A Revision of the New Zealand Landsnails Referred to Allodiscus Pilsbry, 1892 and Pseudallodiscus Climo, 1971, with the Introduc
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Tuhinga 19: 57–167 Copyright © Museum of New Zealand Te Papa Tongarewa (2008) A revision of the New Zealand landsnails referred to Allodiscus Pilsbry, 1892 and Pseudallodiscus Climo, 1971, with the introduction of three new genera (Mollusca: Gastropoda: Charopidae) BrUce A. Marshall1 and Gary M. Barker2 1 MUseUm of New Zealand Te Papa Tongarewa, PO Box 467, Wellington, New Zealand ([email protected]) 2 Landcare Research, Private Bag 3127, Hamilton, New Zealand ([email protected]) ABSTRACT: The New Zealand species referred to Allodiscus Pilsbry, 1892 during the last 60 years are evaluated. A total of 60 species are recognised, of which 37 are described as new. The taxa are described, they are illustrated by colour photographs and scanning electron microscope images, their distributions are described and mapped, and their biology and con- servation are briefly discussed. Pseudallodiscus tataensis Climo, 1971 is referred to Allodiscus [s. lat.]. Three new genera are introduced: Granallodisus (for Helix granum L. Pfeiffer, 1857 and two new species), Costallodiscus (for three new species) and Canallodiscus (for Allodiscus fectoloides Dell, 1955, Phelussa elliottae Gardner, 1968 and a new species). Hirsutodiscus Climo, 1971 is elevated to generic status. Lectotypes are designated for Helix dimorpha Reeve, 1852, Pitys cryptobidens Suter, 1891, Helix venulata L. Pfeiffer, 1857, Nanina tullia Gray, 1850 and Psyra godeti Suter, 1891. Neotypes are designated for Helix granum L. Pfeiffer, 1857 and Charopa miranda Hutton, 1883. KEYWORDS: Mollusca, Gastropoda, Charopidae, Allodiscus, Pseudallodiscus, Granallodisus, Costallodiscus, Canallodiscus, Hirsutodiscus, revision, taxonomy, distribution, new taxa, new combinations, lectotypes, neotypes. recognised but presently undescribed species. This paper Introduction addresses the systematics of New Zealand landsnails, includ- New Zealand has a diverse landsnail fauna. Barker (2005) ing a number of undescribed species, presently assigned to estimated that at least 450 New Zealand species-level taxa in the charopid genus Allodiscus Pilsbry, 1892 (type species the family Charopidae are represented in collections, of Helix dimorpha Reeve, 1852). which only 210 have been formally described. Spencer et al. Allodiscus was founded for New Zealand species, but it (in press) tabulated 207 described and 167 undescribed has also been applied to New Caledonian and Australian charopids in their checklist to New Zealand Mollusca. material. Franc (1957) treated the New Caledonian Many charopids are considered of conservation concern Platyrhytida Pilsbry, 1894 (type species Helix saisseti Gassies, (Hitchmough et al. 2007). However, for many groups in the 1860) as a subgenus of Allodiscus. However, Solem (1961) New Zealand Charopidae, proper assessment of conservation recognised conchological differences between Allodiscus and status has been hindered by the lack of modern systematic Platyrhytida, and accordingly treated the latter at genus level. treatment and the absence of formal taxonomy of many Suter (1913) suggested a possible close relationship to 58 Tuhinga, Number 19 (2008) Monomphalus Ancey, 1882 from New Caledonia (type (1884a), Suter (1913), Gardner (1968), Climo (1969b, species Helix rossiterianus Crosse, 1871), based on concho- 1971a) and Schileyko (2001). We erect several conchologi- logical features, but this genus has subsequently been shown cal form groups in Allodiscus, corresponding to Allodiscus (Schileyko 2001; Starmühlner 1970) to differ anatomically s. str. and several groupings in Allodiscus s. lat. Diagnoses are from Allodiscus. The Victorian species Allodiscus cannfluvi- provided for each as an aid to any subsequent revision, but atilus Gabriel, 1929 and A. marysvillensis Gabriel, 1947 are demonstration that these form groups represent mono- presently classified in Oreomava Kershaw, 1956 and phyletic clades must await molecular genetic studies. Full Pillomena Iredale, 1937, respectively (see Smith 1992). Thus, synonomies are provided for all taxa recognised, inclusive of Allodiscus is evidently a New Zealand endemic genus. all literature records in unrefereed checklists, shell club Since the inception of the genus, a total of 29 New articles and unpublished handouts. Zealand species-group names have hitherto been erected in All measurements were performed with a microscope eye- Allodiscus or secondarily assigned to the genus. Over the past piece graticule. In cited dimensions in plate captions, height 50 years, there has been a gradual evolution in the concept dimension precedes width. The method for counting the of Allodiscus. First, Cumber (1961) transferred Flammulina number of protoconch whorls follows Warén (e.g. 1992) (Allodiscus) tholoides Suter, 1907 to Phenacohelix Suter, 1892 (Fig. 2A). Selected specimens were photographed in colour (type species Helix pilula Reeve, 1852). Then Climo (1971a) using automontage software, and some of these and others erected the monotypic subgenus Hirsutodiscus for Allodiscus were subsequently prepared and mounted, then carbon and (Hirsutodiscus) rakiura Climo, 1971, on the basis of smooth, gold/palladium coated and illustrated by scanning electron glossy protoconch and hair-like processes on the axial ribs microscopy. of the teleoconch. Here, we elevate Hirsutodiscus to generic Unless preceded by the acronyms AIM, BMNH or CM, status. all material examined (2962 lots, which includes material Climo (1971a) introduced Pseudallodiscus (type species additional to that recorded herein) is at NMNZ (registration Pseudallodiscus ponderi Climo, 1971) for three small snails numbers preceded by ‘M.’). All examined material is plotted with Allodiscus characters but differing in the reproductive on distribution maps. Full locality information is provided system and radular dentition. However, Climo (1981) sub- for primary type material only. All locality information for sequently transferred Pseudallodiscus spelaeus Climo, 1971 to examined material is stored with full data in Te Papa’s Mocella Iredale, 1915 (type species Helix corniculum Reeve, database (KE EMu) and is also available on the Global 1852). In the present contribution, we assign Pseudallodiscus Biodiversity Information Facility (GBIF) website. The bulk tataensis Climo, 1971 to Allodiscus. With these transfers, of the material was collected by Pauline Mayhill (845 lots), Pseudallodiscus is secondarily monotypic and we have taken David Roscoe (660 lots), Frank Climo (318 lots) and Bruce the opportunity to redefine the genus. Hazelwood (203 lots). Most recently, Climo & Mahlfeld (1998) reassigned four Information about biology and macroecology was derived species formerly referred to Allodiscus. Charopa planulata from specimen labels, geographic information systems (GIS) Hutton, 1883 (misapplied to a number of species of databases and expert knowledge. We have provided com- Allodiscus s. lat.), Allodiscus rusticus Suter, 1894, and ments on the conservation status of all species, with reference Flammulina (Allodiscus) mossi Murdoch, 1897 were assigned to previous assessments by McGuinness (2001), Brook to Chaureopa Climo, 1985 (type species Chaureopa depressa (2002b), Hitchmough (2002) and Hitchmough et al. (2007), Climo, 1985). Allodiscus stewartensis David, 1934 was the latter three based on the nationally agreed standard assigned to Phacussa Hutton, 1883 (type species Zonites classification criteria of Molloy et al. (2002). For all taxa, helmsi Hutton, 1883). including those recognised for the first time, we have pro- vided our own assessments of conservation status using the Molloy et al. (2002) criteria. Materials and methods In the absence of preserved animal material, we have neces- Abbreviations used in the text sarily based this revision primarily on conchological features. AIM – Auckland Institute and Museum, Auckland In the descriptions we refer, where available, to anatomical BMNH – The Natural History Museum, London information provided by previous authors, including Hutton CM – Canterbury Museum, Christchurch A revision of the New Zealand landsnails referred to Allodiscus and Pseudallodiscus 59 NMNZ – Museum of New Zealand Te Papa Tongarewa, Systematics Wellington NZMS 260 – New Zealand Map Series metric grid reference Superfamily PUNCTOIDEA Morse, 1864 (Infomap: New Zealand Department of Survey and Land Family CHAROPIDAE Hutton, 1884 Information) Genus Allodiscus Pilsbry, 1892 HWR – Shell height/width ratio SH – Shell height Psyra Hutton, 1883b: 532. Type species Helix venulata SW – Shell width L. Pfeiffer, 1857, by original designation and monotypy. (Not Psyra Stal, 1876 in Orthoptera.) Index to taxa Allodiscus Pilsbry, 1892a: 56. Type species Helix dimorpha Reeve, 1852, by original designation; Recent, New abaxoides 141 mayhillae 147 Zealand. absidatus 125 miranda 145 Charopa as applied by various authors. (Not Charopa Albers adriana 80 mirificus 108 albinus 85 morioria 119 & Martens, 1860: 87. Type species Helix coma Gray, Allodiscus 59 negiae 103 1843, by original designation; Charopidae.) alloia 132 occidaneus 129 Endodonta as applied by various authors to New Zealand aurora 109 pallidus 135 material. (Not Endodonta Albers, 1850. Type species Helix austrodimorphus 110 parrishi 153 lamellosa Férussac, 1824, by subsequent designation of basiliratus 126 patulus 108 Albers & Martens, 1860; Endodontidae.) brooki 127 pegasus 153 Flammulina as applied by various authors. (Not Flammulina camelinus 61 planulatus 58 Martens, 1873: 12. Type species Vitrina zebra Le Guillou,