Naesiotus Albers, 1850

Breure, Abraham S. H. & Borrero, Francisco J., 2008, An annotated checklist of the land snail family Orthalicidae (Gastropoda: Pulmonata: Orthalicoidea) in Ecuador, with notes on the distribution of the mainland species, Zootaxa 1768, pp. 1-40 : 10-18

Genus Naesiotus Albers, 1850

Type species: Bulimus nux Broderip, 1832 . Distribution: West Indies, Venezuela, Brazil, Paraguay, Uruguay, Argentina, Bolivia, Peru, Ecuador, Colombia.

Remarks: Although some authors have listed the species of Naesiotus from the Galápagos as Bulimulus (Naesiotus) , we have chosen to follow the classification as given in the most recent revision of the subfamily ( Breure 1979). Most species of the Galápagos are highly endangered and several are considered extinct. See the actual list on http://www.iucnredlist.org (they are all listed under Bulimulus ). There are a number of species described by Coppois (1985), but not according to the ICZN-rules; these are considered to be manuscript names and have not been listed here. Synonyms provided by Richardson (1995) have been mentioned under remarks, although his decisions remain unclear by lack of any explanatory notes. Awaiting a further revision of the Galápagos species we have chosen to give here only the type locality and the depository of the primary type material, if known. The distribution is based on the available data as given by Parent (2003) and Seddon (1996, 2000).

34. Naesiotus achatinellus (Forbes, 1850)

Type locality: Chatham Island. Holotype BMNH 1855.4.5.25.

Distribution: Galápagos, Isla San Cristóbal; Isla Española.

Remarks: According to Köhler (2007: 136) it is unknown on how many specimens the description is based. He stated that Forbes did not explicitly designate a holotype and therefore considered the specimen in the Berlin museum as a syntype. According to the label, the BMNH-specimen is the holotype.

35. Naesiotus adelphus (Dall, 1917)

Type locality: Indefatigable Island. Holotype CAS. Distribution: Galápagos, Isla Santa Cruz.

36. Naesiotus akamatus (Dall, 1917)

Type locality: Indefatigable Island. Holotype CAS. Distribution: Galápagos, Isla Santa Cruz.

37. Naesiotus albemarlensis (Dall, 1917)

Type locality: Galápagos, near Villamil, 700–1000 m. Holotype CAS. Distribution: Galápagos, Isla Isabela, near Puerto Villamil [00° 56' 00" S 091° 01' 00" W].

38. Naesiotus alethorhytidus (Dall, 1917)

Type locality: Indefatigable Island, 105– 120 m. Holotype CAS. Distribution: Galápagos, Isla Santa Cruz.

Distribution: Galápagos, Isla San Cristobal.

40. Naesiotus approximatus (Dall, 1900)

Type locality: Galápagos, Hood Island. Distribution: Galápagos, Isla Española.

41. Naesiotus bauri (Dall, 1893)

Type locality: Galápagos, SW end of Chatham Island. Distribution: Galápagos, Isla San Cristobal.

42. Naesiotus blombergi (Odhner, 1951)

Type locality: Galápagos, Santa Cruz Island, 200– 300m. Holotype NRS. Distribution: Galápagos, Isla Santa Cruz.

Remarks: Richardson (1995) considers Naesiotus deroyi A.G. Smith, 1972 as a synonym.

43. Naesiotus calvus (Sowerby, 1833)

Type locality: Galapagos, James I[sland]. Dall & Ochsner (1928) described the variety caryonis from Charles Island, 230 m. Holotype CAS 1686.

Distribution: Galápagos, Isla San Salvador; Isla Floreana.

45. Naesiotus cavagnaroi A.G. Smith, 1972

Type locality: Galápagos, Isla Santa Cruz, 7 km NE Santa Rosa. Holotype CAS 13738. Distribution: Galápagos, Isla Santa Cruz.

Remarks: Richardson (1995) considers Naesiotus gilderoyi (Van Mol, 1972) as a synonym.

46. Naesiotus chemnitzoides (Forbes, 1850)

Type locality: Galápagos, Chatham Island. Distribution: Galápagos, Isla San Cristobal.

48. Naesiotus cucullinus (Dall, 1917)

Type locality: Galápagos, Hood Island, 60– 180 m. Syntypes CAS. Distribution: Galápagos, Isla Española.

49. Naesiotus curtus (Reibisch, 1892)

Type locality: [Galápagos,] Chatham Island. Distribution: Galápagos, Isla San Cristobal.

50. Naesiotus cymatias (Dall, 1917)

Type locality: Galápagos, Indefatigable Island, 120– 200 m. Syntypes CAS. Distribution: Galápagos, Isla Santa Cruz.

51. Naesiotus darwini (Pfeiffer, 1846)

Type locality: Galapagos Islands.

Distribution: Galápagos, Isla Santiago. According to Parent (2003) five subpopulations are known on the island and total range area is likely to be less than 2000 km 2. The species is found throughout the humid zone on Santiago Island.

53. Naesiotus duncanus (Dall, 1893)

Type locality: Galapagos Island, Duncan Island. Distribution: Galápagos, Isla Pinzón.

55. Naesiotus eos Odhner, 1951

Type locality: Galapagos, Santa Cruz Island. Holotype NRS. Distribution: Galápagos, Isla Santa Cruz.

Remarks: Richardson (1995) considers Naesiotus scalesiana A.G. Smith, 1972 as a synonym.

56. Naesiotus eschariferus (Sowerby, 1833)

Type locality: Galapagos.

Distribution: Galápagos, Isla San Cristóbal. The species is known only from one location with a range size of approximately 3 km 2. The range was probably greater sometime in the past as suggested by the numerous empty shells found on that island (Parent 2003).

57. Naesiotus florschuetzi Breure, 1978

Type locality: Ecuador, Prov. El Oro, 50 km N Santa Rosa. Holotype FLMNH 22772. Distribution: El Oro, 50 km N Santa Rosa ( FLMNH 22772, 22773/45 paratypes; RMNH 55184/10 paratypes)*; Guayas, 4 km N Palmar ( FLMNH 139089)*.

58. Naesiotus galapaganus (Pfeiffer, 1855)

Type locality: Galapagos Islands. Lectotype BMNH 1975146 ( Breure 1979). Distribution: Galápagos, Isla Floreana (see Parent 2003).

60. Naesiotus habeli (Dall, 1892)

Type locality: Galapagos.

Distribution: Galápagos, Isla San Cristóbal (see Parent, 2003).

61. Naesiotus hemaerodes (Dall, 1917)

Type locality: Galapagos, Albemarle Island, Cowley Mountains, 640– 700 m. Syntypes CAS. Distribution: Galápagos, Isla Isabela; Isla Fernandina (see Parent 2003).

62. Naesiotus hirsutus Vagvolgyi, 1977

Type locality: Ecuador, Archipiélago de Colón [Galápagos], Santa Cruz Island, on southern slope, 4 km N Puerto Ayora, 120– 135 m. Holotype USNM 757715.

Distribution: Galápagos, Isla Santa Cruz. It is known from only one population with a range of less 2 km 2. The range was probably greater in the past as suggested by empty shells found on the ground. The total area of suitable habitat available for this species is likely to be less than 100 km 2 (Parent 2003).

63. Naesiotus hoodensis (Dall, 1900)

Type locality: Galapagos, Hood Island. Distribution: Galápagos, Isla Española.

64. Naesiotus indefatigabilis (Dall, 1900)

Type locality: Galapagos, James and Indefatigable Islands.

Distribution: Galápagos, Isla San Salvador. According to Parent (2003) this species is restricted to one small population of Isla Santiago [= San Salvador].

65. Naesiotus jacobi (Sowerby, 1833)

Type locality: Galapagos, James Island. A probable syntype ( ZMB 10283) is figured by Köhler 2007. Distribution: Galápagos, Isla San Salvador. Restricted to two subpopulations, with a range less than 10 km 2, on Isla Santiago [= San Salvador] (Parent 2003).

66. Naesiotus jervisensis (Dall & Ochsner, 1917)

Type locality: Galapagos, Jervis Island, 270– 300 m. Syntypes CAS. Distribution: Galápagos, Isla Rábida.

68. Naesiotus lycodus (Dall, 1917)

Type locality: [Galápagos,] Indefatigable Island, 135– 165 m. Syntypes CAS. Distribution: Galápagos, Isla Santa Cruz.

69. Naesiotus nesioticus (Dall, 1896)

Type locality: [Galápagos,] James Island.

Distribution: Galápagos, Isla San Salvador. Six subpopulations on the island of which the population size fluctuates widely from year to year (Parent 2003).

Remarks: Dall (1920: 121) emended the name to naesioticus.

71. Naesiotus nux (Broderip, 1832)

Type locality: Galapagos, Charles Island. Lectotype BMNH 1975172 ( Breure 1979). Junior synonyms for which type material is known are: Bulimus incrassatus Pfeiffer, 1853 (Lectotype BMNH 1975157; Breure 1979), Bulimus nuciformis Petit, 1853 (Lectotype MNHN; Breure 1976) and Bulimus verrucosus Pfeiffer, 1855 (Lectotype BMNH 1975168; Breure 1979). Dall & Ochsner (1928) described two subspecies for the populations from Isla San Cristóbal: Bulimulus (Naesiotus) nux basiplicatus (Syntypes CAS) and Bulimulus (Naesiotus) nux perchloris (Syntypes CAS). A variety for which no type material is known to exist is Bulimus asperatus Albers, 1857 .

Distribution: Galápagos, Isla San Cristóbal; Isla Floreana. Four subpopulations known, with a total area less than 20 km 2 (Parent 2003).

72. Naesiotus ochseneri (Dall, 1893)

Type locality: [Galápagos,] Indefatigable Island, 60– 200 m. Syntypes CAS. Distribution: Galápagos, Isla Santa Cruz. Only one population known, with a range of less than 10 km 2 (Parent 2003).

73. Naesiotus olla (Dall, 1893)

Dall published it as a new name for Bulimus jacobi Reeve, 1848 not Bulinus jacobi Sowerby, 1833 . Type locality: [Galápagos,] James Island.

Distribution: Galápagos, Isla San Salvador. Two subpopulations, each of less than 200 individuals, with a total area of occupancy probably less than 5 km 2 (Parent 2003).

74. Naesiotus pallidus (Reibisch, 1892)

Type locality: [Galápagos,] Albemarle Island. Lectotype ZMB 47951a (designated and figured by Köhler 2007: 138, Fig. 54).

Distribution: Galápagos, Isla Isabela; Isla Pinta (Parent 2003).

Remarks: Considered by Richardson (1995) to be a synonym of Naesiotus jacobi (Reeve, 1848) .

75. Naesiotus perrus (Dall, 1917)

Type locality: [Galápagos,] Narborough Island, rim of the crater, 60– 150 m. Syntypes CAS. Distribution: Galápagos, Isla Fernandina. Single population at the summit of the island (Parent 2003).

76. Naesiotus perspectivus (Pfeiffer, 1846)

Type locality: Locality unknown. Lectotype BMNH 1975166 ( Breure 1979); “Galapagos” as locality mentioned on label.

Distribution: Galápagos, Isla San Cristóbal; Isla Floreana.

77. Naesiotus pinzonensis Vagvolgyi, 1977

Type locality: Archipiélago de Colón [Galápagos], Pinzón Island, northern and western cliffs of main peak, 425– 455 m. Holotype USNM 757718.

Distribution: Galápagos, Isla Pinzón.

78. Naesiotus pinzonopsis Vagvolgyi, 1977

Type locality: Archipiélago de Colón [Galápagos], Pinzón Island, northern and western cliffs of main peak, 425– 455 m. Holotype USNM 757719.

Distribution: Galápagos, Isla Pinzón.

80. Naesiotus prepinguis Vagvolgyi, 1977

Type locality: Archipiélago de Colón [Galápagos], Pinzón Island, secondary peak, 263 m, north of the crater. Holotype USNM 757720.

Distribution: Galápagos, Isla Pinzón.

81. Naesiotus quitensis (Pfeiffer, 1848)

Type locality: Quito. Lectotype BMNH 1975320 ( Breure 1979). Bulimus catlowiae Pfeiffer, 1853 is a junior synonym of this species; the lectotype is also in the BMNH-collection ( BMNH 1975414). Other synonyms are Bulimus irregularis Pfeiffer, 1848 , also from Quito (syn. nov.) and Bulimus caliginosus Reeve, 1849 (syn. nov.).

Distribution: Carchi, 35 km N Ibarra on the road to El Angel, 1700 m ( RMNH)*; Chimborazo, Riobamba ( FLMNH); S of Cajabamba ( FLMNH 176545)*; Cerro de Altar ( Pilsbry 1897: 33); Imbabura, 9.4 km N Ibarra ( FLMNH 189112, 189113)*; Otavalo (Pilsbry 1896: 159); Cerro Imbabura; Chota ( MCZ 113825);?”Salinas Ibarre, 1659 m, coll. Stübel” ( Pilsbry 1897: 34); Napo , slopes of Mt. Antisana ( FMNH 70905, 107553); Pastaza, Cerros de Abitagua ( FMNH 125462);?Río Pastaza, 3.2 km above Falls Agougou ( ZMH 7373);?banks of Río Pastaza ( ZMH 7374) [both localities not located in the GNS gazetteer]; Pichincha, “near Quito” ( BMNH 1975321/1 paralectotype of Bulimus catlowiae Pfeiffer ). Quito ( CMC C11759, as B. catlowiae Pfeiffer ; FLMNH; MCZ 0 74513, 140390; ZMZ 512411); 14.5 km NW Quito ( FMNH 146936, RMNH); slopes Cerro Pichincha ( FMNH 86651, 86652, 125440, 125442, 125444, 125464, 158902; MCZ 0 65114, 0 65115, 118581); Pomasqui ( FLMNH); 13.9 km W Pifo ( FLMNH 37396)*; Cayambe ( FLMNH 37394, 37395)*; Tumbaco; Guaillabamba (both Pilsbry, 1897: 34); 1 km W Mitad del Mundo, 2400 m ( RMNH)*; 22.6 km NNE Quito at the road to Cayambe, 2200 m ( RMNH)*; Chimba, 3 km E Olmedo ( FMNH 223385, 223386, 223387)*; Cumbayá, 9.6 km NE Quito ( MCZ 083922); above Chillogallo ( Pilsbry 1897: 33); Tungurahua, 8 km N San Antonio ( FLMNH 39904)*; Ambato ( Pilsbry 1897: 35; MCZ 045823); Cerro Cachigani near Ambato ( FMNH 216992); Píllaro ( FLMNH; FMNH 86654, 86686; MCZ 118583); Puñapi ( MCZ 155442); Baños ( FMNH 125482, 126162, 125700, 126207, 216987; MCZ 115666); Río Agoyán falls ( FMNH 72346, 125472, 126183, 126239; MCZ 173383).

Remarks: This is a rather widespread and quite variable species, for which several subspecies have been described:

Naesiotus quitensis ambatensis Rehder, 1940

Type locality: Tunguragua, Ambato. Holotype: USNM 473973, paratypes MCZ 0 70604.

Naesiotus quitensis antisana Rehder, 1942

Type locality: Napo , slopes of Mt. Antisana. Holotype USNM 516940, paratypes FMNH 86659, 125369, 125371, 126188, 126988.

Naesiotus quitensis jacksoni Rehder, 1940

Type locality: Pichincha, Guaillabamba. Holotype USNM 473969, paratypes FMNH 86661, 158913, 306498.? Paratypes MCZ 118582.

Naesiotus quitensis orinus Rehder, 1940

Type locality: Chimborazo, near Riobamba, 3650 m. Holotype USNM 473971, paratypes FMNH 86660, 125402, 126268, 158871.? Paratypes MCZ 118580.

Naesiotus quitensis rufescens (Germain, 1910)

Type locality: Pichincha, Tumbaco. It is also mentioned from “Cujuja”, which we have been unable to locate.

Naesiotus quitensis vermiculatus Rehder, 1940

Type locality: Tungurahua, Agoyán. Holotype MCZ 112473, paratypes FMNH 126269, MCZ 0 64957.

82. Naesiotus rabidensis (Dall, 1917)

Type locality: [Galápagos,] Jervis Island, 275– 300 m. Syntypes CAS. Distribution: Galápagos, Isla Rábida.

84. Naesiotus rugatinus (Dall, 1917)

New name for Bulimulus acutus Reibisch, 1892 not Leach, 1814. Type locality: [Galápagos,] Chatham Island. Lectotype ZMB 47964a (designated and figured by Köhler 2007: 136, Fig. 44). Distribution: Galápagos, Isla San Cristóbal; Isla Floreana (Parent 2003).

85. Naesiotus rugiferus (Sowerby, 1833)

Type locality: [Galápagos,] James Island. Lectotype BMNH 1975178 ( Breure 1979). Distribution: Galápagos, Isla Santa Cruz; San Salvador (Parent 2003). Remarks: Köhler (2007) figures a probable paralectotype.

87. Naesiotus saeronius (Dall, 1917)

Type locality: [Galápagos,] Indefatigable Island. Holotype USNM 274097.

Distribution: Galápagos, Isla Santa Cruz.

Remark: Parent (2003) mentions this species as being a synonym of Naesiotus rugiferus (Sowerby, 1833) in the summary, despite that otherwise she treats saeronius as a distinct species.

88. Naesiotus sculpturatus (Pfeiffer, 1846)

Type locality: Galapagos. Lectotype BMNH 1975174 ( Breure 1979). Distribution: Galápagos, Isla San Salvador; it is only known from one population found in a small area near a natural fresh water source. There are probably no more than 200 individuals on a total range less than 1 km 2 (Parent 2003).

89. Naesiotus simrothi (Reibisch, 1892)

Type locality: [Galápagos,] Albemarle Island.

Distribution: Galápagos, Isla Isabela.

90. Naesiotus snodgrassi (Dall, 1900)

Type locality: [Galápagos,] Hood Island. Distribution: Galápagos, Isla Española.

91. Naesiotus steadmani Chambers, 1986

Type locality: [Galápagos,] Santa Cruz, Cueva de Kubler. Holotype CAS 59358. Distribution: Galápagos, Isla Santa Cruz.

92. Naesiotus tanneri (Dall, 1895)

Type locality: [Galápagos,] Indefatigable Island.

Distribution: Galápagos, Isla Santa Cruz. May already be extinct (Parent 2003).

Naesiotus tanneri bartolomensis Vagvolgyi, 1977

Type locality: [Galápagos], off Santiago Island, Bartolomé Island, at western foot of main peak. Holotype USNM 757717.

Naesiotus tanneri edenensis Vagvolgyi, 1977

Type locality: [Galápagos], off Santa Cruz Island, Eden Island, on southern slope, 30– 45 m. Holotype USNM 757716.

93. Naesiotus tortuganus (Dall, 1893)

Type locality: [Galápagos,] South Albemarle Island, La Tortuga. Distribution: Galápagos, Isla Isabela. It is known from only one population at the top of Darwin Volcano (Parent 2003).

95. Naesiotus unifasciatus (Sowerby, 1833)

Type locality: [Galápagos,] Charles Island. Lectotype BMNH 1975187 ( Breure 1979). Distribution: Galápagos, no further data known (see Seddon 2000).

96. Naesiotus ustulatus (Sowerby, 1833)

Type locality: [Galápagos,] Charles Island. A probable syntype is in the Berlin museum ( ZMB 10239) and figured by Köhler 2007. Another syntype, originating from the Cuming collection, is in the Hamburg museum and labelled “Galapagos” ( ZMH 7377). Syntypes of Bulimulus (Naesiotus) ustulatus pallescens Dall & Ochsner, 1928 are in the CAS collection.

Distribution: Galápagos, no further data known (see Seddon 1996).

98. Naesiotus wolfi (Reibisch, 1892)

Type locality: [Galápagos,] Indefatigable Island. Lectotype ZMB 47950 (designated and figured by Köhler 2007: 140, Fig. 59)

Distribution: Galápagos, Isla Santa Cruz. It is known from one population with a very restricted distribution (less than 2 km 2). Population size is no more than 500 indivduals (Parent 2003).

Remarks: Richardson (1995) considers this taxon to be synonymous with Naesiotus reibischii (Dall, 1895) .