A Reassessment of High Quality Natural Communities on Camp Grayling

Prepared by: Michael A. Kost and Joshua G. Cohen

Michigan Natural Features Inventory P.O Box 30444 Lansing, MI 48909-7944

For: Michigan Department of Military and Veterans Affairs Camp Grayling, MI 49739

June 30, 2005

Report Number 2005-11

Cover photograph: Best Bog on Camp Grayling (M. Kost).

EXECUTIVE SUMMARY

This report presents the results of a reevaluation of 14 natural communities first identified in the early 1990s and discussed in Higman et al. (1994). Several significant changes have occurred since the natural communities were originally documented. Most notably, invasive species present a threat to many of the communities, particularly the mesic sand prairie at the Portage Lake Complex. This site contains a population of the invasive plant, leafy spurge (Euphorbia esula), that threatens to degrade habitat for numerous rare species including Houghton’s goldenrod (Solidago houghtonii), a federally threatened species. Several of the wetlands recently experienced fire, which appears to have bolstered species diversity and habitat heterogeneity in both the wetland and upland habitats. Water levels for many of the wetlands have dropped considerably since first surveyed in the 1990s. This phenomenon is likely the result of regional decreases in water levels tied throughout the Upper Great Lakes Region. Degradation by off road vehicles was observed in two wetlands but was not severe in either, indicating that education, enforcement, and deterrent efforts are having a positive affect. The report contains detailed site summaries and conservation and management recommendations for each natural community with the Results section entitled Site Descriptions and Management Recommendation. In addition, we present seven natural community abstracts in the Appendices (see List of Appendices for list of abstracts).

Grayling Community Assessment - i Grayling Community Assessment - ii TABLE OF CONTENTS

EXECUTIVE SUMMARY ...... i INTRODUCTION...... 1 METHODS ...... 1 RESULTS ...... 1 Site Descriptions and Management Recommendation ...... 1 1. Portage Creek-Howes Lake Complex ...... 2 2. Frog Lake Complex Frog Lake Complex...... 5 3. Lake Margrethe North...... 7 4. The Doughnut...... 10 5. Barker Creek Fen ...... 13 6. C-shaped Depression...... 16 7. Lovells Fen...... 19 8. Best Bog...... 21 9. Lovells Bog ...... 24 10. Crawford Red Pines ...... 26 11. Watson Swamp...... 29 12. Cannon Creek Meadow...... 31 13. Chub Creek Swamp...... 33 14. Beaver Creek...... 35 DISCUSSION ...... 37 Water Level Changes...... 37 Invasive Species...... 37 Fire Suppression...... 37 ORV Impacts ...... 38 CONCLUSION ...... 38 ACKNOWLEDGEMENTS ...... 39 LITERATURE CITED ...... 39 APPENDICES. NATURAL COMMUNITY ABSTRACTS...... 43

LIST OF TABLES

Table 1. Mesic sand prairie species composition by region...... 47

LIST OF PHOTOS

Photo 1. Prairie dropseed, a state threatened grass, dominates portions of the mesic sand prairie at the Portage Creek Complex...... 2

Photo 2. The Frog Lake Complex contains a diverse grouping of nine depressional wetlands that experience fluctuating water levels as evidenced by the concentric rings of differing vegetation in the photo above...... 5

Photo 3. The Lake Margrethe North intermittent wetland contains open Carex oligosperma-dominated sedge flats on organic soil interspersed with raised leatherleaf- and jack pine-dominated islands on acidic sands...... 7

Grayling Community Assessment - iii

Photo 4. The sedge, Carex oligosperma, commonly dominates the sedge flats within acidic wetlands such as intermittent wetland and bog...... 8

Photo 5. The Doughnut is an intermittent wetland that contains a ring of Carex oligosperma, which surrounds a leatherleaf- and tree-dominated center. Degradation by off road vehicles (ORVs) is evident within this wetland...... 10

Photo 6. The frame of an abandoned snowmobile occurs on the edge of the Doughnut intermittent wetland near the parking area and should be removed. Also visible are tracks from ORV use in the wetland...... 11

Photo 7. The Barker Creek northern fen is a level peatland dominated by slender woolly sedge with scattered, raised, peat mounds harboring sphagnum moss and shrubs (leatherleaf, sweet bay, shrubby cinquefoil and sage willow)...... 13

Photo 8. A thick layer of dried, sedge leaf litter blankets the organic soil at Barker Creek Fen and inhibits seed bank expression and species diversity. Prescribed fires conducted on an occasional basis act to reduce leaf litter in wetlands and uplands and foster seed bank expression and species diversity...... 14

Photo 9. Marl from a soil core is shown above along with silverweed, which dominates the exposed marl flats on the east side of the C-shaped depression northern fen...... 16

Photo 10. Following fire, rough fescue (state threatened, foreground) and big bluestem were growing robustly among the burnt snags of jack pine above the C-shaped depression northern fen. Nearby was a robustly flowering population of Hill’s thistle (special concern) also observed growing under burnt snags of jack pine...... 17

Photo 11. A fire recently burned across Lovells Fen and up the pine-dominated slopes, killing many of the pines. The leatherleaf (foreground) resprouted following burning...... 19

Photo 12. A small lake remains within peat-filled depression occupied by Best Bog...... 21

Photo 13. Sundew and sphagnum moss were abundant, floating in 20 cm of standing water within the Carex oligosperma-dominated, open sedge flats at Best Bog...... 22

Photo 14. A small lake surrounded by black spruce and jack pine remain near the center of Lovells Bog. Leatherleaf (foreground) dominates much of the open bog...... 24

Photo 15. Large red pines, approximately 180 years old and measuring 70(±) cm in diameter, dominate the canopy of the dry northern forest at Crawford Red Pines...... 26

Photo 16. At Crawford Red Pines, red maples are outcompeting red pine saplings. Prescription burning and manual removal should be used to reduce the density of red maple at this site...... 27

Photo 17. Cut stumps are common within the northeast portion of Watson Swamp where several living northern white cedars were aged to approximately 80 and 86 years old...... 29

Photo 18. The Cannon Creek Meadow is a northern wet meadow that is dominated by tussock sedge (Carex stricta) and occurs along Cannon Creek, a tributary to the Manistee River. Reed

Grayling Community Assessment - iv canary grass, an invasive species, was observed growing along South Sharon Road on the edge of the wetland and should be removed to prevent it from spreading further into the wet meadow...... 31

Photo 19. The Chub Creek Swamp is comprised of a northern shrub thicket that grades from shrubby wet meadow along Chub Creek to forested conifer swamp (rich conifer swamp) near the edge of the adjacent uplands. Periodic fires and flooding by beaver probably kept this wetland in a continual flux between wet meadow, shrub thicket, and rich conifer swamp...... 33

Photo 20. Tamarack has become very abundant within the Beaver Creek northern shrub thicket. In the absence periodic natural disturbances such as fire, beaver flooding, or insect outbreaks (e.g., larch sawfly infestation), the northern shrub thicket will eventually succeed to forested wetland...... 35

LIST OF APPENDICES

Appendix 1. Mesic Sand Prairie...... 45 Appendix 2. Intermittent Wetland...... 57 Appendix 3. Northern Fen ...... 65 Appendix 4. Poor Fen ...... 81 Appendix 5. Bog ...... 95 Appendix 6. Northern Wet Meadow...... 109 Appendix 7. Northern Shrub Thicket...... 117

Grayling Community Assessment - v Grayling Community Assessment - vi INTRODUCTION

During the summer of 2004 Michigan Natural such as fire suppression, invasive species, off road Features Inventory (MNFI) staff conducted vehicles (ORVs), etc., can cause significant surveys of fourteen high quality natural degradation within this time frame. This report communities that had been previously identified summarizes the findings of MNFI’s surveys, as part of an inventory effort on Camp Grayling in provides management recommendations aimed at the early 1990s (Higman et al. 1994). The surveys protecting biodiversity, and presents natural were initiated because it had been over ten years community abstracts for seven of the community since the high quality communities were identified types that occur on Camp Grayling. and potential threats to their ecological integrity

METHODS

Preparation for field surveys involved anthropogenic disturbances, community studying element occurrence records, aerial boundaries, and landscape context. photographs, topographic maps, and previously Following completion of the field surveys, the recorded site information (Higman et al. 1994). In data was analyzed and transcribed to update the addition, relevant ecological literature was element occurrence records in the statewide reviewed for each community type. A site packet biodiversity conservation database managed by was then assembled for each of the fourteen MNFI. The data from each element occurrence community occurrences, which contained relevant was also used to determine its Alliance according aerial photographs, topographic maps, element to the US National Vegetation Classification occurrence records, natural community forms, and (Faber-Langendoen 2001). Lastly, information pertinent literature. from the field surveys was used to produce site Community surveys were conducted during descriptions and conservation and management the summer of 2004. Each natural community recommendations for each natural community occurrence was visited one or more times over the occurrence, which appear within the Results summer to evaluate potential threats and collect section under Site Descriptions and additional information on soils, species Management Recommendations. composition, water levels, recent natural and

RESULTS

Fourteen community element occurrences from Higman et al. (1994). Invasive species were updated in the statewide biodiversity followed by an asterisk (*) are considered to be conservation database managed by MNFI. In highly invasive and efforts to control their spread addition, seven community abstracts were written and remove them from high quality natural and appear in the appendices. Site descriptions communities are considered a top management and management recommendations were objective for maintaining biodiversity. The highly produced and appear below. invasive species that were identified within the natural communities discussed below include the Site Descriptions and Management following: giant reed (Phragmites australis), reed Recommendation canary grass (Phalaris arundinacea), spotted knapweed (Centaurea maculosa), common St. The site descriptions that appear below are an John’s wort (Hypericum perforatum), and Canada updated version of the original site descriptions bluegrass (Poa compressa).

Grayling Community Assessment - 1 1. Portage Creek-Howes Lake Complex Natural Community Type: mesic sand prairie Rank: G1 S1?, critically imperiled Alliance: Andropogon gerardii - (Calamagrostis canadensis, Panicum virgatum) Herbaceous Alliance (V.A.5.N.a) Location: T26N R04W, T27N R04W, T26N R05W

M. Kost Photo 1. Prairie dropseed, a state threatened grass, dominates portions of the mesic sand prairie at the Portage Creek Complex.

Site Description: This mesic sand prairie is located in a three-mile-long band extending from the west side of Howes Lake to the southwest, parallel to Portage Creek. The prairie is divided into eighteen fragments totaling approximately 77 acres. This is a shrub/grass-dominated wetland that experiences significant water table fluctuation during the year. When the southern portion of this site was first located in June, 1992, surface soils were saturated. By September, the water table had dropped to 75 cm (30 in) below the surface. It is quite likely that these fragments flood most years in the spring. Soil organic matter at this site reaches a depth of 15.5 cm (6 in). Subsoil pH was measured at 7.0 (7.5). This site was first identified due to the presence of the federally threatened (FE) Houghton's goldenrod (Solidago houghtonii), which was previously known from around Howes Lake, and at the intersection of M- 72 and Arrowhead Road. Through analysis of aerial photos, proglacial landforms, and ground searches, it

Grayling Community Assessment - 2 was discovered that these small goldenrod populations were oriented in a linear fashion in what was a prairie-like habitat. It was discovered that these linear patches of prairie habitat were located in historical shoreline deposits from pro-glacial Lake Margrethe. By following these narrow depressions to the southwest, the full extent of this rare plant community was discovered. Jack pine (Pinus banksiana), and in places, tamarack (Larix laricina) are common tree species intermixed with the prairie, forming a widely scattered canopy. Dominant shrubs include speckled alder (Alnus rugosa), leatherleaf (Chamaedaphne calyculata), shrubby cinquefoil (Potentilla fruticosa), chokeberry (Aronia prunifolia), and several willows (Salix spp.). Common ground cover species include several sedges (Carex buxbaumii, C. leptalea, C. stricta, C. viridula), blue-joint grass (Calamagrostis canadensis), hair-grass (Deschampsia cespitosa), big bluestem (Andropogon gerardii), and marsh wild- timothy (Muhlenbergia glomerata). Other rare plant species found here include prairie dropseed (Sporobolus heterolepis, state threatened), Clinton's bulrush (Scirpus clintonii, state threatened), Vasey's rush (Juncus vaseyi, state threatened), and Long-leaved aster (Aster longifolius, special concern). In some fragments, prairie dropseed is the dominant species in the ground cover. Surveys of the community documented 91 vascular plant species at this site, indicating the relatively high diversity of the site. The secretive locust (Appalachia arcana, special concern) was also found within this complex. Surrounding the prairie fragments, spruce and tamarack-dominated swamp, second-growth jack pine forest and jack pine plantations are common. In the 1980s, jack pine was harvested and replanted along M- 72 surrounding the prairie remnants. Several roads pass through the prairie in a number of locations. Protection of this prairie habitat should include the establishment of upland buffer areas where intensive forest management and training activities will be excluded. Where possible, a 100-200 m buffer zone would likely be sufficient. Although existing roads probably pose little additional threat to the prairie fragments, thought should be given to the possibility of closing the dirt road east of the intersection at M-72 and Arrowhead Road. This is an often washed-out road that has Houghton's goldenrod growing along both its margins. Invasive species control at this site, especially for leafy spurge (Euphorbia esula) is a top priority. General Land Office survey notes indicate that wildfires once burned through the community as does its position within a large, level, outwash plain dominated by jack pine (Comer et al. 1995). Thus, maintenance of biodiversity at this site should include prescription burning on an occasional basis. Further research into the nature of the soils, parent material, and water table fluctuations of this site are needed. Monitoring of Houghton’s goldenrod should be reinitiated. For additional information on conservation and management of mesic sand prairies, see the natural community abstract in Appendix 1.

Soil: Information was gathered from thirteen soil cores taken within the dispersed patches of mesic sand prairie that were mapped in 1992. Below is a general description of soils compiled from the thirteen samples.

0 to 10 (20) cm: light-colored-loamy sand (pH 6.5 (7.0)) mixed with black organic matter (may also be described as black loamy sand) 10 to 50 (70) cm: iron-colored sand or occasionally light-colored sand (pH 7.0 (7.5)) with organic streaking and occasionally strong iron mottling 50 (70) cm to 130 cm: iron-colored sand or occasionally light-colored sand (pH 7.0 (7.5)) with strong iron mottling

Water table: No standing water was observed at the surface nor within the visible portions of holes made by the soil auger; however, saturated sands were encountered in many soil cores at depths ranging from 40 cm to 60 cm. Even at depths of 130 cm, saturated sands were not observed in some cores. Conversely, in one swale that was dominated by wetland plants but which also contained Houghton’s goldenrod, saturated, iron-colored sands were encountered at 20 cm.

Grayling Community Assessment - 3 Invasive species Scientific Name Common Name Estimated Abundance Agrostis gigantea redtop uncommon Centaurea maculosa* spotted knapweed* uncommon Euphorbia esula* leafy spurge* locally abundant Hieracium aurantiacum orange hawkweed locally common Hypericum perforatum* common St. John’s wort* occasional Poa compressa* Canada bluegrass* locally common Prunella vulgaris lawn prunella uncommon Tragopogon dubius goat's beard occasional

Conservation and Management: • Monitor for and remove invasive species, especially leafy spurge • Conduct prescribed burns on an occasional basis • Monitor population of Houghton’s goldenrod

Grayling Community Assessment - 4 2. Frog Lake Complex Frog Lake Complex Natural Community Type: intermittent wetland Rank: G3, S2; very rare globally, imperiled in state Alliance: Chamaedaphne calyculata - (Kalmia angustifolia) Seasonally Flooded Dwarf-Shrubland Alliance (IV.A.1.N.f) Location: T27N R02W

M. Kost Photo 2. The Frog Lake Complex contains a diverse grouping of nine depressional wetlands that experience fluctuating water levels as evidenced by the concentric rings of differing vegetation in the photo above.

Site Description: The Frog Lake Complex is a series of small lakes and wet depressions located seven miles north of M- 72 just west of Stephans Bridge Road. It was formed by melting ice blocks on sandy glacial deposits. The intermittent wetlands are formed in the smallest depressions and along several lake margins that experience water level fluctuations. The nine different depressions total approximately 10 acres. The complete complex of immediately adjacent uplands includes approximately 200 acres and occurs within the Pine Barrens Management Opportunity Area identified in Higman et al (1994) and the focus of Kost et al. (2000). Prairie cord grass (Spartina pectinata) typically dominates the edge of these wetlands, with running bog sedge (Carex oligosperma) and leatherleaf (Chamaedaphne calyculata) dominating zones with deeper water. Other common plant species include tall goldenrod (Solidago altissima), blue-joint grass (Calamagrostis canadensis), and wood sage (Teucrium canadense). Sixty eight plant species were documented in these wetlands. Although no listed species of plants or animals were located within the complex, two uncommon

Grayling Community Assessment - 5 leafhoppers (Notus spp. and Cicadula smithi) were found here. Adjacent uplands include pine barrens and second growth forest of jack pine (Pinus banksiana). Hill’s thistle (special concern) occurs within the adjacent pine barrens. Two track roads cross the area. Protection of this complex should involve limiting vehicle use to currently existing roads. Easy access to the south end of Frog Lake resulted in damage from off road vehicles during 1993. There may be no way to avoid this type of damage without closing immediately adjacent access roads. A vegetated upland buffer of at least 100 meters should be maintained around each wetland depression. The ecological integrity of these intermittent wetland communities may be directly related to the maintenance of open pine barren habitat in the immediately adjacent upland. Thus, management of this site should include the use of prescribed burning in the uplands and allowing fire to carry into the wetlands. Invasive species monitoring and control should be implemented within the wetlands and adjacent uplands. For additional information on conservation and management of intermittent wetlands, see the natural community abstract in Appendix 2.

Soil: Soil cores were examined in each of the nine depressional, intermittent wetlands that comprise the Frog Lake Complex. The information presented below is a general description compiled from these soil cores, which were generally very similar in composition and pH.

0 to 150 cm: fibric, sedge peat (pH 5.5)

Note: the sands around Frog Lake were alkaline (pH 8.0) and one very small depression at the south end of Frog Lake contained marl (pH 8.0).

Water table: No standing water was observed in any of the wetlands except for two depressions that contain lakes. No water could be observed within the holes produced by the soil auger; however, the fibric peat extracted from the soil cores was moist in all samples.

Invasive species: Scientific Name Common Name Estimated Abundance Centaurea maculosa* spotted knapweed* uncommon Phalaris arundinacea* reed canary grass* occasional Phragmites australis* giant reed* uncommon Poa compressa* Canada bluegrass* locally common

Conservation and Management: • Monitor for and remove invasive species • Limit vehicle use in area to currently existing roads • Maintain vegetated upland buffer of at least 100 meters around each depression • Conduct prescribed fire in adjacent uplands on an occasional basis and allow it to carry into the wetlands

Grayling Community Assessment - 6 3. Lake Margrethe North Community Type: intermittent wetland Rank: G3 S2, very rare globally, imperiled in state Alliance: Chamaedaphne calyculata - (Kalmia angustifolia) Seasonally Flooded Dwarf-Shrubland Alliance (IV.A.1.N.f) Location: T26N R04W

M. Kost Photo 3. The Lake Margrethe North intermittent wetland contains open Carex oligosperma-dominated sedge flats on organic soil interspersed with raised leatherleaf- and jack pine-dominated islands on acidic sands.

Grayling Community Assessment - 7

M. Kost Photo 4. The sedge, Carex oligosperma, commonly dominates the sedge flats within acidic wetlands such as intermittent wetland and bog.

Site Description: This 237 acre wetland is located one mile north of Lake Margrethe on a poorly drained glacial lake bed. Although surface soils here are medium sands, there is likely a layer of heavy soils several feet below the surface that is responsible for the fluctuation in water levels. According to the original land surveys of 1850, this wetland was mostly open, dominated by shrubs, sedges and grasses. Today, much of the area has closed in with jack pine, and occasionally, white pine and red pine forming a partial overstory canopy. Species such as leatherleaf (Chamaedaphne calyculata), speckled alder (Alnus rugosa), and sheep laurel (Kalmia angustifolia) dominate the shrub layer. Running bog sedge (Carex oligosperma) and sedge (Carex livida) dominate the ground layer, with the latter sedge dominating large, open portions of the complex. Just 22 plant species were noted in this wetland complex. An active bald eagle nest (Haliaeetus leucocephalus, federally threatened (FT)) is located at the north end of the wetland. The secretive locust (Appalachia arcana, special concern) was also found within this wetland. Over the years, this wetland has been impacted by road construction, timber harvest, and pine plantation establishment. Road construction may have caused slight alterations in the hydrology of this system. It is also possible that the area was historically maintained more open by periodic fires spreading from the adjacent, fire prone, jack pine plains. The suppression of wildfires during the past sixty years may also account for the development of an overstory canopy. Most of the tree plantations established at the east end of the complex failed shortly after establishment. Protection of this complex should include maintaining an adequate vegetated upland buffer. Consider limiting access to the community during the bald eagles nesting season. Where the road crosses the wetland, culverts should be installed under the road to allow for water flow. Tire tracks were observed within the

Grayling Community Assessment - 8 wetland during the 2004 surveys. To prevent further degradation of the natural community, vehicles (e.g., ORVs) should be prevented from entering the wetland. Consider conducting prescribed fire in wetland and surrounding upland on an occasional basis. Invasive species monitoring and control should be implemented within the wetland and adjacent uplands. For additional information on conservation and management of intermittent wetlands, see the natural community abstract in Appendix 2.

Soils: Within Carex oligosperma- and Carex aquatilis-dominated sedge flats: 0 to 25 cm: fibric, sedge peat (pH 4.5) 25+cm: coarse-textured loamy sand Water table: 10 cm below soil surface

Within slightly raised areas dominated by leatherleaf and widely scattered jack pine with sheep laurel (Kalmia angustifolia), blueberry (Vaccinium angustifolia), and wintergreen (Gaultheria procumbens): 0 to 5 cm: mull hummus (pH 4.5) 5 to 20m cm: grey-brown sand (pH 5.0). 20 to 50 cm: iron colored sand with large iron mottles 50 to 100 cm: saturated, light brown sand with few iron mottles Water table: 50 cm below soil surface

Invasive species: None observed

Conservation and Management: • Prevent use of ORV’s in wetland • Remove trash from adjacent parking areas • Conduct prescribed burn on an occasional basis • Limit access during bald eagle nesting season • Maintain vegetated upland buffer • Install culverts under roads that cross wetland • Monitor for and remove invasive species

Grayling Community Assessment - 9 4. The Doughnut Natural Community Type: intermittent wetland Rank: G3 S2, very rare globally, imperiled in state Alliance: Chamaedaphne calyculata - (Kalmia angustifolia) Seasonally Flooded Dwarf-Shrubland Alliance (IV.A.1.N.f) Location: T27N R04W

M. Kost Photo 5. The Doughnut is an intermittent wetland that contains a ring of Carex oligosperma, which surrounds a leatherleaf- and tree-dominated center. Degradation by off road vehicles (ORVs) is evident within this wetland.

Grayling Community Assessment - 10

M. Kost Photo 6. The frame of an abandoned snowmobile occurs on the edge of the Doughnut intermittent wetland near the parking area and should be removed. Also visible are tracks from ORV use in the wetland.

Site Description: This nine acre wetland is located just northwest of Howes Lake. It lies in a shallow depression on the wide outwash channel which includes the riverbed of the Manistee River. This site takes its name from its vegetative structure, which includes an outer ring dominated by running bog sedge (Carex oligosperma) and leatherleaf (Chamaedaphne calyculata), surrounding a tree-dominated center, with jack pine (Pinus banksiana) and black spruce (Picea mariana). Other common plant species include blue joint (Calamagrostis canadensis), thin grass (Agrostis perennans), laurel (Kalmia polifolia), and large cranberry (Vaccinium macrocarpon). Only 23 vascular plant species were noted in this wetland. The secretive locust (Appalachia arcana, special concern) was found in and around this wetland. Historically, in the immediately adjacent uplands, jack pine barrens and closed-canopy, jack pine forest was characteristic. Today much of this area remains as second growth jack pine forest and burned openings. A narrow two track road passes along the west side of the wetland, and there has been garbage dumped along the wetland margins. Protection of this wetland should include maintaining an upland vegetated buffer for at least 100 meters. Fires occurring in the adjacent uplands should be allowed to carry into the wetland. Tire tracks were observed within the wetland during the 2004 surveys. To prevent further degradation of the natural community, vehicles (e.g., ORVs) should be prevented from entering the wetland. Consider managing the adjacent uplands with prescribed fire on a periodic basis and allowing fire to carry into the wetland. Invasive species monitoring and control should be implemented within the wetland and adjacent uplands. For additional information on conservation and management of intermittent wetlands, see the natural community abstract in Appendix 2.

Grayling Community Assessment - 11 Soils: 0 to 110 cm: muck (pH 4.5 (4.0)) 110+ cm: coarse-textured, gleyed sand

Water table: 10 to 20 cm of standing water in Carex oligosperma-dominated outer ring

Invasive species: None observed

Conservation and Management Recommendations: • Prevent use of ORV’s in wetland. • Consider installing a barrier at parking area to prevent ORV use of wetland • Conduct prescribed fire in adjacent uplands and allow fire to carry into wetland • Maintain vegetated upland buffer for at least 100 meters • Remove abandoned snowmobile from edge of wetland • Monitor for and remove invasive species

Grayling Community Assessment - 12 5. Barker Creek Fen Natural Community Type: northern fen Rank: G3 S3, very rare and local throughout range Alliance: Carex lasiocarpa Saturated Herbaceous Alliance (V.A.5.N.m) Location: T27N R02W

M. Kost Photo 7. The Barker Creek northern fen is a level peatland dominated by slender woolly sedge with scattered, raised, peat mounds harboring sphagnum moss and shrubs (leatherleaf, sweet bay, shrubby cinquefoil and sage willow).

Grayling Community Assessment - 13

M. Kost Photo 8. A thick layer of dried, sedge leaf litter blankets the organic soil at Barker Creek Fen and inhibits seed bank expression and species diversity. Prescribed fires conducted on an occasional basis act to reduce leaf litter in wetlands and uplands and foster seed bank expression and species diversity.

Site Description: The Barker Creek Fen is a 31 acre wetland located one mile south of North Down River Road at the southeast extreme of North Camp Grayling property. Northern fens are groundwater fed wetlands that are typically located in areas containing porous, calcium and magnesium rich soils. This wetland is located on a poorly drained portion of outwash deposits. Water from this wetland flows south into the Au Sable River. This is a large, sedge-dominated wetland with shrubs and scattered trees located on a few rises. Dominant herbaceous species include slender woolly sedge (Carex lasiocarpa), sedge (Carex livida), slender bulrush (Scirpus hudsonianus), and pitcher plant (Sarracenia purpurea). Common shrub and tree species include shrubby cinquefoil (Potentilla fruticosa), bog birch (Betula pumila), leatherleaf (Chamaedaphne calyculata), sweet bay (Myrica gale), sage willow (Salix candida), and tamarack (Larix laricina). A total of 34 vascular plant species were noted in this wetland. Adjacent uplands are dominated by trembling aspen (Populus tremuloides). Recent clearcuts are evident on the east side of the complex with approximately one hundred meters of natural buffer remaining. These groundwater-fed wetlands are considered to be sensitive to increased nutrient run-off from adjacent land management activities. Protection of this wetland should include the establishment and maintenance of a buffer for 100 to 200 meters on all sides. The wetland contains a thick layer of leaf litter that acts to stifle seedling establishment and seed bank expression. Consider managing the adjacent uplands with prescribed fire and allowing fire to carry into wetland and reduce thickness of leaf litter and stimulate seed bank expression. Invasive species monitoring and control should be implemented within the wetland

Grayling Community Assessment - 14 and adjacent uplands. For additional information on conservation and management of northern fens, see the natural community abstract in Appendix 3.

Soil: 0 to 120 cm: fibric peat (pH 7.0) 120+ cm: coarse-textured gleyed sand (pH 8.0)

Water table: 3 to 5 cm of standing water covers most of sedge-dominated portions of wetland. Depth of standing water is significantly higher near Barker Creek. No standing water occurs on the scattered raised peat mounds, which are dominated by sphagnum moss and shrubs (leatherleaf, sweet bay, shrubby cinquefoil and sage willow).

Invasive species: none observed

Conservation and Management: • Monitor for and remove invasive species • Maintain vegetated buffer of 100 to 200 meters around wetland • Allow fires occurring in the surrounding uplands to carry into wetland • Prevent ORV use in wetland

Grayling Community Assessment - 15 6. C-shaped Depression Community Type: northern fen Rank: G3 S3, very rare and local throughout range Alliance: Carex lasiocarpa Saturated Herbaceous Alliance (V.A.5.N.m) Location: T27N R02W

M. Kost Photo 9. Marl from a soil core is shown above along with silverweed, which dominates the exposed marl flats on the east side of the C-shaped depression northern fen.

Grayling Community Assessment - 16

M. Kost Photo 10. Following fire, rough fescue (state threatened, foreground) and big bluestem were growing robustly among the burnt snags of jack pine above the C-shaped depression northern fen. Nearby was a robustly flowering population of Hill’s thistle (special concern) also observed growing under burnt snags of jack pine.

Site Description: This small eight acre wetland is located three miles north of North Down River Road just west of Stephans Bridge Road. It takes its name from the shape of the 50 foot deep depression where it is located. This site was originally classified as a poor fen but upon closer examination of the alkaline, organic soils, it was reclassified as a northern fen. A narrow band of exposed sand around the margin of this wetland indicates a small degree of regular water level fluctuations. Dominant plant species in this wetland include blue-joint grass (Calamagrostis canadensis), wool grass (Scirpus cyperinus), twig rush (Cladium mariscoides), meadow sedge (Carex stricta), slender woolly sedge (Carex lasiocarpa), and grass-leaved goldenrod (Euthamia graminifolia). In 2004, the east side of the wetland contains an exposed marl flat dominated by silverweed (Potentilla anserina). A total of 34 vascular plant species were noted in this wetland. The surrounding steep slopes above the wetland are dominated by second growth jack pine (Pinus banksiana) and trembling aspen (Populus tremuloides). A fire burned across portions of the wetland and jack pine-forested uplands in the spring of 2004. Although many of the jack pine were killed, two rare plants, rough fescue (Festuca scabrella, threatened) and Hill’s thistle (Cirsium hillii, special concern), and clumps of big bluestem (Andropogon gerardii) appeared to be reaping the benefits of the added sunlight and nutrients as these species were observed growing robustly and in full flower. Protection of this wetland should include monitoring for and removing invasive species. When possible, fire occurring in the adjacent uplands should be allowed to carry into the wetland. A vegetated upland buffer

Grayling Community Assessment - 17 should be maintained for approximately 200 meters on all sides of the wetland. For additional information on conservation and management of northern fens, see the natural community abstract in Appendix 3.

Soil: West side: 0 to 150+ cm: fibric peat (pH 7.0) East side: 0 to 50+ cm: marl (pH 8.0)

Water table: West side: saturated peat throughout but standing water was observed only in one small area that contained a quaking mat of vegetation. East side: marl was moist but no water observed

Invasive species Estimated Abundance Scientific Name Common Name Estimated Abundance Typha angustifolia narrow-leaved cattail uncommon Phalaris arundinacea* reed canary grass* occasional

Conservation and Management: • Monitor for and remove invasive species • Conduct prescribed fire in adjacent uplands on an occasional basis and allow fire to carry into wetland • Maintain vegetated upland buffer for approximately 200 meters

Grayling Community Assessment - 18 7. Lovells Fen Community Type: poor fen Rank: G3 S3, very rare and local throughout range Alliance: Carex oligosperma - Carex lasiocarpa Saturated Herbaceous Alliance (V.A.5.N.m) Location: T27N R02W, T28N R02W

M. Kost Photo 11. A fire recently burned across Lovells Fen and up the pine-dominated slopes, killing many of the pines. The leatherleaf (foreground) resprouted following burning.

Site Description: Lovells fen is a 27 acre poor fen located two miles south of the town of Lovells. It developed in a deep depression formed by a remnant ice block that melted in sandy glacial deposits. Generally, poor fens are partially ground water-fed wetlands where soils are somewhat less rich in calcium and magnesium than those found in northern fens. As a result, they typically include zones of vegetation characteristic of both northern fens and bogs. Lovells fen includes a small open water zone at its southern end, then grades from a sedge-dominated meadow to a more bog-like, shrub-dominated zone at the north end. Dominant plant species include water- lily (Nymphaea odorata), hard-stemmed bulrush (Scirpus acutus), blue-joint grass (Calamagrostis canadensis), slender woolly sedge (Carex lasiocarpa), twig rush (Cladium mariscoides), leatherleaf (Chamaedaphne calyculata), and pitcher plant (Sarracenia purpurea). A total of 55 vascular plant species were noted in this wetland. In the recent past, the steep slopes that border the fen were dominated by second-growth jack pine (Pinus banksiana) and red pine (Pinus resinosa), some of which was established by plantation. However, a fire recently burned across the wetland and up the slopes, killing many of the pines. In addition, the pine plantation along the north and east sides of the wetland was recently clearcut, disked, and planted to red pine. Signs of a small tornado touch down within the deep depression were

Grayling Community Assessment - 19 evident in early 1993 when most of the trees along the north end of the depression were observed blown down in a northeastern direction. Protection of this wetland should include the maintenance of a vegetated upland buffer for approximately 200 meters on all sides of the wetland. Fires occurring within the surrounding uplands should be allowed to spread into the wetland. Invasive species monitoring and control should be implemented within the wetland and adjacent uplands. For additional information on conservation and management of poor fens, see the natural community abstract in Appendix 4.

Soil: fibric sedge peat (pH 6.5)

Water table: ranges from 5 cm below soil surface on north end to a small area of standing water in the far southern portion

Invasive species Estimated Abundance Scientific Name Common Name Estimated Abundance Phalaris arundinacea* reed canary grass* locally common

Conservation and Management: • Monitor for and remove invasive species • Maintain vegetated upland buffer • Allow fires occurring in the surrounding uplands to carry into wetland • Prevent ORV use in wetlands

Grayling Community Assessment - 20 8. Best Bog Community Type: bog Rank: G3 S3, very rare and local throughout range Alliance: Chamaedaphne calyculata Saturated Dwarf-Shrubland Alliance (IV.A.1.N.g) Location: T27N R01W

M. Kost Photo 12. A small lake remains within peat-filled depression occupied by Best Bog.

Grayling Community Assessment - 21

M. Kost Photo 13. Sundew and sphagnum moss were abundant, floating in 20 cm of standing water within the Carex oligosperma-dominated, open sedge flats at Best Bog.

Site Description: This 25 acre bog is located just south of North Down River Road near the southeastern extreme of the camp property. It lies in a poorly drained portion of outwash deposits that drains to the southwest into Barker Creek. The south end of this wetland includes a well developed floating mat and a small area of open water. Throughout the remainder of the wetland is a mosaic of sedge-dominated meadow, shrub-dominated rises, and several islands with black spruce (Picea mariana), and tamarack (Larix laricina). Dominant vascular plant species include leatherleaf (Chamaedaphne calyculata), running bog sedge (Carex oligosperma), white beak rush (Rhynchospora alba), large cranberry (Vaccinium macrocarpon) and, locally, round-leaved sundew (Drosera rotundifolia). Common sphagnum mosses in this wetland include Sphagnum cuspidatum, Sphagnum papillosum, and Sphagnum magellanicum. Twenty five vascular plant species were noted in this bog. When the bog was first document in 1993 the adjacent uplands were dominated by second growth jack pine (Pinus banksiana) and red pine (Pinus resinosa); since then, this forest has been clearcut, disked to create furrows, and planted with jack pine. A road passes along the southwest and western border of the wetland. Protection of this wetland should include maintaining the culverts in the road at the south end of the wetland. A vegetated upland buffer for approximately 200 meters on all sides of the wetland should be maintained. Fires occurring in the adjacent uplands should be allowed to carry into the wetland. Invasive species monitoring and control should be implemented within the wetland and adjacent uplands. For additional information on conservation and management of bogs, see the natural community abstract in Appendix 5.

Grayling Community Assessment - 22

Soil: 50 to 60 cm of fibric peat with pH of peat 4.5 (4.0) over coarse-textured sand

Water table: 20 cm of standing water in sedge meadow areas dominated by Carex oligosperma and floating sphagnum moss and sundew

Invasive species: none observed

Conservation and Management: • Monitor for and remove invasive species • Maintain culverts in road at south end of the wetland • Conducts prescribed fire in adjacent uplands on an occasional basis and allow fire to carry into wetland • Maintain a vegetated upland buffer for approximately 200 meters on all sides of the wetland

Grayling Community Assessment - 23 9. Lovells Bog Community Type: bog Rank: G3 S3, very rare and local throughout range Alliance: Chamaedaphne calyculata Saturated Dwarf-Shrubland Alliance (IV.A.1.N.g) Location: T28N R02W

M. Kost Photo 14. A small lake surrounded by black spruce and jack pine remain near the center of Lovells Bog. Leatherleaf (foreground) dominates much of the open bog.

Site Description: Lovells Bog is located approximately two miles northeast of KP Lake along KP Twin Bridge Truck Trail. It lies in a shallow, poorly drained depression formed by a remnant ice block on sandy glacial deposits. This wetland includes a two acre area of open water near the center, and is surrounded by a mosaic of sedge meadow, shrubs, and islands dominated by black spruce (Picea mariana) and tamarack (Larix laricina). Dominant vascular plant species include bulrush (Scirpus hudsonianus), running bog sedge (Carex oligosperma), sundews (Drosera rotundifolia, and Drosera intermedia), laurel (Kalmia polifolia), bog rosemary (Andromeda glaucophylla), and Labrador tea (Ledum groenlandicum). A total of 24 vascular plant species were noted in this wetland. An active beaver den was observed at the south end of the wetland in 1993. The adjacent uplands surrounding this bog include second growth forest of trembling aspen (Populus tremuloides) and red oak (Quercus rubra). In the early 1990s or late 1980s, much of the upland along the east side of the wetland was clearcut and now supports a dense stand of trembling aspen saplings. Jack pine (Pinus banksiana) has been planted in part of this area.

Grayling Community Assessment - 24 Protection of this wetland should include establishment of a 200 meter-wide upland vegetated buffer. Fires occurring within the adjacent uplands should be allowed to carry into the wetland. Invasive species monitoring and control should be implemented within the wetland and adjacent uplands. For additional information on conservation and management of bogs, see the natural community abstract in Appendix 5.

Soil: 0 to 150+ cm: fibric peat with pH of 4.5 (4.0)

Invasive species: none observed

Conservation and Management: • Monitor for and remove invasive species • Conduct prescribed fire in adjacent uplands on an occasional basis and allow fire to spread into wetland • Maintain vegetated upland buffer for approximately 200 meters

Grayling Community Assessment - 25 10. Crawford Red Pines Community Type: dry northern forest Alliance: Pinus resinosa Forest Alliance (I.A.8.N.b) Rank: G3? S3?, very rare and local throughout range Location: T27N R01W

M. Kost Photo 15. Large red pines, approximately 180 years old and measuring 70(±) cm in diameter, dominate the canopy of the dry northern forest at Crawford Red Pines.

Grayling Community Assessment - 26

M. Kost Photo 16. At Crawford Red Pines, red maples are outcompeting red pine saplings. Prescription burning and manual removal should be used to reduce the density of red maple at this site.

Site Description: This 14 acre tract of old growth red pine (Pinus resinosa) was first identified in 1954, and is considered one of the only remnant stands of its type in Lower Michigan. It is located twelve miles east of Grayling along Dyer Truck Trail at the southeastern extreme of Camp Grayling property. This site is located on a low rise in what is otherwise a poorly drained outwash plain that drains into Barker Creek. Red pine, up to 178 years old, forms a partially closed canopy in this tract. Common plant species within this tract include blueberry (Vaccinium angustifolium), bunchberry (Cornus canadensis), bracken fern (Pteridium aquilinum), rough-leaved rice grass (Oryzopsis asperifolia), and trailing arbutus (Epigaea repens). Immediately surrounding portions of the old growth tract is second growth forest of red pine, northern pin oak (Quercus ellipsoidalis), jack pine (Pinus banksiana), and red maple (Acer rubrum). The area directly to the north was recently clear cut, disked to create furrows, and planted with jack pine. Adjacent wetlands include a bog element occurrence (Best Bog) and second growth hardwood-conifer swamp. A northern fen (Barker Creek Fen) occurs approximately one mile southwest of the site.

Grayling Community Assessment - 27 During the middle 1980s, camp activities were ceased around the vicinity of the tract. Long-term management of the tract should consider the removal of competing vegetation, such as red maple, which impedes the natural regeneration of red pine through creation of dense shade (Abrams 1998). Red maple seedlings were especially abundant in old abandoned trails, roads, and camping areas where sedge (Carex pensylvanica) is sparse and pine needles and oak leaves insulate the soil and trap moisture. Removal of red maple could be accomplished either by appropriately timed prescribed burning or mechanical means. Given the proximity of two adjacent high quality natural community occurrences (Best Bog and Barker Creek Fen), consideration should be given to developing an integrative management plan for this area of Camp Grayling. In addition to prescribed burning, invasive species monitoring and control should be implemented within this site and in adjacent wetlands. For additional information on conservation and management of dry northern forests, see the natural community abstract (Cohen 2002).

Soil: sand, pH 5.5 (5.0)

Invasive species Estimated Abundance Scientific Name Common Name Estimated Abundance Agrostis gigantea redtop uncommon Hypericum perforatum* common St. John’s wort* occasional Poa compressa* Canada bluegrass* locally common

Conservation and Management: • Conduct prescribed fire on an occasional basis • Remove red maple either through cutting and herbicide or repeated prescribed fires • Consider developing an integrated management plan for this portion of Camp Grayling that would encompass Crawford Red Pines, Best Bog, and Barker Creek Fen • Monitor for and remove invasive species

Grayling Community Assessment - 28 11. Watson Swamp Community Type: rich conifer swamp Rank: G4 S4, apparently secure globally Alliance: Thuja occidentalis Saturated Forest Alliance (I.A.8.N.g) Location: T26N R06W

M. Kost Photo 17. Cut stumps are common within the northeast portion of Watson Swamp where several living northern white cedars were aged to approximately 80 and 86 years old.

Site Description: This 305 acre swamp is located eight miles northeast of Sharon along Melum Road. This swamp was formed on a poorly drained outwash plain along the Manistee River. This very diverse swamp is dominated by northern white cedar (Thuja occidentalis), black spruce (Picea mariana), balsam fir (Abies balsamea), and red maple (Acer rubrum). Other common vascular plant species include paper birch (Betula papyrifera), mountain holly (Nemopanthus mucronata), creeping snowberry (Gaultheria hispidula), swollen sedge (Carex intumescens), and marsh marigold (Caltha palustris). Many diverse micro-habitats exist within this swamp, especially along the slow flowing streams found throughout the site. The northern portion of the wetland contains an old logging road. This area also contain many cut stumps of northern white cedar. Cores were taken of several live cedars and aged to 80 years (18.5 cm dbh) and 86 years (26.5 cm dbh) old. A total of 121 one vascular plant species were noted in this swamp. Protection of this wetland should include the creation and maintenance of a vegetated, upland buffer between 100 and 200 meters wide along the east wetland edge. Culverts should be maintained along Mecum Road to maintain natural water flows through the system. Invasive species monitoring and control should be implemented within the wetland and adjacent uplands. For additional information on conservation and management of rich conifer swamps, see the natural community abstract (Kost 2002).

Grayling Community Assessment - 29 Soil: 0 to 30 cm: peat (pH 7.0) 30 to 150+ cm: peat and large coarse woody debris and many wood fragments

Invasive species Estimated Abundance Scientific Name Common Name Estimated Abundance Prunella vulgaris lawn prunella uncommon Solanum dulcamara bittersweet nightshade uncommon Typha angustifolia narrow-leaved cattail uncommon

Conservation and Management: • Monitor for and remove invasive species • Maintain culvert along Mecum Road to allow natural water flow through the wetland • Maintain a vegetated, upland buffer of between 100 and 200 meters wide

Grayling Community Assessment - 30 12. Cannon Creek Meadow Community Type: northern wet meadow Rank: G4 S4, apparently secure globally Alliance: Calamagrostis canadensis Seasonally Flooded Herbaceous Alliance (V.A.5.N.k) Location: T25N R06W

M. Kost Photo 18. The Cannon Creek Meadow is a northern wet meadow that is dominated by tussock sedge (Carex stricta) and occurs along Cannon Creek, a tributary to the Manistee River. Reed canary grass, an invasive species, was observed growing along South Sharon Road on the edge of the wetland and should be removed to prevent it from spreading further into the wet meadow.

Site Description: This 149 acre northern wet meadow is located three miles south of Sharon along South Sharon Road at the southwestern extreme of Camp Grayling property. It lies within a much larger wetland complex on a poorly drained outwash plain. Cannon Creek itself drains to the west into the Manistee River. This is a dense, sedge-dominated meadow, with meadow sedge (Carex stricta) and slender woolly sedge (Carex lasiocarpa) being most common. Scattered tamarack (Larix laricina), bog birch (Betula pumila), shrubby cinquefoil (Potentilla fruticosa), sweet bay (Myrica gale), meadowsweet (Spiraea alba), and blue joint grass (Calamagrostis canadensis) are also characteristic. Forty four vascular plant species were noted in this wet meadow. Surrounding the meadow are zones of willow (Salix)-dominated swamp and hardwood-conifer swamps. The immediately adjacent slopes contain many seeps with a highly diverse flora. Historically, adjacent uplands were dominated by oak and pine-dominated forest. Today, open-canopy, oak- dominated forest is characteristic. Protection of this sensitive wetland complex should include establishing a 200 meter upland buffer where no timber management activities take place. Emphasis should be placed on protecting the seepages

Grayling Community Assessment - 31 adjacent to the wetland complex. Invasive species monitoring and control should be implemented within the wetland and adjacent uplands. In particular, reed canary grass (Phalaris arundinacea), an invasive species, was observed growing along South Sharon Road on the edge of wetland and should be removed to prevent it from spreading further into the wetland. Because fire helps maintain open conditions and species diversity in wetland ecosystems, fires occurring within the uplands should be allowed to carry into the wetland. For additional information on conservation and management of northern wet meadows, see the natural community abstract in Appendix 6.

Soil: 0 to 30 (150+) cm: fibric sedge peat (i.e., depth of peat varies from 30 cm to > 150 cm) 30 to 140 cm: grey, coarse-textured sand with some organics 140 to 150+ cm: marl

Water table: 10 cm standing water above soil surface

Invasive species Estimated Abundance Scientific Name Common Name Estimated Abundance Phalaris arundinacea* reed canary grass* locally abundant

Conservation and Management: • Monitor for and remove invasive species • Maintain a 200 meter wide vegetated buffer around wetland • Protect seepage areas along sloping edges of wetland • Conduct prescribed fire in adjacent uplands on an occasional basis and allow fire to spread into wetland

Grayling Community Assessment - 32 13. Chub Creek Swamp Community Type: northern shrub thicket Rank: G4 S4, apparently secure globally Alliance: Alnus incana Seasonally Flooded Shrubland Alliance (III.B.2.N.e) Location: T29N R02W

M. Kost Photo 19. The Chub Creek Swamp is comprised of a northern shrub thicket that grades from shrubby wet meadow along Chub Creek to forested conifer swamp (rich conifer swamp) near the edge of the adjacent uplands. Periodic fires and flooding by beaver probably kept this wetland in a continual flux between wet meadow, shrub thicket, and rich conifer swamp.

Site Description: This 123 acre wetland is located just northwest of Rosecrans Hill at the northern extreme of Camp Grayling property. It lies on a poorly drained outwash plain at the confluence of Chub creek and the North Branch of the Au Sable River. Approximately half of this wetland, north of the river, is outside of Camp Grayling. Dominant species in this very diverse wetland include speckled alder (Alnus rugosa), bog birch (Betula pumila), sweet bay (Myrica gale), shining willow (Salix lucida), shrubby cinquefoil (Potentilla fruticosa), sedge (Carex aquatilis), sedge (Carex flava), slender woolly sedge (Carex lasiocarpa), wild rice (Zizania aquatica), and tall goldenrod (Solidago altissima). Tamarack (Larix laricina) is scattered throughout the shrub swamp and becomes dominant around the margins. A total of 93 vascular plant species were noted in this wetland. An active bald eagle (Haliaeetus leucocephalus) nest is located at the north end of this complex.

Grayling Community Assessment - 33 Protection of this wetland should include the maintenance of an upland buffer zone for 100 meters along all sides of the wetland. Invasive species monitoring and control should be implemented within the wetland and adjacent uplands. Fires occurring within the adjacent uplands should be allowed to carry into the wetland. For additional information on conservation and management of northern shrub thicket, see the natural community abstract in Appendix 7.

Soil: 0 to 20 cm: fibric peat 20 to 100 cm: coarse woody debris in matrix of peat 100 to 150+ cm: sapric peat (muck)

Invasive species: none observed

Conservation and Management: • Monitor for and remove invasive species • Maintain a 100 meter vegetated buffer around wetland • Allow fires occurring in adjacent upland to carry into wetland

Grayling Community Assessment - 34 14. Beaver Creek Natural Community Type: northern shrub thicket Rank: G4 S4, apparently secure globally Alliance: Alnus incana Seasonally Flooded Shrubland Alliance (III.B.2.N.e) Location: T25N R04W (scale 1:24,000)

M. Kost Photo 20. Tamarack has become very abundant within the Beaver Creek northern shrub thicket. In the absence periodic natural disturbances such as fire, beaver flooding, or insect outbreaks (e.g., larch sawfly infestation), the northern shrub thicket will eventually succeed to forested wetland.

Site Description: This 41 acre wetland is located nearly two miles west of Military Road, and just south of Fletcher Road. It lies in a poorly drained portion of a large, flat outwash plain that is drained by Beaver Creek. Tamarack (Larix laricina) and black spruce (Picea mariana) are found throughout the wetland, while jack pine (Pinus banksiana) is concentrated along the wetland edge and in the adjacent uplands. Dominant plant species in the wetland include speckled alder (Alnus rugosa), leatherleaf (Chamaedaphne calyculata), bog birch (Betula pumila), willow (Salix spp.), meadowsweet (Spiraea alba), black choke berry (Aronia prunifolia), red osier dogwood (Cornus stolonifera), lake sedge (Carex lacustris), and blue joint grass (Calamagrostis canadensis). Sphagnum mosses (Sphagnum spp.) are also abundant in this wetland. A total of 71 vascular plant species were noted. This shrub thicket is part of a larger wetland complex, which extends both to the east and west. To the west, much of the area has been logged and the water level has

Grayling Community Assessment - 35 been impacted by road construction. The adjacent uplands contain numerous oil/gas wells scattered throughout a second growth, jack pine-dominated forest. The steady, natural flow of water through this system is necessary to maintain the integrity of the wetland. Thus, protection of this wetland should include the installation and maintenance of culverts in the two-track road that passes along its west side. This area also contains reed canary grass (Phalaris arundinacea), an invasive species, that should be removed to prevent it from spreading further into the shrub swamp. Invasive species monitoring and control should be implemented within the wetland and adjacent uplands. A 100 meter wide vegetated upland buffer should be maintained on all sides of this wetland. Fires occurring within the adjacent uplands should be allowed to carry into the wetland. For additional information on conservation and management of northern shrub thicket, see the natural community abstract in Appendix 7.

Soil: 0 to 100 cm: fibric peat (pH 6.5) with large coarse woody debris 100 to 110 cm: coarse-textured, light colored sand. pH of sphagnum hummocks: 4.0 to 4.5

Water table: varies from approximately 10 cm above surface on east side of wetland between sedge hummocks to 10 cm below soil surface on west side of wetland. Peat and underlying sand are very cold, indicating contact with flowing groundwater.

Invasive species Estimated Abundance Scientific Name Common Name Estimated Abundance Phalaris arundinacea* reed canary grass* locally abundant

Conservation and Management: • Monitor for and remove invasive species • Consider conducting prescribed fire in some portions of the adjacent uplands on an occasional basis and allowing fire to spread into wetland. Note: active gas wells occur in some portions of the adjacent upland should be protected from fire. • Install and maintain culvert in two-track road along west side of wetland • Maintain 100 meter vegetated upland buffer around wetland

Grayling Community Assessment - 36 DISCUSSION

Several significant changes were observed (Phragmites australis), reed canary grass within the 14 communities we studied. Water (Phalaris arundinacea), spotted knapweed levels are now significantly lower in most (Centaurea maculosa), common St. John’s wort wetlands than when first observed in the early (Hypericum perforatum), and Canada bluegrass 1990s. While invasive species have not yet caused (Poa compressa) currently pose the greatest threat significant degradation in any of the communities because of their ability to invade intact as of yet, most now contain small infestations that communities and quickly dominate an area. In threaten to degrade ecological integrity. The addition to the species listed above, several other continued affects of fire suppression were very highly invasive species have the potential to apparent within many of the communities we become established on Camp Grayling including surveyed. Fortunately, damage by ORVs within garlic mustard (Alliaria petiolata), glossy the high quality natural communities was not buckthorn (Rhamnus frangula), common severe. buckthorn (Rhamnus cathartica), exotic honeysuckle (Lonicera morrowii, L. tatarica L. Water Level Changes Xbella, L. maackii, etc.), autumn olive (Elaeagnus umbellata), oriental bittersweet (Celastrus In most of the wetlands surveyed in 2004 it umbellata), and purple loosestrife (Lathyrus was very apparent that water levels were much salicaria). Because new introductions of invasive, lower than when these communities were first exotic species continue to occur regularly, it is documented in the early 1990s. These imperative that a systematic and sustained effort observations are consistent with decreases in be undertaken to monitor for and control invasive water levels recorded in wetlands throughout the species on Camp Grayling. Upper Great Lakes Region. Regional water levels declined sharply in the late 1990s and have since Fire Suppression risen slightly. Our findings indicate that groundwater is an important source of water input Camp Grayling occurs within the High Plains for most wetlands at Camp Grayling. Subsection, one of the most fire prone regions of Michigan (Albert 1995). Most natural Invasive Species communities that occur within the High Plains are well adapted to fire, which is an ecological Invasive species pose a major threat to species process that maintains the biological integrity of and habitat diversity and ecological integrity the region’s varied ecosystems. Where it can be within several high quality natural communities safely implemented, prescribed fire should be on Camp Grayling. By outcompeting and used in managing both upland and wetland replacing native species, invasives change species ecosystems on Camp Grayling. Because fire is an composition, alter vegetation structure, and reduce important ecological process for maintaining native species diversity, often causing local or species and habitat diversity in both upland and even complete extinction of native species (Harty wetland ecosystems, fires occurring within the 1986). Invasive exotic species can also upset uplands should be allowed to carry into adjacent delicately balanced ecological processes such as wetlands whenever possible. trophic relationships, interspecific competition, The ecological process of fire has played a nutrient cycling, soil erosion, hydrologic balance, critical role in shaping the vegetation of the upper and solar insolation (Bratton 1982). Lastly, exotic Midwest region (Curtis 1959, Davis 1979, Kline invasive species often have no natural predators and Cottam 1979, Dorney 1981, Grimm 1984, and spread aggressively through rapid sexual and Dorney and Dorney 1989). Prior to European asexual reproduction. settlement, the occurrence of fire, whether due to While numerous invasive species occur within lightning strikes or indigenous cultures, was the high quality natural communities, giant reed frequent in many types of ecosystems (Dorney

Grayling Community Assessment - 37 1981, Guyette and Cutter 1991). Since then, the which may be especially important for incidence of fire in all natural communities has maintaining species diversity within wetlands been greatly reduced (Curtis 1959). As a (Leach and Givnish 1996, Kost and De Steven consequence, many natural areas have 2000). experienced significant changes in their species Impacts to faunal communities should also be composition (Cottam 1949, Curtis 1959, Davis considered when planning a prescribed burn. 1979, Grimm 1984, McCune and Cottam 1985, Dividing a large area into smaller burn units that Abrams and Nowacki 1992, McClain et al. 1993, can be burned in alternate years or seasons can Motzkin et al. 1993). In an effort to preserve protect populations of many species. This allows biodiversity and restore natural communities, fire unburned units to serve as refugia for immobile has been reintroduced as a management tool to invertebrates and slow moving amphibian and many natural areas (Vogl 1964, White 1983, Vora reptile species. When burning larger areas it may 1993, Henderson and Statz 1995, Bowles et al. be desirable to strive for patchy burns by igniting 1996). during times of high relative humidity. As Plant communities, whether upland or mentioned above, the unburned patches may then lowland, benefit from prescribed fire in several serve as refugia and thus facilitate recolonization ways. Depending on the season and intensity of a of burned patches by fire-sensitive species. burn, prescribed fire may be used to decrease the Burning under overcast skies and when air cover of exotic, cool-season grasses and woody temperatures are cool (<55°F) can help protect species, and increase the cover of warm-season reptiles, since they are less likely to be found grasses and native forbs (White 1983, Abrams and basking above the surface when conditions are Hulbert 1987, Tester 1989, Anderson and cloudy and cool. Lastly, impacts to reptiles may Schwegman 1991, Collins and Gibson 1990, also be minimized by conducting burns during the Glenn-Lewin et al. 1990). Prescribed fire helps dormant season (late October through March). reduce litter levels, allowing sunlight to reach the soil surface and stimulate seed germination and ORV Impacts enhance seedling establishment (Daubenmire 1968, Hulbert 1969, Knapp 1984, Tester 1989, Continued efforts to restrict ORV use to Anderson and Schwegman 1991, Warners 1997). designated areas are having positive impacts on Important plant nutrients (e.g., N, P, K, Ca, and ecological integrity and should be continued. Mg) are elevated following prescribed fire ORV use was observed in only two (14%) of the (Daubenmire 1968, Viro 1974, Reich et al. 1990, 14 natural communities we studied, both of which Schmalzer and Hinkle 1992), which contributes to were intermittent wetlands (The Doughnut and increased plant biomass, flowering, and seed Lake Margrethe North). Continuing efforts to production (Laubhan 1995, Abrams et al. 1986, educate ORV users and discourage their use in Warners 1997, Kost and De Steven 2000). wetlands and other undesignated areas will be an Prescribed fire can also significantly increase seed important component of any long-term effort to bank expression and help rejuvenate seed banks, maintain biodiversity on Camp Grayling.

CONCLUSION

In the decade or more since the 14 high the Portage Creek Complex. The continued affects quality natural communities were documented of fire suppression have resulted in canopy many have experienced relatively significant closure and the buildup of a thick layer of leaf changes. Water levels have dropped significantly litter, which stifles seed germination and within most wetlands since the early 1990s. establishment, especially in wetlands. Invasive species now pose a significant threat to These results indicate that a systematic and several natural communities. In particular, if left sustained effort should be undertaken to monitor unchecked, leafy spurge is likely to degrade the for and control invasives, especially within the mesic sand prairie and rare species populations at high quality natural communities and in the

Grayling Community Assessment - 38 surrounding landscapes. Because fire is an seed bank expression and native species diversity. important ecological process for maintaining Thanks to ongoing control and education efforts, species and habitat diversity in both upland and ORVs have caused relatively minor damage to the wetland ecosystems, wherever possible, high quality natural communities in the past prescribed fire should be used to manage these decade. Efforts to discourage ORV use in communities and the surrounding landscape. Fires undesignated areas, especially wetlands, will be occurring in the uplands should be allowed to an important component of any long-term effort to carry into the wetlands when possible to bolster maintain biodiversity on Camp Grayling.

ACKNOWLEDGEMENTS

Funding for this project was made available this project and providing guidance and support through a grant from the Michigan Department of throughout its execution. We thank Phyllis Military and Veterans Affairs. We wish to extend Higman for taking the lead role in coordinating our sincere gratitude to the many Camp Grayling efforts between Camp Grayling and MNFI staff staff who provided access to fields sites, logistical and for her expert guidance in the field and support, and kept us safe from harms way on a throughout this project. We are grateful for the very busy military training facility. The staff at additional administrative support provided by the Range Control were consistently helpful and following MFNI staff: Sue Ridge, Lyn Scrimger, accommodating, which was a pleasant contrast to Connie Brinson, and Pat Brown. Jeff Lee the sometimes grueling summer field conditions. conducted final editing and formatting of the Larry Jacobson, Greg Huntington, and John Hunt report. deserve our sincerest appreciation for initiating

LITERATURE CITED

Abrams, M.D. 1998. The red maple paradox: Forest Service – U.S. Department of What explains the widespread expansion of Agriculture. St. Paul, MN. red maple in eastern forests? BioScience 48: 355-363. Anderson, R. C. and J. E. Schwegman. 1991. Abrams, M. D. and G. J. Nowacki. 1992. Twenty years of vegetational change on a Historical variation in fire, oak recruitment, Southern Illinois barren. Natural Areas and post-logging accelerated succession in Journal 11: 100-107. central Pennsylvania. Bulletin of the Torrey Botanical Club 119: 19-28. Bowles, M., J. McBride, N. Stynoff, and K. Johnson. 1996. Temporal changes in Abrams, M. D. and L. C. Hulbert. 1987. Effect vegetation composition and structure in a fire- of topographic position and fire on species managed prairie fen. Natural Areas Journal composition in tallgrass 16:275-278.

Abrams, M. D., A. K. Knapp and L. C. Hulbert. Bratton, S. P. 1982. The effects of exotic plant 1986. A ten year record of aboveground and animal species on nature preserves. biomass in a Kansas tallgrass prairie: Effects Natural Areas Journal 2(3): 3-13. of fire and topographic position. American Journal of Botany 73: 1509-1515. Cohen, J. G. 2002. Natural community abstract for dry northern forest. Michigan Natural Albert, D. A. 1995. Regional landscape Features Inventory, Lansing, MI. 14 pp. ecosystems of Michigan, Minnesota, and Wisconsin: A working map and classification. Collins, S. L. and D. J. Gibson. 1990. Effects of North Central Forest Experiment Station, fire on community structure in tallgrass and

Grayling Community Assessment - 39 mixed grass prairie. Pp. 81-98 In (S.L. Collins (eds.), Fire in North American tallgrass and L.L. Wallace, eds.) Fire in North prairies, University of Oklahoma Press, American Tallgrass Prairies, University of Norman, OK. Oklahoma Press, Norman, OK. Grimm, E. C. 1984. Fire and other factors Comer, P. J., D. A. Albert, H. A. Wells, B. L. controlling the big woods vegetation of Hart, J. B. Raab, D. L. Price, D. M. Kashian, Minnesota in the mid-nineteenth century. R. A. Corner and D. W. Schuen. 1995. Ecological Monographs 54: 291-311. Michigan’s presettlement vegetation, as interpreted from the General Land Office Guyette R. P., and B. E. Cutter. 1991. Tree-ring Surveys 1816-1856. Michigan Natural analysis of fire history of a post oak savanna Features Inventory, Lansing, MI. Digital Map. in the Missouri Ozarks. Natural Areas Journal 11: 93-5. Cottam, G. 1949. The phytosociology of an oak woods in southwestern Wisconsin. Ecology Harty, F. M. 1986. Exotics and their ecological 30: 271-287. ramifications. Natural Areas Journal. 6(4): 20-26. Curtis, J. T. 1959. Vegetation of Wisconsin. The University of Wisconsin Press, Madison, WI. Henderson, R. A. and S. H. Statz. 1995. Bibliography of fire effects and related Daubenmire, R. 1968. Ecology of fire in literature applicable to the ecosystems and grasslands. Advances in Ecological Research species of Wisconsin. Technical Bulletin No. 5: 209-66. 187, Department of Natural Resources, Madison, WI. Davis, A. M. 1979. Wetland succession, fire and the pollen record: a Midwestern example. The Higman, P. J., P. J. Comer, M. R. Penskar, M. L. American Midland Naturalist 102: 86-94. Rabe, D. A. Albert, J. T. Legge, T. R. Leibfreid, L. J. Scrimger, and M. B. Austin. Dorney, J. R. 1981. The impact of Native 1994. Final Report for a Floristic and Natural Americans on presettlement vegetation in Features Inventory of camp Grayling Military Southeastern Wisconsin. Transactions of the Reservation, Grayling Michigan. Report Wisconsin Academy of Science, Arts, and prepared for Michigan Department of Military Letters. 69: 26-36. and Veterans Affairs. Michigan Natural Features Inventory. Lansing, MI. 112 pp. + Dorney, C. H. and J. R. Dorney. 1989. An Appendices 198 pp. unusual oak savanna in northeaster Wisconsin: the effect of Indian-caused fire. Hulbert, L. C. 1969. Fire and litter effects in American Midland Naturalist 122: 103-13. undisturbed bluestem prairie in Kansas. Ecology 50: 874-877. Faber-Langendoen, D., editor. 2001. Plant communities of the Midwest: Classification in Kline, V. M. and G. Cottam. 1979. Vegetation an ecological context. Association of response to climate and fire in the driftless Biodiversity Information, Arlington, VA. area of Wisconsin. Ecology 60: 861-68. 61pp + appendix (705 pp.). Knapp, A. K. 1984. Post-burn differences in Glenn-Lewin, D. C., L. A. Johnson, T. W. Jurik, solar radiation, leaf temperature and water A. Akey, M. Leoschke, and T. Rosberg. stress influencing production in a lowland 1990. Fire in central North American tallgrass prairie. American Journal of Botany grasslands: vegetative reproduction, seed 71: 220-227. germination, and seedling establishment. Pp. 28-45 in S. L. Collins and L. L. Wallace

Grayling Community Assessment - 40 Kost, M. A. 2002. Natural community abstract Bulletin of the Torrey Botanical Club 116: for rich conifer swamp. Michigan Natural 134-144. Features Inventory, Lansing, MI. 9 pp. Viro, P. J. 1974. Effects of forest fire on soil. Pp. Kost, M. A. and D. De Steven. 2000. Plant 7-45 In (T.T. Kozlowski and C.E. Ahlgren community responses to prescribed burning in eds.) Fire and Ecosystems, Academic Press, Wisconsin sedge meadows. Natural Areas New York, NY. Journal 20:36-49. Vogl, R. J. 1964. The effects of fire on a muskeg Kost, M. A., P. J. Higman, D. L. Cuthrell, and J. in northern Wisconsin. Journal of Wildlife L. Cooper. 2000. North Camp Grayling Pine Management 28: 317-29. Barrens Management Plan. Report prepared for Michigan Department of Military and Vora, R. S. 1993. Moquah barrens: Pine Barrens Veterans Affairs. 28 pp. + Appendices 39 pp. Restoration Experiment Initiated in the Chequamegon National Forest. Restoration Laubhan, M. K. 1995. Effects of prescribed fire and Management Notes 11:39-44. on moist-soil vegetation and macronutrients. Wetlands 15:159-166. Warners, D. P. 1997. Plant diversity in sedge Leach, M. K. and T. J. Givnish. 1996. Ecological meadows: Effects of groundwater and fire. determinants of species loss in remnant Ph.D. dissertation, University of Michigan, prairies. Science 273: 1555-1558. Ann Arbor, MI. 231 pp.

McClain, W. E., M. A. Jenkins, S. E. Jenkins, and White, A. S. 1983. The effects of thirteen years J. E. Ebinger. 1993. Changes in the woody of annual prescribed burning on a Quercus vegetation of a bur oak savanna remnant in ellipsoidalis community in Minnesota. central Illinois. Natural Areas Journal 13:108- Ecology 64: 1081-1085. 114.

McCune, B. and G. Cottam. 1985. The successional status of a southern Wisconsin oak woods. Ecology 66:1270-1278.

Motzkin, G., W. A. Patterson and N. E. Drake. 1993. Fire history and vegetation dynamics of a Chamaecyparis thyoides wetland on Cape Cod, Massachusetts. Journal of Ecology 81: 391-402.

Reich, P. B., M. D. Abrams, D. S. Ellsworth, E. L. Kruger & T. J. Tabone. 1990. Fire affects ecophysiology and community dynamics of Central Wisconsin oak forest regeneration. Ecology 71: 2179-2190.

Schmalzer, P. A. and C. R. Hinkle. 1992. Soil dynamics following fire in Juncus and Spartina marshes. Wetlands 12: 8-21.

Tester, J. R. 1989. Effects of fire frequency on oak savanna in east-central Minnesota.

Grayling Community Assessment - 41 Grayling Community Assessment - 42 APPENDICES. NATURAL COMMUNITY ABSTRACTS

Grayling Community Assessment - 43 Grayling Community Assessment - 44 Appendix 1. Mesic Sand Prairie

Overview: Mesic sand prairie is a native which total to 398 acres (161 ha) and range in size grassland community that is typically dominated from 2 to 115 acres (>1 to 47 ha). It is difficult to by little bluestem (Andropogon scoparius), big reliably determine the total acreage of mesic sand bluestem (Andropogon gerardii), Indian grass prairie in Michigan in the 1800s. However, based (Sorghastrum nutans), and/or prairie dropseed on comparisons of the total acreage of all upland (Sporobolus heterolepis) and occurs on sandy prairie element occurrences in Lower Michigan loam, loamy sand, or sand soils on nearly level today (1,463 acres, 592 ha) with that found in the glacial outwash. Sites that support mesic sand early to mid 1800s (provided above), it appears prairie are seasonally wet, with high water tables that only 2% of the original upland prairie remains occurring in the spring followed by drought intact in all of Lower Michigan. conditions in late summer and fall. Thus, the community contains species from a broad range of Landscape and Abiotic Context: Mesic sand moisture classes such as those more typically prairie and lakeplain mesic sand prairie, which has associated with wet-mesic prairie, mesic prairie, been documented on the lakeplain in southeast woodland prairie, and dry sand prairie. Areas Michigan, differ in that the hydrology of lakeplain dominated by native grasses with less than one mesic sand prairie is strongly influenced by mature tree per acre are considered prairie (Curtis changes in the Great Lakes water levels (Albert 1959). and Kost 1998). However, both communities experience seasonal water table fluctuations, with Global and State Rank: S1/G2G3 the wettest conditions occurring in spring and driest periods in late summer and fall. Prolonged Range: Mesic sand prairie occurs in IL, IN, MI, spring inundation may occur in the wettest OH, WI, and southern Ontario. (Faber- portions of some mesic sand prairies. Langendoen 2001). In Michigan, this community has been documented in both the southern and Soils supporting mesic sand prairie are sandy northern Lower Peninsula. In southern Lower loam or occasionally loamy sand, loamy fine Michigan, mesic sand prairie occurs within the sand, or fine sand, with pH ranging form 5.4 to Interlobate Region, where it is found on glacial 7.3 (ave. pH 5.5) and water retaining capacity outwash in Oakland, Washtenaw, and Van Buren ranging from 31% to 62% (ave. 43%) (Chapman counties. Within the northern Lower Peninsula, 1984). The mesic condition of the sandy soils is mesic sand prairie occurs on glacial outwash facilitated by a high water table and, in some sites, (Oceana and Lake counties) and on an old glacial by a relatively high organic content within the lakebed within an outwash plain (Crawford sand matrix, which increases the water holding County) (Higman et al 1995). Historically, the capacity of soil. community likely occurred as small patches within fire prone landscapes on sandy soils with a In the 1800s, mesic sand prairie in Michigan high water table and as an ecotone between occurred as small patches of grassland within and savanna (e.g., oak opening, oak barrens, oak-pine between fire prone communities. The community barrens, bur oak plains,) and non-forested occupied sandy sites with high water tables such wetlands. as those occurring in shallow depressions within outwash plains and on old glacial lakebeds, Rank Justification: In the early to mid 1800s, the abandoned stream channels, and river terraces. southern Lower Peninsula supported Mesic sand prairie also occurred as an ecotone approximately 73,000 acres (29,500 ha) of upland between fire-dependent uplands and open prairie, which included pockets of mesic sand wetlands (e.g., wet-mesic prairie, wet prairie, prairie, mesic prairie, woodland prairie, dry sand prairie fen, northern fen, southern wet meadow, prairie, and hillside prairie. The Michigan Natural northern wet meadow, intermittent wetland, Features Inventory database currently includes coastal plain marsh, emergent marsh). eight element occurrences of mesic sand prairie,

Grayling Community Assessment - 45 Natural Processes: A high water table in the diversity. A recensus of 54 prairie remnants in spring followed by drought-like conditions in the Wisconsin found that 8% to 60% of the original late summer and fall produces a diverse floristic plant species recorded at the sites had been lost composition of species for mesic sand prairie with over time (32 to 52 years) even though the sites species representing a broad range of moisture appeared relatively undisturbed (Leach and tolerances. In addition to seasonal water level Givnish 1996). The authors suggest the decline in fluctuations, longer term changes in the regional diversity was a result of taller vegetation water table also influence the community outcompeting species with small stature, those composition. with small seeds (e.g., orchids), and those that rely on nitrogen-fixing symbioses such as members of As in other prairie and savanna communities, fire the legume family (Fabaceae) (e.g., lupine played a critical role in maintaining open (Lupinus perennis), wild indigo (Baptisia spp.), conditions in mesic sand prairie. The frequency bush clover (Lespedeza spp.), and tick-trefoil and intensity of fire depended on a variety of (Desmodium spp.)). Because fire maintains open factors including the type and volume of fuel, conditions and burns off accumulated leaf litter, topography, presence of natural firebreaks, and species that require open microsites for seedling density of Native Americans (Chapman 1984). In establishment and growth such as those with small general, the probability of wide-ranging fire seeds or small statures, are able to garner enough increases in level topography like large outwash space and light to coexist with taller, denser plains (Chapman 1984). Carried by wind, fires vegetation. In the absence of frequent fires, small moved across the outwash plains, through species are outcompeted by taller and denser graminoid-dominated wetlands, and up slopes of vegetation, and small-seeded species with low end moraines and ground moraines. food reserves have difficulty growing through thick litter. The decline in species diversity is While occasional lightning strikes resulted in fires especially pronounced in mesic and wet that spread across the landscape, Native community types where biomass accumulates Americans were the main sources of ignition. rapidly. Because fire volatilizes much of the There are many early accounts of Native nitrogen stored in combustible vegetation, Americans intentionally setting fires to frequent burning also favors species that form accomplish specific objectives (see Day 1953, nitrogen-fixing symbioses (e.g., legumes and Curtis 1959, Thompson and Smith 1970, rhizobium bacteria) by providing a competitive Chapman 1984, Denevan 1992, Kay 1995). edge not found in unburned sites (Leach and Native Americans intentionally set fires in the fall Givnish 1996). to clear briars and brush and make the land more easily passable. Frequent fires kept the land open, Fire also helps maintain species diversity by increasing both short- and long-range visibility, facilitating expression of the soil seed bank and which facilitated large game hunting and provided promoting seed germination and establishment. a measure of safety from surprise attacks by By consuming accumulated and standing leaf neighboring tribes. Fire was used to increase litter, fire increases light availability to the soil productivity of berry crops and agricultural fields. surface and increases diurnal temperature As a habitat management tool, fires were used to fluctuations, both of which trigger seed maintain high quality forage for deer, elk, germination. Critical microsites for seed woodland caribou, bison and other game species. germination and seedling establishment are also It was also used as a hunting tool to both drive created when litter levels are reduced by fire. and encircle game. During warfare, fire was Through burning accumulated litter and dead, strategically employed to drive away advancing standing vegetation, fire increases the availability enemies, create cover for escape, and for waging of many important plant nutrients (e.g., N, P, K, attacks. Ca and Mg), which are thought to contribute to

higher plant biomass, increased flowering and In addition to maintaining open conditions, fire seed production, and greater palatability to plays a critical role in maintaining species

Grayling Community Assessment - 46 herbivores following a burn (Vogl 1964, creating microsites for seed germination and Daubenmire 1968, Viro 1974, Vogl 1974, Smith seedling establishment and reducing the and Kadlec 1985, Abrams et al. 1986, Collins and dominance of robust perennials (Steuter 1997). Gibson 1990, Reich et al. 1990, Schmalzer and Hinkle 1992, Timmins 1992, Laubhan 1995, Vegetation Description: Unfortunately, no Warners 1997). detailed ecological study of mesic sand prairie was completed in Michigan before the nearly total While this discussion has focused on plants it is demise of the community. What information is important to note that these species serve as host available comes from a detailed study of prairie plants for a variety of insects and the structure of communities in Michigan by Chapman (1984) and open grasslands is critical to a wide variety of data from Michigan Natural Features Inventory animal species, many of which are considered rare element occurrence records. or declining today (see Other Noteworthy Species section). The vegetation of mesic sand prairie supports a sparse to moderately dense growth of low to Ants, particularly the genus Formica, play an medium height vegetation with patches of bare important role in mixing and aerating prairie soils soil evident (Chapman 1984). The community is (Curtis 1959, Trager 1998). Large ant mounds, dominated by the following prairie grasses, which which may measure half a meter in height and can occur in varying degrees of dominance to one over one meter wide and number 40 to 50 per acre another: little bluestem, big bluestem, Indian are especially conspicuous following a prairie fire grass, and prairie dropseed. Pennsylvania sedge (Curtis 1959). Because of their abundance and (Carex pensylvanica) may also be co-dominant frequent habit of abandoning old mounds and where drought-like conditions in late summer and building of new ones, ants overturn large portions fall are a common occurrence. Within Michigan, of prairies in a relatively short time (Curtis 1959). species composition varies across ecoregions. The Other important species contributing soil mixing table below summarizes species composition of and aeration include moles, voles, mice, skunks, mesic sand prairie for the High Plains subsection ground hogs, ground squirrels, and badgers in northern Lower Michigan, the western northern (Curtis 1959). Lower Peninsula (Lake and Oceana counties), and the Interlobate Region of southern Lower Historically, large herbivores such as bison Michigan. While species composition varies significantly influenced plant species diversity in regionally, it is clear that much overlap exists Michigan prairie and oak savanna ecosystems. among regions. Also evident from the list below is The diet of bison consists of 90% to 95% grasses that within each region, the community is and sedges (Steuter 1997). As bison selectively comprised of species with a wide range of wetland forage on grasses and sedges, they reduce the coefficients (e.g., moisture tolerances), indicating dominance of graminoids and provide a that large fluctuations in local and regional water competitive advantage to forb species. The tables strongly influences community activities of bison, which includes wallowing and composition. trampling, promotes plant species diversity by

Table 1. Mesic sand prairie species composition by region.

Western High Lower Southern Scientific Name Common Name Plains Peninsula Interlobate Grasses, Sedges, and Rushes Agropyron trachycaulum slender wheat grass x Andropogon gerardii big bluestem x x x Andropogon scoparius little bluestem grass x x x Aristida purpurascens three awned grass x x

Grayling Community Assessment - 47 Table 1. (continued)

Western High Lower Southern Scientific Name Common Name Plains Peninsula Interlobate Grasses, Sedges, and Rushes Calamagrostis canadensis blue joint grass x x x Danthonia spicata poverty grass; oatgrass x x x Eragrostis spectabilis purple love grass x x Glyceria striata fowl manna grass x x Panicum boreale northern panic grass x x Panicum implicatum panic grass x x Panicum virgatum switch grass x x Sorghastrum nutans Indian grass x x Spartina pectinata cordgrass x x Sporobolus heterolepis prairie dropseed x x x Carex buxbaumii sedge x x Carex flava sedge x x Carex pellita sedge x x Carex pensylvanica sedge x x x Carex stricta sedge x x Eleocharis elliptica golden seeded spike rush x x Schoenoplectus tabernaemontani softstem bulrush x x Scirpus cyperinus wool grass x x Juncus balticus rush x x x Juncus canadensis Canadian rush x x Juncus effusus soft stemmed rush x x Juncus greenei Greene's rush x x Juncus tenuis path rush x x Juncus vaseyi Vasey's rush x x Forbs Artemisia campestris wormwood x x Aster ericoides heath aster x x Aster umbellatus tall flat top white aster x x Campanula rotundifolia harebell x x x Castilleja coccinea Indian paintbrush x x Comandra umbellata bastard toadflax x x x Coreopsis lanceolata sand coreopsis x Coreopsis tripteris tall coreopsis x Euphorbia corollata flowering spurge x x Euthamia graminifolia grass leaved goldenrod x x x Fragaria virginiana wild strawberry x x x Galium boreale northern bedstraw x Helianthemum canadense common rockrose x x Helianthus divaricatus woodland sunflower x x Helianthus giganteus tall sunflower x Helianthus occidentalis western sunflower x x Heuchera americana alum root x Hieracium gronovii hairy hawkweed x Hieracium longipilum long bearded hawkweed x x

Grayling Community Assessment - 48 Table 1. (continued)

Western High Lower Southern Scientific Name Common Name Plains Peninsula Interlobate Forbs Hieracium venosum rattlesnake weed x Houstonia longifolia long leaved bluets x Iris versicolor wild blue flag x Iris virginica southern blue flag x x Krigia biflora false dandelion x Lechea villosa hairy pinweed x x Liatris aspera rough blazing star x x x Lithospermum caroliniense plains puccoon x x Lobelia cardinalis cardinal flower x x Lobelia spicata pale spiked lobelia x x x Lupinus perennis wild lupine x x Lycopus americanus common water horehound x x Maianthemum canadense Canada mayflower x x Monarda fistulosa wild bergamot x x x Oenothera perennis small sundrops x x Phlox pilosa prairie phlox x Polygonum amphibium water smartweed x Potentilla simplex old field cinquefoil x x Pycnanthemum virginianum common mountain mint x x Rubus pubescens dwarf raspberry x x Rudbeckia hirta black eyed susan x x Senecio pauperculus balsam ragwort x x x Sisyrinchium albidum common blue eyed grass x x Solidago canadensis Canada goldenrod x x Solidago houghtonii Houghton's goldenrod x Solidago juncea early goldenrod x x Solidago nemoralis old field goldenrod x x Solidago rigida stiff goldenrod x Solidago uliginosa bog goldenrod x x Tephrosia virginiana goat's rue x x Triadenum fraseri marsh St. John's wort x Vernonia missurica Missouri ironweed x Zigadenus glaucus white camas x Ferns and Fern Allies Thelypteris palustris marsh fern x x Equisetum hyemale scouring rush x x Equisetum laevigatum smooth scouring rush x x Trees and Shrubs Pinus strobus white pine x x x Pinus banksiana jack pine x x x Pinus resinosa red pine x x Prunus serotina wild black cherry x x Larix laricina tamarack x Quercus ellipsoidalis Hill's oak x

Grayling Community Assessment - 49 Table 1. (continued)

Western High Lower Southern Scientific Name Common Name Plains Peninsula Interlobate Trees and Shrubs Quercus macrocarpa bur oak x Quercus alba white oak x x Populus tremuloides quaking aspen x x Quercus velutina black oak x Chamaedaphne calyculata leatherleaf x Hypericum kalmianum Kalm's st. john's wort x x x Rosa carolina pasture rose x x x Rubus flagellaris northern dewberry x x x Rubus hispidus swamp dewberry x x Salix humilis prairie willow x x x Spiraea alba meadowsweet x x x

Michigan Indicator Species: Southern Interlobate Region: prairie dropseed High Plains Subsection: prairie dropseed (SC), (SC), Clinton’s bulrush (Trichophorum clintonii) Vasey’s rush (Juncus vaseyi) (T), Clinton’s (SC), and colic root (Aletris farinosa). bulrush (Trichophorum clintonii) (SC), and Houghton’s goldenrod (Solidago houghtonii) (T, Other Noteworthy Species: Rare plant species LT). associated with mesic sand prairie are listed below along with their status, which is indicated by the Western Northern Lower Peninsula: prairie following abbreviations: X, extirpated from state; dropseed (SC) and Greene’s rush (Juncus E, State Endangered; T, State Threatened; SC, greenei). State Species of Special Concern; LT, Federally Threatened.

Scientific Name Common Name Status Asclepias hirtella tall green milkweed T Baptisia lactea white false indigo SC Eryngium yuccifolium rattlesnake-master T Helianthus mollis downy sunflower T Juncus vaseyi Vasey's rush T Scleria triglomerata tall nut-rush SC Solidago houghtonii Houghton’s goldenrod T, LT Sporobolus heterolepis prairie dropseed SC Strophostyles helvula trailing wild bean SC Trichophorum clintonii Clinton’s bulrush SC Trichostema dichotomum bastard pennyroyal T Viola novae-angliae New England violet T

Rare animal species associated with mesic sand (Asio otus) (T), northern harrier (Circus cyaneus) prairie include the following: (SC), migrant loggerhead shrike (Lanius ludovicianus migrans) (E), Dickcissel (Spiza Grassland birds: Henslow’s sparrow americana) (SC), western meadowlark (Sturnella (Ammodramus henslowii) (SC), grasshopper neglecta) (SC), and barn owl (Tyto alba) (E). sparrow (Ammodramus savannarum) (SC), short- eared owl (Asio flammeus) (E), long-eared owl

Grayling Community Assessment - 50 Insects: American burying beetle (Nicrophorus seedlings, long established trees and tall shrubs americanus) (E/LE), blazing star borer like black cherry (Prunus serotina) and dogwoods (Papaipema beeriana) (SC), Culver’s root borer (Cornus spp.) typically resprout and can reach (Papaipema sciata) (SC), Silphium borer former levels of dominance within two to three (Papaipema silphii) (T), red-legged spittlebug years. Applying herbicide to the cut stumps will (Prosapia ignipectus) (SC), Sprague’s pygarctia prevent resprouting. (Pygarctia spraguei) (SC), grizzled skipper (Pyrgus centaureae wyandot) (SC), phlox moth To reduce the impacts of management on fire- (Schinia indiana) (E), and Spartina moth intolerant species it will be important to consider a (Spartiniphaga inops) (SC). rotating schedule of prescribed burning in which adjacent management units are burned in alternate Mammals: prairie vole (Microtus ochrogaster) years. This is especially important when planning (E). burns in remnant prairies. Insect species that are restricted to these habitats have already Reptiles: eastern massasauga (Sistrurus c. experienced severe losses in the amount of catenatus) (SC and Federal Candidate Species), available habitat due to forest succession brought black rat snake (Elaphe o. obsoleta) (SC), on by years of fire suppression. By burning Kirtland’s snake (Clonophis kirtlandii) (T), and adjacent management units in alternate years, eastern box turtle (Terrapene c. carolina) (SC). insect species from unburned units may be able to Spotted turtle (Clemmys guttata) and Blanding’s recolonize burned areas (Panzer et al. 1995). turtle (Emydoidea blandingii) may nest in mesic Avian species diversity is also thought to be sand prairie when it occurs adjacent to wetlands. enhanced by managing large areas of grassland as a mosaic of burned and unburned patches (Herkert Conservation and Management: Efforts should et al. 1993). be made to identify, protect, and manage remnants of mesic sand prairie. Several studies to identify Prairie ants (Formica) are an extremely important prairie remnants in Michigan have been component of grassland communities and research undertaken and most remnants are very small indicates that they respond with population and/or occur as narrow strips adjacent to railroads increases to restoration activities, especially (Hauser 1953, Scharrer 1972, Thompson 1970, prescribed fire (Trager 1998). Prescribed burning 1975, and 1983, Chapman 1984). The small size precipitates changes in the dominance of ant and poor landscape context of most prairie species from carpenter and woodland ants remnants makes large-scale restoration of existing (Camponotus and Aphaenogaster) to prairie ants prairies nearly impossible. because it reduces woody vegetation and detritus used by the arboreal and litter- and twig-nesting Managing mesic sand prairie requires frequent species in favor of species restricted to grassland burning, from annual to every two to three years. habitats (Trager 1998). Restorations involving Longer burn intervals will result in tree and tall prairie plantings near old fields or remnant shrub encroachment. Prescribed burning is prairies are typically colonized by several species required to protect and enhance plant species of prairie ants within a few years (Trager 1990). diversity and prevent encroachment of trees and tall shrubs, which outcompete light demanding Controlling invasive species is a critical step in prairie plants. In prairie remnants where fire has restoring and managing mesic sand prairie. By been excluded for long periods (e.g., decades), outcompeting native species, invasives alter local extinctions of plant species are common vegetation structure, reduce species diversity, and (Leach and Givnish 1996). upset delicately balanced ecological processes such as trophic relationships, interspecific In addition to prescribed fire, brush cutting competition, nutrient cycling, soil erosion, accompanied by herbicide application to cut hydrologic balance, and solar insolation (Bratton stumps is an important component of prairie 1982, Harty 1986). At present some of the most restoration. While fires frequently kill woody aggressive invasive species that threaten

Grayling Community Assessment - 51 biodiversity of grassland communities include 250 acres in size. Area sensitive species reed canary grass (Phalaris arundinacea), white requiring large patches of grassland (>100 and yellow sweet clover (Melilotus alba and M. acres) include northern harrier (SC), officinalis), autumn olive (Elaeagnus umbellata), bobolink (Dolichonyx oryzivorus), multiflora rose (Rosa multiflora), common savannah sparrow (Passerculus buckthorn (Rhamnus cathartica), Eurasian sandwichensis), Henslow’s sparrow (SC), honeysuckles (Lonicera maakii, L. morrowii, L. short-eared owl (E), and barn owl (E) tartarica, L. x bella.), and black locust (Robinia (Herkert et al. 1993, Sample and pseudoacacia). Mossman 1997). Patches of grassland less than 50 acres will benefit the least area- In addition to reestablishing ecological processes sensitive grassland birds such as northern such as fire, most restoration sites will require the bobwhite (Colinus virginianus), red- reintroduction of appropriate native species and winged black bird (Agelaius phoeniceus), genotypes. Plants can be reintroduced through American goldfinch (Carduelis tristis), both seeding and seedling transplants. Small, Vesper sparrow (Pooecetes gramineus), isolated prairie remnants may harbor plant field sparrow (Spizella pusilla), song populations that have suffered from reduced gene sparrow (Melospiza melodia), dickcissel flow. Restoration efforts at isolated prairie (SC), and common yellowthroat remnants should consider introducing seeds (Geothlypis trichas) (Herkert et al. 1993). collected from nearby stocks to augment and 3. Maximize interior grassland habitat by maintain genetic diversity of remnant plant establishing circular (best) or square populations. The Michigan Native Plant Producers grassland plantings and avoiding long, Association may be a helpful resource for locating narrow plantings, which increase edge sources prairie plants with Michigan genotypes habitat. (http://www.nohlc.org/MNPPA.htm). 4. Where grassland habitats border forests, strive to create a feathered edge by Several helpful guides are available for restoring allowing prescribed fires to burn through prairies and starting prairie plants from seed adjacent forests as opposed to installing (Packard and Mutel 1997, Nuzzo 1976, firebreaks along the forest edge. Schulenberg 1972). See Packard and Mutel (1997) Grasslands with feathered edges for a comprehensive treatment of the subject and experience lower rates of nest predation additional references. than those with sharply contrasting edges (Ratti and Reese 1988). Restoration and management of grasslands such as mesic sand prairie are critically important to Research Needs: Remaining remnants of mesic grassland birds, which have suffered precipitous sand prairie need to be identified, protected, and population declines due to habitat loss and managed. Further research on the historical plant changing agricultural practices (e.g., early species composition of mesic sand prairie in mowing of hay fields). Detailed habitat Michigan would be useful for developing seed management guidelines for grassland birds have mixes for restoration. Studies designed to been developed by Herkert et al. (1993) and compare plant species composition and abiotc Sample and Mossman (1997). Listed below are factors (soils, landscape position, etc.) among several of the recommendations suggested by prairie types in Michigan are needed to improve Herkert et al. (1993) (see publication for complete community classification. In particular, further list of management guidelines). research is needed to elucidate differences 1. Avoid fragmentation of existing between northern wet-mesic prairie and mesic grasslands. sand prairie. Studies aimed at understanding the 2. Grassland restorations aimed at effects of small, isolated populations on plant supporting populations of the most area- species genetic diversity will provide important sensitive grasslands birds should be at information on managing prairie remnants. least 125 acres and preferably more than Research on the utilization of restored and

Grayling Community Assessment - 52 remnant prairies by grassland birds and insects gerardii - Panicum virgatum - Helianthus will provide useful information for understanding grosseserratus Herbaceous Vegetation; Big how mesic sand prairies contribute to biodiversity. Bluestem - Switchgrass - Sawtooth Sunflower Studies on methods of prairie establishment and Herbaceous Vegetation management, including controlling invasive species, will benefit both ongoing and new efforts Related Abstracts: dry sand prairie, woodland to restore mesic sand prairie. Conservation and prairie, mesic prairie, bur oak plains, oak management efforts will benefit from further openings, oak barrens, lakeplain wet-mesic study of how species composition is influenced by prairie, Culver’s root borer, eastern box turtle, fire frequency, intensity, and periodicity. eastern massasauga, Henslow’s sparrow, migrant loggerhead shrike, northern harrier, red-legged Similar Communities: Lakeplain mesic sand spittlebug, prairie dropseed and Houghton’s prairie, northern wet-mesic prairie, wet-mesic goldenrod. prairie, mesic prairie, woodland prairie, dry sand prairie, bur oak plains, and oak openings Literature Cited: Abrams, M.D., A.K. Knapp and L.C. Hulbert. Other Classifications: 1986. A ten year record of aboveground Michigan Natural Features Inventory circa 1800s biomass in a Kansas tallgrass prairie: Effects Vegetation (MNFI): Grassland of fire and topographic position. American Journal of Botany 73: 1509-15. Michigan Department of Natural Resources Albert, D.A. 1995. Regional landscape (MDNR): G ecosystems of Michigan, Minnesota, and Wisconsin: A working map and classification. The Nature Conservancy U.S. National North Central Forest Experiment Station, Vegetation Classification and International Forest Service – U.S. Department of Classification of Ecological Communities (Faber- Agriculture. St. Paul, MN. Langendoen 2001, NatureServe 2004): CODE; ALLIANCE; ASSOCIATION; Albert, D.A. and M.A. Kost. 1998. Natural COMMON NAME: community abstract for lakeplain wet-mesic prairie. Michigan Natural Features Inventory, V.A.5.N.a; Andropogon gerardii – Lansing, MI. 6 pp. (Sorghastrum nutans) Herbaceous Alliance; V.A.5.N.a; Andropogon gerardii – Bratton, S. P. 1982. The Effects of Exotic Plant Sorghastrum nutans – Schizachyrium and Animal Species on Nature Preserves. scoparium – Aletris farinosa Herbaceous Natural Areas Journal. 2(3):3-13. Vegetation; Big Bluestem – Yellow Indiangrass – Little Bluestem– Northern Chapman, K.A. 1984. An ecological investigation White Colicroot Herbaceous Vegetation of native grassland in southern Lower Michigan. M.S. Thesis, Western Michigan V.A.5.N.a; Schizachyrium scoparium - University, Kalamazoo, MI. 235 pp. Sorghastrum nutans Herbaceous Alliance; Schizachyrium scoparium - Sorghastrum Collins, S.L. and DJ. Gibson. 1990. Effects of fire nutans - Andropogon gerardii - Lespedeza on community structure in tallgrass and mixed capitata Sand Herbaceous Vegetation; Little grass prairie. Pp. 81-98 in S.L. Collins and Bluestem - Yellow Indiangrass - Big L.L. Wallace (eds.), Fire in North American Bluestem - Roundhead Bushclover Sand tallgrass prairies, University of Oklahoma Herbaceous Vegetation Press, Norman, OK.

V.A.5.N.a; Andropogon gerardii - Curtis, J.T. 1959. Vegetation of Wisconsin. The (Calamagrostis canadensis, Panicum University of Wisconsin Press, Madison, WI. virgatum) Herbaceous Alliance; Andropogon 657 pp.

Grayling Community Assessment - 53 Daubenmire, R. 1968. Ecology of fire in Wisconsin. Pp. 182-189 in Proceedings of the grasslands. Advances in Ecological Research Fifth Midwest Prairie Conference. Iowa State 5:209-266. University, Ames, Iowa.

Day, G.M. 1953. The Indian as an ecological Packard, S. and C.F. Mutel. 1997. The tallgrass factor in the northeast forest. Ecology 34:329- restoration handbook for prairies savannas 346 and woodlands. Island Press, Washington D.C. 463pp. Denevan, W.M. 1992. The pristine myth: The landscape of the Americas in 1492. Annals of Panzer, R.D., D. Stillwaugh, R. Gnaedinger, and the Association of American Geographers G. Derkowitz. 1995. Prevalence of remnant 83:369-385. dependence among prairie-and savanna- inhabiting insects of the Chicago region. Faber-Langendoen, D. editor. 2001. Plant Natural Areas Journal 15:101-116. communities of the Midwest: Classification in an ecological context. Association for Ratti, J.T. and K.P. Reese. 1988. Preliminary test Biodiversity Information, Arlington, VA. 61 of the ecological trap hypothesis. Journal of pp. + appendix (705 pp.). Wildlife Management 52:484-491. Harty, F. M. 1986. Exotics and their ecological ramifications. Natural Areas Reich, P.B. and M.D. Abrams. 1990. Fire affects Journal. 6(4):20-26. ecophysiology and community dynamics of Central Wisconsin oak forest regeneration. Herkert, J.R., R.E. Szafoni, V.M. Kleen, and J.E. Ecology 71:2179-90. Schwegman. 1993. Habitat establishment, enhancement and management for forest and Sample, D.W. and M.J. Mossman. 1997. grassland birds in Illinois. Division of Natural Managing habitat for grassland birds: A guide Heritage, Illinois Department of for Wisconsin. Bureau of Integrated Science Conservation, Natural Heritage Technical Services, Department of Natural Resources, Publication #1, Springfield, IL, 20pp. Madison, WI. 154pp.

Kay, C.E. 1995. Aboriginal overkill and Native Scharrer, E.M. 1972. Relict prairie flora of burning: Implications for modern ecosystem southwestern Michigan. Pp. 9-12 in J.H. management. Western Journal of Applied Zimmerman (ed.) Proceedings of the Second Forestry 10:121-126. Midwest Prairie Conference, Madison ,WI. 242 pp. Laubhan, M.K. 1995. Effects of prescribed fire on moist-soil vegetation and macronutrients. Schmalzer, P.A. and C. R. Hinkle. 1992. Soil Wetlands 15:159-66. dynamics following fire in Juncus and Spartina marshes. Wetlands 12:8-21. Leach, M.K. and T.J. Givnish. 1996. Ecological determinants of species loss in remnant Schulenberg, R. 1972. Notes on the propagation prairies. Science 273:1555-1558. of prairie plants. The Morton Arboretum, Lisle IL. 15p. NatureServe. 2004. NatureServe Explorer: An online encyclopedia of life [web application]. Smith, L.M. and J.A. Kadlec. 1985. Fire and Version 1.8. NatureServe, Arlington, Virginia. herbivory in a Great Salt Lake marsh. Available Ecology 66:259-65. http://www.natureserve.org/explorer. Steuter, A.A. 1997. Bison. Pp. 339-347 in Nuzzo, V. 1976. Propagation and planting of Packard, S. and C.F. Mutel (eds.), The prairie forbs and grasses in southern Tallgrass Restoration Handbook for Prairies,

Grayling Community Assessment - 54 Savannas, and Woodlands. Island Press, (eds.), Fire and Ecosystems. Academic Press, Washington D.C. 463 pp. New York, NY.

Thompson, D.Q. and R.H. Smith. 1970. The forest Warners, D.P. 1997. Plant diversity in sedge primeval in the Northeast - a great myth? in meadows: Effects of groundwater and fire. Proceedings of the Tall Timbers Fires Ph.D. dissertation, University of Michigan, Ecology Conference. 10:255-265. Ann Arbor, MI. 231 pp.

Thompson, P.W. 1970. The preservation of prairie stands in Michigan. Pp. 13-14 in J.H. Abstract Citation: Zimmerman (ed.) Proceedings of the Second Kost, M.A. 2005. Natural community abstract for Midwest Prairie Conference, Madison ,WI. mesic sand prairie. Michigan Natural Features 242 pp. Inventory, Lansing, MI. 11 pp.

Thompson, P.W. 1975. The floristic composition

of prairie stands in southern Michigan. pp.

317-331 in M.K. Wali (ed.), Prairie: A Copyright 2004 Michigan State University Board multiple view. The University of North of Trustees. Dakota, Grand Fork, N.D.

Michigan State University Extension is an Thompson. P.W. 1983. Composition of prairie affirmative-action, equal-opportunity stands in southern Michigan and adjoining organization. areas. Pp. 105-111 in R. Brewer (ed.),

Proceedings of the Eighth North American Funding for abstract provided by Michigan Prairie Conference. Department of Military and Veterans Affairs.

Timmins, S.M. 1992. Wetland vegetation

recovery after fire: Eweburn Bog, Te Anau,

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30:383-99.

Trager, J.C. 1990. Restored prairies colonized by

native prairie ants (Missouri, Illinois).

Restoration and Management Notes 8:104-

105.

Trager, J.C. 1998. An introduction to ants

(Formicidae) of the tallgrass prairie. Missouri

Prairie Journal 18:4-8.

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Grayling Community Assessment - 55

Grayling Community Assessment - 56 Appendix 2. Intermittent Wetland

Overview: An herb or herb-shrub wetland found Rank Justification: Intermittent wetlands are rare along lakeshores or in depressions and in the Great Lakes region and typically occur as characterized by fluctuating water levels small (e.g. less than 60 acres), isolated seasonally and from year to year. Intermittent depressions. In Michigan, 29 intermittent wetlands occur in depressions in glacial outwash wetlands have been identified, occupying less and sandy glacial lake plains and in kettles on than 1,800 acres in all. Across the Great Lakes pitted outwash. Soils range from loamy sand and basin, fewer than 10,000 acres of intermittent peaty sand to peaty muck and are very strongly wetland persist in the Great Lakes basin acid to strongly acid. Characteristic vegetation (NatureServe 2005). Historically, widespread fires includes Carex spp. (sedges), Juncus spp. following the turn of the century logging (rushes), sphagnum mosses, and ericaceous drastically altered many peatlands, either shrubs. Intermittent wetlands exhibit converting poor conifer swamp to open peatland characteristics of both peatlands and marshes. systems or destroying the peat and converting Some sites were created when fire burned bogs. peatlands to wetlands without organic soils (mineral soil wetlands) (Dean and Coburn 1927, Global and State Rank: G3/S3 Gates 1942, Curtis 1959). It is possible that some of the current intermittent wetlands are the Range: Intermittent wetlands are an uncommon products of intense fires within bogs that feature of glaciated landscapes of the northern destroyed or partially destroyed surface peats. Great Lakes basin, occurring in Michigan, Beginning in the 1920s, effective fire control by Wisconsin, New York, and Ontario (NatureServe the U.S. Forest Service and state agencies reduced 2005). Within Michigan, intermittent wetlands the acreage of fires ignited by man or lightning occur in the Lower Peninsula, almost exclusively (Swain 1973). In landscapes where frequent fire north of the climatic tension zone, and across the was the prevalent disturbance factor, fire Upper Peninsula but primarily in the eastern suppression has caused shrub and tree portion. Intermittent wetlands and other peatland encroachment within intermittent wetlands and systems occur where excess moisture is abundant has likely led to the conversion of some to closed (where precipitation is greater than canopy wetlands (Curtis 1959, Riley 1989). evapotranspiration) (Mitsch and Gosselink 2000). Currently, intermittent wetlands are primarily Conditions suitable for the development of threatened by draining, flooding, filling, intermittent wetlands have occurred in the development, off-road-vehicle (ORV) activity, northern Lake States for the past 3000-5000 years peat mining, logging, and agricultural runoff and following climatic cooling (Heinselman 1970, enrichment (Bedford and Godwin 2003, Boelter and Verry 1977). Several natural peatland NatureServe 2005). Peat mining and cranberry communities that share similarities with farming have degraded numerous peatlands intermittent wetlands also occur in Michigan and throughout the region (Gates 1942, Curtis 1959, can be distinguished from them based on Eggers and Reed 1997, Chapman et al. 2003). comparisons of hydrology, nutrient levels, flora, Michigan, along with Florida and Minnesota, are and distribution. Open wetlands occurring on peat leaders in peat production in the U.S. (Miller include bog, northern fen, and poor fen, all of 1981). In addition to direct impacts to vegetation, which are characterized by stable water levels. alteration of hydrology from road building, ORV Coastal plain marsh, a grass and rush dominated activity, creation of drainage ditches and dams, wetland that also experiences yearly and seasonal and runoff from logging has led to the drastic water level fluctuation, occurs south of the change of intermittent wetland composition and climatic tension zone and is characterized by a structure (Schwintzer and Williams 1974, Riley flora with numerous coastal plain disjuncts (Kost 1989, Chapman et al. 2003). Intermittent wetland and Penskar 2000). vegetation is extremely sensitive to minor changes in water levels and chemistry (Siegel 1988, Riley

Grayling Community Assessment - 57 1989). Succession to more eutrophic wetlands can contain intermittent wetlands support well drained occur as the result of increased alkalinity and and droughty upland communities. The uplands permanently raised water levels, which can cause surrounding intermittent wetlands are dominated the increased decomposition of acidic peats. by fire-dependent conifer systems (i.e., pine Eutrophication from pollution and altered barrens, dry northern forest, and dry-mesic hydrology has detrimentally impacted intermittent northern forest). wetlands by generating conditions favorable for the invasion of exotic species (Riley 1989, Peatlands develop in humid climates where Bedford and Godwin 2003) and dominance by precipitation exceeds evapotranspiration (Boelter aggressive, common natives such as Phalaris and Verry 1977, Gignac et al. 2000). The northern arundinaceae (reed canary grass) and Typha spp. Lake States are characterized by a humid, (cat-tails) (Richardson and Marshall 1986, continental climate with long cold winters and Almendinger and Leete 1998). Permanent short summers that are moist and cool to warm lowering of water tables from drainage can allow (Gates 1942, Boelter and Verry 1977, Damman for tree and shrub encroachment into intermittent 1990, Mitsch and Gosselink 2000). The Michigan wetlands and the eventual succession to closed range of intermittent wetlands falls within the area canopy peatland. The acidity of intermittent classified by Braun (1950) as the Northern wetlands makes them especially susceptible to Hardwood-Conifer Region (Hemlock/White acid rain and air pollution (Siegel 1988, Chapman Pine/Northern Hardwoods Region) and within the et al. 2003). Atmospheric deposition can following regions classified by Albert et al. (1986) contribute nitrogen, sulphur, calcium and heavy and Albert (1995): Region I, Southern Lower metals to intermittent wetlands (Damman 1990, Michigan; Region II, Northern Lower Michigan; Chapman et al. 2003). Dust-fall and atmospheric Region III, Eastern Upper Michigan; and Region deposition from air pollution are particularly IV, Western Upper Michigan. The Northern threats to wetland systems that are surrounded by Hardwood-Conifer Region has a cool snow-forest cultivated land and close to industrial and urban climate with warm summers. The mean number of centers (Damman 1990). freeze-free days is between 90 and 220, and the average number of days per year with snow cover Landscape Context: Intermittent wetlands occur of 2.5 cm or more is between 10 and 140. The on poorly-drained flat areas or mild depressions of normal annual total precipitation ranges from 740 sandy glacial outwash and sandy glacial lake to 900 mm with a mean of 823 mm. The daily plains and in kettle depressions on pitted outwash maximum temperature in July ranges from 24 to (Michigan Natural Features Inventory 2003, 32 °C (75 to 90 °F), the daily minimum NatureServe 2005). This community is found in temperature in January ranges from –21 to –4 °C depressions and along the shores of softwater (-5 to 25 °F) and the mean annual temperature is 7 seepage lakes and ponds where water levels °C (45 °F) (Albert et al. 1986, Barnes 1991). fluctuate both seasonally and yearly. The sandy soils underlying intermittent wetlands are strongly Natural Processes: Water level fluctuations to very strongly acidic and are primarily sands but occur both seasonally and yearly within can also range to loamy sands. Shallow organic intermittent wetlands. Seasonally water levels deposits of peat, muck, or sandy peat may overlay tend to be highest during the winter and spring the sandy substrate. and lowest in late summer and fall. The yearly oscillations are less predictable. Studies of Intermittent wetland may be bordered by several hydrology in related coastal plain marsh systems other wetland communities. For example, have found a pattern of short drawdowns of one to intermittent wetland along lakeshores can three years followed by extensive periods of neighbor submergent marsh, emergent marsh or a inundation (Schneider 1994). Fluctuations of floating bog mat. Along the upper margin of the water level within intermittent wetlands allow for wetland, northern wet meadow, northern shrub temporal variability of the accumulation and thicket or poor conifer swamp may occur. The decomposition of organic matter. Stable periods sandy outwash, pitted outwash and lakeplains that

Grayling Community Assessment - 58 of saturated and inundated conditions inhibit in early spring. This seasonal movement of plant organic matter decomposition and allow for the propagules and organic matter acts to maintain accumulation of peat (Almendinger and Leete diversity and nutrient levels at the upper 1998). Under cool, anaerobic, and acidic elevations of the wetland basin (Schneider 1994). conditions, the rate of organic matter In addition, high water levels can limit tree and accumulation exceeds organic decay (Schwintzer shrub encroachment into intermittent wetlands and Williams 1974, Damman 1990, Mitsch and since prolonged flooding can result in tree and Gosselink 2000). Low levels of oxygen protect shrub mortality. plants from microorganisms and chemical actions that cause decomposition and high levels of Fire is also an important component of the acidity have inhibitory effects on decay organisms disturbance regime of intermittent wetlands. (Heinselman 1963, Miller 1981, Mitsch and Surface fire can contribute to the maintenance of Gosselink 2000). Dam-building activities of open conditions by killing encroaching trees and beaver can result in blocked drainage and flooding shrubs without completely removing the organic which facilitate sphagnum peat development and soils (Curtis 1959, Vitt and Slack 1975). Fire expansion (Heinselman 1963, Heinselman 1970). severity and frequency in intermittent wetlands is High decomposition rates within intermittent closely related to fluctuations in water level. wetlands are correlated with periods of water level Prolonged periods of lowered water table can fluctuation which promote oxidation and the loss allow the vegetation and surface peat to dry out of organic material that would otherwise form enough to burn (Schwintzer and Williams 1974). peat (Miller 1981, Zoltai and Vitt 1995). Water When the surface peat of intermittent wetlands level fluctuations limit the amount of organic burns, the fire releases organic matter from the matter that can accumulate. As noted above, peat, kills seeds and latent buds, stimulates decay, intermittent wetlands often contain a shallow layer and slows peat accumulation (Damman 1990, of organic peat or muck overlaying the sand Jean and Bouchard 1991). Peat fires likely convert substrate. bogs to more graminoid dominated peatlands such as intermittent wetlands and poor fens or if the Water level fluctuation in intermittent wetlands peat is completely destroyed, to mineral soil also facilitates seed germination and seed wetlands such as northern wet meadow (Curtis dispersal, and reduces competition from woody 1959). Because fire has been shown to increase plants. Seasonal drawdowns are critical to the seed germination, enhance seedling establishment, survival of many intermittent wetland species and bolster flowering, fire likely acts as an (especially annuals) that depend on these important mechanism for maintaining plant fluctuations for seed germination. As water levels species diversity and replenishing the seed bank begin to recede in early and mid-summer, direct of intermittent wetlands (Warners 1997). sunlight penetrates the exposed substrate and triggers seed germination (van der Valk 1981). In Vegetation Description: Intermittent wetlands addition, the sunlight warming the soils during the are herb or herb-shrub dominated with a day results in soil temperatures rising during the graminoid dominated herbaceous layer, low day before cooling at night. Diurnal temperature ericaceous, evergreen shrubs, and widely scattered fluctuation also stimulates seed germination for and stunted conifer trees (Michigan Natural many wetland species (Thompson and Grime Features Inventory 2003, NatureServe 2005). The 1983). Season water level fluctuations act as an flora of intermittent wetlands is characteristically important mechanism for seed dispersal dominated by monocotyledons, with annual (Schneider 1994). During the winter and spring species contributing significantly to overall when water levels rise, seeds deposited along the species diversity. For the majority of species, ponds low-water line float to the surface and are flowering and seed set occurs in late summer and carried by wave action to the wetland basin’s fall, when water levels are lowest. However, outer margin. In addition to carrying dormant species with bog affinities found on bog mats seeds, rising water levels also move sprouting within these wetlands tend to be spring flowering seeds and organic matter into the upper shoreline (Curtis 1959). Intermittent wetlands typically

Grayling Community Assessment - 59 contain several vegetation zones, especially when careyi (Carey's smartweed, state threatened), they are adjacent to a lake or pond. The deepest Potamogeton bicupulatus (waterthread pondweed, portion of the depression is usually inundated and state threatened), Pycnanthemum verticillatum supports floating aquatic plants including (whorled mountain mint, state threatened), Brasenia schreberi (water shield), Nuphar Ranunculus cymbalaria (seaside crowfoot, state variegata (bull-head pond-lily), Nymphaea threatened), Schoenoplectus torreyi (Torrey's odorata (sweet-scented water-lily), Potamogeton bulrush, state special concern), and Trichophorum spp. (pondweeds), and Utricularia spp. clintonii (Clinton’s bulrush state special concern). (bladderworts). Occurring along the lower shores and pond margins is a seasonally flooded zone Other Noteworthy Species: Intermittent with sparse graminoid cover with species such as wetlands provide habitat for numerous rare insect Eleocharis olivacea (bright green spike-rush), E. species including Appalachia arcana (secretive robbinsii (Robbin’s spike-rush), Fimbrystilis locust, state special concern), Boloria freija autumnalis (slender fimbry), Juncus spp. (rush), (Freija fritillary, state special concern butterfly), and Scirpus spp. (bulrush). In the saturated soil Boloria frigga (Frigga fritillary, state special further from the shore, where the seasonal water concern butterfly), Erebia discoidalis (red-disked levels typically reach their peak, is a dense alpine, state special concern butterfly), graminoid-dominated zone. This is the most Merolonche dollii (Doll’s merolonche moth, state floristically diverse zone and typically includes special concern), Somatochlora incurvata species such Calamagrostis canadensis (blue-joint (incurvate emerald, state special concern grass), C. stricta (reedgrass), Carex oligosperma dragonfly), and Williamsoni fletcheri (ebony (few-seed sedge), Cladium marisicoides (twig- boghaunter, state special concern dragonfly). Rare rush), Dulichium arundinaceum (three-way herptiles that utilize intermittent wetlands include sedge), Eriocaulon septangulare (seven-angle Clemmys guttata (spotted turtle, state threatened), pipewort), Euthamia graminifolia (flat-topped Elaphe obsoleta obsolete (black rat snake, state goldenrod), Iris versicolor (wild blue flag), special concern), Emys blandingii (Blanding’s Lysimachia terrestris (swamp candles), and turtle, state special concern), Sistrurus catenatus Rhynchospora spp. (beak-rush). Many intermittent catenatus (eastern massasauga, state special wetlands contain a bog mat (often floating) with concern), and Terrapene carolina carolina vegetation typical of an ombrotrophic bog. These (eastern box turtle, state special concern). If bog mats are characterized by sphagnum mosses suitable nesting trees or snags are available, and low, ericaceous shrubs with Chamaedaphne Haliaeetus leucocephalus (bald eagle, state calyculata (leatherleaf) being the most prevalent. threatened), Falco columbarius (merlin, state Trees within intermittent wetlands are either threatened), and Pandion haliaetus (osprey, state absent or widely scattered and stunted. The most threatened) can be found nesting in these systems common canopy dominants are Picea mariana and Ardea herodias (great blue heron, protected (black spruce) and Larix laricina (tamarack). by the Migratory Bird Treaty Act of 1918) can Additional associates include Pinus banksiana establish rookeries. Other rare birds that could (jack pine) and Pinus strobus (white pine). occur in intermittent wetlands are Asio flammeus (short-eared owl, state endangered), Botaurus Michigan Indicator Species: Bartonia lentiginosus (American bittern, state special paniculata (panicled screwstem, state threatened), concern), Circus cyaneus (northern harrier, state Carex nigra (black sedge, state endangered), special concern), Coturnicops noveboracensis Carex wiegandii (Wiegand's sedge, state (yellow rail, state threatened), Gallinula threatened), Eleocharis melanocarpa (black- chloropus (common moorhen, state special fruited spike-rush, state special concern), concern), Ixobrychus exilis (least bittern, state Eleocharis robbinsii, Eriocaulon septangulare, threatened), Phalaropus tricolor (Wilson's Gratiola virginiana (round-fruited hedge hyssop, phalarope, state special concern), and Rallus state threatened), Hemicarpha micrantha (dwarf elegans (king rail, state endangered). Gavia immer bulrush, state special concern), Juncus vaseyi (common loon, state threatened) establish nest (Vasey’s rush, state threatened), Polygonum sites on natural islands and bog-mats. Alces alces

Grayling Community Assessment - 60 (moose, state threatened), Canis lupus (gray wolf, of management to hydrologic regimes, especially state threatened), and Lynx canadensis (lynx, state increased surface flow and alteration of endangered) could utilize intermittent wetland groundwater recharge. This can be accomplished habitat. by establishing a no-cut buffer around intermittent wetlands and avoiding road construction and Intermittent wetlands support a large number of complete canopy removal in adjacent stands. A rare plants, including Bartonia paniculata serious threat to intermittent wetland hydrology (panicled screwstem, state threatened), Carex and species diversity is posed by ORV traffic, nigra (black sedge, state endangered), Carex which can drastically alter the hydrology through wiegandii (Wiegand's sedge, state threatened), rutting and erosion. Soil erosion resulting from Eleocharis melanocarpa (black-fruited spike-rush, ORV use within the wetland or surrounding state special concern), Gentiana linearis (linear- uplands may greatly disturb the seed bank, leaved gentian, state threatened), Gratiola reducing plant density and diversity (Wisheu and virginiana (round-fruited hedge hyssop, state Keddy 1989). For species that depend on threatened), Hemicarpha micrantha (dwarf recruitment from the seed bank such as annuals, bulrush, state special concern), Huperzia selago significant soil disturbance may result in (fir clubmoss, state special concern), Juncus extirpation from the site. Reduction of access to vaseyi (Vasey’s rush, state threatened), Ludwigia wetland systems will help decrease detrimental alternifolia (seedbox, state special concern), impacts from ORVs. Where shrub/tree Lycopodiella margueriteae (northern prostrate encroachment threatens to convert open wetlands clubmoss, state special concern), Lycopodiella to shrub-dominated systems or forested swamps, subappressa (northern appressed clubmoss, state prescribed fire can be employed to maintain open threatened), Polygonum careyi (Carey's conditions. Prescribed fires should be carried out smartweed, state threatened), Potamogeton in intermittent wetlands during droughts or in the bicupulatus (waterthread pondweed, state late summer and fall when water levels are lowest. threatened), Pycnanthemum verticillatum In addition to controlling woody invasion, fire (whorled mountain mint, state threatened), will likely promote seed bank expression and Ranunculus cymbalaria (seaside crowfoot, state rejuvenation. threatened), Sabatia angularis (rose-pink, state threatened), Schoenoplectus torreyi (Torrey's Research Needs: Intermittent wetland is one of bulrush, state special concern), and Trichophorum the least studied wetland community types of the clintonii (Clinton’s bulrush state special concern). Great Lakes region. Classification research is needed that explores the interrelationship between Conservation and Biodiversity Management: floristic composition and structure and Intermittent wetland is an uncommon community physiography, hydrology, and fire. Intermittent type in the Great Lakes region that contributes wetland and related community types (bog, poor significantly to the overall biodiversity of fen, and northern fen) are frequently difficult to Michigan by providing habitat for a unique suite differentiate. Research on abiotic and biotic of plants and wide variety of animal species. indicators that help distinguish related wetlands Numerous rare species are associated with would be useful for field classification. intermittent wetlands. Protection of the regional Systematic surveys for intermittent wetlands and and local hydrologic regime is critical to the related wetlands are needed to help prioritize preservation of intermittent wetlands (Schneider conservation and management efforts. 1994). Stabilization of water levels can allow for the establishment of perennials and woody species Little is known about the hydrologic and fire which can displace less competitive annuals. Even regimes of intermittent wetlands and the small changes in hydroperiod may cause interaction of disturbance factors within these significant shifts in wetland community systems. An important research question to composition and structure. Resource managers address is how so fire and flooding influence operating in uplands adjacent to intermittent species diversity of intermittent wetlands. As wetlands should take care to minimize the impacts noted by Hammerson (1994), beaver significantly

Grayling Community Assessment - 61 alter the ecosystems they occupy. An important Related Abstracts: American bittern, Blanding’s research question to examine is how the wetland turtle, bog, coastal plain marsh, common ecosystems of the Great Lakes have been and moorhen, eastern box turtle, eastern massasauga, continue to be affected by fluctuations in great blue heron rookery, incurvate emerald, king populations of beaver. Experimentation is needed rail, least bittern, merlin, northern appressed to determine how best to prevent shrub and tree clubmoss, northern fen, northern harrier, northern encroachment in intermittent wetlands that are wet meadow, panicled screw-stem, poor fen, threatened by conversion to shrub thicket or prairie fen, rich conifer swamp, secretive locust, conifer swamp. A better understanding is needed spotted turtle, and yellow rail. of the influence of direct and indirect anthropogenic disturbance on intermittent Literature Cited: wetlands. Effects of management within Albert, D.A. 1995. Regional landscape intermittent wetlands should be monitored to ecosystems of Michigan, Minnesota, and allow for assessment and refinement. Monitoring Wisconsin: A working map and classification. should also focus on how intermittent wetland Gen. Tech. Rep. NC-178. St. Paul, MN: U.S. succession and management influence populations Department of Agriculture, Forest Service, of rare species. Given the sensitivity of peatlands North Central Forest Experiment Station. to slight changes in hydrology and nutrient Northern Prairie Wildlife Research Center availability, it is important for scientists to predict Home Page. how peatlands will be affected by global warming http://www.npwrc.usgs.gov/resource/1998/rla and atmospheric deposition of nutrients and ndscp/rlandscp.htm. (Version 03JUN98.) 250 acidifying agents (Heinselman 1970, Riley 1989, pp. Bedford et al. 1999, Gignac et al. 2000, Mitsch and Gosselink 2000). Albert, D.A., S.R. Denton, and B.V. Barnes. 1986. Regional landscape ecosystems of Michigan. Similar Communities: bog, coastal plain marsh, Ann Arbor, MI: University of Michigan, emergent marsh, northern fen, northern wet School of Natural Resources. 32 pp. & map. meadow, poor fen. Almendinger, J.C., J.E. Almendinger, and P.H. Other Classifications: Glaser. 1986. Topographic fluctuations across Michigan Natural Features Inventory Circa 1800 a spring fen and raised bog in the Lost River Vegetation (MNFI): Intermittent Wetland (6228) Peatland, northern Minnesota. Journal of Ecology 74(2): 393-401. Michigan Department of Natural Resources (MDNR): D-treed bog, V-bog, N-marsh Barnes, B.V. 1991. Deciduous forest of North America. Pp 219-344 in E. Röhrig and B. Michigan Resource Information Systems Ulrich, eds., Temperate Deciduous Forests. (MIRIS): 62 (non-forested wetland), 621 (aquatic Elsevier, Amsterdam. 635 pp. bed wetland), and 622 (emergent wetland) Bedford, B.L., and K.S. Godwin. 2003. Fens of The Nature Conservancy National Classification: the United States: Distribution, CODE; ALLIANCE; ASSOCIATION; characteristics, and scientific connection COMMON NAME versus legal isolation. Wetlands 23(3): 608-

629. IV.A.1.N.f; Chamaedaphne calyculata –

(Kalmia angustifolia) Seasonally Flooded Bedford, B.L., M.R. Walbridge, and A. Aldous. Dwarf-Shrubland Alliance; Chamaedaphne 1999. Patterns in nutrient availability and calyculata / Carex oligosperma / Sphagnum plant diversity of temperate North American spp. Dwarf-shrubland; Leatherleaf / Few-seed wetlands. Ecology 80(7): 2151-2169. Sedge / Peatmoss Species Dwarf-shrubland;

Great Lakes Leatherleaf Intermittent Wetland

Grayling Community Assessment - 62 Boelter, D.H., and E.S. Verry. 1977. Peatland and Gignac, L.D., L.A. Halsey, and D.H. Vitt. 2000. A water in the northern Lake States. North bioclimatic model for the distribution of Central Forest Experiment Station. USDA Sphagnum-dominated peatlands in North Forest Service General Technical Report NC- America under present climatic conditions. 31. 26 pp. Journal of Biogeography 27(5): 1139-1151.

Braun, E.L. 1950. Deciduous forests of eastern Hammerson, G. 1994. Beaver (Castor North America. Hafner Press, New York, NY. canadensis): Ecosystem alterations, 596 pp. management, and monitoring. Natural Areas Journal 14(1): 44-57. Chapman, S., A. Buttler, A.-J. Francez, F. Laggoun-Defarge, H. Vasander, M. Schloter, Heinselman, M.L. 1963. Forest sites, bog J. Combe, P. Grosvernier, H. Harms, D. processes, and peatland types in the Glacial Epron, D. Gilbert, and E. Mitchell. 2003. Lake Region, Minnesota. Ecological Exploitation of northern peatlands and Monographs 33(4): 327-374. biodiversity maintenance: A conflict between economy and ecology. Frontiers in Ecology Heinselman, M.L. 1970. Landscape evolution, and the Environment 1(10): 525-532. peatland types, and the environment in the Lake Agassiz Peatland Natural Area, Curtis, J.T. 1959. Vegetation of Wisconsin: An Minnesota. Ecological Monographs 40(2): Ordination of Plant Communities. University 235-261. of Wisconsin Press, Madison, WI. 657 pp. Jean, M., and A. Bouchard. 1991. Temporal Damman, A.H. 1990. Nutrient status of changes in wetland landscapes of a section of ombrotrophic peat bogs. Aquilo Series the St. Lawrence River, Canada. Botanica 28: 5-14. Environmental Management 15(2): 241-250.

Dean, D., and H. Coburn. 1927. An ecological Kost, M.A, and M.R. Penskar. 2000. Natural study of Linne Bog, Cheboygan County, community abstract for coastal plain marsh. Michigan with special reference to Michigan Natural Features Inventory, Nemopanthus mucranata (L.) Trelease. Lansing, MI. 5 pp. Paper’s of the Michigan Academy of Science, Arts, and Letters 8: 87-96. Michigan Natural Features Inventory. 2003. Draft description of Michigan natural community Eggers, S.D., and D.M. Reed. 1997. Wetland types. (Unpublished manuscript revised plants and plant communities of Minnesota March 4, 2003.) Michigan Natural Features and Wisconsin. U.S. Army Corps of Inventory, Lansing, MI. 36 pp. Available: Engineers, St Paul, Minnesota. 263 pp. http://www.msue.msu.edu/mnfi/lists/natural_c ommunity_types.pdf. Faber-Langendoen, D., ed., 2001. Plant communities of the Midwest: Classification in Mitsch, W.J., and J.G. Gosselink. 2000. Wetlands. an ecological context. Association for John Wiley and Sons, Inc, New York, NY. Biodiversity Information, Arlington, VA. 61 920 pp. pp & appendix (705 pp.). Miller, N. 1981. Bogs, bales, and BTU’s: A Gates, F.C. 1942. The bogs of northern Lower primer on peat. Horticulture 59: 38-45. Michigan. Ecological Monographs 12(3): 213-254. NatureServe. 2005. NatureServe Explorer: An online encyclopedia of life [web application]. Version 4.2. NatureServe, Arlington, Virginia.

Grayling Community Assessment - 63 Available: Warners, D.P. 1997. Plant diversity in sedge http://www.natureserve.org/explorer. meadows: effects of ground water and fire. (Accessed: March 03, 2005.) Ph.D. Diss., University of Michigan, Ann Arbor. 231 pp. Richardson, C.J., and P.E. Marshall. 1986. Processes controlling movement, storage, and Wisheu, I.C., and Keddy, P.A. 1989. The export of phosphorous in a fen peatland. conservation and management of a threatened Ecological Monographs 56(4): 279-302. coastal plain plant community in eastern North America (Nova Scotia, Canada). Riley, J.L. 1989. Southern Ontario bogs and fens Biological Conservation 48: 229-238. of the Canadian Shield. Wetlands: Inertia or Momentum: 355-367. Zoltai, S.C., and D.H. Vitt. 1995. Canadian wetlands: Environmental gradients and Schneider, R. 1994. The role of hydrologic regime classification. Vegetatio 118: 131-137. in maintaining rare plant communities of New York’s coastal plain pondshores. Biological Conservation 68:253-260. Abstract Citation:

Schwintzer, C.R, and G. Williams. 1974. Cohen, J.G, and M.A. Kost. 2005. Natural Vegetation changes in a small Michigan bog community abstract for intermittent wetland. from 1917 to 1972. American Midland Michigan Natural Features Inventory, Naturalist 92(2): 447-459. Lansing, MI. 8 pp.

Siegel, D.I. 1988. Evaluating cumulative effects of disturbance on the hydrologic function of bogs, fens, and mires. Environmental Management 12(5): 621-626.

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Thompson, K. and J.P. Grime. 1983. A comparative study of germination responses to diurnally-fluctuating temperatures. Journal of Applied Ecology 20:141-156. van der Valk, A.G. 1986 The impact of litter and annual plants on recruitment from the seed

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Vitt, D.H., and N.G. Slack. 1975. An analysis of the vegetation of Sphagnum-dominated kettle-

hole bogs in relation to environmental gradients. Canadian Journal of Botany 53: 332-359.

Grayling Community Assessment - 64 Appendix 3. Northern Fen

Overview: A sedge and rush dominated wetland (nutrient-poor) (Gignac et al. 2000). The on neutral to slightly alkaline saturated peat hydrology of fens is influenced by ground-water influenced by groundwater rich in calcium and and as a result, fens have higher nutrient magnesium carbonates. Occurs north of the availability, increased alkalinity (less acidity), and climatic tension zone where calcareous bedrock greater species richness compared to bogs, with underlies a thin mantle of glacial drift on flat areas poor fens being most similar to bogs in terms of or mild depressions of glacial outwash and glacial these factors and species composition. Like lake plains. northern fen and poor fen, prairie fens are graminoid dominated and groundwater Global and State Rank: G4G5/S3 influenced, however prairie fens are restricted to south of the climatic tension zone. In addition to a Range: Northern fen is a peatland type of greater importance by graminoids in fens versus glaciated landscapes of the northern Great Lakes bogs, nutrient-rich fens also are less dominated by region, ranging from Michigan west to Minnesota sphagnum mosses (Sphagnaceae) with brown and northward into central Canada (Ontario, mosses (Amblystegiaceae) being more prevalent. Manitoba, and Quebec) (Gignac et al. 2000, Intermittent wetlands are herb or herb-shrub Faber-Langendoen 2001, Amon et al. 2002, dominated wetlands that experience fluctuating NatureServe 2005a). Northern fen may also occur water levels seasonally and yearly and have soils in parts of the northeastern U.S. (i.e., New York that range from loamy sand and peaty sand to and Maine). In Michigan, northern fens occur in peaty muck and are very strongly acid to strongly the northern Lower Peninsula and the Upper acid. Like bogs, muskegs and poor conifer swamp Peninsula, most frequently in close proximity to are nutrient-poor, acidic wetlands. However, these Great Lakes coast. Most documented occurrences ombrotrophic peatlands exhibit a greater degree of from the U.P. are known from the eastern portion. canopy closure than bogs (muskegs having Fens and other peatlands occur where excess clumped and scattered conifers and poor conifer moisture is abundant (where precipitation is swamp being a closed canopy system). Closed greater than evapotranspiration) (Mitsch and canopy, minerotrophic peatlands include rich Gosselink 2000). Conditions suitable for the conifer swamp, a Thuja occidentalis (northern development of fens have occurred in the northern white cedar) dominated system found north of the Lake States for the past 8000 years. Expansion of tension zone, and relict conifer swamp, which is peatlands likely occurred following climatic dominated by Larix laricina (tamarack) and cooling, approximately 5000 years ago occurs primarily south of the tension zone (Kost (Heinselman 1970, Boelter and Verry 1977, Riley 2001). 1989). Rank Justification: Northern fens are uncommon Several other natural peatland communities also features of the northern Great Lakes region, occur in Michigan and can be distinguished from occurring sporadically in Michigan’s northern minerotrophic (nutrient-rich) northern fens, based Lower Peninsula and the Upper Peninsula. The on comparisons of nutrient levels, flora, canopy northern lake states contain over six million closure, distribution, and ground-water influence. hectares (15 million acres) of peatland (Boelter Northern fens are dominated by sedges, rushes, and Verry 1977). What percentage of that area is and grasses (Mitsch and Gosselink 2000). northern fen has yet to be determined. Likewise, Additional open wetlands occurring on peat the current status of fens relative to their historical include bog, poor fen, and prairie fen. Bogs, peat- status is unknown (Bedford and Godwin 2003). covered wetlands raised above the surrounding Peatland scientists concur that fens have always groundwater by an accumulation of peat, receive been localized and not very abundant but have inputs of nutrients and water primarily from suffered from extensive loss, fragmentation, and precipitation and are classified as ombrotrophic degradation (Bedford and Godwin 2003,

Grayling Community Assessment - 65 NatureServe 2005a). Historically, widespread detrimentally impacted fens by generating fires following the turn of the century logging conditions favorable for the invasion of exotic drastically altered many peatlands, either species (Riley 1989, Bedford and Godwin 2003) converting conifer swamp to open fen systems or and dominance by aggressive, common natives destroying the peat and converting peatlands to such as Phalaris arundinaceae (reed canary grass) wetlands without organic soils (mineral soil and Typha spp. (cat-tails) (Richardson and wetlands) (Dean and Coburn 1927, Gates 1942, Marshall 1986, Almendinger and Leete 1998b). Curtis 1959). Logging of cedar and tamarack from Bedford et al. (1999) have noted a widespread peatland systems also favored the conversion of decline in wetland species richness associated forested peatlands to open peatlands (Gates 1942, with the overall eutrophication of the landscape: Dansereau and Segadas-Vianna 1952, Riley nutrient enrichment has converted numerous 1989). Beginning in the 1920s, effective fire species rich wetlands such as northern fen into control by the U.S. Forest Service and state monospecific stands of nitrophilic species. agencies reduced the acreage of fires ignited by Lowering of water tables from drainage has man or lightning (Swain 1973). In landscapes allowed for tree and shrub encroachment into where frequent fire was the prevalent disturbance open fens and the eventual succession to closed factor, fire suppression has led to the conversion canopy peatland (Almendinger and Leete 1998b). of open fens to closed canopy peatlands or shrub Increased shrub and tree canopy cover typically thickets (Curtis 1959, Schwintzer 1981, Riley results in decreased species richness of fen 1989). systems (Bowles et al. 1996). In addition, lowering of the water table can reduce carbonate Currently, fens are primarily threatened by peat deposition at the fen surface and thereby alter the mining, logging, quarrying, agricultural runoff growing conditions, causing a loss in rare and enrichment, draining, flooding, off-road- calciphilic vegetation and an increase in more vehicle (ORV) activity, and development common plants (Almendinger and Leete 1998b). (Bedford and Godwin 2003, NatureServe 2005a). The high alkalinity of fens makes them especially Peat mining and cranberry farming have degraded susceptible to acid rain and air pollution (Siegel numerous peatlands throughout the region (Gates 1988, Chapman et al. 2003). Atmospheric 1942, Curtis 1959, Eggers and Reed 1997, deposition can contribute Nitrogen, Sulphur, Chapman et al. 2003). Michigan, along with Calcium, and heavy metals to fens (Damman Florida and Minnesota, are leaders in peat 1990, Chapman et al. 2003). Dust-fall and production in the U.S. (Miller 1981). In addition atmospheric deposition from air pollution are to direct impacts to vegetation, alteration of particularly threats to fen systems that are peatland hydrology from road building, quarrying, surrounded by cultivated land and close to ORVs, creation of drainage ditches and dams, and industrial and urban centers (Damman 1990). runoff from logging has led to the drastic change of peatland composition and structure (Schwintzer Physiographic Context: Two landscape features and Williams 1974, Schwintzer 1978a, Riley are conducive to the development of peat; poorly- 1989, Bedford and Godwin 2003, Chapman et al. drained, level terrain and small ice-block basins 2003). Fen vegetation is extremely sensitive to (Boelter and Verry 1977). Northern fen occur on minor changes in water levels and chemistry, flat areas or mild depressions of glacial outwash ground water flow, and nutrient availability and glacial lake plains, often in close proximity to (Siegel 1988, Riley 1989). A reduction in the Great Lakes shoreline, and also in kettle groundwater flow and subsequent decrease in depressions on pitted outwash and moraines nutrients in northern fens can result in the shift to (Gates 1942, Verry 1975, Vitt and Slack 1975, less minerotrophic wetlands such as a poor fens or Boelter and Verry 1977, Schwintzer 1978a, Siegel even a bog. Conversion to more eutrophic 1988, Michigan Natural Features Inventory 2003, wetlands has occurred as the result of nutrient NatureServe 2005a). The overall topography of enrichment and raised water levels, which cause fens is flat to gently undulating with increased decomposition of peat. Eutrophication microtopography characterized by hummocks and from pollution and altered hydrology has hollows (Heinselman 1963, Vitt and Slack 1975,

Grayling Community Assessment - 66 Wheeler et al. 1983, Siegel 1988, NatureServe Tomberlin 1982, Riley 1989, Bedford and 2005a). Fens found in kettle depressions are Godwin 2003). Ground-water discharge produces associated with active or extinct glacial lakes that continuously saturated conditions in the rooting are alkaline (Vitt and Slack 1975). Within kettle zone. The water table of fens is stable, typically at depressions, fens can occupy the entire basin or the soil surface with the peat soils saturated but frequently occur as a floating mat on the margin seldom flooded (Heinselman 1970, Schwintzer of the remaining glacial lake (Vitt and Slack 1975, 1978b, Schwintzer 1981, Riley 1989, Amon et al. Schwintzer 1978a, Schwintzer 1978b). When fens 2002, Bedford and Godwin 2003). The cool occur along the edge of large bodies of water, they ground water which enters fens is telluric, having are found in sheltered bays or coves that are moved over or percolated through base-rich protected from wave and ice action, which can bedrock, calcareous glacial deposits, or mineral prevent the development of peat or erode existing soil (Schwintzer 1978b, Bedford and Godwin peat mats (Gates 1942, NatureServe 2005a). Fens 2003). As a result, the ground water discharge into occurring on former glacial lake beds and fens is mineral rich, carrying high concentrations drainageways tend to be more extensive than of calcium and magnesium carbonates (Curtis kettle fens, which are limited in area by the size of 1959, Heinselman 1970, Verry 1975, Boelter and the glacial ice-block which formed the basin Verry 1977, Schwintzer 1978b, Schwintzer 1981, (Lindeman 1941): the large peatlands of Almendinger et al. 1986, Almendinger and Leete lakeplains and outwash plains are often over 100 1998b, Mitsch and Gosselink 2000, Amon et al. acres while fens found in kettle depressions 2002, Bedford and Godwin 2003, NatureServe typically range from 10 to 50 acres. Northern fens 2005a). In addition to high levels of dissolved occurring on glacial outwash and glacial lake minerals, the groundwater of fens is alkaline to plains occur on sapric to fibric peat or marly flats circumneutral and characterized by high specific overlaying calcareous bedrock, typically dolomite conductivity, cool temperature, and a clear color or limestone of Devonian age (Heinselman 1970, resulting from low levels of dissolved organic Schwintzer 1978b, Schwintzer 1981, Amon et al. matter (Verry 1975, Glaser et al. 1981, Wheeler et 2002, NatureServe 2005a). The majority of al. 1983, Riley 1989, Glaser et al. 1990). documented northern fens within Michigan occur Scientists studying minerotrophic fens in the on old glacial lake beds in close proximity to a Great Lakes have reported a wide range of pH Great Lakes shoreline. values (5.0-8.0) (Heinselman 1970, Boelter and Verry 1977, Schwintzer 1978b, Glaser et al. 1981, Northern fens often occur within large wetland Wheeler et al. 1983, Siegel and Glaser 1987, complexes, typically adjacent to other peatland Riley 1989, Glaser et al. 1990). Within northern communities, grading into rich conifer swamp, fens of Michigan, recorded pH values range poor fen, bog, muskeg, and poor conifer swamp. between 5.6 and 8.0. The degree of minerotrophy Northern fens within kettle depressions that of a given fen and within a fen depends on the contain active glacial lakes and ponds often kind and amount of groundwater discharge, border aquatic communities such as submergent degree of dilution from precipitation, the marsh and emergent marsh. Northern fen can also characteristics of the bedrock and/or glacial occur as one of many zones within matrix deposits the groundwater has percolated through communities such as Great Lakes marsh and (i.e., older glacial sediments have less dissolved wooded dune and swale complex. Upland minerals due to prior leaching), the distance the community types are found adjacent to northern water has traveled through the peatland, and the fen include dry-mesic northern forest, dry thickness and character of the peat (Heinselman northern forest, pine barrens, and boreal forest. 1963, Heinselman 1970, Boelter and Verry 1977, Siegel and Glaser 1987, Amon et al. 2002). Hydrology: Northern fens are minerotrophic peatlands, receiving inputs of water and nutrients Soils: The organic soils of fens are composed primarily from nutrient-rich groundwater primarily of peat which frequently forms a (Heinselman 1970, Vitt and Slack 1975, Boelter shallow, continuous mat and is typically 1-3 and Verry 1977, Schwintzer 1981, Schwintzer and meters deep (Glaser et al. 1981). Peat is a fibrous

Grayling Community Assessment - 67 network of partially decomposed organic material Climate: Peatlands develop in humid climates that is formed under anaerobic conditions where precipitation exceeds evapotranspiration (Almendinger et al. 1986, Heinselman 1963). The (Boelter and Verry 1977, Gignac et al. 2000, surface peats of fens are saturated, range from Bedford and Godwin 2003). The northern Lake sapric to fibric peat, and like the surface water, are States are characterized by a humid, continental alkaline to neutral and characterized by high climate with long cold winters and short summers nutrient availability (Curtis 1959, Heinselman that are moist and cool to warm (Gates 1942, 1963, Heinselman 1970, Schwintzer and Williams Boelter and Verry 1977, Damman 1990, Mitsch 1974, Boelter and Verry 1977, Almendinger et al. and Gosselink 2000). The Michigan range of 1986, Swanson and Grigal 1989, NatureServe northern fen falls within the area classified by 2005a). Sapric peat, which is held together by Braun (1950) as the Northern Hardwood-Conifer roots and rhizomes, is highly decomposed with Region (Hemlock/White Pine/Northern occasional fragments of sedge, reed, and shrub. Hardwoods Region) and within the following Fibric peat, which is loosely compacted, contains regions classified by Albert et al. (1986) and partially decomposed mosses with fragments of Albert (1995): Region II, Northern Lower wood and occasionally sedge. Fibric peat has high Michigan; Region III, Eastern Upper Michigan; water retaining capacity and large intercellular and Region IV, Western Upper Michigan. The pores that permit rapid water movement (The rate Northern Hardwood-Conifer Region has a cool of water movement through saturated fibric peat is snow-forest climate with short, warm summers, 1000 times faster than water movement through cold winters, and a large number of cloudy days. sapric peats) (Boelter and Verry 1977). Hemic The mean number of freeze-free days is between peats are intermediate between sapric and fibric 90 and 160, and the average number of days per peats in terms of decomposition and water year with snow cover of 2.5 cm or more is retaining capacity (Boelter and Verry 1977, Miller between 80 and 140. The normal annual total 1981, Swanson and Grigal 1989). Peats of fens precipitation ranges from 740 to 900 mm with a tend to have lower water retaining capacity mean of 823 mm. The daily maximum compared to the peats of bogs (Miller 1981). Peat temperature in July ranges from 24 to 29 °C (75 to composition changes with depth and depending on 85 °F), the daily minimum temperature in January the successional history of a given fen. Generally, ranges from -21 to -9 °C (-5 to 15 °F) and the fiber content and hydraulic conductivity usually mean annual temperature is 7 °C (45 °F) (Albert decreases with depth; deeper peats are more et al. 1986, Barnes 1991). Temperatures vary less decomposed, retain more water, and drain slower in peatlands compared to the surrounding than surface peats (Verry 1975, Boelter and Verry landscape because of ground-water influence, the 1977). In addition to peat, northern fens often insulating effect of fens’ saturated peat carpet in contain or develop over extensive areas of marl, the growing season, and snow cover in the winter an organic soil with smooth silty texture that (Burns 1906, Heinselman 1963, Curtis 1959, develops when Calcium and Magnesium Glaser 1992). Fens are characterized by carbonate aquatically precipitate (Almendinger microclimates that are cooler in the summer and and Leete 1998b, Amon et al. 2002, Bedford and warmer in the winter compared to the regional Godwin 2003, NatureServe 2005a). While levels climate (Heinselman 1963, Bedford and Godwin of available Calcium, Magnesium, and Nitrogen 2003). are typically high within northern fens, Phosphorous can be limiting (Richardson and Natural Processes: Peat establishment requires Marshall 1986, Riley 1989, Bedford et al. 1999, an abundant supply of water: peatlands occur in Mitsch and Gosselink 2000, Amon et al. 2002). regions where precipitation is greater than Low concentrations of Phosphorous can result evapotranspiration producing substantial from co-precipitation with carbonate, microbial groundwater discharge (Dansereau and Segadas- immobilization, reduced aeration of the rooting Vianna 1952, Boelter and Verry 1977, zone, and iron toxicity (Richardson and Marshall Almendinger and Leete 1998b, Mitsch and 1986, Almendinger and Leete 1998, Amon et al. Gosselink 2000). Saturated and inundated 2002, Bedford et al. 1999).

Grayling Community Assessment - 68 conditions inhibit organic matter decomposition from lake to marsh to fen to poor fen or bog and allow for the accumulation of peat (Heinselman 1963, Boelter and Verry 1977, (Almendinger and Leete 1998b, Amon et al. Schwintzer 1981, Swineheart and Parker 2000). 2002). Under cool and anaerobic conditions, the Once Sphagnum mosses become established on rate of organic matter accumulation exceeds fen peat, they maintain and enhance saturated and organic decay (Schwintzer and Williams 1974, acidic conditions which in turn promote continued Damman 1990, Mitsch and Gosselink 2000). Low sphagnum peat development (Heinselman 1963). levels of oxygen protect plants from The ability of sphagnum to absorb and hold microorganisms and chemical actions that cause cations increases the acidity and low nutrient decay (Miller 1981). Fens have greater levels of availability of peatlands (Osvald 1935, Curtis microbial activity compared to bogs because of 1959, Verry 1975, Vitt and Slack 1975, Boelter the lesser acidity and higher base status of and Verry 1977). In addition, accumulating minerotrophic waters. As a result, organic matter sphagnum peat can dilute groundwater influence decay is greater while peat accumulation is lesser by absorbing large amounts of precipitated water, in fens versus bogs (Heinselman 1970). impeding drainage, and increasing the distance of Development and expansion of fens occurs via the rooting zone from telluric water (Dansereau two different processes in kettle depressions and Segadas-Vianna 1952, Vitt and Slack 1975, versus glacial lakeplain and outwash. Fens Schwintzer 1981). Sphagnum moss, which has develop in glacial lakeplain and outwash where numerous pores, partitions, and capillary space, groundwater influence maintains saturated has an enormous water-holding capacity (Osvald conditions which partially inhibit organic matter 1935, Dansereau and Segadas-Vianna 1952, decomposition and allow peat accumulation Curtis 1959): sphagnum peat can hold 15 to 30 (Almendinger and Leete 1998b). Peat develops times its own weight in water (Miller 1981, vertically and spreads horizontally (Boelter and Mitsch and Gosselink 2000). In addition to Verry 1977). Estimates of vertical accumulation sphagnum peat accumulation, beaver dams can of peat range between 100 to 200cm/1000 years also cause blocked drainage in fens and the (Mitsch and Gosselink 2000). Lake-filling or subsequent succession of fens to bogs terrestrialization occurs in small kettle lakes with (Heinselman 1963, Heinselman 1970). minimal wave action where gradual peat accumulation results in the development of a Fens frequently succeed to northern shrub thicket sedge mat that can fill the basin or occur as a or rich conifer swamp. Lowering of the water floating mat in the lake or as a grounded mat table of fens results in the increase in along the water’s edge (Gates 1942, Bay 1967, decomposition rates of organic matter and the Curtis 1959, Heinselman 1963, Mitsch and consequent accumulation of compact peat, which Gosselink 2000). Floating mats of fen sedges (i.e., is more conducive to shrub and tree growth Carex lasiocarpa) pioneer open water or emergent (Schwintzer and Williams 1974, Schwintzer 1981, marsh. Carex lasiocarpa (wiregrass sedge) Riley 1989, Almendinger and Leete 1998b, possesses rhizomes which can grow out into open Gignac et al. 2000). Conversions of bog to fen can water. The interlacing of rhizomes and roots also occur, however with far less frequency forms a floating mat that is buoyed up in water (Glaser et al. 1990). A discharge of alkaline and accumulates organic matter in the form of groundwater at the peat surface of a bog, caused sapric peat (Gates 1942). Over time fen mats are by a change in hydraulic head, can result in the often invaded by ericaceous shrubs and acidifying conversion of bog vegetation to fen vegetation sphagnum mosses (Osvald 1935, Gates 1942, (Siegel and Glaser 1987, Glaser et al. 1990). Schwintzer and Williams 1974, Swineheart and Mixing of as little as 10% groundwater from Parker 2000). underlying calcareous parent material with acid bog water is sufficient to raise the peatland pH The invasion of sphagnum moss into fen systems from 3.6 to 6.8 (Glaser et al. 1990). Fens and bogs often results in the conversion of fens to more are very sensitive to changes in pH and acidic communities such as poor fen or bog. subsequent availability of nutrients: fen vegetation Succession in lake-filled fens typically proceeds

Grayling Community Assessment - 69 can replace bog flora when pH increases above positioned overhead much of the year. More 4.5 (Siegel 1988). cyclones pass over this area than any other area in the continental U.S. (Frelich and Lorimer 1991). Disturbance factors influencing fens include fire, Trees growing in fens are particularly susceptible flooding, windthrow, and insects. Numerous fens to windthrow because peat provides a poor contain charcoal within their peat profile (Curtis substrate for anchoring trees (Burns 1906). The 1959, Heinselman 1963) and many researchers living roots of woody peatland plants occur in a have reported fire as a prevalent part of fen’s shallow rooting zone, generally restricted to the disturbance regime (Gates 1942, Curtis 1959, Vitt uppermost few centimeters where there is and Slack 1975). Surface fire can contribute to the sufficient oxygen to maintain aerobic respiration maintenance of fens by killing encroaching trees (Glaser and Janssens 1986). The superficial and shrubs without completely removing the peat, rooting of trees results in numerous windthrows which is normally saturated (Curtis 1959, Vitt and (Dansereau and Segadas-Vianna 1952). Tree Slack 1975). Graminoid dominance of fen survival in bogs is also limited by insects and systems can be perpetuated by surface fires parasites. Insect outbreaks of the Pristiphora (Bowles et al. 1996). In addition, many of the erichsonii (larch sawfly) cause heavy mortality of ericaceous plants that thrive in fens are fire Larix laricina (tamarack) while, the plant parasite adapted and often grow densely following fire Arceuthobium pusillum (dwarf mistletoe) kills (Wheeler et al. 1983). Fire severity and frequency Picea mariana (black spruce) (Coburn et al. 1933, in fens is closely related to fluctuations in water Gates 1942, Heinselman 1963). level. Prolonged periods of lowered water table can allow the surface peat to dry out enough to Vegetation Description: Northern fens are burn (Schwintzer and Williams 1974). When the characterized by a unique and diverse heliophilus surface peat of fens burns, the fire releases flora with a rich herbaceous layer dominated by organic matter from the peat, kills seeds and latent graminoids, a patchy to continuous moss carpet buds, stimulates decay, and slows peat with brown mosses (Amblystegiaceae) more accumulation (Damman 1990, Jean and Bouchard prevalent than sphagnum mosses (Sphagnaceae), 1991). Such peat fires can result in the conversion low ericaceous, evergreen shrubs, and widely of peatland to mineral soil wetland. scattered and often stunted conifer trees (Gates 1942, Curtis 1959, Vitt and Slack 1975, Mitsch As noted above, flooding can result in the and Gosselink 2000, Amon et al. 2002, Bedford conversion of fens to bogs. Flooding can also and Godwin 2003, NatureServe 2005a). contribute to fen maintenance. Roots of peatland Floristically fens are among the most diverse of trees are physiologically active near the surface all wetland types in the United States, exhibiting and are quickly killed when the water table rises high within-plot species diversity and high site- following flooding (Glaser and Janssens 1986). level species richness, and also supporting Within kettle fens, flooding-induced tree mortality numerous rare and uncommon bryophytes and is likely greater on grounded mats compared to vascular plants, particularly calciphiles free floating mats: free mats float up with rising (Almendinger and Leete 1998a, Almendinger and water table while grounded mats become Leete 1998b, Bedford and Godwin 2003, inundated and have shallower aerobic zones NatureServe 2005a). Species richness of fens is (Schwintzer 1978a, Schwintzer 1978b, Schwintzer related to geographical location, climatic factors, 1979). In addition to flooding, kettle fens can be nutrient availability, and habitat heterogeneity influenced by waves and ice. Wave and ice action (Glaser et al. 1990, Glaser 1992). Floristic can prevent the expansion of fen mats by eroding diversity within northern fens is correlated with the shoreline vegetation (Gates 1942). high levels of available nutrients and microtopography (Riley 1989, Glaser et al. 1990). The natural disturbance regime in fens is also The high degree of small scale environmental influenced by wind. The Great Lakes region is heterogeneity results in strong vegetational one of the most active weather zones in the zonation (Amon et al. 2002, Bedford and Godwin northern hemisphere, with polar jet streams 2003). Vegetational zones that frequently occur

Grayling Community Assessment - 70 within northern fens include sedge lawns, sparsely 1990, NatureServe 2005a). Hummock and hollow vegetated marl flats, and shrub thickets, which microtopography often occurs in northern fens often occur as narrow bands on the upland and allows for high levels of bryophyte diversity margin. Floristic composition is determined by since individual species of moss can occur at gradients in pH, light, soil moisture, and cation specific elevations (Vitt and Slack 1975, Wheeler concentrations (nutrient availability) (Heinselman et al. 1983, Riley 1989). The vertical zonation of 1970, Vitt and Slack 1975, Schwintzer 1978a, species corresponds to gradients in pH and Glaser et al. 1981, Glaser et al. 1990, Siegel 1988, moisture with the hollows being wetter and more Anderson et al. 1996, Bedford et al. 1999). The alkaline than the drier and more acidic tops of the mean number of vascular species per plot in hummocks (Vitt et al. 1975, Wheeler et al. 1983). northern fens in the northern Lower Peninsula of As noted above, ombrotrophic Sphagnum mosses Michigan was found to be 29 by Vitt and Slack can occur on the tops of hummocks (Amon et al. (1975) and 30 by Schwintzer (1978b) with a range 2002). of 10-53. For northern fens within Michigan Natural Features Inventory’s database, the mean Cyperaceous graminoids dominate the herbaceous number of species per northern fen is layer of fens. The most dominant plant in northern approximately 48. Very few introduced, weedy fens is Carex lasiocarpa (wiregrass sedge), which species are able to establish within bogs and fens, can form extensive lawns (NatureServe 2005a). likely because of the unique growing conditions Sedges that are characteristic of northern fens and competition from the adapted flora. Northern include Carex aquatilis, C. chordorrhiza fens are dominated by plants that thrive under (creeping sedge), C. leptalea (bristly-stalked minertrophic conditions. Occasionally sedge), C. limosa (mud sedge), and C. livida (livid ombrotrophic indicators may be present in fens at sedge). Other sedges that often occur in northern low cover. The tops of hummocks can support fens are Carex buxbaumii (Buxbaum’s sedge), C. sphagnum mosses and a more acidic micro- capillaries (hair-like sedge), C. exilis (coastal environment where these ombrotrophic species sedge), C. interior (inland sedge), C. lacustris can occur isolated from the influence of mineral- (lake or hairy sedge), C. rostrata (beaked sedge), rich ground water (Wheeler et al. 1983, Amon et C. sterilis (dioecious sedge), C. stricta (tussock al. 2002). sedge), and C. viridula (little green sedge). Calamagrostis canadensis (blue-joint grass), C. The patchy to continuous surface carpet of mosses stricta (reedgrass), Muhlenbergia glomerata in northern fens is dominated by calcicolous (marsh wild-timothy), and Panicum lindheimeri brown mosses of the family Amblystegiaceae (panic grass) are typical northern fen grasses. (Glaser et al. 1990, Zoltai and Vitt 1995, Additional graminoids that thrive in northern fens Swinehart and Parker 2000, Amon et al. 2002). include Cladium marisicoides (twig-rush), Typical minerotrophic bryophytes of northern fen Dulichium arundinaceum (three-way sedge), include the following brown mosses: Calliergon Eleocharis compressa (flat-stem spike-rush), trifarium, Campylium stellatum, Drepanocladus Eleocharis elliptica (elliptic spike-rush), revolvens, and Scorpodium scorpoides (Crum Eleocharis rostellata (beaked spike-rush), 1983, Riley 1989, Glaser et al. 1990). Sphagnum Eriophorum angustifolium (tall cotton-grass), mosses are either absent from northern fens or Eriophorum spissum (sheathed cotton-grass), subordinate to the Amblystid mosses (Schwintzer Rhynchospora alba (white beak-rush), R. 1978). Bryum pseudotriquetrum can also occur capillacea (needle beak-rush), Scirpus cespitosus within northern fen. Sphagnum teres thrives in (tufted bulrush), Scirpus hudsonianus (Hudson’s alkaline conditions and is often found in Bay bulrush), and Typha latifolia (cattail). association with Carex lasiocarpa (wiregrass Northern fens frequently contain marl flats that sedge) (Vitt and Slack 1975). Other sphagnum are dominated by twig-rush, beak-rushes, spike- mosses that could occur within northern fens rushes, rushes, and bulrushes (i.e., Eleocharis include Sphagnum angustifolium, S. capillifolium, rostellata). The following is a list of prevalent S. centrale, S. magellanicum, S. subsecundum, S. northern fen herbs: Aster borealis (rush aster), warnstorfii (Vitt and Slack 1975, Glaser et al. Campanula aparinoides (marsh bellflower),

Grayling Community Assessment - 71 Decodon verticillatus (whorled loosestrife), Trees within fens are widely scattered and Euthamia graminifolia (flat-topped goldenrod), typically of low stature (ranging from two to ten Iris versicolor (wild blue flag), Lobelia kalmii meters but seldom reaching six meters) (Wheeler (Kalm’s lobelia), Lycopus uniflorus (northern et al. 1983, NatureServe 2005). Tree cover is bugleweed), Lysimachia terrestris (swamp typically below ten percent. The most common candles), Menyanthes trifoliata (bogbean), dominants of the open canopy are Larix laricina Parnassia palustris (grass-of-Parnassus), (tamarack) and Thuja occidentalis (northern white Potentilla anserine (silverweed), Solidago cedar). Infrequent associates include Picea uliginosa (bog goldenrod), Tofieldia glutinosa mariana (black spruce), Pinus banksiana (jack (false asphodel), Triadenum fraseri (marsh St. pine), and P. strobus (white pine). (Above species John’s-wort), and Triglochin maritimum (arrow- lists compiled from Michigan Natural Features grass). Insectivorous plants, Drosera rotundifolia Inventory database, Gates 1942, Curtis 1959, (roundleaf sundew), Drosera intermedia (spoon- Heinselman 1963, Heinselman 1965, Heinselman leaf sundew), Sarracenia purpurea (pitcher- 1970, Schwintzer and Williams 1974, Vitt and plant), and Utricularia intermedia (flat-leaved Slack 1975, Schwintzer 1978a, Glaser et al. 1981, bladderwort), are common features of fens. The Schwintzer 1981, Schwintzer and Tomberlin fern ally Equisetum fluviatile (water horsetail) is 1982, Wheeler et al. 1983, Richardson and also typical. Marshall 1986, Riley 1989, Glaser et al. 1990, Glaser 1992, Eggers and Reed 1997, Mitsch and Northern fens contain both a tall shrub layer and a Gosselink 2000, Swinehart and Parker 2000, low shrub layer. Typically shrub cover is at least NatureServe 2005a.) 25%. Some areas of fen can contain dense thickets of shrubs (over 60% cover), particularly along the Michigan Indicator Species: bog birch, Carex upland margins and where fire and/or flooding chordorrhiza, C. lasiocarpa , C. limosa, C. have failed to limit shrub encroachment. The low leptalea, northern white cedar, shrubby cinquefoil, shrub layer is usually less than three feet high and tamarack, (Heinselman 1970, Wheeler et al. with Potentilla fruticosa (shrubby cinquefoil), 1983, Anderson et al. 1996). Myrica gale (sweet gale), and Betula pumila (bog birch) often being the most prevalent species. Other Noteworthy Species: Northern fens Other important associates of the low shrub layer provide habitat for numerous rare insect species include Hypericum kalmianum (Kalm’s St. including Appalachia arcana (secretive locust, John’s-wort), Potentilla palustris (marsh state special concern), Merolonche dollii (Doll’s cinquefoil), Rhamnus alnifolia (alder-leaved merolonche moth, state special concern), buckthorn), and Spiraea alba (meadowsweet). Phyciodes batesii (tawny crescent, state special Ericaceous shrubs occur within the low shrub concern), Somatochlora hineana (Hine's emerald, layer of northern fens but with far lesser state and federally endangered), and frequency and density than in bogs and poor fens. Somatochlora incurvata (incurvate emerald, state The following are common heath shrubs of special concern dragonfly). Numerous butterflies northern fens: Andromeda glaucophylla (bog- and moths are restricted to bogs and fens because rosemary), Chamadaephne calyculata their food plants occur within these peatland (leatherleaf), Ledum groenlandicum (Labrador- systems (Riley 1989). Numerous tiny land snails tea), and Vaccinium oxycoccos (small cranberry). are associated with calcareous fens (Bedford and The tall shrub layer of northern fens, which is Godwin 2003). Snail populations of northern fens three to six feet tall, is less dense than the low includes numerous rare species such as Catinella shrub layer and is often restricted to the periphery exile (Pleistocene catinella, state special concern), of the fen. Tall shrubs typical of northern fens Euconulus alderi (land snail, state special include Alnus rugosa (speckled alder), Cornus concern), Hendersonia occulta (cherrystone drop, stolonifera (red-osier dogwood), Salix pedicellaris threatened), Planogyra asteriscus (eastern flat- (bog willow), and Salix petiolaris (slender whorl, state special concern), Vertigo elatior willow). Bog birch and meadowsweet can occur (tapered vertigo, state special concern), Vertigo in both the tall and low shrub layers morsei (six-whorl vertigo, state special concern),

Grayling Community Assessment - 72 and Vertigo pygmaea (crested vertigo, state heleonastes (Hudson Bay sedge, state special concern). Rare herptiles that utilize endangered), Carex scirpoidea (bulrush sedge, northern fens include Clemmys guttata (spotted state threatened), Drosera anglica (English turtle, state threatened), Emys blandingii sundew, state special concern), Empetrum nigrum (Blanding’s turtle, state special concern), (black crowberry, state threatened), Erigeron Pseudacris triseriata maculata (boreal chorus hyssopifolius (hyssop-leaved fleabane, state frog, state special concern), Sistrurus catenatus threatened), Juncus stygius (moor rush, state catenatus (eastern massasauga, state special threatened), Pinguicula vulgaris (butterwort, state concern), and Terrapene carolina carolina special concern), Rubus acaulis (dwarf raspberry, (eastern box turtle, state special concern). state endangered), and Solidago houghtonii (Houghton’s goldenrod, state and federally If suitable nesting trees or snags are available, threatened). Falco columbarius (merlin, state threatened), Haliaeetus leucocephalus (bald eagle, state Conservation and Biodiversity Management: threatened), and Pandion haliaetus (osprey, state Northern fen is a widely distributed but threatened) can be found nesting in these systems uncommon community type in the Great Lakes and Ardea herodias (great blue heron, protected region that contributes significantly to the overall by the Migratory Bird Treaty Act of 1918) can biodiversity of northern Michigan by providing establish rookeries. Other rare birds that could habitat for a unique suite of plants and wide occur in northern fens are Asio flammeus (short- variety of animal species. Numerous rare species eared owl, state endangered), Botaurus are associated with fens, including many lentiginosus (American bittern, state special calciphiles which depend on the carbonate concern), Circus cyaneus (northern harrier, state precipitate. In addition to their high levels of special concern), Coturnicops noveboracensis biodiversity, fens also contribute numerous (yellow rail, state threatened), and Picoides ecosystem services. Fens modulate water arcticus (black-backed woodpecker, state special temperature of connecting surface waters, are concern). Alces alces (moose, state threatened), characterized by high rates of denitrification and Canis lupus (gray wolf, state threatened), and Phosphorous sorption, and serve as critical buffers Lynx canadensis (lynx, state endangered) utilize between downstream waters and nutrients and peatland habitat. Fens provide important habitat other pollutants from the surrounding uplands. for small mammals such as Blarina brevicauda Fens maintain water quality and flows to streams (short-tailed shrew), Castor canadensis (beaver), (Bedford et al. 1999, Bedford and Godwin 2003). Microtus pennsylvanicus (meadow vole), Mustela By storing high levels of sequestered carbon and vison (mink), Ondatra zibethicus (muskrat), and functioning as carbon sinks, fens and related Sorex cinereus (masked shrew). Both muskrats peatlands play an important role in global and beaver can profoundly influence the geochemical cycles. Fens also preserve paleo- hydrology of peatlands. Muskrats create open environmental records: a wealth of information is water channels through the peat and beavers can stored in the remains of plants, animals, and cause substantial flooding through their dam- atmospheric particles deposited and stored in fen building activities (Gates 1942, Heinselman peat profiles (Chapman et al. 2003). 1963). The primary mechanism for preserving fens is to Northern fens support a large number of rare maintain their hydrology. As noted, peatland plants, including many calciphilic species systems are sensitive to slight changes in water (Almendinger and Leete 1998, Bedford and chemistry: modifications in fen hydrology result Godwin 2003). Compared to other wetland types, in changes in peatland vegetation. Perhaps the fen systems support a disproportionate number of greatest threat to northern fens comes from off- threatened and endangered rare plant species road-vehicle traffic, which can destroy (Eggers and Reed 1997). Rare plants associated populations of sensitive species and drastically with northern fens include Cacalia plantaginea alter fen hydrology through rutting. Reduction of (Indian plantain, state special concern), Carex access to peatland systems will help decrease

Grayling Community Assessment - 73 detrimental impacts. Resource mangers operating should be monitored to allow for assessment and in uplands adjacent to fens should take care to refinement. Monitoring should also focus on how minimize the impacts of management to fen succession and management influence hydrologic regimes, especially increased surface populations of rare species. Scientific flow and reduction in groundwater recharge. This understanding of the microbes and invertebrates can be accomplished by establishing a no-cut that thrive in the organic soils of fens is lacking. buffer around fens and avoiding road construction More research is needed to elucidate the and complete canopy removal in stands relationship of chemical factors and nutrients to immediately adjacent to fens. Where shrub/tree floristic community structure of peatlands (Amon encroachment threatens to convert open wetlands et al. 2002). Given the sensitivity of peatlands to to shrub-dominated systems or forested swamps, slight changes in hydrology and nutrient prescribed fire or selective cutting can be availability, it is important for scientists to predict employed to maintain open conditions (Bowles et how peatlands will be affected by global warming al. 1996). Silvicutural management of fens to and atmospheric deposition of nutrients and preserve open canopy should be employed during acidifying agents (Heinselman 1970, Riley 1989, the winter to minimize damage to the organic soils Bedford et al. 1999, Gignac et al. 2000, Mitsch and impacts to the hydrologic regime. and Gosselink 2000). Peat deposits are of great scientific interest because they contain historical Research Needs: Northern fen has a broad ecological records in the form of fossils of plants, distribution and exhibits numerous regional, animals, and organic matter that contributed to the physiographic, hydrologic, and edaphic variants. deposit. Stratigraphical analysis of peat cores The diversity of variations throughout its range provides insights into past climatic change and demands the continual refinement of regional associated vegetation change, floristic classifications that focus on the inter-relationships distribution, the development of wetland between vegetation, physiography, and hydrology ecosystems, and the successional pathways of (Barnes et al. 1982, Heinselman 1963). Northern peatlands (Heinselman 1963, Glaser et al. 1981, fens and related community types (i.e., poor fen, Miller 1981, Glaser and Janssens 1986, Riley bog, and intermittent wetland) are frequently 1989, Gignac et al. 2000). difficult to differentiate (Heinselman 1963, NatureServe 2005a). Research on abiotic and Similar Communities: bog, Great Lakes marsh, biotic indicators that help distinguish similar intermittent wetland, muskeg, patterned fen, poor peatlands would be useful for field classification. conifer swamp, poor fen, prairie fen, rich conifer Systematic surveys for northern fens and related swamp, and wooded dune and swale complex. peatlands are needed to help prioritize conservation and management efforts. Other Classifications: Michigan Natural Features Inventory Circa 1800 Little is known about the fire regimes of northern Vegetation (MNFI): Emergent Marsh (6221), Wet fens and the interaction of disturbance factors Meadow (6224), and Inland Wet Prairie (6227) within these systems. As noted by Hammerson (1994), beaver significantly alter the ecosystems Michigan Department of Natural Resources they occupy. An important research question to (MDNR): D-treed bog, V-bog, N-marsh examine is how the wetland ecosystems of the Great Lakes have been and continue to be affected Michigan Resource Information Systems by fluctuations in populations of beaver. (MIRIS): 62 (non-forested wetland) and 622 Experimentation is needed to determine how best (emergent wetland) to prevent shrub and tree encroachment of fens that are threatened by conversion to shrub thicket The Nature Conservancy National Classification: or conifer swamp. A better understanding is CODE; ALLIANCE; ASSOCIATION; needed of the influence of direct and indirect COMMON NAME anthropogenic disturbance on peatlands (Amon et al. 2002). Effects of management within fens

Grayling Community Assessment - 74 III.B.2.N.g; Betula pumila – (Salix spp.) (Carex rostrata) – Equisetum fluviatile Saturated Shrubland Alliance; Alnus incana – Herbaceous Vegetation; Wiregrass Sedge – Salix spp. - Betula pumila / Chamaedaphne (Swollen-beak Sedge) – Water Horsetail calyculata Shrubland; Speckled Alder – Herbaceous Vegetation; Wiregrass Sedge Willow Species – Bog Birch / Leatherleaf Shore Fen Shrubland; Bog Birch-Willow Shore Fen Related Abstracts: American bittern, black- III.B.2.N.g; Betula pumila – (Salix spp.) backed woodpecker, Blanding’s turtle, bog, Saturated Shrubland Alliance; Betula pumila / cherrystone drop, eastern box turtle, eastern Chamaedaphne calyculata / Carex lasiocarpa massasauga, English sundew, great blue heron Shrubland; Bog Birch / Leatherleaf / rookery, Great Lakes marsh, incurvate emerald, Wiregrass Sedge Shrubland; Bog Birch – Indian plantain, intermittent wetland, Hine’s Leatherleaf Rich Fen emerald, Houghton’s goldenrod, merlin, northern harrier, poor fen, prairie fen, rich conifer swamp, III.B.2.N.g; Betula pumila – (Salix spp.) secretive locust, short-eared owl, spotted turtle, Saturated Shrubland Alliance; Betula pumila and wooded dune and swale complex, and yellow – Dasiphora fruticosa spp. floribunda / Carex rail. lasiocarpa –Trichophorum alpinum Shrubland; Bog Birch – Shrubby-cinquefoil / Literature Cited: Wiregrass Sedge – Alpine Cottongrass Albert, D.A. 1995. Regional landscape Shrubland; Bog Birch – Shrubby-cinquefoil ecosystems of Michigan, Minnesota, and Rich Boreal Fen Wisconsin: A working map and classification. Gen. Tech. Rep. NC-178. St. Paul, MN: U.S. IV.A.1.N.g; Chamaedaphne calyculata Department of Agriculture, Forest Service, Saturated Dwarf-shrubland Alliance; North Central Forest Experiment Station. Chamaedaphne calyculata – Myrica gale / Northern Prairie Wildlife Research Center Carex lasiocarpa Dwarf-shrubland; Home Page. Leatherleaf – Sweet Gale / Wiregrass Sedge http://www.npwrc.usgs.gov/resource/1998/rla Dwarf-shrubland; Leatherleaf – Sweet Gale ndscp/rlandscp.htm. (Version 03JUN98.) 250 Shore Fen pp.

V.A.5.N.m; Calamagrostis canadensis – Albert, D.A., S.R. Denton, and B.V. Barnes. 1986. Carex viridula – Cladium marisicoides – Regional landscape ecosystems of Michigan. Lobelia kalmii Saturated Herbaceous Ann Arbor, MI: University of Michigan, Alliance; Calamagrostis canadensis – Carex School of Natural Resources. 32 pp. & map. viridula – Cladium marisicoides – Lobelia kalmii Herbaceous Vegetation; Bluejoint – Almendinger, J.C., J.E. Almendinger, and P.H. Hairy Sedge – Twig-rush – Ontario Lobelia Glaser. 1986. Topographic fluctuations across Herbaceous Vegetation; Great Lakes Sedge a spring fen and raised bog in the Lost River Rich Shore Fen Peatland, northern Minnesota. Journal of Ecology 74(2): 393-401. V.A.5.N.m; Carex lasiocarpa Saturated Herbaceous Alliance; Carex lasiocarpa – Almendinger, J.A., and J.H. Leete. 1998a. Peat Carex buxbaumii – Trichophorum characteristics and groundwater geochemistry caespitosum Boreal Herbaceous Vegetation; of calcareous fens in the Minnesota River Wiregrass Sedge – Brown Bog Sedge – Basin, U.S.A. Biogeochemistry 43: 17-41. Deerhair Bulrush Boreal Herbaceous Vegetation; Boreal Sedge Rich Fen Almendinger, J.A., and J.H. Leete. 1998b. Regional and local hydrogeology of V.A.5.N.m; Carex lasiocarpa Saturated calcareous fens in the Minnesota River Basin, Herbaceous Alliance; Carex lasiocarpa – USA. Wetlands 18(2): 184-202.

Grayling Community Assessment - 75 Amon, J.P., C.A. Thompson, Q.J. Carpenter, and Chapman, S., A. Buttler, A.-J. Francez, F. J. Mines. 2002. Temperate zone fens of the Laggoun-Defarge, H. Vasander, M. Schloter, glaciated Midwestern USA. Wetlands 22(2): J. Combe, P. Grosvernier, H. Harms, D. 301-317. Epron, D. Gilbert, and E. Mitchell. 2003. Exploitation of northern peatlands and Anderson, D.S, R. B. Davis, S.C. Rooney, C.S. biodiversity maintenance: A conflict between Campbell. 1996. The ecology of sedges economy and ecology. Frontiers in Ecology (Cyperaceae) in Maine peatlands. Bulletin of and the Environment 1(10): 525-532. the Torrey Botanical Club 123(2): 100-110. Coburn, H., D. Dean, and G.M. Grant. 1933. An Barnes, B.V. 1991. Deciduous forest of North ecological study of Bryant’s Bog, Cheboygan America. Pp 219-344 in E. Röhrig and B. County. Michigan. Paper’s of the Michigan Ulrich, eds., Temperate Deciduous Forests. Academy of Science, Arts, and Letters 17: 57- Elsevier, Amsterdam. 635 pp. 65.

Barnes, B.V., K.S. Pregitzer, T.A. Spies, and V. Crum, H. 1983. Mosses of the Great Lakes Forest. H. Spooner. 1982. Ecological forest site University of Michigan, Ann Arbor, MI. 417 classification. Journal of Forestry 80(8): 493- pp. 498. Curtis, J.T. 1959. Vegetation of Wisconsin: An Bay, R.R. 1967. Ground water and vegetation in Ordination of Plant Communities. University two peat bogs in northern Minnesota. Ecology of Wisconsin Press, Madison, WI. 657 pp. 48(2): 308-310. Damman, A.H. 1990. Nutrient status of Bedford, B.L., and K.S. Godwin. 2003. Fens of ombrotrophic peat bogs. Aquilo Series the United States: Distribution, Botanica 28: 5-14. characteristics, and scientific connection versus legal isolation. Wetlands 23(3): 608- Dansereau, P., and F. Segadas-Vianna. 1952. 629. Ecological study of the peat bogs of eastern North America. I. Structure and evolution of Bedford, B.L., M.R. Walbridge, and A. Aldous. vegetation. Canadian Journal of Botany 30: 1999. Patterns in nutrient availability and 490-520. plant diversity of temperate North American wetlands. Ecology 80(7): 2151-2169. Dean, D., and H. Coburn. 1927. An ecological study of Linne Bog, Cheboygan County, Boelter, D.H., and E.S. Verry. 1977. Peatland and Michigan with special reference to water in the northern Lake States. North Nemopanthus mucranata (L.) Trelease. Central Forest Experiment Station. USDA Paper’s of the Michigan Academy of Science, Forest Service General Technical Report NC- Arts, and Letters 8: 87-96. 31. 26 pp. Eggers, S.D., and D.M. Reed. 1997. Wetland Braun, E.L. 1950. Deciduous forests of eastern plants and plant communities of Minnesota North America. Hafner Press, New York, NY. and Wisconsin. U.S. Army Corps of 596 pp. Engineers, St Paul, Minnesota. 263 pp.

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pp & appendix (705 pp.).

Grayling Community Assessment - 76 Fitzgerald, S., and R.E. Bailey. 1975. Lake Region, Minnesota. Ecological Vegetational characteristics of a circum- Monographs 33(4): 327-374. neutral bog, Barney’s Lake, Beaver Island, Michigan. The Michigan Academician 7(4): Heinselman, M.L. 1965. String bogs and other 477-488. patterned organic terrain near Seney, Upper Michigan. Ecology 46: 185-188. Frelich, L.E., and C.G. Lorimer. 1991. Natural disturbance regimes in hemlock-hardwood Heinselman, M.L. 1970. Landscape evolution, forests of the Upper Great Lakes region. peatland types, and the environment in the Ecological Monographs 61(2): 145-164. Lake Agassiz Peatland Natural Area, Minnesota. Ecological Monographs 40(2): Gates, F.C. 1942. The bogs of northern Lower 235-261. Michigan. Ecological Monographs 12(3): 213-254. Jean, M., and A. Bouchard. 1991. Temporal changes in wetland landscapes of a section of Gignac, L.D., L.A. Halsey, and D.H. Vitt. 2000. A the St. Lawrence River, Canada. bioclimatic model for the distribution of Environmental Management 15(2): 241-250. Sphagnum-dominated peatlands in North America under present climatic conditions. Kost, M.A. 2001. Natural community abstract for Journal of Biogeography 27(5): 1139-1151. relict conifer swamp. Michigan Natural Features Inventory, Lansing, MI. 6 pp. Glaser, P.H. 1992. Raised bogs in eastern North America – Regional controls for species Lindeman, R.L. 1941. The developmental history richness and floristic assemblages. Journal of of Cedar Creek Bog, Minnesota. American Ecology 80: 535-554. Midland Naturalist 25(1): 101-112.

Glaser, P.H., and J.A. Janssens. 1986. Raised bogs Michigan Natural Features Inventory. 2003. Draft in eastern North America: Transitions in description of Michigan natural community landforms and gross stratigraphy. Canadian types. (Unpublished manuscript revised Journal of Botany 64: 395-415. March 4, 2003.) Michigan Natural Features Inventory, Lansing, MI. 36 pp. Available: Glaser, P.H., G.A. Wheeler, E. Gorham, and H.E. http://www.msue.msu.edu/mnfi/lists/natural_c Wright, Jr. 1981. The patterned mires of the ommunity_types.pdf. Red Lake Peatland, northern Minnesota: Vegetation, water chemistry and landforms. Mitsch, W.J., and J.G. Gosselink. 2000. Wetlands. Journal of Ecology 69(2): 575-599. John Wiley and Sons, Inc, New York, NY. 920 pp. Glaser, P.H., J.A. Janssens, and D.I. Siegel. 1990. The response of vegetation to chemical and Miller, N. 1981. Bogs, bales, and BTU’s: A hydrological gradients in the Lost River primer on peat. Horticulture 59: 38-45. Peatland, northern Minnesota. Journal of Ecology 78(4): 1021-1048. NatureServe. 2005a. NatureServe Explorer: An online encyclopedia of life [web application]. Hammerson, G. 1994. Beaver (Castor Version 4.2. NatureServe, Arlington, Virginia. canadensis): Ecosystem alterations, Available: management, and monitoring. Natural Areas http://www.natureserve.org/explorer. Journal 14(1): 44-57. (Accessed: March 03, 2005.)

Heinselman, M.L. 1963. Forest sites, bog NatureServe. 2005b. International Ecological processes, and peatland types in the Glacial Classification Standard: Terrestrial Ecological Classifications. NatureServe Central

Grayling Community Assessment - 77 Databases. Arlington, VA. U.S.A. Data Siegel, D.I., and P.H. Glaser. 1987. Groundwater current as of January 13, 2005. flow in a bog-fen complex, Lost River Peatland, northern Minnesota. Journal of Osvald, H. 1935. A bog at Hartford, Michigan. Ecology 75(3): 743-754. Ecology 16(3): 520-528. Swain, A.M. 1973. A history of fire and Richardson, C.J., and P.E. Marshall. 1986. vegetation in northeastern Minnesota as Processes controlling movement, storage, and recorded in lake sediments. Quaternary export of phosphorous in a fen peatland. Research 3: 383-396. Ecological Monographs 56(4): 279-302. Swanson, D.K., and D.F. Grigal. 1989. Vegetation Riley, J.L. 1989. Southern Ontario bogs and fens indicators of organic soil properties in of the Canadian Shield. Wetlands: Inertia or Minnesota. Soil Science Society of America Momentum: 355-367. Journal 53: 491-495.

Schwintzer, C.R. 1978a. Nutrient and water levels Swinehart, A.L., and G.R. Parker. 2000. in a small Michigan bog with high tree Palaeoecology and development of peatlands mortality. American Midland Naturalist in Indiana. American Midland Naturalist 100(2): 441-451. 143(2): 267-297.

Schwintzer, C.R. 1978b. Vegetation and nutrient Verry, E.S. 1975. Streamflow chemistry and status of northern Michigan fens. Canadian nutrient yields from upland-peatland Journal of Botany 56: 3044-3051. watersheds in Minnesota. Ecology 65(5): 1149-1157. Schwintzer, C.R. 1979. Vegetation changes following a water level rise and tree mortality Vitt, D.H., and N.G. Slack. 1975. An analysis of in a Michigan bog. The Michigan Botanist 18: the vegetation of Sphagnum-dominated kettle- 91-98. hole bogs in relation to environmental gradients. Canadian Journal of Botany 53: Schwintzer, C.R. 1981. Vegetation and nutrient 332-359. status of northern Michigan bogs and conifer swamps with a comparison to fens. Canadian Vitt, D.H., H. Crum, and J.A. Snider. 1975. The Journal of Botany 59: 842-853. vertical zonation of Sphagnum species in hummock-hollow complexes in northern Schwintzer, C.R, and G. Williams. 1974. Michigan. The Michigan Botanist 14(4): 190- Vegetation changes in a small Michigan bog 200. from 1917 to 1972. American Midland Naturalist 92(2): 447-459. Wheeler, G.A., P.H. Glaser, E. Gorham, C.M. Wetmore, F.D. Bowers, and J.A. Janssens. Schwintzer, C.R, and T.J. Tomberlin. 1982. 1983. Contributions to the flora of the Red Chemical and physical characteristics of Lake Peatland, northern Minnesota, with shallow ground waters in northern Michigan special attention to Carex. American Midland bogs, fens, and swamp. American Journal of Naturalist 110(1): 62-96. Botany 69(8): 1231-1239. Zoltai, S.C., and D.H. Vitt. 1995. Canadian Siegel, D.I. 1988. Evaluating cumulative effects wetlands: Environmental gradients and of disturbance on the hydrologic function of classification. Vegetatio 118: 131-137. bogs, fens, and mires. Environmental Management 12(5): 621-626.

Grayling Community Assessment - 78 Abstract Citation: Cohen, J.G. 2005. Natural community abstract for northern fen. Michigan Natural Features Inventory, Lansing, MI. 15 pp.

Grayling Community Assessment - 79 Grayling Community Assessment - 80 Appendix 4. Poor Fen

Overview: A sedge dominated wetland on very nutrient availability, increased alkalinity (less strongly to strongly acidic, saturated peat acidity), and greater species richness compared to moderately influenced by groundwater. Occurs bogs, with poor fens being most similar to bogs in north of the climatic tension zone in kettle terms of these factors and species composition depressions and on flat areas or mild depressions (Zoltai and Vitt 1995, Bedford and Godwin 2003). of glacial outwash and glacial lakeplain. Both poor fens and northern fens are dominated by sedges, rushes, and grasses, with poor fens Global and State Rank: G3G4/S3 containing a higher coverage of ericaceous shrubs and sphagnum mosses than northern fens (Mitsch Range: Poor fen is a peatland type of glaciated and Gosselink 2000). Like northern fen and poor landscapes of the northern Great Lakes region, fen, prairie fens are graminoid dominated and ranging from Michigan west to Minnesota and groundwater influenced, however prairie fens are northward into central Canada (Ontario, restricted to south of the climatic tension zone. Manitoba, and Quebec) (Gignac et al. 2000, Intermittent wetlands are herb or herb-shrub Faber-Langendoen 2001, Amon et al. 2002, dominated wetlands that experience fluctuating NatureServe 2005a). Poor fens may also occur in water levels seasonally and yearly and have soils parts of the northeastern U.S. (i.e., Maine, New that range from loamy sand and peaty sand to Hampshire, and New York) and range south into peaty muck and are very strongly acid to strongly northern Illinois and Iowa (NatureServe 2005a). acid. Like bogs, muskegs and poor conifer swamp In Michigan, poor fens occur in the northern are nutrient-poor, acidic wetlands. However, these Lower Peninsula and the Upper Peninsula (within ombrotrophic peatlands exhibit a greater degree of the U.P., poor fen occurrences have been canopy closure than bogs (muskegs having documented by Michigan Natural Feature clumped and scattered conifers and poor conifer Inventory in just the eastern portion). Fens and swamp being a closed canopy system). Closed other peatlands occur where excess moisture is canopy, minerotrophic peatlands include rich abundant (where precipitation is greater than conifer swamp, a Thuja occidentalis (northern evapotranspiration) (Mitsch and Gosselink 2000). white cedar) dominated system found north of the Conditions suitable for the development of fens tension zone, and relict conifer swamp, which is have occurred in the northern Lake States for the dominated by Larix laricina (tamarack) and past 8,000 years. Expansion of peatlands likely occurs primarily south of the tension zone (Kost occurred following climatic cooling, 2001). approximately 5,000 years ago (Heinselman 1970, Boelter and Verry 1977, Riley 1989). Rank Justification: Poor fens are uncommon features of the northern Great Lakes region, Several other natural peatland communities also occurring sporadically in Michigan’s northern occur in Michigan and can be distinguished from Lower Peninsula and the Upper Peninsula. The weakly minerotrophic poor fens, based on careful northern lake states contain over six million comparisons of nutrient levels, flora, canopy hectares (15 million acres) of peatland (Boelter closure, distribution, and groundwater influence. and Verry 1977). What percentage of that area is Additional open wetlands occurring on peat poor fen has yet to be determined. Likewise, the include bog, northern fen, and prairie fen. Bogs, current status of fens relative to their historical peat-covered wetlands raised above the status is unknown (Bedford and Godwin 2003). surrounding groundwater by an accumulation of Peatland scientists concur that fens have always peat, receive inputs of nutrients and water been localized and not very abundant but have primarily from precipitation and are classified as suffered from extensive loss, fragmentation, and ombrotrophic (nutrient-poor) (Gignac et al. 2000). degradation (Bedford and Godwin 2003, In comparison, the hydrology of fens is influenced NatureServe 2005a). Historically, widespread by groundwater and as a result, fens have higher fires following the turn of the century logging

Grayling Community Assessment - 81 drastically altered many peatlands, either and dominance by aggressive, common natives converting conifer swamp to open fen systems or such as Phalaris arundinaceae (reed canary grass) destroying the peat and converting peatlands to and Typha spp. (cat-tails) (Richardson and wetlands without organic soils (mineral soil Marshall 1986, Almendinger and Leete 1998). wetlands) (Dean and Coburn 1927, Gates 1942, Bedford et al. (1999) have noted a widespread Curtis 1959). Logging of cedar and tamarack from decline in wetland species richness associated peatland systems also favored the conversion of with the overall eutrophication of the landscape: forested peatlands to open peatlands (Gates 1942, nutrient enrichment has converted numerous Dansereau and Segadas-Vianna 1952, Riley wetlands into monospecific stands of nitrophilic 1989). Beginning in the 1920s, effective fire species. Lowering of water tables from drainage control by the U.S. Forest Service and state has allowed for tree and shrub encroachment into agencies reduced the acreage of fires ignited by open fens and the eventual succession to closed man or lightning (Swain 1973). In landscapes canopy peatland (Almendinger and Leete 1998). where frequent fire was the prevalent disturbance Increased shrub and tree canopy cover typically factor, fire suppression has led to the conversion results in decreased species richness of fen of open fens to closed canopy peatlands or shrub systems (Bowles et al. 1996). The sensitivity of thickets (Curtis 1959, Schwintzer 1981, Riley fens to changes in water chemistry makes them 1989). especially susceptible to acid rain and air pollution (Siegel 1988, Chapman et al. 2003). Currently, fens are primarily threatened by peat Atmospheric deposition can contribute Nitrogen, mining, logging, quarrying, agricultural runoff Sulphur, Calcium and heavy metals to fens and enrichment, off-road-vehicle (ORV) activity, (Damman 1990, Chapman et al. 2003). Dust-fall draining, flooding, and development (Bedford and and atmospheric deposition from air pollution are Godwin 2003, NatureServe 2005a). Peat mining particularly threats to fen systems that are and cranberry farming have degraded numerous surrounded by cultivated land and close to peatlands throughout the region (Gates 1942, industrial and urban centers (Damman 1990). Curtis 1959, Eggers and Reed 1997, Chapman et al. 2003). Michigan, along with Florida and Physiographic Context: Two landscape features Minnesota, are leaders in peat production in the are conducive to the development of peat; poorly- U.S. (Miller 1981). In addition to direct impacts to drained, level terrain and small ice-block basins vegetation, alteration of peatland hydrology from (Boelter and Verry 1977). Poor fen occur on flat road building, ORVs, quarrying, creation of areas or mild depressions of sandy glacial drainage ditches and dams, and runoff from outwash and glacial lakeplains, and in kettle logging has led to the drastic change of peatland depressions on pitted outwash and moraines composition and structure (Schwintzer and (Gates 1942, Verry 1975, Vitt and Slack 1975, Williams 1974, Schwintzer 1978b, Riley 1989, Boelter and Verry 1977, Schwintzer 1978a, Siegel Bedford and Godwin 2003, Chapman et al. 2003). 1988, Michigan Natural Features Inventory 2003, Fen vegetation is extremely sensitive to minor NatureServe 2005a). The overall topography of changes in water levels and chemistry, ground fens is flat to gently undulating with water flow, and nutrient availability (Siegel 1988, microtopography characterized by hummocks and Riley 1989). A reduction in groundwater flow and hollows (Heinselman 1963, Vitt and Slack 1975, subsequent decrease in nutrients in poor fens can Wheeler et al. 1983, Siegel 1988, NatureServe result in the shift to less minerotrophic wetlands 2005a). Poor fens found in kettle depressions are such as bog. Conversion to more eutrophic associated with active or extinct glacial lakes that wetlands has occurred as the result of nutrient are very strongly acidic to strongly acidic (Vitt enrichment and raised water levels, which cause and Slack 1975). Within kettle depressions, fens increased decomposition of peats. Eutrophication can occupy the entire basin or frequently occur as from pollution and altered hydrology has a floating mat on the margin of the remaining detrimentally impacted fens by generating glacial lake above the level of seasonal flooding conditions favorable for the invasion of exotic (Vitt and Slack 1975, Schwintzer 1978a, species (Riley 1989, Bedford and Godwin 2003) NatureServe 2005a). When fens occur along the

Grayling Community Assessment - 82 edge of large bodies of water, they are found in mineralized or poorly buffered ground water sheltered bays or coves that are protected from discharge of poor fens typically contains low wave and ice action, which can prevent the concentrations of dissolved mineral nutrients development of peat or erode existing peat mats (Heinselman 1970, Boelter and Verry 1977, (Gates 1942, NatureServe 2005a). Fens occurring Glaser et al. 1981, Glaser et al. 1990, Bedford and on former glacial lake beds and drainageways Godwin 2003, NatureServe 2005a). The acidity of tend to be more extensive than kettle fens, which poor fens limits the availability and uptake of are limited in area by the size of the glacial ice- essential mineral plant nutrients. Poor fens are block which formed the basin (Lindeman 1941). characterized by low concentrations of available Calcium, Magnesium, Nitrogen, and Phosphorous Poor fens occur adjacent to other peatland in the surface water and peat (Heinselman 1970, communities, often grading into bog, poor conifer Glaser et al. 1981, Schwintzer 1981, Glaser et al. swamp, and muskeg. More minerotrophic systems 1990, Mitsch and Gosselink 2000). Scientists such as northern fen, northern shrub thicket, studying poor fens in the Great Lakes have northern wet meadow, and rich conifer swamp can reported pH values to range between 4.1-5.9 occur along the outer margins of poor fens where (Heinselman 1970, Boelter and Verry 1977, groundwater seepage from the adjacent uplands is Glaser et al. 1981, Glaser et al. 1990, prevalent. Upland community types which Almendinger and Leete 1998, Bedford et al. 1999, neighbor poor fen include dry-mesic northern NatureServe 2005a). The degree of minerotrophy forest, dry northern forest, and pine barrens. of a given fen and within a fen depends on the kind and amount of groundwater discharge, Hydrology: Poor fens are weakly minerotrophic degree of dilution from precipitation, the peatlands, receiving inputs of water and nutrients characteristics of the bedrock and/or glacial from both ion-poor precipitation and low exposure deposits the groundwater has percolated through to nutrient-rich groundwater (Heinselman 1970, (i.e., older glacial sediments have less dissolved Boelter and Verry 1977, Siegel and Glaser 1987, minerals due to prior leaching), the distance the Siegel 1988, Bedford et al. 1999, Gignac et al. water has traveled through the peatland, and the 2000, Bedford and Godwin 2003, NatureServe thickness and character of the peat (Heinselman 2005a). Low-levels of groundwater discharge and 1963, Heinselman 1970, Boelter and Verry 1977, the high water retaining capacity of fibric peat Siegel and Glaser 1987, Amon et al. 2002, produce continuously saturated conditions in the Bedford and Godwin 2003). The ground water rooting zone of poor fens. The water table of poor entering poor fens typically passes through fens is stable, typically at the soil surface with the materials with low solubility or low buffering peat soils saturated but seldom flooded capacity (Bedford and Godwin 2003). Poor fens (Heinselman 1970, Schwintzer 1978b, Riley 1989, often occur in basins with small watersheds which Amon et al. 2002, Bedford and Godwin 2003, minimize the groundwater input from the NatureServe 2005a). The surface waters of poor surrounding uplands (NatureServe 2005a). It is fens are characterized by very strong to strong likely that many poor fens are restricted to areas acidity, low available nutrients, low specific where the bedrock is buried under thick glacial conductivity, cool temperatures, anaerobic deposits. conditions, and moderate levels of dissolved organic matter (Verry 1975, Boelter and Verry Soils: The organic soils of poor fens are 1977, Schwintzer 1978b, Glaser et al. 1981, composed of peat which frequently forms a Glaser et al. 1990, Bedford et al. 1999, Bedford shallow, continuous mat and is typically 1-3 and Godwin 2003, NatureServe 2005a). The meters deep (Glaser et al. 1981). Peat is a fibrous limited amount of ground water which enters poor network of partially decomposed organic material fens is telluric, having moved over or percolated that is formed under anaerobic conditions through base-rich bedrock, calcareous glacial (Almendinger et al. 1986, Heinselman 1963). The deposits, or mineral soil (Heinselman 1970, surface peats of poor fens are saturated, fibric Schwintzer 1978b, Mitsch and Gosselink 2000, peats, and like the surface waters, are very Bedford and Godwin 2003). The poorly strongly to strongly acidic and characterized by

Grayling Community Assessment - 83 low nutrient availability (NatureServe 2005a). °C (45 °F) (Albert et al. 1986, Barnes 1991). Fibric peat, which is loosely compacted, contains Temperatures vary less in peatlands compared to partially decomposed sphagnum moss with the surrounding landscape because of fragments of wood and occasionally sedge. Fibric groundwater influence, the insulating effect of peat has high water retaining capacity and large fens’ saturated peat carpet in the growing season, intercellular pores that permit rapid water and snow cover in the winter (Burns 1906, movement (Boelter and Verry 1977, Miller 1981, Heinselman 1963, Curtis 1959, Glaser 1992). Swanson and Grigal 1989, Amon et al. 2002). Fens are characterized by local climates or Peats of fens tend to have lower water retaining microclimates that are cooler in the summer and capacity and higher levels of organic warmer in the winter compared to the regional decomposition compared to the peats of bogs climate (Heinselman 1963, Bedford and Godwin (Boelter and Verry 1977, Miller 1981). Peats of 2003). poor fens tend to be less decomposed with higher water retaining capacity compared to northern fen Natural Processes: Peat establishment requires peats. Peat composition changes with depth and an abundant supply of water: peatlands occur in depending on the successional history of a given regions where precipitation is greater than fen. Generally, fiber content and hydraulic evapotranspiration producing substantial conductivity usually decreases with depth; deeper groundwater discharge (Dansereau and Segadas- peats are more decomposed, retain more water, Vianna 1952, Boelter and Verry 1977, and drain slower than surface peats (Verry 1975, Almendinger and Leete 1998, Mitsch and Boelter and Verry 1977). Gosselink 2000). Saturated and inundated conditions inhibit organic matter decomposition Climate: Peatlands develop in humid climates and allow for the accumulation of peat where precipitation exceeds evapotranspiration (Almendinger and Leete 1998, Amon et al. 2002). (Boelter and Verry 1977, Gignac et al. 2000, Under cool and anaerobic conditions, the rate of Bedford and Godwin 2003). The northern Lake organic matter accumulation exceeds organic States are characterized by a humid, continental decay (Schwintzer and Williams 1974, Damman climate with long cold winters and short summers 1990, Mitsch and Gosselink 2000). Low levels of that are moist and cool to warm (Gates 1942, oxygen protect plants from microorganisms and Boelter and Verry 1977, Damman 1990, Mitsch chemical actions that cause decay (Miller 1981). and Gosselink 2000). The Michigan range of poor Fens have greater levels of microbial activity fen falls within the area classified by Braun compared to bogs because of the lesser acidity and (1950) as the Northern Hardwood-Conifer Region higher base status of minerotrophic waters. As a (Hemlock/White Pine/Northern Hardwoods result, organic matter decay is greater while peat Region) and within the following regions accumulation is lesser in fens versus bogs classified by Albert et al. (1986) and Albert (Heinselman 1970). Development and expansion (1995): Region II, Northern Lower Michigan; of fens occurs via two different processes in kettle Region III, Eastern Upper Michigan; and Region depressions versus glacial lakeplain and outwash. IV, Western Upper Michigan. The Northern Fens develop in glacial lakeplain and outwash Hardwood-Conifer Region has a cool snow-forest where groundwater influence maintains saturated climate with short, warm summers, cold winters conditions which inhibit organic matter and a large number of cloudy days. The mean decomposition and allow peat accumulation number of freeze-free days is between 90 and 160, (Almendinger and Leete 1998). Peat develops and the average number of days per year with vertically and spreads horizontally (Boelter and snow cover of 2.5 cm or more is between 80 and Verry 1977). Estimates of vertical accumulation 140. The normal annual total precipitation ranges of peat range between 100 to 200cm/1000 years from 740 to 900 mm with a mean of 823 mm. The (Mitsch and Gosselink 2000). Lake-filling or daily maximum temperature in July ranges from terrestrialization occurs in small kettle lakes with 24 to 29 °C (75 to 85 °F), the daily minimum minimal wave action where gradual peat temperature in January ranges from -21 to -9 °C (- accumulation results in the development of a 5 to 15 °F) and the mean annual temperature is 7 sedge mat that can fill the basin or occur as a

Grayling Community Assessment - 84 floating mat in the lake or as a grounded mat table of fens results in the increase in along the water’s edge (Gates 1942, Bay 1967, decomposition rates of organic matter and the Curtis 1959, Heinselman 1963, Mitsch and subsequent accumulation of compact peat that is Gosselink 2000, Swinehart and Parker 2000). more conducive to shrub and tree growth Floating mats of fen sedges (i.e., Carex (Schwintzer and Williams 1974, Miller 1981, lasiocarpa) pioneer open water or emergent Schwintzer 1981, Riley 1989, Almendinger and marsh. The interlacing of rhizomes and roots Leete 1998, Gignac et al. 2000). Conversions of forms a floating mat that is buoyed up in water bog to fen can also occur, however with far less and accumulates organic matter in the form of frequency (Glaser et al. 1990). A discharge of sapric peat (Gates 1942). Over time fen mats are alkaline groundwater at the peat surface of a bog, often invaded by ericaceous shrubs and acidifying caused by a change in hydraulic head, can result sphagnum mosses (Osvald 1935, Gates 1942, in the conversion of bog vegetation to fen Schwintzer and Williams 1974, Swineheart and vegetation (Siegel and Glaser 1987, Glaser et al. Parker 2000). 1990). Mixing of as little as 10% groundwater from underlying calcareous parent material with The invasion of sphagnum moss into fen systems acid bog water is sufficient to raise the peatland often results in the conversion of fens to more pH from 3.6 to 6.8 (Glaser et al. 1990). Fens and acidic communities such as poor fen or bog. bogs are very sensitive to changes in pH and Succession in lake-filled fens typically proceeds subsequent availability of nutrients: fen vegetation from lake to marsh to fen to poor fen or bog can replace bog flora when pH increases above (Heinselman 1963, Boelter and Verry 1977, 4.5 (Siegel 1988). Schwintzer 1981, Swineheart and Parker 2000). Once Sphagnum mosses become established on Disturbance factors influencing poor fens include fen peat, they maintain and enhance saturated and fire, flooding, windthrow, and insects. Numerous acidic conditions which in turn promote continued fens contain charcoal within their peat profile sphagnum peat development (Heinselman 1963). (Curtis 1959, Heinselman 1963) and many The ability of sphagnum to absorb and hold researchers have reported fire as a prevalent part cations increases the acidity and low nutrient of fen’s disturbance regime (Gates 1942, Curtis availability of peatlands (Osvald 1935, Curtis 1959, Vitt and Slack 1975). Surface fire can 1959, Verry 1975, Vitt and Slack 1975, Boelter contribute to the maintenance of fens by killing and Verry 1977, Zoltai and Vitt 1995). In encroaching trees and shrubs without completely addition, accumulating sphagnum peat can dilute removing the peat, which is normally saturated groundwater influence by absorbing large (Curtis 1959, Vitt and Slack 1975). Graminoid amounts of precipitated water, impeding drainage, dominance of fen systems can be perpetuated by and increasing the distance of the rooting zone surface fires (Bowles et al. 1996). In addition, from telluric water (Dansereau and Segadas- many of the ericaceous plants that thrive in fens Vianna 1952, Vitt and Slack 1975, Schwintzer are fire adapted and often grow densely following 1981, Zoltai and Vitt 1995). Sphagnum moss, fire (Wheeler et al. 1983). Fire severity and which has numerous pores, partitions, and frequency in fens is closely related to fluctuations capillary space, has an enormous water-holding in water level. Prolonged periods of lowered water capacity (Osvald 1935, Dansereau and Segadas- table can allow the surface peat to dry out enough Vianna 1952, Curtis 1959): sphagnum peat can to burn (Schwintzer and Williams 1974). When hold 15 to 30 times its own weight in water the surface peat of fens and bogs burns, the fire (Miller 1981, Mitsch and Gosselink 2000). In releases organic matter from the peat, kills seeds addition to sphagnum peat accumulation, beaver and latent buds, stimulates decay, and slows peat dams can also cause blocked drainage in fens and accumulation (Damman 1990, Jean and Bouchard the subsequent succession of fens to bogs 1991). Such peat fires can result in the conversion (Heinselman 1963, Heinselman 1970). of peatland to mineral soil wetland. Peat fires within bogs can also release enough nutrients to Poor fens can also succeed to poor conifer swamp favor succession to more minerotrophic peatlands or northern shrub thicket. Lowering of the water such as poor fen or intermittent wetland.

Grayling Community Assessment - 85 As noted above, flooding can result in the Godwin 2003, NatureServe 2005a). The harsh conversion of fens to bogs. Flooding can also growing conditions of poor fens (strong acidity, contribute to fen maintenance. Roots of peatland low nutrient availability, and saturated peat) trees are physiologically active near the surface results in a distinct flora of low to moderate and are quickly killed when the water table rises diversity: relatively few species have evolved the following flooding (Glaser and Janssens 1986). necessary adaptations to cope with ombrotrophic Within kettle fens, flooding-induced tree mortality and weakly minerotrophic conditions (Siegel is likely greater on grounded mats compared to 1988, Glaser 1992, Mitsch and Gosselink 2000). free floating mats: free mats float up with rising Very few introduced, weedy species are able to water table while grounded mats become establish within bogs and fens because of the inundated and have shallower aerobic zones unique growing conditions and competition from (Schwintzer 1978a, Schwintzer 1978b, Schwintzer the adapted flora (Riley 1989). Poor fen plants 1979). In addition to flooding, kettle fens can be have developed a diversity of adaptations to cope influenced by waves and ice. Wave and ice action with low nutrient availability including plant can prevent the expansion of fen mats by eroding carnivory, evergreen leaves, sclerophylly (thick the shoreline vegetation (Gates 1942). epidermal tissue), and high root biomass (Mitsch and Gosselink 2000). Poor fens are dominated by The natural disturbance regime in fens is also plants that thrive under moderately ombrotrophic influenced by wind. The Great Lakes region is to weakly minerotrophic conditions. Occasionally one of the most active weather zones in the minerotrophic indicators may be present in poor northern hemisphere, with polar jet streams fen at low cover. Plants found typically in more positioned overhead much of the year. More alkaline habitat such as Betula pumila (bog birch), cyclones pass over this area than any other area in Carex aquatilis (water sedge), Carex rostrata the continental U.S. (Frelich and Lorimer 1991). (beaked sedge), and Carex stricta (tussock sedge), Trees growing in fens are particularly susceptible can occur sporadically in poor fen when their to windthrow because peat provides a poor roots extend beneath the surface mat to substrate for anchoring trees (Burns 1906). The minerotrophic peat influenced by groundwater living roots of woody peatland plants occur in a (NatureServe 2005a). The tops of hummocks shallow rooting zone, generally restricted to the support sphagnum mosses and a more acidic uppermost few centimeters where there is micro-environment within poor fens where sufficient oxygen to maintain aerobic respiration ombrotrophic species can occur isolated from the (Glaser and Janssens 1986). The superficial influence of ground water. Species richness of rooting of trees results in numerous windthrows poor fens is related to geographical location, (Dansereau and Segadas-Vianna 1952). Tree climatic factors, nutrient availability, and habitat survival in fens is also limited by insects and heterogeneity (Glaser et al. 1990, Glaser 1992). parasites. Insect outbreaks of the Pristiphora Floristic diversity within poor fens is strongly erichsonii (larch sawfly) cause heavy mortality of correlated with levels of available nutrients and Larix laricina (tamarack) while, the plant parasite microtopography (Riley 1989, Glaser et al. 1990). Arceuthobium pusillum (dwarf mistletoe) kills Small scale environmental heterogeneity can Picea mariana (black spruce) (Coburn et al. 1933, result in vegetational zonation (Amon et al. 2002, Gates 1942, Heinselman 1963). Bedford and Godwin 2003). Gradients in pH, light, soil moisture, and cation concentrations Vegetation Description: Poor fens have a unique (nutrient availability) determine floristic flora that is intermediate between bog and composition of poor fens (Heinselman 1970, Vitt northern fen. Poor fens are characterized by a and Slack 1975, Schwintzer 1978a, Glaser et al. continuous carpet of sphagnum mosses, a 1981, Anderson et al. 1986, Siegel 1988, Glaser et graminoid-dominated herbaceous layer of low to al. 1990, Bedford et al. 1999). moderate diversity, low ericaceous, evergreen shrubs, and widely-scattered and stunted conifer The continuous moss layer of poor fens is trees (Gates 1942, Curtis 1959, Verry 1975, Vitt dominated by sphagnum mosses especially and Slack 1975, Glaser et al. 1991, Bedford and Sphagnum magellanicum, S. angustifolium, S.

Grayling Community Assessment - 86 capillaceum, S. capillifolium, S. recurvum, S. graminifolia (flat-topped goldenrod), Iris papillosum, and S. fuscum (Schwintzer 1978b, versicolor (wild blue flag), Menyanthes trifoliata Crum 1983, Riley 1989, NatureServe 2005a). In (bog bean), Smilacina trifolia (false Solomon- comparison, the moss layer of northern fens is seal), Solidago uliginosa (bog goldenrod), and patchy to continuous and dominated by Triglochin maritimum (arrow-grass). calcicolous brown mosses of the family Insectivorous plants, Drosera rotundifolia Amblystegiaceae (Glaser et al. 1990, Zoltai and (roundleaf sundew), Drosera intermedia (spoon- Vitt 1995, Swinehart and Parker 2000, Amon et leaf sundew), Sarracenia purpurea (pitcher- al. 2002). The hummock and hollow plant), and Utricularia intermedia (flat-leaved microtopography of poor fens allows for high bladderwort), are common features of fens. The levels of bryophyte diversity since individual fern ally Equisetum fluviatile (water horsetail) is species of sphagnum occur at specific elevations often found in poor fens (Vitt and Slack 1975, Wheeler et al. 1983, Riley 1989,). Hollows support S. cuspidatum, S. The shrub layer of poor fens is dominated by low, magellanicum, and S. papillosum (Vitt and Slack ericaceous shrubs with Chamadaephne calyculata 1975, Vitt et al. 1975, Heinselman 1970, Wheeler (leatherleaf) often being the most prevalent. In et al. 1983, Riley 1989). The lower, moist slopes addition to leatherleaf, the following heath shrubs of hummocks often support S. magellanicum and are important: Andromeda glaucophylla (bog- S. recurvum while the drier hummock crests are rosemary), Kalmia polifolia (bog-laurel), Ledum dominated by S. fuscum, S. capillaceum, and S. groenlandicum (Labrador-tea), Vaccinium cappillifolium (Vitt et al. 1975, Wheeler et al. macrocarpon (large cranberry), and V. oxycoccos 1983, Riley 1989). The vertical zonation of (small cranberry). The tall shrub layer of poor species corresponds to gradients in pH and fens is less dense than the low shrub layer and is moisture with the hollows being wetter and more often restricted to the periphery. Tall shrubs alkaline than the drier and more acidic tops of the typical of poor fens include Salix discolor (pussy hummocks (Vitt et al. 1975, Wheeler et al. 1983). willow), S. pedicellaris (bog willow), and Spirea tomentosa (steeplebush). As noted, more Cyperaceous graminoids dominate the species minerotrophic shrubs, like Betula pumila (bog poor herbaceous layer of poor fens. Carex birch), Hypericum kalmianum (Kalm’s St. John’s- oligosperma (few-seed sedge) and Carex wort), Potentilla fruticosa (shrubby cinquefoil), lasiocarpa (wiregrass sedge) are common and P. palustris (marsh cinquefoil), can occur in dominants. Other sedges that are characteristic of poor fens when their roots can extend beneath the poor fens include Carex chordorrhiza (creeping surface mat to minerotrophic peat. sedge), C. exilis (coastal sedge), C. livida (livid sedge), C. pauciflora (few-flower sedge), and C. Trees within poor fens are widely scattered and limosa (mud sedge). Other sedges that often occur stunted (seldom reaching six meters) (Wheeler et in bogs are Carex paupercula (bog or poor sedge), al. 1983, NatureServe 2005a). Tree cover is C. rostrata (beaked sedge), and C. trisperma typically below ten percent (NatureServe 2005a). (three-seeded sedge). Additional graminoids that The most common canopy dominants are Picea thrive in poor fens include Cladium marisicoides mariana (black spruce) and Larix laricina (twig-rush), Dulichium arundinaceum (three-way (tamarack). Additional associates include Pinus sedge), Eriophorum angustifolium (tall cotton- banksiana (jack pine) and Pinus strobus (white grass), Eriophorum spissum (sheathed cotton- pine). (Above species lists compiled from grass), Eriophorum virginicum (tawny cotton- Michigan Natural Features Inventory database, grass), Lysimachia terrestris (swamp candles), Gates 1942, Dansereau and Segadas-Vianna 1952, Rhynchospora alba (white beak-rush), Curtis 1959, Heinselman 1963, Heinselman 1965, Scheuchzeria palustris (rannoch-rush), and Bay 1967, Heinselman 1970, Schwintzer and Scirpus cespitosus (tufted bulrush). The following Williams 1974, Vitt and Slack 1975, Schwintzer is a list of prevalent bog herbs: Aster borealis 1978, Glaser et al. 1981, Schwintzer 1981, (rush aster), Epilobium angustifolium (fireweed), Wheeler et al. 1983, Riley 1989, Glaser et al. E. ciliatum (fringed willow-herb), Euthamia 1990, Glaser 1992, Eggers and Reed 1997, Mitsch

Grayling Community Assessment - 87 and Gosselink 2000, Swinehart and Parker 2000, (yellow rail, state threatened), Falcipennis NatureServe 2005a.) canadensis (spruce grouse, state special concern), and Picoides arcticus (black-backed woodpecker, Michigan Indicator Species: bog birch, Carex state special concern). Alces alces (moose, state chordorrhiza, C. lasiocarpa, C. limosa, C. livida, threatened), Canis lupus (gray wolf, state northern white cedar, shrubby cinquefoil, and threatened), and Lynx canadensis (lynx, state tamarack (Heinselman 1970, Wheeler et al. 1983, endangered) utilize peatland habitat. Poor fens Anderson et al. 1996). Wheeler et al. (1983), in a provide important habitat for small mammals such study of peatland flora in northern Minnesota, as Blarina brevicauda (short-tailed shrew), Castor concluded that Carex livida is a poor fen canadensis (beaver), Microtus pennsylvanicus indicator; the presence of this species readily (meadow vole), Mustela vison (mink), Ondatra distinguishes sites as weakly minerotrophic. zibethicus (muskrat), and Sorex cinereus (masked shrew). Both muskrats and beaver can profoundly Other Noteworthy Species: Poor fens provide influence the hydrology of peatlands. Muskrats habitat for numerous rare insect species including create open water channels through the peat and Appalachia arcana (secretive locust, state special beavers can cause substantial flooding through concern), Atlanticus davisi (Davis's shield-bearer, their dam-building activities (Gates 1942, state special concern), Boloria freija (Freija Heinselman 1963). fritillary, state special concern butterfly), Boloria frigga (Frigga fritillary, state special concern Rare plants associated with poor fens include butterfly), Erebia discoidalis (red-disked alpine, Carex nigra (black sedge, state endangered), state special concern butterfly), Merolonche dollii Carex wiegandii (Wiegand's sedge, state (Doll’s merolonche moth, state special concern), threatened), Eleocharis nitida (slender spike-rush, Phyciodes batesii (tawny crescent, state special state endangered), and Petasites sagittatus (sweet concern), Somatochlora incurvata (incurvate coltsfoot state threatened). emerald, state special concern dragonfly), and Williamsoni fletcheri (ebony boghaunter, state Conservation and Biodiversity Management: special concern dragonfly). Numerous butterflies Poor fen is a widespread community type in the and moths are restricted to bogs and fens because Great Lakes region that contributes significantly their food plants occur within these peatland to the overall biodiversity of northern Michigan systems (Riley 1989). Rare herptiles that utilize by providing habitat for a unique suite of plants poor fens include Clemmys guttata (spotted turtle, and wide variety of animal species. Numerous state threatened), Elaphe obsoleta obsolete (black rare species are associated with poor fens. By rat snake, state special concern), Emys blandingii storing high levels of sequestered Carbon and (Blanding’s turtle, state special concern), serving as Carbon sinks, poor fens and related Sistrurus catenatus catenatus (eastern peatlands play an important role in global massasauga, state special concern), and Terrapene geochemical cycles. Poor fens also preserve carolina carolina (eastern box turtle, state special paleo-environmental records: a wealth of concern). If suitable nesting trees or snags are information is stored in the remains of plants, available, Falco columbarius (merlin, state animals, and atmospheric particles deposited and threatened), Haliaeetus leucocephalus (bald eagle, stored in fen peat profiles. Paleo-ecologists may state threatened), and Pandion haliaetus (osprey, be able to provide crucial information about state threatened) can be found nesting in these restoration of peatland ecosystems (Chapman et systems and Ardea herodias (great blue heron, al. 2003). The primary mechanism for preserving protected by the Migratory Bird Treaty Act of poor fens is to maintain their hydrology. As noted, 1918) can establish rookeries. Other rare birds that peatland systems are sensitive to slight changes in could occur in poor fens are Asio flammeus (short- water chemistry. A serious threat to poor fens is eared owl, state endangered), Botaurus posed by off-road-vehicle traffic, which can lentiginosus (American bittern, state special destroy populations of sensitive species and concern), Circus cyaneus (northern harrier, state drastically alter fen hydrology through rutting. special concern), Coturnicops noveboracensis Reduction of access to peatland systems will help

Grayling Community Assessment - 88 decrease detrimental impacts. Resource mangers direct and indirect anthropogenic disturbance on operating in uplands adjacent to poor fens should peatlands (Amon et al. 2002). Effects of take care to minimize the impacts of management management within fens should be monitored to to hydrologic regimes, especially increased allow for assessment and refinement. Monitoring surface flow and alteration of groundwater should also focus on how fen succession and discharge. This can be accomplished by management influence populations of rare species. establishing a no-cut buffer around poor fens and Scientific understanding of the microbes and avoiding road construction and complete canopy invertebrates that thrive in the organic soils of removal in stands immediately adjacent to poor fens is lacking (Amon et al. 2002). Given the fens. Where shrub/tree encroachment threatens to sensitivity of peatlands to slight changes in convert open wetlands to shrub-dominated hydrology and nutrient availability, it is important systems or forested swamps, prescribed fire or for scientists to predict how peatlands will be selective cutting can be employed to maintain affected by climate change and atmospheric open conditions. Silvicutural management of poor deposition of nutrients and acidifying agents fens to preserve open canopy should be employed (Heinselman 1970, Riley 1989, Bedford et al. during the winter to minimize damage to the peat 1999, Gignac et al. 2000, Mitsch and Gosselink and impacts to the hydrologic regime. 2000, Bedford and Godwin 2003). Peat deposits are of great scientific interest because they contain Research Needs: Poor fen has a broad historical ecological records in the form of fossils distribution and exhibits numerous regional, of plants, animals, and organic matter that physiographic, hydrologic, and edaphic variants. contributed to the deposit. Stratigraphical analysis The diversity of variations throughout its range of peat cores provides insights into past climatic demands the continual refinement of regional change and associated vegetation change, floristic classifications that focus on the inter-relationships distribution, the development of wetland between vegetation, physiography, and hydrology ecosystems, and the successional pathways of (Heinselman 1963, Fitzgerald and Bailey 1975, peatlands (Heinselman 1963, Glaser et al. 1981, Barnes et al. 1982, Amon et al. 2002). Poor fens Miller 1981, Glaser and Janssens 1986, Riley and related community types (bog, northern fen, 1989, Gignac et al. 2000). and intermittent wetland) are frequently difficult to differentiate (Heinselman 1963, NatureServe Similar Communities: bog, intermittent wetland, 2005a). Research on abiotic and biotic indicators muskeg, northern fen, patterned fen, poor conifer that help distinguish related peatlands would be swamp, prairie fen, rich conifer swamp. useful for field classification. Systematic surveys for poor fens and related peatlands are needed to Other Classifications: help prioritize conservation and management Michigan Natural Features Inventory Circa 1800 efforts. More research is needed to elucidate the Vegetation (MNFI): Emergent Marsh (6221), Wet relationship of chemical factors and nutrients to Meadow (6224), and Inland Wet Prairie (6227) floristic community structure of peatlands (Amon et al. 2002). Little is known about the fire regimes Michigan Department of Natural Resources of poor fens and the interaction of disturbance (MDNR): D-treed bog, V-bog, N-marsh factors within these systems. As noted by Hammerson (1994), beaver significantly alter the Michigan Resource Information Systems ecosystems they occupy. An important research (MIRIS): 62 (non-forested wetland) and 622 question to examine is how the wetland (emergent wetland). ecosystems of the Great Lakes have been and continue to be affected by fluctuations in The Nature Conservancy National Classification: populations of beaver. Experimentation is needed CODE; ALLIANCE; ASSOCIATION; to determine how best to prevent shrub and tree COMMON NAME encroachment of fens that are threatened by conversion to shrub thicket or conifer swamp. A III.B.2.N.g; Betula pumila – (Salix spp.) better understanding is needed of the influence of Saturated Shrubland Alliance; Alnus incana –

Grayling Community Assessment - 89 Salix spp. - Betula pumila / Chamaedaphne (Swollen-beak Sedge) – Water Horsetail calyculata Shrubland; Speckled Alder – Herbaceous Vegetation; Wiregrass Sedge Willow Species – Bog Birch / Leatherleaf Shore Fen Shrubland; Bog Birch-Willow Shore Fen Related Abstracts: American bittern, black- III.B.2.N.g; Betula pumila – (Salix spp.) backed woodpecker, Blanding’s turtle, eastern Saturated Shrubland Alliance; Betula pumila / box turtle, eastern massasauga, great blue heron Chamaedaphne calyculata / Carex lasiocarpa rookery, incurvate emerald, intermittent wetland, Shrubland; Bog Birch / Leatherleaf / merlin, northern fen, northern harrier, poor fen, Wiregrass Sedge Shrubland; Bog Birch – prairie fen, rich conifer swamp, secretive locust, Leatherleaf Rich Fen spotted turtle, sweet coltsfoot, and yellow rail.

III.B.2.N.g; Betula pumila – (Salix spp.) Literature Cited: Saturated Shrubland Alliance; Betula pumila Albert, D.A. 1995. Regional landscape – Dasiphora fruticosa spp. floribunda / Carex ecosystems of Michigan, Minnesota, and lasiocarpa –Trichophorum alpinum Wisconsin: A working map and classification. Shrubland; Bog Birch – Shrubby-cinquefoil / Gen. Tech. Rep. NC-178. St. Paul, MN: U.S. Wiregrass Sedge – Alpine Cottongrass Department of Agriculture, Forest Service, Shrubland; Bog Birch – Shrubby-cinquefoil North Central Forest Experiment Station. Rich Boreal Fen Northern Prairie Wildlife Research Center Home Page. IV.A.1.N.g; Chamaedaphne calyculata http://www.npwrc.usgs.gov/resource/1998/rla Saturated Dwarf-shrubland Alliance; ndscp/rlandscp.htm. (Version 03JUN98.) 250 Chamaedaphne calyculata – Myrica gale / pp. Carex lasiocarpa Dwarf-shrubland; Leatherleaf – Sweet Gale / Wiregrass Sedge Albert, D.A., S.R. Denton, and B.V. Barnes. 1986. Dwarf-shrubland; Leatherleaf – Sweet Gale Regional landscape ecosystems of Michigan. Shore Fen Ann Arbor, MI: University of Michigan, School of Natural Resources. 32 pp. & map. V.A.5.N.m; Calamagrostis canadensis – Carex viridula – Cladium marisicoides – Almendinger, J.C., J.E. Almendinger, and P.H. Lobelia kalmii Saturated Herbaceous Glaser. 1986. Topographic fluctuations across Alliance; Calamagrostis canadensis – Carex a spring fen and raised bog in the Lost River viridula – Cladium marisicoides – Lobelia Peatland, northern Minnesota. Journal of kalmii Herbaceous Vegetation; Bluejoint – Ecology 74(2): 393-401. Hairy Sedge – Twig-rush – Ontario Lobelia Herbaceous Vegetation; Great Lakes Sedge Almendinger, J.A., and J.H. Leete. 1998. Rich Shore Fen Regional and local hydrogeology of calcareous fens in the Minnesota River Basin, V.A.5.N.m; Carex lasiocarpa Saturated USA. Wetlands 18(2): 184-202. Herbaceous Alliance; Carex lasiocarpa – Carex buxbaumii – Trichophorum Amon, J.P., C.A. Thompson, Q.J. Carpenter, and caespitosum Boreal Herbaceous Vegetation; J. Mines. 2002. Temperate zone fens of the Wiregrass Sedge – Brown Bog Sedge – glaciated Midwestern USA. Wetlands 22(2): Deerhair Bulrush Boreal Herbaceous 301-317. Vegetation; Boreal Sedge Rich Fen Anderson, D.S, R. B. Davis, S.C. Rooney, C.S. V.A.5.N.m; Carex lasiocarpa Saturated Campbell. 1996. The ecology of sedges Herbaceous Alliance; Carex lasiocarpa – (Cyperaceae) in Maine peatlands. Bulletin of (Carex rostrata) – Equisetum fluviatile the Torrey Botanical Club 123(2): 100-110. Herbaceous Vegetation; Wiregrass Sedge –

Grayling Community Assessment - 90 Barnes, B.V. 1991. Deciduous forest of North County. Michigan. Paper’s of the Michigan America. Pp 219-344 in E. Röhrig and B. Academy of Science, Arts, and Letters 17: 57- Ulrich, eds., Temperate Deciduous Forests. 65. Elsevier, Amsterdam. 635 pp. Crum, H. 1983. Mosses of the Great Lakes Forest. Barnes, B.V., K.S. Pregitzer, T.A. Spies, and V. University of Michigan, Ann Arbor, MI. 417 H. Spooner. 1982. Ecological forest site pp. classification. Journal of Forestry 80(8): 493- 498. Curtis, J.T. 1959. Vegetation of Wisconsin: An Ordination of Plant Communities. University Bay, R.R. 1967. Ground water and vegetation in of Wisconsin Press, Madison, WI. 657 pp. two peat bogs in northern Minnesota. Ecology 48(2): 308-310. Damman, A.H. 1990. Nutrient status of ombrotrophic peat bogs. Aquilo Series Bedford, B.L., and K.S. Godwin. 2003. Fens of Botanica 28: 5-14. the United States: Distribution, characteristics, and scientific connection Dansereau, P., and F. Segadas-Vianna. 1952. versus legal isolation. Wetlands 23(3): 608- Ecological study of the peat bogs of eastern 629. North America. I. Structure and evolution of vegetation. Canadian Journal of Botany 30: Bedford, B.L., M.R. Walbridge, and A. Aldous. 490-520. 1999. Patterns in nutrient availability and plant diversity of temperate North American Dean, D., and H. Coburn. 1927. An ecological wetlands. Ecology 80(7): 2151-2169. study of Linne Bog, Cheboygan County, Michigan with special reference to Boelter, D.H., and E.S. Verry. 1977. Peatland and Nemopanthus mucranata (L.) Trelease. water in the northern Lake States. North Paper’s of the Michigan Academy of Science, Central Forest Experiment Station. USDA Arts, and Letters 8: 87-96. Forest Service General Technical Report NC- 31. 26 pp. Eggers, S.D., and D.M. Reed. 1997. Wetland plants and plant communities of Minnesota Braun, E.L. 1950. Deciduous forests of eastern and Wisconsin. U.S. Army Corps of North America. Hafner Press, New York, NY. Engineers, St Paul, Minnesota. 263 pp. 596 pp. Faber-Langendoen, D., ed., 2001. Plant Burns, G.P. 1906. Bog studies. Field Studies in communities of the Midwest: Classification in Botany. University Bulletin, New Series, an ecological context. Association for 7(14): 3-13. University of Michigan, Ann Biodiversity Information, Arlington, VA. 61 Arbor. pp & appendix (705 pp.).

Chapman, S., A. Buttler, A.-J. Francez, F. Fitzgerald, S., and R.E. Bailey. 1975. Laggoun-Defarge, H. Vasander, M. Schloter, Vegetational characteristics of a circum- J. Combe, P. Grosvernier, H. Harms, D. neutral bog, Barney’s Lake, Beaver Island, Epron, D. Gilbert, and E. Mitchell. 2003. Michigan. The Michigan Academician 7(4): Exploitation of northern peatlands and 477-488. biodiversity maintenance: A conflict between economy and ecology. Frontiers in Ecology Frelich, L.E., and C.G. Lorimer. 1991. Natural and the Environment 1(10): 525-532. disturbance regimes in hemlock-hardwood forests of the Upper Great Lakes region. Coburn, H., D. Dean, and G.M. Grant. 1933. An Ecological Monographs 61(2): 145-164. ecological study of Bryant’s Bog, Cheboygan

Grayling Community Assessment - 91 Gates, F.C. 1942. The bogs of northern Lower Jean, M., and A. Bouchard. 1991. Temporal Michigan. Ecological Monographs 12(3): changes in wetland landscapes of a section of 213-254. the St. Lawrence River, Canada. Environmental Management 15(2): 241-250. Gignac, L.D., L.A. Halsey, and D.H. Vitt. 2000. A bioclimatic model for the distribution of Kost, M.A. 2001. Natural community abstract for Sphagnum-dominated peatlands in North relict conifer swamp. Michigan Natural America under present climatic conditions. Features Inventory, Lansing, MI. 6 pp. Journal of Biogeography 27(5): 1139-1151. Lindeman, R.L. 1941. The developmental history Glaser, P.H. 1992. Raised bogs in eastern North of Cedar Creek Bog, Minnesota. American America – Regional controls for species Midland Naturalist 25(1): 101-112. richness and floristic assemblages. Journal of Ecology 80: 535-554. Michigan Natural Features Inventory. 2003. Draft description of Michigan natural community Glaser, P.H., and J.A. Janssens. 1986. Raised bogs types. (Unpublished manuscript revised in eastern North America: Transitions in March 4, 2003.) Michigan Natural Features landforms and gross stratigraphy. Canadian Inventory, Lansing, MI. 36 pp. Available: Journal of Botany 64: 395-415. http://www.msue.msu.edu/mnfi/lists/natural_c ommunity_types.pdf. Glaser, P.H., G.A. Wheeler, E. Gorham, and H.E. Wright, Jr. 1981. The patterned mires of the Mitsch, W.J., and J.G. Gosselink. 2000. Wetlands. Red Lake Peatland, northern Minnesota: John Wiley and Sons, Inc, New York, NY. Vegetation, water chemistry and landforms. 920 pp. Journal of Ecology 69(2): 575-599. Miller, N. 1981. Bogs, bales, and BTU’s: A Glaser, P.H., J.A. Janssens, and D.I. Siegel. 1990. primer on peat. Horticulture 59: 38-45. The response of vegetation to chemical and hydrological gradients in the Lost River NatureServe. 2005a. NatureServe Explorer: An Peatland, northern Minnesota. Journal of online encyclopedia of life [web application]. Ecology 78(4): 1021-1048. Version 4.2. NatureServe, Arlington, Virginia. Available: Hammerson, G. 1994. Beaver (Castor http://www.natureserve.org/explorer. canadensis): Ecosystem alterations, (Accessed: March 03, 2005.) management, and monitoring. Natural Areas Journal 14(1): 44-57. NatureServe. 2005b. International Ecological Classification Standard: Terrestrial Ecological Heinselman, M.L. 1963. Forest sites, bog Classifications. NatureServe Central processes, and peatland types in the Glacial Databases. Arlington, VA. U.S.A. Data Lake Region, Minnesota. Ecological current as of January 13, 2005. Monographs 33(4): 327-374. Heinselman, M.L. 1965. String bogs and other Osvald, H. 1935. A bog at Hartford, Michigan. patterned organic terrain near Seney, Upper Ecology 16(3): 520-528. Michigan. Ecology 46: 185-188. Richardson, C.J., and P.E. Marshall. 1986. Heinselman, M.L. 1970. Landscape evolution, Processes controlling movement, storage, and peatland types, and the environment in the export of phosphorous in a fen peatland. Lake Agassiz Peatland Natural Area, Ecological Monographs 56(4): 279-302. Minnesota. Ecological Monographs 40(2): 235-261.

Grayling Community Assessment - 92 Riley, J.L. 1989. Southern Ontario bogs and fens Swanson, D.K., and D.F. Grigal. 1989. Vegetation of the Canadian Shield. Wetlands: Inertia or indicators of organic soil properties in Momentum: 355-367. Minnesota. Soil Science Society of America Journal 53: 491-495. Schwintzer, C.R. 1978a. Nutrient and water levels in a small Michigan bog with high tree Swinehart, A.L., and G.R. Parker. 2000. mortality. American Midland Naturalist Palaeoecology and development of peatlands 100(2): 441-451. in Indiana. American Midland Naturalist 143(2): 267-297. Schwintzer, C.R. 1978b. Vegetation and nutrient status of northern Michigan fens. Canadian Verry, E.S. 1975. Streamflow chemistry and Journal of Botany 56: 3044-3051. nutrient yields from upland-peatland watersheds in Minnesota. Ecology 65(5): Schwintzer, C.R. 1979. Vegetation changes 1149-1157. following a water level rise and tree mortality in a Michigan bog. The Michigan Botanist 18: Vitt, D.H., and N.G. Slack. 1975. An analysis of 91-98. the vegetation of Sphagnum-dominated kettle- hole bogs in relation to environmental Schwintzer, C.R. 1981. Vegetation and nutrient gradients. Canadian Journal of Botany 53: status of northern Michigan bogs and conifer 332-359. swamps with a comparison to fens. Canadian Journal of Botany 59: 842-853. Vitt, D.H., H. Crum, and J.A. Snider. 1975. The vertical zonation of Sphagnum species in Schwintzer, C.R, and G. Williams. 1974. hummock-hollow complexes in northern Vegetation changes in a small Michigan bog Michigan. The Michigan Botanist 14(4): 190- from 1917 to 1972. American Midland 200. Naturalist 92(2): 447-459. Wheeler, G.A., P.H. Glaser, E. Gorham, C.M. Siegel, D.I. 1988. Evaluating cumulative effects Wetmore, F.D. Bowers, and J.A. Janssens. of disturbance on the hydrologic function of 1983. Contributions to the flora of the Red bogs, fens, and mires. Environmental Lake Peatland, northern Minnesota, with Management 12(5): 621-626. special attention to Carex. American Midland Naturalist 110(1): 62-96. Siegel, D.I., and P.H. Glaser. 1987. Groundwater flow in a bog-fen complex, Lost River Zoltai, S.C., and D.H. Vitt. 1995. Canadian Peatland, northern Minnesota. Journal of wetlands: Environmental gradients and Ecology 75(3): 743-754. classification. Vegetatio 118: 131-137.

Swain, A.M. 1973. A history of fire and vegetation in northeastern Minnesota as Abstract Citation: recorded in lake sediments. Quaternary Cohen, J.G. 2005. Natural community abstract for Research 3: 383-396. poor fen. Michigan Natural Features Inventory, Lansing, MI. 13 pp.

Grayling Community Assessment - 93 Grayling Community Assessment - 94 Appendix 5. Bog

Overview: A nutrient-poor peatland inputs of nutrients and water primarily from characterized by acidic, saturated peat and the precipitation (Gignac et al. 2000). Additional open prevalence of Sphagnum mosses and ericaceous wetlands occurring on peat include northern fen shrubs. Located in depressions in glacial outwash and poor fen. Fens are minerotrophic (nutrient- and sandy glacial lakeplains and in kettles on rich) wetlands that are dominated by sedges, pitted outwash and moraines; frequently occurring rushes, and grasses (Mitsch and Gosselink 2000). as a floating mat on the margins of lakes and The hydrology of fens is influenced by ponds. Fire occurs naturally during drought groundwater, and as a result, fens have higher periods and can alter the hydrology, mat surface, nutrient availability, increased alkalinity (less and flora. acidity), and greater species richness compared to bogs, with poor fens being most similar to bogs in Global and State Rank: G5/S4 terms of these factors and species composition. In addition to a greater importance by graminoids, Range: Bogs are a frequent peatland type of fens also are less dominated by sphagnum mosses glaciated landscapes of the entire northern (Sphagnaceae) with brown mosses hemisphere and are characterized by remarkably (Amblystegiaceae) being more prevalent. uniform floristic structure and composition across Intermittent wetlands are herb or herb-shrub the circumboreal region (Curtis 1959). In North dominated wetlands that experience fluctuating America, they are found throughout the glaciated water levels seasonally and yearly and have soils Midwest (Michigan, Minnesota, Wisconsin, and that range from loamy sand and peaty sand to northern portions of Illinois, Indiana, and Ohio) peaty muck and are very strongly acid to strongly and the northeastern United States (New York, acid. Like bogs, muskegs and poor conifer swamp New Hampshire, Vermont, and Maine), and range are nutrient-poor, acidic wetlands. However, these from central Canada (Ontario, Manitoba, and ombrotrophic peatlands exhibit a greater degree of Quebec) to the maritime provinces (Nova Scotia canopy closure than bogs (muskegs having and New Brunswick) (Faber-Langendoen 2001, clumped and scattered conifers and poor conifer NatureServe 2005a). Subtle variations in overall swamp being a closed canopy system). Closed species composition and physiognomy occur canopy, minerotrophic peatlands include rich across its range along north-south and east-west conifer swamp, a Thuja occidentalis (northern climatic gradients (Glaser 1992). In Michigan, white cedar) dominated system found north of the bogs are common throughout the northern Lower tension zone, and relict conifer swamp, which is Peninsula and the Upper Peninsula and are less dominated by Larix laricina (tamarack) and common south of the climatic tension zone occurs primarily south of the tension zone (Kost (Amon et al. 2002). Bogs and other peatlands 2001). occur where excess moisture is abundant (where precipitation is greater than evapotranspiration) Rank Justification: Bogs are frequent features of (Mitsch and Gosselink 2000). Conditions suitable the northern Great Lakes region, occurring for the development of bogs have occurred in the throughout the northern Lower Peninsula and the northern Lake States for the past 3000-5000 years Upper Peninsula and sporadically south of the following climatic cooling (Boelter and Verry climatic tension zone. The northern lake states 1977, Heinselman 1970). contain over six million hectares (15 million acres) of peatland (Boelter and Verry 1977). Several other natural peatland communities also Within the southern portion of their range, bogs occur in Michigan and can be distinguished from typically occur as isolated pockets separated by ombrotrophic (nutrient-poor) bogs, based on large expanses of agricultural lands (Amon et al. comparisons of nutrient levels, flora, canopy 2002). Historically, widespread fires following the closure, and groundwater influence. Bogs, peat- turn of the century logging drastically altered covered wetlands raised above the surrounding many peatlands, either converting poor conifer groundwater by an accumulation of peat, receive swamp to open bog systems or destroying the peat

Grayling Community Assessment - 95 and converting bogs to wetlands without organic arundinaceae (reed canary grass) and Typha spp. soils (mineral soil wetlands) (Dean and Coburn (cat-tails) (Almendinger and Leete 1998). 1927, Gates 1942, Curtis 1959). Logging of cedar and tamarack from peatland systems also favored Physiographic Context: Two landscape features the conversion of forested peatlands to open, are conducive to the development of peat; small ombrotrophic bogs (Gates 1942, Dansereau and ice-block basins and poorly-drained, level terrain Segadas-Vianna 1952, Riley 1989). Beginning in (Boelter and Verry 1977). Bogs occur in kettle the 1920s, effective fire control by the U.S. Forest depressions on pitted outwash and moraines and Service and state agencies reduced the acreage of on flat areas or mild depressions of glacial fires ignited by man or lightning (Swain 1973). In outwash and glacial lakeplains (Lindeman 1941, landscapes where frequent fire was the prevalent Gates 1942, Curtis 1959, Bay 1967, Boelter and disturbance factor, fire suppression has led to the Verry 1977, Glaser and Janssens 1986, Siegel conversion of open bogs to closed canopy 1988, Michigan Natural Features Inventory 2003, peatlands (Curtis 1959, Riley 1989). Peat mining NatureServe 2005a). The overall topography of and cranberry farming have degraded numerous bogs is flat to gently undulating with bogs throughout the region (Gates 1942, Curtis microtopography characterized by hummocks and 1959, Eggers and Reed 1997, Chapman et al. hollows (Heinselman 1963, Vitt and Slack 1975, 2003). Michigan, along with Florida and Wheeler et al. 1983, Glaser et al. 1990, Minnesota, are leaders in peat production in the NatureServe 2005a). Many bogs are oriented U.S. (Miller 1981). In addition to direct impacts to northwest to southeast, corresponding to the vegetation, alteration of peatland hydrology from direction of glacial movement (Schwintzer road building, creation of drainage ditches and 1978a). Bogs found in kettle depressions are dams, and runoff from logging has led to the associated with active or extinct glacial lakes drastic change of bog composition and structure (Curtis 1959). Within kettle depressions, bogs can (Schwintzer and Williams 1974, Riley 1989, occupy the entire basin or frequently occur as a Chapman et al. 2003). Bog vegetation is mat (floating or grounded) on the margin of the extremely sensitive to minor changes in water remaining glacial lake (Vitt and Slack 1975, levels and chemistry (Siegel 1988, Riley 1989). Schwintzer 1978a). When bogs occur along the Succession to more minerotrophic wetlands can edge of large bodies of water, they are found in occur as the result of increased alkalinity and sheltered bays or coves that are protected from raised water levels, which can cause the increased wave and ice action, which can prevent the decomposition of acidic peats. Lowering of water development of peat or erode existing peat mats tables from drainage can allow for tree and shrub (Gates 1942). Bogs occurring on former glacial encroachment into open bogs and the eventual lake beds and drainageways tend to be more succession to closed canopy peatland. The natural extensive than kettle bogs, which are limited in acidity of bogs makes them especially susceptible area by the size of the glacial ice-block which to acid rain and air pollution (Siegel 1988, formed the basin (Lindeman 1941). The large Chapman et al. 2003). Atmospheric deposition peatlands of lakeplains and outwash plains are can contribute Nitrogen, Sulphur, Calcium and often over 100 acres while bogs found in kettle heavy metals to bogs (Damman 1990, Chapman et depressions typically range from 10 to 30 acres. al. 2003). Dust-fall and atmospheric deposition from air pollution are particularly threats to bog Bogs within large wetland complexes typically systems in the southern portion of their range, occur adjacent to other peatland communities, where bogs are surrounded by cultivated land and often grading into poor conifer swamp, muskeg, close to industrial and urban centers (Damman or poor fen. More minerotrophic systems such as 1990). Eutrophication from pollution and altered northern fen, prairie fen, shrub thicket, wet hydrology can detrimentally impact bogs by meadow, rich conifer swamp, and relict conifer generating conditions favorable for the invasion of swamp can occur along the outer margins of bogs exotic species (Riley 1989) and dominance by where groundwater seepage from the adjacent aggressive, common natives such as Phalaris uplands is prevalent. Bogs within kettle depressions that contain active glacial lakes and

Grayling Community Assessment - 96 ponds often border aquatic communities such as characterized by low primary productivity which submergent marsh and emergent marsh. Bog mats is correlated with the very low concentrations of can also occur as a vegetative zone within available Calcium, Magnesium, Nitrogen, intermittent wetlands, coastal plain marsh, and Phosphorous, and Potassium in the surface water wooded dune and swale complexes. A wide array and peat (Heinselman 1963, Heinselman 1970, of upland community types can occur adjacent to Schwintzer 1978a, Schwintzer 1981, Schwintzer bogs; some of the more frequent neighboring and Tomberlin 1982, Wheeler et al. 1983, upland systems include dry-mesic northern forest, Richardson and Marshall 1986, Riley 1989, dry northern forest, mesic northern forest, pine Glaser 1992, Bedford et al. 1999, Mitsch and barrens, dry-mesic southern forest, and dry Gosselin 2000). southern forest. Soils: The organic soils of bogs are composed of Hydrology: No apparent inlets or outlets supply peat which forms a continuous mat and can range or drain bogs which are isolated from ground in thickness from 3-30 feet (1-8 meters) but is water influence as the result of peat accumulation typically 2-3 meters deep (Heinselman 1965, Bay (Dean and Coburn 1927, Schwintzer 1978b, Riley 1967, Heinselman 1970, Siegel and Glaser 1987). 1989, Swineheart and Parker 2000, Hoffman Peat is a fibrous network of partially decomposed 2002, NatureServe 2005a). Bogs are ombrotrophic organic material that is formed under anaerobic to weakly minerotrophic peatlands, receiving conditions (Heinselman 1963, Almendinger et al. inputs of water and nutrients primarily from ion- 1986). The surface peats of bogs are dominated by poor precipitation (Heinselman 1970, Verry 1975, saturated fibric peat which is loosely compacted, Boelter and Verry 1977, Schwintzer 1981, contains partially decomposed Sphagnum moss Schwintzer and Tomberlin 1982, Siegel 1988, with fragments of wood and occasionally sedge, Riley 1989, Damman 1990, Glaser et al. 1990, and like the surface water, is extremely acidic, Mitsch and Gosselink 2000, Bedford and Godwin cool, and characterized by low nutrient 2003). The rooting zone is dominated by inputs of availability and oxygen levels (Burns 1906, Curtis atmospheric water and nutrients (Bedford and 1959, Heinselman 1963, Heinselman 1970, Godwin 2003). The water retaining capacity of Schwintzer and Williams 1974, Boelter and Verry sphagnum peat is tremendous and as a result bogs 1977, Almendinger et al. 1986). Fibric peat has are saturated, anoxic systems with water tables high water retaining capacity and large near the surface (Burns 1906, Dansereau and intercellular pores that permit rapid water Segadas-Vianna 1952, Curtis 1959, Heinselman movement (Boelter and Verry 1977, Swanson and 1970, Schwintzer 1978b, Siegel and Glaser 1987, Grigal 1989). Peat composition changes with Glaser 1992, Eggers and Reed 1997). The depth and depending on the successional history stagnant surface waters of bogs are characterized of a given bog. Generally, fiber content and by high acidity, low available nutrients, low hydraulic conductivity usually decrease with specific conductivity, cool temperatures, depth; deeper peats are more decomposed, retain anaerobic conditions, and high levels of dissolved more water, and drain slower than surface peats organic matter that imparts a brown color (Gates (Verry 1975, Boelter and Verry 1977). 1942, Verry 1975, Schwintzer 1978a, Glaser et al. 1981, Wheeler et al. 1983, Riley 1989, Damman Climate: Peatlands develop in humid climates 1990, Glaser 1992). Studies of bog water and peat where precipitation exceeds evapotranspiration across the northern Great Lakes have found pH (Boelter and Verry 1977, Gignac et al. 2000). The measurements to range from 3.2-4.7 (Heinselman northern Lake States are characterized by a 1970, Boelter and Verry 1977, Schwintzer 1981, humid, continental climate with long cold winters Schwintzer and Tomberlin 1982, Wheeler et al. and short summers that are moist and cool to 1983, Riley 1989, Glaser et al. 1990). The high warm (Gates 1942, Boelter and Verry 1977, acidity of bogs limits the availability and uptake Damman 1990, Mitsch and Gosselink 2000). The of essential mineral plant nutrients which are Michigan range of bog falls within the area inherently scarce in these systems because of the classified by Braun (1950) as the Northern lack of groundwater input (Glaser 1992). Bogs are Hardwood-Conifer Region (Hemlock/White

Grayling Community Assessment - 97 Pine/Northern Hardwoods Region) and within the Likewise, high levels of acidity have inhibitory following regions classified by Albert et al. (1986) effects on decay organisms (Heinselman 1963, and Albert (1995): Region I, Southern Lower Mitsch and Gosselink 2000). Once Sphagnum Michigan; Region II, Northern Lower Michigan; mosses become established on the peat mat, they Region III, Eastern Upper Michigan; and Region maintain and enhance saturated and acidic IV, Western Upper Michigan. The Northern conditions, which in turn promote continued peat Hardwood-Conifer Region has a cool snow-forest development. The ability of sphagnum to absorb climate with warm summers. The mean number of and hold cations increases the acidity and low freeze-free days is between 90 and 220, and the nutrient availability of peatlands (Osvald 1935, average number of days per year with snow cover Curtis 1959, Verry 1975, Vitt and Slack 1975, of 2.5 cm or more is between 10 and 140. The Boelter and Verry 1977). Sphagnum moss, which normal annual total precipitation ranges from 740 has numerous pores, partitions, and capillary to 900 mm with a mean of 823 mm. The daily space, has an enormous water-holding capacity maximum temperature in July ranges from 24 to (Osvald 1935, Dansereau and Segadas-Vianna 32 °C (75 to 90 °F), the daily minimum 1952, Curtis 1959). Sphagnum peat can hold 15 to temperature in January ranges from –21 to –4 °C 30 times its own weight in water (Miller 1981, (-5 to 25 °F) and the mean annual temperature is 7 Mitsch and Gosselink 2000). °C (45 °F) (Albert et al. 1986, Barnes 1991). Temperatures vary less in bogs compared to the Development and expansion of peatlands occurs surrounding landscape because of the insulating via two distinct processes: lake-filling and effect of bogs’ saturated peat carpet in the paludification. Lake-filling or terrestrialization growing season and snow cover in the winter occurs in small lakes with minimal wave action, (Burns 1906, Curtis 1959, Heinselman 1963, where gradual peat accumulation results in the Glaser 1992). In Wisconsin, Curtis (1959) development of a bog mat that can fill the basin or observed that at root level, temperatures during occur as a floating mat in the lake or as a the growing season rarely exceed 60°F and are grounded mat along the water’s edge (Burns 1906, usually between 45 and 55 °F. In Minnesota, Gates 1942, Bay 1967, Curtis 1959, Heinselman Heinselman (1963) found that the maximum bog 1963, Mitsch and Gosselink 2000). Succession in lake-filled bogs typically proceeds from lake to temperature in August was 56 °F and the marsh to fen to bog (Heinselman 1963, Boelter minimum temperature in late winter was 30 °F. and Verry 1977, Schwintzer 1981, Swineheart and Bogs are characterized by microclimates that are Parker 2000). Floating mats of fen sedges (i.e., cooler in the summer and warmer in the winter Carex lasiocarpa) pioneer open water and compared to the regional climate. accumulate organic matter in the form of peat,

which is invaded by sphagnum and ericaceous Natural Processes: Peat establishment requires shrubs (Osvald 1935, Gates 1942, Schwintzer and an abundant supply of water. As noted, Williams 1974, Swineheart and Parker 2000). ombrotrophic peatlands occur in regions where precipitation is greater than evapotranspiration Fallen logs in kettle lakes and ponds can also and on sites with blocked drainage (Dansereau provide the substrate for bog vegetation and Segadas-Vianna 1952, Boelter and Verry establishment and invasion of the water: 1977, Mitsch and Gosselink 2000). Saturated and Chamaedaphne calyculata (leatherleaf ) is inundated conditions inhibit organic matter particularly adept at expanding along logs (Dean decomposition and allow for the accumulation of and Coburn 1927, Gates 1942, Dansereau and peat (Almendinger and Leete 1998). Under cool, Segadas-Vianna 1952). Peatland vegetation has anaerobic, and acidic conditions, the rate of been recorded advancing into kettle lakes at a rate organic matter accumulation exceeds organic of 2.1cm/year (Schwintzer and Williams 1974). decay (Schwintzer and Williams 1974, Damman Estimates of vertical accumulation of bog peat 1990, Mitsch and Gosselink 2000). Low levels of range between 100 to 200cm/1000 years (Mitsch oxygen protect plants from microorganisms and and Gosselink 2000). For both lake-filling and chemical actions that cause decay (Miller 1981). paludification, peat accumulates above the water

Grayling Community Assessment - 98 table and the bog becomes isolated from fire (Wheeler et al.1983). Fire severity and groundwater influence (Heinselman 1970, Boelter frequency in bogs is closely related to fluctuations and Verry 1977, Glaser and Janssens 1986, in water level. Prolonged periods of lowered water Mitsch and Gosselink 2000). Paludification is the table can allow the surface peat to dry out enough blanketing of terrestrial systems (often forests) by to burn (Schwintzer and Williams 1974). When the overgrowth of peatland vegetation (Dansereau the surface peat of bogs burns, the fire releases and Segadas-Vianna 1952, Heinselman 1963, organic matter from the peat, kills seeds and latent Mitsch and Gosselink 2000). Paludified peatlands buds, stimulates decay, and slows peat develop on flat areas (typically lakeplain) where accumulation (Damman 1990, Jean and Bouchard peat develops vertically and spreads horizontally 1991). Peat fires can convert bogs to more (Heinselman 1965, Boelter and Verry 1977). graminoid dominated peatlands such as Succession in paludified bogs can proceed from intermittent wetlands or poor fens or if the peat is fen, marsh, or shrub carr to swamp forest to bog completely destroyed, to mineral soil wetlands (Heinselman 1970, Schwintzer 1981). such as northern wet meadow (Curtis 1959).

Once established, bogs can persist for hundreds of Flooding often contributes to the development, years given stable hydraulic conditions. A expansion, and maintenance of bogs. Dam- discharge of alkaline groundwater at the peat building activities of beaver can result in blocked surface of a bog, caused by a change in hydraulic drainage and flooding which facilitate sphagnum head, can result in the conversion of bog peat development and expansion (Heinselman vegetation to fen vegetation (Siegel and Glaser 1963, Heinselman 1970) and can also cause 1987, Glaser et al. 1990). Mixing of as little as grounded bog mats to become loosened from the 10% groundwater from underlying calcareous bottom and float (Gates 1942). Roots of peatland parent material with acid bog water is sufficient to trees are physiologically active near the surface raise the peatland pH from 3.6 to 6.8 (Glaser et al. and are quickly killed when the water table rises 1990). Bogs are very sensitive to changes in pH following flooding (Glaser and Janssens 1986). and subsequent availability of nutrients: fen Within kettle bogs, flooding induced tree vegetation can replace bog flora when pH mortality is greater on grounded bog mats increases above 4.5 (Siegel 1988). However, compared to free floating mats: free mats float up conversions of bog to fen have been seldom with rising water table while grounded mats reported in the literature (Glaser et al. 1990). become inundated and have shallower aerobic More typically, bogs are converted to shrub zones (Schwintzer 1973, Schwintzer 1978a, swamp or swamp forest following the lowering of Schwintzer 1979). In addition to flooding, kettle the water table. Water table lowering results in bogs can be influenced by waves and ice. Wave increased decomposition rates of organic matter and ice action can prevent the expansion of bog and facilitates the invasion of bogs by mats by eroding the shoreline vegetation (Gates opportunistic woody species (Almendinger and 1942). Leete 1998, Gignac et al. 2000). The natural disturbance regime in bogs is also Disturbance factors influencing bogs include fire, influenced by wind. The Great Lakes region is flooding, windthrow, and insects. Numerous bogs one of the most active weather zones in the contain charcoal within their peat profile (Curtis northern hemisphere, with polar jet streams 1959, Heinselman 1963) and many researchers positioned overhead much of the year. More have reported fire as a prevalent part of bog’s cyclones pass over this area than any other area in disturbance regime (Dean and Coburn 1927, the continental U.S. (Frelich and Lorimer 1991). Gates 1942, Curtis 1959). Surface fire can Trees growing in bogs are particularly susceptible contribute to the maintenance of bogs by killing to windthrow because sphagnum peat provides a encroaching trees without completely removing poor substrate for anchoring trees (Burns 1906). the sphagnum (Curtis 1959, Vitt and Slack 1975). The living roots of woody peatland plants occur in Many of the ericaceous plants that thrive in bogs a shallow rooting zone, generally restricted to the are fire-adapted and often grow densely following uppermost few centimeters where there is

Grayling Community Assessment - 99 sufficient oxygen to maintain aerobic respiration (Glaser et al. 1990, Glaser 1992). Species (Glaser and Janssens 1986). The superficial diversity within bogs is strongly correlated to rooting of trees results in numerous windthrows microtopography (Glaser et al. 1990). Within a (Dansereau and Segadas-Vianna 1952, Eggers and bog, floristic composition is determined by Reed 1997). Tree survival in bogs is also limited gradients in pH, light, soil moisture, and cation by insects and parasites. Insect outbreaks of the concentrations (nutrient availability) (Heinselman Pristiphora erichsonii (larch sawfly) cause heavy 1970, Vitt and Slack 1975, Schwintzer 1978a, mortality of Larix laricina (tamarack) while, the Glaser et al. 1981). plant parasite Arceuthobium pusillum (dwarf mistletoe) kills Picea mariana (black spruce) Bogs are dominated by mosses from the (Coburn et al. 1933, Gates 1942, Heinselman Sphagnaceae and shrubs from the Ericaceae; 1963). other well-represented families include the Cyperaceae, Orchidaceae, and Ranunculaceae Vegetation Description: Bogs are characterized (Gates 1942, Curtis 1959, Heinselman 1970). The by a continuous carpet of sphagnum moss, a poor most important primary producers within bogs are herbaceous layer, low ericaceous, evergreen ericaceous shrubs and sedges (Mitsch and shrubs, and widely scattered and stunted conifer Gosselink 2000). Bog flora is predominantly trees (Gates 1942, Curtis 1959, Verry 1975, Vitt spring flowering and heliophitic (Curtis 1959, and Slack 1975, Glaser et al. 1991, NatureServe Dansereau and Segadas-Vianna 1952). Bog plants 2005a). Floristically bogs are homogenous and of have developed a diversity of adaptations to cope limited diversity, exhibiting remarkably uniform with low nutrient availability including plant structure and composition across their wide range carnivory, evergreen leaves, sclerophylly (thick (Curtis 1959, Riley 1989). Slight variations in epidermal tissue), and high root biomass (Mitsch composition, especially within the shrub and tree and Gosselink 2000). While bogs are dominated layer, occur in Michigan bogs along a longitudinal by plants that thrive under ombrotrophic gradient. Southerly bogs have more of a conditions, occasionally minerotrophic indicators deciduous tree and shrub component compared to may be present in bogs at low cover. Plants found the northern coniferous bogs. The harsh growing typically in more alkaline habitat such as Betula conditions of bogs (high acidity, low nutrient pumila (bog birch), Carex aquatilis (water sedge), availability, and saturated peat) results in a unique and Carex stricta (tussock sedge), can occur but depauperate flora: relatively few species have sporadically in bogs when their roots extend evolved the necessary adaptations to cope with beneath the bog mat to minerotrophic peat ombrotrophic conditions (Siegel 1988, Glaser influenced by ground water (NatureServe 2005a). 1992, Mitsch and Gosselink 2000). Very few introduced, weedy species are able to establish The continuous moss layer of bogs is dominated within bogs and fens because of the unique by sphagnum mosses especially Sphagnum growing conditions and competition from the magellanicum, S. angustifolium, and S. fuscum adapted flora (Riley 1989). In a study of bogs (Vitt and Slack 1975, Schwintzer 1978a, across eastern North America, Glaser (1992) NatureServe 2005a). Additional mosses can found the native vascular bog flora to be limited include S. capillaceum, S. capillifolium, S. to only 81 species and the mean number of compactum, S. cuspidatum, S. papillosum, S. species per bog to be below 26. The mean number recurvum, and Drepanocladus aduncus (Gates of species per plot in kettle bogs in the northern 1942, Vitt and Slack 1975, Crum 1983, Riley Lower Peninsula of Michigan was found to be 1989, Glaser et al. 1990). The hummock and fifteen by Vitt and Slack (1975) and fourteen by hollow microtopography of bogs allows for high Schwintzer (1981), with a range of 9-20. For bogs levels of bryophyte diversity since individual within Michigan Natural Features Inventory’s species of sphagnum occur at specific elevations database, the mean number of species per bog is (Vitt and Slack 1975, Wheeler et al. 1983, Riley approximately 30. Species richness of bogs is 1989). Hollows support S. magellanicum, S. related to geographical location, climatic factors, cuspidatum, and S. papillosum (Vitt and Slack nutrient availability, and habitat heterogeneity 1975, Vitt et al. 1975, Heinselman 1970, Wheeler

Grayling Community Assessment - 100 et al. 1983, Riley 1989). The lower, moist slopes shrub layer is typically less than three feet high of hummocks often support S. magellanicum and and usually covers at least 25% of the bog area S. recurvum while the drier hummock crests are (Eggers and Reed 1997, NatureServe 2005a). In dominated by S. fuscum, S. capillaceum, and S. addition to leatherleaf, the following heath shrubs cappillifolium (Vitt et al. 1975, Wheeler et al. are important components of bogs: Andromeda 1983, Riley 1989). The vertical zonation of glaucophylla (bog-rosemary), Gaylussacia species corresponds to gradients in pH and baccata (huckleberry), Kalmia angustifolia moisture with the hollows being wetter and more (sheep-laurel), K. polifolia (bog-laurel), Ledum alkaline than the drier and more acidic tops of the groenlandicum (Labrador-tea), Vaccinium hummocks (Vitt et al. 1975, Wheeler et al. 1983). angustifolium (low sweet blueberry), V. macrocarpon (large cranberry), V. myrtilloides The herbaceous layer of bogs is dominated by (Canada blueberry), and V. oxycoccos (small cyperaceous plants. Fine-leaved sedges are more cranberry). The tall shrub layer of bogs is less prevalent in bogs while broad-leaved sedges dense than the low shrub layer and is often dominate minerotrophic sites (Boelter and Verry restricted to the periphery of the bog. Tall shrubs 1977). Several sedges that are characteristic of typical of bogs include Aronia prunifolia bogs include Carex oligosperma (few-seed (chokeberry), Nemopanthus mucranta (mountain sedge), Carex pauciflora (few-flower sedge), and holly), Salix pedicellaris (bog willow), Spirea Carex lasiocarpa (wiregrass sedge). Other sedges tomentosa (steeplebush), and Viburnum that often occur in bogs are C. limosa (mud cassinoides (wild-raisin). South of the climatic sedge), C. paupercula (bog or poor sedge), C. tension zone, Cephalanthus occidentalis rostrata (beaked sedge), and C. trisperma (three- (buttonbush), Toxicodendron vernix (poison seeded sedge). Additional graminoids that thrive sumac), and Vaccinium corymbosum (highbush in bogs include Cladium marisicoides (twig-rush), blueberry) frequently occur along bog margins. Dulichium arundinaceum (three-way sedge), As noted, bog birch can occur at low cover when Eriophorum angustifolium (tall cotton-grass), E. it roots can extend beneath the bog mat to spissum (sheathed cotton-grass), E. vaginatum minerotrophic peat. (tussock cotton-grass), E. virginicum (tawny cotton-grass), Rhynchospora alba (white beak- Trees within bogs are widely scattered and stunted rush), Scheuchzeria palustris (rannoch-rush), and (seldom reaching six meters) (Wheeler et al. 1983, Scirpus spp. (bulrushes). The following is a list of NatureServe 2005a). Tree cover is typically below prevalent bog herbs: Epilobium angustifolium ten percent (NatureServe 2005a). The most (fireweed), E. ciliatum (fringed willow-herb), Iris common canopy dominants are Picea mariana versicolor (wild blue flag), Menyanthes trifoliata (black spruce) and Larix laricina (tamarack). (bogbean), Smilacina trifolia (false Solomon- Additional associates include Pinus banksiana seal), and Triglochin maritima (arrow-grass). (jack pine), Pinus strobus (white pine), and Acer Insectivorous plants, Drosera rotundifolia rubrum (red maple), with red maple being more (roundleaf sundew), Drosera intermedia (spoon- prevalent south of the climatic tension zone. leaf sundew), Sarracenia purpurea (pitcher- (Above species lists compiled from Michigan plant), and Utricularia intermedia (flat-leaved Natural Features Inventory database, Dean and bladderwort), are common features of fens. Coburn 1927, Coburn et al. 1933, Osvald 1935, Woodwardia virginica (chain-fern) is one of few Gates 1942, Dansereau and Segadas-Vianna 1952, ferns that occur in bogs. Bogs frequently contain Curtis 1959, Heinselman 1963, Heinselman 1965, open pools of water or are surrounded by moats Bay 1967, Heinselman 1970, Schwintzer and that contain emergent vegetation such as Nuphar Williams 1974, Vitt and Slack 1975, Schwintzer spp. (pond-lilies) and Nymphaea odorata (sweet- 1978a, Glaser et al. 1981, Schwintzer 1981, scented water-lily). Wheeler et al. 1983, Riley 1989, Glaser et al. 1990, Glaser 1992, Anderson et al. 1996, Eggers The shrub layer of bogs is dominated by low, and Reed 1997, Mitsch and Gosselink 2000, ericaceous shrubs with Chamadaephne calyculata Swinehart and Parker 2000, NatureServe 2005a.) (leatherleaf) being the most prevalent. The dwarf

Grayling Community Assessment - 101 Michigan Indicator Species: black spruce, Carex establish rookeries. Other rare birds that could oligosperma, leatherleaf, pitcher plant, sphagnum occur in bogs are Botaurus lentiginosus moss, and sundew. (American bittern, state special concern), Circus cyaneus (northern harrier, state special concern), Other Noteworthy Species: Bogs provide habitat Coturnicops noveboracensis (yellow rail, state for numerous rare insect species including threatened), Falcipennis canadensis (spruce Appalachia arcana (secretive locust, state special grouse, state special concern), and Picoides concern), Atlanticus davisi (Davis's shield-bearer, arcticus (black-backed woodpecker, state special state special concern), Calephelis mutica (swamp concern). Gavia immer (common loon, state metalmark, state special concern), Boloria freija threatened) establish nest sites on natural islands (Freija fritillary, state special concern butterfly), and bog-mats. Small mammals associated with Boloria frigga (Frigga fritillary, state special bog habitat include Sorex fumeus (smoky shrew, concern butterfly), Erebia discoidalis (red-disked state special concern) and Cryptotis parva (least alpine, state special concern butterfly), Erynnis shrew, state threatened). Alces alces (moose, state baptisiae (wild indigo duskywing, state special threatened), Canis lupus (gray wolf, state concern), Liodessus cantralli (Cantrall's bog threatened), and Lynx canadensis (lynx, state beetle, state special concern), Merolonche dollii endangered) utilize peatland habitat (Mitsch and (Doll’s merolonche moth, state special concern), Gosselink 2000). Neoconocephalus lyrists (bog conehead, state special concern), Oarisma poweshiek (poweshiek In general, the population of animals is low in skipperling, state threatened), Oecanthus laricis bogs because of the low productivity of bog (tamarack tree cricket, state special concern), plants, the unpalatability of bog vegetation, and Orchelimum concinnum (red-faced meadow the high acidity of bog waters (Mitsch and katydid, state special concern), Paroxya hoosieri Gosselink 2000). Melospiza georgiana (swamp (Hoosier locust, state special concern), sparrow) and M. melodia (song sparrow) are Somatochlora incurvata (incurvate emerald, state typical bog songbirds. Common herptiles that special concern dragonfly), and Williamsoni frequent bogs include Bufo americanus fletcheri (ebony boghaunter, state special concern americanus (Eastern American toad), Rana dragonfly). Numerous butterflies and moths are pipiens (Northern leopard frog), and Thamnophis restricted to bogs and fens because their food sirtalis sirtalis (garter snake) (Riley 1989). Bogs plants occur within these peatland systems (Riley provide important habitat for small mammals such 1989). Rare herptiles that utilize bogs include as Blarina brevicauda (short-tailed shrew), Castor Acris crepitans blanchardi (Blanchard’s cricket canadensis (beaver), Microtus pennsylvanicus frog, state special concern), Clemmys guttata (meadow vole), Mustela vison (mink), Ondatra (spotted turtle, state threatened), Elaphe obsoleta zibethicus (muskrat), and Sorex cinereus (masked obsolete (black rat snake, state special concern), shrew) (Curtis 1959, Mitsch and Gosselink 2000). Emys blandingii (Blanding’s turtle, state special Both muskrats and beaver can profoundly concern), Nerodia erythrogaster neglecta influence the hydrology of bogs. Muskrats create (copperbelly watersnake, state endangered), open water channels through the bog peat and Pseudacris triseriata maculata (boreal chorus beavers can cause substantial flooding through frog, state special concern), Sistrurus catenatus their dam-building activities (Gates 1942, catenatus (eastern massasauga, state special Heinselman 1963). concern), and Terrapene carolina carolina (eastern box turtle, state special concern). If Rare plants associated with bogs include suitable nesting trees or snags are available, Amerorchis rotundifolia (round-leaved orchis, Haliaeetus leucocephalus (bald eagle, state state endangered), Carex wiegandii (Wiegand's threatened), Falco columbarius (merlin, state sedge, state threatened), Empetrum nigrum (black threatened), and Pandion haliaetus (osprey, state crowberry, state threatened), Isotria verticillata threatened) can be found nesting in these systems (whorled pogonia, state threatened), Platanthera and Ardea herodias (great blue heron, protected ciliaris (orange or yellow fringed orchid, state by the Migratory Bird Treaty Act of 1918) can threatened), Platanthera leucophaea (eastern

Grayling Community Assessment - 102 prairie fringed orchid, state endangered and listed Research Needs: Bog has a broad distribution as federally threatened), Rubus acualis (dwarf and exhibits subtle regional, physiographic, raspberry, state endangered), and Sarracenia hydrologic, and edaphic variants. The diversity of purpurea f. heterophylla (yellow pitcher-plant, variations throughout its range demands the state threatened). Eleocharis radicans (spike-rush, continual refinement of regional classifications extirpated in Michigan) was historically known that focus on the inter-relationships between from floating mat bogs in Washtenaw County. vegetation, physiography, hydrology, and the successional history of the peat (Heinselman Conservation and Biodiversity Management: 1963, Fitzgerald and Bailey 1975, Barnes et al. Bog is a widespread community type in the Great 1982). Bogs and related community types (poor Lakes region that contributes significantly to the fen, muskeg, and intermittent wetland) can be overall biodiversity of northern Michigan by difficult to differentiate (Heinselman 1963, providing habitat for a unique suite of plants and NatureServe 2005a). Research on abiotic and wide variety of animal species. Numerous rare biotic indicators that help distinguish similar and unique species are associated with bogs. By peatlands would be useful for field classification. storing high levels of sequestered carbon and Systematic surveys for bogs and related peatlands serving as carbon sinks, bogs and related are needed to help prioritize conservation and peatlands play an important role in global management efforts. Little is known about the fire geochemical cycles. Bogs also preserve paleo- regimes of bogs and the interaction of disturbance environmental records: a wealth of information is factors within these systems. As noted by stored in the remains of plants, animals, and Hammerson (1994), beaver significantly alter the atmospheric particles deposited and stored in bog ecosystems they occupy. An important research peat profiles (Chapman et al. 2003). The primary question to examine is how the wetland mechanism for preserving bogs is to maintain ecosystems of the Great Lakes have been and their hydrology. As noted, peatland systems are continue to be affected by fluctuations in sensitive to slight changes in water chemistry. A populations of beaver. Experimentation is needed serious threat to bog hydrology is posed by off- to determine how best to prevent shrub and tree road-vehicle traffic, which can drastically alter encroachment of bogs that are threatened by bog hydrology through rutting. Reduction of conversion to shrub thicket or conifer swamp. A access to peatland systems will help decrease better understanding is needed of the influence of detrimental impacts. Resource mangers operating direct and indirect anthropogenic disturbance on in uplands adjacent to bogs should take care to peatlands (Amon et al. 2002). Effects of minimize the impacts of management to management within bogs should be monitored to hydrologic regimes, especially increased surface allow for assessment and refinement. More flow. This can be accomplished by establishing a research is needed to elucidate the relationship of no-cut buffer around bogs and avoiding road chemical factors and nutrient levels to floristic construction and complete canopy removal in community structure of peatlands (Amon et al. stands immediately adjacent to bogs. Elevation of 2002). Given the sensitivity of peatlands to slight a bog’s water table and clear-cutting within a bog changes in hydrology and nutrient availability, it can result in the expansion of leatherleaf and is important for scientists to predict how peatlands sphagnum and a subsequent decrease in floristic will be affected by global warming and diversity (Schwintzer 1979). Where shrub/tree atmospheric deposition of nutrients and acidifying encroachment threatens to convert open wetlands agents (Heinselman 1970, Riley 1989, Bedford et to shrub-dominated systems or forested swamps, al. 1999, Gignac et al. 2000, Mitsch and Gosselink prescribed fire or selective cutting can be 2000). Peat deposits are of great scientific interest employed to maintain open conditions. because they contain historical ecological records Silvicutural management of bogs to preserve open in the form of fossils of plants and animals and canopy should be employed during the winter to organic matter that contributed to the deposit. minimize damage to the peat and impacts to the Fossilized humans have even been found in non- hydrologic regime. decomposing bog peat: the bog people of Scandinavia where preserved for approximately

Grayling Community Assessment - 103 2000 years (Mitsch and Gosselink 2000). Few-flower Sedge – Tussock Cottongrass / Stratigraphical analysis of peat cores provides Peatmoss Species Herbaceous Vegetation; insights into past climatic change and associated Open Graminoid / Sphagnum Bog vegetation change, floristic distribution, the development of wetland ecosystems, and the NatureServe Ecological Systems Classification: successional pathways of peatlands (Heinselman CES103.581: Boreal-Laurentian Bog 1963, Glaser et al. 1981, Miller 1981, Glaser and Janssens 1986, Riley 1989, Gignac et al. 2000). Related Abstracts: American bittern, black- backed woodpecker, Blanchard’s cricket frog, Similar Communities: coastal plain marsh, Blanding’s turtle, coastal plain marsh, eastern box intermittent wetland, inundated shrub swamp, turtle, eastern massasauga, eastern prairie fringed muskeg, northern fen, patterned fen, poor conifer orchid, English sundew, great blue heron rookery, swamp, poor fen, prairie fen, relict conifer incurvate emerald, intermittent wetland, merlin, swamp, rich conifer swamp. northern fen, northern harrier, poor fen, prairie fen, relict conifer swamp, rich conifer swamp, Other Classifications: round-leaved orchis, secretive locust, spotted Michigan Natural Features Inventory Circa 1800 turtle, yellow pitcher plant, and yellow rail. Vegetation (MNFI): Bog (6121) Literature Cited: Michigan Department of Natural Resources Albert, D.A. 1995. Regional landscape (MDNR): D-treed bog, V-bog ecosystems of Michigan, Minnesota, and Wisconsin: A working map and classification. Michigan Resource Information Systems Gen. Tech. Rep. NC-178. St. Paul, MN: U.S. (MIRIS): 62 (non-forested wetland) and 622 Department of Agriculture, Forest Service, (emergent wetland) North Central Forest Experiment Station. Northern Prairie Wildlife Research Center The Nature Conservancy National Classification: Home Page. CODE; ALLIANCE; ASSOCIATION; http://www.npwrc.usgs.gov/resource/1998/rla COMMON NAME ndscp/rlandscp.htm. (Version 03JUN98.) 250 pp. IV.A.1.N.g; Chamaedaphne calyculata Saturated Dwarf-Shrubland Alliance; Albert, D.A., S.R. Denton, and B.V. Barnes. 1986. Chamaedaphne calyculata / Carex Regional landscape ecosystems of Michigan. oligosperma –Eriophorum virginicum Dwarf- Ann Arbor, MI: University of Michigan, shrubland; Leatherleaf / Few-seed Sedge – School of Natural Resources. 32 pp. & map. Tawny Cottongrass Dwarf Shrubland; Leatherleaf Kettle Bog Almendinger, J.C., J.E. Almendinger, and P.H. Glaser. 1986. Topographic fluctuations across IV.A.1.N.g; Chamaedaphne calyculata a spring fen and raised bog in the Lost River Saturated Dwarf-Shrubland Alliance; Peatland, northern Minnesota. Journal of Chamaedaphne calyculata – Ledum Ecology 74(2): 393-401. groenlandicum – Kalmia polifolia Bog Dwarf-shrubland; Leatherleaf – Labrador-tea Almendinger, J.A., and J.H. Leete. 1998. –Bog Laurel Bog Dwarf-Shrubland; Regional and local hydrogeology of Leatherleaf Bog calcareous fens in the Minnesota River Basin, USA. Wetlands 18(2): 184-202. V.A.5.N.m; Carex oligosperma – Carex lasiocarpa Saturated Herbaceuos Alliance; Amon, J.P., C.A. Thompson, Q.J. Carpenter, and Carex oligosperma – Carex pauciflora – J. Mines. 2002. Temperate zone fens of the Eriophorum vaginatum / Sphagnum spp. glaciated Midwestern USA. Wetlands 22(2): Herbaceous Vegetation; Few-seed Sedge – 301-317.

Grayling Community Assessment - 104 Anderson, D.S, R. B. Davis, S.C. Rooney, C.S. economy and ecology. Frontiers in Ecology Campbell. 1996. The ecology of sedges and the Environment 1(10): 525-532. (Cyperaceae) in Maine peatlands. Bulletin of the Torrey Botanical Club 123(2): 100-110. Coburn, H., D. Dean, and G.M. Grant. 1933. An ecological study of Bryant’s Bog, Cheboygan Barnes, B.V. 1991. Deciduous forest of North County. Michigan. Paper’s of the Michigan America. Pp 219-344 in E. Röhrig and B. Academy of Science, Arts, and Letters 17: 57- Ulrich, eds., Temperate Deciduous Forests. 65. Elsevier, Amsterdam. 635 pp. Crum, H. 1983. Mosses of the Great Lakes Forest. Barnes, B.V., K.S. Pregitzer, T.A. Spies, and V. University of Michigan, Ann Arbor, MI. 417 H. Spooner. 1982. Ecological forest site pp. classification. Journal of Forestry 80(8): 493- 498. Curtis, J.T. 1959. Vegetation of Wisconsin: An Ordination of Plant Communities. University Bay, R.R. 1967. Ground water and vegetation in of Wisconsin Press, Madison, WI. 657 pp. two peat bogs in northern Minnesota. Ecology 48(2): 308-310. Damman, A.H. 1990. Nutrient status of ombrotrophic peat bogs. Aquilo Series Bedford, B.L., and K.S. Godwin. 2003. Fens of Botanica 28: 5-14. the United States: Distribution, characteristics, and scientific connection Dansereau, P., and F. Segadas-Vianna. 1952. versus legal isolation. Wetlands 23(3): 608- Ecological study of the peat bogs of eastern 629. North America. I. Structure and evolution of vegetation. Canadian Journal of Botany 30: Bedford, B.L., M.R. Walbridge, and A. Aldous. 490-520. 1999. Patterns in nutrient availability and plant diversity of temperate North American Dean, D., and H. Coburn. 1927. An ecological wetlands. Ecology 80(7): 2151-2169. study of Linne Bog, Cheboygan County, Michigan with special reference to Boelter, D.H., and E.S. Verry. 1977. Peatland and Nemopanthus mucranata (L.) Trelease. water in the northern Lake States. North Paper’s of the Michigan Academy of Science, Central Forest Experiment Station. USDA Arts, and Letters 8: 87-96. Forest Service General Technical Report NC- 31. 26 pp. Eggers, S.D., and D.M. Reed. 1997. Wetland plants and plant communities of Minnesota Braun, E.L. 1950. Deciduous forests of eastern and Wisconsin. U.S. Army Corps of North America. Hafner Press, New York, NY. Engineers, St Paul, Minnesota. 263 pp. 596 pp. Faber-Langendoen, D., ed., 2001. Plant Burns, G.P. 1906. Bog studies. Field Studies in communities of the Midwest: Classification in Botany. University Bulletin, New Series, an ecological context. Association for 7(14): 3-13. University of Michigan, Ann Biodiversity Information, Arlington, VA. 61 Arbor. pp & appendix (705 pp.).

Chapman, S., A. Buttler, A.-J. Francez, F. Fitzgerald, S., and R.E. Bailey. 1975. Laggoun-Defarge, H. Vasander, M. Schloter, Vegetational characteristics of a circum- J. Combe, P. Grosvernier, H. Harms, D. neutral bog, Barney’s Lake, Beaver Island, Epron, D. Gilbert, and E. Mitchell. 2003. Michigan. The Michigan Academician 7(4): Exploitation of northern peatlands and 477-488. biodiversity maintenance: A conflict between

Grayling Community Assessment - 105 Frelich, L.E., and C.G. Lorimer. 1991. Natural Heinselman, M.L. 1970. Landscape evolution, disturbance regimes in hemlock-hardwood peatland types, and the environment in the forests of the Upper Great Lakes region. Lake Agassiz Peatland Natural Area, Ecological Monographs 61(2): 145-164. Minnesota. Ecological Monographs 40(2): 235-261. Gates, F.C. 1942. The bogs of northern Lower Michigan. Ecological Monographs 12(3): Hoffman, R. 2002. Wisconsin’s Natural 213-254. Communities. How to recognize them, where to find them. The University of Wisconsin Gignac, L.D., L.A. Halsey, and D.H. Vitt. 2000. A Press, Madison, WI. 375 pp. bioclimatic model for the distribution of Sphagnum-dominated peatlands in North Jean, M., and A. Bouchard. 1991. Temporal America under present climatic conditions. changes in wetland landscapes of a section of Journal of Biogeography 27(5): 1139-1151. the St. Lawrence River, Canada. Environmental Management 15(2): 241-250. Glaser, P.H. 1992. Raised bogs in eastern North America – Regional controls for species Kost, M.A. 2001. Natural community abstract for richness and floristic assemblages. Journal of relict conifer swamp. Michigan Natural Ecology 80: 535-554. Features Inventory, Lansing, MI. 6 pp.

Glaser, P.H., and J.A. Janssens. 1986. Raised bogs Lindeman, R.L. 1941. The developmental history in eastern North America: Transitions in of Cedar Creek Bog, Minnesota. American landforms and gross stratigraphy. Canadian Midland Naturalist 25(1): 101-112. Journal of Botany 64: 395-415. Michigan Natural Features Inventory. 2003. Draft Glaser, P.H., G.A. Wheeler, E. Gorham, and H.E. description of Michigan natural community Wright, Jr. 1981. The patterned mires of the types. (Unpublished manuscript revised Red Lake Peatland, northern Minnesota: March 4, 2003.) Michigan Natural Features Vegetation, water chemistry and landforms. Inventory, Lansing, MI. 36 pp. Available: Journal of Ecology 69(2): 575-599. http://www.msue.msu.edu/mnfi/lists/natural_c ommunity_types.pdf. Glaser, P.H., J.A. Janssens, and D.I. Siegel. 1990. The response of vegetation to chemical and Mitsch, W.J., and J.G. Gosselink. 2000. Wetlands. hydrological gradients in the Lost River John Wiley and Sons, Inc, New York, NY. Peatland, northern Minnesota. Journal of 920 pp. Ecology 78(4): 1021-1048. Miller, N. 1981. Bogs, bales, and BTU’s: A Hammerson, G. 1994. Beaver (Castor primer on peat. Horticulture 59: 38-45. canadensis): Ecosystem alterations, management, and monitoring. Natural Areas NatureServe. 2005a. NatureServe Explorer: An Journal 14(1): 44-57. online encyclopedia of life [web application]. Version 4.2. NatureServe, Arlington, Virginia. Heinselman, M.L. 1963. Forest sites, bog Available: processes, and peatland types in the Glacial http://www.natureserve.org/explorer. Lake Region, Minnesota. Ecological (Accessed: March 03, 2005.) Monographs 33(4): 327-374. NatureServe. 2005b. International Ecological Heinselman, M.L. 1965. String bogs and other Classification Standard: Terrestrial Ecological patterned organic terrain near Seney, Upper Classifications. NatureServe Central Michigan. Ecology 46: 185-188. Databases. Arlington, VA. U.S.A. Data current as of January 13, 2005.

Grayling Community Assessment - 106 Osvald, H. 1935. A bog at Hartford, Michigan. Swain, A.M. 1973. A history of fire and Ecology 16(3): 520-528. vegetation in northeastern Minnesota as recorded in lake sediments. Quaternary Richardson, C.J., and P.E. Marshall. 1986. Research 3: 383-396. Processes controlling movement, storage, and export of phosphorous in a fen peatland. Swanson, D.K., and D.F. Grigal. 1989. Vegetation Ecological Monographs 56(4): 279-302. indicators of organic soil properties in Minnesota. Soil Science Society of America Riley, J.L. 1989. Southern Ontario bogs and fens Journal 53: 491-495. of the Canadian Shield. Wetlands: Inertia or Momentum: 355-367. Swinehart, A.L., and G.R. Parker. 2000. Palaeoecology and development of peatlands Schwintzer, C.R. 1978a. Nutrient and water levels in Indiana. American Midland Naturalist in a small Michigan bog with high tree 143(2): 267-297. mortality. American Midland Naturalist 100(2): 441-451. Verry, E.S. 1975. Streamflow chemistry and nutrient yields from upland-peatland Schwintzer, C.R. 1978b. Vegetation and nutrient watersheds in Minnesota. Ecology 65(5): status of northern Michigan fens. Canadian 1149-1157. Journal of Botany 56: 3044-3051. Vitt, D.H., and N.G. Slack. 1975. An analysis of Schwintzer, C.R. 1979. Vegetation changes the vegetation of Sphagnum-dominated kettle- following a water level rise and tree mortality hole bogs in relation to environmental in a Michigan bog. The Michigan Botanist 18: gradients. Canadian Journal of Botany 53: 91-98. 332-359. Vitt, D.H., H. Crum, and J.A. Snider. 1975. The Schwintzer, C.R. 1981. Vegetation and nutrient vertical zonation of Sphagnum species in status of northern Michigan bogs and conifer hummock-hollow complexes in northern swamps with a comparison to fens. Canadian Michigan. The Michigan Botanist 14(4): 190- Journal of Botany 59: 842-853. 200.

Schwintzer, C.R, and G. Williams. 1974. Wheeler, G.A., P.H. Glaser, E. Gorham, C.M. Vegetation changes in a small Michigan bog Wetmore, F.D. Bowers, and J.A. Janssens. from 1917 to 1972. American Midland 1983. Contributions to the flora of the Red Naturalist 92(2): 447-459. Lake Peatland, northern Minnesota, with special attention to Carex. American Midland Schwintzer, C.R, and T.J. Tomberlin. 1982. Naturalist 110(1): 62-96. Chemical and physical characteristics of shallow ground waters in northern Michigan bogs, fens, and swamp. American Journal of Abstract Citation: Botany 69(8): 1231-1239. Cohen, J.G. 2005. Natural community abstract for bog. Michigan Natural Features Inventory, Siegel, D.I. 1988. Evaluating cumulative effects Lansing, MI. 13 pp. of disturbance on the hydrologic function of bogs, fens, and mires. Environmental

Management 12(5): 621-626.

Siegel, D.I., and P.H. Glaser. 1987. Groundwater flow in a bog-fen complex, Lost River Peatland, northern Minnesota. Journal of Ecology 75(3): 743-754.

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Grayling Community Assessment - 108 Appendix 6. Northern Wet Meadow

Overview: An open, groundwater-influenced, However, in Wisconsin, where 459,000 ha sedge and grass–dominated wetland that occurs in (1,130,000 acres) of sedge meadow are thought to northern Lower Michigan and Upper Michigan have existed circa 1800 (Curtis 1959), it is and typically borders streams but is also found on estimated that less than one percent remain intact pond and lake margins and above beaver dams. (Reuter 1986). It is likely northern wet meadow Soils are nearly always on muck soil, which is acreage has declined similarly in other Midwest strongly acid to slightly acid. Open conditions states, such as Michigan, where similar maintained by seasonal flooding, beaver-induced agricultural methods have been practiced. flooding, and/or fire. Northern wet meadows have fared slightly better than southern wet meadows because agriculture Global and State Rank: G4G5/S4 and development has been less extensive north of the climatic tension zone (Hoffman 2002). Range: Northern wet meadow, which is commonly referred to as sedge meadow, occurs in Northern wet meadows have been extensively Michigan, Minnesota, North Dakota, Wisconsin, utilized for agriculture. Prior to the 1950s mowing and Ontario. In Michigan, northern wet meadow for marsh hay was widely practiced (Stout 1914, is thought to occur in the northern half of the Curtis 1959, Eggers and Reed 1997). Wet Lower Peninsula above the climatic transition meadows were frequently tiled, ditched, drained, zone and throughout the Upper Peninsula and to and converted to pasture and row crops or mined differ from sedge meadows in southern Michigan for peat or muck (Costello 1936, Curtis 1959, (Kost 2001). However, no detailed study of the Reuter 1986, Eggers and Reed 1997). The differences between northern and southern types hydrology of these systems is threatened by the has been undertaken in Michigan. Curtis (1959) reduction and diversion of surface runoff, studied sedge meadows in northern and southern channelization of stream flow, lowering local Wisconsin and found them to be floristically groundwater discharge, and deterioration of water similar but concluded that northern meadows had quality (Reuter 1986). Lowering of the water table consistently lower soil pH values and were has caused the conversion of many sedge frequently wetter and smaller than many southern meadows to shrub thickets (Curtis 1959, Reuter wet meadows. Southern wet meadow is 1986, Eggers and Reed 1997). In addition, fire dominated primarily by Carex stricta (tussock suppression has allowed shrub encroachment with sedge) while northern wet meadows can be many sedge meadows converting to shrub thicket dominated or co-dominated by several additional within ten to twenty years since the interruption of sedges and/or Calamagrostis canadensis (blue- fire disturbance (Curtis 1959, White 1965, Davis joint grass) (Faber-Langendoen 2001, Michigan 1979, Reuter 1986, Jean and Bouchard 1991). Natural Features Inventory 2003, NatureServe This is especially evident where the water table 2005a). Another sedge-dominated natural has been lowered through tiling or ditching and community, poor fen, also occurs in Michigan but where the practice of mowing for marsh hay has differs markedly from sedge meadow because of been abandoned (White 1965, Eggers and Reed its acidic, organic soils, and the prevalence of 1997). Alteration of the fire and hydrologic Carex oligosperma (few-seed sedge) and other regimes has allowed for the invasion of sedge open bog species (Michigan Natural Features meadows by pernicious exotic species, especially Inventory 2003). Lythrum salicaria (purple loosestrife), Rhamnus cathartica (common buckthorn), and Rhamnus Rank Justification: Because northern wet frangula (glossy buckthorn) (Reuter 1986). meadow often occurs as a zone within large wetland complexes, information on its historical Landscape and Abiotic Context: Northern wet extent and present acreage is not readily available. meadow occurs on glacial lakebeds, in channels of

Grayling Community Assessment - 109 glacial outwash, and in depressions on glacial annual temperature is 7 °C (45 °F). The mean outwash and moraines (Curtis 1959, Reuter 1986, number of freeze-free days is between 90 and 160, Faber-Langendoen 2001, Hoffman 2002, and the average number of days per year with NatureServe 2005a, NatureServe 2005b). The snow cover of 2.5 cm or more is between 80 and community frequently occurs along the margins of 140. The normal annual total precipitation ranges lakes, ponds, and streams where seasonal flooding from 740 to 900 mm with a mean of 823 mm or beaver-induced flooding is common (Curtis (Albert et al. 1986, Barnes 1991, Albert 1995). 1959, Reuter 1986, Hoffman 2002). Northern wet meadows are characterized by local climates with lower temperatures and evaporation Wet meadow typically occurs on organic soils rates and shorter growing seasons than the such as muck and well-decomposed sapric peat surrounding uplands (Curtis 1959). (Curtis 1959) but saturated mineral soil may also support the community (Costello 1936, Curtis Natural Processes: Northern wet meadow is a 1959, Faber-Langendoen 2001, NatureServe groundwater-dependent, graminoid-dominated, 2005a). Because of the calcareous nature of the wetland community. Water levels in northern wet glacial drift in the regions occupied by wet meadow fluctuate seasonally, reaching their peak meadow, its wet soils can contain high levels of in spring and lows in late summer (Costello 1936, dissolved minerals such as calcium and Warners 1993). However, water levels typically magnesium. Northern wet meadow occurs on remain at or near the soil’s surface throughout the more acidic soils compared to southern wet year (Costello 1936, Curtis 1959, Warners 1993, meadow which is found on neutral to strongly Eggers and Reed 1997). The community’s alkaline soils (Costello 1939, Curtis 1959, structure may depend on maintaining a Warners 1993). Northern wet meadow soils range consistently high water table. Costello (1936) from strongly acid to slightly acid. states that the Carex stricta tussocks disappeared within 10 years from a meadow where the water Sedge meadows are found adjacent to other levels were reduced to two to four feet below the wetland communities, often in large wetland surface as a result of tiling. In addition to seasonal complexes. Northern shrub thicket and swamp flooding, beaver-induced flooding may also play forest are typically adjacent to northern wet an important role in maintaining the community meadows that occur along streams (Curtis 1959). by occasionally raising water levels and killing On the edges of inland lakes, northern wet encroaching trees and shrubs. Beaver may also meadow often borders emergent marsh and less help create new northern wet meadows by frequently northern fen. It may also occur along flooding swamp forests and northern shrub the Great Lakes shoreline within extensive areas thickets and thus creating suitable habitat for the of Great Lakes marsh. growth of shade-intolerant wet meadow species.

The Michigan range of northern wet meadow falls Evidence from wetland peat cores and circa 1800 within the area classified by Braun (1950) as the vegetation maps indicate that wet meadow is a Northern Hardwood-Conifer Region fire-dependent natural community (Curtis 1959, (Hemlock/White Pine/Northern Hardwoods Davis 1979). Analysis of wetland peat cores Region) and within the following regions shows that charcoal fragments are consistently classified by Albert et al. (1986): Region II, associated with sedge and grass pollen (Davis Northern Lower Michigan; Region III, Eastern 1979). Conversely, charcoal fragments are lacking Upper Michigan; and Region IV, Western Upper from sections of peat cores dominated by shrub Michigan. The Northern Hardwood-Conifer pollen. Fires typically occur in sedge meadows Region has a cool snow-forest climate with short, during dry conditions of early spring or late fall warm summers, cold winters and a large number (White 1965). By reducing leaf litter and allowing of cloudy days. The daily maximum temperature light to reach the soil surface and stimulate seed in July ranges from 24 to 29 °C (75 to 85 °F), the germination, fire can play an important role in daily minimum temperature in January ranges maintaining wet meadow seed banks (Warners from -21 to -9 °C (-5 to 15 °F) and the mean 1997, Kost and De Steven 2000). Fire also plays a

Grayling Community Assessment - 110 critical role in preventing declines in species rostrata, and C. vesicaria can often occur as richness in many community types by creating dominants or co-dominants. micro-niches for small species (Leach and Givnish 1996). Another critically important The most dominant grass species in northern wet attribute of fire for maintaining open sedge meadow is Calamagrostis canadensis (blue-joint meadow is its ability to temporarily reduce shrub grass) (Stout 1914, Kost and De Steven 2000). cover (White 1965, Reuter 1986, Hoffman 2002). Other common grasses include: Bromus ciliatus In the absence of fire or flooding, all but the (fringed brome), Glyceria canadensis (manna wettest sedge meadows typically convert to shrub grass), G. striata (fowl manna grass), thicket and eventually swamp forest (Curtis Muhlenbergia glomerata (marsh wild timothy), 1959). Because many of the species that inhabit Muhlenbergia mexicana (leafy satin grass), and wet meadow are shade-intolerant, species richness Poa palustris (fowl meadow grass). Spike-rushes usually declines following shrub and tree invasion (i.e., Eleocharis erythropoda), cat-tails (Typha (Curtis 1959, White 1965, Reuter 1986). latifolia and Typha angustifolia), Cladium marisicoides (twig-rush), and Scirpus atrovirens Vegetation Description: Northern wet meadow is (green bulrush) are also common graminoids. a sedge dominated system that typically has 100% Sedge meadows disturbed by agricultural use, vegetative cover in the ground layer (Curtis 1959, grazing, drainage, and/or filling are frequently Eggers and Reed 1997). Sedge meadow is often dominated by Phalaris arundinaceae (reed canary dominated by Carex stricta (tussock sedge) (Stout grass) and extremely aggressive grass which 1914, Costello 1936, Curtis 1959, Warners 1997, forms persistent, monotypic stands (Eggers and Kost and De Steven 2000). Because the roots of Reed 1997). Carex stricta form large hummocks or tussocks, the species is often responsible for the A wide variety of wetland forbs occur scattered in community’s hummock and hollow structure. northern wet meadow. Due to the high moisture Individual culms of Carex stricta grow from the conditions during the spring, many of the forbs tussocks, which may reach more than one meter in bloom in the summer and fall (Curtis 1959, height and half a meter in diameter and live for Hoffman 2002). The following are some of the more than 50 years (Costello 1936). The Carex more common species: Anemone canadensis stricta tussocks can occur at very high densities (1 (Canada anemone), Asclepias incarnata (swamp to 4 per m2) and occupy more than 40% of a milkweed), Aster lanceolatus (panicled aster), A. meadow’s area (Costello 1936). Because the lateriflorus (calico aster), A. puniceus (swamp shaded areas between tussocks are often covered aster), A. umbellatus (flat-topped aster), with standing water and leaf litter, many of the Campanula aparinoides (marsh bellflower), shorter species inhabiting sedge meadows grow Cicuta bulbifera (water-hemlock), C. maculata almost exclusively from the sides or tops of Carex (water-hemlock), Cirsium muticum (swamp stricta tussocks. thistle), Epilobium strictum (downy willowherb), Eupatorium maculatum (Joe-pye-weed), Other sedges that commonly occur in northern Eupatorium perfoliatum (boneset), Euthamia wet meadow include: Carex aquatilis (water graminifolia (flat-topped goldenrod), Galium sedge), C. bebbii (Bebb’s sedge), C. buxbaumii asprellum (rough bedstraw), G. trifidum (Buxbaum’s sedge), C. comosa (long-hair sedge), (threepetal bedstraw), Impatiens capensis (spotted C. hystericina (bottlebrush sedge), C. lacustris touch-me-not), Iris versicolor (wild blue flag), (lake or hairy sedge), C. lanuginose (woolly Lathyrus palustris (marsh pea), Lycopus sedge), C. lasiocarpa (wiregrass sedge), C. americanus (American water-horehound), L. rostrata (beaked sedge), C. stipata (saw-beak uniflorus (northern bugleweed), Lysimachia sedge), C. vesicaria (blister sedge), and C. thyrsifolia (tufted loosestrife), Mentha arvensis vulpinoidea (fox sedge). Although most of the (wild mint), Polygonum amphibium (water associated sedge species tend to be randomly smartweed), Rumex orbiculatus (greater water interspersed, Carex lacustris, C. lasiocarpa, C. dock), Sagittaria latifolia (duck-potato or common arrowhead), Scutellaria galericulata

Grayling Community Assessment - 111 (marsh skullcap), Solidago canadensis (Canada idas nabokovi (northern blue butterfly, state goldenrod), S. gigantea (late goldenrod), S. patula threatened). (rough-leaved goldenrod or swamp goldenrod), Thalictrum dasycarpum (purple meadow-rue), Northern wet meadow provide habitat for Triadenum fraseri (marsh St. John’s-wort), numerous herptiles such as Clemmys guttata Verbena hastata (blue vervain), and Viola (spotted turtle, state threatened), Emys blandingii cucullata (marsh violet). Characteristic fern or (Blanding’s turtle, state special concern), fern allies include Dryopteris cristata (crested Glyptemys insculpta (wood turtle, state special woodfern), Equisetum arvense (field horsetail), E. concern), Pseudacris triseriata maculata (boreal fluviatile (water horsetail), Onoclea sensibilis chorus frog, state special concern), and Sistrurus (sensitive fern), and Thelypteris palustris (marsh catenatus catenatus (eastern massasauga, state fern). special concern, federal candidate species). The late blooming composites found in sedge Northern wet meadow can also contain numerous, meadows provide an important food source for scattered shrub and tree species. Shrub and tree songbirds while the hummock provide excellent encroachment is especially pronounced in sites nesting habitat for wetlands birds (Eggers and that have altered flooding or fire regimes. Reed 1997). Rare birds that utilize these wetlands Prevalent shrubs include Alnus rugosa (speckled include Asio flammeus (short-eared owl, state alder), Betula pumila (bog birch), Cornus endangered), Botaurus lentiginosus (American stolonifera (red-osier dogwood), Potentilla bittern, state special concern), Chlidonias niger fruticosa (shrubby cinquefoil), P. palustris (marsh (black tern, state special concern), Circus cyaneus cinquefoil), Salix spp. (willows), Spirea alba (northern harrier, state special concern), (meadowsweet), and S tomentosa (hardhack). Cistothorus palustris (marsh wren, state special Scattered trees and tree saplings are often found concern), Coturnicops noveboracensis (yellow invading northern wet meadow. Typical tree rail, state threatened), Gallinula chloropus species include Acer rubrum (red maple), (common moorhen, state special concern), Fraxinus nigra (black ash), Larix laricina Ixobrychus exilis (least bittern, state threatened), (tamarack), Populus balsamifera (balsam poplar), Phalaropus tricolor (Wilson's phalarope, state Populus tremuloides (quaking aspen), and Thuja special concern), Rallus elegans (king rail, state occidentalis (northern white cedar). (Above endangered), and Sterna forsteri (Forster's tern, species lists compiled from Michigan Natural state special concern). Alces alces (moose, state Features Inventory database, Curtis 1959, Reuter threatened), Canis lupus (gray wolf, state 1986, Eggers and Reed 1997, Hoffman 2002, threatened), and Lynx canadensis (lynx, state Faber-Langendoen 2001, NatureServe 2005a) endangered) utilize sedge meadow habitat.

Michigan Indicator Species: blue-joint grass, Conservation and Biodiversity Management: Carex lacustris, Carex stricta, common boneset, Northern wet meadows contribute significantly to greater water dock, Joe-pye-weed, marsh the overall biodiversity of northern Michigan by bellflower, northern bugleweed, swamp aster, and providing habitat to a wide variety of plant and tufted loosestrife. animal species including numerous rare species. In addition, sedge meadows provide ecosystem Other Noteworthy Species: Several rare plants services, protecting water quality by assimilating can be found in northern wet meadow and nutrients, trapping sediment, and retaining associated open wetlands including Cacalia stormwater and floodwater (Eggers and Reed plantaginea (Indian plantain, state special 1997). Protecting the hydrology of northern wet concern), Carex wiegandii (Wiegand’s sedge, meadow is imperative for the community’s state threatened), Gentiana linearis (linear-leaved continued existence. This may include avoiding gentian, state threatened), Parnassia palustris surface water inputs to meadows from drainage (marsh-grass-of-Parnassus, state threatened), and ditches, agricultural fields, road construction, and Vaccinium cespitosum (dwarf bilberry, state logging in the adjacent uplands, and protecting threatened), which is the host plant for Lycaeides groundwater recharge areas by maintaining native

Grayling Community Assessment - 112 vegetation types in the uplands around the community structure and dramatically reducing community. Resource mangers operating in species richness. Management should strive to uplands adjacent to sedge meadows should take prevent the further spread of these invasive care to minimize the impacts of management to species and implement control measures when hydrologic regimes, especially increased surface possible. Establishment of invasive species can be flow. This can be accomplished by establishing a prevented by maintaining the hydrologic and fire no-cut buffer around wet meadows and avoiding disturbance regimes and avoiding grazing (Reuter road construction and complete canopy removal in 1986). stands immediately adjacent to wetlands. Restoration of degraded northern wet meadows Management for wet meadow should include the depends on the occurrence of water-saturated peat use of prescribed fire (Curtis 1959, White 1965). and muck soils, maintaining waters levels very Prescribed fire can help reduce litter, stimulate near the soil surface throughout the year, seed germination, promote seedling establishment, providing protection from shrub encroachment and bolster grass, sedge, and perennial and annual and invasive species, and the availability of forb cover (Bowles et al. 1996, Warners 1997, appropriate seed stock (Reuter 1986). Finding Kost and De Steven 2000). While prescribed fire viable seed for sedges, the plant group responsible can be an important tool for rejuvenating wet for the overall structure of wet meadow, may be a meadow seed banks, it can also help ensure that difficult task. Costello (1936) reports that in more the community remains in an open condition by than six years of studying Carex stricta- temporarily setting back invading woody species dominated sedge meadows he did not find a single (Reuter 1986). Using prescribed fire to control seedling of the species. Because of the difficulty shrub invasion in sedge meadows has also been of restoring wet meadow in the absence of shown to be 85% less expensive to implement favorable hydrology and intact organic soils, than manual cutting (Reuter 1986). The use of conservation efforts should focus on protecting prescribed fire should be avoided during periods the remaining community occurrences (Reuter of drought to avoid igniting the community’s 1986). organic soils (Curtis 1959, Vogl 1969). Burning in the early spring while the soil moisture is high Research Needs: Further work on community reduces the chances of destroying the organic classification is needed to elucidate differences soils (Reuter 1986), however, growing season among sedge meadow types both within and burns can be more effective at reducing among ecoregions (Reuter 1986). More studies aggressive woody vegetation (Bowles et al. 1996). need to focus on the flooding and fire regimes of If prescribed burning is not feasible, mowing can northern wet meadow and the interaction of be used to simulate fire disturbance but should be disturbance factors. As noted by Hammerson restricted to the winter, when ground frost will (1994), beaver significantly alter the ecosystems reduce disturbance to soils, plants, and hydrology, they occupy. An important research question to or late summer and fall when the meadows are dry examine is how the wetland ecosystems of the (White 1965, Reuter 1986). In situations where Great Lakes have been and continue to be affected shrub encroachment is severe, resource managers by the fluctuating beaver population. may need to cut invading shrubs and herbicide the Experimentation is needed to determine how best cut stumps, especially if the shrubs are stump to prevent shrub encroachment of open wetlands sprouters, like Alnus rugosa (speckled alder) or that are threatened by conversion to northern Rhamnus frangula (glossy buckthorn) (Heidorn shrub thicket (Reuter 1986). In addition, scientists 1991). should gain an understanding of plant and animal community responses to the frequency and Invasive species that can occur in northern wet seasonal timing of prescribed burning and meadow include glossy buckthorn, common anthropogenic flooding. Effects of management buckthorn, purple loosestrife, reed canary grass, within northern wet meadow need to be monitored and Phragmites australis (reed). Each of these to allow for assessment and refinement. Research species is capable of significantly altering is needed on plant and animal community

Grayling Community Assessment - 113 responses to the frequency and seasonal timing of Herbaceous Vegetation; Northern Sedge Wet prescribed burning and flooding. Conservation Meadow. and management of northern wet meadow will be stimulated by research on the importance of the V.A.5.N.k; Carex stricta Seasonally Flooded community for maintaining rare species and Herbaceous Alliance; Carex stricta – Carex regional biodiversity. spp. Herbaceous Vegetation; Tussock Sedge – Sedge Species Herbaceous Vegetation; Similar Communities: emergent marsh, Great Tussock Sedge Wet Meadow. Lakes marsh, intermittent wetland, southern wet meadow, northern fen, northern shrub thicket, NatureServe Ecological Systems Classification: northern wet-mesic prairie, poor fen, and wet CES201.582: Laurentian-Acadian Wet Meadow- prairie. Shrub Swamp

Other Classifications: CES202.701: North-Central Interior Wet Michigan Natural Feature Inventory Circa 1800 Meadow-Shrub Swamp Vegetation (MNFI): wet meadow (6224) Related Abstracts: American bittern, black tern, Michigan Department of Natural Resources Blanding’s turtle, dwarf bilberry, eastern (MDNR): L-lowland brush, N-marsh, V-bog massasauga, floodplain forest, Forster’s tern, Great Lakes marsh, Indian plantain, intermittent Michigan Resource Information Systems wetland, king rail, least bittern, marsh-grass-of- (MIRIS): 622 (emergent wetland) Parnassus, northern blue butterfly, northern fen, northern harrier, northern shrub thicket, poor fen, The Nature Conservancy National Classification: short-eared owl, southern wet meadow, spotted CODE; ALLIANCE; ASSOCIATION; turtle, wood turtle, and yellow rail. COMMON NAME Literature Cited: V.A.5.N.k; Calamagrostis canadensis Albert, D.A. 1995. Regional landscape Seasonally Flooded Herbaceous Alliance; ecosystems of Michigan, Minnesota, and Calamagrostis canadensis – Phalaris Wisconsin: A working map and classification. arundinacea Herbaceous Vegetation; Gen. Tech. Rep. NC-178. St. Paul, MN: U.S. Bluejoint – Reed Canary Grass Herbaceous Department of Agriculture, Forest Service, Vegetation; Bluejoint Wet Meadow. North Central Forest Experiment Station. Northern Prairie Wildlife Research Center V.A.5.N.k; Carex aquatilis Seasonally Home Page. Flooded Herbaceous Alliance; Carex aquatilis http://www.npwrc.usgs.gov/resource/1998/rla – Carex spp. Herbaceous Vegetation; Aquatic ndscp/rlandscp.htm. (Version 03JUN98.) 250 Sedge – Sedge Species Herbaceous pp. Vegetation; Water Sedge Wet Meadow. Albert, D.A., S.R. Denton, and B.V. Barnes. 1986. V.A.5.N.k; Carex lacustris Seasonally Regional landscape ecosystems of Michigan. Flooded Herbaceous Alliance; Carex lacustris Ann Arbor, MI: University of Michigan, Herbaceous Vegetation; Lake Sedge School of Natural Resources. 32 pp. & map. Herbaceous Vegetation; Lake Sedge Wet Meadow. Barnes, B.V. 1991. Deciduous forest of North America. Pp 219-344 in E. Röhrig and B. V.A.5.N.k; Carex (rostrata, urticulata) Ulrich, eds., Temperate Deciduous Forests. Seasonally Flooded Herbaceous Alliance; Elsevier, Amsterdam. 635 pp. Carex rostrata – Carex lacustris – (Carex vesicaria) Herbaceous Vegetation; Swollen- Beak Sedge – Lake Sedge – (Inflated Sedge)

Grayling Community Assessment - 114 Braun, E.L. 1950. Deciduous forests of eastern Jean, M., and A. Bouchard. 1991. Temporal North America. Hafner Press, New York, NY. changes in wetland landscapes of a section of 596 pp. the St. Lawrence River, Canada. Environmental Management 15(2): 241-250. Bowles, M., J. McBride, N. Stynoff, and K. Johnson. 1996. Temporal changes in Kost, M.A. 2001. Natural community abstract for vegetation composition and structure in a fire- southern wet meadow. Michigan Natural managed prairie fen. Natural Areas Journal 16: Features Inventory, Lansing, MI. 5 pp. 275-278. Kost, M.A. and D. De Steven. 2000. Plant Costello, D. F. 1936. Tussock meadows in community responses to prescribed burning in southeastern Wisconsin. Botanical Gazette 97: Wisconsin sedge meadows. Natural Areas 610-48. Journal 20: 36-49.

Curtis, J.T. 1959. Vegetation of Wisconsin: An Michigan Natural Features Inventory. 2003. Draft Ordination of Plant Communities. University description of Michigan natural community of Wisconsin Press, Madison, WI. 657 pp. types. (Unpublished manuscript revised March 4, 2003.) Michigan Natural Features Davis, A. M. 1979. Wetland succession, fire and Inventory, Lansing, MI. 36 pp. Available: the pollen record: A Midwestern example. The http://www.msue.msu.edu/mnfi/lists/natural_c American Midland Naturalist 102: 86-94. ommunity_types.pdf.

Eggers, S.D., and D.M. Reed. 1997. Wetland NatureServe. 2005a. NatureServe Explorer: An plants and plant communities of Minnesota online encyclopedia of life [web application]. and Wisconsin. U.S. Army Corps of Version 4.2. NatureServe, Arlington, Virginia. Engineers, St Paul, Minnesota. 263 pp. Available: http://www.natureserve.org/explorer. Faber-Langendoen, D. (editor). 2001. Plant (Accessed: March 03, 2005.) communities of the Midwest: Classification in an ecological context. Association for NatureServe. 2005b. International Ecological Biodiversity Information, Arlington, VA. 61 Classification Standard: Terrestrial Ecological pp. + appendix (705 pp.). Classifications. NatureServe Central Databases. Arlington, VA. U.S.A. Data Hammerson, G. 1994. Beaver (Castor current as of January 13, 2005. canadensis): Ecosystem alterations, management, and monitoring. Natural Areas Reuter, D.D. 1986. Sedge meadows of the upper Journal 14(1): 44-57. Midwest: A stewardship summary. Natural Areas Journal 6(4): 27-34. Heidorn 1991. Vegetation management guidelines: Buckthorns. Natural Areas Journal Stout, A. B. 1914. A biological and statistical 11: 216-217 analysis of the vegetation of a typical wild hay meadow. Transactions of the Wisconsin Hoffman, R. 2002. Wisconsin’s Natural Academy of Sciences, Arts, and Letters 17: Communities. How to recognize them, where 405-57. to find them. The University of Wisconsin Press, Madison, WI. 375 pp. Vogl, R. J. 1969. One hundred and thirty years of plant succession in a Southeastern Wisconsin Leach, M.K. and T.J. Givnish. 1996. Ecological lowland. Ecology 50: 248-55. determinants of species loss in remnant prairies. Science 273: 1555-1558. Warners, D. P. 1993. Species diversity in southern Michigan sedge meadows. Unpublished report

Grayling Community Assessment - 115 to The Nature Conservancy, Michigan Chapter, East Lansing, MI 35 pp.

Warners, D. P. 1997. Plant diversity in sedge meadows: Effects of groundwater and fire. Ph.D. dissertation, University of Michigan, Ann Arbor, MI. 231 pp.

White, K. L. 1965. Shrub-carrs of southeastern Wisconsin. Ecology 46: 286-304.

Abstract Citation: Cohen, J.G., and M.A. Kost. 2005. Natural community abstract for northern wet meadow. Michigan Natural Features Inventory, Lansing, MI. 8 pp.

Grayling Community Assessment - 116 Appendix 7. Northern Shrub Thicket

Overview: A fairly persistent, shrub-dominated noted by Vincent (1964), shade-intolerant alder wetland located north of the transition zone, develops rapidly following the removal of the typically occurring along streams, but also softwood overstory. In addition, alder can become adjacent to lakes and beaver floodings. Muck or established in multiple treefall gaps. Extensive peat soil is medium acid to neutral and nutrient mortality of Ulmus americana (American elm) rich. Fluctuating water table, beaver, and caused by Dutch elm disease not only eliminated windthrow limit succession to closed canopy elm as a dominant overstory tree in swamp swamp. Northern shrub thickets are ecosystems but also allowed the expansion of overwhelmingly dominated by Alnus rugosa shrub-dominated communities (Huenneke 1983). (speckled alder). The capacity of alder to form dense, impenetrable thickets can retard or prevent tree establishment Global and State Rank: G5?/S5 and regeneration (Huenneke 1983, Huenneke 1987). The extensive logging at the turn of the Range: Alder swamp community types are century and the gradual recovery of beaver widespread in the Midwestern and northeastern following the fur trapping era have likely caused a United States and southern Canada, ranging from rise in the water table in many sites, which have Maine west to Manitoba, south to Iowa, and east caused the expansion of shrub-dominated systems to New Jersey, mostly north of the glacial at the expense of forest. In other areas, historically boundary (Van Deelen 1991, Faber-Langendoen dominated by open, herbaceous wetlands (i.e., 2001, NatureServe 2005a, NatureServe 2005b). northern wet meadow, northern fen, emergent Within Michigan, northern shrub thicket is found marsh), tiling or ditching caused the lowering of in the northern half of the Lower Peninsula above the water table which resulted in the conversion of the climatic transition zone and throughout the these open systems to shrub-dominated wetlands Upper Peninsula (Barnes and Wagner 1981). (Curtis 1959, White 1965, Reuter 1986, Eggers and Reed 1997, Hoffman 2002). Beginning in the Rank Justification: In the Great Lakes region, 1920s, effective fire control by the U.S. Forest northern shrub thicket is a widespread community Service and state agencies reduced the acreage of type that has dramatically increased in acreage fires ignited by man or lightning (Swain 1973). As from its historical extent due to anthropogenic the result of fire suppression and subsequent shrub disturbance (Daly 1966). Interpretation of the encroachment, many open wetlands have notes of the original land surveyors of Michigan converted to shrub-dominated wetlands (Curtis reveal that in the 1800s, alder/willow-dominated 1959, Davis 1979, Reuter 1986, Jean and shrub wetland covered approximately 50,490 ha Bouchard 1991, Eggers and Reed 1997, Hoffman (124,761 ac) or just over 0.32% of the state 2002). This is especially evident where the water (Comer et al. 1995). Based on 2000 remote table has been lowered through tiling or ditching sensing imagery interpretation (MDNR 2003), and where the practice of mowing for marsh hay 385,105 ha (951,600 ac) of lowland shrub type has been abandoned (White 1965, Eggers and occurs in the northern Lower Peninsula and the Reed 1997). Northern shrub thicket has been Upper Peninsula, just under 2.5% of the state, maintained and expanded by wildlife management which constitutes close to an eight-fold increase in geared toward providing favorable habitat for extent. The drastic increase in northern shrub game species of early successional habitat thicket is the result of ubiquitous logging of (particularly white tailed deer, American swamp forests, alteration of hydrologic regimes, woodcock, and ruffed grouse). and fire suppression. Widespread logging of conifer swamp at the end of the 19th century and Physiographic Context: Northern shrub thickets the beginning of the 20th century (Karamanski occur principally along streams, beaver floodings, 1989) resulted in the conversion of many forested lakeshores, and rivers primarily within glacial swamps to northern shrub thicket in the Great outwash channels and less frequently within ice- Lakes region (Curtis 1959, Heinselman 1963). As contact topography and coarse-textured end

Grayling Community Assessment - 117 moraines (Curtis 1959, Schwintzer and Tomberlin Hardwood-Conifer Region has a cool snow-forest 1982, Hoffman 1989, Faber-Langendoen 2001, climate with short, warm summers, cold winters Hoffman 2002, Michigan Natural Features and a large number of cloudy days. The daily Inventory 2003, NatureServe 2005a). Sites are maximum temperature in July ranges from 24 to characterized by little to no slope, can range from 29 °C (75 to 85 °F), the daily minimum small pockets to extensive acreages, and are often temperature in January ranges from -21 to -9 °C (- a narrow band or zone of 20-30 meters within a 5 to 15 °F) and the mean annual temperature is 7 larger wetland complex (Curtis 1959, White 1965, °C (45 °F). The mean number of freeze-free days Faber-Langendoen 2001, NatureServe 2005a, is between 90 and 160, and the average number of NatureServe 2005b). Northern shrub thicket days per year with snow cover of 2.5 cm or more typically occurs adjacent to other wetland is between 80 and 140. The normal annual total communities such as emergent marsh, northern precipitation ranges from 740 to 900 mm with a wet meadow, northern swamp, poor conifer mean of 823 mm (Albert et al. 1986, Barnes 1991, swamp, and rich conifer swamp. Northern shrub Albert 1995). thicket can also be one of many zones within matrix communities such as Great Lakes marsh, Natural Processes: The dominant species of northern fen, and wooded dune and swale northern shrub thicket is speckled alder. Alder complex. Within the Great Lakes, this shrub contains symbiotic nitrogen-fixing bacteria wetland is found on Precambrian Shield bedrock (Frankia) in its root nodules that fix atmospheric that is overlaid with sandy loam soils that are nitrogen (Daly 1966, Van Deelen 1991, typically moderately well drained and deep NatureServe 2005a). Soil nitrogen may (Faber-Langendoen 2001). The soils overlaying accumulate at rates in excess of 85kg/ha (Daly the glacial till are wet to moist, nutrient-rich, well- 1966, Voigt and Steucek 1969). In addition to decomposed muck, peat, or mineral soil (Curtis atmospheric fixation of nitrogen, the rapid 1959, Van Deelen 1991, Faber-Langendoen 2001, decomposition of alder leaves contributes to the Michigan Natural Features Inventory 2003, enrichment of the soil (Barnes and Wagner 1981). NatureServe 2005a). The pH ranges widely from Organic matter accumulates very slowly in these alkaline to acidic (Curtis 1959) with medium systems since litter fall is broken down extremely acidity being the most prevalent condition rapidly (Daly 1966). Leaf litter beneath the shrub (Michigan Natural Features Inventory 2003). canopy is usually thin (often less than one Northern shrub thickets are non-stagnant wetlands centimeter) (Voigt and Steucek 1969). In addition with high levels of dissolved oxygen and soil to increasing the nitrogen supply of wetland soils, nitrogen that are seasonally flooded and range northern shrub thickets input nutrient rich detritus from poorly-drained to well-drained with most into aquatic ecosystems (NatureServe 2005a). sites remaining saturated throughout the growing Northern shrub thickets are most frequently found season (Curtis 1959, Daly 1966, Parker and along streams subject to periodic, seasonal Schneider 1974, Van Deelen 1991, Eggers and flooding. The soils are typically saturated and Reed 1997, NatureServe 2005a). Researchers well-aerated (Curtis 1959). Northern shrub have documented a strong correlation between thickets are typically areas of high primary soil moisture content and nitrogen concentration productivity because of the high nutrient supply in (Voigt and Steucek 1969). addition to normally favorable oxygen and soil moisture levels (Tilton and Bernard 1975). In sites The Michigan range of northern shrub thicket falls subject to periodic flooding, alder stems can slow within the area classified by Braun (1950) as the flood waters and trap sediment. Over time, fine- Northern Hardwood-Conifer Region textured sediments accumulate over coarser (Hemlock/White Pine/Northern Hardwoods alluvial material and the land surface may Region) and within the following regions eventually rises above the flood levels classified by Albert et al. (1986) and Albert (NatureServe 2005a). (1995): Region II, Northern Lower Michigan; Region III, Eastern Upper Michigan; and Region IV, Western Upper Michigan. The Northern

Grayling Community Assessment - 118 Historically, alder was one of the first woody are adapted to periodic flooding but intolerant of species to colonize recently deglaciated prolonged and severe flooding (Knighton 1981, landscapes as an early successional species (Sears Ohmann et al. 1990). 1948). Currently, northern shrub thickets can become established following severe disturbance In the absence of disturbance factors that prevent of swamp forested systems or by invading open tree establishment and survival or conversion to wetlands. Alder is shade intolerant (Barnes and more open conditions, northern shrub thicket Wagner 1981, Huenneke and Marks 1987), typically succeed to closed canopy swamp forest tolerant of seasonal flooding (Knighton 1981, (Curtis 1959). The capacity of alder to condition Ohmann et al. 1990), and moderately tolerant of the soil by increasing the available nitrogen and fire (Van Deelen 1991). Alder often persists in contribute to the accumulation of top soil through swamp forests in light gaps. Flooding (i.e. from sediment trapping creates a suitable soil substrate beaver or fluvial processes), fire, disease, and for tree establishment and growth. Saplings of tree windthrow can result in sufficient mortality of the species grow through the shrub layer and once a swamp forest overstory to allow for the complete tree canopy becomes established, alder are shaded opening of the canopy and the expansion of alder out, unable to continue sprout production, through establishment of seedlings or stump experience a diminished capacity to fix nitrogen, sprouting. Following canopy release, alder can and decline (Daly 1966, Barnes and Wagner 1981, form dense, impenetrable thickets that retard or Huenneke and Marks 1987). Tree species that prevent tree establishment (Huenneke 1983, typically invade northern shrub thickets include Huenneke 1987). Within open wetlands, alder and Abies balsamea (balsam fir), Acer rubrum (red associated shrubs can become established maple), Fraxinus nigra (black ash), Larix laricina following alteration in the fire or hydrologic (tamarack), Picea mariana (black spruce), regime. Prolonged periods without fire or the Populus balsamifera (balsam poplar), Populus lowering of the water table allows for shrub tremuloides (quaking aspen), and Thuja encroachment and conversion to northern shrub occidentalis (northern white cedar) (Parker and thicket (Curtis 1959, White 1965, Jean and Schneider 1974, Van Deelen 1991, Faber- Bouchard 1991, Eggers and Reed 1997, Hoffman Langendoen 2001, Michigan Natural Feature 2002). Inventory 2003, NatureServe 2005a). Northern shrub thickets can be replaced by hardwood Once established northern shrub thicket can swamp, hardwood-conifer swamp, and conifer persist if disturbance factors maintain the open swamp. Succession from shrub swamp to swamp canopy conditions. Windthrow, beaver herbivory, forest can occur within ten years but may take beaver flooding, seasonal flooding, and fire can longer. Curtis (1959) postulated that the minimum limit tree establishment and survival (Michigan life expectancy of shrub-carr (in southern Natural Features Inventory 2003). Alders capacity Wisconsin) is fifty years. to stump sprout following flooding, fire, and herbivory allow it to persist after these Vegetation Description: Northern shrub thicket disturbances (Ohmann et al. 1990, Huenneke and is characterized by an overwhelming dominance Marks 1987). Basal sprout production is critical of speckled alder which forms dense (often for maintenance of alder thickets (Huenneke monotypic) thickets with canopy coverage 1987, Huennek and Marks 1987). Northern shrub ranging between 40-95% and stand height ranging thickets can recover from moderate flooding and from one to eight meters but typically between fire disturbance within five years (White 1965). In one and three meters (Curtis 1959, Vincent 1964, instances where flooding or fire is severe White 1965, Parker and Schneider 1974, Tilton (prolonged flooding above the root crown or and Bernard 1975, Mattson and Winsauer 1986, burning of the mineral soil and root crown), alder Huenneke 1987, Hoffman 1989, Van Deelen fails to stump sprout and the shrub thicket may be 1991, Hoffman 2002, Michigan Natural Features replaced by an open wetland, such as northern wet Inventory 2003, Faber-Langendoen 2001, meadow or northern fen (Knighton 1981, Ohmann NatureServe 2005a). Estimates of stem density et al. 1990, Van Deelen 1991). Alder and willow range widely from 5,000 to 30,000 stems per acre

Grayling Community Assessment - 119 (Vincent 1964, Mattson and Winsauer 1986, Mimulus ringens (monkey-flower), Polygonum Tilton and Bernard 1975). In a study in Upper sagittatum (tear-thumb or smartweed), Rumex Michigan and northern Wisconsin, Mattson and orbiculatus (greater water dock), Scutellaria Winsauer (1986) estimated that on average, alder galericulata (marsh skullcap), Scutellaria thickets produce close to 30 tons of green biomass lateriflora (mad-dog skullcap), Senecio aureus per acre. Stem diameter of alder is typically (golden ragwort), Smilacina trifolia (false between one to five inches (Parker and Schneider Solomon-seal), Solidago canadensis (Canada 1974, Tilton and Bernard 1975, Barnes and goldenrod), S. gigantea (late goldenrod), S. Wagner 1981, Mattson and Winsauer 1986). rugosa (rough-leaved goldenrod), Symplocarpus Alder are monoecious and wind-pollinated with foetidus (skunk cabbage), and Thalictrum germination usually requiring exposed mineral dasycarpum (purple meadow-rue). Where the tall soil (Huenneke 1985, Van Deelen 1991). Within shrub canopy is open, graminoids can become established stands of alder, vegetation dense. The most dominant grass species in reproduction through stump sprouting provides northern shrub thicket is Calamagrostis most of the new stems (Huenneke 1985, canadensis (blue-joint grass). Other common Huenneke 1987, Van Deelen 1991). Alder forms grasses include Bromus ciliatus (fringed brome), clonal clumps where few to many stems arise at Glyceria striata (fowl manna grass), Leersia the root collar (Barnes and Wagner 1981). A fast- oryzoides (cut grass), and Poa palustris (fowl growing, shade intolerant shrub, alder is usually meadow grass). A diversity of sedges is found in short-lived; alder stems typically live 10-30 years these systems including Carex stricta (tussock (Tilton and Bernard 1975, Barnes and Wagner sedge), Carex flava (yellow sedge), Carex 1981, Huenneke 1987, Huenneke and Marks lacustris (lake or hairy sedge), and Carex leptalea 1987). (bristly-stalked sedge). Bulrushes such as Scirpus atrovirens (green bulrush) are also common. Northern shrub thicket exhibits a high degree of Numerous Sphagnum species and ferns thrive in floristic homogeneity due to the dominance of these moist, saturated systems. Characteristic alder (Tilton and Bernard 1975). Floristic ferns and fern allies include Dryopteris cristata diversity is usually correlated with the degree of (crested woodfern), Equisetum arvense (field shrub canopy closure with more diversity being horsetail), Onoclea sensibilis (sensitive fern), found in the more open sites (Eggers and Reed Osmunda cinnamomea (cinnamon fern), Osmunda 1997). The understory, intermediate between regalis (royal fern), and Thelypteris palustris meadow and forest (Curtis 1959), is dominated by (marsh fern). an array of short shrubs, forbs, grasses, sedges, and ferns. The density of the understory varies The understory layer also contains numerous short inversely with the density of the tall-shrub shrubs including Chamaedaphne calyculata canopy. (leatherleaf), Ledum groenlandicum (Labrador- tea), Myrica gale (wax-myrtle or bayberry), Prevalent herbs of the northern shrub thicket Potentilla palustris (marsh cinquefoil), Ribes include: Asclepias incarnata (swamp milkweed), americanum (wild black currant), Rubus hispidus Aster lanceolatus (panicled aster), A. puniceus (swamp dewberry), R. pubescens (dwarf (swamp aster), A. umbellatus (flat-topped aster), raspberry), R. strigosus (wild red raspberry), and Caltha palustris (marsh marigold), Campanula Spirea alba (meadowsweet). Where alder does not aparinoides (marsh bellflower), Chelone glabra form a monospecific shrub layer, associates of the (turtlehead), Clematis virginiana (woodbine), tall shrub layer can include Aronia prunifolia Epilobium coloratum (purple-leaf willow-herb), (chokeberry), Betula pumila (bog birch), Cornus Eupatorium maculatum (Joe-pye-weed), amomum (silky dogwood), C. stolonifera (red- Eupatorium perfoliatum (boneset), Galium osier dogwood), Ilex verticilata (winterberry), asprellum (rough bedstraw), Impatiens capensis Salix bebbiana (Bebb’s willow), S. discolor (spotted touch-me-not), Iris versicolor (wild blue (pussy willow), S. exigua (sandbar willow), S. flag), Lycopus uniflorus (northern bugleweed or petiolaris (slender willow), Viburnum cassinoides water-horehound), Mentha arvensis (wild mint), (wild-raisin), and Viburnum opulus (highbush-

Grayling Community Assessment - 120 cranberry). Scattered trees and tree saplings are Lepus americanus (snowshoe hare). Beaver build often found invading northern shrub thickets. dams and lodges with speckled alder. The buds Typical tree species include balsam fir, red maple, and seeds of alder are eaten by a diversity of black ash, tamarack, black spruce, balsam poplar, birds. Songbirds feed on the seeds and Philohela quaking aspen, and northern white cedar. (Above minor (American woodcock) and Bonasa species lists compiled from Michigan Natural umbellus (ruffed grouse) eat the buds and catkins. Features Inventory database, Curtis 1959, White Thickets of alder provide important hiding cover 1965, Parker and Schneider 1974, Hoffman 1989, for species such as Odocoileus virginianus (white- Van Deelen 1991, Eggers and Reed 1997, Faber- tailed deer), Lutra canadensis (river otter), and Langendoen 2001, Hoffman 2002, NatureServe Mustela vison (mink) (Barnes and Wagner 1981, 2005a.) Van Deelen 1991). Canis lupus (gray wolf, state threatened) and Lynx canadensis (lynx, state Michigan Indicator Species: blue-joint grass, endangered) utilize shrub thicket habitat. Joe-pye-weed, marsh fern, marsh marigold, northern bugleweed, sensitive fern, speckled Conservation and Biodiversity Management: alder, and spotted touch-me-not. Northern shrub thicket is a widespread community type in the Great Lakes region and has Other Noteworthy Species: Northern shrub dramatically increased from its historical extent thicket provide habitat for numerous herptiles due to anthropogenic disturbance. Alder swamps including Emys blandingii (Blanding’s turtle, state contribute significantly to the overall biodiversity special concern), Glyptemys insculpta (wood of northern Michigan by providing habitat to a turtle, state special concern), Pseudacris triseriata wide variety of plant and animal species including maculata (boreal chorus frog, state special several rare species. However, northern shrub concern), and Sistrurus catenatus catenatus thickets have replaced many rare and declining (eastern massasauga, state special concern, federal wetland communities such as rich conifer swamp candidate species). If suitable nesting trees or and northern fen. Where shrub encroachment snags are available, Haliaeetus leucocephalus threatens to convert open wetlands to shrub- (bald eagle, state threatened) and Pandion dominated systems, repeated prescribed fires, haliaetus (osprey, state threatened) can be found mowing, or herbicide application to cut shrub nesting in these systems and Ardea herodias stumps can be employed to maintain open (great blue heron, protected by the Migratory Bird conditions (White 1965, Heidorn 1991). On sites Treaty Act of 1918) can establish rookeries in which northern shrub thicket is succeeding to (Hoffman 1989). swamp forest, allowing succession to proceed unhindered will result in the increase of less Rare plants associated with northern shrub thicket common swamp systems. In situations where the include Listera auriculata (auricled twayblade, management objective is to prevent succession, state special concern), Lonicera involucrata northern shrub thicket can be maintained by (black twinberry, state threatened), Mimulus cutting the overstory (Vincent 1964). Following guttatus (western monkey-flower, state special canopy removal with scarification of the soil and concern), Stellaria crassifolia (fleshy stitchwort, mild intensity burning encourages alder state threatened), and Thalictrum venulosum var. regeneration (Van Deelen 1991). Northern shrub confine (veiny meadow-rue, state special thickets provide ecosystem services, protecting concern). The single collection of Equisetum water quality by assimilating nutrients, trapping telmateia (giant horsetail, state extirpated) in sediment, and retaining stormwater and Michigan was from a “damp alder thicket”. floodwater.

The leaves and twigs of alder provide important Research Needs: Northern shrub thicket has a browse for a wide array of mammals including broad distribution and exhibits numerous regional, Alces alces (moose, state threatened), Ondatra physiographic, hydrologic, and edaphic variants. zibethicus (muskrat), Castor canadensis (beaver), The diversity of variations throughout its range Sylvilagus floridanus (cottontail rabbit), and demands the continual refinement of regional

Grayling Community Assessment - 121 classifications that focus on the inter-relationships Related Abstracts: Blanding’s turtle, eastern between vegetation, physiography, hydrology, and massasauga, floodplain forest, great blue heron soils (White 1965, Barnes et al. 1982, Hoffman rookery, Great Lakes marsh, northern fen, 1989, Faber-Langendoen 2001, NatureServe northern wet meadow, rich conifer swamp, 2005a). Little is known about the flooding and fire wooded dune and swale complex, and wood regimes of northern shrub thickets and the turtle. interaction of disturbance factors within these systems. As noted by Hammerson (1994), beaver Literature Cited: significantly alter the ecosystems they occupy. An Albert, D.A. 1995. Regional landscape important research question to examine is how the ecosystems of Michigan, Minnesota, and wetland ecosystems of the Great Lakes have been Wisconsin: A working map and classification. and continue to be affected by fluctuations in Gen. Tech. Rep. NC-178. St. Paul, MN: U.S. populations of beaver. Experimentation is needed Department of Agriculture, Forest Service, to determine how best to prevent shrub North Central Forest Experiment Station. encroachment of open wetlands that are Northern Prairie Wildlife Research Center threatened by conversion to northern shrub Home Page. thicket. Effects of management within northern http://www.npwrc.usgs.gov/resource/1998/rla shrub thickets need to be monitored to allow for ndscp/rlandscp.htm. (Version 03JUN98.) 250 assessment and refinement. pp.

Similar Communities: floodplain forest, Great Albert, D.A., S.R. Denton, and B.V. Barnes. 1986. Lakes marsh, hardwood-conifer swamp, northern Regional landscape ecosystems of Michigan. fen, northern swamp, northern wet meadow, poor Ann Arbor, MI: University of Michigan, conifer swamp, rich conifer swamp, southern School of Natural Resources. 32 pp. & map. shrub-carr, and wooded dune and swale complex. Barnes, B.V. 1991. Deciduous forest of North Other Classifications: America. Pp 219-344 in E. Röhrig and B. Michigan Natural Features Inventory Circa 1800 Ulrich, eds., Temperate Deciduous Forests. Vegetation (MNFI): Alder, Willow, Bog Birch Elsevier, Amsterdam. 635 pp. Thicket (6122) Barnes, B.V., and W.H. Wagner, Jr. 1981. Michigan Department of Natural Resources Michigan Trees: A Guide to the Trees of (MDNR): L-lowland brush Michigan and the Great Lakes Region. University of Michigan Press, Ann Arbor, MI. Michigan Resource Information Systems 383 pp. (MIRIS): 612 (shrub/scrub wetland) Barnes, B.V., K.S. Pregitzer, T.A. Spies and V. H. The Nature Conservancy National Classification: Spooner. 1982. Ecological forest site CODE; ALLIANCE; ASSOCIATION; classification. Journal of Forestry 80(8): 493- COMMON NAME 498.

III.B.2.N.e; Alnus incana Seasonally Flooded Braun, E.L. 1950. Deciduous forests of eastern Shrubland Alliance; Alnus incana Swamp North America. Hafner Press, New York, NY. Shrubland; Speckled Alder Swamp 596 pp. Shrubland; Speckled Alder Swamp Comer, P.J., D.A. Albert, H.A. Wells, B.L. Hart, NatureServe Ecological Systems Classification: J.B. Raab, D.L. Price, D.M. Kashian, R.A. CES201.582: Laurentian-Acadian Wet Meadow- Corner, and D.W. Schuen. 1995. Michigan’s Shrub Swamp presettlement vegetation, as interpreted from the General Land Office Surveys 1816-1856.

Grayling Community Assessment - 122 Michigan Natural Features Inventory, Huenneke, L.F. 1983. Understory response to Lansing, MI. Digital map. gaps caused by the death of Ulmus americana in central New York. Bulletin of the Torrey Curtis, J.T. 1959. Vegetation of Wisconsin: An Botanical Club 110(2): 170-175. Ordination of Plant Communities. University of Wisconsin Press, Madison, WI. 657 pp. Huenneke, L.F. 1985. Spatial distribution of genetic individuals in thickets of Alnus incana Daly, G.T. 1966. Nitrogen fixation by nodulated ssp. rugosa, a clonal shrub. American Journal Alnus rugosa. Canadian Journal of Botany 44: of Botany 72(1): 152-158. 1607-1621. Huenneke, L.F. 1987. Demography of a clonal Davis, A. M. 1979. Wetland succession, fire and shrub, Alnus incana ssp. rugosa (Betulaceae). the pollen record: A Midwestern example. The The American Midland Naturalist 117(1): 43- American Midland Naturalist 102: 86-94. 55.

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