WETLANDS, Vol. 29, No. 2, June 2009, pp. 520–526 ’ 2009, The Society of Wetland Scientists FIRE IN FLOODPLAIN FORESTS IN THE SOUTHEASTERN USA: INSIGHTS FROM DISTURBANCE ECOLOGY OF NATIVE BAMBOO Paul R. Gagnon Louisiana State University Department of Biological Sciences 202 Life Sciences Building Baton Rouge, Louisiana, USA 70803 Current address: University of Florida Department of Wildlife Ecology and Conservation 110 Newins-Ziegler Hall Gainesville, Florida, USA 32611 E-mail: [email protected] Abstract: Floodplain forests in the southeastern USA have recently been the focus of intensive restoration efforts after centuries of human-caused decline. Many of these restored forests appear to suffer from systemic problems arising from the altered disturbance regime in modern southeastern floodplains. Increasing evidence suggests that fire may be an occasional but important ecosystem component missing from these forests. Most relevant literature mentions fire only in passing, if at all; the literature that does discuss fire is typically either speculative or draws heavily from other ecosystems. This article develops the hypothesis that fire has been an important and recurrent disturbance in southeastern alluvial floodplains for millennia. It first synthesizes research indicating that the expansive monodominant bamboo stands (called canebrakes) once common throughout these floodplain forests were likely fire-obligate and might therefore be used as indicators of recurrent fires. It then examines prehistoric, historic, and recent evidence of fire in bottomland forests from both natural and human sources. Finally, it places these findings into ecological context, proposes an integrated study by which future research might clarify the ecological role of fire in southeastern floodplain forests, and addresses some implications for management. Key Words: Arundinaria gigantea, bottomland hardwood forests, canebrakes, multiple disturbance interactions, fire ecology, hurricanes, tornados, windstorms INTRODUCTION water quality maintenance, flood control, and habitat for diverse wildlife species, BLH have increasingly been the focus of reforestation efforts since then (Stanturf et al. 2001, King et al. 2005, Wilson et al. 2007). As a result, the amount of land reforested with BLH tree species has recently increased substantially, and additional reforestation projects are planned (King et al. 2006). Reforestation cannot ensure fully functioning restoration of these bottomlands (King et al. 2005). Reasoned and sometimes intensive efforts are necessary for effective restoration because of the extent to which humans have changed underlying ecological processes throughout the BLH ecosystem (Stanturf et al. 2001, Wilson et al. 2007). To return full function, it is necessary to restore the relevant disturbance regime (Wilson et al. 2007). Large disturbances in BLH included windstorms, ice storms, droughts, floods, and fires (King et al. 2005, Wilson et al. 2007). After flooding, fire Bottomland hardwood forests (BLH) in the southeastern U.S. are rebounding after two centuries of radical alteration to alluvial floodplains. Prior to European settlement, these forests covered perhaps 40–50 million hectares of what is now the southeastern United States (The Nature Conservancy 1992). That amount has been reduced by 60% overall, much more in certain regions like the lower Mississippi River alluvial valley (Noss et al. 1995, Stanturf et al. 2001). During the 19th and 20th centuries, land-clearing for industrial agriculture, intensive logging and river channelization combined to take a heavy toll on BLH forests (King et al. 2005, Saikku 2005). When high soy prices drove another round of forest clearing in the 1970s, area of BLH hit an all-time low (Stanturf et al. 2001). Because of the many products and ecosystem services BLH provide, including valuable timber, 520 Gagnon, FIRE IN BOTTOMLAND HARDWOOD FORESTS regimes are most likely to have been fundamentally altered, and fire may be the most often overlooked ecological and evolutionary force in BLH. Systemic problems are recognized with regenerating BLH that arise from their altered disturbance regimes. Wilson et al. (2007) note that ‘‘Reduced levels of disturbance acting in concert with unsustainable forest management practices have resulted in homogeneous, closed canopy forests with little structural diversity or understory vegetation.’’ Canopy structure in modern BLH is altered, as evidenced by poor regeneration of several shadeintolerant species valued for their timber and mast, a critical food source for wildlife. Recent research indicates that flooding alone will not ensure the regeneration of young bottomland oaks (King and Antrobus 2005), which need large or recurrent gaps to reach the forest canopy (Oliver et al. 2005, Holladay et al. 2006, Collins and Battaglia 2007). There is also evidence that oak-dominated forests historically established after fire (Aust et al. 1985, Hodges 1998), and that certain other bottomland assemblages were maintained by fire (e.g., Hughes 1957, Ewel 1998, Gagnon and Platt 2008). Most discussions in the literature about fire in BLH are either speculative or extrapolate from other systems (e.g., Sharitz et al. 1992, Myers and Van Lear 1998). In this article I examine the hypothesis that fire has been an important and recurrent disturbance in southeastern alluvial floodplains for millennia. I first synthesize research indicating that expansive monodominant bamboo stands (called canebrakes) once common throughout BLH were likely fire-obligate, and might therefore be used as indicators of recurrent fires. Next, I examine pre-historic, historic, and recent evidence of fire in BLH, from both natural and human sources. Finally, I put these findings into ecological context, propose an integrated study by which future research might clarify the ecological role of fire in BLH, and discuss some related management implications. CANEBRAKES AS DISTURBANCE-OBLIGATE MONODOMINANT COMMUNITIES Canebrakes are monodominant stands of Arundinaria bamboo, a genus endemic to the United States. Of the three recognized species of Arundinaria, it is A. gigantea (Walt.) Muhl. (giant cane or river cane) that frequently occurs on higher ground within southeastern alluvial floodplains (Judziewicz et al. 1999, Triplett et al. 2006). Culms of A. gigantea can exceed 6 m in height and 3 cm in diameter; they 521 attain their largest size in the most fertile soils (Saikku 2005, Stewart 2007). Giant cane today is a common understory and edge component in BLH (Marsh 1977, Kellison et al. 1998, Judziewicz et al. 1999). Like other woody bamboos, A. gigantea is thought to be semelparous, meaning that after decades of vegetative growth, it flowers once and then dies (Judziewicz et al. 1999). Nowadays giant cane grows in small patches along forest edges or diffusely under forest canopy (Marsh 1977, Gagnon et al. 2007). Early European explorers describe vast, dense stands of giant cane frequently growing on levees and ridges in southeastern alluvial floodplains (Platt and Brantley 1997, Stewart 2007). John James Audubon describes how canebrakes were typically too thick to traverse even on foot unless one pushed through hunched-over and backwards (Platt et al. 2002a). Floods influenced giant cane and other species in BLH over time. Floods occurred during most years in southeastern floodplains (Lentz 1931, Hodges 1998, Kellison et al. 1998). Their impact on vegetation varied greatly depending on local hydrology and microtopography (Hodges 1998, Kellison et al. 1998, Saikku 2005). On natural levees and ridges where giant cane was common, floods were occasional, intermittent events (Kellison et al. 1998). Giant cane could tolerate these intermittent floods (Cirtain et al. 2004), although not prolonged inundation, and may have benefited from the new space and sediment that accompanied floods (Gagnon and Platt 2008). Violent storms also shaped giant cane populations in BLH. Tornados, hurricanes, severe thunderstorms, and ice storms all damaged the canopy of BLH and thereby increased light levels in the understory. Some of these events snapped single branches or tipped-over individual trees; others removed forest canopy over large areas (Ker 1816, Sharitz et al. 1992, Gagnon et al. 2007). In the high light environment of large, wind-generated forest gaps, rates of new culm production in A. gigantea could be twice that under forest canopy (Gagnon et al. 2007). Droughts may have interacted with windstormdamaged landscapes to create conditions favorable for fires. Windstorm and fire interactions have been documented for other southeastern ecosystems (Platt et al. 2002b, Liu et al. 2008), and could have also been important in BLH. Fires are reported in BLH during drought years in large re-vegetating gaps (Lentz 1931, Kaufert 1933). Over ecological time, droughts in BLH were probably linked to ENSO weather patterns, which have cycled on a 3–7 year interval for the last 130,000 years (Tudhope et 522 al. 2001, Beckage et al. 2003). High frequency of lightning strikes has been pervasive across the southeastern U.S. (Platt 1999). Lightning from spring storms following a dry period could have ignited fires in windstorm-damaged areas (Myers and Van Lear 1998, Gagnon and Platt 2008). Human-caused fires probably augmented the number of fire events affecting giant cane in southeastern alluvial floodplains. Stand structure in Arundinaria bamboos is largely a function of disturbance regime (Hughes 1957, Gagnon et al. 2007, Gagnon and Platt 2008). Over a few years, stands of Arundinaria growing in large gaps gradually decline, perhaps a result of selfcrowding or self-shading (Hughes 1957, 1966, Gagnon and Platt 2008). A new disturbance like fire will typically reinvigorate declining Arundinaria stands (Hughes 1957, 1966, Wright and Bailey 1982). Without periodic disturbances to open growing space, stands of giant cane will decline and trees of BLH will eventually overtop them (Wright and Bailey 1982, Gagnon et al. 2007), precluding the formation of expansive monodominant canebrakes like those described by so many early European explorers (Platt and Brantley 1997). In open-grown stands of giant cane, fire clears competing vegetation and accelerates new culm production while eliminating senescing older culms (Gagnon and Platt 2008). The link between fire and canebrake-like stand structure (i.e., dense and expansive monodominant bamboo) in Arundinaria is tight enough that historical canebrakes can serve as indicators of recurrent fires (Wright and Bailey 1982, Frost 2000). FIRE DURING THE PRE-HISTORIC PAST Evidence suggests that fires recurred in southeastern alluvial floodplains for millennia. Palynological records of herbaceous Chenopodiaceae and Amaranthus pollen in the Yazoo River floodplain indicate open, potentially burned areas were present in BLH forests since the last glaciation (Saikku 2005). In the Western Lowlands of the Mississippi River alluvial valley, Arundinaria thickets first appeared in the pollen record around 9,500 years ago (Royall et al. 1991, King et al. 2005). Beginning approximately 5,000 years ago, climatic patterns became like modern ones (Saikku 2005), so lightning would have been frequent across the landscape. The pollen record indicates a concurrent shift from oakto pine-dominated plant communities along the northern Gulf Coast, meaning fire frequencies likely increased on the landscape level (Delcourt and Delcourt 1987, King et al. 2005). WETLANDS, Volume 29, No. 2, 2009 Native Americans inhabited and set fires in BLH for thousands of years (Komarek 1974, Sharitz et al. 1992, Saikku 2005, Mann 2006). Evidence of Native American occupancy in the lower Mississippi River alluvial valley first appears 12,000 years ago (Saikku 2005, King et al. 2005). Human populations fluctuated in the floodplains over time, and these early inhabitants survived by hunting different types of large and small animals, and by gathering oak/ hickory mast and various understory flora (Saikku 2005, King et al. 2005). During the early Archaic period (9,000 to 5,000 years ago), drier climatic conditions developed, and more people moved into the bottomlands as resources grew scarcer in adjacent uplands (Saikku 2005). Modern climatic patterns during the middle Archaic period (5,000– 2,500 years ago) coincided with intensifying exploitation of riverine resources, growth of interregional trade, and development of cultigens and ceramic containers (Saikku 2005). The development of Poverty Point in northeastern Louisiana (3,600– 2,600 years ago) is clear evidence that large human populations were living in permanent habitations within and alongside southeastern floodplains during that period (Saikku 2005, Mann 2006). Over time, Native Americans grew increasingly reliant on horticulture (Saikku 2005, King et al. 2005). Cornbased agriculture was introduced in the lower Mississippi River alluvial valley around 400 A.D. and quickly became an important means of subsistence (Saikku 2005). Human activities caused intensifying changes in flora and fauna during this period, mainly from their use of fire and stone wood-cutting tools to clear patches of BLH for garden plots (King et al. 2005). The Mississippian culture arose around 900 A.D. and lasted until the first Europeans arrived (Saikku 2005). These people were reliant on corn-based agriculture and on fire for clearing land. Sedentary riverine villages with sociopolitical organization were located around ceremonial centers that included large temple mounds. For their fields, people preferred the natural levees on recently abandoned channels because the sandy loam there was easily tilled using hand tools, and because these high grounds were the last to flood (Saikku 2005). When fields lost fertility, people cleared new ones by girdling trees and setting fires (Morse and Morse 1983, Saikku 2005). Their primary game animal was the white-tailed deer (Odocoileus virginianus Zimm.), and tribes maximized the edge habitat that deer prefer by setting fires (Saikku 2005). Analyses of faunal remains indicate decreasing arboreal species and increasing edge species during this period (Saikku 2005). Gagnon, FIRE IN BOTTOMLAND HARDWOOD FORESTS The first Europeans brought waves of epidemics that killed perhaps 80–90% of Native American populations, and caused their societies to collapse (Platt and Brantley 1997, Saikku 2005, Mann 2006). Native American occupation of the lower Mississippi River alluvial valley ended during the first half of the 18th century after two centuries of epidemics and slave-raiding. Reforestation occurred naturally after that, and the area remained an occasional hunting ground for dispersed remnant survivors and for upland tribes (Platt and Brantley 1997, Saikku 2005). FIRE AND EUROPEAN SETTLEMENT Accounts by European explorers of southeastern bottomlands strongly suggest that recurrent presettlement fires had shaped that habitat. There are numerous descriptions of large treeless areas in the floodplains (Platt and Brantley 1997, Saikku 2005, Stewart 2007). For example, early French accounts and 19th century surveyor field notes both mention large grasslands or ‘‘prairies’’ on high ground within the Yazoo floodplain (Saikku 2005). Without fire or some other recurrent space-opening disturbance, southeastern alluvial floodplains would have quickly succeeded into BLH forests (Kuchler 1964, Sharitz et al. 1992). According to many early accounts, canebrakes were pervasive on alluvial high ground in the Southeast (Platt and Brantley 1997). They were so common that any mention of canebrakes was typically off-hand (Stewart 2007). For example, Thomas Nutall, described how along the Mississippi River, ‘‘vast tracts of cane land occur in the bends’’ (Saikku 2005). Travelers often sought canebrakes for evening camps because Arundinaria was a highly nutritious fodder for their horses (Wright and Bailey 1982). The naturalist William Bartram made note of canebrakes as he documented vegetation in the southeastern U.S. between 1773 and 1777 (Harper 1958). He mentions ‘‘cane’’ (Arundinaria) dozens of times in his ‘‘Travels,’’ often in reference to expansive canebrakes (Harper 1958). The ubiquity of canebrakes in southeastern floodplains was a strong indicator of recurrent fires (Wright and Bailey 1982, Frost 2000). European settlers used fire to clear land, and they sought out canebrakes for their fields. The first wave of settlers drove their herds of cattle, horses and pigs into the bottomlands, where they feasted on nutritious canebrakes (Platt and Brantley 1997, Stewart 2007). During a second wave, which peaked between 1840 and 1860 in the Mississippi River alluvial valley, would-be cotton growers 523 targeted cane lands as indicators of rich soil and relative safety from flooding (Platt and Brantley 1997, Saikku 2005). Planters systematically eradicated most canebrakes by chopping the culms, pulling up the rhizomes, and then burning several days later once the cane was dry (Stewart 2007). They cleared forested lands by girdling trees and burning, and their fires frequently escaped (Saikku 2005). FIRE AND THE RECENT PAST Alteration of BLH accelerated after the Civil War. In the late 19th century, intensive logging of southern forests began in earnest (Sharitz et al. 1992, Saikku 2005). Ditching and levee building accelerated shortly thereafter, drastically changing the natural flood regime (King et al. 2005, Saikku 2005). Millions of hectares were logged and then permanently converted to cotton, corn, and soy fields (King et al. 2005, Saikku 2005). By the middle of the 20th century, cyclical flooding in many BLH had ceased entirely. Remaining BLH forests were highly fragmented and had greatly altered hydrology. Accounts from several early forest researchers describe how fires occurred in BLH forests in the recent past. G. H. Lentz, a forester from the New York College of Forestry at Syracuse on sabbatical with the Southern Forest Experiment Station (Saikku 2005), describes a recurrent ‘‘fire problem’’ in Mississippi bottomland forests in 1931: The summer of 1924 was characterized by an insufficient amount of rainfall throughout the lower Mississippi Delta. A dry fall and winter followed. Many of the usually moist and wet areas in the bottomlands had dried out by the early spring of 1925 and many serious fires resulted. Although foresters and timbermen generally discount the fire problem in the southern hardwoods, a real problem exists… The past summer, 1930, was decidedly dry and was followed by a fall and winter also quite dry. Many sloughs and brakes usually filled with water have been dry for several months. The spring rains have not been able to wet down the litter and debris and conditions in the woods are just right for fires to spread. Lentz describes fires in a stand that had been clearcut three years prior. He notes that the fire was hot enough to kill virtually all advanced regeneration and to severely damage several full-size trees. He notes other fires burning simultaneously in ‘‘both cut and uncut’’ stands of timber. He concludes by saying that ‘‘If timber growing on an appreciable scale is to be practiced in the bottomlands, the 524 WETLANDS, Volume 29, No. 2, 2009 solution of the fire problem is going to be one of the first essential steps.’’ F. H. Kaufert (1933) followed Lentz’s report with a study of fire scars in BLH of the Mississippi River alluvial valley in Louisiana. Kaufert examined over 500 stumps from three separate logged tracts and documented recurrent fire damage predating the Civil War. He states that ‘‘Information obtained from many ring counts indicates that widespread burning of the bottoms occurred in the years 1898– 99, 1911–12, 1916–17, and 1924–25,’’ and lesswidespread fires in 1931–32, an interval that ranges from 5 to 13 years. Ironically, Kaufert states, ‘‘It seems probable…that widespread burning of the bottoms did not occur until about 1890, for old settlers recall the time (before 1895) when the bottoms were veritable canebrakes,’’ and says that, according to the ‘‘oldtimers’’ in the area, ‘‘Cane is killed by a single burn.’’ This anecdotal information directly contradicts experimental studies of fire effects on Arundinaria performed later, all of which conclude that cane benefits from periodic fire (Hughes 1957, 1966, Wright and Bailey 1982, Gagnon and Platt 2008). If anything, the presence of canebrakes might more accurately be considered evidence that fires did occur. Kaufert attests to the pyrogenic nature of cane when he says that according to these ‘‘oldtimers,’’ fires in the past were often more severe than the more recent fires because ‘‘dry cane made excellent fuel.’’ He says that ‘‘since the passing of the cane the principal fire hazard has been leaf litter.’’ He goes on to decry great damage done to both merchantable timber and young trees by rot that enters via fire scars. He states flatly that, ‘‘To eliminate the serious losses caused by decay and to insure (sic) good stocking in young stands, fire must be kept out.’’ FIRE, WINDSTORM AND DROUGHT INTERACTIONS With today’s modified BLH as our reference, it can be difficult to conceive that fires were likely once a part of this floodplain disturbance regime. Land managers have suppressed fire for decades after its detrimental effects to timber became well publicized (Lentz 1931, Kaufert 1933, Toole and McKnight 1956). Suppressing fires is not difficult now because modern second-growth forests are virtually fireproof – their dense canopies reduce air movement and shade the understory, minimizing plant growth and fuel build-up (Wilson et al. 2007). Fires no longer travel across the landscape as when pyrogenic pine-filled uplands lay adjacent to the floodplains (Platt 1999). And canebrakes that were conduits to fire-spread in BLH have been eradicated in these fragmented forests (Kaufert 1933, Platt and Brantley 1997). Accounts of Lentz (1931) and Kaufert (1933) inform our thinking with several points about the history of fire in BLH. First, while many of the fires they describe occurred in large clear-cut gaps, tornados, hurricanes, and ice-storms similarly created large gaps in these forests over ecological time (Sharitz et al. 1992, Gagnon et al. 2007, Liu et al. 2008). Second, they describe fires that occurred several years after the gap-creating disturbances. Because of the multi-year lag between logging and drought, these fires would have spread through dense and continuous regenerating vegetation rather than through long-decomposed logging slash. This is consistent with the multi-year lag in hurricane/fire interactions described by Liu et al. (2008) from the Alabama Gulf Coast. Third, Lentz and Kaufert describe fires that occurred in late winter or spring after prolonged drought had dried sloughs that might have otherwise served as fire-breaks. Spring is typically the natural fire season throughout the coastal southeastern U.S., including where Kaufert and Lentz made their observations in Louisiana, because fuel conditions are favorable and lightning is prevalent (Olson and Platt 1995, Platt 1999). However, natural fires are also possible later in the growing season whenever lightning coincides with dry fuels (Foti and Glenn 1992). Although Lentz and Kaufert suggest that the fires were humancaused, the pyrogenic power of lightning was often underestimated at the time, and it is likely that the true ignition sources were often unknown. Lentz (1931) notes that early spring rainstorms were insufficient to moisten the dry leaf litter; these same storms likely came with lightning. Fourth, the interval between fires described by Kaufert (1933) ranges from five to 13 years, which fit closely the fire interval suggested by Hughes (1957, 1966) to keep Arundinaria stands healthy. There are some important qualifiers for this fire hypothesis. Fires would not have burned in BLH with the frequency or continuity of adjacent upland pine savannas (Platt 1999). Fires would have ranged from creeping low-intensity fires in hardwood leaf litter, to hot and violent conflagrations that consumed canebrakes and threatened individual trees. Certain areas, especially the ‘‘prairies’’ and canebrakes on higher ground, might have burned during most droughts, while adjacent areas would have been virtually fire-proof just like modern second-growth BLH. Native Americans probably increased the frequency and pervasiveness of fires in the bottomlands, but when conditions were right Gagnon, FIRE IN BOTTOMLAND HARDWOOD FORESTS and disturbances coincided, some areas almost certainly burned without human input. The true extent to which fires influenced BLH remains an open question, but surely the answer is ‘‘somewhat’’ rather than ‘‘not at all’’ as is now commonly practiced by modern BLH land-managers. A multi-pronged study could clarify the extent to which fires shaped BLH. The prevalence of A. gigantea remains in southeastern bottomlands could serve as an indicator of fire regime. Unfortunately, palynological information is sparse because Arundinaria pollen is virtually indistinguishable from that of many other grasses (K-b. Liu, Louisiana State University Department of Oceanography and Coastal Sciences, personal communication). It has recently been shown that phytoliths (precipitating microscopic plant silica bodies) from A. gigantea are unique and could well serve the same purpose (Lu and Liu 2003). An integrated study of macroscopic charcoal, microscopic charcoal, and Arundinaria phytoliths could clarify the extent to which fires shaped given bottomland sites ranging back several millennia (K-b. Liu, personal communication). Bottomland hardwood forests likely burned during the confluence of other interacting disturbances. This is similar to findings by Liu et al. (2008) of interactions between hurricanes and fires on the Alabama Gulf Coast. In BLH, the stage is set for fire when effects of windstorms or other large canopy disturbances coincide in subsequent years with prolonged droughts. Natural fires could have occurred in the following scenario: 1) A powerful windstorm or other disturbance opens a large canopy gap. 2) Over a few years, the gap fills with dense, regenerating vegetation. 3) Eventually, drought renders this new vegetation flammable and increases landscape connectivity for potential fire. 4) With the onset of spring thunderstorms, lightning strikes cause fires in these large, highly flammable canopy gaps. 5) Canebrakes and adjoining upland pine savannas facilitate fire spread into other parts of BLH. 6) Giant cane and other pyrogenic grasses thrive after burning, producing a flammable fuel bed that can burn again during the next drought. As canebrakes expand over time (Gagnon et al. 2007), they increase the area beyond the initial canopy disturbance that is susceptible to burning in subsequent droughts, and thereby set up a positive feedback (Gagnon and Platt 2008). In this way once a large canopy disturbance initiates the sequence, canebrakes and other pyrogenic assemblages within BLH could essentially be self-sustaining. Management plans for comprehensive restoration efforts should consider fire-maintained plant assemblages within BLH. These pyrogenic plant systems 525 likely burned during drought conditions and may not have burned otherwise. Such drought conditions might be outside management prescriptions for controlled burning; therefore, land managers should give consideration to how such areas might be burned within prescription. Areas containing pyrogenic species like A. gigantea could be cut and then burned two weeks later, before new culms resprout. Selected clearcuts or existing large blowdowns containing Arundinaria could be candidate sites for prescribed burning. ACKNOWLEDGMENTS I thank Kevin Robertson and Loretta Battaglia for ideas that inspired this paper. 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