ZoologicalJournal of the I_dnneanSociety (1995), 113:47-91. With 29 figures The anatomy and systematics of the New Caledonian land snail genus Draparnaudia Montrouzier, 1859 (Pulmonata: Orthurethra) SIMON TILLIER Museum National d'Histoire NatureUe (CNRS URA 699), 55, Rue de Buffon 75005 Paris, France AND PETER B. MORDAN The Natural Histo~ Museum, CromwellRoad, London SW7 5BD Received December 1993, revisedand acceptedfor publication J@ 1994 A comprehensive study of the anatomy, ecology and distribution of the orthurethran genus Draparnaudia confirms its family-group status, and suggests a probable sister-group relationship with another Pacific family, the Partulidae; its previous position as a subfamily of the Enidae is refuted. Draparnaudia is endemic to New Caledonia, but records from the New Hebrides are thought to result from introductions. A total of six species are recognized in the taxonomic revision, including two new species. Spermatophores are recorded for the first time. ADDITIONAL KEY WORDS:~permatophores - taxonomy - New Hebrides - Mollusca. CONTENTS Introduction ................... 48 Material and methods ................. 49 Systematic position .................. 49 Endemic status in New Caledonia .............. 52 Ecology .................... 53 Multivariate analysis ................. 54 Systematic treatment ................. 56 Draparnaudia Montrouzier, 1859 .............. 56 D. michaudi Montrouzier, 1859 .............. 57 D. s/ngu/ar/s Reeve, 1854 ............... 65 D. gass~si Pilsbry, 1902 ................ 70 D. anniae sp. nov ................. 76 D. subnecata sp. nov ................ 80 D. walken"Sykes, 1903 .............. 83 Acknowledgements ................ 86 References .................. 86 Appendix ................... 88 47 0024-4082/95/001047+43 $08.00/0 © 1995 The Linnean Society of London 48 S. TILLIER AND P. B. MORDAN INTRODUCTION The enigmatic sinistral New Caledonian land snail genus Draparnaudia was first described by the French missionary Montrouzier (1859). One hundred years later, Solem (1959) was still uncertain of its relationships; even though Moss & Webb (1897) had figured the jaw, radula, central nervous system and terminal genitalia of material from the Loyalty Islands, he wrote: "without the study of the pallial region, however, speculation on affinities is useless, and I have left the species, with some hesitation, in the Camaenidae". In this, Solem was following Pilsbry (1899, 1909-10) who was quite clear that Draparnaudia was closer to Papuina than to any other genus. Draparnaudia was reviewed again by Solem (1961) but he was still able to say no more than that "only dissection of the soft parts can determine whether it is a camaenid, a bulimulid or a tornatellinid". In 1958 two adult specimens referred to Draparnaudia walkeri Sykes were collected from Tadine on the island of Mar~ in the Loyalty Islands and their anatomy, including that of the pallial system, was described by Solem (1962). This clearly placed Daparnaudia within the Orthurethra, and Solem created a new monotypic subfamily of Enidae, the Draparnaudinae, but he still considered the family position to be 'uncertain'. More recently both Tillier (1989) and Mordan (1992) have discussed the- higher relationships of Draparnaudia. Tillier gave detailed figures of the pulmonary complex, kidney internal morphology and nervous system, and, broadly following Solem (1962), included Draparnaudia in the Cerastinae for the purposes of his analysis. Mordan, however, excluded the genus from his cladistic treatment of the cerastines. Draparnaudia does seem to form a discrete, extremely well-defined unit, much like the Partulidae, whose exact relationships remain uncertain, but which are clearly of considerable interest and importance in the context of the origins and evolution of the Pacific orthurethran fauna. To date, all the taxonomic reviews of Draparnaudia have been based on shell characters alone and have shown remarkable consistency in terms of the number of species recognized. Pilsbry (1901-2) recognized five species within the genus, as did Solem (1961). However, the shells of Draparnaudia do not exhibit many taxonomically useful characters and, because of this, no satisfactory taxonomic revision exists. The present account relies heavily on information from an anatomical survey of the genus in New Caledonia, and forms part of a series of papers on the New Caledonian landsnail fauna (Tillier, 1981; Solem, Tillier & Mordan, 1984; Mordan & Tillier, 1986; Tillier and Mordan, 1989). It recognizes six species, including two new ones: Draparnaudia michaudi Montrouzier, D. singularis Reeve, D. anniae sp. nov., D. subnecata sp. nov., D. gassiesi Pilsbry and D. walkeri Sykes. The present paper first considers the relationships of Draparnaudia in the light of recent work on the Orthurethra. There then follows a discussion of the endemic status of the genus in New Caledonia, and some general comments on ecology. The paper concludes with a taxonomic revision of Draparnaudia based principally on an extensive collection from 108 sites made between 1978 and 1988, and housed in the Museum National d'Histoire Naturelle, Paris (MNHN). The results of multivariate analyses based on shell, anatomical and radular characters are presented prior to the formal taxonomic treatment. NEW CALEDONIAN SNAIL GENUS 49 MATERIAL AND METHODS The complete list of collecting stations in New Caledonia is given in the Appendix. Additionally, a list of material of each species is given by station number under the systematic section for that species, anatomical material being indicated by an asterisk. A small amount of preserved material from the Field Museum of Natural History (FMNH), some of which had already been dissected, was also studied but no further dissections made; full details of this are given at the end of the Appendix, and FMNH numbers are also listed under each species. All preserved material in the MNHN collections was relaxed and then preserved in 70% ethanol. Only sexually mature specimens were considered for the anatomical analysis, i.e. those with a clearly developed apertural lip and a hermaphrodite duct containing sperm (sensu Bayne, 1973). For the multivariate analysis, data were collected on mature alcohol-preserved specimens; measurements were taken of maximum shell height and width, maximum apertural height and width, number of whorls, penis length, height of the epiphallar pore above the base of the penis, length of epiphallus, length of vagina, and length of free oviduct. In addition, the following measurements were recorded from SEM preparations of the radulae of these animals: width of mesocone of central tooth, width of basal plate of central tooth, height of basal plate of central tooth, and number of teeth per half row (Table 1). A series of principal component analyses was undertaken on a data set of these 14 metric variables from a total of 42 adults; typically only one living adult specimen was recorded per site, and hence only one was dissected, but where possible more than one was utilized (Table 1). The MVSP multivariate statistics shareware package, Version 2, distributed by Warren L. Kovach, University of Wales, was used for the analyses. SYSTEMATIC POSITION Solem (1962) created a new subfamily, the Draparnaudinae, within the Enidae to hold the single genus Draparnaudia, defining it as follows: Orthurethrous holopods, foot with median groove, spermatheca enlarged in basal portion, penis unforked with subapical epiphallic insertion through a large stimulatory pad. No appendages on spermatheca or penis. Radula enid with central reduced in size, bicuspid laterals and multicuspid marginals. Jaw crescent shaped with very weak striations. Solem (1962: 219) obviously had problems placing this subfamily with any precision: It is quite difficult to know where to classify this group, since the anatomy of the Orthurethra is known to show many parallelisms and convergences. Combinations of the varying characters outlined above exclude Draparnaudia from most family units and certain features show strong similarities to various families. He pointed out that Draparnaudia lacks the penial appendages of the Amastridae, which also differ in having the genital pore well behind the optic tentacle, close to the mantle; and he stated that the Partulidae have a shortened, triangular kidney quite unlike any other orthurethran. He also excluded the Achatinellidae (as the Tornatellinidae) on the basis of their radula and greatly 50 S. TILLIER AND P. B. MORDAN TAnLE 1. Character matrix used in principal component analysis. ST, station; SP, species; SH, maximum shell height; SD, maximum shell diameter; AH, maximum apertural height; AD, maximum apertural diameter; WH, whorls; PL, penis length, EP, height of epiphallar pore above penis base; EL, length of epiphallus; VA, vaginal length; FO, length of free oviduct; MW, width of mesocone of central tooth; PW, width of basal plate of central tooth; PL, height of basal plate of central tooth; TN, number of teeth per half row. Shell measurements in mm.; radular measurements in ~m.; reproductive measurements all to same scale. Species code as for Figs 1,2 ST SP SH SD AId AD WH PL EP EL VA FO IVIW PW PL TN 3 M 10.2 7.8 4.8 5.4 6.7 40 19 90 7 45 8.2 11.1 17.5 24 5 M 12.2 9.7 5.5 7.0 6.7 66 44 118 33 45 9.2 13.5 18.6 25 26b S 9.5 8.4 4.2 5.4 6.8 52 33 204 12 42 6.4 8.0 17.6 24 37a S 10.9 9.9 5.0 6.8 6.3 41 23 228 28 50 7.0 9.0 21.5 22 43a S 10.5 8.8 5.5 6.1 6.8 51 35 207 42 22 6.4 8.9 15.9 16 47 S 11.8 9.3