J. Kevin Hiers

Modeling silviculture after natural disturbance to sustain biodiversity in the longleaf pine (Pinus palustris) ecosystem: balancing complexity and implementation

Modeling silviculture after natural disturbance to maintain biodiversity is a popular concept, yet its application remains elusive. We discuss difficulties inherent to this idea, and suggest approaches to facilitate implementation, using longleaf pine (Pinus palustris) as an example. Natural disturbance regimes are spatially and temporally variable. Variability leads to a range of structural outcomes, or results in different pathways leading to similar structures. In longleaf pine, lightning, hurricanes, surface fires, and windthrow all lead to similar structures, but at different rates. Consequently, a manager can select among various natural disturbance patterns when searching for an appropriate silvicultural model. This facilitates management by providing flexibility to meet a range of objectives. The outcomes of natural disturbances are inherently different from those of silviculture, for example, harvesting always removes boles. It is instructive to think of silvicultural disturbances along a gradient in structural outcomes, reflecting degree of disparity with natural disturbance. In longleaf pine this might involve managing for two-cohort structure, instead of multi-cohort structure characteristic of old growth stands. While two-cohort structure is a simplification over the old growth condition, it is an improvement over single-cohort management. Reducing structural disparity between managed and unmanaged forests is key to sustaining biodiversity because of linkages that exist between structural elements, forest biota, and ecosystem processes. Finally, interactions of frequency, severity, intensity, seasonality, and spatial pattern define a disturbance regime. These components may not have equal weight in affecting biodiversity. Some are easier to emulate with silviculture than are others. For instance, ecologists consider growing-season fire more reflective of the natural fire regime in longleaf pine and critical for maintenance of biodiversity. However, dormant season fire is easier to use and recent work with native plants suggests that seasonality of fire may be less critical to maintenance of species richness, as one component of biodiversity, than is generally believed. Science can advance the goal of modeling silviculture after natural disturbances by better illustrating cause and effect relationships among components of disturbance regimes and the structure and function of ecosystems. Wide application requires approaches that are adaptable to different operational situations and landowner objectives. A key point for managers to remember is that strict adherence to a silvicultural regime that closely parallels a natural disturbance regime may not always be necessary to maintain biodiversity. We outline examples of silvicultural systems for longleaf pine that demonstrates these ideas.

FOREST FLOOR DEPTH MEDIATES UNDERSTORY VIGOR IN XERIC PINUS PALUSTRIS ECOSYSTEMS

Longleaf pine (Pinus palustris) woodlands and savannas are among the most frequently burned ecosystems in the world with fire return intervals of 1-10 years. This fire regime has maintained high levels of biodiversity in terms of both species richness and endemism. Land use changes have reduced the area of this ecosystem by >95%, and inadequate fire frequencies threaten many of the remnants today. In the absence of frequent fire, rapid colonization of hardwoods and shrubs occurs, and a broad-leaved midstory develops. This midstory encroachment has been the focus of much research and management concern, largely based on the assumption that the midstory reduces understory plant diversity through direction competition via light interception. The general application of this mechanism of degradation is questionable, however, because midstory density, leaf area, and hardwood species composition vary substantially along a soil moisture gradient from mesic to extremely xeric sites. Reanalysis of recently reported data from xeric longleaf pine communities suggests that the development of the forest floor, a less conspicuous change in forest structure, might cause a decline in plant biodiversity when forests remain unburned. We report here a test of the interactions among fire, litter accumulation, forest floor development, and midstory canopy density on understory plant diversity. Structural equation modeling showed that within xeric sites, forest floor development was the primary factor explaining decreased biodiversity. The only effects of midstory development on biodiversity were those mediated through forest floor development. Boundary line analysis of functional guilds of understory plants showed sensitivity to even minor development of the forest floor in the absence of fire. These results challenge the prevailing management paradigm and suggest that within xeric longleaf pine communities, the primary focus of managed fire regime should be directed toward the restoration of forest floor characteristics rather than the introduction of high-intensity fires used to regulate midstory structure.

The wildland fuel cell concept: an approach to characterize fine-scale variation in fuels and fire in frequently burned longleaf pine forests

In ecosystems with frequent surface fire regimes, fire and fuel heterogeneity has been largely overlooked owing to the lack of unburned patches and the difficulty in measuring fire behavior at fine scales (0.1-10 m). The diverse vegetation in these ecosystems varies at these fine scales. This diversity could be driven by the influences of local interactions among patches of understorey vegetation and canopy-supplied fine fuels on fire behavior, yet no method we know of can capture fine-scale fuel and fire measurements such that these relationships could be rigorously tested. We present here an original method for inventorying of fine-scale fuels and in situ measures of fire intensity within longleaf pine forests of the south-eastern USA. Using ground-based LIDAR (Light Detection and Ranging) with traditional fuel inventory approaches, we characterized within-fuel bed variation into discrete patches, termed wildland fuel cells, which had distinct fuel composition, characteristics, and architecture that became spatially independent beyond 0.5 m 2 . Spatially explicit fire behavior was measured in situ through digital infrared thermography. We found that fire temperatures and residence times varied at similar scales to those observed for wildland fuel cells. The wildland fuels cell concept could seamlessly connect empirical studies with numerical models or cellular automata models of fire behavior, representing a promising means to better predict within-burn heterogeneity and fire effects.

The effects of fire regime on legume reproduction in longleaf pine savannas: is a season selective?

Abstract. The biodiversity of fire-dependent ecosystems is increasingly threatened by habitat fragmentation and fire suppression. Reducing species loss requires that salient features of natural fire regimes be incorporated into managed regimes. Lightning-season burns have been emphasized as the critical component of disturbance regimes that maintain native biodiversity within endangered longleaf pine savannas, the most diverse plant community in North America. Over evolutionary time, lightning-season fire is thought to have exerted strong selection pressure on plant pollinator interaction, flower production, and seedling establishment. In this study, season of burn had no effect on pollination activity of native legumes. Contrary to the conventional paradigm, we found a wide range of reproductive responses among dominant legumes in response to the season of burn treatments, suggesting that a variable fire season, rather than a single season of burn, is appropriate to maintain a greater variety of native species. We propose that varying the components of fire regimes, rather than selecting a particular fire regime, is likely to be important to conserve biodiversity in this and other fire-dependent communities.

Overstory tree mortality resulting from reintroducing fire to long-unburned longleaf pine forests: the importance of duff moisture

In forests historically maintained by frequent fire, reintroducing fire after decades of exclusion often causes widespread overstory mortality. To better understand this phenomenon, we subjected 16 fire-excluded (ca. 40 years since fire) 10 ha longleaf pine (Pinus palustris Mill.) stands to one of four replicated burning treatments based on volumetric duff moisture content (VDMC): wet (115% VDMC); moist (85% VDMC); dry (55% VDMC); and a no-burn control. Dur- ing the first 2 years postfire, overstory pines in the dry burns suffered the greatest mortality (mean 20.5%); pine mortality in the wet and moist treatments did not differ from the control treatment. Duff reduction was greatest in the dry burns (mean 46.5%), with minimal reduction in the moist and wet burns (14.5% and 5%, respectively). Nested logistic regression using trees from all treatments revealed that the best predictors of individual pine mortality were duff consumption and crown scorch (P < 0.001; R 2 = 0.34). Crown scorch was significant only in dry burns, whereas duff consumption was sig- nificant across all treatments. Duff consumption was related to moisture content in lower duff (Oa; R 2 = 0.78, P < 0.001). Restoring fire to long-unburned forests will require development of burn prescriptions that include the effects of duff con- sumption, an often overlooked fire effect.

Measurements of convective and radiative heating in wildland fires

Time-resolved irradiance and convective heating and cooling of fast-response thermopile sensors were measured in 13 natural and prescribed wildland fires under a variety of fuel and ambient conditions. It was shown that a sensor exposed to the fire environment was subject to rapid fluctuations of convective transfer whereas irradiance measured by a windowed sensor was much less variable intime, increasing nearly monotonically with the approach of the flamefrontandlargelydecliningwithitspassage.Irradiancebeneathtwocrownfirespeakedat200and300kWm � 2 ,peak irradiance associated with fires in surface fuels reached 100kWm � 2 and the peak for three instances of burning in shrub fuels was 132kWm � 2 . The fire radiative energy accounted for 79% of the variance in fuel consumption. Convective heatingatthesensorsurfacevariedfrom15%tovaluesexceedingtheradiativeflux.Detailedmeasurementsofconvective and radiative heating rates in wildland fires are presented. Results indicate that the relative contribution of each to total energy release is dependent on fuel and environment.

Interactions among overstory structure, seedling life-history traits, and fire in frequently burned neotropical pine forests.

Abstract Fire-dependent pine forests in the Caribbean Basin cover extensive areas in the coastal plain of the Caribbean Sea and Gulf of Mexico and on several islands in the Bahamas Archipelago, Cuba, Hispaniola, and the Honduran Bay islands. These forests are high in conservation value but, unfortunately, remain mostly unprotected. Moreover, even though they are fire dependent, the use of fire for forest management often suffers from poor public perception and is prohibited by law in several countries. In this paper, we describe the fundamental links among fire, forest regeneration, and forest persistence in these ecosystems. We identify two general strategies based on the presence or absence of pine seedling adaptations for fire survival and describe management implications of these two strategies. We also introduce conceptual models describing fire, forest structure, and regeneration strategy linkages.

Bird assemblage response to restoration of fire‐suppressed longleaf pine sandhills

The ecological restoration of fire-suppressed habitats may require a multifaceted approach. Removal of hardwood trees together with reintroduction of fire has been suggested as a method of restoring fire-suppressed longleaf pine (Pinus palustris) forests; however, this strategy, although widespread, has not been evaluated on large spatial and temporal scales. We used a landscape-scale experimental design to examine how bird assemblages in fire-suppressed longleaf pine sandhills responded to fire alone or fire following mechanical removal or herbicide application to reduce hardwood levels. Individual treatments were compared to fire-suppressed controls and reference sites. After initial treatment, all sites were managed with prescribed fire, on an approximately two- to three-year interval, for over a decade. Nonmetric multidimensional scaling ordinations suggested that avian assemblages on sites that experienced any form of hardwood removal differed from assemblages on both fire-suppressed sites and reference sites 3-4 years after treatment (i.e., early posttreatment). After >10 years of prescribed burning on all sites (i.e., late posttreatment), only assemblages at sites treated with herbicide were indistinguishable from assemblages at reference sites. By the end of the study, individual species that were once indicators of reference sites no longer contributed to making reference sites unique. Occupancy modeling of these indicator species also demonstrated increasing similarity across treatments over time. Overall, although we documented long-term and variable assemblage-level change, our results indicate occupancy for birds considered longleaf pine specialists was similar at treatment and reference sites after over a decade of prescribed burning, regardless of initial method of hardwood removal. In other words, based on the response of species highly associated with the habitat, we found no justification for the added cost and effort of fire surrogates; fire alone was sufficient to restore these species.

The precision problem in conservation and restoration

Within the varied contexts of environmental policy, conservation of imperilled species populations, and restoration of damaged habitats, an emphasis on idealized optimal conditions has led to increasingly specific targets for management. Overly-precise conservation targets can reduce habitat variability at multiple scales, with unintended consequences for future ecological resilience. We describe this dilemma in the context of endangered species management, stream restoration, and climate-change adaptation. Inappropriate application of conservation targets can be expensive, with marginal conservation benefit. Reduced habitat variability can limit options for managers trying to balance competing objectives with limited resources. Conservation policies should embrace habitat variability, expand decision-space appropriately, and support adaptation to local circumstances to increase ecological resilience in a rapidly changing world.

A dynamic reference model: a framework for assessing biodiversity restoration goals in a fire-dependent ecosystem

The use of reference models as templates of historical or natural conditions to assess restoration progress is inherently logical; however, difficulties occur in application because of the need to incorporate temporal variation in ecosystems caused by disturbance and succession, as well as seasonal, interannual, or decadal variability. The landscape-scale restoration of the globally threatened and fire-dependent longleaf pine ecosystem in the southeastern United States is an example in which restoration efforts are even more complicated by the limited availability of extant reference sites. This study uses the dynamic reference conceptual framework to assess the direction and rate of recovery with respect to biodiversity restoration goals using a 15-year vegetation data set from an experimental restoration treatment in fire-excluded, hardwood-encroached longleaf pine sandhills. We compared ground-cover vegetation response to midstory hardwood removal through herbicide application, mechanical removal, and fire only. Nonmetric multidimensional scaling ordinations and proportional similarity analyses suggest that, while vegetation changed in all treatments over time, no differences in species composition or hardwood density in the ground cover were attributable to hardwood reduction treatments after 15 years with frequent prescribed fire. Furthermore, the results of this study indicate that considerable variability is associated with reference sites over time. Sites identified in 1994 as attainable restoration targets had become a moving target themselves, changing in magnitude consistent with alterations in restoration plots attributable to treatment effects and shaped by the modest increase in fire frequency imposed since 1998. In a broad restoration context, this study demonstrates a conceptual framework to better understand and integrate the range of spatial and temporal variation associated with the best available reference sites. It also illustrates a practical tool for statistically defining reference sites and for measuring restoration success in continually changing conditions that should be widely applicable to other ecosystems and restoration goals.