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Three Undescribed Species of Napaeus (Gastropoda: Pulmonata: Enidae) from La Gomera (Canary Islands), the Richest Centre of Species Radiation for the Genus

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Three Undescribed Species of Napaeus (Gastropoda: Pulmonata: Enidae) from La Gomera (Canary Islands), the Richest Centre of Species Radiation for the Genus JOURNAL OF CONCHOLOGY (2013), VOL.41, NO.3 271 THREE UNDESCRIBED SPECIES OF NAPAEUS (GASTROPODA: PULMONATA: ENIDAE) FROM LA GOMERA (CANARY ISLANDS), THE RICHEST CENTRE OF SPECIES RADIATION FOR THE GENUS 1 2 3 1 4 4 JESÚS SANTANA , MIGUEL ARTILES , YURENA YANES , FRANCISCO DENIZ , MARIA R. ALONSO & MIGUEL IBÁÑEZ 1Las Palmas de Gran Canaria, Gran Canaria, Canary Islands, Spain 2Arinaga, Gran Canaria, Canary Islands, Spain 3Department of Geology, University of Cincinnati, Cincinnati, OH 45221, USA 4Department of Animal Biology, La Laguna University, E–38206 La Laguna, Tenerife, Canary Islands, Spain Abstract Three new species of the genus Napaeus (Albers 1850) are described from La Gomera (Canary Islands). This is one of the smaller islands of the archipelago but it has the highest number (26) of living species of the genus. Shells of Napaeus species on La Gomera include the shortest, the most slender, the longest and the widest in the genus and show significant diversity in shell colour. Moreover, the Napaeus species from La Gomera exhibit more different patterns of genital system anatomy (five) than the whole of the genus in all the other Canary islands together (four patterns, one of which is also present on La Gomera). Hesse’s subgenera Napaeus and Napaeinus correspond to two of the eight anatomical patterns involved. At least adults of N. doloresae n. sp. and juveniles of the other new species described can disguise their shells with a cover of soil, apparently to reduce predation. Key words Napaeus, taxonomy, species radiation, insular endemics, genital anatomy, shells, shell disguise INTRODUCTION Gomera but four times more extensive and 466 Present knowledge of the Canary Islands archi- metres higher, where only 13 known living and pelago (eastern mid-Atlantic) indicates that the one possibly extinct species have been described small rounded island of La Gomera (378 km2 and to date (Webb & Berthelot, 1833; Mousson, 1872; 1484 m altitude, 12 Mya old) is much the rich- Wollaston, 1878; Groh et al., 1992; Alonso et al., est in living species of Napaeus, with 23 species 1995; Yanes, Santana et al., 2011). In the present study three new Napaeus species (Mousson, 1872; Wollaston, 1878; Henríquez, are described from western, southern and east- Ibáñez et al., 1993; Alonso et al., 1995; Alonso, ern areas of La Gomera island. All of them occur Goodacre et al., 2006; Bank et al., 2002; Yanes outside the National Park of Garajonay, which et al., 2009; Yanes, Martin et al., 2011; Holyoak occupies the centre of the island. As in previ- et al., 2011). Each of the Canarian species of ous papers, the new species are not assigned to Napaeus is typically restricted to a small area the Napaeus subgenera (Napaeus and Napaeinus) within a single island (i.e. they demonstrate “sin- described by Hesse (1933) because a phyloge- gle island endemism”). The number of Napaeus netic analysis of the genus has not been carried species from La Gomera is clearly greater than out. that predicted by the most parsimonious species number–island age model for the richness of the main snail families of the Canarian Archipelago MATERIALS AND METHODS (Cameron et al., 2013). Thus, it contrasts with that of the neighbouring Tenerife (8 Mya), which Maps of geographical distribution (Fig. 1) were is five times more extensive and 2227 metres produced using MapViewer software (Golden higher than La Gomera, but accommodates only Software Inc.). The photographic methodology was described by Ibáñez et al. (2006). Drawings 16 known living and one extinct Napaeus spe- of shell outlines (Fig. 2) were obtained semi- cies described to date (Castillo et al., 2006; Yanes automatically, adopting the methods used by et al., 2009), and also with that of Gran Canaria Yanes et al. (2009). Standardised measurements Island (14.5 Mya), which is rounded like La of the shells (Table 1, Fig. 2) were made following Contact author : [email protected] Alonso, Nogales et al. (2006), using the software 272 SANTANA ET AL. Figure 2 Drawings of the holotype shell of N. dolore- sae sp. nov. showing the placement of the measure- ments obtained (in mm or mm2): AB aperture breadth; AH aperture height; AP aperture perimeter; AS, aperture surface area (plane view); BH body whorl height (at columella level); BP body whorl perim- eter; BS body whorl surface area (plane view); FB first whorls breadth; FH first whorls height; FP first Figure 1 Geographical distribution of the three new whorls perimeter; FS first whorls surface area (plane species of Napaeus. view); PB penultimate whorl breadth; PH penulti- mate whorl height; PP penultimate whorl perimeter; PS penultimate whorl surface area (plane view); SB analySIS® (Soft Imaging System GmbH). The shell breadth; SH shell height; SP shell perimeter; SS angle at the upper palatal corner between the shell surface area (plane view); T maximum distance columella and the upper palatal side of the from columella to start of body whorl; U maximum distance from columella to palatal aperture lip; α aperture, as well as the lateral extension of the angle of columella-upper palatal side. aperture beyond the penultimate whorl were also measured, following Yanes, Martin et al. (2011). Abbreviations for shell characters and ABBREVIATIONS measurements are shown in Fig. 2. The number of shell whorls was counted using the methodol- AIT Alonso and Ibáñez collection, ogy described by Kerney & Cameron (1979: 13). Department of Animal Biology, Terminology for the shape and proportions of University of La Laguna, Tenerife, shells is based on the biometric data provided in Canary Islands, Spain Table 1, following Henríquez, Ibáñez et al. (1993; FDGC F. Deniz private collection, Las see also Table 2), and that of parts of the penial Palmas de Gran Canaria, Spain appendix follows Schileyko (1984: 39, fig. 18). ICZN International Commission on “Proximal” and “distal” refer to the position in Zoological Nomenclature relation to the ovotestis. The distinction between JMHGC J. M. Hernández private collection, “epiphallus” and “penis” is based on the inter- Las Palmas de Gran Canaria, Spain nal anatomy of these organs, not the position of JSGC J. Santana private collection, Las insertion of the penial retractor muscle which Palmas de Gran Canaria, Spain has been used by many early authors and some MAGC M. Artiles private collection, recent researchers. Arinaga, Gran Canaria, Spain THREE NEW NAPAEUS SPECIES FROM LA GOMERA 273 Table 1 Data for the shell characters measured (in mm or mm2) for Napaeus spp. sp1: N. doloresae; sp2: N. estherae; sp3: N. magnus; n, number of measured specimens; SD, standard deviation; Min., minimum; Max., maximum; other abbreviations as in Fig. 2. Statistical Character/ sp1 sp2 sp3 Character/ sp1 sp2 sp3 parameter index index Mean SH 13,24 21,03 22,25 AP 18,20 24,52 31,17 SD 0,50 1,39 1,44 0,87 1,18 1,60 Min. 12,59 19,61 19,93 17,39 23,13 28,57 Max. 14,24 23,44 24,23 20,14 26,82 33,71 Mean SB 7,69 10,11 13,00 FH 2,65 5,30 3,92 SD 0,28 0,51 0,67 0,30 0,64 0,74 Min. 7,21 9,59 11,97 2,14 4,27 2,87 Max. 8,27 11,25 14,09 3,20 6,49 5,06 Mean SS 65,40 141,49 191,04 FB 3,61 6,49 6,59 SD 3,62 13,26 18,33 0,16 0,42 0,41 Min. 61,72 127,73 167,80 3,32 5,88 5,92 Max. 72,18 162,36 217,46 3,82 7,18 7,21 Mean SP 33,32 50,77 56,46 FS 6,24 21,72 15,60 SD 1,03 2,91 3,14 0,90 4,03 3,84 Min. 32,14 48,17 51,62 4,73 16,09 11,29 Max. 35,46 55,44 60,69 8,01 29,13 21,73 Mean BH 8,58 12,09 14,83 FP 10,58 19,58 17,86 SD 0,35 0,70 0,76 0,62 1,67 1,59 Min. 8,13 11,20 13,67 9,59 17,04 15,25 Max. 9,22 13,45 16,24 11,69 22,23 20,06 Mean BS 49,28 91,78 145,01 PH 2,12 3,63 3,50 SD 3,06 6,64 13,20 0,35 0,30 0,36 Min. 45,90 83,80 131,09 1,68 3,27 2,99 Max. 55,73 102,06 167,40 3,14 4,25 3,96 Mean BP 27,40 38,87 47,51 PB 5,29 8,00 9,58 SD 0,67 1,67 2,22 0,16 0,35 0,36 Min. 26,48 37,00 45,16 5,02 7,47 9,05 Max. 28,66 41,46 50,62 5,52 8,44 10,02 Mean AH 6,34 8,47 10,91 PS 9,88 27,99 30,35 SD 0,36 0,50 0,49 1,07 3,49 4,51 Min. 5,97 7,74 10,21 7,86 23,33 24,79 Max. 7,05 9,33 11,64 11,57 33,87 36,43 Mean AB 5,46 7,24 9,14 PP 13,81 22,48 24,72 SD 0,25 0,40 0,67 0,53 1,25 1,25 Min. 5,10 6,52 8,07 13,12 20,66 22,80 Max. 6,00 7,84 10,23 14,60 24,24 26,28 Mean AS 24,76 45,04 72,21 SB/SH 0,58 0,48 0,58 SD 2,54 4,31 7,85 BH/SH 0,65 0,58 0,67 Min.
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