Hossack et al. Montana Branchiopoda 4/24/2016

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Fairy shrimp (order Anostraca) and tadpole shrimp (order Notostraca), members of the crustacean class Branchiopoda, are important components of many freshwater and saline wetlands (Dodson and Frey 2001, Brendonk et al. 2008). These animals are especially common in seasonal wetlands, where populations are maintained by banks of resting eggs that may remain dormant for decades or longer (Brendonck 1996). Both orders are distributed worldwide (Brendonk et al. 2008). Fairy shrimp are more prone to endemism, with several species listed as threatened or endangered, and new species are described frequently (e.g., Rogers 2006, Rogers et al. 2006). Most of the comparatively few species of tadpole shrimp are widespread, but there is 1 endemic described from central California (L. packardi) and the recently described Lepidurus cryptus in the Northwest (Rogers 2001).

We report fairy shrimp and tadpole shrimp collected during surveys for amphibians on Department of the Interior lands in western Montana from 2001 to 2008, primarily in Glacier National Park (NP). Glacier NP has an extensive history of limnology research but wetland invertebrates have not been well documented (Newell and Hossack, in review). Although western North America has been surveyed extensively for large branchiopod crustaceans, most effort in Montana has been focused on the Great Plains in the eastern portion of the state. We also provide an updated account of fairy shrimp and tadpole shrimp collected from Montana, as well as species that occur in surrounding areas and may eventually be documented in the state.
Study Area and Methods

Wetlands in Glacier NP have been surveyed for amphibians since 1999 as part of a long-term monitoring and research program by the U.S. Geological Survey (Corn et al. 2005). Glacier NP is a 4082 km² reserve that forms part of the Glacier-Waterton International Peace Park along the U.S.–Canada border. Elevations range from approximately 950 m to 3190 m. Much of the park is characterized by U-shaped valleys that reflect the extensive glaciation of the region during the Pleistocene (Carrara 1989). West of the Continental Divide, the park is dominated by a moist Pacific maritime climate. East of the Divide, the continental-polar air masses result in a colder, drier climate (Carrara 1989). Topographic and climatic variation in the park produce a variety of habitats, ranging from western red cedar (Thuja plicata)-western hemlock (Tsuga hetrophylla) forests typical of the Pacific Northwest, to alpine tundra and grasslands.

Local wildlife refuges, including the National Bison Range and Lost Trail National Wildlife Refuge (NWR), have also been inventoried periodically for amphibians since 2001. The National Bison Range (elevation range: 788 to 1489 m), part of the U.S. Fish and Wildlife Service’s Wildlife Refuge System, is composed of Palouse Prairie grasslands and mountain forest. Lost Trail NWR (elevation range: 1064 to 1403 m) is in an intermountain valley surrounded by public and private forest that is managed for timber production. The valley bottom is covered with prairie grasses and weeds and has extensive wetlands. Most of the wetlands have been created or altered in support of previous livestock operations on the property, but there are also several natural wetlands on the west side of the refuge that dry annually.

We sampled a wide variety of wetlands, ranging from temporary marshes and pools to large cirque lakes, between mid-May and mid-August. With a few exceptions, most water bodies had near-neutral pH and specific conductance < 300 μS/cm. Surveys were timed to coincide with the approximate 6–8 wk period when amphibian larvae were expected to be present and free-swimming. Crews searched shallow areas ( 0.5 m) of each site, using a dip net (3.2 mm mesh) to sweep through vegetation and to capture amphibian larvae. Because our field efforts were directed towards amphibians rather than crustaceans, we only sampled water bodies that were likely to have water long enough for amphibian larvae to metamorphose. By excluding the most ephemeral water bodies, we did not sample many habitats that would have been suitable for branchiopod crustaceans.

Our collections were mostly opportunistic and our will not reflect the true extent of any species documented. The presence of fairy shrimp, especially, was frequently recorded by field technicians in Glacier NP during the early years of the study without the collection of specimens. Since 2001, however, when we first found tadpole shrimp in Glacier NP, crews were instructed to collect specimens whenever they were detected. Similarly, crews were instructed to collect fairy shrimp from all water bodies (374 surveyed) where they were detected in 2007 and 2008. All specimens were fixed in 10% formalin or 95% ethanol and identified at the University of Montana Flathead Lake Biological Station using standard taxonomic keys (Belk 2000, Rogers 2001, Rogers 2002). Species identifications were verified at EcoAnalysts, Inc. in Davis, California.

The list of confirmed and likely species for the state is from personal collections and records of D.C. Rogers. Collection abbreviations are as follows: DCR, D. Christopher Rogers personal collections; DB, Denton Belk collection (now at the U.S. Natural History Museum); U.S.N.M., U.S. Natural History Museum.

We collected 4 species of fairy shrimp (3 in Glacier NP) from 57 wetlands and 1 species of tadpole shrimp from 16 wetlands (Figure 1). All specimens were collected from valley-bottom or hillside water bodies. Fairy shrimp are common in some high elevation (including alpine) ponds elsewhere in the Rocky Mountains (e.g., Anderson 1974, Saunders et al. 1993, Bohonak and Whiteman 1999). We have not found fairy shrimp in high-elevation habitats in Glacier NP, but we have received reports of a high-elevation population near Snyder Lake.

We made the most collections of fairy shrimp in 2002 and 2008. The larger number of records in 2008 partially reflects sampling effort and a greater emphasis on preserving specimens, but similar effort in 2007 resulted in fewer observations. For example, we have sampled 1 wetland near Lake McDonald at about the same time (late May/early June) every year since 1999, and fairy shrimp were present in some years (e.g., 2002 and 2008) and not others.

We never found fairy shrimp at Lost Trail NWR or tadpole shrimp at the National Bison Range. Although co-occurrence within and between orders is common elsewhere (Dodson and Frey 2001), we found co-occurrence by 2 species of fairy shrimp and by a fairy shrimp and tadpole shrimp in only 1 wetland. Both cases of co-occurrence were near Lake Sherburne on the east side of the park. In Glacier NP, we found all 3 species of fairy shrimp only in the lower Lee Ridge area, in the northeast corner of the park where the Great Plains transition into montane forests. Coordinates for all wetlands where we collected specimens identifiable to species are provided in Appendix 1. In addition to those we collected, 9 species of fairy shrimp from 5 genera and 3 species of tadpole shrimp from 2 genera have been documented from Montana (Table 1).

Nine species of fairy shrimp and four species of tadpole shrimp have been previously reported from Montana. Our recent collections add 5 species of fairy shrimp and provide new location records for 16 species of fairy shrimp and tadpole shrimp, detailed below. Additionally, we provide comments on species that have the potential to be found in Montana during future surveys.
Order Anostraca Sars, 1867

Branchinecta paludosa (Müller, 1788)

We collected the circumpolar fairy shrimp (Branchinecta paludosa) from 1 pond in each of 3 drainages (Cutbank, Lake Sherburne, Lee Creek) on the east side of the park (Figure 1). All ponds were in mid-elevation (1708–1900 m), mixed aspen-conifer forests. Branchinecta paludosa is primarily a circumpolar arctic species but in North America its distribution extends southward along the Rocky Mountains (Saunders et al. 1993, Maynard Stern and Belk 1999). This shrimp occupies a wide variety of habitats, including alpine ponds >3000 m elevation in Wyoming (Saunders et al. 1993). It has been collected previously near Glacier NP (Lynch 1958) and from just north of our study sites in Waterton Lakes NP, Canada (Anderson 1974).

We collected the Reading fairy shrimp (Branchinecta readingi) from a single pool in the Ravalli Potholes on the National Bison Range in 2002. Fairy shrimp were also noted from neighboring pools but were not collected. These pools fill only after heavy spring rains and have high specific conductance. When we collected B. readingi, specific conductance exceeded the limits of our meter (1990 μS/cm) and pH was 9.2.

We collected the smoothlip fairy shrimp (Eubranchipus intricatus) from only 2 ponds (1574 m and 1635 m elevation) in the northeast portion of Glacier NP (Figure 1), in similar habitats as B. paludosa. E. intricatus is distributed widely in montane ponds in Alberta and low salinity wetlands of the central prairies of Canada (Heartland-Rowe 1967, Anderson 1974) and has been documented previously east of the Rocky Mountains in Montana (Table 1). Aside from Montana, E. intricatus has been documented only in the states of Massachusetts (where it is a species of concern) and Minnesota (NatureServe 2008).

Collected from 51 sites, the ethologist fairy shrimp (Eubranchipus serratus) seems to be the most common and widely-distributed fairy shrimp in Glacier NP (Figure 1). Eubranchipus serratus was especially common in the Lake McDonald, Marias Pass, and Lee Ridge areas, although it was also found in many other low elevation areas around the perimeter of the park. This species is distributed across much of the continental U.S.A. as well as British Columbia (Belk 2000), and has been documented previouly in the Flathead and Ovando valleys in western Montana (Dexter 1953, Ludden 2000) and in northern Idaho (Rogers et al. 2006).

This species was not encountered during our study, and there are no records for this species in Montana. It is found in central and southern Wyoming (Rogers, 2006) and has since been found in southern Idaho, in rock outcropping type pools at City of Rocks National Reserve, Cassia County) and may possibly be found in Montana at some point in the future. This is the first record of this species from outside Wyoming. Casssia County: City of Rocks National Preserve, 42˚ 4’ 31.08’’, 113˚ 43’ 19.27’’, date unknown, M. Vinson.

The first record of this species comes from J. E. Lynch, with no locality data beyond “Montana”, deposited at the U.S.N.M. under accession number 294523. Cascade County: Near Great Falls, 30 June 1964, J. S. Laurie, U.S.N.M. 294523. Carter County: Rhodes Area ditch, 24 May 1995, D. L. Gustafson, Det. D. C. Rogers, DCR-569. Lewis & Clark County: Alkali Flats ditch, 26 July 1995, D. L. Gustafson, Det. D. C. Rogers, DCR-579. Powder River County: Otter area pool, 25 May1995, D. L. Gustafson, Det. D. C. Rogers, DCR- 572.

This species was not encountered during our study, and there are no records for this species in Montana. It is endemic to the Snake River Plateau in Idaho, but may be found in Montana in the future.

We collected the Couese tadpole shrimp (Lepidurus couesii) from several sites along the east side of Glacier NP and at Lost Trail NWR. Lepidurus couesii is distributed throughout much of western North America, including on the plains of eastern Montana where the type specimen was collected (Packard 1883, Rogers 2001). In the park, L. couesii was found most frequently in the Railroad Creek area and seemed patchily distributed elsewhere along the eastern boundary.

All specimens collected from Glacier National Park, the National Bison Range, and Lost Trail National Wildlife Refuge are archived at the University of Montana, Flathead Lake Biological Station. We are grateful to D. L. Gustafson and C. H. Eriksen for providing specimens. We thank XX and XX for comments that improved the manuscript. Mention of trade names does not imply endorsement or approval by the U.S. Government.

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Figure 1. Locations where we collected 3 species of fairy shrimp and 1 species of tadpole shrimp (Lepidurus couesii) in Glacier National Park (GNP). Detailed collection locations are not shown for Lost Trail National Wildlife Refuge (LTNWR) and the National Bison Range (NBR) because only 1 species was found on each refuge. The black line bisecting the park and the state is the Continental Divide.

Figure 1.

APPENDIX. Location and collection date of Anostraca (fairy shrimp) and Notostraca (tadpole shrimp) collected from the National Bison Range (NBR), Glacier National Park (GNP), and Lost Trail National Wildlife Refuge (LOTR), Montana. If specimens were collected from the same site more than once, only the first collection date is listed. Geographic coordinates are NAD83.