The widespread slug Limax maximus: Limax maximus - variation species boundaries and redescription
Isabel Hyman1, Barbara Klee1, Ulrich Schneppat2, Gerhard Haszprunar1 1 Zoologische Staatssammlung München, Germany 2 Bündner Naturmuseum, Switzerland
What do we know so far about Limax maximus? Limax maximus • Mantle with dark spots • Formal description: Linnaeus, 1758 • Sole of foot unicolour • Relatively short penis • probably native to southwest Europe • Adult body length 10-20 cm • Occurs in the rest of Europe • Introduced into North and South America, Australasia, Hawaii, South Africa
Taxon sampling Why study Limax maximus? • Limax maximus from: • Type species of Limax – Many populations around Europe – Australia, New Zealand • Complex taxonomic history – North America, Hawaii • Extremely variable appearance – South Africa • Very widespread species – Wide variety of colour forms …over 200 specimens • Introduced to many countries • Many other Limax species • Confusion with other Limax species • Outgroups from Limacidae
1 Comparisons Comparisons Other species confused with L. maximus in literature, bioinventories and collections Spotted forms:
Dark forms: – Limax cinereoniger
– Limax sp. nov. MOTU #012 – Limax cinereoniger – Limax MOTU #013
– Limax sp. nov. MOTU #012 – Limax conemenosi
– Limax MOTU #011 – Limax punctulatus
Sole field coloration - a good Comparisons character? Striped forms:
– Limax cinereoniger
– Limax sp. nov. MOTU #012
Limax maximus Pale forms:
– Limax cinereoniger
– Limax sp. nov. MOTU #012 Limax cinereoniger
Comparisons Anatomy
Pale forms: • Important characters include – Penis length – Presence of blind penis tip – Attachment of penis retractor – Limax cinereoniger muscle – Attachment of vas deferens • Limax maximus: short penis, very short blind tip • Some externally similar species – Limax sp. nov. MOTU #012 can be distinguished from L. maximus based on penis length
2 Anatomy - long penis Anatomy - intermediate penis
L. sp. nov. L. conemenosi L. maximus L. punctulatus L. cinereoniger L. maximus MOTU #012 Drawing from A. Wiktor 2001
Anatomy - short penis DNA sampling
• 1334 bp of mitochondrial gene COI
• More than 30 specimens of L. maximus from “all over the world”
• Probably most species sampled that are confused with L. maximus
L. MOTU #011 L. MOTU #013 L. maximus
Results Results • Limax maximus clade well- supported •Blue= short penis • Overall variation 1.4% • Green = intermediate penis • no grouping of colour •Red= long penis variations within L. maximus L. maximus can be distinguished • Most closely related from all other species from Italy externally similar Bayesian reconstruction, GTR + Γ8 species
Bayesian reconstruction, GTR + Γ8
3 Conclusions Acknowledgements • Range of L. maximus now better known • Thank you to those who provided • Colour ranges from black to pale cream; strong samples and other assistance to weak pattern of spots and / or stripes • Thank you for listening! • Always (?): mantle with dark spots and sole uniformly pale Photographs: R. Heim, I. Hyman, A. Klee, B. Klee, R. • Colour variations independent of biogeography Melzer, M. Unsöld or habitat • Penis anatomy as good character in sexually mature/healthy specimens • Can be distinguished from all other species by analysis of COI sequences
Announcement
II. International Meeting of Task-Force-Limax in Val Resia / Udine Italy, Natural Park of Julian Pre-Alps 21. September – 23. September 2007
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