Terrestrial Gastropods of the Upper Fraser Basin of British Columbia

Transcription

1 Terrestrial Gastropods of the Upper Fraser Basin of British Columbia ROBERT G. FORSYTH Research Associate, Royal BC Museum, Victoria 675 Belleville Street, Victoria, British Columbia, Canada V8W 9W2 LIVING LANDSCAPES: UPPER FRASER BASIN

2 Copyright 2005 by the Royal British Columbia Museum. Figures 3 and 5 Robert G. Forsyth On preceding page: Scanning electron photomicrograph of a specimen of Punctum randolphii (see page 10) from Bone Creek, North Thompson River (outside of the upper Fraser drainage) (RBCM ). Library and Archives Canada Cataloguing in Publication Data Forsyth, Robert G. Terrestrial gastropods of the Upper Fraser Basin of British Columbia [electronic resource] Available on the Internet. Living Landscapes: Upper Fraser Basin Includes bibliographical references: p. ISBN Gastropoda British Columbia Fraser River Watershed. 2. Gastropoda British Columbia, Northern. I. Royal BC Museum. II. Title. QL430.4.F C This report can be cited as: Forsyth, Robert G Terrestrial Gastropods of the Upper Fraser Basin of British Columbia. Living Landscapes, Royal BC Museum, Victoria. ufb_snails/index.html [HTML version] or bc.ca/upperfraserbasin/ufb-snails/ufb-snails.pdf [PDF version].

3 Terrestrial Gastropods of the Upper Fraser Basin of British Columbia ROBERT G. FORSYTH* Research Associate, Royal BC Museum, Victoria 675 Belleville Street, Victoria, British Columbia, Canada V8W 9W2 *Mailing address: Box 3804, Smithers, British Columbia, Canada V0J 2N0 ABSTRACT Information on the terrestrial gastropod fauna is compiled from new or recent field collections, museum records and literature for that part of British Columbia encompassing the basin of the Fraser River north of about 52 N. Recent fieldwork (2001) has added many new locality records for a region that has received little study and has significantly improved the region s representation of terrestrial gastropods in the collection of the Royal British Columbia Museum. At least 28 species belonging to 18 genera are recorded from 83 localities. All but three species were previously unreported in the literature from this area of British Columbia. CONTENTS Overview of the Project Materials and Methods Annotated Species List Family Succineidae Family Cionellidae Family Valloniidae Family Vertiginidae Family Punctidae Family Discidae Family Euconulidae Family Gastrodontidae Family Daudebardiidae Family Vitrinidae Family Limacidae Family Agriolimacidae Family Arionidae Conclusions Acknowledgements Literature Cited Distribution Maps Locality Locality Data

4 115 4 LIVING LANDSCAPES: UPPER FRASER BASIN kilometres 300 British Columbia Figure 1 Location of the upper Fraser Basin in the province of British Columbia. 1, Burns Lake; 2, Vanderhoof; 3, Prince George; 4, McBride; 5, Valemount; 6, Quesnel; 7, Williams Lake; 8, 100 Mile House. OVERVIEW OF THE PROJECT This report, as part of Living Landscapes, documents the terrestrial molluscs (snails and slugs) of the upper Fraser Basin. This is a vast region of British Columbia that extends from the headwaters of the Fraser River at the Alberta border (52 34 N, W; N, W at the Yellowhead Pass) west to the Coast Mountains, including the headwaters of the Nechako River. To the north the Fraser drainage extends to Summit Lake on Highway 97 and northwest to the headwaters of the Driftwood River (56 10 N, W) (Figure 1). Existing data on terrestrial molluscs of the upper Fraser Basin consisted mostly of a few historical collections, two published scientific papers, and my own collections and unpublished data. In June 2001 I took part in fieldwork for the purpose of collecting specimens for the Invertebrate Zoology Collection of the Royal BC Museum. Previous to this, the upper Fraser Basin was poorly represented in the museum s collection. The main objectives of the project were: To survey for, and improve our knowledge of, terrestrial molluscs of the upper Fraser Basin; To add value to specimens already part of the Royal BC Museum collection by checking identifications and specimen data; and To amalgamate collection records and data from various sources into a single document (this report).

5 TERRESTRIAL GASTROPODS OF THE UPPER FRASER BASIN 5 The majority of British Columbia s terrestrial molluscs are tiny and usually overlooked in general natural history surveys. Prior to the Royal BC Museum handbook, Land Snails of British Columbia (Forsyth 2004) only two publications (Stanwell-Fletcher & Stanwell-Fletcher 1943; Harris & Hubricht 1982) report any species from the upper Fraser Basin. Even the landmark monograph of Pilsbry ( ) does not list locality records from this large area of British Columbia. This report documents the species of terrestrial molluscs known from the upper Fraser Basin, and gives the results of recent fieldwork. For further information on the species documented here, see Land Snails of British Columbia (Forsyth 2004), which includes full descriptions, figures and keys. MATERIALS AND METHODS Most of the records documented here are the result of a reconnaissance survey of the upper Fraser Basin in July The primary purpose was to obtain specimens and data of terrestrial molluscs for the Royal British Columbia Museum. Time limitations prevented us from surveying the expansive plateau area west of Williams Lake and Quesnel, and our efforts were concentrated in the areas to the east and west of Prince George and around Quesnel. Most of the collections documented in this report are from along or near highways and other roads. Few collections were made at high-elevation sites. Specimens were collected by hand picking from soil, leaf litter, rocks, logs and other dead wood, and were incorporated into the Invertebrate Zoology Collection of the Royal British Columbia Museum, Victoria (RBCM). All slug and some snail material were drowned in water (to relax animal and extend the bodies), then preserved in 70% EtOH (ethyl alcohol). A portion of the shelled snail material was put directly into EtOH, dried later in the lab for the dry collection. Specimens from my personal study collection (RGF) and some material from the Canadian Museum of Nature (CMN) were examined, and their data included in this report. However, the collection of the CMN was not exhaustively studied, as much of the material was inaccessible to me. I also briefly include the catalogue number for one lot of Succineidae that were cited by Harris & Hubricht (1982) from the upper Fraser Basin and now part of the Invertebrate Collection of the Field Museum of Natural History (FMNH, Chicago). I have not attempted, however, to verify these identifications, nor have I tried to Figure 2 Collection localities (from all sources) in the upper Fraser Basin (see Appendix for data). identify to species any of the succineids that I have seen. While in the field, locality geopositions were acquired using a hand held GPS; these readings were subsequently checked on National Topographic Series 1:50,000-scale paper maps or electronic versions of these maps. Some earlier museum collections lack precise position data (most CMN records, for example) and an approximate position was derived from maps for mapping purposes. Elevations above sea level (abbreviated as a.s.l.) presented in the Appendix and elsewhere are approximate, having been secondarily derived from maps and in most cases converted from feet to metres. This report uses the same nomenclature as appears in Land Snails of British Columbia (Forsyth 2004). ANNOTATED SPECIES LIST In June 2001 a total of 53 localities or stations were surveyed, in addition to other localities outside the upper Fraser Basin Living Landscapes region and beyond the scope of this report. They form the majority of the 83 unique localities listed in the Appendix (see also Figure 2). Thirteen families 1, 16 genera and 27 species (exclusive of the Succineidae) are documented here from the 1 The families used here are as presented in Land Snails of British Columbia (Forsyth 2004); many North American authors (for example, Turgeon et al. 1998) recognize fewer families.

6 6 LIVING LANDSCAPES: UPPER FRASER BASIN Table 1 Conspectus of terrestrial gastropods in the upper Fraser Basin. Species Native/Introduced Literature Source or Collection Containing Records (a) Arion circumscriptus Introduced RBCM [LL] Arion subfuscus Introduced RBCM Cochlicopa lubrica Native? (b) RBCM [LL] Columella columella Native RBCM [LL] Columella edentula Native RBCM [LL], RGF Deroceras laeve Native RBCM [LL], CMN Deroceras reticulatum Introduced RBCM [LL], CMN Discus shimekii Native Stanwell-Fletcher & Stanwell-Fletcher (1943); see text Discus whitneyi Native RBCM, RBCM [LL], RGF, CMN Euconulus fulvus Native RBCM, RBCM [LL], RGF, CMN Euconulus praticola Native RBCM [LL], RGF Limax maximus Introduced RBCM [LL] Nesovitrea binneyana Native RBCM [LL], RGF Nesovitrea electrina Native RBCM [LL] Oxyloma nuttallianum Native Harris & Hubricht (1982), FMNH Paralaoma servilis Native RGF Punctum randolphii Native RBCM [LL] Prophysaon andersonii Native RBCM, RBCM [LL], CMN Vallonia excentrica Introduced RBCM [LL] Vallonia pulchella Introduced RBCM [LL] Vertigo arthuri Native RGF Vertigo cristata Native RBCM [LL] Vertigo gouldii Native RBCM [LL] Vertigo modesta Native RBCM [LL], RGF Vertigo ovata Native RBCM [LL] Vitrina pellucida Native RBCM [LL], RGF, CMN Zonitoides arboreus Native RBCM [LL], RGF, CMN Zoogenetes harpa Native RBCM [LL] Unidentified Succineidae Native RBCM [LL], RGF (a) (b) Abbreviations are as follows: CMN: Canadian Museum of Nature; FMNH: Field Museum of Natural History; RBCM: Royal British Columbia Museum (collections made prior to Living Landscapes); RBCM [LL]: Royal BC Museum (collections made in 2001 for Living Landscapes); RGF: author s reference collection. Upper Fraser basin records of Cochlicopa lubrica are clearly synanthropic and may be introduced.

7 TERRESTRIAL GASTROPODS OF THE UPPER FRASER BASIN 7 upper Fraser Basin. Only two species Discus shimekii and O. nuttallianum have been recorded in the malacological literature from this region (Stanwell-Fletcher & Stanwell-Fletcher 1943; Harris & Hubricht 1982). The Living Landscapes survey in 2001 added 12 species not previously recorded from the area. The remaining 15 species were already known from the upper Fraser Basin but unpublished (Table 1). The majority of species in upper Fraser Basin are native, but at least six species Vallonia excentrica, Vallonia pulchella, Arion circumscriptus, Arion subfuscus, Limax maximus and Deroceras reticulatum are introduced. Cochlicopa lubrica, is a special case since it shows a strong synanthropic tendency and may be at least partially introduced in British Columbia. FAMILY SUCCINEIDAE Shells of succineids offer few characteristics for specific identification, and species determinations require that reproductive anatomy be considered. Reliable identifications of western succineids in particular is further inhibited by a very limited knowledge of some nominal species and the range of variation within species. Most of what we know on this group was given by Pilsbry (1948), but Harris & Hubricht (1982) have dealt with some species of Oxyloma from southwestern Canada. Based on their data and the Hubricht collection (now in the Field Museum of Natural History, Chichago, IL) only one species has been recorded from the upper Fraser Basin: Oxyloma nuttallianum (I. Lea, 1841) (1 lot, FMNH 2341, Lac la Hache), but a second species, as Oxyloma groenlandica (Müller, 1774), was recorded from nearby areas outside of the area covered by this report. I have not tried to identify or verify collections of succineids from the upper Fraser Basin. FAMILY CIONELLIDAE Cochlicopa lubrica (Müller, 1774): Glossy Pillar Records. Map 1. RBCM: , In British Columbia, this is a largely synanthropic species, having what appears to be introduced and possibly native populations (Forsyth 2004); in the upper Fraser Basin C. lubrica was found only in disturbed habitats. There are only two records from the upper Fraser Basin, but synanthropic habitats were not extensively sampled for snails and slugs and the species is likely common in towns. FAMILY VALLONIIDAE Vallonia excentrica Sterki in Pilsbry, 1893: Iroquois Vallonia Records. Map 2. RBCM: Vallonia excentrica was living under shrubs and young cottonwoods on a roadside embankment in Quesnel. This is an introduced species in British Columbia, found in disturbed and synathropic habitats. Vallonia pulchella (Müller, 1774): Lovely Vallonia Records. Map 3. RBCM: In the upper Fraser Basin Vallonia pulchella was living under wood on a grassy roadside along Hwy 16 east of Prince George. This species is also an introduction and similarly to V. pulchella is found in disturbed and synathropic habitats. Zoogenetes harpa (Say, 1824): Boreal Top Records. Map 4. RBCM: , , In the upper Fraser Basin Zoogenetes harpa was found living in leaf litter and on dead wood in a mesic spruce forest, a stand of trembling aspen with willow, and a clearcut (burnt in 1986) with willow and young conifers. Elsewhere in the province, this species has also been found living on disturbed roadside habitats and in a gasline right-of-way. Z. harpa appears to form sporadic, discreet colonies, but when found snails are usually abundant. In British Columbia, this species is mostly known from the central and northern half of the province, but there is a single record from Albert Canyon in the Selkirk Mountains (RBCM collection). The known localities for Z. harpa in the upper Fraser Basin are between m a.s.l.; the elevational range for all B.C. records is m a.s.l. FAMILY VERTIGINIDAE Columella columella (von Martens, 1830): Mellow Column Records. Map 5. RBCM: , , , In the upper Fraser Basin Columella columella appears to inhabit only the eastern, mountainous parts of the region, and was found living under dead wood and in leaf litter in mesic forests and wooded floodplains of waterways, at elevations between 7000 m. In the whole of British Columbia, C. columella is known only from the mountainous southeast and north along the Rocky Mountains,

8 8 LIVING LANDSCAPES: UPPER FRASER BASIN The name Columella edentula is widely used in North America for the smaller and shorter shelled Columella, but B.C. material may be another taxon. I use the name Columella edentula in a provisional sense; British Columbia shells and animals should to be compared with examples of C. edentula from Eurasia and the rest of North America, as well as another Eurasian species, C. aspera Waldén 1966 (see Pokryszko 1990 for characters of these species). Figure 3 Vertigo arthuri. Lac La Hache (RGF ) (above); Kiskatinaw Park (RGF ) (below). the adjacent valleys and the Rocky Mountain Foothills. This is the species that Pilsbry (1948) called Columella alticola (Ingersoll, 1875), but some recent North American authors treat this name as a synonym or subspecies of C. columella, which was described from Quaternary fossils from Germany. Pokryszko (1987, 1990), who however chose to not synonymize C. alticola with C. columella, distinguished Polish examples of C. columella from C. edentula by their more cylindrical and strongly irregularly and sharply striate shells and the elongate-oval apertures of mature individuals. British Columbia shells also show these characters, though B.C. specimens of C. edentula (see below) are also rather well-striated. Pokryszko also gave anatomical characters to help separate European Columella, but this has not yet been attempted for B.C. material. Columella edentula (Draparnaud, 1805): Widespread Column Records. Map 6. RBCM: , , , , RGF: In the upper Fraser Basin, Columella edentula was found in mesic forests, at elevations between 670 m and 880 m. Unlike C. columella, this species is more generally distributed in British Columbia (Forsyth 2004). Vertigo arthuri von Martens, 1882: Callused Vertigo Records. Map 7. RGF: In British Columbia, Vertigo arthuri is currently known only from two localities, the one documented here (Lac la Hache, 820 m a.s.l.) and another at Kiskatinaw Park in the Peace River region (Forsyth collection; 590 m a.s.l.) At both localities snails were found under dead wood or sticks. The Kiskatinaw site has young cottonwoods, while the Lac la Hache site was relatively dry, open Douglas-fir forest. Vertigo arthuri was described from North Dakota by von Martens but remained unrecorded from much of its range until recently. This species has since been recognized from Alaska, the Yukon, Alberta, South Dakota, Wyoming, Minnesota, Ontario (T. Frest, S. Harris, W. Grimm, and B. Coles, personal communications; Nekola & Massart 2001). This distinct little Vertigo is recognized by its subcylindrical form, sculpture of rather sharp, almost rib-like axial lines, huge (albeit variable) crest behind the apertural lip, six apertural denticles (angular, parietal, columellar, subcolumellar and upper and lower palatals) and typically a well-developed palatal callus (Figure 3). Also quite distinct in this species is the fact that the columellar denticle appears bilobed, at least in some well-developed individuals; Pilsbry (1948: 977) noted this as having a convex callus against its [the columellar denticle s] lower side. The Kiskatinaw specimen (length, 1.75 mm) is quite typical, but the Lac la Hache specimen (length, 1.75 mm) has a smaller crest and correspondingly weaker palatal callus. Vertigo cristata Sterki in Pilsbry, 1919: Crested Vertigo Records. Map 8. RBCM: , , , , , , , , In the upper Fraser Basin Vertigo cristata was found on dead sticks, under logs and fallen bark in coniferous for-

9 TERRESTRIAL GASTROPODS OF THE UPPER FRASER BASIN 9 ing from V. arthuri by not having the columellar denticle bilobed and by having a less developed crest. Vertigo modesta (Say, 1824): Cross Vertigo Records. Map 10. RBCM: , , , , , , , , RGF: In the upper Fraser Basin Vertigo modesta was found living under dead wood and dead vegetation and in litter in coniferous forests at elevations between m a.s.l. Sites varied from relatively dry, open pine or spruce forests to moist spruce or hemlock forests with diverse vegetation. Elsewhere in British Columbia, this species is also recorded from near treeline, up to about 1700 m. Many examples of B.C. Vertigo modesta have an angular denticle developed to varying degrees, even within a series of shells from one location. Figure 4 Vertigo cristata. Hwy 16, E of Tête Jaune Cache at Mt Terry Fox Rest Area (RBCM ). ests of Douglas-fir, spruce or lodgepole pine, but at two localites it was also collected in groves of trembling aspen. The elevational range of these records is from about m a.s.l. In British Columbia, Vertigo cristata is common along the Rocky Mountains and in the northern part of the province. I use this name provisionally for small Vertigo animals with shells having four apertural denticles (a columellar, a parietal and two palatals), remarkably regular, rib-like axial striae, a small crest, and no palatal callus (Figures 4, 5). Vertigo gouldii (A. Binney, 1843): Variable Vertigo Records. Map 9. RBCM: Vertigo gouldii was found at only one locality in the upper Fraser Basin, near the summit along Hwy 20 between Williams Lake and the Fraser River (RBCM ). Here, this species was found under dead wood in a dry Engelmann spruce forest with an open understory (elevation 900 m a.s.l.). In British Columbia, Vertigo gouldii is mostly known from the southern Rocky Mountains. This name is used provisionally (see Forsyth 2004) for Vertigo with rib-striate shells having six apertural denticles (a columellar, a parietal, an angular, two palatals, and a basal), but differ- Vertigo ovata Say, 1822: Ovate Vertigo Records. Map 11. RBCM: In the upper Fraser Basin Vertigo ovata was found at a single locality, on the marshy shore of Amanita Lake, north of McGregor (670 m a.s.l.). This is a characteristically wetland species found in marshes, wet lakeshore meadows and fens, where it is typically found under dead vegetation and wood. Although restricted to wetlands which tend to be less intensely searched for terrestrial molluscs, this species is likely not uncommon in B.C. in suitable habitats and shows a pattern of scattered distribution records indicative of specialized habitats that tend to be infrequently surveyed for land molluscs. Figure 5 Vertigo cristata. Charlie Lake (outside of the upper Fraser drainage) (RGF ).

10 10 LIVING LANDSCAPES: UPPER FRASER BASIN Figure 6 Punctum randolphii. A specimen from Bone Creek, North Thompson River (outside of the upper Fraser drainage) (RBCM ). FAMILY PUNCTIDAE Paralaoma servilis (Shuttleworth, 1852): Pinhead Spot Records. Map 12. RGF: , In British Columbia Paralaoma servilis is most often seen at disturbed, open sites during the wetter months of the year. In the upper Fraser Basin, this species was collected from under aspen logs at two localities, both with somewhat modified habitat, at elevations of 690 m and 760 m a.s.l. Although this species has an inclination to be synanthropic, it appears to be native to northwestern North America. Among the plethora of synonymous names applied to this widespread, almost cosmopolitan species are Punctum conspectum (Bland, 1865), Punctum (or Toltecia) pusilla (Lowe, 1831), and Paralaoma caputspinulae (Reeve, 1852) (see Falkner et al and references therein). Punctum randolphii (Dall, 1895): Conical Spot Records. Map 13. RBCM: , , In the upper Fraser Basin, this species was living in leaf litter in a cottonwood stand, an open mixed-wood forest, and a grove of trembling aspen; the elevational range of the three sites is m a.s.l. Elsewhere in B.C., the species has been found from sea level to about 1700 m a.s.l. Recent collections made in , but outside the geographic limits of this report, suggest that the species is common in the Quesnel and Shuswap Highlands and along the North Thompson River valley. It may be more infrequent on the eastern margin of the Interior Plateau and appears absent from a large area of the Interior Plateau. P. randolphii is also recorded from the Kootenays, the Skeena and Hazelton mountains (Forsyth 2001a), the Queen Charlotte Islands (Forsyth collection), the north coast (RBCM collection) and from many localities on the south coast. Compared to coastal British Columbia examples, most interior specimens (Figure 6) are generally a little smaller, but shell size and the relative height to width is variable both in coastal and interior examples. Punctum randolphii looks to be much more variable in form and size than most literature on these snails (e.g., Pilsbry 1946) would suggest. Although a single, large shell reported earlier from the Columbia Basin was tentatively called P. californicum Pilsbry, 1898 (Forsyth 1999), I now consider this and all British Columbia material as belonging to one variable species for which I use the name P. randolphii. However, the issue is further complicated because Alberta snails are usually identified as P. minutissimum (Lea, 1841), because they occur east of the Continental Divide. FAMILY DISCIDAE Discus shimekii (Pilsbry, 1890): Striate Disc Records. Map 14; around Lake Tetana (Stanwell-Fletcher & Stanwell-Fletcher 1943). This species is included here on the grounds that Stanwell-Fletcher & Stanwell-Fletcher (1943:36) recorded it from Tetana Lake on the Driftwood River (ca N, W). Except for Discus shimekii, they did not list other species of terrestrial molluscs in brief observations on the common insects and a few other invertebrates that they collected around the lake. Any material, if it existed, would be expected to be in the RBCM, but I have not found it, and the record cannot be verified. In British Columbia, D. shimekii occurs along the Rocky Mountains, in the Peace River region and west to the Haynes Triangle. Discus whitneyi (Newcomb, 1864): Forest Disc Records. Map 15. RBCM: , , , , , , , , , , , , , , , , , , , , , , , , RGF: , , , , , , , , , , , , , , , CMN:

11 TERRESTRIAL GASTROPODS OF THE UPPER FRASER BASIN 11 Discus whitneyi is one of the most common and widespread land snails in the upper Fraser Basin and in British Columbia as a whole, except along the coast. In the upper Fraser Basin, only Euconulus fulvus appears more ubiquitous. D. whitneyi lives in forests and at open sites, under fallen branches, logs, stones, debris and in leaf litter; in the upper Fraser Basin it has been found at elevations between m a.s.l. FAMILY EUCONULIDAE Euconulus fulvus (Müller, 1774): Brown Hive Records. Map 16. RBCM: , , , , , , , , , , , , , , , , , , , , , , , , , , , , , , , , , , RGF: , , , , , , , , , , CMN: 2859, Euconulus fulvus is perhaps the most common species in the upper Fraser Basin and northern B.C. and is a generalist, living in mesic to dry sites, both in forests and nonforested habitats. It occurs under dead wood, leaf litter, stones and vegetation, and it lives in leaf litter. E. fulvus is one of species most often found at or near the treeline. In the upper Fraser Basin, this species was found at elevations up to 2100 m a.s.l. Euconulus praticola (Reinhardt, 1883): Marsh Hive Records. Map 17. RBCM: RGF: While the majority of Euconulus in collections are attributable to Euconulus fulvus, there appears to be a second species that is identifiable by shell characters, body colour, and habitat preference. In British Columbia examples, E. praticola differs from E. fulvus by having a reddish rather than yellowish shell and shining rather than silken microsculpture on the apical surfaces of the whorls; the microsculpture on the base of the shell in E. praticola has relatively deeper spiral striae and stronger axial grooves than E. fulvus. The basal shell surface of E. fulvus varies among British Columbia material; some have the base almost without any microsculpture, while others have clear but relatively shallower spiral striae than E. praticola. There are also usually some irregularly spaced axial grooves present in B.C. examples of E. praticola, which are seldom ever as well developed in the other species. Animals of E. praticola are black throughout, including the mantle, whereas the body of E. fulvus is variously light to dark grey (paler on the sides of the foot) with the mantle spotted with black. Some authors have used the name Euconulus alderi (Gray in Turton, 1840) for the wetland species in Ontario and the U.S. Midwest (e.g., Grimm 1996; Nekola 2001; Nekola & Massart 2001), as has been done in Europe (e.g., Kerney and Cameron 1979). However, Falkner et al. (2002) recently treated E. alderi as a synonym of E. trochiformis (Montagu, 1803), a forest-dwelling species, and resurrected the name E. praticola for the dark-bodied wetland species in Europe. It is surprising that this wetland species has only recently been recognized in North America, as it seems to be quite distinct. However, wetland habitats are generally under-investigated for terrestrial gastropods, and the acceptance of a wetland congener to E. fulvus has been controversial, even in Europe. For example, Schileyko (2002) stated that there was probably only one variable Holarctic species despite several decades of literature stating otherwise. In addition to two records from the upper Fraser Basin, there are nine additional unpublished B.C. localities: two in the Bulkley Valley near Telkwa (Forsyth collection), four from the Columbia Basin (RBCM , , , ), and three from Greater Vancouver and the lower Fraser Valley (Forsyth collection). Unlike Euconulus fulvus, E. praticola is found only at wet sites: marshes, forested floodplains and grassy shorelines of waterways. Elevations of the known sites range from sea level to 760 m a.s.l.; in the upper Fraser Basin, both known localities are at ca 760 m a.s.l. The two species of Euconulus were not sympatric at any of these localities. E. praticola is often found along with characteristically wetland species such as Vertigo ovata, Zonitoides nitidus, various unidentified species of succineids, and amphibious lymnaeids. FAMILY GASTRODONTIDAE Zonitoides arboreus (Say, 1816): Quick Gloss Records. Map 18. RBCM: , , , , , These four lots were included under Euconulus fulvus in Terrestrial Gastropods of the Columbia Basin, British Columbia (Forsyth 1999).

12 12 LIVING LANDSCAPES: UPPER FRASER BASIN 004, , , , , , , , , , , , , , , , , , , RGF: , , , , , , , CMN: Zonitoides arboreus is widespread and common in northern British Columbia, often found along with Discus whitneyi and Euconulus fulvus in coniferous, mixed-wood and deciduous forests. It is typical of dry to mesic forests in the upper Fraser Basin, but also has occasionally been found along the margins of water bodies. In the upper Fraser Basin this species has been found at elevations between 460 m and 1040 m a.s.l. FAMILY DAUDEBARDIIDAE Nesovitrea binneyana (Morse, 1864): Blue Glass Records. Map 19. RBCM: , , , , , , , , , , , , , RGF: , , This species is widespread across B.C., where it is found mostly in mesic deciduous or mixed-wood forests. Occasionally it is found in open or forb-covered, disturbed ground. In the upper Fraser Basin, Nesovitrea binneyana was collected at m a.s.l. Nesovitrea electrina (Gould, 1841): Amber Glass Records. Map 20. RBCM: Nesovitrea electrina is widespread in B.C. but is less frequently encountered as the previous species. In the upper Fraser Basin, it was found only at McBride, where it was in cottonwood woods along the Fraser River. FAMILY VITRINIDAE Vitrina pellucida (Müller, 1774): Western Glass-snail Records. Map 21. RBCM: , , , , , , , , , , , , , , RGF: ; ; CMN: 83140, Vitrina pellucida occurs across the Palearctic and is widespread in western North America where it often goes by the synonym V. alaskana Dall All material collected in July 2001 fieldwork was either juvenile or were empty shells only, but dissections of snails from Dreyskull (ca 4 km N of Glendale; CMN 93333) and the Hazelton Mountains (Forsyth collection) show that the vas deferens is enclosed in a penial sheath. In this regard, their genital anatomy conforms to that of V. pellucida, as described by Forcart (19; see also Bequaert & Miller 1973, Roth & Lindberg 1981, and Roth & Sadeghian 2003). This species is common and widespread throughout British Columbia, living in a wide variety of habitats that are often seasonally dry. In the upper Fraser Basin, Vitrina pellucida the localities where it was found are between 660 m and 1220 m a.s.l., and all were below tree line. However, no alpine or subalpine habitat was surveyed in the upper Fraser Basin; elsewhere in B.C., V. pellucida has been located above 1220 m. For example, this species was living at 1740 m in the alpine tundra on Hudson Bay Mountain near Smithers (Forsyth collection). FAMILY LIMACIDAE Limax maximus Linnaeus, 1758: Giant Gardenslug Record. Map 22. RBCM: Limax maximus is an introduced western European slug, now widespread in southern British Columbia (Rollo & Wellington 1975; Forsyth 2004; RBCM collection). In the upper Fraser Basin, this species was collected a roadside park in Quesnel. Additional searches in urban areas probably would turned further records. FAMILY AGRIOLIMACIDAE Deroceras laeve (Müller, 1774): Meadow Slug Records. Map 23. RBCM: ; ; ; CMN: In the upper Fraser Basin Deroceras laeve was found along the shorelines of waterways and permanently wet forested sites. At Teardrop Lake (RBCM ) slugs were found under logs in a site that was clear-cut in 1985 and burnt in 1986; major vegetation consisted of willows and young conifers. Localities in the upper Fraser Basin are at elevations between 760 m and 880 m a.s.l. This is one of the most widespread species of slugs in the world, occurring on all continents except Antarctica (Wiktor 2000). Although introduced to many parts of the world, it is apparently, at least in part, native to North America. There is no evidence to suggest that the species is not native in the upper Fraser Basin, where it occurs in both disturbed and undisturbed habitats.

13 TERRESTRIAL GASTROPODS OF THE UPPER FRASER BASIN 13 Deroceras reticulatum (Müller, 1774): Grey Fieldslug Records. Map 24. RBCM: , , , CMN: 93332, 93333, 93339, In the upper Fraser Basin Deroceras reticulatum was found at an abandoned farm site, roadside highway pullouts, and a provincial park. This European slug is widely introduced into British Columbia and likely is a common synanthrope in the upper Fraser Basin. FAMILY ARIONIDAE Arion circumscriptus Johnston, 1828: Brown-banded Arion Records. Map 25. RBCM: , In the upper Fraser Basin, and around Canim Lake (outside the scope of this report), the introduced European slug Arion circumscriptus was found in disturbed habitats and coniferous and mixed-wood forests immediately adjacent to roads. Elsewhere in British Columbia, this species is recorded from Greater Vancouver (Rollo & Wellington 1975), southern Vancouver Island, the Queen Charlotte Islands, and the Bulkley Valley at Smithers (RBCM collection). This species was recognized from other species of Arion by having a false keel (present in the subgenus Carinarion) and the presence of black pigment spots on the mantle. These spots are believed to be a reliable character for distinguishing A. circumscriptus from other Carinarion [A. fasciatus (Nilsson, 1823); A. silvaticus Lohmander, 1937]. However, one animal was found at Hwy 16 near the Goat River (RBCM ) that when alive had bright orange-yellow lateral bands, suggestive of Arion fasciatus (Nilsson, 1823), but the dissection of this slug showed that it was juvenile the genitalia were undeveloped and therefore not identifiable with certainty (H. Reise, pers. comm.). Arion fasciatus, A. silvaticus and A. circumscriptus form a group (subgenus Carinarion) of very similar species that are at best only weakly distinguishable by external pigmentation; a recent study has further determined that diet can influence body pigmentation in these species (Jordaens et al. 2001). Thus, the strongly yellow-orange pigmentation of this individual is not conclusive. There seems to be no anatomically verified records of Arion fasciatus from British Columbia, although records of A. fasciatus were cited in some older literature and most (or all) are probably misidentifications of A. circumscriptus. Arion subfuscus (Draparnaud, 1805): Dusky Arion Records. Map 26. RBCM: Arion subfuscus is another introduced European slug in the upper Fraser Basin. Although there is only one record of it from the area (in the campground at Ten Mile Lake Provincial Park), this species is likely both widespread and common. Elsewhere in northern British Columbia A. subfuscus is known from the Bulkley Valley, the Queen Charlotte Islands and Alice Arm (unpublished records, RBCM). Prophysaon andersonii (Cooper, 1872): Reticulate Taildropper Records. Map 27. RBCM: , , , , , , , CMN: 93338, In the upper Fraser Basin, this species was found in open, mixed-wood forests and pure stands of trembling aspen (at m a.s.l.) where it was usually found under logs or openly crawling. This species and Deroceras laeve are the only native species of slugs in the upper Fraser Basin. CONCLUSIONS Four species were most commonly encountered. Although none of the collections made in the upper Fraser Basin employed quantitative methods, a rough idea of common versus less common species can be inferred from the frequency that species are recorded from the region (see maps). The most collected species were Euconulus fulvus, Discus whitneyi, Vitrina pellucida and Nesovitrea binneyana. Wetlands are undersurveyed for terrestrial gastropods. Species listed in this report that are restricted to wetland habitats tend to have fewer locality records than those species that are generalists. While it is true that exclusively wetland species will be spotty in their distribution, wetlands are usually overlooked, and species such as Vertigo ovata and Euconulus praticola are probably under recorded. Exotic species are likely more widespread. Although well known to gardeners and backyard naturalists, exotic species are often overlooked because as everyday occurrences they may not be considered worthy to record. During the 2001 field survey, we did not especially target sites in towns of the upper Fraser Basin, so the relatively few collections of some synanthropic species, such as Limax maximus, Arion subfuscus, Cochlocopa lubrica, result from a lack of collecting effort in urban areas. However,

14 14 LIVING LANDSCAPES: UPPER FRASER BASIN certain exotic species were found more frequently the grey fieldslug, Deroceras reticulatum, was living in many roadside stops that we visited. Species expected but not found. One species, Discus shimekii, although historically recorded from the upper Fraser Basin (Stanwell-Fletcher & Stanwell-Fletcher 1943; Map 14), has not recently been collected. My current understanding of the distribution of this species would suggest that it certainly would not be present over much of the upper Fraser Basin (particuarly the central, southern and western portions). It is possible that this literature record of D. shimekii is based on a misidentification for D. whitneyi. Among native species of terrestrial gastropods, there are several species that have been recorded elsewhere in B.C. that were expected for the upper Fraser Basin but subsequently not found: Vallonia gracilicosta Reinhardt, 1883; Striatura pugetensis (Dall 1895); and Microphysula ingersollii (Bland, 1875). V. gracilicosta is known from scattered localities north, west and south of the upper Fraser Basin. S. pugetensis is known from the valley of the North Thompson River and from the B.C. coast (Forsyth 2001b). M. ingersollii lives in subalpine forests and avalanche tracks up to the treeline and is currently known from southeastern B.C. (south of near Field) and above Pine Pass in the Murray Range (Forsyth 2004). ACKNOWLEDGEMENTS I would like to thank Philip Lambert (Curator of Invertebrates, RBCM) for organizing and participating in the Living Landscapes fieldwork in Kelly Sendall (Senior Collections Manager, Ichthyology, Herpetology and Invertebrate Zoology, RBCM) offered his assistance while sorting, identifying and curating material. Dr Heike Reise (Curator of Molluscs, Staatliches Museum für Naturkunde Görlitz, Germany) helped with and commented on some of the slug identifications. Drs Jean- Marc Gagnon (Chief Collection Manager, Invertebrate Section, Canadian Museum of Nature, Ottawa) and Jochen Gerber (Invertebrates Collections Manager, Field Museum of Natural History, Chicago) allowed for the loan of specimens in their collections. Tammy Forsyth helped collect some of the material reported here. LITERATURE CITED BEQUAERT, J. C. & W. B. MILLER The Mollusks of the Arid Southwest with an Arizona Check List. Tucson: University of Arizona Press. xiv, 271 pp. FALKNER, G., T. E. J. RIPKEN & M. FALKNER Mollusques continentaux de France. Liste de Référence annotée et Bibliographie. Patrimoines naturels 52: FORCART, L. 19. Die nordischen Arten der Gattung Vitrina. Archiv für Molluskenkunde 84(4/6): 1 166, pl FORSYTH, R. G Terrestrial gastropods of the Columbia Basin, British Columbia. Royal British Columbia Museum, Living Landscapes. Available at landscapes.bc.ca/cbasin/molluscs/contents.html (ISBN: ). Accessed 25 June FORSYTH, R. G. 2001a. New records of land snails from the mountains of northwestern British Columbia. The Canadian Field-Naturalist 115(2): FORSYTH, R. G. 2001b. A note on the distribution of Striatura pugetensis in British Columbia. The Festivus 33(5): FORSYTH, R. G Land Snails of British Columbia. Victoria: Royal BC Museum. iv, 192 pp., [8] pls. GRIMM, F. W Terrestrial molluscs. In: Assessment of species diversity in the Mixedwood Plains Ecozone; edited by I. M. Smith. Ecological Monitoring and Assessment Network. Available at watch.ca/mixedwood/landsnai/ and on CD-ROM. Accessed 25 June HARRIS, S. A. & L. HUBRICHT Distribution of the species of the genus Oxyloma (Mollusca, Succineidae) in southern Canada and the adjacent portions of the United States. Canadian Journal of Zoology 60: JORDAENS, K., P. VAN RIEL, S. GEENEN, R. VERHAGEN & T. BACKELJAU Food-induced body pigmentation questions the taxonomic value of colour in the selffertilizing slug Carinarion spp. Journal of Molluscan Studies 67: KERNEY, M.P., & CAMERON, R.A.D A Field Guide to the Land Snails of Britain and North-west Europe. London: Collins. 288 pp., 24 pls. NEKOLA, J. C Distribution and ecology of Vertigo cristata Sterki, 1919 in the Western Great Lakes Region. American Malacological Bulletin 16(1/2): NEKOLA, J. C. & P. A. MASSART Distribution and ecology of Vertigo nylanderi Sterki, 1909 in the Western Great Lakes region. American Malacological Bulletin 16(1/2): PILSBRY, H. A Land Mollusca of North America (north of Mexico). The Academy of Natural Sciences of Philadelphia, Monographs 3, 1(1) [1939]: i xvii, 1 573, i ix. 1(2) [1940]: i viii, 1 994, i ix pp. 2(1) [1946]: frontispiece, i vi, (2) [1948]:

15 TERRESTRIAL GASTROPODS OF THE UPPER FRASER BASIN 15 i xlvii, pp. POKRYSZKO, B. M European Columella reconsidered (Gastropoda, Pulmonata, Vertiginidae). Malakologische Abhandlungen Staatliches Museum für Tierkunde Dresden 12(1): POKRYSZKO, B. M The Vertiginidae of Poland (Gastropoda: Pulmonata: Pupilloidea) a systematic monograph. Annales Zoologici 43(8): ROLLO, C. D. & W. G. WELLINGTON Terrestrial slugs in the vicinity of Vancouver, British Columbia. The Nautilus 89(4): ROTH, B., & D. R. LINDBERG Terrestrial mollusks of Attu, Aleutian Islands, Alaska. Arctic 34(1): ROTH, B. & P. S. SADEGHIAN Checklist of the land snails and slugs of California. Santa Barbara Museum of Natural History Contributions in Science 3: SCHILEYKO, A.A Treatise on Recent terrestrial pulmonate mollucs. Part 8: Punctidae, Helicodiscidae, Discidae, Cystopeltidae, Euconulidae, Trochomorphidae. Ruthenica Supplement 2: STANWELL-FLETCHER, J. F. & T. C. STANWELL-FLETCHER Some accounts of the flora and fauna of the Driftwood Valley region of north central British Columbia. Occasional Papers of the British Columbia Provincial Museum 4: TURGEON, D. D., J. F. J. QUINN, A. E. BOGAN, E. V. COAN, F. G. HOCHBERG, W. G. LYONS, P. M. MIK- KELSEN, R. J. NEVES, C. F. E. ROPER, G. ROSENBERG, B. ROTH, A. SCHELTEMA, F. G. THOMPSON, M. VEC- CHIONE & J. D. WILLIAMS Common and scientific names of aquatic invertebrates from the United States and Canada: Mollusks, 2nd edition. American Fisheries Society, Special Publication 26: i ix, WIKTOR, A Agriolimacidae (Gastropoda: Pulmonata) a systematic monograph. Annales Zoologici (Museum and Institute of Zoology, Polish Academy of Sciences) 49(3):

16 16 LIVING LANDSCAPES: UPPER FRASER BASIN

17 TERRESTRIAL GASTROPODS OF THE UPPER FRASER BASIN 17 DISTRIBUTIONAL MAPS Map 1 Cochlicopa lubrica Map 2 Vallonia excentrica Map 3 Vallonia pulchella Map 4 Zoogenetes harpa

18 18 LIVING LANDSCAPES: UPPER FRASER BASIN Map 5 Columella columella Map 6 Columella edentula Map 7 Vertigo arthuri Map 8 Vertigo cristata Map 9 Vertigo gouldii Map 10 Vertigo modesta

19 TERRESTRIAL GASTROPODS OF THE UPPER FRASER BASIN 19 Map 11 Vertigo ovata Map 12 Paralaoma servilis Map 13 Punctum randolphii Map 14 Discus shimekii (literature record) Map 15 Discus whitneyi Map 16 Euconulus fulvus

20 20 LIVING LANDSCAPES: UPPER FRASER BASIN Map 17 Euconulus praticola Map 18 Zonitoides arboreus Map 19 Nesovitrea binneyana Map 20 Nesovitrea electrina Map 21 Vitrina pellucida Map 22 Limax maximus

21 TERRESTRIAL GASTROPODS OF THE UPPER FRASER BASIN 21 Map 23 Deroceras laeve Map 24 Deroceras reticulatum Map 25 Arion circumscriptus. Open circle: A. fasciatus? Map 26 Arion subfuscus Map 27 Prophysaon andersonii

22 22 LIVING LANDSCAPES: UPPER FRASER BASIN

23 TERRESTRIAL GASTROPODS OF THE UPPER FRASER BASIN 23 APPENDIX Locality data for terrestrial gastropods in the upper Fraser Basin Localities are listed by station number for records from the Royal British Columbia Museum invertebrate collection and from the private collection of the author. Localities from CMN records are listed by lot catalogue number, since station numbers are not incorporated into the catalogue numbering system. Collection, Cat. No. Land Dist. Locality Name Nat. Topo. Ser. Map Lat Deg Lat Min Long Deg Long Min Collector(s) Date Elev. (m) a.s.l. 1 CMN 2859 Cariboo Prince George 093G G.W. Dawson 1875? 2 CMN 83080, Cariboo 8 km SW of Williams Lake 093B CMN 83140, 83141, Cariboo E shore, Fraser River, along Hwy 20????? F. Schueler; A. Schueler F. Schueler; A. Schueler 4 CMN Cariboo ca 4 km N of Cinema 093G F. Schueler 5 CMN Cariboo ca 4 km N of Glendale 093B F. Schueler 6 CMN 93339, Cariboo Hwy 97 at Salmon River 093J F. Schueler 7 CMN Cariboo NE of McLeese 093B F. Schueler 8 RBCM Lillooet near Ruth Lake 092P RBCM Cariboo Amanita Lake 093I RBCM Cariboo 11 RBCM Cariboo Teardrop Lake Rd, S of Tsayaz Lake Bowron Lake road, near Summit Creek 093J H RBCM Cariboo Shesta Lake 093G RBCM Lillooet Lac La Hache Park 092P RBCM Cariboo Pelican Forest Service Road 093G RBCM Cariboo E of Chief Lake 093J RBCM Cariboo 17 RBCM Cariboo Mount Robson Park: Portal Lake Mount Robson Park: Yellowhead Lake 083D D RBCM Cariboo McBride 093H RBCM Cariboo Hush Lake 093G RBCM Cariboo 21 RBCM Cariboo Robson Valley: near Spittal Creek Robson Valley: SE of Hile Creek 083E E RBCM Cariboo Robson Valley: La Salle Lakes 093H Aug Sep 1976? 26 Aug Aug Aug Aug ? Jun Jun Jun Jun Jun Jun Jun Jun Jun Jun Jun Jun Jun Jun Jun