Gastropoda: Arionidae): New Distributional Records and Reproductive Anatomy

Western North American Naturalist

Volume 64 Number 4 Article 15

10-29-2004

Prophysaon coeruleum Cockerell, 1890, blue-gray taildropper (Gastropoda: Arionidae): new distributional records and reproductive anatomy

Kristiina Ovaska Victoria, British Columbia, Canada

William P. Leonard Olympia, Washington

Lyle Chichester Wellington, Florida

Thomas E. Burke Olympia, Washington

Lennart Sopuck Sidney, British Columbia, Canada

See next page for additional authors Follow this and additional works at: https://scholarsarchive.byu.edu/wnan

Recommended Citation Ovaska, Kristiina; Leonard, William P.; Chichester, Lyle; Burke, Thomas E.; Sopuck, Lennart; and Baugh, Jim (2004) "Prophysaon coeruleum Cockerell, 1890, blue-gray taildropper (Gastropoda: Arionidae): new distributional records and reproductive anatomy," Western North American Naturalist: Vol. 64 : No. 4 , Article 15. Available at: https://scholarsarchive.byu.edu/wnan/vol64/iss4/15

This Note is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Western North American Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Prophysaon coeruleum Cockerell, 1890, blue-gray taildropper (Gastropoda: Arionidae): new distributional records and reproductive anatomy

Authors Kristiina Ovaska, William P. Leonard, Lyle Chichester, Thomas E. Burke, Lennart Sopuck, and Jim Baugh

PROPHYSAON COERULEUM COCKERELL, 1890, BLUE-GRAY TAILDROPPER (GASTROPODA: ARIONIDAE): NEW DISTRIBUTIONAL RECORDS AND REPRODUCTIVE ANATOMY

Kristiina Ovaska1, William P. Leonard2, Lyle Chichester3, Thomas E. Burke4, Lennart Sopuck5, and Jim Baugh6

Key words: Prophysaon coeruleum, distribution, phylogeography, reproductive anatomy, British Columbia, Idaho, Pacific Northwest, Rocky Mountains, Washington.

The genus Prophysaon is composed of 9 ical variation is present in Oregon, and the recognized species of slugs native to temper- name P. coeruleum may apply to a species ate forests of western North America (Turgeon complex (Kelley et al. 1999). Herein we report et al. 1998). Differences in the anatomical on previously unknown populations of P. coeru- characteristics of the distal genitalia have been leum from southwestern British Columbia and used to identify 2 species groups (subgenera; northern Idaho. Pilsbry 1948): Mimetarion, which includes P. We collected P. coeruleum during surveys vanattae, P. obscurum, P. fasciatum, and P. for terrestrial gastropods on Vancouver Island, humile; and Prophysaon, which includes the British Columbia, Canada, and in Washington remaining species (P. andersoni, P. boreale, P. and northern Idaho, United States. On Van- coeruleum, P. dubium, and P. foliolatum). The couver Island we carried out time- and area- genus has a disjunct geographic distribution: constrained surveys along 100-m-long and 1- most species occur along the Pacific Coast m-wide transects in different forested habitats from southeastern Alaska to northern Califor- at 3 sites near Victoria (56 transects searched nia, but a few are present in the Rocky Moun- for a total of 72 person-hours) in March and in tains of northern Idaho and western Montana October–November 2002. In northern Idaho (Smith 1943, Pilsbry 1948, Frest and Johannes we visually searched approximately 30 sites for 2000, Leonard et al. 2003). One species (P. a total of ca. 200 person-hours between April humile) is restricted to the Rocky Mountains and October from 1999 to 2002. In Washington portion of the distribution. we conducted surveys on approximately 300 Prophysaon coeruleum Cockerell (1890) is a sites in the eastern Cascade Mountains and on small, poorly known slug of conservation inter- >130 sites between the Cascade Crest and the est with previously known range from western Pacific Ocean. In addition, one of us (TB) had Washington to northwestern California (Pilsbry access to the results of hundreds of surveys 1948, Kelley et al. 1999). The species has been conducted for “survey and manage” mollusks designated as a “survey and manage” species by the U.S. Forest Service on National Forests under the Northwest Forest Plan (USDA For- within those areas. est Service and USDI Bureau of Land Man- To confirm identification, we examined the agement 1994) due to its close association with reproductive system of the specimens with a older forests, few known localities, and lack of stereo-zoom microscope under 7.5–60X mag- information on its natural history and ecology nification (one specimen from site 2; two spec- (McGraw et al. 2002). Considerable geograph- imens from site 3 in Appendix). Specimens

14180 Clinton Place, Victoria, British Columbia V8Z 6M1, Canada. 2223 Foote Street NW, Olympia, WA 98502, USA. 32805 Greenbriar Boulevard, Wellington, FL 33414, USA. 44715-61st Avenue NE, Olympia, WA 98516, USA. 51759 Colburne Place, Sidney, British Columbia V8L 5A2, Canada. 62018 Dry Creek Road, Ellensburg, WA 98926, USA.

538 2004] NOTES 539

Fig. 1. Distribution map for locality records of Prophysaon coeruleum: solid circles = this study (numbers correspond to those in Appendix); open circles = previous localities (Pilsbry 1948, Branson and Branson 1984). were deposited in collections at the Carnegie several years). The species appears to have a Museum (CM), at the Royal British Columbia very restricted distribution in Canada. Museum (RBCM), and in the personal collec- In Idaho we found P. coeruleum at 2 of tion of T. Burke. Some specimens were sent to approximately 30 sites surveyed. The 2 sites Dr. Thomas Wilke at George Washington Uni- were about 30 km apart in the Couer d’Alene versity to be included in a genetic study. Lake watershed. These observations extend In British Columbia we found P. coeruleum the species’ known range approximately 400 along 2 of the 56 transects searched. These km eastward and represent the 1st records for localities were about 500 m apart on Rocky the Rocky Mountains (Fig. 1). Apparently, the Point Peninsula in the District of Metchosin, population in Idaho is disjunct from the more about 20 km southwest of Victoria on Vancou- western populations, as the intervening Colum- ver Island (records 1–2 in Appendix). These bia Basin consists of arid shrub-steppe habitat records extend the known distribution of the unsuitable for forest-dwelling gastropods (Bar- species approximately 120 km northwest from nosky et al. 1987, Franklin and Dyrness 1988). Washington State and represent the 1st records Brunsfeld et al. (2001) and Layser (1980) pro- from Canada (Fig. 1). Since 1999, two of us (KO vided numerous examples of fungi, plants, and and LS) have carried out surveys for forest- animals that have Pacific coastal and interior dwelling gastropods throughout Vancouver Rocky Mountain distributions, which mirror Island and on the southern coastal mainland of the distribution of western hemlock forest. British Columbia without locating this species Leonard et al. (2003) reported on a previously (about 170 sites surveyed, some intensively over undocumented example of the slug Prophysaon 540 WESTERN NORTH AMERICAN NATURALIST [Volume 64 dubium, Cockerell, from Idaho and cited addi- Externally, our specimens conform to the tional examples of invertebrates that have sim- description in Pilsbry (1948): live animals were ilar disjunct distribution patterns. gray with light blue flecking, the prominence In Washington we found P. coeruleum at 2 of which varied among localities. On all of the sites in the Cispus River watershed and at 1 specimens distinct, longitudinal grooves were site on Pin Creek, a tributary to the lower present on the foot behind the mantle, and the Columbia River. The slugs appeared to be foot margin was narrow with a distinct border most abundant at the Kraus Ridge site in the above (Fig. 2). The extended length of the Cispus watershed (record 5 in Appendix; 11 specimens ranged between 23 mm and 30 mm detections). One of us (TB) has found P. when alive and 12.5 mm and 15 mm after coeruleum at this site on several previous preservation. occasions. The Iron Creek site (record 6 in The genitalia of the new specimens from Appendix) is about 4 km southwest of the Idaho and British Columbia resemble those of Kraus Ridge site and on the opposite side of P. coeruleum from Washington illustrated by the Cispus River. In several visits we have Pilsbry (1948; Figure 378d), except that the found only a single specimen of P. coeruleum penis is much larger; in 1 of the 2 Idaho speci- at this site. There is a 3rd site in this area, mens it is almost as large as the muscular por- approximately 7 km northeast of the Kraus tion of the epiphallus (Fig. 3). Most of the long, Ridge site at 442 m elevation, from which a slender portion of the epiphallus is in a tangle single P. coeruleum was collected on 14 Novem- immediately anterior to the muscular portion. ber 2001. The specimen was identified by Tom The ovotestis is large and consists of approxi- Kogut, Cowlitz Ranger District biologist, but mately 24 egg-shaped lobules, with little pig- was not collected (T. Kogut personal commu- ment. The albumen gland is a small, appendix- nication). The species appears to have a very like structure at the terminus of the common restricted distribution in Washington. duct; it is not clearly demarcated from the com- Surveys by the U.S. Forest Service and U.S. mon duct either by color or texture, at least in Bureau of Land Management found no addi- the preserved animals. The duct of the semi- tional sites in Washington but found several nal receptacle is very long and slender and localities for the species in western Oregon loosely adherent to the common duct. The and also extended its range into northern Cali- vagina is slender and long, as is the entire fornia. However, because similar, undescribed common duct. species have also been found in the southern Interestingly, the genitalia of the new spec- Oregon Cascades (Kelley et al. 1999, USDA imens resemble Pilsbry’s (1948) Figure 376c Forest Service and USDI BLM 2000, Nancy of P. boreale from Oregon. However, his brief Duncan personal communication), there is description of the external morphology does some question whether the California speci- not match these specimens. Pilsbry (1948) mens indeed belong to this species. noted much individual variation in the shape All sites with P. coeruleum were in moist and size of the bulbous muscular portion of forest stands with a deciduous component of the epiphallus of P. coeruleum. Kelley et al. bigleaf maple (Acer macrophyllum), black cot- (1999) suggested that, as currently used, the tonwood (Populus balsamifera), or in 1 case name P. coeruleum might apply to a species quaking aspen (Populus tremuloides; Appendix). complex, particularly in Oregon where much One site was in an old-growth stand (Krause variation exits among specimens from differ- Ridge); the remaining sites were in 2nd-growth ent localities. A forthcoming molecular study stands. Sword ferns (Polystichum munitum), in- of P. coeruelum (T. Wilkie personal communi- dicating moist conditions, were typically pres- cation), including specimens from across the ent. The sites ranged from near sea level to species range, will provide important informa- about 650 m in elevation (Appendix). This tion regarding whether the taxa represents a species is thought to be associated with older polyphyletic group and will be critically impor- forests and moist plant communities (Kelley et tant for developing meaningful conservation al. 1999). Although not restricted to old-growth planning strategies. forests, it appears to require old-growth attri- butes, such as abundant coarse woody debris Funding for surveys on Vancouver Island and moist forest floor conditions. came from the Canadian Department of 2004] NOTES 541

Fig. 2. Specimens of Prophysaon coeruleum: A, Vancouver Island, British Columbia (locality 1); B, Kootenai County, Idaho (locality 4); C, Cowlitz County, Washington (locality 7); bar = ca. 5 mm. 542 WESTERN NORTH AMERICAN NATURALIST [Volume 64

last glaciation. Volume K-3. Geological Society of America, Boulder, CO. BRANSON, B.A., AND R.M. BRANSON. 1984. Distributional records for terrestrial and freshwater Mollusca from the Cascade and Coast Ranges, Oregon. Veliger 26: 248–257. BRUNSFELD, S.J., J. SULLIVAN, D.E. SOLTIS, AND P. S . S OLTIS. 2001. Comparative phylogeography of northwestern North America: a synthesis. Pages 319–339 in J. Sil- vertown and J. Antonovics, editors, Integrating eco- logical and evolutionary processes in a spatial con- text, Blackwell Science, Oxford. COCKERELL, T.D.A. 1890. New northwestern slugs. Nauti- lus 3:111–113. FRANKLIN, J.F., AND C.T. DYRNESS. 1988. Natural vegeta- tion of Oregon and Washington. Oregon State Uni- versity Press, Corvallis. 452 pp. FREST, T.J., AND E.J. JOHANNES. 2000. An annotated checklist of Idaho land and freshwater mollusks. Journal of the Idaho Academy of Science 36:1–51. KELLEY, R., S. DOWLAN, N. DUNCAN, AND T. B URKE. 1999. Field guide to survey and manage terrestrial mollusk species from the Northwest Forest Plan. USDI Bureau of Land Management, Oregon State Office, Portland. 114 pp. Layser, E.F. 1980. Flora of Pend Oreille County, Washing- ton. Washington State University Cooperative Exten- sion, Pullman. 146 pp. LEONARD, W.P., L. CHICHESTER, AND K. OVASKA. 2003. Prophysaon dubium Cockerell, 1890, the papillose taildropper (Gastropoda: Arionidae): distribution and anatomy. Nautilus 117:62–67. Fig. 3. Reproductive system of Prophysaon coeruleum MCGRAW, R., N. DUNCAN, AND E. CAZARES. 2002. Fungi and from Idaho (CM 64922; locality 3 in Appendix): EP = other items consumed by the blue-gray taildropper epiphallus, MB = muscular body of epiphallus, OVTES slug (Prophysaon coeruleum) and the papillose tail- = ovotestis, PE = penis, SPOV = spermoviduct, SR = dropper slug (Prophysaon dubium). Veliger 45:261–264. seminal receptacle, VA = vagina, VD = vas deferens. PILSBRY, H.A. 1948. Land Mollusca of North America (north of Mexico). Volume 2, part 2. Monographs of the Academy of Natural Sciences of Philadelphia (3), Philadelphia. 640 pp. National Defense. We thank Arthur Robinson SMITH, A.G. 1943. Mollusks of the Clearwater Mountains, for initiating these surveys. Barb Behan initi- Idaho. Proceedings of the California Academy of ated and funded the mollusk surveys for the Sciences 23(36):537–554. Lower Cispus Watershed. Tom Wilkie, Nancy TURGEON, D.D., J.F. QUINN, JR., E.V. COAN, F.G. HOCHBERG, W. G. L YONS, P.M. MIKKELSEN, R.J. NEVES, ET AL. 1998. Duncan, and Tom Kogut shared unpublished Common and scientific names of aquatic inverte- information. Robert Forsyth provided review brates from the United States and Canada: mollusks. comments. We thank Tara Chestnut, Brent 2nd edition. American Fisheries Society Special Haddaway, Kelley Jorgensen, Tom Kogut, Publication 26. 526 pp. Megan and Vicki Leonard, Laura Matthias, USDA FOREST SERVICE AND USDI BUREAU OF LAND MAN- AGEMENT. 1994. Record of decision for amendments Pat McQueary, Bill Null, Ana Licano, Noriada to the Forest Service and Bureau of Land Manage- Martinez, Brad Moon, Casey Richart, Robin ment Planning Documents within the range of the Shoal, Corey Strom, and Joan Ziegltrum for Northern Spotted Owl. [Portland, OR]: U.S. Depart- help in the field. ment of Agriculture, Forest Service; U.S. Department of Interior, Bureau of Land Management. ______. 2000. Final supplemental environmental impact LITERATURE CITED statement for amendment to the survey and manage, protection buffer, and other mitigation measures BARNOSKY, C.W., P.M. ANDERSON, AND P. J . B ARTLEIN. 1987. standards and guidelines Volume 1, Chapters 1–4. The northwestern U.S. during deglaciation: vegeta- tion history and paleoclimatic implications. Pages Received 28 May 2003 289–321 in W. F. Ruddiman and H.E. Wright, Jr., edi- Accepted 31 December 2003 tors, North America and adjacent oceans during the 2004] NOTES 543

APPENDIX. Localities for Prophysaon coeruleum exam- 4. Beauty Creek, Kootenai County, Idaho, USA; elevation ined for this study: CM = Carnegie Museum; RBCM = 650 m asl; 47°35.65′N, 116°39.0′W; ca. 60-year-old Royal British Columbia Museum; TEB = personal collec- stand of western redcedar (Thuja plicata) and black tion of Thomas E. Burke. cottonwood with an understory of red-osier dogwood (Cornus stolonifera), vine maple (Acer circinatum), and 1. Rocky Point, Vancouver Island, British Columbia, Can- pacific yew (Taxus brevifolia); 21 April 2002 (1 speci- ada; elevation <50 m above sea level (asl); 48°20.77′N, men collected by W. Leonard, T. Burke, and J. Baugh): 123°34.12′W; ca. 80-year-old stand of bigleaf maple TEB 02-080. (Acer macrophyllum) and Douglas-fir (Pseudotsuga men- ziesii) with sword fern (Polystichum munitum) under- 5. Kraus Ridge, Gifford Pinchot National Forest, Lewis story; 28 October 2002 (1 specimen collected by K. County, Washington, USA; elevation 460 m asl; Ovaska and L. Sopuk): RBCM 003-102-001. 46°25.98′N, 121°58.3′W; over 250-year-old stand of western hemlock, Douglas-fir, and bigleaf maple with 2. Rocky Point, Vancouver Island, British Columbia, Can- sword fern understory; 22 June 1995 (1 specimen ada; elevation <50 m asl; 48°20.44′N, 123°34.04′W); recorded by T. Burke); 26 September 1996 (2 speci- ca. 70-year-old stand of quaking aspen (Populus tremu- mens recorded by T. Burke); 29 May 1996 (2 speci- loides) with sword fern understory at the edge of small mens recorded by T. Burke); 13 November 1997 (5 wetland adjacent to a large, old coniferous stand; 18 specimens recorded by T. Burke); 13 December 2002 November 2002 (5 specimens collected by L. Sopuck (1 specimen collected by W. Leonard and T. Burke): and K. Ovaska): RBCM 003-103-001 (1 specimen); 1 sent to Dr. Thomas Wilkie at George Washington Uni- specimen donated for genetic studies by Dr. T. Wilke, versity for genetic studies. George Washington University, Washington, DC, USA; 2 specimens kept alive in the author’s collection. 6. Iron Creek, Gifford Pinchot National Forest, Lewis County, Washington, USA; elevation 350 m asl; 3. Chatcolet Lake, Benewah County, Idaho, USA; elevation 46°25.55′N, 121°59.05′W; 2nd-growth stand of western 650 m asl; 47°21.13′N, 116°46.68′W; ca. 70-year-old hemlock, Douglas-fir, and bigleaf maple with sword stand of western hemlock (Tsuga heterophylla), black fern understory; 7 October 1998 (1 specimen recorded cottonwood (Populus balsamifera), paper birch (Betula by T. Burke). papyrifera), and chokecherry (Prunus virginiana) with a sparse understory; 20 April 2002 (7 specimens detected, 7. Tributary to Pin Creek, 0.8 km east of Carrolls, Cowlitz 1 collected by W. Leonard, T. Burke, and J. Baugh): County, Washington; elevation 80 m asl; 46°04.58′N, TEB 02-059; 15 September 2002 (6 specimens col- 122°50.93′W; ca. 50-year-old stand of bigleaf maple, lected by W. Leonard, T. Burke, and J. Baugh): TEB western redcedar, and bitter cherry (Prunus emargi- 02-176 (1 specimen), CM 64922 (3 specimens). nata) with sword fern understory; 7 March 2003 (1 specimen collected by W. Leonard): CM 64963.