Fire history and the establishment of oaks and maples in second-growth forests

Todd F. Hutchinson, Robert P. Long, Robert D. Ford, and Elaine Kennedy Sutherland


Abstract: We used dendrochronology to examine the influence of past fires on oak and maple establishment. Six study units were located in southern Ohio, where organized fire control began in 1923. After stand thinning in 2000, we collected basal cross sections from cut stumps of oak (n = 137) and maple (n = 204). The fire history of each unit was developed from the oaks, and both oak and maple establishment were examined in relation to fire history. Twenty-six fires were documented from 1870 to1933; thereafter, only two fires were identified. Weibull median fire return intervals ranged from 9.1 to 11.3 years for the period ending 1935; mean fire occurrence probabilities  (years/fires) for the same period ranged from 11.6 to 30.7 years. Among units, stand initiation began ca. 1845 to 1900, and virtually no oak recruitment was recorded after 1925. Most maples established after the cessation of fires. In several units, the last significant fire was followed immediately by a large pulse of maple establishment and the cessation of oak recruitment, indicating a direct relationship between fire cessation and a shift from oak to maple establishment.

Résumé : Nous avons eu recours à la dendrochronologie pour étudier l’influence du feu dans le passé sur l’établissement  du chêne et de l’érable. Six unités expérimentales ont été localisées dans le sud de l’Ohio où la lutte organisée contre les  feux a débuté en 1923. Après que des peuplements eurent été éclaircis en 2000, nous avons collecté des sections radiales  sur des souches de chêne (n = 137) et d’érable (n = 204). Dans chaque unité, l’historique des feux a été établi à partir des  chênes et l’établissement du chêne et de l’érable a été étudié en lien avec l’historique des feux. Vingt-six feux ont été documentés de 1870 à 1933; par la suite, seulement deux feux ont été identifiés. L’intervalle médian de Weibull entre les feux variait de 9,1 à 11,3 ans pour la période se terminant en 1935; la probabilité moyenne d’occurrence de feux (années/ feux) pendant la même période variait de 11,6 à 30,7 ans. Parmi les unités, l’origine des peuplements remonte aux environs de 1845 à 1900 et pratiquement aucun chêne n’a été recruté après 1925. La plupart des érables se sont établis après que les feux eurent cessé. Dans plusieurs unités, le dernier feu important a immédiatement été suivi d’une importante vague d’établissement de l’érable et de l’arrêt du recrutement du chêne, indiquant qu’il y a une relation directe entre la cessation des feux et le changement marqué par l’établissement de l’érable au lieu du chêne.

[Traduit par la Rédaction]

CLICK HERE TO DOWNLOAD AS PDF


Introduction

Across much of the eastern United States, red maple (Acer rubrum L.), sugar maple (Acer saccharum Marsh.), and other mesophytic and (or) shade-tolerant species have become abundant in historically oak-dominated landscapes, threatening the continued dominance of oak (Lorimer 1984; Abrams 1992). Fire-control policies instituted ca. 1910 to 1930 often are considered a primary cause of these successional trends (e.g., Lorimer 1993).

Oaks are considered to be better adapted than maples to a regime of periodic fires primarily because of their relatively thick and, thus, fire-resistant bark; their ability to compartmentalize wounds caused by fire; and the capacity of established seedlings to continue to sprout after being top-killed  repeatedly (Smith and Sutherland 1999; Johnson et al. 2002;  Van Lear and Brose 2002). Periodic anthropogenic fire is widely considered to have promoted and sustained eastern  oak ecosystems throughout their postglacial history  (Abrams 2002). However, specific knowledge of past fire  regimes, which can be obtained by analysis of fire-scarred  trees, is limited to relatively few areas. Several studies  show that fire was frequent in oak ecosystems prior to  (Cutter and Guyette 1994; Guyette et al. 2003; Shumway  et al. 2001) and after Euro-American settlement (Sutherland 1997; Schuler and McClain 2003; Guyette and Stambaugh 2004; Soucy et al. 2005) until fire control was instituted.


Received 8 August 2007. Accepted 8 November 2007. Published on the NRC Research Press Web site at cjfr.nrc.ca on 26 April 2008.
T.F. Hutchinson,1 R.P. Long, and R.D. Ford. USDA Forest Service, Northern Research Station, 359 Main Road, Delaware, OH 43015, USA.
E.K. Sutherland. USDA Forest Service, Rocky Mountain Research Station, 800 Block East Beckwith, P.O. Box 8089, Missoula, MT 59807, USA.

1Corresponding author (e-mail: thutchinson@fs.fed.us).


Much more is known about long-term patterns of tree establishment in old-growth oak forests. These studies often suggest a strong influence of fire cessation on tree recruitment (e.g., Abrams and Downs 1990; Abrams and Copenheaver 1999; Aldrich et al. 2005). Oak recruitment is known to have occurred for up to several hundred years but decreased or ceased around the time that fire control began. Several of these studies also document greatly increased recruitment of maples and other nonoak species during the same period (e.g., Abrams and Downs 1990; Abrams and Copenheaver 1999).

Shumway et al. (2001) were the first to document both the fire history and patterns of establishment for oak and other species in an old-growth oak-dominated stand in western Maryland. The authors showed that fire and oak recruitment were frequent from the early 1600s through the early 1900s. Both the cessation of oak recruitment and the increased recruitment of red maple and black birch (Betula lenta L.) coincided with reduced fire frequency ca. 1930. Soucy et al. (2005) showed that oak-hickory stands in the Arkansas Ozarks originated following harvesting or fire ca. 1900 and that fires were frequent through the 1930s. As fires became much less frequent after ca. 1940, oak recruitment ceased, and other shade-tolerant, nonoak species, such as flowering dogwood (Cornus florida L.), red maple, and blackgum (Nyssa sylvatica Marsh.), became established (Soucy et al. 2005).

The unglaciated ‘‘hill country’’ of southeastern Ohio was dominated by oak forests ca. 1800, just prior to Euro-American settlement (Beatley 1959; Gordon 1969; Dyer 2001). After nearly all of the forests in the region were cut over in the 19th century, many stands regenerated to oak dominance (Goebel and Hix 1997; Dyer 2001; Yaussy et al. 2003), and oak remains abundant in the region today (Griffith et al. 1993). However, as with most areas in the eastern United States that historically were oak dominated, the continued abundance of oak is threatened by increasing densities of maples and other species (e.g., blackgum and beech (Fagus grandifolia Ehrh.)) and poor oak regeneration. Dendrochronology fire histories indicate that fires occurred frequently in the region from ca. 1870 to 1935 (Sutherland 1997; McEwan et al. 2007b). It is hypothesized that this fire regime sustained oak dominance as second-growth forests developed (McEwan et al. 2007b) and that fire control directly facilitated the establishment of the now-abundant maples and other competitors (Sutherland et al. 2003).

To better understand how past fires were related to tree establishment, we conducted a dendrochronology study at the Ohio Hills site of the national Fire and Fire Surrogate Study (FFS). Our study was carried out on three replicate sites, each containing two separate units (*20 ha each) that here thinned. The second-growth forests were dominated by oak in the overstory, but maples and other shade-tolerant species were abundant in the midstory and understory. We collected basal cross sections from cut stumps of both oaks and maples to document stand fire histories and tree establishment. We hypothesized that, within a stand, temporal patterns of oak and maple recruitment would be closely related to the occurrence of past fires. Specifically, we hypothesized that (i) fires were frequent prior to the initiation of fire control, (ii) oak establishment occurred primarily before the initiation of fire control, and (iii) maples established primarily during the fire-control era. By studying units within three spatially separated replicate sites that likely had different fire histories, we also hoped to better understand how variability in past fire regimes affected oak and maple establishment. To our knowledge, this is the first study that directly examines the relationship between specific historic fire events and oak and maple recruitment events. A better understanding of how past fire regimes affected the pattern and pace of recruitment during stand development could provide new insights for the use of prescribed fire to manage oak forests.


Methods

Study area and site descriptions

The study area is in southern Ohio within the Southern Unglaciated Allegheny Plateau (McNab and Avers 1994). The topography is highly dissected, consisting of sharp ridges, steep slopes, and narrow valleys. The bedrock geology is predominantly sandstones and shales that produce well-drained and acidic soils.

The Ohio Hills FFS study site has three replicates (hereafter sites): one each in the Raccoon Ecological Management Area (REMA), Zaleski State Forest, and Tar HollowState Forest. The REMA site (39812’34@N, 82823’07@W) is in Vinton County and within the Vinton Furnace Experimental Forest; owned by Forestland Group, LLC, and comanaged with the USDA Forest Service Northern ResearchStation. The Zaleski site (39821’22@N, 82821’59@W), also in Vinton County, is 19 km north of the REMA site. The Tar Hollow site (39819’ 47@N, 82846’11@W) is in Ross County, 35 km west of the Zaleski site. Soils at both REMA and Zaleski are predominantly Steinsburg and Gilpin series silt loams (Typic Hapludalfs); Tar Hollow soils are predominantly Shelocta-Brownsville complex sandy loams (Typic Hapludalfs and Typic Dystrochrepts, respectively) (Boerner et al. 2007). Both state forests are managed by the Ohio Department of Natural Resources’ (ODNR) Division of Forestry.

Human land use has had a major effect on these forests. Both the REMA and Zaleski sites are located near charcoal iron furnaces that operated in the 1800s. REMA is <2 km from Vinton Furnace and Zaleski is <4 km from Hope Furnace; these were in operation from 1853 to 1883 and from 1854 to 1874, respectively (Stout 1933). Forests at both sites presumably were harvested at least once to provide charcoal for iron smelting. The Tar Hollow site was not affected by the iron industry since it was more than 25 km from the nearest furnace. Land deeds show direct human occupation in small parcels within the Tar Hollow study site until the mid-1930s (ODNR, Division of Forestry, District Office, Chillicothe, Ohio).

The forests generally were similar in structure and composition across the three sites. Prethinning data collected in 2000 showed that mean stand basal area at REMA was 28 m2/ha; white oak (Quercus alba L.) accounted for 21% of the basal area; black oak (Quercus velutina Lam.), 17%; chestnut oak (Quercus montana Willd.), 15%; and scarlet oak (Quercus coccinea Muenchh.), 12% (D.A. Yaussy, USDA Forest Service, Delaware, Ohio, unpublished data).

Mean basal area at Zaleski was 27 m2/ha and was dominated by chestnut oak (31%), followed by white oak (22%), red maple (13%), and black oak (13%). At Tar Hollow, mean basal area was 32 m2/ha, and the dominant species were chestnut oak (33%), white oak (20%), and black oak (16%). Oak site indices (base age 50 years) are variable across the landscape because of the dissected topography, ranging from about 17 m (55 ft) on upper south-facing slopes to 24 m (80 ft) on lower north-facing slopes (D.A. Yaussy, USDA Forest Service, Delaware, Ohio, personal communication). On all sites, the sapling layer (1.4 m tall to 9.9 cm diameter at breast height (DBH)) and the midstory (trees 10-25 cm DBH) were dominated by shade-tolerant trees, the most abundant of which were red maple, sugar maple, blackgum, and beech (Albrecht and McCarthy 2006). Shade-tolerant trees >25 cm DBH occurred at low densities on all sites.

The mean annual temperature and precipitation are 11.3 8C and 1024 mm, respectively. Precipitation is distributed fairly evenly throughout the year with no months averaging <60 mm. Today, most fires occur during the early spring (March and April) and fall (October and November), when vegetation is predominantly dormant; spring dormant season fires are the most frequent (Haines et al. 1975; Sutherland et al. 2003), and nearly all fires are anthropogenic in origin.

At each FFS site, four treatment units (19-26 ha) were established: an untreated control (control), mechanical thinning (thin), prescribed fire (burn), and a combination of thinning and fire (thin+burn). Our dendrochronology study was conducted on the two thinned units (thin and thin+burn) at each site. Midstory thinning occurred from November 2000 to April 2001 and favored the retention of dominant and codominant oaks. However, to meet commercial thinning objectives, some dominant and codominant oaks were harvested. Across sites, stand density (trees ‡10 cm DBH) was reduced by 32% from a mean of 400 to 269 trees/ha, and tree basal area was reduced by 30% from a mean of 29 to 20 m2/ha.

Sampling design and field methods
At the REMA site, the two thinned units (hereafter, REMA 2 and REMA 3) were separated by a triangular wedge of untreated forest that ranged in width from several meters to 275 m. The Zaleski thinned units (Zaleski 2 and Zaleski 3) were contiguous, and the boundary between units was an intermittent stream drainage. The Tar Hollow thinned units (Tar Hollow 2 and Tar Hollow 3) also were contiguous, but the boundary did not follow a major topographic feature. Despite the contiguous units at Zaleski and Tar Hollow, we treated the units separately for summary and  nalyses, because 50% (14 of 28) of the fires that we document were recorded only in a single unit.

Ten 0.1 ha plots were established in each unit to monitor vegetation and soils for the FFS study. Plot corners were georeferenced with global positioning system (GPS) technology. The plots were distributed across the landscape to represent a continuous range of soil moisture conditions from dry (upper south-facing slopes) to mesic (lower north-facing slopes). In 2000, all overstory trees (‡10 cm DBH) were tallied by species and DBH on each plot prior to treatments. We focused our collection of oak and maple basal cross sections on the plots to utilize the tree data and georeferenced locations.

Full basal cross sections were cut from stumps with a chainsaw from December 2000 to May 2001, soon after thinning operations had been completed in each unit. All oak stumps in a plot were examined for the presence of wounds (discoloration, seams, staining, and wound wood ribs) that might indicate a fire event (Smith and  Sutherland 1999). We attempted to locate at least two oak stumps within or adjacent to each plot. We collected 137 oak cross sections across all units and recorded the upslope position on each. Samples included 64 white oak, 40 chestnut oak, 22 black oak, 7 scarlet oak, and 4 northern red oak (Quercus rubra L). All samples were cut at a height of about 5-10 cm aboveground. The mean basal diameter of the oak samples was 47.4 cm and ranged from 23.1 to 98.1 cm. We mapped the approximate location of each sample based on its position within or adjacent to a georeferenced vegetation plot.

To determine the temporal pattern of maple establishment, our objective was to collect basal cross sections from three stumps in and (or) adjacent to each plot. Our goal was to collect three maples for every two oaks, because maples were approximately 1.5 times as abundant as oaks in the midstory, where the thinning treatment was focused. Our first priority was to obtain larger maples to document the time when establishment began, but we sampled across a range of maple stump diameters. We collected 30-39 maples per unit for a total of 204 (142 red maple and 62 sugar maple). Nearly all sugar maple samples were collected in three units: Tar Hollow 2 (n = 27), Tar Hollow 3 (n = 20), and REMA 3 (n = 15). The mean basal diameter of maple samples was 27.5 cm and ranged from 10.1 to 58.4 cm.

Laboratory methods
Cross sections were planed and sanded to enhance ring boundaries and facilitate dating. Each oak sample was crossdated using skeleton plots (Stokes and Smiley 1968) against a previously established master chronology for the region (Sutherland 1997). Maple cross sections ‡70 years old also were skeleton plotted and cross-dated. Younger maple samples were ring-counted along two to four radii and crossdated by identification of key stress years. Several factors contributed to make the exact pith dates for maples somewhat less precise than that for the oaks. Firstly, the crossdating was less clear for maples than for the oaks, i.e., key stress years were not as consistent among the maples. Secondly, a small proportion of the maples had decay or incipient decay in or near the pith, which obscured the ring boundaries. Thirdly, some maples had rings that were locally absent in a portion of their circumference, a common phenomenon documented by Lorimer et al. (1999) for suppressed sugar maple trees. Several samples that we felt could not be reliably dated (primarily small suppressedstems) were omitted from further analyses.

We required at least three scarred samples per unit per year to classify a wound event as a fire scar and, consequently, a year as a fire year. If only two wounds were present in a unit in a given year but the adjacent unit showed evidence of a fire in the same year (three or more samples scarred), we recorded a fire for the unit with only two scars (this occurred once). These criteria were applied to limit the likelihood that wounds caused by other factors (e.g., logging, falling trees and branches, or animals) were recorded as fire scars (see McEwan et al. 2007a). The seasonality of each fire event was determined by examining where the wounds intersected the annual growth ring. For dormant-season scars (located between annual growth rings) it was not possible to determine whether the scar occurred in the fall after the previous growing season or in the late  Hutchinson et al. winter – early spring prior to the upcoming growing season (Sutherland 1997). Because fires in our region are most frequent in the early spring dormant season (March-April), we assigned dormant-season wounds to the calendar year of the upcoming growing season. For example, a dormant-season wound located between the 1922 and 1923 annual growth rings was recorded as a 1923 wound.

To better examine how the relative intensity and (or) extent of fires may have affected tree establishment, we defined fires as ‘‘significant’’ if (i) ‡33.3% of the samples exhibited wounds and (ii) at least five samples had wounds. Since little is known about the relationship between fire intensity and scarring in oaks (see Smith and Sutherland 1999; Guyette and Stambaugh 2004; McEwan et al. 2007a), this definition provides a relative measure of the intensity and extent of the fires within this study. We mapped the location of all samples (oaks and maples) in all fire years at REMA to visualize the spatial pattern of fire scars and tree establishment across the landscape. We used the ArcView version 3.2a geographic information system to map samples based on our field maps that showed locations within or adjacent to a georeferenced vegetation plot. We selected eight of the nine fire years at the REMA study site to illustrate spatial patterns of fire scarring and tree establishment.

Ring widths were measured on the oaks so that growth dynamics and potential release events could be determined. Oak cross sections were scanned and measured using WINDENDRO (Regent Instruments Inc., Ste-Foy, Que.). Two radii approximately 1808 apart were measured in each cross section. Radii were located to minimize the influence of wounds and associated wound wood on growth measurements. Ring-width measurements and crossating were verified with the program COFECHA 2.1 (Holmes 1983; Grissino-Mayer 2001b). The program ARSTAN (Cook and Kairiukstis 1990) from the Lamont Doherty Earth Observatory’s Tree-Ring Laboratory, was used to detrend measurements with a negative exponential curve or linear regression line. This standardization procedure removes the growth trend associated with age and produces dimensionless indices that can be averaged to create a master chronology for a site (Fritts 1976). A master chronology was created for each of the six sampled sites.

Potential releases associated with disturbance events were identified in each master chronology with the JOLTS program (Holmes 1999) from the International Tree-Ring Data Bank Dendrochronology Program Library. Major releases were those where there was a >100% increase in growth expressed as the mean chronology ring-width index over a 15 year period compared with the mean chronology ringwidth index in the preceding 15 year period. Minor releases were those where growth increased 50% over a 10 year period compared with a previous 10 year period (Lorimer and Frelich 1989; Soucy et al. 2005). We report release events only when there were at least 10 trees present in the master chronology. Fire-return interval analyses Data on fire history derived from oak cross sections were analyzed using the FHX2 program (Grissino-Mayer 2001a, 2004). Because fires were so infrequent after 1935, it was not possible to statistically compare fire frequency between pre- and post-fire suppression periods. Instead, for each study unit, we calculated fire intervals from the first fire to 1935 (before and during the early fire-control period) and compared these with fire intervals from the first fire to 2000. Both mean fire intervals (MFI) and Weibull median fire intervals (WMFI) were calculated. The latter is considered a better estimator of central tendency for the typically nonnormal fire-interval distributions (Grissino-Mayer and Swetnam 1997; Grissino-Mayer et al. 2004).

For each unit, we also calculated the mean fire occurrence probability (MFOP) for two periods (stand origination to 1935 and to 2000). Defined by Guyette et al. (2006), MFOP is the number of years divided by the number of fires in a chronological period. Guyette et al. (2006) calculated the MFOP to account for the fire-free period prior to the first recorded fire. For each site, we defined the stand origination as the first year in which at least four samples were present that could potentially record a fire.

Results

Twenty-eight fires were recorded, of which 26 occurred from 1870 to 1933 (Table 1). Twelve fires scarred five or more samples and were classified as significant fires (‡33.3% of the samples were scarred); these fires occurred from 1877 to 1923. Most wounds attributed to fire were recorded on small-diameter trees; for all fires, the basal diameter of oaks at the time of wounding was 12.7 ± 0.7 cm (mean ± SE). In most of the fires (n = 24), all wounds were located between annual growth rings, indicating occurrence in the dormant season (September to early April). In fire years, 147 of the 213 total wounds (69%) were on the uphill portion of the stem (3008 clockwise to 608), based on the uphill position recorded on the sample in the field. Of the 204 maple samples, only one exhibited a fire scar (a wound in a fire year); that maple, from Zaleski 2, had a wound in 1965.

Fire histories of the study units

REMA 2 had the greatest number of fires (n = 7) and significant fires (n = 5) (Table 1); fires were documented from 1877 to 1933. The 1917 significant fire had both dormant and earlywood scars, suggesting an early growing season fire. In the 1933 fire, most wounds were present in the late earlywood; in that fire, all seven scarred trees were young and small, having established in 1923 or 1924 and averaged only 5.1 cm in basal diameter (Table 1). We recorded six fires at REMA 3, three of which were significant, from 1878  1923 (Table 1; Fig. 1). As with REMA 2, the wounds in the 1917 fire indicate an early growing season fire; in the 1906 fire, all three wounds intersected the earlywood.

Zaleski 2 had evidence of six fires; five occurred from 1870 to 1928, and a sixth occurred in 1965 (Table 1). Although only the 1923 fire was classified as significant, it wounded 83% (15 of 18) of the samples; none of the other fires wounded more than three samples. No fires were recorded at Zaleski 2 during a 25 year period from stand origination (1844 to 1869). At Zaleski 3, the chronology was shorter, dating from 1880 to 2000. Although only three fires were recorded, both the 1917 and 1923 fires were significant. Again, the 1923 fire wounded a high percentage (61.9%) of the samples. All fire scars in both Zaleski units were in the dormant season.

Table 1. Summary data for the 28 fires documented on the six study units.


Diameter scarred (cm)

Study site, sample size,and chronologyaFire yearFire seasonbScarred tree (%)No. of scarred treesTotal no of treesMeanRange
REMA 2
n = 29
1855-2000
1877
1885
1895
1900
1917
1923
1933
D
D
D
D
D and E
D
E
50.0
66.7
25.0
50.0
50.0
50.0
20.7
6
8
3
6
6
9
7
12
12
12
12
12
18
29
8.6
13.0
14.4
18.1
22.0
13.1
5.1
7.0-10.7
9.5-17.2
13.5-15.2
15.4-23.1
18.0-28.8
3.3-25.8
2.7-7.5
REMA 3
n = 22
1858-2000
1878
1885
1895
1906
1917
1923
D
D
D
E
D and E
D
23.1
38.5
57.1
23.5
33.3
18.2
3
5
8
3
7
4
13
13
14
17
21
22
7.4
11.0
14.2
18.4
11.7
16.8
4.6-10.5
6.7-14.5
5.5-19.5
12.1-21.5
3.9-21.7
8.6-25.0
Zaleski 2
n = 24
1844-2000
1870
1897
1917
1923
1928
1965
D
D
D
D
D
D
75.0
33.3
16.7
83.3
12.5
12.5
3
3
3
15
3
3
4
9
18
18
24
24
4.7
5.0
13.3
16.0
11.1
22.5
4.2-5.4
1.2-8.3
6.3-18.1
7.5-29.8
4.0-15.6
16.8-33.7
Zaleski 3
n = 25
1880-2000
1914
1917
1923
D
D
D
22.2
35.0
61.9
4
7
13
18
20
21
5.2
8.6
10.3
3.8-6.8
4.9-13.6
6.3-17.2
Tar Hollow 2
n = 22
1844-2000
1883
1900
1926
D
D
D
71.4
26.7
22.7
5
4
5
7
15
22
7.9
4.4
13.4
5.7-13.7
1.5-8.0
8.0-24.0
Tar Hollow 3
N = 17
1899-2000
1900
1912
1984
D
D
D
50.0
25.0
17.6
2
3
3
4
12
17
6.4
7.2
40.7
3.0-6.8
3.7-10.7
36.3-47.0

Note: All data are from the oak samples. Years in bold type indicate fires that scarred five or more trees and ‡33.3% of samples.
aSample size is the total number of oak samples. Chronology period begins with the first year when four or more samples were present to record fires.
bD, dormant season; E, earlywood; D and E, both dormant and earlywood wounds were present in the samples.


We recorded three fires at Tar Hollow 2 from 1883 to 1926 (Table 1). No fires were documented in the 39 years from stand origin (1844) to the 1883 fire; that fire was the only significant fire, wounding five of seven trees. Tar Hollow 3 had the shortest chronology (1899-2000) of all units; fires were documented in 1900, 1912, and 1984. Only two trees were scarred in 1900, but this is included as a fire because of its concordance with the four trees scarred in 1900 in Tar Hollow 2. Only three trees were scarred in the 1912 and 1984 fires at Tar Hollow 3. As with Zaleski, all fire scars were located between annual growth rings, indicating dormant-season fires.

Fire-return intervals
In the period before active fire control and ending in 1935, composite mean fire intervals (MFI) only could be calculated at three of the six units (REMA 2, REMA 3, and Zaleski 2). At these units, MFI ranged from 9.0 to 14.5 years (Table 2). Likewise, the composite Weibull median fire interval (WMFI) ranged from 9.1 years at REMA 2 to 11.3 years at Zaleski 2. For the same pre-1936 period, the mean fire occurrence probabilities (MFOP; Guyette et al. 2006), which also take into account the period of time prior to the first fire, ranged from 11.6 and 12.2 years at REMA 2 and REMA 3, respectively, to 30.7 years at Tar Hollow 2. For the five units originating in 1880 or before (all but Tar Hollow 3), there was a period of at least 20 years from stand origination to the first recorded fire. As only two fires were documented from 1936 to 2000, fire-interval calculations that end in 2000 are longer (Table 2). The WMFI ranged from 12.7 and 13.9 years at REMA 2 and REMA 3, respectively, to 35.4 years at Tar Hollow 2.

Fire and the establishment of oaks and maples
At REMA 2, all oak samples established prior to 1924, and nearly every maple recruited after the 1923 fire (Fig. 2a).

Fig. 1. Fire history diagram for REMA 3. The broken horizontal lines represent the growth years for the 22 oak samples. The solid triangles are wounds that were in a recorded fire year; vertical bars are wounds present in years not recorded as fire years. The six fire years are indicated by the vertical lines located above the timeline.

 


 

Study sitePeriod ending in1935Period ending in 2000
MFIWMFI (87.5%-12.5%)MFOP (years/fires)MFIaWMFI (87.5%-12.5%)aMFOP (years/fires)
REMA 29.39.1 (4.8-14.1)11.6 (81/7)17.612.7 (2.8-36.1)20.9 (146/7)
REMA 39.09.2 (6.7-11.3)12.2 (73/6)20.313.9 (2.6-42.4)23.8 (143/6)
Zaleski 214.511.3 (2.8-28.8)18.4 (92/5)21.718.2 (5.5-40.3)26.2 (157/6)
Zaleski 318.7 (56/3)28.714.0 (1.5-63.2)40.3 (121/3)
Tar Hollow 230.7 (92/3)39.035.4 (13.3-68.2)52.3 (157/3)
Tar Hollow 318.5 (37/2)33.327.3 (7.6-64.2)34.0 (102/3)

Note: A dash indicates that there were an insufficient number of fire events to calculate the interval.
aFire interval calculations in these columns are based on a final incomplete interval ending in 2000.

After an initial period of oak establishment (1852 to 1865), presumably after harvesting for the charcoal iron industry, there was a 51 year period (1866 to 1916) when no oak recruitment was recorded. Thereafter, two pulses of oak establishment were documented immediately after the significant fires of 1917 and 1923. No maples predated the 1917 fire, and several maples established between the 1917 and 1923 fires. In 1923, immediately after the last significant fire, 15 maples recruited. Thereafter, 17 maples established from 1924 to 1938.

At REMA 3, initial oak establishment occurred from 1849 to 1860. As with REMA 2, no establishment was recorded after 1924 (Fig. 2b). After 1860, there were no large pulses of oak establishment, but there were 7 years from 1885 to 1924 in which pith dates were recorded for one or two oaks. The oldest maple dated to 1921, and a pulse of 11 stems established in 1923, immediately after the last fire. An additional 12 maples established from 1924 to 1949.

The temporal patterns of fire and establishment at the Zaleski units were similar to those of REMA, remarkably so for the initiation of maple establishment and the corresponding cessation of oak recruitment. At Zaleski 2, four oaks established in the 1840s (Fig. 2c). There was a period of oak recruitment from 1872 to 1880, with a pulse of eight stems in 1879 and 1880, following the 1879 fire. After 1880, we record virtually no oak recruitment for 42 years (1881 to 1922). Oaks then established in 1923 and 1924, immediately after the 1923 fire which scarred 15 of 18 oaks. Maple establishment began in 1922 (n = 4), just before the 1923 fire; four others dated to 1923. Thereafter, 22 maples established from 1924 to 1965, with a maximum of three stems in a single year.

 

Fig. 2. (af) Temporal establishment of oaks and maples for the six study units. Fires are indicated by vertical lines above the timeline; significant fires, those with ‡33.3% of samples wounded, are indicated by vertical arrows.

 

At Zaleski 3, we recorded 5 oaks that established before 1900 (primarily ca. 1880); then, 10 trees established in 1902 (Fig. 2d). The 1902 pulse of oak recruitment was not associated with a fire. As in unit 2, there was another period of oak establishment (n = 5) in 1923 and 1924, immediately following the 1923 significant fire; thereafter, we recorded only a single oak that established in 1954. As in unit 2, maple recruitment initiated in 1922 (n = 4), and eight trees established in 1923, directly after the significant fire. From 1926 to 1928, 11 maples established, and 11 others had pith dates from 1937 to 1959.

Tar Hollow 2 exhibited an early period of oak establishment (n = 8) from 1835 to 1851, six of the eight trees had pith dates of 1842 and 1843 (Fig. 2e). We recorded no oak establishment from 1852 to 1885; 15 trees established from 1886 to 1919. There was a small pulse (n = 3) of oak recruitment in 1900 after the fire of that year. Unlike REMA and Zaleski, maple establishment began nearly 40 years earlier at Tar Hollow 2. Maples (both red and sugar) recruited for 70 years (1881 to 1951) in a fairly continuous manner but did not exhibit the large pulses recorded at REMA and Zaleski.

The oldest oak recorded in Tar Hollow 3 dated to 1851, but no other samples predated 1894 (Fig. 2f). We record fires in 1900 and 1912. The 1900 fire scarred two of the four samples, and there was a pulse of oak recruitment (n = 6) that year. Thereafter, six oaks had pith dates from 1902 to 1924; no more than one tree was recorded in any single year. Maple recruitment at Tar Hollow 3 spanned from 1897 to 1963. As in Tar Hollow 2, there were no large establishment events. At both Tar Hollow units, despite different patterns of fire and an earlier initiation of maple establishment compared with REMA and Zaleski, oak establishment ceased at the same time at all sites (ca. 1920 to 1925).

Spatial distribution of fire scars and tree establishment at REMA

Fires that occurred in 1885, 1895, 1917, and 1923 were recorded in units 2 and 3 (Fig. 3). The 1885 and 1917 fires were classified as significant in both units. By contrast, the presence of fire-scarred trees was limited to one unit in the other five fire years (1877, 1878, 1900, 1906 [not shown], and 1933). Presumably, these fires did not burn across the intermittent stream drainage separating the two units. Similarly, the stream drainage in the center of unit 2, running southwest, appears to have limited fire spread in several years when trees were scarred only northwest (1895 and 1900) or southeast (1933) of the drainage.

For all fire years, scarred oaks were located near oaks that were not scarred. The trees most prone to exhibiting fire scars were in the northern portion of unit 2, near the top of the ridge; two or more of the seven trees that had established there before the first fire in 1877 were scarred in all unit 2 fires prior to 1933.

Maple establishment is first shown in both units on the 1923 fire map (Fig. 3g); these trees established from 1917 to 1922 and survived the 1923 fire. In unit 3, most of the maples that established before 1923 were in areas where fire scars were not recorded on oaks in 1923, suggesting that those small trees were in unburned patches. In unit 2, all three maples predating 1923 are within 5-20 m of an oak with a 1923 fire scar; however, all three oaks with fire scars were small in 1923, each having established immediately after the 1917 fire. By the time of the 1933 fire (Fig. 3h), maples had established across most of the landscape. All of the maples in the 1933 fire map were relatively near scarred oaks and thus escaped that fire; none of these samples had 1933 wounds. However, the low-intensity and perhaps patchy nature of the 1933 spring growing season fire is suggested by the fact that only small oaks (mean basal diameter 5.1 cm) that established after the 1923 fire were scarred.

Radial growth and releases

The master chronologies show growth that is typical of trees from forest interior sites and show only several sustained release events (Fig. 4). Growth releases were identified at only two of the six units; however, none of these releases coincided with a fire. Zaleski 2 had a major release beginning in 1896, perhaps coinciding with a harvest based on its magnitude. No oak recruitment was associated with this release. A moderate release also was identified at this site for 1906. REMA 3 had a major release in 1864  (Fig. 4), although considerable variability in early growth associated with the small sample size may partially account for this release event. No growth releases were identified at Zaleski 3, REMA 2, or at the Tar Hollow units. At REMA 3, some oak recruitment preceded the 1864 release event, but there is no evidence that these were related (Figs. 2 and 4). These analyses, based on the standardized mean ring width chronologies, indicate that fires were of insufficient intensity to cause standwide mortality and the release of surviving oaks.


Discussion

Historic fire regime

Generally, fires were frequent from ca. 1870 to 1935 as stands developed but were uncommon thereafter, reflecting the regional postsettlement history of anthropogenic fire and its suppression. A 1920-1922 forest survey of 10 southern Ohio counties reported that 25% of all forested land showed visible evidence of having burned at least once within the previous decade (ODNR, Ohio Division of Forestry, Columbus, Ohio). Data from the same survey indicated that 5% to 7% of forested land burned annually (Ohio Experiment Station 1922). Organized fire control was instituted in 1923, and its infrastructure and effectiveness developed rapidly.  By 1935, 19 fire lookout towers had been erected in 8 southern Ohio counties, and 447 fire wardens were employed (Leete 1938). From 1926 to 1935, the mean annual forest acreage burned had been reduced to 0.8% (Leete 1938); from 1950 to 2000, it was further reduced to only 0.1% per year (ODNR, Division of Forestry, Columbus, Ohio).

Our study adds to the growing body of dendrochronological evidence that fire was frequent in the central hardwood region prior to organized fire control; examples include oak and oak-pine community types in the Missouri and Arkansas Ozarks (Cutter and Guyette 1994; Guyette et al. 2002; Guyette and Spetich 2003; Soucy et al. 2005); pine-oak communities in the southern Appalachians (Brose and Waldrop 2006); post oak (Quercus stellata Wang.) barrens in Indiana (Guyette et al. 2003) and Tennessee (Guyette and Stambaugh 2004); and oak forests in southern Ohio (Sutherland 1997; McEwan et al. 2007b), Maryland (Shumway et al. 2001), and West Virginia (Schuler and McClain 2003). The fire-return intervals that we calculated for the REMA and Zaleski sites, ranging from 9.1 years prior to 1936 to 18.2 years overall (WMFI), are within the range reported in those studies (2-24 years), despite our more conservative criteria for classifying fire years. However, the 35 year firereturn interval at Tar Hollow 2 exceeds the range in the other studies.

In the central hardwoods region, dissected topography is known to have limited the spread of fires historically (Guyette et al. 2002). In our study, mapped fire-scarred trees suggest that even relatively small intermittent stream drainages limited fire spread in some years, resulting in some fires that were recorded on, and presumably burned, only a portion of the 20 ha units. By contrast, several of the significant fires spanned two units, scarring trees as far as 900 m apart.

Fig. 3. (ah) Spatial distribution of oaks and maples at REMA in eight fire years. Solid circles indicate oaks scarred by a fire in the year associated with the map, and open circles show oaks that were not scarred in that year. The shaded triangles indicate the location of maples that had established by the time of the 1923 (Fig. 3g) and 1933 (Fig. 3h) fire years. (No maples were documented to have established at the time of the 1917 fire or before.)

 

Fig. 4. Master tree ring chronologies (ARSTAN) showing fire events (vertical arrows) and the point when a minimum of 10 trees were averaged (vertical line) into the mean chronology. Major (M) and moderate releases (m) were only noted in the Zaleski 2 and REMA 3 chronologies.

As other dendrochronological fire-history studies in the region have shown (e.g., Sutherland 1997; Shumway et al. 2001; McEwan et al. 2007b), the great majority of fires occurred in the dormant season (September to early April). Only fires in 1906, 1917, and 1933 at REMA had wounds located in the earlywood. In southern Ohio, radial growth (earlywood production) in oak begins in middle to late April, during bud-swelling and leaf unfolding (Phipps 1961). Oak cross sections collected in early May clearly show earlywood production, whereas samples from mid June show latewood production (R.W. McEwan, Department of Forestry, University of Kentucky, Lexington, Ky., unpublished data). Thus, we estimate that fires exhibiting both dormant and earlywood scars likely occurred in mid-April at the onset of radial growth (e.g., the 1917 REMA fire). The single fire (REMA 2 in 1933) that exhibited wounds intersecting the late portion of the earlywood probably occurred in May.

Sutherland (1997) and McEwan et al. (2007a) showed that historic fire occurrence in this region was not related strongly to monthly climatic conditions. Similarly, we recorded fires in both wet and dry periods. However, for the years in which fires occurred at more than one study site (1900, 1917, and 1923), all exhibited two or more months of drought conditions (Palmer drought severity index more negative than -1.5) the previous fall (1900, 1917, 1923) or also in the spring of the recorded fire year (1900).

The intensity of fires as these stands developed is difficult to determine with certainty because research is lacking that directly relates fire intensity to scarring in oak. However, several studies that have examined patterns of scarring in oak following prescribed fires provide some insight. Smith and Sutherland (1999) found that 14 of 18 small oak trees (4-23 cm DBH) had at least one fire scar after two low-intensity prescribed fires (flame lengths generally <50 cm with no overstory tree mortality). Guyette and Stambaugh (2004) showed that 35% to 65% of mostly small post oak trees (10-25 cm basal diameter) were scarred during three separate prescribed fires in an oak community in Tennessee. These fires burned 72%-93% of the area and reduced stand density (mostly small-diameter trees) by 35%. However, in both studies, only trees with visible bark char were selected for sampling. In our study, we found that, on average, 40% of the oak samples, most of which were small at the time (5 to 25 cm basal diameter) were scarred in the historic fires. These scarring percentages suggest that the fires would have been similar in intensity to the prescribed fires reported by Smith and Sutherland (1999) and Guyette and Stambaugh (2004). Although pulses of oak establishment immediately after some historic fires in our study suggest abundant resprouting after top kill, there is no evidence of high-severity stand-replacement fires even in these relatively young, regenerating stands.

McEwan et al. (2007a) reported that during 15 separate prescribed fires that were similar in intensity to those in Smith and Sutherland (1999), the scarring rate was much lower (12.6%) in white oak. However, because the sample trees in that study were much larger (most were >20 cm DBH), it is difficult to compare those scarring percentages with the historic scarring of small trees in our study.

Five years after a prescribed fire, Wendel and Smith (1986) found that 66% of overstory trees (all species, >12.7 cm DBH) exhibited fire scars visible on the exterior of the stem as exposed wood with callous tissue. The fire in their study was higher in intensity, reducing stand basal area by nearly 20%. The high scarring percentage of larger trees in their study suggests a higher intensity fire than was typical of the historic fires in our study.

Fire, land use, and tree establishment
At REMA and Zaleski, periods of oak and maple establishment were related to specific fire events as these stands developed. The establishment and subsequent survival of maples generally began immediately after the cessation of significant fires, i.e., fires that wounded at least one-third of the oak samples. The final oak establishment event also occurred directly after the last significant fire at three of the four REMA and Zaleski units. Because maples seldom were recorded as witness trees in upland forests just before Euro American settlement (Beatley 1959; Dyer 2001), these results lend support to the hypothesis that organized fire control facilitated the invasion of maples into the uplands from the more fire-protected lowlands (Abrams 1998).

The temporal patterns of fire history and maple establishment were similar at all four units at REMA and Zaleski. All units had periodic fires from ca. 1870 to 1925, and all units burned in both 1917 and 1923; in each of those years, fires were significant in three units. The initiation of maple establishment was similar in that none was documented in any units before the 1917 fires. Limited establishment was documented between the 1917 and 1923 fires, and large pulses occurred immediately after the 1923 fires followed by continuous establishment into the 1960s. The large pulses of maple establishment after the 1923 fires suggest resprouting from previously established individuals. Red maple, which accounted for 89% of the maple samples at REMA and Zaleski, has thin bark and is highly susceptible to top kill by fire (Harmon 1984; Regelbrugge and Smith 1994; Hutchinson et al. 2005); however, it also sprouts prolifically after topkill (Albrecht and McCarthy 2006; Blankenship and Arthur 2006). Maples probably began recruiting into these stands earlier than we document, perhaps much earlier, but presumably were being killed or top-killed until the cessation of fires.

The limited establishment and survival of maples before the 1923 fires (1917 to 1922) in all units may have resulted from several factors. Firstly, wildfires usually burn in a mosaic pattern, particularly in dissected landscapes, resulting in variable fire intensities and including unburned patches. Established maples may have escaped the 1923 fires in unburned patches. We also speculate that the initiation of maple establishment at REMA and Zaleski may have been facilitated by reduced anthropogenic land use, particularly by livestock in open-range woodland livestock grazing (Green 1907). The human population of Vinton County declined steadily with the demise of the iron furnace industry from a maximum of 17 223 in 1880 to 10 287 by 1930 (Vinton County Ohio Genealogy 2005). During the same period, farmland in the county decreased from 93283 to 61559 ha, and most of these lands reverted to forest (Bromley 1934a). Woodland grazing also likely decreased during this period, which would have favored the recruitment of trees, including maples. Brose and Waldrop (2006) showed that the cessation of livestock grazing in the Great Smoky Mountains National Park contributed to increased tree recruitment there in the 1920s and 1930s.

Oak also exhibited pulses of establishment immediately after some fires, suggesting resprouting from previously established stems. However, some fires were not followed by pulses of oak establishment. The final period of oak establishment occurred immediately after the 1923 fires; thereafter, we recorded virtually no additional oak stems. These data suggest that the large increase in maple establishment after the cessation of fires contributed via competition to the lack of subsequent oak recruitment. The absence of fire  after 1923, probably coupled with reduced woodland grazing, also likely facilitated the development of higher stand densities. The resulting closed-canopy conditions that developed would have greatly limited the ability of the relatively shade-intolerant oaks to establish from seed (Beck 1970) but not the shade-tolerant red maple, which can persist for long periods beneath a canopy (Tift and Fajvan 1999). Thus, further oak recruitment from seed, followed by growth and survival, probably was limited by a combination of shading and competition from both overstory trees and understory maples after fires ceased (e.g., Aldrich et al. 2005)

The history of fire and tree establishment at Tar Hollow differed from that of REMA and Zaleski in several aspects. First, at Tar Hollow, there were fewer historical fires (n = 5) and only one was significant. In all, we recorded only 19 historic fire scars at Tar Hollow compared with 48 at Zaleski and 75 at REMA.

The temporal pattern of maple establishment also differed, beginning nearly 40 years earlier (1881) and not exhibiting the distinct pulses after fire cessation that occurred at REMA and Zaleski. Tar Hollow also differed in that there was a long period of fairly continuous oak establishment (ca. 1890 to 1925), that coincided with the continuous recruitment of maples. These differences in fire and regeneration among sites may have resulted from different human land use.

Timber harvesting in the 1800s at Tar Hollow was not associated with charcoal production as at REMA and Zaleski and, thus, may have differed in intensity and extent. Perhaps more important is the evidence of greater and more varied human land use at the Tar Hollow site. In the 1930s, several Land Utilization Project (LUP) areas were established in which the State of Ohio purchased submarginal farmlands and then resettled the occupants (Bromley 1934a, 1934b). The REMA and Zaleski study sites were located near LUP areas while the Tar Hollow site was within the Ross-Hocking LUP. Land titles and appraisals that included detailed ownership and land-use maps from the time of purchase (ca. 1935) indicate that the Tar Hollow site consisted of a number of small parcels of mixed-ownership (ODNR, Division of Forestry, Regional Office, Chillicothe, Ohio). The maps indicate a patchy mixture of cover types: the most abundant was ‘‘forest land (including woodland pasture),’’ but it also included some areas of ‘‘grazing land (grazing or open pasture)’’ and a smaller portion as ‘‘crop land (including orchard and hay meadows).’’ The more varied land uses at Tar Hollow likely would have created a more patchy distribution of disturbances (fire, grazing, and harvesting). In particular, the patchy ownership and land use might have limited fire spread, resulting in the lower observed occurrence of fire. In turn, fewer fires probably facilitated the recruitment and survival of maples beginning much earlier at this site. At Tar Hollow, woodland grazing may have been more prevalent for a longer period with direct human occupation into the mid-1930s, potentially limiting the large pulses of maple establishment that occurred at REMA and Zaleski. Although fires generally were less frequent and wounded fewer trees at Tar Hollow, these units also developed into oak-dominated forests.

Other potential factors affecting tree establishment In addition to fire and human land use (particularly woodland grazing), the regeneration of oak forests was surely influenced by dramatic changes in wildlife populations. The decline and extirpation of the acorn-consuming white-tailed deer (Odocoileus virginianus (Zimmermann)), wild turkeys (Meleagris gallopavo L.), and passenger pigeons (Ectopistes migratorius (L.)) occurred from the mid-1800s to the early 1900s, as these stands were developing (Chapman 1938). Although these declines could have benefited oak regeneration from seed, the consumption of acorns and grazing of seedlings by domestic livestock probably were widespread during this period. No deer were present in Ohio from 1904 to 1922 when a restocking program was initiated (Chapman 1938). By 1938, it was estimated that only 2000 deer were in Ohio (Chapman 1938); the current estimate is 650 000 (ODNR, Division of Wildlife, Columbus, Ohio). Thus, excessive deer browsing clearly would not have contributed to the cessation of oak recruitment ca. 1925. In fact, a lack of deer browsing, the cessation of fire, and a decrease in livestock grazing may have facilitated the dramatic increase in maple establishment during the 1920s and 1930s.

It also is unlikely that American chestnut (Castanea dentata (Marsh.) Borkh.) mortality, caused by the chestnut blight fungus, facilitated the initial recruitment of maple in these stands. Even at REMA and Zaleski where maple recruitment began later (ca. 1920), it predated the arrival of chestnut blight, which caused mortality in Vinton County primarily from 1928 to 1936 (Beatley 1959). Also, chestnut accounted for only 4%-6% of witness trees ca. 1800 (Beatley 1959).

Selective harvesting was common after the mid-1930s on the Zaleski and Tar Hollow State Forests and at the REMA, then owned by D.B. Frampton and Co. (Beatley 1959). Previous work on sites near our REMA sites showed some growth releases suggestive of selective harvesting (Hutchinson et al. 2003). However, our data indicate that, in the absence of periodic fire, canopy disturbances after ca. 1925 did not facilitate oak recruitment. Similarly, during the fire control era, small openings in closed-canopy stands would have favored the growth of maples, which can respond with rapid growth even after long periods of suppression (Tift and Fajvan 1999).

Implications for oak regeneration today
Our results show the past importance of periodic fire in sustaining oak establishment and in limiting maple recruitment as stands developed. However, in many oak forests, there is now an abundance of maples in the midstory that are large enough to be fire resistant. Also, many forests that remain dominated by overstory oaks may be too dense to support oak regeneration even if the maple midstory could be removed with fire. Research has shown that simply returning low-intensity prescribed fires to fully stocked stands does not open the canopy sufficiently to improve the competitive status of oak regeneration (Hutchinson et al. 2005; Blankenship and Arthur 2006). Similar to the past importance of fire in early stand development, oak regeneration has improved when prescribed fire was applied to openstructured stands that developed after partial harvest (Kruger and Reich 1997; Brose and Van Lear 1998; Iverson et al. 2008). However, other studies have shown that the timing and intensity of the mechanical treatments and fire are critical to their success (Franklin et al. 2003; Albrecht and McCarthy 2006). Although fire was important in sustaining oak forests in the past, the legacy of prolonged fire exclusion necessitates research to refine oak regeneration prescriptions that incorporate canopy disturbances, fire, and other tools (Brose et al. 2006).

Acknowledgements

We thank David Hosack, Kristy Tucker, Brad Tucker, Bill Borovicka, Tim Fox, Joan Jolliff, and Zachary Traylor for field and laboratory assistance. We thank Patrick Brose, James Rentch, Ryan McEwan, Marty Jones, and two anonymous reviewers for providing many valuable suggestions on previous drafts of the manuscript. We thank the Ohio Department of Natural Resources, Division of Forestry, for supporting this research on Tar Hollow and Zaleski State Forests; we thank Bob Boyles and Michael Bowden of the Division of Forestry for assistance with historical documents. We also thank Forestland Group, LLC, for supporting this research on the Vinton Furnace Experimental Forest. This is publication No. 172 of the Fire and Fire Surrogate Network Project funded by the Join Fire Sciences Program.

References

Abrams, M.D. 1992. Fire and the development of oak forests. Bioscience, 42: 346-353. doi:10.2307/1311781.

Abrams, M.D. 1998. The red maple paradox: what explains the widespread expansion of red maple in eastern forest? Bioscience, 48: 355-364. doi:10.2307/1313374.

Abrams, M.D. 2002. The postglacial history of oak forests in eastern North America. In Oak forest ecosystems. Ecology and management for wildlife. Edited by W.J. McShea and W.M. Healy. Johns Hopkins University Press, Baltimore, Md. pp. 34-45.

Abrams, M.D., and Copenheaver, C.A. 1999. Temporal variation in species recruitment and dendroecology of an old-growth white oak forest in the Virginia Piedmont, USA. For. Ecol. Manage. 124: 275-284. doi:10.1016/S0378-1127(99)00071-7.

Abrams, M.D., and Downs, J.A. 1990. Successional replacement of old-growth white oak by mixed mesophytic hardwoods in southwestern Pennsylvania. Can. J. For. Res. 20: 1864-1870. doi:10. 1139/x90-250.

Albrecht, M.A., and McCarthy, B.C. 2006. Effects of prescribed fire and thinning on tree recruitment patterns in central hardwood forests. For. Ecol. Manage. 226: 88-103. doi:10.1016/j. foreco.2005.12.061.

Aldrich, P.R., Parker, G.R., Romero-Severson, J., and Michler, C.H. 2005. Confirmation of oak recruitment failure in Indiana old-growth forest: 75 years of data. For. Sci. 51: 406-416.

Beatley, J.C. 1959. The primeval forests of a periglacial area in the Allegheny Plateau (Vinton and Jackson Counties, Ohio). Bull. Ohio Biol. Surv. 1: 1-166.

Beck, D.E. 1970. Effect of competition on survival and height growth of northern red oak seedlings. USDA For. Serv. Southeast. For. Exp. Stn. Res. Pap. SE-5.

Blankenship, B.A., and Arthur, M.A. 2006. Stand structure over 9 years in burned and fire-excluded oak stands on the Cumberland Plateau, Kentucky. For. Ecol. Manage. 225: 134-145. doi:10.1016/j.foreco.2005.12.032.

Boerner, R.E.J., Brinkman, J.A., and Yaussy, D.A. 2007. Ecosystem restoration treatments affect soil physical and chemical properties in Appalachian mixed-oak forests. In Proceedings, 15th Can. J. For. Res. Vol. 38, 2008 Central Hardwood Forest Conference, 27 February – 1 March 2006, Knoxville, Tenn. [CD-ROM.] Edited by D.S. Buckley and W.S. Clatterbuck. USDA For. Serv. Gen. Tech. Rep. SRS-101. pp. 107-116.

Bromley, W.C. 1934a. Preliminary report on the Zaleski rehabilitation and forest project of Vinton County Ohio. Ohio Division of Forestry, Columbus, Ohio.

Bromley, W.C. 1934b. Preliminary report on the submarginal land and state forest project of southeastern Vinton County Ohio. Ohio Division of Forestry, Columbus, Ohio.

Brose, P.H., and Van Lear, D.H. 1998. Responses of hardwood advance regeneration to seasonal prescribed fires in oak-dominated shelterwood stands. Can. J. For. Res. 28: 331-339. doi:10.1139/ cjfr-28-3-331.

Brose, P.H., and Waldrop, T.A. 2006. Fire and the origin of table mountain pine – pitch pine communities in the southern Appalachian Mountains, USA. Can. J. For. Res. 36: 710-718. doi:10. 1139/x05-281.

Brose, P.H., Schuler, T.M., and Ward, J.S. 2006. Responses of oak and other hardwood regeneration to prescribed fire: what we know as of 2005. In Fire in Eastern Oak Forests: Delivering Science to Land Managers. Proceedings of a Conference, 15-17 November 2005, Columbus, Ohio. Edited by M.B. Dickinson. USDA For. Serv. Gen. Tech. Rep. NRS-P-1. pp. 123-135.

Chapman, F.B. 1938. The development and utilization of the wildlife resources of unglaciated Ohio. Ph.D. dissertation, Ohio State University, Columbus, Ohio.

Cook, E.R., and Kairiukstis, L.A. 1990. Methods of dendrochronology. Kluwer Academic Publications, Hingham, Mass.

Cutter, B.E., and Guyette, R.P. 1994. Fire frequency on an oakhickory ridgetop in the Missouri Ozarks. Am. Midl. Nat. 132: 393-398. doi:10.2307/2426595.

Dyer, J.M. 2001. Using witness trees to assess forest change in southeastern Ohio. Can. J. For. Res. 31: 1708-1718. doi:10. 1139/cjfr-31-10-1708.

Franklin, S.B., Robertson, P.A., and Fralish, J.S. 2003. Prescribed burning effects on upland Quercus forest structure and function. For. Ecol. Manage. 184: 315-335. doi:10.1016/S0378-1127(03) 00153-1.

Fritts, H.C. 1976. Tree rings and climate. Academic Press, London. Goebel, P.C., and Hix, D.M. 1997. Changes in the composition and structure of mixed-oak, second-growth forest ecosystems during the understory reinitiation stage of stand development. Ecoscience, 4: 327-340.

Gordon, R.B. 1969. The natural vegetation of Ohio in pioneer days. Ohio State University, Columbus, Ohio. Bull. Ohio Biol. Surv. New Ser. 3. pp. 1-109.

Green, W.J. 1907. First report on forest conditions in Ohio. Bull. Ohio Agric. Exp. Stn. 188: 37-51.

Griffith, D.M., DiGiovanni, D.M., Witzel, T.L., and Wharton, E.H. 1993. Forest statistics for Ohio, 1991. USDA For. Serv. Northeast. For. Range Exp. Stn. Res. Bull. NE-128.

Grissino-Mayer, H.D. 2001a. FHX2—software for analyzing temporal and spatial patterns in fire regimes from tree rings. TreeRing Res. 57: 115-124.

Grissino-Mayer, H.D. 2001b. Evaluating crossdating accuracy: a manual and tutorial for the computer program COFECHA. Tree-Ring Res. 57: 205-221.

Grissino-Mayer, H.D. 2004. FHX2 user’s manual: software for analyzing temporal and spatial patterns in fire regimes from tree rings. University of Tennessee, Knoxville, Tenn.

Grissino-Mayer, H.D., and Swetnam, T.W. 1997. Multi-century history of wildfire in the ponderosa pine forests of El Malpais National Monument. Bull. N.M. Bur. Mines Mineral Resour. 156: 163-172.

Grissino-Mayer, H.D., Romme, W.H., Floyd, M.L., and Hanna, D.D. 2004. Climatic and human influences on fire regimes of the southern San Juan Mountains, Colorado, USA. Ecology, 85: 1708-1724. doi:10.1890/02-0425.

Guyette, R.P., and Spetich, M.A. 2003. Fire history of oak-pine forests in the Lower Boston Mountains, Arkansas, USA. For. Ecol. Manage. 180: 463-474. doi:10.1016/S0378-1127(02) 00613-8.

Guyette, R.P., and Stambaugh, M.C. 2004. Post-oak fire scars as a function of diameter, growth, and tree age. For. Ecol. Manage. 198: 183-192. doi:10.1016/j.foreco.2004.04.016.

Guyette, R.P., Muzika, R.M., and Dey, D.C. 2002. Dynamics of an anthropogenic fire regime. Ecosystems (NY., Print), 5: 472-486. doi:10.1007/s10021-002-0115-7.

Guyette, R.P., Dey, D.C., and Stambaugh, M.C. 2003. Fire and human history of a barren-forest mosaic in southern Indiana. Am. Midl. Nat. 149: 21-34. doi:10.1674/0003-0031(2003) 149[0021:FAHHOA]2.0.CO;2.

Guyette, R.P., Spetich, M.A., and Stambaugh, M.C. 2006. Historic fire regime dynamics and forcing factors in the Boston Mountains, Arkansas, USA. For. Ecol. Manage. 234: 293-304. doi:10.1016/j.foreco.2006.07.016.

Haines, D.A., Johnson, V.J., and Main, W.A. 1975. Wildfire atlas of the northeastern and north central states. USDA For. Serv. Gen. Tech. Rep. NC-16.

Harmon, M.E. 1984. Survival of trees after low-intensity surface fires in Great Smoky Mountains National Park. Ecology, 65: 796-802. doi:10.2307/1938052.

Holmes, R.L. 1983. Computer-assisted quality control in tree-ring dating and measurement. Tree-Ring Bull. 43: 69-78.

Holmes, R.L. 1999. Program JOLTS: finding growth surges or suppression in trees. Laboratory of Tree-Ring Research, University of Arizona, Tucson, Ariz.

Hutchinson, T.F., Rubino, D., McCarthy, B.C., and Sutherland, E.K. 2003. History of forests and land use. In Characteristics of mixed-oak forest ecosystems in southern Ohio prior to the reintroduction of fire. Edited by E.K. Sutherland and T.F. Hutchinson. USDA For. Serv. Gen. Tech. Rep. NE-299. pp 17-27.

Hutchinson, T.F., Sutherland, E.K., and Yaussy, D.A. 2005. Effects of repeated fires on the structure, composition, and regeneration of mixed-oak forests in Ohio. For. Ecol. Manage. 218: 210-228. doi:10.1016/j.foreco.2005.07.011.

Iverson, L.R., Hutchinson, T.F., Prasad, A., and Peters, M.P. 2008. Thinning, fire, and oak regeneration across a heterogeneous landscape in the eastern U.S.: seven-year results. For. Ecol. Manage. In press.  oi:10.1016/j.foreco.2007.09.088.

Johnson, P.S., Shifley, S.R., and Rogers, R. 2002. The ecology and silviculture of oaks. CAB International, New York.
Kruger, E.L., and Reich, P.B. 1997. Responses of hardwood regeneration to fire in mesic forest openings. I. Post-fire communitydynamics. Can. J. For. Res. 27: 1822-1831. doi:10.1139/cjfr-27- 11-1822.

Leete, B.E. 1938. Forest fires in Ohio 1923 to 1935. Bull. Ohio Agric. Exp. Stn. 598.

Lorimer, C.G. 1993. Causes of the oak regeneration problem. In Oak Regeneration: Serious Problems, Practical Recommendations, Symposium Proceedings, 8-10 September 1992, Knoxville, Tenn. Edited by D. Loftis and C.E. McGee. USDA For. Serv. Gen. Tech. Rep. GTR-SE-84. pp. 14-39.

Lorimer, C.G. 1984. Development of the red maple understory in northeastern oak forests. For. Sci. 30: 3-22.

Lorimer, C.G., and Frelich, L.E. 1989. A methodology for estimating canopy disturbance frequency and intensity in dense temperate forests. Can. J. For. Res. 19: 651-663. doi:10.1139/X89-102.

Lorimer, C.G., Dahir, S.E., and Singer, M.T. 1999. Frequency of partial and missing rings in Acer saccharum in relation to canopy position and growth rate. Plant Ecol. 143: 189-202. doi:10.1023/A:1009847819158.

McEwan, R.W., Hutchinson, T.F., Ford, R.D., and McCarthy, B.C.
2007a. An experimental evaluation of fire history reconstruction using dendrochronology in white oak (Quercus alba). Can. J. For. Res. 37: 806-816. doi:10.1139/X06-294.

McEwan, R.W., Hutchinson, T.F., Long, R.P., Ford, R.D., and McCarthy, B.C. 2007b. Temporal and spatial patterns of fire occurrence during the establishment of mixed-oak forests in eastern North America. J. Veg. Sci. 18: 655-664. doi:10.1658/1100 9233(2007)18[655:TASPIF]2.0.CO;2.

McNab, W.H., and Avers, P.E. 1994. Ecological subregions of the United States: section descriptions. USDA For. Serv. Wash. Off. Admin. Publ. WO-WSA-5. Ohio Experiment Station. 1922. 42nd annual report. Bull. Ohio Ag-
ric. Exp. Stn. 373.

Phipps, R.L. 1961. Analysis of five years of dendrometer data obtained within three deciduous forest communities of Neotoma. Neotoma Ecological and Bioclimatic Laboratory, Ohio Agricultural Experiment Station, Wooster, Ohio. Res. Circ. 105. pp. 3-34.

Regelbrugge, J.C., and Smith, D.W. 1994. Postfire tree mortality in relation to wildfire severity in mixed oak forests in the Blue Ridge of Virginia. North. J. Appl. For. 11: 90-97.

Schuler, T.M., and McClain, W.R. 2003. Fire history of a Ridge and Valley forest. USDA For. Serv. Northeast. Res. Stn. Res. Pap. NE-724.

Shumway, D.L., Abrams, M.D., and Ruffner, C.M. 2001. A 400 year history of fire and oak recruitment in an old-growth oak forest in western Maryland, USA. Can. J. For. Res. 31: 1437 1443. doi:10.1139/cjfr-31-8-1437.

Smith, K.T., and Sutherland, E.K. 1999. Fire-scar formation and compartmentalization in oak. Can. J. For. Res. 29: 166-171. doi:10.1139/cjfr-29-2-166.

Soucy, R.D., Heitzman, E., and Spetich, M.A. 2005. The establishment and development of oak forests in the Ozark Mountains of Arkansas. Can. J. For. Res. 35: 1790-1797. doi:10.1139/x05 104.

Stokes, M.A., and Smiley, T.L. 1968. An introduction to tree-ring dating. University of Chicago Press, Chicago, Ill.

Stout, W. 1933. The charcoal iron industry of the Hanging Rock Iron District—its influence on the development of the Ohio Valley. Ohio Archaeol. Hist. Q. 42: 72-104.

Sutherland, E.K. 1997. The history of fire in a southern Ohio second-growth mixed-oak forest. In Proceedings of the 11th Central Hardwoods Forest Conference, 23-26 March 1997, Columbia, Mo. Edited by S.G. Pallardy, R.A. Cecich, H.E. Garrett, and P.S. Johnson. USDA For. Serv. Gen. Tech. Rep. NC-188. pp. 172-183.

Sutherland, E.K., Hutchinson, T.F., and Yaussy, D.A. 2003. Introduction, study area description, and experimental design. In Characteristics of mixed-oak forest ecosystems in southern Ohio prior to the reintroduction of fire. Edited by E.K. Sutherland and T.F. Hutchinson. USDA For. Serv. Gen. Tech. Rep. NE-299, pp. 1-16.

Tift, B.D., and Fajvan, M.A. 1999. Red maple dynamics in Appalachian hardwood stands in West Virginia. Can. J. For. Res. 29: 157-165. doi:10.1139/cjfr-29-2-157.

Van Lear, D.H., and Brose, P.H. 2002. Fire and oak management.  In Oak forest ecosystems. ecology and management for wildlife. Edited by W.J. McShea and W.H. Healy. Johns Hopkins University Press, Baltimore, Md. pp. 269-279.

Vinton County Ohio Genealogy. 2005. Vinton County Ohio population changes 1850-1990 [online]. Available from www.rootsweb.com/~ohvinton/vintpop.gif.

Wendel, G.W., and Smith, H.C. 1986. Effects of prescribed fire in a central Appalachian oak-hickory stand. USDA For. Serv. Northeast. For. Range Exp. Stn. Res. Pap. NE-RP-594.

Yaussy, D.A., Hutchinson, T.F., and Sutherland, E.K. 2003. Structure, composition, and condition of overstory trees. In Characteristics of mixed-oak forest ecosystems in southern Ohio prior to the reintroduction of fire. Edited by E.K. Sutherland and T.F. Hutchinson. USDA For. Serv. Gen. Tech. Rep. NE-299. pp. 99-111.

Managing forests with prescribed fire: Implications for a cavity-dwelling bat species

Justin G. Boylesa, Doug P. Aubreyb,*

a Department of Ecology and Organismal Biology, Indiana State University, Terre Haute, 47809, USA
b USDA Forest Service, Savannah River, P.O. Box 700, New Ellenton, SC 29809, USA
Received 1 June 2005; received in revised form 27 September 2005; accepted 30 September 2005


Abstract

Prescribed burning is used as a restoration and management technique in many deciduous forests of eastern North America. The effects of fire have been studied on habitat selection of many vertebrate species, but no studies have reported the effect of fire on bat roosting habitat. Fire initially leads to an influx of dead and dying trees, an increase of light availability, and a decrease of canopy and sub-canopy tree density. These characteristics are beneficial to many forest-dwelling vertebrates including cavity-roosting bats. We evaluated evening bat (Nycticeius humeralis) roost-site selection at the stand-scale in order to determine roosting preferences as they relate to prescribed burning. Standard radiotelemetry techniques were used to locate evening bat roost trees. Canopy light penetration and overstory tree density were measured in both burned and unburned forests. Sixty-three trees used as roosts by both male and female evening bats were located during both the summer and winter and all 63 roosts were located in the burned portion of the study area. Canopy light penetration was higher and canopy tree density was lower in the burned forest than unburned forest. An increase in light availability may release bats from one of the constraints suggested for many forest-dwelling bat species in roost tree selection—sun-exposure. This should increase the abundance of trees with characteristics suitable for roosting and may allow bats to roost throughout the interior of the forest as opposed to only on forest edges, thereby allowing bats to roost closer to foraging grounds and possibly lessening predation rates. Lower tree density may allow for ease of flight within the forest as well as more efficient locating of roost trees. In addition, there were a significantly higher proportion of dead trees, which evening bats commonly use as roost trees, in burned forests compared to unburned forests. Prescribed burning appears to initially lead to creation or restoration of favorable cavity-dwelling bat habitat and its continual implementation perpetuates an open sub-canopy. Therefore, we suggest that prescribed burning may be a suitable tool for management of roosting habitat for cavity-roosting bats.

# 2005 Elsevier B.V. All rights reserved.

Keywords: Fire ecology; Forest restoration; Nycticeius humeralis; Oak-hickory forest; Prescribed burning; Roost selection

CLICK HERE TO DOWNLOAD AS PDF


1. Introduction

Fire is critical in regulating and maintaining many forest ecosystems (Huddle and Pallardy, 1996; van Lear, 2002). In particular, much of the western portion of North America’s eastern deciduous forest is thought to have been shaped and maintained through fires set by Native Americans prior to European settlement (Cottam, 1949; Ladd, 1991; Pyne, 1982). Prescribed burning has been shown to alter many characteristics of forest habitat potentially affecting forest-dwelling bats including tree mortality, which increases available trees commonly used by wildlife (Huddle and Pallardy, 1996; Arthur et al., 1998; Hartman and Heumann, 2003; Aubrey, 2004); pathogen susceptibility, which expedites cavity formation (Burns, 1955; Paulsell, 1957; Smith and Sutherland, 2001); and canopy light penetration (Anderson and Brown, 1986; McCarty, 1998; Aubrey, 2004), which is known to affect roost suitability (Kurta et al., 1993; Brigham et al., 1997).


* Corresponding author. Tel.: +1 803 725 1758; fax: +1 803 725 0311.
E-mail address: daubrey@fs.fed.us (D.P. Aubrey).

0378-1127/$ – see front matter # 2005 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2005.09.024


Many forests are managed for timber harvest by the use of mechanical thinning or clear-cutting. Prescribed fire differs from these management techniques because fire is generally used to maintain or restore natural forest ecosystems and reduce understory competition, while other silvicultural methods are predominately used to maximize harvest yield and quality. When first implemented, prescribed burning decreases overstory tree density and basal area (Anderson and Brown, 1986; Peterson and Reich, 2001). Following this initial fire with frequent  burns generally reduces fuel accumulation and subsequent burns are therefore relatively cool and somewhat nonuniform (Ladd, 1991). However many fire-sensitive seedlings and saplings are generally eliminated or prevented from regenerating (Lorimer, 1985; Ladd, 1991; Moser et al., 1996; Brose and van Lear, 1997; Barnes and van Lear, 1998). Fire-tolerant trees remain as overstory dominants and canopy recruitment coincides with periods of decreased burn frequency or intensity (Crow et al.,1994). For example, oaks possess thick bark which insulates them from heat associated with burning and are capable of rapid compartmentalization when damaged by fire which inhibits fungal infection and makes mature individuals relatively tolerant of fire (Lorimer, 1985; Abrams, 1985; Crow, 1988; Stearns, 1991; Smith and Sutherland, 1999; Peterson and Reich, 2001). Much of the western portion of the eastern deciduous forest is believed to have resembled the structure and composition of a woodland (i.e. moderate to low canopy coverage) more than a forest (i.e. high canopy coverage) prior to European settlement, which is largely attributed to fire (Cottam, 1949; Curtis, 1959; Nuzzo, 1986; Ladd, 1991). Therefore, managing forests with prescribed fire reintroduces a historic disturbance process that other silvicultural techniques lack, and may provide a more heterogeneous habitat for wildlife that is similar to forested areas prior to wide-spread fire suppression by European settlers. Furthermore, prescribed burning has become increasingly common over the past few decades, as land managers have seen the effects of fire suppression. Specifically, fire has been most commonly reintroduced to systems where conservation of biodiversity and restoration of historic ecosystem processes are paramount.

Numerous studies have focused on vertebrates in forests maintained by prescribed burning (e.g. reptiles, McLeod and Gates, 1998; small mammals, Simon et al., 2002; amphibians, Schurbon and Fauth, 2003; birds, Blake, 2005). However, to our knowledge, there are no studies reporting habitat selection of tree-dwelling bats in burned forests, although the need for such studies has been suggested (Menzel et al., 2001b; Carter et al., 2002). Furthermore, the effects of forest management on roosting habitat of bats are not clear and it has been suggested that wildfires and prescribed burning may have detrimental (Chambers et al., 2002) or beneficial (Carter et al., 2002) impacts on bats. For some bat species, such as the eastern red bat (Lasiurus borealis), fire may pose a direct threat to survival as early spring burns remove litter where occasional winter roosting occurs and can potentially scorch hibernating individuals (Moorman et al., 1999; Rodrigue et al., 2001). Bats roosting in snags (standing dead trees) are also susceptible to direct disturbance from fire if roost trees ignite (Rodrigue et al., 2001), but this is probably not a major cause of mortality unless the roost burns quickly and the bat is in deep torpor (Carter et al., 2002). Availability of suitable roosts likely influences habitat selection (Kunz, 1982) and roosting sites are thought to be of critical importance to conservation of many bat species (Fenton, 1997). Fire generally removes understory saplings and may lead to the creation of suitable roost trees for cavity-roosting bats. Fire may also remove standing snags if fuel loading is high. However, fuel should not accumulate directly under a snag when burning is frequent so snags should persist under these circumstances. As fire increases the heterogeneity of forest structure and composition (Fule et al., 2004), there should be an increase in the diversity of available roosting habitat.

Previous studies have focused on activity (Humes et al., 1999; Patriquin and Barclay, 2003; Tibbels and Kurta, 2003; Mazurek and Zielinski, 2004), demographic parameters (Miller, 2003), and roosting preference (Campbell et al., 1996; Menzel et al., 2002; Elmore et al., 2004) of bats in heavily managed forests, but have not examined fire-based management. With the exception of a few anecdotal reports of L. borealis being driven from their leaf litter  ibernacula during winter burns (Moorman et al., 1999; Rodrigue et al., 2001), there is little empirical data about how fire affects forest dwelling bats.

The objective of this study was to determine evening bat (Nycticeius humeralis) roost-site selection at the stand-scale in a forest heavily managed by fire and to understand those characteristics of the forest that may influence roost tree selection. The evening bat is a locally abundant cavity-dwelling bat species found throughout much of the southeastern United States, but it may be declining in parts of its range (Whitaker et al., 2002; Whitaker and Gummer, 2003). Evening bats are known to roost in large numbers in man-made structures (Watkins, 1969; Watkins and Shump, 1981; Bain and Humphrey, 1986; Wilkinson, 1992) and tree cavities (Wilk-inson, 1992; Bowles et al., 1996; Menzel et al., 1999, 2001a; Boyles et al., 2003; Boyles and Robbins, in press). Evening bats roost mainly in cavities in trees of various stages of decay (Bowles et al., 1996; Boyles and Robbins, in press), but relatively little is known about the formation or microclimate of the roosts. During the summer, evening bats roost mainly in large dead trees, but during the winter, live trees are commonly used (Boyles and Robbins, in press). They have a short wingspan and high wing loading so it has been predicted that they are not highly maneuverable relative to other bat species (Norberg and Rayner, 1987). Due to relatively inefficient flight, this species may avoid long foraging trips (Norberg and Rayner, 1987) and it has been suggested that evening bats forage close to their roosting areas  Duchamp et al., 2004). Evening bats feed heavily on coleopterans, homopterans, and hemipterans (Whitaker, 2004). As with other tree roosting bat species, they spend over half of its time each day in roosts (Brigham et al., 1997), so conserving roosts is important in managing this species. Evening bats are commonly referred to as migratory in middle latitudes (Jones et al., 1967; Humphrey and Cope, 1968; Watkins, 1969; Wilkinson, 1992; Sparks et al., 1999; Geluso et al., 2004), but it appears that the population discussed herein is largely non-migratory (Boyles et al., 2003; Boyles and Robbins, in press) so roost trees refer to trees used throughout the year.

It was predicted that evening bats would prefer roosting in burned forests because of an increase in the density of dead and dying trees, an increase in light penetration and an overall decrease in overstory and understory tree  ensity. These characteristics should benefit forest-dwelling bats and promote use of burned forests as roosting habitat. This study compliments previous work (Boyles and Robbins, in press), which reports the characteristics of roost trees and the surrounding habitat used by this population of evening bats during both the summer and winter. The results presented herein focus on roost selection at the forest stand-level.


2. Methods

2.1. Study area

We conducted this research on the Drury Conservation Area (DCA) in Taney County, Missouri (UTM 40.47.000N, 4.93.000E). DCA is a 1200 ha area located in extreme southwestern Missouri in the Ozark Mountains Region and is bordered on two sides by Bull Shoals Lake. It is actively managed by the Missouri Department of Conservation, which has implemented prescribed burning on approximately 55% of the potential area available for roosting habitat in an attempt to restore historic glades and oak-hickory woodlands and reduce red cedar (Juniperus virginiana) concentrations. Burning was initiated in 1999 after nearly 50 years of fire suppression and the area was then burned on a biennial schedule. All burning was conducted in March or April. The initial 1999 burn was probably more intense than subsequent burns because of accumulated fuels; therefore more overstory tree mortality may have occurred during or as a result of the initial burn (Aubrey, 2004).

Approximately 60% of DCA is dominated by oak-hickory forest with the remainder of the area being glades, wildlife food plots, ponds, and riparian areas (Missouri Department of Conservation, 1991). Elevation ranges from 185 to 335 m on DCA. Several gravel roads facilitate access to the interior of much of the forest and one large gravel road serves as the firebreak between burned and unburned portions of the forest. The canopy of the area consists almost entirely of deciduous trees from the white and red oak groups (Quercus spp.), hickories (Carya spp.), elms (Ulmus spp.), and ashes (Fraxinus spp.).

2.2. Location of roosting sites

Bats were captured from March 2003 to March 2004 using mist nets (Avinet, Dryden, NY, USA) of various lengths (6, 9, 12, or 18 m) placed across ponds or forest roads. The majority of netting sites were on the gravel road that acts as a firebreak between the burned and unburned forest, but one pond and two roads in the burned forest and one pond and one creek bed in the unburned forest were also netted. Approximately 55% of the available roosting area for evening bats was located in an area treated with prescribed fire. Thus, if evening bats were roosting at random, we would expect 55% of roost trees to be located in burned forest and 45% of roost trees to be located in unburned forest. Twenty-three evening bats were fitted with radio- transmitters (0.52 g; Model LB-2N, Holohil, Carp, Canada) by clipping fur to the skin in the interscapular region and affixing the transmitter with surgical adhesive (Skin Bond, Smith and Nephew Inc., Largo, FL, USA). The range of weights of bats fitted with transmitters was 7.5-13.5 g; therefore the transmitter represented 3.9-6.9% of the individual’s body mass.

Each bat’s roost tree was located every day following attachment of the transmitter and tracking was discontinued when the transmitter expired, was shed by the bat, or the bat remained outside of the study area for more than 5 days. This study was conducted year-round so roost trees include those used during both summer and winter by males and females in all reproductive classes (pregnant, lactating, post-lactating, and non-reproductive). All animal handling methods follow guidelines of the American Society of Mammalogists (Animal Care and Use Committee, 1998).

2.3. Canopy light penetration sampling

Leaf area index (LAI) was used as a measure of canopy light penetration to determine if three biennial prescribed burns had resulted in a more open canopy. LAI is an estimate of the ratio of overstory leaf area relative to ground area. A LAI of 0 indicates complete canopy light penetration whereas a LAI of 12 indicates no canopy light penetration (Hyer and Goetz, 2004). Here we define canopy light penetration (CLP) as: 12 LAI.

Two blocks each containing three transects in burned and unburned forests were monitored throughout the 2003 growing season. Unburned and burned forests were adjacent to one another with a gravel road acting as a fire buffer between them. LAI was obtained indirectly at each transect using an AccuPAR PAR-80 light interception device (Decagon Devices Inc., Pullman, WA, USA). Individual light measurements were collected 1.2 m above ground-level at five randomly spaced points along each transect. These five measurements were then averaged to calculate one LAI value, and therefore, one CLP value for each transect per sample period. Measurements were collected monthly throughout the 2003 growing season (April through September). The majority of canopy trees were deciduous species; thus, CLP was high in both burned and unburned forests during winter and was not measured from October through March.

2.4. Tree density

Canopy tree density was estimated in 0.05 ha circular plots centered on 25 randomly selected trees in each of the burn treatments. A random number generator was used to select UTM coordinates for the trees. Coordinates were located using a GPS unit (eTrex, Garmin International Inc., Olathe, Kansas) and selected the tree nearest that point to serve as the center of the plot. All trees in the plot greater than 10 cm diameter at breast height (dbh) were considered overstory trees and were counted and classified as either live or dead.

2.5. Statistical analysis

A binomial probability distribution was used to determine if bats roosted at random or at a proportion different from what would be expected given the amount of burned and unburned forest available. The effect of prescribed fire on CLP was assessed using a multi-factorial analysis of variance (ANOVA). The experiment was a nested block design with repeated measures. Month (April-September) and habitat (burned forest and unburned forest) were treated as fixed factors. Block (n = 2) and transect (n = 12) were treated as random factors, with transect nested under treatment. Mean separations were performed using a post hoc Tukey test. Tree density was analyzed using a t-test. In addition to absolute densities, the arcsine of the proportion of dead trees compared to total trees was calculated and analyzed using a t-test (Zar, 1984). All statistical analyses were conducted in Minitab 14. Alpha is 0.05 for all analyses.


3. Results

3.1. Roost tree location

Fig. 1. Seasonal trends (mean S.E.) in canopy light penetration (CLP) calculated as 12 leaf area index in burned and unburned forest on Drury Sixty-three roost trees were used by 23 (11 females and 12 males) evening bats from 9 March 2003 to 31 March 2004. All 63 roost trees were located in the portion of the area that was subjected to prescribed burning, although many of the bats were captured on the road that serves as the break between burned and unburned forest or within 200 m of that road. Bats roosted in the burned forest exclusively and significantly more often than expected if they selected burned or unburned forest randomly (P < 0.001). In addition, nearly all the trees used as roosts were located more than 50 m from any forest edge (62 of 63). The one exception was a large white oak Quercus alba (L.) used by a male evening bat in October that was located at the forest edge less than 50 m from where that individual was captured. This tree was not used the first day after capture; it was used on the second and fourth days, so it appears that this tree was actively selected and was not used in response to stress caused by handling.

3.2. Canopy light penetration

Both the main effects of treatment (P < 0.001) and month (P < 0.001) as well as their interaction (P < 0.001) signifi-cantly affected CLP. Maximum leaf expansion occurred in May for the burned forest and July in the unburned forest. Averaged over the 6-month sample period, canopy light penetration was significantly greater in burned forest as compared to unburned forest (P < 0.001). CLP was high in both the burned and unburned forests in April (Fig. 1). Pairwise comparisons suggest that there was no difference in canopy light penetration between burned and unburned forests in April or May but differences were significant throughout the rest of the growing season (June-September, P < 0.05 in all comparisons). As expected, measurements collected prior to leaf expansion (April) suggest that there was no difference in CLP between the burned and unburned forests; therefore, there was likely no difference in light availability between the treatments during the winter.

Fig. 1.  Seasonal  trends  (mean    S.E.)  in  canopy  light  penetration  (CLP) calculated as 12    leaf area index in burned and unburned forest on Drury Conservation Area, Taney County, Missouri during the 2003 growing season. CLP, as defined herein, is measured on a scale of 0-12, with 12 indicating complete light penetration and 0 indicating no light penetration. All measurements were collected mid-month. Asterisks indicate significant differences (at alpha = 0.05) between burned and unburned forests in each month

3.3. Tree density

Mean overstory tree density per hectare was significantly higher in unburned forest (612.0 25.0) relative to burned Conservation Area, Taney County, Missouri during the 2003 growing season. CLP, as defined herein, is measured on a scale of 0-12, with 12 indicating complete light penetration and 0 indicating no light penetration. All measurements were collected mid-month. Asterisks indicate significant differences (at alpha = 0.05) between burned and unburned forests in each month.

 


forest (513.6 24.2; t = 2.83, P = 0.007) and live tree density was significantly higher in unburned forest (576.0 24.6) than burned forest (468.0 24.1; t = 3.13, P = 0.003). The density of dead trees was not significantly different between the burned (45.6 7.3) and unburned forests (36.0 4.3; t = 1.14, P = 0.263). However, the proportion of dead trees compared to total trees was significantly higher in burned forest (0.092 0.014) relative to unburned forest (0.060 0.007; t = 2.08, P = 0.045).

 


4. Discussion

In our study area, evening bats showed a strong preference for forests managed with prescribed fires. Although there are no data on roost tree selection by evening bats before prescribed burning was initiated, it is likely that the forest characteristics were similar between what are now burned and unburned forests (Aubrey, 2004). All roost trees used by both males and females throughout the year were in the burned portion of the study area. For at least a few years after the initial burn, prescribed fire enhances roosting habitat for evening bats in several ways. First, burning increases the abundance of dead trees in some forests (Burns, 1955; Paulsell, 1957; Huddle and Pallardy, 1996; Peterson and Reich, 2001; Fule et al., 2004) and therefore increases the number of tree cavities formed by decay. We did not directly test the effect of fire on overstory tree mortality; however, dead trees were found in higher densities and at significantly higher proportions in the burned forest. Initial burns with high fuel accumulation can result in the formation of dead trees (Paulsell, 1957; Scowcroft, 1966; Anderson and Brown, 1986; White, 1986; Peterson and Reich, 2001) and leads to a forest with a large number of trees in various stages of decay. Ambient temperatures vary widely during the winter in southwestern Missouri so a large number of trees in varying stages of decay may provide sufficient options for evening bats to meet thermoregulatory needs simply by roost-switching. It has also been suggested that fire-scars allow entry for fungal pathogens, which can lead to heartrot and possibly mortality, which should promote cavity formation (Smith and Sutherland, 2001; Parsons et al., 2003). This population of evening bats is known to roost in trees in all decay stages (Boyles and Robbins, in press) and frequent burning should increase options for roost tree selection.

Second, CLP is significantly higher in burned forest than the unburned forest during the growing season. Although evening bats are not known to preferentially roost on forest edges, many tree-dwelling bats are thought to select roost trees that receive high levels of sun-exposure during the summer months (Kurta et al., 1993); therefore, trees selected as roosts are commonly taller than the canopy (Vonhof and Barclay, 1996; Britzke et al., 2003), near forest edges (Grindal, 1999) or areas with an open canopy (Menzel et al., 2001a). High light exposure is known to improve roost suitability (Kurta et al., 1993; Brigham et al., 1997) so high light penetration into the interior forest will allow bats to roost in trees that would not receive adequate sunexposure to facilitate thermoregulation in unburned forests. The ability to roost in interior forest trees that would not receive adequate sun-exposure in unburned forest may yield many benefits for evening bats. For example, the opportunity to use interior forest trees as roosts increases the availability of suitable roost trees in the habitat relative to populations that are forced to roost near forest edges or openings. This increase in suitable roost trees may allow bats to roost closer to their preferred foraging area and therefore reduce commuting distance and energy expenditure. Predation by meso-carnivores (Sparks et al., 2003) should also be lessened because the roosting sites are not concentrated in small forest patches or along forest edges, which may be easily found by and accessible to predators. An abundance of suitable roost trees may also facilitate frequent roost switching, which in turn may lessen ecto-parasite loads (Lewis, 1995).

Finally, prescribed burning may improve foraging habitat, thereby encouraging bats to roost in the vicinity. High fire frequency of low to moderate intensity prevents the regeneration of a sapling layer if preceded by a high intensity fire that removes this layer (Peterson and Reich, 2001). Exclusion of this stratum will reduce obstructions and make navigation easier in burned forest compared to unburned forest (Boyles and Robbins, in press). Evening bats forage in open areas and wooded habitats (Duchamp et al., 2004) and it has been noted that activity is higher for some bat communities in thinned forest compared to unthinned forest (Humes et al., 1999). It has been predicted that evening bats are not highly maneuverable and have relatively inefficient flight (Norberg and Rayner, 1987). This may prohibit evening bats from roosting in unburned areas because of dense understory that would make straight flight difficult. Other studies have also noted a preference for roosting in areas with a more open understory (Castleberry et al., 2005). Previous work has suggested that evening bats forage close to their roosting areas (Duchamp et al., 2004); therefore roost selection in burned forest may simply be an artifact of selection of burned habitat for foraging. In addition, the major food source of evening bats, coleopterans (Whitaker, 2004), have been found at both higher abundances and greater species richness in burned forests than unburned forests (Saint-Germain et al., 2004). Although it is difficult to distinguish if roosting habitat or foraging habitat is more important in observed habitat selection, anecdotally there does appear to be a close association between the two in this population of evening bats.

It is unlikely that any one of these reasons alone can adequately explain the preferential habitat selection seen in evening bats. Combinations of any or all of these broad forest characteristics may contribute to suitable habitat for this species and each may be of fluctuating importance throughout the year. For example, the difference in light penetration between burned and unburned forest is probably only important during the summer months. Because of the deciduous nature of the majority of the trees in southwestern Missouri, the difference in light penetration between burned and unburned forest will be lessened during the winter. Therefore, it is likely that evening bats roosted exclusively on the burned portion of the area during the winter because of prey availability, lessened clutter, or simple roost fidelity. However, it should be noted that many tree-dwelling bat species in the eastern United States only roost in trees during the summer months, so this may be inconsequential in reference to those species.


5. Conclusions

Prescribed burning has the potential to provide habitat for evening bats and possibly other species and should be taken into consideration when constructing management plans for these species. Fire may initially increase both the quantity and quality of roosting habitat for evening bats by creating an influx of dead and dying trees as well as facilitating disease and decay in live trees. This study deals only with the first 6 years following the initial burn, and long-term effects of burning may not be as beneficial to evening bats. For example, frequent burning may lead to a decline in roost trees by felling suitable snags and creating a canopy of fire-tolerant species that are not generally killed by fire (Curtis, 1959; Pyne, 1982; Crow, 1988; Ladd, 1991). However, anecdotal evidence suggests that snags persist through these low intensity burns, possibly because leaf litter fails to accumulate underneath a snag (personal observation). The same type of anecdotal evidence exists in southeastern long-leaf pine systems where biennial fire leaves snags intact because of reduced fuel accumulation near the base of the tree (S. Castleberry, University of Georgia, personal communication). To our knowledge, no studies have addressed the fate or residence time for snags suitable for roosting in a frequently burned system. Fire frequency should not remain a static part of management strategy. If biennial burns are continued, eventually the only trees left in the canopy will be fire-tolerant. It is  necessary to withhold fire from areas to allow a sufficient fuel source to accumulate, which will allow for highintensity fires that can cause canopy tree mortality and create snags. However, benefits of managing with prescribed fire may surpass those of mechanical thinning techniques because of the random and delayed mortality imposed on canopy trees (Loomis, 1974). Furthermore, prescribed burning is a more cost-effective management tool than mechanical methods (Wade and Lunsford, 1988; Dubois et al., 1999).

This study demonstrates the preference of evening bats to periodically burned forest, but fire can also be beneficial to other bat species that utilize trees as roosts. For example, the Indiana bat (Myotis sodalis), listed as federally endangered in the United States, is known to use a large number of dead and dying trees as roosts because these trees often offer the exfoliating bark that this species uses as a roost-site (Kurta et al., 1993). Through mortality and damage to bark, fire will increase the abundance of trees with exfoliating bark (Burns, 1955; Paulsell, 1957; Smith and Sutherland, 1999) and thereby increase available roosting habitat for species such as the Indiana bat. Other bat species could benefit from fire as a management tool and it should be investigated in other regions with other bat species.

Fire does have the potential to negatively affect forest dwelling bat habitat. For species that preferentially roost in trees in late stages of decay, frequent fire may destroy roost trees before they become suitable for roosting. Frequent fire may also cause the loss of winter habitat for litter-roosting species such as L. borealis (Moorman et al., 1999; Boyles et al., 2003) by continually removing accumulated leaf litter from the forest floor. Long-term studies are necessary to understand the patterns and processes of snag residence time in a periodically burned forest. Resource managers should consider the different habitat needs and life history traits of both tree-dwelling and litter roosting bats when creating management plans. Furthermore, forests respond differently according to the season when burning occurs. For example, summer burns have been shown to cause increased overstory tree mortality relative to burns conducted in other seasons (van Lear and Waldrop, 1991; Brose and van Lear, 1997). Fires in this season would therefore benefit tree-dwelling species as well as allow leaf litter to accumulate in fall benefiting litter-roosting species. More research examining the effect of burn frequency and season of burning on a variety of taxa is necessary to properly manage forests for biodiversity.

Acknowledgements

Funding for this project was provided in part by the Missouri Department of Conservation, the City of Springfield, MO, Dickerson Park Zoo, and Southwest Missouri State University. We would like to thank L. Robbins and D.A. Wait for assisting in obtaining funding and providing equipment for this project. The Missouri Department of Conservation and the Bull Shoals Field Station provided vehicles, equipment, and access to their property. J. Timpone, M. Milam, B. Mormann, P. Brown, and many others assisted in capturing and tracking bats and conducting vegetation analysis. M. Milam, M. McKnight, M. Coleman, J. Orrock, and anonymous reviewers provided many helpful comments on earlier versions of this manuscript.

References

Abrams, M.D., 1985. Fire history of oak gallery forests in northeast Kansas tallgrass prairie. Am. Midl. Nat. 114, 188-191.

Anderson, R.C., Brown, L.E., 1986. Stability and instability in plant communities following fire. Am. J. Bot. 73, 364-368.

Animal Care and Use Committee, 1998. Guidelines for the capture, handling, and care of mammals as approved by The American Society of Mammalogists. J. Mamm. 79, 1416-1431.

Arthur, M.A., Paratley, R.D., Blakenship, B.A., 1998. Single and repeated fires affect survival and regeneration of woody and herbaceous species in an oakpine forest. J. Torrey Bot. Soc. 125, 225-236.

Aubrey, D.A., 2004. Savanna restoration through prescribed fire: demographic and physiological responses of oak and hickory seedlings and saplings to a changing light environment. Master’s Thesis, Southwest Missouri State University.

Bain, J.R., Humphrey, S.R., 1986. Social organization and biased primary sex ratio of the evening bat Nycticeius humeralis. Fla. Sci. 49, 22-31.

Barnes, T.A., van Lear, D.H., 1998. Prescribed fire effects on advanced regeneration in mixed hardwood stands. South. J. Appl. For. 22, 138-142.

Blake, J.G., 2005. Effects of prescribed burning on distribution and abundance of birds in a closed-canopy oak-dominated forest, Missouri, USA. Biol. Conserv. 121, 519-531.

Bowles, J.B., Howell, D., Van Zee, J.W., Wilson, G.M., 1996. Use of alternate roost trees by the evening bat, Nycticeius humeralis, in Iowa. In: Genoways, H.H., Baker, R.J. (Eds.), Contributions in Mammalogy: A Memorial Volume Honoring Dr. J. Knox Jones, Jr. Museum of Texas Tech University, Lubbock, Texas, pp. 217-224.

Boyles, J.G., Timpone, J.C., Robbins, L.W., 2003. Late-winter observations of red bats, Lasiurus borealis, and evening bats, Nycticeius humeralis, in Missouri. Bat Res. News 44, 59-61.

Boyles, J.G., Robbins, L.W. Characteristics of summer and winter roost trees used by evening bats (Nycticeius humeralis) in southwestern Missouri. Am. Midl. Nat., in press.

Brigham, R.M., Vonhof, M.J., Barclay, R.M.R., Gwilliam, J.C., 1997. Roosting behavior and roost-site preferences of forest-dwelling California bats (Myotis californicus). J. Mamm. 78, 1231-1239.

Britzke, E.R., Harvey, M.J., Loeb, S.C., 2003. Indiana bat, Myotis sodalis, maternity roosts in the southern United States. Southeast. Nat. 2, 235 242.

Brose, P.H., van Lear, D.H., 1997. Effects of seasonal prescribed fires on density of hardwood advanced regeneration in oak-dominated shelterwood stands.

In: Proceedings of the 25th Annual Hardwood Symposium, ‘‘25 Years of Hardwood Silviculture: A Look Back and a Look Ahead’’, 7-10 May, 1997, Cashiers, NC. National Hardwood Lumber Association, Memphis, TN, pp.
139-146.

Burns, P.Y., 1955. Fire scars and decay in Missouri oaks. University of Missouri College of Agriculture, Agricultural Experiment Station Bulletin 642.

Campbell, L.A., Hallett, J.G., O’Connell, M.A., 1996. Conservation of bats in managed forests: use of roosts by Lasionycteris noctivagans. J. Mamm. 77, 976-984.

Carter, T.C., Ford, W.M., Menzel, M.A., 2002. Fire and bats in the Southeast and Mid-Atlantic: more questions than answers? In: Ford, W.M., Russell, K.R., Moorman, C.E. (Eds.), The Role of Fire for Nongame Wildlife Management and Community Restoration: Traditional Uses and New Directions. United States Department of Agriculture, General Technical Report NE-288. pp. 139-143.

Castleberry, S., Miles, A., Miller, D., Conner, L., 2005. Habitat ecology of bats at multiple scales in natural longleaf and intensively managed loblolly pine forests in Georgia. In: Abstract from 90th Annual Ecological Society of America Meeting, Montreal, Canada.

Chambers, C.L., Alm, V., Sider, M.S., Rabe, M.J., 2002. Use of artificial roosts by forest-dwelling bats in northern Arizona. Wildl. Soc. Bull. 30, 1085-1091.

Cottam, G., 1949. The phytosociology of an oak woods in south-western Wisconsin. Ecology 30, 271-287.

Crow, T.R., 1988. Reproductive mode and mechanisms for self-replacement of northern red oak (Quercus rubra): a review. Forest Sci. 34, 19-40.

Crow, T.R., Johnson, W.C., Adkisson, C.S., 1994. Fire and recruitment of Quercus in a postagricultural field. Am. Midl. Nat. 131, 84-97.

Curtis, J.T., 1959. The vegetation of Wisconsin. University of Wisconsin Press, Madison.

Dubois, M.R., Straka, T.J., Crim, S.D., Robinson, L.J., 1999. Costs and cost trends for forestry practices in the South. Forest Landowner 50, 28 32.

Duchamp, J.E., Sparks, D.W., Whitaker Jr., J.O., 2004. Foraging-habitat selection by bats at an urban-rural interface: comparison between a successful and less successful species. Can. J. Zool. 82, 1157-1164.

Elmore, L.W., Miller, D.A., Vilella, F.J., 2004. Selection of diurnal roosts by red bats (Lasiurus borealis) in an intensively managed pine forest in Mississippi. Forest Ecol. Manage. 199, 11-20.

Fenton, M.B., 1997. Science and the conservation of bats. J. Mamm. 78, 1-14.

Fule, P.Z., Cocke, A.E., Heinlein, T.A., Covington, W.W., 2004. Effects of an intense prescribed forest fire: is it ecological restoration. Restor. Ecol. 12, 220-230.

Geluso, K.N., Benedict, R.A., Kock, F.L., 2004. Seasonal activity and reproduction in bats of east-central Nebraska. Trans. Nebraska Acad. Sci. 29, 33 44.

Grindal, S.D., 1999. Habitat use by bats, Myotis spp., in western Newfoundland. Can. Field-Nat. 113, 258-263.

Hartman, G.W., Heumann, B.H., 2003. Prescribed fire effects in the Ozarks of Missouri: the Chilton Creek project 1996-2001. In: Proceedings of the Second International Wildland Fire Ecology and Fire Management Congress, 16-22 November, 2003, Orlando, FL.

Huddle, J.A., Pallardy, S.G., 1996. Effects of long-term annual and prescribed burning on tree survival and growth in a Missouri Ozark oak-hickory forest. Forest Ecol. Manage. 82, 1-9.

Humes, M.L., Hayes, J.P., Collopy, M.W., 1999. Bat activity in thinned, unthinned, and old-growth forests in western Oregon. J. Wildl. Manage. 63, 553-561.

Humphrey, S.R., Cope, J.B., 1968. Records of migration of the evening bat Nycticeius humeralis. J. Mamm. 49, 329.

Hyer, E.J., Goetz, S.J., 2004. Comparison and sensitivity analysis of instruments and radiometric methods for LAI estimation: assessments from a boreal forest site. Agric. Forest Meteorol. 122, 157-174.

Jones Jr., J.K., Fleharty, E.D., Dunnigan, P.B., 1967. The distributional status of bats in Kansas. Misc. Publ. Mus. Nat. Hist. Univ. Kansas 46, 1-33.

Kunz, T.H., 1982. Roosting ecology of bats. In: Kunz, T.H. (Ed.), Ecology of Bats. Plenum Press, New York, pp. 1-55.

Kurta, A., King, D., Teramino, J.A., Stribley, J.M., Williams, K.J., 1993. Summer roosts of the endangered Indiana bat (Myotis sodalis) on the northern edge of its range. Am. Midl. Nat. 129, 132-138.

Ladd, D., 1991. Reexamination of the role of fire in Missouri oak woodlands. In: Buger, G.V., Ebinger, J.E., Wilhelm, G.S. (Eds.), Proceedings of the Oak Woods Management Workshop, Eastern Illinois University, Charleston, pp.
67-80.

Lewis, S.E., 1995. Roost fidelity of bats—a review. J. Mamm. 76, 481-496.

Loomis, R.M., 1974. Predicting the losses in sawtimber volume and quality from fires in oak hickory forests. USDA Forest Service Research Paper NC- 104.

Lorimer, C.G., 1985. The role of fire in the perpetuation of oak forests. In: Johnson, J.E. (Ed.), Challenges in Oak Management and Utilization. Cooperative Extension Service, University of Wisconsin, Madison, pp. 8-25.

Mazurek, M.J., Zielinski, W.J., 2004. Individual legacy trees influence vertebrate wildlife diversity in commercial forests. Forest Ecol. Manage. 193, 321-334.

McCarty, K., 1998. Landscape-scale restoration in Missouri savannas and woodlands. Restor. Manage. Notes 16, 22-32.

McLeod, R.F., Gates, J.E., 1998. Response of herpetofaunal communities to forest cutting and burning at Chesapeake farms, Maryland. Am. Midl. Nat.  139, 164-177.

Menzel, M.A., Krishon, D.M., Carter, T.C., Laerm, J., 1999. Notes on tree roost characteristics of the northern yellow bat (Lasiurus intermedius), the Seminole bat (L. seminolus), the evening bat (Nycticeius humeralis), and the eastern pipistrelle (Pipistrellus subflavus) Fla. Sci. 62, 185-193.

Menzel, M.A., Carter, T.C., Ford, W.M., Chapman, B.R., 2001a. Tree roost characteristics of subadult and female adult evening bats (Nycticeius humeralis) in the upper coastal plain of South Carolina. Am. Midl. Nat. 145, 112-119.

Menzel, M.A., Menzel, J.M., Carter, T.C., Ford, W.M., Edwards, J.W., 2001b. Review of the forest habitat relationships of the Indiana bat (Myotis sodalis).United States Department of Agriculture General Technical Report NE 284.

Menzel, M.A., Owen, S.F., Ford, W.M., Edwards, J.W., Wood, P.B., Chapman, B.R., Miller, K.V., 2002. Roost tree selection by northern long-eared bat (Myotis septentrionalis) maternity colonies in an industrial forest of the
central Appalachian mountains. Forest Ecol. Manage. 155, 107 114.

Miller, D.A., 2003. Species diversity, reproduction, and sex ratios of bats in managed pine forest landscapes of Mississippi. Southeast. Nat. 2, 59 72. Missouri Department of Conservation, 1991. Natural features inventory of Drury-Mincy Wildlife Area.

Moorman, C.E., Russell, K.R., Menzel, M.A., Lohr, S.M., Ellenberger, J.E., Van Lear, D.H., 1999. Bats roosting in deciduous leaf litter. Bat Res. News 40,  74-75.

Moser, W.K., Ducey, M.J., Ashton, P.M.S., 1996. Effects of fire intensity on competitive dynamics between red and black oaks and mountain laurel. North. J. Appl. For. 13, 119-123.

Norberg, U.M., Rayner, J.M.V., 1987. Ecological morphology and flight in bats (Mammalia; Chiroptera): wing adaptations, flight performance, foraging strategy and echolocation. Philos. Trans. R. Soc. London B: Biol. Sci. 316, 335-427.

Nuzzo, V., 1986. Extent and status of midwest oak savanna: presettlement and 1985. Nat. Areas J. 6, 6-36.

Parsons, S., Lewis, K.J., Psyllakis, J.M., 2003. Relationships between roosting habitat of bats and decay of aspen in the sub-boreal forests of British Columbia. Forest Ecol. Manage. 177, 559-570.

Patriquin, K.J., Barclay, R.M.R., 2003. Foraging by bats in cleared, thinned and unharvested boreal forest. J. Appl. Ecol. 40, 646-657.

Paulsell, L.K., 1957. Effects of burning on Ozark hardwood timberlands. University of Missouri College of Agriculture, Agricultural Experimental Station Bulletin 640.

Peterson, D.W., Reich, P.B., 2001. Prescribed fire in oak savanna: fire frequency effects on stand structure and dynamics. Ecol. Appl. 11, 914 927.

Pyne, S.J., 1982. Fire in America. Princeton University Press, Princeton, NJ. Rodrigue, J.L., Schuler, T.M., Menzel, M.A., 2001. Observations of bat activity during prescribed burning in West Virginia. Bat Res. News 42, 48-49.

Saint-Germain, M., Drapeau, P., Hebert, C., 2004. Comparison of coleoptera assemblages from a recently burned and unburned black spruce forests of northeastern North America. Biol. Conserv. 118, 583-592.

Schurbon, J.M., Fauth, J.E., 2003. Effects of prescribed burning on amphibian diversity in a southeastern U.S. National Forest. Conserv. Biol. 17, 1338 1349.

Scowcroft, P.G., 1966. The effects of fire on hardwood forests of the Missouri Ozarks. M.S. Thesis, University of Missouri Columbia, MO, pp. 126.

Simon, N.P.P., Stratton, C.B., Forbes, G.J., Schwab, F.E., 2002. Similarity of small mammal abundance in post-fire and clearcut forests. Forest Ecol. Manage. 165, 163-172.

Smith, K.T., Sutherland, E.K., 1999. Fire-scar formation and compartmentalization in oak. Can. J. Forest Res. 29, 166-171.

Smith, K.T., Sutherland, E.K., 2001. Terminology and biology of fire scars in selected central hardwoods. Tree-Ring Res. 57, 141-147.

Sparks, D.W., Choate, J.R., Winn, R.J., 1999. Observations on reproduction in three species of bats. Prairie Nat. 31, 245-248.

Sparks, D.W., Simmons, M.T., Gummer, C.L., Duchamp, J.E., 2003. Disturbance of roosting bats by woodpeckers and raccoons. Northeast. Nat. 10, 105-108.

Stearns, F., 1991. Oak woods: an overview. In: Buger, G.V., Ebinger, J.E., Wilhelm, G.S. (Eds.), Proceedings of the Oak Woods Management Workshop, Eastern Illinois University, Charleston, pp. 1-7.

Tibbels, A.E., Kurta, A., 2003. Bat activity is low in thinned and unthinned stands of red pine. Can. J. Forest Res. 33, 2436-2442.

van Lear, D.H., 2002. Upland oak ecology and management. In: Spetich, M.A. (Ed.), Upland Oak Ecology  Symposium: History, Current Conditions, and Sustainability.

van Lear, D.H., Waldrop, T.A., 1991. Prescribed burning for regeneration. In: Duryea, M.L., Dougherty, P.M. (Eds.), Forest Regeneration Manual. Kluwer Academic Publishers, The Netherlands, pp. 235-250.

Vonhof, M.J., Barclay, R.M.R., 1996. Roost-site selection and roosting ecology of forest-dwelling bats in southern British Columbia. Can. J. Zool. 74, 1797-1805.

Wade, D.D., Lunsford, J.D. 1988. A guide for prescribed fire in southern forests. Revised Edition. United States Department of Agriculture, Forest Service, TechnicalPublicationR8-TP 11,Southern Region. Atlanta,Georgia, pp. 56.
Watkins, L.C., 1969. Observations on the distribution and natural history of the evening bat (Nycticeius humeralis) in northwestern Missouri and adjacent Iowa. Trans. Kansas Acad. Sci. 72, 330-336.

Watkins, L.C., Shump Jr., K.A., 1981. Behavior of the evening bat Nycticeius humeralis at a nursery roost. Am. Midl. Nat. 105, 258-268.

Whitaker Jr., J.O., 2004. Prey selection in a temperate zone insectivorous bat community. J. Mamm. 85, 460-469.

Whitaker Jr., J.O., Gummer, S.L., 2003. Current status of the evening, Nycticeius humeralis, in Indiana. Proc. Indiana Acad. Sci. 112, 55- 60.

Whitaker Jr., J.O., Brack Jr., V., Cope, J.B., 2002. Are bats in Indiana declining? Proc. Indiana Acad. Sci. 111, 95-106.

White, A.S., 1986. Prescribed burning for oak savanna restoration in central Minnesota. USDA Forest Service, North Central Forest Experiment Station; Research Paper NC-266.

Wilkinson, G.S., 1992. Information transfer at evening bat colonies. Anim. Behav. 44, 501-518.

Zar, J.H., 1984. Biostatistical Analysis, 2nd ed. Prentice-Hall, Englewood Cliffs, NJ.

Sparks, Jeffrey C.1, Masters, Ronald E.1*, Engle, David M.2,
Palmer, Michael W.3 & Bukenhofer, George A.4

1Department of Forestry, 2Department of Agronomy, 3Department of Botany, Oklahoma State University, Stillwater,
OK 74078, USA; 4U.S. Forest Service, Ouachita National Forest, Heavener, OK 74937, USA;
*Corresponding author; Tel.+1 405 744 6432; Fax +1 405 744 9693; E-mail: rmaster@okway.okstate.edu


Abstract. We compared the effects of late dormant-season and late growing-season prescribed fires on herbaceous species in restored shortleaf pine- (Pinus echinata) grassland communities in the Ouachita Highlands of western Arkansas.Herbaceous species richness, diversity, and total forb and legume abundance increased following fire. Late growing season burns reduced distribution and abundance of panicums (primarily Panicum boscii, P. dichotomum, and P. lineari folium) while late dormant-season burns increased Panicum distribution and abundance. Density of legumes (such as Stylosanthes biflora) increased following frequent or annual dormant-season fires. However, season of fire influenced the distribution and abundance of fewer than 10 % of the species. Fire plays an essential role in pine-grassland communities by creating and maintaining open canopy conditions that perpetuate understory herbaceous plant communities.

Keywords: Arkansas; Fire ecology; Fire frequency; Fire season; Ouachita Mountains; Restoration ecology.

NomenclatFure: Smith (1988).

CLICK HERE TO DOWNLOAD AS PDF


Introduction

Fire played an important role in shaping formerly abundant pine- (Pinus spp.) grassland communities in the southeastern United States (Buckner 1989; Platt et al. 1988; Waldrop et al. 1992; Masters et al. 1995). Historical accounts before settlement describe these pine-grassland communities as open ‘park-like’ pine stands with a distinct grass-dominated herbaceous layer and recurrent woody layer, depending on fire frequency (James 1823; Featherstonhaugh 1844; Komarek 1965; Nuttall 1980; Waldrop et al. 1992; Masters et al. 1995). The accumulation of herbaceous material provided adequate fuels for frequent fires of aboriginal and lightning origin which maintained the open structure of these pine-grassland communities (Komarek 1965; Buckner 1989; Foti & Glenn 1991; Waldrop et al. 1992; Masterset al. 1995).

Similar to other forest communities of the World, settlement in the southeastern United States (18th to mid-19th century) altered these landscapes by removing or changing much of the natural vegetation, resulting in fragmented and dissected landscapes (Cottam 1949; Stearns 1949; Curtis 1956; Forman & Godron 1986; Kreiter 1995). The frequency and scale of fires in the region declined after settlement because of aboriginal displacement, fragmentation of habitats causing artificial fire breaks, and fire suppression by settlers (Pyne 1982). This decline in fire frequency caused once open pine-grassland communities to become much more densely forested. Dense forests minimize light reaching the forest floor, thus reducing the herbaceous plant community, understory forage, and habitat quality for many species of wildlife (Lewis & Harshbarger 1976; Masters 1991a; Wilson et al. 1995). The endangered redockaded woodpecker (Picoides borealis), an endemic of southeastern pine forests, is one example of a species that has declined, in part, as a result of increased forest density in the southeastern United States.

The U.S. Forest Service has begun to reconstruct or restore shortleaf pine- (Pinus echinata) grassland communities to benefit both plant and wildlife species dependent on these systems. In the Ouachita National Forest of western Arkansas, the Forest Service uses a program known as Wildlife Stand Improvement (WSI) that consists of thinning midstory and codominant pine and hardwood trees to near pre-settlement basal areas. Currently, WSI treated stands are burned during the dormant season on 3-yr intervals to maintain open structure. However, recent studies in the Ouachitas suggest that the historical fire regime was one of predominantly late growing-season fires and to a lesser extent dormant season burns (Foti & Glenn 1991; Masters et al. 1995). To effectively restore this system, knowledge of the effects of both growing-season and dormant-season prescribed burns is necessary (Masters et al. 1995, 1996).

 

Fig. 1. Restored pine-grassland in the Ouachita Mountains with Pinus echinata

Numerous studies have compared the effects of growing-season and dormant-season fires on vegetation in Coastal Plain regions of Florida, Louisiana, and South Carolina (Grelen 1975; Hughes 1975; Lewis & Harsh-barger 1976; Platt et al. 1988; Waldrop et al. 1992; Glitzenstein et al. 1995). Masters (1991a, b) and Masters et al. (1993) described the effects of dormant season burns of varying frequency on vegetation under a variety of overstory conditions in interior highlands. Masters et al. (1996) described the effects of WSI and dormant-season burns on restored pine-bluestem communities. However, no information is available on the effects of growing-season burns in the Ouachita Mountains. Our main objective was to compare the effects of growing-season and dormant-season burns on herbaceous vegetation richness, diversity, and abundance in WSI-treated stands.

Study area

Our study focused on stands under active management for the endangered red-cockaded woodpecker within the 40 000-ha Pine-bluestem Ecosystem Renewal Area, on the Poteau Ranger District of the Ouachita National Forest (ONF) in Scott County Arkansas. The ONF lies in the 2 280 000 ha Ouachita Mixed Forest  Meadow Province and comprises 648 000 ha throughout the Ouachita Mountains in Arkansas and Oklahoma (Neal & Montague 1991; Bailey 1995). The Ouachita mountains are east-west trending, strongly dissected and range in elevation from 150 – 790 m (Fenneman 1938: 669). South-facing slopes tend to be dominated by shortleaf pine and more mesic north-facing slopes tend to be dominated by oaks (Quercus spp.), hickories (Carya spp.) and other hardwoods (Johnson 1986; Foti & Glenn 1991). Ouachita Mountain soils developed from sandstone and shales and are thin and drought prone. A semi-humid to humid climate prevails with hot summers and mild winters (Smith 1989).

Pinus echinata was the dominant overstory species in all stands (Fig. 1). Codominant and intermediate over story species included Quercus stellata, Q. marilandica, alba, Q. rubra, Q. velutina, Carya texana and C. tomentosa. Tree heights in our study stands ranged from 15 – 23 m ( x = 18.3 m; S.D. = 3.1). Canopy cover ranged  from 68 – 93 % (x = 84.1% ; S.D. = 7.5). Woody sprouts  (≤ 3 m tall) dominated the understory of these stands. The dominant understory woody species and vines included Toxicodendron radicans, Vaccinium pallidum, Quercus stellata, Carya tomentosa, Rubus spp., Parthenocissus quinquefolia, Ceanothus americanus, Vitis rotundifolia, Quercus alba and Pinus echinata (Sparks 1996).

Methods

Experimental design

Our experimental design encompassed two studies (Study 1 and Study 2) and was completely randomized. In these studies we used 12 stands (13.8 to 26.7 ha) that had been previously subjected to WSI and prescribed fire at 3-yr intervals (≥ 3 prescribed fire cycles). Overstory pine density and basal area was similar across all stands (Sparks 1996). Study 1 consisted of three treatments with four replications of each treatment (n = 12). Study 2 used the control and dormant-season fire stands from Study 1 (n = 8). Treatments are as follows:

Study 1

(1) No-burn control (CON1; n = 4);

(2) Late growing-season burn, September 1994 (GS1; n = 4);

(3) Late dormant-season burn March-April 1995 (DS1; n = 4);

Study 2

(4) Late growing-season burn, October 1995 (GS2; n = 2);

(5) Late dormant-season burn, March 1996 (DS2; n = 2);

(6) Frequent dormant-season fire, March-April 1995 and March 1996

(FDS; n = 2);

(7) Infrequent dormant-season fire, burned March-April 1995, no-
burn 1996 (IFDS; n = 2).

Study 1 and Study 2 dormant-season and growing season fire treatments differed in that prescribed burns were applied after three vs. four growing seasons, respectively, following previous dormant-season fire. Study 2 used the dormant-season fire treatments from Study 1 to determine the effects of fire frequency on the herbaceous community. In both studies, late growing-season fires were performed because of poor burning conditions (primarily fuel moisture, presence of live vegetation and high relative humidities) earlier in the season.

Vegetation sampling

We sampled herbaceous vegetation during a two week period in late July 1994 (Study 1 pre-treatment), July 1995 (Study 1 post-treatment; Study 2 pre-treatment), and July 1996 (Study 2 post-treatment). In each stand, we established 30, 1 m × 1 m permanent plots (after Oosting 1956) at 30-m intervals on two to four randomly spaced lines perpendicular to the contour (after Masters 1991a, b). To avoid bias from surrounding stands, we did not sample within 50 m of any edge (Mueller-Dombois & Ellenberg 1974: 123). For each herbaceous species, we recorded percent frequency of occurrence and stem density within plots. Percent cover for vascular plant groups and objects such as rocks, tree boles, and logs was also estimated. Voucher specimens were collected, verified and deposited in the Oklahoma State University Herbarium.

Data analysis

We calculated species richness and diversity (Shannon-Weaver H’) after Ludwig & Reynolds (1988) at the sample (m2) and stand scales. In both studies, we summarized herbaceous species by mean density and percent frequency of occurrence for each year and treatment. All plant species were classified according to plant growth form (e.g., forb, legume, grass, etc.) and season of growth (cool vs. warm). Season of growth was determined by flowering dates described by the Great Plains Flora Association (1986) with cool-season species flowering from November to mid May, and warm season species flowering from mid May through October. To account for pre-treatment differences, we determined the percent change [(post-treatment – pre-treatment / pre-treatment) × 100] in density and frequency of occurrence caused by treatments. All variables were tested for homogeneity of variance using Levene’s test (Snedecor & Cochran 1980). These tests indicated homogeneity of variances, so we tested for treatment differences in percent change using a one-way GLM in which treatment was the factor of interest (Anon. 1985).

In Study 1, we used orthogonal contrasts (burn vs noburn and growing-season fire vs. dormant-season fire) and separated treatment means (P ≤ 0.05) with the protected least significant difference test (Steel & Torrie 1980; Conover & Iman 1981).

We performed Detrended Correspondence Analysis (DCA) using CANOCO (ter Braak 1988), to analyze the species composition data. We checked the results for instability caused by a bug in the program (Oksanen & Minchin 1997). DCA is a multivariate indirect gradient analysis that uses variation in species abundance data to display species and stand locations in a two-dimensional ordination space (ter Braak 1986). DCA axes are in units of constant beta-diversity, where one unit is equal to one standard deviation of species turnover (Hill & Gauch 1980). In DCA, changes in location of a stand over time indicate corresponding changes in real or relative species composition of the stand (Wyant et al. 1991). DCA was used to analyze importance values relative density + relative frequency) to determine changes in stand composition from pre-treatment to post-treatment (after Mueller-Dombois & Ellenberg 1974; Smith 1990). We square-root transformed species abundances before analysis.

Results

Response to fire and fire season

We observed more than 150 herbaceous species during these two studies. Fewer than 10% of these species were influenced (P ≤ 0.05) by season of fire. Late dormant-season fires produced a greater frequency of occurrence of Panicum dichotomum (Study 1: F = 26.9; P = 0.0006, Study 2: F = 29.7, P = 0.0320) and Scleria triglomerata (Study 1: F = 15.3; P = 0.0035, Study 2: F = 19.9, P = 0.0467) than late growing-season fires. Density of Panicum dichotomum (F = 54.5; P = 0.0001) and Scleria triglomerata (F = 5.6; P = 0.0416) was less after late growing-season fires than after late dormant season fires in Study 1.

Although few species were influenced by season of fire, differences (P ≤ 0.05) in density and frequency of major plant categories were apparent (Tables 1 and 2). Late dormant-season fires increased panicum density (primarily Panicum boscii, P. dichotomum, and P. linearifolium) while late growing-season fires greatly reduced total panicum density (Tables 1 and 2). Panicum frequency also declined after late growing-season fires in Study 1 (Table 1). Grasses showed a tendency to decrease in percent cover following fire (Table 3), and a tendency for further decline in density following late growing-season fires (Table 2).


Table 1. Study 1, herbaceous stem density (stems/m2) and percent frequency of occurrence response to season of fire in restored pine-grassland communities on the Ouachita National Forest, Arkansas, summer 1994 and 1995.1

1Row means followed by different letters are different (P < 0.05, Least Significant Difference); 2 Percent change = [(post treatment (1995) – pretreatment (1994) / pre-treatment (1994)) × 100] , presented P > F values are for this category; 3 Contrasts: C = Control; B = Burned stands regardless of season; D = Dormant-season fires; G = Growing-season fires.

Table 2. Study 2, herbaceous stem density (stems/m2) and percent frequency of occurrence response to season of fire in restored pine-grassland communities on the Ouachita National Forest, summer 1995 and 1996.1

1 Row means followed by different letters are different (P < 0.05, Least Significant Difference); 2Percent change = [(post treatment (1996) – pretreatment (1995) / pre-treatment (1995)) × 100] presented P > F values are for this category.

Table 3. Post-treatment percent cover in restored red-cockaded woodpecker clusters on the Ouachita National Forest in 1995.1

1Row means followed by different letters are different (P < 0.05, Least Significant Difference); 2Contrasts: C = Control; B = Burned stands regardless of season; D = Dormant-season fires; G = Growing-season fires.

Regardless of season, fire increased density of legumes, however legume frequency did not increase with burning (Tables 1 and 2). Legume species such as Stylosanthes biflora increased in density (F = 16.9; P = 0.0026) after fire, while other legumes such as Desmodium ciliare (F = 6.58; P = 0.0334) and Lespedeza procumbens (F = 8.37; P = 0.0179) increased in frequency of occurrence after fire. Fire also increased density and frequency of occurrence of numerous forbs such as Coreopsis tinctoria, Polygala alba, and Erechtites hieraciifolia, resulting in an increase in total forb density in Study 1 (Table 1). We found that forbs after late dormant-season fires occurred more frequently than after late growing-season fires and generally increased with fire, although it was not biologically significant (Table 1). Cover of herbaceous vegetation was similar for all treatments, but stands burned during the late dormant season had more bare ground and exposed rock (Table 3). Warm season species had lower densities in response to late growing-season burns than late dormant-season burns (Tables 1 and 2).

Response to frequent fire

Panicum frequency increased with frequent late dormant-season burns (Table 4). However, density of Chasmanthium sessiliflorum declined (F = 35.6; P = 0.0270) after frequent late dormant-season fire. Legume density was greater after frequent late dormant-season fires (Table 4). Density of Lespedeza procumbens (F = 124.0, P = 0.0080) and Stylosanthes biflora (F = 124.9; P = 0.0079) was greater after frequent late dormant-season fires. Helianthus hirsutus (F = 33.7; P = 0.0284) frequency of occurrence was also greater after frequent fire. Stand species richness in frequently burned stands remained stable, but declined in control stands (Fig. 2).

Fig. 2. a. Stand species richness by study and treatment; b. Net-change and standard errors in stand species richness by study and treatment. Means followed by different letters are different (P ≤ 0.05, Least Significant Difference). CON1 = Study 1, no-burn control; GS1 = Study 1, growing-season burn; DS1 = Study 1, dormant-season burn; GS2 = Study 2, growing-season burn; DS2 = Study 2, dormant-season burn; FDS = Study 2, frequent dormant-season burn; and IFDS = Study 2, infrequent dormant-season burn.

 


 

Fig. 3. a. Sample (m2) species richness by study and treatment; b. Net-change and standard errors in sample species richness by study and treatment. Means followed by different letters are different (P ≤ 0.05, Least Significant Difference). CON1 = Study 1, no-burn control; GS1 = Study 1, growing-season burn; DS1 = Study 1, dormant-season burn; GS2 = Study 2, growing-season burn; DS2 = Study 2, dormant-season burn; FDS = Study 2, frequent dormant-season burn; and IFDS = Study 2, infrequent dormant-season burn.


Table 4. Study 2, herbaceous stem density (stems/m2) and percent frequency of occurrence response to frequent fire in restored pine-grassland communities on the Ouachita National Forest, summer 1995 and 1996.1

1 Row means followed by different letters are different (P < 0.05, Least Significant Difference); 2 Percent change = [(post treatment (1995) -pre-treatment (1994) / pre-treatment (1994)) × 100] , presented P > F values are for this category.

Community response to fire

Fire dramatically influenced community composition in restored pine-grassland stands. Species diversity when compared to unburned stands was greater (P ≤ 0.05) after both late growing-season and late dormant season prescribed fires. Stand species richness increased after both late growing-season and late dormant-season fires, while declining in unburned stands (Fig. 2b). Furthermore, post-treatment stand species richness after late dormant-season and late growing-season fires was greater than the unburned controls in Study 1 (Fig. 2a). Sample (m2) species richness increased dramatically after late dormant-season fires with net change in stand species richness being greatest after late dormant-season fires (Fig. 3b).

Detrended Correspondence Analysis illustrated the nature of change in these stands over time and in response to fire (Fig. 4). Axis 1 indicated that year-to-year variation may be the most important factor in determining species composition of these stands (Fig. 4). Axis 2 indicated that geographical location of stands also determines species composition. Axis 3 was tentatively interpreted as an indicator of stand openness with species assemblages more characteristic of prairies being grouped together versus those more characteristic of closed forest being grouped together. Axis 4 may be interpreted as a treatment axis (Fig. 4). We included Axis 4 because treatment effect was our primary interest.

Control stands shifted to the right on Axis 1 and upward on Axis 4 indicating a year and treatment effect, while late dormant-season fire stands shifted right on Axis 1 and down on Axis 4 also indicating a year and treatment effect (Fig. 4). Late growing-season fire stands shifted directly to the right on Axis 1 indicating that year effects had an overriding influence on treatment (Fig. 4). The shift in stands after treatment indicates a similar change in species composition among the treatments. Axes 1 through 4 had eigenvalues of 0.161, 0.081, 0.060, and 0.040 respectively. Together all axes account for 26.7% of the total variation in species data. With an eigenvalue of only 0.042, and the fact that the apparent ‘treatment axis’ is the 4th axis, it is obvious that the effects of treatment, while highly significant, is minor compared with year-to-year effects and site location effects.

Fig. 4. Detrended Correspondence Analysis of stand importance values by treatment, Ouachita National Forest. Stands are connected by vectors to indicate change from pre-treatment sampling to post-treatment sampling. CON1 = Study 1, no-burn control; GS1 = Study 1, growing-season burn; DS1 = Study 1, dormant-season burn; GS2 = Study 2, growing-season burn; DS2 = Study 2, dormant-season burn; FDS = Study 2, frequent dormant-season burn;and IFDS = Study 2, infrequent dormant season burn.

 


Discussion

Treatment response

Burned stands had higher stand species richness and diversity than no-burn controls (Fig. 2). These results are similar to many studies that indicate an initial in crease in species diversity and richness following fire (Trabaud & Lepart 1980; Armour et al. 1984; Thanos et al. 1996). In Study 2, stands in the 2nd growing season since late dormant-season fire (DSC), declined in species richness, indicating that the initial increase in stand richness after fire is short lived (Figs. 2 and 3) and probably influenced by environmental conditions during a given year (Fig. 4). The majority of individual species in both studies did not respond favorably to any one treatment, but were common in all treatments. We believe this is because species present (e.g. Andropogon spp. and various legume species) in restored pine-grassland communities are well adapted to fire, and community changes are small and of short duration. Waldrop et al. (1992) noted that the pine-grassland ecosystem once common throughout the southeastern U.S. was fire derived and fire maintained. Herbaceous species in these restored pine-grassland communities were likely present in pre-settlement communities that developed under a periodically frequent fire regime during both the dormant and growing seasons (Masters et al. 1995).

Fire does not drastically alter species composition in stands with a recent history of fire. Pre-fire composition is a major factor in determining post-fire composition (Armour et al. 1984; Stickney 1986; Rego et al. 1991). Adjacent forests without WSI treatment have dense midstories minimizing light from reaching the forest floor, so species richness and abundance of herbaceous species is much less than in WSI-treated areas (Masters et al. 1996). We also suggest that post-fire species richness and composition is influenced by fire intensity, which is related to litter consumption and reduction in the stature of woody species (Masters et al. 1993; Sparks 1996).

Stand structure

Prescribed fire plays a major role in determining the vegetation structure and composition in restored pinegrassland communities (Wilson et al. 1995). Understories of stands treated with WSI are characteristically dominated by woody sprouts (> 50 000 stems/ha) that restrict light from reaching the forest floor. Late dormant-season fires in these stands on average produce greater fireline intensity than growing-season fires (1300 Kw/m versus < 300 Kw/m), and are more effective at maintaining an open forest structure by reducing the stature of woody sprouts (Sparks 1996).

The effect of a disturbance such as fire on any community or ecosystem depends on the intensity, scale, and frequency (Sousa 1984; Perry 1994; Sparks 1996). Late dormant-season fires in these stands act as more intense disturbances than late growing-season fires, by more effectively reducing stature of the woody community and reducing the litter layer. Increased light penetration due to the reduced stature of the woody understory and reduction of litter after fire provides an opportunity for new herbaceous species to become established, thereby significantly increasing species richness and diversity (Sousa 1984; Masters 1991a, b; Masters et al. 1993). But, fire in either season increases light and allows species already present to prosper, thus the increase in density and percent frequency of occurrence of forbs after fire.

Species composition

Herbaceous species actively growing at the time of a fire in grassland systems are more susceptible to injury than species that are dormant or in early stages of development (Towne & Owensby 1984). Fires during the dormant season reduce cool-season species while favoring many warm-season species (Owensby & Anderson 1967; Hover & Bragg 1981; Towne & Owensby 1984; Hulbert 1988; Howe 1994a). In contrast, growing-season fires reduce warm season species while favoring cool-season species (Hover & Bragg 1981; Ewing & Engle 1988; Biondini et al. 1989; Howe 1994a). Our results in Study 2 showed an increase in density of warm-season species after burning, Study 1 also showed an increase, but not significantly. Neither study showed an increase for cool-season species when late dormant- and late growing-season burns were compared. Growing-season burns may have increased coolseason species had our growing-season fires been conducted earlier in the growing season and before coolseason species initiated new growth. It is important to note that in both studies we attempted to burn earlier in the growing season, but burning conditions (primarily fuel moisture, presence of live vegetation and high relative humidities) were not conducive to fire until later in the growing season.

Several studies have noted that growing-season fires when compared to dormant-season fires and unburned areas increase diversity and richness by increasing the number of annuals and promoting cool-season grasses and forbs (Biondini et al. 1989; Howe 1994b). Platt et al. (1988) noted that growing-season fires produced more flowering stems than fires in other seasons. Many warm season grasses such as wiregrass (Aristida stricta) and little bluestem (Schizachrium scoparium) flower profusely after growing-season fires (Lewis 1964; Robbins & Myers 1992). Hodgkins (1958) noted that composites and legumes increase in response to growing-season fires. Our results indicate an aggressive response from legumes and forbs (Tables 1 and 2), and a larger increase in species richness after dormant-season fires (Fig. 2 and 3). Other studies have found similar results (Grelen & Lewis 1981; Landers 1981; White et al. 1991). Legumes in particular are adapted to fire and benefited because of a hard seed coat and subsequent persistence in the soil seed bank (White et al. 1991; Arianoutsou & Thanos 1996; Thanos et al. 1996).

In Study 2, legume density (primarily Amphicarpa bracteata, Clitoria mariana, Lespedeza repens, and Stylosanthes biflora) was 2 × greater after two frequent late dormant-season fires, similar to Masters et al. (1993), who found legume biomass > 4 × greater after 5 years of annual burning compared to unburned controls. White et al. (1991) found that 43 years of annual winter fires increased legumes by > 25 × over periodic summer and winter burns or annual summer burns. The lack of an increase in legume frequency in our study may be due to relatively high frequency for legumes (80 – 88 %), especially since we observed that most of these species had a tendency towards aggregation. Stem densities within aggregations or sample plots can increase without increasing frequency (Mueller-Dombois & Ellenberg 1974). Further, initial high legume frequency may have been related to the previous fire history within these stands.


Conclusions

Because pine-grassland communities developed under a fire regime that included both dormant and growing season fire, both seasons of fire should be used as management tools in a restoration context. Fire in either season increased species richness, diversity, and total abundance of forbs and legumes, while herbaceous species abundance and richness declined in no-burn controls. Fire reduces woody structure, which influences herbaceous plant composition in restored pine-grassland ecosystems. Increased light and presence of bare ground after fire provide the opportunity for many herbaceous species to become established. Change in species composition and abundance is linked to change in stand structure. Late dormant-season fires are more effective than late growing-season fires at reducing woody sprouts in the understory and at providing bare ground for colonization. As a result, herbaceous species abundance and richness was greater after late dormant season fires.


Acknowledgements. We thank M. E. Payton for assistance with experimental design and statistical analysis, J. Kulbeth, S. Farley, S. Crockett, and D. Gay, for assistance in collecting data and W. Montague for assistance with experimental burns. This project was funded by the U.S. Department of Agriculture, Forest Service, Department of Forestry at Oklahoma State University and was a cooperative effort with Oklahoma Agricultural Experiment Station. This article was published with authorization by the director of the Oklahoma Agricultural Experiment Station.


References

Anon. 1985. SAS users’ guide: statistics, version 5 edition. SAS Institute Inc., Cary, SC.

Arianoutsou, M. & Thanos, C.A. 1996. Legumes in the fireprone Mediterranean regions: an example from Greece. Int. J. Wildland Fire 6: 77-87.

Armour, C.D., Bunting, S.C. & Neuenschwander, L.F. 1984. Fire intensity effects on the understory in ponderosa pine forests. J. Range Manage. 37: 44-49.

Bailey, R.G. 1995. Description of the ecoregions of the United States. 2nd ed. USDA, For. Serv., Misc. Publ. 1391 (revised), Washington, D.C.

Biondini, M., Steuter A.A. & Grygiel, G.E. 1989. Seasonal fire effects on diversity patterns, spatial distribution and community structure of forbs in northern mixed prairie, USA. Vegetatio 85: 21-31.

Buckner, E. 1989. Evolution of forest types in the Southeast. In: Waldrop, T.A. (ed.) Proc. Pine-Hardwood Mixtures: A Symposium on Management and Ecology of the Type, pp. 27-33. USDA, Forest Service, Southeastern Forest Ex-
perimental Station, General Technical Report SE-58. Asheville, NC.

Conover, W.J. & Iman, R.L. 1981. Rank transformations as a bridge between parametric and nonparametric statistics. Am. Stat. 35: 124-129.

Cottam, G. 1949. The phytosociology of an oak woods in southwestern Wisconsin. Ecology 30: 271-287.

Curtis, T.J. 1956. The modification of mid-latitude grasslands and forests by man. In: Thomas, W.L. (ed.) Man’s role in changing the face of the earth, pp. 721-736. University of Chicago Press, Chicago, IL.

Ewing, A.L. & Engle, D.M. 1988. Effects of late summer fire on tallgrass prairie microclimate and community composition. Am. Midl. Nat. 120: 212-223.

Featherstonhaugh, G.W. 1844. Excursion through the slave states. Harper Bros., New York, NY.

Fenneman, N.M. 1938. Physiography of eastern United States.McGraw-Hill Book Co., New York, NY.

Forman, R.T.T. & Godron, M. 1986. Landscape ecology. John Wiley & Sons, New York, NY.

Foti, T.L. & Glenn, S.M. 1991. The Ouachita Mountain landscape at the time of settlement. In: Henderson, D. &

Hedrick, L.D. (eds.) Restoration of old growth forests in the interior highlands of Arkansas and Oklahoma: Proc. of the Conference, pp. 49-68. Morrilton, AR.

Glitzenstein, J.S., Platt, W.J. & Streng, D.R. 1995. Effects of fire regime and habitat on tree dynamics in North Florida longleaf pine savannas. Ecol. Monogr. 65: 441-476.

Great Plains Flora Association. 1986. Flora of the Great Plains. Univ. Press of Kansas, Lawrence, KS.

Grelen, H.E. 1975. Vegetation response to twelve years of seasonal burning on a Louisiana longleaf pine site. USDA, For. Serv., South. For. Exp. Stn., Res. Note, SO-192, New Orleans, LA.

Grelen, H.E. & Lewis, C.E. 1981. Value of range data on fire effects to the wildlife manager. In: Wood, G.W. (ed.) Prescribed fire and wildlife in southern forests, pp. 155-161. The Belle W. Baruch Forest Science Institute, Clemson University, Clemson, SC.

Hill, M.O. & Gauch Jr., H.G. 1980. Detrended correspond ence analysis, an improved ordination technique. Vegetation 42: 47-58.

Hodgkins, E.J. 1958. Effects of fire on undergrowth vegetation in upland southern pine forests. Ecology 39: 36-46.
Hover, E.I. & Bragg, T.B. 1981. Effect of season of burning and mowing on an eastern Nebraska Stipa-Andropogon prairie. Am. Midl. Nat. 105: 13-18.

Howe, H.F. 1994a. Response of early- and late-flowering plants to fire season in experimental prairies. Ecol. Appl. 4: 121-133.

Howe, H.F. 1994b. Managing species diversity in tallgrass prairie: assumptions and implications. Conserv. Biol. 8: 691-704.

Hughes, R.H. 1975. The native vegetation in South Florida related to month of burning. USDA, For. Serv., Southeast. For. Exp. Stn., Res. Note, SE-222, Asheville, NC. Hulbert, L.C. 1988. The causes of fire effects in tallgrass prairie. Ecology 69: 46-58.

James, E. 1823. Account of an expedition from Pittsburgh to the Rocky Mountains, performed in the years 1819 and 1820. H.C. Carey, Philadelphia, PA.

Johnson, F.L. 1986. Woody vegetation of southeastern Leflore County, Oklahoma, in relation to topography. Proc. Okla. Acad. Sci. 66: 1-6.

Komarek, E.V. 1965. Fire ecology – grasslands and man. Proc. Tall Timbers Fire Ecol. Conf. 4: 169-220.

Kreiter, S.D. 1995. Dynamics and spatial pattern of a virgin old-growth hardwood-pine forest in the Ouachita Mountains, Oklahoma, from 1896 to 1994. M.Sc. Thesis, Oklahoma State University, Stillwater, OK.

Landers, J.L. 1981. The role of fire in bobwhite quail management. In: Wood, G.W. (ed.) Prescribed fire and wildlife in southern forests, pp. 73-80. The Belle W. Baruch Forest Science Institute, Clemson University, Clemson, SC.

Lewis, C.E. 1964. Forage response to month of burning. USDA Forest Service,Southeastern Forest Experimental Station, Research Note SE-35, New Orleans, LA.

Lewis, C.E. & Harshbarger, T.J. 1976. Shrub and herbaceous vegetation after 20 years of prescribed burning in South Carolina Coastal Plain. J. Range Manage. 29: 13-18.

Ludwig, J.A. & Reynolds, J.F. 1988. Statistical ecology. John Wiley & Sons, New York, NY.

Masters, R.E. 1991a. Effects of timber harvest and prescribed fire on wildlife habitat and use in the Ouachita Mountains of eastern Oklahoma. Ph.D. Thesis, Oklahoma State University, Stillwater, OK. kansas Press, Fayetteville, AR.

Masters, R.E. 1991b. Effects of fire and timber harvest on vegetation and cervid use on oak-pine sites in Oklahoma Ouachita Mountains. In: Nodvin, S.C. & Waldrop, T.A. (eds.) Fire and the environment: ecological and cultural perspectives, pp. 168-176. USDA, Forest Service, General Technical Report, SE-69, Asheville, NC.

Masters, R.E., Lochmiller, R.L. & Engle, D.M. 1993. Effects of timber harvest and prescribed fire on white-tailed deer forage production. Wildl. Soc. Bull. 21: 401-411.

Masters, R.E., Skeen, J.E. & Whitehead, J. 1995. Preliminary fire history of McCurtain County Wilderness Area and
implications for red-cockaded woodpecker management. In: Kulhavy, D., Hooper, B. & Costa, R. (eds.) Red cockaded woodpecker: species recovery, ecology and management, pp. 290-302. Center for Applied Studies, Stephen F. Austin University, Nacogdoches, TX.

Masters, R.E., Wilson C.W., Bukenhofer, G.A. & Payton, M.E. 1996. Effects of pine-grassland restoration for red-
cockaded woodpeckers on white-tailed deer forage production. Wildl. Soc. Bull. 25: 77-84.

Mueller-Dombois, D. & Ellenberg, H. 1974. Aims and methods of vegetational ecology. John Wiley & Sons, New York, NY.

Neal, J.C. & Montague, W.G. 1991. Past and present distribution of the red-cockaded woodpecker Picoides borealis and its habitat in the Ouachita Mountains, Arkansas. Proc. Ark. Acad. Sci. 45: 71-75.

Nuttall, T. 1980. A journal of travels into the Arkansas Territory during the year of 1819. University of Oklahoma Press, Norman, OK.

Oksanen, J. & Minchin, P.R. 1997. Instability of ordination results under changes in input data order: explanations and remedies. J. Veg. Sci. 8: 447-454.

Oosting, H.J. 1956. The study of plant communities. W.H. Freeman & Co., San Francisco, CA.

Owensby, C.E. & Anderson, K. L. 1967. Yield responses to time of burning in the Kansas Flint Hills. J. Range Manage. 20: 12-16.

Perry, D.A. 1994. Forest ecosystems. John Hopkins Univ. Press, Baltimore, MD.

Platt, W.J., Evans, G.W. & Davis, M.M. 1988. Effects of fire season on flowering forbs and shrubs in longleaf pine forests. Oecologia (Berl.) 76: 353-363.

Pyne, S.J. 1982. Fire in America: a culture history of wildland and rural fire. Princeton University Press, Princeton, NJ.

Rego, F.C., Bunting, S.C. & da Silva, J.M. 1991. Changes in understory vegetation following prescribed fire in mari-
time pine forests. For. Ecol. Manage. 41: 21-31.

Robbins, L.E. & Myers, R.L. 1992. Seasonal effects of prescribed burning in Florida: a review. Tall Timbers Research, Inc., Miscellaneous Publication Number 8. Tallahassee, FL.

Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, 2nd ed. University of Arkansas, Fayettville, AR.

Smith, R.L. 1990. Ecology and field biology. Harper & Row, New York, NY.

Smith, R.M. 1989. The atlas of Arkansas. University of Ar

Snedecor, G.W. & Cochran, W.G. 1980. Statistical methods. 7th ed. Iowa State University Press, Ames, IA.

Sousa, W.P. 1984. The role of disturbance in natural communities. Annu. Rev. Ecol. Syst. 15: 353-391.

Sparks, J.C. 1996. Effects of growing-season and dormant season prescribed fire on vegetation in red-cockaded woodpecker clusters. M.Sc. Thesis, Oklahoma State University, Stillwater, OK.

Stearns, F.W. 1949. Ninety years change in a northern hard wood forest in Wisconsin. Ecology 30: 350-358.

Steel, R.G.D. & Torrie, J.H. 1980. Principles and procedures of statistics, a biometrical approach. 2nd ed. McGraw-
Hill Book, New York, NY.

Stickney, P.F. 1986. First decade plant succession following the Sundance Forest Fire, northern Idaho. USDA, Forest

Service, Interior Forest & Range Experimental Station, General Technical Report,INT-197, Ogden, UT. ter Braak, C.J.F. 1986. Canonical correspondence analysis: a new eigenvector technique for multivariate direct gradient analysis. Ecology 67: 1167-1179.

ter Braak, C.J.F. 1988. CANOCO – a FORTRAN program for Canonical Community Ordination. Microcomputer Power, Ithaca, NY.

Thanos, C.A., Daskalakou, E.N. & Nikolaidou, S. 1996. Early post-fire regeneration of a Pinus halepensis forest on Mount Parnis, Greece. J. Veg. Sci. 7: 273-280.

Towne, G. & Owensby, C. 1984. Long-term effects of annual burning at different dates in ungrazed Kansas tallgrass prairie. J. Range Manage. 37: 392-397.

Trabaud, L. & Lepart, J. 1980. Diversity and stability in garrigue ecosystems after fire. Vegetatio 43: 49-57.

Waldrop, T.A., White, D.L. & Jones, S.M. 1992. Fire regimes for pine – grassland communities in the southeastern United States. For. Ecol. Manage. 47: 195-210.

White, D.L., Waldrop, T.A. & Jones, S.M. 1991. Forty years of prescribed burning on the Santee fire plots: effects on understory vegetation. In: Nodvin, S.C. & Waldrop, T.A. (eds.) Fire and the environment: ecological and cultural perspectives, pp. 51-59. USDA, Forest Service, South eastern Forest Experimental Station., General Technical Report, SE-69, Asheville, NC.

Wilson, C.W., Masters, R.E. & Bukenhofer, G.A. 1995. Breeding bird response to pine-grassland community restoration for red-cockaded woodpeckers. J. Wildl. Manage. 59: 56- 67.

Wyant, J.G., Ralph, J.A. & Bechold, W.A. 1991. Physiographic position, disturbance and species composition in North Carolina coastal plain forests. For. Ecol. Manage. 41: 1- 19.


Received 11 March 1997;
Revision received 4 August 1997;
Accepted 13 August 1997;
Final revision received 9 January 1998.