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Archbold Biological Station, PO Box 2057, Lake Placid, FL 33870
The Nature Conservancy, Department of Biology, PO Box 118526, University of Florida,
Gainesville, FL 32611
Biological Sciences




Archbold Biological Station, PO Box 2057, Lake Placid, FL 33870
The Nature Conservancy, Department of Biology, PO Box 118526, University of Florida,
Gainesville, FL 32611

ABSTRACT: Mechanical treatments such as roller-chopping, mowing, chain-sawing, and logging,

and herbicide application, are increasingly used to manage fire-maintained Florida ecosystems. Goals
include achieving or restoring desired vegetation structure and composition, providing habitat for listed
species, and allowing the reintroduction of fire. We review studies evaluating mechanical treatments
and herbicide effects on Florida’s plant and animal communities. Mechanical treatments and herbicide
often accelerated vegetation structure changes, but ecological benefits were generally greatest when
they were combined with fire. Soil disturbances, weedy species increases, and rapid hardwood
resprouting were sometimes problems with mechanical treatments. Fire itself was crucial for
maintenance of individual species and species diversity. When feasible, mechanical and herbicide
treatments should be used as pretreatments for fire rather than as fire surrogates. Managers should
segue to fire-only approaches as soon as possible. The effects of removing fire, the most evolutionarily
significant disturbance in Florida, on fire-adapted plants and animals requires more study. If we
increase the use of mechanical agents, how will this affect ecosystem resilience? We suggest caution in
using these evolutionarily novel treatments, with close monitoring of their effects. More information on
the long-term effects of repeated non-fire treatments is needed before such approaches are adopted

Key Words: Ecological restoration, mowing, roller-chopping, logging,

Florida scrub, sandhill, flatwoods, dry prairie, pine rockland

MOST upland ecosystems in Florida are affected by large-scale natural

disturbances, principally fire and tropical storms. Lightning ignitions between
May and October have dominated Florida for thousands of years, with the
largest proportion of land area burned at the end of the dry season (May–July;
Robbins and Myers, 1992). In addition, ignitions during winter months by
Native Americans and European settlers have likely had significant effects on
Florida’s ecosystems (Robbins and Myers, 1992). Because of the long history
of fire in Florida, many of its ecosystems and species are likely adapted to
characteristics of the fire regime (Platt et al., 1988; Menges, 2007; Slapcinsky et
al., 2009). Prescribed fire is currently used as a management tool by public and
private land managers for restoration and maintenance of Florida ecosystems
(e.g., Wade et al., 1980; Duncan et al., 1999; Ruth et al., 2007).

* Corresponding author: Email:


Prescribed fire is used to achieve many goals, including restoring and

maintaining vegetation composition and structure, reducing wildfire hazard,
promoting the viability of rare plants and animals, enhancing wildlife, and
encouraging tree growth for forestry. Fire is also used in grazed and hunted
systems to maintain relatively open habitats and stimulate new growth for
grazers and browsers. Fire regimes, including such factors as fire frequency,
intensity, size, and patchiness, can be manipulated to encourage certain
vegetation structure and enhance particular species (Menges, 2007). Fire
managers may also try to mimic the baseline fire regime, reflecting conditions
pre-settlement or at some later time. Depending on the organization and their
goals, land managers may concentrate burns during the lightning season or
spread ignitions throughout most of the year. For example, spring burns are
used to promote groundcover productivity without impact to ground-nesting
birds or trees. In Florida, fall burns are rarely used in forested areas because of
increased pine mortality (Menges and Deyrup, 2001). Fire frequencies vary by
ecosystem, ranging from more frequent than every five years for flatwoods,
sandhills and prairies, to fires occurring every 15–30 years or longer in certain
types of Florida scrub (Menges, 2007).
Despite the clear desirability of prescribed fire as a management tool, fire
can be challenging to apply. This is particularly true for areas of the wildland/
urban interface and where fuels have accumulated to high levels (Long et al.,
2005). Florida’s landscape is dominated by areas where increasing human
populations and suburban sprawl have encroached upon wildlands. The risks
of escaped fires and smoke are now known to all Floridians, and land
managers are under incredible pressure to conduct burns that will not affect
homeowners and businesses. The legacy of fire suppression complicates
matters, as burning these areas may create difficult control problems, wildfire
hazards, additional smoke, and result in greater tree mortality (Long et al.,
2005). In many cases, land managers are turning to alternatives or pre-
treatments to fire. Such tools as roller-chopping, mowing, logging, and
herbicide applications have moved from the spheres of forestry and range
management to conservation areas that are primarily managed for natural
values such as biodiversity and rare species protection.
Mechanical treatments are now being widely used by land managers in the
western half of the United States, particularly to reduce fuel loads and the
impact of wildfires. These fuel treatments may not necessarily improve
ecosystem health (Reinhardt et al. 2008) although they often are touted as
being valuable for ecosystem restoration. These non-fire treatments generally
are recommended in combinations with fire, but can have potential
disadvantages (Schwilk et al., 2009; Stephens et al., 2009).
Likewise, mechanical treatments and herbicides are used as tools to restore
and maintain ecosystems in Florida. In particular, these treatments are aimed
at restoring vegetation structure and composition, including providing habitat
for endangered and threatened species such as the federally threatened Florida
Scrub-Jay (Aphelocoma coerulescens) and Red-cockaded Woodpecker (Picoides

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

(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.

RESULTS—Scrub—Florida’s pyrogenic scrub systems are on xeric, upland,

sandy soils and are dominated by evergreen hardwood shrubs (Quercus spp.,
Ericaceae, Ceratiola ericoides, Sabal etonia, and Serenoa repens), sometimes
with a pine canopy (Myers, 1990; Menges, 1999). Numerous rare and endemic
TABLE 1. Summary of major studies of restoration using mechanical treatments and herbicides in upland Florida ecosystems (see text for specific
citations). Columns labeled Q1–Q8 provide answers to the following questions: Q1: Did the studies include a fire-only treatment? Q2: What was the length of the
study in years? Q3: What response variables were measured? Q4: Was there replication of treatments? Q5: Were reference/control sites sampled for comparison?
No. 2 2010]

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
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
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.
Menges et al. 2007, Rickey et al. SFX Yes 7 VR No Yes Yes 2–4 ha Yes
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

Watts and Tanner 2004; Watts

et al. 2006
Flatwoods USDA Forest Service 2008 FCMX Yes 3 V Yes Yes Yes .510 ha Yes
Wilkens et al. 1993a H No 2 V Yes Yes Yes 0.09 ha NA
Pine Rockland Maschinski et al. 2005 LH No 2 V Yes Yes Yes 13 ha NA4
Treatments: F5fire; H5herbicide; C5chop (includes soil disturbance); M 5 mow, brown tree cutter, etc; L 5 Log, X5combinations
Response Variables: V5vegetation; R5rare plants; H 5 herps; B5 birds, A5arthropods, etc.
Watts et al. (2006) study only.
Not applicable.

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

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.

and that mechanical treatments be used carefully, monitored closely, and

followed up with prescribed burns. Logging was not recommended as a
restoration tool because of the significant soil disturbance involved.
In a long-unburned sand pine scrub at Ocala National Forest in north-
central Florida, vegetation (Greenberg et al., 1995b), arthropods (Greenberg
and McGrane, 1996), reptiles (Greenberg et al., 1994), and birds (Greenberg et
al., 1995a) were compared among two logging and site preparation treatments
or wildfire with salvage logging (no fire-only treatment). All treatments had
higher herbaceous species richness, herb diversity, sand pine (Pinus clausa var.
clausa) density, and bare ground than did the long-unburned mature forest
control, which was dominated by a sand pine canopy and ground lichens. Rare
scrub plant species and post-fire herbaceous species often proliferated in open
conditions associated with fire or ground disturbance. Overall, vegetation
richness, diversity, and composition in the logged and re-seeded treatments
were similar to that in the burned and salvage-logged treatment (Greenberg et

al., 1995b). However, negative impacts of roller-chopping on Serenoa repens,

which can be critical for carrying fire in scrub, resulted in the suggestion that
managers be cautious about implementing this mechanical management.
Overall, the results suggested that high intensity silvicultural management has
similar impacts on the vegetation as wildfire followed by salvage logging.
Total numbers and dry weight of surface-active, ground-dwelling
arthropods were equivalent in all treated and untreated plots, though numbers
per size class varied (Greenberg and McGrane, 1996). Greater numbers and
biomass of arthropods , 5mm in width were found in the burned and salvaged
logged than in either of the logged or mature forest, and more litter-dwelling
species were in the mature forest. Species distribution varied across the plots,
though no clear patterns were identified. Overall, all treatments provided
relatively equivalent arthropod habitat (Greenberg and McGrane, 1996).
Reptile richness, evenness, and diversity were not different across the
treatments or long-unburned mature forest site in this study, but reptile
community composition varied significantly (Greenberg et al., 1994). In all
treatment plots, the reptile community was characterized by open scrub species
and correlated with percent cover of bare sand and other microhabitat
features. More generalist forest reptiles were found in the long-unburned
forest. As a result, silvicultural management appeared to mimic conditions
created by wildfire followed by salvage logging; however, the authors suggest
that the spatial distribution of silviculture may fragment a historically
landscape-scale fire regime, with long-term reptile population impacts (Green-
berg et al., 1994).
Migratory breeding birds and forest birds (canopy and cavity nesters,
canopy and bark foragers) were significantly more common in the mature
forest than in any of the other treatments, while migratory winter residents
were not treatment-dependent in habitat use. These combined patterns resulted
in higher richness and diversity in the mature forest avian community than the
logging or wildfire and salvage log communities (Greenberg et al., 1995a). The
endangered Florida Scrub-Jay was found in similar abundance in all
treatments except the mature forest, where it was absent. High-intensity
logging and site preparation treatments mimicked fire and salvage logging
effects on birds, as it did for the other communities studied (Greenberg et al.,
In the Greenberg studies, no fire-only treatment was available for
comparison. The authors concluded that ‘‘Due to the absence of a ‘‘virgin’’
(unsalvaged) burn treatment or pre-treatment data and the short-term scope of
this study, interpretation of results should be made with caution’’ (Greenberg
et al., 1995, p. 149).
A project by Williges and co-workers (2006; also see Berish et al., 2002)
considered various mechanical treatments (mowing, roller-chopping, logging),
with and without fire, on Florida scrub at three locations (Fig. 2). No fire-only
treatment was included and some of the prescribed fires following mechanical
treatments were patchy. Vegetation was sampled for up to five years post-

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.

treatment. A combination of mechanical treatments and burning produced

more bare ground than mechanical treatments alone. Herb richness was higher
in the mow & burn combination. The study design could not evaluate any
treatment-dependent responses of endemic plants. Increases of weedy species
such as broomsedge (Andropogon virginicus) and the non-native natalgrass
(Rhyncheletrum repens) were attributed to soil disturbance from chopping or
logging and were cause for concern. Treatment effects generally diminished
after five years as the scrub recovered, and mow-only treatments especially
returned to ‘‘an impenetrable thicket’’ by three years. The authors concluded
that there was no evidence of negative long-term effects of mechanical
treatments (Williges et al., 2006).
Scrub restoration techniques have been investigated at Kennedy Space
Center for over 16 years (Schmalzer et al., 2003; Schmalzer and Foster, 2006;
2008). A variety of machines have been used as mechanical treatments (Fig. 3),
and some treatments have been followed by fire. Frequent monitoring showed
that recovery after these treatments was similar to post-fire recovery, with
rapid resprouting and growth of dominant shrubs. However, shrub regrowth
was sometimes faster after mechanical treatments than in periodically burned

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

Sandhill—Sandhill systems are pyrogenic pine savannas that are wide-

spread across the southeastern coastal plain (Peet, 2006). In Florida, they are
dominated by an open longleaf pine (Pinus palustris) overstory, open midstory
with deciduous oaks (primarily Quercus laevis) and other scattered hardwoods,
and a diverse understory of wiregrass (Aristida stricta) and other perennial
grasses and forbs (Myers, 1990; Kirkman et al., 2004). Rare and endemic plant

FIGURE 4. Restoration of sandhill vegetation studied by Provencher and collaborators (2001,

2002). Photographs by Louis Provencher. a. Long-unburned sandhill vegetation dominated by
turkey oak (Quercus laevis). b. Vegetation four years after spring burn. c. Vegetation four years
after herbicide use and fire. d. Vegetation four years after felling and fire.

species are characteristic of sandhill ecosystems (Hardin and White 1989).

Frequent (1–5 year), low intensity fires favor herbs, constrain hardwood
density and cover, and encourage continued frequent fires (Kirkman et al.,
2004). In Florida, sandhill soils are primarily xeric coarse sands, similar to
those that support scrub systems. The fire-dependence of plant (e.g., longleaf
pine, wiregrass) and animal (e.g., gopher tortoise [Gopherus polyphemus])
species integral to this habitat has been well documented (Myers, 1990; Platt et
al., 1988).
Although fire is the preferred tool for sandhill restoration (Provencher et
al., 2001a), multiple fires may be required to achieve these objectives
(Glitzenstein et al., 1995; Drewa et al., 2002). Multiple fires can be effective
in increasing grass and forb cover, reducing litter and lichens, and reducing the
cover of taller woody plants (Reinhart and Menges, 2004). Mechanical
treatments and herbicides have been used in combination with fire in sandhill
Provencher and co-workers (2001a; b) examined the different effects of
mechanical treatments and herbicide application on long-unburned xeric
sandhills in northwest Florida (Fig. 4). The sandhill was overgrown with a
well-developed turkey oak (Quercus laevis) midstory. The goals for vegetation
management included substantial reduction of oak density and cover, increases
in groundcover richness and cover, and increases in longleaf pine seedling
establishment. Three hardwood reduction treatments were tested: growing
season prescribed fire, granular hexazinone herbicide application (1.68 kg a.i./

ha) followed by fire, chainsaw felling or girdling of hardwoods followed by fire,

and a control with no treatment. Responses of vegetation (Provencher et al.,
2001a; b), arthropods (Provencher et al., 2002a), herpetofauna (Litt et al.,
2001), and birds (Provencher et al., 2002b; 2002c), were monitored one year
before treatments and then annually for six years. Each of six blocks had four-
200 ac plots in a completely randomized block design. Reference sites were
frequently burned sandhill, disjunct from the blocked treatments.
The felling/girdling treatment immediately top-killed the most oaks: (93%
versus 69% in the herbicide treatment, and 18% in fire; Provencher et al.,
2001a; b). However, juvenile oak density was reduced the most in all years by
the herbicide treatment (Provencher et al., 2001b). Following herbicide with
fire most reduced oaks in all size classes due to low rates of resprouting. Felling
and girdling treatments followed by fire had 63% oak mortality, while the
single fire treatment had 41% oak mortality. During the same period, 27% of
oaks were top-killed in the frequently burned reference sandhills. The authors
concluded that both fire alone and mechanical treatments followed by fire
required subsequent fire management for hardwood control. For longleaf
pines, seedling mortality was highest after fire but juvenile survival was
minimally influenced by management treatment (Provencher et al., 2001b).
Herbaceous species richness and density were reduced by the herbicide
treatments and remained significantly lower than that in the fire treatment until
they, too, were burned (Provencher et al., 2001a). Fire increased understory
similarity in the treatment plots relative to the reference plots because of
reductions in fine litter and lichens and increases in bare ground cover.
Arthropod communities, like plant species richness, were most similar to
reference sites 1.5 yrs post-fire, regardless of prior treatment (Provencher et al.,
2002a). The arthropod community in the unburned treatments differed from
treatments and the frequently burned reference plots (Provencher et al., 2002a).
The authors suggest that the response of arthropods to fire is critical for
management of sites with the endangered Red-cockaded Woodpeckers because
their diet depends on this group during the breeding season.
Breeding and winter migrant birds responded to structural change; all
hardwood reduction techniques increased the similarity of breeding birds with
that in reference sites (Provencher et al., 2002c). Winter migrant richness was
independent of treatment, but numbers and flock size increased with the degree
of hardwood density reduction (Provencher et al., 2002b).
Herptofauna were only examined post-fire in all treatments; this
community was more similar to the reference condition in all treatment plots
than to the control plots (Litt et al., 2001). As found by Greenberg and co-
workers (1994), discussed above, the herptofauna was dependent on hardwood
density and litter cover, both of which were highest in the control plots.
Overall, fire, applied either alone or in combination with the other
management methods, was necessary to accomplish sandhill management
goals of reducing hardwoods, opening the midstory, and increasing the
similarity of plant and animal communities to the reference condition

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).

(Provencher et al., 2003). Fire increased plant species richness, increased

herbaceous vegetation densities, increased herb-layer arthropods, increased
longleaf pine-associated breeding bird detection rates, and created favorable
(patchy) habitat for herps. In addition, fire was more than eight times less
expensive per acre to apply than either of the other treatments (Provencher et
al., 2001a). The felling/girdling and herbicide treatments sped up restoration by
more rapidly reducing hardwoods, but fire was necessary for the response of
other floral and faunal components (Provencher et al., 2003).
Another study in sandhill, on central Florida’s Lake Wales Ridge, tested
the effectiveness of chainsaw felling in restoring structure and composition of a
long-unburned site (Rickey et al., 2007; Menges et al., 2007). The seven-year
study compared the effects of prescribed fire with (saw & burn treatment) and
without (burn-only treatment) prior felling of the oak subcanopy to an
untreated control (Fig. 5). Restoration goals included retaining canopy pines,
reducing subcanopy and shrub layers, creating more bare sand, decreasing
litter, increasing herb diversity and abundance, and maintaining rare species
Subcanopy felling brought fuels to the surface, where they increased fire
intensity and coverage relative to a burn-only treatment. Longleaf pine survival
was lowest in the saw & burn treatment, intermediate with burn-only, and
highest in control. The oak subcanopy was eliminated by the saw & burn

treatment and reduced with burn-only. Shrub stem densities decreased

temporarily, but then increased in both burn treatments. Both treatments
(but especially saw & burn) increased bare sand and decreased litter, but had
limited effects on herbaceous plants and graminoids. Although the treatments
altered the structure of the sandhill community, species composition was not
dramatically changed. Treatments had variable effects on the demography of
introduced and extant populations of Florida scrub endemics. Menges and co-
workers (2007) suggested the saw & burn pre-treatment could be useful in
‘‘speeding up’’ restoration and segueing to a frequent fire management regime,
but care should be taken to avoid high fire intensities, especially near mature
longleaf pines.
A series of studies in central Florida sandhills also investigated the effects
of a range of management treatments in long-unburned sandhills (Wilkins et
al., 1993a; b; Brockway et al., 1998; Brockway and Outcalt, 2000). Goals
included reducing oaks and releasing wiregrass in areas where insufficient fine
fuels rendered initial fire reintroduction impossible (Wilkins et al., 1993a).
Hexazinone was applied in liquid formulation at 0, 0.42, 0.84, and 1.64 kg/ha
in a randomized block (n53) design, with no fire treatments for comparison.
Oak density declined while cover of wiregrass and other grasses increased with
increasing hexazinone concentration. Oaks .14 cm dbh and saw palmettos
were not affected at any of the applied rates. In a second study applying
granular hexazinone at 0, 1.7, 3.4, and 6.8 kg/ha, again without a fire
treatment, woody cover and diversity and herbaceous species diversity declined
with increasing hexazinone concentration (Wilkins et al., 1993b). The
herbaceous community began to recover by the second year, however, and
differences in cover and diversity among treatments diminished.
Brockway and co-workers (1998) also studied hexazinone application
(granular broadcast at 1.1 kg/ha and liquid spot treatment at 1.1 and 2.2 kg/
ha) as a potential replacement for fire, collecting data pre-treatment and up to
3 years post-treatment, again with no fire treatment. As observed elsewhere,
oak cover decreased (by 83–92%) across the treatments while increasing in the
control plots. Foliar cover of wiregrass and other graminoids increased in all
plots regardless of treatment, although there was a stronger trend in treated
plots than controls. Understory species richness initially declined in the
herbicide treatment plots because of loss of forbs, but then recovered to control
levels by the second post-treatment growing season. Such decline was not
observed in plots treated with a spot-grid (2 m 3 2 m) of liquid hexazinone,
since the active ingredient did not contact the entire plot surface (Brockway,
2009, personal communication).
Four years after the original herbicide treatments, Brockway and Outcalt
(2000) experimentally burned all the study plots of Brockway and co-workers
(1998). Hexazinone followed by fire also resulted in lower cover of all oaks
than fire alone, but no significant differences in cover of woody plants, longleaf
pines, wiregrass, graminoids, or forbs. While fire alone reduced oak and shrub
cover in the first post-fire year, both cover types were significantly higher than

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.

Dry prairie—Dry prairies in Florida were once quite widespread

(Abrahamson and Hartnett, 1990) but have been greatly reduced by
agriculture. They are generally treeless systems dominated by graminoids,
forbs, and sparse patches of shrubs (including Serenoa repens) on acidic,
sandy soils with a spodic horizon that are generally dry in the winter and
flooded in the summer (Watts et al., 2006). The interaction of hydrology and
frequent (1–3 year), low intensity fire is hypothesized to structure this system
(Abrahamson and Hartnett, 1990). Dry prairies are similar to flatwoods, and
pines can colonize dry prairies that are unburned for several years. Dry
prairies in south-central Florida support the endangered Florida Grasshop-
per Sparrow, Ammodramus savannarum floridanus (Shriver and Vickery,
Dry prairie at Myakka River State Park in west-central Florida was
subjected to an experiment that involving repeated roller chopping (every six
years) and fire (every three years) independently, and in combination in two
seasons (summer or winter treatments). The experiment was a randomized
complete block design with four blocks and six treatments. Fitzgerald and
Tanner (1992) found that woody vegetation and bird communities were
affected by season of burn and roller chopping. After one year post-treatment,
they found no differences in shrub richness, but shrub cover in chopped plots
was 50% less than in burned or control plots, regardless of season. Shrubs in
control plots were taller than in any treatment, but there were no significant
differences among treatment shrub heights. More birds were found in control
and burned than chopped treatments. Highest richness and numbers of
shrubland birds were found in the more vertically complex control plots; the
opposite was true for the prairie birds that were the management focus.
Chopping increased numbers of prairie birds ‘‘indicating a trend toward prairie
restoration’’ (Fitzgerald and Tanner, 1992, p. 396).
Watts and colleagues (Watts and Tanner, 2004; Watts et al., 2006) re-
examined the vegetation in these plots after 12–15 years of treatment, with

comparisons to consistently burned reference sites. Overall, density of species

in the burned and chop & burn treatments was most similar to that in the
reference plots regardless of season of treatment. However, forb richness and
grass and forb density were higher in the chop & burn treatments than the
burn-only treatment. Shrub densities were lowest and shrub height was highest
in the chop-only treatment, meeting the restoration objective of reducing
shrubs, but not the objective of increasing the abundance of the herbaceous
species. These results were generally independent of season of treatment,
though responses of individual species varied by both treatment and season.
Treatments also had significant effects on individual species (Watts and
Tanner, 2004). For example, wiregrass cover was favored by a combination of
chopping and burning but occurred at lowest densities following chopping
only. Saw-palmetto densities declined with chopping or chopping and burning
in winter months. Gallberry (Ilex glabra) had lowest densities in burn-only
plots. The authors concluded (Watts et al., 2006, p. 299) that ‘‘a combination
of fire and roller chopping is more effective at restoring fire-suppressed dry
prairie than burning alone.’’

Flatwoods—Pine flatwoods are the most extensive ecosystem in Florida

(Abrahamson and Hartnett, 1990) and vary widely from mesic to hydric. These
open pine savannas usually have a sparse overstory of longleaf pines and may
have understories dominated by wiregrass, palmettos, or shrubs maintained
under 2 m in height through relatively frequent fire (Abrahamson and
Hartnett, 1990). Pine flatwoods are compositionally similar to dry prairies
and can be structurally similar to sandhill vegetation (Glitzenstein et al., 1995).
Frequent, relatively complete fires (Abrahamson and Hartnett, 1990) create
rapid resprouting and recovery in shrubs and herbs (Abrahamson, 1984a; b).
Without fire, grassy vegetation is disadvantaged and shrubs increase (Maliakal
et al., 2000). Restoration of fire-suppressed flatwoods usually involves
returning fire, but may be accompanied by mechanical treatments to reduce
palmettos and limit pine mortality by reducing fire intensity. Drainage of
flatwoods can create opportunities for invasion by xeric shrubs (Yahr et al.,
Also at Myakka River State Park, flatwoods have been treated with fire,
roller-chopping, mowing, and various combinations (Fig. 6) as part of the U.S.
Forest Service fire and fire surrogate study (USDA Forest Service, 2008)
( After two years, growing season burns
had reduced cover and heights of saw palmetto. Combined chop & burn or
mow& burn treatments were even more successful at reducing palmetto cover
and height. Treatments had temporary effects of increasing forb cover in the
first year post-treatment, but these differences were insignificant by the second
year. Grass cover was increased most notably in the chopping treatment.
Treatments including chopping had the greatest effects in reducing shrub
cover, but all treatments reduced shrub cover relative to the control. Chopping

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.

Pine rocklands—Pine rocklands is a south Florida pine forest or savanna

growing on oolitic limestone outcrops. The overstory pine is south Florida
slash pine (P. elliottii var. elliottii). The diverse understory includes shrubs and
herbs of tropical and temperate origins and includes several dozen endemic

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).

DISCUSSION—Inconsistent methods and monitoring—The studies reviewed

here used a variety of mechanical methods and herbicides, applied at different
times and in different combinations. The sites used vary considerably in abiotic
and biotic properties. The overall designs of these studies vary, for example by
the inclusion (or not) of reference sites, a burn-only treatment, and pre-
treatment data. The methods used to measure responses also differ (Table 1).
Sample size is variable, so the finding of no differences in some studies may
reflect inadequate sample sizes. Because of these differences, many of the
reported results are idiosyncratic and not conducive to meta-analysis. Some of
the particular nature of responses is probably inevitable due to spatial and
temporal heterogeneity in soils, herbivores, weather, etc. All this variation
argues for monitoring the effects of mechanical treatments, herbicide
application, and fire at individual sites whenever possible.

What we have learned—Nonetheless, there are several generalities that

emerge from this review.
First, mechanical treatments and herbicides were often successful at
speeding structural changes that would be unlikely with fire alone. These
include situations where fire could not be safely applied with results likely to
meet the management goals, such as under high fuel loads and in areas where
canopy fuels were discontinuous with surface fuels. The non-fire treatments
were especially good at reducing hardwood and palmetto abundance, usually
best in combination with fire. For animals that respond primarily to vegetation

structure, such as birds, responses to mechanical or herbicide treatments were

positive whenever observations were made.
Second, some components of Florida ecosystems responded only to fire.
These include arthropods, herps, and specific plant species. Fire was deemed
essential in all upland ecosystems examined: scrub, sandhill, dry prairie,
flatwoods, and pine rocklands.
Third, most authors agree that mechanical treatments and herbicides are
best used in combination with subsequent fire. Longer-term studies (e.g.,
Provencher et al,. 2001a; Brockway et al., 2005, Watts et al., 2006) found that
fire, subsequent to earlier herbicide or mechanical treatments, continued to
speed restoration. The combination of herbicides and fire also produced the
greatest shift to herbaceous plant dominance in the understory of pine stands
in Louisiana (Miller and Chamberlain, 2008). This result appears consistent in
the fire-adapted communities across the U.S. included in the U.S. Forest
Service fire and fire surrogate study (Schwilk et al., 2009). A recent argument
for using Florida Scrub-Jay as an umbrella species also suggests that
mechanical treatments won’t have the same effects as fire and should be used
largely in combination with fire (Kent and Kindell 2009).
Fourth, mechanical treatments carry risks. Excessive soil disturbance is an
undesirable outcome that has the potential to facilitate the invasion of non-
native plants such as Cogon grass (Imperata cylindrica) (Schmalzer and Foster,
2006; Lippincott, 2000). Exotic invasions have several potential impacts on
native plants, either directly through competition, or indirectly by altering
nutrient and fire regimes (Gordon, 1998; Lippincott, 2000). Soil compaction is
another potential problem with mechanical treatments, but this appears to be
little studied.
Fifth, the useful nature of mechanical pre-treatments to fire in early stages,
as well as the potential problems with repeated mechanical treatments, argues
for their use in a discrete time frame and in the initial stages of restoration. The
prudent goal will be to transition to a fire-only management strategy as soon as
possible. In addition, frequent fires may need to follow up initial mechanical or
herbicide treatments in order to solidify initial progress. For example,
mechanical treatments that are not followed by periodic fires may lose their
effectiveness within a few years or decades (Battaglia et al., 2008; Albrecht and
McCarthy, 2006; Freeman and Jose, 2009).

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.

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.,
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).

Investigations of Florida plants have also found species, including the

endangered sandhill herb Lewton’s polygala (Polygala lewtonii), that show
smoke-stimulated germination (Lindon and Menges, 2008). Vital rates of
several rare plant species in fire-adapted habitats across Florida increase post-
fire (Slapcinsky et al., 2009). The loss of fire from Florida ecosystems in favor
of alternative treatments could alter the population dynamics and viability of
fire-specialist species.
Other unknown issues should affect our restoration efforts. We have
insufficient data to identify differences in fire surrogate effects on sites with
higher versus lower soil moisture. Wetter sites may be more vulnerable to weed
invasion and soil compaction. We also don’t know if mechanical treatment or
herbicide effects vary with the scale of the experiment. Many of the
experiments have necessarily been conducted at smaller scales than occur
under routine ecosystem management (Table 1). Consequently, the results may
not scale-up to the landscape level.
The various unknown effects of fire pre-treatments and surrogates suggest
that simplistic, single-species approaches to management should not be used in
Florida upland ecosystems. Various species, strata, and trophic levels respond
independently to the details of management. For example, many bird species
appear to respond to structural changes regardless of the cause (e.g.,
Provencher et al., 2002b; c), while other groups respond differentially to
different management treatments, indicating that one type of treatment will not
meet the needs of all species. Fire was necessary for recovery of some plant,
arthropod, and herp species (Provencher et al., 2001a; b; c; Litt et al., 2001).
This finding requires caution when management objectives focus on single
species (e.g., longleaf pine, Florida Scrub-Jay, Bobwhite Quail).
As mechanical treatments and herbicide use become more popular, and as
natural areas become more greatly impacted by adjacent suburban and urban
developments, the prevalence of prescribed fire as a management tool may be
diminished in many areas. What would be the effect of losing his basic
ecosystem process? Will Florida ecosystems lose key components and will this
have a cascading effect on ecosystem processes like decomposition? Will the
systems lose resilience in the face of pressure from exotic species and climate
From this review, we make the following recommendations on the use of
mechanical treatments and herbicides to restore natural ecosystems in Florida:
1. More research is needed on repeated mechanical treatment and herbicide
effects on ecosystems, especially the long-term effects of these treatments,
before these approaches are uncritically applied to large areas of Florida
natural areas. These research efforts should collect pre-treatment data,
monitor treatment intensities, and use reference sites to provide context.
2. Mechanical and herbicide treatments should be used only in the initial
phases of ecosystem restoration, and should be followed by prescribed
fire. If repeated prescribed fires are not used, then the positive effects of

the initial mechanical treatments or herbicides may be lost due to vigorous

recovery of resprouting species.
3. When fire alone is feasible and will accomplish restoration goals, repeated
fire within the full range of the natural regime should be implemented.
4. Management based on conservation of a single species is not recom-
mended. As a result, recommendations that species like the Florida Scrub-
Jay should be considered as an umbrella species for management of
peninsular Florida scrub (Kent and Kindell, 2009) may be interpreted too
simplistically. As several studies show that different groups (e.g., birds vs.
vegetation) respond differently to various management treatments,
management for an umbrella species needs to be demonstratively
beneficial for other species as well. Spatial and temporal heterogeneity
in management (including variation in the fire regime) are likely to benefit
a greater variety of species than a single-species, invariant approach. Such
an approach also provides a bet-hedging strategy against our ignorance of
management effects.

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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Florida Scient. 73(2): 147–174. 2010

Accepted: September 1, 2009
Florida Academy of Sciences. 2010