Taxonomy and Distribution of Enigmatic “Helicoid”
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Zoosyst. Evol. 96 (1) 2020, 91–101 | DOI 10.3897/zse.96.51047 Taxonomy and distribution of enigmatic “helicoid” Polygyratia Gray, 1847 (Gastropoda, Stylommatophora) Rodrigo Brincalepe Salvador1, Daniel Caracanhas Cavallari2 1 Museum of New Zealand Te Papa Tongarewa, Wellington, New Zealand 2 Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo. Ribeirão Preto, SP, Brazil http://zoobank.org/E87CE884-99A0-4900-96C1-C3F15E3B01A3 Corresponding author: Rodrigo Brincalepe Salvador ([email protected]) Academic editor: Frank Köhler ♦ Received 12 February 2020 ♦ Accepted 1 March 2020 ♦ Published 13 March 2020 Abstract Herein, we present a taxonomic revision of the genus Polygyratia Gray, 1847, with a new systematic placement in Scolodontidae and containing only the species Polygyratia polygyrata (Born, 1778). We offer an updated morphological description and geographical distribution, based on museum specimens and occurrence data gathered from literature and online database iNaturalist. We synony- mise P. charybdis Mörch, 1852 with P. polygyrata. The species is known only from Atlantic Forest areas in Bahia state, eastern Bra- zil. We exclude three other species from the genus Polygyratia, classifying them as: Systrophia (Systrophia) heligmoida (d’Orbigny, 1835) and S. (Entodina) reyrei (Souverbie, 1858), based on conchological features; and S. (E.) pollodonta (d’Orbigny, 1835), though tentatively, based on scant published data. Finally, we present the first report ofS. (S.) heligmoida (d’Orbigny, 1835) from Brazil. Key Words Atlantic Forest, Brazil, Eupulmonata, Helicoidea, iNaturalist, Polygyratia polygyrata (Born, 1778), Scolodontidae Introduction Material and methods Polygyratia polygyrata (Born, 1778) was one of the first The main museum collections worldwide that could terrestrial gastropods to be described from Brazil. After harbour specimens of Polygyratia were visited or con- its original description, which did not provide a locality tacted. Specimens were analysed for conchological of origin, all subsequent major lists of terrestrial gastro- characters (see below for molecular analysis) and the pods included the species, pinpointing its occurrence to available locality data was compiled for a distribution the forests of Bahia (e.g. Beck 1837; Albers 1860). De- map. Collection dates are unknown for most of the his- spite being an instantly-recognisable species and a favou- torical specimens, but at least a minimum age can be rite item of shell collectors of old, the species remained given to those that belonged to private collections by scarcely studied after the 19th century, with a single addi- using the owner’s date of death (Coan and Kabat 2018). tional species described (P. charybdis Mörch, 1852) and The owners’ names were cross-checked with the report a brief revision presented by Pilsbry (1894). of Dance (1966), which lists where these former private Given that the knowledge about this species advanced collections are presently housed. Further photographical very little since its description, we decided to reassess records of the species were extracted from the online da- here all the information available regarding its taxonomy tabase iNaturalist (https://www.inaturalist.org) and shell and geographical distribution, analysing museum speci- collector websites. Additional locality data was extract- mens from worldwide collections, as well as online data- ed from literature. bases. We present here a formal re-description of the ge- The only specimens with soft parts that could be nus and its species, alongside venues for future research. sampled for molecular studies (MZSP 62060, 62061 Copyright Rodrigo B. Salvador, Daniel C. Cavallari. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 92 Salvador, R.B. & Cavallari, D.C.: Taxonomy and distribution of Polygyratia and 98742) had small sections of their foot clipped Systematics for DNA extraction. The specimens, however, were in poor condition: largely decomposed, indicative of the Family Scolodontidae ethanol having dried up at some point. Different types of DNA extraction kits (and protocols) were tried: Genus Polygyratia Gray, 1847 DNeasy Blood & Tissue Kit (QIAGEN), NucleoSpin DNA RapidLyse (Macherey-Nagel) and E.Z.N.A.Mol- Polygyratia Gray, 1847: 173; Gude 1920: 59; Thiele 1931: 679; Mor- lusc DNA Kit (Omega Bio-tek). Unfortunately, none of retes 1949: 163; Zilch 1960: 603; Richardson 1985: 262; Parkinson the trials resulted in good quality DNA for amplifica- et al. 1987: 65; Salgado and Coelho 2003: 172; Schileyko 2006: tion and sequencing. 1844; Simone 2006: 247; Breure and Araujo 2017: 121; Mollus- Selected specimens were used for micro-computed caBase 2018; Salvador 2019: 95. tomography analysis (CT scan), scanned using a GE Ophiogyra Albers, 1850: 91; Albers 1860: 94. Phoenix v|tome|x s 240 industrial high-resolution CT & x-ray system at the Centro para Documentação da Bi- Type species. Helix Polygyrata Born, 1778, by monotypy. odiversidade (FFCLRP). This system is equipped with a DXR250RT 1,000 × 1,000 pixels detector; a diamond Included species. P. polygyrata (Born, 1778). target was used with the following parameters: source operating at 60 kV, 100 μA, 1 × 1 binning, 333 ms expo- Diagnosis. Shell planispiral, multi-whorled and large for sure time, 1,200 projections, 1 frames skipped, 4 frames family. Body whorls marked by three angulations: one averaged, default offset and gain corrections. Image pro- dorsal, one median and one basal. Aperture large (for cessing and 3D volume reconstruction were obtained family), rounded, with reflected peristome. Presence of using GE phoenix datos|x 2 and the subsequent volume short internal (not visible from aperture) parietal and pal- analysis was carried out in VGSTUDIO 3.0 (Volume atal lamellae on body whorl. Aperture without barriers. Graphics, Germany). Abbreviations: Institutions: AMNH = American Description. As P. polygyrata below. Museum of Natural History (New York, USA); ANSP = Academy of Natural Sciences of Drexel University Remarks. The name Ophiogyra was introduced by Al- (Philadelphia, USA); CMNH = Cleveland Museum bers (1850), who likely was not aware of the new name of Natural History (Cleveland, USA); FMNH = Field established by Gray just a few years prior. The only spe- Museum of Natural History (Chicago, USA); MNB = cies included by Albers (1850) in the new genus was Muzeul Național Brukenthal, Muzeul de Istorie Natu- Ophiogyra polygyrata. Thus, Ophiogyra is an objective rală (Sibiu/Hermannstadt, Romania); MNCN = Museo junior synonym of Polygyratia. Nacional de Ciencias Naturales (Madrid, Spain); MNZ = Museum of New Zealand Te Papa Tongarewa (Wel- lington, New Zealand); MZSP = Museu de Zoologia de Polygyratia polygyrata (Born, 1778) Universidade de São Paulo (São Paulo, Brazil); NHMD Figs 1, 2 = Statens Naturhistoriske Museum (Copenhagen, Den- mark); NHMUK = Natural History Museum (London, Helix Polygyrata Born, 1778: 382; Wilhelm 1802: 356. UK); NHMW = Naturhistorisches Museum Wien (Vi- Helix polygyrata: Born, 1780: 373, pl. 14, figs 19–20; Schröter 1784: enna, Austria); NMR = Natuurhistorisch Museum Rot- 266; Chemnitz 1786: 98, pl. 127, figs 1124–1125; Dillwyn 1817: terdam (Rotterdam, The Netherlands); RMNH = Nat- 908; Férussac 1819: 4, pl. 69A, figs 7–9, pl. 69B, fig. 5); Férussac uralis Biodiversity Center (former Rijksmuseum van 1821: 40; Deshayes in Lamarck 1838: 98; Moricand 1846: 151, pl. Natuurlijke Histoire; Leiden, The Netherlands); SMF = 5, figs 1–3; Pfeiffer 1848: 405; Mörch 1852: 7; Reeve 1854: species Senckenberg Naturmuseum (Frankfurt am Main, Ger- 541, pl. 98; Hupé 1857: 12; Hidalgo 1870: 32; Hidalgo 1893: 80. many); SNSD = Senckenberg Natural History Collec- Helix polygyra [sic] Gmelin, 1788: 3624. tions Dresden (Dresden, Germany); USNM = Smithso- Helix Elongata Röding, 1798: 107. nian National Museum of Natural History (Washington, Helix (Helicella) polygyrata: Moricand, 1836: 422. D.C., USA); ZMA = Naturalis Biodiversity Center Polygyra? Polygyrata: Beck 1837: 23. (former Zoölogisch Museum Amsterdam; Leiden, The Polygyratia polygyrata: Gray 1847: 173; d’Orbigny 1849: pl. 19, figs Netherlands); ZMB = Museum für Naturkunde, Lei- 14–16; Mörch 1852: 7; Thiele 1931: 679; Zilch 1960: 603, fig. 2119; bniz Institute for Evolution and Biodiversity Science Richardson 1985: 262; Parkinson et al. 1987: pl. 13, fig. 6; Abbott (Berlin, Germany). Specimens: col = collector; colln 1989: 142, textfig.; Salgado and Coelho 2003: 172; Schileyko 2006: = collection; D = greatest shell width (perpendicular 1844, fig. 2359; Simone 2006: 247, fig. 946; Schileyko 2016: fig. 4; to H); H = shell length (parallel to columellar axis); Breure and Araujo 2017: 121, figs 47D–F; Sei et al. 2017: table 9, obs = observer; sh = shell(s); spm = specimen(s); W = fig. 5n; MolluscaBase 2019a; Salvador 2019: 95. number of whorls. Ophiogyra polygyrata: Albers 1850: 91; Albers 1860: 94. zse.pensoft.net Zoosyst. Evol. 96 (1) 2020, 91–101 93 Polygyratia charybdis Mörch, 1852: 7. Habitat. Atlantic Forest. Polygyratia (detrita.): Mörch 1852: 7. Helix charybdis: Mörch 1852: 170. Diagnosis. As for genus above. Helix Charybdis: Hupé 1857: 12. Helix (Anchistoma) polygyrata: Brauer 1878: 181. Description. Shell (Fig. 1) planispiral, multiwhorled. Polygyra (Polygyratia) polygyrata: Tryon 1887: