Napaeus 属是大西洋加纳利群岛 (Canary Islands) 特有的一类陆生蜗牛，属艾纳螺科 (Enidae)。在这些岛屿上生活着约 65 种 Napaeus 属蜗牛，曾经还有 2 种现已灭绝。这些蜗牛都生活在非常狭小的区域，比如说某一个非常小的岛屿，却有着很高的物种多样性，是研究物种适应性辐射的典型材料。在 La Gomera 这一小岛上就生活着 23 种 Napaeus 属蜗牛，是加纳利群岛上 Napaeus 属蜗牛种数最多的一个岛屿，Tenerife 岛上生活有 16 种，Gran Canaria 岛上生活有 13 种，最西边的 El Hiero 岛和 La Palma 岛上有各有 5 种，最东部的 Lanzarote 岛上有 2 中，Fuerteven-tura 岛上有 1 种 (Yanes et al. 2009, Yanes et al. 2011)。然而，虽说加纳利群岛名义上有数十种 Napaeus 蜗牛，但是它们都是根据壳形的微小差异进行分类的，并缺乏分子系统分析的证据。Yanes (2009) 曾将 Napaeus 的 56 个种总结起来归为 3 个概念组：N. baeticatus, N. variatus, N. nanodes。后来的研究中则仍旧沿用了之前的分类标准。

Napaeus bajamarensis Ibáñez and Alonso (N. baeticatus group)

Type material: Holotype (Yanes et al. 2009, Figures 2, 4B, 7A): TFMC (MT 0397).
Type locality: Barranco de Vargas, Bajamar, Anaga massif (Tenerife; UTM coordinates: 28RCS7058; 300 m altitude); Leg. Manuel J. Valido and Fátima C. Henríquez, 1 December 1989.

(Yanes et al. 2009) Fig. 7A, Holotype, N. bajamarensis sp. nov. Fig. 9. N. bajamarensis sp. nov., paratypes from Barranco de Vargas, Bajamar. (A) Genital system; (B) inner anatomy of distal epiphallus and proximal penis, showing the penial papilla.

Napaeus tafadaensis Yanes (N. baeticatus group)

Type material: Holotype (Figures 3N, 7B): TFMC (MT 0398).
Type locality: Montaña Tafada, Anaga massif (Tenerife; UTM coordinates: 28RCS8761; 600m altitude). Leg. M. Ibáñez, 6 April 1991.

(Yanes et al. 2009) Fig. 7B, Holotype, N. tafadaensis sp. nov. Fig. 11A, N. tafadaensis sp. nov., paratype from Montana Tafada.

Napaeus ornamentatus Moro (N. variatus group)

Type material: Holotype (Yanes et al. 2009, Figures 4F, 7C): TFMC (MT 0400).
Type locality: Mirador de Taguluche (La Gomera; UTM coordinates: 28RBS7113; 800 m altitude). Leg. Javier Martín and Leopoldo Moro, 5 March 2008.

(Yanes et al. 2009) Fig. 7C, Holotype, N. ornamentatus sp. nov. Fig. 11B, N. ornamentatus sp. nov., paratype from Mirador de Taguluche.

Napaeus teobaldoi Martín (N. variatus group)

Type material: Holotype (Figures 4S, 7D): TFMC (MT 0399).
Type locality: Barranco Hondo, in the southeastern slope of La Culata mountain (Tenerife; UTM coordinates: 28RCS6742; 270 m altitude). Leg. Javier Martín, 7 May 2008

(Yanes et al. 2009) Fig. 7D, Holotype, N. teobaldoi sp. nov. Fig. 11C, N. teobaldoi sp. nov., paratype from Barranco Hondo

Napaeus josei Santana, Alonso & Ibáñez

Type material: Holotype (Yanes et al. 2011, Fig. 3A) TFMC (MT 0415). Leg. J. Santana, 26 January 2009.
Type locality: Barranco del Lechugal (Gran Canaria; UTM: 28RDR3075, 100 m altitude).

(Yanes et al. 2011) Fig. 3A, Holotype, Napaeus josei n. sp. Fig. 6, Napaeus josei n. sp., paratype from Barranco del Lechugal, Gran Canaria (AIT).

Napaeus venegueraensis Artiles, Santana & Deniz

Type material: Holotype (Yanes et al. 2011, Fig. 3D) TFMC (MT 0416). Leg. M. Valido and M. Ibáñez, 29 December 1988.
Type locality: Barranco de Veneguera (Gran Canaria; UTM: 28RDR2583, 100 m altitude).

(Yanes et al. 2011) Fig. 3D, Holotype, N. venegueraensis n. sp. Fig. 8A, N. venegueraensis n. sp., paratype from Barranco de Taurito, Gran Canaria (AIT).

Napaeus arinagaensis Artiles, Deniz & Martín

Type material: Holotype (Yanes et al. 2011, Fig. 3N) TFMC (MT 0417). Leg. M. Valido and M. Ibáñez, 9 July 1989.
Type locality: North slope of Arinaga Mountain (Gran Canaria; UTM: 28RDR6182, 180 m altitude).

(Yanes et al. 2011) Fig. 3N, Holotype, Napaeus arinagaensis n. sp.

Napaeus validoi Ibáñez, Alonso & Martín

Type material: Holotype (Yanes et al. 2011, Fig. 3F) TFMC (MT 0418). Leg. M. Nogales, 10 November 1984.
Type locality: Ñameritas (Gran Canaria; UTM: 28RDR3389, 1000 m altitude).

(Yanes et al. 2011) Fig. 3F, N. validoi n. sp. Fig. 8B, N. validoi n. sp., paratype from Monte de Inagua, Gran Canaria (AIT).

Napaeus grohi Alonso, Ibáñez & Santana

Type material: Holotype (Yanes et al. 2011, Fig. 3J) TFMC (MT 0419, shell and dissected soft body). Leg. M. Ibáñez, 5 January 1993.
Type locality: Pista del Derrabado (El Hierro; UTM: 28RDR6182, 850 m altitude).

(Yanes et al. 2011) Fig. 3J, Napaeus grohi n. sp. Fig. 10, Napaeus grohi n. sp., holotype. A. Genital system. B. Detail of the flagellum. C. Internal anatomy of penis.

Napaeus doloresae Santana

Type material: Holotype (Santana Benítez et al. 2013, Fig. 4A) TFMC (MT 0843); Leg. J. Santana, 23 July 2010.
Type locality: West side of Tagamiche mountain (La Gomera; UTM: 28RBS8609; 879 m altitude).

(Santana Benítez et al. 2013) Fig. 4A, N. doloresae sp. nov. Fig. 7A, N. doloresae sp. nov., from Tagamiche, La Gomera.

Napaeus estherae Artiles

Type material: (Santana Benítez et al. 2013, Fig. 8D) 1 sh, TFMC (MT 0842); Leg. J. Santana, 7 December 2010.
Type locality: La Mérica, north slope (La Gomera; UTM: 28RBS7012; 820 m altitude).

(Santana Benítez et al. 2013) Fig. 8D, N. estherae sp. nov. Fig. 7B, N. estherae sp. nov., from La Mérica, La Gomera.

Napaeus magnus Yanes, Deniz, Alonso & Ibáñez

Type material: Holotype (Santana Benítez et al. 2013, Fig. 8G) 1 sh, TFMC (MT 0844); Leg. M. Ibáñez, January 1999 (Fig. 8G).
Type locality: Monte Calvario (La Gomera; UTM: 28RBS7012; 750 m altitude).

(Santana Benítez et al. 2013) Fig. 8G, N. magnus sp. nov. Fig. 9A-C, Genital systems of N. magnus sp. nov. C detail of the penis internal anatomy.

N. bertheloti (L. Pfeiffer 1846) (N. baeticatus group)

(Santana Benítez et al. 2013) Fig. 8A, Shell from Jerduñe, La Gomera. Fig. 9, D, genital system and E detail of the internal anatomy of the epiphallus distal part and the penis proximal part.

Napaeus nanodes (N. nanodes group)

(Yanes et al. 2009, Yanes et al. 2011, Fig. 3B) Shell from Barranco del Agua, Tenerife (AIT).

Napaeus badiosus (N. baeticatus group)

(Yanes et al. 2009, Yanes et al. 2011, Fig. 3C) Shell from Barranco de Valle Seco, Tenerif.

Napaeus obesatus (N. baeticatus group)

(Yanes et al. 2009, Yanes et al. 2011, Fig. 3G) Shell from Barranco del Pagador, Gran Canaria (AIT)

Napaeus chrysaloides (N. variatus group)

(Yanes et al. 2009, Yanes et al. 2011, Fig. 3E) Shell from Tamadaba, Gran Canaria (AIT).

Napaeus osoriensis (N. variatus group)

(Yanes et al. 2009, Yanes et al. 2011, Fig. 3J) Shell from Osorio, Gran Canaria (AIT).

Napaeus myosotis (N. variatus group)

(Yanes et al. 2009, Yanes et al. 2011, Fig. 3L) Shell from Montaña de Guía, Gran Canaria (AIT).

Napaeus exilis (N. variatus group)

(Yanes et al. 2009, Yanes et al. 2011, Fig. 3M) Shell from Altos de Gáldar, Gran Canaria (AIT).

Napaeus moquinianus (N. variatus group)

(Yanes et al. 2009, Yanes et al. 2011, Fig. 3O) Shell from Brezal del Palmital, Gran Canaria (AIT).

Napaeus isletae (N. variatus group)

Holotype. (Yanes et al. 2009, Yanes et al. 2011, Fig. 3P)

Napaeus subsimplex (Wollaston 1878) (N. baeticatus group)

(Santana Benítez et al. 2013, Fig. 8B) Shell from Valle del Cabrito, El Hierro

Napaeus consecoanus (Mousson 1872) (N. baeticatus group)

(Santana Benítez et al. 2013, Fig. 8C) Shell from Túnel de Agulo, La Gomera.

Bulimus savinosa Wollaston 1878

(Santana Benítez et al. 2013, Fig. 8E) NHMUK 1895.2.2.223; photo by Jonathan Ablett.

Bulimus savinosa var. inflatiuscula Wollaston 1878

(Santana Benítez et al. 2013, Fig. 8F) Shell from the Melvill-Tomlin collection (NMW.1955.158.25066; photo by James Turner)

* Napaeus severus* (J. Mabille 1898) (N. baeticatus group)

(Santana Benítez et al. 2013, Fig. 8H) Shell from Hurona, La Gomera.

Napaeus tenoensis Henríquez 1993 (N. baeticatus group)

(Santana Benítez et al. 2013, Fig. 8J) Shell from Las Tablas, Tenerife.

Napaeus baeticatus (Webb & Berthelot 1833) (N. baeticatus group)

(Santana Benítez et al. 2013, Fig. 8K) Shell from Casas de Afur, Tenerife.

Abbreviations of Genitalia

Usual Abbreviation	Name	中文名称
A1-A5	parts of the penial appendix	阴茎牵引器官的不同部分
ag	albumen gland	蛋白腺
bc	bursa copulatrix	交配囊
bd	bursa copulatrix duct	雌雄生殖管
d	diverticulum
dp	distal penis	阴茎远端
e	epiphallus	阳茎基背板
ec	epiphallar caecum	阳茎基背板盲突
f	flagellum	鞭器
go	genital orifice	生殖孔
o	free oviduct	输卵管
p	penis	阴茎
pa	penial papilla
pp	proximal penis	阴茎近端
pr	penis retractor	阴茎牵引肌
pw	penis wall	阴茎壁
r	retractor muscles	牵引肌
t	body tegument
v	vagina	阴道
vd	vas deferens	输精管

Abbreviations of Museum

Abbreviations	Museum
CML	J. Martín private collection, La Laguna, Tenerife, Canary Islands, Spain
FMNH	Field Museum of Natural History, Chicago, Illinois, USA
ICZN	International Commission on Zoological Nomenclature
NHM	Natural History Museum, London, UK
NMW	National Museum of Wales, Cardiff, UK
TFMC	Museo de Ciencias Naturales de Tenerife, Canary Islands, Spain
UTM	Universal Transverse Mercator, cartographic projection system
ZMZ	Zoologisches Museum der Universität, Zürich, Switzerland
AIT	Alonso and Ibáñez collection, Department of Animal Biology, University of La Laguna, Tenerife, Canary Islands, Spain

References

Yanes, Y., J. Barrios, L. Moro, M. Alonso, and M. Ibanez. 2009. On the relationships of the genus Napaeus (Gastropoda: Pulmonata: Enidae) with the descriptions of four new species from the Canary Islands. Journal of Natural History - J NATUR HIST 43: 2179-2207.
Yanes, Y., J. Barrios, J. Delgado, M. Alonso, and M. Ibanez. 2010. Active disguise in land snails: Napaeus badiosus (Gastropoda, Pulmonata, Enidae) from the Canary Islands. Journal of Conchology 40: 143-148.
Yanes, Y. et al. 2011. Five new Napaeus species (Gastropoda: Pulmonata: Enidae) from Gran Canaria and El Hierro (Canary Islands). Zootaxa 2901: 35-51.
Santana Benítez, J. et al. 2013. 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 41: 271-286.

Genus Napaeus