Zootaxa, an Annotated Checklist of the Land Snail Family Orthalicidae

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An annotated checklist of the land snail family Orthalicidae (Gastropoda: Pulmonata: Orthalicoidea) in Ecuador, with notes on the distribution of the mainland species

ABRAHAM S.H. BREURE¹ & FRANCISCO J. BORRERO²,³ ¹National Museum of Natural History / Naturalis, P.O.Box 9517, Leiden, the Netherlands; E-mail: [email protected] ²Cincinnati Museum Center, 1301 Western Avenue, Cincinnati, Ohio 45203, USA; ³Cincinnati Country Day School. E-mail: [email protected]

Table of contents

Abstract ...... 1 Introduction ...... 2 Methods ...... 3 Checklist of species ...... 4 SUBFAMILIA BULIMULINAE ...... 5 Genus Plekocheilus Guilding, 1828 ...... 6 Genus Thaumastus Albers, 1860 ...... 8 Genus Bostryx Troschel, 1847 ...... 10 Genus Bulimulus Leach, 1814 ...... 10 Genus Naesiotus Albers, 1850 ...... 11 Genus Scutalus Albers, 1850 ...... 19 Genus Stenostylus Pilsbry, 1898 ...... 19 Genus Drymaeus Albers, 1850 ...... 20 Genus Simpulopsis Beck, 1837 ...... 26 SUBFAMILIA ORTHALICINAE ...... 26 Genus Sultana Shuttleworth, 1856 ...... 26 Genus Orthalicus Beck, 1837 ...... 27 Genus Corona Albers, 1850 ...... 28 Genus Porphyrobaphe Shuttleworth, 1856 ...... 29 Genus Hemibulimus Martens, 1885 ...... 30 Incertae sedis ...... 30 Discussion ...... 31 Distribution of genera—modelled by Maxent software ...... 32 Directions for further research ...... 37 Acknowledgements ...... 38 Literature cited ...... 38

Abstract

The family Orthalicidae is wide-spread in the Neotropics and a major member of the land snail fauna of most South American countries. This checklist presents the species known to exist in Ecuador. It is based on data from relevant literature and museums world wide. There are 168 species in total, 63 of which occur on the Galápagos; 9 species are listed for the first time from this country. New synonymies are: Bulimus abscissus Pfeiffer, 1855 = Drymaeus (D.) fallax Pfe-

Accepted by B. Ruthensteiner: 4 Apr. 2008; published: 14 May 2008 1 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. iffer, 1853; Bulimus caliginosus Reeve, 1849 = Bulimus irregularis Pfeiffer, 1848 = Naesiotus quitensis (Pfeiffer, 1848). In the geographical analysis only the mainland species have been taken into account. The species were sampled at 128 mainland localities, 46 of which have been sampled after 1950, indicating that some parts of the country are undersampled. The potential distribution has been analysed at (sub-)generic level, using Maxent and GIS software. Finally four directions for future research are suggested: strengthening the locality database, improved sampling for better modelling, analysis of models on a greater spatial scale and phylogeographic analyses.

Key words: Mollusca, Bulimulinae, Orthalicinae, systematics, new synonym, endemism, distribution modelling.

Introduction

With the world wide decline of floras and faunas, especially in the tropics, biodiversity research has gained more interest in recent years. Land snails are relatively poorly represented in this type of research. For a better understanding of overall patterns of biodiversity, rates of extinction, areas of endemism and conservation issues would need more attention of malacologists. Malacological papers dealing with the Neotropics mainly concern taxonomy at species level and there are very few comprehensive works that cover higher taxa or country levels. A positive exception is the recent work of Simone (2006) for Brazil. Of the 612 land snails listed for this country, 305 belong to the family Orthalicidae (nearly 50%). Similar rates are known or expected for other South American countries, making this family a major component of the land snail fauna of this continent. The family is relatively well studied at the genus level (Breure 1979 and references therein), but is in need of phylogenetic studies to clarify the relationships between the subfamilies hitherto recognized. Within South America, Ecuador is generally recognized as one of the countries with a high biodiversity (Myers et al. 2000, Brooks et al. 2006). The country has 22 provinces (including the Galápagos Islands), cov- ering a total area of 269.939 km2. The mainland is divided into three regions: 1) La Costa, the low-lying coastal plain in the western part of the country; 2) La Sierra, the high-altitude belt running north to south along the center of the country is part of the Andes mountain range; and 3) El Oriente, the Amazon rainforest areas in the eastern part of the country (Fig. 1A). This checklist enumerates the taxa known to (possibly) exist in Ecuador belonging to the land snail family Orthalicidae Albers, 1860 [= Familia Bulimulidae Crosse & Fischer, 1873 (sensu lato) = Superfamilia Orthal- icoidea Martens, 1860]. The list is based on the taxa mentioned in Breure (1979), on unpublished data from several collections—either personally checked or based on available databases—and on data taken from publications. All taxa are attributed to species following the works of Pilsbry and recent authors. Species are clas- sified in (sub-)genera following the last revision for the subfamily Bulimulinae Tryon, 1867 (Breure 1979) and that of Zilch (1959–1960) for the subfamily Orthalicinae Thiele, 1931. The classification at (sub-)familiar level follows Bouchet & Rocroi (2005). A critical review of this list will undoubtedly lead to synonymization of several taxa, thus this document can also be viewed as a longlist of the representatives of this family in the Ecuadorian malacofauna. The land snail fauna of Ecuador has been sampled mainly during the 19th and 20th centuries. Old localities on the mainland were predominantly in the “Valley of the Volcans” and less often in the coastal area or in the eastern part of the country. The only known comprehensive survey of the continental Ecuadorian malacofauna is the one by Cousin (1887). He lists 96 taxa of what is here considered to be Orthalicidae. Over the past 50 years collections on the mainland were mainly made by K. Campell, R.W. McDiarmid, F.G. Thompson and N.H. Williams, who deposited their material all in museums in the USA. The only known active Ecuadorian malacologist today is M. Correoso from Quito (Correoso 2002). The Galápagos Islands are a special case. Famous for their unique fauna which inspired Charles Darwin’s “Origin of species”, its land snails belong mainly to the Bulimulinae. They have not been listed before. These snails are resulting from a remarkable radiation which has led to many species occupying a niche habitat on one or a few islands. Many of these are now on the IUCN Red List of endangered species (see Parent 2003).

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Species distribution modelling has developed quite rapidly over the past 10 years, providing a useful tool in a range of fields including biogeography, ecology, palaeo-ecology and conservation biology. Also the link between phylogenetics and environmental niche models has been explored (e.g. Graham et al. 2004a, b). Examples with malacological applications, however, are scarce (Hugall et al. 2002) and there are several challenges for further development of the methodology (Araujo & Guissan 2006). There is a range of models available for use with presence-only data. Elith et al. (2006) have evaluated 16 approaches using test data from different regions and biological groups. In their study the Maxent software model performed better than well-known methods like BIOCLIM (Nix 1986; Busby 1991; implemented in DIVA-GIS) and GARP (Stock- well & Peters 1999). This was corroborated by Hernandez et al. (2006), who compared four models that use presence occurrence data and produce distribution models without explicit quantification of absence data. Although there are some methodological caveats when only limited distributional information is available, using environmental niche models may be suitable to identify regions that have similar environmental conditions to where species are known to occur (Pearson et al. 2007).

Methods

For each species the locality is given as stated in the original publication. As usual with records from the 19th century the locality is often very imprecise (“Quito”, “Andes of New Grenada”, etc.). If a more precise inter- pretation could be given, it is added between square brackets. The distribution is given with data from collections or publications, arranged according to the provinces (bold in text) in alphabetical order. The type locality is added when available, derived from different sources, and references have been added when types have been designated or figured by modern authors. Where data from museum collections are available they have been mentioned (with the exception of species from the Galápagos, see also remarks on Naesiotus). In general we have followed the identifications given on the labels and in the databases (museums) that have been con- sulted, unless it was clearly a misidentification. Doubtful identifications are mentioned under “remarks”. Those species that are known from specific localities (mainland only) have been analyzed according to the occurrence in ecoregions (see Tables 2 and 4). To explore possible threats, endemism has been indicated as well as the occurrence at a single locality. It must be noted, however, that more sampling may reveal that some species actually occupy a broader range of ecoregions than is evident here. Species used for distributional analyses, are listed in Table 4. The altitudinal range and the ecoregion(s) (National Geographic Society 2008; Fig. 1B) to which the species can be attributed are also provided therein. Maxent software has been applied to produce maps showing areas where species are likely to occur. Geo- referenced locality data with longitude and latitude coordinates derived from the GNS website (National Geospatial-Intelligence Agency 2007), were used as input in Maxent 3.0.8 (Phillips et al. 2007), together with environmental data layers. Given the limited number of localities for the various species, the focus of analysis is at the generic level unless stated otherwise. The results (Fig. 2) are expressed as logistic values, represented by colours. These maps suggest avenues for additional research, while at the same time it may be noted that explorations of potential distributions at the species level are currently not possible due to the very incomplete data available. As default settings in Maxent software the following variables were used: annual diurnal temperature range; annual frost frequency; January, April, July, October and annual precipitation; minimum, maximum and mean annual temperature; annual vapor pressure; elevation and ecoregions (all based on data as resam- pled by Phillips et al. 2006, who also discusses the use of environmental variables). Models were generated using the following settings: random test percentage 25, regularization multiplier 1, maximum iterations 500 and convergence treshold 0.00001 (see also Phillips et al. 2006). The resulting distribution was a grid file with logistic values, used as input for DIVA-GIS 5.4 software (Hijmans et al. 2007).

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Within DIVA-GIS a multi-layered data analysis was performed combining the grid file output from Max- ent as basis, plus shape files of the provinces of Ecuador and the relevant localities of species. The result was viewed using the Design tab of DIVA-GIS and saved as a tiff-file for further processing.

FIGURE 1. Maps of Ecuador. A. Altitudinal map. B. Ecoregions of Ecuador according to National Geographic Society (2008). NT0115: Chocó-Darién moist forests; NT0121: Eastern Cordillera real montane forests; NT0142: Napo moist forests; NT0145: Northwestern Andean montane forests; NT0178: Western Ecuador moist forests; NT0214: Ecuadorian dry forests; NT0232: Tumbes-Piura dry forests; NT0905: Guayaquil flooded grasslands; NT1004: Cordillera Central páramo; NT1006: Northern Andean páramo; NT1405: Belizean Coast mangroves. C. Localities of endemic, range- restricted species; see text for further explanation.

The following abbreviations have been used to indicate the museum collections: ANSP—Academy of Natural Sciences, Philadelphia, USA; BMNH—The Natural History Museum, London, UK; CAS—Califor- nia Academy of Sciences, Dept. Geology, San Francisco, USA; CMC—Cincinnati Museum Center, Malacol- ogy collection, Cincinnati; CMNH—Carnegie Museum of Natural History, Pittsburgh, USA; EPN—Escuela Politécnica Nacional, Quito, Ecuador; FLMNH—Florida Museum of Natural History, Gainesville, USA; FMNH—Field Museum of Natural History, Chicago, USA; HMNH—Hungarian Museum of Natural History, Budapest, Hungary; IML—Instituto Miguel Lillo, Tucumán, Argentina; IRSN—Institut Royal des Sciences Naturales, Brussels, Belgium; MCZ—Museum of Comparative Zoology, Cambridge (Mass.), USA; MNHN—Muséum National d’Histoire Naturelle, Paris, France; NRS—Naturhistoriska Riksmuseet, Stock- holm, Sweden; RMNH—Nationaal Natuurhistorisch Museum / Naturalis, Leiden, the Netherlands; SMF— Natur-Museum Senckenberg, Frankfurt am Main, Germany; USNM—National Museum of Natural History, Washington, USA; UVZ—Universidad del Valle, Zoología, Cali, Colombia; ZMA—Zoölogisch Museum, Amsterdam, the Netherlands; ZMB—Museum für Naturkunde, Humboldt Universität, Berlin, Germany; ZMH—Zoologisches Museum der Universität Hamburg, Hamburg, Germany; ZMZ—Zoologisches Museum, Zürich, Switserland; ZSM—Zoologische Staatssammlung München, Germany.

Checklist of species

The list gives the current classification of the taxa that are mentioned in all known literature on Ecuadorian Orthalicoids. Subspecies are not numbered separately but are listed under the nominal species. New records for Ecuador are indicated by ** after the species name. An asterisk (*) has been added if the material has been collected after 1950. In species distributions, we have used a question mark (?) anterior to the locality to indicate records based solely on citations from the literature, but for which there is no supporting voucher material. Imprecise locality records are given between quotation marks (“”) to indicate text taken from labels.

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SUBFAMILIA BULIMULINAE

Remarks: For a key of the genera and a bibliography, see Breure 1979.

Genus Plekocheilus Guilding, 1828

Subgenus Plekocheilus (Plekocheilus) Guilding, 1828 Type species: Voluta aurissileni Born, 1780. Distribution: West Indies (St. Vincent, Barbados), Venezuela, Ecuador, Colombia.

1. Plekocheilus (Plekocheilus) cardinalis (Pfeiffer, 1853) Type locality: Quito. Lectotype ZMB 112721 (designated and figured by Köhler 2007). Distribution: ?Napo (Pilsbry 1899: 77); Pastaza, Mera (FLMNH 109841; FMNH 86698, 125478); Pichincha, near Mindo, circa 1000 m (Weyrauch 1967: 460); Nanegal; valley of Río Pilatón (both according to Pilsbry 1899: 77). “New Grenada” (RMNH). “Ecuador” (ZMA). Remarks: New Grenada was used in the 19th century and is referring to an area comprising Ecuador, Colombia and parts of Venezuela and Panama.

Subgenus Plekocheilus (Eurytus) Albers, 1850 Type species: Helix pentadina d’Orbigny, 1835. Distribution: West Indies (St. Lucia, St. Vincent), Vene- zuela, Brazil, Bolivia, Peru, Ecuador, Colombia.

2. Plekocheilus (Eurytus) aristaceus (Crosse, 1869) Type locality: Quito, reipublicae Aequatoris. Distribution: Chimborazo, Bucay (IRSN); Cotopaxi, Páramo de Sigchos (IRSN); Pichincha, near Mindo (FMNH 216871)*; Río Pilatón (IRSN). “New Grenada” (ZMB). Remarks: These are the first precise localities for this species.

3. Plekocheilus (Eurytus) aureonitens (Miller, 1878) Type locality: Pilatón valley, 1000 m. Distribution: Pichincha, Río Pilatón. “Ecuador” (ANSP 78577). Remarks: This species may prove to be a synonym of Plekocheilus (Eurytus) taylorianus (Reeve, 1849).

4. Plekocheilus (Eurytus) corydon (Crosse, 1869) Type locality: Quito. Distribution: ? [no specific locality known].

5. Plekocheilus (Eurytus) doliarius (Da Costa, 1898) Type locality: Paramba. Lectotype BMNH 1907.11.21.110 (Breure 1979; see also Neubert & Janssen 2004: 208, who figure a paralectotype (SMF 9513) in Pl. 1 Fig. 1). Distribution: Carchi, Hacienda Paramba (BMNH, IRSN, SMF). Remarks: As this species is also recorded from Colombia, the locality is most likely in Carchi, which is also the only province where Paramba is attributed to in the GNS-gazetteer.

6. Plekocheilus (Eurytus) eros (Angas, 1878) Type locality: Ecuador. Holotype BMNH 1879.1.21.2. Distribution: ? [no specific locality known].

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7. Plekocheilus (Eurytus) floccosus (Spix, 1827) Type locality: Brazil. Holotype ZSM (Simone 2006: 149). Distribution: Morona-Santiago, Cusuimi (FMNH 170620); Río Macuma, ca. 10 km from Río Morona (FMNH 187606)*; Napo, Río Napo, 500 m (SMF 156306; see Weyrauch 1967: 462); Tungurahua, Topo (FMNH 86667); 0.7 km W Río Negro (RMNH)*. “Ecuador” (RMNH). Remarks: This is a broad ranged species that is also known from Peru, Brazil and Bolivia. Weyrauch (1967: 462) placed Helix pentadina d’Orbigny, 1835 in the synonymy of this species.

8. Plekocheilus (Eurytus) jimenezi (Hidalgo, 1872) Type locality: San José (according to Weyrauch 1967: 463 this is located 50 km E Baeza; it is most probably San José de Suno [00° 32' 00" S 077° 25' 00" W], also known as San José Viejo). Plekocheilus (Eurytus) gibbonius (Hidalgo, 1872), described from the same locality, is a synonym according to Pilsbry (1895: 87). Distribution: Orellana, San José de Suno. “Oriente Prov.” (FMNH 72336).

Plekocheilus (Eurytus) jimenezi oligostylus Pilsbry, 1939 Type locality: [Ecuador, between Baeza and Archidona, Nachiyacu, see Clench & Turner 1962: 109]. Holotype: ANSP 170696. Distribution: Napo, Nachiyacu; Sarayacu (see Weyrauch 1967: 463); valley of Río Quijos (FMNH 216872); Pastaza, Puyo (FLMNH 109855; FMNH 42168, 126934); Mera (FLMNH 184702).

9. Plekocheilus (Eurytus) lynciculus (Deville & Hupé, 1850) Type locality: Peru, Loreto, Sarayacu. Distribution: Napo, Nachiyacu (ANSP 170694: Holotype of the synonym Plekocheilus (Eurytus) jacksoni Pilsbry, 1939; according to Weyrauch 1967: 463); Orellana, Loreto, 550 m [SMF 156302; see Weyrauch 1967: 463]; Pastaza, Chichirota (FMNH 78742).

10. Plekocheilus (Eurytus) mcgintyi ‘Pilsbry’ H.B. Baker, 1963 Type locality: Rio Napo, northeastern boundary of Ecuador. Holotype ANSP 227455. Distribution: Napo, Río Jatunyacu [= Río Napo] (ANSP).

11. Plekocheilus (Eurytus) nocturnus Pilsbry, 1939 Type locality: Ecuador, Puyo. Holotype ANSP 170695. Distribution: Pastaza, Puyo (FMNH 86644).

12. Plekocheilus (Eurytus) phoebus (Pfeiffer, 1863) Type locality: Ecuador. Lectotype BMNH 1975143 (Breure 1979). Distribution: ? [no specific locality known].

13. Plekocheilus (Eurytus) pulicarius (Reeve, 1848) Type locality: New Grenada. Lectotype BMNH 1975281 (designated and figured by Breure 1978: 15, Pl. 11 Fig. 6). Distribution: ? [no specific Ecuadorian locality known; the species is better known in Colombia]. “Ecua- dor” (FMNH 202082). “New Grenada” (BMNH 1975282/2 paralectotypes).

14. Plekocheilus (Eurytus) roseolabrum (E.A. Smith, 1877) Type locality: Malacatos. Lectotype BMNH 1975135 (Breure 1978). Distribution: Loja, Malacatos (BMNH 1877.3.28.2/1 paralectotype); Zamora-Chinchipe, Tapichalaca*

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(B. Harris, personal communication and photograph). Remarks: This is the first record and precise locality after the original description in 1877.

15. Plekocheilus (Eurytus) taylorianus (Reeve, 1849) Type locality: Environs of Quito. Lectotype BMNH 1874.12.11.271 (Here corrected for BMNH 1975142 as given by Breure 1978). Pilsbry (1895) placed Eurytus taylorioides Miller, 1878 in the synonymy of this species; its type locality is: “circa Quito, Chimborazo”. Distribution: Pastaza, Mera (FMNH 202059); Pichincha, Milpe (FLMNH 166225); Mindo, near Rio Blanco (FLMNH 166226); Nanegal (EPN); Pacto (EPN); Pintag (EPN); Gualea (FMNH 61138, 31173). “Ecuador” (CMC C11056, C11057; FMNH 100722; RMNH; ZMA). Remarks: Köhler (2007) gives a figure of a paralectotype (ZMB 112722).

16. Plekocheilus (Eurytus) tricolor (Pfeiffer, 1853) Type locality: New Grenada, Gualea. According to Pilsbry (1901: 131) Bulimus semipictus Hidalgo, 1869 is a junior synonym is, for which the type locality is Baeza. Distribution: Cotopaxi, Sigchos (FMNH 70912); Imbabura, Ibarra (FMNH 70911); Napo, Baeza; Orel- lana, Ávila (FMNH 31171); Pichincha, Gualea. “Ecuador” (RMNH).

Subgenus Plekocheilus (Aeropictus) Weyrauch, 1967 Type species: Bulimus veranyi Pfeiffer, 1848. Distribution: Venezuela, Brazil, Ecuador, Colombia.

17. Plekocheilus (Aeropictus) tenuissimus (Weyrauch, 1967) Type locality: 20 km W Quito, Tandayapa, 2500 m. Holotype IML 3364. Distribution: Pichincha, Tandayapa* (IML); near Nanegal (RMNH)*.

Plekocheilus sp. Distribution: El Oro, 13.1 km WNW Pinas (FLMNH 36133)*; Napo, Archidona (FMNH 31133), labelled as P. blainvilleanus loveni (Pfeiffer), which is a Venezuelan species; Pichincha, 15 km NE Las Pal- mas, Río Faisanes (FLMNH 22917)*; 14.3 km NE La Palma, Río Faisanes (FLMNH 26611)*.

Genus Thaumastus Albers, 1860

Subgenus Thaumastus (Kara) Strebel, 1910 Type species: Bulimus thompsonii Pfeiffer, 1845. Distribution: Venezuela, Brazil, Bolivia, Peru, Ecuador.

18. Thaumastus (Kara) thompsonii (Pfeiffer, 1845) Type locality: Quito. Lectotype BMNH 1975464. Junior synonyms of this species are Orphnus thompsoni nigricans Cousin, 1887 (type locality: Cuenca), Orphnus thompsoni olivacea Cousin, 1887 (type locality: Cuenca) and Orphnus thompsoni zebra Cousin, 1887 (type locality: near Azogues). Distribution: Azuay, Cuenca (ANSP 168228; CMC C11040; EPN; FMNH 78736); Cañar, near Azogues; Pichincha, Nanegal (ZMB). “Ecuador” (CMC C11000, C11041, C11126; RMNH; ZMA; ZMH 7528). “Quito” (BMNH 1975465/2 paralectotypes; FMNH 103304).

19. Thaumastus (Kara) thompsonoides Oberwimmer, 1931 Type locality: Ecuador, Loja. Holotype SMF 5144. Distribution: Loja, ? [no specific locality known].

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Subgenus Thaumastus (Scholvienia) Strebel, 1910 Type species: Bulimus bitaeniatus Nyst, 1845. Distribution: Peru, Ecuador.

20. Thaumastus (Scholvienia) jaspideus (Morelet, 1863)** Type locality: Peru, Huancavelica. Distribution: Tungurahua, Baños (FLMNH 109807). Remarks: This is the first record—if correctly identified—from outside the known distribution in Peru.

Subgenus Thaumastus (Thaumastiella) Weyrauch, 1956 Type species: Bulimulus sarcochrous Pilsbry, 1897. Distribution: Peru, Ecuador.

21. Thaumastus (Thaumastiella) sarcochrous (Pilsbry, 1897)** Type locality: Peru. Holotype ANSP 4705. Distribution: Pichincha, 24 km N Quito (FMNH 106151, 106152). Remarks: This subgenus is known from northern Peru (Depts. La Libertad, Cajamarca) and if the specimens have been correctly identified, this is an extension of the known range into Ecuador.

Subgenus Thaumastus (Thaumastus) Albers, 1860 Type species: Bulimus hartwegi Pfeiffer, 1846. Distribution: Brazil, Bolivia, Peru, Ecuador.

22. Thaumastus (Thaumastus) brunneus Strebel, 1910 Type locality: Ecuador. Distribution: ? [no specific locality known].

23. Thaumastus (Thaumastus) buckleyi (Higgins, 1872) Type locality: San Lucas. Syntypes BMNH 1872.5.22.6. Distribution: Loja, San Lucas (BMNH). “Ecuador” (ANSP 81285).

24. Thaumastus (Thaumastus) flori (Jousseaume, 1897) Type locality: Machala. Distribution: El Oro, Machala; Pichincha, Nanegal (see Weyrauch 1967: 472).

25. Thaumastus (Thaumastus) hartwegi (Pfeiffer, 1846) Type locality: “El Catamaija” near Loja. Syntype BMNH 1975126. Distribution: Loja, Catamayo (BMNH); Pichincha, probably vicinity of Nanegal (ANSP 195217). “Ecuador” (CMC C11020; RMNH).

26. Thaumastus (Thaumastus) indentatus (Da Costa, 1901) Type locality: Ecuador. Holotype BMNH 1907.11.21.115. Distribution: ? [no specific locality known]. “Ecuador” (BMNH 1907.11.21.116/1 paralectotype).

27. Thaumastus (Thaumastus) integer (Pfeiffer, 1855) Type locality: Quito. Lectotype BMNH 1975244 (Breure 1979). Distribution: ? [no specific locality known]. “Quito” (BMNH 1975245/1 paralectotype).

28. Thaumastus (Thaumastus) loxostomus (Pfeiffer, 1853) Type locality: Andes of New Granada [Judging by the name this taxon may originate from the province of Loja].

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Distribution: ?Loja [no specific locality known].

29. Thaumastus (Thaumastus) orcesi Weyrauch, 1967 Type locality: Ecuador, basin of río Esmeraldas, 35 km NW Quito, region of Nanegal, 1500 m. Holotype IML 3165. Distribution: Pichincha, Nanegal (IML).

Thaumastus sp. Distribution: Zamora-Chinchipe, road to Valladolid (FLMNH 157166)*; Tapichalaca* (B. Harris, personal communication and photograph).

Genus Bostryx Troschel, 1847 Type species: Bulimus (Bostryx) solutus Troschel, 1847. Distribution: Venezuela, Argentina, Bolivia, Chile, Peru, Ecuador.

30. Bostryx bilineatus (Sowerby, 1833) Type locality: Santa Elena and in western Colombia [Although there are more places with this name, we suppose that here is meant Santa Elena in Prov. Guayas]. Syntypes ZMB 10261 (Köhler 2007: 131, Fig. 21). Distribution: Guayas, 17 km WNW Progreso [= Gómes Rendón] (FLMNH 139140)*; 11 km W Progreso (FLMNH 139079)*; 1 km ENE Progreso (FLMNH 139047)*; 5 km NNE Progreso (FLMNH 139049)*; 4 km N Palmar (FLMNH 139078)*; 5 km N Palmar (FLMNH 139056)*; 4 km N Monteverde (FLMNH 139077)*; 3 km E Zapotal (FLMNH 139106)*; 9 km S Manglaralto (FLMNH 139114)*; Santa Elena (RMNH); 11.2 km NE Santa Elena (FMNH 106230)*; 32 km NE Santa Elena (FMNH 106153)*;. Manabí, Manta (FMNH 125507, 125216); 11 km WNW Jipijapa (FLMNH 139038)*; 1 km S Ayampe (FLMNH 139048)*. Remarks: There are according to the GNS two places named Progreso in Guayas province [01° 46' 00" S 079° 45' 00" W and 01° 53' 00" S 079° 52' 00" W respectively]. Given the record for Zapotal, it is most probably that a place named Gómes Rendón was meant [02° 24' 00" S 080° 22' 00" W] for which the GNS lists as a variant name Progreso.

31. ?Bostryx ceroplastus (Pilsbry, 1896) Type locality: Peru, Río Marañon, Balsas. Lectotype ANSP 25468 (Breure 1979). Distribution: Tungurahua, 32 km SE Ambato (FMNH 106156). Remarks: This is a species from the eastern slopes of the Andes in northern Peru and its disjunct distribution—both geographically and in altitude—makes a confirmation of this record necessary.

32. Bostryx juana (Cousin, 1887) Type locality: Gualacco, Prov. Cuenca. Lectotype MNHN (designated and refigured by Breure 1976, Pl. 6 Fig. 5). Distribution: Azuay, Gualeco (MNHN).

Genus Bulimulus Leach, 1814 Type species: Helix exilis Gmelin, 1791. Distribution: West Indies, Venezuela, Guyana, Surinam, French Guiana, Brazil, Paraguay, Uruguay, Argentina, Bolivia, Peru, Ecuador, Colombia, Panama, Costa Rica, Nica- ragua, Guatemala, Belize, Mexico.

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33. Bulimulus fontainii (d’Orbigny, 1837) Type locality: Near Guayaquil. Holotype BMNH 1854.12.4.165. Distribution: Guayas, near Guayaquil (FMNH 31339). Remarks: This is the only species of Bulimulus s.str. and is well outside the range of this genus. Once revised this taxon may prove to be a synonym, possibly of Drymaeus cactivorus (Broderip).

Genus Naesiotus Albers, 1850 Type species: Bulimus nux Broderip, 1832. Distribution: West Indies, Venezuela, Brazil, Paraguay, Uru- guay, 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.

39. Naesiotus amastroides (Ancey, 1887) Type locality: Galapagos Islands. Dall described in 1893 a variety, Bulimulus (Naesiotus) amastroides anceyi, from “Chatham Island, 488 m”.

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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.

44. Naesiotus canaliferus (Reibisch, 1892) Type locality: [Galápagos,] Chatham Island. Lectotype ZMB 47948a (designated by Köhler 2007: 137, Fig. 46). Distribution: Galápagos, Isla San Cristobal.

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.

47. Naesiotus cinerarius (Dall, 1917) New name for Bulimulus (Naesiotus) cinereus Reibisch, 1892 not Bulimus cinereus Reeve, 1848. Type locality: [Galápagos,] James Island. Distribution: Galápagos, Isla San Salvador.

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.

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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 km2. The species is found throughout the humid zone on Santiago Island.

52. Naesiotus deroyi A.G. Smith, 1972 Type locality: Galápagos, Isla Santa Cruz, on the NW side, 264 m. Holotype CAS 13730. Distribution: Galápagos, Isla Santa Cruz.

53. Naesiotus duncanus (Dall, 1893) Type locality: Galapagos Island, Duncan Island. Distribution: Galápagos, Isla Pinzón.

54. Naesiotus elaeodes (Dall, 1917) Type locality: Galapagos, Albemarle Island, near Banks Bay, 455–700 m. Syntypes CAS. Distribution: Galápagos, Isla Isabela.

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 km2. The range was probably greater sometime in the past as suggested by the numer- ous 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).

59. Naesiotus gilderoyi (Van Mol, 1972) Type locality: Galápagos, Santa Cruz, à proximité du Cerro Coralon [near Cerro Coralon]. Holotype IRSN. Distribution: Galápagos, Isla Santa Cruz. Restricted to a single highland valley (Anonymous 2006).

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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 km2. 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 km2 (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 km2, 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.

67. Naesiotus kublerensis Chambers, 1986 Type locality: [Galápagos,] Santa Cruz, Island, Cueva de Kubler. Holotype USNM 842298. Distribution: Galápagos, Isla Santa Cruz.

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.

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70. Naesiotus nucula (Pfeiffer, 1854) Type locality: Galapagos Islands. Distribution: Galápagos, Isla San Cristobal; Isla Floreana (Parent 2003).

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 km2 (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 km2 (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 km2 (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.

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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.

79. Naesiotus planospira (Ancey, 1887) Type locality: [Galápagos,] Chatham Island. Distribution: Galápagos, Isla San Cristóbal; Isla Floreana; Isla Gardner; Isla Campion (Parent 2003).

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]; Pichin- cha, “near Quito” (BMNH 1975321/1 paralectotype of Bulimus catlowiae Pfeiffer). Quito (CMC C11759, as B. catlowiae Pfeiffer; FLMNH; MCZ 074513, 140390; ZMZ 512411); 14.5 km NW Quito (FMNH 146936, RMNH); slopes Cerro Pichincha (FMNH 86651, 86652, 125440, 125442, 125444, 125464, 158902; MCZ 065114, 065115, 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 070604. 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.

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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 064957.

82. Naesiotus rabidensis (Dall, 1917) Type locality: [Galápagos,] Jervis Island, 275–300 m. Syntypes CAS. Distribution: Galápagos, Isla Rábida.

83. Naesiotus reibischii (Dall, 1895) Type locality: [Galápagos,] Indefatigable Island. Distribution: Galápagos, Isla Santa Cruz. Single population with a range of probably not more than 10 km2 (Parent 2003).

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.

86. Naesiotus rugulosus (Sowerby, 1838?) Type locality: Galapagos. Lectotype BMNH 1975176 (Breure 1979); “Chatham Island” as locality on the label. Distribution: Galápagos, Isla San Cristóbal; Isla Floreana; Isla Pinta (Parent 2003).

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 km2 (Parent 2003).

89. Naesiotus simrothi (Reibisch, 1892) Type locality: [Galápagos,] Albemarle Island.

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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).

94. Naesiotus trogonius (Dall, 1917) Type locality: [Galápagos,] Albemarle Island, 400 m. Holotype USNM 274096. Distribution: Galápagos, Isla Isabela.

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).

97. Naesiotus ventrosus (Reibisch, 1892) Type locality: Barrington Island. Lectotype ZMB 47949a (designated and figured by Köhler 2007: 138, Fig. 57) Distribution: Galápagos, Isla Santa Fe?

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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 km2). Population size is no more than 500 indivduals (Parent 2003). Remarks: Richardson (1995) considers this taxon to be synonymous with Naesiotus reibischii (Dall, 1895).

Genus Scutalus Albers, 1850 Remarks: Breure (1979) placed Thaumastus umbilicatus Miller, 1879 (type locality: Prov. Loja, Cata- mayo, 2000–3000 ft.) under Nomina inquirenda, doubtfully suggesting that it might be a Scutalus.

Subgenus Scutalus (Vermiculatus) Breure, 1978 Type species: Bulimus bicolor Sowerby, 1835. Distribution: Bolivia, Peru, Ecuador.

99. Scutalus (Vermiculatus) aequatorius (Pfeiffer, 1853) Type locality: Mountain Schinchulagua. Lectotype BMNH 1975377 (Breure 1979). Distribution: Chimborazo, Cerro Chanchalahua, a foothill of Cerro Chimborazo (BMNH 1975178/1 paralectotype); “Chimborazo” (ZMZ 512209); Cotopaxi, Cerro Cotopaxi (ZMB); Napo, Antisana (FMNH 124573; MCZ 193333; ZMB); Pastaza, Cerros de Abitagua (FMNH 129020); “Quito” (CMC C11763, ZMH 7625).

100. Scutalus (Vermiculatus) anthisanensis (Pfeiffer, 1853) Type locality: Mountain Anthisana. Lectotype BMNH 1975372. Bulimus cotopaxensis Pfeiffer, 1853 (type locality: Mountain Cotopaxi, lectotype BMNH 1975370) and Bulimus subfasciatus Pfeiffer, 1853 (type locality: Anthisana, lectotype BMNH 1975368) are considered synonyms, following Weyrauch 1967: 385 [see Breure 1978: 185]. Distribution: Chimborazo, Cerro Altar (ZMB); Cotopaxi, Cerro Cotopaxi (BMNH 1975371/1 paralectotype); Sigchos (RMNH); Napo, Cerro Antisana (BMNH 1975373/1 paralectotype of Bulimus anthisanensis Pfeiffer; BMNH 1975369/1 paralectype of Bulimus subfasciatus Pfeiffer; ZMB); Pichincha, Cayambe (BMNH); San Francisco (HMNH)*. “Quito” (CMC C11762; ZMZ 512211, as B. subfasciatus Pfeiffer). “Ecuador” (ZMZ 512218, 512219; ZMZ 512213, as B. subfasciatus Pfeiffer; ZMZ 512228, as B. cotopaxien- sis Pfeiffer). Remarks: As Cerro Cotopaxi forms the cross of intersection of the provinces of Cotopaxi, Pichincha and Napo, we have arbitrarily attributed the locality to Cotopaxi, not knowing on which slope the type material was taken.

101. Scutalus (Vermiculatus) cousini (Jousseaume, 1887) Type locality: “Concha”. Holotype MNHN. Distribution: ? Morona-Santiago, Conchay (MNHN). Remarks: Conchay is the only name that turns up in the GNS, that is closely resembling the original locality.

Scutalus sp. Distribution: Azuay, Paute (FLMNH 151259); Pichincha, near Pucará (HMNH).

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Genus Stenostylus Pilsbry, 1898 Type species: Bulimus nigrolimbatus Pfeiffer, 1854. Distribution: Venezuela, Peru, Ecuador, Colombia. 102. Stenostylus colmeiroi (Hidalgo, 1872) Type locality: Baeza. Holotype MNHN (refigured by Breure 1976). Distribution: Napo, Baeza. Remarks: Material in the Florida museum has been found under the name of Simpulopsis colmeiroi. How- ever, compared to the figure of the type, these specimens lack the whitish axial streaks on the last whorls and have the aperture slightly more elongate. They proved to be S. citrinovitrea (Moricand). See also Breure (2008).

103. ? Stenostylus guttulus (Pfeiffer, 1854) Type locality: Gualea. Distribution: Pichincha, Gualea. Remarks: Breure (1979: 102) mentioned that this taxon may prove to belong to Simpulopsis (Eudioptus) citrinovitrea (Moricand, 1836).

Genus Drymaeus Albers, 1850 Subgenus Drymaeus (Drymaeus) Albers, 1850 Type species: Helix hygrohylaea d’Orbigny, 1835. Distribution: Venezuela, Brazil, Uruguay, Argentina, Bolivia, Peru, Ecuador, Colombia, Panama, Costa Rica, Nicaragua, Honduras, Guatemala, Mexico.

104. Drymaeus (Drymaeus) aequatorianus (E.A. Smith, 1877) Type locality: Ecuador. Lectotype BMNH 1975137 (Breure 1979). Distribution: Carchi, 1,5 hour walk W Chical (FLMNH 26617)*; San Nicolás (FMNH 31409). “Ecua- dor” (ANSP 25752, 78535; BMNH 1975137/1 paralectotype; ZMZ 512312). Remarks: These are the first exact localities reported for this species.

105. Drymaeus (Drymaeus) albolabiatus (E.A. Smith, 1877) Type locality: Malacatos. Holotype BMNH 1877.3.28.3. Distribution: Loja, Malacatos. “Oriente Prov.” (FMNH 72339).

106. Drymaeus (Drymaeus) ambustus (Reeve, 1849) Type locality: No locality given. Syntypes BMNH 1975441, 1975442; labelled “between Tucunga and Ambato”. Bulimus chamaeleon Pfeiffer, 1855 is considered a junior synonym (Syntypes BMNH 1975443; labelled as “near Quito”). Distribution: Pichincha, Pomasqui (FLMNH 176052)*; 3 km NE Cayambe (FLMNH 37397)*; Coto- paxi/Tungurahua: between Jacunga [Latacunga] and Ambato (BMNH 1975441, 1975442). “Quito” (CMC C11760; FMNH 31480, 61126; ZMA). “Ecuador” (ZMA).

107. Drymaeus (Drymaeus) andai (Jousseaume, 1898) Type locality: Tena. Lectotype MNHN (designated and figured by Breure 1976). Distribution: Napo, Tena (MNHN).

108. Drymaeus (Drymaeus) baezensis (Hidalgo, 1869) Type locality: Baeza. Distribution: Napo, Baeza. Without locality (RMNH).

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109. Drymaeus (Drymaeus) bourcieri (Pfeiffer, 1853) Type locality: Pichincha. Lectotype BMNH 1975446 (Breure 1979). Distribution: Pichincha [no specific locality] (BMNH 1975447/1 paralectotype); Tungurahua, Chaupi (FMNH 72599). “Ecuador” (ANSP 5249; RMNH; ZMA; ZMZ 512161). Remarks: This is the first precise locality for this species.

110. Drymaeus (Drymaeus) buckleyi (Sowerby, 1895) Type locality: Ecuador. Lectotype BMNH 1907.11.21.48 (designated, redescribed and figured by Breure & Eskens 1981). Distribution: ? [no specific locality known]. “Ecuador” (BMNH 1907.11.21.49-50/2 paralectotypes).

111. Drymaeus (Drymaeus) chimborasensis (Reeve, 1848) Type locality: Chimborazo. Syntypes BMNH 1975460. Distribution: Chimborazo [probably on the slopes of Mt. Chimborazo, which could also be in provs. Bolívar or Tungurahua].

112. Drymaeus (Drymaeus) decoratus (Lea, 1838) Type locality: Colombia, near Carthagena. Distribution: Napo, Llanganate [Cordillera de Llanganatis] (NRS).

113. Drymaeus (Drymaeus) elegantissimus (Mousson, 1873)** Type locality: Northern part of South America. Distribution: Morona-Santiago, Cusuimi (FMNH 173042, 173043)*. Remarks: This is the first Ecuadorian locality for this species, which has been reported from Colombia (Pilsbry 1899: 211).

114. Drymaeus (Drymaeus) expansus (Pfeiffer, 1848) Type locality: Peru, Huallaga. Distribution: Tungurahua, Topo (ANSP 306773)*. “Ecuador” (ZMZ 511962).

Drymaeus (Drymaeus) expansus altorum (Weyrauch, 1958) Type locality: Peru, Río Chanchamayo, Quirimi Sur, above Puente Herreria. Holotype SMF 156295. Distribution: Morona-Santiago, Cusuimi (FMNH 173050)*. Drymaeus (Drymaeus) expansus orcesi (Weyrauch, 1958) Type locality: Montalvo, Río Bobonaza, 340 m. Holotype SMF 156292 (refigured by Neubert & Janssen 2004). Distribution: Napo, Sarayacu; Orellana, Loreto; Pastaza, Montalvo (all Weyrauch 1958).

115. Drymaeus (Drymaeus) fallax (Pfeiffer, 1853) Type locality: Tungurahua. Bulimus abscissus Pfeiffer, 1855 is now considered a junior synonym (syn. nov.). The type material is labelled “Quito” (ex Cuming collection, BMNH 1969142). Distribution: Pastaza, Puyo (ANSP 170713); Pichincha, Quito (ANSP 5247; ZMZ 512145; ZMZ 512413, as Bulimus abscissus Pfeiffer), Nono [Mitad del Mundo] (CMNH 86212); near Nanegal (ANSP 195213; ZMB); Santo Domingo de los Colorados (ANSP 306782)* (FMNH 85376). “Ecuador” (MCZ 227645). Remarks: There are two places called Puyo, one is in Azuay province at 2850 m altitude, the other is in Pastaza province at 1000 m. Given the type locality and the altitudinal range of the other localities (up to 1500

20 · Zootaxa 1768 © 2008 Magnolia Press BREURE & BORRERO TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. m), it is probable that the Pastaza locality is meant, rather than the centrally located Puyo in Azuay province.

116. Drymaeus (Drymaeus) fordii Pilsbry, 1898 Type locality: None given. Lectotype ANSP 72368a (Breure 1979). Distribution: Morona-Santiago, Miasal; Orellana, Loreto (both according to Weyrauch 1958).

117. Drymaeus (Drymaeus) fucatus (Reeve, 1849) Type locality: [Colombia] Sebundoi. Lectotype BMNH 1874.12.11.224 (Breure 1979); labelled “New Granada”. Distribution: Pichincha, near Nanegal (ZMB). “Ecuador” (ZMZ 512131). Remarks: Sebundoi, often referred to as an Ecuadorian locality, is most likely Sibundoy in Colombia, Department of Putumayo [01° 11' 00" N 76° 55' 00" W]. We have been unable to locate the other localities mentioned by Pilsbry (1898: 234): “Lumaco”; “near San Florencio and on the way to Manabe” [Manabí?].

118. ?Drymaeus (Drymaeus) fusoides (d’Orbigny, 1835) Type locality: Bolivia, Dept. La Paz, Prov. Yungas. Distribution: ? [no specific locality known]. “Quito” (ZMZ 512101). “Ecuador” (ANSP 78538). Remarks: The occurrence of this Bolivian species remains to be corroborated.

119. Drymaeus (Drymaeus) hidalgoi (Da Costa, 1898) Type locality: Ecuador. Holotype BMNH 1907.11.21.28. Distribution: Napo, Nachiyacu (ANSP 170712); Pastaza, Mera (FMNH 125503, labelled as “Meva”). “Ecuador” (BMNH 1907.11.21.29-30/2 paratypes). Remarks: These are the first exact localities for this species.

120. Drymaeus (Drymaeus) inaequalis (Pfeiffer, 1857) Type locality: Peru, banks of the Maranhon [Río Marañon]. Distribution: Napo, Nachiyacu (ANSP 170711).

121. Drymaeus (Drymaeus) membielinus (Crosse, 1867) Type locality: Ecuador. Distribution: Napo (see Pilsbry 1898: 210); Orellana, Loreto (Weyrauch 1958). “Ecuador” (ZMH 7269).

122. Drymaeus (Drymaeus) murrinus (Reeve, 1848) Type locality: Colombia, Bogotá. Lectotype BMNH 1975213 (Breure 1979). A synonym is Bulimus convexus Pfeiffer, 1855 (type locality: New Granada; lectotype BMNH 1975192, figured by Breure & Eskens 1981: Pl. 8 Fig. 7). Distribution: ? [no specific locality known]. “Ecuador” (BMNH, type of Bulimus convexus Pfeiffer; ZMZ 512156).

123. Drymaeus (Drymaeus) nystianus (Pfeiffer, 1853) Type locality: Ecuador. Lectotype BMNH 1975573 (Breure 1979). Distribution: Napo, Cerro Antisana (ANSP 186859; FMNH 125506; MCZ 193332); Pichincha, Pomas- qui (ANSP 25804; BMNH 1975574/1 paralectotype, labelled “Valley of Pomasqui”; ZMA); Machachi (FLMNH 22973)*; Tumbaco (Pilsbry 1898: 263). “near Quito” (ANSP 76001); “Quito” (CMC C11761; FMNH 31326, 78753; RMNH; ZMH 7271; ZMZ 512148). “Ecuador” (RMNH; ZMA).

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124. Drymaeus (Drymaeus) ochrocheilus (E.A. Smith, 1877) Type locality: Malacatos. Holotype BMNH 1877.3.28.4. Distribution: Loja, Malacatos.

125. Drymaeus (Drymaeus) orthostomus (E.A. Smith, 1877) Type locality: Ecuador. Holotype BMNH 1975132. Distribution: ? [no specific locality known]. “Ecuador” (BMNH 1975133/2 paratypes; ZMZ 512105). Remarks: This species is only known from the type material, a lot in the ZMZ (both with the imprecise locality) and a lot in FMNH without locality (FMNH 31469). E.A. Smith (1877) compared this species with Drymaeus albolabiatus, which is said to have a slightly different colour pattern and a somewhat smaller aperture. The two taxa may prove to be synonyms.

126. Drymaeus (Drymaeus) peeliii (Reeve, 1859) Type locality: Peruvian side of Amazonas. Distribution: Pastaza, Canelos (Pilsbry 1898: 205). “Ecuador” (FMNH 77540; ZMZ 512573).

127. Drymaeus (Drymaeus) petasites (Miller, 1878) Type locality: Nanegal, Sebondoi, Pilatón valley. Distribution: Los Rios, 35 mi E of Quevedo (FMNH 106260)*; Pichincha, Nanegal; valley of Río Pilatón; ?, “Los Puentes” (FMNH 31083). “Ecuador” (ZMA). Remarks: As the type locality as given by Miller (1878) is a mixture of localities from Ecuador and Colombia [see also the remarks under Drymaeus (D.) fucatus], it is here restricted to Nanegal.

128. Drymaeus (Drymaeus) planibasis Pilsbry, 1932 Type locality: above Chunchi, Pagma forest. Holotype ANSP 148436a. Distribution: Chimborazo, Chunchi (ANSP).

129. Drymaeus (Drymaeus) quadrifasciatus (Angas, 1878) Type locality: Ecuador. Lectotype BMNH 1879.1.21.3. Bulimus napo Angas, 1878 (type locality: Ecua- dor. Holotype BMNH 1879.1.21.4) is considered a junior synonym by Weyrauch (1958: 128). Distribution: Pichincha, Nanegal (EPN, according to Weyrauch 1958); Tungurahua, Cerro Tungurahua near Baños (FMNH 72597).

130. Drymaeus (Drymaeus) rabuti (Jousseaume, 1898) Type locality: Tena. Lectotype MNHN (designated and figured by Breure 1976). Distribution: Napo, Tena.

131. Drymaeus (Drymaeus) rhoadsi Pilsbry, 1932 Type locality: Pachijal. Holotype ANSP 148440. Distribution: Pichincha, Río Pachijal (ANSP).

132. Drymaeus (Drymaeus) rubrovariegatus (Higgins, 1868)** Type locality: Peru, Huamachuco. Syntypes BMNH 1868.4.3.1. Distribution: El Oro, 6 km N Zaruma on road to Malvas (FLMNH 37402)*.

133. Drymaeus (Drymaeus) sachsei (Albers, 1854) Type locality: Peru, Río Marañon. Lectotype ZMB 10290a (designated and figured by Köhler 2007).

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Distribution: Chimborazo, Huigra (FMNH 78757); Loja, Catamayo (type locality of the junior synonym Mormus catamayensis Miller, 1879); Loja (ZMH 9768). ”Ecuador” (ZMA).

134. ?Drymaeus (Drymaeus) scitulus (Reeve, 1849)** Type locality: Peru, Chachapoyas. Lectotype BMNH 1975217 (designated by Breure & Eskens 1981). Distribution: ? [no specific locality known]. ”Ecuador” (FLMNH 109353, 109354). Remarks: The occurrence in Ecuador of this easily recognizable Peruvian species needs corroboration. If so, it may be expected in the southern part of the country.

135. Drymaeus (Drymaeus) strigatus (Sowerby, 1833) Type locality: Peru, Huallaga. Distribution: Morona-Santiago, Cusuima (FMNH 173048). “Ecuador” (FMNH 77563; ZMZ 512121).

136. Drymaeus (Drymaeus) subeffusus (Philippi, 1869)** Type locality: Peru, Huancayo, Coyllorbamba. Distribution: Loja, Macará (FMNH 31317).

137. Drymaeus (Drymaeus) tigrinus (Da Costa, 1898) Type locality: Ecuador. Holotype BMNH 1907.11.21.55. Distribution: ? [no specific locality known]. Remarks: Pilsbry (1898: 231) suggests that this taxon may be a junior synonym of Drymaeus strigatus (Reeve, 1848), which occurs in northern Peru. If he is correct, this species might be expected to occur in southern Ecuador.

138. Drymaeus (Drymaeus) violaceus (Mousson, 1873) Type locality: In the northern part of South America. Distribution: Pichincha, hill N of Tandapi, 71.7 km SW Quito on the road to Santo Domingo, 1600 m (RMNH)*.

139. Drymaeus (Drymaeus) volsus Fulton, 1907 Type locality: Ecuador. Holotype BMNH 1907.5.3.162. Distribution: ? [no specific locality known]. “Ecuador” (BMNH). Remarks: Known from the type material only.

Subgenus Drymaeus (Mesembrinus) Albers, 1850 Type species: Helix virgulata Férussac, 1821. Distribution: West Indies, Venezuela, Guiana, Surinam, French Guyana, Brazil, Peru, Ecuador, Colombia, Panama, Costa Rica, Nicaragua, El Salvador, Honduras, Guatemala, Belize, Mexico, USA.

140. Drymaeus (Mesembrinus) cactivorus (Broderip, 1832) Type locality: Colombia “ad montem Chris”. A junior synonym is Mormus occidentalis Miller, 1879, type locality: Guyaquil. Distribution: Guayas, 3 km E Zapotal (FLMNH 139105)*; 11 km W Progreso (FLMNH 139108)*. Remarks: The type locality might be Cerro Montecristi in Manabí [01° 03' 00" S 080° 40' 00" W] instead of a place in Colombia, where this species has not been found so far. According to the GNS there are two places named Progreso in Guayas province [01° 46' 00" S 079° 45' 00" W and 01° 53' 00" S 079° 52' 00" W respectively]. Given the record for Zapotal, it is most probably that a place named Gómes Rendón was meant

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141. ?Drymaeus (Mesembrinus) fidustus (Reeve, 1849) Type locality: [Colombia, Sibundoy]. Lectotype BMNH 1975517 (Breure 1979). Distribution: ? [no specific locality known]. “New Grenada, Sebundoi” (BMNH 1975518/1 paralectotype). Remarks: This species is tentatively referred to Ecuador on the authority of Miller (1878). The type locality is just across the border from northern Ecuador, in southwestern Colombia.

142. Drymaeus (Mesembrinus) flavidulus (E.A. Smith, 1877) Type locality: South Ecuador, Zarama. Lectotype BMNH 1975134. Distribution: El Oro, “Zarama” [sic, Zaruma] (BMNH 1877.3.28.1/1 paralectotype).

143. Drymaeus (Mesembrinus) loxanus (Higgins, 1872) Type locality: Loja. Lectotype BMNH 1975552. Distribution: ?Loja [no specific locality known]. “New Granada” (ZMB).

144. Drymaeus (Mesembrinus) loxensis (Pfeiffer, 1846) Type locality: Ecuador, El Catamaija prope Loxa. Lectotype BMNH 1975553 (Figured by Breure & Eskens 1981). Distribution: Loja, Catamayo (BMNH 1975554/1 paralectotype). “Quito” (ZMZ 512331). “Ecuador” (CMNH 42426).

145. ?Drymaeus (Mesembrinus) nigrofasciatus elongatulus Pilsbry, 1898 Type locality: Not given. Distribution: ? [no specific locality known]. “Ecuador” (ZMZ 512303, as Thaumastus nigrofasciatus “Reeve 1849 or Pfeiffer”). Remarks: Pilsbry (1898: 308) mentions this subspecies from “Tequendano” on the authority of Strebel. We have been unable to locate this place in Ecuador. As this species is quite common around Bogotá, it is possible that Tequendama (W Bogotá) is meant.

146. ?Drymaeus (Mesembrinus) serenus (Philippi, 1867)** Type locality: Peru, Dept. Cajamarca, Hacienda Sunchubamba. Distribution: ? [no specific locality known]. “Ecuador” (FMNH 31279). Remarks: The occurrence of this Peruvian species needs to be corroborated. It could possibly be expected in the southern part of the country.

147. Drymaeus (Mesembrinus) subpellucidus (E.A. Smith, 1877) Type locality: Ecuador. Holotype BMNH 1872.5.22.19. Distribution: ? [no specific locality known].

148. Drymaeus (Mesembrinus) wintlei Finch, 1929 Type locality: Ecuador. Holotype BMNH 1929.6.11.1. Distribution: ? [no specific locality known].

Drymaeus sp. Carchi, 2 km E Maldonado, 1550 m (FLMNH 26610)*; Loja, Loja (ZMH 9771) [labelled as Drymaeus

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Genus Simpulopsis Beck, 1837

Subgenus Simpulopsis (Eudioptus) Albers, 1860 Type species: Helix pseudosuccinea Moricand, 1836. Distribution: Brazil, Paraguay, Argentina, Peru, Ecuador, Colombia.

149. Simpulopsis (Eudioptus) citrinovitrea (Moricand, 1836) Type locality: Brazil, Bahia, near Salvador. Syntypes MNHN. Distribution: Napo, Baeza; 5.5 km SSE Baeza (FLMNH 37384)*; 19.3 km NNE Baeza (FLMNH 37385)*; Pichincha, 59 km by road W Machachi, 1250m (RMNH)*. Remarks: The material from the Florida Museum was labelled as Simpulopsis colmeiroi (Hidalgoi). This appears to be a misidentification, comparing the material with other specimens of S. citrinovitrea at hand, both from Ecuador and from Brazil.

SUBFAMILIA ORTHALICINAE

Remarks: The most recent revision of this subfamily is from Strebel (1909), who apparently saw all the material in the major European collections at that time.

Genus Sultana Shuttleworth, 1856

Subgenus Sultana (Sultana) Shuttleworth, 1856 Type species: Helix sultana Dillwyn, 1817. Distribution: Guiana, Brazil, Bolivia, Peru, Ecuador.

150. Sultana (Sultana) sultana (Dillwyn, 1817)** Type locality: New Zealand [sic]. Distribution: Los Rios, Centro Científico Río Palenque (FLMNH 181509)*; Morona Santiago, 59 km SSE Patuca (FLMNH 139123)*; Cusuimi (FMNH 173044); Río Macuma, ca. 10 km from Río Merona (FMNH 170626); Orellana, Loreto (ANSP 195216). Remarks: This is the first record of the nominal subgenus for Ecuador. There is also a lot collected by Cousin and labelled “Rio Napo” (FLMNH 166126). In the Philadelphia museum is a lot of Orthalicus sultana angustior Preston, which is a taxon described from eastern Peru, and labelled “Topo, Ecuador” [prov. Tun- gurahua] (ANSP 306772).

Subgenus Sultana (Metorthalicus) Pilsbry, 1899 Type species: Bulimus yatesi Pfeiffer, 1855. Distribution: Peru, Ecuador.

151. Sultana (Metorthalicus) augusti (Jousseaume, 1887) Type locality: Ecuador. Distribution: Azuay, Quebrada Machai (Pilsbry, 1899: 195); Pastaza, Mera (ANSP 306612); Puyo (FMNH 86638); Tungurahua, Topo (FMNH 86639).

152. Sultana (Metorthalicus) deburghiae (Reeve, 1859) Type locality: Peruvian side of the Amazon.

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Distribution: ?Morona-Santiago, Mirador (Strebel 1909: 173); Napo, 6.5 km SSE Baeza (FLMNH 36103)*; Nachiyacu (ANSP 170702; FLMNH 166252); Pastaza, Cerros de Abitagua (FMNH 126171, 125467); Mera (FMNH 86674; MCZ 193330); Porvenir (Strebel 1909: 173); Pichincha, Nanegal (ZMB); Tungurahua, Topo (FMNH 86673); Baños (FMNH 126172, 125514, 124582); Río Negro (MCZ 271635). “Ecuador” ( FLMNH 166253; RMNH). Remarks: Although there are different localities named “Mirador”, there is only one in the eastern part of the country. “Porvenir”, which is also a locality mentioned by Strebel, could be either in Prov. Orellana or Pastaza. The latter place is located in the easternmost part of the country, close to the border with Peru.

153. Sultana (Metorthalicus) kellettii (Reeve, 1850) Type locality: Ecuador? Holotype BMNH 1975241. Distribution: Azuay, Cuenca (ZMH 7637); Nabón (Strebel 1909: 160); Loja, Malacatos (Strebel 1909: 159); Pastaza, Mera (FLMNH 109372); Cerros de Abitagua (FMNH 125477); Tungurahua, Rio Negro (MCZ 271634); Zamora-Chinchipe, Tapichalaca* (B. Harris, personal communication and photograph). “Ecuador“ (CMNH 86211; FLMNH 166255; RMNH; ZMH 7636, 7641).

Subgenus Sultana (Trachyorthalicus) Strebel, 1909 Type species: Bulimus fraseri Pfeiffer, 1858. Distribution: Ecuador. Remarks: We have retained here the classification of Trachyorthalicus as a subgenus of Sultana, although we are doubtful if it is warranted given the slight differences with the previous one.

154. Sultana (Trachyorthalicus) fraseri (Pfeiffer, 1858) Type locality: Prov. Cuenca. Holotype ANSP 78573, variety brevispira Pilsbry, 1899. Distribution: Azuay, near Cuenca; Loja (Strebel 1909: 154, as forma brevispira; no specific locality mentioned); Morona-Santiago, Gualaquiza (SMF 90606). “Ecuador” (ANSP 78573, as type of variety brevispira Pilsbry, 1899; FLMNH 109360, 109358; RMNH; ZMH 7654).

Genus Orthalicus Beck, 1837 Type species: Buccinum zebra O.F. Müller, 1774. Distribution: West Indies, Venezuela, Guiana, Surinam, French Guyana, Brazil, Peru, Ecuador, Colombia, Panama, Costa Rica, Nicaragua, El Salvador, Honduras, Guatemala, Belize, Mexico, USA. Remarks: The species of this genus are generally very variable, with different colour forms and transitions between them. Strebel (1909) has made a useful revision, although this does not prevent that there are obviously quite some misidentifications in collections.

155. Orthalicus bensoni (Reeve, 1849) Type locality: Banks of the Amazon. Syntypes BMNH 1975582, labelled “Brazil”. Distribution: Napo, Sarayacu (Pilsbry 1899: 148); Río Napo (CMC C10806).

156. Orthalicus bifulguratus (Reeve, 1849) Type locality: Andes of Colombia. Distribution: El Oro, 10.2 km W Pinas (FLMNH 26616)*; Pichincha, Río Pilatón (Pilsbry 1899: 143 mentions Pilaton Valley on the basis of specimens collected by Wolf); Tungurahua, Topo (FMNH 86649).

157. Orthalicus maracaibensis (Pfeiffer, 1856) Type locality: Venezuela, Maracaibo.

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Distribution: Guayas, Guayaquil (ZMH 9883); San Juan (Pilsbry 1899: 140 mentions “Ecuador, San Juan de la Costa” as locality for his colour form imitator. From among the many places named San Juan this is the only one located near the coast).

158. Orthalicus mars (Pfeiffer, 1861) Type locality: Ecuador. Distribution: ? [no specific locality known]. Remarks: This taxon is known from the original description only, which has been translated by Pilsbry (1899: 143).

Orthalicus sp. Napo, 14.7 km NE Río Salado (FLMNH 22929)*; Río Napo (FMNH 72330), labelled as Orthalicus phlo- gerus (d’Orbigny, 1835), which is a Bolivian species.

Genus Corona Albers, 1850 Type species: Helix regina Férussac, 1821. Distribution: Venezuela, Guiana, Brazil, Bolivia, Peru, Ecua- dor, Colombia.

159. Corona pfeifferi (Hidalgo, 1869) Type locality: Canelos. Distribution: Pastaza, Canelos; Tungurahua, Topo (ANSP 306620).

160. Corona regalis (Hupé, 1857) Type locality: Brazil. Distribution: Tungurahua, Baños (FLMNH 109374 [labelled “Banes”]); “Ecuador” (FLMNH 166132, 166132).

161. Corona regina (Férussac, 1823) Type locality: Not given. Lectotype MNHN (Simone 2006). Distribution: “Ecuador” (FLMNH 109375). Remarks: The attribution of authorship and year of publication of this species has been confusing, but can be summarized as follows: Helix regina Férussac, 1821: 40 [nomen nudum]. Helix regina Férussac, 1823 in Férussac & Deshayes, 1819–1851: pl. 119. not Helix regina Fér., Bowdich, 1823: Pl. 8 Fig. 26 [= Liguus virgineus Linné, 1766; see Pilsbry, 1901: 165]. The confusion was probably due to the complex publication history of Férussac’s Histoire... which was continued after his death by Deshayes; Kennard (1942) has sorted out the publication dates of the different volumes in which this work was issued. Helix regina was originally mentioned by Férussac in 1821, but not validly published according to the ICZN-rules. It was referred to by Simone (2006: 160) as Corona regina (Férussac in Bowdich, 1822).

162. Corona rosenbergi Strebel, 1909** Type locality: Not given. Distribution: Orellana, Loreto (ANSP 195205). Remarks: This is the first precise locality for this species.

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Genus Porphyrobaphe Shuttleworth, 1856

Subgenus Porphyrobaphe (Porphyrobaphe) Shuttleworth, 1856 Type species: Bulinus iostoma Sowerby, 1824. Distribution: Peru, Ecuador.

163. Porphyrobaphe (Porphyrobaphe) iostoma (Sowerby, 1824) Type locality: Not given. Distribution: El Oro, near Machala (FMNH 72337); Chacras ; Santa Rosa (both Pilsbry 1899: 151); Esmeraldas, 9 km SW Atacames (FLMNH 139140)*; San Lorenzo (FLMNH 109870, 166241); ravine W of Camarones (FLMNH 182642)*; Camarones, near Esmeraldas (CMC C10825, 10826); Esmeraldas [without further indication] (FMNH 94931, 127535; ZMH 9886); Guayas, 3.1 miles [5 km] N Santa Elena (FLMNH 21274)*; 5 km N Palmar (FLMNH 139104, 139116, 139117)*; 4 km N Palmar (FLMNH 139117)*; 5 km NNE Progreso [= Gómez Rendón] (FLMNH 139141)*; 4 km N Monteverde (FLMNH 139144)*; Isla Puna (FMNH 146951); Colonche (Pilsbry 1899) (ANSP 266744); Cerro Colorado, 15 km N Guayaquil (ANSP 195221); Guayaquil (ZMB); Chongón (Strebel 1909: 104); Manabí, Manta (CMC C11029, 11030; FLMNH 109869, 182643; FMNH 72331, 86701; ZMB); Machalilla (ZMB; ZMH 28452, 28453); Quebrada Balsa- Maragua, N of Jaramijo (ANSP 195222; FMNH 216972); 6 km SSE Jipijapa (FLMNH 139103, 139134)*; 11 km WSW Jipijapa (FLMNH 139136)*; 9 km WSW Paján (FLMNH 139135)*; 8 km NNW Cascol [= Casca- jal] (FLMNH 139143)*; Portoviejo (Pilsbry 1899); Manabi [without further details] (ZMH 7634). “Ecuador” (CMC C10823, 10824, 11121; CMC C11125, as P. iostomus phasianella Valenciennes; FLMNH 48927, 48928; ZMA). Remarks: In the ANSP-collection is a lot (ANSP 306623) labelled “Mindo” [Prov. Pichincha], on the western slopes of the Andes but at an altitude of 1100 m. This might be P. i r ro ra tu s , which is a species of higher altitudes and has also been reported from the same locality. Another lot (ANSP 142458) identified as P. iostoma is labelled “Punta Ayancay”, which suggests a location on the coast. The only name in the GNS-database is a place called Ayancay, which is in Prov. Cañar at 2375 m on the eastern slopes of the Andes.

164. Porphyrobaphe (Porphyrobaphe) saturnus (Pfeiffer, 1860) Type locality: Pallatanga. Distribution: Chimborazo, Pallatanga (FLMNH 109376); Riobamba (ZMB); El Oro, 10 km S Piñas (FLMNH 36106, 37404)*; 6 km N Zaruma on road to Malvas (FLMNH 36125)*; Loja, Malacatos (Strebel 1909: 107, as “Malacates”). “Ecuador” (CMC C10819, 11028, 11122, 11123). Remarks: Besides the village Pillatanga in Prov. Chimborazo on the western slopes of the Andes at 1700 m, there is a Hacienda Pallatanga in Prov. Azuay on 2475 m altitude [02° 44' 00" S 078° 38' 00" W].

Subgenus Porphyrobaphe (Oxyorthalicus) Strebel, 1909 Type species: Bulimus irrorata Reeve, 1849. Distribution: Ecuador, Colombia. Remarks: Richardson (1993) lists this subgenus as synonym of the nominal one, but as he gives no argu- ments we have retained the classification of Zilch (1959–1960).

165. Porphyrobabphe (Oxyorthalicus) irrorata (Reeve, 1849) Type locality: Brazil? New Granada?. Syntypes BMNH 1975248, labelled “Quito, Ecuador”. Distribution: Napo, “Oriente” (FMNH 126177, 125476, 125499); Pastaza, Puyo (ANSP 170707); Mera (FLMNH 109894; FMNH 126254, 127526); Pichincha, hill N of Río Tandapi, 71.7 km SW Quito on road to Santo Domingo, 1600 m, secundary forest, in leaf litter (RMNH)*; Santo Domingo (ZMB); Mindo (ANSP 337840; FLMNH 166234; FMNH 86681, 216973); near Nanegal (ANSP 195218, 195219); Gualea (CMNH 86210); Río Cinto (ZMB); valley of Río Pilatón; (Pilsbry 1899:156); Tungurahua, Topo (ANSP 306611;

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FMNH 86690). “Ecuador” (CMC C11739). Remarks: Bulimus grevillei Pfeiffer, 1876—type locality: Quito—is considered a subjective junior synonym.

166. Porphyrobaphe (Oxyorthalicus) subirroratus (Da Costa, 1898) Type locality: Paramba. Holotype BMNH 1907.11.21.114. Distribution: Carchi, Hacienda Paramba; Pastaza, Mera (ANSP 220422/1 paratype; FMNH 86675, 125481); ?Pichincha, Mera (FLMNH 166243). Remarks: The GNS-gazetteer lists two localities in Pichincha named “Hacienda Mera”, one directly south of Quito [00° 25' 00" S 078° 33' 00" W], the other now being part of that city. It is more likely—also in view of the material from the FMNH—that Mera in Pastaza is meant. The specimens are labelled Porphyrobaphe subirroratus yeronta Miller, which is an unpublished name.

Genus Hemibulimus Martens, 1885 Subgenus Hemibulimus (Hemibulimus) Martens, 1885 Type species: Liguus (Hemibulimus) excisus Martens, 1885. Distribution: Ecuador, Colombia.

167. Hemibulimus (Hemibulimus) excisus (Martens, 1885) Type locality: Colombia, near Popayán. Distribution: ? Azuay, Macas (Strebel 1909: 108, as “Maccas”). Remarks: There are two places called Macas, one in Prov. Azuay the other in Prov. Morona-Santiago which we think is too low in altitude compared with the Colombian type locality (2400 m). Besides, there are two places called Quebrada de Macas, one in Prov. Chimborazo (near Riobamba), the other in Prov. Cotopaxi. We have tentatively attributed the locality, which is based on material traded by Rolle, to the one in Azuay, near Cuenca. The species has thus a remarkably disjunct distribution, which may be in part a reflection of insufficient sampling.

168. Hemibulimus (Hemibulimus) magnificus (Pfeiffer, 1848) Type locality: Quito. Distribution: ? [no specific locality known]. Remarks: This species is only known from the original description, translated by Pilsbry (1899: 185). Type specimen refigured by E.A. Smith (1907).

Incertae sedis Thaumastus alausiensis was described by Cousin (1887: 228, Pl. 4 Fig. 13) from “le versant du mont Hacu, entre Achapallas et la rivière Sula, Alausi [02° 12' 00" S 078° 50' 00" W], province de Chimborazo”. Only the type material in the MNHN is known. Breure (1976: 1141, Pl. 6 Fig. 4) placed this taxon in Peronaeus (Lis- soacme) Pilsbry, 1896 = Bostryx Troschel, 1847. In his revision of the Bulimulinae (Breure 1979) the taxon was placed under nomina inquirenda and doubtfully assigned to either Scutalus Albers, 1850 or Naesiotus Albers, 1850.

Discussion

Geographical distribution Orthalicidae have been collected at 128 localities in Ecuador. These sites, however, are not evenly spread

CHECKLIST ORTHALICIDAE IN ECUADOR Zootaxa 1768 © 2008 Magnolia Press · 29 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. throughout the country (Table 1). When compared to the area size of each province (adapted after Lahmeijer, 2007), it is clear that the Galápagos is outstanding in its biodiversity, while some provinces (notably Bolívar, Sucumbíos and Zamora-Chinchipe) are clearly undersampled.

TABLE 1. Number of localities and the number of taxa per province, and province size (in km2) in total and percentage. 22 taxa for which no specific localities are known, unidentified specimens for which specific localities were given in collections and questionable records are omitted.

Province km2 % localities % taxa % Azuay 8125 3% 7 6% 5 2,3% Bolívar 3940 1% 0 0% 0 0,0% Cañar 3122 1% 1 1% 1 0,5% Carchi 3605 1% 5 4% 4 2,0% Chimborazo 6072 2% 7 6% 7 3,4% Cotopaxi 6569 2% 3 3% 3 1,5% El Oro 5850 2% 5 4% 6 2,9% Esmeradas 15239 6% 3 3% 1 0,5% Guayas 20503 8% 10 9% 5 2,3% Imbabura 4559 2% 2 2% 2 1,0% Loja 11207 4% 4 3% 11 5,4% Los Ríos 7175 3% 2 2% 2 1,0% Manabí 18879 7% 8 7% 2 1,0% Morona-Santiago 25690 10% 7 6% 7 3,4% Napo114314%109%2210,7% Orellana 22500 8% 3 3% 8 4,0% Pastaza 29774 11% 7 6% 16 7,8% Pichincha 12915 5% 25 21% 26 12,7% Sucumbíos 18328 7% 0 0% 0 0,0% Tungurahua 3335 1% 7 6% 14 6,8% Zamora-Chinchipe 23111 9% 1 1% 0 0,0% Galápagos 8010 3% 63 30,7% TOTAL 269939 100% 117 100% 100,0%

Ecoregions and endemism To enable the study of biodiversity and development of conservation strategies, Olson et al. (2001) and Olson & Dinerstein (2002) have systematically compared representative samples of ecosystems and mapped them according to ecoregions. An ecoregion is defined (see also http://worldwildlife.org/science/ecoregions.cfm) as an area that contains a geographically distinct assemblage of natural communities that (a) share a large majority of their species and ecological dynamics; (b) share similar environmental conditions; and (c) interact ecologically in ways that are critical for their long-term persistence.

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TABLE 2. Occurrence of genera in different mainland ecoregions. In bold type the ecoregions have been indicated where more than 50% of the specific (sub-)genus localities occur.

Genus Subgenus Ecoregion Plekocheilus Plekocheilus NT0145 Plekocheilus Eurytus NT0121, NT0142, NT0145 Plekocheilus Aeropictus NT0145 Thaumastus Kara NT0121 Thaumastus Scholvienia NT0121 Thaumastus Thaumastus NT0121 Bostryx NT0121, NT0178, NT0214 Naesiotus NT0121, NT0145, NT0214, NT0232, NT1006 Scutalus Vermiculatus NT0121, NT0142 Drymaeus Drymaeus NT0121, NT0142, NT0145, NT1006 Drymaeus Mesembrinus NT0121, NT0241 Simpulopsis Eudioptus NT0121, NT0145 Sultana Sultana NT0121, NT0142, NT0178 Sultana Methorthalicus NT0121, NT1006 Sultana Trachyorthalicus NT0121 Orthalicus NT0121, NT0145 Corona NT0121, NT0142 Porphyrobaphe Porphyrobaphe NT0115, NT0214 Porphyrobaphe Oxyorthalicus NT0121, NT0145, NT1006

TABLE 3. Endemism and range-restrictedness arranged according to mainland ecoregions. e: ecological endemic (i.e., present only on a certain ecoregion); E: endemic to Ecuador; SL: known only from single locality. For explantion of Ecoregions see legend of Figure 1.

Ecoregions e e+E SL e+E+SL NT0115 0 0 0 0 NT0121 31 19 17 10 NT0142 9 7 6 4 NT0145 9 4 5 2 NT0178 0 0 0 0 NT0214 1 0 0 0 NT0232 0 0 0 0 NT1006 1 1 1 1 TOTALS 52 31 29 17

In mainland Ecuador, the following ecoregions are present (Fig. 1B): Chocó-Darién moist forests (NT0115); Eastern Cordillera real montane forests (NT0121); Napo moist forests (NT0142); Northwestern Andean montane forests (NT0145); Western Ecuador moist forests (NT0178); Ecuadorian dry forests (NT0214); Tumbes-Piura dry forests (NT0232); Guyaquil flooded grasslands (NT0905); Cordillera Central páramo (NT1004); Northern Andean páramo (NT1006); Esmeraldas-Pacific Colombia mangroves (NT1409); Gulf of Guayaquil-Tumbes mangroves (NT1413); Manabí mangroves (NT1418). See http://worldwildlife.org/ science/ecoregions/neotropic.cfm for detailed description and photographs. The dominant ecosystems are

CHECKLIST ORTHALICIDAE IN ECUADOR Zootaxa 1768 © 2008 Magnolia Press · 31 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. moist and montane forests, with smaller areas of dry forests on the coast and patches of páramo at high altitude. Dominant threats are deforestation, agriculture, grazing and increased colonization in the eastern part of the country. An endemic species is one that is restricted to a particular geographic area. This area can be restricted by political boundaries (e.g. endemic to Ecuador) or by ecological boundaries (e.g. restricted to páramos). Thus the context is important. A related concept is a range-restricted species, which are species with a small distribution. This also is contextually-framed, viz. varying across different groups of animals. Whereas some groups (e.g. birds) are sufficiently known to establish a threshold expressed in km2, distributional data of Neo- tropical land molluscs is too scarce to allow this. Hence a species is here considered to be range-restricted if it is known from a single locality. When we exclude the species with unknown distribution, Ecuador has 150 species belonging to the family Orthalicidae, of which 119 (79%) are only known from this country (including the Galápagos). Of the 72 mainland species analysed, 39 are endemic to Ecuador (54%) and 52 are restricted to one ecoregion. From the mainland species 29 are known only from a single locality, and 10 of these species have not been recollected after their original description. The number of endemic, range-restricted species is 17, of which 12 are confined to montane forests. The Eastern Cordillera montane forests is the ecoregion with the highest number of endemic range-restricted species (10 species; Table 3). This contrasts with the 63 taxa known from the Galápagos, of which many are known from very small areas. Parent & Crespi (2006) have studied the sequential colonization and diversification of these species. They conclude that the diversification has been driven by a combination of geographical factors that affect colonization patterns, and ecological factors that foster within-island speciation. Geographically, these endemic range-restricted species cluster in three areas and some scattered localities (Fig. 1C): a) five species on the northwestern Andean slopes, with Nanegal accounting for two species; b) five species on the eastern slope of the Andes, centering around Tena with two species; c) six species in southern Ecuador, centering around Malacatos with three species. These areas are currently best seen as containing concentrations of endemic, restricted-range species, which may be of interest for future conservation work once further research corroborates the present findings. Herbert & Moussalli (2008) pointed to the knowledge gathered by botanists about areas of endemism as a guide for malacological field work. In Ecuador areas of plant endemism are mainly based on the work of Borchsenius (1997) and Gentry et al. (1995). From their work it also appears that especially the areas identified here on eastern slopes of the Andes coincide with areas of plant endemism (Gentry et al. 1995: Huancabamba region, Gran Sumaco and Upper Napo River region respectively).

Distribution of genera—modelled by Maxent software

Plekocheilus Guilding, 1828 (Fig. 2A) This genus is clearly occurring at both fringes of the Andes, with the notable exception of Plekocheilus (Aeropictus) tenuissimus which lives on the plateau near Quito. The nominal subgenus is known only from localities on the western slopes of the Andes, while Plekocheilus (Eurytus) occurs mainly on the eastern side. The altitudinal range is from 450–2750 m, with an average of 1245 m.

Thaumastus Albers, 1860 (Fig. 2B) This genus has a patchy distribution throughout the Andes, having its main distribution in the Andean parts of Peru and southern and eastern states of Brazil (see Simone 2006). Most of the Ecuadorian species occur at higher altitudes (1200–2625 m; average 1975 m), with the only exception being the record of Thau- mastus (T.) flori from Machala in El Oro province, which is at sea level. This figure indicates that Thaumastus

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FIGURE 2. Potential distribution range of selected genera using Maxent modelling; squares are localities where mem- bers of genera have been found. Colours ranging from very unlikely (0–0.2, dark green) to very likely (0.8–1.0, red). A. Plekocheilus. B. Thaumastus. C. Bostryx. D. Naesiotus. E. Drymaeus (D.). F. Drymaeus (Mesembrinus). G. Simpulopsis. H. Sultana. I. Orthalicus. J. Corona. K. Porphyrobaphe (P.). L. Pophyrobaphe (Oxyorhtalicus).

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TABLE 4. Data on species used for distributional analysis. Alt. range: altitudinal range; SL: single locality—marked species are only known from a single locality (x) or by the original type material (*); Endemic: Endemic to Ecuador.

Genus Subgenus species Alt.range in m Ecoregion Endemic SL Plekocheilus Plekocheilus cardinalis 1000–1500 NT0145 Plekocheilus Eurytus aristaceus 850–2750 NT0145 x Plekocheilus Eurytus aureonitens 1150 NT0145 x x Plekocheilus Eurytus doliarus 700 NT0145 x Plekocheilus Eurytus floccosus 200–1350 NT0121, NT0142 Plekocheilus Eurytus jacksoni 250–750 NT0142 x Plekocheilus Eurytus jimenezi 750–2000 NT0121, NT0142 x Plekocheilus Eurytus lynciculus 500–750 NT0121, NT0142 Plekocheilus Eurytus mcginteyi 450 NT0142 x * Plekocheilus Eurytus nocturnus 1000 NT0121 x x Plekocheilus Eurytus oligostylus 750–1000 NT0121, NT0142 Plekocheilus Eurytus roseolabrum 1500 NT0121 x Plekocheilus Eurytus taylorianus 900–2500 NT0121, NT0145 x Plekocheilus Eurytus tricolor 750–2750 NT0121 x Plekocheilus Aeropictus tenuissimus 1250–1650 NT0145 x Thaumastus Kara thompsonii 2500–2625 NT0121 x Thaumastus Scholvienia jaspideus 2000 NT0121 Thaumastus Thaumastiellus sarcochrous 2950 NT0121 x Thaumastus Thaumastus buckleyi 2500 NT0121 x * Thaumastus Thaumastus flori 0–1500 NT0121 x Thaumastus Thaumastus hartwegi 1350 NT0121 x Thaumastus Thaumastus orcesi 1500 NT0121 x x Bostryx bilineatus 5–300 NT0178, NT0214 Bostryx juana 2400 NT0121 x * Naesiotus florschuetzi 5–25 NT0214, NT0232 x Naesiotus quitensis 1500–3800 NT0121, NT0145, NT1006 Scutalus Vermiculatus aequatorius 1475–4300 NT0121 Scutalus Vermiculatus anthisanensis 3500–4300 NT0121 x Drymaeus Drymaeus aequatorianus 900–3000 NT0145 Drymaeus Drymaeus albolabiatus 1500 NT0121 x * Drymaeus Drymaeus ambustus 2400–2900 NT0121, NT1006 x Drymaeus Drymaeus andai 500 NT0142 x * Drymaeus Drymaeus baezensis 1950 NT0121 x * Drymaeus Drymaeus bourcieri 3400 NT0121 x x Drymaeus Drymaeus decoratus 2700 NT0121 x Drymaeus Drymaeus elegantissimus 225 NT0142 x Drymaeus Drymaeus expansus 225–1400 NT0121, NT0142 Drymaeus Drymaeus fallax 600–1500 NT0121, NT0145 Drymaeus Drymaeus fordii 250–400 NT0142 x Drymaeus Drymaeus fucatus 1500 NT0145 x Drymaeus Drymaeus hidalgoi 750–1100 NT0121 x

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Drymaeus Drymaeus inaequalis 750 NT0121 x Drymaeus Drymaeus membielinus 400 NT0142 x Drymaeus Drymaeus nystianus 2350–4300 NT0145, NT1006 x Drymaeus Drymaeus ochrocheilus 1500 NT0121 x * Drymaeus Drymaeus petasites 1500 NT0121, NT0145 x Drymaeus Drymaeus planibasis 2950 NT1006 x * Drymaeus Drymaeus quadrifasciatus 1500–2000 NT0121, NT0145 x Drymaeus Drymaeus rabuti 500 NT0142 x * Drymaeus Drymaeus rubrovariegatus 1200 NT0121 x Drymaeus Drymaeus rhoadsi 600 NT0145 x * Drymaeus Drymaeus sachsei 1300–1500 NT0121 Drymaeus Drymaeus strigatus 225 NT0142 x Drymaeus Drymaeus subeffussus 450 NT0121 x Drymaeus Drymaeus violaceus 1600 NT0145 x Drymaeus Mesembrinus cactivorus 25–150 NT0214 Drymaeus Mesembrinus flavidulus 1200 NT0121 x x Drymaeus Mesembrinus loxensis 1350 NT0121 x x Simpulopsis Eudioptus citrinovitrea 1250–1950 NT0121, NT0145 Sultana Sultana sultana 25–1425 NT0142, NT0178 Sultana Metorthalicus augusti 1100–2700 NT0121, NT0142 x Sultana Metorthalicus deburghiae 750–2000 NT0121, NT0142 Sultana Metorthalicus kellettii 1100–2700 NT0121 Sultana Trachyorthalicus fraseri 1250–2550 NT0121 Orthalicus bensoni 1500 NT0121 x Orthalicus bifulguratus 950–1350 NT0121 Corona pfeifferi 450–1350 NT0121 x Corona regalis 2000 NT0121 x Corona rosenbergi 400 NT0142 x x Porphy- Porphyrobaphe iostoma 0–300 NT0115, NT0121, robaphe NT0178, NT0214 Porphy- Porphyrobaphe saturnus 950–1300 NT0121 x robaphe Porphy- Oxyorthalicus irrorata 1000–1600 NT0121, NT0145 robaphe Porphy- Oxyorthalicus subirrorata 700–1100 NT0121, NT0145 robaphe

The southernmost locality in Ecuador, Malacatos in prov. Loja, is on the limits of its range as modeled by Maxent. The distribution of Plekocheilus, predominantly influenced by annual frost frequency and April precipitation, extends northward into Colombia. South of Ecuador the genus is known from isolated areas in Cen- tral Peru, western Brazil and Bolivia.

Bostryx Troschel, 1847 (Fig. 2C) This genus is widely distributed in Peru, and ranges southward into Bolivia and Argentina. It reaches its northernmost range limit in Ecuador, although ongoing studies show that it also occurs at isolated spots in

CHECKLIST ORTHALICIDAE IN ECUADOR Zootaxa 1768 © 2008 Magnolia Press · 35 TERM OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website site is prohibited. southwestern Colombia (Borrero & Breure, in preparation). The majority of the locations are at altitudes up to 300 m (average 83 m). The only exception is the single record for Bostryx juliani from Azuay province, Gua- leco, which is also well beyond the modeled distribution by Maxent. Raxworthy et al. (2007) suggested the use of ecological niche modelling for recognition of spatial error localities, which could also be the case with this record. It is suggested here that outlier localities could also point to incorrect identifications. As this species has not been recollected since its original description in 1887, its status needs to be corroborated. The dominant environmental variable that influences the distribution model is the October precipitation level.

Naesiotus Albers, 1850 (Fig. 2D) The mainland species are distributed clearly in two distinct groups. Naesiotus florschuetzi is a lowland species, occurring on the coast. In contrast, Naesiotus quitensis is a high-altitude species, which ranges from 1850–4300 m (average 2836 m). Heuristic estimates of the relative contributions of variables show that April precipitation and elevation are dominant.

Drymaeus Albers, 1850 (Fig. 2E–F) This genus is wide-spread in the Neotropics and very rich in species. In Ecuador the species mainly belong to the nominal subgenus, which ranges from 250–4300 m (average 1689 m). Drymaeus (Mesembri- nus) shows some overlap with Drymaeus (Drymaeus) in distribution and in altitudinal range (25–1950 m, average 692 m). There are three localities where species of both subgenera are sympatric, ranging in altitude from 1200–1950 m. The same overlap, in additional but distinct centers of gravity in distribution may be observed when the Maxent models for the two taxa are compared. These models also indicate that the subgenera differ in their ecological constraints: Drymaeus (D.) is influenced by April precipitation and the annual diurnal temperature range, whereas the model of Drymaeus (Mesembrinus) is explained by a comparatively high contribution of minimum annual temperature and ecoregions. It should be noted that for Drymaeus (D.) two localities are somewhat outside the modelled range, which might be due to over-fitting of the model or the reasons explained before and by Raxworthy et al. (2007). The influence of the different variables and the settings to obtain the best fit for modelling species in this group will be subject for further research.

Simpulopsis Beck, 1837 (Fig. 2G) This genus is currently represented by a few locality records on the fringes of the Cordillera; it may prove to be present in some other areas, especially in the north of Ecuador, connecting to the distribution in southwestern Colombia. Figure 2G shows the Maxent modelled distribution that was obtained by including distribution data from other parts of South America as well. The maximum annual temperature and October precipitation are relatively important variables in this model.

Sultana Shuttleworth, 1856 (Fig. 2H) Species of this genus have an altitudinal range extending from 25–2700 m (average 1133 m). The nominal subgenus ranges from 25–1450 m (average 465 m), while S. (Metorthalicus) and S. (Trachyorthalicus) range from 200–2700 m (average 1467 m). This difference in altitudinal range is partly reflected in the distribution models of the subgenera (not shown), where Sultana (S.) is mainly confined to “El Oriente”, while the other subgenera exhibit a patchy distribution on the western slopes of the Andes. Figure 2H shows the Maxent model for the genus as a whole, with the annual diurnal temperature range making the predominant contribution among the variables.

Orthalicus Beck, 1837 (Fig. 2I) This genus shows more or less the same distribution compared to Sultana in the Maxent model, with the same relatively high contribution of the annual diurnal temperature range.

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The altitudinal range where species of this genus have been found is 450–1500 m (average 1110 m). This genus is probably distributed at localities across the whole country, being particularly likely to occur in the eastern part.

Corona Albers, 1850 (Fig. 2J) The distribution of this genus has a striking resemblance to Sultana, both in altitudinal range (400–2000 m; average 1050 m) and in distribution pattern as modelled by Maxent. Also the environmental variable that has the highest relative contribution is the same, viz. the annual diurnal temperature range.

Porphyrobaphe Shuttleworth, 1856 (Fig. 2K-L) There is a striking difference in the two subgenera. Porphyrobaphe (Oxyorthalicus) ranges from 710– 1600 m (average 1146 m), while the nominal subgenus ranges from sea level to 2750 m, but with an average of 368 m. This difference is reflected in the Maxent modelling: Porphyrobaphe (P.) shows a coastal distribution and October precipitation is the main variable (Figure 2K), whereas P. (Oxyorthalicus) shows a patchy distribution on the fringes of the Andes and in “El Oriente” (Figure 2L). The April precipitation and the maximum annual temperature are relatively important environmental variables for the latter taxon.

Directions for further research

Comprehensive studies on Ecuadorian land snails are absent so far and this paper can only fill a small part of the gap that obviously exists. Given that context a number of directions are here suggested for further research. One important direction is sampling poorly known areas. Distribution maps such as Figure 2 may provide some guidance in achieving that aim. In addition, it would be especially useful to recollect species only known from single localities or ancient type material. This would allow for establishing distribution ranges more precisely, give more insight in potential areas of endemism and could underpin future conservation activities. Recently Guisan & Thuiller (2005) have critically reviewed the limitations of species distribution models. An important limitation of this study is its reliance on museum collections and the fact that only part of the records are based on recent (defined as after 1950) sampling. It may be assumed that (nearly) all samples have been collected near urban dwellings and roads, these areas being more easily accessible, and are therefore biased in relation to the true spatial or environmental distribution of species. Transect-based sampling together with the collection of data on a wide range of environmental variables, may allow for improved modelling of species distributions. Whereas this study is limited by its focus to one family, it would be interesting to use the same approach on a broader scale, both systematically and geographically. Recently there are some comprehensive checklists that are forthcoming or have been published for a wider range of land snail taxa for Colombia, Peru and Brazil (Borrero & Breure in preparation; Ramirez et al. 2003; Simone 2006). If the underlying data were analysed, this could lead to better insights into biogeographical patterns within the Neotropical land snail fauna. A well established link between ecology and evolution is population biology. However, there is rapidly evolving a new fusion between these two disciplines, with inputs from ecologicla data on the one hand and of phylogenetic trees on the other hand (Westoby 2006). Recent studies (Brumfield & Edwards 2007; Carstens & Richards 2007; Graham et al. 2004a; Knouft et al. 2006; Richards et al. 2007; Wiens & Graham 2005) indicate that this is a promising field. Further research could deepen our knowledge on speciation processes and the co-evolution of different groups in past, present-day and even future environments.

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Acknowledgements

First of all the senior author would like to thank G. Cuno Tisalema (Tisaleo), who has encouraged him to focus again on the Ecuadorian malacofauna after a gap of 30 years. We are very indebted to W. Blankenhorn (Zur- ich), P. Callomon (Philadelphia), J. Gerber (Chicago), B. Hausdorf (Hamburg), F. Köhler (Berlin), T. Pearce and P. Robb (Pittsburgh), for kindly supplying data on relevant material from their collections, and/or assist- ing us in using their collection during our studies. B. Harris (Fundación Jocotoco, Quito) kindly provided information on snails collected in one of the nature reserves in southern Ecuador. Thanks are also due to D. van Bruggen, B. Hausdorf and J. Hemmen, for their helpful comments. K. Way (London) critically read the manuscript and improved the English, for which we are much indebted. Finally, we like to thank B. Ruthen- steiner, J. Ablett and an anonymous referee for their suggestions, allowing us to make useful improvements in the final version of the manuscript.

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