Basis for Listing

The winged mapleleaf was thought to be extinct until a small population was discovered in the St. Croix River in 1987 (Doolittle 1988). Extensive surveys in the St. Croix River over the last 15 years indicate that the species is restricted to a 20 km (12 mi.) stretch south of Taylors Falls, Minnesota (Hornbach et al. 1996; Hove et al. 2003). This extant population occurs in runs over clean gravel, sand, and rubble substrates and in clear areas of high water quality. Hornbach (1992) concluded that the winged mapleleaf is only found under conditions that would be considered high quality for mussel habitat. The age structure of the individuals sampled indicates that recruitment to this population has been low in recent years. The youngest sizeable cohort is from 1984. Given it's extreme rarity, the winged mapleleaf was listed as federally endangered by the U.S. Fish and Wildlife Service in 1991. It has been state listed as endangered since 1996.

  Description

The shell of the winged mapleleaf is squarish, thick, moderately inflated, and up to 10 cm (4 in.) long. Externally it is brown, and young mussels have faint rays. The shell has a well-developed posterior wing with radiating ridges, and 2 rows of pustules or larger tubercles separated by a sulcus (depression or furrow) extending from the beak to the ventral margin. The pseudocardinal and lateral teeth are well-developed, and the inner shell is white. The winged mapleleaf is very similar to the mapleleaf (Quadrula quadrula), but the wing and 2 rows of pustules or larger tubercles on the winged mapleleaf's shell usually distinguish it from the mapleleaf.

  Habitat

Historically, the winged mapleleaf has been described as a large river species (Baker 1928). It has been found in the St. Croix River in riffles dominated by gravel, sand, and rubble substrates in water averaging about 1 m (3 ft.) deep (Hornbach et al. 1996). In general, winged mapleleafs have similar habitat requirements as the other mussel species residing in the St. Croix River mussel community. Three species of mussels, the deertoe (Truncilla truncata), the fawnsfoot (T. donaciformis), and the monkeyface (Quadrula metanevra), have been found to be significantly associated with winged mapleleafs (Hornbach et al. 1996).

  Biology / Life History

Mussels are long-lived animals. Members of many species may live for several decades and in rare instances, a century or more. The maximum lifespan of the winged mapleleaf is not known, but individuals up to 22 years of age have been collected from the St. Croix River (U.S. Fish and Wildlife Service 1997).

Mussels spend most of their lives buried in the bottom sediments of permanent water bodies, and often live in multi-species communities called mussel beds. They are primarily sedentary, but they can move around with the use of their foot, which is a hatchet shaped muscle that can be extended out between the valves (shells). A mussel will burrow its foot into the sediment and then contract it to pull itself slowly along the bottom of its aquatic habitat (Sietman 2003).

Mussels eat by filtering bacteria, protozoans, algae, and other organic matter out of the water. They draw water into their body through their incurrent siphon, remove food and oxygen with their gills, and then expel the filtered water through their excurrent siphon. Food particles are carried to the mussel's mouth by tiny hairlike cilia located on the gills. Waste is expelled through the excurrent siphon (Sietman 2003).

Mussels have a complex and distinctive reproductive cycle. Males release sperm into the water, which are drawn in by females through their incurrent siphon. Fertilized eggs are brooded in the female's gills, where they develop into tiny larvae called glochidia. The winged mapleleaf is presumed to brood between late May and July, and females contain glochidia in their gills in late September and early October (Heath et al. 2001). Once the glochidia are expelled from the female's gills, they attach to fish gills or fins by clamping onto them with their valves. The glochidia live as parasites on the host fish until they develop into juvenile mussels, at which point they detach from the fish and fall to the streambed as free-living mussels. The blue catfish (Ictalurus furcatus) and the channel catfish (I. punctatus) are potential fish hosts for the glochidia of the winged mapleleaf (Steingraeber et al. 2004).

  Conservation / Management

Endangered throughout its range, the following factors potentially threaten the continued existence of the winged mapleleaf: narrow range, sparse population and low reproduction, probability of inbreeding which could weaken the species genetically, low streamflow episodes, high streamflow variations on the St. Croix River caused by a hydroelectric dam operating on a seasonal peaking regime, and the threat of non-point and point source water and sediment pollution (U.S. Fish and Wildlife Service 1997). Dams, channelization, and dredging increase siltation, physically alter habitat conditions, and block the movements of fish hosts. The winged mapleleaf is also being impacted by the infestation of non-native zebra mussels (Dreissena polymorpha) in the Mississippi River and its tributaries. Zebra mussels can attach themselves in large numbers to the shells of native mussels, eventually causing death by suffocation. The health of winged mapleleaf host fish populations is also crucial to the recovery of this species.

The St. Croix population was thought to be the only extant population of this species until recently, when live specimens were found in the Ouachita River and tributaries in Arkansas, and the Bourbeuse River in Missouri. Surveys of the Ouachita River and tributaries in 2005 documented significant numbers of individuals, however the St. Croix River population is the only one known to be reproducing. Consequently, a single catastrophic event in the St. Croix River could effectively eliminate this species.

  Conservation Efforts in Minnesota

A recovery team consisting of biologists from the U.S. Fish and Wildlife Service, Minnesota and Wisconsin DNRs, the U.S. Geological Survey, and Macalaster College completed a winged mapleleaf recovery plan in 1997. The plan identifies and prioritizes actions necessary for the recovery of wing mapleleaf populations and provides specific criteria for delisting (U.S. Fish and Wildlife Service 1997). In 1992, the MDNR Division of Ecological Services conducted an instream flow study that resulted in stream flow recommendations aimed at protecting and conserving the winged mapleleaf and other mussel populations residing in the St. Croix River (Johnson 1995). Consequently, the operator of the dam just upstream of the the winged mapleleaf population is now coordinating with the Minnesota and Wisconsin DNRs and the U.S. Fish & Wildlife Service to insure that dam discharge and resultant instream flow levels are high enough to be protective of this rare mussel. Research is also being conducted by personnel from the Minnesota DNR, the Wisconsin DNR, the University of Minnesota, and Macalester College to determine habitat preferences and population dynamics of the winged mapleleaf. Since 2003, an interagency team has been conducting propagation work for this species. Channel catfish are being infested with glochidia and placed in cages in the Mississippi River, following a methodology developed for the Higgins eye mussel (Lampsilis higginsii). Juveniles have been documented in some cages, and have been left to develop to a larger size. Research is also being conducted by federal agency staff to develop tests of infestation of wild catfish. The National Park Service has posted signs at Interstate State Park prohibiting the handling of mussels. Lastly, multiple state and federal agencies are working to keep zebra mussels out of habitat occupied by winged mapleleafs in the St. Croix River.

  References and Additional Information

Baker, F. C. 1928. The fresh water mollusca of Wisconsin: part II: Pelecypoda. Wisconsin Geological and Natural History Survey Bulletin No. 70, Part II. University of Wisconsin, Madison. 495 pp.

Doolittle, T. C. J. 1988. Distribution and relative abundance of freshwater mussels in the Saint Croix National Scenic Riverway. Final report submitted to the Natural Heritage and Nongame Research Program, Minnesota Department of Natural Resouces. Unpaged.

Heath, D. J., R. L. Benjamin, M. B. Endris, R. L. Kenyon, and M. C. Hove. 2001. Determination of basic reproductive characteristics of the Winged Mapleleaf (Quadrula fragosa) relevant to recovery. Job 1: determination of gravidity period. Ellipsaria 3(1):18-19.

Hornbach, D. J. 1992. An examination of the population structure, community relationships and habitat characteristics for the Winged Mapleleaf mussel (Quadrula fragosa) at Interstate Park, Saint Croix River, Wisconsin and Minnesota. Final report submitted to the Minnesota Department of Natural Resources. 17 pp. + tables, figures, appendices.

Hornbach, D. J., J. G. March, T. Deneka, N. H. Troelstrup, Jr., and J. A. Perry. 1996. Factors influencing the distribution and abundance of the endangered Winged Mapleleaf Mussel Quadrula fragosa in the St. Croix River, Minnesota and Wisconsin. The American Midland Naturalist 136:278-286.

Hove, M., M. Berg, J. DeMarre, K. Dietrich, C. Gonzalez, K. Westberg, and D. Hornbach. 2003. Historical range expansion of Winged Mapleleaf provides possibilities for additional reintroduction sites. Ellipsaria 5(3):17-18.

Johnson, S. J. 1995. Instream flow requirement of Quadrula fragosa and the aquatic community in the Lower St. Croix River downstream from the Northern States Power hydroelectric dam at St. Croix Falls, Wisconsin. Report submitted to the Bureau of Endangered Resources, Wisconsin Department of Natural Resources. Madison, Wisconsin. 100 pp.

Sietman, B. E. 2003. Field guide to the freshwater mussels of Minnesota. Minnesota Department of Natural Resources, St. Paul, Minnesota. 144 pp.

Steingraeber, M., M. Hove, M. Bartsch, D. Hornbach, C. Nelson, T. Newton, J. Kalas, A. Kapuscinski, and E. Simonsen. 2004. Two fish species identified as hosts for Winged Mapleleaf (Quadrula fragosa). Ellipsaria 6(1):7-8.

United States Fish and Wildlife Service. 1997. Winged Mapleleaf Mussel (Quadrula fragosa) recovery plan. United States Fish and Wildlife Service, Ft. Snelling, Minnesota. 359 pp.