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Profissional Documentos
Cultura Documentos
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148 FLORIDA SCIENTIST [VOL. 73
borealis). These treatments are also used to allow the re-establishment of the
natural disturbance regime, fire. Treatments such as roller-chopping and
mowing can bring fuels into the zone where they will be consumed, promoting
shifts in vegetation structure from tree-dominated to shrub- or herb-
dominated. Finally, herbicides and mechanical treatments are used (along
with fire) to reduce fire intensity and fire severity in subsequent fires, e.g., in
Florida’s pine flatwoods (Brose and Wade, 2002).
As pre-treatments for fire, mechanical treatments and herbicide use can
allow fires to more effectively carry and consume fuels, and may make control
of fires easier. However, such restructuring of the fuel may alter fire intensities
(Busse et al., 2005) and spatial patterns (Rickey et al., 2007). The effect of
repeated fire surrogate pre-treatments on vegetation is not well known. Finally,
the risks of soil compaction and disturbance are a worry even among strong
advocates of fire pre-treatments (Long et al., 2005) because such disturbances
could favor non-native plant invasions (Catling et al., 2002; Dodson and
Fiedler, 2006; Hobbs and Atkins, 2006 but see Maschinski et al., 2005). The
combination of mechanical disturbance and fire has also been shown to
exacerbate non-native invasions in the western U.S. (Keeley, 2001). The
vulnerability of treated sites to invasion certainly can depend on the landscape
context: in small urban fragments, large populations of non-native species may
lurk along the edges of protected areas (Yates et al., 2004). In Florida, a
combination of many invasive species and a high degree of landscape
fragmentation suggests that management decisions may exacerbate existing
threats from invaders. Fundamental alterations of ecosystem processes caused
by non-native species have already been documented in Florida (Gordon, 1998;
Lippincott, 2000).
As surrogates for fire, mechanical treatments and herbicide use avoid all
the problems with escapes and smoke while being feasible under broader
weather conditions than fire. In some cases, mechanical treatments are used in
treating overgrown vegetation that is deemed unsafe to burn, to deal with
situations where thick vegetation or unfavorable fuel structure will not carry
fire, or where ignition of high fuel loads may threaten desirable resources on
site. However, fire surrogates may create additional problems. These include
alteration of carbon, nutrient, and microbial dynamics (Gundale et al., 2005;
Giai and Boerner, 2007; Boerner et al., 2008) and failure to provide direct fire
germination cues such as heat or smoke (e.g., Rokich and Dixon, 2007; Lindon
and Menges, 2008; Reyes and Trabaud, 2009).
Many managers of Florida’s uplands share common goals of ecological
restoration, whether the tools used include fire, roller-chopping, mowing,
herbicide application, large-scale logging, individual tree felling, or other
approaches. For example, restoration of overgrown sandhill usually will
include a goal of opening up the midstory vegetation and increasing the species
richness of the ground layer (Brockway et al., 1998; 2005; Provencher et al.,
2001a). In Florida scrub, increasing bare sand to 10–40% cover will help
recover scrub endemic plants (Menges and Hawkes, 1998), scrub herpetofauna
150 FLORIDA SCIENTIST [VOL. 73
(Greenberg et al., 1994) and benefit the Florida Scrub-Jay (Breininger and
Schmalzer, 1990; Fitzpatrick et al., 1991). Such bare ground levels can typically
be found just after fires in scrub (e.g., Outcalt and Greenberg, 1998), are found
for only a few years after fire in coastal scrub (Schmalzer, 2003), but may
persist for years or decades in interior scrub (Greenberg, 2003; Hawkes and
Menges, 1996). The Florida Scrub-Jay is also sensitive to vegetation height
(preferring shrubs , 3 m tall; Breininger and Oddy, 2004), shrub species
composition (preferring open oak areas; Breininger et al., 1995) and areas
distant from forests (Fitzpatrick et al., 1991; Breininger et al., 1995). For
prairies and flatwoods, reduction of palmetto cover is a common management
objective (Kalmbacher et al., 1983). On the other hand, palmetto cover may be
important in encouraging fire spread in Florida scrub (Greenberg et al., 1995b;
Schmalzer et al., 2003).
Finally, at a larger scale, landscape heterogeneity may be a goal. For
example, Greenberg and Simons (1999) argued that patches of older oaks are a
valuable landscape feature within the longleaf pine (Pinus palustris) landscape.
While fire is thought to have historically maintained these various conditions
across the landscape, we know little about whether fire surrogates will be able
to similarly restore the landscape structure and function.
The potential of management using fire surrogates for conservation and
restoration goals leads us to several key questions. First, in defining targets for
ecosystem management, should we focus more on structure or function? If fire,
the evolutionarily significant natural disturbance agent, is reduced or
eliminated in Florida’s ecosystems, what will be the consequences? When is
it appropriate to use alternative management treatments to fire, and when
should fire be combined with mechanical treatments and/or herbicide use?
How much do we know about the long-term effects of alternative management
on Florida’s ecosystems?
METHODS—We reviewed the published literature, including agency reports, that dealt with
mechanical treatments or herbicide use, with or without fire, and their effects on aspects of
Florida’s ecosystems. These studies varied widely in a number of ways, precluding a formal meta-
analysis. We paid particular attention to the following issues (summarized for major studies in
Table 1):
1.) Did the studies include a fire-only treatment? 2.) What was the length of the study? 3.) What
response variables were measured? 4.) Was there replication of treatments? 5.) Were reference
sites sampled for comparison? 6.) Were pre-treatment data collected? 7.) What was the size of
the treatment block? 8.) Were fire intensities quantified?
We summarize the results from individual studies here by community type, moving from most
to least xeric.
Q6: Were pre-treatment data collected? Q7: What was the size of the treatment plot (hectares)? Q8: Were fire intensities quantified?
Treat- Q1 Q2 Q3 2 Q4 Q5 Q6 Q7 Q8
Habitat Citations ments1 Fire-only Yrs. Vars. Reps Ref Pre-T Size Fire Int
Scrub Rickey et al. 2007, Weekley et al. MLFX Yes 6 VR No Yes Yes 1–20 ha Yes
2008
Greenberg et al. 1994, 1995a, L, F+L No 7 VHBA Yes Yes No .8 ha No
1995b, 1996
Williges et al. 2006, Berish et al. CMLFX No 4 VR No No No . 10 ha No
2002
Schmalzer et al. 2003, Schmalzer MFX Yes 13 VR Yes Yes Yes variable No
and Foster 2006, 2008
Sandhill Provencher et al. 2001a, 2001b, FHMX Yes 7 VHBA Yes Yes Yes 81 ha Yes
2002a, 2002b, 2002c, Litt et al.
2001
Menges et al. 2007, Rickey et al. SFX Yes 7 VR No Yes Yes 2–4 ha Yes
2007
Wilkins et al. 1993a, b, H No 1–2 V Yes Yes Yes 0.04–0.09 ha NA
Brockway et al. 1998; Brockway HF Yes 7 V Yes Yes Yes 0.25 ha No
and Outcalt 2000
Dry Prairie Fitzgerald and Tanner 1992; FCX Yes 12–13 VB Yes Yes3 No 6 ha No
MENGES AND GORDON—FIRE SURROGATES
plant and animal species are found in these habitats (Christman and Judd,
1990; Estill and Cruzan, 2001). In contrast to several other pyrogenic
communities discussed here, scrub fires are generally longer interval (Menges,
2007), high-intensity, and stand-replacing, resulting in top-kill of shrubs and
overstory trees. Fire reduces aboveground biomass and releases herbaceous
species from competition (Quintana-Ascencio and Morales-Hernández, 1997).
Some of these herbaceous species are killed by fire, but populations are rapidly
restored by recruitment from seeds in a persistent seed bank (Menges and
Kohfeldt, 1995).
Scrub restoration using mechanical treatments, either alone or preceding
fire, and in comparison with fire, have been studied on the Lake Wales Ridge
(LWR) in central peninsular Florida (Weekley et al., 2008; Rickey et al., 2007).
A series of large-scale, field experiments compared the effects of logging,
mowing (with a brown-tree cutter), and chopping (with a gyrotrack) to restore
the structure of overgrown (. 2m tall hardwood layer) scrub. Detailed
vegetation data were collected before and up to five-years post-treatment, but
treatments were not replicated. Restoration objectives included reducing
canopy, subcanopy, shrub, litter, and lichen cover, increasing both cover of
bare sand and abundances of rare plants.
In the logging experiment, all treatments dramatically reduced canopy pine
and subcanopy hardwood densities, but no treatment was successful in
reducing stem densities in the shrub layer (Weekley et al., 2008). Logging, with
or without fire, created greater bare sand cover than seen in any scrub site not
subjected to mechanical treatments. Although logged areas in this study were
not invaded by exotic species, logged areas with excessive bare sand cover may
have increased risk of such invasion and may have insufficient fuels to carry
subsequent prescribed fires.
The mowing versus fire study (Fig. 1) was conducted at two scrub sites
(Weekley et al., 2008). The two burn treatments were more effective than
mowing alone in reducing woody cover, increasing bare sand and decreasing
litter cover. All three treatments were effective in reducing shrub height and
these reductions persisted for five years. Litter depth was reduced by both burn
treatments, but actually increased in the mow-only treatment relative to the
control. The burn treatments (and particularly the burn-only treatment) were
the most successful in increasing rare species densities at both sites. Many
effects of burning persisted for at least five years after treatments.
In the chopping study (Weekley et al., 2008), burning (with or without
prior chopping) was more effective than chopping alone in promoting
conditions required for the recruitment of rare and imperiled Florida scrub
herbs. While all three management treatments reduced canopy, subcanopy,
and shrub cover relative to the control site, only the burn-only and chop &
burn treatments increased bare sand and reduced litter and lichen cover,
outcomes favorable for rare herb recruitment.
Summarizing these three studies, Weekley and co-workers (2008)
recommended that prescribed fire be the primary scrub management tool
No. 2 2010] MENGES AND GORDON—FIRE SURROGATES 153
FIGURE 1. Archbold Biological Station worked with Florida Fish and Wildlife Conservation
Commission to study effects of mowing and burning at Royce Ranch (Weekley et al. 2008). a.
Overgrown Florida scrub with Florida rosemary (Ceratiola ericoides) and scrub oak (Quercus inopina)
in foreground, and sand pine (Pinus clausa) in background. Photograph by Eric Menges. b. Field crew
sampling Florida scrub vegetation in 2004, two years post-mowing. Photograph by Carl Weekley.
FIGURE 2. Guana River SRA is one of several study sites part of the study by Williges and
co-workers (2006). Photographs by Eric Menges. a. Overgrown scrub adjacent to treatment areas.
b. Chopped Florida scrub shortly after treatment.
FIGURE 3. A variety of mechanical treatments have been combined with fire in studies by
Paul Schmalzer and colleagues (e.g., Schmalzer and Foster, 2008) on Merritt Island scrub.
Photographs by Paul Schmalzer. a. A K-G Blade. b. A brown tree cutter.
scrub in the same area (compare with Schmalzer, 2003). Bare ground was
scarce in all treatments (Schmalzer and Foster, 2006). In other mechanically
treated sites (both cut sites and roller-chopped sites), loss of palmetto cover
after cutting was hypothesized to reduce the ability of these sites to carry fire
(Schmalzer et al., 2003; Schmalzer and Foster, 2006; 2008), although this effect
has not yet been clearly shown (Schmalzer, 2009, personal communication).
Brazilian pepper invasion was a problem on mechanically treated coastal scrub
on more alkaline soils (Schmalzer and Foster, 2008). Mechanical treatments were
recommended by these authors as a one-time pre-treatment to be followed by
prescribed burning (Schmalzer et al., 2003; Schmalzer and Foster, 2006; 2008).
Other studies in scrub are also relevant. Campbell and Christman (1982)
found similarities in vegetation structure following mechanical treatments and
high intensity fire. Kenner (1994) reported higher plant diversity (especially
grasses and legumes) and increases in key animals in a sand pine scrub stand
that was clear-cut and then burned, but no other treatments were available for
comparison.
FIGURE 5. Subcanopy felling has been used as a pre-treatment for fire in sandhill on the Lake
Wales Ridge, as studied by Menges and co-workers (2007). Photographs by Eric Menges. a. Fire
moving through sandhill plots with areas with a subcanopy felling pre-treatment or burn-only. b.
Post-fire view of a sandhill area pre-treated with subcanopy felling (foreground) and burn-
only (background).
the pre-fire cover after two years. The cover of graminoids and forbs was
equivalent across the herbicide & fire and fire-only treatments after two years.
The authors found no significant differences in plant species richness or other
diversity indices between the herbicide & fire and the fire-only treatments.
Species richness was generally highest one year after burning. Spot application,
rather than broadcast application of herbicides, had lower impacts on forbs
(Brockway et al., 2005).
After reviewing several studies, these authors concluded that herbicide
treatment followed by fire provided the most expedient treatment for
restoration of xeric longleaf pine wiregrass ecosystems (Brockway et al.,
2005). While fire was slow to reduce hardwood density, it was critical for
stimulating increased cover and recruitment of herbaceous species (Brockway
et al., 2005). These conclusions were consistent with those drawn by
Provencher and co-workers (2001a), above.
FIGURE 6. A variety of treatments have been used in Fire and Fire Surrogate study at
Myakka River State Park. The vegetation is typical south Florida flatwoods with a scattered pine
overstory and saw palmetto and gallberry understory. This project was reported by Outcalt and
colleagues from the US Forest Service (USDA Forest Service, 2008). Photographs by Kenneth W.
Outcalt, US Forest Service. a. Flatwoods shortly after chopping, 2001. b. Prescribed fire in burn-
only treatment.
here was applied carefully with a small chopper in an effort to reduce saw
palmetto without damage to desirable grasses.
When granular hexazinone (no fire treatment) was applied to mesic pine
flatwoods at 0, 1.7, 3.4, and 6.8 kg/ha in a randomized blocked design with
three replicates (Wilkins et al., 1993a), the results for woody species were
similar to results reported above for sandhills. Hardwood and herbaceous
species cover and diversity were reduced in proportion to the concentration of
the herbicide used, but differences among treatments were no longer significant
for herbaceous species in the second year. The 69% reduction in oak cover in
the 1.7 kg/ha hexazinone treatment was significantly lower than the 97%
reduction in the xeric sandhill experiment (Wilkins et al., 1993a). Specific
responses of other species differed as well.
taxa (Snyder et al., 1990; Possley et al., 2008). Pine rocklands are fire-
maintained, with typical fire return intervals 2–15 (Snyder et al., 1990) or 10–20
(Carlson et al., 1993) years. Post-fire, shrubs resprout vigorously and the
growth of herbs is enhanced (Snyder et al., 1990, Carlson et al., 1993). For
example, population viability of one narrowly endemic pineland herb is
favored by fires every 5–7 years (Liu et al., 2005). With several decades of fire
suppression, pine rocklands develop into tropical hammocks with a residual
overstory of pine (Snyder et al., 1990). Fire suppressed, isolated, and small pine
rockland fragments may harbor significant densities of invasive non-native
plants (Gordon, 1998) which may have negative effects on native plant
diversity (Possley and Maschinski, 2006). Pineland overstories are affected by
periodic hurricanes, but understories are little affected (Loope et al., 1994).
Restoration of pine rocklands in Florida is very challenging in developed
areas of Miami-Dade County, where many fire-suppressed pine rockland
fragments remain. Thinning dense thickets of south Florida slash pine reduced
tree and shrub cover and increased grass cover (Maschinski et al., 2005). More
importantly, thinning increased species richness without introducing exotic
species. No concurrent fire treatment was available for comparison. Intensive
site manipulation and introduction of new propagules may be necessary
adjuncts to restoration of fire in some pine rockland fragments (Wendelberger
et al., 2008).
What we still don’t know—One weakness in most of these studies (but see
Watts et al., 2006) is that they are short- term and do not consider the effects of
repeated fire surrogate treatments. Negative responses to various treatments,
such as losses of some native species, increased invasion of non-native plant
species with soil disturbance, and alterations in litter and nutrient cycling,
could be exacerbated with repetition. Insufficient research and monitoring on
these unintended consequences of management limits our ability to understand
long-term implications of these management approaches.
166 FLORIDA SCIENTIST [VOL. 73
There may be some ecosystems or some conditions under which fire alone
will not effectively achieve restoration goals. Repeated fire may not reduce
palmetto and shrub cover in long-unburned uplands. However, fire is an
effective management tool in other situations. For example, in scrub on the
Lake Wales Ridge, fire alone has been used to successfully restore some
overgrown scrub areas. At The Nature Conservancy’s (TNC) Saddle Blanket
Preserve, long-unburned scrub was burned with a series of prescribed burns
over an 11 year period, with special attention to the geometry of subsequent
burns. Most burns were prescribed to burn into blackened areas that had been
burned relatively recently. Qualitative vegetation data and photographic
evidence shows desirable changes in shrub height and cover (B. Pace-Aldana,
The Nature Conservancy, unpubl. data).
Fire-alone has been used in other areas to restore Florida scrub. In
northwest Florida, fire significantly reduced woody plant densities, although
repeated fires were recommended to restore herbaceous plants (Ruth et al.,
2007). A single fire in sand pine scrub at Ocala National Forest restored
structure and resulted in increases in rare scrub herbs (Custer and Thorsen,
1996). However, a single fire may fail to regenerate herbaceous species in very
long-unburned areas, probably because of declines in species representation in
the vegetation and soil seed bank (Abrahamson and Abrahamson, 1996). In
such cases, as on former agricultural sites, the difficult process of reestablishing
Florida scrub from plantings will need to be considered (Schmalzer et al.,
2002).
Repeated fires in sandhill have also been successful in restoration efforts.
Three successive growing season fires successfully reduced oak density
(Glitzenstein et al., 1995). Data from repeated fires over an eight year period
in sandhill at TNC preserves in central and north Florida suggest that fire
alone can make progress toward composition and structure goals (Slapcinsky
and Gordon, 2001a; b). Starting conditions clearly influence the variables that
show increased similarity to those in local reference sites. However, several of
the burn units that have received two or three fires are showing recovery in
over 50% of the 16 variables monitored. Repeated fires in a south-central
Florida sandhill also produced many favorable changes, including increased
grasses, reduced litter, and suppressed large shrubs (Reinhart and Menges,
2004). In the western U.S., some have argued that fire-alone is not receiving
enough attention as an alternative to more intensive silvicultural treatments in
restoring overgrown ponderosa pine forests (Allen et al., 2002).
We also lack evidence of whether fire in Florida has critical effects that are
not mimicked by mechanical or herbicide treatments. One possibility is that fire
creates heat pulses or provides chemicals through smoke that are critical to
some species. Wiregrass seed production and viability is greatest subsequent to
growing season fires (Seamon and Myers, 1992). Effects of heat and smoke in
stimulating germination of recalcitrant seeds is well known from many parts of
the world, including Australia (Rokich and Dixon, 2007), South Africa
(Newton et al., 2006), and the American West (Preston and Baldwin, 1999).
No. 2 2010] MENGES AND GORDON—FIRE SURROGATES 167
ACKNOWLEDGMENTS—We thank the authors of the various studies for their hard work and
Dale Brockaway, Katie Greenberg, Ken Outcalt, Louis Provencher, Paul Schmalzer, George
Tanner, Adam Watts, and Neal Wilkins for checking excerpts from this manuscript for factual
accuracy. We also thank Tricia Martin, Reed Bowman, Carl Weekley, and Hilary Swain for
encouraging us to develop this review, as well as the Florida Fish and Wildlife Conservation
Commission (FFWCC) and the Lake Wales Ridge Ecosystem Working Group for providing
venues for discussion of this work. We appreciate the support of the agencies that have facilitated
our own work on their properties on mechanical treatments, herbicides, and fire, including
FFWCC, U.S. Fish and Wildlife Service (USFWS), The Nature Conservancy, Florida Division of
Forestry, and Eglin Air Force Base. Funding for this research was provided by USFWS, FFWCC,
and Department of Defense. The views and conclusions contained herein are those of the authors
and should not be interpreted as necessarily representing the official policies or endorsements of
any of the above organizations.
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