Response of Vegetation in Open and Partially Wooded Fens 

to Prescribed Burning at Seney National Wildlife Refuge

Scientific Investigations Report 2018–5168

U.S. Department of the Interior

U.S. Geological Survey
Cover. Photographs of prescribed burns at Seney National Wildlife Refuge. 

Photographs by J.E. Austin, U.S. Geological Survey
Response of Vegetation in Open and 

Partially Wooded Fens to Prescribed 

Burning at Seney National Wildlife Refuge

By Jane E. Austin and Wesley E. Newton

Scientific Investigations Report 2018–5168

U.S. Department of the Interior

U.S. Geological Survey
U.S. Department of the Interior

DAVID BERNHARDT, Acting Secretary

U.S. Geological Survey

James F. Reilly II, Director

U.S. Geological Survey, Reston, Virginia: 2019

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Suggested citation:

Austin, J.E., and Newton, W.E., 2019, Response of vegetation in open and partially wooded fens to prescribed burning 

at Seney National Wildlife Refuge: U.S. Geological Survey Scientific Investigations Report 2018–5168, 62 p., 

https://doi.org/10.3133/sir20185168.

ISSN 2328-0328 (online)
iii

Acknowledgments

Seney National Wildlife Refuge provided logistical support, historical information, and spatial 

data. We are particularly appreciative of D. Olson, refuge biologist, and G. Lindsay, fire management officer, at Seney National Wildlife Refuge for their support and insightful discussions 

about wetland and fire ecology and management. 

Funding was provided by U.S. Geological Survey’s Northern Prairie Wildlife Research Center. 

This study was not possible without field assistance provided by botanists S. Danneman, 

A. Sabai, and S. Swanson, and other field assistance from N. Carle, J. Edwards, A. Given, 

C. Hanson, K. Marinoff, and B. Pauly. We thank D. Olson, G. Lindsay, K. Kowalski, and 

B. Middleton for their comments on earlier drafts.

v

Contents

Acknowledgments........................................................................................................................................iii

Abstract...........................................................................................................................................................1

Introduction.....................................................................................................................................................1

Study Area.......................................................................................................................................................2

Methods...........................................................................................................................................................4

Sampling Design and Site Selection .................................................................................................4

Data Collection......................................................................................................................................4

Prescribed Burn Treatments...............................................................................................................8

Data Analyses......................................................................................................................................12

Results ...........................................................................................................................................................13

Environmental and Habitat Variables ..............................................................................................13

Plant Community .................................................................................................................................18

Species Richness and Diversity..............................................................................................18

Percent Cover by Taxa—Year and Treatment Effects .........................................................18

Changes in Plant Community Indicated by Ordinations ...............................................................29

Woody Height, Patchiness, and Cover............................................................................................32

Vertical Cover......................................................................................................................................33

Discussion.....................................................................................................................................................39

Changes in Plant Community ............................................................................................................40

Woody Cover and Structure..............................................................................................................42

Fire Management and History in Fens ............................................................................................42

Rethinking Fire Management for Controlling Woody Encroachment in Fens....................................43

References Cited..........................................................................................................................................44

Glossary.........................................................................................................................................................49

Appendix 1. Crosswalk Table of Taxonomy of Plant Species. ..............................................................50

References Cited..........................................................................................................................................53

Appendix 2. Fire Conditions During Prescribed Burns at Marsh Creek, July 2007 

and 2008, and C3, May 2008..........................................................................................................54

References Cited..........................................................................................................................................54

Appendix 3. Frequency of Occurrence of Plant Taxa by Block in C3 and Marsh Creek, 

Seney National Wildlife Refuge, 2006–10...................................................................................55

Appendix 4. Frequency of Occurrence and Percent of Points (Summed Across 

Sampling Years) of Plant Taxa by Block in Marsh Creek, Seney National 

Wildlife Refuge, Michigan, 2006–10. ...........................................................................................59

Figures

1. Map showing location of C3 and Marsh Creek study areas in Seney National 

Wildlife Refuge, in the upper peninsula of Michigan .............................................................3

2. Map showing location of study blocks at the C3 study area relative to cover 

types in Seney National Wildlife Refuge, Michigan ...............................................................5

3. Map showing location of study blocks at the Marsh Creek study area relative 

to cover types in Seney National Wildlife Refuge, Michigan................................................6
vi

4. Diagram showing layout of transects and sampling points, spaced 25 meters 

apart, and associated identifying numbers, within each experimental block....................7

5. Photographs of conditions after the prescribed burn on the C3 study area, 

May 2008 ........................................................................................................................................9

6. Photographs of conditions after the prescribed burn on the MC–2 block of the 

Marsh Creek study area, July 2007..........................................................................................10

7. Photographs of conditions after prescribed burn on the Marsh Creek study area, 

July 2008.......................................................................................................................................11

8. Graphs showing summer water depths in control and burn blocks in C3 and 

Marsh Creek blocks, 2006–10, at Seney National Wildlife Refuge, Michigan..................15

9. Graphs showing litter depths in control and burn blocks in C3 and Marsh 

Creek blocks, 2006–10, at Seney National Wildlife Refuge, Michigan ..............................16

10. Graphs showing graminoid height in control and burn blocks in C3 and 

Marsh Creek blocks, 2006–10, at Seney National Wildlife Refuge, Michigan..................17

11. Graphs showing measures of biological diversity detected in control and 

burn blocks in C3 and Marsh Creek study areas, 2006–10, at Seney National 

Wildlife Refuge, Michigan.........................................................................................................19

12. Graph showing percent open area affected by year × treatment interactions 

in control and burn blocks in C3 and Marsh Creek blocks, 2006–10, at Seney 

National Wildlife Refuge, Michigan.........................................................................................21

13. Graph showing percent cover of sedges affected by year × treatment 

interactions in control and burn blocks in C3 and Marsh Creek blocks, 

2006–10, at Seney National Wildlife Refuge, Michigan........................................................22

14. Graph showing percent cover of sedges and rushes affected by year × treatment 

interactions in control and burn blocks in C3 and Marsh Creek blocks, 2006–10, 

at Seney National Wildlife Refuge, Michigan........................................................................23

15. Graphs showing percent cover of grasses affected by separate year and 

treatment effects in control and burn blocks in C3 and Marsh Creek blocks, 

2006–10, at Seney National Wildlife Refuge, Michigan........................................................24

16. Graphs showing percent cover of forbs affected by year × treatment 

interactions in control and burn blocks in C3 and Marsh Creek blocks, 

2006–10, at Seney National Wildlife Refuge, Michigan........................................................25

17. Graphs showing percent cover of forbs affected by year effects in control 

and burn blocks in C3 and Marsh Creek blocks, 2006–10, at Seney National 

Wildlife Refuge, Michigan.........................................................................................................27

18. Graph showing percent cover of earth loosestrife affected by separate year 

and treatment effects in control and burn blocks in C3 and Marsh Creek 

blocks, 2006–10, at Seney National Wildlife Refuge, Michigan..........................................28

19. Graph showing percent cover of mosses affected by separate year and 

treatment effects in control and burn blocks in C3 and Marsh Creek blocks, 

2006–10, at Seney National Wildlife Refuge, Michigan........................................................28

20. Graph showing percent cover of leatherleaf affected by year × treatment 

interactions in control and burn blocks in C3 and Marsh Creek blocks, 

2006–10, at Seney National Wildlife Refuge, Michigan........................................................29

21. Graphs showing percent cover of woody species affected by year in control 

and burn blocks in C3 and Marsh Creek blocks, 2006–10, at Seney National 

Wildlife Refuge, Michigan.........................................................................................................30

22. Graphs showing average annual nonmetric dimensional scaling scores for 

plant community in control and burn blocks in C3 and Marsh Creek, Seney 

National Wildlife Refuge, Michigan.........................................................................................31
vii

23. Graphs showing visual obstruction readings in control and burn blocks for 

33-centimeter strata in C3, Seney National Wildlife Refuge, Michigan, 2006–10............38

24. Graphs showing visual obstruction readings in control and burn blocks for 

33-centimeter strata in C3, Seney National Wildlife Refuge, Michigan, 2006–10............39

Tables

1. Fire history, status, and timing of prescribed burning on study blocks in the 

C3 and Marsh Creek study areas, Seney National Wildlife Refuge, Michigan, 

2006–10 ...........................................................................................................................................8

2. Correlations among mean and variability of water depth, litter depth, and 

graminoid height measures in study blocks in C3 and Marsh Creek study 

areas, Seney National Wildlife Refuge, Michigan, 2006–10................................................13

3. Results of analysis of variance for habitat and environmental variables on 

study blocks in C3 and Marsh Creek study areas, Seney National Wildlife 

Refuge, Michigan, 2006–10 .......................................................................................................14

4. Results of analysis of variance for species richness, species diversity index, 

nonmetric dimensional scaling scores on 3 axes, and percent cover of open, 

moss, and 27 plant taxa on study blocks in C3 and Marsh Creek study areas, 

Seney National Wildlife Refuge, Michigan, 2006–10 ............................................................20

5. Pearson correlation coefficients for 27 most common plant taxa with nonmetric 

dimensional scaling axes in study blocks in C3 and Marsh Creek study areas, 

Seney National Wildlife Refuge, Michigan, 2006–10 ............................................................32

6. Results of analysis of variance for woody taxa height and standard deviation 

of woody height on study blocks in C3 and Marsh Creek study areas, Seney 

National Wildlife Refuge, Michigan, 2006, 2007, and 2010...................................................33

7. Mean shrub height, variation of height, number of patches, and percent cover 

for six shrub taxa and total shrub cover at Seney National Wildlife Refuge, 

Michigan, 2006, 2007, and 2010.................................................................................................34

8. Results of analysis of variance for percent cover and number of patches of 

woody taxa on study blocks in C3 and Marsh Creek study areas, Seney 

National Wildlife Refuge, Michigan, 2006, 2007, and 2010...................................................37

9. Results of analysis of variance for visual obstruction readings for six height 

strata on study blocks in C3 and Marsh Creek study areas, Seney National 

Wildlife Refuge, Michigan, 2006–10.........................................................................................38

Appendix Tables

1.1. Crosswalk table of taxonomy of plant species following the Flora of North 

America (2003) and U.S. Department of Agriculture National PLANTS Database..........50

2.1. Fire conditions during prescribed burns at Marsh Creek, July 2007 and 2008, 

and C3, May 2008 ........................................................................................................................54

3.1. Frequency of occurrence and percent of points summed across sampling years of 

plant taxa by block in C3 of Seney National Wildlife Refuge, Michigan, 2006–10 ...........55

4.1. Frequency of occurrence and percent of points of plant taxa by block in Marsh 

Creek of Seney National Wildlife Refuge, Michigan, 2006–10 ............................................59
viii

Conversion Factors

International System of Units to U.S. customary units

Multiply By To obtain

Length

centimeter (cm) 0.3937 inch (in.)

meter (m) 3.281 foot (ft) 

Area

hectare (ha) 2.471 acre

square kilometer (km2) 247.1 acre

hectare (ha) 0.003861 square mile (mi2) 

square kilometer (km2) 0.3861 square mile (mi2)

Temperature in degrees Celsius (°C) may be converted to degrees Fahrenheit (°F) as follows:

°F = (1.8×°C) + 32

Temperature in degrees Fahrenheit (°F) may be converted to degrees Celsius (°C) as follows:

°C = (°F-32) / 1.8

Abbreviations

ANOVA analysis of variance

BACI before-after-control-impact

NMS nonmetric multidimensional scaling

VOR visual obstruction reading
Response of Vegetation in Open and Partially Wooded 

Fens to Prescribed Burning at Seney National Wildlife 

Refuge

By Jane E. Austin and Wesley E. Newton

Abstract

The health and function of northern peatlands, particularly for fens, are strongly affected by fire and hydrology. 

Fens are important to several avian species of conservation 

interest, notably the yellow rail (Coturnicops noveboracensis). Fire suppression and altered hydrology often result 

in woody encroachment, altering the plant community and 

structure. Woody encroachment and its effects on biodiversity 

have become an increasing concern in the conservation and 

management of plant communities. This study evaluated the 

effects of spring and summer prescribed burns on the plant 

community, cover, and structure in open and partially wooded 

fens at Seney National Wildlife Refuge, Michigan, using a 

before-after-control-impact design. Paired, 1-hectare blocks 

were established in two fen areas, C3 and Marsh Creek, and 

data were collected for 2 years before burning (2006–7) and 

3 years after burning (2008–10). We used generalized linear 

mixed models and ordination to assess differences among 

four treatments: C3 control, C3 spring burn (May 2008), 

Marsh Creek control, and Marsh Creek summer burn (July 

2008); results from a block burned under drier conditions in 

July 2007 also are reported. Variables include water depth; 

litter depth; graminoid height; species richness and diversity; 

percent cover of plant taxa, mosses, and open area; shrub 

height, number of patches, and cover; and visual obstruction readings. The 2008 prescribed burns were done under 

moderate fire conditions, whereas the 2007 summer burn 

on one block was done under high fire conditions because 

of prolonged drought. We identified 104 plant taxa over the 

5 years and noted differences between C3 and Marsh Creek 

communities. We examined data for effects of treatment, year, 

and year × treatment interactions for percent open and the 

28 most common taxa. Most differences among treatments 

were related to natural differences in the plant community 

and hydrology between the two areas rather than fire effects; 

year effects were likely related to annual differences in water 

conditions. We detected few effects of spring burning in 

C3, even in the same year of burning. In Marsh Creek, most 

treatment effects were in 2008, when data were collected 

within 3 weeks of burning. Some fire effects there, however, 

persisted one to two growing seasons (2009, 2010) and two to 

three growing seasons in the block burned in the more intense 

summer 2007 fire. Effects of burning on shrub measures were 

more apparent on summer-burned blocks, but most measures 

returned to preburn conditions by 2010. Our results demonstrate the heterogeneity of plant community and environmental 

conditions of fens within and among years and the interactions 

of water conditions with burning. The results also demonstrate 

that neither single spring nor summer burning under moderate fire conditions are effective in setting back woody cover. 

Maintaining more open conditions in fens may require different approaches to water management, more frequent fires, 

more aggressive fire management, or a combination of tools to 

control woody cover.

Introduction

Peatlands are one of the dominant ecosystem types in the 

upper Great Lakes, covering roughly 60,000 square kilometers 

(km2) and more than 10 percent of total land area (Boelter and 

Verry, 1977). They exist in the landscape as part of a matrix of 

wetland and upland forested communities. Peatlands encompass a variety of habitats, including marsh, bog, open fen, 

partially wooded fen (intermix of grasses, sedges [Carex], and 

woody species), and lowland forest. Peatlands are of significant ecological importance because of their role in hydrology, 

water quality, carbon sequestration, and habitat for wildlife. 

The health and function of peatlands are strongly affected by 

fire and hydrology (Vogl, 1969; Rydin and Jeglum, 2006). Fire 

suppression and altered hydrology often result in the encroachment of trees and shrubs into open fens and bogs, altering 

vegetation structure and community (White, 1965; Vogl, 1969; 

Middleton, 2002; Brisson and others, 2006). Increased shrub 

cover also can disrupt the fire regime by altering microclimate 

and suppressing development of fine fuels such as graminoids 

that would carry fire through the system. Alternatively, during 

dry conditions, the high woody fuel loads in areas of heavy 

shrub cover can result in fires of higher intensity and greater 

severity, which are disruptive at site and landscape levels.
2 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Open and partially wooded fens are important to several 

species of conservation concern in North America. The yellow 

rail (Coturnicops noveboracensis) is a rare species associated with shallowly flooded wetlands, most commonly open 

fens dominated by sedges. Knowledge about its population 

status in most areas, its basic ecology, and its response to 

habitat management is limited (Goldade and others, 2002; 

Austin and Buhl, 2013; Leston and Bookhout, 2015). The 

species is considered endangered in Michigan and is listed as 

a priority species in Partners in Flight for the Boreal Hardwood Transition Area (Bird Conservation Area 20) and is a 

species of conservation concern by the U.S. Fish and Wildlife 

Service–Region 3 (U.S. Fish and Wildlife Service, 2008). 

Yellow rail populations are believed to be limited by loss or 

degradation of wetland habitat because of drainage, altered 

hydrology, and fire suppression (Committee on the Status of 

Endangered Wildlife in Canada, 2009; Leston and Bookhout, 

2015). Habitat differences relative to fire history, and comparisons between sites with and without yellow rails indicated 

that yellow rails used areas with the deepest litter and highest 

ground cover, and low shrub cover and heights, and used landscapes having greater sedge-graminoid cover and less lowland 

woody or upland cover types (Austin and Buhl, 2013). Fens 

also are important habitats for LeConte’s sparrows (Ammodramus leconteii) and sedge wrens (Cistothorus platensis), both 

considered priority species for Bird Conservation Area 20. 

Most information on these two species is based on grassland 

ecosystems (Dechant and others, 2002a, b), and little data exist 

regarding their abundance or response to fire, hydrology, or 

vegetative structure in fens. Understanding what fire regimes 

or treatments result in preferred habitat conditions is key to 

managing these peatland habitats for species of conservation 

concern and for the larger biological community.

Large tracts of fens are in the eastern upper peninsula 

of Michigan and are exemplified by the habitats within the 

Seney National Wildlife Refuge (hereafter referred to as “the 

Refuge”). These habitats commonly host substantial numbers 

of breeding yellow rails, LeConte’s sparrows, and sedge 

wrens, among other avian species (Bookhout and Stenzel, 

1987; J. E. Austin, unpub. data, 2007–09). However, many 

years of altered fire regimes and hydrology have resulted in 

encroachment of shrubs into once extensive open fens and 

bogs and have altered the plant community (Kowalski and 

Wilcox, 2003; Bork and others, 2013). A key priority of the 

Refuge, as identified in their Comprehensive Conservation 

Plan (U.S. Fish and Wildlife Service, 2009), is to restore 

these habitats back to their historical conditions, primarily 

through fire management, for the benefit of species such as 

the yellow rail and LeConte’s sparrow. Previous research on 

burning at the Refuge and elsewhere provided limited data 

as to the effectiveness of burning to reduce woody encroachment in open fens because they usually examined only spring 

burns, included only 1 year of data postburn, and had limited 

data on shrub and community structure (Hanowski and 

others, 1999; Middleton, 2002; Brisson and others, 2006). 

Fire conditions and severity were rarely reported, limiting 

the ability to discern the effects (or lack of effect) of fire on 

habitat conditions (such as litter) or vegetation community and 

structure. An earlier yellow rail study at the Refuge indicated 

that rails avoided areas the first growing season after burning 

but seemed to prefer them the second growing season after 

burning, but sample design and sizes were limited (Burkman, 

1993). Hence, more detailed investigation of effects of fire 

on habitat conditions, at least 2 years of data after burning, 

were needed to begin to evaluate appropriate fire management 

practices on the plant and shrub community and, in turn, for 

avian species.

This study was part of a larger project to investigate the 

avian community of fen habitats in the Refuge in the context 

of fire (Austin and Buhl, 2013; J. E. Austin, unpub. data, 

2007–10). At the heart of the project was this experimental 

component that examined the effects of prescribed burns 

on fen habitats using a before-after-control-impact (BACI) 

design. Our objective was to evaluate effects of prescribed 

burning in spring versus summer on plant community, plant 

cover, and vegetative structure in open and partially wooded 

fens at the Refuge to better inform fire management practices.

Study Area

The Refuge (lat. 46°15′N, long. 86°04′W) is in Schoolcraft County on the eastern upper peninsula of Michigan 

(fig. 1). This region once had about 185,000 hectares (ha) of 

open sedge-, forb-, and moss-dominated communities before 

European settlement (Comer and others, 1995; Slaughter and 

Cohen, 2010). The Refuge’s extensive peatlands are primarily 

open fens (dominated by sedge and other graminoid species), 

scrub/shrub lowland (wooded fen), and patterned bogs, 

interspersed with forested sand ridges and knolls (U.S. Fish 

and Wildlife Service, 2009). The open fens have a diverse 

community of sedges, grasses, rushes, forbs, Sphagnum and 

other mosses, and low shrubs, with scattered, sparse patches 

of broadleaf cattail (Typha latifolia) in some areas. Speckled 

alder (Alnus incana spp. rugosa), bog birch (Betula pumila), 

and leatherleaf (Chamaedaphne calyculata) are the most 

common shrub species. The primary land cover types of 

nonforested lowlands in the Refuge are scrub/shrub lowland 

(27 percent), sedge/bluejoint (Carex/Calamagrostis canadensis; 10 percent), Sphagnum/leatherleaf (4 percent), and mixed 

emergents/grasses/forbs (3 percent) (U.S. Fish and Wildlife 

Service, 2009). Water conditions in wetlands are affected by 

natural flow patterns, which move northwest to southeast, 

and water management of man-made pools, which were 

constructed in the 1930s. Fens are usually shallowly flooded 

in spring and remain saturated into the fall. Soils are generally mucks and peats 1–3 meters (m) in depth, overlaid on 

wet sand, with interspersed sand dunes. Natural and modified 

drainages drain the area to the southeast into the Manistique 

River (not shown). More detailed descriptions of the Refuge 

landscape and habitats are provided in U.S. Fish and Wildlife 

Service (2009).
Study Area 3

Seney National Wildlife Refuge Natural Area

Pool

Seney National Wildlife Refuge boundary

Stream

Road

C–3 study block

Marsh Creek study block

EXPLANATION

C3 Pool

Marsh Creek

C–3 Pool

Marsh Creek

Pool

Sweeney Creek

Base map modified from U.S. Fish and Wildlife Service, 1:24,000

Universal Transverse Mercator projection, zone 16

North American Datum of 1983

0 5 MILES

0 1 2 3 4 5 KILOMETERS

1 2 3 4

86°20’ 86°15’ 86°10’ 86°05’ 86°00’ 85°55’

46°20’

46°15’

46°10’

Figure 1. Location of C3 and Marsh Creek study areas in Seney National Wildlife Refuge, in the upper peninsula of Michigan.

Our study focused on two areas in the Refuge that have a 

long history of research related to yellow rails (Stenzel, 1982; 

Bart and others, 1984; Burkman, 1993), have extensive fire 

histories, and were delineated as operational burn units by 

the Refuge: the area south of C3 Pool (hereafter referred to as 

“C3”) and south of Marsh Creek Pool (hereafter referred to 

as “Marsh Creek”) (fig. 1). These areas were predominantly 

sedge-bluejoint and scrub/shrub lowland of leatherleaf, speckled alder, and bog birch, with scattered wooded knolls. 

The climate of the Refuge is strongly affected by its 

location between Lake Superior and Lake Michigan (not 

shown). Average annual precipitation at the Refuge is 78 centimeters (cm); 32 cm fall during May–September (Michigan 

State Climatologist’s Office, 2012). Minimum and maximum 

temperatures range from −2 to 11 degrees Celsius (°C) in 

April and from 12 to 27 °C in July, respectively. The median 

growing season is 125 days (May 27 to September 25). During 

April–September 2006–10, the region experienced moderate 

to severe drought in 2006 (Palmer Drought Severity Index, 

range from −3.88 to −1.72), moderate drought to normal 

conditions in 2007 (−3.81 to 0.16), and moderately wet conditions in 2008 (+2.55 to +2.70). In 2010, the spring started 

in moderate to severe drought during March–May (−4.49 to 

−2.46), but heavy rains in June followed by consistent rains in 

July–September resulted in increasingly wet conditions (+1.56 

to +3.49) for the remainder of the growing season.
4 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Methods

Sampling Design and Site Selection

We used detailed National Land Cover Database data, 

based on interpretation of 2004 color-infrared aerial images 

(U.S. Fish and Wildlife Service, unpub. data, 2004), to 

select locations for eight 1-ha blocks in both C3and Marsh 

Creek. We first overlaid a 150-×150-m grid (1-ha core plus 

25-m buffer around the perimeter) over each area, oriented 

so that the grid lines were parallel and perpendicular to the 

predominant flow of water through the area (northwest to 

southeast). This grid delineated possible 150-×150-m blocks, 

identified by the Universal Transverse Mercator coordinates 

of the northernmost corner. Because a primary interest was 

how fire affected shrubs, we identified blocks that were in 

lowland shrub (LS–B, 25–60 percent shrub cover; and LS–C, 

60–90 percent shrub cover) and that excluded upland types. 

In each area, 12 to 16 of these blocks were selected at random 

for field inspection. During preliminary field visits in summer 

2006, we visually estimated cover of sedges, grasses, and 

shrubs and paired blocks based on estimated cover characteristics and proximity. Block pairs were less than (<) 150 m 

from one another to provide some flexibility in fire operations 

but close enough to be similar in soil and hydrology. After a 

second round of field inspections, we made the final selection of 8 paired blocks (4 pairs in each area) based on field 

characteristics: 3 pairs of sedge-dominated blocks with low 

shrub cover, 3 pairs dominated by sedges and grasses with 

high shrub cover, and 2 pairs with high density of old shrubs 

(representing extreme conditions of shrub cover) (figs. 2–3). 

We randomly selected one block in each pair to be burned and 

the other to serve as a control, as part of the BACI design. 

We established our sampling blocks in summer 2006. 

Each 1-ha block consisted of an outer 25-m buffer area and 

25 sampling points distributed in a 5×5 grid, 25 m apart 

(fig. 4). Each sampling point was marked with a short stake 

topped with an aluminum washer, which allowed field personnel to relocate points each year with a metal detector. We 

marked the outer four corners of the buffer area and the beginning and end of each of the five transects with tall quaking 

aspen (Populus tremuloides) poles and flagging. We recorded 

locations of the four outer corners and each sampling point 

using a global positioning system receiver (plus or minus [±] 

2–3 m). Marking the outer buffer helped fire operations and 

other field activities to avoid trampling in the block except 

during data collection and postfire inspection.

Because we chose to fit our experiment within the context 

of normal fire operations at the Refuge, season of burning was 

area specific and, thus, was affected by each area’s existing 

plant community and hydrology. Both C3 and Marsh Creek, 

however, have similar effects from managed pools and levees 

and similar fire histories; only one pair of blocks had been 

burned within the previous 5 years (table 1).

Data Collection

We collected full plant data on all eight blocks in Marsh 

Creek and six blocks in C3 during 2006–10. This BACI design 

provided for data collection two growing seasons before the 

burn (2006 and 2007) and three growing seasons (2008, 2009, 

and 2010) after burning; the unburned plot of each pair served 

as the control, and the burned plot of each pair the treatment 

(table 1). Data for MC–2, the only block burned in 2007, were 

excluded from analyses, but results are presented for comparison. We collected data during mid-July to early September 

each year, when plants had likely reached their maximum 

growth and before killing frosts. Data collection within that 

period was alternated between the two areas to minimize 

possible temporal effects. Information of water depths in May 

was obtained from a concurrent study (Austin and Buhl, 2013) 

at locations within 50 m of each block.

Data on plant cover (to species), litter depth, and graminoid height were recorded at each sample point using a modified point-intercept method (Elzinga and others, 1998). Field 

identification followed Crow and Hellquist (2000a, b), Chadde 

(2002), and Flora of North America Editorial Committee 

(2003). Taxonomy used here follows the National PLANTS 

database (U.S. Department of Agriculture, Natural Resources 

Conservation Service, 2017; see appendix 1 for a crosswalk 

table of scientific and common names of plant taxa found, for 

the PLANTS database as of the date accessed and Flora of 

North America Editorial Committee, 2003). We merged some 

species to genera where we were unable to consistently identify to species (Eriophorum, Equisetum, Galium, Rubus, Salix, 

and Vaccinium). Sphagnum was the dominant moss in both 

areas, but we did not distinguish this group from other mosses. 

We included creeping bentgrass (Agrostis stolonifera) with 

bluejoint because the two grass species were often difficult to 

separate without seed heads. A 2-m transect was 1 m southeast from each sample point. Each species touching along a 

single edge of a transect was recorded within each of 40 5-cm 

segments. For each segment, we also recorded the presence/

absence of Sphagnum and other mosses and open area (no 

vegetation or litter). At every fifth segment (eight measures per 

sampling point), we also recorded litter depth (in centimeters), 

height of litter above the solid substrate) and graminoid height 

(in centimeters) by firmly setting the end of a meter tape onto 

the substrate below any litter. Metrics were expressed as a 

percentage (number of hits per 40 segments) for each sample 

point. Graminoid height reflected actual plant height; points 

with no graminoids were excluded from analyses.

Shrubs in this community often have multiple stems, and 

the line-intercept method used for herbs (as described above) 

would not accurately reflect their canopy cover. We instead 

used shrub patch number and shrub patch size (Harrell and 

Fuhlendorf, 2002) to provide a measure of shrub occurrence, 

cover, and height by species as well as a measure of horizontal patchiness. These measures are similar to those used by 
Methods 5

White (1965) to measure importance of shrub species. We 

located a 5-m transect 1 m northwest of each sampling point, 

laid adjacent to the baseline transect to minimize trampling 

effects. We used standard line-transect methods (Elzinga and 

others, 1998) to record the length of each shrub patch (to the 

nearest centimeter) along each transect for each species. A 

distinct shrub patch was defined as greater than or equal to 

10 cm long and continuous (canopy break <10 cm). We also 

recorded lengths of overlap of cover between shrub species 

and maximum height of each shrub patch (to the nearest 

5 cm). For each sampling point, we summarized shrub data 

as average patch height, size, and number of patches and 

86°09’ 86°08’30” 86°08’ 86°07’30” 86°07’

46°18’

46°17’30”

46°17’

Land cover type

EXPLANATION

Upland

Lowland shrub-B

Lowland shrub-C

Lowland shrub-D

Marsh-C

Marsh-D

Open water

Sedge-bluejoint-B

Sedge-bluejoint-C

Sedge-bluejoint-D

Sphagnum-Leatherleaf-B

Spaghnum-Leatherleaf-C

Sphagnum-Leatherleaf-D

Speckled alder-D

Willow-B

C–3 Pool

C3–3

C3-4

C3–5

C3–8

C3–1

C3–7

C3–2 C3–6

Marsh Creek

Sweeney Creek

Base map modified from U.S. Fish and Wildlife Service, 

National Land Cover data, 2004, 1:24,000

Universal Transverse Mercator projection, zone 16

North American Datum of 1983

0 1 MILE

0 0.5 1 KILOMETER

0.5

Figure 2. Location of study blocks at the C3 study area relative to cover types in Seney National Wildlife Refuge, Michigan. 

For cover types, B indicates 25–60 percent cover of the dominant vegetation; C indicates 60–90 percent cover of the 

dominant vegetation, and D indicates 100 percent cover of the dominant vegetation. Land cover data from the U.S. Fish 

and Wildlife Service.
6 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge 86°04’ 86°04’30” 86°05’ 86°05’30” 86°06’ 46°11’ 46°10’30” 46°10’ Upland Land cover types Open water Lowland shrub-B Lowland shrub-C Lowland shrub-D Marsh-D Sedge-bluejoint-B Sedge-bluejoint-C Sedge-bluejoint-D Submerged vegatation-B Speckled alder-B Speckled alder-C Speckled alder-D Marsh Creek Pool MC–3 MC–4 MC–5 MC–8 MC–1 MC–7 MC–2 MC–6 Base map modified from U.S. Fish and Wildlife Service, National Land Cover data, 2004, 1:24,000 Universal Transverse Mercator projection, zone 16 North American datum of 1983 EXPLANATION

0 0.5 0.25

0 0.5 0.25 0.75 KILOMETER

0.75 MILE

Figure 3. Location of study blocks at the Marsh Creek study area relative to cover types in Seney National Wildlife Refuge, Michigan. For cover types, B indicates 25–60 percent cover of the dominant vegetation; C indicates 60–90 percent cover of the dominant vegetation, and D indicates 100 percent cover of the dominant vegetation. Land cover data from U.S. Fish and Wildlife Service.
Methods 7

percent cover (summed length per 500 cm) by species. For 

each species, we summarized data as average patch height, 

size, and number, and percent cover (summed length/500 cm). 

We also calculated total percent shrub cover (summed lengths 

of each species minus overlaps/500 cm) and overall average 

height and number of shrub patches (expressed as number per 

5-m transect). 

Vertical cover or visual obstruction of the vegetation was 

measured using a cover board (Nudds, 1977), 30 cm wide, 

1.5 m tall, and marked in six 33-cm vertical increments, or 

“strata,” demarked by alternating white and red strips. The 

observer placed the cover board 1 m beyond the 2-m plant 

sampling line and estimated the percent of each stratum not 

visible (visual obstruction reading [VOR]) from 5 m away in 

1–1 1–2 1–3 1–4 1–5

2–1 2–2 2–3 2–4 2–5

3–1 3–2 3–3 3–4 3–5

5–1 5–2 5–3 5–4 5–5

4–1 4–2 4–3 4–4 4–5

N

EXPLANATION

Buffer area

Outer corner marking post

Transect-point identifier

Shrubline transect

Sample point

2-meter point-intercept line

Cover board point

5–5

Figure 4. Layout of transects and sampling points, 

spaced 25 meters apart, and associated identifying 

numbers, within each experimental block. Shaded 

area represents buffer area (25 meters wide) around 

sampling transects.
8 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

each of the four cardinal directions. The four readings for each 

sampling point were averaged for each stratum.

Prescribed Burn Treatments

Prescribed burns in each study area (Refuge burn unit) 

were implemented following normal Refuge operations. 

Blocks assigned to the control treatment were prevented 

from burning by compacting the vegetation around the block 

(outside the buffer area) into existing surface moisture using 

a large tracked, amphibious vehicle that exerted low ground 

pressure (1.0–1.5 pounds per square inch). The compaction 

line was monitored during the fire, and creeping fire was 

extinguished with water as needed to prevent fire intrusion. 

Burned blocks were allowed to burn normally as part of the 

larger fire operation. The burns were completed using ground 

ignition using drip torches to create a backing and flanking fire 

around the perimeter followed by a head fire, with additional 

fire application applied as needed. Weather parameters during 

the burns (appendix table 2.1) were recorded hourly on site 

and at the Refuge’s remote automated weather station (available at Great Lakes Fire and Fuels, https://glff.mesowest.org); 

additional observations were recorded periodically in the field.

Within 7 days after each burn, we recorded fire effects to 

vegetation, Sphagnum, and litter and noted moisture conditions. We visually estimated burn severity and percent of the 

block affected for the main fuel types (graminoids and forbs, 

leatherleaf, speckled alder, bog birch, willow [Salix], tamarack 

[Larix laricina], Spirea, and other). Burn severity for each 

type was categorized as unburned, scorched, lightly burned, 

moderately burned, or heavily burned (National Park Service, 

2003). We also visually estimated how much of the litter layer 

was consumed ([1] all of that year’s growth and last year’s 

litter, [2] also consumed 2-year-old litter [partially degraded 

material but stems still visible], [3] only undefined peat mat 

remained, [4] ashed down to nonpeat substrate). We rated 

how the burn affected Sphagnum ([1] surface singed or lightly 

burned but green still visible on surface; [2] surface burned but 

interior still green; [3] burned, no green detected in interior, 

at least partially consumed). Average water conditions in the 

block were recorded for water depth (in centimeters above the 

surface) for pooled water; water conditions of coarse surface 

peat (0–10 cm below surface) and fine-particle subsurface 

peat (greater than [>] 10 cm below surface) were categorized 

as saturated to dry. We also noted the phenological status of 

vegetation (for example, bud development, leave emergence, 

and height of new nonwoody growth in spring; stage of senescence or leaf fall in autumn) immediately before the burn.

The C3 burn unit was burned in early spring under 

moderate fire conditions for spring (appendix table 2.1). At 

the time of the fire, new growth of graminoids was 3–10 cm 

high, leaf buds of shrubs were beginning to break, and willows 

had partially opened catkins. Surface and subsurface peat 

were saturated, and some areas had surface water as much as 

5 cm deep. Postburn inspections of the burned blocks (fig. 5) 

indicated 95–98 percent of current nonwoody vegetation and 

Table 1. Fire history, status, and timing of prescribed burning on study blocks in the C3 and Marsh 

Creek (MC) study areas, Seney National Wildlife Refuge, Michigan, 2006–10.

[Pre, pretreatment; numbers in 2008–10, the number of growing seasons postburn]

Block Year of previous 

fire 2006 2007 2008 2009 2010

C3–1 2003 Pre Pre Control Control Control

C3–2 2003 Pre Pre Burned May 8 1.5 2.5

C3–3 1976 Pre Pre Burned May 8 1.5 2.5

C3–4 1976 Pre Pre Control Control Control

C3–5 1976 Pre Pre Burned May 8 1.5 2.5

C3–6 1976 Pre Pre Control Control Control

MC–1 1992 Pre Pre Control Control Control

MC–2 1992 Pre Burned July 17 1 2 3

MC–3 1991 Pre Pre Control Control Control

MC–4 1976 Pre Pre 1Control Control Control

MC–5 1976 Pre Pre Burned July 24 1 2

MC–6 1976 Pre Pre Control Control Control

MC–7 1976 Pre Pre Burned July 24 1 2

MC–8 1976 Pre Pre Control Control Control

1

MC–4 was originally designated as a burn block in the study design but would not burn during the 2008 fire and, 

therefore, was reclassified as a control block for analyses.
Methods 9

A B

C D

E F

Figure 5. Conditions after the prescribed burn on the C3 study area, May 2008. A, Aerial photograph of the C3 study area and pool; 

B, alder brush torching; C, fire front moving through sedge and cattail; D, transect line in C3 showing effect of foot traffic on litter 

consumption; E, scorched shrubs in C3 the day after the burn; F, technician examining conditions in an unburned block 1 month after the 

C3 burn. Aerial photograph by the Michigan Department of Natural Resources; other photographs by J.E. Austin, U.S. Geological Survey.
10 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

50–75 percent of the previous year’s litter were consumed, 

80–100 percent of shrubs were scorched, and most Sphagnum

hummocks were only singed or the top burned. Fire did not 

carry into C3–7 because of heavy shrub cover, low amount of 

fine fuels, and damper microclimate. Because we were unsuccessful in that burn treatment and constrained by man-hours, 

we dropped the paired C3–7 and C3–8 blocks from subsequent 

data collection and data analyses.

Two summer burns were completed. The Marsh Creek 

burn unit was planned as a summer 2007 burn. A 21-ha area 

encompassing MC–2 was burned on July 17, 2007, under high 

fire danger conditions for summer (table 1, fig. 6; appendix 

table 2.1). Surface and subsurface peat were damp but not 

saturated (some water drops emitted when a sample was 

compressed), and no surface water was present; pooled water 

was detected 12 cm below the surface. Postburn inspection 

indicated a burn of moderate severity: 90–100 percent of 

the nonwoody vegetation and the previous year’s litter were 

consumed, 50–80 percent of shrubs were scorched or lightly 

burned, and 60 percent of Sphagnum clumps were burned. 

Most Sphagnum hummocks were either surface burned 

(30 percent) or burned (60 percent). The Refuge was unable 

to safely burn the rest of the Marsh Creek unit as planned 

that year because of intensification of drought conditions and 

large wildfires in the region. The remaining parts of that burn 

unit were burned on July 24, 2008, under moderate conditions 

for summer (table 1, fig. 7; appendix table 2.1). Surface and 

subsurface peat were saturated, and some areas had surface 

water as much as 7 cm deep. Postburn inspection of blocks 

indicated that MC–5 and MC–7 experienced light to moderate burns, with variable burn severity. An estimated 80 to 

90 percent of current nonwoody vegetation and the previous 

year’s litter were consumed, and 70–80 percent of Sphagnum

clumps were surface burned. Most shrubs were unburned 

or scorched, but 25–30 percent of leatherleaf was severely 

burned compared to 0–10 percent for alder, birch, and willow. 

A B

C D

Figure 6. Conditions after the prescribed burn on the MC–2 block of the Marsh Creek study area, July 2007. A, Fire line between 

burned and unburned vegetation, established using an amphibious track vehicle; B, botanist examining results of newly burned MC–2 

block; C, burned Sphaghum clump; D, green-up of vegetation by September. Photographs by J.E. Austin, U.S. Geological Survey.
Methods 11

A B

C D

E Figure 7. Conditions after 

prescribed burn on the 

Marsh Creek study area, July 

2008. A, Fire crew using drip 

torches to ignite vegetation 

the burn perimeter; B, initial 

fire conditions along the east 

fire-line perimeter of the burn; 

C, patchy consumption of green 

vegetation; D, fire advancing 

through shrubs and graminoids; 

E, color-infrared aerial 

photograph of Marsh Creek 

about 1 week after burning; 

reddish colors indicate green 

vegetation (unburned areas). 

Aerial photograph by the 

U.S. Fish and Wildlife Service; 

other photographs by J.E. 

Austin, U.S. Geological Survey.
12 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

MC–4, in contrast, was burned lightly only on its western edge 

despite the absence of surface water and repeated efforts to 

ignite fuels in the block. In that block, 90 percent of graminoids and forbs and 85–100 percent of shrubs were unburned, 

and only 10 percent of the 2-year-old litter was consumed; 

5 percent of Sphagnum hummocks were singed and 5 percent 

were surface burned; hence, this block was redesignated as a 

control block for analyses.

Data Analyses

In general, the study design would be considered a 

repeated-measures design with various levels of subsampling completed within each block depending on the specific 

response variable quantified (Milliken and Johnson, 2009). 

The study also is in a BACI design (Underwood, 1991). Here, 

the 1-ha blocks (that is, study plots) are considered random 

and nested within each of the four treatments (C3 unit spring 

treatment includes C3 control and C3 spring burn; Marsh 

Creek unit summer treatment includes MC control and MC 

summer burn). As noted above, MC–2 was excluded from 

analyses because it was burned in 2007 rather than with the 

other burn treatments in 2008, MC–4 was reclassified as 

a control because it was only slightly burned on one edge, 

and the paired C3–7 and C3–8 blocks were dropped because 

fire did not penetrate C3–7. The following blocks were used 

in analyses:

• C3 spring burn includes C3–2, C3–3, C3–5 (number of 

samples [n] =3)

• C3 control includes C3–1, C3–4, C3–6 (n=3)

• MC summer burn includes MC–5, MC–7 (n=2)

• MC control includes MC–1, MC–3, MC–4, MC–6, 

MC–8 (n=5)

We considered the 5 baseline transects (which ran 

perpendicular to the general water-flow gradient), the 

5 sampling points along each transect, and the 40 5-cm 

segments within transect-point locations (where applicable) 

as subsamples within each study block. With the exception 

of the environmental variables water depth, litter depth, and 

graminoid height, all response variables and analyses were 

summarized across these subsamples to compute means and 

standard deviations at the block level because the blocks are 

considered the experimental unit for the burn and control 

treatments. Transect was incorporated as a factor in analyses 

for water depth, litter depth, and graminoid heights because 

we were interested in assessing a gradient effect within the 

study areas. The pairing of study blocks within C3 and MC 

was not used in analyses. Year was considered the repeatedmeasures factor because each block was measured across 

5 years (2006–10): 2 pretreatment years (2006–7) and 3 postburn years (2008–10). Because the VOR was quantified at six 

height levels at each sampling point, height was considered a 

third factor in a split-plot structure.

For analyses, we combined data for some closely related 

or similar species, resulting in 96 taxa in analyses:

• CAREX includes Carex lasiocarpa, C. oligosperma, 

C. interior, and four narrow-leaved Carex species that 

could not be identified to species (recorded in 2006 and 

2007); hereafter referred to collectively as “narrowleaved sedges.”

• CARAQU includes Carex aquatilis and an unknown 

wider-leaved Carex found in 2006. 

• CALCANAG includes Calamogrostis canadensis and 

Agrostis stolonifera.

• ERIOPH includes cottongrasses Eriophorum angustifolium and E. viridicarinatum.

• GALIUM includes bedstraws Galium labradoricum

and G. trifidum.

• RUBUS includes all Rubus berries (Rubus acaulis, 

R. hispidus, R. pubescens, and unidentified Rubus

species).

• SALIX includes all willows (Salix).

• VACCIN includes all blueberries (Vaccinium macrocarpon, V. myrotillides, and unidentified Vaccinium

species).

• VIOLAS includes violets Viola nephrophylla and 

V. macloskeyi.

We developed specific contrasts for a priori planned 

comparisons among the treatments while utilizing all data 

from all blocks (Milliken and Johnson, 2009). We analyzed 

only those taxa that were detected more than 100 times in the 

65,000 total sampling points-by-year combinations. For each 

area, we did within- and across-year comparisons of control 

and burn blocks when analysis of variance (ANOVA) results 

indicated significant effects or interactions. To accommodate 

the repeated measures on the blocks, we assumed an autoregressive lag 1 error structure in generalized linear models for 

ANOVAs (Littell and others, 2007). When numerical convergence was present, we used the Kenward-Roger correction to 

degrees of freedom to accommodate the unequal number of 

blocks per treatment and the random effects for all ANOVAS 

(Stroup, 2012); no adjustments were made when numerical 

computations would not converge. Preliminary analyses did 

not indicate consistent spatial correlation within and among 

blocks, so we did not make any accommodations for this in 

the ANOVAs (Gbur and others, 2012). We used SAS software 

(SAS Institute, Inc., 2013) to complete all data processing and 

analyses, in particular, the generalized linear mixed model 

procedure (PROC GLIMMIX). We assumed a beta distribution for all percentage response variables and normality for 

all others (Gbur and others, 2012). Because of inconsistencies 
Results 13

in methods of shrub data collection in 2008 and 2009, those 

2 years were excluded from analyses of shrub cover data. 

We also examined simple correlations among the means and 

standard deviations of the three habitat variables (water depth, 

litter depth, and graminoid height).

Shifts in plant community composition were examined 

using nonmetric multidimensional scaling (NMS) ordinations 

of mean plant cover from each block-by-year combination. We 

reduced the effect of very uncommon plant species by only 

using species that were detected in more than 3 percent of total 

sampling points within each block. As with other response 

variables, NMS axis scores also were used in ANOVAs to 

assess burn effects on the plant community in general; correlations with individual plant taxa were used to guide interpretation of the axes. For descriptive purposes, we plotted the NMS 

scores of three axes and connected succeeding years of each 

of the four treatments with arrows to examine trends over 

time in plant communities. Because NMS axes are linear, we 

could estimate changes from year to year by subtracting NMS 

scores from one year to the succeeding year (for example, 

2007, before burning, to 2008, after burns). The unit measures 

of scores or the change in scores themselves do not carry 

meaning; rather, we were interested in the magnitude of the 

change from zero and in comparing that magnitude relative 

to that of idle control plots. We used the program PC–ORD 

(McCune and Mefford, 2011) with Sorensen distance measure 

in the NMS ordination. We followed guidelines in Peck (2010) 

to determine dimensionality of the NMS by utilizing default 

settings in PC–ORD from multiple runs under medium autopilot settings.

Data analyzed during this study are available as a 

U.S. Geological Survey data release (Austin, 2018).

Results

Environmental and Habitat Variables

As expected, means of water depth, litter depth, and 

graminoid height were strongly correlated with their respective standard deviations (r; r=0.48 to 0.92); however, only 

moderate correlations were detected among these variables 

(table 2). Notably, mean and variability of litter depth were 

correlated with mean and variability of water depth (r=0.34 

and 0.53, respectively; p<0.01).

The year × treatment interaction was significant for both 

means and within-block standard deviations (hereafter referred 

to as “variability”) of water depth, litter depth, and graminoid 

height (table 3), indicating differences among the treatments varied with year. The water-flow gradient within each 

block (across the five baseline transects) was not significant 

(p>0.05), nor did it interact with year or treatment.

Water depth patterns in both areas were generally similar 

among treatments, with little standing water in summer 2006, 

2007, or 2009; some standing water in 2008; and highest 

levels of standing water in 2010. Similarly, May water conditions, measured in each study area during yellow rail surveys 

(J.E. Austin, unpub. data, 2009), were highest in 2008 (7.2 and 

6.6 cm for C3 and Marsh Creek, respectively) and 2009 (4.4 

and 7.3 cm for C3 and Marsh Creek, respectively) and lowest 

in 2007 (3.0 and 1.1 cm for C3 and Marsh Creek, respectively). Ancillary information indicates May water depths in 

2006 were similar to those in 2007, and those in 2010 were 

similar to or higher than those measured in 2008. Mean and 

variability of water depth within years and areas did not differ 

between the burn and the control blocks (fig. 8). In 2010, 

summer-burned blocks in Marsh Creek had a somewhat higher 

mean water depth (4.85±0.44 versus 3.09±0.28 cm) and variability (3.63±0.25 versus 2.45±0.16) than control blocks there. 

Table 2. Correlations among mean and variability (standard deviation) of water depth, litter depth, and graminoid height measures 

in study blocks in C3 and Marsh Creek study areas, Seney National Wildlife Refuge, Michigan, 2006–10.

[The probability (p-value) is in parentheses; the number of samples is 325 block by treatment by transect combinations.*, significant at α=0.05; --, no data]

Mean litter depth Mean graminoid height Standard deviation, 

water depth

Standard deviation, 

litter depth

Standard deviation, 

graminoid height

Mean water depth

0.34 (<0.01)* −0.06 (0.26) 0.92 (<0.01)* 0.49 (<0.01)* 0.08 (0.13)

Mean litter depth

-- 0.11 (0.04)* 0.32 (<0.01)* 0.71 (<0.01)* 0.27 (<0.01)*

Mean graminoid height

-- -- −0.08 (0.17) 0.12 (0.03)* 0.48 (<0.01)*

Standard deviation, water depth

-- -- -- 0.53 (<0.01)* 0.17 (<0.01)*

Standard deviation, litter depth

-- -- -- -- 0.29 (<0.01)*
14 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Table 3. Results of analysis of variance for habitat and environmental variables on study blocks in C3 and Marsh Creek study 

areas, Seney National Wildlife Refuge, Michigan, 2006–10.

[p, probability; *, significant at α=0.05]

Source of variation Numerator degrees 

of freedom

Denominator degrees 

of freedom F-statistic p-value

Mean water depth

Treatment 3 9 1.13 0.39

Transect 4 36 0.34 0.85

Treatment×transect 12 36 0.25 0.99

Year 4 180 82.80 <0.01*

Year×treatment 12 180 6.30 <0.01*

Year×transect 16 180 0.30 0.99

Year×treatment×transect 48 180 0.16 0.99

Standard deviation, water depth

Treatment 3 9 0.83 0.51

Transect 4 36 0.66 0.63

Treatment×transect 12 36 0.22 0.99

Year 4 180 196.10 <0.01*

Year×treatment 12 180 5.93 <0.01*

Year×transect 16 180 0.39 0.98

Year×treatment×transect 48 180 0.19 0.99

Mean litter depth

Treatment 3 9 13.02 <0.01*

Transect 4 36 0.14 0.97

Treatment×transect 12 36 0.30 0.98

Year 4 180 80.59 <0.01*

Year×treatment 12 180 21.54 <0.01*

Year×transect 16 180 0.41 0.98

Year×treatment×transect 48 180 0.42 0.99

Standard deviation, litter depth

Treatment 3 9 5.92 0.02*

Transect 4 36 0.92 0.46

Treatment×transect 12 36 0.18 0.99

Year 4 180 160.84 <0.01*

Year×treatment 12 180 5.50 <0.01*

Year×transect 16 180 0.68 0.81

Year×treatment×transect 48 180 0.37 0.99

Mean graminoid height

Treatment 3 9 9.83 <0.01*

Transect 4 36 0.53 0.72

Treatment×transect 12 36 0.49 0.91

Year 4 180 85.28 <0.01*

Year×treatment 12 180 9.41 <0.01*

Year×transect 16 180 0.85 0.63

Year×treatment×transect 48 180 0.86 0.72

Standard deviation, graminoid height

Treatment 3 9 5.97 0.02*

Transect 4 36 1.57 0.20

Treatment×transect 12 36 1.40 0.21

Year 4 180 29.06 <0.01*

Year×treatment 12 180 14.69 <0.01*

Year×transect 16 180 0.67 0.82

Year×treatment×transect 48 180 0.72 0.91
Results 15

In MC–2, mean water depth in summer increased from 0 cm in 

2006 and 2007 to >7 cm in 2008–10 and was markedly higher 

during 2008–10 than in other experimental blocks.

In C3, mean and variability of litter depth did not differ 

between control and burn blocks within each year, although 

mean litter depths were somewhat lower in spring-burned 

blocks in 2008 (fig. 9). In Marsh Creek, mean litter depth 

did not differ between control and burn blocks in 2006 but 

was 43 percent higher in control blocks in 2007. Litter depth 

in control blocks remained higher than in the burn blocks in 

the years after burning; by 2010, litter depth was 66 percent 

higher than 2008 levels in both treatments. Both Marsh Creek 

treatments had greater litter depths by 2010 than recorded 

in 2007. Litter-depth variability was 47–51 percent lower in 

summer-burned blocks in the first 2 years after burning than 

in control blocks (2008, 1.81±0.49 versus 3.45±0.31; 2009, 

A

B Water depth, in centimeters Standard deviation of water depth, in centimeters

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

Year

2006 2007 2008 2009 2010

8

6

4

2

0

10

3

2

1

0

4

Preburn Postburn

Preburn Postburn

C3 control

C3 spring burn

MC control

MC summer burn

EXPLANATION

Figure 8. Summer water depths in control and burn blocks in C3 and Marsh 

Creek (MC) blocks, 2006–10, at Seney National Wildlife Refuge, Michigan. 

A, Least-squares means (standard error) of water depth across years; B, 

standard deviation (standard error) of water depths across years. Asterisks 

indicate significant difference (probability <0.05) between burn and control 

blocks within an area and year.
16 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

C3 control

MC–2 summer burn

C3 spring burn

MC control

MC summer burn

EXPLANATION

Year

2006 2007 2008 2009 2010

15

10

5

0

20

8

6

4

2

0

10

Preburn Postburn

Preburn Postburn

A

B

Litter depth, in centimeters Standard deviation of litter depth, in centimeters

C3 control

C3 spring burn

MC control

MC summer burn

EXPLANATION

Figure 9. Litter depths in control and burn blocks in C3 and Marsh Creek (MC) blocks, 2006–10, at 

Seney National Wildlife Refuge, Michigan. A, Least-squares means (standard error) of water depths 

across years; B, Standard deviation (standard error) of water depths across years. C3 was burned in 

May 2007 and Marsh Creek in July 2007. The MC–2 block was burned in July 2007. Asterisks indicate 

significant difference (probability <0.05) between burn and control blocks within an area and year.

2.22±0.49 versus 4.51±0.31) but similar thereafter; differences 

were significant only in 2009. Mean and variability of litter 

depths in all four treatments indicated broad similarities across 

years, generally decreasing to 2008 then increasing again. 

In MC 2, litter was entirely removed by the July 2007 burn, 

remained minimal (0.12 cm) in 2008, and then increased to 

17.0 cm by 2010.

Trends in graminoid heights across the years were generally similar in all treatments except the summer-burned blocks 

in Marsh Creek. No differences were detected in mean or 

variability of graminoid heights between control and springburned blocks in C3 within any of the 5 years but trended 

lower in posttreatment years (fig. 10). In Marsh Creek, mean 

graminoid heights did not differ between treatments in the 

pretreatment years (2006, 2007), but they were 84 percent 
Results 17

C3 control

MC–2 summer burn

C3 spring burn

MC control

MC summer burn

EXPLANATION

Year

2006 2007 2008 2009 2010

50

40

20

30

10

0

60

30

20

10

0

40

Preburn Postburn

Preburn Postburn

C3 control

C3 spring burn

MC control

MC summer burn

EXPLANATION

A

B

Graminoid height, in centimeters Standard deviation of graminoid height, in centimeters

Figure 10. Graminoid height (mean and standard error) in control and burn blocks in C3 and Marsh 

Creek (MC) blocks, 2006–10, at Seney National Wildlife Refuge, Michigan. A, Least-squares means 

(standard error) of water depths across years; B, standard deviation (standard error) of water depths 

across years. C3 was burned in May 2007 and Marsh Creek in July 2007. The MC–2 block was burned in 

July 2007. Asterisks indicate significant difference (probability <0.05) between burn and control blocks 

within an area and year.

lower in summer-burned blocks than in control blocks in 

2008. They remained 54 percent lower in 2009 but were not 

significantly different that year. No treatment differences 

were detected in 2010, 2 years after the burn. Variability in 

graminoid heights showed similar trends in Marsh Creek, with 

61 percent lower variability in summer-burned blocks in 2008 

and 26 percent lower variability in 2009 but no significant 

difference between treatments in 2009 or 2010. The marked 

decline in graminoid height and variability in summer-burned 

blocks between 2007 and 2008 reflects the short (3-week) 

period between burning and data collection and a combination 

of a patchy burn (some remaining graminoids) and rapid early 

regrowth. Graminoid height in MC–2 fell to 0 cm after the 

summer burn in 2007 (a complete burn), remained 21–27 cm 

lower than the 2006 heights in the 2 years after burning, and 

was 11 cm lower 3 years after burning.
18 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Plant Community

We identified a total of 104 plant taxa (excluding bryophytes) on the 14 blocks over the 5 years (90 identified taxa 

in C3 and 81 in Marsh Creek; see glossary and appendix 1 

for plant taxa scientific and common names). For combined 

areas, 38 percent of identified taxa were forbs; 25 percent 

were woody species; 21 percent were sedges, rushes, or 

cattail; 9 percent were grasses; and 7 percent were nonangiosperms (for example, ferns). Only one introduced species was 

detected (marsh thistle, Cirsium palustre); it was detected 

only in MC–7 sampling points but also was noted scattered 

throughout C3 outside of the experimental blocks in 2010. 

Most species (57 percent) were obligate wetland hydrophytes, 

35 percent were facultative wetland hydrophytes, 4 percent 

were facultative hydrophytes, 3 percent were facultative 

upland nonhydrophytes, and 1 percent was upland nonhydrophyte (Canada mayflower, Maianthemum canadense).

The plant communities of the two areas were generally similar, but some broad differences in taxa occurrence 

(appendix 3) are of note. C3 contained a greater diversity of 

woody taxa, whereas leatherleaf was detected almost entirely 

in Marsh Creek. Although we had difficulties in identifying 

some narrowed-leaved Carex to species in 2006 and 2007 

(hence the combination into narrow-leaved sedge), speciesspecific results indicate fewseed sedge (C. oligosperma) was 

common in Marsh Creek and rarely detected in C3, whereas 

woollyfruit sedge (C. lasiocarpa) seemed to be present mainly 

in C3; only in 2010 was it identified in Marsh Creek. Another 

narrow-leaved sedge, inland sedge (C. interior), was equally 

common in both areas and was likely among the unidentified 

taxa in 2006, particularly as unidentified CAREX2. Among the 

more common species, creeping sedge (C. chordorrhiza), livid 

sedge (C. livida), bedstraws (Galium spp.), harlequin blueflag 

(Iris versicolor), bog goldenrod (Solidago uliginosa), and 

northern bog aster (Symphyotrichum boreale) were detected 

more often in Marsh Creek, whereas spike muhly (Muhlenbergia glomerata) was detected more frequently in C3. White 

beak-rush (Rhynchospora alba) was detected only in C3–1 and 

C3–2. Broadleaf cattail, a species of concern in some areas for 

its potential to develop into monotypic stands, was detected at 

low frequency and only in C3–3 and C3–4, near marsh (fig. 2).

Species Richness and Diversity

In 2007, the year before burn treatments were applied, 

species richness per block averaged 24.7±2.4 in C3 control 

plots, 26.0±2.4 in C3 burn plots, 25.6±2.4 in Marsh Creek 

control plots, and 31.0±3.0 in Marsh Creek burn plots 

(fig. 11A). Species richness in C3 control and burn blocks and 

in Marsh Creek control blocks was somewhat lower in 2009 

than other years but generally recovered to or near pretreatment levels by 2010. Significant year × treatment interaction (table 4) for species richness was largely due to annual 

changes in Marsh Creek. Species richness there declined from 

2007 to 2008 in both control and summer-burned blocks by 19 

and 65 percent, respectively. Not surprisingly, species richness 

differed significantly between treatments there in 2008, when 

data were collected just a few weeks after burning. Species 

richness in MC–2 followed a similar pattern to other blocks 

(fig. 11A).

Similarly, the Shannon diversity index (H) differed 

marginally between burn and control treatments only in Marsh 

Creek in 2008 (year × treatment interaction; table 4, fig. 11B). 

After one full growing season after the 2008 burn at Marsh 

Creek, the diversity index in burned blocks had recovered to 

levels similar to those of 2007 in the burned blocks. The diversity index in C3 remained relatively constant across years.

Percent Cover by Taxa—Year and Treatment 

Effects

We examined data for effects of treatment, year, and 

year × treatment interactions for percent open and the 28 most 

commonly detected taxa. Of those taxa, 2 showed no effects 

of treatment, year, or their interaction, whereas 16 showed no 

differences between burn and control blocks within an area 

and year (table 4). We describe results by plant forms (sedges/

rushes, grasses, forbs, moss, woody plants) and for open area.

We detected a year × treatment interaction for percent 

cover of open area (OPEN; table 4). Open area in burned 

treatments was 300 percent higher than in control treatments 

in Marsh Creek in 2008 but did not differ for other years there; 

no treatment differences were detected in C3 (fig. 12). Burn 

and control blocks in both areas showed a gradual upward 

trend in open area from 0 percent in 2006 to 21–33 percent in 

2010. Open area in MC–2 changed little after recovering from 

the 2007 summer burn. Similar to other blocks, percent open 

in MC–2 was 0 in 2006 and higher in postburn years.

Year × treatment interactions were detected for six of 

the eight sedges and rushes (table 4). Percent cover of the 

narrow-leaved sedges (CAREX, fig. 13A) was highest in 

C3 but declined 44 and 49 percent in C3 control and burn 

blocks, respectively, between 2007 and 2010. In 2009, percent 

cover of narrow-leaved sedges in C3 was 59 percent higher 

in spring-burned blocks than in control blocks. In Marsh 

Creek, narrow-leaved sedge cover values were consistently 

lower in summer-burned blocks than control blocks, but 

differences were significant only in 2008 and 2009, when 

cover was 67–68 percent lower on summer-burned blocks. 

Cover of creeping sedge (CARCHO, fig. 13B) also indicate 

year × treatment interactions and was particularly variable in Marsh Creek. Cover trended higher in 2008 in all 

but Marsh Creek summer-burn blocks; in C3, percent cover 

across years declined to or below 2006 levels by 2010. Livid 

sedge (CARLIV, fig. 13C), detected mainly in Marsh Creek, 

was rarely detected in 2006, but we detected no changes in 

percent cover across years or differences between burn and 

control blocks there. Its absence in 2006 may have been due 

to difficulties in sedge identification the first year. Buxbaum’s 

sedge (Carex buxbaumii) was detected only in 2010 in Marsh 
Results 19

Creek, with little difference between burned and control 

blocks (CARBUX, fig. 13D); it was most abundant in MC–2 

in 2010. Cover of cottongrasses (ERIOPH, fig. 14A) was 

highest in Marsh Creek in 2006, but in later years, few differences were detected in percent cover among years or between 

treatments except immediately after the summer burn in 2008. 

By 2010 in Marsh Creek, three growing seasons later, its 

cover remained 76 percent below the 2007 levels in summerburned blocks and 68 percent below in control blocks. Elliptic 

spikerush (Eleocharis elliptica; ELEELL, fig. 14B) was most 

common in C3 control blocks across all years, but its cover 

was highly variable within blocks. We detected only a year 

effect; in 2008, percent cover in all blocks declined to less 

than 5 percent but largely recovered to 2006–7 levels by 2010. 

The steepest decline and recovery were detected in C3 control 

blocks. For the less common Robbins’ spikerush (Eleocharis 

robbinsii; ELEROB, fig. 14C), percent cover was higher in 

Marsh Creek burn blocks than control blocks in 2006 and 

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

Year

2006 2007 2008 2009 2010

35

30

25

15

10

20

5

0

40

2.5

2.0

1.5

1.0

0.5

0

3.0

Preburn Postburn

Preburn Postburn C3 control

C3 spring burn

MC control

MC summer burn

EXPLANATION

A

B

Number of species

H diversity

Figure 11. Measures of biological diversity detected in control and burn blocks in C3 and Marsh Creek 

study areas, 2006–10, at Seney National Wildlife Refuge, Michigan. A, Species richness; and B, Shannon 

diversity index (H). No data were collected in MC–2 in 2007, the year it was burned. Whiskers indicate 

standard errors. Asterisks indicate significant difference (probability <0.05) between burn and control 

blocks within an area and year.
20 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Table 4. Results of analysis of variance for species richness, species diversity index (Shannon diversity index, H), nonmetric 

dimensional scaling (NMS) scores on 3 axes, and percent cover of open, moss, and 27 plant taxa on study blocks in C3 and 

Marsh Creek study areas, Seney National Wildlife Refuge, Michigan, 2006–10.

[See appendix 1 for taxa codes. p, probability; *, significant at α=0.05]

Variable

F-statistic (p-value)

Treatment Year Year × treatment

Species richness 0.24 (0.87) 39.98 (<0.01)* 5.72 (<0.01)*

Shannon diversity index (H) 4.42 (0.04)* 12.45 (<0.01)* 3.26 (<0.01)*

NMS—Axis 1 9.39 (<0.01)* 13.52 (<0.01)* 5.62 (<0.01)*

NMS—Axis 2 18.59 (<0.01)* 36.66 (<0.01)* 527 (<0.01)*

NMS—Axis 3 2.49 (0.13) 51.19 (<0.01)* 3.79 (<0.01)*

Open 5.91 (<0.01)* 67.03 (<0.01)* 9.16 (<0.01)*

Moss 3.32 (0.07) 17.00 (<0.01)* 1.46 (0.19)

ANDGLA 1.78 (0.22) 2.79 (0.04)* 1.14 (0.36)

BETPUM 1.40 (0.30) 11.07 (<0.01)* 1.56 (0.15)

CALCANAG1 0.53 (0.68) 4.01 (<0.01)* 1.78 (0.09)

CAMAPA 1.49 (0.28) 14.35 (<0.01)* 2.66 (0.01)*

CARBUX 18.75 (<0.01)* 52.53 (<0.01)* 18.75 (<0.01)*

CARCHO 0.93 (0.46) 5.89 (<0.01)* 1.99 (0.05)*

CAREX 15.54 (<0.01)* 25.36 (<0.01)* 2.83 (<0.01)*

CARLIV 7.74 (<0.01)* 19.89 (<0.01)* 3.02 (<0.01)*

CHACAL 82.28 (<0.01)* 14.76 (<0.01)* 7.63 (<0.01)*

ELEELL 2.09 (0.17) 20.28 (<0.01)* 1.56 (0.15)

ELEROB 5.54 (0.02)* 11.42 (<0.01)* 4.26 (<0.01)*

ERIOPH 1.02 (0.43) 19.17 (<0.01)* 2.98 (<0.01)*

GALIUM 12.11 (<0.01)* 5.34 (<0.01)* 2.37 (0.02)*

HYPKAL 1.43 (0.30) 3.13 (0.03)* 1.29 (0.27)

IRIVER 1.96 (0.19) 1.70 (0.17) 2.41 (0.02)*

LYSTER 4.73 (0.03)* 8.44 (<0.01)* 1.31 (0.26)

MUHGLO 18.00 (<0.01)* 0.88 (0.48) 0.74 (0.70)

POTFRU 0.18 (0.91) 11.52 (<0.01)* 1.89 (0.07)

RHYALB 0.88 (0.49) 2.03 (0.11) 0.86 (0.59)

RUBUS 0.23 (0.88) 4.73 (<0.01)* 1.16 (0.35)

SALIX 8.13 (<0.01)* 17.69 (<0.01)* 1.56 (0.15)

SOLULI 1.22 (0.36) 5.74 (<0.01)* 2.62 (0.01)*

SYMBOR 1.65 (0.25) 124.89 (<0.01)* 2.20 (0.03)*

SYMPUN 0.27 (0.84) 54.09 (<0.01)* 5.90 (<0.01*)

TRIFRA 0.76 (0.54) 9.57 (<0.01)* 0.27 (0.99)

VACCINS 2.20 (0.16) 1.74 (0.16) 0.95 (0.51)

VIOLAS 1.29 (0.34) 69.85 (<0.01)* 1.30 (0.26)

1

PROC GLIMMIX would not converge using beta distribution; assumed distribution was normal.
Results 21

2007, but its cover declined to 0 after the summer burn and 

remained at low levels in all blocks through 2010. In contrast 

to other sedges and rushes, we detected no year, treatment, or 

year × treatment interactions for white beak-rush (RHYABL, 

fig. 14D), which was detected only in C3–1 (detected at 

75 percent of sample points) and C3–2 (14 percent of sample 

points); it was not detected in those blocks in 2010.

The two grasses examined differed in effects. Percent 

cover of bluejoint (CALCANAG) varied among years, with 

highest percent cover detected in MC–2 in 2006 (table 4, 

fig. 15A); its cover was lowest in 2008 after the summer burn. 

Spike muhly (MUHGLO, fig. 15B) was affected by treatment 

because of its greater abundance in C3 (table 4).

Among forbs, we detected year × treatment interactions 

for six taxa (table 4). Cover of marsh bellflower (Campanula 

aparinoides; CAMAPA, fig. 16A) trended upward from 2006 

to 2008 in all but Marsh Creek burn blocks, then declined to 

2006 levels by 2010. In C3, its cover was consistently higher 

on control than burn blocks, but differences were significant 

only in 2007. In Marsh Creek, its cover also tended to be 

higher on control than burn blocks across years, but differences were significant only in 2008. Cover in summer-burned 

blocks was 96 percent lower in control blocks that year, but 

by 2010, percent cover was similar between burn and control 

blocks. In MC–2, marsh bellflower cover was highest the 

year after burning but quickly declined to below preburn 

levels. Bedstraw (GALIUM; fig. 16B) cover was highest in 

Marsh Creek before burning; cover in burn blocks there did 

not recover to preburn levels by 2010. Harlequin blueflag 

(IRIVER, fig. 16C) was somewhat more common in Marsh 

Creek control blocks through 2008 but declined to levels 

similar to other blocks in 2009 and 2010. It was most abundant 

in MC–2 but also showed a decline in postburn years. Cover 

of bog goldenrod (SOLULI, fig. 16D) tended to be higher in 

Marsh Creek than in C3 before 2008 but declined thereafter to 

equivalent levels. We also detected year × treatment interactions for percent cover of the two asters examined, but the 

species showed different responses across years. Cover of 

northern bog aster (SYMBOR, fig. 16E) was higher in the 

preburn years, fell to near zero in 2008 and 2009 in all blocks, 

and remained 73–94 percent below 2007 levels by 2010. 

By 2010, its cover in summer-burned blocks in Marsh Creek 

was higher than in control blocks, but differences were not 

significant. In contrast, cover of purplestem aster (Symphyotrichum puniceum; SYMPUN, fig. 16F) in Marsh Creek spiked 

one growing season after the summer burn (2009) before 

largely disappearing in 2010 in all blocks. In C3, it was most 

abundant in 2007 and 2009 and was higher in burn blocks than 

in control blocks in 2007.

Of the forb taxa, four were affected only by year. Cover 

of Kalm’s St. Johnswort (Hypericum kalmianum; HYPKAL, 

fig. 17A), detected primarily in Marsh Creek, was somewhat 

lower in 2008 and 2009 than 2007. The high variability in the 

cover of shrubby cinquefoil (Potentilla fruticosa, POTFRU, 

fig. 17B) limited any ability to detect differences among treatments. Cover of Fraser’s marsh St. Johnswort (Triadenum 

fraseri; TRIFRA, fig. 17C) declined in both areas after 2007. 

Few violets (Viola spp.; <0.1 percent cover) were detected 

in 2006 and 2009, but cover was higher in most treatments 

in 2007 and 2010, with similar cover in spring- and summerburned blocks (VIOLA, fig. 17D). Within areas and years, 

violet cover was somewhat higher in C3 burn blocks in 2007 

and significantly higher in spring-burned blocks in 2008 

and 2010.

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

2006 2007 2008 2009 2010

90

80

70

60

40

30

20

50

10

0

100

OPEN Preburn Postburn

Mean percent cover

Year

Figure 12. Percent open area (leastsquares means, standard error) affected 

by year × treatment interactions in control 

and burn blocks in C3 and Marsh Creek 

(MC) blocks, 2006–10, at Seney National 

Wildlife Refuge, Michigan. Whiskers indicate 

standard errors. Asterisks indicate significant 

difference (probability <0.05) between burn 

and control blocks within an area and year.
22 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge Preburn Postburn Preburn Postburn Preburn Postburn C3 control C3 spring burn MC control MC summer burn MC–2 summer burn EXPLANATION A. CAREX C. CARLIV Preburn Postburn B. CARCHO D. CARBUX

Mean percent cover Mean percent cover

55504540353025201510

50

65601510

50

302520

Mean percent cover Mean percent cover

4540353025201510

50

50

6543210

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

Year

2009 2010 2008 2007 2006

Year

2009 2010 2008 2007 2006

Figure 13. Percent cover of sedges (least-squares means, standard error) affected by year × treatment interactions in control and burn blocks in C3 and Marsh Creek (MC) blocks, 2006–10, at Seney National Wildlife Refuge, Michigan. Whiskers indicate standard errors. 

A, Fine-leaved sedges (CAREX); 

B, creeping sedge (CARCHO); 

C, livid sedge 

(CARLIV); 

D, Bebb’s oval sedge (CARBUX). Asterisks indicate significant difference (probability <0.05) between burn and control blocks within an area and year.
Results 23

A. ERIOPH

C. ELEROB

Preburn Postburn

B. ELEELLD. RHYALB

Mean percent cover Mean percent cover

201510

50

25

543210

Mean percent cover Mean percent cover

3025201510

50

35

1.51.00.5

0

3.02.52.0

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

Year

2009 2010 2008 2007 2006

Year

2009 2010 2008 2007 2006

Preburn Postburn

Preburn Postburn Preburn Postburn

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

Figure 14. Percent cover of sedges and rushes (least-squares means, standard error) affected by year × treatment interactions in control and burn blocks in C3 and Marsh Creek (MC) blocks, 2006–10, at Seney National Wildlife Refuge, Michigan. Whiskers indicate standard errors. A, Cottongrasses (ERIOPH); B, elliptical spikerush (ELEELL); C, Robbins’ spikerush (ELEROB); D, white beak-rush (RHYALB). Asterisks indicate significant difference (probability <0.05) between burn and control blocks within an area and year.
24 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Year

2006 2007 2008 2009 2010

8

10

6

4

2

0

12

12

10

8

6

4

2

0

16

Preburn Postburn

Preburn Postburn

14

A. CALCANAG

B. MUHGLO

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

Mean percent cover Mean percent cover

Figure 15. Percent cover of grasses (leastsquares means, standard error) affected 

by separate year and treatment effects in 

control and burn blocks in C3 and Marsh 

Creek (MC) blocks, 2006–10, at Seney National 

Wildlife Refuge, Michigan. Whiskers indicate 

standard errors. A, Bluejoint (CALCANAG); 

B, spike muhly (MUHGLO). Asterisks indicate 

significant difference (probability <0.05) 

between burn and control blocks within an 

area and year.

We detected separate year and treatment effects for earth 

loosestrife (Lysimachia terrestris; table 4; LYSTER, fig. 18), 

indicating a mixed response to burning. Its cover increased 

from near 0 in 2006 to peak levels in 2008 in all but Marsh 

Creek burn blocks, then declined to near 0 in all blocks by 

2010. It was most abundant in MC–2 during 2008–10 but 

also declined there after 2008. In C3, its cover was higher 

in control than burn blocks the year before spring burns but 

not thereafter, whereas in Marsh Creek, cover was higher in 

control than in summer-burned blocks in 2008 and 2009.

Percent cover of moss indicated a year × treatment 

interaction (table 4; MOSS, fig. 19). Moss cover ranged from 

4 to 13 percent in 2006, 2007, and 2009. In C3, moss cover 

was 314 percent higher in spring-burned blocks in 2008 and 

164 percent higher in 2010 than in control blocks. Moss cover 

in Marsh Creek in 2010 also increased from earlier levels 

on both burned and control blocks; its cover tended to be 

higher on control blocks than burned blocks during 2008–10, 

but differences were not significant. Often, when the sample 

segment seemed devoid of vascular plants, moss was detected 

growing on decaying litter or peat.

A year × treatment interaction affected the percent cover 

of leatherleaf (table 4). Leatherleaf was rarely detected in 

C3 or MC–2 but was abundant elsewhere in Marsh Creek 

(CHACAL, fig. 20). Leatherleaf cover was consistently higher 

in blocks assigned to summer burns than in the control blocks, 

except after burning in 2008. Leatherleaf was 116 percent 

higher in burn blocks in 2007, fell 93 percent in burned blocks 

in 2008, but recovered to near preburn levels by 2010.
Results 25

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

A. CAMAPAC. IRIVER

Preburn Postburn

B. GALIUMD. SOLULI

Mean percent cover Mean percent cover

10

86420

12

543210

Mean percent cover Mean percent cover 2.5 2.0 1.5 1.0

0.5

0

2.52.01.51.00.53.0

0

3.0

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

Year

2009 2010 2008 2007 2006

Year

2009 2010 2008 2007 2006

Preburn Postburn

Preburn Postburn Preburn Postburn

Figure 16. Percent cover of forbs (least-squares means, standard error) affected by year × treatment interactions in control and burn blocks in C3 and Marsh Creek (MC) blocks, 2006–10, at Seney National Wildlife Refuge, Michigan. Whiskers indicate standard errors. A, Marsh bellflower (CAMAPA); B, bedstraws (GALIUM); C, harlequin blueflag (IRIVER); D, bog goldenrod (SOLULI); E, northern bog aster (SYMBOR); F, purplestem aster (SYMPUN). Asterisks indicate significant difference (probability <0.05) between burn and control blocks within an area and year.
26 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

E. SYMBOR

F. SYMPUN Preburn Postburn

Mean percent cover

9

8

7

6

5

3

2

1

4

0

10

Mean percent cover

6

5

4

3

2

1

0

7

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

Preburn Postburn

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

Year

2006 2007 2008 2009 2010

Figure 16. Percent cover of forbs (leastsquares means, standard error) affected by 

year × treatment interactions in control and 

burn blocks in C3 and Marsh Creek (MC) 

blocks, 2006–10, at Seney National Wildlife 

Refuge, Michigan. Whiskers indicate 

standard errors. A, Marsh bellflower 

(CAMAPA); B, bedstraws (GALIUM); 

C, harlequin blueflag (IRIVER); D, bog 

goldenrod (SOLULI); E, northern bog aster 

(SYMBOR); F, purplestem aster (SYMPUN). 

Asterisks indicate significant difference 

(probability <0.05) between burn and control 

blocks within an area and year.—Continued

We detected no year × treatment interactions for other 

woody taxa (table 4). Percent cover of bog rosemary (Andromeda polifolia var. glaucophylla) (ANDGLA, fig. 21A), raspberries (Rubus spp.; RUBUS, fig. 21B), and bog birch (BETPUM, 

fig. 21C) were affected only by year, whereas willows 

(SALIX, fig. 21D) were affected by separate year and treatment interactions. Cover of bog birch was consistently higher 

in C3 burn blocks than in control blocks and significantly 

higher in 2007 and 2010. Willows were primarily detected in 

C3; cover there peaked in control blocks in 2007 and 2010, but 

cover did not differ between burned and control blocks within 

the area. Vaccinium (VACCIN, fig. 21E) was rarely found in 

Marsh Creek blocks and somewhat more abundant in C3.
Results 27

A. HYPKALC. TRIFRA

Preburn Postburn B. POTFRU

D. VIOLA

Mean percent cover Mean percent cover

2.52.01.51.00.5

0

3.00.40.2

0

1.00.80.6

Mean percent cover Mean percent cover

543210543210

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

Year

2009 2010 2008 2007 2006

Year

2009 2010 2008 2007 2006

Preburn PostburnPreburn Postburn Preburn Postburn

C3 controlC3 spring burnMC controlMC summer burnMC–2 summer burn

EXPLANATION

Figure 17. Percent cover of forbs (least-squares means, standard error) affected by year effects in control and burn blocks in C3 and Marsh Creek (MC) blocks, 2006–10, at Seney National Wildlife Refuge, Michigan. Whiskers indicate standard errors. A, Kalm’s St. Johnswort (HYPKAL); B, shrubby cinquefoil (POTFRU); C, Fraser’s St. Johnswort (TRIFRA); D, violets (VIOLA). Asterisks indicate significant difference (probability <0.05) between burn and control blocks within an area and year.
28 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

Year

2006 2007 2008 2009 2010

2.5

2.0

1.5

1.0

0.5

0

3.0

LYSTER Preburn Postburn

Mean percent cover

Figure 18. Percent cover of earth 

loosestrife (LYSTER) (least-squares 

means, standard error) affected 

by separate year and treatment 

effects in control and burn blocks 

in C3 and Marsh Creek (MC) 

blocks, 2006–10, at Seney National 

Wildlife Refuge, Michigan. 

Whiskers indicate standard errors. 

Asterisks indicate significant 

difference (probability <0.05) 

between burn and control blocks 

within an area and year.

C3 control

MC–2 summer burn

C3 spring burn

MC control

MC summer burn

EXPLANATION

Year

2006 2007 2008 2009 2010

45

40

35

30

20

15

25

10

5

0

50

MOSS Preburn Postburn

Mean percent cover

Figure 19. Percent cover of 

mosses (MOSS) (least-squares 

means, standard error) affected 

by separate year and treatment 

effects in control and burn 

blocks in C3 and Marsh Creek 

(MC) blocks, 2006–10, at Seney 

National Wildlife Refuge, 

Michigan. Whiskers indicate 

standard errors. Asterisks 

indicate significant difference 

(probability <0.05) between burn 

and control blocks within an 

area and year.
Results 29

Year

2006 2007 2008 2009 2010

45

40

35

30

20

15

25

10

5

0

50

55

CHACAL Preburn Postburn

Mean percent cover

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

Figure 20. Percent cover of 

leatherleaf (CHACAL) (least-squares 

means, standard error) affected 

by year × treatment interactions in 

control and burn blocks in C3 and 

Marsh Creek (MC) blocks, 2006–10, 

at Seney National Wildlife Refuge, 

Michigan. Whiskers indicate 

standard errors. Asterisks indicate 

significant difference (probability 

<0.05) between burn and control 

blocks within an area and year.

Changes in Plant Community Indicated by 

Ordinations

The NMS results indicate that two to three axes were 

adequate for reducing the dimensionality of the plant cover 

data; axis 1 explained 71 percent, axis 2 explained an additional 12 percent, and axis 3 explained 8 percent. Combined, 

the three axes explained 91 percent of the plant cover data. 

NMS scores clearly differentiated C3 and Marsh Creek blocks 

(fig. 22), with generally more positive scores for C3 on axes 1 

and 2 and positive scores for Marsh Creek on axis 3 in all 

but 2008. Positive scores on axes 1 and 2 were most highly 

correlated with elliptic spikerush, spike muhly, and narrowleaved sedges (both axes); white beak-rush (axis 1); and 

Fraser’s marsh St. Johnswort (axis 2). Negative scores were 

most highly correlated with leatherleaf, creeping sedge, Harlequin blueflag (axis 1), and livid sedge (axis 2; table 5). On 

axis 3, positive NMS scores were most highly correlated with 

leatherleaf, bog rosemary, Robbins’ spikerush, and livid sedge 

and negative scores with marsh bellflower, spike muhly, earth 

loosestrife, and creeping sedge.

ANOVA of NMS scores detected year × treatment 

interactions for all three axes (table 4). Statistical comparisons 

between burn and control blocks within years revealed no 

differences in NMS scores for C3 in any year. In Marsh Creek, 

NMS scores were 70 percent lower in summer-burned blocks 

than control blocks on axis 2 in 2008 and 67 percent lower on 

axis 1 in 2009; differences between burn and control blocks 

on axis 3 also were apparent in 2007 and 2008 but were not 

statistically significant.

Shifts in the plant community across years are apparent 

by the trajectories of the NMS scores (fig. 22). Comparison of 

2007 and 2010 NMS means and confidence intervals indicate 

NMS axis 1 scores increased 62 percent for C3 control and 

87 percent for Marsh Creek burned blocks. For axis 2, scores 

decreased 58 percent for C3 burned blocks and (nonsignificantly) 44 percent for Marsh Creek burned blocks. These 

shifts represent changes after three growing seasons in C3 and 

two in Marsh Creek. Similar comparisons between 2007 and 

2009 NMS scores, representing two and one growing seasons, 

respectively, also indicated axis 1 scores were 53 percent 

higher in 2009 for C3 control blocks and axis 2 scores were 

114 percent higher in 2009 for Marsh Creek burned blocks.
30 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Year

2006 2007 2008 2009 2010

Year

2006 2007 2008 2009 2010

A. ANDGLA

C. BETPUM D. SALIX

Mean percent cover Mean percent cover

2.5

2.0

1.5

1.0

0

0.5

10

3.0

0.8

0.6

0.4

0

0.2

1.0

3

2

1

0

9

8

5

4

7

6

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

B. RUBUS

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

Preburn Postburn Preburn Postburn EXPLANATION

Preburn Postburn Preburn Postburn

2.5

2.0

1.5

1.0

0

0.5

3.0

E. VACCIN Preburn Postburn

2.5

2.0

1.5

1.0

0

0.5

3.0

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

C3 control

C3 spring burn

MC control

MC summer burn

MC–2 summer burn

EXPLANATION

Mean percent cover

Figure 21. Percent cover of woody species (least-squares 

means, standard error) affected by year in control and burn 

blocks in C3 and Marsh Creek (MC) blocks, 2006–10, at 

Seney National Wildlife Refuge, Michigan. Whiskers indicate 

standard errors. A, Bog rosemary (ANDGLA); B, raspberries 

(RUBUS); C, bog birch (BETPUM); D, willows (SALIX); 

E, Vacinnium species (VACCIN). Asterisks indicate significant 

difference (probability <0.05) between burn and control blocks 

within an area and year.
Results 31

EXPLANATION

C3 Control 2007C3 Spring burn 2007MC Control 2007MC Spring burn 2007C3 Control 2008C3 Spring burn 2008MC Control 2008MC Summer burn 2008C3 Control 2009C3 Spring burn 2009MC Control 2009MC Summer burn 2009C3 Control 2010C3 Spring burn 2010MC Control 2010MC Summer burn 2010

1.00.5

0

–0.5–1.0–1.5–2.0

1.5

Nonmetric dimensional

scaling axis 2

Nonmetric dimensional scaling axis 1

CAREXMUHGLOELEELLTRIFRA

CARLIV

CHACALCARCHOIRIVER

ELEELLRHYALBMUHGLOCAREX

–1.5 –0.5 0 0.5 1.0 1.5 –1.0

Nonmetric dimensional scaling axis 1 CHACAL

CARCHOIRIVER

CAREXMUHGLOELEELLTRIFRA

–1.5 –2.0 –0.5 0 0.5 1.0 1.5 –1.0

0.60.40.2

0

–0.2–0.4–0.6–0.8

0.8

Nonmetric dimensional

scaling axis 3

Nonmetric dimensional scaling axis 2

CHACALANDGLAELEROBCARLIV

CAMAPAMUHGLOLYSTERCARCHO

CARLIV

ELEELLRHYALBMUHGLOCAREX

–1.5 –0.5 0 0.5 1.0 1.5 –1.0

0.60.40.2

0

–0.2–0.4–0.6–0.8

0.8

Nonmetric dimensional

scaling axis 3

CHACALANDGLAELEROBCARLIV

CAMAPAMUHGLOLYSTERCARCHO

A

B

C

Figure 22.

Average annual nonmetric dimensional 

scaling (NMS) scores for plant community in control and burn blocks in C3 and Marsh Creek (MC), Seney National Wildlife Refuge, Michigan. Arrows indicate changes in average NMS scores in treatments from one year to the next, 2007–10. 

A, Axes 1 and 

2 scores; 

B, axes 2 and 3 scores; 

C, axes 1 and 3 

scores. Plant taxa noted along the axes are those with significant correlations (probability <0.05) to NMS score and Pearson’s rho <–0.40 or >0.40. ANDGLA, bog rosemary; CAMAPA, marsh bellflower; CAREX, narrow-leaved sedges; CARLIV, livid sedge; CARCHO, creeping sedge; CHACAL, leatherleaf; ELEELL, elliptic spikerush; ELEROB, Robbins’ spikerush; IRIVER, Harlequin blueflag; LYSTER, earth loosestrife; MUHGLO, spike muhly; RHYALB, white beak-rush; TRIFRA, Fraser’s marsh St. Johnswort.
32 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Table 5. Pearson correlation coefficients for 27 most common plant taxa with nonmetric dimensional scaling axes in study blocks in C3 

and Marsh Creek study areas, Seney National Wildlife Refuge, Michigan, 2006–10.

[See appendix 1 for taxa codes. r, Pearson correlation coefficient; p, probability]

Plant taxa

Axis 1 Axis 2 Axis 3

r p-value r p-value r p-value

ANDGLA −0.360 0.003 −0.190 0.129 0.450 <0.001

BETPUM −0.020 0.874 0.106 0.402 0.262 0.035

CALCANAG −0.041 0.748 0.265 0.033 −0.237 0.057

CAMAPA −0.089 0.479 −0.037 0.768 −0.620 <0.001

CARBUX −0.136 0.279 −0.189 0.132 0.257 0.038

CARCHO −0.661 <0.001 −0.114 0.367 −0.473 <0.001

CAREX 0.453 0.001 0.857 <0.001 −0.297 0.016

CARLIV −0.322 0.009 −0.778 <0.001 0.433 <0.003

CHACAL −0.705 <0.001 −0.321 0.009 0.546 <0.001

ELEELL 0.722 <0.001 0.484 <0.001 −0.235 0.060

ELEROB −0.315 0.011 −0.094 0.455 0.455 <0.001

ERIOPH −0.299 0.016 0.250 0.045 0.335 0.006

GALIUM −0.375 0.002 −0.104 0.409 0.276 0.026

HYPKAL −0.331 0.007 −0.106 0.400 0.290 0.019

IRIVER −0.466 <0.001 −0.249 0.046 0.102 0.420

LYSTER 0.145 0.249 0.050 0.694 −0.490 <0.001

MUHGLO 0.511 <0.001 0.593 <0.001 −0.496 <0.001

POTFRU −0.067 0.594 0.058 0.648 0.277 0.026

RHYALB 0.615 <0.001 0.189 0.131 −0.100 0.428

RUBUS 0.130 0.303 0.024 0.847 −0.172 0.170

SALIX 0.358 0.003 0.452 <0.001 −0.182 0.147

SOLULI −0.309 0.012 −0.030 0.813 0.359 0.003

SYMBOR −0.282 0.023 0.310 0.012 0.280 0.024

SYMPUN −0.154 0.221 −0.282 0.023 −0.095 0.452

TRIFRA 0.040 0.752 0.432 <0.001 −0.017 0.891

VACCINS 0.038 0.765 0.047 0.712 0.012 0.927

VIOLAS 0.055 0.666 0.120 0.340 0.257 0.039

Woody Height, Patchiness, and Cover

Comparisons of woody cover, number of patches, and 

height were limited to the 2 pretreatment years (2006, 2007) 

and the final posttreatment year (2010). We detected year and 

treatment effects for mean and standard deviation of height for 

leatherleaf, bog birch, and tamarack (table 6); however, the 

only significant difference between burn and control blocks 

within years was in C3 (greater leatherleaf height in control 

blocks in 2006 and 2010) (table 7). We detected no effects of 

year, treatment, or their interaction for heights of alder, willow, 

and other woody taxa.

Woody taxa varied in the number of woody patches by 

year × treatment interaction (leatherleaf and willow) and year 

(bog birch, other woody species) (table 8); however, we again 

detected no significant difference between burn and control 

blocks within years for either area. The greater prevalence 

of leatherleaf and other woody taxa in Marsh Creek drove 

the year × treatment results, whereas patches of willow were 

detected mainly in C3.

Percent cover differed among treatments for leatherleaf and tamarack; by year for bog birch; and by the year × 

treatment interaction for willow, other woody taxa, and total 

woody cover (tables 7 and 8). Once again, however, no withinyear differences between burn and control blocks were significant for either area. Leatherleaf cover tended to be higher in 

Marsh Creek burn blocks than in control blocks in pre- and 

posttreatment years. Treatment effect for tamarack was due 

to its absence in Marsh Creek. Cover of bog birch tended to 

be higher in 2007 and in C3. Willow and other woody taxa 
Results 33

were uncommon, but willow cover tended to be higher in C3, 

whereas the other woody taxa were more prevalent in Marsh 

Creek. Total woody cover was consistently but nonsignificantly higher in Marsh Creek burn blocks than in control 

blocks and somewhat higher in 2007 and 2010 than in 2006. 

In MC–2, total shrub cover was initially less than 4 percent 

and showed little change between 2007 and 2010, although 

number of patches measured declined from 13 to 7.

Vertical Cover

We detected significant interactions of year, treatment, and strata for mean and variability of VORs (table 9). 

VORs for strata 3–6 (>66 cm) tended to be higher in C3 

than in Marsh Creek, reflecting differences in shrub species 

and stature (figs. 23–24). No significant differences were 

detected between control and spring-burned plots in C3 

for any year, but across years, stratum 2 (34–66 cm) and 3 

(67–99 cm) seemed to be affected by the spring burn and 

trended downward in 2008 and 2009. In Marsh Creek, after 

the 2008 summer burn, the VOR of stratum 1 (0–33 cm) in 

burned blocks was 42 percent lower than in control blocks; it 

then increased 61 percent in 2009 before fully recovering to 

original levels in 2010. We also detected small but significant 

differences in mean VORs for stratum 5 (134–166 cm) and 6 

(167–200 cm) between control and burn blocks in the 2 years 

before burning (2006, 0.007±0.004 versus 0.001±0.001, 

respectively; 2007, 0.008±0.004 versus 0.001±0.002) (fig. 24). 

Changes in other strata after burning were nonsignificant. 

VOR variability differed between Marsh Creek control and 

burn blocks only in stratum 6, 2 years after burning (2009; 

control, 0.014±0.006; burn, 0.002±0.003).

In MC–2, mean VORs of strata 1 and 2 declined 50 and 

94 percent, respectively, before recovering by 2010 to levels 4 

and 41 percent higher than 2006, respectively (fig. 23). VORs 

for higher strata showed little change over the years (fig. 24).

Table 6. Results of analysis of variance for woody taxa height and standard deviation of woody height on study blocks in C3 and Marsh 

Creek study areas, Seney National Wildlife Refuge, Michigan, 2006, 2007, and 2010.

[p, probability; *, significant at α=0.05]

Source of 

variation

Numerator 

degrees 

of freedom

Denominator 

degrees 

of freedom

Mean height 

F-statistic 

(p-value)

Numerator 

degrees 

of freedom

Denominator 

degrees 

of freedom

Standard deviation 

of height

F-statistic (p-value)

ALDNIC

Treatment 3 4 2.20 (0.231) 2 3 2.27 (0.251)

Year 2 7 2.51 (0.151) 2 3 0.12 (0.893)

Year×treatment 4 7 2.39 (0.149) 4 3 0.91 (0.552)

BETPUM

Treatment 3 9 0.54 (0.669) 3 9 4.07 (0.044)*

Year 2 17 10.29 (0.001)* 2 16 2.35 (0.127)

Year×treatment 6 17 3.37 (0.023)* 6 16 1.08 (0.414)

CHACAL

Treatment 3 5 10.90 (0.012)* 3 5 5.15 (0.055)

Year 2 10 0.35 (0.710) 2 10 7.44 (0.100)

Year×treatment 5 10 1.05 (0.441) 5 10 2.17 (0.139)

LARLAR

Treatment 1 4 0.56 (0.495) 1 3 0.24 (0.661)

Year 2 6 9.37 (0.014)* 2 4 13.55 (0.016)*

Year×treatment 2 6 0.29 (0.760) 2 4 3.28 (0.144)

SALIX

Treatment 3 9 0.72 (0.567) 3 9 2.63 (0.114)

Year 2 12 3.04 (0.085) 2 8 0.76 (0.498)

Year×treatment 6 12 1.49 (0.263) 5 8 0.81 (0.576)

Other

Treatment 3 9 0.99 (0.440) 3 7 0.66 (0.601)

Year 1 4 0.11 (0.752) 1 3 0.97 (0.398)

Year×treatment 1 4 0.02 (0.887) 1 3 0.61 (0.491)
34 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Table 7. Mean shrub height (least-squares mean), variation (standard deviation) of height, number of patches, and percent cover for 

six shrub taxa and total shrub cover at Seney National Wildlife Refuge, Michigan, 2006, 2007, and 2010.

[Means are expressed as least-square means; percent cover is back transformed. C3, C3 study area; *, significant differences within year between control and 

burn treatments at α=0.05; MC, Marsh Creek study area; nd, not detected; nc, not calculated]

Year Treatment Mean height 

(standard error)

Standard deviation height 

(standard error)

Mean number of patches 

(standard error)

Mean percent cover 

(standard error)

Alder

2006 C3—Control 121.2 (29.5) 73.3 (33.7) 6.7 (3.1) 1.1 (1.1)

2006 C3—Burn 146.0 (41.7) 66.6 (33.7) 1.0 (3.1) 0.3 (0.3)

2007 C3—Control 154.3(29.5) 99.5 (33.7)* 5.7 (3.1) 0.9 (0.9)

2007 C3—Burn 213.0 (41.7) 0.0 (33.7)* 0.7 (3.1) 0.5 (0.6)

2010 C3—Control 121.2 (21.7) 73.3 (33.7) 6.7 (3.1) 1.1 (0.1)

2010 C3—Burn 146.0 (41.7) 66.6 (33.7) 1.0 (3.1) 0.3 (0.3)

2006 MC—Control 53.0 (23.2) 19.0 (19.5) 3.6 (2.4) 0.6 (0.5)

2006 MC—Burn nd nd 0.0 (3.9) 0.1 (0.2)

2007 MC—Control 85.0 (20.8) 42.8 (16.9) 2.6 (2.4) 0.7 (0.5)

2007 MC—Burn nd nd 0.0 (3.9) 0.1 (0.2)

2010 MC—Control 134.7(20.8) 37.8 (23.9) 2.8 (2.4) 0.7 (0.6)

2010 MC—Burn 65.0 (41.7) nd 0.5 (3.9) 0.1 (0.2)

Bog birch

2006 C3—Control 102.1 (18.0) 45.5 (6.1) 34.7 (10.9) 13.2 (9.5)

2006 C3—Burn 93.5 (18.0) 41.2 (6.1) 61.7 (10.9) 22.6 (14.4)

2007 C3—Control 117.0 (18.0) 39.9 (6.1) 28.7 (10.9) 15.6 (10.9)

2007 C3—Burn 117.5 (18.0) 42.1 (6.1) 48.0 (10.9) 28.1 (16.7)

2010 C3—Control 102.1 (18.0) 45.5 (6.1) 34.7 (10.9) 13.2 (9.5)

2010 C3—Burn 93.5 (18.0) 41.2 (6.1) 61.7 (10.9) 22.5 (14.4)

2006 MC—Control 67.0 (13.9) 22.3 (4.7) 31.8 (8.4) 6.4 (3.8)

2006 MC—Burn 75.5 (22.0) 15.7 (7.5) 34.0 (13.3) 5.2 (5.1)

2007 MC—Control 89.6 (14.2) 25.4 (4.9) 23.0 (8.4) 9.8 (5.7)

2007 MC—Burn 93.0 (22.0) 18.1 (7.5) 24.5 (13.3) 8.8 (8.2)

2010 MC—Control 100.0 (13.9) 28.7 (4.9) 28.8 (8.4) 8.7 (5.1)

2010 MC—Burn 70.6 (22.0) 23.4 (7.5) 26.5 (13.3) 6.3 (6.1)

Leatherleaf

2006 C3—Control 71.0 (6.1)* 5.3 (2.2) 1.0 (19.4) 0.2 (0.2)

2006 C3—Burn 40.4 (6.1)* 15.5 (2.2) 2.7 (19.4) 0.3 (0.3)

2007 C3—Control 75.5 (6.1) 2.1 (2.2) 0.7 (19.4) 0.4 (0.3)

2007 C3—Burn nd nd 0.0 (19.4) 0.1 (0.1)

2010 C3—Control 71.0 (6.1)* 5.3 (2.2) 1.0 (19.4) 0.2 (0.2)

2010 C3—Burn 40.4 (6.1)* 15.5 (2.2) 2.7 (19.4) 0.3 (0.3)

2006 MC—Control 37.3 (2.7) 8.9 (1.0) 129.0 (15.0) 24.3 (6.2)

2006 MC—Burn 39.0 (4.3) 12.4 (1.6) 191.5 (23.8) 56.6 (13.0)

2007 MC—Control 37.0 (2.7) 7.5 (1.0) 56.4 (15.0) 24.1 (6.2)

2007 MC—Burn 39.0 (4.3) 8.8 (1.6) 105.5 (23.8) 58.9 (12.8)

2010 MC—Control 42.7 (2.7) 12.6 (1.0) 63.4 (15.0) 32.7 (7.4)

2010 MC—Burn 40.0 (4.3) 10.7 (1.6) 81.5 (23.8) 63.2 (12.3)
Results 35

Table 7. Mean shrub height (least-squares mean), variation (standard deviation) of height, number of patches, and percent cover for 

six shrub taxa and total shrub cover at Seney National Wildlife Refuge, Michigan, 2006, 2007, and 2010.—Continued

[Means are expressed as least-square means; percent cover is back transformed. C3, C3 study area; *, significant differences within year between control and 

burn treatments at α=0.05; MC, Marsh Creek study area; nd, not detected; nc, not calculated]

Year Treatment Mean height 

(standard error)

Standard deviation height 

(standard error)

Mean number of patches 

(standard error)

Mean percent cover 

(standard error)

Tamarack

2006 C3—Control 111.8 (55.4) 70.6 (46.1) 3.3 (2.9) 1.3 (1.3)

2006 C3—Burn 157.2 (59.4) 84.0 (38.6) 10.7 (2.9) 1.8 (1.8)

2007 C3—Control 188.2 (55.4) 94.1 (46.1) 4.0 (2.9) 1.8 (1.8)

2007 C3—Burn 266.0 (55.4) 153.0 (37.7) 9.0 (2.9) 4.0 (4.0)

2010 C3—Control 111.8 (55.4) 70.6 (46.1) 3.3 (2.9) 1.3 (1.3)

2010 C3—Burn 157.2 (59.4) 84.0 (38.6) 10.7(2.9) 1.8 (1.8)

2006 MC—Control nd nd 0.0 (2.3) 0.1 (0.1)

2006 MC—Burn nd nd 0.0 (3.8) 0.1 (0.1)

2007 MC—Control nd nd 0.0 (2.3) 0.1 (0.1)

2007 MC—Burn nd nd 0.0 (3.8) 0.1 (0.1)

2010 MC—Control nd nd 0.0 (2.3) 0.1 (0.1)

2010 MC—Burn nd nd 0.0 (3.8) 0.1 (0.1)

Willows

2006 C3—Control 73.2 (22.7) 36.0 (12.5) 25.3 (2.8) 3.9 (1.8)

2006 C3—Burn 68.3 (22.7) 48.8 (12.5) 13.7 (2.8) 2.4 (1.2)

2007 C3—Control 81.7 (22.7) 37.5 (12.7) 5.7 (2.8) 2.1 (1.0)

2007 C3—Burn 96.6 (22.7) 49.2 (12.5) 6.7 (2.8) 2.2 (1.1)

2010 C3—Control 73.3 (22.7) 36.1 (12.5) 25.0 (2.8) 3.9 (1.8)

2010 C3—Burn 68.3 (22.7) 43.8 (12.5) 13.7 (2.8) 2.4 (1.1)

2006 MC—Control 42.0 (17.6) 9.3 (9.6) 6.6 (2.2) 0.5 (0.3)

2006 MC—Burn 35.0 (27.8) 14.0 (15.3) 10.0 (3.5) 0.8 (0.5)

2007 MC—Control 58.2 (21.8) 6.5 (11.8) 0.6 (2.2) 0.1 (0.1)

2007 MC—Burn 35.0 (27.8) nd 1.0 (3.5) 0.8 (0.1)

2010 MC—Control 54.7 (18.3) 9.7 (10.6) 1.0 (2.2) 0.2 (0.1)

2010 MC—Burn 38.7 (27.8) 0.0 (16.2) 1.5 (3.5) 0.1 (0.1)

Other taxa

2006 C3—Control nd nd 0.0 (11.0) 0.1 (0.1)

2006 C3—Burn nd nd 0.0 (11.0) 0.1 (0.1)

2007 C3—Control 69.7 (12.7) 18.4 (8.1) 3.0 (11.0) 1.0 (0.9)

2007 C3—Burn 43.2 (12.7) 17.6 (4.7) 10.7 (11.0) 2.1 (1.2)

2010 C3—Control nd nd 0.0 (11.0) 0.1 (0.1)

2010 C3—Burn nd nd 0.0 (11.0) 0.1 (0.1)

2006 MC—Control nd nd 0.0 (8.50) 0.0 (0.0)

2006 MC—Burn nd nd 0.0 (13.5) 0.1 (0.1)

2007 MC—Control 46.5 (9.9) 8.4 (4.0) 20.4 (8.5) 2.0 (1.3)

2007 MC—Burn 43.3 (15.6) 12.4 (5.7) 35.0 (13.5) 6.2 (5.7)

2010 MC—Control 47.9 (9.8) 17.2 (4.0) 33.0 (8.5) 3.4 (2.1)

2010 MC—Burn 43.8 (15.6) 13.3 (5.7) 36.5 (13.5) 7.1 (6.5)
36 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Table 7. Mean shrub height (least-squares mean), variation (standard deviation) of height, number of patches, and percent cover for 

six shrub taxa and total shrub cover at Seney National Wildlife Refuge, Michigan, 2006, 2007, and 2010.—Continued

[Means are expressed as least-square means; percent cover is back transformed. C3, C3 study area; *, significant differences within year between control and 

burn treatments at α=0.05; MC, Marsh Creek study area; nd, not detected; nc, not calculated]

Year Treatment Mean height 

(standard error)

Standard deviation height 

(standard error)

Mean number of patches 

(standard error)

Mean percent cover 

(standard error)

Total woody cover

2006 C3—Control nc nc nc 22.4 (10.1)

2006 C3—Burn nc nc nc 27.1 (11.4)

2007 C3—Control nc nc nc 24.1 (10.6)

2007 C3—Burn nc nc nc 37.0 (13.4)

2010 C3—Control nc nc nc 22.4 (10.1)

2010 C3—Burn nc nc nc 27.1 (11.4)

2006 MC—Control nc nc nc 34.1 (10.5)

2006 MC—Burn nc nc nc 64.6 (16.1)

2007 MC—Control nc nc nc 43.3 (10.9)

2007 MC—Burn nc nc nc 74.3 (13.5)

2010 MC—Control nc nc nc 47.7 (11.1)

2010 MC—Burn nc nc nc 74.2 (13.5)
Results 37

Table 8. Results of analysis of variance for percent cover and number of patches of woody taxa on study blocks in C3 and Marsh 

Creek study areas, Seney National Wildlife Refuge, Michigan, 2006, 2007, and 2010.

[p, probability; *, significant at α=0.05; --, no data; nc, not calculated]

Source of 

variation

Numerator 

degrees 

of freedom

Denominator 

degrees 

of freedom

Percent cover

F-statistic

(p-value)

Numerator 

degrees 

of freedom

Denominator 

degrees 

of freedom

Number of patches

F-statistic

(p-value)

ALDNIC

Treatment 3 10.25 0.66 (0.593) 3 9 0.77 (0.538)

Year 2 18.63 0.04 (0.964) 2 18 0.28 (0.761)

Year×treatment 6 18.63 0.73 (0.632) 6 18 0.08 (0.997)

BETPUM

Treatment 3 8.46 0.66 (0.600) 3 9 1.77 (0.222)

Year 2 17.46 16.18 (<0.001)* 2 18 13.56 (<0.001)*

Year×treatment 6 17.46 1.26 (0.324) 6 18 1.00 (0.456)

CHACAL

Treatment 3 9 29.89 (<0.001)* 3 9 11.74 (0.002)*

Year 2 13.35 0.10 (0.909) 2 18 18.84 (<0.001)*

Year×treatment 6 20.34 0.47 (0.823) 6 18 6.37 (0.001)*

LARLAR

Treatment 3 11.71 10.08 (0.001)* 3 9 2.95 (0.091)

Year 2 27 0.03 (0.966) 2 18 0.12 (0.885)

Year×treatment 6 20.78 1.00 (0.449) 6 18 0.50 (0.797)

SALIX

Treatment 3 13.31 16.76 (<0.001)* 3 9 14.83 (0.001)*

Year 2 27 1.70 (0.202) 2 18 18.21 (<0.001)*

Year×treatment 6 20.6 2.78 (0.038)* 6 18 3.95 (0.011)*

Other

Treatment 3 13.03 0.91 (0.464) 3 9 1.51 (0.278)

Year 2 17.94 26.18 (<0.001)* 2 18 5.64 (0.013)*

Year×treatment 6 17.94 3.86 (0.012)* 6 18 1.68 (0.183)

All woody cover

Treatment 3 8.98 2.02 (0.182) -- -- nc

Year 4 17.97 10.61 (<0.001)* -- -- nc

Year×treatment 6 17.98 3.00 (0.033)* -- -- nc
38 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Table 9. Results of analysis of variance for visual obstruction readings for six height strata on study blocks in C3 and Marsh Creek 

study areas, Seney National Wildlife Refuge, Michigan, 2006–10.

[VOR, visual obstruction reading; p, probability; *, significant at α=0.05]

Source of variation Numerator degrees 

of freedom

Denominator degrees 

of freedom

Mean VOR Standard deviation of VOR

F-statistic p-value F-statistic p-value

Treatment 3 9 2.18 0.16 3.64 0.06

Strata 5 45 244.15 <0.01* 20.91 <0.01*

Treatment×strata 15 45 2.52 0.01* 3.93 <0.01*

Year 4 216 51.45 <0.01* 27.66 <0.01*

Year×treatment 12 216 8.81 <0.01* 3.49 <0.01*

Year×strata 20 216 6.30 <0.01* 3.56 <0.01*

Year×treatment×strata 60 216 2.80 <0.01* 2.18 <0.01*

C3 Control

C3 Spring burn

MC Control

MC Summer burn

MC–2 Summer burn

EXPLANATION

Year

2006 2007 2008 2009 2010

A

Strontium 1 visual obstruction reading

1.0

0.2

0

0.8

0.4

0.6

Year

2006 2007 2008 2009 2010

B

Strontium 2 visual obstruction reading

1.0

0.2

0

0.8

0.4

0.6

Year

2006 2007 2008 2009 2010

C

Strontium 3 visual obstruction reading

0.5

0.1

0

0.4

0.2

0.3

Figure 23. Visual obstruction readings (VORs, least-squares 

means) in control and burn blocks for 33-centimeter strata 

in C3, Seney National Wildlife Refuge, Michigan, 2006–10. 

A, Stratum 1; B, stratum 2; C, stratum 3. C3 was burned in May 

2008 and VOR data were collected 2 months later; Marsh Creek 

was burned in July 2008 and VOR data were collected 3 weeks 

later. Whiskers indicate standard errors.
Discussion 39

C3 Control

C3 Spring burn

MC Control

MC Summer burn

MC–2 Summer burn

EXPLANATION

Year

2006 2007 2008 2009 2010

A

Strontium 4 visual obstruction reading

0.5

0.1

0

0.4

0.2

0.4

Year

2006 2007 2008 2009 2010

B

Strontium 5 visual obstruction reading

Year

2006 2007 2008 2009 2010

C

Strontium 6 visual obstruction reading

0.20

0.05

0

0.15

0.10

0.20

0.05

0

0.15

0.10

Figure 24. Visual obstruction readings (VORs, least-squares 

means) in control and burn blocks for 33-centimeter strata 

in C3, Seney National Wildlife Refuge, Michigan, 2006–10.

A, Stratum 4; B, stratum 5; C, stratum 6. C3 was burned in May 

2008 and VOR data were collected 2 months later; Marsh Creek 

was burned in July 2008 and VOR data were collected 3 weeks 

later. Whiskers indicate standard errors.

Discussion

Burning is considered an important tool in wet grasslands 

for maintaining biodiversity, but results of our study indicated 

that summer burns had a greater effect on vegetation than 

more commonly applied spring burns. We detected few effects 

of spring burning in C3, even in the same year of burning. 

Most vegetation was in early stages of growth at the time of 

burning and had 11–14 weeks to recover before data collection 

that year. In Marsh Creek, not surprisingly, most treatment 

effects were in 2008, when data were collected within 3 weeks 

of burning and vegetation was just beginning to regrow. Some 

fire effects there, however, persisted one to two growing 

seasons (2009, 2010). These results concur with other fen 

studies that reported recovery to preburn conditions within one 

to two growing seasons after light or moderate fires (Reuter, 

1986; Burkman, 1993; Kost and De Steven, 2000; Middleton, 

2002). For MC–2, which was burned in summer 2007 under 

higher fire conditions, some measures did not recover to 

preburn levels for two to three growing seasons.

The goal of this study was to understand the effects 

of spring and summer burns on the plant community and 

structure of fens, and in particular how they change woody 

structure. Because we wanted treatments to reflect normal 

prescribed-fire operations, only one burn treatment was 

applied to an area (that is, Refuge burn unit); hence, differences between spring and summer burns are partially 

confounded by natural differences in the plant community 

and hydrology between the two areas, as noted above. Indeed, 

most of the significant year × treatment effects were related to 

differences by area and infrequently by differences between 

burn and control treatments within an area in the years after 

burning; moreover, the heterogeneity of these areas is apparent 

in many variations within and among blocks.

The fire treatments applied in this study in 2008 were 

under moderate fire conditions for each season and resulted 
40 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

in light to moderate burns. Such fire conditions often result 

in patchy fuel consumption and are not severe enough to 

burn into the organic substrate to affect the seed bank or root 

systems. Temperatures of about 60 °C are considered lethal 

to most living tissue, but soil temperature is quickly attenuated by moisture, so only the top few centimeters of soil may 

reach lethal temperatures during light to moderate fires, with 

minimal effect to species that have deeper root crowns or 

rhizomes (Flinn and Wein, 1977). The general absence of fire 

effects in C3 reflects the cooler and wetter substrate, more 

moderate spring fire conditions, and low fire severity results; 

however, this fire was deemed successful in terms of broader 

fire management goals, which was to open up the brush and 

promote more sedge to set the stage for a hotter summer burn 

14–15 months later (see further discussion of this in the “Fire 

Management and History in Fens” section). The fire treatments 

applied in this study differed most markedly between 2007 

and 2008. MC–2 was burned in July 2007 under much drier 

soil conditions and high fire danger rating (table 1), yielding 

moderate fire severity effect, extensive top kill of shrubs, and 

longer-lasting effects than in the other Marsh Creek blocks 

burned 1 year later under more moderate fire conditions, 

yielding light to variable fire severity effects. The prolonged 

drought and resulting very high to extreme fire danger conditions in 2007 led to the Sleeper Lake Fen wildfire, 45 km from 

the Refuge, just 3 weeks after the MC–2 burn; it burned more 

than 7,200 ha over 3 months (Bess and others, 2014).

A predominant effect of fire is its consumption of standing vegetation and litter, which exposes the substrate, thereby 

altering light and temperature conditions, removing a filter for 

nutrients, and recycling nutrients. The resulting higher albedo 

and soil temperatures and increased light conditions enhance 

seed germination and growth and can alter the competitive 

abilities of species in the seed bank or remaining root systems 

(Auclair and others, 1973; Smith, 1973; Rooney, 1990; 

Johnson and Knapp, 1995a, b; Warners, 1997). Change in litter 

depth was one of the most marked fire effects in this study. 

Although the 2008 fires did not entirely consume litter, they 

resulted in reduced and often patchy levels that year, with a 

lesser decline in C3. Litter depth took two growing seasons to 

recover after the 2008 summer burn and three growing seasons 

to recover after the more severe 2007 summer burn. Although 

litter loss from fire may have played a role in the increasing 

levels of percent open during the study, the absence of fire 

effects within year and area, except in the summer-burned 

blocks in 2008, indicates rapid regeneration of vegetation. 

Surface water, contributing to litter decomposition, also 

may have played a role in the trends of percent open. Litter 

compaction and decomposition were likely less during the 

dry years of 2005–7 than during the wetter conditions in later 

years (Neckles and Neill, 1994).

Linked to litter is graminoid height, indicative of aboveground production and future litter deposition. Burning can 

stimulate aboveground biomass production and reproduction 

(flowering or seed production) in the first season after burning 

(Young, 1987; Bernard and others, 1988; Warners, 1997; Kost 

and De Steven, 2000; Main and Barry, 2002). However, we 

detected no evidence of stimulated graminoid growth after 

spring burning in C3, and graminoid height took two to three 

growing seasons to recover after summer burning in Marsh 

Creek and MC–2. Results instead indicate that other factors 

such as water conditions and competition from other vegetation contributed to changes in graminoid height. Burkman 

(1993) also noted annual differences in herbaceous vegetation on both burn and control plots in Marsh Creek related to 

wetter conditions.

Changes in Plant Community

Overall, effects of treatment or year × treatment interactions largely represented spatial differences of species within 

and among study blocks, and few of the 27 species examined 

showed clear effects of burning within areas (excluding the 

expected changes in summer-burned blocks in 2008 because 

of the timing of sampling). The minimal effect on species richness and diversity in the first growing seasons after burning 

is not surprising given nearly all taxa characteristic of fens 

are long-lived perennials such as sedges and leatherleaf that 

are often clonal and rely on rhizomatous root systems and, 

therefore, can quickly recover from light to moderate burning. 

Other studies have similarly reported little change in plant 

community after burning fens or wet meadows (Cornely and 

others, 1983; Smith and Kadlec, 1985; Boyd and others, 1993; 

Austin and others, 2007), but some have reported increased 

cover of annual forbs in the first year after burning (Kost 

and De Steven, 2000; Bartels and Wilson, 2001; Middleton, 

2002). In the fens examined here, however, all but one forb 

in the study areas were perennials. Similarly, Kowalski and 

Wilcox (2003) reported only 1 of 71 plant taxa was an annual 

in their study of vegetation along the Marsh Creek above and 

below the C3 levee and water-control structure. The ordination 

results also demonstrated that the plant communities tended 

to return to preburn conditions within two growing seasons of 

burning.

The fens in this study are composed mostly of obligate or 

facultative-wet perennial species, which vary in their hydrological niche. Similar trends between control and burn blocks 

across years for several measures, such as species richness, 

graminoid height, and NMS scores, indicate seasonal water 

conditions were an important factor influencing the changes 

we detected across years and, in some situations, interacted 

with fire effects. The study began during the second year of a 

moderate drought and concluded under moderately wet conditions. There was little standing water in the study blocks by 

July each year except in 2010, when summer rains led to more 

surface water, particularly on the western end of Marsh Creek 

(MC–1 and MC–2), which is farther from the levee and other 

upland areas. Spring flooding conditions were more variable, 

with greatest depths in 2008 and 2009.
Discussion 41

Sedges comprised more than 40 percent of herbaceous 

cover, with the narrow-leaved sedges (mainly fewseed and 

woollyfruit sedge) dominating on most sites. Year × treatment interactions indicate interactions between local water 

conditions and fire season—no effects in the first growing 

season after the spring burn in C3 (2008) but higher cover 

1 year later, but sedge cover was lower in both the first and 

second growing seasons after the Marsh Creek summer burn 

(2009–10). Some year-to-year differences in narrow-leaved 

sedge cover and frequency were likely due to different field 

personnel, but the dry conditions during the pretreatment 

years resulted in little seed production, which was particularly 

important for differentiating the narrow-leaved sedges. Flowering and seed production improved in the later, wetter years, 

but less than 15 percent of stems of woollyfruit and fewseed 

sedges produced seeds in 2008–10 (J.E. Austin, unpub. data, 

2008–10).

Water table and groundwater connectivity are dominant 

factors driving plant communities of fens and other wetlands 

(Warners, 1997; Weltzin and others, 2003; Henszey and others, 

2004; Little and others, 2010). Seasonal water levels also 

may contribute to a lag in community responses (Wilcox and 

Nichols, 2008); for example, bluejoint tends to produce more 

biomass under drier conditions (Fraser and Karnezis, 2005) 

and may decline or disappear under persistently flooded conditions (Gates, 1942). For some species, germination, growth, 

and reproduction after wetter conditions may have contributed 

to the detection in later years (for example, Buxbaum’s sedge 

in Marsh Creek in 2010). Alternatively, declines in species 

such as Robbins’ spikerush, bedstraws, and northern bog aster 

may have been related to the shift from drier to wetter conditions during the study.

Some graminoids such as creeping sedge, elliptic spikerush, and spike muhly were common but patchily distributed, 

which contributed to high variability within treatments and 

limited our ability to discern any fire effects. Anderson (1982) 

reported cottongrasses colonized burned fens the spring after 

the fall 1976 Seney fire, but we detected no difference between 

burned and control blocks one to two growing season after 

burning; its earlier abundance in Marsh Creek burn blocks 

never recovered after 2008. Robbins’ spikerush disappeared 

from Marsh Creek burn blocks after summer burning, but it 

had been rare in control blocks there. The distinctive white 

beak-rush was detected only in the westernmost blocks of 

C3 and was not detected in 2010. Wet conditions may have 

contributed to the apparent disappearance of this sedge after 

summer burning. 

Fire may stimulate the reappearance of less common 

species. Marsh bellflower and earth loosestrife had markedly 

higher abundance one growing season after the summer 2007 

fire in MC–2; their abundance in other blocks seemed more 

driven by annual or site conditions than fire effects. Similarly, Middleton (2002) reported marsh bellflower appearing 

after a long absence after a winter burn in a Wisconsin sedge 

meadow. A similar brief stimulus was detected with marked 

spike in purplestem aster cover in Marsh Creek in 2009, one 

growing season after the summer burn. In contrast, northern 

bog aster largely disappeared from MC–2 after 2006 and in 

all blocks after 2007, indicating water conditions were a key 

driver. Mosses (predominantly Sphagnum) also seemed to 

respond to spring burning in the first growing season but likely 

also were responding to wetter conditions in 2008 and 2010.

Marsh thistle and broadleaf cattail were the only two 

species of invasive concern detected, but both were detected at 

low levels and were sparsely distributed. Kowalski and Wilcox 

(2003) detected marsh thistle along the Marsh Creek drainage 

above the C3 pool but not below the pool; we detected it thinly 

scattered throughout the C3 study area. Fire may contribute 

to its occurrence or spread, because it requires open substrate 

for germination. Although cattail can develop into monotypic 

stands, in these fens their distribution seems largely adjacent 

to sources of groundwater seepage (for example, near some 

areas of greater seepage below the levee along the south side 

of C3 pool, closer to Sweeney Creek), as noted also by Kowalski and Wilcox (2003); it was not detected in Marsh Creek. 

The systems’ pH levels likely are an important factor limiting 

the species’ distribution. 

Earlier studies at the Refuge reported woollyfruit sedge 

as the dominant sedge in the Marsh Creek area (Stenzel, 1982; 

Bart and others, 1984), whereas our study found fewseed 

sedge to be most common there and woollyfruit sedge to 

be dominant in C3. Kowalski and Wilcox (2003) also noted 

woollyfruit sedge as abundant below C3, in an area west of 

our study blocks and along the Marsh Creek drainage. We 

suspect this discrepancy of narrow-leaved taxa occurrence 

among studies over 28 years relates to spatial and hydrological 

differences. Stenzel’s 1980 study was completed west of the 

Marsh Creek pool levee (Stenzel, 1982), and therefore, under 

different hydrological effects, and just 4 years after the 1976 

Seney fire (Anderson, 1980). The study area of Bart and others 

(1984) included western parts of our study area; methods do 

not indicate how they distinguished Carex species, so they 

may have assumed the same species as reported in Stenzel’s 

study. Of the six plots in Burkman’s (1993) study, four were 

overlapped with our Marsh Creek area, and two were west 

of the Marsh Creek pool, similar to that of Stenzel’s’ (1982) 

study area. Burkman (1993) reported mean pH as 6.08 (burned 

plots) and 6.32 (control plots). Fewseed sedge has a much 

narrower range of pH tolerance and is more common in more 

acidic areas (pH ~4) than woollyfruit and interior sedge 

(pH>6; Wheeler and others, 1983; Gignac, 1994; Anderson 

and others, 1996), which indicates the dominant narrowleaved sedge west of Marsh Creek pool was woollyfruit sedge, 

as reported by the earlier studies. We did not measure pH in 

our study areas, but our results indicate more alkaline conditions immediately below the Marsh Creek pool. Both woollyfruit and fewseed sedges, however, share the narrow-leaved 

characteristics that provide preferred habitat characteristics of 

yellow rails.
42 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Woody Cover and Structure

We detected no significant effects of either spring or 

summer burn within years for any of the shrub metrics; rather, 

treatment effects were largely due to differences between the 

two areas or among blocks in an area even before burning. 

Some differences across years may be related to different 

observers among years (for example, slight shifts in how shrub 

transects were laid out each year). An earlier fire-effects study 

in the western part of Marsh Creek in 1991–92 reported total 

shrub cover was substantially reduced for 2 years after a single 

dormant-season (spring or fall) burn, primarily because of the 

reduction of speckled alder (Burkman, 1993). A closer examination of percent cover data (appendices A and B of Burkman, 

1993) showed partial recovery of woody cover 2 years after 

burning in two of the burned plots but no change in woody 

cover in one of the spring-burned plots. Most shrub taxa in 

fens have strong resprouting abilities, and fire may only briefly 

reduce woody dominance (Bowles and others, 1996; Pendergrass and others, 1998; Middleton, 2002; Brisson and others, 

2006). Most studies, however, involved surface fires of low 

intensity and a single fire during the dormant season (often 

early spring), resulting in high survival of shrubs, resprouting, increased stem densities, and only temporary reduction 

in cover or height, as observed in this study. Higher fire 

frequency or more intense fire conditions seem necessary for 

more lasting control of woody species through increased stress 

and direct mortality.

Although cover and height metrics showed little fire 

effects, data for VORs demonstrated some effects of fire and 

not just at the lowest strata, which was mostly herbaceous 

vegetation. VORs of intermediate strata (34–99 cm) in Marsh 

Creek trended downward in 2008 and 2009, indicating less 

dense growth and likely lower stem densities at these heights 

one growing season after burning. VORs, however, had 

recovered to preburn conditions two growing seasons after 

summer burning. The lower VOR measures indicate shading 

was temporarily reduced around woody patches, which could 

alter competition and growth among species. Burkman (1993) 

reported lower mean vertical cover 1 year after dormantseason burning than in the paired control plots, but she did 

not differentiate results between the early spring and early 

fall burns. Keddy (1989) noted that only one-quarter of plant 

species responded to removal of the shrub sweetgale (Myrica 

gale); smaller-statured species with high seedbank densities 

increased after removal, but tall species like bluejoint and 

common spikerush (Eleocharis palustris), which are more 

shade tolerant, declined. Overall, Keddy (1989) noted that 

changes in the plant community took several years to become 

apparent.

Fire Management and History in Fens

Most prescribed fires in northern wetlands are implemented under light to moderate conditions, often tied to 

concerns of safety and controllability; many are therefore done 

around the dormant season (spring or fall), when temperatures are cooler and fuel moisture is higher. Dormant-season 

burning also minimizes effects on nesting birds (Beule, 1979; 

Kantrud, 1986). Such fires are often of low fire severity; leave 

a mosaic of burned and unburned patches; and have shortterm effects (<3 years), mainly reducing litter and temporarily 

increasing species diversity with germination of light-sensitive 

and nonnative invasive species (Thompson and Shay, 1985, 

1989; Mallik and Wein, 1986; Shay and others, 1987; Ailstock 

and others, 2001; Brisson and others, 2006). Single light to 

moderate burns are ineffective in setting back woody taxa 

such as alder and willow for more than 1–2 years (Kost and 

De Steven, 2000; Middleton, 2002; Brisson and others, 2006; 

this study). Burns at the Refuge were specifically planned 

and implemented to avoid starting peat fires; thus, killing 

the below-ground part of vegetation (plant tissue) was never 

considered achievable. A primary objective of both spring and 

summer burns on the Refuge was to top-kill the brush in the 

wetlands. We noted that flame lengths as short as 0.3 m would 

top kill the wetland brush; this was observed when the fire 

was carried under the brush canopy or girdled the brush stem. 

Experienced fire managers in the region assume that, under 

a frequent enough fire-return interval, brush density will be 

reduced because of repeated stress of the fires (G. Lindsay, 

personal commun., 2018); however, the duration of the project 

was not long enough to validate this assumption.

Although spring burns on their own may have shortterm effects (<3 years), they are valuable for broader fire 

management goals. Under moderate fire conditions, fire often 

does not carry into heavy brush because of a lack of fine 

fuels (primarily grasses and sedges) and moister microclimate. This problem was encountered during a summer burn 

attempted in C3 in 2004; hence, the goal of the C3 spring fire 

in 2008 was to top kill the brush and promote more sedge 

(and therefore, fine fuels) to set the stage for a hot summer 

burn 14–15 months later, which would be more likely to kill 

shrubs. The C3 spring fire achieved those objectives for fire 

management and set the stage for a followup summer burn. 

As sometimes happens in fire management, unfortunately, that 

followup burn was not attempted because of local weather 

conditions during the summer fire window and national wildfire activity.

Relatively few wetland fire studies have compared results 

from different burning seasons (Diiro, 1982; Reuter, 1986; 

Thompson and Shay, 1985, 1989; Shay and others, 1987; 

Mallik and Wein, 1986; Main and Barry, 2002). Summer 

burns are more effective than spring or fall burns at reducing biomass and the subsequent year’s growth of dominant 

perennials such as phragmites (Phragmites australis) and 

broadleaf cattail. With the reduced dominance of such robust 

perennials, species diversity and richness were higher after 
Rethinking Fire Management for Controlling Woody Encroachment in Fens 43

summer burns (Mallik and Wein, 1986; Thompson and Shay, 

1989). Fewer studies have considered the timing of burning 

relative to target species’ growth or flowering phenology 

(Beule, 1979; van der Toorn and Mook, 1982; Cross, 1983; 

Krusi and Wein, 1988), despite indications of its importance 

in early wetland literature (Ward, 1942) and results from 

upland studies (Knapp and others, 2009). Burning at the end 

of the growing season after plants have translocated nutrient 

reserves below ground, during the winter, or in early spring 

before shoot emergence will have little effect on below-ground 

reserves and, hence, lessened effect on the subsequent year’s 

growth. Prescribed burns applied in these seasons are largely 

to remove aboveground biomass to expose more substrate and 

promote greater germination and growth the following spring. 

More lasting effects to perennial taxa can be achieved when 

burns are done when nutrient reserves are aboveground (flowering, active growth) and it is too late in the season for plants 

to resprout and recoup reserves for the subsequent growing 

season. As noted in the review by Knapp and others (2009), 

however, seasonal differences in fuel consumption and fire 

intensity may be a more important factor in fire effects than 

plant phenology at the time of the fire. In our study, our goal 

for summer burns was to top kill shrubs before they began to 

translocate reserves belowground. However, although some 

effects of summer burning lasted two growing seasons (three 

seasons for some measures in MC–2), we saw little effect on 

woody cover or height, and only some reduction in VORs, 

indicating this strategy is not effective or requires greater 

fire intensity to set back woody cover than we were able to 

achieve.

Most wildfires in temperate wetland systems occur during 

drier conditions of summer and early fall and can be most 

intense during drought—conditions that are avoided by fire 

managers but are more likely to set back woody encroachment. The prolonged drought in 2007 prevented the Refuge 

fire staff from fully implementing the Refuge burn plan and 

our BACI design as planned; however, results for the area we 

did burn that summer (MC–2) indicated longer-lasting effects 

than the areas burned under the more moderate fire conditions 

and wetter landscape the following year. In the areas of the 

Refuge burned more intensely during the summer and fall of 

1976, bog birch and willow did not recolonize the area until 

nearly 3 years after the fire (Anderson, 1982). Photographs of 

the Marsh Creek study area taken in June 1982, 6 years after 

the fire, show a low density of low-stature shrubs (Stenzel, 

1982). More recently, the land cover type derived from 2004 

aerial imagery delineated this study area as lowland shrub, 

25–60 percent cover, similar to the types encompassed in most 

of our study areas. 

Before European settlement, fire was probably an important natural disturbance factor in the fens of Wisconsin (Curtis, 

1959) and elsewhere in the north-central region (Prince, 

1997). Droughts and logging during the late 1800s to the early 

1900s contributed to the severe wildfires that burned both 

uplands and wetlands. Before settlement (1840–56), General 

Land Office surveys indicated 23 percent of the U.S. Forest 

Service’s Seney Subdistrict of Michigan’s upper Peninsula 

was large patches of wetlands, including emergent and shrubdominated wetlands (Comer and others, 1995). By the 1950s, 

fire suppression on uplands, combined with wetland drainage 

efforts, led to reduced wetland size and woody encroachment 

into wetlands (Curtis, 1959; Jahn and Hunt, 1964). Jahn and 

Hunt (1964:54) reported that “[u]nder present-day, strict fire 

protection, new depressions are rarely burned in meadows, 

and sedges and grasses give way to shrubs and trees.” By 

1990, wetlands and water bodies in the Seney Subdistrict had 

declined to 10 percent (Zhang and others, 2000). 

Prescribed fire programs on public lands in this region 

began in the 1990s, primarily to restore fire to the region’s 

fire-dependent forest ecosystems; the fire program began at the 

Refuge in 2000. Prescribed burning is recognized as an important management tool used to restore and manage fens in this 

Refuge for yellow rails, sedge wrens, Le Conte’s sparrows, 

and other birds (U.S. Fish and Wildlife Service, 2009), as well 

as restoring fens on other conservation lands (Middleton and 

others, 2006). Given the short-term effects of light to moderate spring or summer burns, such fires will have little effect 

on breeding birds other than temporary loss of ground-nesting 

conditions within the first growing season after burning. Fire 

can, however, be useful for creating more diverse habitat 

conditions spatially and temporally (Anderson, 1982).

Rethinking Fire Management for 

Controlling Woody Encroachment in 

Fens

The open and partially wooded fens at Seney National 

Wildlife Refuge are dominated by perennial wetland plants 

that are largely resistant to surface fires. Light to moderate 

spring burns had minor effects on plant community, vegetative 

cover, and woody vegetation in even the first growing season 

after burning, whereas summer burns had somewhat longerlasting effects, mainly reducing fine fuels (litter, lower VORs). 

The summer 2007 burn completed under drought conditions 

was closer to achieving habitat objectives, but all measures 

recovered within two to three growing seasons. Some changes 

in plant cover indicate seasonal water conditions were important and interacted with burning. Our results demonstrated the 

heterogeneity of plant community and environmental conditions within and among areas. Differences that were detected 

largely represented spatial differences in the plant community 

within and among the two study areas.

Maintaining or restoring more open conditions in fens 

may require different approaches to water management, more 

aggressive fire management (for example, shorter fire-return 

interval, more intense fires, or burning during drier conditions), or the combination of fire with other tools such as 

cutting. Although of little apparent effect on their own, spring 

burns can be useful where woody cover limits the effective 
44 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

use of summer burns by setting up conditions for later summer 

burns that can be more effective in setting back woody cover. 

Important to understanding fire effects is the documentation 

of preburn environmental conditions, such as water, substrate, 

and fuel conditions, and phenology of primary taxa of interest; weather and fuel conditions during the fire; and fire 

severity immediately after the burn. Few fire studies report 

these measures, which limits ability to compare or interpret 

effects among fires or studies. Longer-term (>3 years) or 

more substantial changes to the plant community, particularly 

reducing the dominance of woody plants, will likely require 

targeting the timing and intensity of fires to reduce the vigor 

or kill perennial plants. Longer-term studies or monitoring are 

needed to detect interactions between fire disturbances and 

water or other environmental conditions and to detect longterm shifts in the plant community and structure.

References Cited

Ailstock, M.S., Norman, C.M., and Bushmann, P.J., 2001, 

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Glossary 49

Glossary

ALDNIC speckled alder, Alnus incana spp. 

rugosa

ANDGLA bog rosemary, Andromeda polifolia var. glaucophylla

BETPUM bog birch, Betula pumila

CALCANAG bluejoint, Calamagrostis 

canadensis, and creeping bentgrass, Agrostis 

stolonifera

CAMAPA marsh bellflower, Campanula 

aparinoides

CARAQU water sedge, Carex aquatilis, and 

unknown wide-leaved Carex species

CARBUX Buxbaum’s sedge, Carex buxbaumii

CARCHO creeping sedge, Carex chordorrhiza

CAREX narrow-leaved sedges combined 

(woollyfruit sedge, Carex lasiocarpa; fewseed 

sedge, C. oligosperma; inland sedge, C. interior, and unidentified narrow-leaved Carex

species)

CARLIV livid sedge, Carex livida

CHACAL leatherleaf, Chamaedaphne calyculata

ELEELL elliptic spikerush, Eleocharis elliptica

ELEROB Robbins’ spikerush, Eleocharis 

robbinsii

ERIOPH cottongrasses (Eriophorum angustifolium, E. viridicarinatum)

GALIUM bedstraws (northern bog bedstraw, 

Galium labradoricum; Threepetal bedstraw, 

G. trifidum) 

HYPKAL Kalm’s St. Johnswort, Hypericum 

kalmianum

IRIVER harlequin blueflag, Iris versicolor

LARLAR tamarack, Larix laricina

LYSTER earth loosestrife, Lysimachia terrestris

MC Marsh Creek

MUHGLO spike muhly, Muhlenbergia glomerata

POTFRU Shrubby cinquefoil, Potentialla 

fruticosa

RHYALB white beak-rush, Rhynchospora 

alba

RUBUS berries (arctic blackberry, Rubus 

arcticus ssp. acaulis; bristly dewberry, R. hispidus; dwarf red blackberry, R. pubescens) 

SALIX willows (sageleaf willow, Salix candida, and unidentified Salix species)

SD standard deviation

SOLULI bog goldenrod, Solidago uliginosa

spp. species

ssp. subspecies

SYMBOR Northern bog aster (Symphyotrichum boreale)

SYMPUN Purplestem aster (Symphyotrichum puniceum)

TRIFRA Fraser’s marsh St. Johnswort (Triadenum fraseri)

VACCINS Vaccinium species (cranberry, 

Vaccinium macrocarpon; velvetleaf huckleberry, V. myrtilloides; and unidentified Vaccinium species)

VIOLAS violets (small white violet, Viola 

macloskeyi; northern bog violet, V. nephrophylla)
50 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Appendix 1. Crosswalk Table of Taxonomy of Plant Species.

Table 1.1. Crosswalk table of taxonomy of plant species following the Flora of North America (2003) and U.S. Department of Agriculture 

National PLANTS Database (U.S. Department of Agriculture, Natural Resources Conservation Service, 2003).

[List includes species that may have been found only in blocks C3–7 or C3–8, which were sampled in 2006 and 2007 but not included in analyses. Codes 

are those used in the field and analyses. Wetness, an indicator of a species’ tolerance to wet soils, was from Chadde (2002) to be specific to the study region. 

Some codes differ from the PLANTS scientific name because the names in the regional guide used (Chadde, 2002) differ from the most recent taxonomy in the 

PLANTS Database. Wetness, indicator of plant tolerance to wet soils; --, no data; OBL, obligate wetland hydrophyte; FACW, facultative wetland hydrophyte; 

spp., species; FAC, facultative hydrophyte;(NI), no indicator; **, introduced; UPL, upland nonhydrophyte; ssp., subspecies; var., variety]

Code National PLANTS Database taxonomy Flora of North American taxonomy Wetness

Genus and species Common name Genus and species Common name

Sedges/rushes/cattail

CAREX1 Sedge, unknown 2006 #1 (Probably C. oligosperma) -- -- --

CAREX2 Sedge, unknown 2006 #2 (Probably C. interior) -- -- --

CAREX3 Sedge, unknown 2007 -- -- -- --

CAREX4 Sedge, unknown 2006 #4 (Probably C. lasiocarpa) -- -- --

CAREX9 Sedge, unknown 2007 -- -- -- --

CARAQU Carex aquatilis Water sedge Carex aquatilis Water sedge OBL

CARBEB Carex bebbii Bebb’s oval sedge Carex bebbii Bebb’s oval sedge FACW+

CARBUX Carex buxbaumii Buxbaum’s sedge Carex buxbaumii Buxbaum’s sedge OBL

CARCHO Carex chordorrhiza Creeping sedge Carex chordorrhiza Cordroot sedge OBL

CAREXI Carex exilis Coastal sedge Carex exilis Coast sedge OBL

CARHYA Carex hyalinolepis Shoreline sedge Carex hyalinolepsis Shoreline sedge OBL

CARINT Carex interior Inland sedge Carex interior Inland sedge OBL

CARLAS Carex lasiocarpa Woollyfruit sedge Carex lasiocarpa Slender sedge (wiregrass) OBL

CARLEP Carex leptalea Bristlystalked sedge Carex leptalea Slender sedge OBL

CARLIV Carex livida Livid sedge Carex livida Livid sedge OBL

CAROLI Carex oligosperma Fewseed sedge Carex oligosperma Running bog sedge OBL

CARROS Carex rostrata Beaked sedge Carex rostrata Beaked sedge OBL

CARUTR Carex utriculata Northwest Territory sedge Carex utriculata Beaked sedge OBL

DULARU Dulichium arundinaceum Three-way sedge Dulichium arundinaceum Three-way sedge OBL

ELEELL Eleocharis elliptica Elliptic spikerush Eleocharis elliptica Elliptic spikerush FACW

ELEROB Eleocharis robbinsii Robbin’s spikerush Eleocharis robbinsii Robbin’s spike-rush OBL

ERIANG Eriophorum angustifolium Tall cottongrass Eriophorum angustifolium Tall cottongrass OBL

ERIVIR Eriophorum viridicarinatumThinleaf cottonsedge Eriophorum viridicarinatumDark-scale cotton-grass OBL

JUNCAN Juncus canadensis Canadian rush Juncus canadensis Canada rush OBL

RHYALB Rhynchospora alba White beak-rush Rhynchospora alba White beak-rush OBL

SCICYP Scirpus cyperinus Woolgrass Scirpus cyperinus Wool-grass, bulrush OBL

TYPLAT Typha latifolia Common/broadleaf cattail Typha latifolia Common/broad-leaved 

cattail

OBL

Grasses

GRASS Grass, unknown spp. -- -- -- --

AGRHYE Agrostis hyemalis Winter bentgrass Agrostis hyemalis Ticklegrass FAC−

AGRSTO Agrostis stolonifera Redtop Agrostis gigantea Redtop (NI)

BROCIL Bromus ciliatus Fringed brome Bromus ciliatus Fringed brome FACW

CALCAN Calamagrostis canadensis Bluejoint, reedgrass Calamagrostis canadensis Blue-joint, reedgrass OBL
Appendixes 51

Table 1.1. Crosswalk table of taxonomy of plant species following the Flora of North America (2003) and U.S. Department of Agriculture 

National PLANTS Database (U.S. Department of Agriculture, Natural Resources Conservation Service, 2003).—Continued

[List includes species that may have been found only in blocks C3–7 or C3–8, which were sampled in 2006 and 2007 but not included in analyses. Codes 

are those used in the field and analyses. Wetness, an indicator of a species’ tolerance to wet soils, was from Chadde (2002) to be specific to the study region. 

Some codes differ from the PLANTS scientific name because the names in the regional guide used (Chadde, 2002) differ from the most recent taxonomy in the 

PLANTS Database. Wetness, indicator of plant tolerance to wet soils; --, no data; OBL, obligate wetland hydrophyte; FACW, facultative wetland hydrophyte; 

spp., species; FAC, facultative hydrophyte;(NI), no indicator; **, introduced; UPL, upland nonhydrophyte; ssp., subspecies; var., variety]

Code National PLANTS Database taxonomy Flora of North American taxonomy Wetness

Genus and species Common name Genus and species Common name

Grasses—Continued

GLYCAN Glyceria canadensis Rattlesnake mannagrass Glyceria canadensis Rattlesnake-mannagrass OBL

MUHGLO Muhlenbergia glomerata Spike muhly Muhlenbergia glomerata Marsh muhly FACW+

PANBOR Dichanthelium boreale Northern panicgrass Panicum boreale Northern panic-grass --

POASPP Poa spp. Bluegrass Poa spp. Bluegrass --

Forbs

FORB Forb, unknown spp. -- -- -- --

ASTNEM Oclemena nemoralis Bog aster Aster nemoralis Leafy bog-aster OBL

ASTUMB Doellingeria umbellata Parasol whitetop Aster umbellatus Tall flat-topped white aster FACW

CAMAPA Campanula aparinoides Marsh bellflower Campanula aparinoides Marsh-bellflower OBL

CHALUT Chamaelirium luteum Fairywand Chamaelirium luteum Fairywand FACU

CHEGLA Chelone glabra White turtlehead Chelone glabra White turtlehead OBL

CICBUL Cicuta bulbifera Bulblet-bearing water 

hemlock

Cicuta bulbifera Bulblet-bearing waterhemlock

FACW+

CIRMUT Cirsium muticum Swamp thistle Cirsium muticum Swamp thistle OBL

CIRPAL Cirsium palustre** Marsh thistle Cirsium palustre** Marsh thistle (NI)

COPTRI Coptis trifolia Threeleaf goldthread Coptis trifolia Threeleaf goldthread (NI)

DROROT Drosera rotundifolia Roundleaf sundew Drosera rotundifolia Round-leaved sundew OBL

EUPMAC Eupatoriadelphus maculatusSpotted joe pye weed Eupatorium maculatum Spotted joe-pye weed --

EUPPER Eupatorium perfoliatum Common boneset Eupatorium perfoliatum Boneset FACW+

EUTGRA Euthamia graminifolia Flat-topped goldenrod Euthamia graminifolia Common flat-topped 

goldenrod

FACW−

GALLAB Galium labradoricum Northern bog bedstraw Galium labradoricum Labrador bedstraw OBL

GALTRI Galium trifidum Threepetal bedstraw Galium trifidum Northern three-lobed 

bedstraw

FACW+

GENRUB Gentiana rubricaulis Closed gentian Gentiana rubricaulis Great Lakes gentian --

HYPKAL Hypericum kalmianum Kalm’s St. Johnswort Hypericum kalmianum Kalm’s St. Johns-wort FACW+

IRIVER Iris versicolor Harlequin blueflag Iris versicolor Blue flag OBL

LATPAL Lathyrus palustris Marsh pea Lathyrus palustris Marsh-pea FACW

LYCUNI Lycopus uniflorus Northern bugleweed Lycopus uniflorus Northern water-horehound OBL

LYSTER Lysimachia terrestris Earth loosestrife Lysimachia terrestris Swamp-candles OBL

LYSTHY Lysimachia thyrsiflora Tufted loosestrife Lysimachia thyrsiflora Swamp loosestrife OBL

MAICAN Maianthemum canadense Canada mayflower Maianthemum canadense Canada mayflower UPL/

FAC

MENARV Mentha arvensis Wild mint Mentha arvensis Field mint FACW

PLAHYP Platanthera hyperborea Northern green orchid Platanthera hyperborea Northern bog-orchid FACW+

POTFRU Dasiphora fruticosa Shrubby cinquefoil Potentilla fruticosa Shrubby-cinquefoil FACW

POTPAL Comarum palustre Purple marshlocks Potentilla palustris Marsh cinquefoil OBL

RUMORB Rumex orbiculatus Greater water dock Rumex orbiculatus Great water-dock OBL

SMITRI Maianthemum trifolium Threeleaf false lily-of-thevalley

Smilacina trifolia False Solomon’s seal --
52 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Table 1.1. Crosswalk table of taxonomy of plant species following the Flora of North America (2003) and U.S. Department of Agriculture 

National PLANTS Database (U.S. Department of Agriculture, Natural Resources Conservation Service, 2003).—Continued

[List includes species that may have been found only in blocks C3–7 or C3–8, which were sampled in 2006 and 2007 but not included in analyses. Codes 

are those used in the field and analyses. Wetness, an indicator of a species’ tolerance to wet soils, was from Chadde (2002) to be specific to the study region. 

Some codes differ from the PLANTS scientific name because the names in the regional guide used (Chadde, 2002) differ from the most recent taxonomy in the 

PLANTS Database. Wetness, indicator of plant tolerance to wet soils; --, no data; OBL, obligate wetland hydrophyte; FACW, facultative wetland hydrophyte; 

spp., species; FAC, facultative hydrophyte;(NI), no indicator; **, introduced; UPL, upland nonhydrophyte; ssp., subspecies; var., variety]

Code National PLANTS Database taxonomy Flora of North American taxonomy Wetness

Genus and species Common name Genus and species Common name

Forbs—Continued

SOLCAN Solidago canadensis Canada goldenrod Solidago canadensis Canada goldenrod FACU

SOLGIG Solidago gigantea Giant goldenrod Solidago gigantea Smooth goldenrod FACW

SOLIDAG Solidago spp. Goldenrod spp. Solidago spp. Goldenrod spp. --

SOLULI Solidago uliginosa Bog goldenrod Solidago uliginosa Northern bog goldenrod OBL

SYMBOR Symphyotrichum boreale Northern bog aster Aster borealis Northern bog aster OBL

SYMPUN Symphyotrichum puniceum Purplestem aster Aster puniceus Purple-stem aster OBL

TRIFRA Triadenum fraseri Fraser’s marsh St. Johnswort

Triadenum fraseri Marsh St. John’s wort OBL

URTDIO Urtica dioica Stinging nettle Urtica dioica Stinging nettle FAC+

UTRINT Utricularia intermedia Flatleaf bladderwort Utricularia intermedia Flatleaf bladderwort OBL

VIOMAC Viola macloskeyi Small white violet Viola macloskeyi Smooth white violet OBL

VIONEP Viola nephrophylla Northern bog violet Viola nephrophylla Northern bog violet FACW+

Shrubs/woody taxa

ALDINC Alnus incana spp. rugosa Speckled alder Alnus incana Speckled alder OBL

ANDGLA Andromeda polifolia var. 

glaucophylla

Bog rosemary Andromeda glaucophylla Bog rosemary OBL

AROMEL Aronia melanocarpa Black chokecherry Aronia melanocarpa Black chokecherry FACW−

BETPAP Betula papyrifera White birch Betula papyrifera White birch --

BETPUM Betula pumila Bog birch Betula pumila Bog birch OBL

CHACAL Chamaedaphne calyculata Leatherleaf Chamaedaphne calyculata Leatherleaf OBL

ILEVER Ilex verticillata Common winterberry Ilex verticillata Winterberry, black-alder FACW+

LARLAR Larix laricina Tamarack Larix laricina Tamarack FACW

LEDGRO Ledum groenlandicum Bog Labrador tea Ledum groenlandicum Labrador tea OBL

ORTSEC Orthilia secunda Sidebells wintergreen Orthilia secunda Sidebells wintergreen FAC+

PICGLA Picea glauca White spruce Picea glauca White spruce FACU

PICMAR Picea mariana Black spruce Picea mariana Black spruce FACW

PYRASA Pyrola asarifolia Liverleaf wintergreen Pyrola asarifolia Pink shinleaf FACW

ROSPAL Rosa palustris Swamp rose Rosa palustris Swamp rose OBL

RUBARC Rubus arcticus ssp. acaulis Arctic blackberry Rubus acaulis Dwarf raspberry OBL

RUBHIS Rubus hispidus Bristly dewberry Rubus hispidus Swamp dewberry FACW

RUBPUB Rubus pubescens Dwarf red blackberry Rubus pubescens Dwarf raspberry FACW+

RUBUS Rubus spp. Raspberry spp. Rubus spp. Raspberry spp. --

SALCAN Salix candida Sageleaf willow Salix candida Sage-leaf willow OBL

SALIX Salix spp. Willow spp. -- -- --

SALPED Salix pedicellaris Bog willow Salix pedicellaris Bog-willow OBL

SPIALB Spiraea alba Meadowsweet Spiraea alba Meadowsweet FACW+

SPITOM Spiraea tomentosa Steeplebush Spiraea tomentosa Steeple-bush FACW

VACCIN Vaccinium spp. Vaccinium spp. Vaccinium spp. Vaccinium spp. --

VACMAC Vaccinium macrocarpon Cranberry Vaccinium macrocarpon Large cranberry OBL

VACMYR Vaccinium myrtilloides Velvetleaf huckleberry Vaccinium myrtilloides Velvetleaf-blueberry FACW
Appendixes 53

Table 1.1. Crosswalk table of taxonomy of plant species following the Flora of North America (2003) and U.S. Department of Agriculture 

National PLANTS Database (U.S. Department of Agriculture, Natural Resources Conservation Service, 2003).—Continued

[List includes species that may have been found only in blocks C3–7 or C3–8, which were sampled in 2006 and 2007 but not included in analyses. Codes 

are those used in the field and analyses. Wetness, an indicator of a species’ tolerance to wet soils, was from Chadde (2002) to be specific to the study region. 

Some codes differ from the PLANTS scientific name because the names in the regional guide used (Chadde, 2002) differ from the most recent taxonomy in the 

PLANTS Database. Wetness, indicator of plant tolerance to wet soils; --, no data; OBL, obligate wetland hydrophyte; FACW, facultative wetland hydrophyte; 

spp., species; FAC, facultative hydrophyte;(NI), no indicator; **, introduced; UPL, upland nonhydrophyte; ssp., subspecies; var., variety]

Code National PLANTS Database taxonomy Flora of North American taxonomy Wetness

Genus and species Common name Genus and species Common name

Nonangiosperms

DRYCRI Dryopteris cristata Crested woodfern Dryopteris cristata Crested wood-fern OBL

EQUSPP Equisetum spp. Horsetail Equisetum spp. Horsetail --

ONOSEN Onoclea sensibilis Sensitive fern Onoclea sensibilis Sensitive fern FACW+

OPHVUL Ophioglossum vulgatum Southern adderstongue Ophioglossum vulgatum Adder’s tongue FACW

OSMREG Osmunda regalis Royal fern Osmunda regalis Royal fern OBL

PTEAQU Pteridium aquilinum Western bracken fern Pteridium aquilinum Bracken fern FACU

THEPAL Thelypteris palustris Eastern marsh fern Thelypteris palustris Eastern marsh fern --

References Cited

Chadde, S.W., 2002, A Great Lakes wetland flora (2d ed.): 

Laurium, Mich., PocketFlora Press, 648 p.

Flora of North America Editorial Committee, eds., 2003, Flora 

of North America North of Mexico, v. 4: New York, Oxford 

University Press, 512 p.

U.S. Department of Agriculture, Natural Resources Conservation Service, 2003, The PLANTS Database: U.S. 

Department of Agriculture, Natural Resources Conservation Service web page, accessed October 24, 2017, at 

https://plants.sc.egov.usda.gov/java/.
54 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge

Appendix 2. Fire Conditions During Prescribed Burns at 

Marsh Creek, July 2007 and 2008, and C3, May 2008.

Table 2.1. Fire conditions during prescribed burns at Marsh Creek, July 2007 and 2008, and C3, 

May 2008. Data were obtained from Seney National Wildlife Refuge remote area weather station 

(MesoWest, 2008) and field observations.

[The Keetch-Byram Drought Code is a numerical rating representing the net effect of evapotranspiration and precipitation in producing cumulative moisture deficiency in deep duff and upper soil layers; the index ranges from 0, the point 

of no moisture deficiency, to 800, the maximum drought that is possible. Fuel moisture readings were obtained from 

the remote area weather station at 13:00 hours. The fire adjective rating is from the Canadian Forest Fire Danger Rating 

System, as used by Michigan Department of Natural Resources and Seney National Wildlife Refuge (Stocks and others, 

1989). Water depth was depth below soil surface to pooled water. ha, hectare; kph, kilometer per hour; cm, centimeter]

Marsh Creek C3

Burn date July 17, 2007 July 24, 2008 May 9, 2008

Burn area (ha) 21 155 341

Relative humidity (percent) 50–30 74–40 70–32

Air temperature 24–28 22–28 8–18

Wind speed (kph) 6–18 5–13 0–9

Cloud cover (percent) 30–60 40–80 80–100

Keetch-Byram Drought Code 207 116 4.5

1-hour fuel moisture 5.8 8.6 7.4

10-hour fuel moisture 6.2 11.6 9.9

100-hour fuel moisture 17.8 16 15.4

Fire adjective rating High 

(summer criteria)

Moderate 

(summer criteria)

Moderate 

(spring criteria)

Water depth (cm) −12 1–7 0–6

References Cited

MesoWest, 2008, Great Lakes fire and fuels: MesoWest digital data, accessed July 2008 at 

https://glff.mesowest.org/.

Stocks, B.J., Lynham, T.J., Lawson, B.D., Alexander, M.E., Wagner, C.V., McAlpine, R.S., 

Dube, D.E., 1989, Canadian forest fire danger rating system—An overview: The Forestry 

Chronicle, v. 65, no. 6, p. 450–457.
Appendixes 55

Appendix 3. Frequency of Occurrence of Plant Taxa by Block in C3 and Marsh 

Creek, Seney National Wildlife Refuge, 2006–10. Table 3.1. Frequency of occurrence and percent of points summed across sampling years of plant taxa by block in C3 of Seney National Wildlife Refuge, Michigan, 2006–10. Blocks C3–1 to C3–6 were sampled 5 years (total 125 points); C3–7 and C3–8 were sampled 2 years (total 50 points). [See appendix 1 for definition of taxa codes. No., number; %, percent] Taxa code C3–1 C3–2 C3–3 C3–4 C3–5 C3– 6 C3–7 C3– 8 C3, 1 to 6 No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

%

AGRHYE 5 4.0 14 11.2 17 13.6 27 21.6 16 12.8 5 4.0 5 10 2 4 84 11.2AGRSTO 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 1 0.1ALNINC 0 0.0 0 0.0 1 0.8 0 0.0 4 3.2 13 10.4 1 2 5 10 18 2.4ANDGLA 0 0.0 0 0.0 1 0.8 0 0.0 1 0.8 0 0.0 1 2 0 0 2 0.3AROMEL 0 0.0 0 0.0 1 0.8 0 0.0 1 0.8 1 0.8 0 0 0 0 3 0.4ASTNEM 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 0 0 2 0.3ASTUMB 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 0 0 2 4 1 0.1BETPAP 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0 0 0 1 0.1BETPUM 17 13.6 52 41.6 43 34.4 24 19.2 66 52.8 25 20.0 12 24 7 14 227 30.3BROCIL 0 0.0 0 0.0 2 1.6 0 0.0 13 10.4 10 8.0 10 20 0 0 25 3.3CALCAN 42 33.6 40 32.0 35 28.0 43 34.4 54 43.2 72 57.6 49 98 45 90 286 38.1CAMAPA 38 30.4 14 11.2 49 39.2 102 81.6 75 60.0 83 66.4 23 46 26 52 361 48.1CARAQU 2 1.6 0 0.0 0 0.0 0 0.0 2 1.6 8 6.4 4 8 11 22 12 1.6CARBEB 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 2 1 2 0 0.0CARBUX 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0CARCHO 32 25.6 36 28.8 85 68.0 94 75.2 89 71.2 109 87.2 36 72 20 40 445 59.3CAREX1 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0CAREX2 25 20.0 25 20.0 25 20.0 25 20.0 25 20.0 25 20.0 24 48 22 44 150 20.0CAREX3 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 1 0.1CAREX4 22 17.6 8 6.4 8 6.4 8 6.4 2 1.6 1 0.8 4 8 1 2 49 6.5CAREX9 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0CAREXI 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0CARHYA 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 0 0 1 0.1CARINT 27 21.6 55 44.0 84 67.2 48 38.4 67 53.6 40 32.0 10 20 8 16 321 42.8CARLAS 98 78.4 100 80.0 98 78.4 97 77.6 86 68.8 92 73.6 24 48 22 44 571 76.1CARLEP 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 0 0 1 0.1CARLIV 0 0.0 4 3.2 0 0.0 4 3.2 1 0.8 10 8.0 0 0 0 0 19 2.5CAROLI 1 0.8 3 2.4 1 0.8 0 0.0 0 0.0 0 0.0 0 0 0 0 5 0.7CARROS 3 2.4 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 3 0.4CARUTR 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 3 2.4 0 0 0 0 4 0.5
56 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge Table 3.1. Frequency of occurrence and percent of points summed across sampling years of plant taxa by block in C3 of Seney National Wildlife Refuge, Michigan, 2006–10. Blocks C3–1 to C3–6 were sampled 5 years (total 125 points); C3–7 and C3–8 were sampled 2 years (total 50 points).—Continued [See appendix 1 for definition of taxa codes. No., number; %, percent] Taxa code C3–1 C3–2 C3–3 C3–4 C3–5 C3– 6 C3–7 C3– 8 C3, 1 to 6 No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

%

CHACAL 1 0.8 1 0.8 0 0.0 0 0.0 1 0.8 3 2.4 0 0 6 12 6 0.8CHALUT 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 0 0 1 0.1CHEGLA 0 0.0 0 0.0 0 0.0 0 0.0 3 2.4 1 0.8 0 0 1 2 4 0.5CICBUL 0 0.0 0 0.0 2 1.6 1 0.8 2 1.6 2 1.6 0 0 2 4 7 0.9CIRMUT 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 1 0.1CIRPAL 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0COPTRI 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0DROROT 7 5.6 7 5.6 0 0.0 1 0.8 9 7.2 0 0.0 0 0 0 0 24 3.2DRYCRI 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 1 2 1 0.1DULARU 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0ELEELL 119 95.2 49 39.2 96 76.8 99 79.2 10 8.0 33 26.4 0 0 0 0 406 54.1ELEROB 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0EQUSPP 0 0.0 0 0.0 5 4.0 0 0.0 0 0.0 0 0.0 0 0 0 0 5 0.7ERIANG 5 4.0 7 5.6 4 3.2 1 0.8 2 1.6 3 2.4 0 0 0 0 22 2.9ERIVIR 86 68.8 65 52.0 33 26.4 41 32.8 78 62.4 73 58.4 19 38 28 56 376 50.1EUPMAC 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 1 0.8 0 0 0 0 2 0.3EUPPER 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 1 2 0 0.0EUTGRA 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0FORB 4 3.2 8 6.4 3 2.4 0 0.0 6 4.8 8 6.4 10 20 8 16 29 3.9GALLAB 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0GALTRI 0 0.0 0 0.0 0 0.0 1 0.8 2 1.6 2 1.6 4 8 5 10 5 0.7GENRUB 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0GLYCAN 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 6 4.8 0 0 0 0 6 0.8GRASS4 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0HYPKAL 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 1 0.1ILEVER 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0IRIVER 0 0.0 2 1.6 3 2.4 0 0.0 6 4.8 25 20.0 28 56 24 48 36 4.8JUNCAN 8 6.4 20 16.0 2 1.6 3 2.4 0 0.0 0 0.0 0 0 0 0 33 4.4LARLAR 0 0.0 3 2.4 7 5.6 6 4.8 12 9.6 2 1.6 0 0 0 0 30 4.0LATPAL 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 1 0.8 0 0 0 0 2 0.3
Appendixes 57

Table 3.1. Frequency of occurrence and percent of points summed across sampling years of plant taxa by block in C3 of Seney National Wildlife Refuge, Michigan, 2006–10. Blocks C3–1 to C3–6 were sampled 5 years (total 125 points); C3–7 and C3–8 were sampled 2 years (total 50 points).—Continued[See appendix 1 for definition of taxa codes. No., number; %, percent]

Taxa code

C3–1 C3–2 C3–3 C3–4 C3–5 C3– 6 C3–7 C3– 8 C3, 1 to 6

No. % No. % No. % No. % No. % No. % No. % No. % No. %

LEDGRO 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0LYCUNI 0 0.0 0 0.0 0 0.0 2 1.6 2 1.6 5 4.0 2 4 3 6 9 1.2LYSTER 18 14.4 20 16.0 17 13.6 13 10.4 3 2.4 18 14.4 14 28 14 28 89 11.9LYSTHR 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0MAICAN 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 0 0 1 0.1MENARV 0 0.0 0 0.0 1 0.8 0 0.0 1 0.8 2 1.6 3 6 0 0 4 0.5MOSS 66 52.8 91 72.8 75 60.0 58 46.4 83 66.4 50 40.0 34 68 24 48 423 56.4MUHGLO 84 67.2 97 77.6 92 73.6 111 88.8 64 51.2 71 56.8 1 2 1 2 519 69.2ONOSEN 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 6 4.8 4 8 5 10 6 0.8OPEN 92 73.6 96 76.8 96 76.8 84 67.2 96 76.8 98 78.4 25 50 25 50 562 74.9OPHVUL 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 0 0 1 0.1ORTSEC 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 0 0 1 0.1OSMREG 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0PANBOR 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0PICGLA 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 0 0 0 0 1 0.1PICMAR 0 0.0 0 0.0 1 0.8 0 0.0 1 0.8 0 0.0 0 0 0 0 2 0.3PLAHYP 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0 0 0 1 0.1POASP 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0POTFRU 29 23.2 32 25.6 3 2.4 12 9.6 52 41.6 7 5.6 0 0 0 0 135 18.0POTPAL 0 0.0 0 0.0 2 1.6 0 0.0 0 0.0 2 1.6 1 2 3 6 4 0.5PTEAQU 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0PYRASA 0 0.0 0 0.0 0 0.0 0 0.0 2 1.6 0 0.0 0 0 0 0 2 0.3RHYALB 92 73.6 18 14.4 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 110 14.7ROSPAL 0 0.0 0 0.0 0 0.0 0 0.0 2 1.6 0 0.0 0 0 0 0 2 0.3RUBARC 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 6 12 2 4 1 0.1RUBHIS 0 0.0 0 0.0 2 1.6 0 0.0 2 1.6 6 4.8 6 12 4 8 10 1.3RUBPUB 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 0 0 1 0.1RUBUS 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 2 1.6 0 0 0 0 3 0.4RUMORB 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 2 1.6 0 0 1 2 3 0.4SALCAN 0 0.0 0 0.0 1 0.8 2 1.6 0 0.0 0.0 1 2 0 0 3 0.4
58 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge Table 3.1. Frequency of occurrence and percent of points summed across sampling years of plant taxa by block in C3 of Seney National Wildlife Refuge, Michigan, 2006–10. Blocks C3–1 to C3–6 were sampled 5 years (total 125 points); C3–7 and C3–8 were sampled 2 years (total 50 points).—Continued [See appendix 1 for definition of taxa codes. No., number; %, percent] Taxa code C3–1 C3–2 C3–3 C3–4 C3–5 C3– 6 C3–7 C3– 8 C3, 1 to 6 No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

%

SALIX 23 18.4 10 8.0 17 13.6 26 20.8 7 5.6 18 14.4 3 6 4 8 101 13.5SALPED 2 1.6 4 3.2 5 4.0 3 2.4 1 0.8 1 0.8 0 0 0 0 16 2.1SCICYP 0 0.0 2 1.6 0 0.0 0 0.0 0 0.0 0 0.0 0 0 3 6 2 0.3SMITRI 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 2 1.6 5 10 3 6 2 0.3SOLCAN 0 0.0 0 0.0 1 0.8 0 0.0 6 4.8 1 0.8 0 0 0 0 8 1.1SOLGIG 0 0.0 2 1.6 0 0.0 0 0.0 4 3.2 6 4.8 10 20 14 28 12 1.6SOLIDAG 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 2 0 0 0 0.0SOLULI 3 2.4 4 3.2 0 0.0 3 2.4 6 4.8 7 5.6 0 0 0 0 23 3.1SPIALB 0 0.0 2 1.6 0 0.0 0 0.0 0 0.0 0 0.0 1 2 4 8 2 0.3SPITOM 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0SYMBOR 38 30.4 24 19.2 32 25.6 7 5.6 24 19.2 42 33.6 9 18 18 36 167 22.3SYMPUM 3 2.4 10 8.0 14 11.2 4 3.2 13 10.4 17 13.6 21 42 12 24 61 8.1THEPAL 0 0.0 0 0.0 0 0.0 0 0.0 11 8.8 4 3.2 3 6 3 6 15 2.0TRIFRA 8 6.4 8 6.4 16 12.8 14 11.2 10 8.0 16 12.8 11 22 15 30 72 9.6TYPLAT 0 0.0 0 0.0 25 20.0 22 17.6 0 0.0 8 6.4 0 0 0 0 55 7.3URTDIO 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 0 0 1 0.1UTRINT 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0 0 0 0 0.0VACCIN 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0 0 0 1 0.1VACMAC 0 0.0 2 1.6 8 6.4 0 0.0 3 2.4 0 0.0 0 0 0 0 13 1.7VACMYR 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0 0 0 1 0.1VIOMAC 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0 0 0 1 0.1VIONEP 26 20.8 29 23.2 13 10.4 5 4.0 28 22.4 14 11.2 9 18 9 18 115 15.3
Appendixes 59

Appendix 4. Frequency of Occurrence and Percent of Points (Summed Across 

Sampling Years) of Plant Taxa by Block in Marsh Creek, Seney National Wildlife 

Refuge, Michigan, 2006–10. Table 4.1. Frequency of occurrence and percent of points (summed across sampling years) of plant taxa by block in Marsh Creek of Seney National Wildlife Refuge, Michigan, 2006–10. Blocks were sampled 5 years (total 125 points). [See appendix 1 for definition of taxa codes. No., number; %, percent] Taxa code MC–1 MC–2 MC–3 MC–4 MC–5 MC–6 MC–7 MC–8 MC–1, 3 to 8 No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

%

AGRHYE 15 12.0 13 10.4 16 12.8 10 8.0 6 4.8 6 4.8 6 4.8 3 2.4 75 7.7AGRSTO 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0ALNINC 1 0.8 0 0.0 1 0.8 2 1.6 0 0.0 1 0.8 3 2.4 0 0.0 8 0.8ANDGLA 0 0.0 0 0.0 0 0.0 1 0.8 31 24.8 5 4.0 4 3.2 29 23.2 70 7.2AROMEL 1 0.8 0 0.0 0 0.0 0 0.0 3 2.4 2 1.6 2 1.6 4 3.2 12 1.2ASTNEM 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0ASTUMB 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.1BETPAP 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0BETPUM 0 0.0 1 0.8 42 33.6 29 23.2 8 6.4 16 12.8 68 54.4 38 30.4 202 20.7BROCIL 0 0.0 1 0.8 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 2 0.2CALCAN 58 46.4 57 45.6 68 54.4 83 66.4 20 16.0 39 31.2 48 38.4 17 13.6 390 40.0CAMAPA 59 47.2 39 31.2 58 46.4 80 64.0 32 25.6 93 74.4 46 36.8 36 28.8 443 45.4CARAQU 0 0.0 0 0.0 5 4.0 0 0.0 0 0.0 3 2.4 0 0.0 1 0.8 9 0.9CARBEB 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 1 0.1CARBUX 7 5.6 9 7.2 14 11.2 18 14.4 19 15.2 2 1.6 8 6.4 14 11.2 91 9.3CARCHO 83 66.4 73 58.4 121 96.8 116 92.8 98 78.4 119 95.2 116 92.8 79 63.2 805 82.6CAREX1 20 16.0 25 20.0 10 8.0 14 11.2 12 9.6 1 0.8 5 4.0 13 10.4 100 10.3CAREX2 19 15.2 16 12.8 25 20.0 25 20.0 26 20.8 25 20.0 29 23.2 25 20.0 190 19.5CAREX3 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0CAREX4 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0CAREX9 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.1CAREXI 1 0.8 1 0.8 0 0.0 1 0.8 2 1.6 0 0.0 4 3.2 7 5.6 16 1.6CARHYA 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0CARINT 28 22.4 54 43.2 68 54.4 73 58.4 24 19.2 68 54.4 21 16.8 45 36.0 381 39.1CARLAS 7 5.6 18 14.4 20 16.0 20 16.0 16 12.8 12 9.6 10 8.0 11 8.8 114 11.7CARLEP 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0CARLIV 78 62.4 44 35.2 15 12.0 63 50.4 56 44.8 60 48.0 90 72.0 90 72.0 496 50.9CAROLI 84 67.2 67 53.6 79 63.2 81 64.8 69 55.2 70 56.0 47 37.6 75 60.0 572 58.7CARROS 0 0.0 2 1.6 6 4.8 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 9 0.9CARUTR 0 0.0 1 0.8 11 8.8 0 0.0 1 0.8 0 0.0 0 0.0 1 0.8 14 1.4
60 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge Table 4.1. Frequency of occurrence and percent of points (summed across sampling years) of plant taxa by block in Marsh Creek of Seney National Wildlife Refuge, Michigan, 2006–10. Blocks were sampled 5 years (total 125 points).—Continued [See appendix 1 for definition of taxa codes. No., number; %, percent] Taxa code MC–1 MC–2 MC–3 MC–4 MC–5 MC–6 MC–7 MC–8 MC–1, 3 to 8 No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

%

CHACAL 47 37.6 5 4.0 101 80.8 46 36.8 103 82.4 101 80.8 96 76.8 116 92.8 615 63.1CHALUT 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0CHEGLA 0 0.0 0 0.0 0 0.0 0 0.0 2 1.6 0 0.0 1 0.8 0 0.0 3 0.3CICBUL 4 3.2 0 0.0 1 0.8 3 2.4 0 0.0 2 1.6 3 2.4 0 0.0 13 1.3CIRMUT 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.1CIRPAL 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 4 3.2 0 0.0 4 0.4COPTRI 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 1 0.1DROROT 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.1DRYCRI 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 1 0.1DULARU 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.1ELEELL 48 38.4 9 7.2 34 27.2 17 13.6 43 34.4 32 25.6 43 34.4 21 16.8 247 25.3ELEROB 3 2.4 0 0.0 3 2.4 0 0.0 8 6.4 1 0.8 8 6.4 17 13.6 40 4.1EQUSPP 0 0.0 0 0.0 0 0.0 5 4.0 0 0.0 1 0.8 0 0.0 0 0.0 6 0.6ERIANG 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.1ERIVIR 92 73.6 70 56.0 103 82.4 98 78.4 68 54.4 60 48.0 77 61.6 84 67.2 652 66.9EUPMAC 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0EUPPER 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0EUTGRA 0 0.0 1 0.8 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 2 0.2FORB 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0FORB1 5 4.0 0 0.0 3 2.4 3 2.4 3 2.4 8 6.4 6 4.8 6 4.8 34 3.5GALLAB 0 0.0 0 0.0 6 4.8 13 10.4 3 2.4 12 9.6 2 1.6 0 0.0 36 3.7GALTRI 32 25.6 1 0.8 18 14.4 35 28.0 15 12.0 10 8.0 15 12.0 11 8.8 137 14.0GENRUB 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 1 0.1GLYCAN 2 1.6 5 4.0 3 2.4 0 0.0 0 0.0 4 3.2 1 0.8 0 0.0 15 1.5GRASS4 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 1 0.1HYPKAL 2 1.6 0 0.0 8 6.4 1 0.8 4 3.2 43 34.4 14 11.2 47 37.6 119 12.2ILEVER 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 3 2.4 0 0.0 3 0.3IRIVER 42 33.6 49 39.2 15 12.0 23 18.4 13 10.4 18 14.4 20 16.0 17 13.6 197 20.2JUNCAN 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0LARLAR 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0
Appendixes 61

Table 4.1. Frequency of occurrence and percent of points (summed across sampling years) of plant taxa by block in Marsh Creek of Seney National Wildlife Refuge, Michigan, 2006–10. Blocks were sampled 5 years (total 125 points).—Continued[See appendix 1 for definition of taxa codes. No., number; %, percent]

Taxa code

MC–1 MC–2 MC–3 MC–4 MC–5 MC–6 MC–7 MC–8 MC–1, 3 to 8

No. % No. % No. % No. % No. % No. % No. % No. % No. %

LATPAL 3 2.4 9 7.2 5 4.0 10 8.0 0 0.0 0 0.0 6 4.8 0 0.0 33 3.4LEDGRO 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 3 2.4 0 0.0 3 0.3LYCUNI 19 15.2 4 3.2 0 0.0 4 3.2 0 0.0 0 0.0 3 2.4 3 2.4 33 3.4LYSTER 8 6.4 19 15.2 3 2.4 9 7.2 0 0.0 6 4.8 1 0.8 9 7.2 55 5.6LYSTHY 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 1 0.1MAICAN 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0MENARV 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 1 0.8 3 0.3MOSS 28 22.4 7 5.6 82 65.6 62 49.6 68 54.4 75 60.0 57 45.6 81 64.8 460 47.2MUHGLO 40 32.0 11 8.8 29 23.2 53 42.4 11 8.8 40 32.0 19 15.2 13 10.4 216 22.1ONOSEN 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0OPEN 102 81.6 75 60.0 93 74.4 97 77.6 93 74.4 96 76.8 97 77.6 97 77.6 750 76.9OPHVUL 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0ORTSEC 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0OSMREG 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 2 0.2PANBOR 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 2 1.6 0 0.0 4 0.4PICGLA 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0PICMAR 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0PLAHYP 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0POASP 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 1 0.8 0 0.0 2 0.2POTFRU 2 1.6 0 0.0 0 0.0 0 0.0 27 21.6 67 53.6 11 8.8 51 40.8 158 16.2POTPAL 3 2.4 1 0.8 0 0.0 4 3.2 8 6.4 6 4.8 3 2.4 1 0.8 26 2.7PTEAQU 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 2 0.2PYRASA 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0RHYALB 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0ROSPAL 0 0.0 0 0.0 4 3.2 0 0.0 0 0.0 0 0.0 2 1.6 2 1.6 8 0.8RUBARC 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0RUBHIS 5 4.0 1 0.8 4 3.2 0 0.0 1 0.8 0 0.0 5 4.0 0 0.0 16 1.6RUBPUB 2 1.6 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 2 0.2RUBUS 2 1.6 0 0.0 0 0.0 1 0.8 0 0.0 1 0.8 1 0.8 0 0.0 5 0.5RUMORB 0 0.0 0 0.0 1 0.8 3 2.4 0 0.0 0 0.0 0 0.0 2 1.6 6 0.6
62 Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge Table 4.1. Frequency of occurrence and percent of points (summed across sampling years) of plant taxa by block in Marsh Creek of Seney National Wildlife Refuge, Michigan, 2006–10. Blocks were sampled 5 years (total 125 points).—Continued [See appendix 1 for definition of taxa codes. No., number; %, percent] Taxa code MC–1 MC–2 MC–3 MC–4 MC–5 MC–6 MC–7 MC–8 MC–1, 3 to 8 No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

% No.

%

SALCAN 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0SALIX 1 0.8 0 0.0 4 3.2 6 4.8 7 5.6 5 4.0 4 3.2 7 5.6 34 3.5SALPED 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0SCICYP 1 0.8 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 2 0.2SMITRI 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0SOLCAN 2 1.6 0 0.0 0 0.0 0 0.0 0 0.0 8 6.4 0 0.0 1 0.8 11 1.1SOLGIG 1 0.8 0 0.0 2 1.6 0 0.0 0 0.0 2 1.6 0 0.0 0 0.0 5 0.5SOLIDAG 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 2 0.2SOLULI 3 2.4 0 0.0 3 2.4 5 4.0 15 12.0 26 20.8 7 5.6 29 23.2 88 9.0SPIALB 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 2 0.2SPITOM 1 0.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 2 0.2SYMBOR 40 32.0 21 16.8 45 36.0 46 36.8 43 34.4 38 30.4 51 40.8 30 24.0 314 32.2SYMPUM 10 8.0 6 4.8 22 17.6 9 7.2 17 13.6 8 6.4 18 14.4 5 4.0 95 9.7THEPAL 1 0.8 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 4 3.2 0 0.0 6 0.6TRIFRA 10 8.0 8 6.4 11 8.8 11 8.8 9 7.2 2 1.6 5 4.0 2 1.6 58 5.9TYPLAT 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0URTDIO 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0UTRINT 0 0.0 6 4.8 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 6 0.6VACCIN 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 0 0.0 0 0.0 1 0.1VACMAC 9 7.2 0 0.0 0 0.0 7 5.6 0 0.0 4 3.2 0 0.0 6 4.8 26 2.7VACMYR 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0VIOMAC 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0VIONEP 12 9.6 0 0.0 11 8.8 18 14.4 15 12.0 20 16.0 19 15.2 11 8.8 106 10.9
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Austin and Newton—Response of Vegetation in Open and Partially Wooded Fens to Prescribed Burning at Seney National Wildlife Refuge —SIR 2018–5168

ISSN 2328-0328 (online)

https://doi.org/10.3133/sir20185168