And the Phylogenetic Relationships of the Subfamily Plutoniinae (Gastropoda: Limacoidea: Vitrinidae)*

MALACOLOGIA, 2000, 42(1-2): 39-62

PLUTONIA {CANARIVITRINA), NEW SUBGENUS, FROM THE CANARY ISLANDS, AND THE PHYLOGENETIC RELATIONSHIPS OF THE SUBFAMILY PLUTONIINAE (GASTROPODA: LIMACOIDEA: VITRINIDAE)*

1 1 2 1 **

Maria R. Alonso , Manuel J. Valido , Klaus Groh & Miguel Ibáñez

ABSTRACT

Plutonia (Canarivitrina), n. subgen., and four new species of this subgenus are described from three islands of the mid-Atlantic Canarian archipelago. The genus Plutonia Morelet, in Stabile,

1864 (of which Insulivithna Hesse, 1923, is a new synonym), restricted to the mid-Atlantic islands, is characterized by the autapomorphy of the special course of the penial retractor muscle, rounding the right optic nerve. Canarivitrina, n. subgen., has a penis with two easily distinguishable portions: a distal one, short and slightly widened, containing a globular to cylindrical and perforated penial papilla level with the opening of the vas deferens (far from the penial apex), and a proximal one, long and slender, with apical insertion of the penial retractor muscle and two parallel longitudinal inner structures: a torus (thick fold, with roundish cross section) and, opposite, a laminar crest (velum). Torus and penial papilla are coated by glandular tissue, the penial gland. The autapomorphic character state of this subgenus is the proximal portion of the penis, long and equipped with internal torus and velum. New data are given about the sheath that surrounds the spout of the glándula amatoria in Ca- narivitrina, n. subgen., and in other supraspecific Plutoniinae taxa demonstrating the homology of the glándula amatoria and its spout sheath with the sarcobelum of the genus Semilimax, a basal branch of the Vitrinidae: the subfamily Plutoniinae has the "sarcobelum" fused with the proximal part of the vagina to form the glándula amatoria.

A phylogenetic scenario of the supraspecific taxa of the subfamily Plutoniinae is presented. The cladistic analysis shows the Plutoniinae as the sister group of Semilimax, and Phenacolimax

as the sister group of the remaining Plutoniinae: the genera Phenacolimax, Gallandia (which is almost certainly a junior synonym of Oligolimax), Arabivitrina and Plutonia; but this analysis does

not resolve for the other taxa because the data are still too few to apply to a cladistic study of the group.

The genus Plutonia includes the subgenera Plutonia, s. str. , Guerrina, Insulivitrina, Madeirovit- rina and Canarivitrina, n. subgen. The presence of this genus in the archipelagoes of the Azores, the Madeiras and the Canary Islands is probably as old as the humid "laurisilva" laurel-forest, a Tertiary relict that colonized these archipelagoes before the impact of the Pleistocene glaciations. Key words: Gastropoda, Vitrinidae, Plutoniinae, Canarivitrina, phylogeny, biogeography.

INTRODUCTION were described: G. christinae Groh, 1993; /.

eceroensis Alonso & Ibáñez, 1987; /. tubercu-

The Canarian Vitrinidae were studied only lata Ibáñez & Alonso, 1987; /. gomerensis superficially between 1821-1954; eight spe- Alonso & Ibáñez, 1988; /. emmersoni Morales, cies were described -one doubtful: Vitrina 1988; /. oromii Ibáñez & Alonso, 1988; /. fasciolataA. Férussac, 1832; one of Guerrina machadoi Ibáñez & Alonso, 1990; /. nogalesi

Odhner, 1954: Helix cutícula Shuttleworth, Alonso & Ibáñez, 1990; /. tamaranensis

1852; and six grouped in Insulivitrina: Helico- Valido, 1990; and /. mascaensis Morales, limax lamarckii A. Férussac, 1821; V blauneri 1 987 (Alonso et al., 1 987; Ibáñez et al., 1 987;

Shuttleworth, 1852; V latebasis Mousson, Morales et al., 1 988; Valido et al., 1 990, 1 993). 1872; V canariensis Mousson, 1872; V retic- The last species, from Tenerife Island, has a ulata Mousson, 1872; and V parryi Gude, peculiar penis, with a very long and slender 1896. In the last years, ten additional species proximal portion having a remarkable distance

Department of Animal Biology, University of La Laguna, E-38206 Tenerife, Canary Islands, Spain 2 Mainzer Strasse 25, D-55546 Hackenheim, Germany "Notes on the Malacofauna of the Canary Islands, 38 "To whom correspondence should be addressed; [email protected] 39 40 ALONSO ETAL. between the vas deferens entrance to the Abbreviations penis and the insertion of the penial retractor. We have recently found four additional new a species with the same penial characteristics: three from La Gomera and one from La Palma

(Fig. 5); these five species are assigned to a new subgenus, Canarivitrina, n. subgen. In this paper, we describe these taxa and at- tempt a phylogenetic study of the entire subfamily Plutoniinae.

METHODS

The introduction of Canarivitrina, n. subgen., is the consequence of our anatomical studies of the European Phenacolimax major (A. Férussac, 1807), Arabivitrina jansseni Neubert, 1988 (from the Asir province, Saudi Arabia), Madeirovitrina nitida (Gould, 1848) and M. marcida (Gould, 1848) from the Madeira Archipelago, and the Canarian species above mentioned. Moreover, biblio- graphic data on the Plutoniinae have been ob- tained from Forcart (1 959), Hubendick (1 953), Mermod (1930), Zilch (1979), and several additional papers (Table 1). Calculation of number of shell whorls follows Kerney et al. (1 979: 1 3). The terms "shell" and "specimen" in the enumerations of the material studied refer to empty shells and live specimens respectively, and "proximal" and "distal" refer to the position in relation to the gonad. The term "semislugs" refers to the classification of Tillier (1984), that is, the animal usually cannot withdraw completely into the shell, the posterior edge of the foot cavity is lower and further forward than the most posterior part of the digestive tract, and the stomach is retained in the upper visceral cavity. The cladistic analysis involves the eight genus-group taxa of the Plutoniinae and was performed using PAUP 4.0 Beta 2 for Windows

(Swofford, 1 999) and the Treeview application (Page, 1996). Tree searches were performed using the exhaustive search command. Zero- legth branches were collapsed, MulTrees was activated and the accelerated transformation option (Ace Tran) was used. The characters and character states used are listed in Table 2 and the data matrix in Table 3 mainly based on the type species of the single supraspecific taxa. Character polarity was determined by outgroup comparison using Semilimax as outgroup for the reasons indicated in the comments after the family diagnosis. All characters have equal weight. Two out of nine characters CANARIVITRINA AND THE PLUTONIINAE PHYLOGENY 41

TABLE 1. Genus-group taxa, type species and bibliographical sources of anatomical data.

Genus-group taxa Type species Sources

Arabivitrina Thiele, 1931 Vitrina arabica Thiele, 1910 Thiele, 1931; Forcart, 1957; Zilch, 1959; Neubert, 1998 Canarivitrina, n. subgen. P. (Canarivitrina) taburientensisn. this paper sp.

Oligolimax Fischer, in Paulucci, Vitrina paulucciae Fischer, in Manganelli et al., 1995 1878 Paulucci, 1878 Gallandia Bourguignat, 1880 Vitrina conoidea, sensu Schileyko, 1986; Hausdorf, 1995, (= Trochovitrina O. Boettger, Bourguignat, 1880 [non and personal communication 1Í Martens, 1874] = Gallandia o/ymp/ca Hausdorf, 1995

Guerrina Odhner. 1954 Helix cutícula Shuttleworth, 1852 Odhner, 1954; Ibáñez et al., 1987; Valido et al., 1993; personal unpublished data Insulivitrina Hesse, 1923 Helicolimax lamarckii A. Hesse, 1923; Odhner, 1937;

Férussac, 1821 Forcart, 1957; Ibáñez et al., 1987; Schileyko, 1986; personal unpublished data Madeirovithna Groh & Hemmen, Vitrina nitida Gould, 1848 Groh & Hemmen, 1986 1986 Phenacolimax Stabile, 1859 Helicolimax major A. Férussac, Forçait, 1944, 1949, 1957; 1807 Hausdorf, personal communication; personal unpublished data Plutonia Morelet, in Stabile, 1864 Viquesnelia atlántica Morelet, Wiktor & Backeljau, 1995; Mordan, 1860 personal communication Semilimax Agassiz, 1845 Helix semilimax J. Férussac, Eckhardt, 1914; Forcart, 1944; 1802 Kerney et al., 1979; Grossu, 1983; Schileyko, 1986; Hausdorf, personal communication

(1,7) had three character states that were ing whorls. Protoconch generally spirally dot- treated as nonadditive; character 1 was ted with numerous pits, which sometimes are treated as unordered, and 7 as ordered for the also present on the teleoconch. Animal with reasons indicated in the explanation of the penial gland and without penial nerve passing cladistic analysis; four characters (3, 5, 8, 9) through the cerebral ganglion (see Hausdorf, were autapomorphies with an additional au- 1998). tapomorphy in each multi-state character.

Comments RESULTS Schileyko (1986) divided the Vitrinidae into TAXONOMIC DESCRIPTIONS three subfamilies -Vitrininae Fitzinger, 1833; Phenacolimacinae Schileyko, 1986; and Family Vitrinidae Fitzinger, 1833 Semilimacinae Schileyko, 1986. Vitrina Dra-

Type genus: Vitrina parnaud, 1 801 , and Calidivitrina Pilsbry, 1 91 9, Draparnaud, 1801 were included in the subfamily Vitrininae (sarcobelum and glándula amatoria absent; Diagnosis vagina very reduced). Semilimacinae, characterized by the short vagina and by the pres- Limacoidea, sensu stricto (semislugs and ence of a distinct, atrial sarcobelum, included slugs with vas deferens running along the in- Semilimax Agassiz, 1845, Oligolimax Fischer, side of the penial sheath; without sper- in Paulucci, 1878, Vitrinobrachium Kunkel, matophore, epiphallus and flagellum), with 1929, and dubitatively Eucobresia Baker, shell very thin, glossy, pale and translucent, 1 929. This last genus lacks a sarcobelum and usually with fewer than three rapidly expand- has a vaginal papilla near the atrium possibly 42 ALONSO ETAL.

TABLE 2. Characters used on the phylogenetic Odhner, 1954; Arabivitrina Thiele, 1931; In- analysis. The numeration refers to the explanations sulivitrina Hesse, 1923; Phenacolimax Sta- in the text. bile, 1859; Trochovitrina O. Boettger, 1880; and Plutonia Morelet, in Stabile, 1864. (1) Shell Hausdorf (1998) regarded this system as External, the animal cannot withdraw into it = unacceptable, the Vitrininae being poly- External, the animal can withdraw into it = 1 Internal, reduced = 2 phyletic, the Semilimacinae paraphyletic, and

(2) Teleoconch ornamentation only the Phenacolimacinae monophyletic. Glossy = Moreover, the name Phenacolimacinae is a Entirely and very densely ribbed = 1 junior synonym of Plutoniinae Cockerell (3) Radular type (1893), established for Plutonia. Dichoglossan = According to Schileyko (1986), the Vitrini- Beloglossan = 1 nae and the Phenacolimacinae evolved inde- Location of the glándula amatoria — stimulator (4) pendently from an unknown common ances- organ tor, while the Semilimacinae derived from the Distinct from vagina (Sarcobelum) = Phenacolimacinae by the transformation of Fused with the proximal part of vagina = 1

(5) Proximal portion of penis long, equipped with the glándula amatoria ("vaginal gland") into a torus and velum sarcobelum. We assume the contrasting the- Absent = ory, sketched by Simroth (1889) and sup-

= 1 Present ported by Hausdorf (1997, 1998), is the most (6) Penial sheath probable: the Phenacolimacinae derived from Strong = a Semilimax-Uke ancestor by the fusion of the Rudimentary or absent = 1 sarcobelum with the proximal vagina so as to (7) Arrangement of the penial retractor muscle give the glándula amatoria. Free of the right optic nerve = Passing rounding the right optic nerve (Fig. 3G) The sarcobelum of the type species of

= 1 Semilimax (Fig. 1) consists of a voluminous Penial retractor absent = 2 oblong oval or club-shaped stimulator not (8) Vaginal retractor muscle covered by soft tissue, with a thick glandular Absent = layer wrapping a longitudinal duct, which Present = 1 opens distally into the atrium through a spout, Oviducal retractor muscle (9) the duct being upholstered (internally with re- Absent = spect to the glandular layer) by a thin layer of Present = 1 circular muscles and a horny tube, the latter

missing in all the other Vitrinidae. The wall of the distal portion of the sarcobelum invagi- homologous to the spout of the sarcobelum nates sheathing the spout; at the invagination

(Schileyko 1 986: figs. 11,12). Phenacolimaci- edge, the sarcobelum wall is connected with nae, characterized mainly by the long vagina the base of the spout by a very thin layer of and the presence of a glándula amatoria connective tissue. on the proximal vagina, includes Guerrina As Hausdorf (1998) has shown in his hypo-

TABLE 3. Original data matrix used for cladistic analysis. All characters with three character states are '7' treated as nonadditive; the character T is treated as unordered and the character as ordered. The keys and numeration refer to the explanations in the text. The variable or unapplicable character states were coded as a '?'.

Key CANARIVITRINA AND THE PLUTONIINAE PHYLOGENY 43

FIG. 1. Genital system of Semilimax showing the sarcobelum without the inner horny tube; redrawn from Schileyko (1986: figs. 6-7) without scale, as the original drawings.

thetical explanation of stimulator evolution in also possible: the sarcobelum could be origi- the Stylommatophora, the Semilimax the ancestral loss of portions and genus nated by A2 may be seen as a basal branch of the Vit- A , the shortening of the A portion (the 3 4 rinidae, the atrial sarcobelum possibly being spout), and the invagination of A into the A 4 1 homologous to the penial appendix of the Or- portion (Giusti, pers. comm.). We assume the thurethra, with portions A and A absent and last theory because it explains favourably the 3 4 and fused. But another explanation is of the spout sheath. A2 A5 presence 44 ALONSO ET AL.

Subfamily Plutoniinae Taxa Belonging to Plutoniinae Cockerell, 1893 Schileyko (1986: 147) referred Guerrina, (junior synonym: Phenacolimacinae Schi- Arabivitrina, Insulivitrina, Phenacolimax, Tro- leyko, 1986) chovitrina, and PlutoniaXo his Phenacolimaci- Type genus: Plutonia Morelet, in Stabile, 1864 nae. The type species of Trochovitrina, T. lederi (O. Boettger, 1878) and Phenacolimax annularis (Studer, 1820), studied by Schileyko Diagnosis (1986) as examples of the respective genera, were later grouped with Gallandia olympica Vitrinidae with a thin shell variable in shape, Hausdorf, 1995, in the genus Gallandia Bour- normally with a narrow basal periostracal guignat, 1880, by Hausdorf (1995). Hausdorf fringe. Distinct atrial stimulator (sarcobelum) recognized this genus to be characterized by absent. Vagina long, its proximal portion the following autapomorphies: shell conoid- housing a muscular glándula amatoria with a globose, shell and mantle lobes reduced, free distal spout (see Schileyko, 1986). oviduct short, pedunculus [of the bursa copu- The autapomorphic character state of this latrix] short and penis short. subfamily is the fusion of the sarcobelum with Ibáñez et al. (1987) grouped the taxa in- the proximal vagina to give the glándula ama- cluded by Forcart (1957) in Phenacolimax, s. toria (Hausdorf, 1995). I. (Phenacolimax, s. str., Arabivitrina, Oligoli- max, Insulivitrina, Plutonia, and Guerrina) and Madeirovitrina in the Phenacolimacinae, Comments mainly based on the same characters as Schi- leyko (1986). Plutoniinae established The name was by The type species of Oligolimax, Vitrina Cockerell for Viquesnelia atlántica (1893) paulucciae Fischer, in Paulucci, 1878, was Morelet, 1860, an Azorean carnivorous slug considered by Forcart (1965) and Giusti feeding on such animals as earthworms and (1971) to be a synonym of V. bonelli Targioni slugs, with subterranean and an almost way Tozzetti, 1873, which belongs to the genus of life, sheltered in moist places, in moss or Semilimacella Sobs, 1917, anatomically not detritus; to its under stones and due very spe- far from Semilimax. Recently, Manganelli et cialized features for a predatory life style, Wik- al. (1995) indicated that V. paulucciae is not a tor & Backeljau proposed to remove it (1995) synonym of Semilimacella bonelli and that its from the family Vitrinidae and assign it to a genital system anatomy is so similar to that of separate family. the species grouped in Insulivitrina and Gal- Cockerell the possibility (1893) mentioned landia, that these last two names could be that Plutonia had also been used for a genus seen as junior synonyms of Oligolimax. of trilobites, and Collinge (in Cockerell, 1893) Gallandia is almost certainly a junior syn- of the proposed an eventual change name onym of Oligolimax, but the complete descrip- Plutoniinae Vitriplutoniinae. to However, as tion of O. paulucciae has not yet been pub- Backhuys (1975) indicated, the name Pluto- lished (Giusti et al., in prep.). We therefore nia for a genus of the Trilobita published was continue to utilize the name Gallandia. On the later: Plutonia Hicks, 1871, Invalid Generic other hand, Insulivitrina is clearly distinguish- Name by homonymy (ICZN, 1997, Opinion able from Gallandia (and Oligolimax) because 1880). of the peculiar course of its penial retractor. Shelley & Backeljau (1995) recognized a case of between Plutoniinae Boll- homonymy Genus Plutonia Morelet, in Stabile, 1864 man, 1893 (Arthropoda, Chilopoda, type- genus: Plutonium Cavanna, 1881) and Plu- Type species by monotypy: Viquesnelia at- toniinae Cockerell, 1893, and proposed the lántica Morelet, 1860 emendation to Plutoniainae Cockerell, 1893. (Junior synonym: Insulivitrina Hesse, 1923, However, Backeljau & Shelley (1996) modi- n. syn.) fied their original application and proposed to amend the myriapod name to Plutoniuminae Diagnosis and to retain the molluscan name unchanged as Plutoniinae; this proposition has been ac- Plutoniinae with the autapomorphic charac- cepted by the ICZN (1997, Opinion 1880). ter state "penial retractor muscle (starting CANARIVITRINA AND THE PLUTONIINAE PHYLOGENY 45 from the penis) moving forward, forming loop one of the Azorean "Insulivitrina" due to re- around the optic nerve, passing below the duction of the penial retractor (see Com- right ommatophore retractor, then running ments). backwards (left of the same right om- Nervous system as described by Tillier matophore retractor) to end on left side of di- (1989: 64) for the family. Palliai complex as aphragm" (Fig. 3G). described by Wiktor & Backeljau (1995) in Plutonia atlántica; lung small, kidney sig- murethrous, bean-shaped, secondary ureter Description closed like a tube, renal opening near anus and pneumostome. Slugs (Plutonia atlántica), semislugs (the Genital atrium, penis and distal vagina be- "Insulivitrina group") or Vithna-Wke helicoid come completely evaginated when protruded snails (the "Guerrina group"). Mantle border (in some preserved specimens and possibly with well-developed mantle and shell lobes also at mating; Fig. 2J). Genital atrium vari- (the "Insulivitrina group"), with lobes small to able in length, connected with adjacent part of very small (the "Guerrina group"), or with shell animal integument by short atrial muscles. lobes absent {Plutonia atlántica, with a vesti- Penial complex without epiphallus and flagel- gial shell entirely hidden in the mantle cavity). lum. Penial sheath thin, inconspicuous, rudi- Right side of mantle generally with distinct mentary or absent. darker coloured "lateral band" above pneu- Bursa copulatrix duct and free oviduct in- mostome. Foot aulacopod. Sole tripartite, with sert side by side at proximal end of glándula distinct sole furrows, the lateral fused at short amatoria. Glandula amatoria variable in di- distance from tail end. Foot dorsally flattened mensions, wider than distal vagina in which it beneath the shell, bordered by a pair of dis- opens through a spout. It can be partially to tinct lateral crests extending for about half the completely coated by a layer of soft tissue that tail length; tail tip dorsally keeled. can easily be stripped off and it is formed by Shell thin, greenish, translucent, glossy, thick outer muscular layer and inner glandular with 2-4 whorls. Shell variable in shape, from layer wrapping the longitudinal duct. slightly depressed, keeled, conical above and Distal vagina variably long, its wall invagi- with small aperture, to auricular with lower nated to envelop the spout of the glándula side very little developed, very large aperture amatoria, to its tip where it fuses with the and with narrow periostracal fringe. Proto- same spout tip. Invaginated vaginal wall and conch pitted, sometimes with radial ribs. spout wall entirely connected by thin connec- Jaw oxygnathous, smooth, with median rib tive fibres easily removable without destroy- and central denticle. Radula mainly di- ing the vaginal wall (allowing the invagination choglossan (Jungbluth et al., 1985); central to be eliminated in the laboratory); at the in- tooth little smaller than first lateral teeth, with vagination edge, the vaginal wall is also con- long, slender mesocone having sharp pointed nected with the base of the spout by a very tip and two basal ectocones reaching half the thin layer of connective tissue (as found in the mesocone length; lateral teeth with long, wide sarcobelum of Semilimax; Fig. 1). mesocone, one basal ectocone and one small Spermoviduct long, about one-quarter of laterodistal endocone; marginal teeth variable total genital system length. Prostate poorly in shape. distinguishable. Albumen gland relatively System of retractor muscles basically as small and oval in shape. Hermaphrodite duct described by Odhner (1937) for Insulivitrina: long. Gonad follicular, located at apex of vis- right ommatophore retractor free of penis and ceral sac. vagina. Penial retractor, long and slender; soon after it leaves the penis wall, it moves forward passing below some nerves arising Comments from the right pedal ganglion, then above the nerve and the retractor of the lower right ten- The autapomorphic character state of the tacle; soon after, it forms loop around the optic diagnosis was firstly evidenced in Plutonia nerve and, passing below the right om- (Insulivitrina) lamarcki, two other Plutonia (In- matophore retractor, moves backwards (left of sulivitrina) species and Plutonia (Guerrina) the same right ommatophore retractor) to end cutícula by Hoffmann (1929) and Odhner on left of diaphragm. This situation is secun- (1937, 1954). It is shared by all the Canahan daria lost in Plutonia atlántica and in at least and Madeiran Plutoniinae and was also evi- 46 ALONSO ETAL. denced in the Azorean Plutoniinae (those erens into the same penis (far from the penial which retain a penial retractor; Mordan, pers. apex). Penial papilla perforated, its central comm.). It is interesting to note that one channel continuous with vas deferens. Penial Azorean species (different from Plutonia at- papilla and torus coated by glandular tissue, lántica) has been recently discovered to have the "penial gland". Distal portion of penis with a very vestigial or perhaps missing retractor two very thin penial folds, which join level with (Mordan & Frias Martins, in prep.), which the penial papilla, a small pit remaining be- hence has been interpreted as a possible tween them and the papilla itself. Thin vaginal synapomorphy of this species and P. at- fold sometimes present, ending before reach- lántica. Thus, all the Macaronesian Plutoni- ing genital atrium. inae would belong to the same genus, Pluto- nia Morelet, in Stabile, 1864, of which Type Material Insulivitrina Hesse, 1923, will become a subgenus. The relations of P. atlántica with the Holotype (Fig. 2A): collected by K. Groh other Azorean Plutoniinae taxa is supported and M. Ibáñez (8-1-1988) in the Caldera de by the cladistic analysis given below. Taburiente (La Palma Island; UTM: The adult animal (with the exception of the 28RBS1879), at 750 m altitude; deposited in Guerrina species), cannot withdraw com- AIT. Paratypes: 144 specimens and 22 shells pletely into the shell. This may happen only collected between January 1988 and Febru- (or) after a long period of desiccation and ary 1 997, in various localities of the island and without feeding, a situation that has been ver- deposited in ANSP (A 18852/2), CGH (32 ified in some probably senile adult specimens. paratypes, 29 specimens and 3 shells), CRT Forcart attributed glandular func- (1957) a (8 paratypes), CGS (1 paratype), SMF tion to the soft tissue coating the glándula am- (311875/5), TFMC (MT 0314/2), ZMH atoria. Other authors believe that despite it (2745/2) and AIT. glandular appearance, it has a different function, as yet undetermined (Schileyko, 1986). Type Locality The glandular function is feasible, but un- proven since this tissue wraps the thick mus- Caldera de Taburiente, La Palma Island. cular layer of the glándula amatoria externally, and there is no evident communication with Etymology the central lumen of the glándula amatoria. The Plutonia species are endemic to three The specific name refers to the Caldera de mid-Atlantic archipelagoes located at the Taburiente, a geological feature of La Palma. western fringe of the Palaearctic: Azores, Madeira and Canary Islands. The alpine Vit- Habitat and Distribution rina glacialis Forbes, 1837, once assigned to Insulivitrina (Forcart, 1944: 654), has been re- A species endemic to La Palma, occurring cently presumed as belonging to Eucobresia in wide areas climatologically different, with Baker, 1929 (H. Nordsieck, in Falkner, 1991: pinewood and sometimes with lowland vege- 103); data on the course of its penial retractor tation, at an altitude between 300 and 1,800 are nevertheless still pending. m (Fig. 5). Plutonia taburientensis Groh & Valido, n. sp. Animal

Diagnosis Body whitish-beige, with numerous small pale-beige spots, more dense in number at

Plutonia with penis as long as vagina (in- head. Mantle and tail with darker grey spots cluding its glándula amatoria portion), divided and a short blackish-grey lateral band. Foot into two portions: proximal penis long, slen- sole sometimes with marginal grey spots. der, with penial retractor muscle inserted at apex and two internal parallel longitudinal Shell structures, a torus and, opposite, a wide velum. Distal penis short, slightly widened, Oblong, with about 2.5 whorls. Protoconch containing a globular to cylindrical penial pitted; starting from beginning of second papilla level with the entrance of the vas def- whorl, the pits are less deep and more scat- FIG. 2. A-D. Holotype shells. A. P. taburientensis, n. sp. . P. ripkeni, n. sp. P. dianae, n. sp. D. P. falcifera, n. sp. E. Protoconch of P. taburientensis, n. sp. F-G. Protoconch and pits detail of P. ripkeni, n. sp. H-l. Radula of P. ripkeni. n. sp. H, central tooth and lateral teeth; I, marginal teeth. J. A specimen of P. mascaensis with the distal part of the genital system everted, showing the penis with the penial papilla and the vagina with the glándula amatoria (by transparency). Scale: A-D, J, 5 mm; E, F, 200 ; G, 100 \ivn; H, I, 20 . 48 ALONSO ETAL.

FIGS. 3, 4. Genital system and anatomical details. A. P. taburientensis. n. sp., paratype; Caldera de Taburi- ente. B. P. ripkeni, n. sp., paratype; Jerduñe. . P. dianae, n. sp.. paratype; Montaña Bejira. D. P. falcifera. n. sp., paratype; Tamolde. E. P. mascaensis (redrawn from Ibáñez et al., 1987, fig. 14). F. Internal penial details of P. taburientensis, n. sp. G. Arrangement of the Insulivitrina penial retractor muscle. Scale: A-E, 1 mm; F-G, without scale. CANARIVITRINA AND THE PLUTONIINAE PHYLOGENY 49

FIG. 4. See figure 3 for legend. 50 ALONSO ETAL. tered, disappearing after first half of second Gomera Island; UTM: 28RBS85 10), at 950 m whorl (Fig. 2A, E). altitude; deposited in AIT. Paratypes: 71 spec-

imens collected between December 1 985 and Radula March 1997 in various localities of the island and deposited in ANSP (A 18853/2 and Formula: 31 -33M + 11-12L + C; marginal 400849/2), CGH (2 paratypes), CRD (2 teeth with only one cusp. paratypes), CRT (6 paratypes, 3 specimens + 3 shells), NNM (55868/1), SMF (311874/5 Genital System and 311876/2), TFMC (MT 0313/2), ZMH (2746/2) and AIT. Shown in Figures , F, 4A; 12 specimens dissected. Penis as long as vagina (including Type Locality its glándula amatoria portion). Laminar crest inside the proximal portion of penis wide and Tagamiche, La Gomera Island. same length as torus, the latter distally smaller and thinner, as a laminar fold, joins Etymology the penial papilla; this fold partially surrounds the penial papilla and extends into the distal The specific name derives from the family portion of penis; opposite the papilla, another name of Mr. Theodor "Theo" E. J. Ripken, small laminar fold present inside distal penis; Delft, The Netherlands, to whom the species the two folds join level with the penial papilla, is dedicated. a small pit remaining between them and the papilla itself; finally, the folds extend sepa- Habitat and Distribution rately into the atrium. Distal vagina twice as long as the glándula A species endemic to La Gomera, occurring amatoria portion, nearly one-third shorter than in wide areas with lowland vegetation, border- the free oviduct and 2-2.5 times longer than ing the evergreen "lauhsilva" forest of the Na- the bursa copulatrix duct; sometimes with one tional Park of Garajonay, at an altitude be- thin internal longitudinal fold ending before tween 200 and 1 ,025 m (Fig. 5). the atrium. Glandula amatoria with a protruding spout. Animal

Comments Body whitish-beige-grey, with numerous small darker grey spots, more dense on man- Of all the described Plutonia species, P. tle and tail areas. Lateral band blackish-grey. taburientensis, n. sp., shares only with P. mascaensis the presence of a proximal penis Shell with the penial retractor muscle inserted at apex and internal torus and velum; the main Oblong, with 2.25 whorls. Pits in spiral rows differences between them are listed in the Di- on protoconch and teleoconch; from begin- agnosis and Comments on the latter. The ning of second whorl, the pits of some rows or proximal penis is eversible (Fig. 2J) and it is of adjacent rows frequently fuse forming an ir- not a flagellum. There is no spermatophore. regular drawing. Pits disappear after first half of second whorl (Fig. 2B, F, G). Plutonia ripkeni Alonso & Ibáñez, n. sp. Radula

Diagnosis Formula: 29M + 12L + C; first marginal teeth with very small ectocone and last mar-

As for P. taburientensis, n. sp., but the vagi- ginal teeth with only one cusp (Fig. 2H, I). nal fold, always present, extends to fuse with a penial fold into the atrium. Genital System

Type Material Shown in Figures 3B and 4B; 1 5 specimens

dissected. Penis similar or little longer than Holotype (Fig. 2B): collected by K. Groh vagina (including its glándula amatoria por- and M. Ibáñez (3-1 -1 988) from Tagamiche (La tion). Laminar crest inside proximal penis CANARIVITRINA AND THE PLUTONIINAE PHYLOGENY 51

wide and same length as torus, the latter dis- Valido de Armas, to whom this species is ded- tally thinner, as a fold, joins the penial papilla; icated. this fold partially surrounds the penial papilla and extends into the distal portion of penis; Habitat and Distribution opposite the papilla, another fold is present inside distal penis; the two folds join level with A species endemic to northwest La the penial papilla, a small pit remaining be- Gomera, occurring in a small area with low- tween them and the papilla itself; finally, the land vegetation, at an altitude between 350 folds extend separately into the atrium, the and 530 m (Fig. 5). fold deriving from torus fuses with the vaginal longitudinal fold. Animal Distal vagina nearly three times longer than the glándula amatoria portion, as long as free Body whitish-beige, with numerous small

oviduct and 1 .5-2 times as long as bursa cop- pale-beige spots, more dense at head. Mantle ulatrix duct; with one internal longitudinal fold and tail with darker grey spots. Lateral band which fuses with the penial fold inside the short, blackish-grey, interrupted level with atrium. Glandula amatoria with small spout. pneumostome. Foot sole sometimes paler than body. Comments Shell

Other than the characters listed in the diagnosis, P. ripkeni, n. sp., differs from P. taburi- Very oblong, with slightly more than 2.75 entensis, n. sp., in vaginal folds more devel- whorls. Pits in spiral rows on protoconch and oped; distal portion of penis, penis and vagina teleoconch, disappearing at beginning of sec- longer; glándula amatoria smaller; and spout ond whorl (Fig. 2C). less protruding. Genital System Plutonia dianae Shown in Figures 3C and 4C; five speci- Valido & Alonso, n. sp. mens dissected. Penis approximately half as long as vagina (including its glándula amato- Diagnosis ria portion), flattened, tongue-shaped. Proxi-

mal penis finger-like, with rounded tip. Lami- As for P. tabuhentensis, n. sp., but the penis nar crest inside proximal penis wide and is approximately half as long as the vagina same length as torus; distal torus slightly (including its glándula amatoria portion). smaller, partially surrounding the penial papilla and extending thicker into distal por- Type Material tion of penis; opposite the papilla, another similar fold present inside distal penis; the two Holotype (Fig. 2C): collected by R. Hutterer folds join level with penial papilla, a deep pit and M. Ibáñez (3-11-1994) from Montaña Be- remaining between them and papilla. jira (La Gomera Island; UTM: 28RBS7320), at Distal vagina very long, up to four times 530 m altitude; deposited in AIT. Paratypes: longer than the glándula amatoria portion, lit- 1 specimens and 8 shells collected the same tle longer than free oviduct and 1.5-2 times day in the type locality and the nearby Argua- longer than bursa copulatrix duct, with internal mul and deposited in ANSP (A 1 8854/1 ), CGH longitudinal fold that ends before reaching (1 paratype), TFMC (MT 0312/1) ZMH atrium. Glandula amatoria very short, with (2747/1, shell) and AIT. small spout.

Type Locality Comments

Montaña Bejira, La Gomera Island. Other than the characters listed in the diagnosis, P. dianae, n. sp., differs from P. taburi- Etymology entensis, n. sp., and P. ripkeni, n. sp., in the distal penis with deeper pit between penial The specific name derives from that of the papilla and the two thicker penial folds. Pluto- young daughter of the junior author, Diana nia dianae, n. sp., differs from P. taburienten- 52 ALONSO ETAL. sis, n. sp., also in the spout less protruding blackish-grey. Foot sole sometimes with mar- and from P. ripkeni, n. sp., in the vaginal fold, ginal grey spots. which ends before genital atrium.

Shell

Plutonia falcifera Oblong, with 2.75 whorls. Pits in spiral rows Ibáñez & Groh, n. sp. on protoconch and up to the end of second whorl where they become smaller to disap- Diagnosis pear at the beginning of third whorl (Fig. 2D).

As for P. taburientensis, n. sp., but the laminar crest of the proximal portion of penis is Genital System narrow or even vestigial; distal portion of penis without pit near penial papilla; the pit is Shown in Figure 4D; eight specimens dis- substituted by a pilaster connected proximally sected. Penis as long as vagina (including its with the papilla and ending distally in a small glándula amatoria portion), with proximal por- slope with transversal furrows, reminiscent of tion curved, entire penis being sickle shaped a fingerprint. when extended after dissection; proximal penis with an empty zone at its beginning,

which is narrower than the rest. Laminar crest Type Material narrow, vestigial in some specimens. Torus big, starting after the empty zone at the be- Holotype (Fig. 2D): collected by R. Hutterer ginning of proximal penis. Distal torus thin, and M. Ibáñez (31-1-1994) from Tamolde (La partially surrounding the penial papilla and ex- Gomera Island; UTM: 28RBS9112), at 200 m tending into the distal penis as a thick pilaster, altitude; deposited in AIT. Paratypes: 23 spec- which ends in a small slope with transversal imens collected between January 1988 and furrows, reminiscent of a fingerprint. March 1997 in various localities from the is- Distal vagina with thin internal longitudinal land and deposited in ANSP (A 18855/1), fold, nearly twice as long as glándula amato-

CGH (7 paratypes), SMF (311979/1) TFMC ria portion and little longer than both free (MT 0311/2), ZMH (2748/1) and AIT. oviduct and bursa copulatrix duct. Glandula amatoria with spout well protruding.

Type Locality Comments

Tamolde, La Gomera Island. Other than the characters listed in the diag-

nosis, P. falcifera, n. sp., differs from all the Etymology other species herein described for: penis sickle shaped when extended after dissec- The specific name derives from the Latin tion; at its beginning the proximal penis has falcifer ("sickle carrier"), due to the shape of an empty zone. The vagina has a thin internal penis. longitudinal fold.

Plutonia mascaensis Habitat and Distribution (Morales, 1987)

A species endemic to east La Gomera, oc- Insulivitrina mascaensis Morales, 1987, in curring in a small area with lowland vegetation Ibáñez et al.: 133-135, 139, figs. 7, 14, and pinewood, at an altitude between 200 and 26-28, 36, 47, 48, 51 [loe. typ.: Barranco 600 m (Fig. 5). de Masca, Tenerife].

Animal Diagnosis

Body whitish-beige, with numerous small As for P. falcifera, n. sp., but the proximal beige spots, more dense at head. Mantle and penis ends pointed and lacking an empty tail with darker grey spots. Lateral band short, zone. CANARIVITRINA AND THE PLUTONNNAE PHYLOGENY 53

28R UTM zones CANARY ISLANDS UTM grid: 100 km Lanzarote^J^ 18°00': meridian

Fuerteventura

Gran Canaria

Africa

FIG. 5. Geographical distribution of the Canarivitrina. n. subgen, species; the symbols represent 1 x 1 km squares. 54 ALONSO ETAL.

Type Material Comments

Holotype (AIT) and 114 paratypes, de- Other than the characters listed in the diag- posited in ANSP (A1 8856/2), CGH (2 nosis, P. mascaensis differs from the other paratypes), FMNH (205917/1), MHNG species described herein because the penis, (984647), MNHN, NHM (1986130/2), NNM coiled in natural position, is fringed on one

(55862/1), SMF (305945/2), TFMC (MT side; from P. falcifera, n. sp., it also differs in 0310/2), ZMH (2749/2) and AIT. not having a sickle-shaped penis, its penis in- stead having a shorter distal portion and a Habitat and Distribution longer proximal portion.

A species endemic to west Tenerife, occurring in areas with lowland vegetation or DISCUSSION pinewood, at an altitude between 90 and 1,100 m (Fig. 5). The five species are cohesive and probably represent a distinct supraspecific taxon, ST, Animal due to the following synapomorphies: genital system with long penis divided into two por- Body whitish-grey, with numerous small tions; proximal penis long and slender, with darker grey spots, more dense on mantle, penial retractor muscle inserted at apex and head and tail areas. Lateral band grey, with two internal parallel longitudinal struc- blurred. Some specimens have foot sole uni- tures, a torus and, opposite, a velum; distal formly whitish. penis short and slightly widened, containing a globular to cylindrical and perforated penial Shell papilla level with the entrance of the vas deferens into the same penis (far from the penial Slightly oblong, with 2.5 whorls. Pits in spi- apex); penial papilla and torus coated by glan- ral rows on protoconch and teleoconch, be- dular tissue (the "penial gland"). coming smaller after first whorl to disappear at ST differs from all the other Plutoniinae in end of second whorl. the following autapomorphy: proximal penis long and equipped with two internal parallel Radula longitudinal structures, torus and velum. Fol- lowing the praxis of traditional taxonomy, this Formula: 29M + 10L + (Ibáñez et al., autapomorphy is sufficient to support inclu- 1987: figs. 26-28); marginal teeth with very sion of ST in a distinct supraspecific taxon small ectocone. closely related to the other Macaronesian Plu- toniinae. Genital System The structure of the glándula amatoria in ST and in the other Plutoniinae supports the hy- Shown in Figures 2J, 4E; 18 specimens dis- pothesis of a homology with the sarcobelum of sected. Penis longer than vagina (including its Semilimax, even if the latter is distinct from the glándula amatoria portion), generally coiled in proximal vagina. The homology was advanced natural position, fringed on one side. Proximal by Simroth (1886, 1889), rejected by Wieg- penis very long, proximally pointed. Distal mann (1886), and later accepted by several penis short, slightly widened. Laminar crest authors, such as Hesse (1923), Schileyko narrow, sometimes vestigial. Distal torus thin, (1986), and Hausdorf (1995). partially surrounding penial papilla and ex- To verify the status (genus/subgenus) of tending into distal penis as a thick pilaster, ST, that of the other Macaronesian Plutoni- which ends in a small distal slope with trans- inae and their relationships we applied a versal furrows, reminiscent of a fingerprint. cladistic approach. Vagina twice as long as the glándula amatoria portion, one-third longer than free Plutoniinae Phylogenetic Relationships oviduct and little longer than bursa copulatrix duct, with two thick internal longitudinal folds Explanations of the characters used in the which converge near the atrium. Glandula phylogenetic analysis; the numeration refers amatoria with small spout, almost blunt. to Table 2 and 3 and the cladogram (Fig. 6). In CANARIVITRINA AND THE PLUTONIINAE PHYLOGENY 55

these explanations and Table 3, we utilize the imax, have the penial retractor muscle with a name Plutonia in the old sense, that is, only normal course between its insertions, free for P. atlántica. from the right optic nerve. (1) Shell. Semilimax, as well as ST, Arabi- In ST, Guerhna, Insulivitrina, and Madei- vitrina, Insulivitrina, Phenacolimax, and Ma- rovitrina the penial retractor muscle has a pe- deirovitrina have an external shell with more culiar course, rounding the right optic nerve than 1 .5 whorls, into which the animal cannot (Fig. 3G). withdraw. Plutonia has no penial retractor muscle, Gallandia and Guerhna have a conical-de- probably due to a secondary reduction origi- pressed shell, keeled in Guerhna and Gallan- nating independently from that of S. semili- dia lederi, with a small aperture through which max.

the animal can withdraw entirely. (8) Vaginal retractor muscle. ST, Arabivi- Plutonia has a reduced, capuliform, internal thna, Gallandia, Guerhna, Insulivitrina, Phe- oval shell, with about half a whorl. nacolimax, and Plutonia have no vaginal re- (2) Teleoconch ornamentation. ST, Insulivi- tractor muscle, like Semilimax. trina, Madeirovitrina, and Phenacolimax have Madeirovitrina, with the exception of M. al- a glossy teleoconch, like Semilimax. bopalliatus (see above), has a vaginal retrac- The Arabivithna teleoconch ornamentation tor muscle.

is variable (see Appendix). (9) Oviducal retractor muscle. ST, Arabivi- Gallandia lederi and G. olympica have an thna, Gallandia, Guerhna, Insulivitrina, Ma- entirely ribbed teleoconch, with the ribs rami- deirovitrina, and Phenacolimax have no ovid- fied and interconnected in G. olympica; G. an- ucal retractor muscle, like Semilimax. nularis has a glossy teleoconch. Therefore Plutonia has a short oviducal retractor with the character state is variable within the a branch inserting at the bursa copulatrix genus. duct. Guerhna has an entirely and very densely ribbed teleoconch, with a pattern different Cladistic Analysis from that of Gallandia lederi and Gallandia olympica. The Plutoniinae phylogenetic relationships

Plutonia has the teleoconch with concentric are not easy to establish, because it is not yet growth lines but the shell is internal and its or- possible to determine the state of several namentation is not comparable with that of the characters which initially appear to be taxo- other Plutoniinae. nomically interesting, due to insufficient (3) Radula. Semilimax, as well as almost all knowledge, intrageneric variability or redun- the Vithnidae, has the radula of the dichoglos- dancy (see Appendix). san type. Character 7 was treated as ordered by the The radula of Plutonia belongs to the bel- possible synapomorphy of one Azorean In- oglossan type. sulivitrina species and Plutonia atlántica, re- (4) Location of the glándula amatoria. In ferred to the absence of the penial retractor Semilimax, this is a club-like atrial diverticu- muscle (as indicated in the Comments after lum distinct from vagina. the description of the genus Plutonia); a simi- All the Plutoniinae have a glándula amato- lar scenario was found also in the outgroup. ria fused with proximal vagina. {Semilimax semilimax does not have a penial (5) Penis. Semilimax, as well as Arabivit- retractor muscle but one is present in S. kotu- hna, Gallandia, Guerhna, Insulivitrina, Ma- lae, free from the right optic nerve: see Ap- deirovitrina, Phenacolimax, and Plutonia, pendix.) have a penis without a long proximal portion Character 4 justifies the monophyly of the equipped with torus and velum. Plutoniinae with respect to the outgroup. Un-

ST has a penis with a long proximal portion fortunately, only three other characters (1, 6 equipped with torus and velum. and 7) are phylogenetically informative; the

(6) Penial sheath. Semilimax and remaining characters (2, 3, 5, 8, and 9) are Phenacolimax have a strong penial sheath. only found in one taxon. In the other Plutoniinae, the penial sheath is Eight informative (but not resolutive) trees rudimentary or absent. from the matrix (Table 3) were found. How- (7) Penial retractor muscle. Semilimax, as ever, the strict consensus of these trees well as Arabivithna, Gallandia, and Phenacol- shows the Plutoniinae as the sister group of 56 ALONSO ETAL.

Semilimax (character 4) and Phenacolimax Semilimax as the sister group of the remaining Plutoni- inae (character 6). Moreover, the eight trees Phenacolimax show ST (key C), Insulivitrina (key F), and Madeirovitrina (key G) grouped together at Arabivitrina the same level. The trees are as follows: 1-1 tree 1 = (A((B(CFG)(D(EI)))H)) Gallandia tree2 = (A((B(C(DEI)FG))H)) tree3 = (A((B(CDEFGI))H)) P. (Canarivitrina) tree4 = (A((B(CEFGI)D)H)) n. subgen. tree5 = (A((B(CFGI)(DE)H)) 1-1 P. (Guerrina) tree6 = (A((BC(DEI)FG)H)) tree 7 = (A((B(C(DE)FGI))H)) tree8 = (A((BC(DE)FGI)H)) P. (Insulivitrina) Strict consensus tree = (A((BCDEFGI)H)) P. (Madeirovitrina) Figure 6 represents tree number 4 of the analysis, with a consistency index (CI) of 1-2 0.917, a retention index (Rl) of 0.800 and a P. (Plutonia) rescaled CI index (RC) of 0.733. This tree shows a probable phylogenetic scenario with FIG. 6. Hypothesis of phylogeny of Plutoniinae. Numbers refer to characters listed within the the Macaronesian taxa included in the genus Table 2. Plutonia (of which Insulivitrina is a new syn- onymy), with a single conflict: the character state 1.1 ('shell external, the animal can withdraw into it') appears in Gallandia and Guer- CONCLUSIONS rina, but it could be homoplasic. This supposi- Plutoniinae includes tion is based on: (1) The subfamily the (A) Guerrina clearly differs from Gallandia genera Phenacolimax, Gallandia (which is al- certainly junior of Oligoli- and belongs to the genus Plutonia. This is most a synonym max,) Arabivitrina, and Plutonia. demonstrated by the fact that it shares the same synapomorphy (the peculiar course of (2) The genus Phenacolimax is the sister the penial retractor) which links all the other group of the remaining Plutoniinae. taxa of the Plutonia group. (3) The supraspecific taxon ST has a sub- (B) The capacity of the animal to withdraw generic category in the genus Plutonia, which Plutonia, s. str., into the shell is related to the respective di- also includes the subgenera mensions and the shell form; the suitable re- Guerrina, Insulivitrina, and Madeirovitrina. lation could be acquired independently in Gal- (4) We describe the supraspecific taxon ST landia and Guerrina. as follows: (C) The independent action of the Vitrinidae limacization process, found in Semilimax, In- Subgenus Canarivitrina sulivitrina, and Madeirovitrina, as well as in Valido & Alonso, n. subgen. other Vitrinidae; these taxa have some species with the ventral side of the shell well Type species: P. (C.) taburientensis Groh & developed and other species with the shell Valido, . sp. auricular with very wide aperture and very reduced ventral side (see Appendix). The Guer- Diagnosis rina shell form could have originated as a phase opposite to the auricular form in the li- Plutonia with a long penis divided into two macization process of the genus Plutonia. portions. Proximal portion long, slender, with (D) Gallandia and Guerrina live at opposite penial retractor muscle inserted at apex and ends of the geographical range of the Plutoni- two internal parallel longitudinal structures: inae. torus and, opposite, velum. Distal portion

After excluding character 1 of the analysis, short, slightly widened, containing a globular only one tree was found: the same one repre- to cylindrical and perforated penial papilla sented in Figure 6, with a consistency index level with entrance of the vas deferens into (CI) of 1.0000. penis (far from proximal apex). Penial papilla CANARIVITRINA AND THE PLUTONIINAE PHYLOGENY 57

and torus coated by glandular tissue, the "pe- Waiden (1984) concluded that "the most con- nial gland". spicuous difference between the mollusc fau- nas of Madeira and the Canary Islands is the complete absence of taxa with NW African affinities on Madeira". BIOGEOGRAPHY The different faunistic composition of these archipelagoes is probably due, at least par- The monophyly of the Macaronesian Plu- tially, to the different origin of the colonizers toniinae has biogeographical implications. that landed on them after the Pleistocene The three archipelagoes housing these Plu- glaciations. The Azorean fauna and flora are toniinae, plus the Cape Verde Islands (which much less diverse than those of the other also house a vitrinid, probably a Plutoniinae: archipelagoes (Ashmole et al., 1996). Groh, 1983) are often considered to form a A conspicuous example of the different fau- biogeographic unit called Macaronesia (= nistic composition is offered by the Bulim- "fortunate islands"), located in the north-east- inidae, a taxon present with different subfam- ern Atlantic at the western fringe of the ilies on the Azores and the Canaries (Hesse, Palaearctic. The Azores are the most remote 1933; Henriquez et al., 1993). The Madeiran from continental land, nearly in the centre of archipelago lacks Buliminidae, their niches the Atlantic, and are situated at about 850 km occupied by the Clausiliidae, a group absent NW of Madeira, the latter lying about 650 km from the Azores and the Canaries (Neubert & from Morocco. Groh, 1998). The Canary Islands lie about 450 km south Eason & Ashmole (1992) evidenced isola- of Madeira, about 500 km north of the Tropic tion of the Azores to be severe and that it de- of Cancer, and their easternmost island is pends not only on their distance from conti- only 100 km distant from Morocco. Land nents, but also on meteorological and bridges with Morocco have been hypothe- océanographie conditions, which are not con- sized in the past but today the oceanic origin ducive to the arrival of animals or plants by of all the Canary Islands is currently attributed sea from western Europe or Africa. Surface to the activity of a hotspot in the Earth's man- currents are, in fact, basically from the west tle (Holik et al., 1991; Carracedoetal., 1998). throughout the year. The same authors also The oldest island of the Azores dates back pointed out that water circulation in the North nearly 5.5 Ma (Nunn, 1994), the oldest part of Atlantic was in a highly dynamic state around the Madeiran archipelago has an age be- the end of the last glaciation and earlier in the tween 13-20 Ma (Mitchell-Thomé, 1976) and Pleistocene, and that colonization by the an- that of the Canaries, 22.5 Ma (Ancochea et cestors of the modern Azorean forms might al., 1993). have occurred during a relatively short phase The present subtropical flora of the three in one of the glaciation cycles during which northern archipelagoes, which includes the oceanic currents were more conducive to col- evergreen laurel-forest or "laurisilva," is onization than they are at present. closely related and also with that of the As for the Madeiran fauna, Baez (1993)

Miocene and Pliocene of the Mediterranean suggested it must have been originated region. Nevertheless, the fauna is different, mainly by anemochore or idrochore trans- although it is predominantly of a Palaeartic oceanic colonization. Nevertheless, the sea- origin (the Azores have also Neartic affinities; mounts along the Madeira-Iberian crest may Eason & Ashmole, 1 992). The flora and fauna have provided stepping stone islands in early of the Cape Verde Islands show a remarkably times. strong Ethiopian influence, especially of the Finally, the Canary Islands colonization has region called Saharo-Sindian, (Lobin, 1982), probably taken place from northwest Africa, but it also include some relicts of the laurel- which in the past was covered by dense forest. With respect to the malacofauna there forests and had mighty rivers (Baez et al., is a mixture of Palaeartic, Ethiopian and Mac- 1983). In the Miocene-Pliocene, the Atlantic aronesian elements in equal portions (Groh, Moroccan rivers would drag vegetation islets, 1983). As Baez (1993) indicated, most of the floating rafts, that would be transported to the Madeiran faunal groups present more affini- nearby coasts of the Canarian archipelago by ties with those of Central Europe than with the trade winds and the marine streams. The those of the Mediterranean area. The contrary wadi Drâa was the most probable colonizing is true for the neighbouring Canary Islands. source because its mouth is located opposite 58 ALONSO ETAL. to Lanzarote and Fuerteventura, the eastern- Chilopoda) and Plutoniinae Cockerell, 1893 (Mol- most Cañarían Islands. lusca, Gastropoda) (2). Bulletin of Zoological Nomenclature, We consider that the presence of Plutonia 53(2): 120. BACKHUYS, W., 1975, Zoogeography and taxon- on the three (or four) archipelagoes is not omy of the land and freshwater molluscs of the recent, Put probably as old as the humid lau- Azores. 350 pp., 97 maps, 32 pi. Amsterdam rel-forest, that is, existing long before the (Backhuys & Meesters). glaciations, Pleistocene as Waiden (1984) BAEZ, M., 1993, Origins and affinities of the fauna suggested, and that the evolution of the of Madeira. Boletim do Museu Municipal do Fun- genus took place in all archipelagoes in a pe- chal, Suppl. 2 (Proceedings of the Manchester riod preceding the Pleistocene glaciations. North Atlantic Islands Conference): 9-40. The Plutonia species survived the glaciations BAEZ, M., T BRAVO & J. F. NAVARRO, 1983, Ca- narias. Origen poblamiento. (Cír- probably associated to the Tertiary relict forest y 97 pp. Madrid culo de Estudios Sociales de Canarias & and later on they conquered some others of Queimada, eds.). their present biotopes, whereas their extinc- CARRACEDO, J. O, S. J. DAY, H. GUILLOU, E. tion in the continental lands may also be as- RODRIGUEZ, J. A. CANAS & F. J. PÉREZ, 1 998, sociated to the extinction of the laurel-forest. Origen y evolución del volcanismo de las Islas Canarias. In: Ciencia y Cultura en Canarias: ACKNOWLEDGEMENTS 67-89. Santa Cruz de Tenerife (OACIM). COCKERELL, T D. A., 1893, A check-list of the slugs (with appendix and notes by W. E. Our special thanks go to Dr. B. Hausdorf Collinge). The Conchologist, 2(8): 185-232. (Hamburg), Dr. P. Mordan (London), and Dr. EASON, E. H. & N. P. ASHMOLE, 1992, Indigenous E. Neubert (Frankfurt), well for the critical as centipedes (Chilopoda: Lithobiomorpha) from review of the earlier versions of the manu- Azorean caves and lava flows. Zoological Journal script and their very valuable informations and of the Linnean Society, 105: 407-429. suggestions as for disposing us some of their ECKHARDT, E., 1914, Beiträge zur Kenntnis der very important unpublished data; to Dr. F. einheimischen Vitrinen. Jenaische Zeitschrift für Giusti (Siena) and an anonymous referee for Naturwissenschaften, 51: 213-376. 112- the exhaustive review of the paper; addition- FALKNER. G., 1989, Binnenmollusken. Pp. 280, in: G. steinbach, ed.. Weichtiere. München ally to Dr. Giusti, Dr. G. Manganelli (Siena), (Mosaik Verlag). and Dr. Jorge Crisci (La Plata) for their com- FALKNER, G., 1991, Vorschlag für eine Neufas- puterized cladistic analysis of the data matrix; sung der Roten Liste der in Bayern vorkom- to Dr. Neubert for the gift of specimens of Ara- menden Mollusken (Weichtiere). Schriftenreihe bivitrina jansseni; to Dr. W. Rähle (Tübingen) Bayerisches Landesamt für Umweltschutz, for contributing his data on P. taburientensis, München, 97:61-112. n. sp., and P. ripkeni, n. sp.; to Mr. T. E. J. Rip- FALKNER, G., 1992, Binnenschnecken und ken (Delft), for his collaboration in the collect- Süßwassermuscheln. In: j. . reichholf & G. steinbach, eds., Die grosse Bertelsmann Lexiko- ing of the P. ripkeni, n. sp., material; and to thek, Naturenzyklopädie Europas, 6: 238-321. Mrs. P. Agnew for linguistic revision. München (Mosaik Verlag).

FITZINGER. L. I., 1833. Systematisches Verzeich- LITERATURE CITED ne der im Erzherzogthume Oesterreich vorkom- menden Weichthiere, als Prodrom einer Fauna ALONSO, M. R„ M. IBÁÑEZ & P. MORALES, 1987. desselben. Beiträge zur Landeskunde Oester-

La familia Vitrinidae en Canarias. II. Revisión de reichs, Enns (Verein für Vaterländische Ge- las especies de La Palma y El Hierro, con de- schichte, Wien), 3: 88-122. scripción de una especie nueva (Gastropoda: FORCART, L., 1944, Monographie der schwei- Pulmonata). Archiv für Molluskenkunde, 118: zerischen Vitrinidae (Moll. Pulm.). Revue Suisse 63-76. de Zoologie, 5(29): 629-678 + pl. 2. ANCOCHEA, E., J. L. BRÄNDLE, . R. CUBAS, F. FORCART, L, 1949, Die Erektion der Kopulations-

HERNÁN & M. J. HUERTAS, 1993, La Serie I de organe und der Kopulationsmodus von Phe- la isla de Fuerteventura. Memorias de la Real nacolimax major (Fér.). Archiv für Mollusken- Academia de Ciencias Exactas, Físicas y Natu- kunde, 77: 115-119. rales de Madrid (Ciencias Naturales). 27: 1-151. FORCART, L, 1957, Journey to the High Simien ASHMOLE N. P., P. OROMÍ, M. J. ASHMOLE & J. (Northern Ethiopia), 1952-3. Three: Species of L. MARTÍN, 1996, The invertebrate fauna of early Phenacolimax (Gastropoda. Vitrinidae). with successional volcanic habitats in the Azores. Bo- notes on the taxonomy of the genus. Journal of letim do Museu Municipal do Funchal, 48: 5-39. the Linnean Society, 43: 113-122. pl. 4. BACKELJAU, T & R. M. SHELLEY, 1996. Com- FORCART, L. 1959, Revision Nordafrikanischer ments on the proposal to remove the homonymy Vitrinidae. Archiv für Molluskenkunde. 88: 1-6, between Plutoniinae Bollman, 1893 (Arthropoda, pl. 1. CANARIVITRINA AND THE PLUTONIINAE PHYLOGENY 59

FORCART, L, 1965, Rezente Land- und Süss- with PLUTONIINAE Cockerell, 1893 (Mollusca, wassermollusken der süditalienischen Land- Gastropoda). Bulletin of Zoological Nomencla- schaften Apulien, Basilicata und Calabrien. Ver- ture, 54(3): 197-199.

handlungen der Naturforschenden Gesellschaft JUNGBLUTH, J. H., I. M. LIKHAREV & A. WIKTOR, in Basel, 76: 59-184. 1985, Vergleichend morphologische Unter-

GIUSTI, F., 1971, Notuale malacologicae 16. I mol- suchungen an der Radula der Landnackt-

luschi terrestri e di acqua dolce viventi sul mass- schnecken. II. Arionoidea und Trigonochlamy- iccio dei monti Reatini (Appennino Centrale). La- doidea (Gastropoda: Pulmonata). Archiv für vori délia Società Italiana di Biogeografia, (Nuova Molluskenkunde, 116:25-45. Serie) 2: 423-576, 7 pi. KERNEY, M. P, R. A. D. CAMERON & G. RILEY,

GROH, ., 1983, Revision der Land- und 1 979, A field guide to the land snails of Britain and Süßwassergastropoden der Kapverdischen In- north-west Europe. 288 pp. London (W. Collins). seln. Archiv für Molluskenkunde, 113: 159-223. LOBIN, W., 1982, Untersuchungen über Flora, Veg- GROH, K. & J. HEMMEN, 1986, Zur Kenntnis der etation und biogeographische Beziehungen der Vitriniden des Madeira-Archipels (Pulmonata: Kapverdischen Inseln. Courier Forschungsinsti-

Vitrinidae). Archiv für Molluskenkunde, 116: 183- tut Senckenberg, 53: 1 - 1 1 2.

217. MANGANELLI, G., M. BODON, L. FAVILLI & F.

GROSSU, A. V., 1983, Ordo Stylommatophora. GIUSTI, 1995, Gastropoda Pulmonata. In: a. Gastropoda Romaniae. 4. 563 pp. Litera, Bu- minelli, s. RUFFO, & s. la posta, eds., Checklist caresti. delle specie delta fauna italiana, 16. Bologna HAUSDORF, ., 1995, The Vitrinidae of Turkey, (Calderini). with remarks on the phylogeny of Gallandia (Gas- MERMOD, G., 1930, Catalogue des invertébrés de tropoda: Stylommatophora). Zoologischer Anzei- La Suisse. 18. Gastéropodes. Muséum d'Histoire ger, 234: 63-74. Naturelle, Genève. HAUSDORF, ., 1997, Phylogeny of the Lima- MITCHELL-THOMÉ, R. O, 1976, Geology of the

coidea sensu lato (Gastropoda: Stylommato- Middle Atlantic Islands. In: f. bender, v. jacobs-

phora). Heidia, 4(Sonderheft 5): 42-43. hagen, j. D. Jong & G. luttig, ed., Beiträge zur HAUSDORF, ., 1998, Phylogeny of the Lima- regionalen Geologie der Erde, 12: IX + 382 pp. coidea sensu lato (Gastropoda: Stylommato- Berlin & Stuttgart (Bomtraeger). phora). Journal of Molluscan Studies. 64: 35-66. MORALES, P., M. IBÁÑEZ & M. R. ALONSO, 1988,

HENRIQUEZ, F., M. IBÁÑEZ & M. R. ALONSO, La familia Vitrinidae en Canarias. III. 3 nuevas es- 1993, Revision of the genus Napaeus Albers, pecies de La Gomera (Gastropoda: Pulmonata).

1850 (Gastropoda Pulmonata: Enidae). I. The Archiv für Molluskenkunde, 118: 1 53- 1 66.

problem of Napaeus (Napaeinus) nanodes (Shut- NEUBERT, E., 1 998, Annotated checklist of the ter- tleworth, 1852) and description of five new restrial and freshwater molluscs of the Arabian species from its conchological group. Journal of Peninsula. Fauna of Arabia, 17: 333-461, Molluscan Studies, 59: 147-163. Riyadh/Basle. HESSE, P., 1923, Beiträge zur näheren Kenntnis NEUBERT, E. & K. GROH, 1998, Contributions to der Familie Vitrinidae. Archiv für Mollusken- the nomenclature and phylogeny of Boettgeria O. kunde. 55 (Systematischer Teil): 81-115. Die Vit- Boettger, 1863, with description of Loosjesiella n. rinen der atlantischen Inseln: 129-145. subgen. (Gastropoda: Pulmonata: Clausiliidae). HESSE, P, 1933. Zur Anatomie und Systematik der Basteria, 62: 157-168. Familie Enidae. Archiv für Naturgeschichte, NUNN, P. D., 1994, Oceanic Islands. 413 pp. Lon- (Neue Folge) 2(2): 145-224. don (Blackwell). HOFFMANN, H., 1929, Die Vitrinen der Atlanti- ODHNER, N. H., 1937, Little-known land Mollusca schen Inseln. Senckenbergiana, 11(4): 218-235. from Madeira and La Palma (Canary Islands). HOLIK, J. S., P D. RABINOWITZ & J. A. AUSTIN, Proceedings of the Malacological Society of Lon- 1991, Effects of Canary hostpot volcanism on don, 22: 353-364. structure of oceanic crust off Morocco. Journal of ODHNER, N. H., 1954, Vitrina (Guerrina n. sect.) Geophysical Research, 96(B7): 12039-12067. cutícula (Shuttleworth) and its relations. Proceed-

HUBENDICK, ., 1953, The relationships of the ings of the Malacological Society of London. 31 :

East African Vitrinidae with notes on the Taxon- 56-63, pi. 4. omy of Vitrina. Arkiv for Zoologi, (2)6(5): 83- PAGE, R. D. M., 1996, Treeview: an application to 96. display phylogenetic trees on personal comput- IBÁÑEZ, M., P. MORALES & M. R. ALONSO, 1987, ers. Computer Applications in the Biosciences.

La familia Vitrinidae en Canarias. I. Revisión de 12:357-358 las especies de Tenerife, con descripción de 2 (http://taxonomy.zoology.gla.ac.uk/rod/rod.html). especies nuevas (Gastropoda: Pulmonata). SCHILEYKO, A. A., 1984, Nazemnye mollyuski

Archiv für Molluskenkunde, 1 1 7: 1 1 7- 1 49. podotrada Pupillina fauny SSSR (Gastropoda, INTERNATIONAL COMMISSION ON ZOOLOGI- Pulmonata, Geophila). Fauna SSSR, Mollyuski,

CAL NOMENCLATURE [shortened ICZN], 1997. III (3) (= N. S. 130), 399 pp. Leningrad (Nauka). Opinion 1880. PLUTONIINAE Bollman, 1893 SCHILEYKO, A. A., 1986, The system and the phy- (Arthropoda, Chilopoda): spelling emended to logeny of Vitrinidae (Gastropoda Pulmonata) (in

PLUTONIUMINAE, so removing the homonymy Russian). In: y. i starobogatov & A. A. SCHI- 60 ALONSO ETAL.

LEYKO, eds., Morphological and ecological bases ZILCH, A., 1 979, Die Typen und Typoide des Natur- of the molluscan taxonomy. Proceedings of the Museums Senckenberg, 62: Mollusca: Zoni- Zoological Institute, Leningrad, 148: 124-157. tacea: Vitrinidae. Archiv für Molluskenkunde, SHELLEY, R. M. & T. BACKELJAU, 1995, Plutoni- 110:81-101, pl. 6-8. inae Bollman, 1893 (Arthropoda, Chilopoda) and

Plutoniinae Cockerell, 1893 (Mollusca, Gas- Revised ms. accepted 1 June 1999 tropoda): proposed removal of homonymy. Bul-

letin of Zoological Nomenclature, 52(2): 1 50- 1 52. SIMROTH, H., 1886, Über den Liebespfeil der Vitri- APPENDIX. TAXONOMIC COMPARISON nen. Sitzungsberichte der Naturforschenden OF THE GENUS-GROUP TAXA

Gesellschan zu Leipzig, 12 (5): 6-7. SIMROTH, H., 1889, Beiträge zur Kenntnis der Semilimax (anatomical data taken mainly der Kaiserlichen Nacktschnecken. Nova Acta from Schileyko, 1986). Shell variable in Leopoldinisch-Carolinischen Deutschen Akade- shape: auricular with very wide aperture

mie der Naturforscher, 54 (1 ): 1 -91 , 4 pis. and reduced ventral side (S. semilimax) STABILE, J., 1864, Mollusques terrestres vivants or with ventral side well developed (Vit- du Piémont. Atti délia Société Italiana di Scienze rina carniolica O. Boettger, 1 884: Grossu, Naturali, 7: 1-141 + 2 pis. SWOFFORD, D. L, 1999, PAUP*. Phylogenese 1983). Animal cannot withdraw into shell. Analysis Using Parsimony (*and Other Methods). Teleoconch glossy. Right shell lobe Version 4.0 beta 2'. Sinauer Associates, Sunder- rather narrow, twisted backwards to land, Massachusetts. cover shell apex. Radula dichoglossan THIELE, J., 1931, Gastropoda: Opisthobranchia (Jungbluth et al., 1985). Genital atrium and Pulmonata. Handbuch der systematischen short (Fig. 1). Penis short, without long Weichtierkunde, 1 377-788. Stuttgart (G. (2): proximal portion (torus and velum conse- Fischer) [reprint: A. Asher and Co., Amsterdam, quently absent). The presence of a perfo- 1963. Translated to English (R. bieler & p. m. rated penial papilla needs verification. mikkelsen, eds.): Handbook of systematic mala- Semilimax semilimax does not have a cology. 1189 pp. Smithsonian Institution Li- braries, Washington, 1992]. penial retractor muscle, but one is pres-

TILLIER, S., 1984, Patterns of digestive tract mor- ent in S. kotulae (Weste rl und, 1883), free phology in the limacisation of helicarionid, suc- from the right optic nerve. "Glandula am- cineid and athoracophorid snails and slugs (Mol- atoria" voluminous, oblong oval or club- lusca: Pulmonata). Malacologia, 25: 173-192. shaped, not covered by soft tissue, lo- TILLIER, S., 1989, Comparative morphology, phy- cated in atrial sarcobelum, which wall logeny and classification of land snails and slugs invaginates sheathing the spout (no data (Gastropoda: Pulmonata: Stylommatophora). available about connection between in- Malacologia, 30: 1 -303. vaginated sarcobelum wall and spout VALIDO, M. J., M. R. ALONSO & M. IBÁÑEZ, 1 990, La familia Vitrinidae en Canarias. IV Revisión de wall). Vagina short. las especies de Gran Canaria, con descripción Arabivitrina (anatomical data taken mainly de 3 especies nuevas (Gastropoda: Pulmonata). from Neubert, 1998). Shell medium to Archiv für Molluskenkunde, 120: 95-114. large, auricular to subglobose, with 3-3.5 VALIDO, M., . GROH, M. IBÁÑEZ & M. R. whorls. Animal cannot withdraw com- 1 993, La familia Vitrinidae en Canarias. ALONSO, pletely into shell (at least in A. jansseni). V El género, Guerrina (Gastropoda: Pulmonata). Teleoconch ornamentation variable, al- 7 Archiv für Molluskenkunde, 1 2 1 : 1 1 - 1 24. most smooth, sometimes with blunt axial WALDÉN, H. W., 1984. On the origin, affinities, and riblets in several species (as in type evolution of the land Mollusca of the mid-Atlantic islands, with special reference to Madeira. Bole- species, Vitrina arabica Thiele, 1910), or tim do Museu Municipal do Funchal, 36(158): without blunt axial riblets. Aperture bor- 51-82. dered by a small periostracal fringe. WIEGMANN, F., 1886, Der sogenannte Liebespfeil Right shell lobe rather narrow, twisted der Vitrinen. Jahrbücher der Deutschen Malako- backwards to cover shell apex. Radula 74-94. zoologischen Gesellschaft, 13: dichoglossan. Genital atrium long. Penis WIKTOR, A. & T BACKELJAU, 1995. Redescrip- as a broad tube, about as long as vagina, tion of the Azorean endemic slug Plutonia at- without long proximal portion (torus and lántica (Morelet, 1860) (Gastropoda terrestria velum are consequently absent); with a nuda). Bulletin de l'Institut Royal des Sciences at least of Naturelles de Belgique, (Biologie)65: 69-82. longitudinal pilaster consisting ZILCH, A., 1959, Gastropoda, Euthyneura. In: w. an apical glandular part followed by a wenz, ed., Handbuch der Paläozoologie, 6(2): narrow to broadened part with transverse 202-400. Berlin (Borntraeger). lamellae. Typical penial papilla missing. CANARIVITRINA AND THE PLUTONIINAE PHYLOGENY 61

Penial retractor free from right optic vaginal wall and spout wall entirely con- nerve. Vas deferens entering penis sub- nected by thin connective fibres. apically, then turning towards retractor in- Insulivitrina (anatomical data taken mainly

sertion area to run inside glandular tissue from Ibáñez et al., 1 987). Shell variable in of apical organ and open at apex. Distal shape, as in Semilimax. Teleoconch vagina and glándula amatoria similar in glossy. Animal cannot withdraw into shell. length. Glandula amatoria covered with Right shell lobe well developed, covering soft tissue, large, club-shaped to oblong almost entire shell in undisturbed condi- oval, but A. darnaudi has a short glándula tions, but disturbed specimens (Ibáñez et amatoria. Invaginated vaginal wall and al., 1987: figs. 6, 8) show this lobe similar spout wall entirely connected by strong to those published for Semilimax semili- connective fibres in A. jansseni. max, S. pyrenaicus (A. Férussac, 1821), Gallandia (anatomical data taken mainly from and Phenacolimax major (Falkner, 1989:

Hausdorf, Shell 1 1 1995). conoid-globose, 73; 992: 270-271 ; Kerney et al., 1 979, with 2.5-3.25 whorls and small aperture. pis. 6, 7). Radula dichoglossan with vari- Teleoconch entirely ribbed or glossy. Ani- able marginal teeth (finger-shaped with mal can withdraw more-or-less entirely or without ectocones, or multicuspid). into shell. Shell and mantle lobes re- Genital atrium variable in length. Penis duced (as Hausdorf, 1995, stated, this re- short, without long proximal portion (torus duction is connected with the fact speci- and velum consequently absent) and mens can withdraw completely into the with perforated penial papilla. Vas defer- shell). Radula dichoglossan. Genital ens entering penis apically to subapically. atrium long. Penis short, without long Penial retractor rounding right optic proximal portion (torus and velum conse- nerve. Glandula amatoria covered with quently absent). Vas deferens entering soft tissue, usually voluminous, club- subapically at penis, under penial gland. shaped to oblong-oval, but Insulivitrina Penial retractor free from right optic canariensis (Mousson, 1872) has a small nerve. Glandula amatoria small, club- glándula amatoria. Distal vagina similar shaped to oblong-oval, sometimes par- in length or even shorter than glándula tially covered by soft tissue, arranged like amatoria portion. Invaginated vaginal a flower calyx (no data available about wall and spout wall entirely connected by connection between invaginated vaginal thin connective fibres. wall and spout wall). Madeirovitrina (anatomical data taken mainly Guerrina (anatomical data taken mainly from from Groh & Hemmen, 1986). Shell vari- Ibáñez et al., 1987, and Valido et al., able in shape, as in Semilimax. Teleo- 1993). Shell slightly depressed, keeled, conch glossy. Animal cannot withdraw conical above, with 3.25-3.75 whorls and into shell. Right shell lobe well developed, small aperture. Teleoconch entirely and covering almost entire shell in undis- densely ribbed. Animal can withdraw en- turbed conditions. Radula dichoglossan. tirely into shell. Shell lobes and right body Genital atrium medium to long. Penis lobe of mantle very small, not evident in large and proximally globe-shaped, with live specimens; left body lobe small. perforated penial papilla subapical and Radula dichoglossan. Penis short, with without long proximal portion (torus and perforated penial papilla and two small velum consequently absent). Vas defer- portions, the homologies of which are un- ens entering penis subapically. Penial re- clear because of different location of in- tractor rounding right optic nerve. Glan- sertion of penial retractor in G. cutícula dula amatoria voluminous, club-shaped and G. christinae; without long proximal to oblong-oval, covered with soft tissue. portion (torus and velum consequently Invaginated vaginal wall and spout wall

absent). Vas deferens entering penis lat- entirely connected by thin connective fi- erally, between the two portions. Penial bres. Distal vagina wide and shorter than retractor rounding right optic nerve. Glan- glándula amatoria portion, with a strong dula amatoria small, spheroidal, with vaginal retractor muscle. (Madeirovitrina proximal end covered with soft tissue, albopalliatus Groh & Hemmen, 1986, dif-

arranged like a flower calyx. Distal vagina fers from the other species in its tubular very long, up to four times longer than penis and the absence of a vaginal re- glándula amatoria portion. Invaginated tractor muscle; this species should there- 62 ALONSO ETAL.

fore be referred to another genus, possi- between invaginated vaginal wall and bly Insulivitrina). spout wall). Distal vagina and glándula Phenacolimax (anatomical data taken mainly amatoria similar in length. from Forcart, 1957; Hausdorf, pers. Plutonia (referred to P. atlántica; anatomical comm., and our own unpublished data). data taken mainly from Wiktor & Backel- Shell globose-depressed, with 2.5-3.25 jau, 1995). Shell entirely hidden by the whorls and small aperture. Teleoconch mantle (shell lobes absent), oval, vesti- glossy. Animal cannot withdraw into shell. gial, whitish, transparent, with about half Right shell lobe rather narrow, twisted a whorl and a wide aperture. Teleoconch backwards to cover shell apex. Radula with growth lines. Radula beloglossan, dichoglossan. Genital atrium very short. with dagger-shaped teeth. Vas deferens Penis long, tubular, with a clear constric- entering penis apically. Penis short, with- tion wrapped by a short but strong penial out long proximal portion (torus and sheath (similar to those shown by Schi- velum consequently absent), penial leyko, 1986: figs. 11-12, in Eucobresia); papilla and penial retractor. Glandula am- without long proximal portion (torus and atoria voluminous, club-shaped to ob- velum consequently absent) and with long-oval (no data available about con- perforated penial papilla. Vas deferens nection between invaginated vaginal wall running inside penial sheath and entering and spout wall). Distal vagina similar in penis subapically. Penial retractor free length or even longer than glándula ama- from right optic nerve. Glandula amatoria toria portion. Short oviducal retractor with small, not completely covered by soft tis- one branch inserted into bursa copulatrix sue (no data available about connection duct.