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Mollusca, Pulmonata BULLETIN DE L'I NSTITUT ROYAL DES SCI ENCES NATURELLES DE BELGIQUE, BIOLOGIE, 60: 35-68. 1990 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUU RWETENSCHAPPEN, BIOLOGIE, 60: 35-68, 1990 On the infrageneric systematics of the genus Arion F:ERussAc, 1819 (Mollusca, Pulmonata) by Thierry BACKELJAU and Luc DE BRUYN Summa1·y 1819 in particular, have always been controversial. No one so far has been able to present a convenient and generally This paper deals with the infrageneric systematics of the genus Arion. It accepted generic and infrageneric classification or phylo­ presents a historical account, after which the infrageneric position of A. geny of these slugs. We propose and discuss here some intermedius is discussed. This species is included in the subgenus Kobe/tia, reflections and hypotheses on this subject. For the author­ of which the type species is reconsidered. Next, a discussion on the in frage­ neric allocation of A. /usitanicus and A. jlagellus is provided. Both species ship of the arionid genus- and speciesgroup names used are provisionally placed in the subgenus Mesarion, which in tum may be in the text, we refer to appendix 1. grouped with Arion s.s. in the "chromosomal group" Lochea (n:S27). Kobe/tia and Carinarion may then constitute the "chromosomal group" Prolepis (n2:27). This classification relies on chromosomal, electrophoretic, Historical account genital and morphological characters. Yet, the currently available data do not allow a reliable phylogenetic analysis, so that no taxonomic ranks are attributed to the retained groupings. Nevertheless, some tentative phyloge­ Before LIN NAEus (1758), slugs were usually referred to as netic and evolutive hypotheses are presented. Thereby special attention is "Limax" or "Cochleis nudis" (OKLAND, 1922). The former paid to I) chromosome numbers, 2) electrophoretic data, 3) breeding sys­ of these names was subsequently selected by LINNAEUS tems, 4) spermatophores and 5) the complexity of genitalia and mating ( 1758) to denote five terrestrial slugs, including Lim ax ater behaviour. Key-words : Arion, breeding systems, chromosomes, electrophoresis, geni­ and L. rufus. Later, LrNNAEUS (1761, 1767) extended his talia, spennatophores, systematics, type species. "genus" Limax to include some marine species too. FERUSSAC (1819) divided Limax into several "genera", one of which was referred to as Arion. It contained the species Resume with the following characters (original language) : "Cui­ rasse partielle ou nulle; tentacules retractiles. Tetraceres. Cette contribution est consacree a Ia systematique infragenerique du genre Cuirassees anterieurement. Cavite pulmonaire anterieure; Arion. Un a per~ u historique du sujet est presente, apres quoi Ia position infragem!rique d 'A. intermedius est discutee. Cette espece est classee dans orifice au bord droit de Ia cuirasse, anterieurement. Orifice le sous-genre Kobeltia, dont l'espece-type est commentee. La discussion du rectum, pres celui de Ia respiration. Un pore muqueux se poursuit par !'attribution infragenerique d'A. /usitanicus et d'A.jlagellus. terminal. Organes de Ia generation reunis : Orifice sous Provisoirement, on propose de placer les deux especes dans Ie sous-genre celui de Ia respiration. Point de levres ou tentacules buc­ Mesarion. Puis, on presente une tentative de classification du genre Arion caux : des papilles; des macho ires. Une couche de pous­ en retenant deux grandes unites chromosomiques (Lochea et Prolepis). Chacune d'elles est composees de deux groupes d'especes (Arion s.s. et siere calcaire et graveleuse, interieurement dans Ia cuiras­ Mesarion pour Lochea, Kobeltia et Carinarion pour Prolepis). Cette classi­ se." (FERussAC, 1821). This genus appeared to be so fication est fondee sur des caracteres chromosomiques, electrophoretiques, peculiar, that GRAY (in TuRTON, 1840) created a separate genitaux et morphologiques. Cependant, Ia connaissance actuelle de Ia family for it, viz. the Arionidae. systematique du genre Arion n 'etant pas suffisante pour permettre une Several authors attempted to split the genus Arion into analyse phylogenetique, cela ne permet pas de definir le niveau taxonomi­ que des groupements distingues. Neanmoins, on propose quelques hypothe­ smaller taxa. MoQUIN-TAN DON (1855) distinguished two ses phylogenetiques et evolutives concernant I) les nombres chromosomi­ "subgenera", namely Lochea for the species with a shell­ ques, 2) les donnees electrophoretiques, 3) les systemes reproducteur, 4) plate consisting of loose, discrete granulae, and Prolepis les spermatophores et 5) Ia complexite de l'appareil genital. for the species having a shellplate consisting of an irregu­ Mots-c1efs : Arion, systemes reproducteur, chromosomes, electrophorese, lar, but coherent aggregate of granulae. The fom1er group appareil genital, spermatophores, systematique, espece-type. contained "A. subfuscus" and "A. rufus" (i.e. A. lusitani­ cus), while the latter group contained "A. hortensis". General introduction MAB ILLE (1868) constituted the genus Baudonia to include B. timida and B. montana. These species would differ from The systematics of the terrestrial slugs of the family Ario­ Arion by 1) the anteriorly broadened and depressed body, nidae in general, and those of the genus Arion FERUSSAC, 2) the clearly delimited head, 3) the smooth mantle and 36 Thierry BAC KELJ AU and Luc DE BRUYN 4) the short tentacles. HEYNEMANN (1 870) also used Bau­ the genital atrium in arionids is too variable to be used donia, although he considered it, together with Locl·rea and for systematics above the species level. This could not be Prolepis, as a subgenus of Arion. Most authors, however, refuted and hence SI MROTH (1 887, 1891) himself, aban­ rejected Baudonia, because they argued that the characters doned his classification. mentioned by M AB ILLE ( 1868) were much too variable to PaLLO ERA ( 1887) distinguished four species-groups. define higher taxa (COLLINGE, 1897). These groups were classified in two larger units, which M AB ILLE (1870) described still another arionid genus, viz. were related to MoQU IN-TAN DON' s ( 1855) Lochea and Pro­ Carinel!a, for which A. bourguignati (= A. fascia/us s. l. lepis (fig. 1). However, in later publications, Po LLONERA see CocKERE LL, 189 1a, b ; CoLLI NGE, 1892a,b, 1893) acted (1 889, 1890), avoided to link his species-groups with these as type species. Carinella included the species w ith a "subgenera" (appendix 2). "dorsal keel", but this is usually an optical effect caused --:- 1 A'ni male grande , muco / by the mediodorsal tubercle row. The new genus was 1 • Gruppo .8 ] ]' \ incoloro o leggermente . therefore rejected by CoLLI NGE (1897). Later it was also 1 dell' A. n 1{us. shown that even if Carinella was accepted as a taxon, the 8 ~ ~ gi allo"nolo. ~ . :0 1 \t 0 name Carinella itself was not avail able because of the ~ ~ · ~ • 0 o ~ "i Amma1e med10cre, muco 1 2" Gruppo homonymy with Carinella Jo HNSTON, 1833 (Nematomor­ ol ~ giallo vivo. \ dell 'A. subfuscus. pha) and Carine/la So wERBY, 1839 (Mollusca; error pro Cancinelle AoANSON, 1757) (SH ERBORN, 1924). Animale piccolo , d01·so 1 3" Gruppo SEIBERT (1873) created the genus Kobeltia for a species rotondo. I dell'.A. hoTfcusis. he referred to as A. hortensis. Unfo rtunately, SEIBERT ( 1873) did not support his opinion with arguments. So, Animale piccolo , d01·so 1 4" Gruppo carenat o. subsequent authors refrained from using the name Kobel­ \ dell'A. B otwgui_qun ti. tia, except for mentioning it in synonymy lists (e.g. LES­ Fig. 1. - Infrageneric systematics of the genus Ari on as pro­ SONA, 188 1 ; LESSON A & POLLONERA, 1882; COCKERELL, posed by f OLLONERA (1887) . 1892; T AYLOR, 1907), until HESSE (1926) employed it to denote a "section" in the genus Arion. Nowadays, the name This system was adopted by CoLLINGE ( 1892a), who shor­ Kobeltia is commonl y used throughout malacological lite­ tly aft erwards, splitted PoLLONERA's A. hortensis group ratu re. imo two separate units : one including A. hortensis, the LESSON A (188 1) and LESSON A & POLLONERA (1882) ini­ other containing A. intermedius (COL LI NGE, 1897). Hence, ti all y adopted the two subgenera of MoQU IN-TAN DON CoLLINGE (1 897) proposed a classification involving five (1855) (Lochea and Prolepis). In this system, A. bourgui­ infrageneric species-groups (appendix 3). gnati was assigned to Prolepis, but later PoLLONERA ( 1885) This an angement was formali zed by HESSE ( 1926), who transfen ed A. bourguignati and A . subcarinatus to MAB IL­ distinguished three subgenera in Arion, viz. Letourneuxia, LE's (1870) Carinella, which received a rank comparable Ariunculus and Arion s.s. This latter comprised five "sec­ to that of Lochea and Prolepis. tions" corresponding to CoLLI NGE's (1897) species-groups. SJ MROTH (1 885a) refuted MoQU IN-T ANDON's ( 1855) arran­ HESSE (1 926) attached names to these "sections", selected gement by argueing that the degree of aggregati on of the type species for them and provided diagnoses (appendix 4). granulae under the mantle is an unreliable character. Alter­ TORRES MI NGUEZ (1 925) did not accept COL LI NGE'S (1897) natively, SI MROTH (1885a) divided the genus by using the ideas, but relied on PoLLONERA's work (1887, 1889, 1890) structure of the genital atrium and so he distingui shed the to define three infrageneric units viz. the Majorium group Monoatriidae and the Diatri idae. In the former group, the (= A. empiricorum group), the Mediocrum group (= A . genital atrium would consist of a single vestibule, whereas subfuscus group) and the Parvorum group (= A. hortensis in the latter it would be composed of two vestibules : an + A . bourguignati groups). This latter group was further upper (distal) one comprising the terminations of the geni ­ di vided in a subcylindricus secti on for the species with a tal ducts and a lower (proximal) one containing the genital rounded back (e.g. A. hortensis) and a pseudocarinatus porus. In this scheme, S IMROT H ( l885a) assigned A. mini­ section for the species with a "keeled" back (e.g.
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