R.B. Salvador a, C.M. Cunha a & L.R.L. Simone a a Museu de Zoologia da, Universidade de São Paulo, São Paulo,

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1 This article was downloaded by: [R.B. Salvador] On: 29 March 2013, At: 08:08 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: Registered office: Mortimer House, Mortimer Street, London W1T 3JH, UK Journal of Natural History Publication details, including instructions for authors and subscription information: Taxonomic revision of the orthalicid land snails (Pulmonata: Stylommatophora) from Trindade Island, Brazil R.B. Salvador a, C.M. Cunha a & L.R.L. Simone a a Museu de Zoologia da, Universidade de São Paulo, São Paulo, Brazil Version of record first published: 25 Jan To cite this article: R.B. Salvador, C.M. Cunha & L.R.L. Simone (2013): Taxonomic revision of the orthalicid land snails (Pulmonata: Stylommatophora) from Trindade Island, Brazil, Journal of Natural History, 47:13-14, To link to this article: PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.

2 Journal of Natural History, 2013 Vol. 47, Nos , , Taxonomic revision of the orthalicid land snails (Pulmonata: Stylommatophora) from Trindade Island, Brazil Rodrigo B. Salvador*, Carlo M. Cunha and Luiz Ricardo L. Simone Museu de Zoologia da, Universidade de São Paulo, São Paulo, Brazil (Received 3 August 2012; final version received 20 November 2012; first published online 25 January 2013) Land snails (stylommatophoran pulmonates, shells only) were collected on Trindade Island, 1140 km off Vitória, Brazil, by the Marion Dufresnei Expedition (MD-55) and more recently by our team. Trindade s endemic snails are Bulimulus brunoi and Naesiotus arnaldoi (Orthalicidae), and Oxyloma beckeri and Succinea lopesi (Succineidae). As some non-native species have also been found, a taxonomic revision of the native fauna was needed. Here, the revision is focused on orthalicids, presenting updated descriptions and diagnosis. Moreover, the supposed native orthalicid Bulimulus trindadensis is deemed here to be a Subulinidae, and the new genus Vegrandinia is erected to accommodate it. Trindade s insular environment is delicate and very susceptible to invasive species. No living specimens of these taxa, native or non-native, have been collected since before the MD-55 Expedition. However, it is still premature to assume that the introduced species have failed to establish themselves or that the native species are now extinct. B508EAEB0A4B Keywords: Bulimulinae; MD-55 Expedition; non-native species; Subulinidae; Vegrandinia new genus Introduction Trindade Island (Ilha da Trindade) lies about 1140 km away from the Brazilian coast, off the city of Vitória (Espírito Santo state), at S, 29 0 W (Figure 1). It is the top of a volcanic cone rising 5500 m from the ocean floor, and is part of the Vitória Trindade Seamount Chain, which results from a magma plume. It forms an archipelago with its eastern neighbour, the Island of Martin Vaz (Asmus 1978; Morgan 1983). Trindade Island was discovered in 1501 by Portugal and later claimed by England, but has been part of Brazil since its independence, in 1822 (Almeida 2002; Cruz 2006). Ever since the island s discovery, many military vessels and scientific expeditions have travelled there, coming from Portugal, the UK, France, the USA and, naturally, Brazil (Almeida 2002). The latest great expedition was the Marion Dufresnei Expedition (MD-55), in 1987, a joint project of the Muséum National d Histoire Naturelle (Paris, France) and the Universidade Santa Úrsula (Rio de Janeiro, Brazil), having on board three malacologists: Philippe Bouchet, José H. Leal and Bernard Metivier. They managed to recover many pulmonate snail specimens (shells only) from the island. Later, the same was achieved by smaller expeditions by our research team. *Corresponding author. salvador.rodrigo.b@gmail.com 2013 Taylor & Francis

3 950 R.B. Salvador et al. Figure 1. Map showing the location of Trindade Island, off Vitória (20 30 S29 20 W). As some non-native species have been identified in this material (described elsewhere; Cunha et al. forthcoming), there was the suspicion that some of the native snails described for the island were in fact exotic species. This called for an immediate taxonomic revision of Trindade s native land snail fauna, which is presented here for the orthalicid snails (Bulimulus brunoi, B. trindadensis and Naesiotus arnaldoi), making use of the extra material found since their original description. All holotypes are figured herein and formal diagnoses and updated descriptions are offered. Moreover, B. trindadensis was deemed to belong to Subulinidae and the new genus Vegrandinia is presented here to accommodate it. Trindade s other two endemic species, Oxyloma beckeri Lanzieri, 1966 and Succinea lopesi Lanzieri, 1966 (Succineidae), are not revised here. Material and methods All land snails (Pulmonata: Stylommatophora) collected by the MD-55 and later expeditions are only empty shells, most in a good state of conservation. For detailed examination, samples were mounted on stubs, coated with a gold-palladium alloy and observed under a Zeiss DSM 940 scanning electron microscope. Measurements were taken with a digital caliper. The list of material examined follows each species description. Abbreviations used in the text are as follows: MNHN, Muséum National d Histoire Naturelle, Paris, France; MNRJ, Museu Nacional, Rio de Janeiro, Brazil; MZSP, Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil; and for shell measurements: H = shell length; D = greatest width of shell; S = spire length (excluding aperture); S = spire length (excluding body whorl); h = aperture height; d = aperture width. Systematics Family ORTHALICIDAE Subfamily BULIMULINAE Genus Bulimulus Leach, 1814 Bulimulus brunoi (Ihering, 1917) (Figures 2A F)

4 Journal of Natural History 951 Figure 2. (A C) Bulimulus brunoi (holotype, MNRJ 1128; H = 19.4 mm). (D) B. brunoi (MNRJ 1129; H = 20.7 mm). (E, F) B. brunoi, protoconch and teleoconch sculpture (MZSP ). (G) Naesiotus arnaldoi (holotype, MNRJ 3105; H = 9.5 mm). (H J) N. arnaldoi (MZSP ; H = 10.8 mm). (K) N. arnaldoi (paratype, MNRJ 3105; H = 10 mm). (L) N. arnaldoi, protoconch sculpture (MZSP ). (M O) Vegrandinia trindadensis (MZSP ; 5.2 mm). (P, Q) V. trindadensis (holotype, MNRJ 3711; H = 7 mm). (R, S) V. trindadensis (MZSP ), (R) protoconch sculpture, (S) broken aperture showing the lamellae.

5 952 R.B. Salvador et al. Loboa brunoi Ihering 1917: 121 (pl. 4, fig. 7). Bulimulus (Protoglyptus) brunoi: Bartsch 1918: 53 (pl. 4, fig. 7). Protoglyptus brunoi: Morretes 1949; Lanzieri, 1966: 169. Bulimulus brunoi: Breure 1978: 142 (figs ); Breure 1979; Salgado and Coelho 2003; Simone 2006: 117 (fig. 356). Holotype MNRJ 1128 (examined; Figure 2A C). Type locality Trindade Island. Distribution Endemic to Trindade Island. The species seems to occur next to or on the beach (Lanzieri 1966; personal observations). Etymology Species dedicated to Prof. Bruno Lobo (MNRJ). Diagnosis Weak callus on parietal region of aperture. Teleoconch sculptured by fine prosocline axial riblets bearing tiny round and regularly spaced nodules, giving impression of checked pattern. Mixed protoconch sculpture: regularly spaced parallel axial riblets (first 1 1 / 2 whorl) and anastomosing striae (next 1 / 2 whorl). Redescription Shell medium-sized, conical-turreted, with acuminated apex; spire high; seven whorls. Colour dirty white. Spire angle 40. Greatest width on body whorl; width about twofifths shell length. Protoconch sculptured by regularly spaced parallel axial riblets for first 1 1 / 2 whorls, and by fine anastomosing striae for another 1 / 2 whorl; transition to teleoconch unclear. Teleoconch sculpture by fine prosocline and irregular axial riblets and by fine spiral striae, their encounter forming tiny round and regularly spaced nodules and looking like checked pattern. Suture well-marked, slightly oblique (diagonal) to columellar axis. Whorl profile convex. Aperture weakly prosocline, medium-sized, oval, slightly anteroposteriorly elongated; about two-fifths of shell length, about onehalf of shell width. Weak callus on parietal region of aperture. Lamellae or teeth absent. Lip thin, simple, non-reflected, with exception of upper columellar region (reflected and covering umbilicus). Body whorl about three-fifths of shell length. Umbilicus rimate, almost imperforate. Measurements (in mm) Holotype. H = 19.4; D = 7.5.

6 Journal of Natural History 953 Mean (n = 32). H = 18.0 ± 1.2 (max 20.6; min 16.1); D = 7.8 ± 0.4 (max 8.8; min 7.3); S = 11.4 ± 1.0 (max 14.3; min 10.0); S = 8.4 ± 0.7 (max 10.5; min 7.0); h = 6.5 ± 0.4 (max 7.5; min 5.7); d = 4.0 ± 0.3 (max 4.8; min 3.4). Material analysed BRAZIL. Trindade Island: MNRJ 1128 (one shell, holotype), 1129 (one shell). Southwest Trindade Island: MNHN (four shells); MZSP (one shell). Andradas Beach (Praia dos Andradas): MZSP (35 shells). Tartarugas Beach (Praia da Tartarugas): MZSP (one shell), MZSP (one shell). Remarks Only the shell of B. brunoi is known; unfortunately, no complete animal has so far been recovered. The overall high-spired shell profile, albeit not being very common in the genus, is not unknown, as testified by the very similar shell shapes of some other Brazilian species, like, for instance, B. felipponei Marshall, 1930 and B. sporadicus (d Orbigny, 1835). It can be differentiated from these species by its sculpture pattern (Figure 2F) and the callus on the parietal region of the aperture. This last feature closely resembles another widespread Brazilian bulimuline genus, Rhinus Martens in Albers, 1860, in which it is commonly found. The protoconch sculpture of B. brunoi (Figure 2E) is somewhat unusual in the genus, but some species, such as B. guadalupensis (Bruguière, 1789), show similar patterns (Breure 1979). A similar protoconch sculpture can be found in some species of Thaumastus Albers, 1860, specially of the subgenus Thaumastiella Weyrauch, 1956 (Breure 1979); moreover, the overall shell shape of B. brunoi is also reminiscent of some species of this genus. Nevertheless, the number of whorls, their convex profile, the umbilicus and the aperture lacking a lamella, all point to a closer affinity with Bulimulus. Genus Naesiotus Albers, 1850 Naesiotus arnaldoi (Lanzieri and Rezende 1971) (Figures 2G L) Protoglyptus arnaldoi Lanzieri and Rezende 1971: 255 (figs 1 45). Naesiotus arnaldoi: Breure and Coppois 1978; Breure 1979; Salgado and Coelho 2003; Simone 2006: 125 (fig. 395). Holotype MNRJ 3105 (examined; Figure 2G). Paratypes MNRJ 3105 (examined; Figure 2K). Type locality Desejado Peak (Pico do Desejado), Trindade Island.

7 954 R.B. Salvador et al. Distribution Endemic to Trindade Island. Etymology Species dedicated to Prof. Arnaldo C.S. Coelho (MNRJ). Diagnosis Shell narrow. Aperture more anteroposteriorly elongated. Presence of a weak callus on parietal region of aperture. Teleoconch sculptured by fine prosocline and irregular axial ribs. Redescription Shell small, conical to fusiform; five whorls. Spire angle 55. Greatest width on body whorl; width about one-half shell length. Colour dirty white to light brown, somewhat translucent. Protoconch of 1 1 / 2 whorl, sculptured by weak regularly spaced undulating parallel axial riblets. Teleoconch smooth, except for growth lines. Suture weakly marked, slightly oblique (diagonal) to columellar axis. Whorls profile slightly convex. Aperture slightly prosocline, large, oval, somewhat anteroposteriorly elongated; about two-fifths shell length, about one-half shell width. Weak callus on parietal region of aperture. Lamellae or teeth absent. Lip thin, simple, non-reflected, with exception of upper columellar region (reflected and covering umbilicus). Body whorl about two-thirds shell length. Umbilicus rimate, almost imperforate. Measurements (in mm) Holotype. H = 9.5; D = 5.5; S = 4.0; S = 3.2; h = 5.1; d = 3.2. Mean (n = 68). H = 11.0 ± 0.6 (max 12.4; min 9.7); D = 5.7 ± 0.3 (max 6.3; min 5.1); S = 5.7 ± 0.4 (max 6.6; min 5.0); S = 3.6 ± 0.3 (max 4.6; min 2.9); h = 5.1 ± 0.4 (max 6.3; min 4.4); d = 3.2 ± 0.2 (max 3.8; min 2.6). Material analysed BRAZIL. Southwest Trindade Island: MNHN (eight shells); MZSP (two shells). Andradas Beach (Praia dos Andradas): MZSP (80 shells). Desejado Peak (Pico do Desejado): MNRJ 3105 (holotype and paratypes, 73 specimens, 145 shells). Remarks Naesiotus arnaldoi falls well within the morphological range of the genus, both with regard to its shell, specially the sculpture pattern of the protoconch (Figure 2L), which is typical of the genus (Breure 1979), and to its radula and genitalia (described by Lanzieri and Rezende 1971), which compare fittingly with the anatomical work of Breure and Coppois (1978) on the genus Naesiotus. Lanzieri and Rezende (1971) presented a thorough comparison of the shell of N. arnaldoi to other species in the genus, highlighting its similarities with N. eudioptus (Ihering in Pilsbry, 1897) from Southern Brazil and Paraguay (Simone 2006). Naesiotus arnaldoi can be differentiated from other Naesiotus species mainly by its narrower shell

8 Journal of Natural History 955 profile and its large and slightly anteroposteriorly elongated aperture. Other diagnostic features include a weak callus on the parietal region of the aperture and the sculpture pattern of the teleoconch, but these features can also be found in some other species, like N. eudioptus and N. lopesi (Rezende, Lanzieri and Inada, 1972), respectively. Family SUBULINIDAE Subfamily SUBULININAE Vegrandinia gen. nov. (Figures 2M S) Type species Bulimulus trindadensis Breure and Coelho, Included species Vegrandinia trindadensis (Breure and Coelho 1976). Etymology Allusion to the small size of the species, the tiniest native species on Trindade Island. From the Latin word vegrandis, meaning tiny, diminutive. Grammatical gender: feminine. Diagnosis Shell elongated-oval; whorls profile flattened; protoconch sculptured by sinuous axial striae; presence of columellar fold reaching aperture and palatal fold on inner portion of body whorl. Description Shell diminutive, elongated-oval tending towards conical. Suture weakly marked. Whorls profile flattened. Protoconch rounded, large, sculptured by sinuous axial striae; transition to teleoconch unclear. Teleoconch smooth. Aperture elliptical, prosocline; callus on parietal region. Lip thin, simple, non-reflected, with exception of upper columellar region, reflected and covering umbilicus. Columellar fold on median portion of aperture s columellar region. Palatal fold on inner portion of body whorl, on median portion of whorl s palatal region. Umbilicus rimate. Remarks Breure and Coelho (1976) described some animals collected at Trindade Island as an endemic new species, Bulimulus trindadensis. Interestingly, these authors made clear in their work that the species does not resemble any of the other species of Bulimulus (Breure and Coelho 1976: p. 5). The authors also stated that B. trindadensis resembles the fossil genus Itaborahia Maury, 1935 from the Palaeocene of Rio de Janeiro (Breure and Coelho 1976: p. 5), but the differences between the two are overwhelming: its much smaller size, the oval shell shape and the shape, position and features of the aperture and lip are widely disparate from the single species in the genus, Itaborahia lamegoi

9 956 R.B. Salvador et al. Maury, 1935; the only feature both species share is the columellar fold (Maury 1935; Salvador and Simone forthcoming), but this can also be found in many other orthalicid genera (Simone 2006). The following features distinguish B. trindadensis from the genus Bulimulus: small size, thin and fragile shell, prosocline aperture, weakly marked suture, the presence of a weak callus on the parietal region of the aperture, the presence of the columellar fold and the palatal fold and the overall shell shape. The combination of shell characters present in B. trindadensis is more closely allied to Subulinidae than to Orthalicidae: a minute, thin and delicate shell; a large and rounded protoconch; and a weakly marked suture. The presence of a columellar fold is also common in Subulininae, but not ubiquitous. However, because of its particular combination of characters, there is no single subulinid genus that can properly house this species; so the new genus is proposed. Vegrandinia seems to be closely allied to the genus Leptinaria Beck, 1837, but differs from it by its more oval shell shape, flattened whorl profile and by the presence of a palatal fold on the inner portion of the body whorl. Vegrandinia trindadensis (Breure and Coelho, 1976) comb. nov. (Figure 2M S) Bulimulus trindadensis Breure and Coelho 1976: 3 (figs 1 4); Breure 1979; Salgado and Coelho 2003; Simone 2006: 120 (fig. 372). Holotype MNRJ 3711 (examined; Figure 2P Q). Paratypes MNRJ 3712 (many shells, examined). Type locality Cabritos Beach (Praia dos Cabritos), Trindade Island: found imbedded in the loose sediment on the terrace near the beach, away from the tideline (Breure and Coelho 1976). Distribution Endemic to Trindade Island. Etymology Related to the species type locality, Trindade Island. Diagnosis Shell an elongated oval; whorl profile flattened; protoconch sculptured by sinuous axial striae; presence of columellar fold reaching aperture and palatal fold beginning 5 mm away from aperture.

10 Journal of Natural History 957 Redescription Shell diminutive, elongated-oval tending towards conical; about seven whorls; shell width about two-fifths of shell length. Colour white. Spire angle 40. Suture weakly marked. Whorl profile flattened. Protoconch ( 1 1 / 2 whorl) rounded, large, sculptured by sinuous axial striae; transition to teleoconch unclear. Teleoconch smooth. Aperture elliptical, prosocline; callus on parietal region; about one-third shell length, about onehalf shell width. Lip thin, simple, non-reflected, with exception of upper columellar region (reflected and covering umbilicus). Columellar fold on median portion of aperture s columellar region, beginning 1 mm inside aperture or reaching aperture. Palatal fold beginning 5 mm from aperture, on median portion of whorl s palatal region. Umbilicus rimate. Measures (in mm) Holotype. H = 7.0; D = 2.8; S = 4.1; S = 3.6; h = 2.5; d = 1.7. Mean (n = 7; types excluded). H = 4.8 ± 0.3 (max 5.0; min 4.5); D = 2.0 ± 0.1 (max 2.2; min 1.9); S = 3.1 ± 0.2 (max 3.4; min 2.8); S = 2.5 ± 0.3 (max 2.8; min 2.1); h = 1.7 ± 0.1 (max 1.8; min 1.6); d = 1.1 ± 0.8 (max 1.3; min 1.0). Material analysed BRAZIL. Southwest Trindade Island: MNHN (seven shells); MZSP (three shells), (one shell, metallized). Cabritos Beach (Praia dos Cabritos): MNRJ 3711 (one shell, holotype), 3712 (many shells, paratypes). Remarks The species is allocated here to the new monotypic genus Vegrandinia, under the new combination V. trindadensis. Unfortunately, no living specimen was found, so the species anatomical features remain unknown. Curiously, the shell length of the adult specimens analysed by Breure and Coelho (1976) averaged from 8 to 9 mm (which was an overestimation, our measures of the same specimens average 7 mm), while the ones collected by the MD-55 and later expeditions average 4.8 mm; the shell morphology, though, is the same. It is suspected that this reduction in size reflects inappropriate environmental conditions due to the Island s much degraded environment (see Discussion below). Discussion Five non-native species have been reported for Trindade Island (Alves 1998; Cunha et al. forthcoming): Bradybaena similaris (Férussac, 1821) (Bradybaenidae), Discus rotundatus (OF Müller 1774) (Discidae), Pupisoma dioscoricola (CB Adams 1845) (Valloniidae), Gastrocopta pellucida (Pfeiffer 1840) and Vertigo substriata Jeffreys 1833 (Vertiginidae). With the detection of introduced species on the island, the suspicion that some supposedly native land snails could also be introduced appeared. Fortunately, based on the exclusivities of the studied species, the present work supports the idea that the above-revised species are indeed native to Trindade Island. The two Succineidae species, Succinea lopesi and Oxyloma beckeri, were not revised here, for their type localities and distributions are restricted to the Island s two peaks (Desejado

11 958 R.B. Salvador et al. Peak for S. lopesi and São Bonifácio Peak for O. beckeri); so they are very probably also native, because introduced species do not usually expand their area towards such places (Gargominy 2008). As previously stated, many ships, carrying both military and scientific expeditions, have visited the island throughout the last centuries so there have been many opportunities for the introduction of non-native species (some of them possibly from secondary sources; Cunha et al. forthcoming). Land snails have been unintentionally introduced in many regions around the globe, impacting native vegetation and preying upon or outcompeting native species (Hadfield et al. 1993; Byers 1999; Cowie 2011). The environment in oceanic islands is extremely disturbance-sensitive so these problems are greatly aggravated, for introduced species can ultimately lead to the extinction of the native ones (Hadfield et al. 1993; Robinson 1999; Lydeard et al. 2004; Chiba and Roy 2011). Unfortunately, it is not yet possible to tell whether the native snails from Trindade Island were affected by the non-native species (Cunha et al. forthcoming). But the real problem for native snails on Trindade may lie somewhere else: the introduction of goats and house mice (Alves 1998; Silva and Alves 2011). On oceanic islands, introduced mammals are extremely destructive to the native fauna and flora (Hadfield et al. 1993; Cowie 2000; Lydeard et al. 2004; Campbell and Donlan 2005; Chiba and Roy 2011), and goats are indicated as the main cause of extinctions in such environments (Campbell and Donlan 2005). They were responsible for a drastic deforestation on Trindade, leading to the extinction or near extinction of many endemic plant species (Alves 1998; Serafini et al. 2010). Alarmingly, no living land snail, native or non-native, had been collected for long before the MD-55 Expedition. The extinction of native plants caused by the goats, and the resulting loss of habitats and food source, might have played a major role in the disappearance of the native snail species. Furthermore, an interesting fact was observed in V. trindadensis: the shell length of the specimens used in the original description of 1976 measured 7 mm, whereas those collected by the MD-55 expedition, in 1987, and more recent expeditions measure 4.8 mm. This corresponds to a significant reduction of 30% in shell size in 11 years; moreover, this small size was maintained until now. The lack of proper ecological knowledge does not allow much discussion, but we can point here to two possible causes to this phenomenon: (1) the extinction of native plants, caused by the goats, resulted in a loss of habitats and food source and in such a degraded environment the snails could not attain their normal size (although not documented for land snails, something similar was observed for intertidal gastropods by Roy et al. 2003); (2) the presence of non-native species on the island (Cunha et al. forthcoming) and the possible competition for food might have been detrimental to the growth of V. trindadensis. Finally, after a great campaign to eradicate the goats, concluded in 2005, the flora seems to be recovering with the help of some researchers (Martins and Alves 2007; Silva and Alves 2011). With the recovery of the native flora, snail populations should be able to re-establish themselves, provided that some have managed to survive in places where no collecting effort was undertaken. Still, this is valid for both native and non-native species alike. The only introduced mammals remaining on the island are house mice (with a population estimated at tens of thousands; Silva and Alves 2011), and they might also have had a part in the snails disappearance. It is well known that mice can voraciously prey upon land snails, especially if other food sources are scarce (Hadfield et al. 1993;

12 Journal of Natural History 959 Allen 2004); they will only eat medium-sized to large snails (Allen 2004), i.e. the native orthalicids Naesiotus arnoldoi and Bulimulus brunoi and perhaps also the native succineids: Succinea lopesi and Oxyloma beckeri (provided the mice have already reached the peaks on the innermost part of the island). Conclusions (1) There are at least five native species in Trindade, taxonomically reviewed herein. (2) Two of them are bulimuline orthalicids: Naesiotus arnoldoi and Bulimulus brunoi. (3) A supposed third orthalicid revealed itself as a subulinid, in a new genus herein described: Vegrandinia trindadensis. (4) The other native snails, the succineids Succinea lopesi and Oxyloma beckeri, are not reviewed here. (5) No living land snails have been collected on Trindade, which could indicate an extinction event. A possible cause would be the intense environmental degradation, including the introduction of non-native snails and mammals. However, it is still premature to embrace this scenario. (6) Despite this, no measures so far have been implemented towards the protection of this malacofauna and the avoidance of future invasions. Acknowledgements We are grateful to Philippe Bouchet for the invitation to study the MD-55 material housed at MNHN; to Philippe Maestrati and Virginie Heros for their help and hospitality during our visit; and to José Coltro Jr (Femorale), for supporting part of the trip to Paris in the first phase of this project. We are grateful to Lara Guimarães (MZSP) and Phillip Lenktaitis (USP) for helping with the scanning electron microscopy and to Alexandre D. Pimenta (MNRJ) for lending the material housed at his home institution. The present study was partly supported by FAPESP (Fundação de Amparo à Pesquisa do Estado de São Paulo), proc. #2010/ (to CMC), and partly by CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico), proc. #557166/ (to LRLS). References Allen JA Avian and mammalian predators of terrestrial gastropods. In: Barker GM, editor. Natural enemies of terrestrial molluscs. Wallingford (UK): CABI Publishing, p Almeida FFM Ilha de Trindade Registro de vulcanismo cenozóico no Atlântico Sul. In: Schobbenhaus C, Campos DA, Queiroz ET, Winge M, Berbert-Born MLC, editors. Sítios geológicos e paleontológicos do brasil. Vol. 1. Brasília (Brazil): Comissão Brasileira de Sítios Geológicos e Paleobiológicos (SIGEP), DNPM/CPRM, p Asmus HE Hipóteses sobre os sistemas de zonas de fraturas oceânicas/alinhamentos continentais que ocorrem nas regiões Sudeste e Sul do Brasil. In: PETROBRÁS. Aspectos estruturais da margem continental leste e sudeste do Brasil. Série Projeto REMAC 4. Rio de Janeiro (Brazil): PETROBRÁS/CENPES/DINTEP, p Alves RJV Ilha da Trindade & Arquipélago Martin Vaz: um ensaio geobotânico. Niterói (Brazil): Serviço de Documentação da Marinha, DHN.

13 960 R.B. Salvador et al. Bartsch P The status of Loboa brunoi von Ihering. Nautilus 32: Breure ASH Notes on and descriptions of Bulimulidae (Mollusca, Gastropoda). Zool Verhandel. 164: Breure ASH Systematics, phylogeny and zoogeography of Bulimulidae (Mollusca). Zool Verhandel. 168: Breure ASH, Coelho ACS Notes on Bulimulidae (Gastropoda, Euthyneura), 3. Bulimulus trindadensis sp. n. from Ilha da Trindade, Brazil. Basteria. 40:3 6. Breure ASH, Coppois G Notes on the genus Naesiotus Albers, 1850 (Mollusca, Gastropoda, Bulimulidae). Netherland J Zool. 28(2): Byers JE The distribution of an introduced mollusc and its role in the long-term demise of a native confamilial species. Biol Invas. 1: Campbell C, Donlan CJ Feral goat eradications on islands. Conserv Biol. 19: Chiba S, Roy K Selectivity of terrestrial gastropod extinctions on an oceanic archipelago and insights into the anthropogenic extinction process. Proc Natl Acad Sci USA 108(23): Cowie RH Non-indigenous land and freshwater molluscs in the islands of the Pacific: conservation impacts and threats. In: Sherley G, editor. Invasive species in the Pacific: a technical review and draft regional strategy. Apia (Samoa): SPREP, p Cowie RH Snails and slugs. In: Simberloff D, Rejmanek M, editors. Encyclopedia of biological invasions. Berkeley (CA): University of California Press, p Cruz CLMC Para uma historiografia sobre a Questão da Ilha da Trindade ( ). Navigator 4: Cunha CM, Salvador RB, Simone LRL. Forthcoming Non-native land snails on Trindade Island, Brazil. Nautilus Gargominy O Beyond the alien invasion: a recently discovered radiation of Nesopupinae (Gastropoda: Pulmonata: Vertiginidae) from the summits of Tahiti (Society Islands, French Polynesia). J Conchol. 39(5): Hadfield MG, Miller SE, Carwile AH The decimation of endemic Hawaiian tree snails by alien predators. Am Zool. 33: Ihering H Loboa brunoi n. g., n. sp. a land shell from the Brazilian island of Trinity. Nautilus 30(11): Lanzieri PD A família Succineidae (Gastropoda, Pulmonata) da Ilha de Trindade, Costa do Brasil. Papéis Avuls Dept Zool. 19(14): Lanzieri PD, Rezende HEB Protoglyptus arnaldoi, novo Bulimulidae da ilha da Trindade, Brasil (Mollusca, Gastropoda, Pulmonata). Arquiv Mus Nacl. 54: Lydeard C, Cowie RH, Ponder WF, Bogan AE, Bouchet P, Clark AS, Cummings KS, Frest TJ, Gargominy O, Herbert DG, et al The global decline of nonmarine mollusks. BioScience 54(4): Martins LSG, Alves RJV Regeneração natural do Morro Vermelho, Ilha da Trindade. Rev Brasil Biociênc. 5: Maury CJ New genera and new species of fossil terrestrial Mollusca from Brazil. Am Mus Novitat. 764:1 15. Morgan WJ Hotspot tracks and the early rifting of the Atlantic. Tectonophysics 94(1 4): Morretes FL Ensaio de catálogo dos moluscos do Brasil. Arquiv Mus Paranaense. 7: Robinson DG Alien invasions: the effects of the global economy on nonmarine gastropod introductions into the United States. Malacologia 41: Roy K, Collins AG, Becker BJ, Begovic E, Engle JM Anthropogenic impacts and historical decline in body size of rocky intertidal gastropods in southern California. Ecol Let. 6:

14 Journal of Natural History 961 Salgado NC, Coelho ACS Moluscos terrestres do Brasil (gastrópodes operculados ou não, exclusive Veronicellidae, Miladicae e Limacidae). Rev Biol Tropic. 51(suppl.3): Salvador RB, Simone LRL. Forthcoming Taxonomic revision of the fossil pulmonate mollusks of Itaboraí Basin (Paleocene). Brazil: Papéis Avulsos de Zoologia. Serafini TZ, França GB, Andriguetto-Filho JM Ilhas oceânicas brasileiras: biodiversidade conhecida e sua relação com o histórico de uso e ocupação humana. Rev Gestão Costeira Integrada 10(3): Silva NG, Alves RJV The eradication of feral goats and its impact on plant biodiversity a milestone in the history of Trindade Island, Brazil. Rodriguésia 62(3): Simone LRL Land and freshwater mollusks of Brazil. São Paulo (Brazil): EGB, FAPESP.