Bryophyte Community Response to Prescribed Fire and Thinning in Mixed-Oak

Home , Atrichum, Cephaloziaceae, Fissidens dubius, Funaria hygrometrica, Hylocomiaceae, Hypnaceae

Forests of the Unglaciated Allegheny Plateau

A thesis presented to

the faculty of

the College of Arts and Sciences of Ohio University

In partial fulfillment

of the requirements for the degree

Master of Science

John J. Wiley Jr.

May 2013

This thesis titled

Bryophyte Community Response to Prescribed Fire and Thinning in Mixed-Oak

Forests of the Unglaciated Allegheny Plateau

JOHN J. WILEY JR.

has been approved for

the Department of Environmental and Plant Biology

and the College of Arts and Sciences by

Brian C. McCarthy

Professor of Environmental and Plant Biology

Robert Frank

Dean, College of Arts and Sciences 3

ABSTRACT

WILEY, JOHN J., JR., M.S., May 2013, Environmental and Plant Biology

Bryophyte Community Response to Prescribed Fire and Thinning in Mixed-Oak Forests of the Unglaciated Allegheny Plateau (94 pp.)

Director of Thesis: Brian C. McCarthy

Silvicultural treatments are applied to forests to meet a variety of management objectives including timber production and wildlife habitat management. However, such methods may also profoundly affect other non-timber forest resources and negatively impact biodiversity. Modern forest science has largely neglected the role of forest management activities on bryophytes; thus, we have little insight as to how bryophyte communities respond to stand level treatments. The goal of this investigation was to explore changes in mixed-oak forest bryophyte communities associated with the common silvicultural methods of prescribed fire and thinning. Study sites were within the design of the USDA Forest Service Fire and Fire Surrogate (FFS) Research Program located in three southeastern Ohio forests. Each of these forests contain four treatments: untreated control, prescribed fire only, thinning only, and combined prescribed fire and thinning.

Bryophyte occurrence and associated environmental variables were estimated in five 2 ×

5 m quadrats along nine linear transects stratified by an integrated moisture index (IMI) in each treatment. I found a total of 116 bryophyte taxa (97 mosses, 19 liverworts). Of these, only 65 were found in more than 5% of the transects. Burning in xeric sites clearly altered bryophyte community richness and composition as burned sites had a 50% 4

reduction in mean richness and a disproportionate decrease in corticolous species. Mesic moisture conditions appear to mitigate species loss and community change. Thinning strongly altered woody substrate availability; however, the increased availability of these substrates did not significantly alter the bryophyte community in relation to their abundance. The use of fire on xeric sites may have a profound long-term negative impact on bryophyte diversity. Thinning may have a multi-decadal legacy effect on bryophyte species that does not appear to significantly degrade the bryophyte community.

ACKNOWLEDGMENTS

I would like to express my gratitude to all those individuals and organizations that were integral to the development of this research. Foremost, I would like to thank my thesis advisor, Dr. Brian C. McCarthy. I gained much from his guidance and experience in academia, and on a several notable occasions, in life. I would also like to thank my committee members Dr. Morgan L. Vis and Dr. Jared L. DeForest for their input and advice during the development of this thesis and my graduate career.

Perhaps one of the most challenging and rewarding aspects of this research was immersing myself in the intricate and often cryptic world of bryophytes. My initial introduction was guided by Dr. Nancy Slack at the Humboldt Field Research Institute,

Maine. Later collaborations with the Ohio Moss and Lichen Association, especially Dr.

Barbara Andreas, Diane Lucas, and Dr. Robert Klipps, and the Kent State Herbarium proved to be invaluable in my understanding of the Ohio bryophyte flora.

The field work for this research was extensive and, often times, onerous, but would not have been possible without the access and information provided by the USFS

Vinton Furnace Experimental Station. Also, I would like to thank my field assistants,

Sarah Gutzwiller and Pualani Wiley. They both endured long days in less than ideal situations, and I am grateful for their companionship and dedication to detail.

Funding for this research was provided by the Ohio Biological Survey, Hiram

Roy Wilson Fund in Environmental and Plant Biology, Ohio University Graduate Student

Senate, Ohio Moss and Lichen Association, and Association of Southeastern Biologists. 6

TABLE OF CONTENTS

Page

Abstract ...... 3

Acknowledgments...... 5

List of Tables ...... 8

List of Figures ...... 12

Introduction ...... 15

Methods...... 22

Study Sites ...... 22

Study Design ...... 23

Field Methods ...... 25

Substrates ...... 26

Site Conditions ...... 29

Bryophytes ...... 29

Data Analysis ...... 30

Results ...... 35

Bryophyte Diversity ...... 35

Site Condition and Substrate Changes ...... 37

Bryophyte Community Response ...... 42

Taxonomic Relationships ...... 44

Treatments and Substrates ...... 45 7

Discussion ...... 48

Floristics ...... 48

A Synthesis ...... 50

Management Implications ...... 55

Conclusions ...... 59

Tables ...... 60

Figures...... 74

References ...... 83 8

LIST OF TABLES

Page

Table 1. Bryophyte species, general growth form (L, Leafy; T, Thalloid; A,

Acrocarpous; P, Pleurocarpous), coefficient of conservatism (C of C; higher is

more habitat specific) (Andreas et al. 2004), forest of occurrence (R, Raccoon

Ecological Management Area; T, Tar Hollow; Z, Zaleski;*,county

record;`,verified historical record), and accession number from the

Cooperrider Herbarium, Kent State University collected in three mix-oak

forests of southeastern Ohio...... 60

Table 2. Most commonly encountered (> 10% of all transects, n = 108) bryophytes in

three mixed-oak forests in southeastern Ohio...... 64

Table 3. Forest diversity metrics: number of species (S), Shannon-Weiner diversity

(H’), and evenness (J) for the bryophyte community occurring in three mixed-

oak forests in southeastern Ohio...... 65

Table 4. Mean (± S.E.) of site condition variables and substrate cover values within 2

× 5 m plots established in xeric, intermediate, and mesic moisture classes

under a 2 × 2 factorial of silvilcultural thinning and prescribed fire in three

mixed-oak forest in southeastern Ohio. Cover of substrate types represent the

2-D coverage within the plot as visually mapped onto a gridded representation

of the plot. Cover types are: exposed soil (SOIL), leaf litter (LEAF), coarse

woody debris (CWD, diameter ≥ 10.0 cm, < 45° from the ground), fine woody 9

debris (FWD, diameter ≥ 1 cm and < 10 cm), tree base cover (TREE, all stems

>DBH in height), stump base (STUMP, dead standing stems ≥ 45° from the

ground and < diameter at breast height [DBH] in height), and rock (ROCK)...... 66

Table 5. ANOVA model results (F-value, P-value) for environmental and substrate

responses to silvilcultural thinning and prescribed fire in mixed-oak forests of

southeastern Ohio. ANOVA models were formulated as Forest + Burn × Thin

× Moisture and evaluated for additive effects of cover of ROCK or Slope as

covariates. Burn and Thin are 1 df factors whether yes or no, and Moisture

was split into two 1 df orthogonal contrasts comparing intermediate vs. the

xeric and mesic moisture classes (IvXM) and the xeric vs. mesic moisture

classes (XvM). MANOVA was conducted on the subset of variables indicated

below the MANOVA results and separate ANOVAs were calculated for each

variable within the MANOVA and the remainder in the table. Only significant

(P < 0.05 for main effects, P < 0.10 for interactions) and marginally

significant (P < 0.10, for main effects only) results are shown. Cover types

are: exposed soil (SOIL), leaf litter (LEAF), coarse woody debris (CWD,

diameter ≥ 10.0 cm, < 45° from the ground), fine woody debris (FWD,

diameter ≥ 1 cm and < 10 cm), tree base cover (TREE, all stems >DBH in

height), stump base (STUMP, dead standing stems ≥ 45° from the ground and

< diameter at breast height [DBH] in height), and rock (ROCK)...... 67 10

Table 6. MANOVA analysis of forest substrate cover change after burning and

thinning across xeric (X), intermediate (I), and mesic (M) moisture types as

determined by an integrated moisture index (IMI) in three southeastern Ohio

forests. Substrates included in this analysis were exposed soil (SOIL), coarse

woody debris (CWD), fine woody debris (FWD), and tree base cover (TREE).

Cover of rock (ROCK) and slope were included as covariates, and orthogonal

1 df contrasts for moisture are represented...... 68

Table 7. Mean (± S.E.) density (pieces ha-1), volume (m3 ha-1), relative density

(RDEN), relative volume (RVOL), and relative importance (RIV) of coarse

woody debris (CWD) taxa in three mixed-oak forests in southeastern Ohio...... 69

Table 8. Mean (± S.E.) density (stems ha-1), volume (m3 ha-1), relative density

(RDEN), relative volume (RVOL), and relative importance (RIV) of tree

species in three mixed-oak forests in southeastern Ohio...... 70

Table 9. A permutational multivariate analysis of variance using distance matrices

(adonis::vegan (Oksanen et al. 2012)) of the bryophyte communities within

the Fire and Fire Surrogate (FFS) study design in three southeastern Ohio

forests. Data were permuted 9999 times within blocks (forests) as strata using

the Bray-Curtis distance matrix...... 71

Table 10. Multivariate analysis of variance (MANOVA) of bryophyte substrate

occurrence change by burning and thinning across moisture types in three

southeastern Ohio forests. Substrates included in this analysis were exposed 11

soil (SOIL), coarse woody debris (CWD+STUMP), fine woody debris

(FWD), and tree base cover (TREE). Orthogonal 1 df contrasts for moisture

are represented...... 72

Table 11. Multivariate analysis of variance (MANOVA) of analysis of bryophyte

substrate occurrence change incorporating co-variations in the substrates,

themselves by burning and thinning across moisture types in three

southeastern Ohio forests. Substrates included in this analysis were exposed

soil (SOIL), coarse woody debris (CWD+STUMP), fine woody debris

(FWD), and tree base cover (TREE). Cover of rock (ROCK) and slope were

included as covariates, and orthogonal 1 df contrasts for moisture are

represented...... 73

LIST OF FIGURES

Page

Figure 1. Location map of the three forests of the Ohio Hills Site of the USDA Fire

and Fire Surrogate (FFS) Research Program: Raccoon Ecological

Management Area (REMA), Tar Hallow State Forest (TAR), and Zaleski

State Forest (ZAL). Forest cover is shown in green...... 74

Figure 2. Treatment unit representation. Nine transects determined by a stratified

selection based on long-term moisture conditions derived from an integrated

moisture index (IMI) and slope orientation. Stratified by moisture conditions

ranging from xeric (dry, red), intermediate (intermediate, green), and mesic

(moist, blue) such that three transects were in each the xeric, intermediate, and

mesic moisture classes. Diagonal transects involved only the central 50 m

between grid points. Grid spacing is 50 m, contour interval is 6 m, and north is

at the top of the page...... 75

Figure 3. Representation of selected 2 5 m quadrats along established 50 m

transects. Quadrat “1” was placed 2.5 m (orthogonal) or 10.4 m (diagonal)

from the grid point. Additional quadrats were spaced 5 m apart. Hatched areas

represent grid point buffers, light grey areas are unsampled areas, and dark

circles indicate grid points...... 76

Figure 4. Number of transects of occurrence (n = 108) for bryophyte species (n =

112) observed in three southeastern Ohio forests...... 77 13

Figure 5. Bryophyte richness (n = 112) of individual transects (n = 108) from three

southeastern Ohio forests...... 78

Figure 6. Non-metric multidimensional scaling (NMDS) ordination and

environmental correlates of the bryophyte community a) as a whole and b-d)

divided out by moisture classes in three mixed-oak forests in southeastern

Ohio. Transects are labeled by treatment within a 2 × 2 factorial of

silvicultural thinning and prescribed burning with control (C), thin only (T),

burn only (B) and thin and burn together (X). Moisture classes are indicated in

a) by the colors associated with b-d). The grey arrows are linear predictors of

site condition (Veg, understory vegetation; Open, canopy openness) and

substrate cover (LEAF, leaf litter; MSOIL, exposed soil) within each moisture

class. The length of the arrow indicates the strength of the relationship with

the ordination, and only relationships with p > 0.05 were plotted. Note a) was

not fitted with linear predictors and c) had no significant predictors...... 79

Figure 7. Mean richness (±S.E.) of bryophytes across a 2 × 2 factorial design of

thinning and burning with control (C), thin only (T), burn only (B) and thin

and burn together (X) within xeric (X), intermediate (I), and mesic (M)

moisture classes in three southeastern forests. Significance is indicated within

moisture class as '***’ 0.001, ‘**’ 0.01, ‘*’ 0.05, ‘.’ 0.1...... 80

Figure 8. Non-metric multidimensional scaling (NMDS) ordination and species

correlates of the bryophyte community a) as a whole and b-d) divided out by 14

moisture classes in three mixed-oak forests in southeastern Ohio. Transects

are labeled by treatment within a 2 × 2 factorial of silvicultural thinning and

prescribed burning with control (C), thin only (T), burn only (B) and thin and

burn together (X). Moisture classes are indicated in a) by the colors associated

with b-d). The grey arrows are linear predictors of species abundance within

each moisture class. The length of the arrow indicates the strength of the

relationship with the ordination, and only species occurring in > 10% of the

transects having relationships with p > 0.05 were plotted. Species symbols are

the first four letters of the genus and species epithets presented in Table 1.

Note a) was not fitted with linear predictors...... 81

Figure 9. Example of differential response bryophytes by substrates and the

substrates, themselves. Mean (± S.E.) of species occurrence on coarse woody

debris (CWD) and percent of CWD cover across a 2 × 2 factorial design of

thinning and burning with control (C), thin only (T), burn only (B) and thin

and burn together (X) within xeric (X), intermediate (I), and mesic (M)

moisture classes in three southeastern forests...... 82

INTRODUCTION

Modern forestry is one of the most widespread anthropogenic disturbances in the eastern deciduous forest. Nearly one million ha of timberlands are harvested either through clear-cutting or partial-cutting each year (Smith et al. 2009). Disturbances play an important role in determining the diversity (Mackey and Currie 2001), stability (Van der Maarel 1993), and functional capacity (Bengtsson et al. 2000) of an ecosystem.

Forested ecosystems, in particular, are strongly determined in these respects by disturbances ranging from small-scale, endogenous disturbances such as individual tree fall (Runkle 1982) to large-scale, exogenous disturbances such as windstorms and wildfires (Pickett and White 1985, Oliver and Larson 1996). The nature and intensity of these disturbances dictate the nature and composition of a forested ecosystem upon successional initiation and through time (Bormann and Likens 1994). Forestry management practices often aim to simulate the natural disturbance regimes that formed and maintain these ecosystems in order to couple regenerative process with extractive uses (Attiwill 1994).

The mixed-oak forests of the unglaciated Allegheny Plateau are largely dominated by Quercus spp. and Carya spp., especially in, but not limited to, drier, less productive sites. The mixed-oak forest composition has apparently been prevalent in the region prior to European settlement (Dyer 2001). The dominance of shade-intolerant species in this ecosystem has been linked to large-scale disturbance events that create canopy openings and suppress shade-tolerant tree species including, wind throw (Runkle 1982), ice 16

damage (Rebertus et al. 1997), passenger pigeon (Ectopistes migratorius L.) roost damage (Ellsworth and McComb 2003, Buchanan and Hart 2012), and fire (Abrams

1992, Brose et al. 2001). The physical, and, ultimately demographic, changes resulting from the cumulative effects of these disturbance events is evident in historical analyses of tree ring release, damage, and fire scar events, even in old-growth forests (Shumway et al. 2001, Rentch et al. 2003, Buchanan and Hart 2012). There are strong indications that these disturbances are necessary for the development and persistence of mixed-oak forests in the eastern deciduous forest and are necessary for the maintenance of these ecosystems (Buchanan and Hart 2012).

Notable changes in the frequency and intensity of disturbance events since

European settlement have altered disturbance regimes in mixed-oak forests and subsequently altered dynamics in this ecosystem. In southeastern Ohio, the modern, second growth mixed-oak forests originated after widespread forest clearing for the iron industry in the mid-1800s (Hutchinson et al. 2003). Frequent, low to moderate-intensity fires with a return interval of 5 – 10 years also occurred during this period and until fire suppression polices instituted in the 1930s essentially eliminated fire disturbance events

(Sutherland 1997, McCarthy et al. 2001, McEwan et al. 2007). The loss of extensive flocks of passenger pigeons (Ectopistes migratorius) that may have created small-scale to stand-level canopy openings also occurred during this same period (Ellsworth and

McComb 2003). Ultimately, it has been widely noted that contemporary mixed-oak forests contain a higher abundance of shade-tolerant and disturbance sensitive species, 17

including Acer L. spp. and Liriodendron tulipifera L,. than pre European forests

(McCarthy et al. 1987, Goebel and Hix 1997, Dyer 2001). Additionally, oak-regeneration in these forests has been found generally inadequate to maintain the stand composition in the long term. Although there are a variety of interacting factors including changing climate, changing uses by native peoples, and increases in deer herbivory that likely play an important role in these changes in species composition (McEwan et al. 2011), the integral role disturbance has played in the maintenance of these ecosystems cannot be overlooked as possible management tools.

Perhaps the most easily implemented disturbance regimes to maintain the mixed- oak forest ecosystem are fire and mechanical canopy manipulation. Apparent historical relationships between fire, native peoples, and oak abundance (Abrams 1992) formed the basis for several large studies focused on utilizing prescribed fire to simulate the effects of natural fire disturbances in oak forests (Loftis and McGee 1993, Brose et al. 2006). In southeast Ohio, repeated low-intensity fires were not sufficient to restore the composition and regenerative capacity of mixed-oak forests (Hutchinson et al. 2005). After consideration of more complex and integrated natural disturbance cycles that often involved increased fuel loads following canopy disturbance events with fires in subsequent years, a shelterwood method that removed a portion of the canopy, increased light levels, stimulated shade-intolerant regeneration followed by prescribed fire(s) that decreased shade-tolerant, disturbance-sensitive tree species was developed (Brose et al.

1999, 2001). Consequently, the Fire and Fire Surrogate Study was initiated by the USDA 18

Forest Service as a regional-scale, replicated study examining oak regeneration after silvicultural thinning and periodic prescribed burning (Yaussy 2001). If effective, these methods could be widely employed on public and private forest lands; however, their effects on ecosystem function and non-timber species including herbaceous plants

(Phillips et al. 2007) and breeding birds (Streby and Miles 2010) are currently being examined.

The bryophyte community is known to be highly susceptible to environmental change and could be expected to respond directly to silvicultural disturbances (Glime

2001). Even in areas where bryophytes represent little of the biomass within a system, these species can have disproportionate impacts on moisture, temperature, and nutrient dynamics (Glime 2001). They are often used as indicators of old-growth and minimally- impacted ecosystems (Keddy and Drummond 1996). Bryophytes also provide structural complexity and suitable conditions at micro-scales that harbor considerable invertebrate diversity representing multiple additional trophic levels (Meininger et al. 1985, Suren

1991). Moreover, the bryophyte invertebrate community may harbor life stages of species that parasitize forest pests that negatively impact timber production, although this is an active area of research (G. McGee, SUNY ESF, personal comm.).

The effects of management activities associated with forestry, such as silvicultural thinning and prescribed fire, on bryophyte diversity and abundance are variable, but, in general, communities adapted to fire regimes and open canopy environments typically tend to maintain or increase diversity (e.g., Taylor 1920, Attiwill 1994), and those not 19

adapted will decrease (Vandermast et al. 2004). There are conflicting results of previous studies that found both increases or no change (Moul and Buell 1955, de Grandpré et al.

1993, McGee and Kimmerer 2002) and decreases (Duncan and Dalton 1982, Vellak and

Ingerpuu 2005, Bradbury 2006) in bryophyte diversity and/or abundance after disturbance by fire or other stand manipulation. It is apparent that community dynamics vary depending on the specific silvicultural system and ecosystem involved. However, there is a notable gap in the literature in relation to the ecological understanding of disturbance induced bryophyte dynamics, especially in relation to substrate specific effects, in the temperate deciduous forest of eastern North America (Ryoma and Laaka-

Lindberg 2005).

Many bryophyte species are highly specialized with regard to substrate selection, and the presence of suitable substrates may have a disproportionally important influence on the bryophyte community (Evans et al. 2012). Woody substrate colonization by bryophytes represents a vital ecological process in forested ecosystems, especially in temperate forests in which a large proportion of bryophyte species occur on these substrates (McGee and Kimmerer 2002). Changes in the pH, humidity, temperature, and solar radiation on bark surfaces after disturbance may also limit recolonization by bryophyte species (Duncan and Dalton 1982, Ryoma and Laaka-Lindberg 2005,

Bradbury 2006). This has important consequences in relation to forestry practices that remove or alter woody substrates during harvest and leave large amounts of rapidly degraded slash in that bryophyte substrate availability may be altered considerably and 20

may not be suitable long-term. Possible concomitant decreases in abundance and diversity may be the result (Uhl et al. 1981, Bradbury 2006). Non-woody substrates are also often exposed during disturbance. As different guilds of bryophytes often colonize these substrates, the response of the bryophyte community may be notably different among substrate types. Thus, silvicultural thinning and prescribed fire may have strong effects on woody and non-woody substrates.

Alteration of moisture levels below species tolerances due to changes in forest structure can limit photosynthetic activity to the point of exclusion (Shaw and Goffinet

2000). Bryophytes lack subterranean structures and depend on localized moisture conditions for photosynthesis. Bryophyte communities generally differ strongly along moisture gradients, with more mesic conditions favoring the development of bryophyte communities in deciduous forests (Rubino and Vis 2001, Tng et al. 2009). Forestry practices that dramatically decrease canopy cover such as clear cutting and heavy selective cutting can dramatically increase light and moisture stress on bryophyte species

(Patiño et al. 2010); however, mesic refugia within disturbed landscapes have been shown to limit bryophyte community declines (Hylander et al. 2002, Hylander and

Johnson 2010). The degree to which forestry practices unequally affect differing moisture conditions can have important implications for changes in the bryophyte community after management.

Tree-fall is commonly regarded as the dominant natural disturbance regime in the eastern deciduous forests of North America (Runkle 1982), with fire being relatively less important on a regional scale (McCarthy et al. 2001), especially in highly dissected terrain (Kline and Cottam 1979). Modern forestry practices attempt in many cases to mimic the natural (pre-European) disturbance regimes to stimulate regrowth of valuable tree species (Perera et al. 2004). In southeastern Ohio, where such natural disturbances are thought to have occurred at long intervals, common implementation of these practices may have a profound effect on non-adapted species.

This study presents an analysis of the bryophyte community response in mixed- oak forests of the unglaciated Alleghany Plateau to thinning and prescribed burning methods utilized in the Fire and Fire Surrogate Study. Forestry practices are widespread disturbances that can profoundly affect bryophyte communities and these effects may be strongly associated with moisture conditions and substrate type and availability.

Therefore, the purpose of this study was to address four central questions: (1) What is the composition and diversity of the bryophyte community in mixed-oak forests? (2) How does the diversity and composition of the bryophyte community across moisture conditions in mixed-oak forests respond to thinning and prescribed fire utilized for silvilcultural ecosystem management treatments? (3) What effect do these treatments have on site conditions and substrates within these forests? and (4) are the effects on the bryophyte community dependent on the changes or alterations in these conditions and substrates due to the treatments? 22

METHODS

Study Sites

Study sites were within the Ohio Hills Site of the United States Department of

Agriculture (USDA) Fire and Fire Surrogate (FFS) Research Program

(http:\\frames.nbii.gov/ffs), which was established in three southeastern Ohio forests: The

Raccoon Ecological Management Area (REMA), Tar Hollow State Forest (TAR), and

Zaleski State Forest (ZAL) (Figure 1). These forests are currently managed by the Ohio

Department of Natural Resources (ODNR), and are being used for research purposes under a special use permit and a Memorandum of Understanding between ODNR and the

USDA Forest Service, Northeastern Research Station. All forests are located in the unglaciated Allegheny Plateau of southeastern Ohio, which is characterized by a dissected plateau of moderate to moderately-high relief underlain by sandstones, siltstones, and shales (McNab and Avers 1994, Brockman 1998). Braun (1950) included this region in the Low Hills Belt of the Mixed Mesophytic Forest Region, which is characterized by oak-hickory forests on ridgetops and more mixed forests at lower elevations. Forests in this region are predominately maturing second-growth due to extensive clear-cutting during the 1800s for timber as well as charcoal production associated with the early iron industry in Ohio (Sutherland 1997). Forests are composed primarily of Quercus montana Willd. on ridgetops, with Q. alba L., Q. velutina Lam.,

Acer rubrum L., and Carya spp. Nutt. midslope blending with Liriodendron tulipifera L. and Aesculus flava Aiton in mesic areas. 23

The three forests exhibit a similar tree species composition and distribution and are relatively similar in topographical characteristics. REMA (39° 11' 57" N, 82° 23' 46"

W, WGS-84) (6,880 ha) and ZAL (39° 21' 16" N, 82° 22' 5" W, WGS-84) (10,860 ha) are both located within Vinton Co., Ohio, ~17 km apart (Figure 1). Elevations at these sites range from 215 – 315 m with ~60 m of local relief on moderate to moderately steep slopes. Soils are largely Typic Hapludults, Typic Dystrudepts, and Aquic-Auquultic

Hapludalfs that vary from sandy loams and silt loams on ridgetops to silt loams and channery silt loams on slopes and were formed largely in residuum of the local parent material (Soil Survey Staff 2011). TAR (39° 19' 56" N, 82° 45' 57" W, WGS-84) (6,526 ha) is ~35 km from either REMA and ZAL and is located in extreme eastern Ross Co.,

Ohio, immediately west of Vinton Co., Ohio (Figure 1). Elevation ranges from 250 – 350 m with ~90 m of local relief on moderately steep-to-steep slopes. Soils are largely Typic

Hapludults, Typic Dystrudepts, and Oxyaquic Hapludalfs that consist of silt loams and silty clay loams formed in loess over residuum and channery to very channery silt loams formed in residuum on the ridgetops with silt loams and channery to very channery silt loams formed in colluvium on slopes (Soil Survey Staff 2011). Climate varies for all forests from mean monthly temperatures of 23 °C in July to 0 °C in February with a mean annual precipitation of 107 cm (Western Regional Research Center 2009).

Study Design

In accordance with the standard design of the FFS Research Program

(Weatherspoon 2000), each forest (statistical block) contained four completely 24

randomized treatments. The four treatments were silvicultural thinning only (T), prescribed burn only (B), silvicultural thinning and prescribed burn together (X), along with an untreated control (C). Thus, there were three replicates of each of four treatments for a total of twelve experimental units in the FFS design. Experimental units were forest stands (~26 – 35 ha) or portions of larger stands all having irregular boundaries with a 10 m established buffer along the boundary of each treatment unit. Experimental units were selected such that a range of landscape-scale moisture regimes from dry ridgetops to moist valleys were represented throughout each site. Silvicultural thinning from below

(removal concentrated in the smaller size classes) was conducted in the autumn and winter of 2000 – 2001 and reduced stand basal area to approximately 13.75 m2∙ha-1

(~30% reduction in basal area) (Yaussy 2001). Skid roads were established predominately along ridgetops, but smaller feeder systems were established to access lower slopes. Residual slash material was left on the ground after bole removal.

Prescribed burns were conducted in March and April of 2001 and 2005. Burn intensities were low as fuels consisted predominately of leaf litter and small woody fuels (1-hr and

10-hr), resulting in flame heights that were generally less than 2 m (Iverson et al. 2004).

A 50 50 m grid of geo-referenced points were established in each forest with a Global

Positioning System (GPS) to provide reference for navigation, placement of study plots, and assessment of ecological change. 25

Field Methods

Sampling was conducted during the summer of 2007 and utilized the 50 50 m grid system present at the FFS study sites in order to establish 50 m transects as the basis for sampling (9 per experimental unit, 36 per forest, 108 total). The primary benefit of this design was limiting sampling bias during transect location selection. Transects were established between grid points either orthogonally or diagonally such that transects approximately paralleled slope contours in order to limit within sample variability due to changes in elevation or topography (Figure 2). Additionally, transects were selected such that they did not cross any large rock outcroppings, as these habitats contain an often unique bryophyte flora unrepresentative of the majority of the forest.

As bryophyte communities can change drastically across different moisture conditions, another fully replicated treatment factor was added to the FFS treatment design: three distinct moisture classes. The nine transects in each of the experimental units in this study were stratified equally such that three transects were established in each of three moisture classes: xeric (X), intermediate (I), and mesic (M). The segregation of classes was based on Integrated Moisture Index (IMI) values. The IMI values are GIS-derived indices of long-term moisture availability based on several landscape features (e.g., a slope-aspect shading index, water-holding capacity of the soil, landform curvature) that has been shown to predict tree species composition reasonably well (Iverson et al. 1997). Distinct moisture classes were constructed by defining a buffer between each class such that transitions between classes were unambiguous. Transect 26

locations were also limited such that no transect passed through any portion of the established 10 m buffer in each treatment unit to prevent associated edge effects (Figure

2).

The framework for sampling along each transect was based on the establishment of five 2 5 m quadrats per transect oriented with the long axis parallel to and centered on each transect (Figure 3; total area for study, 0.54 ha). All data were collected at the scale of the plot to better capture the small-scale occurrences of bryophyte species. Data were then pooled across transects to limit zero data and capture larger-scale occurrence of substrate data. To limit spatially correlative effects, all quadrats were evenly spaced 5 m apart along transects.

Substrates

Substrates were recorded among seven classes representing the available classes of substrates in mixed-oak forests: leaf litter (leaf litter cover), exposed soil (exposed soil cover), rock (rock cover), fine woody debris (FWD, diameter ≥ 1 cm and < 10 cm), coarse woody debris (CWD, diameter ≥ 10.0 cm, < 45° from the ground), stump (stump base cover, dead standing stems ≥ 45° from the ground and < diameter at breast height

[DBH] in height), and tree (tree base cover, all stems >DBH in height). These substrates reflect the dominant classes on which bryophytes occur in regional forests (e.g.,Rubino and Vis 2001) as well as divisions within these classes that would be expected to respond to silvicultural practices independently (Harmon and Sexton 1996, Graham and

McCarthy 2006). Percent cover of substrate classes were recorded in each plot by 27

mapping all substrate types onto a 1:40 scale gridded representation of each plot with decimeter plot-level divisions. Substrate cover was estimated visually as the proportion of grid cells filled by each substrate type rounded to the nearest percent. Cover values often were > 100% due to the three-dimensional structure of woody debris.

Coarse woody debris metrics were recorded based on the guidelines established in

Harmon and Sexton (1996). The diameters of the largest and smallest ends, as well as, the midpoint of all CWD in each plot were measured to the nearest 0.1 cm using tree calipers. The length of each piece was measured with a reel tape to the nearest 1 cm. If the piece extended outside of the quadrat, then the diameter and length measurements were taken at the intersection of the quadrat boundary. If a piece of CWD tapered to <

10.0 cm in diameter, then only that portion ≥ 10.0 cm diameter was measured. The remainder was considered FWD. The volume of woody debris was determined based on circular areas and the Newton formula:

V = L (Ab + 4Am + At) / 6 where V is the volume, L is the length, and Ab, Am, and At are the areas of the largest, midpoint, and smallest diameters, respectively (Harmon and Sexton 1996). Stumps were included in CWD measurements. For each stump, the upslope height and midpoint diameter were measured analogous to CWD measurement. The volume of stumps was determined based on circular areas and the Huber formula:

V = Am L 28

where V is the volume, Am is the area at the midpoint, and L is the upslope height

(Harmon and Sexton 1996). A decay class (I-V) was assigned to each piece of CWD and each stump based on the following criteria utilized in other regional CWD studies (e.g.,

McCarthy and Bailey 1994, Rubino and McCarthy 2003a): (I) bark intact, small branches present; (II) bark loose or sloughing, small branches not present, no sapwood degradation; (III) little to no bark, sapwood degradation, not punky; (IV) no bark, distinct sapwood degradation, punky; (V) no bark, loss of circular shape, portions of log incorporating into humus layer, high fragmentation. In cases where multiple classes were present on an individual piece, the most prevalent class became the class for the entire piece. Within each plot, total volume of CWD and a weighted mean decay class was calculated. Weighted mean decay class per plot was calculated as

vi d i i 1 Dw n

vi i 1 where Dw is the weighted mean decay and vi and di are the volume and decay class of the ith piece of CWD, respectively. Each piece of CWD and each stump was identified to the lowest possible taxonomic level (Record 1934, Panshin and de Zeeuw 1980, White 1980,

Hoadley 1990). Voucher specimens of CWD species were deposited in the Ohio

University Xylarium, Department of Environmental and Plant Biology. 29

Living trees and dead snags greater than 10 cm in diameter within the quadrats were measured at breast height (1.37 m) on the upslope side to the nearest 0.1 cm and identified to species. If a snag did not reach breast height, it was considered a stump.

Site Conditions

Canopy opening was measured to the nearest percent for each plot with a spherical densitometer. Openness was calculated as the mean of four measurements from plot center facing north, east, south, and west. The prevalence of understory vegetation <

2 m in height was recorded on a 1 – 10 scale with 1 indicating no understory vegetation and a 10 indicating impenetrable vegetation. Additionally, data were recorded for plot aspect, slope angle, and slope position using a delineated compass and clinometer. Slope position was determined by calculating a landform index (LI) as described by (McNab

1993) which is the mean of eight equally spaced angles to the horizon from plot center.

Thus, plots with the lowest LI would be located on ridgetops, and plots with the highest

LI would be located in valleys. LI was calculated in order to objectively assess slope position on a continuous scale.

Bryophytes

Bryophyte species presence was recorded in each quadrat and these data were pooled across transects as a measure of abundance. An attempt was made to record per plot cover for each bryophyte species, but the difficulty of assessing species identity in the field and cover values across a relatively large area made these data unreliable. For all bryophyte sampling, bryophyte specimens were identified to the lowest taxonomic level 30

possible in the field, but all specimens were also collected for microscopic identification at a later time. Generally, within each plot only one sample was collected for each species, but a sample was collected for each species in every plot in order to limit possible lumping of similar taxa across larger areas.

Collected specimens were placed in brown paper bags in the field, air dried in the lab, and stored for later identification. After proper identification, voucher specimens were placed in herbarium packets with a detailed label for deposit in the Bryophyte

Collection of the Cooperrider Herbarium, Kent State University. Nomenclature followed that of the Flora of North America (Flora of North America Editorial Committee 2007) and Tropicos (Missouri Botanical Garden 2012).

Data Analysis

Bryophyte diversity was assessed through examination of richness values of the bryophyte taxa observed, the Shannon-Weiner index (H’), and evenness index (J)

(vegan::diversity; Oksanen et al. 2012). Comparisons were made across forests to illustrate differential diversity gradients across the region of study. Within forests, richness was used as the sole measure of diversity as H’ and J were strongly associated with richness at these scales.

All data were collected at the plot-level; however, since the bryophyte data were pooled to the transect-level, environmental variables were also pooled to provide a mean estimate for each transect. These data were analyzed using ANOVA as a 2 × 2 × 3 factorial treatment of burn, thin, and moisture as fixed effects organized within a 31

randomized complete block design with subsampling. Transects were subsamples within the forest by treatment experimental units (EU). Orthogonal contrasts were calculated within the moisture treatment level comparing intermediate versus xeric and mesic (I vs.

X&M) and xeric versus mesic (X vs. M). These contrasts were developed a priori as intermediate moisture conditions were expected to share characteristics of both xeric and mesic moisture classes while the largest differences were expected between the xeric and mesic moisture classes. For these analyses, alpha was set at 0.05 for treatment main effects, but a more conservative alpha of 0.10 was set for significance of interaction terms as Type II error would be more serious in these cases (Stehman and Meredith 1995). Data were analyzed in the R statistical package (R Core Team 2012).

The Moisture treatment factor was derived from IMI values, which were calculated based on landscape-level metrics that could be interrelated with slope, landform index (LI), and aspect recorded within each plot. The latter were recorded as covariates for treatment-level analyses. However, these potential covariates likely contributed to the construction of the IMI values, so it was necessary to evaluate their relationship to the Moisture variable in this study design. CART regression (tree::tree;

Ripley 2012) of Slope, LI, cos(Aspect) [north vs. south], and sin(Aspect) [east vs. west] as predictors of Moisture found that a four node model using LI and cos(Aspect) had the lowest cost-complexity relationship after cross-validation. As would be expected, xeric moisture classes corresponded to ridgetop, south-facing positions; mesic moisture classes corresponded to valley bottom, north-facing positions; and intermediate moisture classes 32

corresponded to ridgetop, south-facing as well as valley bottom, south-facing positions.

Neither aspect relative to east-west nor percent slope increased the CART model explanatory power.

Due to the strong relationship between Moisture, LI and cos(Aspect), the latter two variables were not used as covariates in the treatment models throughout the remainder of the paper. Upon further inspection, the aspects of the plots within this study were found to be biased west due to the overall orientation of the treatment units. The imbalance in east-west aspects made incorporating sin(Aspect) into the treatment models inappropriate; however, the overall design of the study provided for balance of macro- scale moisture through the use of integrated moisture index metrics. As moisture was felt to be the main driver in these systems the underrepresentation of eastern aspects was not considered a detriment to the generality of the study. Slope was apparently indicative of additional data within the landscape and was checked for usefulness as a covariate in the

ANOVA models.

Changes in substrates were analyzed through a multivariate ANOVA / ANCOVA

(MANOVA / MANCOVA) procedure (car::Manova; Fox and Weisberg 2011) to control for simultaneous changes and robustly compare the effect of the Burn, Thin, and

Moisture treatments on potential bryophyte substrates. Rock substrates did not change significantly in abundance after the FFS treatments; therefore, the rock substrate type was not considered a dependent variable in this study. Rock substrates were viewed as an important covariate to changes in overall substrate composition and were included in 33

models where it was found to improve the variance explained. All substrates were log- transformed to conform to assumptions of normality and equality of variance.

Discussions involving substrates assume this transformation for the remainder of this paper. Leaf litter cover was found to be highly correlated with exposed soil cover (r = -

0.80, df = 36, P << 0.001) and was not analyzed within the MANOVA structure.

Comparisons of Mahalanobis distances against Χ2 distributions for outliers and multivariate normality were found acceptable for MANOVA assumptions. Pillai's trace was utilized as the MANOVA test statistic.

Subsequent to the MANOVA analysis, separate ANOVAs (car::Anova; Fox and

Weisberg 2011) were run for each of the substrate types to evaluate treatment effects on each substrate individually and in conjunction with the MANOVA results. The univariate

ANOVAs were conducted on all substrate types, excluding rock, which was used as a covariate (as described above). Normality and equality of variance assumptions were checked against model residuals and were found acceptable for ANOVA assumptions.

No covariates were assessed for the stump base cover substrate type as stumps were encountered in only 35% of all transects resulting in the sump data assuming a zero- inflated, highly non-normal distribution. ANOVA was conducted on the stump substrate type under the assumption that the robustness of the sampling design and equal sampling would still provide relatively informative, unbiased results.

A permutational multivariate analysis of variance (PERMANOVA) using distance matrices (vegan::adonis; Oksanen et al. 2012) was conducted on the community matrix in 34

order to determine treatment effects across all species. A limitation of this method is that it cannot accommodate mixed-model designs, therefore, the community data were pooled within EUs to remove the secondary sampling error term associated with subsampling multiple transects per EU. Adonis can accommodate blocking structures through selection of a stratum within which to permute the data, and, in this analysis, a stratum of

“Forest” was defined to capture the three forest blocks. Non-metric multidimensional scaling (metaMDS::vegan; Oksanen et al. 2012) was implemented on the bryophyte community matrix in order to elucidate community separation. Environmental and species occurrence variables were then correlated with ordination plots (envfit::vegan;

Oksanen et al. 2012).

Substrate species occurrences were tabulated for each transect as the number of species occurring on each substrate type within each transect. During sampling, CWD and stump occurring bryophytes were recorded together, so the occurrences on these two substrates were pooled and evaluated against a pooled CWD+stump base cover class. All substrate species occurrence data were log-transformed to conform to assumptions of normality and equality of variance. Discussions involving substrate species occurrences assume this transformation for the remainder of this paper. MANOVA and MANCOVA analyses were conducted similarly to those for substrate cover values; however, for these analyses rock cover was included as a covariate as it significantly influenced the bryophyte community response analyzed for these models. 35

RESULTS

Bryophyte Diversity

In total, 116 bryophyte species representing 71 genera and 37 families were collected (Table 1). Of these, 97 species were in the class Bryophyta (mosses) and 19 were in the class Marchantiophyta (liverworts). No members of the class

Anthocerotophyta (hornworts) were observed. Within the Bryophyta, species were closely divided between acrocarpous (upright, sporophytes terminal) and pleurocarpous

(procumbent, sporophytes lateral) species with 44 and 53 species, respectively. All species of Marchantiophyta were leafy except for one thallose species, Pallavicinia lyellii

(Hook.) Carruth. One species, Fissidens hyalinus Wils. & Hook., is listed as Endangered in Ohio.

Two genera of taxonomically confounding species were grouped for community- level analysis: Brachythecium and Cephaloziella. Specimens from both genera were often poor, sterile collections that could only be reliably labeled to the generic level. Four

Brachytheium and two Cephaloziella species were collected for which the species in each genus have similar ecological niches (woody substrates and as secondary taxa among other bryophytes, respectively) but are difficult to separate: B. campestre, B. oxycladon,

B. plumosum, B. salebrosum, C. hampeana, and C. rubella. These species were grouped into Brachythecium spp. and Cephaloziella spp., respectively. Note that two species of

Brachythecium, B. acuminatum and B. rivulare, were not included in the generic level classification because of their distinct niche separation (tree bases and seepy places, 36

respectively) and reliable identification when compared to the rest of the genus. The resulting community-level species data analyzed in this study contained 112 distinct taxa.

Occurrence of individual taxa varied greatly with the scale of observation. Of the taxa recorded, 48% occurred in all three forests, while 24% and 36% occurred in two and one forest, respectively. The majority of taxa occurred infrequently with a few exceptions. The mean number of transects per taxon was 14.4 ± 1.8, and only 39 taxa

(35%) were observed in more than 10% of transects (Figure 4). The two most common species, Platygyrium repens and Steerecleus serrulatus, were present in 99% and 69% of all transects, respectively (Table 2). As a result of the sporadic distribution of the majority of the species observed, the overall transect diversity was relatively low when compared to the total number of species. The mean diversity among transects was 14.9 ±

6.9 species (Figure 5). Pooling within transects limited the effect of extremely diverse microsites that occurred on the level of plots but not transects. Although efforts were made to limit secondary taxa within collected specimens, occasionally there were smaller species interwoven among the primary specimen. When pooled across transects, secondary taxa accounted for < 5% of all transect-species occurrences, and only four species were found solely as secondary taxa.

The three forests studied were found to have varying levels of diversity (Table 3).

The highest species richness and diversity were found at REMA (S = 95, H’ = 3.70).

ZAL and TAR had progressively lower richness, but had inversely related diversities with TAR (S = 74, H’ = 3.57) being more diverse than ZAL (S = 79, H’ = 3.55). This 37

result was due to the presence of more rare species at ZAL that decreased its evenness compared to Tar Hollow (Table 3). The rare species at ZAL were largely individual occurrences of liverworts encountered in the rocky mesic habitats which only occurred at

ZAL.

Site Condition and Substrate Changes

There were notable changes to the measured site conditions and substrates present within the forest blocks of the FFS study (Table 4). The silvicultural thinning applied in

2001 and the prescribed fires applied in 2001 and 2005 increased canopy openness and understory vegetation, although there was some variation in the responses depending on location, year, and moisture conditions (Table 5). The more intense second burning at xeric to intermediate sites at REMA (Iverson et al. 2008) increased canopy openness by

51 to 67% more than in TAR and ZAL, respectively. The canopy at REMA also remained 53 to 64% more open in thinned-only sites compared to ZAL and TAR. These later two forests had returned to openness levels comparable to control stands when viewed from breast height (1.37 m). Overall, burning strongly increased canopy openness

51% over unburned areas (P < 0.001), especially in areas with severe burning. There was some indication that the increase in openness resulting from the thinning treatment was still observable (P = 0.06) and that moist sites had become generally less open than the driest sites (P = 0.06). Thinning strongly increased vegetation cover by 36% compared to unthinned stands (P < 0.001). Xeric sites had 17% greater vegetation cover than more 38

mesic sites (P = 0.04). Tar Hollow had more vegetation cover (P = 0.04) predominantly due to stronger understory vegetation responses to thinning.

MANOVA analysis of substrate changes showed a significant relationship in changes in exposed soil cover, CWD, FWD, and tree base cover under treatments of thinning and burning (Table 6). However, intermediate moisture conditions had an overall decrease in cover relative to xeric and mesic moisture conditions. Rock cover was included as a covariate as it reduced residual variation, while Slope was found to decrease explanatory power and was not included as a covariate. Separate ANVOAs within the substrate types indicated the drivers of the overall substrate responses.

Burning significantly increased exposed soil cover 55% more than unburned stands (P = 0.02) primarily due to the consumption of leaf litter cover by low-intensity burning. Although the effect of burning on exposed soil cover could be pronounced, the input of two years of leaf fall since the 2005 burn resulted in most areas being largely recovered by litter and decreased the overall significance of the effect of burning. The effect of burning on exposed soil cover was less significant than expected as control stands in xeric moisture conditions often had exposed soil cover that approached those in burned stands. Thinning, overall, did not alter soil cover; however, when comparing xeric sites to mesic sites, there was some indication that thinning decreased exposed soil cover in xeric sites when compared to xeric sites in unthinned stands. These effects were not significant within the thin or unthinned stands (P = 0.13, P = 0.66, respectively). 39

Leaf litter cover was excluded from the overall MANOVA as it was strongly correlated with exposed soil cover and responded similarly with regard to burning.

Burning did significantly decrease leaf litter cover; however, this effect was only significant in xeric and intermediate moisture conditions (P < 0.001). Mesic moisture conditions had equal litter cover regardless of burning (P = 0.80). This effect depended on a significant covariate with percent cover rock (P = 0.02) in that transects in areas with higher rock cover near valley bottoms had correspondingly low leaf litter cover.

Thinning operations left considerable residual slash material on the ground that significantly increased CWD 48% over unthinned stands (P < 0.001). Burning also increased CWD; however, this effect depended on whether the stand had been thinned. In thinned stands, burning did not increase CWD (P = 0.84), while in unthinned stands burning did increase CWD 49% over unburned stands (P = 0.01). Xeric and mesic conditions had higher CWD than intermediate conditions (P < 0.001). The later response clearly drove the overall effect of intermediate vs. xeric and mesic moisture conditions in the MANOVA results. Looking at the individual CWD piece metrics, 19 CWD taxa were found within in the CWD substrate class (Table 7). Quercus spp. were the most abundant, followed by Acer rubrum, Carya spp., and Liriodendron tulipifera, while nine taxa were found infrequently (< 5% of all transects). One specimen was identified as a Pinus sp.,

(“hard southern yellow pine”; Panshin and de Zeeuw 1980) based on wood anatomy. This specimen was almost certainly P. virginiana as it was observed growing in the immediate vicinity of the collection. Quercus spp. can only be broadly classified into the “red oaks” 40

(Sect. Erythrobalanus) and “white oaks” (Sect. Leucobalanus) based on wood anatomy alone (Record 1934), but canopy dominants suggest that these specimens were predominately Q. velutina, Q. rubra, Q. coccinea and Q. montana, Q. alba, respectively.

All CWD treatment responses were consistent when compared to changes in CWD volume, indicating that cover estimates of CWD are highly representative of CWD volume. Burning significantly decreased individual CWD decay class (P = 0.003), regardless of thinning, and more mesic conditions had significantly higher CWD decay class (P = 0.02).

Both FWD and CWD were significantly correlated (r = 0.48, P = 0.003), and could be expected to respond similarly across the treatments and moisture conditions.

Thinning did significantly increase FWD 21% over unthinned stands (P = 0.02), but in contrast to CWD, burning increased FWD regardless of thinning (P = 0.05). Also in contrast to CWD, moisture condition did not affect FWD within the stands. ZAL was found to have a significantly greater cover of FWD than either TAR or REMA largely due to a greater increase with thinning. ZAL also had a marginally, though not significantly, higher cover of CWD.

Thinning significantly decreased tree base cover by 58% compared to unthinned areas (P < 0.001). The decrease in tree base cover due to thinning was significantly more pronounced in intermediate moisture conditions when compared to xeric and mesic moisture conditions (P = 0.04). The differential effect of moisture on tree base cover likely contributed to the significant effect of increased CWD at intermediate moisture 41

conditions and the overall MANVOA result. The inclusion of slope within the model improved explanatory power as live tree cover tended to decrease with increasing slope.

Looking at the individual tree stem metrics, 17 tree species were found within the tree substrate class (Table 8). Quercus spp. were the most abundant, followed by Acer rubrum, Liriodendron tulipifera, and Carya spp., while nine species were found infrequently (<5% of all transects). All tree base cover treatment responses were consistent when compared to changes in tree volume, indicating that cover estimates of tree base cover are highly representative of tree volume. Twenty-one snags of 9 taxa were encountered with a total volume of 6.49 ± 0.03 m3 ha-1. Quercus spp. and Acer spp. represented 38% and 36% of the total importance of all species, respectively. Quercus spp. snag importance decreased compared to Quercus spp. importance in the CWD and tree classes likely because of the Acer spp. were strongly negatively impacted by prescribed fire. Snag decay classes ranged from 1 to 3 with a mean value of 1.95 ± 0.19.

A total of 44 stumps were observed in 38 transects (35% of all transects). The distribution of stump data was significantly non-normal, and although model residuals were normal and variances were equal among groups, the patchy nature of the stump data makes conclusions beyond the major treatment effects questionable. As a result, covariates were not assessed for stump cover. Thinning significantly increased percent cover stump base cover (P = 0.014). Forty-four stumps of 9 taxa were encountered with a total volume of 4.12 ± 0.11 m3 ha-1. Quercus spp. represented 43% of the total importance of all stump species. Quercus spp. stump importance also decreased 42

compared to Quercus spp. importance in the CWD and tree classes likely because of the selective removal of more mesic species in the thinning treatments. Stump decay classes ranged from 2 to 5 with a mean value of 3.17 ± 0.12.

Rock cover was evaluated as a covariate in the overall MANOVA and for separate ANOVAs of other substrates. The choice of rock cover as a covariate was confirmed by the finding that mesic sites were 87% rockier than drier sites (P = 0.02), but thinning and burning had no effect on rock cover.

Bryophyte Community Response

Permutational MANOVA results of the bryophyte community response to silvicultural thinning, prescribed fire, and moisture conditions showed strong changes in the bryophyte community (Table 9). The bryophyte community was most strongly associated with changing moisture conditions largely in the difference between xeric and mesic moisture conditions (P < 0.001). In addition, the bryophyte community in intermediate moisture conditions was found to be significantly different than the xeric and mesic moisture conditions (P = 0.023). Burning also strongly altered the bryophyte community composition (P < 0.001); however, the strength of this effect differed between xeric and mesic moisture conditions. There was also some support for an effect of thinning on the bryophyte community (P = 0.063).

The NMDS plot of the bryophyte community showed a clear moisture gradient along Axis 1 as well as the effect of burning on the same axis (Figure 6, a). The co- location of moisture and burning along the same axis confounded interpretation of 43

community dynamics, so separate NMDS ordinations were derived for each moisture class (Figure 6, b-d).

For the xeric transects, there were marked separations among the treatments.

Burning significantly separated the bryophyte communities along Axis 1 (P < 0.001).

Thinning did have a clear effect on the bryophyte community, but only for the thin only treatment, which is clearly separated from the other treatments (Figure 8). The control transects formed a relatively distinct grouping, except for one transect with only two species that grouped out high on the burn side of the ordination. Thin and burn transects were largely associated with burn only transects. The fitted environmental variables indicated that the relationship between exposed soil cover and leaf litter cover as well as the relationship between understory vegetation and canopy openness were significant predictors in this moisture class. Canopy openness increased in conjunction with burning along Axis 1; however, the other three variables did not closely associate with a particular treatment effect. They indicate that non-woody substrates and the amount of understory vegetation are also partitioning the bryophyte community in xeric sites.

For the intermediate transects, there was much less of a clear treatment effect across transects. Burning still had marginal support along Axis 1 (P = 0.057); however, the effect of thinning was no longer evident. The relationship between exposed soil cover and leaf litter cover was again significantly related to the ordination results and also partitioned the bryophyte community not in association with any clear treatment effects. 44

For the mesic transects, there were again noticeable effects of the treatments on the bryophyte community. Both thinning and burning significantly explained partitioning in the bryophyte community (P = 0.006, P = 0.005); however, the size of the effect for these treatments was ~25% less than in the xeric sites. There were no environmental variables that were significantly related to the bryophyte community ordination in mesic sites.

Taxonomic Relationships

Mean bryophyte richness was found to decrease with burning across treatments of thinning and burning within moisture classes, although this effect was only significant within the xeric moisture class (P = 0.014) and represented a decrease in richness of 46% when compared to unburned, xeric sites. There was marginal support for a 12% decrease in richness with burning in intermediate moisture conditions in the burn only stands (P =

0.067). Bryophyte richness strongly increased 51% in mesic moisture conditions compared to xeric moisture conditions (P < 0.001). Bryophyte species frequency exhibited similar responses to richness, excepting that there was marginal support of a decrease with burning regardless of moisture class (P = 0.062).

Examination of the response of the Bryophyta and Marchantiophyta separately found a differential response between these two divisions. The Bryophyta closely followed the overall responses of all bryophytes with burning decreasing richness in the xeric moisture classes (P = 0.016) and a strong increase with increasing moisture (P <

0.001). In contrast, the Marchantiophyta decreased significantly and more dramatically 45

with burning, regardless of moisture class (P = 0.003) as well increased with moisture (P

< 0.001). Responses of the Bryophyta and Marchantiophyta frequency were also similar to the richness trends.

Within the Bryophyta, both acrocarpous and pleurocarpous species responded similarly to the overall response of all bryophytes with burning decreasing richness and frequency in the xeric moisture classes and a strong increase with increasing moisture.

Acrocarpous and pleurocarpous frequency responses were again similar to richness trends.

Treatments and Substrates

Individual species responded strongly to the treatments and substrate gradients across moisture conditions (Figure 8). In xeric moisture conditions, burn stands were only associated with increases in one species, Ditrichum pallidum. This species is a small acrocarpous, shuttle species (During 1979) often associated with disturbed sites and open soil. Indeed, D. pallidum, while associated with burning, also followed the strong exposed soil gradient present in xeric moisture conditions. Unburned stands were distinct in their bryophyte communities between control and thinned stands. Platygyrium repens, the most abundant species in this study, was correlated with unburned stands, as a whole.

Control stands were correlated with two species strongly associated with tree bases (>

90% of all occurrences), Frullania eboracensis and Dicranum montanum as well as a larger acrocarpous species associated with open soil, Polytrichastrum ohioense. Thinned stands were correlated with two species also associated with woody substrates, Entodon 46

seduxtrix and Steerecleus serrulatus. These species were more abundant under the thick understory and on the more decayed and uncharred woody debris found in thinned stands.

In intermediate moisture conditions, burn stands were again not strongly correlated with any species other than Ditrichum pallidum. The exposed soil cover gradient in the ordination correlated with three soil occurring species. Many other species were correlated with unburned stands to some degree, although they do not seem to relate to any specific treatment or substrate.

In mesic stands, there were no clear gradients associated with substrates. There were strong correlations among species with the treatments in the ordination; however, the treatments themselves were not as strongly differentiated as in more xeric conditions.

There was a clear thinning gradient among the bryophyte community with species associated with tree bases such as Dicranum montanum occurring in unthinned stands and species associated with woody debris such as Plagiomnium cuspidatum and P. ciliare occurring in thinned stands. Atrichum angustatum and Dicranella heteromalla, two small, soil dwelling, shuttle species appeared to relate to burning, and Brachythecium spp., Hypnum imponens, two large, pleurocarpous, woody species, appeared to correlate with unburned stands.

Species occurrence frequencies on particular substrates provided insight into the confounding effects of treatments and their effects on substrates across moisture classes.

Actual species occurrence changes by substrates showed differing responses to the silvilcultural thinning and prescribed fire than those found for changes in cover of the 47

substrates, themselves. A clear example of this effect was seen in the CWD data (Figure

9). For the CWD data, thinning and burning both increased CWD percent cover; however, the bryophytes occurred no more frequently in unburned stands than the availability of CWD substrate would suggest. MANOVA of the bryophyte species occurrence data found that burning significantly decreased species occurrence across all substrates (P < 0.001), thinning increased species occurrence among woody substrates, although the increase was less pronounced in intermediate moisture classes (P = 0.049), and that species occurrence increased across all substrates with increasing moisture class

(P < 0.001). These trends closely mirrored changes in overall substrate availability (Table

6), although there was a stronger effect of moisture on the bryophytes community.

Partitioning out the effect of changing substrates on the occurrence of bryophyte species by substrates through MANCOVA, eliminated the effect of thinning (P = 0.482) and found that burning, while still significant, decreased species occurrence less in mesic conditions. Additionally, the substrate types included as covariates were found to be significantly related to bryophyte substrate occurrence, except for CWD (including stump base cover). 48

DISCUSSION

Floristics

A comprehensive survey of the mosses of Ohio (Snider and Andreas 1996) noted

125 and 54 moss species in Ross (Tar Hollow) and Vinton (REMA and Zaleski) counties, respectively. Ross County had been comparatively well studied with records of nearly one third of the 385 species reported for Ohio. In this study, 69 mosses were observed in

Ross County, 25 of which were new records for the county bringing the total number of moss species reported in the literature to 150. Additionally, 33 species had unverified historical records confirmed. Vinton County had been comparatively understudied relative to physiographically similar adjacent counties (including Hocking County with

285 spp.) that were known to contain a higher number of species relative to the remainder of the state. Therefore, it was not unexpected that 60 of the 90 mosses found in Vinton

County were new records for the county bringing the total number of moss species reported in the literature to 114. Additionally, 30 species had unverified historical records confirmed.

Fissidens hyalinus was collected during this study and, as it was the only species of conservation concern recorded, it is of particular interest. The collected specimen was found on bare soil on a steep, north-facing slope above an intermittent stream under a relatively full canopy in an area with small interbeds of limestone. Risk (2002) found that

F. hyalinus almost exclusively occurs in this habitat, and may, in fact, be more common 49

than historical records suggest. A thorough survey in this habitat across southeastern

Ohio may provide data leading to this taxon being delisted in the future.

In this study, eight liverworts were found in Ross County, and all nineteen liverworts species noted in this study were collected in Vinton County. Miller (1964) published a review of the known liverwort species of Ohio in which he noted three and nineteen liverwort species in Ross and Vinton counties, respectively. Excepting taxonomic revisions, Miller found 122 liverwort species in Ohio. In Ross County, seven species collected during this study were new records for the county, and fourteen species collected were county records for Vinton County. The distribution of liverwort species in

Ohio is poorly cataloged, the most recent reference is dated, and future efforts should focus on increasing the knowledge of liverwort distributions across the state.

It is interesting to note that this study recorded 116 bryophyte species in a highly restricted set of habitat types and across a roughly one-half ha area. It was not uncommon to observe unrecorded species immediately outside of or en route to sampling locations, and typically diverse bryophyte habitats including shaded ravines, wetlands, and rocky outcrops were not included or uncommonly encountered within the sampling regime.

This study did not fully represent the total bryophyte diversity in these stands or Ross and

Vinton counties, and further bryophyte floristic endeavors will likely be rewarded with additional new species.

A Synthesis

Understanding the bryophyte community response within the Ohio Hills FFS study design requires understanding the sequence of events and resulting changes in site conditions, substrates, and species all in relation to a strong moisture gradient across the study area. It is important to note that data on the pretreatment values of these metrics in these stands are not available, and it may be possible that the bryophytes were different among the FFS treatments prior to treatment application. The similarity of these results among disconnected stands across a relatively wide geographic area indicates that a prior bias was unlikely. However, there was a clear trend in bryophyte community composition between xeric and mesic sites prior to and after the study was implemented. Intermediate sites contained community elements of both moisture conditions, and exhibited a range of responses from xeric-like, intermediate to both, and mesic-like. Moist sites were consistently more diverse and contained a greater proportion of perennial pleurocarpous mosses. This strong segregation of bryophyte community by moisture class was the reason moisture class was explicitly incorporated into the treatment model.

Thinning in 2001 initially removed stems and opened the canopy, increasing understory light conditions; additionally large inputs of CWD and FWD were added to the forest floor. Subsequent burning in 2002 removed the leaf layer and exposed the soil surface. Litter is generally unsuitable for bryophyte establishment; therefore, any reduction in litter cover through burning or input of additional substrates such as logging slash will tend to increase bryophyte abundance. However, burning removes biomass 51

from the system from both living and dead carbon pools, so losses of both woody substrates and bryophyte biomass happened in opposition to any increases or possible increases of woody substrates or bryophytes in burned stands. Burning also chemically altered woody substrates by creating carbon-rich surfaces that are lower in moisture and decay more slowly than unburned woody substrates (Schimmel and Granstrom 1996).

Burned woody substrates have been shown to be poorly suited to bryophyte establishment post-fire and over time (Bradbury 2006). However, it was notable that more mesic moisture conditions appear to have limited the effect of burning in mesic areas. Mesic stands consistently burned cooler and incompletely on north-facing slopes may explain limited fire effect in these sites (Iverson et al. 2004).

After the initial 2001-2002 thin and burn, the understory vegetation increased in the thinned stands in response to increased light conditions, and leaf cover began returning to the burned stands with annual leaf fall inputs. The increase in openness in the

2001 thinned stands allowed for the development of dense understory vegetation composed primarily of early-successional, shade intolerant species (Albrecht and

McCarthy 2006). Although the thinning treatment increased canopy openness upon implementation, the limited thinning from below allowed for relatively rapid canopy closure to control treatment levels (Iverson et al. 2008). It was originally observed that the most significant presence of bryophyte occurrence on living woody vegetation was on stems >10 cm DBH, but in stands with a dense subcanopy and many small stems, there was often a regular occurrence of bryophytes at the basal 2 cm. The dense understory that 52

formed in thinned stands quickly reduced light levels at ground level and mitigated adverse environmental condition of highly light and low humidity that can negatively impact bryophyte communities (Fenton et al. 2003).

A second burning in 2005 resulted in canopy opening, and in 2007 the open canopy was still prevalent at these sites. More intense burning in 2005 opened up the canopy more dramatically in 2005 in burned stands, allowing for the development of a relatively dense understory in 2007. This change was most notable in dry to intermediate sites. The decrease in canopy openness observed from 2001 to 2007 resulted in no relationship between 2007 openness and vegetation density in thinned stands.

Burning did not cumulatively increase CWD with thinning, likely due to burning of slash CWD material that had become good fuel five years post-thinning, in spite of increases from overstory mortality due to burning (Graham and McCarthy 2006, Iverson et al. 2008). Mesic and xeric sites had increased CWD cover relative to intermediate sites likely due to downhill movement of CWD from intermediate sites, but there are also complex relationships in southeastern Ohio between environmental variation across landscape positions interacting with changing species compositions that could be strongly influencing this trend (Rubino and McCarthy 2003b). FWD trends mirrored CWD responses in that thinning and burning increased FWD. However, burning cumulatively increased FWD cover in thinned stands.

The mechanism behind the differential response of CWD and FWD is again was likely a balance between the amount of woody debris produced by the 2001 thinning, 53

consumed by the 2002 burning, produced during the 2002-2005 lapse in burns, consumed by the 2005 burn, and produced in burned stands until 2007. The large inputs of CWD from thinning had begun to approximate the decay characteristics of the control stands after five years. Burning of both thinned and unthinned stands both charred down CWD and consumed more decayed pieces (Stephens and Mogahaddis 2005, Graham and

McCarthy 2006). Additionally, burning tended to increase CWD more gradually than thinning through inputs from fire-killed trees. CWD of lower decay classes were increasing in 2007 from the 2005 fires. Increases in CWD decay class in more mesic conditions are to be expected as decay rates are generally higher, taxa are generally less recalcitrant, and burning impacts were less pronounced in mesic conditions

The removal of trees with thinning removed a complex and important substrate for bryophytes. However, the impact on the bryophyte community was not pronounced due likely to the fact that the most basal portions of the tree have the highest prevalence of bryophytes in these forests. Although live tree cover was decreased with thinning, stump cover subsequently increased in thin stands. Therefore, there still remains a substrate derived from live tree bases even though the majority of the biomass was removed. Stumps were not found to have dramatically different bryophyte species compositions than live trees; however, relatively few were encountered so a full assessment of the impact on stem removal cannot be made. There may be some effects as stumps will be more short-lived than tree bases, but they also become more suitable 54

substrates when they begin to decay (Kimmerer 1993). Further study is needed to determine these long-term effects.

During the period of this study the response of the bryophyte community necessarily involved understanding the effects of the treatments and the effects of the treatments on substrate availability. It was found that moisture class was interacting with the treatments themselves, the effects of those treatments on the bryophyte community and substrate changes, and the changes in both of the later over time. Looking at substrate species occurrence alone it was apparent that both thinning and burning strongly altered the bryophyte community in conjunction with changes in moisture. However, if the changes in substrates from the treatments and across moisture conditions are removed, only burning strongly affected the bryophyte community.

Ultimately, burning affected the bryophyte community in two ways: 1) it reduced the richness and prevalence of bryophyte species by destroying them and 2) it decreased the suitability of any additional substrates that may have been introduced by increasing their surface carbon content (charring). Only one species was positively associated with burning. Ditrichum pallidum was reported as being uncommon at an undisturbed, regionally adjacent, forested preserve (Snider and He 1990); however, this species was prevalent in burned sites in this study. The destruction of burning did not widely occur in mesic sites, so these communities were relatively unaffected by burning.

The mechanical process of thinning did not strongly alter the richness or prevalence of the bryophyte community in general; however, thinning increased the 55

availability of decomposing woody substrates that in turn increased the prevalence of corticolous components of the bryophyte community. Interestingly, the effect of thinning was not nearly as pronounced in mesic sites as would be expected. This result is understandable after examining the substrate species occurrence data. The relatively fresh woody material added to the stands, was not yet ideal substrate for bryophyte colonization. There were still relatively few highly decayed woody substrates in any moisture class. The input of woody substrates from thinning did not necessarily increase richness or frequency in mesic stands as high richness and frequency of bryophytes was already present. The changes brought about by thinning are that bryophytes are slowly shifting toward the woody substrates as they become suitable. The effect of thinning on the bryophyte community was not fully realized at the time of sampling. Subsequent decomposition of the increased woody substrates found in thinned stands will increase habitat for a variety of corticolous species over time.

Management Implications

Forestry management plans often incorporate the bryophyte community response as a component of the ecosystem response to silvilcultural activities (e.g., Swanson and

Franklin 1992, Vanderpoorten et al. 2001). Activities that can accomplish harvest objectives and maintain the bryophyte community can help to preserve the ecological functions related to nutrient cycling, moisture dynamics, trophic diversity, and habitat quality bryophytes provide. Bryophytes as a guild are generally limited by large-scale disturbances (Grime et al. 1990). Within the guild, there are differing strategies such that 56

many taxa actually benefit from some disturbances. In fact, small-scale disturbances have been shown to increase diversity in bryophyte-dominated systems (Kimmerer 1993) and may be the reason for their persistence. However, it should be noted that forest bryophytes generally cannot survive intensive physical damage or alteration of habitat

(Shaw and Goffinet 2000). Therefore, it seems a logical conclusion that areas that are often precluded from the effects of destructive disturbance will tend to contain the highest abundance and diversity of bryophytes in a system.

In this study, it was found that mesic valleys were difficult to burn and contained the highest abundance and diversity of bryophytes that was not lost in disturbed stands.

Species of Marchantiophyta were generally uncommon outside of mesic sites, except for two epixylic species. In contrast, upland, xeric sites initially had low abundance and diversity of their bryophyte communities, but most notably these values were strongly decreased by the disturbances induced by fire. Consequently, either upland sites represent areas where regular fire disturbance has kept bryophyte communities at bay or areas where bryophyte communities are being affected by an exogenous disturbance event for which they are not adapted (Moul 1952).

In this study, exposed rock outcroppings were avoided to limit sample variability, but in those sites where large rock outcroppings were sampled and otherwise observed, they often exhibited a unique and abundant bryophyte flora. Often these communities were dominated by larger acrocarpous species, and burning seems to significantly decrease the occurrence of larger, more habitat dependent species especially in upland 57

sites. These species were significantly negatively affected by burning in xeric sites within this study. There may, in fact, be a similar strongly negative effect on bryophyte communities occurring on rock outcroppings when compared to the remainder of the forest floor, but this has yet to be examined within the literature.

It has been observed in this region that south-facing, more xeric sites, tend to have a lower bryophyte diversity than north-facing, more mesic sites, but soil scarification associated with walking trails tends to create more available habitat for colonization

(Rubino and Vis 2001). Logging roads were frequently encountered during sampling, and, in their centers, were one often one of the least diverse sites for bryophytes due to the establishment of a thick turf of grasses. However, several ruderal species including

Pogonatum pensilvanicum, were strongly associated with the scarified margins of the logging roads.

In the eastern deciduous forests, the bryophyte community is ubiquitous at the scale of forestry practices. Bryophytes are found predominately on bare rock, bare soil, fallen logs and branches, and live or dead tree bases. The presence of leaf litter covers suitable substrates and prevents bryophyte establishment. Prescribed fire removes leaf litter and may increase available epigeous substrates; however, these substrates will tend to be short-lived without repeated burning. Thinning removes live tree substrate, but also introduces abundant woody substrates for colonization.

If silvilcultural methods are necessary to augment the regeneration of Quercus spp. in the mixed oak forests of southeastern Ohio, thinning from below and prescribed 58

fire have strongly differing effects on the bryophyte community. Burning clearly negatively affects the bryophyte community by reducing diversity and the suitability of available substrates. It does seem apparent that the effects of burning can be mitigated in moist sites as long as moist sights are indeed burned less severely than more xeric sites.

Burning will also likely have long-lasting legacies with regard to the bryophyte community due to chemical changes in woody debris substrates (Bradbury 2006).

Thinning from below does not appear to negatively affect the bryophyte community, and as woody substrate inputs decay, thinning may actually benefit the bryophyte community over the period of their persistence (Lõhmus and Lõhmus 2008).

The management recommendations from this study in order to promote the bryophyte community in mixed-oak forest of southeastern Ohio are 1) Use thinning only when possible as this treatment had little negative effect on the bryophyte community and may, in fact benefit the bryophyte community, 2) If burning is necessary, restrict burning to xeric and intermediate sites to limit the strongly negative effects of burning where the largest diversity and abundance of bryophytes are found 3) Avoid burning of exposed bedrock areas as they often contain a high abundance of highly burn sensitive species and can occur in more xeric conditions. Mesic and other types of refugia are common practices to conserve bryophytes in burned stands (Hylander and Johnson 2010) and would similarly benefit the bryophyte community in these forests. Lastly, it is important to note that the bryophyte community does not respond in an identical manner to the 59

herbaceous community (Phillips et al. 2007). Bryophyte and herbaceous species cannot be viewed as similar entities or managed similarly and promote both their communities.

Conclusions

Substrates were found to strongly determine bryophyte dynamics. The modification of wood and non woody substrates influenced the prevalence of the guilds of bryophytes particular to them. Burning negatively impacted bryophyte species by decreasing their abundance and diversity, predominately in xeric conditions. Burning decreased the quality of woody substrates by charring their surfaces and removing bryophytes species that may have been established. Thinning increased woody substrate availability, and while the bryophyte was not found to respond dramatically to thinning, it is expected that these substrates will slowly become suitable for establishment and serve to increase the abundance of corticolous species in the long-term. 60

TABLES

Table 1. Bryophyte species, general growth form (L, Leafy; T, Thalloid; A, Acrocarpous; P, Pleurocarpous), coefficient of conservatism (C of C; higher is more habitat specific) (Andreas et al. 2004), forest of occurrence (R, Raccoon Ecological Management Area; T, Tar Hollow; Z, Zaleski;*,county record;`,verified historical record), and accession number from the Cooperrider Herbarium, Kent State University collected in three mix-oak forests of southeastern Ohio. Species Family Forma C of Cb Forestsc Accessiond Marchantiophyta Calypogeia fissa subsp. neogaea R. M. Schust. Calypogeiaceae L NA R* T* 11223 Calypogeia muelleriana (Schiffn.) K. Müller Calypogeiaceae L NA R* T* Z* 11226 Cephaloziella cf. hampeana (Nees) Schiffn. ex Loeske Cephaloziellaceae L NA R* 11222 Cephaloziella cf. rubella (Nees) Warnst. Cephaloziellaceae L NA R* Z* 11219, 11220 Chiloscyphus polyanthos (L.) Corda Geocalycaceae L NA R* 11229 Cololejeunea biddlecomiae (Austin ex Pearson) A. Evans Lejeuneaceae L NA R T* Z 9772 Frullania eboracensis Gottsche Jubulaceae L NA R* T* Z* 10648 Jamesoniella autumnalis (DC.) Steph. Jungermanniaceae L NA Z 11234 Jungermannia crenuliformis Austin Jungermanniaceae L NA Z* 11235 Jungermannia leiantha Grolle Jungermanniaceae L NA R* 11233 Jungermannia pumila With. Jungermanniaceae L NA Z* 11236 Lophocolea bidentata (L.) Dumort. Geocalycaceae L NA R* T* 11228 Lophocolea heterophylla (Schrad.) Dumort. Geocalycaceae L NA R* T Z* 11224, 11225, 11227 Nowellia curvifolia (Dicks.) Mitt. Cephaloziaceae L NA R T* Z 10616 Odontoschisma prostratum (Sw.) Trevis. Cephaloziaceae L NA R 11232 Pallavicinia lyellii (Hook.) Gray Pallaviciniaceae T NA R* Z* 10643 Porella platyphylloidea (Schwein.) Lindb. Porellaceae L NA R* T* 10658 Scapania nemorea (L.) Grolle Scapaniaceae L NA Z 10596, 11237 Scapania undulata (L.) Dumort. Scapaniaceae L NA Z* 10593 Bryophyta Amblystegium varium (Hedw.) Lindb. Amblystegiaceae P 2 R* T Z* 10664 Anacamptodon splachnoides (Froel. ex Brid.) Brid. Fabroniaceae P 5 R* 10693 Anomodon attenuatus (Hedw.) Huebener Thuidiaceae P 3 R* T* Z* 10688 Anomodon rostratus (Hedw.) Schimp. Thuidiaceae P 4 R` T` Z` 10663 Atrichum altecristatum (Renauld & Cardot) Smyth & L. C. R. Smyth Polytrichaceae A 3 R* T* Z* 10652 61

Table 1 (continued) Atrichum angustatum (Brid.) Bruch & Schimp. Polytrichaceae A 2 R` T* Z` 10644 Atrichum crispulum Schimp. ex Besch. Polytrichaceae A 5 R* T* Z* 10629 Atrichum tenellum (Röhl.) Bruch & Schimp. Polytrichaceae A 3 R* T* 10599 Aulacomnium heterostichum (Hedw.) Bruch & Schimp. Aulacomniaceae A 5 R` T` Z` 10669 Bartramia pomiformis Hedw. Bartramiaceae A 5 R` Z` 10603 Brachythecium acuminatum (Hedw.) Austin Brachytheciaceae P 4 R* T* Z* 10646 Brachythecium campestre (Müll. Hal.) Schimp. Brachytheciaceae P 5 R* T` Z* 10608 Brachythecium oxycladon (Brid.) A. Jaeger Brachytheciaceae P 2 R* T* Z* 10686 Brachythecium plumosum (Hedw.) Schimp. Brachytheciaceae P 5 T* 10613 Brachythecium rivulare Schimp. Brachytheciaceae P 5 R* 10620, 10635, 10649, 10990, 10992 Brachythecium salebrosum (Hoffm. ex F. Weber & D. Mohr) Schimp. Brachytheciaceae P 2 R* T Z* 10993, 10994 Brotherella recurvans (Michx.) M. Fleisch. Sematophyllaceae P 5 R* T` Z* 10679 Bryhnia novae-angliae (Sull. & Lesq.) Grout Brachytheciaceae P 3 Z* 10983, 10984, 10995 Bryoandersonia illecebra (Hedw.) H. Rob. Brachytheciaceae P 3 R* T* Z* 10985, 10987, 10988, 10991 Bryum caespiticium Hedw. Bryaceae A 1 R* T* 10592 Bryum capillare Hedw. var. capillare Bryaceae A 3 R` T` Z` 10659 Bryum lisae var. cuspidatum (Bruch & Schimp.) Margad. Bryaceae A 1 R* T` Z* 11230, 11231 Callicladium haldanianum (Grev.) H. A. Crum Hypnaceae P 4 R* T* Z* 10600 Campylium chrysophyllum (Brid.) Lange Amblystegiaceae P 3 R* T` Z* 10637 Campylium hispidulum (Brid.) Mitt. Amblystegiaceae P 3 R* T` Z* 10660 Ceratodon purpureus (Hedw.) Brid. subsp. purpureus Ditrichaceae A 1 R* T Z* 10647 Ctenidium malacodes Mitt. Hypnaceae P 4 R` Z` 10612 Cyrto-hypnum minutulum (Hedw.) W. R. Buck & H. A. Crum Thuidiaceae P 5 R* 10670 Cyrto-hypnum pygmaeum (Schimp.) W. R. Buck & H. A. Crum Thuidiaceae P 6 T` 10653 Dicranella heteromalla (Hedw.) Schimp. Dicranaceae A 2 R` T` Z` 10619, 10632, 10675 Dicranum flagellare Hedw. Dicranaceae A 3 R* T` Z* 10665 Dicranum fulvum Hook. Dicranaceae A 7 R` 10650 Dicranum montanum Hedw. Dicranaceae A 3 R` T* Z` 10668 Dicranum scoparium Hedw. Dicranaceae A 3 R` T` Z` 10639 Dicranum viride (Sull. & Lesq.) Lindb. Dicranaceae A 4 R* T* Z* 10606 62

Table 1 (continued) Diphyscium foliosum (Hedw.) D. Mohr Diphysciaceae A 7 R* Z* 10590 Ditrichum pallidum (Hedw.) Hampe Ditrichaceae A 2 R` T* Z` 10609 Ditrichum rhynchostegium Kindb. Ditrichaceae A 2 T* Z* 10607 Entodon brevisetus (Hook. & Wilson) Lindb. Entodontaceae P 7 R* 10677 Entodon cladorrhizans (Hedw.) Müll. Hal. Entodontaceae P 5 R* T` 10667 Entodon seductrix (Hedw.) Müll. Hal. Entodontaceae P 2 R` T Z` 10605 Eurhynchium hians (Hedw.) Sande Lac. Brachytheciaceae P 3 R* T` Z* 10601 Eurhynchium pulchellum (Hedw.) Jenn. var. pulchellum Brachytheciaceae P 3 T` Z* 10989, 10996 Fissidens bryoides Hedw. Fissidentaceae A 6 R* T` Z* 10611 Fissidens bushii (Cardot & Thér.) Cardot & Thér. Fissidentaceae A 5 R* T* Z* 10624, 10625 Fissidens dubius P. Beauv. Fissidentaceae A 4 R* T Z* 9852 Fissidens exilis Hedw. Fissidentaceae A 4 Z* 10638 Fissidens hyalinus Wilson & Hook.f. Fissidentaceae A 9 Z* 10651 Fissidens subbasilaris Hedw. Fissidentaceae A 5 R* T* Z* 10641 Fissidens taxifolius Hedw. Fissidentaceae A 3 R* T Z* 10623, 10628 Forsstroemia trichomitria (Hedw.) Lindb. Leucodontaceae P 5 R` 10694 Funaria hygrometrica Hedw. Funariaceae A 1 R* 10617 Haplocladium microphyllum (Hedw.) Broth. Leskeaceae P 3 R* Z* 10666 Haplohymenium triste (Ces.) Kindb. Thuidiaceae P 6 R* T` Z* 10674 Homomallium adnatum (Hedw.) Broth. Hypnaceae P 3 T` 10604 Hookeria acutifolia Hook. & Grev. Hookeriaceae P 10 Z* 10690 Hygroamblystegium tenax (Hedw.) Jenn. var. tenax Amblystegiaceae P 2 Z` 10662 Hypnum curvifolium Hedw. Hypnaceae P 5 R* T` Z* 10610 Hypnum imponens Hedw. Hypnaceae P 5 R` T` Z` 10642 Hypnum lindbergii Mitt. Hypnaceae P 6 Z* 10640 Hypnum pallescens (Hedw.) P. Beauv. Hypnaceae P 3 R* T* 10636 Isopterygium tenerum (Sw.) Mitt. Hypnaceae P 5 R* T* 10630 Leskea gracilescens Hedw. Leskeaceae P 3 R* T Z* 10621 Leucobryum albidum (Brid. ex P. Beauv.) Lindb. Leucobryaceae A 3 R* T* Z* 10695 Leucobryum glaucum (Hedw.) Ångstr. Leucobryaceae A 4 R` T` Z` 10661 Leucodon julaceus (Hedw.) Sull. Leucodontaceae P 5 R` 10672 63

Table 1 (continued) Lindbergia brachyptera (Mitt.) Kindb. Leskeaceae P 3 R* 10633, 10671 Orthotrichum ohioense Sull. & Lesq. Orthotrichaceae A 4 R` T` 10676 Orthotrichum stellatum Brid. Orthotrichaceae A 4 T` 10680 Physcomitrium pyriforme (Hedw.) Hampe Funariaceae A 1 R* T* Z* 10591, 10602 Plagiomnium ciliare (Müll. Hal.) T. J. Kop. Mniaceae A 5 R` T` Z` 10655 Plagiomnium cuspidatum (Hedw.) T. J. Kop. Mniaceae A 2 R` T Z` 10657 Plagiomnium ellipticum (Brid.) T. J. Kop. Mniaceae A 7 R* Z* 10627, 10685 Plagiothecium cavifolium (Brid.) Z. Iwats. Plagiotheciaceae P 5 T* 10597, 10614 Plagiothecium denticulatum (Hedw) Schimp. Plagiotheciaceae P 4 R* 10634 Plagiothecium laetum Schimp. Plagiotheciaceae P 4 T* Z* 10692 Platygyrium repens (Brid.) Schimp. Hypnaceae P 3 R* T Z* 10598 Pleurozium schreberi (Willd. ex Brid.) Mitt. Hylocomiaceae P 3 R* 9851 Pogonatum pensilvanicum (Bartram ex Hedw.) P. Beauv. Polytrichaceae A 4 T` 10595 Pohlia annotina (Hedw.) Lindb. Bryaceae A 2 Z* 11221 Pohlia nutans (Hedw.) Lindb. Bryaceae A 1 Z` 10645 Polytrichastrum ohioense (Renauld & Cardot) G. L. Sm. Polytrichaceae A 2 R` T` Z` 10691 Pseudotaxiphyllum elegans (Brid.) Z. Iwats. Plagiotheciaceae P 6 R* Z* 10622, 10626 Pylaisiadelpha tenuirostris (Bruch & Schimp. ex Sull.) W. R. Buck Sematophyllaceae P 4 R* T Z* 10687 Pylaisiella selwynii (Kindb.) Crum et al. Hypnaceae P 5 R* 10594 Rhizomnium punctatum (Hedw.) T. J. Kop. var. punctatum Mniaceae A 5 R* Z* 10631 Sematophyllum adnatum (Michx.) E. Britton Sematophyllaceae P 4 R* T* Z* 10678 Sematophyllum demissum (Wilson) Mitt. Sematophyllaceae P 6 R` T* Z` 10684 Steerecleus serrulatus (Hedw.) H. Rob. Brachytheciaceae P 3 R` T` Z` 9850 Taxiphyllum deplanatum (Bruch & Schimp. ex Sull.) M. Fleisch. Hypnaceae P 6 R* T` 10654 Tetraphis pellucida Hedw. Tetraphidaceae A 4 R` 10673, 10986 Thelia asprella (Schimp.) Sull. & Lesq. Theliaceae P 5 R` T` Z` 10689 Thelia hirtella (Hedw.) Sull. & Lesq. Theliaceae P 5 R* T` 10615 Thuidium delicatulum (Hedw.) Schimp. Thuidiaceae P 3 R` T Z` 10656 Tortella humilis (Hedw.) Jenn. Pottiaceae A 3 R` T` Z` 10682 Ulota crispa (Hedw.) Brid. Orthotrichaceae A 5 R` T` 10683 Weissia controversa Hedw. Pottiaceae A 1 R` T` Z` 10681 64

Table 2. Most commonly encountered (> 10% of all transects, n = 108) bryophytes in three mixed-oak forests in southeastern Ohio. Taxon Transects (%) Platygyrium repens 99.1 Steerecleus serrulatus 69.4 Dicranum montanum 58.3 Plagiomnium cuspidatum 54.6 Lophocolea heterophylla 52.8 Dicranella heteromalla 50.9 Atrichum angustatum 50.0 Thuidium delicatulum 46.3 Atrichum altecristatum 43.5 Ditrichum pallidum 43.5 Brachythecium spp. 42.6 Frullania eboracensis 42.6 Polytrichastrum ohioense 41.7 Entodon seductrix 40.7 Amblystegium varium 39.8 Pylaisiadelpha tenuirostris 34.3 Hypnum imponens 30.6 Sematophyllum demissum 30.6 Leucobryum glaucum 29.6 Anomodon attenuatus 27.8 Hypnum curvifolium 27.8 Dicranum flagellare 25.9 Tortella humilis 25.9 Calypogeia muelleriana 23.1 Anomodon rostratus 21.3 Campylium hispidulum 19.4 Dicranum scoparium 19.4 Nowellia curvifolia 17.6 Eurhynchium hians 15.7 Fissidens bushii 15.7 Campylium chrysophyllum 13.9 Fissidens bryoides 13.9 Leucobryum albidum 13.0 Leskea gracilescens 12.0 Plagiomnium ciliare 12.0 Sematophyllum adnatum 12.0 Aulacomnium heterostichum 11.1 Bryum lisae var. cuspidatum 11.1 Cololejeunea biddlecomiae 10.2 65

Table 3. Forest diversity metrics: number of species (S), Shannon-Weiner diversity (H’), and evenness (J) for the bryophyte community occurring in three mixed-oak forests in southeastern Ohio. Forest S H’ J REMA 91 3.66 0.81 Tar Hollow 74 3.57 0.83 Zaleski 79 3.55 0.81

Table 4. Mean (± S.E.) of site condition variables and substrate cover values within 2 × 5 m plots established in xeric, intermediate, and mesic moisture classes under a 2 × 2 factorial of silvilcultural thinning and prescribed fire in three mixed-oak forest in southeastern Ohio. Cover of substrate types represent the 2-D coverage within the plot as visually mapped onto a gridded representation of the plot. Cover types are: exposed soil (SOIL), leaf litter (LEAF), coarse woody debris (CWD, diameter ≥ 10.0 cm, < 45° from the ground), fine woody debris (FWD, diameter ≥ 1 cm and < 10 cm), tree base cover (TREE, all stems >DBH in height), stump base (STUMP, dead standing stems ≥ 45° from the ground and < diameter at breast height [DBH] in height), and rock (ROCK). Canopy Vegetation Cover of substrate types (%) Moisture Treatment Slope (%) openness (%) cover (1-10) SOIL LEAF CWD FWD TREE STUMP ROCK Control 17.6 ± 0.24 6.4 ± 0.22 3.1 ± 0.01 6.7 ± 0.75 87.3 ± 0.86 1.7 ± 0.02 2.7 ± 0.04 1.5 ± 0.03 0.02 ± 0.00 0.9 ± 0.12 Thin 16.4 ± 0.30 9.3 ± 0.69 4.1 ± 0.08 1.3 ± 0.05 89.5 ± 0.28 4.8 ± 0.18 3.4 ± 0.14 1.0 ± 0.06 0.22 ± 0.03 0.8 ± 0.13 Xeric Thin×Burn 13.6 ± 0.24 22.0 ± 0.95 4.7 ± 0.14 8.0 ± 0.51 84.1 ± 0.44 5.1 ± 0.24 3.9 ± 0.21 0.7 ± 0.03 0.44 ± 0.03 0.1 ± 0.00

Burn 13.5 ± 0.69 21.4 ± 2.03 3.2 ± 0.13 10.4 ± 0.71 82.0 ± 0.80 3.8 ± 0.10 3.5 ± 0.09 1.5 ± 0.13 0.14 ± 0.01 0.1 ± 0.01

Control 18.6 ± 0.14 6.3 ± 0.07 2.0 ± 0.02 3.6 ± 0.36 91.6 ± 0.31 0.6 ± 0.05 2.0 ± 0.06 2.4 ± 0.14 0.00 ± 0.00 0.5 ± 0.08 Thin 17.2 ± 0.38 11.5 ± 0.47 4.2 ± 0.05 3.1 ± 0.07 91.1 ± 0.02 2.8 ± 0.01 3.6 ± 0.09 0.6 ± 0.02 0.50 ± 0.03 0.1 ± 0.00 Intermediate Thin×Burn 20.1 ± 0.31 14.4 ± 0.52 3.4 ± 0.10 5.3 ± 0.50 86.9 ± 0.72 4.0 ± 0.16 4.7 ± 0.12 0.4 ± 0.04 0.43 ± 0.04 0.8 ± 0.01

Burn 17.7 ± 0.22 14.8 ± 1.00 2.6 ± 0.04 15.1 ± 1.47 78.4 ± 1.37 2.2 ± 0.14 3.8 ± 0.15 1.4 ± 0.03 0.23 ± 0.02 0.5 ± 0.09

Control 17.4 ± 0.27 5.0 ± 0.10 2.0 ± 0.05 2.3 ± 0.04 86.0 ± 0.76 2.2 ± 0.09 2.5 ± 0.10 1.3 ± 0.05 0.16 ± 0.03 6.6 ± 0.87 Thin 18.3 ± 0.43 7.9 ± 0.40 3.9 ± 0.16 4.6 ± 0.18 84.7 ± 0.42 6.3 ± 0.35 3.5 ± 0.08 0.7 ± 0.07 0.14 ± 0.02 2.1 ± 0.30 Mesic Thin×Burn 22.5 ± 0.57 12.1 ± 0.48 3.8 ± 0.26 6.6 ± 0.75 86.1 ± 1.04 4.4 ± 0.21 2.9 ± 0.06 0.5 ± 0.03 0.19 ± 0.02 1.9 ± 0.28 Burn 20.9 ± 0.48 10.0 ± 0.62 2.8 ± 0.01 3.8 ± 0.28 86.4 ± 0.61 3.3 ± 0.18 2.7 ± 0.05 1.1 ± 0.02 0.01 ± 0.00 4.0 ± 0.65

Table 5. ANOVA model results (F-value, P-value) for environmental and substrate responses to silvilcultural thinning and prescribed fire in mixed-oak forests of southeastern Ohio. ANOVA models were formulated as Forest + Burn × Thin × Moisture and evaluated for additive effects of cover of ROCK or Slope as covariates. Burn and Thin are 1 df factors whether yes or no, and Moisture was split into two 1 df orthogonal contrasts comparing intermediate vs. the xeric and mesic moisture classes (IvXM) and the xeric vs. mesic moisture classes (XvM). MANOVA was conducted on the subset of variables indicated below the MANOVA results and separate ANOVAs were calculated for each variable within the MANOVA and the remainder in the table. Only significant (P < 0.05 for main effects, P < 0.10 for interactions) and marginally significant (P < 0.10, for main effects only) results are shown. Cover types are: exposed soil (SOIL), leaf litter (LEAF), coarse woody debris (CWD, diameter ≥ 10.0 cm, < 45° from the ground), fine woody debris (FWD, diameter ≥ 1 cm and < 10 cm), tree base cover (TREE, all stems >DBH in height), stump base (STUMP, dead standing stems ≥ 45° from the ground and < diameter at breast height [DBH] in height), and rock (ROCK).

Moisture BurnxMoi sture ThinxMoisture Forest Burn Thin IvXM XvM BurnxThin IvXM XvM IvXM XvM ROCK Slope Open 7.23, 0.004 19.02, <0.001 3.81, 0.06 3.8, 0.06 Veg 3.44, 0.05 20.61, <0.001 4.04, 0.06 MANVOA 3.33, 0.03 8.93, <0.001 3.99, 0.02 SOIL 6.32, 0.02 3.23, 0.09 5.21, 0.03 CWD 4.47, 0.05 20.32, <0.001 16.03, <0.001 3.33, 0.08 FWD 3.16, 0.06 4.23, 0.05 5.88, 0.02 TREE 18.99, <0.001 4.82, 0.04 3.71, 0.07 LEAF 5.04, 0.04 3.02, 0.097 3.69, 0.07 16.96, 0.001 STUMP 7.123, 0.014 ROCK 6.59, 0.02

Table 6. MANOVA analysis of forest substrate cover change after burning and thinning across xeric (X), intermediate (I), and mesic (M) moisture types as determined by an integrated moisture index (IMI) in three southeastern Ohio forests. Substrates included in this analysis were exposed soil (SOIL), coarse woody debris (CWD), fine woody debris (FWD), and tree base cover (TREE). Cover of rock (ROCK) and slope were included as covariates, and orthogonal 1 df contrasts for moisture are represented. Pillai's trace approximate Numerator Denominator Error: EU df F df df Pa Forest 2 0.94 8 36 0.497 Burn 1 3.91 4 17 0.020 * Thin 1 12.58 4 17 <<0.001 *** Moisture 2 1.81 8 36 0.106 I vs. X&M 1 3.10 4 19 0.040 * X vs. M 1 0.43 4 19 0.788 Rock 1 0.89 4 17 0.494 Slope 1 1.78 4 17 0.180 Burn:Thin 1 0.84 4 17 0.517 Burn:Moisture 2 1.24 8 36 0.307 I vs. X&M 1 0.82 4 19 0.531 X vs. M 1 0.79 4 19 0.548 Thin:Moisture 2 1.56 8 36 0.170 I vs. X&M 1 1.71 4 19 0.190 X vs. M 1 0.46 4 19 0.765 Burn:Thin:Moisture 2 0.35 8 36 0.941 I vs. X&M 1 0.38 4 19 0.823 X vs. M 1 0.14 4 19 0.963 Residuals 20 Total 35 Error: Transect Rock 1 0.98 4 67 0.424 Slope 1 0.93 4 67 0.453 Residuals 70 Total 72 a '***’ 0.001, ‘**’ 0.01, ‘*’ 0.05, ‘.’ 0.1 69

Table 7. Mean (± S.E.) density (pieces ha-1), volume (m3 ha-1), relative density (RDEN), relative volume (RVOL), and relative importance (RIV) of coarse woody debris (CWD) taxa in three mixed-oak forests in southeastern Ohio. Species Symbol Density Volume RDEN RVOL RIV Acer rubrum L. ACRU 151.85 ± 29.96 3.95 ± 0.84 14.34 ± 2.83 9.00 ± 0.08 11.67 ± 1.45 Acer saccharum Marshall ACSA 29.63 ± 14.08 1.25 ± 0.67 2.80 ± 1.33 2.84 ± 0.06 2.82 ± 0.70 Castanea dentata (Marshall) Borkh. CADE 1.85 ± 1.95 0.01 ± 0.02 0.17 ± 0.18 0.03 ± 0.00 0.10 ± 0.09 Carya spp. Nutt. CARYA 66.67 ± 20.09 4.13 ± 2.65 6.29 ± 1.90 9.43 ± 0.25 7.86 ± 1.07 Cornus florida L. COFL 7.41 ± 3.85 0.16 ± 0.10 0.70 ± 0.36 0.36 ± 0.01 0.53 ± 0.19 Fagus grandifolia Ehrh. FAGR 3.70 ± 2.75 0.13 ± 0.11 0.35 ± 0.26 0.29 ± 0.01 0.32 ± 0.14 Fraxinum americana L. FRAM 35.19 ± 20.72 1.17 ± 0.64 3.32 ± 1.96 2.67 ± 0.06 3.00 ± 1.01 Juglans nigra L. JUNI 1.85 ± 1.95 0.06 ± 0.06 0.17 ± 0.18 0.14 ± 0.01 0.16 ± 0.10 Liquidambar styraciflua L. LIST 1.85 ± 1.95 0.04 ± 0.05 0.17 ± 0.18 0.10 ± 0.00 0.14 ± 0.09 Liriodendron tulipifera L. LITU 59.26 ± 16.83 1.63 ± 0.53 5.59 ± 1.59 3.72 ± 0.05 4.66 ± 0.82 Nyssa sylvatica Marshall NYSY 16.67 ± 8.82 0.36 ± 0.23 1.57 ± 0.83 0.82 ± 0.02 1.20 ± 0.43 Hard pine PINHAR 1.85 ± 1.95 0.22 ± 0.23 0.17 ± 0.18 0.49 ± 0.02 0.33 ± 0.10 Populus grandidentata Michx. POGR 3.70 ± 2.75 0.09 ± 0.07 0.35 ± 0.26 0.21 ± 0.01 0.28 ± 0.13 Prunus serotina Ehrh. PRSE 1.85 ± 1.95 0.24 ± 0.25 0.17 ± 0.18 0.55 ± 0.02 0.36 ± 0.10 Quercus sect. Leucobalanus Engelm. QUAL 322.22 ± 33.44 11.78 ± 1.56 30.42 ± 3.16 26.86 ± 0.15 28.64 ± 1.65 Quercus sect. Erythrobalanus Spach QURU 279.63 ± 39.48 16.59 ± 2.97 26.40 ± 3.73 37.82 ± 0.28 32.11 ± 2.00 Robinia pseudoacacia L. ROPS 46.30 ± 17.19 1.24 ± 0.49 4.37 ± 1.62 2.83 ± 0.05 3.60 ± 0.83 Sassafras albidum (Nutt.) Nees SAAL 24.07 ± 9.06 0.53 ± 0.24 2.27 ± 0.85 1.22 ± 0.02 1.75 ± 0.44 Ulmus rubra Muhl. ULRU 3.70 ± 3.91 0.27 ± 0.28 0.35 ± 0.37 0.61 ± 0.03 0.48 ± 0.20

Quercus spp. L. 601.85 ± 51.67 28.37 ± 3.21 56.82 ± 4.88 64.68 ± 0.30 60.75 ± 2.59 Non Quercus spp. 457.41 ± 62.36 15.49 ± 3.22 43.18 ± 5.89 35.32 ± 0.30 39.25 ± 3.10

Total 1059.26 ± 79.95 43.85 ± 4.5 70

Table 8. Mean (± S.E.) density (stems ha-1), volume (m3 ha-1), relative density (RDEN), relative volume (RVOL), and relative importance (RIV) of tree species in three mixed-oak forests in southeastern Ohio. Species Symbol Density Volume RDEN RVOL RIV Acer rubrum L. ACRU 50.00 ± 12.45 4.95 ± 2.00 17.88 ± 4.45 4.40 ± 0.71 11.14 ± 2.58 Acer saccharum Marshall ACSA 16.67 ± 7.00 1.43 ± 0.75 5.96 ± 2.50 1.27 ± 0.27 3.62 ± 1.39 Carpinus caroliniana Walter CACA 3.70 ± 4.42 0.25 ± 0.29 1.32 ± 1.58 0.22 ± 0.10 0.77 ± 0.84 Carya cordiformis (Wangenh.) K. Koch CACO 1.85 ± 2.21 0.20 ± 0.24 0.66 ± 0.79 0.18 ± 0.08 0.42 ± 0.44 Carya glabra (Mill.) Sweet CAGL 12.96 ± 7.81 3.60 ± 2.87 4.64 ± 2.79 3.19 ± 1.03 3.91 ± 1.91 Carya ovata (Mill.) K. Koch CAOV 3.70 ± 3.10 0.80 ± 0.67 1.32 ± 1.11 0.71 ± 0.24 1.02 ± 0.67 Fagus grandifolia Ehrh. FAGR 9.26 ± 5.74 2.28 ± 2.33 3.31 ± 2.05 2.02 ± 0.83 2.67 ± 1.44 Liriodendron tulipifera L. LITU 25.93 ± 8.73 14.23 ± 7.19 9.27 ± 3.12 12.63 ± 2.57 10.95 ± 2.85 Nyssa sylvatica Marshall NYSY 7.41 ± 4.33 1.08 ± 1.01 2.65 ± 1.55 0.96 ± 0.36 1.81 ± 0.95 Oxydendrum arborum(L.) DC. OXAR 3.70 ± 3.10 0.28 ± 0.24 1.32 ± 1.11 0.25 ± 0.08 0.79 ± 0.60 Quercus alba L. QUAL 50.00 ± 12.45 30.89 ± 9.89 17.88 ± 4.45 27.41 ± 3.54 22.65 ± 3.99 Quercus coccinea Münchh. QUCO 9.26 ± 5.74 5.74 ± 3.70 3.31 ± 2.05 5.09 ± 1.32 4.20 ± 1.69 Quercus prinus L. QUPR 44.44 ± 11.85 25.72 ± 7.10 15.89 ± 4.24 22.82 ± 2.54 19.36 ± 3.39 Quercus rubra L. QURU 12.96 ± 6.42 8.22 ± 4.45 4.64 ± 2.30 7.29 ± 1.59 5.96 ± 1.94 Quercus velutina Lam. QUVE 20.37 ± 8.14 11.16 ± 5.05 7.28 ± 2.91 9.91 ± 1.80 8.60 ± 2.36 Sassafras albidum (Nutt.) Nees SAAL 3.70 ± 3.10 0.75 ± 0.64 1.32 ± 1.11 0.67 ± 0.23 0.99 ± 0.67 Tilia americana L. TIAM 3.70 ± 3.10 1.10 ± 1.20 1.32 ± 1.11 0.97 ± 0.43 1.15 ± 0.77

Quercus spp. L. 137.04 ± 19.11 81.73 ± 13.73 49.01 ± 6.83 72.53 ± 4.91 60.77 ± 5.87 Non Quercus spp. 142.59 ± 21.20 30.95 ± 8.27 50.99 ± 7.58 27.47 ± 2.96 39.23 ± 5.27

Total 279.63 ± 22.33 112.68 ± 14.12

Table 9. A permutational multivariate analysis of variance using distance matrices (adonis::vegan (Oksanen et al. 2012)) of the bryophyte communities within the Fire and Fire Surrogate (FFS) study design in three southeastern Ohio forests. Data were permuted 9999 times within blocks (forests) as strata using the Bray- Curtis distance matrix.

Bryophyte community df SS MS F Pa Burn 1 0.68 0.68 6.62 0.001 *** Thin 1 0.19 0.19 1.82 0.063 . Moisture 2 1.42 0.71 6.88 0.001 *** I vs. M&X 1 0.24 0.24 2.35 0.023 * X vs. M 1 1.17 1.17 11.41 0.001 *** Burn:Thin 1 0.11 0.11 1.08 0.297 Burn:Moisture 2 0.33 0.17 1.62 0.071 . I vs. M&X 1 0.08 0.08 0.76 0.569 X vs. M 1 0.26 0.26 2.48 0.025 * Thin:Moisture 2 0.20 0.10 0.99 0.416 I vs. M&X 1 0.10 0.10 1.02 0.346 X vs. M 1 0.10 0.10 0.96 0.411 Burn:Thin:Moisture 2 0.28 0.14 1.34 0.185 I vs. M&X 1 0.14 0.14 1.41 0.137 X vs. M 1 0.13 0.13 1.28 0.171 Residuals 24 2.47 0.10 Total 35 5.68 a '***’ 0.001, ‘**’ 0.01, ‘*’ 0.05, ‘.’ 0.1

Table 10. Multivariate analysis of variance (MANOVA) of bryophyte substrate occurrence change by burning and thinning across moisture types in three southeastern Ohio forests. Substrates included in this analysis were exposed soil (SOIL), coarse woody debris (CWD+STUMP), fine woody debris (FWD), and tree base cover (TREE). Orthogonal 1 df contrasts for moisture are represented. Pillai's trace approximate Numerator Denominator Error: EU df F df df P Forest 2 1.21 10 38 0.314 Burn 1 9.67 5 18 <0.001 *** Thin 1 4.98 5 18 0.005 ** Moisture 2 4.77 10 38 <0.001 *** I vs. X&M 1 4.10 5 18 0.012 * X vs. M 1 19.71 5 18 <0.001 *** Burn:Thin 1 1.88 5 18 0.147 Burn:Moisture 2 1.65 10 38 0.130 I vs. X&M 1 1.80 5 18 0.164 X vs. M 1 1.52 5 18 0.234 Thin:Moisture 2 1.88 10 38 0.079 . I vs. X&M 1 2.80 5 18 0.049 * X vs. M 1 1.55 5 18 0.223 Burn:Thin:Moisture 2 0.83 10 38 0.603 I vs. X&M 1 1.48 5 18 0.246 X vs. M 1 0.35 5 18 0.874 Residuals 22 Total 35 a '***’ 0.001, ‘**’ 0.01, ‘*’ 0.05, ‘.’ 0.1

Table 11. Multivariate analysis of variance (MANOVA) of analysis of bryophyte substrate occurrence change incorporating co-variations in the substrates, themselves by burning and thinning across moisture types in three southeastern Ohio forests. Substrates included in this analysis were exposed soil (SOIL), coarse woody debris (CWD+STUMP), fine woody debris (FWD), and tree base cover (TREE). Cover of rock (ROCK) and slope were included as covariates, and orthogonal 1 df contrasts for moisture are represented.

Pillai's trace approximate Numerator Denominator Error: EU df F df df P Forest 2 1.73 10 28 0.122 Burn 1 8.21 5 13 0.001 ** Thin 1 0.95 5 13 0.482 Moisture 2 3.21 10 28 0.007 ** I vs. X&M 1 1.17 5 13 0.375 X vs. M 1 14.22 5 13 <0.001 *** CWD+STUMP 1 1.10 5 13 0.405 FWD 1 3.31 5 13 0.038 * TREE 1 6.35 5 13 0.003 ** MSOIL 1 2.89 5 13 0.057 . ROCK 1 8.93 5 13 0.001 *** Burn:Thin 1 1.50 5 13 0.255 Burn:Moisture 2 1.95 10 28 0.081 . I vs. X&M 1 1.06 5 13 0.424 X vs. M 1 2.74 5 13 0.067 . Thin:Moisture 2 1.27 10 28 0.294 I vs. X&M 1 1.13 5 13 0.392 X vs. M 1 1.85 5 13 0.171 Burn:Thin:Moisture 2 0.82 10 28 0.613 I vs. X&M 1 1.78 5 13 0.187 X vs. M 1 0.15 5 13 0.977 Residuals 17 Total 35 a '***’ 0.001, ‘**’ 0.01, ‘*’ 0.05, ‘.’ 0.1

FIGURES

Figure 1. Location map of the three forests of the Ohio Hills Site of the USDA Fire and Fire Surrogate (FFS) Research Program: Raccoon Ecological Management Area (REMA), Tar Hallow State Forest (TAR), and Zaleski State Forest (ZAL). Forest cover is shown in green. 75

Figure 2. Treatment unit representation. Nine transects determined by a stratified selection based on long- term moisture conditions derived from an integrated moisture index (IMI) and slope orientation. Stratified by moisture conditions ranging from xeric (dry, red), intermediate (intermediate, green), and mesic (moist, blue) such that three transects were in each the xeric, intermediate, and mesic moisture classes. Diagonal transects involved only the central 50 m between grid points. Grid spacing is 50 m, contour interval is 6 m, and north is at the top of the page. 76

Figure 3. Representation of selected 2 5 m quadrats along established 50 m transects. Quadrat “1” was placed 2.5 m (orthogonal) or 10.4 m (diagonal) from the grid point. Additional quadrats were spaced 5 m apart. Hatched areas represent grid point buffers, light grey areas are unsampled areas, and dark circles indicate grid points. 77

100

Number of Transects (n=108) Transects of Number

20 0

0 20 40 60 80 100

Rank Species Order

Figure 4. Number of transects of occurrence (n = 108) for bryophyte species (n = 112) observed in three southeastern Ohio forests. 78

Number of Species of Number

10 5

0 20 40 60 80 100

Rank Transect Number

Figure 5. Bryophyte richness (n = 112) of individual transects (n = 108) from three southeastern Ohio forests. 79

Figure 6. Non-metric multidimensional scaling (NMDS) ordination and environmental correlates of the bryophyte community a) as a whole and b-d) divided out by moisture classes in three mixed-oak forests in southeastern Ohio. Transects are labeled by treatment within a 2 × 2 factorial of silvicultural thinning and prescribed burning with control (C), thin only (T), burn only (B) and thin and burn together (X). Moisture classes are indicated in a) by the colors associated with b-d). The grey arrows are linear predictors of site condition (Veg, understory vegetation; Open, canopy openness) and substrate cover (LEAF, leaf litter; MSOIL, exposed soil) within each moisture class. The length of the arrow indicates the strength of the relationship with the ordination, and only relationships with p > 0.05 were plotted. Note a) was not fitted with linear predictors and c) had no significant predictors.

. Richness(S) 10 * *

0 C T X B C T X B C T X B X I M

Treatment

Figure 7. Mean richness (±S.E.) of bryophytes across a 2 × 2 factorial design of thinning and burning with control (C), thin only (T), burn only (B) and thin and burn together (X) within xeric (X), intermediate (I), and mesic (M) moisture classes in three southeastern forests. Significance is indicated within moisture class as '***’ 0.001, ‘**’ 0.01, ‘*’ 0.05, ‘.’ 0.1.

Figure 8. Non-metric multidimensional scaling (NMDS) ordination and species correlates of the bryophyte community a) as a whole and b-d) divided out by moisture classes in three mixed-oak forests in southeastern Ohio. Transects are labeled by treatment within a 2 × 2 factorial of silvicultural thinning and prescribed burning with control (C), thin only (T), burn only (B) and thin and burn together (X). Moisture classes are indicated in a) by the colors associated with b-d). The grey arrows are linear predictors of species abundance within each moisture class. The length of the arrow indicates the strength of the relationship with the ordination, and only species occurring in > 10% of the transects having relationships with p > 0.05 were plotted. Species symbols are the first four letters of the genus and species epithets presented in Table 1. Note a) was not fitted with linear predictors. 82

3.0 2.0

2.5

1.5

2.0

1.5 1.0 CWD (%) CWD(%)

1.0

Mean Species Occurence on CWD on SpeciesOccurence Mean 0.5

0.5

0.0 0.0 C T X B C T X B C T X B C T X B C T X B C T X B X I M X I M

Treatment Treatment

Figure 9. Example of differential response bryophytes by substrates and the substrates, themselves. Mean (± S.E.) of species occurrence on coarse woody debris (CWD) and percent of CWD cover across a 2 × 2 factorial design of thinning and burning with control (C), thin only (T), burn only (B) and thin and burn together (X) within xeric (X), intermediate (I), and mesic (M) moisture classes in three southeastern forests. 83

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