The Anatomy and Systematics of the New Caledonian Land Snail Genus Draparnaudia Montrouzier, 1859 (Pulmonata: Orthurethra)

Home , Amastridae, Eua (gastropod), Orthurethra, Papuina, Partulidae, Pupilloidea

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 4.3 6.0 6.7 40 13 240 23 38 7.0 10.5 17.0 23 56 M 9.0 7.9 4.2 5.1 6.4 45 25 98 21 52 - - - 69 M 9.2 8.1 4.5 5.5 6.8 46 30 125 17 38 7.2 10.9 20.0 23 73 M 9.7 8.0 4.6 5.4 6.5 56 14 151 8 27 6.7 9.0 20.0 19 76 M 10.8 8.1 5.0 5.9 6.7 72 50 103 42 35 7.3 11.6 18.9 25 87 S 8.1 7.2 3.8 4.5 6.3 61 42 148 21 36 6.0 9.9 17.0 19 87 S 8.1 7.2 3.8 4.0 6.3 60 42 156 22 36 6.0 9.0 17.0 19 91b S 8.8 8.0 4.1 5.2 6.1 48 40 165 14 46 7.3 9.4 15.6 23 101a S 8.9 8.4 4.1 4.9 7.0 54 38 216 13 42 7.9 10.2 16.4 22 1011) S 7.4 7.3 3.0 4.2 6.3 38 23 [60 19 35 6.2 8.5 17.5 [8 127 S 10.2 9.3 4.7 5.6 6.3 45 27 191 22 36 7.6 9.6 20.7 24 131 S 9.2 7.6 4.1 5.1 6.5 41 25 205 8 26 6.3 9.0 16.0 22 140a S 11.0 9.5 4.4 6.4 6.8 50 35 144 37 51 6.7 9.0 20.0 19 140e S 9.0 7.4 4.2 5.1 6.1 50 36 154 12 50 7.0 11.0 17.0 19 166 W 9.4 8.2 4.4 5.5 6.5 46 26 110 23 31 - - 173 M 13.7 10.0 6.6 5.8 6.6 61 42 159 20 67 - - 201 M 8.5 6.6 4.2 4.4 6.1 35 18 68 15 33 5.1 8.6 19.9 24 206 G 4.4 3.8 2.1 2.6 5.2 28 15 100 9 14 3.2 5.5 11.5 16 223 S 8.4 7.6 4.0 4.6 6.4 55 37 190 12 37 6.7 8.8 17.1 22 224 G 4.5 4.0 2.1 2.5 5.1 30 21 58 12 18 4.5 8.0 9.5 17 234 S 9.4 8.4 4.3 5.7 6.5 41 19 186 22 45 6.0 10.4 18.3 22 238 S 9.1 7.6 3.9 5.5 6.5 26 20 149 7 35 6.2 7.5 16.5 21 244 S 10.0 9.6 5.0 6.9 6.0 32 10 249 21 37 6.4 10.8 18.8 20 246 S 7.3 6.6 3.2 4.2 5.8 53 39 135 10 43 5.5 7.7 18.3 23 247 S [0.9 9.0 6.5 5.0 6.6 60 43 255 29 30 7.7 10.7 20.5 21 248 S 10.0 10.0 5.0 6.8 6.9 74 50 245 28 54 8.5 13.5 21.2 24 253a A 9.4 7.7 3.8 5.0 6.0 67 55 164 65 27 6.5 12.0 15.0 25 253a M 11.9 8.4 4.8 6. l 6.9 71 52 160 40 67 7.0 11.0 19.0 22 262 W 7.5 7.0 3.7 4.8 6.0 40 22 124 24 20 5.7 7.8 15.2 24 284a SUB 11.7 8.1 5.8 6.0 6.6 35 12 97 17 60 6.5 12.0 17.5 23 307 M [ 1.3 7.8 4.5 5.7 6.7 50 13 100 18 53 7.5 10.0 18.5 20 319 S 9.3 8.2 3.9 5.7 6.5 35 25 163 14 32 8.3 10.0 19.4 23 326 W 7.3 6.5 3.4 4.1 6.1 40 29 213 12 35 7.5 11.0 13.5 23 326 W 7.5 6.4 3.6 4.3 6.1 45 35 210 10 36 7.0 10.5 14.0 22 330 M 11.2 8.2 5.0 5.5 6.4 55 32 120 22 41 7.0 7.5 19.0 21 330 M 11.2 8.9 6.0 6.5 6.1 76 45 131 21 81 6.4 7.3 18.7 21 330 M 11.0 8.6 5.5 5.6 6.4 54 35 124 23 71 6.5 7.0 17.7 20

shortened uterus and prostate complex. Solem concluded that "Draparnaudia is thus excluded .... from any of the Pacific Ocean orthurethran families". The kidney of Draparnaudia is of the typical orthurethran type. It lacks the characteristic renal and rectal folds of the Cerastinae (Mordan, 1984). Solem's (1962) exclusion of the Partulidae, a wide-spread Pacific taxon (Cowie, 1992), NEW CALEDONIAN SNAIL GENUS 51 on the basis of the shortened kidney is not justified. Kondo described the kidney of Samoana burchi as being "the usual orthurethran elongate triangular organ" and his figure (Kondo, 1973: fig. 3D) is indeed of the typical orthurethran type, as are those of Partula sp. indet, figured by Pilsbry (1909-10: P1. 42, fig. 6) and P. caledonica figured by Tillier (1989: figs 30, 31). Clearly there is nothing unique in the kidney morphology of the Partulidae, which differs from that of most orthurethrans only in having a relatively short kidney in reladon to lung length, a character shared at least with some Chondrinidae (Tillier, 1989: fig. 66). The only detailed review of the orthurethran alimentary system is by Tillier (1989), who includes information on Draparnaudia. In terms of the character states used by Tillier for his phylogenetic reconstruction, Draparnaudia differs from the amastrids, achatinellids and partulids in the intestinal loops, which are longer in Draparnaudia; it also has a funnel-shaped crop, widening from the oesophagus to the stomach, unlike the partulids in which it is cylindrical, but does have a differentiated gastric crop without any constriction as in the partulid Eua. In general terms the alimentary system would seem closest to the enid/cerasdne condition, but this itself is somewhat variable within that group. Tillier (1989: 13) noted an interesting observation on the buccal mass of Partula, which is not withdrawn through the nerve ring on retraction of the body, but which is pulled back above the nerve ring by means of sliding the oesophagus forward through the ring. We have also observed this phenomenon in Draparnaudia, as well as enids, cerastines, achatinellids and amastrids, and it may well be the normal situation in orthurethrans. Tillier (1989) has also dealt with the central nervous system (CNS) of orthurethrans, and illustrates that of Draparnaudia michaudi in his figures 143, 144, page 129. In terms of the character states utilized by Tillier, Draparnaudia falls within the range of the 'enid' clade, but also shares the entire suite of nine characters with the various Partulidae, all three genera of which were analysed. Of these three partulid genera, Draparnaudia most closely resembles Samoana, differing only in the relative lengths of the left and right cerebro-pedal connectives; these are subequal in length (and therefore more primitive) in Draparnaudia, whereas in Eua the left is markedly longer. Both the achatinellids and amastrids differ significantly in key aspects of their CNS. Achatinellids differ in the position of the right pleural ganglion (more advanced), visceral ganglion (less advanced) and left parietal ganglion (less advanced); amastrids in the cerebral commissure (less advanced), length of the right cerebro-pedal connective (less advanced), the ratio of the lengths of the left and right cerebro- pedal connectives (more advanced) and position of the visceral ganglion (more advanced). These characters are complex, and the only point that can reasonably be made is that, on the basis of CNS anatomy, Draparnaudia appears to be a member of the partulid/enid clade sensu Tillier. The reproductive system of Draparnaudia is characteristic, with its lack of penial appendages, greatly thickened spermatheca leading directly to the vagina, with the free oviduct as a discrete insertion on the side, and the long, tubular epiphallus. The only orthurethran family with a similar system is the Partulidae, which differs only in lacking the epiphallus. Pilsbry (1909-10: pl. 43, fig. 5) illustrates the reproductive system of Partula rosea Broderip, where the spermatheca and vagina form a continuous thickened tube with the free oviduct inserting 52 S. TILLIER AND P. B. MORDAN laterally; this is less evident in Samoana (Kondo, 1973: fig. 3A). However, loss of penial appendages is generally accepted to be an advanced character within the Orthurethra, so whilst it may be considered a synapomorphy of these two groups, it could equally well be convergent. The Achatinellidae are quite distinct from the amastrid-cochlicopid lineage (e.g. Tillier, 1989). The achatinellids were treated by Cooke & Kondo (1960), and differ from Draparnaudia in numerous respects. The penis, when developed, normally possesses a terminal appendix. According to Cooke and Kondo only one group, the Auriculelfinae, has an epiphallus (viewed by them as an advanced character) but as others such as the Pitysinae produce spermatophores, this can hardly be the case; the epiphallus of AuriculeUa is a short bulbous enlargement of the vas deferens adjacent to the penis. The spermatheca is of the typical orthurethran type, not thickened as in Draparnaudia and Partula. Having excluded the possibility of a relationship with any of the endemic Pacific groups, Solem (1962) concluded that "association of Draparnaudia with the Enidae is based on the radula, the toothless, relatively smooth shell, and the similarity of the apical genitalia to that of the enid Pseudochondrula" (= Imparietula Hesse, 1933) (Forcart, 1940; Zilch, 1959). The genitalia of Imparietula are well-known, and exhibit the hermaphrodite duct culs-de-sac as in all northern Enidae (Mordan, 1986), a structure lacking in Draparnaudia, and it is difficult to understand the basis of Solem's comparison. Solem (1973) examined under the SEM the radulae of three genera of Enidae sensu Solem (the two cerastine genera Rhachistia and Amimopina, and Draparnaudia) and two of Partulidae (Partula and Samoana) and concluded that the radulae of the 'enids' and partulids were quite distinct from each other, and hence could not be used as an argument for their relationship. Subsequently Solem (1988: 517) reinterpreted the same data to show how the cerastine radula differed from that of Draparnaudia: "both cerastine genera have a two-pronged catching cusp, a spade- like main cusp, and lack significant inter-row basal plate support systems, whereas Draparnaudia has a single-pronged catching cusp, a blunt-pointed main cusp, and prominent inter-row basal support system". Solem (1992) later stressed the conservative nature of the various inter-row support mechanisms in pulmonates, and hence their value as taxonomic characters. Of Solem's three reasons for including the Draparnaudinae within the Enidae only the 'smooth, toothless shell' remains, a very weak basis for association. Solem was right to emphasize the difficulty of positioning Draparnaudia within the Orthurethra, but on the basis of the above discussion it would appear to fall in the 'enid/partulid' lineage of Tillier (1989). In respect of the genital system it may possibly be more closely associated with the Partulidae than with either the cerastines or the northern Enidae sensu Mordan (1986). Conversely, the alimentary system appears closer to the latter group. Clearly, a further, more comprehensive cladistic analysis is required to clarify the relationships of the various monophyletic groups of Pacific orthurethrans whose anatomy is now becoming much better known.

ENDEMIC STATUS IN NEW CALEDONIA The type localities of two species and one subspecies of Draparnaudia, singularis Reeve and walkeri Sykes, 1903, and singularis diminuta Ancey, 1905 are given as NEW CALEDONIAN SNAIL GENUS 53 Aneiteum or Espiritu Santo, both islands in the New Hebrides group, but in the last 90 years there appear to have been only three further published records of shells of Draparnaudia from the New Hebrides in the literature, despite intensive collecting: Solem (1959) lists a specimen from the Academy of Natural Sciences, Philadelphia from 'New Hebrides' (ANSP 31512) which he attributes to singularis, and one collected by Robert E. Kuntz from Espiritu Santo which was placed in walkeri, as was the third from Espiritu Santo collected by Borys Malkyn and now in the collections of the Field Museum of Natural History (FNHM 109410) (Solem, 1962). Whilst these later records cannot reasonably be questioned, some of the earlier type localities might possibly be open to doubt; the problem relating to the type locality of singula~is is discussed under that species. What is clear is that the presence of Draparnaudia on the New Hebrides is based upon a handful of shell records, and cannot compare with the abundant material available from New Caledonia, even in old museum collections. The most reasonable explanation for this discrepancy is that Draparnaudia is an endemic New Caledonian genus, of which there has been a very limited artificial introduction into perhaps two of the islands of the New Hebrides. There is no evidence that the New Hebridean shells differ at the specific level from known New Caledonian taxa, nor, therefore, that there has been any endemic radiation in the New Hebrides. In Solem's review of the systematics and zoogeography of the New Hebridean land snail fauna (Solem, 1959: 122) he cites Draparnaudia singularis as "the only species of land snail found in both New Caledonia and the New Hebrides which has not been obviously introduced by man". This view no longer seems tenable. The New Hebrides appear to share no land snail species with New Caledonia, other than those introduced anthropogenically. This is not altogether surprising in view of the geological histories of these two island groups, which are on different tectonic units, separated by a deep oceanic trench, and have only become geographically close in recent geological time. Moreover, the New Hebrides are much younger, being of Tertiary origin, and had a much smaller land surface than at present (possibly less than one-tenth • present area) until the Pleistocene (Mallick, 1975).

ECOLOGY Draparnaudia is found in nearly all kinds of primary environments in New Caledonia where trees occur; like most, if not all, New Caledonian land snails, it is only rarely found in true secondary environments. In rainforests the two larger and commoner species of Draparnaudia (singularis and michaudz) are most commonly encountered on both living and dead tree trunks, which have not fallen on the ground. The individuals seem to occur in aggregates, and most often cover their shells with particles from the trunk surface: wood dust on dead tree trunks (observed in st. 140e), various kinds of debris, or even apparently live moss where this covers the trunk. As a result, it is generally very difficult to see the live animals on the trunks. Although they are predominantly arboreal, Draparnaudia may exceptionally be found alive in rainforest leaf litter, but this was noted only once during the course of 26 systematic samples of 25 m 2 each made between 1986 and 1987 in Rivi~re Bleue (sts 250, 251). 54 S. TILLIER AND P. B. MORDAN In drier environments (mesophil and xerophil forests), D. michaudi and singularis may also be found on tree trunks, but are generally not covered with debris. In the western part of the mainland, they have occasionally been collected crawling on calcareous rock surfaces (st. 217 and 326). The two flammulate species, D. anniae and D. subnecata, are each endemic to the summital region of one of the isolated massifs in the northwest of the mainland. Both occurred in aggregates on Araucaria trunks, shared in one case with D. michaudi, and live in sparse populations in a vegetation consisting of low, sometimes burnt, maquis. The two smaller species, D. walkeri and D. gassiesi, occur mainly in relatively dry environments and, in the case of the latter, only on the mainland. The few specimens collected alive were crawling on rock surfaces (st. 326, 224) or in leaf litter, but records are too sparse to allow any definitive conclusion and these species could just as well be predominantly trunk-dwellers as are the larger ones. The four species which are not highly restricted in distribution seem to show different tolerances to moisture, as expressed by mean annual rainfall of the collecting stations: D. singula~s is commonly found in wet rainforests; with one exception (discussed below) D. michaudi is not found where rainfall exceeds c. 3200 ram, and D. walken" seems to be a lowland species of still-drier environments, which was not found where rainfall exceeds 2100 mm. The fourth widespread species, D. gassiesi, was found too infrequently to allow any useful conclusions, but possibly covers the whole rainfall range of the genus, or most of its lower part, in a niche which probably does not overlap with other species owing to its much smaller size. These varying abilities to support moisture may correspond to real moisture preferences in microhabitat, but at the present broad level of analysis we can only describe the gradation observed, as it may be easier to find a moist microhabitat in a relatively dry environment than vice versa.

MULTIVARIATE ANALYSIS The data for the multivariate analyses comprised five measurements of the shell, five of the reproductive system, and four of the radula taken from the SEM micrographs (Table 1). The data were untransformed, but standardized within MVSP, and of the material dissected; michaudi from Khtdeigne, Nanawa and Ntc6 (all Loyalty Islands) were lacking radula data and hence were not used in the radula analysis. The main analysis was made on shell and reproductive characters. The plots of axes 1 and 2, and axes 2 and 3 are shown in Figures 1 and 2, and the various eigenvalues and eigenvectors (component loadings) for each of the first three axes are given in Tables 2 and 3. The first three axes account for over 80% of the total variation. It is clear that axis one, with consistently positive eigenvectors and uniformly high loadings from all the conchological characters, represents a size factor. It accounts for 53% of the variation. This explains the separation of the two gassiesi stations (Tonghout, St. 224; Mtnazi, St. 206), gassiesi being much smaller than the other species. Axis two is dominated by high positive loadings for penis length, height of insertion of the epiphallar pore on the penis, and vagina length. Axis three, on the other hand, has a very high positive loading for epiphallar length, and also a large negative contribution for the length of the NEW CALEDONIAN SNAIL GENUS 55

0.59 '

0.48 -

0.37 M

O. 26 S S M S G S 0.15 S M G S oJ S M u) W S S S 0.04 x W < M S S M -0.07 MW M S S -0.18 S M S M M -0.29 S S

S -0.39 SUB -1.08 -0:93 -0:78 -0:63 -0:48 -0~34 -0:19 -0.04 0[11 0'.26 0141 0.56 AXIS 1 Figure I. Plot of axes one and two from Principal Components Analysis utilizing shell and reproductive characters of Draparnaudia. M, michaudi; S, s/ngu/ar&,, G, gasst~s/; A, annh2e; SUB, subnecata; W, walked. I

0.37-

O. 28- S / S,, i 0.19- S S S w 0.11 S S S S W W

0.02 S M W S S -0.07- M S S -0.16- M M M M -0.24 SUB M M -0.33

-0.42 -0.39 -0'.31 -0"22 -0'.13 -0'.04 0'.05 0'.14 0'.23 0'.32 0'.41 0150 0.59 AXIS 2 Figure 2. Plot of axes two and three from Principal Components Analysis utilizing shell and reproductive characters of Draparnaudia. M, m~:haud/; S, szhgu/arfi';, G, gass/es/; A, anmoe; SUB, subnecata; W, walken'. free oviduct. This third axis successfully separates singularis, with its relatively long epiphallus and its generally rather short free oviduct, predominantly into the positive half of the plot, and michaudi, with its shorter epiphallus and typically longer free oviduct, into the negative area, but there is a degree of 56 S. TILLIER AND P. B. MORDAN TABLE 2. Eigenvalues and percentage of of total variance of axes 1-3 of principal component analysis

Percentage of Axis Eigenvalue total variance Cumulative total

1 5.345 53.45 53.45 2 1.642 16.42 69.87 3 1.190 11.90 81.77

TABLE 3. Eigenvectors (component loadings) for each character, axes 1-3 of principal component analysis

Character Axis I Axis 2 Axis 3

Shell height 0.400 -- 0.181 - 0.086 Shell diameter 0.401 -- 0.180 0.161 Apertural height 0.375 -0.155 -0.132 Apertural diameter 0.379 --0.247 0.052 No. whorls 0.336 --0.148 0.127. Penial length 0.288 0.496 -0.198 Position of epiphallar pore 0.208 0.642 -0.065 Length epiphallus 0.166 0.01 I 0.715 Length vagina 0.237 0.399 0.247 Length free oviduct 0.267 -0.103 -0.562 overlap. D. walkeri, represented by four specimens from three localities, falls broadly in the middle of the scatter of singularis in the plot of axes 2 and 3, but occupies the negative extremity of the singularis scatter along the first axis in the plot of axis 1 on 2 (Fig. 1), denoting its small size relative to most singularis. In the axis 2 and 3 plot the two gassiesi specimens lie close together, demonstrating their similarity in respect of reproductive structures, but anniae and subnecata are at opposite ends of axis 2, showing greater variation along this axis than throughout the entire range of either singularis or michaudi. D. anniae has a high positive component score along axis 2 (0.588), reflecting its relatively much longer penis and high epiphaller insertion; subnecata, on the other hand, has a high negative value (--0.395) resulting from its short penis and low epiphallar insertion. There are minor differences between these two species along the third axis, but these are of the sort of magnitude found within both the singularis and michaudi scatters along this axis. A second principal components analysis of radular characters was undertaken separately, but other than separating gassiesi on the first (size) axis, it simply demonstrated the generally poor value of the radula in species descrimination in stylommatophorans (see, for example, Kemperman, 1992).

SYSTEMATIC TREATMENT Draparnaudia Montrouzier, 1859 Type species. Draparnaudia michaudi Montrouzier, 1859 [Nomenclatural note. Draparnaudia was first published in synonymy, having previously been a manuscript name. According to Article 11 e of the ICZN Code, a NEW CALEDONIAN SNAIL GENUS 57 name published in synonymy dates from its first publication as a synonym; the type species is that nominal species first directly associated with it under an available species-group name (Article 67 1), i.e. michaudi Montrouzier, 1859.]

Draparnaudia michaudi Montrouzier, 1859 Synonymy Helix sinistrorsa Deshayes, 1840: 24, pl. 161, figs 19-21 (non Serres, 1838) [New Caledonia]. Bulimus sinistrorsus var. B. castaneofasciatus Montrouzier, 1859: 287, pl. VIII, fig. 3. [Ile Art]. Draparnaudia michaudi Montrouzier, 1859:288 [New Caledonia]. Bulimus theobaldianus Gassies, 1870: 143. (non theobaldianus Benson, 1857). [Lifu, Loyalty Islands]. Bulimus sinistrorsus Deshayes. Pfeiffer, 1853: 322; Gassies, 1871: 92. Bulimus sinistrorsus Var. castaneozonulatus Gassies, 1871: 92. Bulimus sinistrorsus Var. albidozonulatus Gassies, 1871: 92. Bulimus theobaldianus Gassies. Gassies, 1871: 93, pl. III, fig. 8. Pseudopartula sinistrorsa (Deshayes). Ancey, 1882: 87; Crosse, 1894: 247. Pseudopartula theobaldiana (Gassies). Ancey, 1882: 87; Crosse, 1894: 248. Pseudopartula sinistrorsa Var. beta. castaneofasciata (Montrouzier). Crosse, 1894: 247. Pseudopartula sinistrorsa Var. gamma, albidozonulata (Gassies). Crosse, 1894: 247. Draparnaudia lifuana Pilsbry, 1901:17, pl. 3, figs 7-9. (nora. nov. pro theobaldianus Gassies) [-Lifu, Loyalty Islands]. Draparnaudia crossei Pilsbry, 1901: 17, pl. 3, figs 10-11. [New Caledonia]; Franc, 1956: 164, pl. 21, fig. 219. Draparnaudia lifouana Pilsbry. Dautzenberg, 1923: 143; Franc, 1956: 164, pl. 21, fig. 220; Solem, 1962: 220. Draparnaudia sinistrorsa (Deshayes). Ancey, 1898: 147; Dautzenberg, 1923: 143; Franc, 1956: 163, pl. 21, fig. 218. Draparnaudia michaudi Montrouzier. Solem, 1961 : 483.

Material examined Type. One type of theobaldianus Gassies, BMNH Reg. no. 1883.11.10.1156. This specimen here selected as the lectotype (Fig. 3E, F). Other. Stations 3", 4, 5", 6, 7, 11, 13, 16", 16a, 20, 56", 69", 73", 76", 90, 97, 102, 1.08, 156, 157, 161, 165, 166", 167, 168, 169, 172, 173", 174, 175, 176, 177, 178, 201", 253a, 253a, 255, 262, 285, 306, 307", 326", 330*. FMNH material: 159229, 159322, 159328, 159350.

Shell (Fig. 3) Shell sinistral, open but narrow umbilicus, approximately one-tenth the maximal shell diameter, globose-conic, thick, opaque colourless to mid-brown, becoming paler with wear, often with darker central band. Surface matt, with irregular radial oblique, often coarse growth lines. Profile of early whorls carinated; canna rarely extending to the body whorl, but if so, becoming progressively lost towards the aperture; sutures wide, with the lower whorl T58 S. TILLIER AND P. B. MORDAN

Figure 3. Shells of D. michaudi. A & B, Specimen labelled sinistrorsus Deshayes, ex. Deshayes Coll. MNHN; C & D, Station 16a. Bobeito. E & F, Lectotype of theobaldianus Gassies. BMNH Reg no. 1883.11.10.1156. Gassies Coll. G & H, Mart, Loyalty Isles, BMNH Reg. no. 1928.5.5.152. Brazier Coll. I & J, Station 307, Koniambo. K,L. Specimen labelled sinistrorsus Deshayes, ex Deshayes Coll. MNHN. NEW CALEDONIAN SNAIL GENUS 59 having a distinct shoulder, giving a somewhat rounded profile to the sides of the spire. Aperture downwardly directed, normally as wide or only slightly wider than high; lip flared and paler than rest of body whorl. Only rarely covered with adherent debris.

Radula (Fig. 4) Typically orthurethran. Central with broad, pointed mesocone and no (or only extremely weak) ectocones. Laterals with rather broader and slightly outwardly directed mesocone, well-developed pointed ectocone slighdy narrower than the mesocone, and quadrate basal plate; endocone lacking. Marginal teeth with short, wide basal plate and progressively finer and more pointed mesocone; ectocone becoming more divided towards the ribbon margin into as many as six fine, sharp denticles. Tooth size varied broadly with shell size. Number of teeth per row varied from 16 to 25 per half-row.

Foot (in preserved specimens) uniformly pale cream coloured, or sole pale with sides pigmented pale brown to brown-grey, often becoming darker towards the back. Animals with darker shells tending to have darker bodies.

Pallial system (Fig. 5) Typically orthurethran; kidney elongate, extending approximately ninetenths the length of the lung cavity, which in turn occupies between 0.9 and 1.3 whorls. Kidney pore subterminal, opening to the left into a short, backwardly directed grove delimited by a weak ridge running parallel to the kidney. Mantle gland lacking. Internal structure figured by Tillier, 1989, Fig. 142.

Reproductive system (Figs 6-8) Ovotestis with 4-5 clavate clumps of acini embedded in the columellar side of the upper whorls of the digestive gland. Hermaphrodite duct expanding into a convoluted seminal vesicle (sensu Bayne, 1973). Talon a simple fold at the base of the duct. Albumen gland small and triangular. Free oviduct appears to insert onto the much thicker spermatheca (which is continuous with the vagina) opposite the opening of the atrium. Vagina attached to the body wall with numerous small muscles. Internally the vagina/spermatheca complex exhibits a series of 8-9 well-defined straight pilasters, which run the full length of the stalk but become weaker as the lumen narrows and are lost before reaching the terminal bulb. Free oviduct equal to or longer than the vagina. Penis with stout terminal retractor muscle inserting on the inner lung wall (diaphragm), and with a lateral epiphallar insertion; epiphallus short, between 1.5 and 2.5 times the length of the penis, and inserting at almost any point along the side of the penis; cylindrical and of broadly uniform diameter throughout its length, sharply narrowing at the junction with the vas deferens which at this point is loosely attached to the base of the penis with connective tissue. Internally the penis has a series of longitudinal pilasters which are thickened below the epiphallar pore, and become thinner above (Fig. 8). The central pilaster, which lies opposite the atrial opening, expands rapidly around the pore to form a prominent pad; two lateral pilasters run between this and the opening to the 60 S. TILLIER AND P. B. MORDAN

Figure 4. Radula of D. michaudi. A, Central and lateral, and B, marginal teeth, station 76, Mt. Kaala; C, Half row, station 69, Mt. Mandjelia. Scale lines: A, B = 10 lain; C = 50 /tm. atrium, and numerous less-pronounced pilasters lie between these larger ones. Penial nerve from the left cerebral ganglion. Atrium not a distinct structure, but simply the area where the free oviduct and penis meet; atrial retractor NEW CALEDONIAN SNAIL GENUS 61

Figure 5. Pallial system of D. michaudi, Station 253a, Mt. Kaala. Abbreviations: AU, auricle; K, kidney; KO, kidney orifice; PN, pneumostome; R, rectum; RF, renal fold; VE, ventricle. shares a branch with the left optical retractor (which passes through the angle of the penis and free oviduct), actually inserting on the body wall next to the gonopore. The gonopore is a simple longitudinal silt, just behind and below the left optic tentacle. Spermatophore Not recorded, but a number of hollow, cylindrical fragments have been found in some of the spermathecae. Geographic variation Although there was no obvious linear cline evident for adult shell size (Table 1), shell height was always greater at the western stations than at stations at the east of the island (Mandjelia, st. 69; Ignambi, st. 73; Cap Bocage, st. 201). The largest dissected specimen was recorded from Lifou, but other dissected material from the Loyalty Islands was in no way exceptional. Although epiphallus length varied by a factor of about two over the entire range, no clear pattern 62 S. TILLIER AND P. B. MORDAN

Figure 6. Reproductive systems of D. michaudi. A, Station 307, Mt. Koniambo; B, Station 56, Nect, Mar& Scale bar = 1 mm. Abbreviations: AG, albumen gland; E, epiphallus; FO, free oviduct; HG, hermaphrodite gland; P, penis; S, spermatheca; V, vagina; VD, vas deferens.

¥" "~.:E'~.. ". " • •

:~" ~, ":?-:' ".... .:"

Figure 7. Reproductive system of D. michaudi, station 69, Mandjtlia. Scale bar = 1 mm. Abbreviations as in Fig. 6. of variation was evident; the same was true of the length of the free oviduct. The vaginal length varied considerably but was always greater at the west- coast stations (5, 76, 253a, 307, 330) than in Mandjelia (st. 69), Ignambi (st. 73) and Cap Bocage (st. 201) near the east coast (Table 1). This may, however, be pardy a function of overall size (see above). NEW CALEDONIAN SNAIL GENUS 63

Figure 8. Penis of D. michaudi, station 56, Nec6, Mar6. Scale bar = 1 mm. Abbreviations: E, epiphallus; EP, ephiphallar pore; PR, penial retractor muscle; V, vagina.

Distribution (Fig. 9) D. michaudi is restricted to the north-western part of the mainland, north of a line running from the valley of the To Ndeu (st. 285) to Nassirah, approximating the transversal Bouloupari/Thio, but extending on the south coast a little further down to Presqu'ile Montagn~s (st. 326). It reaches north to the Belep Islands (Isles Daos du Nord, st. 3), and is also widespread on the Loyalty Islands of Ouv6a, Lifou and Mar6. In addition to being absent from the south-eastern part of the mainland, it has not been recorded from the Isle of Pines. On the mainland it occupies an extremely wide range of habitats from dry calcareous littoral forest (e.g. Pindai, st. 303) to high-altitude rainforest (Ignambi, st. 73) at an elev~ition of 1200-1300 m, as well as high maquis (Kaala, st. 253; 255). At these sites rainfall varies from 700 mm (Presqu'ile Montagn~s, st. 326) to as high as 7000 mm in a single case at Ignambi (st. 73). However, 42 of the 43 collecting stations from which michaudi was recorded have less than 3200 mm annual rainfall, and rise above 500 m in altitude only on the dry massifs of the west coast: the population of the Ignambi, one of the summits of the northeastern isolated Pani6 massif, may represent a differentiated offshoot which we are unable to distinguish morphologically; D. michaudi is basically a species of medium-dry lowland environments. The range of michaudi overlaps with that of singularis in the centre of the mainland, and michaudi is found sympatrically with singularis at Nassirah, Col de Tango (st. 64 S. TILLIER AND P. B. MORDAN

0 ~-1 a .•-'" • °

..0 ~.Q . I I I ~ e

" . Z "'°

~;° U E

? =E p

r--

Z \ NEW CALEDONIAN SNAIL GENUS 65 16), and Bogui (st. 102). Both species are also found on Mt Koniambo, but at different elevations (sts 87 & 307). At Koumac (st. 6) and Presqu'ile Montagnts (st. 326), as well as at Nanawa (st. 166) and Wtdoumel (st. 262) on Lifou, it is found together with the dry lowland species, walkeri, and at Kaala (sts 76 & 253a) with the endemic species anniae.

Comments D. michaudi can generally be distinguished from singularis by both shell and reproductive characters. In michaudi the shell whorl is more rounded with deeper sutures, producing a less flat profile; the body whorl in particular is only weakly carinated, if at all, and never has the sharp carina of typical singularis. The aperture is only slightly wider than high, and more downwardly deflected than in other species. The shell is rarely covered with debris, and. is often pale with, in the Loyalty and Belep islands, a thick spiral brown stripe; this stripe was also present on specimens from Cap Bocage (st. 201) on the mainland. The principal differentiating feature of the genitalia of michaudi is the short length of the epiphallus relative to that of the penis. The shell size of dissected adults (Table 1) was on average higher than in singularis, ranging from a height of 8.5 mm at Cap Bocage (st. 201) to a maximum of 12.25 mm on the mainland (Le Cresson, st. 5) and 13.7 mm on Lifou (Khedtigne, st. 173).

Draparnaudia singularis Reeve, 1854 Synonymy Helix singularis Reeve, 1854: species 1407. [Koondah Mountains, near Calicut, India]. Helix singulalis Pfeiffer, 1855:289 [Isle of Aeiteum, New Hebrides]. Pfeiffer, 1859: 255. Bulimus sinistrorsus Deshayes. Gassies, 1863: 255, pl. II, fig. 3. Bulimus sinistrorsus Var. carinatusmagisstriatus Gassies, 1871: 92. [Ile des Pins]. Pseudopartula sinistrorsa Var. delta, carinatamagisstriata (Gassies). Crosse, 1894: 247. Pseudopartula singularis (Pfeiffer). Ancey, 1882: 87; Crosse, 1894: 248. Draparnaudia singularis (Pfeiffer). Ancey, 1898: 147; Dautzenberg, 1923: 143; Franc, 1956: 162, pl. 21, fig. 216; Solem, 1961: 483. Draparnaudia singularis major Ancey, 1898:147 [New Caledonia]. [Nomenclatural note. Pfeiffer's description of this species was originally read at the Zoological Society of London meeting of 12 December 1854, but not published until 8 May 1855 (Duncan, 1937). Reeve's description dates from December 1854 and thus takes precedence over Pfeiffer's. Reeve gives the type locality as Koondah Mountains, near Calicut, India, which is clearly erroneous; Pfeiffer gives a more credible location of Aneityum in the New Hebrides. Interestingly, Koonda Mountains is the type locality of Helix vaUicola Pfeiffer which is the species immediately following singularis in Pfeiffer's paper, and this may well have been whence Reeve's confusion arose. The original material from the 66 S. TILLIER AND P. B. MORDAN

Figure 10. Shells of D. singularis. A & B, Lectotype of s/ngu/am Pfeiffer. New Hebrides, H. Cuming Coll., BMNH Reg. no. 198098. C & D, Prony Bay, leg. J. Brazier, 1873. BMNH Reg. no. 1928.4.27.173. E & F, Station 25, Adio, dry valley, MNHN.

Cuming Collection, utilized by both Reeve and Pfeiffer, is in the BMNH, and is clearly marked with Pfeiffer's published locality.]

Material examined Types. Three syntypes of singularis Pfeiffer. One here selected as lecto-type, BMNH Reg. no. 198098 (Fig. 10A, B). Other. Stations 16, 22, 25, 26b*, 37a*, 39, 43", 47", 48, 50, 52a, 53, 87", 91b*, 93, 98, 101", 101b*, 102, 107, 112, 127", 131", 140a*, 142, 143, 181, 210, 212, 216, 217, 223", 228, 234", 235, 238", 241, 244", 246", 247", 248", 250, 251, 256a*, 258, 312, 319". FMNH material: 144237, 144238, 144265, 158315, 159270, 159302, 159328. SheU (Fig. 10) Shell sinistral, with open but narrow umbilicus, turbinate-conic, medium thickness, dark brown, becoming paler with wear. Surface matt glossy, with irregular radial oblique growth lines, and weak wavy spiral striae on the upper whorls. Whorl profile usually, but not invariably, sharply carinated, often with distinct narrow pale carinal ridge, carina extending to the body whorl but becoming weaker towards the aperture; sutures very shallow, giving a straight profile to the sides of the spire. Aperture rather downwardly directed, distincdy wider than high; lip flared and paler than rest of body whorl. Typically covered with adherent soil particles and debris. NEW CALEDONIAN SNAIL GENUS 67

Figure 11. Radula of D. sb~gularis. A, Central and lateral teeth and B, marginal teeth, station 238, Pic NE of Mt. Tonta. Scale bar = 10 Itm. Abbreviations as in Fig. 6.

Radula (Fig. 11) As for michaudi; tooth number ranging from 19-24 per half-row.

Varies much as in michaudi, but tends towards a darker grey-brown pigmentation.

PaUial system As for michaudi.

Reproductive system (Figs 12-15) As for michaudi, the main difference being the greater length of the epiphallus which is between 3 and 7 times the length of the penis. The epiphallus inserted on the penis between I/3 and 4/5 up from the base. The vagina is almost always shorter than the free oviduct. 68 S. TILLIER AND P. B. MORDAN

---~E

Figure 12. Reproductive systems of D. dngu/ar/s. A, Station 87, Mt. Koniambo; B, Station 101c, Mt. Me Maoya. Scale bar = lmm. Abbreviations as in Fig. 6.

Spe atophore (Fig. 14) A single spermatophore was found in the dissected specimen from Dent de St. Vincent (st. S19). It was situated in the base of the spermatheca and extended into the vagina. The tip of the open end just entered the atrial region, but was bent back so that the opening was directed away from the atrium. The spermatophore itself was approximately 3.5 mm long, cylindrical, tapering towards the two ends which were blunt and rounded. The wall was soft and smooth. More-or-less at the centre of the spermatophore was a rounded outgrowth, situated at the level of the oviducal opening but pointing away from it. This does not seem to correspond to an equivalent structure in the epiphallus. Sperm was exuding from the open end, and filled the penis, which might explain why this was distended and smooth-walled inside. NEW CALEDONIAN SNAIL GENUS 69

Figure 13. Reproductive systems of D. singularis. A, Station 248, between Wapan and Oup6, Isle of Pines; B, Station 91b, Mt. Aoupini6. Scale bar = lmm. Abbreviations as in Fig. 6.

Geographic variation As with michaudi, size (as shell height of dissected adult snails; Table 1) varied considerably throughout the range of the species, although it seemed relatively constant in animals from the same population. There was a clear change across a line running roughly north/south separating Mt. Ouin (st. 127) and Mt. Dzumac (st. 37a) from Mt. Kouv~l~ (sts. 246 and 223), and passing through Mts. Koghis (sts. 104a and e); to the west height was lower (range 7.3-9.3 mm) than to the east (range 9.4-11.9). A similar line separates populations to the west with relatively short vaginas from those to the east where the vagina is longer; whilst in isolation this could be viewed simply as a function of size (see above), no equivalent variation was evident in the length of the free oviduct which varied by a factor of over two. Island populations showed extremely long (Isle of Pines) or short (W6doumel, Lifou) measurements for the free oviduct. 70 S. TILLIER AND P. B. MORDAN

Fig. 14. Reproductive system and spermatophore of D. singularis, station 319, Dent de St Vincent. Scale bars = lmm. Abbreviations as in Fig. 6 plus SP, spennatophorc. lh'stribution (Fig. 16) D. singularis has been recorded from a very large number of sites in the south-eastern two-thirds of the mainland of New Caledonia, below a line running approximately along the valleys of the Congo and Tipindje. It is also abundant on the Isle of Pines, but has not been recorded from the Loyalty Islands. As with michaudi, singulaTis is found in a wide variety of habitats ranging from partly secondary lowland vegetation (e.g. Goro, st. 50, and Nindah, st. 22) with rainfall around 100 mm to high-altitude rain-forest on Mt. Ningua (st. 181) and Mt. Ouin (st. 127) where precipitation reaches as much as 5000 mm per year. In comparison to D. michaudi, D. singulans is a wet-environment species: two michaudi against 13 singularis stations (out of 43 and 47, respectively) exceed 3000 mm in annual rainfall (;(2=8.67, P<0.01). Sympatric occurrences with rnichaudi have been discussed under that species (above), but additionally singula~is is found with the dry lowland species walkeri at Adio (st. 25) on the mainland, and at Ouro (st. 53) on the Isle of Pines.

Comments In comparison with michaudi, the shell of singulaTis is typically carinated over most of the body whorl in fully grown animals, is normally of a deep horn- brown colour, has a relatively straight-sided profile to the spire, and a wider, more laterally expanded aperture. The shell is also more usually covered with adherent debris, mostly particles from the surface of the tree trunk. The epiphallus is relatively long compared with the penis length.

Draparnaudia gassiesi Pilsbry, 1902 Synonymy Bulimus turgidulus Gassies, 1871: 188. (non Bulimus turgidulus Deshayes, 1864) [L'ile Nou, New Caledonia]. Gassies, 1873: 49, pl. II, fig. 4. NEW CALEDONIAN SNAIL GENUS 71

A .[.... D .. :." i

Fig. 15. Penis and spermatheca of D. singulari~. A, penis, station [01c, Nit. Me Maoya; B, penis, station 87, Mt. Koniambo; C, penis, station 248, between Wapan and Oup6, Isle of Pines; D, spermatheca, station 87, Mt. Koniambo. Scale bar = l mm. Abbreviation: E, epiphallus; EP, epiphallar pore; OP, oviducal pore; PR, penial retractor muscle.

Helix turgidula (Gassies). Pfeiffer, 1876: 388. Pseudopartula tu~gidula (Gassies). Ancey, 1882: 87; Crosse, 1894: 248. Draparnaudia turgidula (Gassies). Pilsbry, 1901-2: 16, pl. 3, figs 5, 6. Franc, 1956: 163, pl. 21, fig. 217. Bulimus gassiesi Pilsbry, 1902: lxxi. (nom. nov. pro Bulimus turgidulus Gassies, 1871. Solem, 1961: 484. Bulimus gassiesi microumbilicata Solem, 1960: 6, fig. 8. [River drift at sea coast a few miles from Bourail, New Caledonia]. Solem, 1961: 484.

Material examined Types. Gassies (1871) described six syntypes of turgidulus from his own collection, collected by P6re Lambert. Two of these are in the BMNH (Reg. No. 1883.11.10.185-6), and a further two are in the MNHN. The likely figured specimen from the latter (Fig. 17E, F) is here selected as the lectotype. Other. Stafons 84, 206*, 224*, 310.

Shell (Fig. 17E-H) Shell sinistral, openly umbilicate, turbinate-conic, thin, light brown. Surface matt glossy, with fine, regular oblique ridges which have a pale crest, giving the fresh shell a whitish sheen. Aperture rounded-oval, peristome expanded with outer and columellar margins connected by a strong callus. Whorls c. 5, 72 S. TILLIER AND P. B, MORDAN

x o~ 0 ~o Q

0 0

-r.

p re 0

Z .r'~ ~ -~"~~~ ~m

• ,."

-r"

04 ,.6 eq .,,

z~ NEW CALEDONIAN SNAIL GENUS 73

Fig. 17. Shells of D. wa'lken" and D. gassk.n'. A & B, Lectotype of walken" Sykes, Renee River, Espiritu Santo, New Hebrides, leg. Jd. Walker, BMNH Reg. no. 1902.12.1.159; C & D, D. walkeri, station 25, Adio, dry valley, MNHN; E & F, Lectotype of turgidulus Gassies, New Caledonia, Gassies CoN, BMNH Reg. no. 1883. 11.10.185; G & H, D. gass#.n', station 301, Mt. Taom, MNHN. rounded in profile, producing deep, even sutures. Fresh shells have adherent soil particles. Size much smaller than other species (see Table 1).

Radula (Fig. 18) The radulae of the two specimens from stations 206 and 204 were examined. Tooth morphology as for michaudi. Basal plates either quadrate (Tonghout) or much more elongate (Mtnazi). Marginal teeth with multicusped ectocones and blunter, inwardly directed mesocone. Both tooth number (16-17 per half-row) and size smaller than in other species (see Table 1).

PaUial system Pallial complex as for D. michaudi. Extending 0.7 whorls in a body length of 4.1 whorls (Kaala) and 0.8 whorls in a body of 4.4 whorls (Tonghour).

Reproductive system (Figs 19, 20) Considerably smaller than other species, with epiphallus between 2 and 3 times the length of the penis, entering penis laterally at, or slightly above, the half-way point. Internally the penis has one strong longitudinal pilaster running the full length of the penis and expanding at the point of entry of the epiphallus to form a rounded protrusion, and four shorter pilasters at the base, again convoluted, and extending above the level of the epiphallar pore as simple, weak folds. These pilasters were much weaker and less convoluted in the Tonghou6 specimen, which may well have only recently become mature (Fig. 20B). Free oviduct about 1.5 times as long as the vagina, both of which 74 S. TILLIER AND P. B. MORDAN

/~ .. ?- ' t/X l....efN

z , , , *

u ~ .,#~--~ ..... " -:.A ~_Z ! 7 ;.j- L-: Fig. 18. Radula of D. gass/esi. A, central and lateral teeth, and B, marginal teeth, station 224, Tonghou~. Scale bar = 10pm. have five longitudinal pilasters. Hermaphrodite gland with four or five clumps of ascini. No spermatophore found.

Geographic variation Insufficient study material was available for any useful comments to be made on variation. However, there are some differences in the reproductive anatomy of the two specimens dissected, notably in the internal penial ornamentation which was much more pronounced in M6nazi than Tonghou6, although the broad pilaster pattern of both is typical of the genus. The two animals were collected at almost the same time of year so that annual changes in sexual maturity are unlikely to account for the differences, but it may well be that the M6nazi specimen was older.

Distribution (Fig. 21) D. gassiesi is known only from a few widely separated sites on the mainland and has not been recorded from any of the islands. No particular pattern is evident; the sites range from high-altitude rainforest such as Tchingou (st. 84) NEW CALEDONIAN SNAIL GENUS 75

E S

P VD ~'~AG

Fig. 19. Reproductive systems of D. gassiesi. A, station 206. M6nazi; B, station 224, Tonghou6. Scale bar= lmm. Abbreviations as in Fig. 6 plus DG, digestive gland. and Taorn (st. 310), to dry, partly secondary forest with around 1000 mm annual precipitation such as Tonghou6 (st. 224). However, rainfall does not exceed 3500 mm at M6nazi and 3100 mm at Tchingou, the two wetter stations, and then drops to 1700 mm on Mr. Taom: as far as may be concluded from such limited data, D. gassiesi seems to be basically a medium- dry to dry-environment species. The type locality of gassiesi is given as 'Nou Island, New Caledonia'; this is more likely to be the island off the capital Noum6a (now connected to it by bridge) than the island of the same name situated at the extreme south-eastern tip of the mainland, as the former was inhabited in the time of P6re Lambert. Additionally there is old Sav6s material in the collections of the MNHN from the Pointe de l'Artillerie, Noum6a, which directly faces the eastern extremity of the Ile Nou; clearly gassiesi also occurred in coastal habitats in the past. Comments This species differs from other Draparnaudia primarily in its very small size at maturity. Both tooth number and adult whorl number are also lower than 76 S. TILLIER AND P. B. MORDAN

Fig. 20. Penis of D. gassiesi. A, station 206. M6nazi; B, station 224, Tonghou6. Scale bar = l mm. Abbreviations as in Fig. 8.

in other species. The shell sculpture of the M6nazi specimen, which was particularly fresh, had a highly characteristic whitish sheen, quite unlike other species. This sculpture may become worn in more mature specimens as there is some evidence from the genital anatomy (see above) that this animal was only recently mature, gassiesi is likely to be considerably more widespread within New Caledonia than recorded here, and might also occur in the Loyalty Islands.

Draparnaudia anniae sp. nov. Type locality. St. 253a, Mt. Kaala, south slope, 810 m, leg. A & S Tillier, 8.x. 1986.

Material examined Holotype: Figs 22C,D. Station 253a, 810 m, 8.x.1986. Paratypes: 5 alcohol specimens from the same lot plus 10 shells; 2 alcohol specimens from St. 76, Mt. Kaala, 770-850 m, 30.vi.1979.

Etymology This beautiful species is appropriately dedicated to Annie Tillier, in recognition of her great skills as a field collector, and her important contribution to the knowledge of the New Caledonian land mollusc fauna. NEW CALEDONIAN SNAIL GENUS 77

0 p. ~3 0 0

• . °-

• °

..Q .r'

I o o

J E

xO 78 S. TIT.T.IER AND P. B. MORDAN

.] /

f tr,

/ '2: ~=r

Fig. 22. Holotypes of A, B. D. ~lmecata sp. nov., station 284, Mt. Ouazangou; C & D, D. anniae sp. nov., station 253a, Mt. Kaala.

Shell (Fig. 22) Rounded conical, embryonic whorls colourless, subsequent 1-1.5 whorls brown, but remainder marked with alternating flammulate bands of white and brown, each brown band becoming slightly paler towards the apertural side. Flammulations not extending to the lip which is white. Body whorl distinctly but not sharply carinated. Aperture wider than high, but with distinct downward deflection.

Radula (Fig. 23) 25 teeth per half-row. B04y Distinct brown pigmentation on the body, which is stronger on the sides. PaUial system Cavity extended 1.2 whorls in an animal with a body total of 5.4 whorls. Kidney extending 7/8th of lung length.

Reproductive system (Figs 24A, 25A) Epiphallus 2.5 times the length of the penis and inserting near the top. Vagina over twice the length of the free oviduct. No spermatophore found. NEW CALEDONIAN SNAIL GENUS 79

Fig. 23. Radula ~l + D. am+me. A, central and lateral teeth, and B, marginal teeth, statism 253a. Nit. Kaala. Scah" bar= IOlml.

Dis//bulion (Fig. 21) D. anniae is restricted to Mt. Kaala, where it co-occurs with michaudi at both sites from which it has been recorded. The sites are at elevations between 770 and 850 m. As in the other northwestern uhrabasic massifs, the summit of Mt. Kaala forms a small island, slighly wetter than the surrounding emdronment, and characterized by a natural vegetation consisting of sparse Arauca,~a in low maquis, which is partly burnt periodically.

Commenls At Kaala the flalmnulate form anniae co-occurred with a brown form, considered to be michaudi; this had similar male reproductive characteristics to am~iae (relative epiphallar length, high insertion of the epiphallus), but the vag-ina was shorter than the fi'ee oviduct (about two-thirds the length), and thus closer to the configuration ill subnecata at Ouazangou (see below). This suggests that character displacement ill anniae at Kaala cannot explain the difference in the male systems between Ouazangou and Kaala, as any displacement would have been expected to operate in the opposite direction, i.e. to produce a lower 80 S. TILLIER AND P. B. MORDAN

B s

Fig. 24. Reproductive systems of A., D. anniae sp. nov., station 235a, Mr. Kaala; B, D. subnecata sp. nov., station 284a, Mt. Ouazangou. Scale bar = Imm. Abbreviations as in Fig. 6.

epiphallar insertion, closer to the Ouazangou situation in subnecata. It also suggests that in sympatry, the female system is more likely to be the isolating mechanism, as this was where differences between the two species at Kaala were most evident.

Draparnaudia subnecata sp. nov. Type locality. St. 284a, Mt. Ouazangou, 850 m. leg. A & S Tillier, 7.x.1986.

Material examined Holotype: Fig. 22A, B. Station 284a. Paratypes: 1 mature plus 3 immature alcohol specimens and 45 shells from the same lot. NEW CALEDONIAN SNAIL GENUS 81

Fig. 25. Penis of A, D. anniae sp. nov., station 235a, Mt. Kaala; B, D. subnecata sp. nov., station 284a, Mt. Ouazangou. Scale bar = lmm. Abbreviations as in Fig. 8.

E~ymolog~ D. subnecata derives its name from its endangered status, resulting from the high likelihood of the loss of its very restricted habitat as a result of mining activity.

Shell (Fig. 22A, B) Ovate conic. Pattern and colouring as for anniae. Body whorl lacking carination, sutures relatively weak. Aperture slightly wider than high, with distinct downward deflection. The shell does not have adherent debris.

Radu (F g. 26) 23 teeth per half-row.

Brown pigmentation on the body, which appears markedly stronger on the sides.

Pallial system Cavity extending 1.1 whorls in an animal of 5.1 whorls. Kidney extending 9/10th of lung length.

Reproductive system (Figs 24B, 25B) As for michaudi. EpiphaUar insertion was near the central point of a very much shorter penis than anniae. Epiphallar length about 2.5 times the penis length. Vagina one-third the length of the free oviduct. No spermatophore found. 82 S. TILLIER AND P. B. MORDAN

. r

i..~., d I "X : . , _

• . ' A '

Fig. 26. Radula of D. subnecata sp. nov. A, central and lateral teeth, and B, marginal teeth. station 284, Nit. Ouazangou. Scale bar= 101ma.

Distribution (Fig. 21) Only known from the type locality, the appropriate habitat consisting of an island of low maquis wdth few Araucarias, which by 1986 had been reduced in area to no more than a few hectares.

Commenls Like D. anniae, D. subnecata is restricted in distribution to a single summital island which is wetter than its surroundings, and separated from similar environments by several kilometres of maquis. There is no doubt that it is totally isolated from other species. Despite superficial resemblence it differs significantly from anniae in both its reproductive system and shell characters. The penis and epiphallus are much shorter than in anniae, even though their relative lengths are similar; epiphallar insertion is much lower down on the penis, as is the insertion of the spermatheca on the free oviduct. The shell is more rounded in profile than anniae, closer to the michaudi shape, as is that of the aperture. NEW CALEDONIAN SNAIL GENUS 83

,._(h ;() f>, C,,, r. /~. ' t f t~ ,,.'>-;\-k.f !4 ;'-~L" ~,/~,~. ",

,,, ,.(:A t~.._.kk.f\_ ~-, f ~ -. ,.

Fig. 27. Radula of D. walked. A, central and lateral teeth, station 262, Wedoumel, Lifou, and B, marginal teeth, station 326, Presq'ile Montagn~s. Scale ba,'= 10pm.

Drapamaudia walken" Sykes, 1903 Draparnaudia walkeri Sykes, 1903: 197, fig. II [Renee River, Espiritu Santo]; Solem, 1959: 123, P1. 8, Fig. 8; Solem, 1962: 220.

Material examined Type Syntype of D. walke~ Sykes, herein selected as lectotype, BMNH 1902.12.1.159, Renee River, Espiritu Santo, LegJJ Walker (Fig. 17A, B). The single type was collected by Mr Walker during the cruise of HMS Ringarooma in the New Hebrides between June and October, 1900. Sykes comments on the 'very careful localization of the species' in Walker's collection. Other Stations 6, 13, 19, 25, 31, 53, 166", 262", 326*.

Sh~Zl (Fig. SZA, D) Uniformly brown in colour. Shape conical, ~th only moderately to weakly carinated body whorl, aperture rounded, but somewhat wider than high. Sutures deep, but this dependent on the degree of carination of the whorls. Frequently with adherent debris on shell. 84 S. TILLIER AND P. B. MORDAN

Fig. 28. Reproductive system of D. walketi, station 262, 4Km SE W6doumel, Lifou. Scale bar = lmm. Abbreviations as in Fig. 6.

Radula (Fig. 27) Radula between 22 and 24 teeth per half-row. B0 y Uniformly pale, unpigmented sole; top and sides of foot pale grey.

PaUial system As for michaudi; lung extending 0.8 whorls.

Reproductive system (Fig. 28) Epiphallus 4 to 5 times the length of the penis, and inserting on it at, or just above, the mid-point. Length of vagina equal to or shorter than the free oviduct.

Distribution (F~. 29) In New Caledonia, D. walkeri has been recorded from a small number of sites scattered throughout the mainland, as well as two on Lifou and one on the Isle of Pines. It is essentially a species of relatively dry lowland forest, with NEW CALEDONIAN SNAIL GENUS 85

8~ o ~o °° .~ ,~

c =.~ Oo Z ,e,e • •.~.== ~ ~--,~ ,~ ~.~~o:~-,0 ~ ~:~.--~

~ ~ |l I i I/ ~ ~

._~ ~-~N z . r~oz ",O [

• .."

o r" ~oL

O "B. "~ .< J 86 S. TILLIER AND P. B. MORDAN rainfall not exceeding about 200 mm per year. At Koumac (st. 6), Pombei (st. 13) and Presqu'ile Montagn6s (st. 326) on the mainland, and on Lifou (W6doumel, st. 262; Nanawa, st. 166) it occurred sympatrically with michaudi. At Adio caves (st. 25) on the mainland, and Ouro (st. 53), Isle of Pines, it was found together with singular#.

Comments D. walkeri resembles a small singular#. However, as well as small size (which is never as small as gassiesi, but approximates the smallest shells of singular#) there are certain conchological characters which distinguish it: the carination of the body whorl is never as sharply marked as in singular#, and is more prominent than in michaudi; moreover, the aperture is never as widely inflated as singular#, nor as downwardly deflected as michaudi, giving it a distinctly rounded appearance compared with both these species.

ACKNOWLEDGEMENTS This work is part of the projects 'Evolution et Vicariance en Nouvelle- Cal~donie' and 'Biodiversit~ terrestre en Nouvelle-Cal+donie', supported since 1984 by the Museum National d'Histoire Naturelle and since 1991 by the- Direction de la Recherche et des Etudes Doctorales, Minist6re de l'Education Nationale. In 1986-1987 collections were supported entirely by the project 'Caract~risation faunistique et floristique des forfits et maquis non anthropis~s' of the UR MAA-3H, ORSTOM. The authors, and especially S.T., wish to express their gratitude to their colleagues of the ORSTOM centre in Noum~a, and to thank Jean Chazeau in particular for his efforts to ensure successful stays in New Caledonia and for his assistance in the field. Joel Danloux (UR DEC-2A) kindly provided rainfall data for the collecting stations. Philippe Bouchet and Annie Tillier participated in most of the field work, and it is mainly due to their efforts that both the quality and quantity of these collections are so high. The distribution maps (Figures 9, 16, 21 and 29) were prepared by Philippe Maestrati (MNHN), and Harry Taylor (BMNH) took the photographs in Figures 3, 10, 17 and 22.

REFERENCES

Ancey CF. 1882. Classification des formes hSlicoides de la Nouvelle-CalSdonie. Le Naturaliste 2: 85-87. Ancey CF. 1898. Notes sur quelques coupes g6n6riques ou sous-g~nSriques de mollusques. Bulletin Music Histoire Naturelle Marseille 1: 147-148. Bayne CJ. 1973. Physiology of the pulmonate reproductive tract: location of spermatozoa in isolated selfo fertilizing succineid snails (with a discussion of pulmonate tract terminology). Vel~er 16: 169-175. Cooke CM, Kondo Y. 1960. Revision of Tornatellinidae and Achatinelfidae (Gastropoda, Pulmonata). Bernice P. Bishop Museum Bulletin 221" 1-303. Cowle RH. 1992. Evolution and extinction of Partulidae, endemic Pacific island snails. Philosophical Transactions of the Royal Society of London, B 335: 167-191. Crosse H. 1891. Faune malacologique terrestre et fluviatile de la Nouvelle Cal~donie et ses d~pendances. Journal de Conchyliologie 42: 161-473. DautTenberg P. 1923. Mollusques terrestres de la Nouvelle-Cal~donie et des lies Loyalty. In: Sarasin F, Roux J, eds. Nova Caledonica, A. Zoologic 3: 133-156. Deshayes G-P. 1810. Histoire naturelle des pulmon~s sans opercule. Genre Bulime. In: F~russac D, Deshayes G-P. 1820-1851. Histoire naturelle gdnerale et particuliare des mollusques terrestres et fluviatiles. Paris: Bailliere, 127pp. Duncan 1[~[. 1937. On the dates of publication of the Society's Proceedings, 1859-1926. With an appendix NEW CALEDONIAN SNAIL GENUS 87 containing the dates of publication of Proceedings, 1830-1858, compiled by the late FH Waterhouse and of the Transactions, 1833-1869, by the late Henry Peavot, originally published in P.Z.S. 1893, 1913. Proceedings of the Zoological Society of London 107: 71-84. Forcart L. 1940. Monographie der turkischen Enidae (Moll., Pulm.). Verhandlungen der Naturforschenden Gesellschaft in Basel 51: 106-263. Franc A. 1956. Mollusques terrestres et fluviatiles de I'Archipel N~o-Calrdonien. Mhnoires de la Museum National d'histoire Naturelle, Paris, n.s., A, ,~oology 13: 1-200. Gassies JIB. 1863. Faune conchyliologique terrestre et fluviolacustre de la Nouvelle Cal~donie, lere partie. Actes de la Sociit~ Linnienne de Bordeaux 24: 211-330. Gassies JB. 1870. Diagnoses d'esp+ces inrdites provenant de la Nouvelle-Calrdonie. Journal de Conchyliologie 18: 140-150. Gassies JB. 1871. Faune conchyliologique terrestre et fluviolacustre de la Nouvelle Calrdonie. 2eme partie. Acres de la Soci~t~ Linn~enne de Bordeaux 28:I-212. Gassies JB. 1873. Description de mollusques terrestres provenant de la Nouvelle Cal~donie. Journal de Conchyliologie 21: 46-54. Kemperman TCM. 1992. Systematics and evolutionary history of the Albinaria species from the Ionian Islands of Kephallinia and Ithaka. Doctoral Thesis, University of Leiden. 251pp. Kondo Y. 1973. Samoana of the Society Islands (Pulmonata: Partulidae). Malacological Review 6: 19-33. Mallick DIJ. 1975. Development of the New Hebrides Archipelago. Philosophical Transactions of the Royal Society of London, Series B 272: 277-285. Montrouzier RP. 1859. Description d'esprces nouvelles. Journal de Conchyliologie 7: 286-289. Mordan PB. 1984. Taxonomy and biogeography of southern Arabian Enidae sensu lato (Pulmonata; Pupillacea). In: Solem A, van Bruggen AC, eds. World-wide snails: biogeographicalstudies on non-marine MoUusca. Leiden, EJ Brill. Mordan PB. 1986. A taxonomic revision of the southern Arabian Enidae sensu lato (Mollusca; Pulmonata). Bulletin of the British l~laseum (Natural History), Zoology 50: 207-271. Mordan PB. 1992. The morphology and phylogeny of the Cerastinae (Pulmonata: Pupilloidea). Bulletiu of the British Museum (Natural Histo!y), Zoology 58: 1-20. Mordan PB, Tillier S. 1986. New Caledonian charopid landsnails. 1. Revision of the genus Pararhytida (Gastropoda; Charopidae). Malacologia 27: 203-241. Moss W, Webb WM. 1897. On the anatomy of Bulimas sinistrorsas, Deshayes. "/7reoTournal of Malacology 6: 1-2. Pfeiffer L. 1853-76. Monographia heliciornm viventum. Vol. III, 1853, 711pp.; Vol. IV, 1859; Vol. V, 1868, 565pp.; Vol. VI, 1868, 598; Vol. VII, 1876, 674pp. Lipsiae: FA Brockhaus. Pfeiffer L. 1855. Descriptions of fifty-seven new species of Helicacea from Mr. Cuming's collection. Proceedings of the Zoological Sodety of London, 1854: 286-298. Pilsbry HA. 1899. On the anatomy of Bulimns sinistrorsns Desh. Nautilus, 12: 95. Pilsbry HA. 1901-2. Manual of Conchology, Second Series. Volume XIV. Oriental bulimoid Helicidae; Odontostominae; Cerionidae. 302pp. Pilsbry HA. 1902. Manual of Conchology, Second Series. Classification of Bulimulidae and index to volumes X, XI, XII and XIV. 99pp. Pilsbry HA. 1909-10. Manual of Conchology, Second Series. Volume XX. Caecilioides, Glessula and Partulidae. 336pp. Reeve LA. 1854. Conchologica Iconica. VII. Monograph of the genus HelLr. London: Lovell Reeve. Solem A. 1959. Systematics and zoogeography of the land and fresh-water Mollusca of the New Hebrides. FMdiana: Zoology 43: 1-359. Solem A. 1960. New caledonian snails collected by TDA Cockerell in 1928. Notulae Naturae 338: 1-9. Solem A. 1961. New caledonian land and fresh-water snails. An annotated check list. Fieldiana: Zoology 41: 413-501. Solem A. 1962. Notes on, and descriptions of New Hebridean land snails. Bulletin of the British Museum (Natural Histoo,), Zoology 9: 215-247. Solem A. 1973. Convergence in pulmonate radulae. Vel~ger 15: 165-171. Solem A. 1988. Non-camaenid land snails of the Kimberley and Northern Territory, Australia. 1 Systematics, affinities and ranges. Invertebrate Taxonomy 2: 455-604. Solem 1992. Are the camaenids helicoids and a monophyletic group? Proceedings of the tenth International Malacological Congress Tiibingen 2:617-620. Solem A, Tillier S, Mordan PB. 1984. Pseudo-operculate land snails from New Caledonia. Vel~ger 27: 193-199. Sykes ER. 1903. On a collection of non-marine shells, formed by Mr jJ Walker, in the New Hebrides. Proceedings of the Malacolngical Society of London 5: 196-200. Tillier S. 1981. Clines, convergence and character displacement in New Caledonian diplommatinids (land prosobranchs), iVlalacologia 21:177-208. Tillier S. 1989. Comparative morphology, phylogeny and classification of land snails and slugs (Gastropoda: Pulmonata: Stylommatophora). Malacologia 30: 1-303. Tillier S, Mordan PB. 1989. Gast~ropodes terrestres Charopidae de Nouvelle Calrdonie: Revision du genre Pararhytida. Rossiniana 44: 3-7. 88 S. TILLIER AND P. B. MORDAN Zilch A. 1959-60. Gastropoda, Euthyneura. In: Schindenwolf OH, ed. Handbuch der Palaozoologie 6. Berlin: Borntraeger, 1-834.

APPENDIX

List of sampling stations. Station numbers belong to the same series as used by Tillier (1981) and Mordan & Tillier (1986). Locality, UTM grid reference, altitude, habitat, collector(s), date, annual precipitation and species present are given where possible for each site; the abbreviation s is for shell, a for alcohol-preserved specimen, and number of individuals precedes each category for each species. Material from the Field Museum of Natural History is listed at the end of the appendix.

3. Ni~nane, Iles Daos du Nord, 163°25'34"E 19°49"28"S, 2 m, dry littoral forest, P Bouchet & C Ch~rel, 23.viii. 1978, 1190 mm, 3s 4a michaudi. 4. Mt. Ti6baghi, 164°11'59"E 20°26'54"S, 520 m, dry forest, ultrabasic rock, A & S Tillier, 30.vii. 1979, 1200 mm, 2s michaudi. 5. Le Cresson, 164°18'36"E 20°29'00"S, 100 m, dry, calcareous forest, A & S Tillier, 30.vi.1979, 1300 mm, l ls 2a michaudi. Ditto Mordan, A & S Tillier, 29.i. 1981, 2s 2a michaudi. 6. Koumac caves, 164°20'27"E 20'~31'52"S, 80 m, dry, calcareous forest, A & S Tilfier, 30.vi. 1979, 1350 mm, 2s michaudi; 3s walkeri. 7. Mt. Mandj6fia, 5 km N saw mill, 164°30'06"E 20°22'29"S, 400 m, rainforest, A & S Tillier, 2.vii. 1979, 1900 mm, 2s michaudi. 11. Lind~ralique, 164°58'22"E 20°41'34"S, 20 m, humus in cavities, calcareous rocks, P Bouchet, 25.xi. 1978, 2150 mm, ls michaudi. 13. Pombei, 165°10'13"E 20°53'21"S, 100 m, rainforest, P Bouchet & S Tillier, vii. 1979, 2100 mm, 1~ wa/ken'. 13a. Pombei, 165°10'13"E 20°53'21"S, 80-120 m, rainforest, P Mordan, A & S Tillier, 18.i.1981, 2100 ram, 2s michaudi. 16. Col de Tango, 165°00'27"E 20°57'13"S, 300-350 m, rainforest, P Bouchet, 24.xii. 1978, 1450 mm, Is 2a juv s/ngu/at/s;, Is michaudi. 16a. Bobeitio, 165°01'01"E 20°57'13"S, 350 m, rainforest, A & S Tillier, 17.xi. 1988, 1450 mm, 3s michaudi. 19. Forfit Plate, 165°06'23"E 21°08'57"S, 450 m, rainforest, P Bouchet & A & S Tillier, 15.vii.1979, 1700 m, 3s walken'. 20. Mt. Pa~oua, 165°06'27"E 21°10'48"S, 950-1000 m, altitude rainforest, A & S TiUier, 5.vii.1979, 2000 mm, ls michaudi. 22. Nindiah, 165°35'21"E 21°18'53"S, 50 m, secondary vegetation, calcareous, P Bouchet, 30.xii. 1978, 1900 mm, Is s/ngu/ar/s. 25. Adio Caves, Vail~e s~che, 165°14'46"E 21°14'44"S, 180 m, dry calcareous forest, P Bouchet, 6.v. 1979, 1400 ram, 4s singularis; 15s walk~; 5s juv. 26. Bogui-Col des Roussettes, 165°26'44"E 21°24'53"S, 150 m, rainforest, P Bouchet, 15.v. 1978, 1600 ram, 3s singu/ar/s. 26b. Bogui-Col des Roussettes, 165°26'49"E 21°24'55"S, 270 m, rainforest, P Mordan, A & S Tillier, 4.v. 1987, 1600 mm, Is 3a ~ngu/ar/s. 31. Oua-Ou~, 165°35'07"E 21°37'00"S, 50 m, dry calcareous forest, P Bouchet, 31.xii. 1978, 1275 mm, 3s td3t7/ko'i. 37. Mt. Dzumac, 166°27'33"E 22°02'25"S, 950-1000 m, rainforest, P Bouchet & S Tillier, 4.vi.1979, 2200 mm, 2s singularis. 37a. Mt. Dzumac, 166°27'33"E 22°02'25"S 910 m, rainforest, A & S Tillier, 29.v. 1987, 2200 mm, la singularis. 39. Nd~, 166°18'00"E 22°08'37"S, 60 m, secondary forest, P Bouchet, 2. vii, 1978, 950 mm, 3s n'ngu/an~. 43a. Rivi~re Bleue, 166°39'25"E 22°05'47"S, 160 m, rainforest, ultrabasic rock, P Mordan & A & S Tillier, 29.ix. 1981, 3700 mm, 3a n'ngu/ar/s. 47. Mt. Guemba, 166°56'10"E 22°10'22"S, 450 m, rainforest on peridotite, P Bouchet, 16.ii. 1979, 4200 mm, 2a singu/ar/s. 48. Touaourou, route to Ni Mine, 166°58'00"E 22°12'00"S, 10 m, humid forest, raised coral, P Bouchet, 8.xii. 1978, 2500 mm, 6s singu/ar/s. 50. Goro, 167°00'25"E 22°19'27"S, 5-20 m, secondary forest, ultrabasic rock, A & S Tillier, 25.vii. 1986, 2700 ram, ls singularis. 52a. Kwa N~i~, Mt. Oungon~, 166°55'12"E 22°19'09"S, 470 m, very wet maquis, A & S Tillier & J Chazeau, 25.x. 1988, 3000 m, 2a ~ngubar/s. 53. Ouro, Ile des Pins, 167°26'17"E 22°39'29"S, 15 m, dry forest, raised coral, P Bouchet, 21.x.1978, 1800 mm, 4s 3a singu/ar/s; 2s walken'. 56. N+c~, Mar~, Loyalty Islands, 167°41'00"E 21°30"45"S, 15-20 m, dry forest, raised coral, P Bouchet, 5.iv.1979, 1500 mm, 6a michaudi. NEW CALEDONIAN SNAIL GENUS 89 69. Mt. Mandj61ia, 164°30'05"E 20°22'29"S, 500 m, rainforest, A & S Tillier, 2.vii. 1979, 1950 mm, la michaudi. 73. Mt. Ignambi, 164°36'00"E 20°27'37"S, 1200-1300 m, altitude rainforest, Bouchet, 27.xii. 1987, 7000 mm, 9a michaudi. 76. Mt. Kaala, 164°23'36"E 20°37'25"S, 770-850 m, high maquis with araucarias, ultrabasic rock, A & S Tillier, 30.vi.1979, 3a michaudi; 2a anniae. 84. Mt. Tchingou, S slope, 165°00'00"E 20°54'27"S, 1250 m, altitude rainforest, P Bouchet, A & S Tillier, 13.vii. 1979, 3100 ram, 2s gass~s/. 87. Mt. Koniambo, 164°48'46"E 21°01'31"S, 660 m, high maquis on ultrabasic rock, P Mordan, A & S Tillier, 28.i. 1981, 1400 mm, 12a singularis. 90. Mou, east coast, 165°25'00"E 21°06'00"S, forest, P Bouchet, 15.v. 1978, 2400 mm, Is michaudi. 9lb. Mt. Aoupini~, 165°17'59"E 21°10'47"S 600-700 m, rainforest, P Mordan, A & S Tillier, 5.v.1987, 2700 mm, la singularis. 93. Mt. Aoupini6, 165°16'23"E 21°11'00"S, 850-950 m, rainforest, P Bouchet, 10.ii. 1979, 2500 mm, la singularis. 97. Between Petit and Grand Boulinda, 165°08'57"E 21°14'44"S, 980-1020 m, altitude rainforest, A & S Tillier, 6.vii.1979, 2000 mm, 3s michaudi. 98. Adio Caves, 165°14'18"E 21°15'36"S, 180 m, dry calcareous forest, P Bouchet & C Ch~rel, 20.viii. 1978, 1400 mm, ls s/ngu/ar/s. 101a. Mt. M~ Maoya, 165°20'21"E 21°22'09"S, 1250 m, rainforest on uhrahasic rock, P Bouchet & S Tilfier, 11.vi. 1979, 1900 mm, la singu/ar/s. 101c. Mt. M~ Maoya, 165°20"21"E 21°22'09"S, rainforest, ultrabasic rock, P Bouchet & S Tillier, 12.vi. 1979, 2200 mm, la singularis. 10lb. Ditto, 13.vi. 1979, 2200 mm, 3a ~ngu/ar/s. 102. Bogui, 165°26'30"E 21°23'21"S, 150 m, rainforest, some secondary, P Bouchet, 15.v. 1978, 1600 mm, ls singularis; Is michaudi. 107. Nessa-Kouya, 165°31'09"E, 21°16'18"S, 250 m, old coffee plantation, P Mordan, A & S Tillier, 14.i.1981, 2250 mm, Is ~ngu/ar/s. 108. Montagnes Blanches, 165°45'55"E 21°32'17"S, 200 m, dry forest, P Bouchet, S Tillier, A War, n, 10.vi.1979, 1000 mm, ls 2a michaudi. 112. Between Mt. M~ Ori and Mt. Table Unio, 165°41"34"E 21°34'09"S, 850-1000 m, P Bouchet, 7.v. 1979, 2000 ram, is singular. 127. Mt. Ouin, 166°27'26"E 22°00'39"S, 1020-1040 m, rainforest, P Mordan, A & S Tillier, 4.ii.1981, 5000 mm, la la singularis. 131. Mt. Mou, 166°19'45"E 22°04'28"S, 270-420 m, rainforest, P Mordan, A & S Tillier, 23.i. 1981, 1600 mm, 5a juv. nngu/ar/s. 137a. Montagne des Sources, 166°36'10"E 22°08'11"S, 850 m, E slope, high maquis with araucarias, P Bouchet, 29.vii.1979, 1800 mm Is s/ngu/ar/s. 140a. Mts Koghi, 166°30'21"E 22°10'35"S, 900 m, rainforest on uhrabasic rock, P Bouchet, 20.viii. 1985, 1800 mm, la ~ngu/ar/s. 140c. Ditto, 720 m, P Bouchet, 27.i. 1979, 1800 mm, la singu/ar/s. 140d. Ditto, 450-520 m, P Mordan, A & S Tillier, 10.i.1981, 1800 mm, la singularis. 140e. Ditto, 450 m, A & S Tillier, 22.v. 1987, 1800 mm, 50a ~ngu/ar/s. 142. Col de Mouirange, 166°39'00"E 22°12'00"S, 180-250 m, rainforest, P Mordan, A & S Tillier, 11.i.1981, 2500 mm, Is la ~ngu/ads 143. Col de Mouirange, 166°40'14"E 22°13'19"S, 200 m, rainforest, S Tillier, 5.vi. 1979, 2200 mm, ls

156. R6, Mard, Loyalty Islands, 167°51'52"E 21°26'20"S, dry forest on raised coral, P Bouchet, 6.iv. 1979, 2a (Is 5a) michaudi. 157. Wako.nd, Mard, Loyalty Islands, 167°06'44"E 21°31'51"S, rainforest, raised coral, P Bouchet, 7.iv. 1979, la 5s michaudi. 161. Pdd~, Mard, Loyalty Islands, 167°57'26"E 21°25'32"S, dry forest, P Bouchet, 7.vii. 1979, 3a michaudi. 165. Natchaom, Lifou, Loyalty Islands, 167°11'37"E 20°42'35"S, dry calcareous forest, A & S Tillier, 20.vii. 1979, la michaudi. 166. Nanawa, Lifou, Loyalty Islands, 167°10'27"E 20°42'39"S, forest on raised coral, A & S Tillier, 20.vii. 1979, 2a michaudi; la walkeri. 167. Doudoulou, Lifou, Loyalty Islands, 167°06'51"E 20°55'39"S, P Lespes, 2a 24s michaudi. 168. Near Hap~tra, Lifou, Loyalty Islands, 167°09=8'47=E, 20°56'18"S, forest on raised coral, P Bouchet, A & S Tillier, 21.vii. 1979, 2a michaudi. 169. Loundou, Lifou, Loyalty Islands, 167°19'31"E 20°56'18"S, forest of endemic palms, P Bouchet, A & S Tillier & A War,n, 17.vii. 1979, (8a 25s) michaudi. 172. Luengoni, Lifou, Loyalty Islands, 167°23'15"E 21°03'36"S, forest, P Bouchet, A & S Tillier, 19.vii. 1979, 3a michaudi. 173. Khddeigne, Lifou, Loyalty Islands, 167°13'29"E 21°04'41"S, araucaria maquis, raised coral, P Bouchet, W Ponder, A & S Tillier, A Wardn, 21.vii. 1979, 30a michaudi. 90 S. TILLIER AND P. B. MORDAN 174. Mou, Lifou, Loyalty Islands, 167°23'22"E 21°05'40"S, rain forest, raised coral, A & S Tillier, 19.vii. 1979, 2a michaudi. 175. Ni6mak, Ouv~a, Loyalty Islands, 166°34'32"E 20°25'29"S, dr), calcareous forest, P Bouchet, 27.vii. 1978, Is midmudi. 176. Gossana-Ognat, Ouv6a, Loyalty Islands, 166°38'15"E 20°27'26"S, rainforest on raised coral, P Bouchet, 6.v. 1978, Is michaudi. 177. Ti6ou, Ouv+a, Loyalty Islands, 166°37'47"E 20°27"59"S, P Bouchet, 6.v. 1978, 7s rnichaudi. 178. Fayaou6, Ouv~a, Loyalty Islands, 166°33'22"E 20°38'50"S, P Bouchet, 6.v. 1978, 2s michaudi. 181. Mr. Ningua, 166°09"25"E 21°45'17"S, 950-1050 m, rainforest on ultrabasic rock, S Tillier, vi. 1979, 4500 mm, l a singular& 201. Cap Bocage, 165°33'38"E 21°09'33"S, 0-50 m, high maquis, ultrabasic rock, P Bouchet, S Tillier & MP Triclot, 20.xi.1984, 5s la mi chaudi. 202. Cap Bocage, 165°35'38"E 22°11'24"S, 320 m, rainforest in sink-hole, peridotite, P Bouchet, S TiUier & MP Trielot, 2hxi. 1984, 2600 mm, 4a midmudi. 206. Mr. M6nazi, E of the summit, 165°44'57"E 21°26'37"S, 850 m, rainforest, araucarias, uhrabasic rock, S Tillier, P Bouchet & MP Triclot, 18.x. 1984, 3500 mm, Is la gassiesi. 210. Barrage of the N6aoua, 165°32'35"E 21°21'57"S, 500 m, P Bouchet, S Tillier & MP Triclot, 20.xi.1984, la sir~daris. 212. Col de Nassirah, 166°04'14"E 21°46'44"S, 250 m, dry, calcareous forest, S Tillier, P Bouchet & MP Triclot, 8.xi.1984, t400 mm, 4s singularis. 216. Valley of the Nimbo, 166"22'49"E 21°43'02"S, 20 m, rainforest, P Bouchet, S Tillier & MP Triclot, 3.xi. 1984, 2400 mm, la juv. sir~ularis. 217. Ouinan6, 166°09'24"E 21°55'00"S, 100 m, secondary dry tbrest, S Tillier, P Bouchet & MP Triclot, 31 .x. 1984, 1150 mm, 14a shtgulam'. 223. Mts. Kouv616, 166°26'56"E 22°05"26"S, 660 m, rainforest, P Bouchet, S Tillier & MP Triclot, 25.xi.1984, 1600 mm, 18a singularis. 224. Hill SW of Tonghou6, 166°27'24"E 22°11'50"S, secondary dr5, forest, S Tilfier, P Bouchet & MP * Tficlot, I.xi.1984, 1150 mm, 3a gassesi. 228. Mt. Kouakou6, 166°31'38"E 22°01'18"S, 1000 1100 m, altitude rainforest, P Bouchet, S Tillier & MP Triclot, 28.x. 1984, 3500, I s J41~daris. 234. North of Grand Lac, 166°55'03"E 22°13'09"S, 310-320 m, rainforest on ultrabasic rock, S Tillier, P Bouchet & MP Triclot, 26.xi. 1984, 3400 ram, 4a singularis. 235. South of Grand Lac, 166°54'00"E 22°16'31"S, 280 m, rainforest on uhrabasic rock, P Bouchet, A & S Tillier, 26.xi.1984, 3100 mm, la singldaris. 238. Peak north-east of Mr. Tonta, 166°17'06"E 21°24'4["S, 900 m, rainforest, P Bouchet, A & S Tillier, 13.xii. 1984, 1600 mm, Is 8a ~ngularis. 241. Vall6e de la Ouinn6, flanc S, 166°32"28"E 22°02'23"S, 650 m, humid forest, with kaoris, P Bouchet, S Tillier, MP Triclot, 16.xi.1984, 4000 ram, 2s singularis. 244. Piste de la Pourina, 166°41'45"E 22°35'00"S, 520 m, rainforest, P Bouchet & MP Triclot, 11.xi.1984, 4500 ram, Is, 4a sit~ularis. 246. Mrs Kouv61~, 166°37'53"E, 22°04'01"S, 750-800 m, maquis paraforestier, P Bouchet, 24.viii. 1985, 1750 mm, 6a singularis. 247. Haute Pourina, 166°37'53"E 22°04'01"S, 750-800 m, rainforest, P Bouchet, 17.x. 1985, 5500 mm, 2a singu laris. 248. Between Wapan and Oup6, Isle of Pines, 167°28'00"E 22°33'56"S, 15 m, P Bouchet, 22.x.1,985, 2s 12a sit~ularis. 250c. Rivi6re Bleue, parcelle VI S, 166°39'16"E 22°06'13"S, rainforest, alluvium, 160 m, A & S Tillier, 3.xi.1987, 3700 mm, Is singular& 250.f Rivi6re Bleue, parcelle VI R, 166"39'16"E 22°06'13"S, 160 m, rainforest on alluvium, A & S Tillier, 5.ii.1987, 3700 mm, Is singularis. 250i. Rivi6re Bleue, parcelle VI W, 166°39'16"E 22°06'13"S, 160 m, rainforest, alluvium, P Mordan, A & S Tillier, 30.iv. 1987, 3700 mm, ls singular& 251. Rivi6re Bleue, parcelle VII V, 166°40'01"E 22°05'59"S, 170 m, ralnforest, slope, ultrabasic rocks, A & S Tillier, 15.viii. 1986, 3700 mm, Is singularis. 2511". Rivi~re Bleue, parcelle VII Z, 166°40'01"E 22°05'59"S, 170 m, rainforest, slope, ultrabasic rocks, A & S Tillier, 12.ii.1987, 3700 mm, Is singularis. 251k. Rivi6re Bleue, parcelle VI X, 166°39'16"E 22°06'13"S, 160 m, ralnforest on alluvium, A Tillier, 7.vii. 1987, 3700 mm, la si1~ularis. 253. Mt. Kaala, 164°23"26"E 20°38'18"S, 810 m, low maquis, araucarias, ultrabasic rocks, J Chazeau, A & S Tillier, 27.viii. 1986, 1450 ram, Is michaudi. 253a. Mt. Kaala, S slope, 164°23'26"E 20°38'18"S, 810 m, araucarias, ultrabasic rock, A & S Tillier, 8.x. 1986, 1450, 3a michaudi; lls 7a anniae. 255. Mt Kaala, pente S, 164°23'23"E 20°38'48"S, 340 m, dry forest pente, ultrabasic rocks, J Ghazeau, A & S Tillier, 27.viii. 1986, 1300 mm, 8s michaudi. 256a. Parc /l Cagous, Rivi~re Blanche, 166°38'20"E 22°08'09"S, 160-170 m, rainforest on ultrabasic rock, Y Letocart, 1.iv.1987, 3300 ram, la singular& NEW CALEDONIAN SNAIL GENUS 91

258. Valley of the Tontouta, 166"18'27"E 21°55'59"S, 50 m, maquis on brown soil, ultrabasic rock, A & S Tillier, 16.ix. 1986, 1350 mm, 2s sil~ularis. 262. 4 Km SE W6doumel, Lifou, Loyalty Islands, 167°08'24"E 22°00'35"S, wet calcareous forest, A & S Tillier, 22.vii. 1979, 6a juv 12s rnidtaudi; 2a walked. 284a. Mr. Ouazangou, 164°29'30"E 20°44'43"S, 850 m, araucarias, ultrabasic rocks, A & S Tillier, 7.x. 1986, 1250 mm, 40s 4a subnecata. 285. Valley of the To Ndeu, 166°17'16"E 20042'19"S, 120 m, high maquis on ultrabasic rock, A & S Tillier, 29.x. 1986, 3100 mm, ls michaudi. 306. Mr. Kop6to, 165°00'38"E 21°10'39"S, 800 m, rainforest with araucarias, ultrabasic, A & S Tillier, 23.iii. 1987, 3s michaudi. 307. Mt. Koniambo, 164°49'05"E 20°59"51"S, 870 m, maquis with low araucarias, ultrabasie rocks, A & S Tillier, 28.i.1981, 1750 ram, 34s, 21a michaudi. 310. Mt. Taom, 164°34'47"E 20°47'10"S, 1000 m, rainforest, araucarias, ultrabasic rocks, A & S Tillier, 24.iii. 1987, 1700 mm, Is gassiesi. 312. Mt. Do, 165°59'33"E 21°45'37"S, 840 m, rainforest with araucarias, ultrabasic rock, A & S Tillier & C & F Monniot, 2.iv. 1987, 1450 mm, Is singular& 319. Dent de St Vincent, 166°13'02"E 21°52'12"S, 1150 m, rainforest and maquis, A & S Tillier, L Bonnet de Larbogne, Y Letocart, 6.viii. 1987, 2100 mm, 2a singularis. 326. Presque'ile Montagn+s, 166°07'23"E 22°02'21"S, 30 m, dry calcareous forest, J Chazeau, A & S Tillier, 9.xi. 1988, 700 mm, la michaudi; 20s 86a walken'. 330. Pindai, 164°58'10"E 21°20'16"S, 20 m, dry calcareous forest, A & S Tillier, 15..,d.1988, 1100 mm, 9s 26a michaudi.

FMNH MATERIAL: 144237. Route les Dalmates, Monts Kouanenoa, E of St Louis, 166°04'E 22°12'S, 24.i.1962, A & B Solem! la singularis. 144238. As above, la a'ngu/ar/s. 144265. Col de la Pirogue, 450 m, Mont Mou, 166°20'E 22°05'S, 23.i. 1962, A & B Solem, 3a singularis. 158315. N.C. 25 - hillside near pierced rock, l mile in and 5 miles W of Bourail, 200', 29.x. 1967, Laurie Price! la a'ngu/ar/s, body only. 159270. N.C. 6 - Mont Mou, 1000-1500' elevation near Paita, 12.xi. 1967, Laurie Price! la singularis. 159302. as above, 6.x. 1967, la singularis. 159322. N.C. 13 - Hiengh~ne, N.E. coast, 13.x. 1967, Laurie Price! la michaudi. 159299. as above, 13.x. 1967, 2s michaudi; one dissected A Solem. 159328. N.C. 8 - east side of Col des Rousettes, 1200' 8.x. 1967, Laurie Price! la singularis, la michaudi. 159350. N.C. 13 - 10 miles ex Poya, inland ex Nrkliai, midwest coast, 7.xi.1967, Laurie Price! la michaudi. Other site not given a NMHN number: Nassirah, valley of the Karaka, Bonnet de Larbogne, 23.ix.1986, ls michaudi.