): 118–124. doi:10.1016/j.zool.2009.08.003. PMID 20202803.
  63. ↑ Cameron, R. (2014). "Speedy snails (and speedier slugs)". Mollusc World. 36: 3–4.
  64. ↑ Rae, R. (2023). "Avoidance and attraction behaviour of slugs exposed to parasitic nematodes". Journal of Invertebrate Pathology. 197: 107896. doi:10.1016/j.jip.2023.107896. PMID 36758665. S2CID 256679186.
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