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Living Landscapes Terrestrial Gastropods of the Columbia Basin, by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

Terrestrial Gastropods of the Columbia Basin, British Columbia Robert G. Forsyth, Volunteer, Royal British Columbia Museum

Acknowledgements

CIP data

Introduction Terrestrial Gastropods in British Columbia What are Terrestrial Gastropods? Terrestrial Gastropods in the Columbia Basin Distribution of Terrestrial Gastropods Introduced Identification of Species Collecting, Preserving and Dissecting Terrestrial Gastropods Morphology The Shell The : External Features Some Internal Anatomy Format of this Report

Systematic Checklist of the Terrestrial Gastropods of the Columbia Basin

Key to Terrestrial Gastropods of British Columbia Explanation to the key

Species Accounts Family Cionellidae Family Family Family Family Family Family Family Family Family Family Family Family Family Family Family

Glossary Literature Cited

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 , British Columbia http://www.royalbcmuseum.bc.ca V8W 9W2 1 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

Acknowledgements

Many people helped either directly or indirectly with this report. Barry Roth (Research Associate, Museum of Paleontology, University of California, Berkeley) identified Babine Mountain specimens of Pristiloma chersinella and some other material to which the Kootenay specimens were subsequently compared. He also kindly reviewed this paper. Heike Reise (Staatliches Museum fur Naturkunde Grlitz, Germany) and John M.C. Hutchinson (School of Biological Sciences, University of Bristol, Bristol, England) made many suggestions on the manuscript, provided literature and specimens. H. Reise also arranged the loan of speci- mens from Staatliches Museum fur Naturkunde Grlitz, which she and J. Hutchinson collected in 1998. Frederike Verspoor (RBCM librarian), Richard E. Petit (North Myrtle Beach, South Carolina), Harry G. Lee (Jacksonville, Florida) and Rachel Collin (Field Museum of Natural History, Chicago, Illinois) assisted in locating hard-to-find literature. Suzanne Beauchesne (Victoria, B.C.) and Drew Skinner (Bremerton, ) offered advice, information or specimens, and Tammy Forsyth helped collect specimens and search out literature.

The key was improved immensely by the suggestions offered by Larry Williams (Burnaby, B.C.). Thanks to Charles Salekin who donated the first Columbia Basin record of nemoralis from Nelson, B.C. Thanks to Phil Lambert, who made editorial suggestions to improve the text. Finally, I would like to thank Peter Newroth and Philip Lambert, Royal British Columbia Museum, who gave me the opportunity to take on this project and who, along with Kelly Sendall, provided the facilities, equipment and other support, and collected specimens.

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 2 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 3 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

Terrestrial Gastropods in British Columbia

Terrestrial, or land, gastropods -- hereafter referred to as and -- occur almost everywhere in British Columbia, from alpine meadows, screes, coniferous mountain forests and valley lowlands to urban gardens and parks. We notice larger species during wet weather in particular, but as a rule, slugs and snails are small, shun light and seek cool, moist places under logs, rocks and vegetation, making them difficult to find. The hard, spiral, calcareous shell of snails capture the interest of naturalists, and gardeners are likely to be very familiar with those species regarded as pests because of their insatiable appetites. Several species of terrestrial molluscs also serve as intermediate hosts for parasites of ungulates (Blood 1963; Boag & Wishart 1982).

The snails and slugs of British Columbia have never been adequately documented. The primary source of information for this region is Pilsbry's four-part monograph (1939, 1940, 1946, 1948). This work, how- ever, provides little information pertaining to British Columbia and the information is dated and often difficult to use. Publications since Pilsbry are scattered in various scientific journals and deal only with a few taxa or other geographic areas. Popular literature on the group is almost nonexistent.

This report is part of the Living Landscapes Project (1997 to 1999), sponsored by the Royal BC Mu- seum (RBCM) and the Columbia Basin Trust to study the natural and human history of the Basin (also known as the Kootenays). Given the paucity of information on terrestrial gastropods in British Columbia, a study of this kind seemed timely. The RBCM's collections revealed several species of snails and slugs not found elsewhere in the province. This Website provides identification aids to the species in the region and brings together information on their biology, ecology and distribution.

What are Terrestrial Gastropods? Snails and slugs belong to the phylum , a large and diverse group of non-segmented living in the ocean, freshwater and on land. The largest subdivision of the Phylum Mollusca, Class (the gastropods), includes several well-known examples: snails, slugs, , conchs and nudibranchs. Gastropods have a distinct head with tentacles and eyes, and a broad, flat foot for locomo- tion. There is a visceral mass dorsally, which is at least partially covered by a . The mantle encloses a mantle cavity that either contains for respiration or is modified to form a "". The visceral mass is usually protected by a calcareous shell.

The three subclasses of gastropods are based on the form and position of the respiratory organs, as well as on other anatomy. Members of the subclass have gills located in front of the heart. Prosobranchs, such as limpets, and whelks, live in marine, freshwater and terrestrial environments, but the majority are marine. There are relatively few terrestrial prosobranchs and none occur in British Columbia. The subclass has gills located behind the heart. All opisthobranchs, including the shell-less sea slugs, and some shelled forms, are marine. The subclass include mostly land snails and slugs, but a few are found in freshwater or in the sea. The mantle cavity of terrestrial pulmonates is modified into an organ capable of breathing air. All the terrestrial molluscs in British Columbia are pulmonates.

The three Orders within Pulmonata are defined by the number of pairs of tentacles, the position of the eyes on the head, and other anatomical details. The have one pair of tentacles with an eye at

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 4 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum the base of each. The group occurs mostly in freshwater but a few are terrestrial. is the only basommatophoran in British Columbia. The largest order, , has two pairs of tenta- cles with the eyes at the tip of the upper pair. The tips of Stylommatophoran tentacles can be retracted inside the tentacle. Most terrestrial molluscs belong to the Order Stylommatophora. is a relatively small group of slugs with no representatives in British Columbia. Like stylommatophorans, this group has a second pair of tentacles with eyes at their tips; however, the tentacles can be contracted but not retracted.

Terrestrial slugs are gastropods with a reduced shell that in most groups is internal. Slugs belong to several unrelated lineages, since shell reduction and a tendency towards a sluglike form has evolved several times within the gastropods.

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 5 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

Terrestrial Malacology in the Columbia Basin, British Columbia

The snails and slugs in British Columbia have received little stud. Four brief accounts (Whiteaves 1905; Vanatta 1906; Berry 1922; Carl & Hardy 1945) and the monograph of Pilsbry (1939-1948) are the only literature having individual records of terrestrial gastropods of our study region: the Columbia Basin. The remoteness and ruggedness of much of B.C., the lack of knowledgeable individuals, and the difficulty of observing, collecting and studying what are mostly small species, have likely all contributed to the current situation. Many of the published records were made many years ago and some need verifying.

The primary source of information for the terrestrial molluscs of North America is the comprehensive monograph Land Mollusca of North America (North of Mexico) by H.A. Pilsbry (1939-1948). After 1948, no additional information on the terrestrial molluscs of the Columbia Basin of British Columbia has appeared, although some taxonomic or distributional innovations have appeared from time to time (e.g. Vagvolgyi 1968; Bequaert & Miller 1973; Emberton 1995) for species known from the region.

While not specifically concerned with the fauna of the Columbia Basin in British Columbia, there are several publications that are significant because they deal with adjacent areas or are important for other reasons. Dawson (1875) reported on the molluscs along the 49th parallel between the Lake of the Woods and the Rockies, and although Dawson did not collect or report from the Kootenay region of British Columbia, he did describe a form of subrudis, as " limitaris" from Waterton Lake in the Rocky Moun- tains (now ). Other reports from Alberta include Taylor (1895a, 1895b) listing species from Laggan (now Lake Louise, Alberta), and Mozley (1926, 1931, 1933) and Platt (1980) for Jasper. Vanatta (1914) and Berry (1919) reported on terrestrial molluscs of Montana. In 1905 the pre-eminent American malacologist of the day, William Healey Dall published Land and freshwater mollusks of Alaska and adjoining regions (Dall 1905). This is historically important as an early attempt to enumerate terrestrial molluscs from Canada and Alaska. The last major compilation of the molluscs of Canada is the checklist by La Rocque (1953). The annotated checklist of Bequaert & Miller (1973) is important for its taxonomic innovations and summaries of species' distributions in western North America, although the checklist is for Arizona.

La Rocque (1962) and Drake (1963) give brief accounts of some of the personalities responsible for pioneering terrestrial malacology (study of molluscs) in British Columbia.

This report for the Columbia Basin was adapted from a partially completed manuscript by R.G. Forsyth covering all the known British Columbia terrestrial molluscs. Included are illustrations being prepared for the British Columbia manuscript. Collections of terrestrial molluscs were made by Kelly Sendall, Philip Lambert and others in September 1998, and this material was identified by the author and added to the report under "Records". To these RBCM records were added records of material collected by the author and his wife, Tammy Forsyth, in 1997 and 1998, and those collected by Heike Reise and John Hutchinson in 1998 (now deposited in Staatliches Museum fr Naturkunde Grlitz, Germany).

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 6 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

Conspectus of Terrestrial Gastropods in the Columbia Basin, BritishColumbia

Previously Unrecorded from the Previously Recorded and Previously recorded but not Columbia River Basin, B.C. Found in this Study Found in this Study

Arion rufus ptychophora elatior species kochi cyclophorella Cionella lubrica edentula1 whitneyi mullani fulvus laeve ? species3 ingersolli Discus shimekii binneyana maximus Nesovitrea electrina andersoni Oreohelix strigosa Pristiloma chersinella Oreohelix subrudis cf. californicum Vertigo gouldii Vallonia excentrica2 Vallonia pulchella2 pellucida Zoogenetes

1Whiteaves (1905) reported " edentulum" from Tobacco Plains, but this record was later re- identified as Columella alticola (now C. columella) by Berry (1922).

2Forsyth (in press) also discusses the Columbia River Basin records of V. pulchella and V. xcentrica.

3See text. Carl & Hardy (1945) have reported H. camelus from the one locality in the Columbia Basin region. This report records of an Hemphillia not unlike H. camelus but requiring verification.

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Distribution of Terrestrial Gastropods

Snails and slugs are commonly found in a wide variety of habitats but many species are very small and easily overlooked because they seek moisture and typically make their home away from direct sunlight. Seasonal differences in weather and moisture affect the presence and numbers of species. Some land snails and slugs are large and easy to find but small and medium sized species require a much more careful search. Look under logs, rocks, roots, fallen branches and fallen leaves or under rubbish or anything which offers some protection from exposure. At times of heavy rainfall, many species crawl up vegetation, finding shelter under the loose top layer of newly fallen leaf litter or on the undersides of fern fronds or other vegetation. Very small species are best found during such times; a handful of loose leaf litter may contain a number of minute snails.

More is known about the coastal faunas than those inland, but the generalizations probably hold true for the fauna of the Kootenay region. Myers (1972) determined that the amount of organic matter in the soil and soil texture, as related to water retention, corresponds directly to the diversity of terrestrial gastropods in alder, cedar and fir habitats in western Washington. These studies and Cameron (1986) found that the greatest diversity and abundance of terrestrial gastropods were under cedar, and least in Douglas-fir habi- tats. Karlin (1961) had earlier observed a strong association between terrestrial gastropods and deciduous trees in many localities from northern Montana south to Colorado.

Cameron (1986) investigated the coastal faunas on Island and the Fraser and Chilliwack Valleys and found that the type of litter, soil and associated vegetation influence diversity and abundance of snails. He characterised litter as either mull - a granular forest humus that forms a layer of mixed organic matter that merges gradually into the mineral soil beneath; or mor - a forest humus forming a layer of largely or- ganic matter abruptly distinct from the mineral soil beneath. High diversity and abundance of terrestrial molluscs is associated with mull litter, which is richer in nutrients and damper than mor litter. Soil, litter and vegetation types are usually correlated. Localities with mull litter are dominated by cedar and/or various deciduous trees, such as bigleaf maple. Localities with mor litter are dominated by Douglas-fir, spruce, or pine. Cameron found that faunas on mor litter and litter intermediate between mor and mull are impover- ished. This is congruent with observations made by others; Boag & Wishart (1982) and Karlin (1961) found that coniferous forests (characteristically having a mor litter) have about the same number of species as deciduous and mixed forests but with significantly lower densities.

Cameron also determined that disturbed sites, usually dominated by deciduous forests and therefore having corresponding mull litter, are often rich in species, and may also contain additional non-indigenous species. He also noted that burning and logging does not significantly reduce diversity.

The distributions of species are dependent upon several factors, including climate, geography, topography and their ancestral migration. Distributions of terrestrial molluscs do not correspond to the geopolitical boundaries of British Columbia but rather follow natural limitations. Along the coast, the ocean moderates the climate. In the east, the form a natural barrier for migration. Towards the north, climatic conditions become progressively more severe.

As a rule, British Columbia has considerably fewer species of terrestrial gastropods than southern Washing- ton, Oregon, Idaho and other areas south. British Columbia has no known endemics (species confined to a

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 8 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum particular geographic area). In British Columbia several species of snails and slugs occur only in the Kootenay region, their main distribution being south into the United States. After the retreat of the last glaciers, the Columbia Basin was repopulated by terrestrial gastropods from the south. The genera Oreohelix, Anguispira andAllogona have their origins south of B.C. The greatest species diversity is consequently in the southern part of the Columbia Basin region. The northern portion of the Columbia Basin lacks the distinctive elements of the southern fauna, and has fewer species, but these are more widespread.

Introduced Species

Approximately 25% of the species known to occur in British Columbia are exotic (intro- duced from another geographic area). All are ultimately European in origin, except perhaps and which are native to Eastern North America as well as . For the vast majority of these exotic species, the time, place and method of introduction is not known. However, the spread of exotic species subsequent to their initial introduction appears to be with and related material (i.e. compost or soil) (Forsyth in press), and earlier Getz & Chichester (1971) noted that for all European slugs introduced into North America transport was with material. In the Columbia Basin region, there have been six species of non-native terrestrial gastropods identified to date. They are Arion rufus, Deroceras reticulatum, ,Vallonia pulchella, Vallonia excentrica, and Cepaea nemoralis. Another exotic species yet to be identified to species, but known to belong to the complex of species, has also been recorded. Other species could also be present in and near towns, cities and agricultural areas, but our collecting did not focus on such areas.

Important papers on exotic terrestrial gastropods of British Columbia are Hanna (1966),Rollo & Wellington (1975), Forsyth (in press) and Reise et al. (in press).

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Identification of Species

To aid in identifying species, a good hand lens of 10-20 magnification or ideally a low-power microscope for observing minute details and small specimens is required. A set of callipers or a ruler (in mm) is needed for measuring. If a microscope is available, then a scale in the eyepiece is ideal for measuring small species.

Almost all of our species of snails can be identified by shell characters, but it is especially important to have fresh, mature shells. Worn shells without a , which can be diagnostic, make unsuitable material as they may be difficult or impossible to identify. Juvenile shells can be a problem, to the novice, because some juveniles look quite different from the adults, and often lack some diagnostic characters. With experi- ence, recognition of adults and corresponding juveniles will become easier, and with practice on fresh material old worn shells may be recognisable.

For a few species of snails, the colour of the body is useful for identification. For example, Zonitoides nitidus and Z. arboreus have similar shells but the animals are markedly different in colour. The former is nearly jet black, and the latter is greyish on the head, tentacles and back and lighter on the sides. Make a habit of noting the colour of all species you collect. The colour is given in most of the species descriptions.

Slugs are generally more difficult to identify than snails. The shell of slugs is rudimentary and either internal or at least partially external. Little information about a slug's identification can be derived from a slug's shell, except perhaps that it belongs to a certain family or genus. For example, the presence of a partially exposed flat shell midway on the back of a slug indicates that it is a Hemphillia. Useful external characters in identi- fying slugs to genus are the: size of the animal, position of the , texture of the mantle, extent to which the keel on the tail is developed and presence and placement of an external shell. Size, shape, band- ing and overall colour, tubercles, sole, and stickiness and colour of the are other important charac- ters. Internal anatomy (particularly of the distal genital system) is diagnostic, and dissection may be required to confirm identifications of some species. Important features of the of slugs are given below.

Variation in nature is commonplace and should always be expected. Variation in colour and form will likely be encountered, but in most cases subtleties in form or colour are of no taxonomic significance. When known, common variants are described.

Morphology

The two colloquial terms, -like and sluglike, refer to the outward appearance of the animal, but do not imply any phylogenetic relationships. Snail-like gastropods have a coiled shell into which the animal can completely or almost completely withdraw. Sluglike gastropods have the appearance of a typical slug, either with no shell, an internal shell, or with a small shell partially embedded in the posterior part of the mantle, but into which the animal cannot fully retract. For this report, snails are identified by external char- acters of the shell and the body. Ideally, slug species should be confirmed by internal anatomy -- external characters are sufficient for identification to genus. For many species, considerable variation in characters can complicate identification.

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 10 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum The Shell The majority of terrestrial molluscs have a coiled, external calcareous shell. In most sluglike gastropods, the reduced shell is usually internal and platelike, but in Arion, the shell is very rudimentary and consists of only a few irregular, discrete calcareous granules. The following section introduces and defines many of the specialized terms for describing gastropod shells.

Internal shell of the slug Limax maximus

The platelike shell of a slug is convex above and concave below. The earliest part of a slug's shell is the nucleus; the nucleus is positioned near the midline or to the side. Fine incremental lines are present on the surface of the shell.

As indicated by the many terms used to describe it, the coiled shell of a snail is much more complex in form than the shell of a slug. The revolutions or turns of the tube that makes up a coiled shell are called whorls. The whorls spiral around an imaginary central axis, and the axis runs through the central pillar of the shell known as the columella. Whorls may enlarge rapidly or slowly, and are separated from one another by a single, continuous line called the . Some thin-shelled species have a second line or a broad, more opaque band next to the suture giving the appearance of being a double suture. In almost all examples, the largest of a shell is the last whorl (=), and the second to last is the penulti- mate whorl. The consists of all the whorls except the last. The tip of the spire is called the , and the opposite end is the base. The animal extends out of its shell through an opening called the [=].

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 11 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

Shells can be viewed in apical, basal, apertural and abapertural views. (The "usual" orientation of a shell is an apertural view, with the aperture toward the viewer and with the apex pointing up.)

Width (= breadth, diameter) is the maximum measurement of the shell at right angles to the central axis. Width does not include the outwardly flared at the edge of the aperture, if this condition is present. Height (=length) is the maximum measurement along the central axis. In the species accounts, the width is only given when it exceeds the height, and height is given when it exceeds the width. These two terms refer to the shell, not the orientation of the shell on the living animal.

In basal view, some coiled shells have a hole or depression, the , which may be wide enough to show the basal surfaces of all or some whorls of the spire. In some shells, there is no umbilicus, but when it is present, its size is expressed as a percentage of the maximum shell width (e.g. umbilicus 25% of the shell width). Seen from the side (i.e. in apertural or abapertural view) the whorls may be flat or convex in profile. The outermost extent of a whorl is its periphery, which may be placed at, above, or less frequently below, the midline of the whorl. For example, if the periphery is above the midline, then the periphery is closer to the suture than if the periphery was below the midline of the whorl.

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At the apex of the spire, the , formed at a very young age, typically has different surface sculp- ture from the other whorls which are termed the teleoconch.

The general form of coiled shells may be described by the following terms (in each case the shell is in apertural view): heliciform (resembling the genus Helix, with the width and height approximately equal and the whole shell somewhat rounded); depressed-heliciform (heliciform, but shell is short and wide); conic-heliciform (heliciform but with the spire more cone-shaped); cylindric (with sides nearly parallel and ends blunt); subcylindric (close to cylindric in form, but with sides not nearly so parallel); elongate-taper- ing (with sides of the spire distinctly tapering to a point and apex pointed); subovate (more or less egg- shaped, with sides of the spire more or less convex); conic-ovate (basally ovate, apically conic); and elongate-ovate (more or less elliptical).

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 13 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

The shell surface is usually sculptured or coloured in a spiral or axial pattern. Spiral or colour patterns run parallel to the suture, and axial patterns are more or less parallel to the central axis of the shell (or perpendicular to the spiral direction).

Colour patterns occur in some species of snails. Narrow or broad spiral bands of colour or axial streaks may be present. Shells without pattern are said to be unicolour. The shell may be opaque or translucent, and glossy or dull. Glassy indicates a combination of both translucent and glossy, like a piece of glass.

Textural ornamentation of the surface of a shell is called sculpture, which may be either raised or incised. Even apparently smooth shells usually have very fine sculpture when viewed under magnification. Striae are fine, usually incised but sometimes raised, lines or ridges that may be regular in size and spacing or

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 14 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum irregular and wrinklelike. Ribs are larger, raised, angular or rounded ridges that may be regular in size and spacing, or not. Riblike elements with very narrow, bladelike edges are lamellar ribs. Striae, lamellar ribs and ribs may be either spiral or axial. Beads may be formed where striae intersect with ribs or striae. In some species, shallow, irregular depressions are referred to as malleations.

The edge of the aperture is the lip, which is divided into four regions: outer lip, basal lip, columellar lip and parietal lip. In most shells, the parietal lip is indistinct, being more or less fused to the preceding whorl and with only a thin layer of shell or . The outer and basal lips in particular may be thickened, flared, or reflected (hooked back). The aperture may have one or more projections within called denticles (=teeth), which are designated by their position (e.g., a parietal denticle is situated on or adjacent to the parietal lip).

Lip of the aperture: outer lip (o), basal lip (b), columellar lip (c) and parietal lip (p) with denticle.

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The families Vertiginidae and possess denticles and some unique features on the outer lip. Terms for these are adapted from Pilsbry (1948), but for brevity, two letter codes for identifying denticles are utilised here. The plicae (singular, plica) are denticles located on the outer lip; these have also been termed folds. Lamellae (singular, lamella) are denticles located on the columellar lip and parietal wall. The four major denticles present in most of our Vertigo species, are the columellar lamella (Lc), parietal lamella (Lp), upper palatal plica (Pu) and lower palatal plica (Pl). Accessory to these, and either present or wanting -- depending upon the species -- are the angular lamella (La), infraparietal lamella (Li), suprapalatal plica (Ps), infrapalatal plica (Pi) and subcolumellar lamella (Ls).

L(.s)

The method used for counting whorls is that of Kerney & Cameron (1979), Gerber (1996) and others, but not Pilsbry (1936). With a hand lens or microscope, orient the shell in apical view. On the protoconch determine where the suture first begins, then imagine a line as in the figure below. Count the whorls from this line, estimating the number of whorls to the nearest one-quarter turn. (Each crossing of the line will be one-half whorl.)

In the descriptions, shells are described as very small, small, medium, large, or very large (refer to the table), which is then followed by a range or a maximum dimension in millimeters (either width or height, whichever is the greatest).

The Animal: External Features The external colour of snails and slugs can sometimes be helpful in identifying species. Many slugs, how- ever, are quite variable in colour and patterns. Animals without spots or bands are said to be unicolour. Darker lateral bands may be present on the back of many slugs. Thehead is at the leading end of the body. In the basommatophorans, there is one pair of sensory tentacles with eyes near the bases. In the stylommatophorans, there are sensory tentacles and occular tentacles. The occular tentacles, bearing eyes, are longer than the sensory tentacles and situated above (or posterior) to them. The mouth is located on the underside of the head.

The mantle is a conspicuous raised area on the dorsal surface behind the head of a slug. The mantle of snails is less conspicuous, usually only visible around the edge of the shell. The mantle of slugs may be

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The body is the dorsal surface of the animal exclusive of the head and mantle. The skin may be smooth or roughened by tubercules or wrinkles. The tail is the posterior end of the body and either tapers gradually to a point or is truncate (abruptly cut off). Many slugs show a sharp ridge, or keel, down the midline of the body. It may extend from the tip of the tail to the mantle or be incomplete and not reach the mantle. Some groups have a caudal mucus gland at the tip of the tail that secretes slime or mucus. The opening of this gland is termed the caudal mucous pore (=caudal mucous gland), which may bear a mucus plug. The genital pore, located on the right side of the animal behind the right occular tentacle, is smaller than the pneumostome and leads into the reproductive tract of the animal.

The flattened ventral surface of the body is the sole of the foot. The sole may be white or pigmented. If it appears to be divided into three longitudinal bands, the sole is said to be tripartite. The edge of the foot may be expanded up the sides of the body and may be developed into a foot fringe (=skirt). In some slugs, the foot fringe may be brightly pigmented.

Mucus, secreted from the sole and body, can be clear, milky, or pigmented (yellow or orange), and sticky, or thin and watery. The colour of the mucus can be useful for identification of some slugs.

In the species accounts, a general statement of size (small, medium-sized, large and very large) for sluglike gastropods is given (refer to table). These and the actual length (in millimeters) are in reference to the length of the entire animal when it is fully extended.

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Some Internal Anatomy The reproductive tract is a significant taxonomic trait, and cannot be avoided in identifying some of our species. Pulmonates are simultaneous (both sexes in the same animal) and therefore pos- sess a complex the reproductive tract. At the proximal end of the reproductive system is the ovotestis (=gonad, gland) where gametes are produced. The ovotestis is connected to the hermaph- rodite duct, which runs forward to connect, along with the albumen gland, to the common duct (=spermoviduct). The male and female gametes are separated before entering the common duct, which has various male and female glands associated with it. The reproductive system distal to the common duct is

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The male portion of the distal reproductive tract consists of the vas deferens, and structures associ- ated with these, although the penis is absent in some genera (Arion, for example). The vas deferens is a long, narrow duct connecting the common duct to the penis or epiphallus. The epiphallus is a dilated distal portion of the vas deferens and is present in some genera and not in others. The vas deferens or epiphallus then usually connects to the penis, but in Arion the penis is absent.

The penis is an eversible saclike organ. Attached to the inside of the penis in some groups is a stimulator (=sarcobellum). Also present in some is a penial flagellum (=appendage, appendix, ). The penis (or epiphallus when the penis is absent) opens into the genital atrium.

The female portion of the reproductive tract consists of the oviduct, which transfers eggs from the common duct to the genital atrium.

In Arion, the atrium is further subdivided into an upper and lower atrium. A stimulating organ called the ligula can be present in the atrium. Also entering the atrium is the spermathecal duct (=bursa copulatrix duct), which leads to the spermatheca (=bursa copulatrix), a sac for storing spermatozoa. Finally, the atrium opens to the outside of the animal through a single hole, the genital pore. The genital pore is located on the animal slightly behind and below the right upper tentacle on the head.

A detailed account of the reproductive system of slugs can be found in Terrestrial Slugs by South (1992).

Although the structure of the jaw, , digestive tract and other organ systems are considered useful in pulmonate systematics, they are not considered here.

A complete list of anatomical and conchological terns in the glossary.

Collecting, Preserving and Dissecting Terrestrial Gastropods

Because so much of British Columbia is relatively unstudied, collections of specimens supplied with data, can be valuable to science. Basic specimen data, as given by Solem et al. (1981), consists of:

1. Date of collection and collector. 2. Locality, so indicated that the exact place of collection can be found by another person in the future (latitude and longitude data and reference to permanent, mapped geographic features). 3. Basic ecological occurrence: major vegetation type; any associations with other organisms; whether collected on trees, in leaf litter, under rocks, logs, bark or wood, etc.

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Specimens of slugs should be preserved by first drowning the slug in water overnight, then placing it in 70% ethanol. It is also best that snails are preserved in the same fashion, otherwise larger snails will have to have the animal removed from the shell and very small species can simply be placed into alcohol for a few days, then taken out and left to dry.

For how to dissect slugs and snails, refer to Kerney & Cameron (1979).

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Format of this Report

In the species descriptions the format is as follows: species are arranged alphabetically within their respec- tive genera and genera are arranged alphabetically within subfamilies or families. All higher categories are arranged in systematic or phylogenetic order, following the systematic checklist. No diagnoses of higher groups are given, and only important literature references, other than Pilsbry (1939-1948), are given for supraspecific taxa. For each species, the Latin name with author and date is given. Synonyms likely to be encountered in other publications and particularly those made since Pilsbry (1939-1948) are given. Special emphasis is placed on synonymous names used in connection to British Columbia or immediately adjacent areas. Species accounts emphasize the shells of snails and the bodies of slugs. The colour and outward appearance of the body of snails is also noted when known.

Following the description, either a full comparison with similar species or a referral to another species is given. In the latter case, the full comparison will be made under the referred species.

The habitat of each species is noted with special emphasis on habitats of British Columbia populations. The species' world-wide range, or distribution, is given, and its occurrence in British Columbia.

Notes on , problematic records, etc., and finally derivations of the currently valid scientific name are presented. For genera with multiple species, the derivation of the generic name is given under the first species only. Records based on museum or private collections, list locality, geoposition and a catalogue number. For literature records, locality, geoposition and reference(s) are noted. Unless given in the original reference or collection data, latitudes and longitudes of records were secondarily obtained from a gazetteer or other sources. Records are ordered south to north and east to west.

The distribution maps show only the Columbia Basin records. A filled circle indicates that the record is based on museum or private collections. An open circle indicates a literature record. A circle may represent more than one record when localities are too close to plot separately, and when a collection record coin- cides with a literature record, the collection record takes precedence.

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Systematic Checklist of the Terrestrial Gastropods of the Columbia Basin, British Columbia

This list includes all species of terrestrial molluscs known from the Columbia Basin of British Columbia, but two cautionary notes are necessary. Firstly, the classification is not final and is still undergoing change. This report does not attempt any taxonomic innovations toward a better system, and the classification used here was adopted from several sources, which means that the systematics of this checklist are not uniformly grounded in modern techniques, since most groups have not been revised in many years. Secondly, this checklist is probably not complete, given that the Columbia Basin region (and much of British Columbia) has not been thoroughly explored for terrestrial molluscs.

Several introduced species are included in this list and are indicated by an asterisk.

Following the main list is a second list of species that have been recorded from areas adjacent to the Kootenays, but which have not yet been collected there.

Superfamily Cionelloidea Kobelt, 1880 Family CIONELLIDAE Kobelt 1880 Genus Cionella Jeffreys, 1829 Cionella lubrica (Mller, 1774) Superfamily Turton, 1831 Family VERTIGINIDAE Stimpson, 1851 Genus Columella Westerlund, 1878 Columella columella (Martens, 1830) (Draparnaud, 1805) Genus Vertigo Mller, 1774 Subgenus Vertigo Mller, 1774 Vertigo elatior Sterki, 1894 Vertigo gouldii (A. Binney, 1843) Vertigo modesta (Say, 1824) Vertigo ovata Say, 1822 Family VALLONIIDAE Pilsbry 1900 Genus Vallonia Risso, 1826 Sterki, 1892 *Vallonia excentrica Sterki, 1893 *Vallonia pulchella(Mller, 1774) Genus Zoogenetes Morse, 1864 Zoogenetes harpa (Say, 1824) Superfamily Gray in Turton, 1840 Family PUNCTIDAE Morse, 1864 Genus Punctum Morse, 1864 Punctum californicum (Pilsbry, 1898) Family DISCIDAE Thiele, 1931 Genus Anguispira Morse, 1864 Subgenus Zonodiscus Pilsbry 1948

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Subfamily Gastrodontinae Tryon, 1866 Genus Zonitoides Lehmann, 1862 Subgenus Zonitoides Lehmann, 1862 Zonitoides arboreus (Say, 1816) Zonitoides nitidus (Mller, 1774) Family VITRINIDAE Fitzinger, 1833 Genus Vitrina Draparnaud, 1801 Subgenus Vitrina Draparnaud, 1801 (Mller, 1774) Family LIMACIDAE Gray, 1824 Genus Limax Linnaeus, 1758 Subgenus Limax Linnaeus, 1758 *Limax maximus Linnaeus, 1758 Family AGRIOLIMACIDAE Wagner, 1935 Genus Deroceras Rafinesque, 1820 Subgenus Deroceras Rafinesque, 1820 (Mller, 1774) Subgenus Agriolimax Mrch, 1855 *Deroceras reticulatum (Mller, 1774) Superfamily Polygyroidea Pilsbry, 1895 Family POLYGYRIDAE Pilsbry, 1895 Subfamily Polygyrinae Pilsbry, 1895 Tribe Allogonini Emberton, 1994 Genus Cryptomastix Pilsbry 1939 Subgenus Cryptomastix Pilsbry 1939 (Bland & Cooper, 1881) Genus Allogona Pilsbry 1939 Subgenus Dysmedoma Pilsbry 1939 Allogona ptychophora (A.D. Brown, 1870) Family THYSANOPHORIDAE Pilsbry 1926 Genus Microphysula Cockerell & Pilsbry 1926 Microphysula ingersollii (Bland, 1875) Superfamily Rafinesque, 1815 Family HELICOIDEA Rafinesque, 1815 Genus Cepaea Held, 1837 Subgenus Cepaea Held, 1837 *Cepaea nemoralis (Linnaeus, 1758)

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There are several additional terrestrial gastropods that have not yet been collected from the Columbia Basin of British Columbia, but have been recorded from adjacent areas. These are recorded below to ensure awareness of these potential species.

Allogona lombardii A.G. Smith, 1943: Idaho (Smith 1943). holzingeri (Sterki, 1889): Alberta (Taylor 1895); Montana (Pilsbry 1948). (Say, 1822): Montana (Vanatta, 1914); "Laggan" [=Lake Louise], Alberta (Taylor 1893). vancouverense (I. Lea, 1839): N.W. Montana (Brunson & Osher 1957), N.E. Oregon and N. Idaho (Pilsbry 1946; Binney & Bland 1869); also along the coast from the Aleutian Islands (Hanna 1925) and the Alexander Archipelago, Alaska, to northwestern California. Known from the interior of British Columbia, only by a single record from Salmon Arm (RBCM 998-00018-002). Not yet reported from the Kootenay region, but potentially it could be found there. Hemphillia camelus Pilsbry & Vanatta, 1897: Idaho (Pilsbry 1948). Hemphillia danielsi Vanatta, 1914: Montana (Vanatta 1914; Pilsbry 1948). ovalis (Say, 1817) [as Succinea ovalis]: Alberta (Taylor 1893). Oxyloma nuttalliana (I. Lea, 1841) [as Succinea nuttalliana]: Montana (Vanatta 1914). caputspinulae (Reeve, 1854) (as Punctum conspectum (Bland, 1965): Montana (Vanatta 1914, Pilsbry 1948). Idaho (Pilsbry 1948). Recorded from many places throughout British Columbia. Pristiloma idahoense (Pilsbry, 1902): Idaho (Pilsbry 1946). Punctum minutissimum (I. Lea, 1841): An eastern Nearctic snail (Bequaert & Miller 1973) reported from Idaho (Pilsbry 1948) and Kananaskis, Alberta (Berry 1922) as P. pygmaeum (fide Bequaert & Miller 1973). Punctum randolphii (Dall, 1895): Reported from Idaho (Pilsbry 1948); also coastal BC, Washington, Oregon (Pilsbry 1948). blandi (Morse, 1865): Alberta (Taylor 1893, 1895; Pilsbry 1948). Montana (Pilsbry 1948). Pupilla hebes (Ancey, 1881): Idaho (Pilsbry 1948). British Columbia. (Linnaeus, 1758): Alberta. Pupilla syngenes (Pilsbry, 1890): Montana (Pilsbry 1948). species: Jasper, Alberta (Platt 1980). Radiodiscus abietum H.B. Baker, 1930: Idaho (Baker 1940); Montana (Brunson & Russell 1967) ferrea E.S. Morse, 1864: Jasper, Alberta (Platt 1980). Striatura pugetensis (Dall, 1895): Montana (Berry 1919); Thunder River, B.C. (H. Reise, personal communication); also along the south Coast of British Columbia. Succinea oregonensis I. Lea, 1841: Montana (Vanatta 1914). Succinea grosvenori I. Lea, 1864: Alberta (Taylor 1895). Reinhardt, 1883: Alberta (Taylor 1893, 1895; Gerber 1996). Idaho (Pilsbry 1948). Montana (Gerber 1996). Vitrina angelicae Beck, 1837 [as Vitrina limpida Gouldin Agassiz, 1850]: Alberta (Taylor 1895; Mozley 1926; Pilsbry 1946; Platt 1980). Zacoleus idahoensis (Pilsbry, 1903): Idaho (Pilsbry 1948).

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Key to Terrestrial Gastropods of British Columbia

The following is an identification key to all terrestrial gastropods known from British Columbia. The species and their relative systematic position are given in the Systematic Checklist. The key relies on shell charac- ters, if present, and external characters of the body, and keys out directly to species. A key to families and other higher categories of snails and slugs based solely on external features of the shell or animal is impracti- cal, given that the classification of the pulmonate gastropods is based upon anatomy of the animals.

This key is for adult snails and slugs. Adults of many species of snails can be recognised by a thickened or reflected lip, a downward turning of the last whorl at the outer lip, or by the development of teeth around the aperture. In others, where these features do not occur at maturity, recognition of adult specimens may be more difficult. Ideally several specimens should be studied. Notes on the body colour of the living animal are sometimes helpful. Weathered or eroded shells may not show surface sculpture or colouration well, and may prove difficult to key out. Consult fresh specimens whenever possible.

Many species of slugs are difficult or impossible to distinguish by external characters, as are members of the family Succineidae. Reference to internal anatomy is therefore unavoidable in identifying these gastropods. As a result, slugs and succineid snails are not fully keyed out, and some anatomical distinctions are pre- sented in the text only for these groups.

The snails of the genus Vertigo and Nearctula are not keyed out to species. Significant characteristics of those species recorded from the Columbia Basin region are presented in a table preceding the species descriptions.

For completeness and as an aid to the beginner who may on occasion find certain aquatic and semi-marine snails living in close association with terrestrial molluscs, several species are included here that are not truly terrestrial. These are not included in the text and are marked either as "semi-marine" or "freshwater".

Finally, those gastropods known from the Kootenay region of British Columbia are set in bold type.

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Key to Terrestrial Gastropods of British Columbia

1a Sluglike gastropods: shell absent or much reduced; shell not in definite coils and internal or small and partially external on the posterior part of the mantle -69 1b Snail-like gastropods: shell coiled into which the animal can more or less retract -2

2a Shell height greater than width -3 2b Shell width greater than or equal to height -17

3a Aperture with 1 or more denticles -4 3b Aperture without denticles -9

4a Adults typically with 4 or more denticles within the aperture -8 4b Adults with 1 or 2 denticles -5

5a Aperture greater than shell height; usually associated with marine marshes - myosotis - semi-marine 5b Aperture less than shell height; fully terrestrial -6

6a Shell white; spire gradually tapering -Carychium occidentale 6b Shell brownish; spire bluntly pointed -7

7a Adults with a single denticle on the parietal wall, connected to the top of the outer lip and sometimes a columellar denticle (obscured by the columellar lip); juveniles with apertural dentition; no crest - cylindracea Introduced to British Columbia (Forsyth in press) 7b Adults usually only with a columellar denticle (obscured by the columellar lip); rarely with a small parietal denticle, but not connected to the top of the outer lip; juveniles without apertural denticles; crest hardly or only moderately developed -Pupilla hebes

8a Shell whitish or translucent horn-coloured (denticles at least 5) -Gastrocopta pentodon 8b Shell brown, reddish brown or yellowish brown (if yellowish, then with only 4 denticles) -Vertigo species A key to Vertigo and Nearctula species is not given the amount of variation exhibited in some spe cies. Four Vertigo are known from the Kootenay region: V. elatior, V. gouldii, V. modesta and V. ovata.

9a Shell with well-spaced lamellar axial ribs Zoogenetes harpa 9b Shell smoothish or with axial striae only -10

10a Shell more or less cylindrical -11 10b Shell not cylindrical -15

11a Early whorls of the spire lost in adults; present; associated with marine marshes Cecina manchurica semi-marine

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11b Early whorls of the spire not lost; no operculum; fully terrestrial -12

12a Shell surface very smooth with a high gloss; umbilicus absent; lip with a narrow riblike thickening within Cionella lubrica 12b Shell surface not glossy or only somewhat glossy; umbilicus present; lip without a riblike thickening within -13

13a Outer lip narrowly reflected and slightly thickened; columella with a small denticle obscured by the columella except when viewed from a slight angle Pupilla hebes 13b Outer lip neither reflected nor thickened; no columellar denticle -14

14a Shell tapered-cylindrical; axial striae fine; length 1.8-2.5 mm (5-6 whorls) -Columella edentula 14b Shell very cylindrical; axial striae rather coarse; length 2.5-3.0 mm (6-7 whorls) -Columella columella

15a Whorls about 3; columella never strongly twisted or thickened; eyes at the tips of ocular tentacles. Succinea species, Oxyloma species, Catinella species Identification of species (and to a lesser extent, identification of genera) by shell characters is unreliable 15b Whorls 4-6; columella often strongly twisted or thickened; no ocular tentacles (eyes at the base of sensory tentacles) -16

16a Operculum present; associated with marine marshes -Assiminea californica semi-marine 16b Operculum absent; living in, or amphibious near, freshwater - freshwater

17a Parietal denticle present -18 17b Parietal denticle absent -20

18a Periostracal hairs usually persistent in the adult; shell width to about 7 mm -Cryptomastix germana 18b Periostracal hairs not usually persistent in the adult; shell width greater than 12 mm -19

19a Shell width 12-26 mm; basal lip denticle or swelling (at its thickest point) located relatively closer to the columella than the periphery; southwest coastal British Columbia - 19b Shell width 12-19 mm; basal lip denticle or swelling (at its thickest point) located relatively closer to the periphery than to the columellar; in southeast British Columbia -Cryptomastix mullani

20a Umbilicus absent -21 20b Umbilicus open, although often slitlike or just a tiny pit -26

21a Shell width greater than 15 mm and usually with colour spiral bands, but sometimes without -22 21b Shell width less than 5 mm and without colour bands -23

22a Aperture height less than its width; unicolour or typically banded, with bands continuous, seldom interrupted by axial streaks - Cepaea nemoralis

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axial streaks - Helix aspersa

23a Aperture broadly and deeply rounded; whorls about 3 -Vitrina pellucida 23b Aperture narrowly crescent-shaped; whorls 3½ or more -24

24a Whorls with distinct axial grooves between the periphery and the suture in addition to very fine axial striae -Pristiloma stearnsii 24b Whorls with fine axial striae (and sometimes spiral striae) only -25

25a Shell waxy white or colourless; height of shell less than or equal to ½ of its width -Pristiloma johnsoni 25b Shell brownish; height of shell greater than ½ its width -Pristiloma lansingi

26a Shell width greater than or equal to 6 mm -27 26b Shell width less than 6 mm -47

27a Shell with colour bands -28 27b Shell without colour bands -31

28a Columellar lip expanded to almost close the umbilicus, leaving only a narrow slitlike opening -22 28b Columellar lip overhanging the umbilicus, but not enough to leave only a narrow slitlike opening -29

29a Shell brown with a narrow, pale, somewhat yellowish band at the periphery, a narrow, dark brown band above this and a dark brown base; shell rarely yellow with pale banding; animal with a reddish tinge; coastal British Columbia - fidelis 29b Shell greyish to brown, and banding not as above; animal not with reddish tinge; SE British Columbia -30

30a Shell dark brown, generally with a lighter peripheral band obscurely bordered by a darker band on either side; this light band visible within the aperture -Anguispira kochi 30b Shell generally greyish or brownish grey with two or more (often interrupted) dark bands (but band ing is highly variable and may be absent) and speckling; dark bands often visible within the aperture - Oreohelix strigosa & O. subrudis O. subrudis usually has a more elevated spire than O. strigosa, but identification by this shell charac ter can be unreliable.

31a Periostracum with minute hairs -32 31b Periostracum without hairs - 33

32a Lip flared, sometimes a little recurved, thin-edged - columbianus 32b Lip flared, strongly recurved, edge appearing thick due to the tight scroll-like folding back of the edge -Cryptomastix mullani

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Cross-section of the lip of Vespericola columbianus (top) and Cryptomastix mullani (bottom)

33a Shell with prominent, sharply defined regular axial ribs (at least on the upper surface of the whorls of the spire) -34 33b Shell without axial ribs, or axial ribs weak or not regular -37

34a Outer lip straightened or concave at shoulder in apertural view -35 34b Outer lip more or less convex at shoulder in apertural view -36

35a Beaded sculpture present on all whorls of the shell; width to 18 mm - sportella 35b Beaded sculpture obsolete on the penultimate and/or last whorl; width to 27 mm -Ancotrema hybridum

36a Axial ribs about as strong on the base as on the apical surface of the shell -Discus whitneyi 36b Axial ribs absent or significantly weaker on the base than on the apical surface of the shell -Discus shimekii

37a Lip strongly expanded and curved back to form a more or less broad, flattened face throughout its length -38 37b Lip thin or slightly thickened but not greatly expanded into a flat face; the lip of Cepaea nemoralis is somewhat expanded at the periphery but not elsewhere along its length -40

38a Sculpture of irregular, rounded axial ribs, strongest near the suture and usually disappearing below the periphery; width greater than 19 mm -39 38b Sculpture of fine axial striae only; width 9-17 mm -40

39a Shell width 21-31 mm; surface sculpture often malleated; apparently restricted in British Columbia to the vicinity of the Chilliwack Valley - 39b Shell width 19-24 mm; sculpture seldom malleated; southeast British Columbia -Allogona ptychophora

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40a Whorls about 3; shell pale, greenish, yellowish-green or nearly colourless -Vitrina pellucida 40b Whorls about 4-6; shell -coloured or pale brown, often paler around the umbilicus -41

41a Columellar lip expanded over to almost close the umbilicus, leaving only a narrow slitlike opening - Cepaea nemoralis 41b Columellar lip may overhang the umbilicus, but not enough to leave only a narrow slitlike opening -42

42a Axial sculpture rather rough and irregular and/or shell width usually greater than 16 mm -43 42b Shell smoothish; width less than or equal to 16 mm -44

43a Shell yellowish, olive green or darker; outer lip straightened at shoulder in apertural view - 43b Shell greyish white or occasionally more brownish; outer lip convex at shoulder -Oreohelix strigosa & O. subrudis O. subrudis usually has a more elevated spire than O. strigosa, but identification by this shell charac ter can be unreliable.

44a Umbilicus about 25% of the shell width; shell with very fine, faint spiral striae - nitidula Introduced to British Columbia (Forsyth et al. in prep.) 44b Umbilicus about 15% of the shell width; shell without spiral striae -45

45a Animal with a strong garlic odour when disturbed; shell to about 7 mm - alliarius Introduced to British Columbia (Forsyth in press) 45b Animal greyish, bluish-grey or nearly black, never with a garlic odour; shell greater than or equal to 7 mm -46

46a Shell to about 10 mm in width; last whorl regularly expanding; shell usually very glossy and with only very weak striae if any; animal pale grey with minute brown specks on the edge of the mantle (evident near the pneumostome on the right side, and on the left side through the shell) - Introduced to British Columbia (Forsyth in press) 46b Shell larger, to 12-16 mm in width; last whorl more rapidly expanded near the aperture so that it appears more than twice as wide as the previous whorl; shell usually less glossy and more striate than above species; animal dark bluish grey - Introduced to British Columbia (Forsyth in press)

47a Major axial sculpture of more or less regularly spaced grooves, with smooth, flattened, raised areas between (see figure below) -48 47b Shell smoothish or with axial sculpture not as above -49

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Cross-section (perpendicular to the axial grooves) of the shell wall of Nesovitrea showing characteristic grooves and flattened areas between.

48a Shell nearly colourless with a greenish tinge and smaller, 3.5-4.3 mm (3-4 whorls) -Nesovitrea binneyana 48b Shell amber-coloured and larger, 4.6-5.2 mm (3-4 whorls) -Nesovitrea electrina

49a Shell with widely and more or less regularly spaced (at least on the whorls of the spire), sharply angular or lamellar axial ribs -50 49b Shell smoothish or with axial sculpture closely and either regularly or irregularly spaced -55

50a Shell white, grey or yellowish grey -51 50b Shell yellowish brown, brown or reddish brown -53

51a Lip flared but thin -52 51b Lip at least slightly thickened -Vallonia gracilicosta

52a Shell width to about 2.2 mm -Vallonia cf. perspectiva 52b Shell width 2.5-3.4 mm -Vallonia cyclophorella

53a Spire almost flat - clappi 53b Spire moderately raised -54

54a Umbilicus about 20-25% of the width of the shell; protoconch initially smooth, then with rather widely spaced spiral striae -Paralaoma caputspinulae 54b Umbilicus larger, greater than 30% of the width of the shell; protoconch smooth -36

55a Lip thickened within by a callus -56 55b Lip not thickened within -57

56a Last whorl near the aperture distinctly increasing in width much more than previously; outline of the shell ovate; umbilicus elongate, elliptically spiral; outer lip gradually expanded, not at a right angle -Vallonia excentrica Introduced to British Columbia (Forsyth in press) 56b Last whorl near the aperture increasing in width only a little more than previously; outline of the shell almost circular; umbilicus regularly spiral; outer lip abruptly flared outward at a right angle Vallonia pulchella Introduced to British Columbia (Forsyth in press)

57a Whorls rapidly enlarging; umbilicus minute -Vitrina pellucida

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57b Whorls enlarging more slowly; umbilicus about 15-30% of the width of the shell -58

58a Shell whitish or colourless -59 58b Shell yellowish, yellowish brown, brown or reddish brown -63

59a Umbilicus about 15% of the shell width - contracta Introduced to British Columbia (Forsyth in press) 59b Umbilicus about 25-30% of the shell width -60

60a Axial sculpture of fine, close riblets -Striatura pugetensis 60b Axial sculpture more or less non-existent; fine incremental striae only -61

61a Umbilicus about 30% of the shell width; spire moderately elevated - minuscula 61b Umbilicus about 25% of the shell width; spire very low -62

62a Width 4-5 mm; 5-5 whorls; SE British Columbia -Microphysula ingersolli 62b Width 3.6-4.4 mm; 4-4 whorls; Coast and Cascade Mountains and Range - Microphysula cookei

63a Whorls rapidly enlarge in width: last whorl at least twice the width of the penultimate whorl one-half coil back from aperture -64 63b Whorls less rapidly enlarge in width: last whorl one-half coil back from aperture last whorl less than twice the width of the penultimate whorl -65

64a Axial sculpture of fine, close riblets; protoconch with minute, close spiral striae; shell width to 1.7 mm (2-3 whorls); width of umbilicus about 30% of the width of the shell -Striatura pugetensis 64b Axial sculpture weak and shell almost smooth; protoconch smooth; shell width greater than 5 mm (5- 6 whorls); width of umbilicus about 17% of the width of the shell - Introduced to British Columbia (Forsyth in press)

65a Shell width up to 1.4 mm or 1.8 mm (3-4 whorls) -P. californicum & Punctum randolphii P. californicum is generally slightly larger than P. randolphii. 65b Shell larger -66

66a Aperture narrow and crescent-shaped -67 66b Aperture broader, more rounded -68

67a Umbilicus tiny, pit-like -Euconulus fulvus 67b Umbilicus larger, about 10% of the width of the shell -Pristiloma chersinella

68a Animal bluish grey on tentacles and on back, lighter on sides and tail; shell width 5-6 mm (4 whorls); shell with faint and extremely fine, microscopic spiral striae; generally inhabiting drier areas than the next species -Zonitoides arboreus 68b Animal blackish; shell width 6-7 mm (4 whorls); microscopic spiral striae absent; generally inhabiting marshy areas -Zonitoides nitidus The spiral striae present on the shell surface of Z. arboreus are exceptionally fine. They should Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 33 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

be looked for, using reflective light and a minimum of 30 magnification, between the larger axial riblets; however, not all specimens show striae clearly, even at this magnification, and in some specimens striae are apparently absent. Pigmentation of the animal is the most reliable distinguishing feature and should be used whenever known. Zonitoides arboreus is characteris tically more depressed and with a wider umbilicus than Z. nitidus.

69a Animal with an exposed or partially exposed shell, either plate-like or ear-shaped -70 69b Animal without external shell -72

70a Shell ear-shaped, at the posterior end of the animal haliotidea Introduced to British Columbia (RBCM specimens). 70b Shell plate-like, near the longitudinal middle of the animal -71

71a Body immediately posterior to the visceral pouch depressed to receive the visceral mass, then pro duced into a high, compressed keel; caudal "horn" present Hemphillia glandulosa 71b Body posterior to the visceral pouch not depressed and not produced into a keel; horn absent Hemphillia species One or possibly two species in addition to H. glandulosa are present in British Columbia

72a Pneumostome at or behind the posterior 1/3 of the mantle on the right side (see figure) -73 72b Pneumostome at or in front of the midline of the mantle on the right side -77

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73a Slug very narrow and wormlike (length about 15 width when fully extended); mantle with a groove present on its right side Introduced to British Columbia (Reise et al. in press) 73b Slug not wormlike; mantle groove absent -74

74a Mantle smooth; caudal mucous pore conspicuous Ariolimax columbianus 74b Mantle with concentric wrinkle-like folds (somewhat like a finger print); caudal mucus pore absent -75

75a Mantle with distinct, dark lateral bands cf. valentiana Introduced to British Columbia (RBCM specimens) 75b Mantle unicolour, spotted or marbled, but without bands -76

76a In side view, tail tapering to a point; mantle with its posterior edge angular; concentric rings on the mantle centred on the midline Limax maximus 76b In side view, tail abruptly truncated; mantle with its posterior edge rounded; concentric rings on the mantle centred to the right of the midline, near the pneumostome - Deroceras species Four species are known to occur in British Columbia: Deroceras laeve; ; Deroceras reticulatum; and [=caruanae]. Most species are not easily differentiated by external characters.

77a Tail often having an oblique constriction which marks the site of self-amputation; caudal mucous pore absent -78 77b Tail never having an oblique constriction; caudal mucous pore present -80

78a Body with a conspicuous black band on either side running from the mantle to tip of the tail and enclosing a lighter, wedge-shaped central area which may enclose another darker median stripe Prophysaon vanattae 78b Body without black bands behind the mantle (but mantle may be banded) -79

79a Length to about 50 mm; mantle without yellow border; constriction usually marked by a grey line on the sole 79b Length 50-100 mm; mantle often with a narrow, bright yellow border; constriction usually not marked by a grey line on the sole Prophysaon foliolatum

80a Adult animal very large, greater than 70 mm in length when extended Arion rufus 80b Adult animal smaller, less than 70 mm in length -81

81a When animal contracted, tubercles have a prickly appearance; length of adult (fully extended) to about 15 mm 81b When contracted, tubercles without a prickly appearance; length of adult greater than 15 mm -82

82a Body with ground colour of reddish brown, rusty-orange or occasionally yellow; body mucus yellow or orange; slugs cannot contract into a hemispherical shape (in lateral view) 82b Body with ground colour of light greyish, brownish grey, dark grey or bluish-grey; body mucus

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colourless; slugs can contract into a hemispherical shape -83

83a Ground colour of light greyish or brownish grey; a pale line present on back, forming a "false keel"; body of contracted animal bell-shaped in cross-section (see figures below) Arion fasciatus species complex Species in this complex are not always reliably distinguishable by external characters alone. Dissection may be required. 83b Ground colour of dark dark grey or bluish grey (sometimes tinged with reddish); no "false keel"; body of contracted animal not bell-shaped in cross-section (see figure) species complex Species in this complex are not reliably distinguishable by external characters alone. Dissec tion is required.

Cross-section of body: Arion fasciatus species complex (left) and Arion hortensis species complex (right)

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Family CIONELLIDAE Kobelt, 1880

Genus Cionella Jeffreys, 1829

Cionella lubrica (Mller, 1774): Glossy Pillar . subcylindrica auctt.

Description: Shell small (height to 6.4 mm), subcylindrical, glossy, translucent and amber coloured; spire tall and tapering; apex bluntly rounded and a little paler in colour from the rest of the shell; whorls moder- ately convex and numbering 5.5; periphery rounded; suture deep and bordered by a more opaque band; protoconch smooth; teleoconch very glossy and smooth, with only very fine, weak axial striae; aperture subovate and without denticles; outer lip with an opaque pinkish or whitish riblike thickening within; umbili- cus absent.

Juveniles differ from adults in shape, have an angled periphery and lack the thickened rim inside the outer lip. Quick (1954) noted that the head, neck, sole and tail of specimens from England are dark grey or bluish-black.

Similar species: Adults are easily distinguished from all other land snails in British Columbia by their very glossy, cigar-shaped shells, and by the uniquely thickened outer lip.

Habitat: Cionella lubrica is a common snail in urban and agricultural areas and less frequent in natural areas. It lives in moist places under logs, rocks, leaf litter, living vegetation and miscellaneous debris. Caesar (1946) reported hundreds of individuals congregating on an old concrete house foundation at Centre, near Vernon.

Range: Holarctic. Point Barrow, Alaska to Mexico and east to Labrador and Washington, D.C. (Pilsbry 1948); Europe and North Africa east to Siberia, China and Japan (Dall 1905, Habe 1964, Kerney & Cameron 1979); Cape Verde Islands and the Azores (Backhuys 1975). Introduced to Madeira (Cameron & Cook 1996), New Zealand (Barker 1982), Venezuela (Adam 1960, Backhuys 1975) and South Africa (van Bruggen 1980).

Distribution: Widespread in British Columbia but more common in the south near human settlement. Although native, Cionella lubrica is often associated with and dispersed by human activity (Roth & Pearce 1984). The majority of the Columbia Basin records are from sites that are humanly disturbed.

Name: Genus name meaning a "little pillar". Species name meaning "slippery", which aptly describes the difficulty often encountered when picking up this snail.

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Records: Creston Valley Wildlife Management Area (4907.19'N, 11637.82'W) (RBCM 998-00282- 001); Creston Valley Wildlife Management Area (4907.34'N, 11637.24'W) (RBCM 998-00264-006); McDonalds Landing, West Arm, , near Six Mile, NE of Nelson (4934.75'N, 11712.99'W) (Forsyth Coll. 97-094-1081); E shore of Kootenay Lake, near boat launch S of ferry, (4940.4'N, 11652.3'W) (Forsyth Coll. 97-097-1095); (4955'N, 11655'W) (Staatliches Museum fur Naturkunde Grlitz); N end of Trout Lake, E of Upper Arrow Lake (5038.75'N, 11732.32'W) (RBCM 998-00285-002); along Hwy 1, at junction with Hwy 23, W of Revelstoke (510.2'N, 11813.4'W) (Forsyth Coll. 98-096-3318); Hunter Creek Forest Recreation Site, along Kicking Horse River, ESE of Golden (5115.95'N, 11644.69'W) (Forsyth Coll. 98-092-3296); Dogtooth Forest Service Road near bridge over Columbia River, Golden (5118.7'N, 11659.5'W) (Forsyth Coll. 98- 093-3056).

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Family VERTIGINIDAE Stimpson, 1851

The complex dentition of the aperture and unique features of the outer lip of some vertiginids are described using special terminology, which is adapted from Pilsbry (1948):

•Sinulus - The inwardly projecting point of the outer lip present in some species. It may be well- developed as a sharp, inwardly directed point or moderately developed and rounded, or weak or absent. •Crest - A raised, axial ridge on the last whorl behind the outer lip, typically set off from the outer lip by a constriction. •Denticles - Collectively, both lamellae and plicae.

The denticles are of two types, designated by their placement within in the aperture:

•Plica (pl. plicae) - A denticle located on the outer and basal lips. Also known as a fold. •Lamella (pl. lamellae) - A denticle located on either the columellar or parietal lips.

The four major denticles present in all our Vertigo, except some forms of V. modesta, are the columellar lamella (Lc), parietal lamella (Lp), upper palatal plica (Pu) and lower palatal plica (Pl). Additional denticles are often present. These are the angular lamella (La), infraparietal lamella (Li), suprapalatal plica (Ps), infrapalatal plica (Pi) and subcolumellar lamella (Ls). The relative positions of these denticles are illustrated below. Designation of the type and placement of denticles by a two-letter code is an innovation of this report and is used here for brevity.

Position of denticles in pupillids

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Genus Columella Westerlund, 1878

Columella columella (Martens, 1830): Mellow Column Synonym. Columella alticola (Ingersoll, 1875).

Description: Shell small (height 2.9 mm), cylindric and brown, with a silky lustre; spire very high; apex blunt; whorls convex and numbering 6-7; often either the last whorl or the penultimate whorl is a little smaller than the adjacent whorls; periphery rounded; suture deep; teleoconch coarsely striate (much more coarse and raised than in C. edentula); aperture ovate and without denticles; outer lip thin, unexpanded and without the sinulus and crest of many Vertigo; umbilicus small.

Similar Species: C. edentula is smaller, more tapering and with fewer whorls which are less coarsely sculptured. The peculiarity of coiling that reduces the size of the penultimate or last whorl in some individuals is never present in C. edentula.

Habitat: This is an arctic-alpine species usually collected at high elevations (Wayne 1959, Bequaert & Miller 1973). In the Columbia Basin region, specimens have been collected on wet ground under a log near the shore of Kootenay Lake and from under dead wood and fallen leaves in a spruce forest at Field.

Range: Holarctic (Forcart 1959). In North America, Aleutian Islands (Hanna 1925) and Port Clarence, Alaska (Bequaert & Miller 1973), British Columbia and Alberta, east to Churchill, Manitoba (Wayne 1959) and the northern coast of Ontario (Oughton 1948), and south in the United States in New Mexico, Arizona, Colorado, Wyoming, Kansas, Iowa, Illinois (Pilsbry 1948, Karlin 1961), and Washington (Branson 1977).

Distribution: Currently this species is only known from the southern Columbia Basin in British Columbia, but it may eventually be discovered elsewhere at higher elevations.

Notes: North American populations of this snail were long known as Columella alticola (Ingersoll, 1875), but Forcart (1959) combined C. columella and C. alticola into one species. He and Bequaert & Miller (1973) retained alticola as the North American.

Name: Genus and specific names meaning "a column."

Records: Tobacco Plains* (4902'N, 11506'W) (Whiteaves 1905; Berry 1922); E shore of Kootenay Lake, near boat launch S of ferry, Kootenay Bay (4940.4'N, 11652.3'W) (Forsyth Coll. 97-097-1094); along 2nd Avenue, 200 m SW of First Street East, Field (5123.75'N, 11629.3'W) (Forsyth Coll. 98-091- 3071); Field (5124'N, 11629'W) (Berry 1922, Pilsbry 1948).

*Whiteaves reported "Sphyradium edentulum" from Tobacco Plains; Berry later reidentified the speci- mens as Columella alticola.

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Columella edentula (Draparnaud, 1805): Toothless Column Synonym. Columella simplex Gould, 1841.

Description: Shell small (height 2.5 mm); subcylindric and a little tapering; brown, with a silky lustre; spire high; apex blunt; whorls convex and numbering about 5 or 6; last whorl always larger than the penultimate whorl; periphery rounded; suture deep; teleoconch with irregular, wrinklelike, rather low, axial striae; aperture ovate and without denticles; outer unthickened and unexpanded; sinulus and crest absent; umbilicus small.

Similar Species: C. columella. Small juveniles of C. edentula resemble Punctum in size and shape except for a more angular periphery and more rapidly enlarging whorls. The undeveloped shells of juvenile Vertigo may be difficult to separate from C. edentula.

Habitat:Under logs, dead wood, and on vegetation, and in leaf litter.

Range: Aleutian Islands, Alaska, east to Labrador (Dall 1905) south to New Jersey, Pennsylvania, Iowa, Montana, and Oregon; Alabama (Pilsbry 1948). In the Palearctic realm, widespread, from Iceland to Japan (Bequaert & Miller 1973).

Distribution: Common and widespread throughout the province.

Notes: North American populations are sometimes considered specifically distinct from C. edentula of Europe and are called Columella simplex Gould, 1841.

Name: Species name meaning toothless, likely named in contrast to Vertigo and other toothed members of the family.

Records: Summit Creek, Blazed Creek Rest Area, Hwy 3, Salmo-Creston Highway (4908.03'N, 11647.85'W) (RBCM 998-00267-002); Schroeder Creek, near highway 31, N of Kaslo (5001.95 N, 11654.55'W) (RBCM 998-00263-007); W of snow sheds, Glacier National Park (circa 5115.5'N, 11728'W) (Staatliches Museum fr Naturkunde Grlitz); Redgrave Rest Area, Hwy 1 between Roger's Pass and Golden (5129.54'N, 11716.18'W) (RBCM 998-00274-004).

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Genus Vertigo Mller, 1774 In some species of this genus, numerous form, varietal and subspecies names have been used for shells having additional or fewer denticles than typical, but the number of denticles (and often sculpture or strength of the crest) is assumed to be variable and form names and unqualified subspecies are not considered here. Named forms or 'subspecies', which are or could be applied to British Columbia material, are listed and defined under each species for reference. Identification is difficult in this genus, which is in need of revision. In place of a key, the most distinctive characters of the four species of Vertigo known from the Columbia Basin are presented in the following table (see the figure for denticle nomenclature and their relative posi- tions):

Vertigo Shape No. of Lp Lc La Li Ls Pu Pl Pi Ps Crest Sinulus denticles (Usual no.) elatior subovate 5-7 (5) + + ± - + + + - ± usually prominent weak gouldii subcylindric 4-6 (4-5) + + ± - ± + + - - absent/ weak/ weak moderate modesta subcylindric 4-4 (4) + + ± - - + + - - yes absent/very weak ovata subovate 7-9 (9) + + + + + + + ± ± yes prominent

Comparison of the four species of Vertigo known from the Columbia Basin, British Columbia.

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Redrawn after Pilsbry 1948

Vertigo elatior Sterki, 1894: Tapered Vertigo Synonym. Vertigo gouldii lagganensis Pilsbry, 1899.

Description: Shell small (height, 2.1 mm); subovate; spire with sides somewhat convex; whorls conves and numbering about 5; periphery rounded; suture deep; aperture irregularly subovate, with 5-7 (usually 5) denticles (Lp, Lc, Ls, Pu and Pl present; La occasionally present; Ps often present); sinulus prominent, turned inward to form a point; crest usually weak. (Specimens of this species have not been seen. The description is based on Pilsbry [1948].)

Similar Species: V. ovata is less conical in form and has additional denticles.

Habitat: Not reported for British Columbia.

Range: Ontario and Maine, west to Montana and Field, British Columbia; New Mexico (Pilsbry 1948).

Distribution: Known in British Columbia only from Field (Vanatta 1906; Pilsbry 1948).

Name: Genus name meaning "whorl". Species name meaning "high".

Record: Field (5124'N, 11629'W) (Vanatta 1906; Pilsbry 1948).

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Vertigo gouldii (Binney, 1843): Variable Vertigo Synonyms. coloradensis Cockerell, 1891; Vertigo columbiana utahensis Sterki in Pilsbry & Vanatta, 1900; V. coloradensis basidens Pilsbry & Vanatta, 1900; V. c. arizonensis Pilsbry & Vanatta, 1900.

Description: Shell small (height, 1.9 mm), subcylindric; reddish brown or brown; spire long, with straight or slightly convex sides; apex blunt; whorls 5, convex; periphery rounded; suture deep; teleoconch with sharp, close axial striae, espe- cially on the middle whorls; aperture subovate, with 4-6 (usually 4 or 5) denticles (Lp, Lc, Pl and Pu present La occasionally present, Ls occasionally present); outer lip slightly thickened and slightly flared; sinulus slight or moderate; crest absent or weak to rather prominent; umbilicus very small, slitlike, or closed.

Similar Species: Vertigo gouldii is smaller than V. modesta, with a stronger crest and sharper, more regularly striate.

Habitat: Under logs, rocks, , lichen and fallen leaves.

Range: Qubec, Ontario and Prince Edward Island to Missouri, Kentucky, Tennessee and Alabama; Colorado; Utah; New Mexico; Montana; British Columbia; Arizona; Kansas; Illinois (Pilsbry 1948). (Combined range of Vertigo gouldii and all its named subspecies or forms.)

Distribution: Widespread throughout mountainous regions of eastern and northern British Columbia, although previously recorded in the province only from the Rocky Mountains at Field (Vanatta 1906; Pilsbry 1948).

Notes: Doubtfully distinct western 'subspecies' of V. gouldii have been distinguished by the presence and absence of certain apertural denticles. Vertigo gouldii coloradensis (Cockerell, 1891) has 4 denticles (Lp, Lc, Pu, Pl) and a crest that is not well-developed. V. g. basidens Pilsbry & Vanatta, 1900 has 5 denticles (Lp, Lc, Ls, Pu, Pl), and a crest that is rather broad and sometimes prominent. V. g. arizonensis Pilsbry & Vanatta, 1900 has 5 denticles (Lp, Lc, La, Pu, Pl), and a crest that is very weak, and V. g. inserta Pilsbry 1919 has 6 denticles (Lp, La, Lc, Ls, Pu, Pl). Populations may consist of a single form or a mixture of forms.

Name: Species name in honour of Augustus Addison Gould (1805-1866), malacologist and Boston physician.

Records: Crowsnest Pass Rest Area (4939.06'N, 11441.76'W) (RBCM 998-00297-003); Hunter Creek Forest Recreation Site, along Kicking Horse R., ESE of Golden (5115.95'N, 11644.69'W) (Forsyth Coll. 98-092-3300); Along 2nd Avenue, 200 m SW of First Street East, Field (5123.75'N, 11629.30'W) (Forsyth Coll. 98-091-3074); Field (5124'N, 11629'W) (Vanatta 1906; Pilsbry 1948); Sherbrooke Trail head, N of Wapta Lake, Yoho National Park (circa 5126.4'N, 11621.5' W) (Forsyth Coll. 98-090-3387).

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Vertigo modesta (Say, 1824): Cross Vertigo Synonyms. Pupa decora Gould, 1848; Isthmia corpulenta Morse, 1865; Pupa corpulenta parietalis Ancey, 1887.

Description: Shell small (height, 2.5 mm), subcylindric, glossy; reddish brown or brown; spire long, with slightly convex sides; apex blunt; whorls 4-5, convex; periphery rounded; suture deep; teleoconch with fine, irregular, weak to strong axial striae, which are more distinct on the middle whorls; aperture subovate, regularly with 4-5 (typically 4) denticles (Lp, Lc, Pu and Pl regularly present; La sometimes present) or with various or all denticles absent; outer lip rather thickened, scarcely flared; sinulus absent or only a slightly flattened area on the outer lip; crest weak or moderately strong; umbilicus very small, slitlike or closed.

Similar Species: Some forms of V. gouldii with only 4 denticles or with 4 denticles and an angular lamella may be mistaken for V. modesta. V. gouldii is smaller, however, with stronger and more regular sculpture. A subcolumellar plica is never present in V. modesta.

Habitat: In British Columbia, V. modesta has been found living under rocks, leaves and moss.

Range: Holarctic: in North America from New England west to California and northward (Dall 1905); Aleutian Islands (Hanna 1925); in Europe, known from Norway, Finland, Sweden, Switzerland, Bavaria and Iceland (Kerney & Cameron 1979).

Distribution: Vertigo modesta is particularly widespread in the northwest and north-central interior of the province. It was reported from Field by Vanatta (1906).

Notes: Specimens bearing an angular lamella (as in figure above) have been given the name Vertigo modesta parietalis (Ancey, 1887). These often occur with typical V. modesta.

Name: Species name meaning "unassuming".

Records: Cummings Creek, N of Sparwood (4946.32'N, 11454.95'W) (RBCM 998-00293-005); Lake Revelstoke, km distant from Mica Dam (5103.81'N, 11833.42'W) (RBCM 998-00259-001); Field (5124'N, 11629'W) (Vanatta 1906; Pilsbry 1948); Redgrave Rest Area, Hwy 1 between Roger's Pass and Golden (5129.54'N, 11716.18'W) (RBCM 998-00274-003) .

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Vertigo ovata Say, 1822: Ovate Vertigo

Description: Shell small (length, 2.5 mm), subovate; somewhat translucent; rather glossy; reddish brown; spire rather short with sides very convex; apex obtuse; whorls 4, convex; periphery rounded; suture deep; teleoconch with fine axial striae; aperture irregularly subovate, with 7-9 (usually 9) denticles (Lp, Lc, La, Li, Ls, Pu, Pl present; Pi and Ps usually present); outer lip unthickened but flared; sinulus prominent, sharply turned inward to form a point; crest prominent; umbilicus slitlike or closed.

Similar Species: This species is similar to Vertigo binneyana Sterki, 1890, and V. elatior. V. ovata is easily recognised by the large number of denticles around the , its very much ovate form, and well-developed crest and sinulus.

Habitat: Marshes and other wet places along the edges of waterways under decaying vegetation, leaf litter and logs.

Range: Ungava Bay, northern Qubec, south to the Florida Keys and the West Indies; in the west from Kodiak and Tigalda Islands, Alaska, south to Utah and Arizona (Pilsbry 1948; Bequaert & Miller1973); perhaps in Sonora, Mexico (Naranjo-Garca 1991). It has been introduced to New Zealand (Barker1982).

Distribution: Widespread in the province, both inland and on the coast.

Name: Species name describing the egg-shaped shell.

Records: Creston Valley Wildlife Management Area (4907.19’N, 11637.82’W) (RBCM 998-00282- 003); Parson Bridge, Hwy. 95, S of Golden, near branch of Columbia River (5103.85’N, 11638.77’W) (RBCM 998-00273-002).

Juvenile pupillids not identified to species:

•Vertigo species: Schroeder Creek, near highway 31, N of Kaslo (5001.95’N, 11654.55’W) (RBCM 998-00263-006). •Vertigo species: Lodgepole Road at Kisoo Pass, SE of Fernie; (4917.03’N, 11442.72’W) (RBCM 998-00288-003) •Pupillid species: Meadow Creek near Duncan Dam, N end of Kootenay Lake (5013.97’N, 11657.37’W) (RBCM 998-00262-002).

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Family VALLONIIDAE Pilsbry 1900

Genus Vallonia Risso, 1826

Gerber (1996) has recently published a worldwide review of genus Vallonia.

Vallonia cyclophorella Sterki, 1892: Silky Vallonia

Description: Shell small (width, to about 2.9 mm), depressed-heliciform, thin-shelled, somewhat translu- cent and greyish or brownish yellow with a slight silky lustre; spire very low or a little more elevated; whorls convex and numbering about 3-3.5; periphery rounded; suture very deep; protoconch with indistinct spiral striae; teleoconch with regular and closely spaced bladelike axial ribs (approximately 40-55 on the last whorl), with finer axial striae between; aperture distinctly wider than high, ovate and without denticles; last whorl descending at the aperture; outer lip abruptly flared outward but not thickened within; umbilicus slightly elliptic and wide, about 33% or more of the width of the shell.

Similar Species: Vallonia pulchella and V. excentrica lack the regular and prominent ribbing of this species, and do not have the final portion of the last whorl descending in adults. Another ribbed species, V. gracilicosta Reinhardt, 1883, is similar, but in that species, the lip is thickened within by an opaque white rib.

Habitat: Living at altitudes of 1,400-3,250 m (Gerber 1996), under leaf litter and dead wood, sometimes sympatric with (Bequaert & Miller 1973; Roth & Reynolds 1990), and in British Columbia known from dry habitats. It was sympatric with V. gracilicosta in Kamloops, and with V. cf. perspectiva Sterki, 1893 near .

Range: Pyramid Island, Alaska, to British Columbia, east to Saskatchewan (Gerber 1996), and south to Arizona, New Mexico and Texas (Pilsbry 1948; Bequaert & Miller 1973; Hubricht 1985).

Distribution: British Columbia records are few and scattered: Skookumchuck (Gerber 1996); Ashnola River (Blood 1963); near Lytton; and Kamloops.

Name: Genus name said to honour the goddess of the valleys (Pilsbry 1948); species name meaning "bearing small circles", likely in reference to the ringlike axial ribs.

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Record: Skookumchuck (Gerber 1996), likely in reference to the locality by that name on the (4955'N, 11544'W).

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Vallonia excentrica Sterki, 1893: Excentric Vallonia

Description: Shell small (width, 2.2 mm), depressed-heliciform, somewhat translucent and glossy white; spire moderately low; whorls convex and numbering about 3; periphery rounded; suture deep; protoconch smooth; teleoconch with very fine irregular axial striae and occasional low wrinkles; last quarter of the last whorl expands rapidly and gives the shell a distinctive elliptic outline in apical view; last whorl not descend- ing at the aperture; aperture subcircular and without denticles; outer lip rather broad, flat and thickened within; lip flared gradually; umbilicus about 25% of the width of the shell and particularly elongatedly elliptic in outline.

Similar Species: Vallonia excentrica is closely similar to V. pulchella, to which it differs by the following: generally smaller size; broadly expanded last one-quarter of last whorl (seen in apical view); gradually flared outer lip; and more elliptic shape of the entire shell and umbilicus.

Habitat: Under rocks, logs, debris and in grass, leaf litter and other decaying vegetation, closely associated with human settlement. Although Vallonia excentrica is generally regarded to live in drier areas than V. pulchella (Kerney & Cameron 1979), both species often occur together.

Range: Holarctic. Europe, east to the Caucasians and the west coast of the Caspian Sea; Azores, Madeira and North Africa; eastern North America from southern Ontario east to Newfoundland and south in the United States to Mississippi, northern Georgia and Wyoming and Colorado. Introduced to the western United States and Canada, and many places worldwide (Gerber 1996).

Distribution: Introduced populations in populated and agricultural areas; probably synanthropic through- out the province; the most northern locality known in British Columbia is Smithers (Forsyth in press).

Name: Species name likely referring to the off-centre appearance of the umbilicus.

Records: McDonalds Landing, West Arm, Kootenay Lake, near Six Mile, NE of Nelson (4934.75'N, 11712.99'W) (Forsyth Coll. 97-094-1083; Forsyth in press); S of ferry, Kootenay Bay, E side of Kootenay Lake (4940.4'N, 11652.3'W) (RBCM 998-00118-001; Forsyth Coll. 97-097-1096; Forsyth in press). View the map.

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Vallonia pulchella (Mller, 1774): Lovely Vallonia

Description: Shell small (width, 2.5 mm); depressed-heliciform; somewhat translucent, somewhat glossy; white; spire moderately low; whorls 3.25, convex; periphery rounded; suture deep; protoconch smooth; teleoconch with very fine irregular axial striae and occasional low wrinkles; in apical view the whorls regu- larly spiral to the adult lip, thereby giving the shell a more or less circular outline; last whorl not descending at the aperture; aperture subcircular and without denticles; outer lip broad, flat, and heavily thickened in adults; lip flared abruptly; umbilicus medium-sized, about 25% of the width of the shell and not especially elliptic in outline.

Similar Species: Vallonia excentrica.

Habitat: Often abundant in damp conditions under logs, debris, rocks, decaying vegetation and in leaf litter or grass, in disturbed areas and in vacant lots of urban areas and in agricultural areas. Sometimes sympatric with Vallonia excentrica and Cionella lubrica.

Range: Holarctic. Europe, North Africa, the Canaries and Cape Verde Islands, east to through Asia (Gerber 1996), to Siberia (Dall 1905); eastern North America from Manitoba to Newfoundland and south into the eastern United States (Pilsbry 1948). Introduced to many places worldwide (Gerber 1996) and to the western U.S. and Canada: California, Oregon (Hanna 1966), Colorado, Utah, Texas (Pilsbry 1948), British Columbia and Washington.

Distribution: Populations of this introduced species appear limited to urban and agricultural areas, and are more common in the south. The most northern record for British Columbia is in Smithers (Forsyth in press). Like Vallonia excentrica, it is probably throughout the province in populated areas.

Name: Species name meaning "lovely".

Records: S of ferry, Kootenay Bay, E side of Kootenay Lake (4940.4'N, 11652.3'W) (RBCM 998- 00118-002; Forsyth Coll. 97-097-1565; Forsyth in press); Kaslo (4955'N, 11655'W) (Staatliches Mu- seum fr Naturkunde Grlitz); Dogtooth Forest Service Road near Columbia River crossing, Golden (5118.7'N, 11659.5'W) (Forsyth Coll. 98-093-3058; Forsyth in press). View the map.

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Genus Zoogenetes Morse, 1864 Zoogenetes harpa (Say, 1824): Boreal Top

Description: Shell small (length 3.3 mm), ovate-conic, higher than wide; subtranslucent and poorly calcified; dark brown; spire moderately el- evated; apex blunt; whorls convex and numbering about 4; periphery rounded; suture deep; protoconch with microscopic granulation; teleoconch with delicate, widely spaced, lamellar axial riblets (about 40 on the last whorl) and axial striae; axial riblets become more irregular and less defined on the last part of the last whorl; aperture ovate and without denticles; outer lip unthickened; umbilicus narrow and partially occluded by the columellar lip.

The body and head is greyish with the larger, occular tentacles darker, the foot is whitish, and the mantle is darker grey. The sides of the foot are crenulated (Binney & Bland 1869; Pilsbry 1948).

Similar Species: None.

Habitat: In British Columbia, Zoogenetes harpa has been found living in cottonwood and trembling aspen leaf litter, and under rocks and stones, and on plants.

Biology: Zoogenetes harpa is viviparous (Pilsbry 1948).

Range: Holarctic. Alaska east to Newfoundland and south to southern Colorado and Massachusetts (La Rocque 1953, Karlin 1961); Siberia, Kamtschatka and the Commander Islands west to Scandinavia, Finland and Switzerland (Likharev & Rammel'meier 1952, Kerney & Cameron 1979).

Distribution: Populations are sporadic in British Columbia where it is known from several locations in northern British Columbia and from the Columbia Basin.

Name: Genus name meaning "animal-birth", signifying live-born young (Pilsbry 1948). Species name meaning "a harp".

Records: Albert Canyon Road, near Albert Canyon Hotsprings Campground, E of Revelstoke (5107.94'N, 11751.61'W) (RBCM 998-00284-001).

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Family PUNCTIDAE Morse, 1864

Genus Punctum Morse, 1864

Punctum cf. californicum Pilsbry, 1898: Ribbed Spot

Description: Shell very small (width to about 1.8 mm), light chestnut brown and with a shining lustre; spire moderately elevated; whorls convex and about 3.75 in number; periphery rounded; suture deep; protoconch smooth; teleoconch with fine, closely spaced axial riblets (bearing very narrow lamellar edges) and with finer axial and spiral striae between; aperture rounded and without denticles; outer lip thin and unexpanded; umbilicus about 20% of the width of the shell.

Similar Species: Punctum californicum is very similar to P. randolphii (Dall, 1895) and is said to differ by being slightly larger and having a lower spire and less developed axial riblets (Pilsbry 1948), but see Notes below.

Another punctid not yet known from the Columbia Basin but likely to be discovered there is Paralaoma caputspinulae (Reeve, 1852); Punctum conspectum (Bland, 1865) is a synonym (see Roth 1985, 1987b). It is larger (width to 2.4 mm) than P. californicum,and the protoconch is sculptured with granules initially and then widely spaced spiral striae. The teleoconch has larger, more widely spaced axial riblets.

Habitat: In Arizona, Punctum californicum has been collected at elevations up to 2,743 m [9,000 ft] (Pilsbry 1948). Berry (1919) reported specimens collected from under sticks, bark, small logs and loose rock at elevations of 1,113-1,768 m [3,650-5,800 ft] in and adjacent to Glacier National Park, Montana. The British Columbia record is that of a dead shell collected from the foot of wet rock.

Range: Previously known from northern Montana and South Dakota, Colorado, Arizona and California (Pilsbry 1948), Punctum californicum is now also recorded from British Columbia for the first time.

Distribution: Punctum californicum is known in British Columbia only by a single, broken specimen (see Records).

Notes: Pilsbry (1948) noted that this species and Punctum randolphii may possibly be the same species. The coastal Punctum, considered to be P. randolphii, is a much more variable shell than credited by Pilsbry and in other literature.In the single, damaged specimen coming from the Kootenay region, and here provisionally referred to as P. californicum, the spire is as elevated as typical P. randolphii.

Name: Genus name meaning "a dot", in reference to the small size of this snail. Species named for Califor- nia.

Records: W of snow sheds, Hwy 1, Glacier National Park (circa 5115.5'N, 11728'W) (Staatliches Museum fr Naturkunke Grlitz p5959). View the map.

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Family DISCIDAE Thiele, 1931

Genus Anguispira Morse, 1864 Anguispira kochi (Pfeiffer,1845): Banded Tigersnail Synonym. Patula solitaria occidentalis Von Martens, 1892.

Description: Shell large (width to 25.3 mm), subheliciform; surface matt; opaque, dark brown, and usually with a lighter periphery band bordered by two obscure darker bands; spire moderately elevated; whorls about 6 and very convex; periphery rounded; suture deep; protoconch smoothish; early teleoconch with regular, low axial riblets becoming coarser and less regular on subsequent whorls, with fine axial striae; aperture deeply rounded; outer lip only very slightly thickened and then only in very mature individuals; no denticles within the aperture; base very convex; umbilicus medium-sized, about 20% of the width. Juveniles less than about half grown have a somewhat angular periphery. The animal is pale brownish or brownish-cream with a tinge of ochre; tentacles are darker, greyish-brown.

Similar species: Juveniles can be recognised from juvenile Allogona ptychophora by the larger umbilicus and less angular periphery. A. kochi differs from most specimens of Oreohelix by having a darker pigmented shell, but separation of eroded or long dead shells of A. kochi from some Oreohelix is often difficult.

Habitat: In British Columbia, it is known from moist, wooded, well-vegetated areas, often near the shores of lakes and streams.

Range: There are two disjunct populations: southeast British Columbia, south through Washington, Idaho and Montana to Oregon (Pilsbry 1948) east of the Cascade Mountains (Smith 1943); and Lake Erie south and west to Kentucky, Missouri and Arkansas (Pilsbry 1948).

Distribution: This species is restricted the southern Columbia-Kootenay River valleys in British Columbia.

Notes: The eastern and western populations were judged by Pilsbry (1948) to be subspecifically distinct. He called the western population Anguispira kochi occidentalis (Von Martens, 1882).

Name: Genus name meaning "snake-spire".

Records: Near Syringa Creek, Lower Arrow Lake (circa 4921.1'N, 11754.6'W) (Forsyth Coll. 97-113- 4241); Boswell, Kootenay Lake (4928.46'N, 11646.0'W) (Forsyth Coll. 97-100-863); Grohman Nar

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Genus Discus Fitzinger, 1833

Discus whitneyi (Newcomb, 1864): Forest Disc Synonyms. Helix cronkhitei Newcomb, 1865; H. striatella Anthony, 1840; cronkhitei anthonyi Pilsbry 1906.

Discus whitneyi

Description: Shell small (width, 6.7 mm), depressed-heliciform; subtranslucent brown or occasionally pale coloured; spire low; whorls 4.5, convex or a little angular (especially in juveniles); suture deep; protoconch without riblets; teleoconch with nearly equally spaced axial riblets, extending onto the base, and fine axial striae; aperture rounded and without denticles; outer lip unthickened; umbilicus rather large, about 33-40% of the width of the shell.

Animals are pale grey on the sides and tail and dark grey or blackish on the back, head and tentacles (Pilsbry 1948).

Similar species: Discus shimekii is similar, but the axial ribs are absent or weak on the base. Paralaoma caputspinulae (Punctum comspectum is a synonym) has the appearance of small D. whitneyi, but the umbilicus is narrower and the protoconch sculptured in the first species.

Habitat: This is an ubiquitous species living in wooded areas under leaf litter, under logs, and under vegeta- tion around the shores of lakes as well as in dry, exposed areas, at low and high elevations. In Montana this gastropod has been found at 3,048 m [10,000 ft] (Vanatta 1914).

Range: Attu Island, Aleutian Islands (Roth & Lindberg 1981), Alaska to Labrador, south to California, Arizona, New Mexico and Texas (Pilsbry 1948).

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Distribution: D. whitneyi occurs throughout most of the province, but is not common on the coast.

Notes: This widely spread and well-known species was previously called Discus cronkhitei, but Discus whitneyi predates that name (Roth 1987a). Dall (1905), Bequaert & Miller (1973) and Roth & Lindberg (1981) have alluded to the strong resemblance of this snail to D. ruderatus (Frussac, 1821) of the northern Europe and Asia.

Populations of this species are variable, with some shells having the periphery more angled than others; these have been given the name anthonyi.

Name: Species name after Professor Whitney, patron to the California Academy of Natural Sciences, San Francisco.

Records: Creston Valley Wildlife Management Area (4907.19'N, 11637.82'W) (RBCM 998-00282- 006); Creston Valley Wildlife Management Area (4907.34'N, 11637.24'W) (RBCM 998-00264-005); Wigwam River (4915'N, 11506'W) (Berry 1922); Lake (4922.3'N, 11551.0'W) (Forsyth Coll. 97- 102-1044); Coal Creek on E side of Cokato Road, Fernie (4929.72'N, 1153.75'W) (Forsyth Coll. 97- 108-1046); Crowsnest Pass Rest Area, Crowsnest Pass (4939.06'N, 11441.76'W) (RBCM 998-00297- 002); S of ferry, Kootenay Bay, E side of Kootenay Lake (4940.4'N, 11652.3'W) (Forsyth Coll. 97-097- 1097); along road to Cody Caves, hillside near Ainsworth (4945.7'N, 11654.7'W) (Forsyth Coll. 97-096- 4238); Cummings Creek, N of Sparwood (4946.32'N, 11454.95'W) (RBCM 998-00293-003); Schroeder Creek, near Hwy 31, north of Kaslo (5001.95'N, 11654.55'W) (RBCM 998-00263-003); N end of Trout Lake, E of Upper Arrow Lake; (5038.75'N, 11732.32'W) (RBCM 998-00285-005); Road- side, Hwy. 1, Eagle Pass, E of Clanwilliam, W of Revelstoke (5058.2'N, 11821.0'W) (Forsyth Coll. 98- 097-3282); Along Hwy 1, W of Revelstoke, B.C., at junction with Hwy 23 (5100.2'N, 11813.4'W) (Forsyth Coll. 98-096-3319); Sale Creek Road, Revelstoke (5109.1'N, 11811.0'W) (RBCM 998- 00260-003); Hwy 1, 4 km E of Canyon Hot Spring (circa 5109.5'N, 11749'W) (Staaliches Museum fr Naturkunde Grlitz); W of snow sheds, Hwy 1, Glacier National Park (circa 5115.5'N, 11728'W) (Staaliches Museum fr Naturkunde Grlitz); Hunter Creek Forest Recreation Site, along Kicking Horse R., ESE of Golden (5115.95'W, 11644.69'N) (Forsyth Coll. 98-092-3298); Field (5124'N, 11629'W) (Vanatta 1906); Sherbrooke Trail head, N of Wapta Lake, Yoho National Park (circa 5126.4'N, 11621.5'W) (Forsyth Coll. 98-090-3388); Doyle Rest Area, Hwy 1, 20 km N of Golden (5126.55'N, 11705.31'W) (RBCM 998-00275-004); unnamed creek, W of Brewster Creek (5138.32'N, 11832.32'W) (RBCM 998-00257-002); Lake Revelstoke, Hwy 37 to Mica Dam (5139.56'N, 11833.73'W) (RBCM 998-00258-002); Lake Revelstoke, Hwy 23 to Mica Dam (5139.56'N, 11833.73'W) (RBCM 998-00261-002).

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Discus shimekii (Pilsbry,1890): Striate Disc Synonyms. Pyramidula cockerelli Pilsbry, 1898; Zonitoides randolphi Pilsbry, 1898.

Description: Shell small (width to 6.5 mm), more or less depressed-heliciform, subtranslucent, yellowish brown; spire moderately elevated or flattened; whorls 4.5, convex; periphery rounded; protoconch smooth; teleoconch with regular, strong axial riblets with fine axial striae between; riblets rarely extending onto the base and becoming lower and irregular on the last whorl; aperture typically rounded, or more ovate in the flattened form of the species; aperture without denticles; outer lip unthickened; umbilicus rather large, about 30% of the width of the shell.

Similar Species: Compare with Discus whitneyi.

Habitat: Discus shimekii is a characteristically montane species living under rocks and dead wood. In Arizona it has been found at altitudes of 2,195-3,658 m [7,200-12,000 ft] (Bequaert & Miller 1973). Discus shimekii is sympatric with the more common and widespread D. whitneyi.

Range: Lindeman Lake, British Columbia (Pilsbry 1948, Forsyth 1999), east and south to Jasper (Platt 1980) and Kananaskis, Alberta (Berry 1922), south to New Mexico and Arizona (Pilsbry 1948, Bequaert & Miller 1973).

Distribution: Discus shimekii is not as widespread as D. whitneyi. In British Columbia, it is known from records at Lindeman Lake (Randolph 1899; Pilsbry 1948; Forsyth 1999) and near Atlin in the North- west and southeast along the Rocky Mountain range.

Notes: Somewhat smoother, more depressed shells with a wider umbilicus have been named Discus shimekii cockerelli (Pilsbry 1898). However, Beetle (1957) noted that a series of specimens from Wyo- ming showed a gradual transition from low to high spired individuals, and variation in the size of the umbili- cus and sculpture was also evident. Pilsbry (1948), and more recently Bequaert & Miller (1973), treated cockerelli as a synonym of D. shimekii.

Name: Genus name meaning "a disc". Species name honouring Bohumil Shimek who collected the speci- mens under Pilsbry's study (Pilsbry 1890). Shimek was a long-time friend of H.A. Pilsbry, with whom he began the study of shells as a boy (Pilsbry 1948).

Records: Cummings Creek, N of Sparwood (4946.32'N, 11454.95'W) (RBCM 998-00293-006).

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Family OREOHELICIDAE Pilsbry, 1939

Genus Oreohelix Pilsbry, 1904

There are two species known in British Columbia. Species and populations of Oreohelix have been differ- entiated by size, proportions, colour and sculpture; however, a great range of variation exists, perhaps due to subtle differences in ecology between individual populations (Solem 1975). Pilsbry (1939) grouped allied species and "subspecies" of Oreohelix. He noted that the shell ofO. strigosa has, in general, a taller, more conical form than the shell of O. subrudis and defined these groups using male reproductive morphology. The penis sac is divided into an internally muscular, ribbed part near the proximal end and a distal papillose section. In the O. strigosa group of species the muscular, ribbed portion of the penis is decidedly less than half of the entire length. In the O. subrudisgroup of species, the internally ribbed portion is half, or more than half, of the total length of the penis. Recently, however, Rees (1988) found that the morphology of the penis sac was highly variable and suggested that Pilsbry's methods of classifying individuals is unclear using both reproductive morphology and relative height of the shell. O. subrudis and O. strigosa may be geo- graphically segretated from each other.

Identification of Oreohelix into species is provisional and based wholly upon shell characters.

Oreohelix strigosa (Gould, 1846): Rocky Mountain Snail Synonyms. Helix cooperi W. G. Binney, 1858; O. strigosa canadica Berry, 1922.

Description: Shell medium-sized to large (width, 16-26 mm), variable in form, but typically depressed- heliciform, opaque, of rough texture; pale, greyish-white to dark brownish; often with darker bands of brown; bands variable, typically with 1 prominent band above and another just below the periphery; addi- tional narrower, fainter bands below the latter band on base; sometimes only the bands on the base show as faint traces; a pale brownish/yellowish periostracum is mostly eroded away; spire low-conical to more raised and extremely variable within populations; apex blunt and spire therefore somewhat domed; whorls convex and about 6 in number, with the periphery weakly to sharply angular (juveniles have angulate pe- riphery, a character that is retained in adults of some individuals or in some populations); suture deep; protoconch with fine, close, raised spiral striae crossed by more or less evenly spaced axial riblets; teleoconch with coarse, irregular axial riblets and striae; there may be faint traces of spiral sculpture in some; aperture ovate to rounded and without denticles; outer lip scarcely thickened or only slightly thick Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 71 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum ened within by a low ridgelike callus; lip slightly flared in most mature shells; umbilicus moderately open and dependent upon the amount that a shell is depressed, about 20-25% of the shell width.

Similar Species. See the comments above, for the genus.

Habitat: Vegetated rock slides. Rees (1988) indicated that Oreohelix subrudis and O. strigosa "are separated geographically: O. subrudis predominates in the streamside habitat, whereas O. strigosa occurs among the rock-slide rubble." Both Pilsbry (1939) and Branson (1980) remarked that this species has a markedly discontinuous range in Washington, and that it is restricted mostly the dry side of the mountains where it lives mostly under talus.

Biology: Members of the genus Oreohelix are viviparous.

Range: Cypress Hills, southeast Alberta; southeast British Columbia and adjacent parts of Alberta, south through Idaho, Montana, eastern Washington, eastern Oregon, Wyoming, South Dakota, Utah, Colorado, Arizona and New Mexico (Dall 1905, Pilsbry 1939, Russell 1951). This is the range of Oreohelix strigosa as a whole, including all named subspecies, forms and mutations.

Distribution: In British Columbia, Oreohelix strigosa is restricted to the Columbia Basin region and occurs as far north as Donald Station, the type locality for Oreohelix strigosa canadica Berry, 1922, and the northernmost locality for the genus.

Name: Genus name meaning "mountain snail"; species name meaning "lean".

Records: S end of Creston Valley Wildlife Management Area (4905.64'N, 11637.31'W) (RBCM 998- 00281-002); Creston Valley Wildlife Management Area (4907.34’N, 11637.24’W) (RBCM 998-00264- 002); Rest Area, Hwy 3A (4917.11'N, 11639.17'W) (Forsyth Coll. 97-101-1008); Lodgepole Creek Road to Flathead (4920.01'N, 11456.67'W) (RBCM 998-00296-003); Near Sand Creek, Betania Road, W of Galloway (4922.23'N, 11514.59'W) (Forsyth Coll. 97-105-908); Elk River valley, along Hwy 3, S of Ski Hill Road, S of Fernie (circa 4927.5'N, 11504.2'W) (Forsyth Coll. 97-109- 1017); Bull River bridge, Wardner-Fort Steele Rd., SE of Fort Steele (4928.39'N, 11526.95'W) (Forsyth Coll. 97-110-906); Boswell, Kootenay Lake, B.C. (4928.46'N, 11646.0'W) (Forsyth Coll. 97-100-870); Kokanee Landing (4936'N, 11706'W) (RBCM 996-00010-001, 996-00010-004); Kokanee Creek Park, West Arm, Kootenay Lake (circa 4936.2'N, 11707.7'W) (Forsyth Coll. 97-095-1088); Schroeder Creek near Hwy 31, N of Kaslo (5001.95'N, 11654.55'W) (RBCM 998-00263-001); Donald Station (5129.5'N, 11710.6'W) (Berry 1922, Pilsbry 1939).

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Oreohelix subrudis (Reeve, 1854): Subalpine Mountainsnail Synonyms. Helix limitaris Dawson, 1875; Helix cooperi auctt., in part; O. c. apiarium Berry, 1919.

Oreohelix subrudis

Description: Shell medium-sized to large (width, 16-23 mm), variable in form, but typically heliciform or almost bee-hive-shaped, opaque, rough textured; pale, greyish-white to dark brownish; often with darker bands of brown variable as in Oreohelix strigosa; periostracum pale brownish/yellowish, mostly eroded off; spire moderately raised in typical shells to distinctly raised but extremely variable and often complexingly similar to O. strigosa; apex blunt; spire rather domed; whorls about 6, convex and with the periphery weakly angular (juveniles have a distinctly angulate periphery; suture deep; protoconch with fine, close, raised spiral striae crossed by more or less evenly spaced axial riblets; teleoconch with coarse, irregular axial riblets and striae; there may be traces of spiral sculpture in some; aperture ovate to rounded and without denticles; outer lip scarcely thickened or only slightly thickened within by a low, ridgelike callus; lip slightly flared in most mature shells; umbilicus about 20-25% of the shell width.

Similar Species: See the comments above, for the genus.

Habitat: Along streams and in swamps.

Range: Cypress Hills, Saskatchewan-Alberta (Russell 1951); Southeast British Columbia and southwest Alberta through Idaho, Montana, Wyoming, Utah, Colorado, Arizona and New Mexico Pilsbry (1939).

Distribution: In British Columbia, this species is limited to the Columbia Basin.

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Name: Species name meaning "somewhat rough".

Records:Wigwam River (circa 4915'N, 11506'W) (Berry 1922); Moyie Lake (4920'N, 11550'W) (Berry 1922); Coal Creek valley near Fernie (circa 4930'N, 11504'W) (Pilsbry 1939); near Hosmer, Hwy 3 between Fernie and Sparwood (4934.94'N, 11458.05'W) (RBCM 998-00299-001); Crowsnest Pass (4938'N, 11504'W) (Pilsbry, 1939); Crowsnest Pass Rest Area ( 4939.06'N, 11441.76'W) (RBCM 998- 00297-001).

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Family ARIONIDAE Gray in Turton, 1840

Genus Arion Frussac, 1821

There are at least six introduced European slugs of the genus Arion in British Columbia.

Arion rufus (Linnaeus, 1758): Chocolate Arion Synonym. Arion ater auctt., in part.

Description: A large slug (extended length to about 180 mm); brown, reddish-brown, greenish brown or black; also yellow or orange; juveniles show a broader range of colour than adults; lateral band usually absent in adults but present in juveniles; no external shell; mantle with a granular surface; pneumostome in front of the mid-point of the mantle; tubercles on the back large, coarse and elongate; keel absent; foot fringe red, orange, yellow or black; sole of foot whitish or greyish white, all black or with broad black bands on either side of a light central area; caudal mucus pore present, located just above the tip of the tail; mucous colourless.

When contracted, the animal is bell-shaped in cross-section and has a tendency to rock side to side when disturbed.

Similar species: The Arion ater complex is comprised of two species, A. ater (Linnaeus, 1758) and A. rufus, which can only be distinguished by reproductive anatomy. In A. ater, the atrium is generally more slender with its proximal portion smaller and shorter than the distal part. In A. rufus, the atrium is less slender, with the proximal portion larger and much wider than the distal portion. Colour of the animal is not reliable for determination of these two species.

Other species of Arion could be confused for young A. rufus. Refer to the key, or Kerney & Cameron (1979) for further figures and descriptions of other species.

Habitat: Gardens, fields, other disturbed sites and wooded areas near human activity.

Range: Western and central Europe (Kerney & Cameron 1979).

Distribution: This slug is widespread in southern British Columbia. Rollo & Wellington (1975) recorded Arion "ater" (sensu lato) throughout the lower and , but did not adequately differentiate between A. rufus and A. ater. A. rufus is con- firmed by dissection from , Greater Vancouver, near Duncan (H. Reise, pers. comm.) and from the Kootenay localities given below.

Name: Genus named after a Greek poet and musician. Species name meaning "red" or "reddish".

Records: Box Lake, SE of (5012.4'N, 12342.7'W) (RBCM 998-00265-008 ); Blanket Creek Provincial Park, S of Revelstoke (5050'N, 11805'W) (Staatliches Museum fr Naturkunde Grlitz).

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Arion species [subgenus Carinarion - Arion fasciatus group]

Records: Blanket Creek Provincial Park, S of Revelstoke (5050'N, 11805'W) (Staatliches Museum fr Naturkunde Grlitz p5961).

Genus Prophysaon Bland & Binney, 1873

Prophysaon andersoni (J.G. Cooper, 1872): Reticulate Taildropper Synonyms. Prophysaon hemphilli Bland & Binney, 1873; P. pacificum Cockerell, 1890; P. flavum Cockerell, 1890; P. andersoni pallidum Cockerell, 1891; P. a. marmoratum Cockerell, 1892; P. a. suffusum Cockerell, 1893.

Prophysaon andersoni

Description: A medium-sized slug (extended length to about 60 mm), pale brownish, reddish grey or yellowish, somewhat clouded with darker tones and with a dia- mond mesh pattern on the back; the mantle is often paler, usually with a pair of dark lateral bands; shell internal and oblong; head pale brown, tentacles darker; mantle with a granular surface; pneumostome close to the middle (or anterior to the middle when the animal is fully extended); keel absent; foot fringe pale; caudal mucus pore absent; body mucus lemon yellow to orangish when disturbed (Kozloff 1976).

Similar Species: Prophysaon vanattae known in British Columbia from southwestern Vancouver Island (pers. observ.; Cameron 1986) has black bands on the mantle, tail and body, and has orangish pigment; it also differs significantly in its anatomy (Pilsbry 1948). There are six additional species of Prophysaon not yet reported from British Columbia.

Habitat: Prophysaon andersoni occurs at all elevations, usually in woods, but sometimes in gardens and other unnatural sites, and mostly in damp conditions. In Manning Park, B.C. it is recorded from 1,829 m [6,000 ft.] (Carl et al. 1952). It lives under logs, leaf litter, rocks and plants, or may be seen crawling out in the open during wet weather.

Biology: This slug is capable of self-amputation and regeneration of the tail, presumably as a defence against predators (Hand & Ingram 1950).The diagonal constriction on the tail marks the site where ampu- tation occurs.

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This species was observed in Vancouver by Rollo & Wellington (1975); in the spring only juveniles were found, but by late summer and fall, all individuals were large. It may be an annual species.

Range: Aleutian Islands (Hanna 1925), Alaska to California and east to Idaho (La Rocque 1953).

Distribution: Probably widespread throughout B.C.; a common species.

Name: Genus name meaning "forward breathing" and describing the placement of the pneumostome in front of the midline of the mantle. Species name honours Dr C.L. Anderson of Santa Cruz, California (Pilsbry 1948).

Records: Creston Valley Wildlife Management Area (4907.34'N, 11637.24'W) (RBCM 998-00264- 003); Glenmarry Creek Road, near Nancy Greene Provincial Park (4915.94'N, 11756.36'W) (RBCM 998-00280-003); W side of Slocan River, just S of (4935.66'N, 11735.47'W) (RBCM 998- 00279-003); N end of , near hills (4954.04'N, 11723.25'W) (RBCM 998-00266-006); S end of Trout Lake near bridge over Lardeau, E of Upper Arrow Lake (5030.52'N, 11716.08'W) (RBCM 998-00286-001); Blanket Creek Provincial Park, S of Revelstoke (5050'N, 11805'W) (Staatliches Museum fr Naturkunde Grlitz); Albert Canyon Road, near Albert Canyon Hotsprings Campground, E of Revelstoke (5107.94'N, 11751.61'W) (RBCM 998-00284-004); Doyle Rest Area, Hwy. 1, 20 km N of Golden (5126.55'N, 11705.31'W) (RBCM 998-00275-002).

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Genus Hemphillia Bland & Binney, 1872

Branson (1975) presented a key to the seven named species of Hemphillia.

Hemphillia species

A species of Hemphillia has been collected from the Columbia Basin and is currently being studied by Heike Reise (Staatliches Museum fr Naturkunde Grlitz, Germany). Slug pale yellowish-grey closely speck- led with darker pigment on the body, especially on the posterior half; on the mantle a pair of lateral bands are formed by coalescing of the speckles; shell partially external, platelike; mantle smoothish, without papillae; pneumostome behind the mid-point of the mantle; body not depressed to receive the dorsal visceral pouch; keel not developed; caudal "horn" absent. Hemphillia camelus

Similar species: The coastal species, Hemphillia glandulosa, is darker in colour and with a distinctly papillose mantle in life (becoming smoother in alcohol). Additionally, the body of H. glandulosa is strongly depressed to receive the visceral pouch (compare figures 396c, d and 398d of Pilsbry, 1948, pages 741 and 743 respectively) and the tail has a "horn". The Columbia Basin specimens externally resemble H. camelus Pilsbry & Vanatta, 1890 and to a lesser extent Hemphillia danielsi Vanatta, 1914. Critical distinctions between these and other Hemphillia rely on reproductive anatomy (see Pilsbry 1948, Branson 1975).

Habitat: In the Kootenays, this slug has been collected from around mossy stumps and rocks, under logs and in leaf litter in forests.

Distribution: In British Columbia, this Hemphillia species is known only from the Columbia Basin region. Carl & Hardy (1945) reported H. camelus from Paradise Mine, west of Windermere; this likely is the same.

Name: The genus name honours Henry Hemphill (1830-1914), early Californian malacologist (Coan & Roth 1987).

Records: Creek Marsh, S end of Creston Valley Wildlife Management Area; Creek Marsh (4905.64'N, 11637.31'W) (RBCM 998-00281-003); Summit Creek, Blazed Creek Rest Area, Salmo-Creston Hwy.; Hwy 3 (4908.03'N 11647.85'W) (RBCM 998-00267-001); Kid Creek, at bridge; dirt track from Cranbrook to Creston, Moyie Creek Campsite (4914.58'N, 11608.92'W) (RBCM 998-00287-001); W of snow sheds, Hwy 1, Glacier National Park (circa 5115.5'N, 11728'W) (Staatliches Museum fr Naturkunde Grlitz); Redgrave Rest Area, Hwy 1, between Roger's Pass and Golden (5129.54'N, 11716.18'W) (RBCM 998-00274-001).

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Family SUCCINEIDAE Beck, 1837

The shell of succineids offer little indication of generic and specific identity, and identification is difficult. Genera are differentiated anatomically and many nominal species are poorly understood. Patterson (1971), and more recently Hoagland & Davis (1987), have partially revised the classification of the succineids.

Various succineid shells. After Pilsbry (1948)

Genus Catinella Pease, 1871 Shell generally short with a relatively large, conical spire; aperture rounded-ovate. Catinella is anatomically defined by the absence of a penial sheath and a distinctly differentiated epiphallus; the penis also has a prominent appendix midway on its length (Pilsbry 1948). Quickella Boettger, 1939 is a synonym (Patterson 1971). One species has been reported in British Columbia: Catinella avara (Say, 1824); however, Succinea avara is a nomen dubiumand the name Catinella vermeta (Say, 1829) has been applied to some populations (Patterson & Burch 1966; Hoagland & Davis 1987).

Genus Oxyloma Westerlund, 1885 Shell elongate-ovate generally with whorls elongate and somewhat flattened between the periphery and the suture; spire usually short; aperture more or less elongate-ovate and rather flared posteriorly. The genus is anatomically differentiated by the presence of an epiphallus and penial sheath and with the penis not having an appendix (Pilsbry 1948). Two or three species have been reported for British Columbia: Oxyloma hawkinsii Baird, 1863; Oxyloma nuttallianum (I. Lea, 1841); and (I. Lea, 1834).

Genus Succinea Draparnaud, 1801 Shell generally short to elongate-ovate with a small to moderate spire; aperture rounded to elongate-ovate. An epiphallus and penial shealth are present, and the penis also has a small, rather narrow appendix en- closed in the penial sheath (Pilsbry 1948). One species has been reported from British Columbia: Succinea rusticana Gould, 1846.

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Family EUCONULIDAE H.B. Baker, 1928

Genus Euconulus Reinhardt, 1883

Euconulus fulvus (Mller, 1774): Brown Hive Synonym. Conulus fulvus alaskensis Pilsbry, 1899.

Description: Shell small (width, 3.2 mm), subconic-heliciform, width only a little greater than height; surface with a silky, shining texture; translucent brown; spire moderately elevated; apex obtuse; whorls slightly convex and numbering 5.5; periphery rounded or a little angular; suture deep; protoconch smooth; teleoconch with fine, close axial striae and exceedingly fine spiral striae; axial striae obsolete and spiral striae more apparent on the base; aperture narrowly crescent-shaped and without denticles; outer lip unthickened; umbilicus closed or almost so.

The body is greyish with darker tentacles and the mantle has large, black blotches. The foot is very long and slender (Pilsbry 1946).

Similar Species: Compare also with similarly shaped species of Pristiloma, which have shells with pro- portionally lower spires and different sculpture.

Habitat: Common and widespread, living under logs and debris in dead grass and leaf litter in a variety of habitats, both dry and moist areas at all elevations. Reported up to 3,048 m [10,000 ft] in Montana (Vanatta 1914).

Range: Holarctic. Throughout Europe, Asia, North Africa, North America, in the Azores (Backhuys 1975) and Madeira (Cameron & Cooke 1996). In North America it was recorded throughout most of the U.S. by Hubricht (1985) and known as far south as northern Mexico (Bequaert & Miller 1973; Naranjo- Garca 1991).

Distribution: Throughout British Columbia, and one of the most generally spread species.

Notes: The form E. fulvus alaskensis was said to differ from typical fulvus by having one less whorl in shells of similar size, and by minor sculptural differences (Pilsbry 1910, 1946), but Henderson (1936) and Berry (1937) expressed difficulties with the distinctiveness of alaskensis. Branson et al. (1966)

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Name: Genus name in reference to the cone-shaped shell. Species name meaning "tawny" or "reddish yellow".

Records: Creston Valley Wildlife Management Area (4907.19'N, 11637.82'W) (RBCM 998-00282- 005); Creston Valley Wildlife Management Area (4907.34'N, 11637.24'W) (RBCM 998-00264-008); Lodgepole Road at Kisoo Pass, SE of Fernie (4917.03'N, 11442.72'W) (RBCM 998-00288-002); near Weaver Creek, along logging road following Moyie River, S of Cranbrook (4924.50'N, 11559.95'W) (RBCM 998-00298-001); Grohman Narrows Provincial Park, near Nelson (4929.8'N, 11720.8'W) (Forsyth Coll. 97-112-3002); Rampart Rest Area, Hwy 3/93, NW of Wardner (4932.91'N, 11538.67'W) (Forsyth Coll. 97-104-1041); Kokanee Creek Park, West Arm, Kootenay Lake (4936.2'N, 1177.7'W) (Forsyth Coll. 97-095-1870); Cummings Creek, N of Sparwood (4946.32'N, 11454.95'W) (RBCM 998-00293-004); N end of Trout Lake, E of Upper Arrow Lake (5038.75'N, 11732.32'W) (RBCM 998-00285-004); Schroeder Creek, near Hwy 31, N of Kaslo (5001.95'N, 11654.55'W) (RBCM 998-00263-004); Hwy. 1, Eagle Pass, E of Clanwilliam, W of Revelstoke (5058.2'N, 11821.0'W) (Forsyth Coll. 98-097-3283); Parson Bridge, Hwy 95, S of Golden (5103.85'N, 11638.77'W) (998-00273-003); Hwy 1, 4 km E of Canyon Hot Spring (circa 5109.5'N, 11749'W) (Staaliches Museum fur Naturkunde Grlitz); Hunter Creek Forest Recreation Site, along Kicking Horse R., ESE of Golden (5115.95'N, 11644.69'W) (Forsyth Coll. 98-092-3299); Dogtooth Forest Service Rd near bridge over Columbia River, Golden (5118.7'N, 11659.5'W) (Forsyth Coll. 98-093-3059); Along 2nd Avenue, 200 m SW of First Street East, Field (5123.75'N, 11629.3'W) (Forsyth Coll. 98-091-3072); Field (5124'N, 11629'W) (Vanatta 1906); Sherbrooke Trail head, N of Wapta Lake, Yoho National Park (circa 5126.4'N, 11621.5'W) (Forsyth Coll. 98-090-3390); Redgrave Rest Area, Hwy 1 between Rog- er's Pass and Golden (5129.54'N, 11716.18'W) (RBCM 998-00274-002); Waitabit Creek Forest Rec- reation Site, near Donald Station (5130.2'N, 11711.11'W) (Forsyth Coll. 98-094-3309); Bluewater Creek Forest Recreation Site, near Donald Station (5131.84'N, 11715.27'W) (Forsyth Coll. 98-095-3150); Lake Revelstoke, Hwy 37 to Mica Dam (5139.56'N, 11833.73'W) (RBCM 998-00258-001); Lake Revelstoke, Hwy 23 to Mica Dam (5139.56'N, 11833.73'W) (RBCM 998-00261-003).

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Family ZONITIDAE Mrch, 1864

Genus Nesovitrea C.M. Cooke, 1921

Nesovitrea binneyana (E.S. Morse, 1864): Blue Glass Synonym. binneyana occidentalis H. B. Baker, 1930.

Description: Shell small (width to about 3.7 mm), depressed-heliciform, thin, translucent, glossy and almost colourless with a greenish or greenish-yellow tinge; spire low, convex; whorls scarcely convex on the spire and numbering about 3.5; whorls rapidly enlarging in width; periphery rounded; suture deep and appearing to be bordered by an opaque band; protoconch smooth; teleoconch with rather regularly spaced axial riblets with shallow, equally sized grooves between and with exceedingly fine (and sometimes obso- lete), closely spaced spiral striae visible in some shells at 50 magnification; aperture ovate, oblique and without denticles; outer lip unthickened; umbilicus 14-19% of the width of the shell.

Similar Species: The shell is smaller and more greenish-yellow in colour than Nesovitrea electrina and with exceedingly faint spiral striae that may sometimes be evident with magnification.

Habitat: Leaf litter and under rocks and logs.

Range: British Columbia and Alberta; Montana, Colorado and California; Ontario, Maine, New York, Pennsylvania, Ohio and Michigan (Pilsbry 1946).

Distribution: Widespread, both inland and on the coast.

Notes: A western 'subspecies', Nesovitrea binneyana occidentalis, is said to differ from the typical N. binneyana by having stronger spiral striae (Pilsbry 1946).

Name: "Neso" meaning island, referring to the Hawaiian Islands from where the type species of the genus originates, and "Vitrea", a European genus of snails. Species name in honour of William G. Binney, American malacologist (1833-1909).

Records: Betania Road, near Sand Creek, W of Galloway (4922.23'N, 11514.59'W) (Forsyth Coll. 97- 105-860); Kaslo (4955'N, 11655'W) (Staatliches Museum fr Naturkunde Grlitz); Albert Canyon Road, near Albert Canyon Hotsprings Campground, E of Revelstoke (5107.94'N, 11751.61'W) (RBCM 998- 00284-002); Hwy. 1, 4 km E of Canyon Hot Spring (circa 5109.5'N, 11749'W) (Staatliches Museum fr

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Naturkunde Grlitz); Field (5124'N, 11629'W) (Vanatta 1906). "East Kootenay District" (Baker 1930; Pilsbry 1946).

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Nesovitrea electrina (Gould, 1841): Amber Glass Synonym. Retinella hammonis auctt.

Nesovitrea electrina

Description: Shell small (width, to 4.6 mm), de- pressed-heliciform; very thin, translucent, glossy; cor- neous or brown; spire low, convex; whorls slightly convex and numbering about 3.25-3.75; whorls rapidly enlarging in width; periphery rounded; suture moderately deep; protoconch smooth; teleoconch with sharply incised, deep axial grooves that are often more or less regularly spaced; between these major grooves are fine axial striae; base smoother; aperture nearly round or ovate, oblique and without denticles; outer lip not thick- ened; umbilicus 16% of the width of the shell. The animal is darkly pigmented. The head, back and tenta- cles are almost black, the edges of the foot and the sole is very dark grey (Pilsbry 1946).

Similar Species: Nesovitrea binneyana.

Habitat: Under rocks and dead wood and in leaf litter. Reported at 3,048 m [10,000 ft] elevation in Montana by Vanatta (1914).

Range: Kodiak Island, Alaska, to Labrador and south to Arizona (Pilsbry 1946).

Distribution: Widespread in British Columbia.

Notes: Some authors have considered N. electrina to be the same as the Eurasian N. hammonis (Strm, 1765). Bequaert & Miller (1973) accepted electrina as a North American subspecies of N. hammonis.

Name: Species name meaning "amber", describing the translucency and colour of the shell.

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Records: Creston Valley Interpretation Centre; opposite side of marsh (4907`34 N, 11637`24) (RBCM 998-00264-009); W side of Slocan River, just S of Winlaw (4935.66'N, 11735.47'W) (RBCM 998-00279-005); North end of Slocan Lake (4954.04'N, 11723.25'W) (RBCM 998-00266-004); Kaslo (4955'N, 11655'W) (Staatliches Museum fr Naturkunde Grlitz); Columbia Lake, adjacent to Canal Flats Park (5010.65'N, 11549.03'W) (RBCM 998-00294-003); Dogtooth Forest Service Rd. near bridge over Columbia River, Golden (5118.7'N, 11659.5'W) (Forsyth Coll. 98-093-3060); Field (5124'N, 11629'W) (Vanatta 1906); Bluewater Creek Forest Recreation Site, near Donald Station (5131.84'N, 11715.27'W) (Forsyth Coll. 98-095-3151); Lake Revelstoke, Hwy 37 to Mica Dam (5139.56'N, 11833.73'W) (RBCM 998-00258-005); unnamed creek, W of Brewster Creek (5138.32'N, 11832.32'W) (RBCM 998-00257-003). "East Kootenay District" (Baker 1930; Pilsbry 1946).

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Genus Pristiloma Ancey, 1887 Pristiloma chersinella (Dall, 1866): Black-foot Tightcoil

Description: Shell small (width, 3.3 mm), depressed-heliciform, glossy, slightly translucent, pale yellowish; spire moderately raised; whorls convex and numbering 4.5-5; periphery rounded; suture moderately deep; teleoconch with fine axial striae and exceedingly minute spiral striae (at high magnification); aperture narrowly crescent-shaped and without denticles; outer lip unthickened and not expanded; umbilicus small, about 10% of the width of the shell. The animal has black pigmentation on the foot and edge of the mantle (Baker 1931). In life, the shell appears dark with the animal showing through.

Similar Species: There are no other species of umbilicate Pristiloma known from British Columbia. Euconulus fulvus is similar in form but is brown rather than yellowish and the umbilicus is proportionately smaller.

Habitat: P. chersinella has been reported from northern Montana at altitudes of 3859-6000 ft. [1176- 1828 m], under sticks, bark and small logs (Berry 1919). In the Babine Range, Mountains, near Smithers, this species has been found under rocks, dead wood and moss, at and below the tree line at altitudes of 1158-1524 m, in old slide areas, melt-water run-off areas and spruce forests. At an altitude of 1740 m in the Kisoo Pass, British Columbia, P. chersinella was collected from under stones.

Range: Oregon and California (Pilsbry 1946) and British Columbia.

Distribution: In British Columbia, presently known only from the Babine Mountains, near Smithers, and Kisoo Pass. These records represent a significant range extension north from Oregon.

Name: Genus name meaning "sawed edge" and referring to the denticulate lip rib often present in Pristiloma lansingi. Species name the diminutive of chersina, meaning "dry land".

Notes: The correct date of this name is 1866, not 1886 as generally given.

Records: Lodgepole Road at Kisoo Pass, SE of Fernie (4917.03'N, 11442.72'W) (RBCM 998-00288- 001).

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Genus Zonitoides Lehmann, 1862 Zonitoides arboreus (Say, 1816): Quick Gloss

Description: Shell small (width to 5.6 mm), depressed-heliciform; translucent brown, somewhat glossy; spire convex; whorls moderately convex and numbering 4-4.5; periphery rounded; suture moderately deep; protoconch of 1 smooth whorls; teleoconch with irregularly placed, weak growth wrinkles and extremely fine, close spiral striae; base smoother; aperture ovate and without denticles; outer lip unthickened and not expanded; umbilicus about 16-25% of the width of the shell. The animal is bluish grey on the tentacles and back and lighter on the sides and tail.

Similar Species: Z. nitidus is larger, with a more elevated spire. It also lacks the exceeding fine spiral striae which are usually evident under magnification in Z. arboreus, and the colour of the animals' bodies are markedly different. The shells of introduced European snails of the genus Oxychilus are also similar but are glossier, flatter spired and less striate. Of the introduced Oxychilus (Forsyth in press), O. alliarius is the most similar of these, but the animal exudes a pungent odour of garlic when irritated.

Habitat: This is an ubiquitous species living in a variety of habitats in lowlands and in higher elevations, where it occurs under rocks, dead wood, fallen leaves and vegetation. Z. arboreus is capable of living in a gravely substrate, but also in wetter habitats, and in general, this species prefers drier conditions than the next species. Vanatta (1914) reported Zonitoides arboreus at 1,471 m [4,825 ft] elevation in Montana.

Range: Great Slave Lake, Northwest Territories, and British Columbia east to Newfoundland and south in the United States, Mexico and Central America to Costa Rica; West Indies (Pilsbry 1946; Bequaert & Miller 1973). Introduced to Kamtschatka, Japan, eastern and northwest Europe, Iceland, Israel, Madeira, South Africa, Australia and Hawaii (Dall 1905; Likharev & Rammel'meier 1952; Bequaert & Miller 1973; Mienis 1977; Kerney & Cameron 1979; Cameron & Cooke 1996).

Distribution: Very widespread throughout the province and the Columbia Basin. This species is much more generally distributed and more common than Z. nitidus.

Name: Genus name derived from the resemblance to , a genus of snails in Europe. Species name meaning "of the trees".

Records: Creston Valley Wildlife Management Area (4907.19'N, 11637.82'W) (RBCM 998-00282- 004); Creston Valley, Creston Valley Interpretation Centre (4907.34'N, 11637.24'W) (RBCM 998-

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00264-010); Glenmarry Creek Road, near Nancy Greene Provincial Park (4915.94'N, 1756.36'W) (RBCM 998-00280-002); Fairmount Hotsprings, S end of Windermere Lake, off Hwy. 95 (4919.88'N, 11550.34'W) (RBCM 998-00300-002); Moyie Lake Park, Moyie Lake (4922.3'N, 11551.0'W) (Forsyth Coll. 97-102-1052); Near Weaver Creek, along logging road following Moyie River, S of Cranbrook (4924.50'N, 11559.95'W) (RBCM 998-00298-002); Elk River valley, along Hwy. 3, S of Fernie (circa 4927.5'N, 1154.2'W) (Forsyth Coll. 97-109-655); Rampart Rest Area, Hwy 3/93, NW of Wardner (4932.91'N, 11538.67'W) (Forsyth Coll. 97-104-1040); McDonalds Landing, West Arm, Kootenay Lake, near Six Mile, NE of Nelson (4934.75'N, 11712.99'W) (Forsyth Coll. 97-094-1093); W side of Slocan River, just S of Winlaw (4935.66'N, 11735.47'W) (RBCM 998-00279-006); Kokanee Creek Park, West Arm, Kootenay Lake (4936.2'N, 1177.7'W) (Forsyth Coll. 97-095-1087); E shore of Kootenay Lake, near boat launch S of ferry, Kootenay Bay (4940.4'N, 11652.3'W) (Forsyth Coll. 97- 097-1098); Columbia Lake, adjacent to Canal Flats Provincial Park (5010.65'N, 11549.03'W) (RBCM 998-00294-002); N end of Trout Lake, E of Upper Arrow Lake (5038.75'N, 11732.32'W) (RBCM 998-00285-007); Blanket Creek Provincial Park, S of Revelstoke (5050'N, 11805'W) (Staatliches Mu- seum fr Naturkunde Grlitz); Albert Canyon Road, near Albert Canyon Hotsprings Campground, E of Revelstoke (5107.94'N, 11751.61'W) (RBCM 998-00284-003); Sale Creek Road, Revelstoke (5109.1'N, 11811.0'W) (RBCM 998-00260-004); Hwy.1, 4 km E of Canyon Hot Spring (circa 5109.5'N, 11749'W) (Staatliches Museum fr Naturkunde Grlitz); W of snow sheds, Glacier National Park (circa 5115.5'N, 11728'W) (Staatliches Museum fr Naturkunde Grlitz); Hunter Creek Forest Recreation Site, along Kicking Horse Rive, ESE of Golden (5115.95'N, 11644.69'W) (Forsyth Coll. 98-092-3297); Field (5124'N, 11629'W) (Vanatta 1906; Pilsbry 1946); Doyle Rest Area, Hwy. 1, 20 km N of Golden (5126.55'N, 11705.31'W) (RBCM 998-00275-001); Bluewater Creek Forest Recreation Site, near Donald Station (5131.84'N, 11715.27'W) (Forsyth Coll. 98-095-3149).

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Zonitoides nitidus (Mller, 1774): Black Gloss

Description: Shell small to medium sized (width, 5.9 mm), depressed-heliciform, brown; translucent with opaque patches; spire moderately raised; whorls slightly convex and numbering 4.75-5; suture moderately impressed; protoconch smooth; teleoconch with irregular, low wrinklelike axial ribs and axial striae but without the exceedingly fine spiral striae of Z. arboreus; base smoother than Z. arboreus as well; aperture subovate and without denticles; outer lip unthickened; umbilicus approximately 20% of the width of the shell. The animal is black throughout. Pilsbry (1946) noted some lighter flecks along the edges of the foot. There is a pale, somewhat orangish spot on the mantle which can be seen through the shell back from the outer lip between the suture and the periphery.

Similar Species: Z. arboreus is slightly smaller, more depressed and more widely umbilicate. Additionally, the extremely fine, microscopic spiral lines present in Z. arboreus are lacking in Z. nitidus. The pigmenta- tion of the animal, as well as the habitat preference, also differ.

Habitat: Usually in wet places, under wood, rocks and vegetation, in marshes along the edges of rivers, sloughs, lakes and ponds.

Range: Holarctic: Alaska east to Hudson Bay and south to southern California (Pilsbry 1946); Iceland (Kerney & Cameron 1979), most of Europe, north Africa and Asia east to Japan (Ellis1969). Introduced to Madeira (Cameron & Cooke 1996) and Australia (Ellis 1969).

Distribution: Z. nitidus is not as common as Z. arboreus in British Columbia. In the Columbia Basin, the species was collected at the north end of Slocan Lake.

Name: Species name emphasizing the black colour of the animal.

Record: N end of Slocan Lake (4954.04' N, 11723.25' W) (RBCM 998-00266-003). View the map.

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Family VITRINIDAE Fitzinger, 1833

Genus Vitrina Draparnaud, 1801 For taxonimic changes since Pilsbry (1946), see the publications of Forcart (1955) and Bequaert & Miller (1973).

Vitrina pellucida (Mller, 1774): Western Glass-snail Synonym. Vitrina alaskana Dall, 1905.

Vitrina pellucida

Description: Shell small (width to 6.2 mm, but usually smaller), heliciform; shiny, translucent greenish, yellowish green or nearly colourless; very thin-shelled and fragile; spire small, low and convex; whorls 2.5-3, convex, regularly and rapidly increasing in width; periphery rounded; suture shallow and bordered by a more opaque band; teleoconch with irregular, low axial wrinkles that are a little more evident near the suture; aperture very large and subovate, without denticles; outer lip unthickened and with a very narrow, membranous edge; umbilicus narrow, formed by a fold in the columellar lip next to the base.

When the animal is extended from its shell, a small lobe of the mantle on the right side encroaches slightly over the shell and covers the termination of the suture. The animal is brownish, especially near the tentacles and on the head.

Similar species: Vitrina is easily distinguished from all other species occuring in the Kootenays by the combination of relatively few whorls, glassy shell and capacious aperture. However, Vitrina angelicae Beck, 1837 (synonym: V. limpida Gould in Agassiz, 1850), of eastern North America, has been reported from Alberta (Taylor 1895; Platt 1980). The shell is indistinguishable from V. pellucida and dissection is required for distinguishing the two species. In V. angelicae, the vas deferens is mostly free from the penis, but in V. pellucida, the vas deferens is enclosed in a large sheath of connective tissue surrounding part of the penis; additionally, the spermathecal duct of V. pellucida has a large swelling at its base adjacent to the atrium (Forcart 1955).

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Habitat: Under rocks, logs, and in grass and leaf litter. Vitrina is a cold-weather snail seen in the fall and early spring, often in seasonally dry situations from sea level to mountain tops above the tree-line, and dead shells are usually more often found. Carl & Hardy (1945) reported this species at 2,377 m [7,800 ft] at Paradise Mines near Windermere.

Biology: The life-span of this snail is probably one year (Boag & Wishart 1982). V. pellucida is usually seen alive in the spring and often there is still remnants of snow and ice on the ground. It is carnivorous and "feeds on almost anything but vascular plants" (Ellis 1969).

Range: Palearctic and western Nearctic. In North America: Alaska to Wyoming and South Dakota, south to California, Arizona and New Mexico (Bequaert & Miller 1973). Additionally, Russell (1951) reported "Vitrina alaskana", from the Cypress Hills of southwestern Saskatchewan.

Distribution: Throughout British Columbia, along the coast and inland, but more common in the north.

Notes: Forcart (1955) reviewed the northern species of the genus Vitrina, but did not treat V. alaskana because the reproductive anatomy was unknown to him. Later however, Bequaert & Miller (1973) placed V. alaskana in the synonymy of Vitrina pellucida stating that the reproductive anatomy of V. alaskana in examples they dissected agreed with that of Eurasian V. pellucida as illustrated and de- scribed by Forcart. Bequaert & Miller retained 'alaskana' as the North American subspecies of V. pelluc- ida on the basis of its geographical separation.

Identifications of Vitrina in this report were based upon geography. None have been dissected.

Name: Both genus and species names are in reference to the glassy, translucent shell.

Records: Olsen Park, Rock Creek (4903.37'N, 11859.90'W) (Forsyth Coll. 97-093-4237); Lodgepole Road at Kisoo Pass, SE of Fernie (4917.03'N, 11442.72'W) (RBCM 998-00288-004); Elk River valley, along Hwy. 3, S of Fernie (circa 4927.5'N, 1154.2'W) (Forsyth Coll. 97-109-634); Coal Creek on E side of Cokato Road, Fernie (4929.72'N, 11503.75'W) (Forsyth Coll. 97-108-1047); Cummings Creek, N of Sparwood (4946.32'N, 11454.95'W) (RBCM 998-00293-001); Paradise Mine, at source of Springs Creek, W of Windermere (circa 5028'N, 11614'W) (Carl & Hardy 1945); Sale Creek Road, Revelstoke (5109.1'N, 11811.0'W) (RBCM 998-00260-002); W of snow sheds, Glacier National Park (circa 5115.5'N, 11728'W) (Staatliches Museum fr Naturkunde Grlitz); Lake Revelstoke, Hwy. 37 to Mica Dam (5139.56'N, 11833.73'W) (RBCM 998-00258-004).

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Family LIMACIDAE Gray, 1824

Genus Limax Linnaeus, 1758

Limax maximus

Limax maximus Linnaeus, 1758: Giant Gardenslug

Description: A large slug (extended length, 100 mm or more), typically yellowish grey or brown spotted or striped with black, or uniformly dark brown; spots often coalesced into 2 or 3, often interrupted, pairs of lateral bands; mantle with spotted or mottled brown, well anterior and with its posterior edge somewhat angular from above, skin of the mantle having a pattern of fine concentric wrinkles, somewhat like a fingerprint, with this pattern centred on the midline of the mantle; pneumostome behind the middle of the mantle; head pale, tentacles reddish brown; foot fringe pale; keel incomplete, near the tip of the tail only; tail tapering more or less gradually to a point; mucus colourless, sole creamy white. Internal shell oblong with the nucleus terminal and left of centre.

Similar Species: Limax maximus is comparable in size to the native, coastal (Ariolimax columbianus) (Gould, 1851), but has a wrinkled mantle and different colour markings.

Habitat: In British Columbia Limax maximus occurs in gardens, fields and woods; in damp, shaded places, under rocks, wood and vegetation.

Range: Europe, Asia Minor and Algeria. Introduced North America, South America, Australia, Hawaii and elsewhere (Pilsbry 1948).

Distribution: Previously reported from Greater Vancouver, the Fraser Valley and Vernon (Rollo & Wel- lington 1975), but now known from southern Vancouver Island and the Columbia Basin.

Name: Genus, Latin for "slug". Species name in reference to the large size.

Records: W side of Slocan River, just S of Winlaw (4935.66'N, 11735.47'W) (RBCM 998-00279-001); Creston Valley Wildlife Management Area (4907.34'N, 11637.24'W) (RBCM 998-00264-001); N end of Slocan Lake (4954.04' N, 11723.25' W) (RBCM 998-00266-001).

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Family AGRIOLIMACIDAE Wagner, 1935

Genus Deroceras Rafinesque, 1820

These small slugs can be difficult to identify and may require dissection. There are two native and two introduced European species in British Columbia.

Deroceras laeve (Mller, 1774): Meadow Slug Synonyms. Limax campestris Binney, 1842; L. montanus Ingersoll, 1875; Limax hyperboreus Westerlund, 1877.

Description: A small slug (extended length, to about 25 mm), amber or dark brown without spots or other markings; mantle skin with a pattern of concentric rings that is centred to the right of the midline; pneumostome behind the middle of the mantle; keel not complete and very short, only at the end of the tail; trail abruptly truncate, rather than gradually tapering to a point; mucous thin, watery and clear; sole pale brownish. Internal shell oblong, with its nucleus terminal and to the left of centre.

Similar species: Deroceras reticulatum is larger than D. laeve, measuring 35-50 mm long when ex- tended; anatomically D. reticulatum has a complex flagellum (=penial appendage) having 1-4 knobby or serrated processes (Kerney & Cameron 1979). When disturbed, D. reticulatum secretes a milky white body mucous (otherwise it is clear); the mucous of D. laeve is clear and never whitish.

Habitat: Deroceras laeve favours very wet places (Kerney & Cameron 1979). In British Columbia this slug has been found living in moist habitats, in leaf litter, grass and sedges.

Range: Holarctic. In North America, from the Arctic south throughout Canada and the United States and into Central America (Pilsbry 1948).

Distribution: The distribution of this species in British Columbia is not well known, but it is likely through- out the province.

Name: Genus name meaning "neck-horn", referring to the tentacles on the head. Species name meaning "smooth", which was probably a comparison to Arion species (some of which have coarse turbercules on the back).

Records: Hwy 1, 4 km E of Canyon Hot Spring (circa 5109.5'N, 11749'W) (Staatliches Museum fr Naturkunde Grlitz p5951). RBCM records (not dissected, but presumed to be this species): Creston Valley Wildlife Management Area (4907.19'N, 11637.82'W) (RBCM 998-00282-007); Bull River near junction of Bull River and Kootenay River (4928.35'N, 11527.04'W) (RBCM 998-00301-001); W side of Slocan River, just S of Winlaw (4935.66'N, 11735.47'W) (RBCM 998-00279-002); N end of Slocan Lake (4954.04'N, 11723.25'W) (RBCM 998-00266-005).

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Deroceras reticulatum (Mller, 1774): Gray Fieldslug Synonym. Agriolimax agrestis auctt.

Description: A small slug (extended length, 35-50 mm), ground colour cream and usually with darker brown flecks; mantle skin with a pattern of concentric rings that is centred to the right of the midline; pneumostome behind the middle of the mantle; keel not complete and very short, only at the end of the tail; trail abruptly truncate, rather than gradually tapering to a point; when irritated, body mucous thick, white and sticky; sole whitish. Internal shell oblong, with its nucleus terminal and to the left of centre.

Similar species: Deroceras laeve, and additional European species; see Kerney & Cameron (1979).

Habitat: Gardens and other disturbed sites.

Range: Europe (Kerney & Cameron 1979). Widely introduced. Distribution. Probably widely introduced into many places throughout British Columbia.

Notes: Deroceras reticulatum was until recently included under the name Agriolimax agrestis (Linnaeus, 1758) in older literature. , however, is a separate species and presumably not introduced into British Columbia, although reported from British Columbia as early as the 1880s (Taylor 1892b), but records of D. agreste in B.C. probably refer to D. reticulatum. Rollo & Wellington (1975) failed to find D. agreste in the Vancouver area.

Name: Species name meaning "netted".

Records: Kaslo (4955'N, 11655'W) (Staatliches Museum fr Naturkunde Grlitz p5949); Columbia Lake adjacent to Canal Flats Provincial Park (5010.65'N, 11549.03'W) (RBCM 998-00294-001). View the map.

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Family POLYGYRIDAE Pilsbry, 1895

For taxonomic changes since Pilsbry (1939), see Emberton (1995).

Genus Cryptomastix Pilsbry, 1939

Cryptomastix mullani (Bland & Cooper, 1881): Coeur d'Alene Oregonian Synonym. () mullani olneyae Pilsbry, 1891.

Description: Shell medium-sized (width to 16 mm), more or less depressed-heliciform and reddish or yellowish brown, with a matt or polished surface; spire low-conical to almost flat; whorls convex and numbering 4-5; whorls slowly enlarging in width; periphery rounded; suture deep; last whorl not descending at the aperture; protoconch with rows of minute granules; teleoconch with low, rounded, irregular axial ribs and fine striae, and sometimes with weak spiral striae; occasional, small, weak malleations may be present on the last portion of the last whorl; periostracum with minute hairs in juveniles and some adults, but most adults without hairs; aperture subovate; parietal denticle short and white, but occasionally absent in some adults; columellar and basal lip thickened, strongly recurved and white; basal lip with a ridge like callus and medially located swelling; outer lip white, rather thickened, very strongly recurved, broadly expanded and with a slight posterior swelling; last whorl very slightly contracted behind outer lip; a rounded crest is present behind the outer lip; umbilicus small, 12-15% of the width of the shell and partially overhung by the expanded outer lip.

Similar Species: The shell of Cryptomastix mullani is larger and generally flatter-spired than the coastal C. germana (Gould in W.G. Binney, 1851). The other coastal Cryptomastix, C. devia (Gould, 1846), is larger than C. mullani and has the thickest point of the swelling on the basal lip located relatively closer to the columella than to the periphery. Specimens of C. mullani lacking a parietal denticle are likely to be confused with Vespericola columbianus (I. Lea, 1838), which however, has the outer lip strongly flared outward but not rolled over.

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Comparison of cross-sections of the outer lip of Vespericola columbianus (top) and Cryptomastix mullani (bottom). (Diagrammatic.)

Habitat: In British Columbia, Cryptomastix mullani has been found in the vicinity of streams, under rocks, logs, leaf litter and vegetation. According to Vagvolgyi (1968) predominantly between 457-1,829 m [1,500-6,000 ft], but sometimes as low as 152 m [500 ft ].

Range: Southeast British Columbia; Washington east of the Cascade Mountains to northeast Oregon and east through Idaho to western Montana (Pilsbry 1940).

Distribution: In British Columbia, previously reported only from Vernon (Pilsbry 1940). This species has been collected in the Kootenay, Columbia, Arrow Lake valleys, as far north as Eagle Pass west of Revelstoke.

Notes: Pilsbry (1940) referred shells from Vernon to the 'subspecies' olneyae, of which the type locality is Spokane, Washington; he recognised a number of named forms or 'subspecies', including olneyae, which have since been placed in the synonymy of Cryptomastix mullani mullani by Vagvolgyi (1968).

Name: Genus name derived from anatomical distinctions with related genera, and meaning hidden flagellum. Species name in honour of American Captain Mullan, who built a road over the Bitter Root and Rocky Mountains in 1853-1854 (Pilsbry 1940).

Records: Kuskanook Rest Area, Hwy 3A, E side of Kootenay Lake (4917.11'N, 11639.17'W) (Forsyth Collection 97-101-1006); Lodgepole Creek Road, SE of Fernie (4920.01'N, 11456.67'W) (RBCM 998- 00296-001); Syringa Creek, Lower Arrow Lake (circa 4921.1'N, 11754.6'W) (Forsyth Coll. 97-113- 4259); Nelson (4930'N, 11717'W) (RBCM 991-00095-005); Lockhart Creek, E shore of Kootenay Lake (4930.6' N, 116 47.1' W) (Forsyth 97-099-1039); W side of Slocan River, just S of Winlaw (4935.66'N, 11735.47'W) (RBCM 998-00279-004); N end of Slocan Lake, near hills (4954.04'N, 11723.25'W) (RBCM 998-00266-002); Roadside, Hwy 1, Eagle Pass, E of Clanwilliam, W of Revelstoke (5058.2'N, 11821.0'W) (Forsyth Coll. 98-097-3284).

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Genus Allogona Pilsbry, 1939

Allogona ptychophora (A.D. Brown, 1870): Idaho Forestsnail Synonyms. Helix ptychophorus castaneus Hemphill, 1890; Polygyra solida Vanatta, 1924.

Allogona ptychophora Description: Shell rather large (width to 23.7 mm), heliciform; surface matt or a little shiny, brownish or straw yellow; spire moderately elevated, but somewhat variable; whorls 5.25-5.75, convex; periphery rounded; suture deep; last part of the last whorl descending at lip; protoconch smoothish; teleoconch with fine axial striae and irregular, lighter-coloured wrinklelike axial riblets; exceedingly fine, wavy spiral striae throughout, but not visible on eroded shells; surface rarely malleated; periostracum not hirsute and sometimes mostly eroded away; aperture subovate, without a parietal denticle; columellar and basal lip white; basal lip heavy, thickened, and with a slight bulging callus at its junction with the columellar lip; outer lip thickened within and strongly reflected; last whorl scarcely contracted behind lip; crest very low or absent; umbilicus small, about 10% of the width of the shell, and partially concealed by the expanded and reflected columellar lip.

Juveniles lack the reflected, thickened outer lip and have the periphery slightly angled. Within populations of this snail a few are paler, of a yellowish straw-colour, and in this respect, is similar to the coastal species Allogona townsendiana (I. Lea, 1838), which also shows variation in colour.

Similar Species: The shell chiefly differs from A. townsendiana (I. Lea, 1838) by its slightly smaller size and generally less malleated ornamentation. A. lombardii A.G. Smith, 1943, is also similar (see Notes below).

Habitat: Under leaf litter, grass, rocks and logs, and on ground in open or in woods. Sometimes very abundant. In Montana, reported at 2,590 m [8,500 ft] (Vanatta 1914).

Range: Southern British Columbia south through Montana and Idaho and west down the Columbia River valley of Washington and Oregon (Pilsbry 1940).

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Distribution: Allogona ptychophora is common and generally spread in the southern Kootenay, Colum- bia and Elk river drainages, at least as far north as Revelstoke. Elsewhere in British Columbia A. ptychopora is known from Vernon (Pilsbry 1940) and Salmon Arm (RBCM 998-00018-001).

Notes: Smith (1943) suggested that specimens of Allogona from Boswell, British Columbia were Allogona lombardii A.G. Smith, 1943, which is said to differ from A. ptychophora by its larger size, heavier texture and more prominent ribbing. This British Columbia record of the A. lombardii is, however, unsubstantiated; Smith stated that the Boswell shells were similar to A. lombardii in sculpture but not in size, and that the specimens should probably take that name. All Allogona from southeast British Columbia are here considered to be A. ptychophora pending a more reliable means of separating A. lombardii and A. ptychophora.

Name: Genus name in reference to the anatomy and meaning "different genitalia". Species name meaning "bearing folds" and describing the sculputure of the shell.

Records: Creek Marsh, S end of Creston Valley Wildlife Management Area (4905.64'N, 11637.31'W) (RBCM 998-00281-001); Trail (4906'N, 11742'W) (Whiteaves 1906); Creston Valley Wildlife Manage- ment Area (4907.34'N, 11637.24'W) (RBCM 998-00264-004); Glenmarry Creek Road, near Nancy Greene Provincial Park (4915.94'N, 11756.36'W) (RBCM 998-00280-001); Lodgepole Creek Road (4917.03'N, 11442.72'W) (RBCM 998-00296-004); Kuskanook Rest Area, Hwy. 3A, E side of Kootenay Lake (4917.11'N, 11639.17'W) (Forsyth Coll. 97-101-1009); Syringa Creek Provincial Park, Lower Arrow Lake (circa 4921.1'N, 11754.6'W) (Forsyth Coll. 97-113-4240); Betania Road near Sand Creek, E end of Betania Road, S side of Hwy 3/93, W of Galloway (4922.23'N, 11514.59'W) (Forsyth Coll. 97-105-907); Boswell (4927'N, 11646'W) (Pilsbry 1940); Elk River valley, along Hwy 3, S of Fernie (circa 4927.5'N, 11504.2'W) (Forsyth Coll. 97-109-1016); Hepher Road at Hwy 3A, Boswell (4928.46'N, 11646.0'W) (Forsyth Coll. 97-100-864); Coal Creek at Cokato Road, Fernie (4929.72'N, 11503.75'W) (Forsyth Coll. 97-108-1048); "Cold Creek valley" [likely in error for Coal Creek (circa 4930'N, 11504'W)] near Fernie (Pilsbry 1940); Cranbrook (4930'N, 11546'W) (Pilsbry 1940); Nelson (4930'N, 11717'W) (RBCM 991-00095-006); Hwy 3 near Hosmer (4934.94'N, 11458.05'W) (RBCM 998-00299-002); Kokanee Landing (4936'N, 11706'W) (RBCM 996-00010-005, 996-00010-006); Kokanee Creek Provincial Park, Kootenay Lake (4936.2'N, 11707.7'W) (Forsyth Coll. 97-095-1084); Crowsnest Pass (4938'N, 11441'W) (Whiteaves 1906, Pilsbry 1940); Cummings Creek, N of Sparwood (4946.32'N, 11454.95'W) (RBCM 998-00293-002); Blanket Creek Provincial Park, S of Revelstoke (5050'N, 11805'W) (Staatliches Museum fr Naturkunde Grlitz p5958); along Hwy 1, W of Revelstoke, B.C., at junction with Hwy 23. (510.2'N, 11813.4'W) (Forsyth Coll. 98-096-3320). "Sproat" (Whiteaves 1906), possibly a locality in the vicinity of Mt. Sproat (5045'N, 11754'W) at the north of the north end of Upper Arrow Lake.

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Family THYSANOPHORIDAE Pilsbry, 1926

Genus Microphysula Cockerell & Pilsbry, 1926

Microphysula ingersollii (Bland, 1875): Spruce Snail

Description: Shell small (width, to 5.0 mm), depressed-heliciform, thin shelled, silky and translucent whitish; spire very low and very slightly convex; whorls convex and about 5.5 in number; whorls tightly coiled and slowly enlarging; periphery rounded; suture very deep; protoconch smoothish; teleoconch with very low, weak axial riblets and fine, close, evenly spaced spiral striae that are even finer and more closely spaced on the base and that are not evident on the early whorls of the teleoconch; aperture subovate and without denticles; lip unthickened and not expanded; umbilicus about 25% of the width of the shell.

Similar Species: A coastal species, Microphysula cookei Pilsbry 1922, is very similar. The shell is smaller and has fewer, less tightly coiled whorls (Pilsbry 1940).

Habitat: In British Columbia, Microphysula ingersollii has been recorded living under mossy boulders, rocks and dead wood in moist places. Beetle (1957) recorded this snail from an aspen grove in Wyoming. In Arizona, M. ingersollii is reported at altitudes of 981-3,353 m [6,500-11,000 ft] (Bequaert & Miller 1973). Vannatta (1914) reported M. ingersollii in Montana from 2,591 m [8,500 ft]. Berry (1919) also recorded the species in Montana, from altitudes of 1,082-1,829 m [3,550-6,000 ft] from under sticks, loose rocks, bark and small logs.

Range: Rocky Mountain region; southeast British Columbia through Washington, Oregon, Idaho, Mon- tana, Wyoming, Utah, Colorado, New Mexico and Arizona (Bequaert & Miller 1973).

Distribution: In British Columbia, Microphysula ingersollii is limited to the mountain of the southeastern British Columbia where it is known from a few localities in the Columbia Basin region. Sherbrooke Trailhead in Yoho National Park is the most northern locality for this species (circa 5126.4'N, 11621.5'W).

Name: Genus name meaning "little bubble". Species named after Ernest Ingersoll, U.S. Geological Survey naturalist, who discovered the species (Binney & Bland 1885).

Records: E of Hwy bridge, S bank of Elk River (4910.76'N, 1159.76'W) (Forysth Coll. 97-107-4236); Lockhart Creek, Kootenay Lake (4930.6'N, 11647.1'W) (Forysth Coll. 97-099-1045); Cummings Creek, N of Sparwood (4946.32'N, 11454.95'W) (RBCM 998-00293-007); along 2nd Avenue, 200 m Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 113 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

SW of First Street East, Field (5123.75'N, 11629.3'W) (Forysth Coll. 98-091-3073); Field (5124'W, 11629'W) (Vannatta 1906; Pilsbry 1940); Sherbrooke Trailhead, N of Wapta Lake, Yoho National Park (circa 5126.4'N, 11621.5'W) (Forysth Coll. 98-090-3389).

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Family Helicidae Rafinesque, 1815

Genus Cepaea Held, 1837

Cepaea nemoralis (Linnaeus, 1758): Grovesnail

Description: Shell large (width to 25 mm), heliciform; opaque, not very glossy; brightly coloured, brown, orange or yellow, with 1-5 blackish or dark brown spiral bands that may be coalesced or absent; spire moderately elevated; whorls 4.5-5.25, convex; periphery rounded; suture deep; last whorl strongly de- scending at the lip; protoconch smooth; teleoconch with axial striae and wrinkles; aperture wider than high, and without denticles within; columella slightly thickened and usually dark coloured; basal lip thickened, rather straight and not evenly curved into the outer and columellar lips; outer lip moderately thickened and a slightly recurved; umbilicus absent in adults.

Juveniles in which the columellar lip is still poorly formed are narrowly umbilicate. Variation in the colour of the shell is remarkable with the number of bands varying from zero to five, but three-banded individuals appear to be most common. Banding is influenced both by genetics and environmental conditions (Brussard 1975; Clarke et al. 1978).

The body of this snail is cream coloured/pale brown, becoming darker towards the head and on the tenta- cles.

Similar Species: The locally introduced snail Helix aspersa Linnaeus, 1758 is larger, with a more rounded aperture and with a different colour pattern consisting of pale axial streaks over top of darker spiral bands. The native, coastal snail, (J.E. Gray, 1838) is also banded, but larger and with a distinctly open umbilicus.

Habitat: Gardens and parks in urban areas. On and under plants and sometimes up trees, well off the ground.

Biology: C. nemoralis breeds in the spring and summer, the eggs hatch in two or three weeks and the young grow to adult size the subsequent year (Ellis 1969). In Britain, this snail takes three years to reach adult size (Williamson 1979).

Range: Central and western Europe. Introduced to North America, presumably with plants (Pilsbry 1936; Hanna 1966).

Distribution: Introduced to southern Vancouver Island, Greater Vancouver, the Fraser Valley, the Okanagan (Westbank) (Forsyth in press) and the Columbia Basin.

Name: Genus name meaning "an onion". Species name meaning "sylvan".

Records: Nelson (ca. 49o30'N, 117o17'W) (RBCM 999-278-001).

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Glossary

Specialised terminology is necessarily used in malacology. In this report, terminology is utilised in the de- scriptions of animals, their shells, natural history, habitat and nomenclature. Terms are standardised and adapted several sources. Shell terminology is adapted from Pilsbry (1936-1948), Cox (1960), Arnold (1966) and Keen & Coan (1974). Anatomical terminology is adapted from Pilsbry (1936-1948), Ellis (1969a, b), Kerney & Cameron (1979), South (1992) and other sources.

adult a shell that is fully grown, having all the characteristics of adult individuals; a sexually mature animal angulate having an angulation angulation an angular formation, ridge, keel or edge where two surfaces meet anterior in front, near the head apertural of or in the aperture aperture the opening in a spirally coiled shell through which the animal extends and into which the animal can retract (in most) apex the tip of the spire; that part of the shell (often pointed) that is first formed apical of or at the apex atrium the chamber to which the oviduct, penis (or epiphallus) and spermathecal duct are connected to, and which itself is connected to the outside by way of the genital pore auctt. auctorum, of authors: indicating a misuse of a name by several authors axial in the same direction as the axis of the coiling of a shell axial striae fine lines (usually incised) on the surface of the shell indicating growth by increments axis imaginary line around which the whorls of a coiled shell are formed basal of or at the base basal lip that part of the lip from the end of the columella to the future suture base the rounded lower part of the last whorl of a coiled shell

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MS (pl. MSS) manuscript: an unpublished work mull granular forest humus with the layer of mixed organic matter and mineral soil merging gradually into the mineral soil beneath; see alsomor. multispiral with many whorls, especially when whorls are slowing enlarging in size and tightly wound mucus a slippery or sticky secretion, often called slime obsolete obscure, not very distinct, atrophied ocular of the eye opaque not transparent or translucent outer lip the edge of the aperture from suture to the base; see alsobasal lip ovate egg-shaped, oval ovate-conic ovate but with a somewhat conic apex oviduct the female duct through which eggs past from the common duct to the atrium palatal that part of the lip or inner surface of the shell from the suture to the base parietal of the parietal area parietal area that part of the inner surface of the shell between columella and the suture formed by the preceding whorl uniting with the lip paucispiral with few (usually rapidly enlarging) whorls penial appendage an outgrowth from the proximal part of the penis (in Deroceras), either simple or branched

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sensu lato in the broad sense sensu stricto in the strict sense shagreened resembling shagreen; having a rough, granular surface shell a hard, calcified or chitinous structure, either external and able to encase all or part of the withdrawn animal, or internal shell height the maximum measurement taken along the line of the axis from the apex to the base or basal lip; equivalent to length of some authors shell width maximum measurement taken perpendicular to the axial of a coiled shell; equivalent to diameter of some authors shoulder that part of the whorl directly below the suture and above the periphery, either angulate or rounded sinistral opposite of dextral; coiled in a left-handed spiral; the aperture is on the left side of the shell when viewed with the apex up sinulus an identation of the outer lip in some species of Vertigo and Gastrocopta s.l. see 'sensu lato' sole the flat, muscular undersurface of the foot of a gastropod species the basic category of biological classification, ranking below the genus; a species consists of related organisms or populations potentially capable of interbreeding; a species is designated by a two part name consisting of its genus and a specific epithet; see also subspecies spermatheca a saclike part of the genital system for storing spermatozoa received from another indi vidual; (=seminal receptacle, bursa copulatrix) spermathecal a tube leading from the spermatheca to the atrium (=seminal duct) duct spiral in the direction of the coiling of the shell spire the whorls of a coiled shell except for the last whorl s.s. see 'sensu stricto'

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striae very fine lines or furrows in the shell surface; usually considered only to be incised, but here both incised and raised lines are considered striae striate sculptured with striae sub- below, under; rather, somewhat, almost, nearly subspecies a division of a species, based upon a isolation (either geographically or ecologically) from the main species' stock suture the continuous seam between two adjacent whorls of a coiled shell (the form "sutures" is incorrect, as there is only one suture in a coiled shell) sympatric occurring together in the same area or region synonym one of two or more names given to a singe species (or genus, family, etc.); the name not considered valid is often referred to as a synonym of the valid name, but more correctly, both are synonyms of each other taxon (pl. taxa) any unit of classification teleoconch the entire except for the protoconch tentacles more or less slender, contractile or retractile, sensory organs on the head of gastropods; see also occular tentacles and sensory tentacles terrestrial living on land thread a long, raised sculptural element translucent allowing the passage of light, but not transparent tripartite said of the sole of the foot when it appears to be divided into three longitudinal bands tubercle a swelling, hump or knob tubercular shaped like tubercle type locality the locality from where the holotype specimen of a species originated umbilicate having the umbilicus open umbilicus in some coiled shells, the basal opening through which passes the central axis of the shell around which the whorls are formed

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vas deferens the narrow duct connecting the prostrate (section of thecommon duct) with either the penis or epiphallus. ventral on or of the lower plane of the body; opposite to dorsal viviparous producing live young, rather than eggs whorl one complete revolution of a coiled shell; see also last whorl, penultimate whorl width see shell width wrinkles irregular furrows and/or ridges

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Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 132 Living Landscapes Terrestrial Gastropods of the Columbia Basin, British Columbia by Robert G. Forsyth Copyright 1999 Royal British Columbia Museum

SNAILS maximum dimension general size (length or width of shell, whatever the greatest)

< 2 mm very small 2-5 mm small 6-20 mm medium-sized 21-30 mm large > 30 mm very large

SLUGS maximum length general size (fully extended, crawling animals)

< 25 mm small 25-60 mm medium-sized 61-100 mm large > 100 mm very large

Royal British Columbia Museum (250) 356-RBCM (7226) 675 Belleville Street 1-888-447-7977 Victoria, British Columbia http://www.royalbcmuseum.bc.ca CANADA V8W 9W2 133