On the relationships of the genus Napaeus (Gastropoda: Pulmonata: Enidae) with the descriptions of four new species from the Canary Islands Author Yanes, Yurena Author Martín, Javier Author Moro, Leopoldo Author Alonso, María R. Author Ibáñez, Miguel text Journal of Natural History 2009 2009-08-26 43 35 - 36 2179 2207 http://dx.doi.org/10.1080/00222930903094621 journal article 10.1080/00222930903094621 1464-5262 5217087 Napaeus bajamarensis Ibáñez and Alonso sp. nov. Type material Holotype ( Figures 2 , 4B , 7A ): TFMC ( MT 0397 ). Type locality: Barranco de Vargas , Bajamar , Anaga massif (Tenerife; UTM coordinates: 28 RCS7058 ; 300 m altitude); Leg. Manuel J. Valido and Fátima C. Henríquez , 1 December 1989 . Paratypes : 13 specimens and 160 shells, collected between 1982 and 2009, from the northeast of Tenerife ( UTM coordinates: 28 RCS6658 , 28 RCS6757 , 28 RCS6758 , 28 RCS6958 , 28 RCS6959 , 28 RCS7058 , 28 RCS7157 , 28 RCS7259 , 28 RCS7260 , 28 RCS7460 and 28 RCS8160 ), deposited in AIT and the following collections: CGH (31 shells), CHB (24 shells), CML (2 shells), NHM (1992154/1), FMNH (158200/6), NMW (Z 1992.072.01/1) and TFMC (24 shells, in 5 lots: MT 0118 , 0119 , 0135 , 0151 and 0152) . Table 3. Arrangement of all the Napaeus species described in the respective conchological groups. For synonymies, see Alonso, Goodacre, et al. (2006: 172).

Conchological	Species name	Island/islet	Illustration
group
N. baeticatus	N. badiosus (Webb and Berthelot, 1833)	Tenerife	Figure 3P
N. baeticatus	N. baeticatus (Webb and Berthelot, 1833)	Tenerife	Figure 3G–J
N. baeticatus	N. bajamarensis sp. nov.	Tenerife	Figure 4B
N. baeticatus	N. bertheloti (L. Pfeiffer, 1846)	La Gomera	Figure 3B
N. baeticatus	N. boucheti Alonso and Ibáñez, 1993	La Palma	Figure 4C
N. baeticatus	N. consecoanus (Mousson, 1872)	La Gomera	Figure 3A
N. baeticatus	N. inflatiusculus (Wollaston, 1878)	La Gomera	Figure 3E
N. baeticatus	N. interpunctatus (Wollaston, 1878)	Gran Canaria	Figure 3K
N. baeticatus	N. obesatus (Webb and Berthelot, 1833)	Gran Canaria	Figure 4D
N. baeticatus	N. savinosa (Wollaston, 1878)	El Hierro	Figure 3D
N. baeticatus	N. servus (Mousson, 1872)	La Gomera	Figure 3L
N. baeticatus	N. severus (J. Mabille, 1898)	La Gomera	Figure 3F
N. baeticatus	N. subsimplex (Wollaston, 1878)	El Hierro	Figure 3C
N. baeticatus	N. tabidus (Shuttleworth, 1852)	Tenerife	Figure 3M
N. baeticatus	N. tafadaensis sp. nov.	Tenerife	Figure 3N, O
N. baeticatus	N. tenoensis Henríquez, 1993	Tenerife	Figure 4A
N. variatus	N. avaloensis Groh, 2006	La Gomera	Figure 4K
N. variatus	N. barquini Alonso and Ibáñez, 2006	La Gomera	Figure 5O
N. variatus	N. beguirae Henríquez, 1995	La Gomera	Figure 5Q
N. variatus	N. chrysaloides (Wollaston, 1878)	Gran Canaria	Figure 5B
N. variatus	N. elegans Alonso and Ibáñez, 1995	Tenerife	Figure 5E
N. variatus	N. encaustus (Shuttleworth, 1852)	La Palma	Figure 4Q
N. variatus	N. esbeltus Ibáñez and Alonso, 1995	Tenerife	Figure 5F
N. variatus	N. exilis Henríquez, 1995	Gran Canaria	Figure 5G
N. variatus	N. gruereanus (Grasset, 1857)	El Hierro	Figure 5N
N. variatus	N. helvolus (Webb and Berthelot, 1833)	Tenerife	Figure 4M
N. variatus	N. huttereri Henríquez, 1991	Alegranza islet	Figure 5D
(Lanzarote)
N. variatus	N. indifferens (Mousson, 1872)	Gran Canaria	Figure 5C
N. variatus	N. isletae Groh and Ibáñez, 1992	Gran Canaria	Figure 4O
N. variatus	N. lajaensis Castillo, Yanes, Alonso and	Tenerife	Figure 4T
Ibáñez, 2006 (fossil)
N. variatus	N. lichenicola Alonso and Ibáñez, 2007	Fuerteventura	Figure 5M
N. variatus	N. maculatus Goodacre, 2006	La Gomera	Figure 4L
N. variatus	N. maffioteanus (Mousson, 1872)	Gran Canaria	Figure 5A
N. variatus	N. moquinianus (Webb and Berthelot, 1833)	Gran Canaria	Figure 4R
N. variatus	N. myosotis (Webb and Berthelot, 1833)	Gran Canaria	Figure 5L
N. variatus	N. ocellatus (Mousson, 1872)	El Hierro	Figure 4H
N. variatus	N. orientalis Henríquez, 1995	La Gomera	Figure 4G
N. variatus	N. ornamentatus sp. nov.	La Gomera	Figure 4F
N. variatus	N. osoriensis (Wollaston, 1878)	Gran Canaria	Figure 5J
N. variatus	N. palmaensis (Mousson, 1872)	La Palma	Figure 5K
N. variatus	N. procerus Emerson, 2006	La Gomera	Figure 4U
N. variatus	N. propinquus (Shuttleworth, 1852)	Tenerife	Figure 4E

( Continued ) Table 3. ( Continued )

Conchological	Species name	Island/islet	Illustration
group
N. variatus	N. roccellicola (Webb and Berthelot, 1833)	Tenerife	Figure 5P
N. variatus	N. rufobrunneus (Wollaston, 1878)	Lanzarote	Figure 4V
N. variatus	N. rupicola (Mousson, 1872)	La Gomera	Figure 4J
N. variatus	N. subgracilior (Wollaston, 1878)	La Palma	Figure 5H
N. variatus	N. teobaldoi sp. nov.	Tenerife	Figure 4S
N. variatus	N. variatus (Webb and Berthelot, 1833)	Tenerife	Figure 4P
N. variatus	N. voggenreiteri Hutterer, 2006	La Gomera	Figure 4N
N. nanodes	N. bechi Alonso and Ibáñez, 1993	Tenerife	Figure 5T
N. nanodes	N. doliolum Henríquez, 1993	Tenerife	Figure 5S
N. nanodes	N. nanodes (Shuttleworth, 1852)	Tenerife	Figure 5R
N. nanodes	N. tagamichensis Henríquez, 1993	La Gomera	Figure 5X
N. nanodes	N. taguluchensis Henríquez, 1993	La Gomera	Figure 5V
N. nanodes	N. texturatus (Mousson, 1872)	La Gomera	Figure 5U
N. nanodes	N. pygmaeus Ibáñez and Alonso, 1993	La Gomera	Figure 5Y

Etymology The specific name derives from Bajamar, the main area of the species distribution. Geographic range and habitat Endemic to Tenerife, occupying an area of about 20 km 2 along the north slope of the Anaga massif (northeast of the island), associated with lowland vegetation and humid subtropical scrub, within an altitudinal range between 20 and 750 m ( Figure 1B ). The highest locality includes a pine forest. This species is a ground dweller and is the only living Napaeus species found also in the Pleistocene Aeolian area of Mancha de La Laja, Bajamar (see arrow in Figure 1B ), where the oldest species of this genus (>130 kyr BP) has been collected ( Castillo et al. 2006 ). Diagnosis Shell robust, ovate, pale brown to tan brown coloured, with first whorls darker than the rest. Teleoconch almost smooth. Epiphallus with epiphallar caecum. Bursa copulatrix duct exceptionally long, with a diverticulum. Description Shell ( Figure 7A ) dextral, robust, obese (shell breadth (SB)/shell height (SH) index), ovate, cone-shaped above, 6¾–7½ convex whorls (almost flat), marked suture. Body whorl intermediate (body whorl height at columella level (BH)/SH index), protoconch smooth, shiny, with 1½–1¾ whorls. Aperture long, wide (aperture height (AH)/SH and aperture breadth ( AB )/SB indices, respectively), with elliptical section at palatal side, slightly angular at union of columellar and palatal edges. Upper palatal side forms angle slightly less than 90º with parietal side, aperture juts out only slightly from body whorl beginning. Whitish discontinuous peristome expanded as small lip, more developed in lower part of palatal edge, reflected in columellar edge, where it partially covers umbilical slit. Older specimens with callosity between peristome edges, nodule at union of parietal–palatal area. Figure 1. (A) Geographical location of the Azores, Madeira and the Canaries; arrows indicate possible colonization patterns used by land snails; position of sea mounts comes from Van den Broeck et al. 2008 ; (B) geographical distribution of the Napaeus species ; symbols represent 1× 1 km Universal Transverse Mercator squares; arrow points to the Pleistocene deposit from Mancha de La Laja-Bajamar, where the oldest Napaeus species ( N. lajaensis ) has been collected ( Castillo et al., 2006 ). Figure 2. Drawings of the N. bajamarensis sp. nov. holotype shell, showing the placement of the measurements obtained. AB, aperture breadth; AH, aperture height; AP, aperture perimeter; AS, aperture surface (plane view); BH, body whorl height (at columella level); BP, body whorl perimeter; BS, body whorl surface (plane view); FB, first whorls breadth; FH, first whorls height; FP, first whorls perimeter; FS, first whorls surface (plane view); PB, penultimate whorl breadth; PH, penultimate whorl height; PP, penultimate whorl perimeter; PS, penultimate whorl surface (plane view); SB, shell breadth; SH, shell height; SP, shell perimeter; SS, shell surface (plane view). Shell colour pale brown to tan brown, with first whorls darker than rest. Ornamentation almost smooth, characterized by numerous weak, radial oblique ribs, regularly placed on almost entire shell ( Figures 7A , 10A ). Shell has some shine despite ribs. Jaw aulacognathous, with numerous grooves, lateral extremities almost smooth. Radula ( Figure 8 ) with 110–126 teeth rows, formula: C+14–15L+16– 19M. Central tooth triangular–ovate, mesocone with rounded tip and two small basal ectocones. First lateral teeth bigger and stronger than central with rounded mesocone and sharp ectocone. Towards laterals, length and width of mesocone diminish as ectocone increases, last broader than mesocone. Marginal teeth with finger-shaped mesocone, much broader ectocone divided into four to six denticles of decreasing size with serrated appearance. Genital system ( Figure 9A–B ; five specimens dissected). Atrium very short. Penis about three times shorter than epiphallus, with two portions. Distal portion tubular, located between atrium and insertion of penial retractor muscle. Proximal portion short and swollen, with well-developed penial papilla ( Figure 9B ). Epiphallus tubular, opening distally on penis, two regions defined by presence of epiphallar caecum; proximal region widened, slightly longer than slender distal region. Flagellum very short. Vas deferens opens laterally on proximal end of epiphallus. Penial appendix arises in distal penis portion near retractor muscle insertion, similar in length or slightly longer than penis and epiphallus together. Part A 1 of penial appendix clearly differentiated from globular part A 2 , slightly longer than penis. Appendicular retractor muscle inserts laterally, joined to penial retractor on diaphragm walls at certain distance of proximal A 1 top; appendicular insertion delimits two A 1 portions, proximal larger than distal. Part A 3 more slender, shorter than proximal A 1 portion. Parts A 3 and A 4 clearly separated from each other, A 4 long, very thin distally, passing gradually into expanded A 5 . Figure 3. Napaeus shell drawings. (A) N. consecoanus ; (B) N. bertheloti ; (C) N. subsimplex ; (D) N. savinosa (syntype, NHM 1895.2.2.223); (E) N. inflatiusculus ; (F) N. severus ; (G) N. baeticatus ; (H) N. baeticatus morphotype subbeticatus ; (J) N. baeticatus morphotype halmyris ; (K) N. interpunctatus ; (L) N. servus ; (M) N. tabidus ; (N) N. tafadaensis sp. nov. (holotype, TFMC MT 0398); (O) N. tafadaensis sp. nov. morphotype “irregular” ( N. tafadaensis paratype, AIT); (P) N. badiosus . Figure 4. Napaeus shell drawings. (A) N. tenoensis (holotype, AIT); (B) N. bajamarensis sp. nov. (holotype, TFMC MT 0397); (C) N. boucheti (holotype, AIT); (D) N. obesatus ; (E) N. propinquus ; (F) N. ornamentatus sp. nov. (holotype, TFMC MT 0400); (G) N. orientalis (holotype, AIT); (H) N. ocellatus (syntype, ZMZ 513767); (J) N. rupicola ; (K) N. avaloensis (holotype, TFMC MT 0382); (L) N. maculatus (holotype, TFMC MT 0386); (M) N. helvolus ; (N) N. voggenreiteri (holotype, TFMC MT 0384); (O) N. isletae (holotype, AIT); (P) N. variatus ; (Q) N. encaustus ; (R) N. moquinianus ; (S) N. teobaldoi sp. nov. (holotype, TFMC MT 0399); (T) N. lajaensis (holotype, TFMC MT 0381); (U) N. procerus (holotype, TFMC MT 0385); (V) N. rufobrunneus . Figure 5. Napaeus shell drawings. (A) N. maffioteanus ; (B) N. chrysaloides ; (C) N. indifferens ; (D) N. huttereri (holotype, AIT); (E) N. elegans (holotype, AIT); (F) N. esbeltus (holotype, AIT); (G) N. exilis (holotype, AIT); (H) N. subgracilior ; (J) N. osoriensis ; (K) N. palmaensis (syntype, NHM 1895.2.2.219); (L) N. myosotis ; (M) N. lichenicola (holotype, TFMC MT 389); (N) N. gruereanus ; (O) N. barquini (holotype, TFMC MT 0383); (P) N. roccellicola ; (Q) N. beguirae (holotype, AIT); (R) N. nanodes ; (S) N. doliolum (paratype, AIT); (T) N. bechi (holotype, AIT); (U) N. texturatus ; (V) N. taguluchensis (holotype, AIT); (X) N. tagamichensis (paratype, AIT); (Y) N. pygmaeus (holotype, AIT). Free oviduct longer than vagina. Vagina firmly fixed to the body tegument by short connective fibres. Bursa copulatrix complex with a well-developed diverticulum. Bursa duct exceptionally long. Two spermatophores found in diverticulum and basal stalk of bursa copulatrix complex of one specimen . Spermatophore rigid, long, narrow, twisted, with two parts delimited by strong, spur-like hook curved towards distal tip, moulded in internal pit of epiphallar caecum. Proximal area compressed near tip with semilunar section; proximal tip probably moulded into flagellum, with oval orifice for sperm transfer; proximal edge reflected as collar in ellipsoidal lip with notch on one side. Remainder proximal area tubular, with two well-developed, non-striated longitudinal lamellae running from proximities of the lip up to near the hook. Distal area tubular, narrowing gradually towards tip (first part to be inserted in other specimen during mating), ending in rounded point. Spermatophore similar to that of N. moquinianus (Webb and Berthelot) from Gran Canaria, photographed in Alonso et al. (1995: 311 , figures 22–24). Figure 6. Scatter plots of some shell measurements for the new species and the nearest species of their respective conchological group. AH, aperture height; AS, aperture surface (plane view); FH, first whorls height; FP, first whorls perimeter; FS, first whorls surface (plane view); SB, shell breadth; SH, shell height. Figure 7. Holotype shells (A–D) and a specimen (E). (A) N. bajamarensis sp. nov. ; (B) N. tafadaensis sp. nov. ; (C) N. ornamentatus sp. nov. ; (D) N. teobaldoi sp. nov. ; (E) N. ornamentatus , paratype from Mirador de Taguluche (L. Moro photograph). Remarks Shell morphology . Within the conchological group of N. baeticatus , the shell of N. bajamarensis sp. nov. ( Figure 4B ) is comparable in shape and size mainly with those of N. boucheti ( Figure 4C ), N. obesatus ( Figure 4D ) and N. tenoensis ( Figure 4A ). The shell of N. bajamarensis is more slender than that of N. tenoensis and N. boucheti and is slightly more obese than that of N. obesatus ( Figure 6A ). The first whorls of N. bajamarensis are similar but slightly more obese than those of N. obesatus and bigger than those of N. tenoensis and N. boucheti ( Figure 6C ). The shell aperture of N. bajamarensis is bigger than that of N. obesatus and smaller than those of N. tenoensis and N. boucheti ( Figure 6B ). The shell ornamentation is almost smooth in N. bajamarensis ( Figure 10A ) and well-developed in the other species ( Figure 12 A–C ). Figure 8. Radula of N. bajamarensis sp. nov. (A) Central tooth and first lateral teeth; (B) lateral teeth; (C) outermost lateral teeth and first marginal teeth; (D) outermost marginal teeth. Genital system anatomy . N. bajamarensis has an exceptionally long bursa copulatrix duct; it is clearly shorter in the other three species. N. bajamarensis and N. obesatus have a well-developed diverticulum, which is lacking in N. tenoensis and N. boucheti .