Archiv für Molluskenkunde 147 (1) 101–128 Frankfurt am Main, 29 June 2018

Terrestrial gastropods from Pedra Talhada Biological Reserve, Alagoas state, Brazil, with the description of a new species of Radiodiscus (: )

Rodrigo B. Salvador1, 2, Laurent Charles3, Luiz R.L. Simone4 & Philippe Maestrati5

1 Museum of New Zealand Te Papa Tongarewa, 169 Tory Street, 6011, Wellington, New Zealand ([email protected]). 2 Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191, Stuttgart, Germany. 3 Muséum de Bordeaux, 5 Place Bardineau, 33000, Bordeaux, France ([email protected]). 4 Museu de Zoologia da Universidade de São Paulo, Av. Nazaré 481, 04263-000, São Paulo, SP, Brazil ([email protected]) 5 Muséum national d’Histoire naturelle, 57 Rue Cuvier, 75005 Paris, France ([email protected]). • Corresponding author: P. Maestrati.

Abstract. A thorough collection effort recently took place at Pedra Talhada Biological Reserve (Reserva Biológica de Pedra Talhada, in Portuguese) in Alagoas state, one of the largest fragments of Atlantic Forest in northeastern Brazil. The expedition recovered hundreds of land snails and this material is fully reported and described herein. Forty-two species were found in the reserve (47 counting historical records), com- prised mostly of stylommatophoran snails, although some species of Neritimorpha and were also found. Many of these species are recorded here for the first time from Alagoas state, extending the geographical range of some or filling distribution gaps of others. The generaBothriopupa and Sterkia, both vertiginids, are reported here from Brazil for the first time. One important find is Megalobulimus cardosoi, which was deemed extinct in the IUCN Red List. Furthermore, a new species of Charopidae

is described, Radiodiscus ubtaoi sp. nov.eschweizerbartxxx sng- Despite being a reserve, signs of depredation were observed in Pedra Talhada by our team, underlining the urgency of strengthening and enforcing the protection of this area. Key words. Neritimorpha, Caenogastropoda, Radiodiscus ubtaoi sp. nov., . DOI. https://doi.org/10.1127/arch.moll/147/101-128

Introduction aim of this survey was to compile a preliminary check- list of Pedra Talhada’s molluscan fauna based on field The Pedra Talhada Biological Reserve (Reserva Biológica observations and photographs (very few specimens were de Pedra Talhada; henceforth “Pedra Talhada”) is a rem- collected). This visit resulted in an unpublished report nant of the coastal Atlantic Forest at the border of the with an illustrated list of 50 taxa bearing provisory iden- Alagoas and Pernambuco states in northeastern Brazil. tifications (typically to only, sometimes separated The Atlantic Forest, the second largest biome in South as distinct morphospecies). A summary of this report was America, covered more than 1 million km² in the 15th eventually published (Maestrati et al. 2015) in a general century, but today is severely fragmented, with less than volume about the of the reserve (Studer et 15% of its original area remaining (Câmara 2005). The al. 2015). This publication, also preliminary in character, conservation of the forest’s remnants is largely managed was intended to raise awareness and to support the request through parks and reserves, such as Pedra Talhada, which to the Brazilian authorities for an official collection effort was decreed a state park in August 1985 and a federal to produce an updated inventory of the reserve’s mollus- reserve in December 1989. can fauna. This led to an expedition during September In 1998, a first visit to the reserve was proposed to Prof. and October of 2015 to collect and photograph Pedra (Muséum national d’Histoire naturelle, Talhada’s land snails. Paris, France) and financially supported by Nordesta, a The material collected during this expedition was Swiss NGO created in May 1985 for educational and deposited in the malacological collection of the Museu environmental projects in and around Pedra Talhada. The de Zoologia da Universidade de São Paulo (São Paulo,

© E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller) and Senckenberg Gesellschaft für Naturforschung, 2018, ISSN 1869-0963 Archiv für Molluskenkunde · 147 (1) 2018

Brazil) and is studied here. Forty-seven species are reported in the present work: 42 from the 2015 expedi- tion and 5 additional species from the 1998 expedition that were not found again. Several of the species records are new for Alagoas state and the 2 vertiginid genera found represent the first records for Brazil. Moreover, a new species of Charopidae is described here and 1 spe- cies (Megalobulimus cardosoi (Morretes, 1952)) that had been deemed extinct was found alive. Another species, Biotocus turbinatus (L. Pfeiffer, 1845), supposed to be extinct earlier but recorded alive in 1998 (Maestrati et al. 2015), was not found again in 2015.

Figure 1. Map showing the location of the Pedra Talhada Biolo­ gical Reserve. Our collecting took place only in the portion of the reserve belonging to Alagoas state. Image adapted from Materials and Methods Tscharner et al. (2015).

Abbreviations. The following abbreviations are used. Shell dimensions: H = shell height (parallel to coiling the most humid (and hence favorable to land snails) and axis); D = greatest shell width (perpendicular to H). Insti- varied environments when compared to the more homo- tutions: ANSP = Academy of Natural Sciences of Drexel geneous plateau towards Pernambuco state. University (Philadelphia, USA); ICMBio = Instituto Chico A total of 49 collection stations were established inside Mendes de Conservação da Biodiversidade (Brasília, Bra- the reserve (Fig. 2, Table 1; with 3 additional stations zil); MNHN = Muséum national d’Histoire naturelle on the outer limits of the reserve, see below). Stations (Paris, France); MZSP = Museu de Zoologia da Universi- and field outings were decided based on vegetation and dade de São Paulo (São Paulo, Brazil); NHMUK = Natu- pedo-geomorphological maps (Tscharner et al. 2015, ral History Museum (London, UK); SMNS = Staat­liches complemented with Nusbaumer et al. 2015a, 2015b) Museum für Naturkunde Stuttgart (Stuttgart, Germany). and the knowledge of the reserve’s guides. For each sta- Study area. Pedra Talhada is situated in northeastern tion, photographs were taken (both of the snails and the

Brazil, on the parallel 09° 15′ S, about 90 km fromeschweizerbartxxx sng- the landscape), GPS coordinates were recorded and noted coast (Fig. 1). It encompasses about 4400 ha of rainforest together with other pertinent observations, and 4–6 L of between 450 and 900 m of altitude. The reserve belongs leaf litter were sampled. to the Borborema geological province, with a basement The fieldwork consisted of daily early-morning sam- of igneous rocks, mainly granites (Tscharner et al. pling conducted by groups of 2 or 3 people, totaling 70 2015), which are clearly observable as inselbergs. For person-days. Afternoons were used to dry, sieve, and sort more information about the reserve’s environment and the leaf litter samples (totaling c. 400–500 L); the latter vegetation, refer to Studer et al. (2015). was done first by the naked eye and then under the micro- The reserve is divided between 2 Brazilian states: Per- scope. During this phase, field notes were transferred to nambuco to the north (with 40% of the reserve’s area) and electronic files, further observations were noted when Alagoas to the south. The southern portion of the reserve, necessary, and live were photographed. in Alagoas, consists of a steep slope that continues north- After sorting, the total number of specimens recovered ward to an extended plateau. The slope is difficult to was rather low, but this is not surprising in neotropical access, by trails, but the northern plateau allows easy metamorphic contexts. The majority of the specimens access by vehicle. consisted of empty shells, which were first preserved in 70% ethanol and then dried a few days later. Live spec- Collection methodology. This work was possible by a imens were preserved in 95% ethanol to allow future permit from the ICMBio (Sisbio #48925-2). The 2015 molecular studies. expedition to Pedra Talhada lasted 3 weeks, from Sep- During the expedition, some localities in the immedi- tember 22 to October 6. It was originally planned to take ate vicinity of the reserve were also visited (stations 1, place a couple of weeks earlier to coincide with the end of 19 and 42; Fig. 2, Table 1), resulting in some additional the region’s wet season; however, it had to be postponed. species recorded. These species were not included in the Regardless, the rainy season was notably short in 2015, systematical part of this work as we intend to focus only with no consistent rainfall during August. As such, the on the undisturbed area of the reserve; however, they are sampling was done on a rather dry soil, with nearly no included in the Discussion. rain during fieldwork. The collection effort did not cover evenly the whole Vouchers and identification. All material (c. 1800 spec- reserve, which was a consciously informed decision. The imens) was deposited in the malacological collection of south portion (Alagoas state) was favored because it has the MZSP: lots MZSP 13358 to 134001. Material from

102 Salvador et al. · Land snails from Pedra Talhada the 1998 expedition is housed in the malacological col- Live specimens were photographed by LC; photo- lection of the MNHN, but the lots do not have accession graphs of preserved specimens were taken by RBS; SEM numbers (for specific details about this expedition, see the images were acquired in the SMNS by Christina G. Mar- Appendix). The identifications were made in comparison tin. Measurements were made either with digital calipers, with the comprehensive catalogue of Simone (2006), the for large shells, or with the aid of the computer software original descriptions and type material whenever possible, Leica Application Suite (LAS, v. 3.8.0) for minute shells. and, if necessary, additional material housed in the collec- tions of the MZSP, MNHN, and NHMUK. Nevertheless, several specimens were either juvenile or fragmentary. These could typically be identified to either family or Systematics genus level, but could not be confidently assigned to a Neritimorpha species; they very likely do not represent additional taxa and belong to a species already listed (it is not possible to Superfamily Helicinoidea decide which). Family Helicinidae All species are briefly discussed and figured below, with notes on their geographical distribution, range Genus Lamarck, 1799 extensions and, when necessary, their and/or morphology. A list of species occurrence by collection Helicina angulifera A.J. Wagner, 1910 station is presented in the Appendix. All the species col- Figure 3A–E lected live are figured as such, showing the animals’ soft Helicina angulifera Wagner 1910: 279, pl. 55, figs 6–8. body, as this information is hardly ever available in the literature, especially for the smallest animals. Type locality. Brazil, Bahia state.

Table 1. Collection stations in the Pedra Talhada Biological Reserve, with coordinates, altitude, and date when the collection took place. *Stations located just outside of the reserve.

Station Latitude (S) Longitude (W) Alt. (m) Coll. date Station Latitude (S) Longitude (W) Alt. (m) Coll. date PT2015-1* 09°15.703' 036°25.719' 536 22/ix/2015 PT2015-27 09°15.366' 036°25.060' 566 28/ix/2015

PT2015-2 09°15.582' 036°25.722' 564eschweizerbartxxx sng- 22/ix/2015 PT2015-28 09°15.095' 036°24.581' 627 29/ix/2015 PT2015-3 09°15.212' 036°25.742' 717 22/ix/2015 PT2015-29 09°15.271' 036°24.611' 581 29/ix/2015 PT2015-4 09°15.169' 036°25.673' 758 22/ix/2015 PT2015-30 09°15.267' 036°24.706' 542 29/ix/2015 PT2015-5 09°15.180' 036°25.670' 759 22/ix/2015 PT2015-31 09°14.860' 036°25.206' 686 30/ix/2015 PT2015-6 09°15.245' 036°25.630' 785 22/ix/2015 PT2015-32 09°13.648' 036°25.058' 869 30/ix/2015 PT2015-7 09°15.410' 036°25.837' 644 22/ix/2015 PT2015-33 09°13.903' 036°25.174' 890 30/ix/2015 PT2015-8 09°15.239' 036°25.604' 801 23/ix/2015 PT2015-34 09°14.318' 036°25.040' 870 30/ix/2015 PT2015-9 09°15.119' 036°25.735' 727 23/ix/2015 PT2015-35 09°13.996' 036°27.407' 749 01/x/2015 PT2015-10 09°15.268' 036°25.797' 679 23/ix/2015 PT2015-36 09°13.958' 036°27.257' 760 01/x/2015 PT2015-11 09°14.415' 036°25.089' 856 24/ix/2015 PT2015-37 09°14.625' 036°26.333' 731 02/x/2015 PT2015-12 09°14.424' 036°25.070' 855 24/ix/2015 PT2015-38 09°14.539' 036°26.498' 707 02/x/2015 PT2015-13 09°14.455' 036°25.079' 846 24/ix/2015 PT2015-39 09°14.695' 036°26.207' 725 02/x/2015 PT2015-14 09°14.953' 036°25.215' 641 24/ix/2015 PT2015-40 09°14.928' 036°25.962' 685 02/x/2015 PT2015-15 09°14.913' 036°25.309' 658 25/ix/2015 PT2015-41 09°15.229' 036°25.846' 660 02/x/2015 PT2015-16 09°15.170' 036°25.216' 588 25/ix/2015 PT2015-42* 09°15.746' 036°26.107' 526 04/x/2015 PT2015-17 09°15.261' 036°25.135' 570 25/ix/2015 PT2015-43 09°15.509' 036°25.956' 571 04/x/2015 PT2015-18 09°15.571' 036°25.275' 575 25/ix/2015 PT2015-44 09°15.498' 036°25.964' 580 04/x/2015 PT2015-19* 09°15.937' 036°24.814' 507 26/ix/2015 PT2015-45 09°15.367' 036°25.101' 558 05/x/2015 PT2015-20 09°15.548' 036°25.018' 487 26/ix/2015 PT2015-46 09°14.600' 036°26.008' 768 06/x/2015 PT2015-21 09°15.446' 036°25.032' 531 26/ix/2015 PT2015-47 09°14.318' 036°25.830' 860 06/x/2015 PT2015-22 09°15.419' 036°25.045' 548 26/ix/2015 PT2015-48 09°14.265' 036°25.763' 860 06/x/2015 PT2015-23 09°15.501' 036°25.169' 585 26/ix/2015 PT2015-49 09°14.297' 036°25.914' 864 06/x/2015 PT2015-24 09°15.504' 036°25.908' 584 27/ix/2015 PT2015-50 09°14.952' 036°24.313' 689 07/x/2015 PT2015-25 09°15.526' 036°25.923' 567 27/ix/2015 PT2015-51 09°14.988' 036°24.361' 703 06/x/2015 PT2015-26 09°15.357' 036°25.083' 562 28/ix/2015 PT2015-52 09°14.934' 036°24.310' 694 06/x/2015

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Figure 2. Map showing the area of the Pedra Talhada Biological Reserve, its vegetation profile and the location of the collection stations.

Distribution. Brazil (Bahia state) (Simone 2006). coastal states (Simone 2006, Salvador & Simone per- sonal observation). The present record slightly extends Remarks. There is some shell color variation in the present material, from nearly entirely brownish shells the species’ distribution to the north. (excluding the whitish columellar region and a whitish band below the keel; Fig. 3E) to specimens with alternate Helicina rotundata A.J. Wagner, 1910 colors (a brownish spire, a yellowish/whitish band above the keel, a brown band below the keel, and a yellowish/ Figure 3F, G whitish portion of the whorl, including the columellar Helicina rotundata Wagner 1910: 280, pl. 55, figs 15, 16. area; Fig. 3C, D). A similar variation in color pattern is Type locality. Brazil. known from a congener, H. variabilis J.A. Wagner, 1827, which is larger overall and inhabits the central Brazilian Distribution. Brazil (Paraíba state) (Simone 2006).

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Figure 3. A–E. Helicina angulifera. Scale bar = 5 mm. A, B. Live specimen (not to scale). C. MZSP 133717 (H = 7.1 mm, D = 9.5 mm). D. MZSP 133725, spm. #1 (H = 7.0 mm, D = 9.5 mm). E. MZSP 133725, spm. #2 (H = 7.3 mm, D = 10.2 mm). F, G. Helicina rotundata. Scale bar = 5 mm. F. Live specimen (not to scale). G. MZSP 133698 (H = 7.6 mm, D = 10.0 mm). H–J. Helicina schereri. Scale bar = 1 mm. H. Live specimen (not to scale). I. MZSP 133715 (H = 4.0 mm, D = 5.0 mm). J. Juvenile, MZSP 133721 (H = 2.5 mm, D = 3.4 mm).

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Remarks. This species can be distinguished from H. an- (Simone 2006, Simone & Salvador 2016), but its sculp- gulifera above by its more rounded shell profile and lack ture (prominent axial ribs and fine spiral cords) is easily of a keel on the body whorl. The present record slightly seen in fresh, well-preserved specimens (Fig. 4G, H). extends the species’ distribution to the south. The present record extends the species’ distribution to the north. Photographs of live specimens are published here Helicina schereri F.C. Baker, 1914 for the first time. Figure 3H–J Family Helicina schereri Baker 1914: 625, pl. 21, figs 1, 2. Genus Adelopoma Doering, 1885 Type locality. Brazil, Rio Grande do Norte state, Ceará- Mirim municipality. Adelopoma cf. brasiliense Morretes, 1954 Distribution. Brazil (Ceará, Rio Grande do Norte, Pa­ Figure 5A–C raíba, Tocantins and possibly Santa Catarina states) (Bickolz et al. 2016). Type locality. Brazil, São Paulo state, Iguape munici- pality, Jipovura neighborhood (“Jupuvura” in Morretes Remarks. The present specimens compare well to 1954). the lectotype (ANSP 109341) and topotype figured by Si­mone (2006). This species is identifiable by its small Distribution. Brazil (São Paulo and Paraná states) (Mor- size, round shell profile, small and round aperture, and retes 1954). the presence of hairs on the shell. The latter character, Remarks. In comparison with other Brazilian congeners, in particular, might indicate that this species actually A. brasiliense can be identified by its proportionately belongs in another genus, such as Alcadia J.E. Gray, 1840 smaller apex and the strong widely spaced ribs. Further- (see also Richling 2004). The present record is the first more, many specimens also show a small columellar from Alagoas state. lamella (e.g., Fig. 5B, C). The present specimens com- pare very well to this species, but given its distribution, Caenogastropoda they could belong to a distinct lineage and should be the Superfamily focus of future comparative studies.

Family Neocyclotidae

eschweizerbartxxx sng- Genus Aperostoma Troschel, 1847

Aperostoma blanchetiana (S. Moricand, 1836) Superfamily Figure 4A–F Family Cyclostoma Blanchetianum Moricand 1836: 442, pl. 2, figs 21–23. Genus Sarasinula Grimpe & Hoffmann, 1924 Type locality. Brazil, Bahia state, Cachoeira municipality. Sarasinula sp. Distribution. Brazil (Pará and Bahia states) (Simone 2006). Figure 5D–F Remarks. The present record of this easily recognizable Remarks. By the size and pigmentation, especially of species fills a gap in its distribution. the ventral portion, the present specimens seem to belong to Sarasinula. Veronicellidae identification is difficult Genus Cyclopomops Bartsch & Morrison, 1942 without very fine anatomical details and the family’s classification is in dire need of revisionary work based Cyclopomops moricandi (L. Pfeiffer, 1852) preferentially on molecular techniques. Therefore, we prefer to take our identification only to genus level. Figure 4G–L Cyclostoma disjunctum Moricand 1846 [non Matheron 1832]: Genus Latipes Colosi, 1922 158, pl. 5, figs 26–29. Cyclostoma Moricandi Pfeiffer 1852b: 64. Latipes sp. Brazil, Bahia state. Type locality. Figure 5G–I Distribution. Brazil (Bahia and Minas Gerais states) Remarks. A clear second morph of veronicellid (Birckolz et al. 2016). occurs in Pedra Talhada. Judging by the small size (1/2–2/3 Remarks. This species is often figured in an eroded of Sarasinula sp.) and the pigmentation pattern, includ- (and thus nearly unsculptured) state in the literature ing the dark ventral region, the present specimen likely

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Figure 4. A–F. Aperostoma blanchetiana. Scale bar = 5 mm. A–C. Live specimen (not to scale). D–F. MZSP 133738 (H = 13.5 mm, D = 22.8 mm). G–L. Cyclopomops moricandi. Scale bar = 2 mm. G, H. Live specimen (not to scale). I. MZSP 133732 (H = 3.7 mm, D = 5.5 mm). J–L. MZSP 133727 (H = 3.9 mm, D = 6.2 mm).

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Figure 5. A–C. Adelopoma cf. brasiliense. Scale bar = 0.5 mm. A. Live specimen (not to scale). B. MZSP 133653 (H = 2.2 mm). C. MZSP 133651 (H = 2.3 mm). D–F. Sarasinula sp., live specimen. G–I. Latipes sp., live specimen.

108 Salvador et al. · Land snails from Pedra Talhada belongs to the genus Latipes. Nevertheless, Simrothula forms might occur together, and thus, considered them Thomé, 1975 cannot be completely excluded. synonymous. To our knowledge, this is the first record for Alagoas state. Stylommatophora Family Superfamily Succineoidea Genus Bothriopupa Pilsbry, 1898 Family Succineidae

Genus Omalonyx d’Orbigny, 1837 Bothriopupa breviconus Pilsbry, 1917 Figure 6G, H Omalonyx sp. Bothriopupa breviconus Pilsbry 1917: 230, pl. 28, figs 9, 10. Figure 6A–C Type locality. Guatemala, mountains west of Livingston. Remarks. There are 2 species of Omalonyx reported from northeastern Brazil, including Alagoas state (Arruda et Distribution. Guatemala and French Guiana (Pilsbry al. 2009, 2016), O. geayi Tillier, 1980, and O. matheroni 1917, Massemin et al. 2009). (Potiez & Michaud, 1835). As these species are very Remarks. This species is mainly diagnosable, especially widespread and South American Omalonyx still lack a from the similar B. conoidea (Newcomb, 1853) from comprehensive taxonomic work defining and diagnosing Venezuela, Suriname, and French Guiana, by the pres- all species, we prefer to take the identification of the pres- ence of the upper palatal lamella (Massemin et al. 2009). ent specimens to genus level only. However, this lamella may be only very weakly marked in some specimens (e.g., Fig. 6G), and thus, an absent Superfamily lamella may not necessarily signify a distinct species. Family Strobilopsidae Other vertiginids and gastrocoptids are known to vary intraspecifically in the presence and strength of their Genus Strobilops Pilsbry, 1893 apertural lamellae. The present record is the first for this species (and genus) in Brazil and greatly extends its geo- Strobilops cf. brasiliana F.C. Baker, 1914 graphical distribution. Figure 6D, E

eschweizerbartxxx sng- Genus Sterkia Pilsbry, 1898 Type locality. Brazil, “City of Pará” (Baker 1914).

Distribution. Brazil (Pará state) (Simone 2006). Sterkia eyriesii (Drouët, 1859) Remarks. The 2 available specimens, in their overall Figure 6I, J conical-discoid shell shape and the axial ribs (strongly Pupa Eyriesii Drouët 1859: 367, pl. 2, figs 16, 17. prosocline and widely spaced), are consistent with the single strobilopsid reported from Brazil, S. brasiliana. Type locality. French Guiana, îlets de Rémire, îlet la Nevertheless, as the present shells are juveniles and Mère. apertural characters are taxonomically important, it is cur- Distribution. Florida to Suriname and French Guiana rently impossible to achieve a more precise identification. (Massemin et al. 2009).

Family Remarks. Similar to Bothriopupa breviconus, S. eyrie- sii is known from northern areas of South America, but Genus Pupisoma Stoliczka, 1873 the present record is the first of the genus for Brazil. The species can be diagnosed by the shape of its whorls (the Pupisoma dioscoricola (C.B. Adams, 1845) first whorls are seemingly flattened and truncated when Figure 6F compared to the rest of the shell) and by the shape and positioning of the apertural dentition. Helix dioscoricola Adams 1845: 16. Type locality. Jamaica. Superfamily Distribution. Circumtropical (Hausdorf 2007). Family Remarks. This species can be recognized by its propor- Genus Bulimulus Leach, 1814 tionately large first whorls and the delicate spiral striae on the teleoconch (Hausdorf 2007). Specimens with axial Bulimulus tenuissimus (d’Orbigny, 1835) riblets, such as those from Pedra Talhada, were tradition- ally regarded as a subspecies, P. d. insigne Pilsbry, 1920. Figure 7A Hausdorf (2007), however, demonstrated that both Helix tenuissima d’Orbigny 1835: 11.

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Figure 6. A–C. Omalonyx sp. Scale bar = 0.5 mm. A. Live specimen (not to scale). B, C. MZSP 133984 (specimen length = 10.4 mm, width = 4.5 mm). D, E. Strobilops cf. brasiliana (MZSP 133864; H = 2.1 mm, D = 2.8 mm). Scale bar = 1 mm. F. Pupisoma dioscoricola (MZSP 133985; H =2.1 mm, D = 1.9 mm). Scale bar = 1 mm. G, H. Bothriopupa breviconus. Scale bar = 0.5 mm. G. MZSP 133992 (H = 1.7 mm). H. MZSP 133997 (H = 1.8 mm). I, J. Sterkia eyriesii. Scale bar = 0.5 mm. I. MNHN spm. #1 (H = 1.4 mm). J. MNHN spm. #2 (H = 1.6 mm).

Type locality. Brazil, Rio de Janeiro. Remarks. The present record stems from the 1998 expe- Distribution. Lesser Antilles (B. t. eyriesii Drouët dition and it is the first report of this species for Alagoas 1859), Suriname, French Guiana, Brazil, Bolivia and state. Bulimulus tenuissimus has been introduced to vari- Uruguay (Massemin et al. 2009, Birckolz et al. 2016). ous localities, with its northernmost record being North

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Figure 7. A. Bulimulus tenuissimus (MNHN; H = 14.5 mm). Scale bar = 5 mm. B, C. Drymaeus flexilabris. Scale bar = 10 mm. B. MZSP 133540 (H = 28.8 mm). C. Live specimen (not to scale). D–F. Drymaeus papyraceus. Scale bar = 10 mm. D. MZSP 133525 (H = 39.5 mm). E, F. Live specimen (not to scale). G, H. Drymaeus rufolineatus. Scale bar = 10 mm. G. MZSP 133551 (H = 18.0 mm). H. Live specimen (not to scale). I. Naesiotus sp. (MZSP 133556; H = 15.3 mm). Scale bar = 5 mm. J, K. Oxychona bifasciata, live specimens.

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Carolina, USA (Robinson & Slapcinsky 2005). Given cioi (Rezende & Lanzieri, 1963), known from central Bra- the species distribution and the status of Pedra Talhada as zil (Tocantins and Goiás states and the Federal District; a reserve, the present record would more likely represent Birckolz et al. 2016), but have a much taller aperture. a natural occurrence of the species. They also differ in the shell’s overall coloration, which seems to show some sort of light brown pattern. This pat- Genus Drymaeus Albers, 1850 tern is not fully discernible due to the poor preservation of the shells, but is reminiscent of N. rivasii (d’Orbigny, Drymaeus flexilabris (L. Pfeiffer, 1853) 1837), a species known from Bolivia and Argentina Figure 7B, C (Simone 2006, Breure & Ablett 2014). The present material could belong to a still undescribed species, but Bulimus flexilabris Pfeiffer 1853: 50 the damaged condition of the shells calls for a more cau- Type locality. Brazil. tionary identification as simplyNaesiotus sp. Distribution. Brazil (Pernambuco and Alagoas states) (Simone 2006, Salvador personal observation for Per- Genus Oxychona Mörch, 1852 nambuco: NHMUK 1888.6.30.5-7). Pilsbry (1898) also indicated, with doubts, the region of the Amazon river. Oxychona bifasciata (Burrow, 1815) Remarks. In some specimens, there is a vestige of a Figures 7J, K, 8A–C light-red color on the apex of the spire (first 3–4 whorls). Trochus bifasciatus Burrow 1815: 177, pl. 27, fig. 2.

Drymaeus papyraceus (Mawe, 1823) Type locality. Brazil, Pernambuco state. Figure 7D–F Distribution. Brazil (Pernambuco, Bahia, and Minas Gerais states) (Simone 2006). Helix Papyracea Mawe 1823: 168, frontispiece fig. 7. Remarks. There is some variation in the color pattern Type locality. Brazil, Rio de Janeiro. of the shells in the present material: the diagnostic twin Distribution. Brazil (widely distributed), Paraguay, Uru- spiral bands vary in width down to very narrow stripes guay and Argentina (Simone 2006). (Fig. 8A); the upper (adapical) band may be absent (Fig. 7J); the typically pink peristome may be white (Fig.

Drymaeus rufolineatus (Droüet, 1859) eschweizerbartxxx sng- 8A–C); and the whole shell may have a more brownish hue instead of a whitish one (Fig. 7K). This variation is Figure 7G, H also observed in specimens from museum collections. Bulimus rufolineatus Droüet 1859: 357, pl. 1, figs 10, 11. The present record fills a gap in the species’ distribution. Type locality. French Guiana, îlets de Rémire, îlet la Mère. Family Distribution. Guyana, Suriname, French Guiana, and Brazil (Pernambuco state) (Simone 2006, Massemin et Genus Biotocus Salgado & Leme, 1990 al. 2009). Remarks. This species can be recognized by its propor- Biotocus turbinatus (L. Pfeiffer, 1845) tionately wide aperture, the somewhat translucent shell, Figure 8D–H and the parallel brown stripes that give the species its name. Similar to the vertiginid species listed above, this Tomogeres turbinatus Pfeiffer 1845: 45. Guianan species also occurs in Brazil, with a record from Type locality. Brazil. Pernambuco state (Simone 2006). The present record in Alagoas state extends the species’ range to the south. Distribution. Brazil (Alagoas and Bahia states) (Simone 2006). Genus Naesiotus Albers, 1850 Remarks. This species was considered extinct (with- out further explanation) in the IUCN Red List (Mansur Naesiotus sp. 1996b). However, it was found alive in Pedra Talhada Figure 7I under humid tree trunks during the 1998 expedition; empty shells were collected on that occasion and are the Remarks. The few specimens available are in poor con- specimens reported here. Neither live animals nor empty dition, but the protoconch is observable in one of them, shells were found in the 2015 campaign. consisting of closely placed sinuous axial riblets with fine spiral lines between them. This feature is consis- Family Simpulopsidae tent with the genus Naesiotus (Breure 1979, Breure & Ablett 2014). The present specimens resemble N. calu- Genus Rhinus E. von Martens in Albers, 1860

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Figure 8. A–C. Oxychona bifasciata. Scale bar = 10 mm. A, B. MZSP 133523 (H =21.0 mm, D = 21.1 mm). C. MZSP 133548 (H = 20.0 mm, D = 21.3 mm). D–H. Biotocus turbinatus. Scale bar = 5 mm. D–G. MNHN spm. #1 (H = 9.9 mm, D = 11.4 mm). H. MNHN spm. #2 (H = 9.8 mm, D = 11.8 mm). I, J. Rhinus botocudus. Scale bar = 5 mm. I. Live specimen (not to scale). J. MZSP 133841 spm. #1 (H = 21.0 mm, D = 16.3 mm).

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Rhinus botocudus Simone & Salvador, 2016 and more defined spire. The present record is from the Figure 8I, J 1998 expedition and expands the species’ distribution to the north. Rhinus botocudus Simone & Salvador 2016: 20, figs 28–52, 68–76. Superfamily Type locality. Brazil, Minas Gerais state, Nanuque munic- ipality (c. 120 m of elevation, c. 17° 51′ S, 040° 23′ W). Family Ferussaciidae Distribution. Known only from type locality (Simone & Genus Cecilioides A. Férussac, 1814 Salvador 2016). Cecilioides consobrina (d’Orbigny, 1841) Remarks. This species has been recently described (including a detailed account of its anatomy) from a frag- Figure 9H, I ment of Atlantic Forest in northern Minas Gerais state Achatina consobrina d’Orbigny 1841: 170. (Simone & Salvador 2016). The present record extends the species’ distribution circa 1,000 km to the northeast. Type locality. Cuba, near Matanzas. Distribution. From Mexico to Argentina (Sandoval Genus Simpulopsis Beck, 1837 1997, Miquel et al. 2007b, Salvador et al. 2017). Remarks. Despite the wide distribution of this species, Simpulopsis atrovirens (S. Moricand, 1836) up to our knowledge this is the first specific record from Figure 9A–C Alagoas state. The large range of C. consobrina might be Helix (Cochlohydra) atrovirens Moricand 1836: 416, pl. 2, fig. 1. a sign that this is actually a species complex. Their small size, fragile shells, subterranean habits, and low abun- Type locality. Brazil, Bahia state, Portas (see also Pils- dance make the species hard to study; cases such as this bry 1899). are where molecular studies are most helpful. Distribution. Known only from the type locality (Si­mone 2006). Family Subulinidae

Remarks. This species can be distinguished from the Genus Allopeas H.B. Baker, 1935 other 2 Simpulopsis from Pedra Talhada by its small size, yellowish coloration and quickly expanding whorls.eschweizerbartxxx sng- The Allopeas micrum (d’Orbigny, 1835) present record extends the species’ distribution to the north. Figure 9J Simpulopsis rufovirens (S. Moricand, 1846) Helix micra d’Orbigny 1835: 9 Bulimus micra d’Orbigny 1837: 262, pl. 41, figs 18, 19. Figure 9D, E Type locality. Brazil, Rio de Janeiro. Helix (Succinea) rufovirens Moricand 1836: 147, pl. 5, fig. 4. Distribution. Florida to southern Brazil, but likely intro- Type locality. Brazil, Bahia state. duced in many places (Delannoye et al. 2015). Distribution. Brazil (Bahia, Espírito Santo and Rio de Remarks. The presence and strength of the axial sculp- Janeiro states) (Simone 2006). ture is known to vary widely in this species (Pilsbry Remarks. Compared to other Simpulopsis species from 1946); all specimens from Pedra Talhada display axial Pedra Talhada, this species can be diagnosed by its much sculpture. larger size, orange-brown coloration and large aperture. The present record, from the 1998 expedition, slightly Genus H.B. Baker, 1961 expands the species’ range to the north. (L. Pfeiffer, 1846) Simpulopsis sulculosa (A. Férussac, 1821) (Figs 9K–M) Figure 9F, G Bulimus Beckianus Pfeiffer, 1846: 82. Helix (Cochlohydra) sulculosa Férussac 1821: 31, pl. 11A, fig. 6. Type locality. Polynesia, Opara Island (Pfeiffer 1846), Type locality. Brazil. but Pilsbry (1906) considered this is a mistaken attribu- Distribution. Brazil (Minas Gerais, Rio de Janeiro, São tion, stating that Pfeiffer’s specimens are very similar to Paulo, Paraná, Santa Catarina, and Rio Grande do Sul those from Central America. states) (Birckolz et al. 2016). Distribution. From Mexico to Venezuela, West Indies, Remarks. In comparison with the two Simpulopsis above, Guyana, Suriname, Peru, Brazil (Roraima, Pará, Rio this species can be diagnosed by its proportionately taller Grande do Norte, Bahia, Rio de Janeiro, and São Paulo

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Figure 9. A–C. Simpulopsis atrovirens. Scale bar = 5 mm. A, B. Live specimen (not to scale). C. MZSP 133842 (H = 9.4 mm, D = 9.9 mm). D, E. Simpulopsis rufovirens (MNHN; H = 17.2 mm, D = 16.0 mm). Scale bar = 5 mm. F, G. Simpulopsis sulculosa (MNHN; H = 11.7 mm, D = 10.1 mm). Scale bar = 5 mm. H, I. Cecilioides consobrina. Scale bar = 0.5 mm. H. Live specimen (not to scale). I. MZSP 133689 (H = 2.0 mm). J. Allopeas micrum (MZSP 133955; H = 4.9 mm). Scale bar = 2 mm. K–M. Beckianum beckianum. Scale bar = 2 mm. K. MZSP 133957 (H = 6.8 mm). L, M. Live specimens (not to scale).

115 Archiv für Molluskenkunde · 147 (1) 2018 states; Fernando de Noronha Archipelago) (Birckolz et Obeliscus agassizi Pilsbry 1906: 249, pl. 36, fig. 76. al. 2016). Type locality. Brazil (Agassiz expedition). Remarks. This widespread species is here recorded from Distribution. Brazil (Paraíba state) (Simone 2006). Alagoas state for the first time. Remarks. The present record extends the species’ distri- Genus Leptinaria Beck, 1837 bution slightly to the south.

Leptinaria cf. mamoreensis F.C. Baker, 1926 Genus Stenogyra Shuttleworth, 1854 Figure 10A Stenogyra octogyra (L. Pfeiffer, 1856) Type locality. Brazil, Rondônia state, 284 km from Porto Figure 10I, J Velho along the Madeira–Mamoré Railroad. Bulimus octogyrus Pfeiffer 1856: 45. Distribution. Venezuela and Brazil (Rondônia state) (Simone 2006). Type locality. Venezuela, Caracas. Remarks. The overall shell profile, aperture shape, tele- Distribution. Venezuela and Brazil (Pará, Ceará, Rio oconch sculpture, umbilicus and reflexed peristome are Grande do Norte, Goiás, Bahia, Mato Grosso, and Minas consistent with the original description and illustration of Gerais states) (Salvador et al. 2017, Salvador & Simone L. mamoreensis (Baker 1926). However, since this spe- personal observation from Minas Gerais). cies’ range is geographically very disconnected from Pedra Remarks. The present records fill a gap in the species’ Talhada (and within the Amazonian Rainforest biome), distribution. the present identification must remain tentative for now. The present material stems from the 1998 expedition. Superfamily Streptaxoidea

Leptinaria ritchiei Pilsbry, 1907 Family

Figure 10B–D Genus Streptaxis Gray, 1837 Leptinaria ritchiei Pilsbry 1907: 304, pl. 46, fig. 12. Streptaxis iheringi (Pilsbry, 1930)

Type locality. Brazil(?), Ituchy, on Purus river eschweizerbartxxx(Pilsbry sng- 1907). Figure 10K–M Distribution. Brazil (Alagoas state) (Simone 2006). Artemon iheringi Pilsbry 1930: 362, pl. 32, figs 6, 6a, 6b. Remarks. Despite the uncertainty regarding the type Type locality. Brazil, “from the state of São Paulo or locality, the present specimens compare very closely southward” (Pilsbry 1930). to the lectotype (ANSP 24106). Simone (2006) already Distribution. Known only from the holotype (ANSP reported the species for Alagoas state, from Barra de São 151896, Simone 2006). Miguel municipality, an area of Atlantic Forest. Remarks. By the shell size, the regular and somewhat Leptinaria lamellata (Potiez & Michaud, 1835) step-like spire and the more central position of the aper- ture, the present specimens compare very closely to the Figure 10E, F holotype of S. iheringi. Even though there is uncertainty Helix unilamellata d’Orbigny 1835: 9 [nomen nudum]. about the type locality of this species, the present record Achatina lamellata Potiez & Michaud 1835: 128, pl. 11, figs 7, 8. represents a large range extension northwards. Type locality. Undefined. Superfamily Rhytidoidea Distribution. West Indies (introduced), Colombia, Vene- zuela, Guyana, French Guiana, Brazil and Bolivia (Simone Family 2006, Massemin et al. 2009). Genus Happia Bourguignat, 1889 Remarks. A widespread species in South America, reach- ing southern Brazil (Simone 2006). The parietal lamella Happia vitrina (J.A. Wagner in Spix, 1827) is usually not present in specimens from Pedra Talhada. Figure 11A–C Genus Obeliscus Beck, 1837 Helix vitrina J.A. Wagner in Spix 1827: 25, pl. 17, fig. 6. Solarium imperforatum Spix 1827: pl. 17, fig. 6. Obeliscus agassizi Pilsbry, 1906 Type locality. Brazil, “southern provinces” (Wagner in Figure 10G, H Spix 1827).

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Figure 10. A. Leptinaria cf. mamoreensis (MNHN; H = 7.1 mm). Scale bar = 2 mm. B–D. Leptinaria ritchiei. Scale bar = 5 mm. B. MZSP 133929 (H = 11.9 mm). C. MZSP 133905 (H = 9.5 mm). D. Live specimen (not to scale). E, F. Leptinaria lamellata. Scale bar = 2 mm. E. Live specimen (not to scale). F. MNHN (H = 9.8 mm). G, H. Obeliscus agassizi. Scale bar = 10 mm. G. Live specimen (not to scale). H. MZSP 133507 (H = 34.0 mm). I, J. Stenogyra octogyra. Scale bar = 2 mm. I. MZSP 133966 (H = 9.1 mm). J. Live specimen (not to scale). K–M. Streptaxis iheringi (MZSP 133857; H = 13.3 mm, D = 15.6 mm). Scale bar = 5 mm.

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Figure 11. A–C. Happia vitrina (MZSP 133787; H = 4.4 mm, D = 2.3 mm). Scale bar = 2 mm. D–G. banghaasi. Scale bar = 2 mm. D–F. MZSP 133769 (H = 2.2 mm, D = 4.8 mm). G. Live specimen (not to scale). H–J. bounobaena (MZSP 133785; H = 3.7 mm, D = 6.1 mm). Scale bar = 2 mm. K–O. Scolodonta spirorbis. Scale bar = 0.5 mm. K. Live specimen (not to scale). L. MZSP 133803 (H = 0.8 mm, D = 1.6 mm). M. MZSP 133773 (H = 1.0 mm, D = 2.0 mm). N. MZSP 133766 (D = 1.9 mm). O. MZSP 133832 (D = 1.8 mm).

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Distribution. Brazil (Bahia, Rio de Janeiro, São Paulo, Superfamily Acavoidea Paraná, and Santa Catarina states) (Birckolz et al. 2016). Family Strophocheilidae Remarks. The present record slightly extends the spe- Genus Megalobulimus K. Miller, 1878 cies’ range to the north.

Megalobulimus cardosoi (Morretes, 1952) Genus Tamayoa H.B. Baker, 1925 Figure 12A, B

Tamayoa banghaasi (Thiele, 1927) Strophocheilus (Megalobulimus) cardosoi Morretes 1952: 119, pl. 3, fig. 3, pl. 4, fig. 3. Figure 11D–G Type locality. Brazil, Alagoas state, Murici. Happia banghaasi Thiele 1927: 319, pl. 26, fig. 13a, b, text-fig. 4. Distribution. Known only from the type locality (Simone Type locality. Brazil, Pernambuco state, Olinda munici- 2006). pality; and Espírito Santo state. Remarks. This is the first record of the species outside its Distribution. Brazil (Paraíba, Pernambuco, Minas Gerais, type locality (also an area of Atlantic Forest), extending Espírito Santo and Rio de Janeiro states) (Santos & its distribution a few km to the west. However, more than Monteiro 2001, Simone 2006, personal observation merely extending the range, the present specimens are from Minas Gerais). important because M. cardosoi was considered extinct, without explanation, in the IUCN Red List (Mansur Remarks. Similar to what is reported by Santos & 1996a); in the Brazilian official list of threatened fauna Monteiro (2001), in Pedra Talhada this species can be (MMA 2014), however, the species is listed as critically very abundant and predominant. The present record fills endangered. Megalobulimus cardosoi was not found alive a small gap in the species’ known distribution. during the 2015 expedition, but the good preservation of the shells in a metamorphic acidic environment suggests Genus Scolodonta Doering, 1875 freshly dead animals. As such, at least the population of M. cardosoi from Pedra Talhada seem still to be thriving, Scolodonta bounobaena (d’Orbigny, 1835) but further studies focused on this species are necessary Figure 11H–J to know its actual distribution and the state of the popula-

eschweizerbartxxx sng- tion according to the IUCN guidelines. Helix bounobæna d’Orbigny 1835: 7.

Type locality. Bolivia, “Chiquitensi Province” (d’Orbigny Megalobulimus oliveirai (Bequaert, 1948) 1835), which probably refers to “Chiquitos”, the colonial Figure 12C, D name of a part of what is nowadays the Chiquitania region (see Breure & Ablett 2014 for similar cases). Strophocheilus terrestris oliveirai Bequaert 1948: 117, pl. 5, fig. 2. Distribution. Bolivia (Chiquitos province) and Brazil Type locality. Brazil, Sergipe state, Propriá municipality. (Morretes 1949, Simone 2006). Distribution. Known only from the type locality (Simone 2006). Scolodonta spirorbis (A. Férussac in A. Férussac & Remarks. This species is easily distinguishable from M. Deshayes, 1850) cardosoi by its larger size and much taller spire. The pres- Figure 11K–O ent record slightly expands the species’ distribution to the Helix spirorbis Férussac 1850 in Férussac & Deshayes 1850: 83, northeast. pl. 82A, figs 1–3. Genus Strophocheilus Spix, 1827 Type locality. Brazil, Rio de Janeiro.

Distribution. Brazil (Rio de Janeiro state) (Simone Strophocheilus debilis Bequaert, 1948 2006). Figure 12E–H Remarks. The present specimens compare very closely Strophocheilus pudicus debilis Bequaert 1948: 34, pl. 16, fig. 1, to the syntype (MNHN-IM-2000-31791) in the convex pl. 17, fig. 2. whorl profile, depressed spire, well-marked suture and Type locality. Brazil, Bahia state, Itapicuru river. a very delicate sculpture consisting of prosocline spiral ribs. Nevertheless, in the present specimens the aperture Distribution. Brazil (Alagoas and Bahia states) (Simone 2006). is less laterally elongated and thus more rounded. The present record greatly extends the species’ distribution to Remarks. This species can be diagnosed from its con- the north. geners by its narrower shell, a more bulbous penultimate

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Figure 12. A, B. Megalobulimus cardosoi (MZSP 133892; H = 55.3 mm, D = 33.7 mm). Scale bar = 20 mm. C, D. Megalobuli- mus oliveirai (MZSP 133898; H = 99.5 mm, D = 62.1 mm). Scale bar = 20 mm. E–H. Strophocheilus debilis. Scale bar = 20 mm. E, F. Live specimen (not to scale). G, H. MZSP 133883 (H = 51.1 mm, D =24.6 mm). I–L. Lilloiconcha gordurasensis. Scale bar = 1 mm. I. Live specimen (not to scale). J, K. MZSP 133638 (D = 2.9 mm). L. MZSP 133645 (H = 2.0 mm, D = 3.1 mm).

120 Salvador et al. · Land snails from Pedra Talhada whorl and a shorter and less conical spire. The shell color Radiodiscus ubtaoi sp. nov. ranges from yellowish brown to strong reddish brown. Figure 13I–L During the 2015 expedition, this species was the most common macrogastropod found living in the reserve. The ZooBank registration. urn:lsid:zoobank.org:act:F7784 species typically inhabits the Atlantic Forest, but its distri- E37-0728-4F90-B904-15FE26936C61 bution in Bahia might include areas of the Caatinga biome. Type material. Holotype: MZSP 133514 (Figs 13I, K). Paratypes: MZSP 133510 (Fig. 13J), 2 shells, from sta. 23 Superfamily (PT2015-23); MZSP 133514b (Fig. 13L), 5 shells, from Family Charopidae type locality.

Genus Lilloiconcha Weyrauch, 1965 Type locality. Brazil, Alagoas state, Pedra Talhada Bio- logical Reserve (Reserva Biológica de Pedra Talhada), Lilloiconcha gordurasensis (Thiele, 1927) sta. 31 (PT2015-31), 09° 14.318′ S, 036° 25.040′ W, 870 m a.s.l. Figure 12I–L Distribution. Known only from Pedra Talhada Biologi- Endodonta gordurasensis Thiele 1927: 322, pl. 26, fig. 19. cal Reserve. Type locality. Brazil, Minas Gerais state, Gorduras. Etymology. The name refers to Ubtao, a fictional god Distribution. Brazil (NE region and Minas Gerais state), from the Faerûnian pantheon of the Forgotten Realms Bolivia, Paraguay and Argentina (Miquel et al. 2004, campaign setting of the Dungeons and Dragons role- 2007a, Cunha et al. 2015). playing game. Ubtao is a god of nature who presides over the untamed jungles; his symbol is a circular maze seen Remarks. Dubious records include Colombia and Peru from above, which is reminiscent of the strongly sculp- (Hausdorf 2005) and a Middle Miocene record from tured shell of this new species, when seen in apical view Patagonia (Miquel & Bellosi 2007). The record from under SEM. Trindade Island, southeast Brazil (Cunha et al. 2015: figs 2A–C), is not L. gordurasensis, as seen by the presence of Diagnosis. Protoconch with 7–9 widely spaced spi- protoconch sculpture in their specimens. ral cordlets. Teleoconch with closely spaced and not very sinuous axial ribs. Aperture proportionately small, Lilloiconcha superba (Thiele, 1927) D-shaped.

eschweizerbartxxx sng- Figure 13A–D Description. Shell minute (D c. 1.5 mm), discoid, with slightly raised spire. Whorl profile greatly convex; suture Endodonta superba Thiele 1927: 322, pl. 26, fig. 23. deeply marked. Protoconch (c. 1¼ whorl) round, sculp- Type locality. Brazil, Rio de Janeiro state, Teresópolis tured with 7–9 widely spaced spiral cordlets; transition municipality. to teleoconch clearly marked by change to teleoconch sculpture. Teleoconch sculptured with closely spaced and Distribution. Brazil (NE region to Rio de Janeiro state) weakly sinuous axial ribs; space between ribs typically (Simone 2006). 2× width of rib (but up to 4× in some specimens); space Remarks. This species is easily diagnosable by its high between ribs filled by axial riblets and spiral cordlets, and conical shell and lightly marked angulation in the forming a reticulated pattern. Aperture D-shaped; peri- median portion of the whorl (a more prominent keel-like stome simple. Umbilicus very wide. angulation is seen in juvenile specimens; Fig. 13C). Remarks. The presence of protoconch sculpture con- sisting of spiral cordlets place this species in the genus Genus Radiodiscus Pilsbry in Pilsbry & Ferriss, 1906 Radiodiscus (Miquel et al. 2007a). The teleoconch sculpture consisting of strong axial ribs and a reticulate Radiodiscus amoenus (Thiele, 1927) pattern between the ribs is also very usual in the genus Figure 13E–H (Miquel et al. 2007a). The present specimens are suf- ficiently distinct in conchological characters (and also Endodonta amoena Thiele 1927: 323, pl. 26, fig. 27. geographically distant) from its congeners, that we are Type locality. Brazil. confident in describing them as belonging to a new spe- cies: Radiodiscus ubtaoi sp. nov. Distribution. Brazil, Argentina and possibly Uruguay In overall shell shape, size, whorl profile and aperture (Miquel et al. 2007a). shape, Radiodiscus ubtaoi sp. nov. is very reminis- Remarks. This species can be distinguished from its cent to R. sanchicoensis Miquel et al., 2007, a species congeners by its taller spire, narrower umbilicus, a some- known only from its type locality in southernmost Bra- what more globular shell, and a protoconch sculptured by zil (Miquel et al. 2007a). Nevertheless, Radiodiscus about 10–12 fine spiral cordlets. ubtaoi sp. nov. can be easily distinguished from it by the

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Figures 13. A–D. Lilloiconcha superba. Scale bar = 1 mm. A. MNHN spm. #1 (H = 3.2 mm, D = 3.1 mm). B. MNHN spm. #2 (D = 2.5 mm). C. MZSP 133517, juvenile (D = 2.3 mm). D. Live specimen (not to scale). E–H. Radiodiscus amoenus. Scale bar = 0.5 mm. E. Live specimen (not to scale). F. MZSP 133640 (H = 1.7 mm, D = 2.4 mm). G. MZSP 133641 (D = 1.8 mm). H. Same, magnified view of the protoconch. Scale bar = 0.1 mm. I–L. Radiodiscus ubtaoi sp. nov. Scale bar = 0.5 mm. I. Holotype, MZSP 133514, magnified view of the protoconch. Scale bar = 0.1 mm.J. Paratype, MZSP 133510 (H = 0.7 mm, D = 1.3 mm). K. Holotype (D = 1.3 mm). L. Paratype, MZSP 133514b (D = 1.1 mm).

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Figure 14. A–C. Pseudoguppya semenlini. Scale bar = 2 mm. A. Live specimen (not to scale). B. Juvenile, MZSP 133667 (H = 3.0 mm, D = 3.9 mm). C. MZSP 133666 (H = 3.9 mm, D = 4.8 mm). D–F. Solaropsis pascalia (MZSP 133744; H = 16.6 mm, D =32.4 mm). Scale bar = 10 mm. G–J. Solaropsis cearana. Scale bar = 5 mm. G–I. MNHN (H = 5.8 mm, D = 10.6 mm). J. Live specimen (not to scale). K–M. Examples of shells bearing signs of predation. Scale bar = 10 mm. K. Drymaeus flexilabris, MZSP 133535 spm. #2 (H = 29.1 mm). L. Oxychona bifasciata, MZSP 133534 spm. #1 (D =22.7 mm). M. Oxychona bifasciata, MZSP 133524 spm. #2 (D =20.9 mm).

123 Archiv für Molluskenkunde · 147 (1) 2018 protoconch sculpture: it bears 7–9 spiral cordlets (Fig. Solaropsis cearana (F.C. Baker, 1914) 13I), while R. sanchicoensis bears circa 25 (Miquel et Figure 14G–J al. 2007a). Moreover, Radiodiscus ubtaoi sp. nov. has a more rounded whorl profile overall, a proportionately Psadara derbyi cearana Baker 1914: 634, pl. 22, fig. 19. smaller and more D-shaped aperture, and less sinuous Type locality. Brazil, Ceará state, Maranguape munici- axial ribs on the teleoconch. pality. Radiodiscus ubtaoi sp. nov. can be easily distin- guished from other congeners mainly by its D-shaped Distribution. Brazil (Ceará state) (Simone 2006). aperture and closely spaced and less sinuous axial ribs. Remarks. A second species of Solaropsis can be found Actually, the present specimens show some variation in in Pedra Talhada, easily distinguishable by its strongly teleoconch sculpture: usually the shell is sculptured by depressed spire and comparatively smaller size. These closely spaced axial ribs, which are only faintly sinu- were considered diagnostic features of the genus (or ous, but some specimens show more sinuous and widely subgenus) Psadara K. Miller, 1878, which is presently spaced ribs, with nearly double the spacing of typical considered a synonym of Solaropsis (Cuezzo 2002, shells—a configuration that is more typical of the genus; 2003). The present specimens display shells of a single e.g., Simone 2006, Miquel et al. 2007a. color (i.e., without the common color patterns seen in the genus), with a large circular aperture, features consistent Superfamily Gastrodontoidea with S. cearana (ANSP 109344, lectotype). However, the only adult shell available has a slightly more elevated Family Euconulidae spire and the aperture a little more abapically positioned. Genus Pseudoguppya H.B. Baker, 1925 Live adults were only observed during the 1998 expe- dition. The present record greatly expands the species’ Pseudoguppya semenlini (S. Moricand, 1846) distribution to the southeast. Figure 14A–C Helix Semen-lini Moricand 1846: 149, pl. 5, fig. 17. Discussion Type locality. Brazil, Bahia state. Distribution. Brazil (Bahia, Rio de Janeiro, São Paulo, In total, 47 species of land snails were found in the Pedra Paraná, Santa Catarina, and Rio Grande do Suleschweizerbartxxx states), sng- Talhada Biological Reserve, distributed among 19 fam- Paraguay, Uruguay, Argentina (Simone 2006). ilies, mainly of “”. The Subulinidae are the most abundant and diverse group (Table 2), followed in Remarks. The present record slightly extends the spe- abundance by the Scolodontidae and Charopidae, but in cies’ distribution to the north. species richness by the Bulimulidae. As many specimens were juveniles and could not be confidently assigned to Superfamily Helicoidea a species (especially in the Charopidae and Scolodonti- Family dae), we refrained from calculating diversity indices, as they would inevitably be biased. Five species present in Genus Solaropsis Beck, 1837 the material from the 1998 expedition could not be found in the 2015 expedition; they are: Biotocus turbinatus, Solaropsis pascalia (Cailliaud, 1857) Bulimulus tenuissimus, Leptinaria cf. mamoreensis, Sim- Figure 14D–F pulopsis rufovirens and Simpulopsis sulculosa. This is likely due to the collections taking place during different Helix pascalia Cailliaud 1857: 102, pl. 2, fig. 3. seasons, but in the case of Biotocus turbinatus this might Type locality. “Brazilian countryside” (Cailliaud 1857). not be so: this species has been listed as extinct (Man- sur 1996b). Finally, it is also worthwhile to note that Distribution. Brazil (Pará and Alagoas states) (Simone some species were observed only on the outskirts of the 2006). reserve, but not within it, namely: Gastrocopta oblonga Remarks. In this species, the shell commonly displays a (L. Pfeiffer, 1852), Orthalicus prototypus Pilsbry, 1899, creamy light-brown coloration, with darker brown spiral and Streptaxis cf. costulosus (L. Pfeiffer, 1852). bands adjacent to the suture and keel (e.g., Simone 2006: Several of the species were reported here for the first fig. 928). In the present material, however, the shell pos- time from Alagoas state, extending the geographical sesses an overall darker brown color and even darker spiral range of some or filling distributional gaps of others. This bands. Despite not finding live specimens, all of the fresh should not come as a surprise, as the terrestrial fauna of shells were collected between the leaves of Bromeliaceae Alagoas has scarcely been scrutinized by malacologists. or on the floor, on the inselbergs. The very few specimens The most notable new records are the genera Bothrio­ occurring on the slopes of the reserve were always worn. pupa and Sterkia (Vertiginidae), reported here for the first

124 Salvador et al. · Land snails from Pedra Talhada

Table 2. Total and relative abundance of each family of ter- Table 3. Shells of the larger species showing sign restrial gastropod in the Pedra Talhada Biological Reserve, of predation (including material from the 1998 expedition); the alongside the number of species. This table considers only values represent the total number of specimens for each case the material from the 2015 collection (including specimens and the proportion. identified only to family or genus level); the material from the 1998 expedition would add 1 species of Bulimulidae, 1 of Predation mark Species Intact shells Odontostomidae, 2 of Simpulopsidae, and 1 of Subulinidae Apex Columella (but no abundance data can be extracted from it). Bulimulidae Number of Relative Number of Drymaeus flexilabris 7 [20%] 8 [23%] 20 [57%] Family specimens abundance (%) species Drymaeus papyraceus 8 [30%] 8 [30%] 11 [40%] Subulinidae 449 24.5 6 Oxychona bifasciata 7 [17.5%] 10 [25%] 23 [57.5%] Scolodontidae 394 21.5 4 Strophocheilidae Charopidae 223 12.2 4 Strophocheilus debilis 2 [4%] 0 53 [96%] Euconulidae 194 10.6 1 Megaobulimus cardosoi 1 [5%] 0 21 [95%] Helicinidae 173 9.4 3 Bulimulidae 100 5.5 5 Vertiginidae 80 4.4 2 More than 30 years after Pedra Talhada became a state Streptaxidae 69 3.8 1 park, the preservation of its environment is still tenta- Strophocheilidae 67 3.7 3 tive and unstable. Unfortunately, during our stay in the Neocyclotidae 25 1.4 2 reserve, it was not rare to observe depredations (e.g., Pleurodontidae 19 1.0 2 traps, woodcutting, wheel rims along the trails) despite Simpulopsidae 17 0.9 2 the daily presence of guards from the ICMBio. Needless Cyclophoridae 8 0.4 1 to say, these activities are incompatible with any program Succineidae 5 0.3 1 of environmental protection. The value of the diverse molluscan fauna reported here should be highlighted and Veronicellidae 3 0.2 2 taken into account in future conservation efforts, as land Strobilopsidae 2 0.1 1 snails are the most threatened group and good Ferussaciidae 2 0.1 1 indicators of well-preserved areas (Lydeard et al. 2004; Valloniidae 1 0.1 1 Régnier et al. 2009). Up to now, there are only six ani- TOTAL 1831 100 eschweizerbartxxx sng- 42 mal species considered protected in Pedra Talhada, all of them birds (ICMBio 2017). We argue that at least the species Megalobulimus cardosoi, considered extinct in time in Brazil. Moreover, some of the species listed here the IUCN Red List (Mansur 1996a), should also receive have surprisingly broad distributions, sometimes span- protected status. ning more than one major type of biome; some of these species (e.g., Pseudoguppya semenlini) also bear shells with few available features for taxonomy, so they could Acknowledgements actually represent a species complex. This could eventu- ally be resolved with molecular data and/or anatomical This work was made possible through the ICMBio permit features such as the radula and the reproductive tract. (Sisbio #48925-2), requested by Eduardo Colley, granting Among the larger, usually tree-dwelling species, of a 1-year license for access and collection in the Biologi- Bulimulidae, it was very common to find shells bearing cal Reserve of Pedra Talhada (PT), Alagoas/Pernambuco. signs of predation: a broken columellar region near the The NGO Nordesta (Switzerland), represented by its aperture (Figs 14K, L) or a broken spire apex (Fig. 14M). President Anita Studer, covered the travel and accom- These types of predation marks are typically assigned to modation of P.M. and L.C. for the collection expedition, birds (but small monkeys can also be responsible; Simone and awarded a small research grant to R.B.S. to study the personal observation). A little above 40% of adult Dry- material. We are deeply grateful to Anita Studer for finan- maeus flexilabris and Oxychona bifasciata and 60% of cial support and unswerving will to defend Quebrangulo, Drymaeus papyraceus show some type of predation PT and the local people; to our colleagues Eduardo Col- (Table 3); the latter could possibly be a preferred target ley and Silvio F.B. Lima for help with the expedition; to due to the visibly weaker shell (as the species name also the park guards (Flavio dos Santos Pereira, Adalto Este- implies). Furthermore, a few Strophocheilidae (< 5%, all vão Gomes da Silva and Fabiano Barros Carvalho), who juveniles; Table 3) also bear similar signs of predation; not only accompanied the team during the collection, their large adult size and sturdier shells, perhaps coupled but also turned out to be skilled snail collectors; to all with their typically ground-dwelling habitat, might dis- the people who helped us during fieldwork, Jailton Fer- courage predators. nandes (ICMBio’s head officer in PT), Claiton M. Silva

125 Archiv für Molluskenkunde · 147 (1) 2018

(Nordesta’s agronomist), Quiteria Valentim, Simonio N. Delannoye, R., Charles, L., Pointier, J.P. & Massemin, D. (2015) Silva, Luis B. Freitas Filho, Dante Buzzetti, and Louis Mollusques continentaux de la Martinique. Paris: Muséum Nusbaumer; to Christina G. Martin (SMNS) for the SEM national d’Histoire naturelle. Drouët, H. (1859) Essai sur les mollusques terrestres et fluviatiles images; to Gilberto Marani (MNHN) for making Figure de la Guyane française. Mémoires de la Société d’agriculture, 2; to Jon Ablett for granting access to the material under des sciences, arts et belles-lettres du département de l’Aube his care in the NHMUK collection; to Carlos Birckolz, (deuxième série) 10: 303–412. Heike Reise, and an anonymous reviewer for their help- Férussac, A.E.J.P.J.F. d’Audebard de (1819–1821) Tableaux systé- ful comments. 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in: Küster, H.C. & Kobelt, W. (Eds) Systematisches Conchy- Naesiotus sp.: station PT2015-29. Oxychona bifasciata: lien-Cabinet von Martini und Chemnitz. Neue Folge. Ersten stations PT2015-18, 26, 27, 28, 31, 32, 33. Biotocus tur- Bandes, Achtzehnte Abtheilung. Nuremberg: Bauer & Raspe. binatus: MNHN material. Rhinus botocudus: stations PT2015-18, 24, 27, 28, 29, 52. Simpulopsis atrovirens: station PT2015-47. Simpulopsis rufovirens: MNHN Appendix material. Simpulopsis sulculosa: MNHN material. Simpulopsis sp.: stations PT2015-18, 31. Orthalicoi- Here are listed the collection stations of Pedra Talhada dea indet.: stations PT2015-18, 22, 24, 31. Cecilioides (see also Table 1) in which each species occurred. The consobrina: stations PT2015-21, 27. Allopeas micrum: species are listed in the same order as they appear in the stations PT2015-18, 24, 29. Beckianum beckianum: sta- main text. It is worthwhile to note that several specimens tion PT2015-18. Leptinaria cf. mamoreensis: MNHN were either juvenile or fragmentary and could only be material. Leptinaria ritchiei: stations PT2015-17, 18, 21, identified to family or genus level (they are listed sepa- 22, 24, 25, 26, 27, 28, 29, 31, 32, 33, 35, 36, 43, 45, 46, rately below). These should belong to one of the species 47, 49, 50, 52. Leptinaria lamellata: stations PT2015- listed here rather than to distinct taxa, but they could not 18, 27, 42. Obeliscus agassizi: stations PT2015-22, 26, be confidently assigned to any particular species; there- 27, 39. Stenogyra octogyra: stations PT2015-18, 21, 24, fore, some species might in reality be more widespread 26, 28, 37, 50. Subulinidae indet.: stations PT2015-7, inside the reserve. 18, 19, 24, 26, 29, 31, 45, 50. Streptaxis iheringi: sta- Furthermore, 5 species could not be found in the 2015 tions PT2015-18, 20, 24, 25, 26, 27, 28, 32, 45, 46, 49, expedition, but were available in the collection of the 52. Happia vitrina: station PT2015-32. Tamayoa bang- MNHN, stemming from the 1998 expedition; they are haasi: stations PT2015-12, 17, 18, 20, 24, 25, 26, 27, 29, indicated as such in the list below. All lots collected dur- 31, 32, 39, 46, 47, 49, 51, 52. Scolodonta bounobaena: ing this expedition do not have accession numbers and stations PT2015-24, 26, 27, 29, 36. Scolodonta spiror- only bear a generalized entry of locality data: “Pedra bis: stations PT2015-18, 24, 25, 26, 27. Scolodontidae Talhada Reserve, 550–850 m a.s.l., 09° 14′ 14.5″ S, 036° sp.: stations PT2015-7, 12, 17, 18, 21, 24, 26, 27, 28, 29, 25′ 25.9″ W”. The expedition took place from May 23 to 31, 32, 33, 35, 36, 37, 39, 46, 49, 50. Megalobulimus June 7, 1998, at the beginning of the rainy season (Mae- cardosoi: stations PT2015-18, 27, 28. Megalobulimus strati et al. 2015). oliveirai: stations PT2015-27, 29, 32, 33, 36, 37, 38, 49. Helicina angulifera: stations PT2015-18, 22, 24, 25, Megalobulimus sp.: stations PT2015-27, 36. Stropho-

26, 27, 28, 29, 31, 32, 33, 35, 47, 48. Helicinaeschweizerbartxxx sng-rotun- cheilus debilis: stations PT2015-18, 19, 24, 27, 32, 35, 36, data: stations PT2015-18, 27. Helicina schereri: stations 49, 52. Lilloiconcha gordurasensis: stations PT2015-7, PT2015-24, 28. Helicina sp.: stations PT2015-18, 21, 12, 18, 20, 21, 24, 25, 26, 27, 28, 29, 33, 46, 49. Lil- 25, 27, 29, 32, 36. Aperostoma blanchetiana: station loiconcha superba: stations PT2015-12, 20, 24, 25, 31. PT2015-28. Cyclopomops moricandi: stations PT2015- Radiodiscus amoenus: station PT2015-18. Radiodiscus 7, 15, 22, 27, 49. Adelopoma cf. brasiliense: stations ubtaoi sp. nov.: stations PT2015-23, 31. Charopidae PT2015-24, 46. Sarasinula sp.: stations PT2015-24, indet.: stations PT2015-18, 21, 24, 26, 31, 33, 35, 37, 36, 35. Latipes sp.: station PT2015-46. Omalonyx sp.: 39, 46. Pseudoguppya semenlini: stations PT2015-7, 12, station PT2015-42. Strobilops cf. brasiliana: station 17, 18, 20, 21, 23, 24, 26, 27, 29, 31, 32, 33, 46, 49, 52. PT2015-18. Pupisoma dioscoricola: station PT2015-18. Solaropsis pascalia: stations PT2015-24, 25, 27, 36, 48. Bothriopupa breviconus: stations PT2015-12, 14, 24, Solaropsis cearana: stations PT2015-18, 27, 45. 27, 29, 49. Sterkia eyriesii: station PT2015-27. Bulimu- lus tenuissimus: MNHN material. Drymaeus flexilabris: stations PT2015-18, 27, 28, 29, 32, 33, 36, 47, 49. Dry- maeus papyraceus: stations PT2015-18, 25, 27, 28, 31, Manuscript submitted 28 November 2017 36, 26. Drymaeus rufolineatus: station PT2015-29. Revised manuscript accepted 6 March 2018

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