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AN ILLUSTRATED GUIDE TO THE GENERA OF THE LAND MOLLUSCA OF NEW ZEALAND.

by W. F. Ponder and T. P. Warren.

INTRODUCTION

Where Land Mollusca Live. The majority of N. Z, land snails are restricted to the isolated remnants of native bush. The bush is called a macrohabitat but the places where the individual animals live are referred to as microhabitats. These can be made into a very lengthy list, but for our purposes the more important microhabitats only will be mentioned. The bush consists essentially of three layers: - trees, bushes and shrubs, and the ground cover. The former provides a home for relatively few snails, though by beating the foliage some species may be found, while others hide beneath loose bark and in the dead leaves nestling between branches. Shrubs and bushes provide a similar habitat to trees. Palms (the nikau) harbour snails in the large leaf bases. The ground cover is, however, the richest habitat, both in the variety of species encountered and in the diversity of microhabitats available. Small ferns, mosses and creeping plants provide shelter for some species while others prefer the dampness and protection found beneath rotting logs on the forest floor. Leaf litter is a rich habitat providing a subaquatic environment for many tiny snails and other animals.

Not only the native bush provides a habitat for snails. Coastal scrub and flax is often rich in species and a few are found in tussock grassland and others beneath sand dune plants.

Subfossil land snails are occasionally encountered in limestone caves or on sand dunes. Several subspecies of the large land snail Placostylus ('Maoristylus) ambagiosus Suter are known only as subfossils.

How to Collect and Store Land Mollusca. Only simple equipment is needed to become a student of land molluscs, though to study small forms adequately a microscope is needed. A pair of forceps, a small paint brush, stoppered tubes and labels are all that is required. A torch may come in handy as light is often very poor in the bush. To collect foliage - living species, a piece of white cloth spread out on a wire frame is all that is needed. This is held beneath a tree while the branches and foliage are shaken or beaten with a stick.

A bag of leaf mould sorted under magnification yields many of the small species that one would not normally see in the field. Other debris involving a lot of fine material should be treated in a similar manner.

Slugs should be drowned in fresh water to extend them and then preserved in 75% ethyl alcohol or in 5% neutralised formalin. Snails can be preserved by the same method, or after relaxing, the animal 22 -

can be removed from the shell. The large Paryphanta and Placostylus should not be left too long as air bubbles may form under the perlostracum of the shell. Paryphanta shells should be plugged with cotton wool to prevent their collapse when dried out. The smaller species can be preserved as above, or, if only the shell is required, soaked for a day or two in 75% alcohol and then dried. Some ribbed or bristled species may be covered in dirt. This can be easily removed by stirring them in a dish containing 1-1 mixture of alcohol and water, and then drying on clean blotting paper.

Large snails should be kept in trays with a label giving the species name, locality, date, data about the habitat, etc. Small shells are best kept in glass vials or short lengths of glass tubing plugged at both ends with cotton wool, not corks. Lables (as above) should be placed inside the tube.

Dissection of the animal is an important part of the study of land snails, but it is too large a topic to be discussed here. The morphology of the jaw; and radula can, however, be studied without much difficulty, and observations of the external features of the living animal present few problems. The radula and jaw can be either dissected out of the head, or the buccal mass can be heated in a 10% solution of caustic potash to dissolve away all non-chitinous material. The radula should then be mounted in a suitable medium (e. g., glycerine jelly or polyvinyl lactophenol) after staining (if necessary) with a suitable dye or chemical (e. g., picric acid). The jaw can usually be examined without mounting and staining, but if this is necessary a similar method can be used as for the radula.

History. The New Zealand terrestrial molluscs have attracted the attention of malacologists for over a hundred years. The early workers in the field, such as Gray, Pfeiffer, and Reeve, were men who lived in Europe and studied material collected by several pioneer explorers to New Zealand. Much valuable work was done locally in the latter half of last century by Hutton, Suter, Cheeseman, Webster and Murdoch. In more recent years the tradition has been carried on chiefly by Iredale, Powell, Dell, Cumber and Burton.

Part of the studies of Hutton, Suter, Webster and Murdoch concerned the radula, genitalia and other anatomical details, which has proved a valuable basis for later workers to continue from Much, however, remains to be done on the anatomy of the smaller species particularly and a rich field of research is available here.

At the present time there are 56 genera and subgenera recognised in New Zealand and these contain some 373 species and subspecies. Further revision and research will undoubtedly lead to increases in both of these figures. HOW TO USE THE GUIDE

In the illustrated guide to the native New Zealand terrestrial molluscan genera, each genus is briefly described under its appropriate family. Rather than use the more common types of key for this rather difficult group, a pictorial guide is considered by us to be more generally useful. A specimen should be readily placed in a genus by comparing it with the figures and checking with the appropriate description of the genus. Specific identification can then be attempt• ed by referring to the checklist in "Shells of New Zealand" by A. W. B. Powell. This reference will give sources for full descriptions of each species. The most useful reference in this respect is Suter's "Manual of the New Zealand Mollusca".

The shells are described and figured, the approximate maximum size given, The distribution, and number of species noted for each genus. The animal, radula, and jaw are described only when shell characters are considered inadequate for positive identification. In most cases the species illustrated is the type species for the genus, though other members are also figured in those instances where there is a wide range of variation within the genus. Few of the figures are original, the majority being taken from Suter (1915).

The following guide concerns the native snails and slugs but there are also a number of introduced snails and slugs that may be encountered, most of which have originated from Europe. A list of these species can be found in Suter's Manual (p. 1071).

Class GASTROPODA Order PROSOBRANCHIA Suborder ARCHAEOGASTROPODA Superfamily NERITACEAE Family HYDROCENIDAE

Animal with no gill, but a pulmonary chamber. Tentacles short, large, eyes prominent and at the outer bases of the tentacles. Foot oval, short. Radula rhipidoglossan - formula » + 1 + (1 + 1 + 1) + 1 + «. Shell imperforate, conic and globular. Whorls convex, spire short. Columella callous. Operculum calcareous, ornamented with concentric striae. Inner side with a prominent apophysis. - 24 -

Genus OMPHALORISSA Iredale, 1913 (Georissa multilirata Brazier). Shell minute, imperforate, translucent. In very moist and dark parts of the bush. North and South Islands. 2x1 mm. Fig. l, Omphalorissa purchasi (Pfeiffer); A, operculum.

Suborder MESOGASTROPODA Superfamily LITTORINACEA Family CYCLOPHORIDAE Animal with long, cylindrical tentacles, the eyes at their outer bases on short peduncles. Foot long, attentuate behind. No gill, but a pulmonary chamber. Jaw reticulate, radula taenioglossan (2+1+1+1+2). Shell conical or depressed, usually covered with a horny periostracum, aperture circular, peristome simple or reflexed. Operculum spiral, testaceous or horny with many to few whorls.

Genus CYTORA Kobelt and Moellendorff, 1897 (Cyclophora cytora Cray) (= Murdochia Ancey). Shell small, umbilicated, with 5-6 whorls. Protoconch 2 whorls, first smooth, second spirally striate. Moist parts of the bush among fallen leaves and fronds. 3. 25 x 5 mm. Three Kings, North and South Islands, Stewart Island. 18 species and subspecies. Fig. 2, Cytora cytora Gray, 2A, Cytora pannosa (Hutton), operculum

Family LIAREIDAE Animal with long cylindrical tentacles, eyes at their outer bases. Foot elongate, tapering behind. No jaws, radula taenioglossan with multicuspate teeth. Shell of moderate size, tall spired, smooth or vertically ribbed, body whorl often angled or keeled. Aperture oval, peristome continuous, often flanged. Operculum horny, spiral.

Genus LIAREA Pfeiffer, 1853 (Realia egea Gray). Shell small, subperforate, operculate, often with zig-zag markings. Protoconch of 2 smooth whorls. peristome continuous, with a labial flange. Mostly under decaying leaves of Bielschmedia taraire. Adult shell (7-9 whorls) approx. 3x9 mm. 12 species and subspecies. North Island, mainly North Auckland. Fig. 3, Liarea egea tesselata Powell, 3 A, Liarea turriculata (Pfeiffer), operculum. 25

Subclass PULMONATA Order STYLOMMATOPHORA Suborder HOLOGNATHA Superfamily ENDODONTACEA Family OTOCONCHIDAE

Animal elongate, much too large to withdraw into the shell. Mantle extended over shell. No posterior mucous gland. Jaw distinctly furrow• ed. Shell very flat, transparent, partly external, of few rapidly increasing whorls. Genus OTOCONCHA Hutton, 1884 (Vitrina dimidiata Pfeiffer). Shell very thin, pale yellow, with minutely striated growth lines, whorls 2£r. Protoconch of 1_- smooth whorls. Base of shell completely cut away. Animal sluglike, yellowish mottled brown with large anterior visceral hump containing the shell. Tail long. Animal 25 mm., shell 5 mm. Under decaying timber in the bush. North and South Islands. 1 species. Fig. 4, Otoconcha dimidiata (Pfeiffer); B, shell.

Genus MAORICONCHA Dell, 1952 Helicarion oconnori Powell). Shell thin, yellowish, with numerous spiral lirae crossed by radial growth lines. Whorls 2^ protoconch l£ smooth whorls, flatten• ed. Base of shell only partly cut away. Animal much larger than shell, tail long and narrow, similar to Otoconcha* Colour light brown with darker lateral stripes. 2 species. Animal 27 mm., shell 8mm. South Island only. Fig. 5, MAORICONCHA o'connori (Powell), A, B, shell.

Genus FLAMMOCONCHA Dell, 1952. (Helicarion (Peloparion) cumberi Powell). Shell hyaline, pale yellow, radial growth striae crossed by linear grooves. Whorls 2_\ Protoconch of l£ spirally striated whorls. Base of shell only partly cut away. Animal somewhat larger than the shell, a much larger portion is capable of being retracted than in MAORICONCHA* Tail quite short, rapidly tapering. Shell 4. 1 mm. 2 species. South and Stewart Islands. Fig. 6, Flammoconcha cumberi (Powell); A, B, shell. - 26 -

Genus RANFURLYA, Suter, 1903 (R. constanceae Suter). Shell ear-shaped, yellowish, very thin and transparent, all base of shell completely cut away. One whorl only, with sparse micro• scopic growth lines. Animal slug-like, black, mantle covering the shell. Tail, short, rounded. Animal 6x3 mm., shell 2. 75 x 1. 5 mm. 1 species. Auckland Islands only. Fig. 7, Ranfurlya constanceae Suter; A, shell.

Families FLAMMULINIDAE, CHAROPIDAE, LAOMIDAE and ZONITIDAE. These four families, together with the Otoconch-idae, are all closely allied and not sharply differentiated on shell characters. The little that is known about their anatomy also suggests similarity, their reproductive organs all being simple. The families are separated on jaw and radula characters. The shells can, with some experience, be placed with a high degree of accuracy in a given family. However, the conchological features that are important such as texture, translucency, lustre, etc. are difficult to describe. A brief diagnosis is set out below, but it is suggested that the genera in all four families be checked in cases where there is any uncertainty.

Super family ZONITACEA Family ZONITIDAE Shell depressed, aperture sharp, no apertural teeth. Animal usually with a posterior pedal gland, sole often tripartite. Radula with marginal teeth narrow, basal plates elongated, unicuspid (in Suterella) and jaw (smooth in Suterella ) membranous.

Genus SUTERELLA Iredale, 1915 (Helix novarae Pfeiffer). Shell flattish, narrowly umbilicated, thin and shining. Pale yellowish, sculptured with fine growth striae more pronounced at the suture. 6. 5 x 2. 2 mm. North Auckland. 1 species. Fig. 8, Suterella novarae (Pfeiffer), A, B, shell, C, jaw.

This shell has been included in Powell's Check list for many years but is unknown to local collectors. It is possible that an introduced mollusc was misidentified as a constituent of the N. Z. fauna, or else there has been confusion with one of the several similar species of small flammulinids or laomids. - 27

Superfamily ENDODONTACEA Family FLAMMULINIDAE Shell variable, smooth or ribbed, aperture without teeth. Jaw usually flatly plaited. Radula variable, cutting points long. Mostly living in less dense parts of the bush, and often on the trunks and leaves of trees.

This group as a whole is not easy to define on shell characters alone as the main distinctions of genera are in their radulae. However, the flammulinid shells are on the whole larger than those of the Charopidae and Laomidae and are mostly of smooth and pellucid appearance with the animal showing clearly through the shell.

Exceptions are Phenaochelixt Therasiella, Allodiscus and Suteria, which have pronounced axial ribbing sometimes produced into sutural bristles or plates.

Genus PHELUSSA Iredale, 1915 (Helix hypopolia Pfeiffer). Shell depressed, umbilicated, axially costate with microscopic spiral striae. Periphery rounded; mostly with zigzag or tesselated colour markings. Protoconch 1-J- smooth whorls. Dentition: central and lateral teeth tricuspid, marginals with a mesocone only. 6 species. 12 x 7 mm. North, South and Stewart Islands. Fig. 9, A, B, Phelussa hypopolia (Pfeiffer).

Genus ALLODISCUS Pilsbury, 1892 (Helix dimorpha Pfeiffer). Shell thin, depressed with low or flat spire. Periphery rounded, and having narrow, sub-imperforate or closed umbilicus. Closely axially ribbed, with microscopic spiral striae. Colour light brown, often with dark brown splashes or tesselations. Protoconch of l_f spirally striated whorls. Dentition: central tooth tricuspud, laterals bi- or tri• cuspid, marginals with 3-5 cutting points. 23 species and subspecies. 14. 5 x 10 mm. Throughout New Zealand and. the Subantarctic Islands. Fig. 10, Allodiscus spiritus Powell. - 28 -

Genus PHENACOHELIX Suter, 1892 (Flammulina (P. ) ponsonbyi Suter). Shell depressed, narrowly umbilicated, the spire somewhat convex. Periphery broadly rounded, whorls finely axially ribbed and with fine spiral striae. Colour yellowish- brown with darker spots or streaks. Protoconch of 1* distinctly spirally striated whorls. Dentition: central tooth narrow, with or without side cusps. Laterals bicuspid, no entocone. Marginals multicuspid, inner cusp large. 3 species. 6 x 3. 25 mm. North, South, Stewart and Campbell Islands. Fig. 11, A, B, Phenaochelix ponsonbyi Suter.

Subgenus (Neophenacohelix) Cumber, 1961 (Phenaochelix giveni Cumber). This subgenus differs from Phenaochelix in that the protoconch of Ijr whorls is smooth or at most occasionally very indistinctly spirally striated. Other shell and radula features are essentially the same as in Phenaochelix, 5 species. 7. 5 x 4 mm. North, South and Auckland Islands.

Genus Flammulina von Martens, 1873 (Helix phlogophora Pfeiffer). Shell narrowly umbilicated or imperforate, globose or depressed, thin and pellucid with few rapidly widening whorls which can be (i) smooth, glossy and greenish olive; or (ii) ribbed and light or dark brown. Aperture large and rounded; protoconch 1 to 1_- whorls, in group (i) smooth, and in (ii) spirally striate. Dentition: central and lateral teeth usually tricuspid; marginals with 3 or 4 cutting points, often coalescing on the outer teeth. 16 species and subspecies. 10 x 4 mm. North, South, Stewart and Subantarctic Islands. Fig. 12, A, Flammulina olivacea Suter; B (a), F, olivacea Suter, sculpture; B (b), P. chiron Gray, sculpture.

Genus SERPHO Hutton, 1904 (Nanina kivi Gray). Shell conical, greyish with purple colour markings. Subperforate or imperforate; axially and spirally striated, periphery angled. Protoconch 1_- convex whorls; spirally striated. Dentition: central and lateral teeth with a broad blunt mesodont. Marginals with several cutting points, the inner bidentate. 2 species. 10 x 8 mm. North Island, particularly in the North Auckland region. Fig. 13, Serpho kivi Gray. - 29 -

Genus THERMIA Hutton, 1904 (Pfeifferia cressida Hutton). Shell subglobose, thin, translucent. Axially striated and often with spiral striae; subperforate. Colour olive green, sometimes with darker base. Protoconch 12' whorls, smooth or striate. Dentition: central tooth with mesocone only; laterals bicuspid, no entocone; marginals with several cusps. 3 species. 13 x 9 mm. Southern half of North Island; South and Stewart Islands. Fig. 14, A, B, Themia virescens (Suter).

Genus GERONTIA Hutton, 1883 (G, pantherina Hutton). Shell depressed, openly umbilicated; thin and pellucid. Striated with close axial riblets and microscopic spirals. Colour olive- green to brown on spire whorls. Protoconch of 1J£ whorls, finely radially and microscopically spirally striate. Dentition: central tooth tricuspid, laterals similar but the ectocone larger; marginals with a broad bifid cusp. 1 species. 10 x 4. 5 mm. North Westland are« only. Fig. 15, A, B, G, pantherina Hutton.

Genus THALASS0EEL1X Pilsbry, 1892 (Helix zelandiae Gray). Shell depressed conical, rarely flattish. Striate or smooth, sometimes hairy. One species imperforate, remainder umbilicate. Protoconch lg- whorls, smooth or striate. Dentition: central tooth with obsolete ectocones; laterals with a short ectocone; marginals with a mesocone only. 9 species and subspecies. 15 x 9 mm. North, South, Stewart, Chatham and Auckland Islands. Fig. 16, A, Thalassohelix zelandiae (Gray); B, T, zigzac (Gould); C (a), T. zigzac (Gould) sculpture; C (b), T, zelandiae (Gray) sculpture.

Genus 1HERASIA Hutton, 1883 (T, thaisa Hutton). Shell moderately large (8x4 mm), depressed - conical, thin, umbilicate. Periphery angular or subangular. Surface axially or spirally striate. Pale brown with darker spots or splashes. Protoconch l_r spirally striate whorls. Dentition: central tooth narrow, tricuspid; inner lateral without side cusps, outer ones tricuspid. Marginals multicuspid. 5 species. North, South and Stewart Islands. Fig, 17, Therasia thaisa Hutton. - 30 -

Genus THERASIELLA Powell, 1948 (Nanina (?) celinde Gray). Shell smaller (4 x 2. 5 mm. ) than Therasia, sharp - spired, periphery acutely angled. Whorls with fine axial threads produced into membranous triangular plates at the periphery, and minute spiral threads. Colour uniformly olive brown. Protoconch 1-g- spirally striate whorls. 3 species. Three Kings, North and South Islands. Dentition as in Therasia, except that the marginals are bicuspid in Therasiella. Fig. 18, A, B, Therasiella celinde (Gray).

Genus OBANELLA Dell, 1952 (Egestula spectabilis Powell). Shell small, subdiscoidal, angled or subangled on the periphery. Whorls flattened above the periphery. Protoconch of l^r spirally striate whorls. Post nuclear sculpture of rather regularly spaced radial ribs produced into broad plates at the periphery, with fine axial riblets in between, the interstices sculptured with fine spiral striae. Umbilicus rather wide. Colour light or yellowish brown. 3 species. 2x1 mm. Southern part of North Island, South and Stewart Islands. Fig. 19, A, Obanella spectabilis (Powell).

Genus SUTERIA Pilsbry, 1892 (Helixide Cray). Shell discoid, openly umbilicated. Periphery rounded with low spirals and radial plates; points of intersection produced into bristles. Colour light brown with darker axial stripes. Protoconch 1_- radially striate, convex whorls. Dentition: central teeth and laterals tricuspid, the ectocone of the latter larger than the ectocone; inner marginals with one bifid cusp, outer ones with several subequal cusps. 1 species. 8x4 mm. North and South Islands mostly under fallen tree fern fronds. Fig. 20, A, Suteriaide (Gray).

Note: - Many of the Flammulinidae bear a superficial resemblance to the small paryphantids Delos and Delouagapia (q. v. ). However, species of the two latter groups are of more robust appearance, have a flatter spire and are more widely and deeply umbilicated. In addition, the outline of the aperture is nearly circular. The flammulinid aperture is, typically, semi-circular or crescent- shaped.

20 - 31

Family CHAROPIDAE Shell ribbed, variable in shape but usually umbilicated and depressed. Periphery usually rounded. Aperture with or without teeth. Foot without posterior pedal pore. Jaw thin, membranous, vertically striated. Radula with the central and lateral teeth tricuspid, marginals with 3 to 4 cusps.

Genus PTYCHODON Ancey, 1888 (Strobilia leiodus Hutton). Shell discoidal with numerous axial riblets, spiral striae present in most species. Umbilicated. Protoconch l-j whorls. Apertural plates present on outer lip and/or columella. 4. 25 x 2 mm. 23 species and subspecies. Throughout New Zealand in leaf mould. Fig. 21, Ptychodon leioda (Hutton); Fig. 22, P. monoplax Suter, Fig. 23, P. tau (Pfeiffer).

Genus PHENACHAROPA Pilsbry, 1893 (Pupa novoseelandica Pfeiffer). Shell with cylindrical spire, height about double width. Closely axially costate, reticulated with dense spiral striae. Protoconch 1^ smooth whorls, umbilicus narrow. 5x2 mm. 1 species. Southern half of North Island. Fig. 24. P. novo8eelandiaa (Pfeiffer),

Genus AESCHRODOMUS Pilsbry, 1892 (Helix stipulata Reeve). Shell conical, perforated, the height equal to the width. Axially costate, the riblets produced into bristles at the sutures. Protoconch l^r whorls, smooth or spirally ribbed. Mostly South Island, but one species in southern half of North Island. 3 species. 4x4 mm. Fig. 25, Aeschrodcmu8 stipulata (Reeve).

Genus CHAROPA Albers, 1860 (Helix coma Gray). Shell with spire nearly flat, widely umbilicated, closely axially costate. Some species with microscopic spiral sculpture. Protoconch l_r whorls, mostly smooth. Colour light or dark brown often with darker blotched or zigzag colour pattern. Aperture slanting backwards owing to the ribs being posteriorly oblique. 12 species and subspecies. 7x3 mm. Throughout New Zealand, Auckland and Antipodes Islands. Fig. 26, A, B, C. coma (Gray). 32 -

Subgenus (PSEUDEGESTULA) Dell, 1954 (Endodonta (Charopa) transenna Suter). Shell minute, depressed. Axial ribs slightly anteriorly oblique producing a slight sinus on the outer lip. Protoconch l"s smooth whorls. Spiral sculpture on base distinct. 2 species and 1 subspecies. 3x1 mm. North, South and Stewart Islands. Fig. 27, Charopa (Pseudegestula) transenna (Suter).

Genus EGESTULA Iredale, 1915 (Helix egesta Gray). Shell depressed, with radial riblets produced into membranous plates; distinctly spirally sculptured. Umbilicus wide, perspective. Protoconch I5 spirally costate whorls. 2 species. 4. 5 x 2. 5 mm. North Island and Three Kings Islands. Fig, 28, Egestula Egesta (Gray).

Genus FECTOLA Iredale, 1915 (Helix infecta Reeve). Shell dark or light brown with anteriorly oblique radial costae and usually with microscopic spiral sculpture. Aperture with distinct sutural sinus. Spire flat to slightly raised, widely umbilicated, Protoconch l_r radially striate whorls, 17 species, 4. 5 x 2. 5 mm. Fig. 29, A, B, Fectola infecta (Reeve); C, protoconch. North, South and Stewart Islands.

Genus SUBFECTOLA Powell, 1939 (Helix caputspinulae Reeve). Shell minute, white, moderately umbilicated. Protoconch 1-g-radially striate whorls. The straight radial ribbing gives the aperture a straight edge. North South and Stewart Islands. 2 species. 2. 8 x 1. 3 mm. Fig, 30, A, B, Subfectola caputspinulae (Reeve).

Genus MOCELLA Iredale, 1915 (Helix cornioulvm Reeve). Shell sub discoidal, qf silky appearance, with close, straight, axial costae. Peristome straight. Protoconch 1^ whorls finely spirally sculptured. Umbilicus moderately wide. Axial ribs straight and therefore aperture with straight edge. 6 species. 3 x 1. 5 mm. Three Kings, North and South Islands. Fig. 31, A, B, Mocella cogitata Iredale; C, protoconch. - 33 -

Genus CAVELLIA Iredale, 1915 (Helix biconcava Pfeiffer). Shell small discoidal, with concave spire. Axial ribbing closely anteriorly oblique forming a weak sinus; no spirals, Protoconch l£ radially striate whorls. Umbilicus broad, perspective. 1 species. 5x2 mm. Southern part of North Island, Fig. 32, A, E, Cavellia biconcava (Pfeiffer).

Genus GEMINOROPA Iredale, 1933 (Helix antialba Beddome), Shell minute, discoidal, deeply concave above and below. Finely radially ribbed, ribs straight, no spirals. Protoconch 1? whorls, spirally striate. Umbilicus very large and deep. 1 species. 1. 6 x 0. 75 mm. Southern parts of North Island. Fig. 33, A, B, Geminoropa vortex (Murdoch).

Subgenus (CAVELLIOROPA) Dell, 1952 (Diplomphalus subantialba Suter). Shell small, discoidal, closely radially costate, ribs straight, with microscopic spiral striae. Sunken spire. Protoconch of 1~ smooth whorls. Umbilicus rather broad and deep. 7 species, 2, 7 x 1. 5 mm. Southern North Island and South Island in leaf mould. Fig. 34, A, B, Geminoropa (Cavellioropa) huttoni (Suter).

Family LAOMIDAE

Foot without posterior mucous pore. Jaw thin, composed of separate rhomboidal plates, more or less imbricating. Radula with tricuspid or unicuspid central teeth; lateral and marginal teeth bicuspid.

Genus LAOMA Gray, 1849 Bulimus (?) (L. ) leimonias Gray). Shell very small, somewhat glossy, narrowly perforated, with plates within the aperture. Protoconch lijr smooth whorls. 7 species and subspecies. 4 x 3. 25 m.. North Island in bush debris and on ferns and tree trunks. Fig. 35, Laoma leimonias (Gray), Genus PHRIXGNATHUS Hutton, 1883 (Helix fatua Hutton » P. celia Hutton). Shells very small, of fragile and glossy appearance; light or reddish brown with colour markings of white blotches or streaks. Protoconch of lg- whorls smooth or spirally sculptured. Three main groups can be recognised: -

(i) Sharply angled at periphery, base flat or concave, some species imperforate, others umbilicated. Spire tall or depressed. Fig. 36, Phrixgnathus murdoohi (Suter).

(ii) Periphery rounded, base concave, some species imperforate, others umbilicated. Fig. 37, Phrixgnathus oriel Hutton.

(iii) Shell sub discoidal, periphery rounded, with microscopic sculpture. Mostly umbilicated but a few species imperforate. Fig. 38, Phrixgnathus clabriusculus (Pfeiffer)•

47 named species and subspecies which range in diameter from 1, 5 mm. to 8 mm. North, South, Stewart, Three Kings, Antipodes, Campbell and Macquarie Islands.

Subgenus (TAGUAHELIX) Powell, 1955 (Thermia expeditionis Suter). Animal with Phrixgnathus dentition but shell features more resembling miniature Phenaoohelix or Allodiscus (q. v. ) Spire moderately high periphery rounded. Axial riblets numerous and crisp, sometimes produced into hairlike processes. Densely spirally lirate. Umbilicus open and deep. Protoconch of 1^- whorls with crowded spiral striations. 2. 5 x 1. 5 mm. 4 species. Subantarctic Islands only. Fig. 39 Phrixgnathus (Taguahelix) campbellicus (Filhol). - 33 -

Genus LAOMAREX Powell, 1948 (L. sericea Powell). No apertural plates, small depressed shell, periphery angled, umbilicus deep and narrow. Protoconch 3_r whorls of dense beaded spirals followed by half whorl of closely spaced radials. Sculptured with close axial riblets and microscopic spiral threads. Dull brown with darker markings and lacks the typical translucent texture of Phrixgnathus. Three Kings Islands only, 1 species. 3. 25 x 2 mm. Fig. 40, Laomarex sericea Powell.

Genus PARALAOMA Iredale, 1913 (P. raoulensis Iredale). Very small shells, shining, flattish. Having widely separated axial riblets with reticulated sculpture in the interstices. Broadly umbilicated. Protoconch of l|- whorls. 8 species. 2x1 mm. Three Kings, North, South and Stewart Islands. Fig. 41, Paralaoma lateumbilicata (Suter).

Superfamily BULIMULACEA Family BULIMULIDAE Shell large, spire tall.

Genus PLACOSTYLUS Beck, 1837 (P. fibratus Martyn).

Subgenus (MAORISTYLUS) Haas, 1935 Bulimus hongii Lesson). Shell large, 3 to 4 inches tall, dark brown. Aperture reddish-brown or white with one or more tubercles. Protoconch 2 whorls. Whangarei to the Far North. 17 species and subspecies. Fig. 42 Placostylus (Maoristylus) hongii (Lesson).

Subgenus (BASILEOSTYLUS) Haas, 1935 (F. bollonsi Suter), Shell long, aperture white or reddish, no tubercles, Protoconch 3 whorls. Three Kings Islands only, 3 species and subspecies Fig. 43, Placostylus (Basileostylus) bollonsi Suter.

Superfamily ACHATINELLACEA Family ELASMATINIDAE Shell very small, thin, tall-spired.

Genus TORNATELLINOPS Pilsbry, 1915 (Tornatellina novoseelandica Pfeiffer), Shell very small, many whorled, smooth and shining, light brown, - 36 -

imperforate. Columella twisted with prominent fold. Parietal wall with distinct tooth. Three Kings Islands, North Island and northern South Island. 1. 5 x 3. 5 mm. 1 species. Fig. 44, Tornatellinops novoseelandica (Pfeiffer).

Genus ELASMIAS Pilsbry, 1910 (Tornatellina aperta Pease). Shell very small, dark brown, subperforate. Columella vertical, not twisted. Parietal wall with small teeth. North Auckland, mostly sand dune associations. 2. 2 x 3. 5 mm. 1 species. Fig. 45, Elasmias subperforata (Suter).

Suborder AGNATHA Superfamily STREPTAXACEA Family PARYPHANTIDAE Shell large or small, depressed, umbilicate, Radula with long teeth with one cusp. No jaw.

Genus RHYTIDA Albers, 1860 (Helix greenwoods Gray). Shell up to 1^ inches in diameter, light brown or olive green, sometimes shining, mostly with wrinkled appearance. Protoconch of radially sculptured whorls. Periphery of body whorl mainly rounded but is keeled in most North Auckland species. Widely umbilicated. Dentition 12 + 0 + 12 to 18 + 1 + 18, 19 species and subspecies. Fig. 46, Rhytida greerw)oodi (Gray); Fig. 47, Rhytida dunniae (Gray). North and South Islands, Stewart Island.

Subgenus (RHYTIDAREX) Powell, 1948 (Rhytida (R. ) johnsoni Powell. ). Shell larger, periphery of body whorl rounded. Protoconch with less distinct radial sculpture. Dentition 33 + 1 + 33. 2 species. Three Kings Islands only. Fig. 48, Rhytida (Rhytidarex) johnsoni Powell.

48 - 37 -

Genus DELOS Hutton, 1904 (Zonites coresia Gray). Shell small, flattish, glossy. Light yellowish-brown with darker colour pattern. Widely umbilicated. Smooth to naked eye, but finely sculptured with microscopic striae, Protoconch of Ig- convex whorls, with fine spirals and/or axials, 3x7 mm, 2 species. North Island. Fig. 49, Deles coresia (Gray).

Subgenus (DELOUAGAPIA) Powell, 1952 (Gerontia Cordelia Hutton). Shell like Delos but larger, up to 8 mm, in diameter, not glossy, dark brown with pale blotches or streaks. Sculptured with very coarse axial growth lines, umbilicus wide and deep, Protoconch l?r whorls with a shallow perforation at the centre, and microscopic spiral lines. North Auckland only. 1 species. Fig. 50, Delos (Delouagapia) cordelia (Hutton).

Genus PARYPHANTA Albers, 1850 (Nanina busbyi Gray). Shell 2g inches in diameter, uniformly dark olive green, widely umbilicate,

Protoconch 2 whorlse North Auckland only, 1 species and 1 subspecies. Fig. 51, Paryphanta busbyi (Gray).

Subgenus (POWELLIPHANTA) 1945 (Helix hochstetteri Pfeiffer). Shell larger (in most species), with axial or spiral colour patterns, Protoconch 2 whorls, Manawatu, Nelson, Marlborough and Westland. 42 species and subspecies. Fig. 52 Paryphanta (Powelliphanta) hochstetteri obscura Beutler.

Genus WAINUIA Powell, 1930 (Helix urnula Pfeiffer) Shell chitinous, shining, dark brown or olive green. Protoconch lif whorls. Southern half of North Island and northern half of South Island. 30 x 20 mm. 5 species and subspecies. Fig. 53, Wainuia urnula (Pfeiffer).

53 - 38 -

Genus SCHIZOGLOSSA Hedley, 1902 (Daudebardia novoseelandica Pfeiffer) Shell up to half an inch long, glossy brown to dark olive, ear-shaped and incapable of containing the large animal which is dark brown with a yellow foot. North Island, Great Barrier and Cuvier Islands. 5 species and subspecies. Fig. 54 Schizoglossa novoseelandica (Pfeiffer) animal; A, shell (dorsal and ventral aspects).

Suborder ELASMOGNATHA Superfamily SUCCINEACEA Family SUCCINEIDAE Anterior tentacles absent or small. Shell internal or external, thin, spiral, last whorl large. Jaw elasmognathic.

Genus PAPUSUCCINEA Iredale, 1941 (Succinea strubelli Strubell) (-Austro succinea Powell). Shell small, external, fragile, semi-transparent. Whorls 5, aperture large, entire, imperforate. Protoconch 1^ smooth whorls. An annual snail living among sand dunes, North Auckland East Coast and Bay of Plenty. 12 x 8 mm. 1 species. Fig. 55, Papusuccinea archeyi (Powell).

Superfamily TRACHEOPULMONATA Family ATHORACOPHORIDAE Slugs with internal, rudimentary shell usually composed of many calcareous deposits; a small dorsal mantle area defined by grooves and bearing a pulmonary cavity with numerous radiating diverticula; jaw elasmognathic. Anterior tentacles absent. Anus on right side, lateral to, or included within the mantle area.

Genus ATH0RAC0PH0RUS Gould, 1852 ( Limax bitentaculatus Quoy and Gaimard). Mantle area extends to perinotum, with a straight posterior lateral margin. Head shield triangular, not extending to mantle area. Skin thin, lacking prominent papillae, with not more than 15 shallow grooves on either side. 3 species. 38 mm. North, South and Stewart Islands. Fig. 56, Athoracophorus bitentaculatus (Quoy and Gaimard). Genus REFLECTOPALLIUM Burton, 1963 (i?. pseudophyllum Burton). Mantle area extends to perinotum, posterior lateral margin runs diagonally forward before recurving to run back to the perinotum. Head shield extending to mantle area. Skin thick, often with prominent papillae... not less than 1A lateral grooves on either side. A species. AO mm. South Island, Stewart Island. Fig. 57, Refleotopallium pseudophyllum Burton.

Genus PSEUDANEITEA Cockerell, 1891 (Janella papillata Hutton). Mantle area small, triangular, or rarely quadrangular, not extending to perinotum. Skin thick, finely granulate, usually with papillae. Lateral grooves 1A - 22 on either side. Anus not included in mantle area. 1A species. 100 mm. North, South, Subantarctic Islands. Fig, 58, Pseudaneitea papillata (Hutton).

Genus PALLI0P0DEX Burton, 1963 (Athoracophoru verrucosus Simroth). Mantle area broader than long, triangular, not extending to perinotum. 20 lateral grooves on either side, all unbranched. Anus included in mantle area. 1 species. 30 mm. Auckland Islands. Fig. 59, Palliopodex verrucosus (Simroth). Fig. 60. A generalised snail shell - apertural view. Aperture. Outer lip. Columella. Inner lip. Parietal wall Protoconch Umbilicus. Axial ribs. Spiral ribs. Lamella. Reticulate sculpture. Parietal tooth. Columella tooth. Labial tooth.

Fig. 61. A generalised snail shell - dorsal view. 1. Protoconch. 2. Limit of body whorl. 3. Body whorl. A. Sinus, 5. Anteriorly oblique axial ribs, 6. Straight axial ribs, -s 7. Posteriorly oblique axial ribs. 3 % Fig. 62. 61 A generalised slug (Athoracophorid) - a dorsal view. 1. Genital opening. 2. Anus, 3. Renal opening. 4. Pulmonary aperture. 5. Mantle area. 3 — 6. Median groove, 7. Lateral groove. 8. Perinotum. 9. Head shield. - 41 -

GLOSSARY

Anterior...... The front end. The anterior end of a snail is the lower part of the aperture. Anus The external opening of the intestine. Aperture...... The opening in a spiral shell. Axial Markings or sculpture extending from the upper to the lower part of a whorl. Base...... Lower part of shell. Body whorl The last whorl of a spiral shell - from the outer lip back to a point immediately above the upper lip. Buccal mass..... A muscular mass behind the mouth containing the radula and jaws. Central tooth (of radula).... The middle tooth of a radula tooth row. Columella...... Lower portion of axis of shell. Costate Ribbed. Cusp...... Cutting point on radula tooth. Granulated. •... Covered with minute grains or beads. Growth lines • •.. Lines on surface of shell not indicating true sculpture. Elasmognathic •... A type of jaw - a process emerges from the upper edge, the lower edge concave. Ectocone...... A sharp cutting point on the» outer side of a radula tooth. Entocone...... A sharp cutting point on the inner side of a radula tooth. Holognathic..,.. A type of jaw - upper edge simple, lower edge concave. Imperforate..... Having no umbilicus. Inner lip.. • «.. Inner part of aperture. Interstices..... Spaces between ribs. Jaw.. •»••<< A chitinous structure at the anterior end of the buccal cavity. Usually flattened, convex above and concave below. Often made up of a series of plates or rods.

Labial..... 0. Pertaining to the lips of the aperture. Lamellae...... Thin plates. Lateral groove... Groove running from median groove to perinotum in slugs. Lateral teeth.. •. (of radula). Teeth bordering the central tooth and usually larger and more complex than the outer (marginal) teeth. Lirate...... Fine sculpture. Mantle area •... • (in slugs)i The area containing the pulmonary aperture. - 42 -

Marginal teeth... (of radula) Teeth outside the laterals, usually simple. Median groove... Central longitudinal groove along the back of slug. Mesocone...... Sharp cutting point in middle of radula tooth Mesodont...... Blunt cutting point in middle of radula tooth Operculum...... A horny plate secreted by the foot and sealing off the aperture. Outer lip... 0. 4 Outer edge of aperture. Parietal wall.... Broader upper portion of inner lip. Peristome...... Edge of aperture. Perinotum • Groove between edge of dorsal surface and foot in slugs. Periostracum.... The external horny covering or "epidermis" of a shell. Perforate...... With an umbilicus Periphery...... The edge of the body whorl. Protoconch The embryonic shell or apex which forms the first portion of the spire. Radula The teeth of molluscs - usually very numerous and on a long ribbon. Reticulated..... Fine sculpture crossing at right angles. Rhipidoglossan radula...... Many similar teeth in each tooth row, though a central, laterals and marginals can usually be distinguished. Sinus Embayment in upper part of outer lip near suture. Spire All coils of the shell above the aperture. Striate . Fine sculpture. Subperforate.... Umbilicus very small. Suture...... Junction between whorls. Taenioglossan radula „...... With 7 teeth in each tooth row. The teeth clearly differentiated into 1 central, 2 laterals and 4 marginals. Umbilicus Hollow axis in middle of base. Whorl...... One complete spiral turn of the shell.

REFERENCES

Further references may be found in Suter's Manual heading the genera and species concerned. Most of the references given are available in local reference libraries.

Several articles written by local amateur conchologists regarding the distributionc ecology, etc9 of New Zealand mollusca have appeared in the bulletins of the Auckland Museum Conchology Section and in the Section's official publication "Poirieria, but are not included in this list 43 -

ADAMS, H. & A*, 1858. The genera of recent Mollusca. 3 vols. J. van Voort, London.

ADAMS, J«, 1886. The land and freshwater Molluscs of the Thames gold fields. Trans. N. Z. Inst., 19. pp. 177-181.

BAKER, H. P., 1938. The endodont genus Otoconcha, Proc. Mal. Soc., London, 23. pp. 89-91.

BURTON, D. W., 1963. A revision of the New Zealand and Subantarctic Athoracophoridae. Trans. Roy. Soc. N. Z., 3 (6). pp. 48-75.

______1962. New Zealand Land Slugs. Pt. l. Tuatara 9 (3). pp. 87-97.

1963. New Zealand Land Slugs. Pt. 2. Ibid. 11 (2). pp. 90-96.

COOKE, A. H., 1927. Molluscs. The Cambridge Natural History. 3. MacMillan & Co. Ltd. London.

CUMBER, R. A,, 1962. Palaeogegraphic history reflected in speciation trends of Charopa coma (Gray). Trans. Roy. Soc. N. Z., 88. (4). pp. 365-371.

______1960. Riblet frequency as a taxonomic character in New Zealand terrestrial Mollusca. Ibid. 88. (1) pp. 99-103.

______1961. A revision of the genus Phenaoohelix Suter, 1892. Ibid. 88 (1). pp. 163-196.

DELL, R. K., 1950. Two new species of land snails from the Snares Islands. Dominion Mus. Rec, 1 (3) pp. 55-58.

______1952. OTOCONCHA and its allies in New Zealand. Ibid. 1 (7). pp. 59-69. 1952. New Species and genera of New Zealand land snails with a revision of the genus Cavellia* Ibid. 1 (9) pp. 87-97.

______1954. The land Mollusca of Stewart Island and Solander Island. Trans. Roy, Soc. N. Z., 82 (5) pp. 137-156.

______1955, The land Mollusca of Fiordland, South-west Otago. Ibid. 82 (5) pp. 1135-1148.

______1955. Some New Zealand sub-fossil land Mollusca. Rec. Dominion Mus., 2 (3) pp. 135-144.

______1957. Natural areas in New Zealand land snails. Proc. N. Z. Ecol. Soc., 4 pp. 13-14. - 44 -

FLEMING, C. A., 1949. The geological history of New Zealand (with reference to the origin and history of the fauna and flora). Tuatara, 2 pp. 72-90.

HUTTON, F. W., 1873. Catalogue of the land shells of New Zealand. Colonial Museum and Geological Survey Dept., Govt. Printer, Wellington.

1880. Manual of New Zealand Mollusca. Govt. Printer, Wellington.

1882. Descriptions of some new land shells. Trans. N. Z. Inst., 15, pp. 134-141.

1883. Notes on some New Zealand land shells. Ibid. 16, pp. 161-186,

1883. Revision of the land Mollusca of New Zealand. Ibid. 16 pp. 186-212.

IREDALE, T., 1915. A commentary on Suter*s Manual of the New Zealand Mollusca. Ibid. 47 pp. 417-497.

______1915. A comparison of the land Mollusca faunas of the Kermadec Group and Norfolk Island. Ibid. 47 pp. 498-508.

MORTON, J. E., 1952. A preliminary study of the land operculate Murdochia pallidum. Trans. Roy. Soc. N. Z., 80 (1) pp. 69-79.

______1958. Molluscs. Hutchinson University Library, London,

MURDOCH, R,, 1847. Descriptions of new species of Endodonta and Flammulina from New Zealand. Proc. Malac. Soc. London, 2 pp. 161-162.

______1902. The anatomy of Paryphanta busbyi. Trans. N. Z. Inst., 35 pp. 258-262. O'CONNOR, A. C, 1954. Notes on the eggs of New Zealand Paryphantidae with a description of a new subgenus. Trans. Roy. Soc. N. Z., 75 (1) pp. 54-56.

POWELL, A. W. B., 1928. Descriptions of five new land shells. Trans. N. Z. Inst., 59 pp. 365-367.

1930. The Paryphantidae of New Zealand. Part 1. Rec. Auck. Inst. Mus,, 1 pp. 17-56,

1932. The Paryphantidae of New Zealand. Part 2. Ibid. 1 pp. 155-162, 45

POWELL, A. W. B., 1933. Two new land snails from New Zealand. Proc. Malac. Soc. London, 20 pp, 191-194.

______„. _____. 1935. Land Mollusca of the Three Kings Islands. Ibid. 21 pp. 243-248.

______1936, The Paryphantidae of New Zealand. Part 3. Rec. Auck. Inst. Mus., 2 pp. 29-41.

______1938. The Paryphantidae of New Zealand. Part 4, and the genus Placostylus in New Zealand. Ibid. 2 pp. 133-150,

______1939. The Mollusca of Stewart Island. Ibid. 2 pp. 211-238.

______1941. Seven new species of New Zealand land Mollusca. Ibid. 2 pp. 260-264.

. 1946. The Paryphantidae of New Zealand. Part 5. Ibid. 3 pp. 99-136.

______1947. Distribution of Placostylus snails in Northern most New Zealand. Ibid. 3 pp. 137-188.

______1948. Land Mollusca of the Three Kings Islands. Ibid. 3 pp. 273-290.

______1949. The species problem in New Zealand land snails. Trans. Roy. Soc. N. Z., 77 (5) pp. 202-208.

______1949. The Paryphantidae of New Zealand. Part 6. Rec. Auck. Inst. Mus., 3 pp. 347-372.

______1950. Life history of Austrosuccinea archeyi. Ibid. 4 pp. 61-72.

______1951. Land Mollusca from the four islands of the Three Kings Group. Ibid. 4 pp. 127-133.

______1951. On further colonies of Placostylus snails. Ibid. 4 pp. 134-140.

______1952, Four new species of New Zealand land snails. Ibid. 4 pp. 163-168.

______1954. The Molluscan land operculate genus Liarea. Ibid. 4 (5) pp. 271-293.

______1955. Mollusca of the Southern Islands of New Zealand, Cape Expedition series. Bull. 15, D. S. I. R., Govt. Printer, Wellington. pp. 1-293. - 46 -

POWELL, A. W. B., 1961, Shells of New Zealand. Whitcombe and Tombs Ltd., Auckland.

______1965. Land snails of the Auckland area. Science in Auckland. Handbook for eleventh Science Congress. D. S. I. R., Papatoetoe, Auckland.

PUNGA, M. T. Te., 1953. The Paryphantidae and a Cook Strait land bridge. N. Z. Journ. Sc. Tecn., 35 pp. 51-63.

SOLEM, A., 1959. Zoogeography of the land and fresh water Mollusca of the New Hebrides, Fieldiana Zoology, 43 (2) pp. 241-359.

SUTER, H., 1892. Contributions to the Molluscan fauna of New Zealand. Trans, N. Z, Inst., 24 pp. 270-278.

______1897. Notes on some New Zealand Flammulinidae. Proc. Malac. Soc. London, 2 pp. 284-285.

______1907. Descriptions of new non-marine shells from New Zealand. Ibid, 7 pp. 236-240,

______1909. The Mollusca of the Subantarctic Islands of New Zealand, in "The Subantarctic Islands of New Zealand". 2 vols, p. 36, Govt. Printer, Wellington.

______1913. Manual of the New Zealand Mollusca, with atlas of plates (1915). 1120 pp. Govt. Printer, Wellington.

SMITH, E. A., 1884. An account of the land and freshwater Mollusca collected during the voyage of the "Challanger", Dec. l872-May, 1876. Proc. Zool. Soc., 6 p. 275.

THIELE, J., 1929-1935. Handbuch der Systematischen Weichtierkunde. Jena.

TRYON, G. W. & PILSBRY, H. A., 1879-1913. Manual of Conchology. Philadelphia.

WENZ, W., 1938. Gastropoda. Handbuch der Palaezoologie, 6.

WEBSTER, W. H., 1904. New Mollusca from New Zealand. Proc. Malac. Soc London, 6 (2) pp. 106-108.

^^^^^^^__ 1907. Additions to the New Zealand Molluscan fauna. Trans. N. Z. Inst., 40 pp. 254-259.