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On South Georgia: Some Implications of Shell Size, Shell Shape, and Site Isolation in a Singular Sub-Antarctic Land Snail P.J.A

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On South Georgia: Some Implications of Shell Size, Shell Shape, and Site Isolation in a Singular Sub-Antarctic Land Snail P.J.A Antarctic Science 23(5), 442–448 (2011) & Antarctic Science Ltd 2011 doi:10.1017/S0954102011000289 Notodiscus (Charopidae) on South Georgia: some implications of shell size, shell shape, and site isolation in a singular sub-Antarctic land snail P.J.A. PUGH1 and R.I. LEWIS SMITH2 1Department of Life Sciences, Anglia Ruskin University, East Road, Cambridge CB1 1PT, UK 2Centre for Antarctic Plant Ecology and Diversity, Alton Road, Moffat DG10 9LB, UK [email protected] Abstract: Multivariate analysis shows that shells of Notodiscus sp. (Charopidae: Pulmonata) reported from South Georgia are smaller and proportionately taller than, but otherwise similar to, populations of Notodiscus hookeri (Reeve) from Iles Crozet and Iles Kerguelen. The origin of this solitary, and spatially limited, South Georgia population is enigmatic. It is confined to a remarkably small coastal lowland site which was glaciated at Last Glacial Maximum, precluding a Tertiary relict origin, and on the leeward north-east coast, ruling out post- glacial ocean rafting. The site is close to the King Edward Point settlement, yet the absence of any logistics connections with the Iles Crozet or Iles Kerguelen mitigates against anthropogenic introduction. The close proximity of the population to nests of blue-eyed shag (Phalacrocorax atriceps), Dominican gull (Larus dominicanus) and light-mantled sooty albatross (Phoebetria palpebrata) could imply the snail was originally introduced to South Georgia via these ocean transiting seabirds. Received 15 September 2010, accepted 14 February 2011, first published online 15 April 2011 Key words: colonization, Gastropoda, Notodiscus, Southern Ocean islands, zoochory Introduction Amsterdam, Marion and possibly Prince Edward Island (Prince Edward Islands) (Fig. 1), and containing a single The islands of the Southern Ocean (Fig. 1) harbour only variable species, N. hookeri Reeve, 1854 (Solem 1968, 41 species of endemic and introduced terrestrial slugs and Smith 1992, Madec & Bellido 2007). The primary focus of snails (Gastropoda: Mollusca) (Pugh & Scott 2002). Twenty- this study is a morphometric comparison of the snails on seven species occur on the South Pacific Province (SPP) South Georgia with those of N. hookeri collected on the Iles Antipodes, Bounty, Campbell, Chatham, Macquarie and Crozet and Iles Kerguelen by Madec & Bellido (2007). Snares islands, while the South Indian Province (SIP) Amsterdam, Crozet, Heard, Kerguelen and Prince Edward islands are colonized by two exotics and a single endemic Methods species - Notodiscus hookeri (Reeve) (Charopidae) (Fig. 1). Field observations The impoverished South Atlantic Province (SAP) fauna is confined to three slugs and snails introduced to the Falkland The snail occurs c. 1 km north of Hope Point, King Edward Islands, and a single snail on South Georgia. Cove, Cumberland East Bay, South Georgia (54816'S, This South Georgian snail, discovered by RILS in January 36823'W) (Fig. 2). The site, c. 25 m above sea level and 1970, was first reported as an unidentified terrestrial mollusc 15–20 m inland from a narrow pebble beach, was the lower (Smith & Walton 1975), and subsequently tentatively ascribed 5 m of a high, vertical shaded cliff with a south-east aspect. to the genus Stephanoda (Endodontidae) or Notodiscus Ledges and crevices on this cliff face were colonized (Charopidae). Stephanoda (sensu Block 1984, Headland by liverworts (including Pachyglossa dissitifolia), lichens 1984), was based on a 1978 personal communication from (Cladonia spp., Rhizocarpon geographicum, Tephromela F.C. Naggs (British Museum, London) and Notodiscus (sensu atra), mosses (Andreaea spp., Bartramia patens, Grimmia, Pugh & Scott 2002), on further personal communications Pohlia nutans, Racomitrium spp.), and small vascular plants from F.C. Naggs (British Museum, London 1981), M. Vogel (Cystopteris fragilis, Deschampsia antarctica, Festuca (Philipps-Universita¨t, Hamburg 1985), the late A. Solem contracta, Phleum alpinum). Moist areas of rock were (Field Museum of Natural History, Chicago 1986), and coated by a film of microalgae and cyanobacteria, and P. Mordan (Natural History Museum, London 1999). We the snails appeared to be feeding on these organisms. The subsequently confirmed Notodiscus (Pugh et al. unpublished), soil and scree at the base of the cliff were covered by a genus previously known only from the SIP sub-Antarctic Acaena magellanica and A. tenera, scattered Festuca and islands of Kerguelen, Possession (Iles Crozet), Heard, mosses (especially Syntrichia robusta). Many empty shells 442 Downloaded from https://www.cambridge.org/core. Open University Library, on 22 Jan 2020 at 00:05:42, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0954102011000289 NOTODISCUS ON SOUTH GEORGIA 443 Fig. 1. Location of South Georgia (SG) relative to the South Pacific and South Indian Ocean islands of Amsterdam (Am), Crozet (Cr), Heard (Hd), Kerguelen (Kr), Prince Edward (PE) and St. Paul (SP). were observed at the base of the cliff face while live snails were confined to c.30m2 of the cliff and particularly abundant at 1–3 m above the cliff base on mosses, in Fig. 3. Notodiscus sp. a. Dorsal. b. Lateral. c. Ventral. Scale bars: 1 mm. crevices and pockets of soil on ledges. Associated invertebrates included Enchytraeidae (Oligochaeta), mites (Acarina) and beetles (Coleoptera: Hydromedion and Perimylops spp.). Similar habitat, flora and fauna occur Fig. 2. Offset: South Georgia showing Cumberland East Bay throughout much of the north-east coast of South Georgia (arrow). Main map: Cumberland East Bay showing collection yet many years of detailed ecological surveys and searches site (K) and the settlements of Grytviken whaling station (G) specifically for the snail have so far failed to locate and King Edward Point (K). evidence of other populations. Downloaded from https://www.cambridge.org/core. Open University Library, on 22 Jan 2020 at 00:05:42, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0954102011000289 444 P.J.A. PUGH & R.I. LEWIS SMITH Table I. Simple comparison of shells from Iles Kerguelen, Iles Crozet wet conditions. Specimens kept for several weeks in a Perspex (after Madec & Bellido 2007) and South Georgia (current study). container at laboratory temperature (c. 15–188C) at South Measurements (after Madec & Bellido 2007) Georgia appeared to be more active at night. RILS fixed a Island AB CD BG BE EG EF total of 40 live specimens in 70% laboratory ethanol for Kerguelen mean 4.32 1.62 2.47 0.10 2.35 0.39 anatomical study, although some subsequently dried out. ± SD 0.37 0.15 0.19 0.04 0.18 0.05 Crozet mean 4.29 1.56 2.59 0.18 2.41 0.47 Specimen preparation ± SD 0.63 0.30 0.38 0.06 0.35 0.12 We cleaned several of the dried snails collected in March 1981 South Georgia mean 3.11 1.30 2.00 0.22 1.78 0.36 and prepared them for scanning electron microscopy. Shells ±SD 0.24 0.17 0.21 0.07 0.19 0.09 were soaked in absolute ethanol for 48 hours and cleaned Note: notable South Georgia values are italic (low)orbold (high). of adherent debris in low viscosity ethanol, to minimize mechanical risk for fragile shells, in an ultrasound bath at Little can be gleaned from the published literature about 47 kHz delivered in 15 second ‘bursts’ to prevent alcohol the ecology of Nothodiscus hookeri throughout its sub- ignition. Loose debris was removed with a trimmed no. 3 Antarctic distribution. It is widespread from near sea level artist’s brush. Cleaned individual shells were mounted on to . 700 m altitude in the sub-Antarctic islands of the conventional aluminium SEM stubs via double-sided adhesive Indian Ocean. Madec & Bellido’s (2007) samples from Iles carbon disks (a clean background for imaging) and a small Crozet were collected between 10 and 70 m, while those spot of colloidal silver (for optimum grounding), then gold- from Iles Kerguelen were from c. 190–700 m altitude. From coated in a Biorad SC502 sputter-coating unit. We imaged Dall’s (1876) and Dell’s (1964) accounts the Kerguelen the shells in a Leica-Cambridge S360 scanning electron populations appear to be associated with stony habitats and microscope configured for maximum depth of field at moss (i.e. fellfield) and also with the megaherb Pringlea a 10 kV acceleration voltage, 120 pA probe current and a antiscorbutica in wetter sheltered habitats. On Marion 20 mm working distance, capturing secondary electron Island N. hookeri typically frequents fellfield habitats at images via an electronic frame store (Fig. 3). lower altitudes (RILS, personal observation). However, no accounts refer to the snails occurring on cliff faces, as on Morphometric analysis South Georgia, although such a habitat could be construed as ‘‘vertical fellfield’’. We briefly dried the 36 ethanol fixed specimens, collected Subsequent summer (January 1981 and February 2000) in 2000, on laboratory tissue, mounted them aperture observations by RILS suggested the snails aggregated uppermost on a small bead of ‘Blu-Tack’ pressure sensitive (5–20 per dm2 over several square metres) in crevices and adhesive and measured via a dissection microscope fitted beneath overhangs during sunny, dry weather but dispersed with calibrated eyepiece graticule at x30. Our analysis focused onto open rock and compact moss cushions during overcast or on six principal measurements of maximal diameter (AB), Table II. Correlation matrix. Similarity matrix ln AB ln CD ln BG ln
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