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TECHNICAL REPORT
Landscape and Watershed Processes

Differential Effects of Surface and Peat Fire on Soil Constituents in a Degraded Wetland of the Northern Florida Everglades

Everglades Dep., Watershed Research and Planning Division, South Florida Water Management District, 3301 Gun Club Road, West Palm Beach, FL 33406

* Corresponding author (smsmith{at}sfwmd.gov)

Received for publication January 22, 2001.

	ABSTRACT

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION SUMMARY AND CONCLUSIONS REFERENCES

 
The effects of surface (aboveground) and peat (belowground) fire on a number of soil constituents were examined within a hydrologically altered marsh in the northern Florida Everglades. Peat fire resulted in losses of total carbon (TC), total nitrogen (TN), and organic phosphorus (P_o), while inorganic phosphorus (P_i) and total calcium (TCa) concentrations increased. In addition, peat fire led to a more pronounced vertical gradient in constituent concentrations between upper and lower soil layers. Surface fire also affected soil constituents, but impacts were small relative to peat fire. The effects of physical versus chemical processes during burning were assessed using ratios of constituent to TCa concentrations. This measure indicated that increases in the levels of total phosphorus (TP) in peat-burned areas were due primarily to the physical reduction of soil, while decreases in TN and TC were the result of volatilization. Increases in concentrations of P_i fractions arose from both chemically and physically mediated processes. In an ecological context, the observed soil transformations may encourage the growth of invasive plant species, such as southern narrow-leaved cattail (Typha domingensis Pers.), which exhibits high growth rates in response to increased P availability.

Abbreviations: BD, bulk density • CaMg-P_i, calcium- and magnesium-bound phosphorus • FeAl-P_i, iron- and aluminum-bound phosphorus • L-P_i, labile inorganic phosphorus • P_i, inorganic phosphorus • P_o, organic phosphorus • RWMA, Rotenberger Wildlife Management Area • TC, total carbon • TCa, total calcium • TN, total nitrogen • TP, total phosphorus

	INTRODUCTION

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION SUMMARY AND CONCLUSIONS REFERENCES

 
FIRE plays an important role in both the preservation and evolution of wetland ecosystems (Cohen, 1994; Kushland, 1990; Wade et al., 1980). However, studies on the effects of fires in wetlands are few compared with terrestrial habitats. Moreover, a disproportionate amount of research on wetland fires has focused on the recovery of flora and fauna relative to abiotic changes that can subsequently influence these processes (Abrahamson, 1984; Lee et al., 1995; Smith and Kadlec, 1985; van Arman and Goodrick, 1979; Vogl, 1973).

Typically, fire alters the concentrations of soil nutrients (Faulkner and de la Cruz, 1982; Vogl, 1969; Wilbur and Christensen, 1983), although the nature of these alterations can be quite variable. In some cases, nutrient levels increase after fire (Kutiel and Shaviv, 1993; Marion et al., 1991; Tomkins et al., 1991), while in others no enrichment occurs (Laubhan, 1995; Weiss, 1980). Alternatively, nutrients may be lost through volatilization, leaching, and/or export of ash particles by updrafts during the burning process and wind (Marion et al., 1991; Turner et al., 1997). The extent of soil modification varies with fire severity, fire temperature, fuel moisture levels, fuel loads in the soil, plant tissue nutrient concentrations, plant tissue combustibility, and other environmental factors (Gunderson and Snyder, 1994; Kutiel and Shaviv, 1989; Robichaud and Waldrop, 1994; Scott and Van Wyk, 1990).

In wetlands, fires often consume only a portion of aboveground vegetation, while moisture-laden roots and soils layers remain unharmed (i.e., surface fire). Upon soil dry-out, however, the combustion of soil organic matter can occur (i.e., peat fire), which can completely destroy the existing vegetation and eliminate some portion of the resident seedbank. In the Florida Everglades, surface fire has contributed to the persistence of sawgrass (Cladium jamaicense Crantz) prairies, the dominant habitat within the marsh, by periodically eliminating successional species (Loveless, 1959). However, the frequency and severity of fire events in the Everglades are thought to have been altered by changes in hydrology over the last several decades (Alexander and Crook, 1974; Gunderson and Snyder, 1994; Robertson, 1953). In certain regions, diversion of surface water toward agricultural and urban areas has resulted in soil desiccation and an increased occurrence of peat fire, followed by the establishment and expansion of opportunistic plant species such as southern narrow-leaved cattail (Christensen and Burrows, 1986; Davis, 1943; Gunderson and Snyder, 1994).

During peat fire, the combustion of soil organic matter lowers ground elevations and increases water depth and hydroperiod, which may impart a competitive advantage to species (e.g., cattail) with a high degree of flood tolerance (Grace, 1989; Newman et al., 1996). However, changes in soil nutrient concentrations may play an even greater role in determining subsequent plant community composition (Newman et al., 1998). Unfortunately, it is difficult to predict the time and location of fire events. Thus, the interpretation of their effects is often constrained by a lack of baseline (i.e., pre-fire) data. Additionally, comparisons of pre- and post-fire data that have been collected from different locations can be confounded by spatial variability in soil properties. In the summer of 1999, we had the opportunity to compare the effects of peat and surface fire on soils within a northern Everglades marsh from which baseline data had been previously obtained. Our objective was to determine the extent of soil transformation (particularly to major nutrients) caused by each type of fire. Based on soil chemical properties, we suspected that total P would be conserved, although mineralization to inorganic forms would occur after both surface and peat fire. Conversely, nitrogen and carbon would probably decrease through volatilization.

	MATERIALS AND METHODS

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION SUMMARY AND CONCLUSIONS REFERENCES

 
Study Area
The Rotenberger Wildlife Management Area (RWMA) encompasses approximately 120 km² of land in the northwest Everglades (Fig. 1) and was originally characterized as a sawgrass-plains landscape (Davis, 1943). Since the mid-1950s, however, the area has been cut off from natural surface water flows by the construction of perimeter levees. Consequently, water levels in the RWMA are dependent upon direct rainfall, which is insufficient to prevent the severe dry-out that now occurs on an annual basis (Newman et al., 1998; Smith et al., 2000). These conditions have facilitated the gradual enrichment of RWMA soils through oxidation and compaction—a process that is implicated in cattail invasion of the area, despite its short hydroperiod. Furthermore, the distribution of cattail in the RWMA is correlated with the boundaries of previous peat-burns (Newman et al., 1998; Sasse, 1999).

View larger version (31K): [in this window] [in a new window]   Fig. 1. Map of South Florida showing the Rotenberger Wildlife Management Area (RWMA) and sampling locations (closed circles) (major canals and levees of South Florida's water management system are included).

On 23 May 1999, a lightning strike ignited a fire in the marsh. Both surface and peat burning was evident across most of the RWMA, including the locations where soils had previously been sampled. Areas that experienced peat fire were light gray in color, completely devoid of vegetation, and lower in elevation. Surface-burned areas were covered with the charred shoot bases of plants and were gray-black in color. On 2 June 1999, the pre-fire sampling sites were relocated by their remnant aluminum markers and soils were collected and analyzed in the manner described above. No standing water was present at the time of pre- or post-fire collections.

Data Analyses
The sampling sites were located in areas of sawgrass, cattail, and mixed graminoid vegetation. Both types of fire occurred in each of these community types. Accordingly, the data (both pre- and post-fire) were grouped according to burn type (i.e., surface-burned, n = 27; peat-burned, n = 34). As such, the complete set of samples was comprised of soils from the following categories: (i) pre-fire, surface-burned area, (ii) post-fire, surface-burned area, (iii) pre-fire, peat-burned area, and (iv) post-fire, peat-burned area. The number of post-fire samples analyzed for specific P fractions was reduced by 10 in both surface- (n = 17) and peat-burned (n = 24) soils due to analysis costs. The constituent data were expressed on a volume basis (µg/cm³), which allows interpretation from an ecological perspective since plant root systems occupy soil in three-dimensional space rather than by weight.

The physical concentration of soil layers into smaller volumes (observed as soil loss) occurs during peat fire. Without an accurate measure of the amount (depth) of soil lost, it is impossible to determine the quantity of nutrients volatilized or to account for any concentration increases caused by the consolidation of remnant soil (ash) into a smaller volume. Therefore, we compared nutrients of interest with TCa since this element did not have a depth gradient within the top 20 cm of soil (Smith et al., 1999) and has a boiling point of 1440°C (Merck, 1989). Thus, TCa would not be lost through volatilization and would increase in the upper layers as it is retained within a smaller volume of soil, essentially behaving as a conservative tracer. Constituent to TCa ratios were calculated to provide a relative measure of how concentration changes were related to physical (i.e., concentrating mechanism of soil loss) and chemical (i.e., volatilization or mineralization) processes.

For statistical analysis, the data were transformed logarithmically to improve normality and heteroscedasticity and subjected to analysis of variance (split-plot design). Spatial (surface- vs. peat-burned) and temporal (pre- vs. post-fire) comparisons between mean values were examined using Statistical Contrasts (SAS Institute, 1989).

	RESULTS AND DISCUSSION

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION SUMMARY AND CONCLUSIONS REFERENCES

 
Pre-Fire Spatial Variation
In both core layers, the majority of pre-fire constituent concentrations were similar between areas that subsequently experienced surface or peat burning (Fig. 2 and 3) . Only TP and CaMg-P_i in the 2- to 10-cm layer were significantly different, in that pre-fire concentrations were lower in soils that underwent peat burning.

View larger version (31K): [in this window] [in a new window]   Fig. 2. Comparisons of bulk density (BD), ash content (% Ash), total nitrogen (TN), total carbon (TC), and total calcium (TCa) in the 0- to 2- and 2- to 10-cm layers of surface-burned (open histograms) and peat-burned (shaded historgams) soils (bars are means + 1 standard error; histograms with shared letters are not significantly different).

View larger version (29K): [in this window] [in a new window]   Fig. 3. Comparisons of total P and P fractions in the 0- to 2- and 2- to 10-cm layers of surface-burned (open histograms) and peat-burned (shaded historgams) soils (bars are means + 1 standard error; histograms with shared letters are not significantly different). CaMg-Pi, calcium- and magnesium-bound phosphorus; FeAl-Pi; iron- and aluminum-bound phosphorus; L-Pi, labile inorganic phosphorus; Po, organic phosphorus; TP, total phosphorus.

Peat Fire Effects
In the 0- to 2-cm layer, BD, percent ash, TCa, TP, L-P_i, and CaMg-P_i showed significant increases from peat burning, whereas FeAl-P_i remained unchanged and P_o was reduced nearly threefold (Fig. 2 and 3). Total nitrogen and TC concentrations also decreased, although the changes were not statistically significant (Fig. 2). In the 2- to 10-cm layer, post-fire BD, percent ash, L-P_i, and CaMg-P_i were significantly higher than pre-fire values. Conversely, major reductions in TC, TN, FeAl-P_i, and P_o were observed and TCa and TP concentrations remained unaltered (Fig. 2 and 3).

Constituent to Calcium Ratios in the 0- to 2-cm Layer
The soil TN to TCa ratio was only slightly reduced from 0.86 to 0.80 following surface fire (Table 1). The TC to TCa ratio responded similarly with only a small decrease from 11.94 to 10.71. In contrast, large reductions occurred from peat burning. For example, the TN to TCa ratio decreased from 1.13 to 0.31 and TC to TCa ratio from 15.28 to 3.55. The TP to Ca ratios showed little response to either surface or peat fire (Table 1). However, the P_o to TCa ratio decreased from 15.10 to 1.36 after peat burning. In surface-burned soils, this reduction was less pronounced, with values declining from 11.92 to 6.37. The CaMg-P_i to TCa ratio exhibited a large increase from 5.97 to 18.50 in response to peat burning but changed little after surface burning. The L-P_i to TCa ratio increased from 0.03 to 0.18 in surface-burned soils but was virtually unchanged by peat fire.

View larger version (31K): [in this window] [in a new window]   Fig. 4. Proportions of inorganic and organic P fractions in soils (0- to 2-cm layer) from pre-fire surface-burn, post-fire surface-burn, pre-fire peat-burn, and post-fire peat-burn areas. CaMg-Pi, calcium- and magnesium-bound phosphorus; FeAl-Pi; iron- and aluminum-bound phosphorus; L-Pi, labile inorganic phosphorus; Po, organic phosphorus.

Although interpretation of the data is complicated by the relative influence of chemical vs. physical transformations, constituent to TCa ratios provided a means by which these two aspects of change could be resolved. The decrease in soil TN to TCa and TC to TCa ratios in the 0- to 2-cm layer following peat fire indicates that both N and C were volatilized. In contrast, the lack of change in TP to Ca ratio indicates that TP was conserved during the fire and that upper-layer concentration increases were the result of soil loss and consolidation. However, the form of P was affected chemically. Specifically, peat fire resulted in the conversion of P_o to P_i, as evidenced by decreases in P_o to TCa ratio and increases in CaMg-P_i to TCa ratio. Similar responses have been observed in other habitats affected by fire (Saa et al., 1993). In surface-burned soils, there was a much smaller decrease in P_o to TCa ratio, and the P_i to TCa ratio changed very little given that L-P_i, which increased, was such a small component of the total P_i pool. Furthermore, the TN to TCa and TC to TCa ratios showed no evidence of loss from surface burning. Interestingly, mass of TCa (in grams) in the 0- to 2-cm layer of post-fire peat-burned samples divided by the amount in the 0- to 10-cm layer (0- to 2-cm layer + 2- to 10-cm layer) of pre-fire peat-burned samples produced a mean value of 1.05. This indicates that on average there was just slightly more TCa in the 0- to 2-cm layer after peat fire as there was in the 0- to 10-cm layer before the fire. In other words, roughly 10 cm of soil collapsed into 2 cm.

As a more general characteristic, relative differences between upper- and lower-layer concentrations of many constituents increased considerably after peat fire. This was due to (i) increases in post-MB upper layer concentrations from chemical conversion and/or physical consolidation and (ii) decreases in post-MB lower-layer concentrations (e.g., TP), which, as a result of soil loss, may reflect preexisting depth gradients. Regardless of the mechanism, from an ecological standpoint the transformed soil profile represents the new environment in which revegetation must occur. In surface-burned areas, much smaller differences between layers were observed (with the exception of L-P_i). Accordingly, vertical heterogeneity of this nature may be useful in characterizing the fire history of soils.

In the Everglades, P availability is a critical factor regulating plant species distributions (Davis, 1994). The natural peat soils of this ecosystem are characterized by extremely low concentrations of P, most of which is present in organic forms (Craft and Richardson, 1997; Reddy et al., 1998). This favors plants like sawgrass that have low P requirements and do not respond strongly to fluctuations in P availability (Davis, 1989, 1991; Miao and Sklar, 1998; Steward and Ornes, 1975). Thus, increasing the amount of bioavailable P (i.e., inorganic forms) through mineralization and physical concentration provides an advantage for species with high affinities for this nutrient. Under laboratory conditions, Rivard and Woodard (1989) reported that the nutrients in common cattail (Typha latifolia L.) ash stimulated seed germination whereas sawgrass seed is apparently unresponsive to nutrient enrichment (Lorenzen et al., 2000; Ponzio, 1998). Smith and Newman (2001) recently conducted bioassays with seedlings of southern narrow-leaved cattail, which revealed that peat-burned soils (from the RWMA) were a better growth medium compared with surface- or non-burned soils as evidenced by higher biomass and tissue P concentrations.

Typha spp. distributions have also been positively correlated with Ca concentrations (Adriano et al., 1980; Volk et al., 1975). In this study, TCa became highly concentrated in the upper layers of peat-burned soils. Thus, peat fire can redistribute other soil elements (besides P) in a way that may ultimately benefit the growth of cattail. The added influence of lowered ground elevation and the absence of plant competition following peat burning may further optimize conditions for cattail establishment.

	SUMMARY AND CONCLUSIONS

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION SUMMARY AND CONCLUSIONS REFERENCES

 
The relative value of this information is enhanced by the fact that (i) pre-fire baseline data were available for comparison, (ii) confounding spatial variability was reduced by sampling at the same locations, (iii) two distinct fire types (surface and peat) could simultaneously be compared, and (iv) analysis of TCa concentrations as a tracer allowed the resolution of physically vs. chemically mediated concentration changes. In general, surface fire resulted in minor changes to soil constituents while peat fire physically concentrated large amounts of TCa and TP into the upper soil layer. Peat fire also increased the proportion of bioavailable P (P_i) while volatilizing TN and TC. These kinds of changes in soil properties, especially nutrients, may play an important role in the reestablishment and eventual character of plant communities following fire disturbance.

	REFERENCES

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION SUMMARY AND CONCLUSIONS REFERENCES

 

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