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A Revision of New Zealand Landsnails of the Genus Cytora Kobelt & Möllendorff, 1897 (Mollusca: Gastropoda: Pupinidae)

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A Revision of New Zealand Landsnails of the Genus Cytora Kobelt & Möllendorff, 1897 (Mollusca: Gastropoda: Pupinidae) Tuhinga18Final 28/5/07 10:23 AM Page 49 Tuhinga 18: 49–113 Copyright © Te Papa Museum of New Zealand (2007) A revision of New Zealand landsnails of the genus Cytora Kobelt & Möllendorff, 1897 (Mollusca: Gastropoda: Pupinidae) 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 endemic New Zealand cyclophoroidean genus Cytora is revised. A total of 42 species are recognised, of which 23 are described as new. All taxa are described and illustrated with colour photographs and scanning electron microscope images. KEYWORDS: Mollusca, Gastropoda, Pupinidae, Cytora, revision, taxonomy, distribution, new taxa. Introduction Materials and methods The New Zealand landsnail fauna is depauperate at the Taxonomy family level, yet secondarily is extraordinarily rich at the Unless preceded by acronyms AIM, BMNH or CM (see species level. Most of the species diversity is contained below), all material examined (3406 lots) was at Museum within the families Pupinidae, Athoracophoridae, of New Zealand Te Papa Tongarewa (registration numbers Rhytididae and, especially, Charopidae and Punctidae, as preceded by ‘M.’). All material with good locality data is the consequence of extensive endemic cladogenesis and plotted on distribution maps. Full details on locality data speciation (Barker 2005). There is concern about the con- are provided in the text only for type material; all other servation status of this unique fauna, as extensive habitat examined material is referred to only by registration loss and degradation have accompanied human settlement numbers. However, comprehensive details for all examined of the archipelago over the past millennium. However, accessions are stored in Te Papa’s collections information progress in formulating robust conservation strategies has system and are available on request (B.A. Marshall) or via been severely constrained by lack of modern taxonomic the Internet (http://collections.tepapa.govt.nz). Much of the treatments. Indeed, a high proportion of the New Zealand material was collected by Pauline C. Mayhill (1026 lots), landsnails remain undescribed (Barker 2005). David Roscoe (611 lots) and Bruce Hazelwood (536 lots). The Architaenioglossa are represented in the New Selected specimens were photographed in colour using Zealand terrestrial landsnail fauna by the cyclophoroidean AutoMontage™ software. These or others were subse- genera Liarea Gray, 1852 and Cytora Kobelt & quently mounted, carbon- and gold/palladium-coated, and Möllendorff, 1897. In this paper we describe the Cytora examined by scanning electron microscopy to enable the fauna. documentation of microscopic character states. Tuhinga18Final 28/5/07 10:23 AM Page 50 50 Tuhinga, Number 18 (2007) The following conchological measurements were made using microscope graticules for shells of 30 mature animals Index to Taxa from the type locality or collection stations in its vicinity: ampla 84 kerrana 83 protoconch diameter; shell height; shell diameter; spire annectens 54 kiama 102 angle (degrees); umbilicus width; aperture width perpen- aranea 58 lignaria 84 dicular to shell axis; aperture height parallel to shell axis; brooki 59 malleata 85 spire height. From these measurements the following calva 60 maui 87 ratios were calculated: shell height to diameter; umbilicus chiltoni 60 mayhillae 89 width to shell diameter; aperture width to shell diameter; climoi 63 minor 90 spire height to aperture height. Additional measurements Cytora 51 motu 91 were made from the selected individual(s) used in light cytora 63 pakotai 92 and scanning electron microphotography. depressa 66 pallida 92 Information about biology and macroecology was fasciata 67 pannosa 94 derived from specimen labels, Geographic Information filicosta 68 paparoa 97 Systems (GIS) databases and expert knowledge. We gardneri 68 parrishi 99 have provided comments on the conservation status of goulstonei 69 rakiura 100 all species, with reference to previous assessments by hazelwoodi 71 septentrionalis 100 McGuinness (2001), Brook (2002a) and Hitchmough hedleyi 71 solitaria 102 (2002), the latter two based on the nationally agreed stan- hirsutissima 75 taipa 103 dard classification criteria of Molloy et al. (2002). For all hispida 75 tawhiti 105 taxa, including those recognised for the first time, we have houhora 76 tepakiensis 105 provided our own assessments of conservation status using jamiesoni 77 tokerau 106 the Molloy et al. (2002) criteria. kahurangi 79 torquillum 107 kakano 79 tuarua 108 Abbreviations used in the text kamura 81 AIM – Auckland Institute and Museum, Auckland BMNH – Natural History Museum, London CM – Canterbury Museum, Christchurch NMNZ – Museum of New Zealand Te Papa Tongarewa, Systematics Wellington NZMS 260 – New Zealand Map Series metric grid refer- Superfamily CYCLOPHOROIDEA. ence (Infomap: New Zealand Department of Survey Family PUPINIDAE L. Pfeiffer, 1853. and Land Information) PUPINIDAE L. Pfeiffer, 1853: 98. Type genus: Pupina SEM – scanning electron microscopy Vignard, 1829. Subfamily LIAREINAE Powell, 1946. LIAREIDAE Powell, 1946: 70. Type genus: Liarea L. Pfeiffer, 1853. CYTORIDAE Climo, 1969: 227. Not available: no diagnosis. REMARKS: Liarea Gray, 1852 and Cytora Kobelt & Möllendorff, 1897 are the only cyclophoroidean genera in the extant New Zealand landsnail fauna. The systematic affinities of these genera have not been fully resolved, reflecting in large part the poor understanding of the evolutionary history and unstable higher systematics of cyclophoroideans globally (Barker 2001). Suter (1913) Tuhinga18Final 28/5/07 10:23 AM Page 51 A revision of New Zealand landsnails of the genus Cytora 51 recognised affinities of Cytora with Cyclophoridae sensu DIAGNOSIS: Shell dextrally coiled, less than 7 mm in height, stricto, and Liarea with snails now classified not as cyclo- ranging in shape from tall conical to depressed; last adult phoroideans but rather as omphalotropine Assimineidae. whorl of mature animal generally constricted (i.e. more In Cyclophoridae sensu lato, Thiele (1929) assigned Cytora narrow than expected if it were the geometric continuation to Cyclophorinae and Liarea to Pupininae. However, of the spire whorls); aperture more or less circular, with Morton (1952) and Climo (1971, 1973) have shown that single or double peristome, at most weakly reflexed. anatomically these genera have more features in common Protoconch 410–930 µm in diameter and of 1.60–2.25 than either have with extralimital taxa. Thus Climo (1975) whorls. Teleoconch of 2.2–5.2 whorls. Last part of proto- and Powell (1979) treated these New Zealand genera as conch and entire teleoconch ornamented with collabral belonging to an endemic family. riblets. Periostracum on teleoconch smooth, or produced Nonetheless, Ponder & Warén (1988) have assigned at summits of riblets as thin lamellae, and sometimes Cytora and Liarea to Liareinae in Pupinidae. This assign- elaborated as hair-like processes. No apparent sexual ment to Pupinidae was followed by Spencer & Willan dimorphism in shell characters. Operculum thin, horny, (1996), Spencer et al. (2002), Barker (2005) and Bouchet not calcareous, with central to subcentral nucleus and few & Rocroi (2005). For nomenclatural stability, and pending whorls. Animal possessing short tentacles, each rounded at a molecular-based phylogenetic study, in the present work the apex, and with eyes situated on a lobe at the outer base. we regard Cytora and Liarea as pupinids. Pallial cavity lacking a ctenidium or other specialised respi- ratory structures; its roof composed of smooth vascularised Genus Cytora Kobelt & Möllendorff, epithelium; with broad hypobranchial gland extending 1897 along the right side of the roof. Buccal organ equipped with a pair of chitinous jaw plates appressed to the roof of Cytora Kobelt & Möllendorff, 1897: 85. Type species (by the buccal cavity. Radula taenioglossate, having formula tautonomy): Cyclophorus cytora Gray, 1850; Recent, 2+1+1+1+2; central tooth with basal plate contracted near northern New Zealand (gender feminine). its base, broadly rounded to a head bearing five to seven Murdochia Ancey, 1901: 24. Nomen nudum. Introduced cusps, of which the median one is often largest; lateral and collectively for New Zealand species referred to Lago- inner marginal teeth of similar structure; marginal teeth chilus Blanford, 1864, none of which, however, were elongated, obliquely oriented, more slender than laterals, associated by name: fails to conform to ICZN (1999) the basal plate characterised by a rectangular lateral reflec- Article 12. tion separated from the cuspid head by a distinct notch. Cyclophorus as applied by various authors to New Zealand Dioecious. Males possessing a tubular vas deferens, arising material. (Not Montfort, 1810: 290. Type species Helix from the testes located in the shell apical whorls, emergent volvulus O.F. Müller, 1774, by monotypy). from the visceral organs at the posterior of the pallial Cyclostoma as applied by various authors to New Zealand material. (Not Lamarck, 1799: 74. Type species Nerita cavity to run forward as a prostatic organ just below the elegans O.F. Müller, 1774, by monotypy). rectum, then penetrates the length of the cephalic penis Leptopoma as applied by various authors to New Zealand and its apical intromittant organ; penis with one or more material. (Not L. Pfeiffer, 1847: 47. Type species
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