Replacement patterns and species coexistence in an Andean Araucaria‐Nothofagus forest

Questions: Fire appears to affect both replacement patterns and coexistence of Araucaria araucana‐Nothofagus pumilio forests in the Andean Araucarian region of south‐central Chile. A quantitative assessment of coexistence in the absence of recent fires, however, is lacking. In this study, we considered the life‐history attributes, time of recruitment and spatial pattern of individuals of both tree species to address the following questions. How regular has recruitment of both species been in time? Is there any temporal niche differentiation? Are the two species positively or negatively associated in space and, if so, at what scale and for what age and size classes? Is there any spatial niche differentiation? Location: Andean Araucarian region of Chile, Villarrica National Park (39°35′S, 71°31′W; 1300 m a.s.l.). Methods: We stem‐mapped and cored a total of 1073 trees in a 1‐ha plot in a late‐successional post‐fire stand to examine spatiotemporal patterns of establishment. We used semivariogram modelling and the pair‐correlation function to distinguish between regeneration modes and describe species interactions. Results: The two species differ in their regeneration mode: whereas A. araucana appeared to recruit more continuously in time and space, episodic pulses of establishment were dominant for N. pumilio. At small scales, younger age‐class stems of A. araucana were randomly distributed, while older age‐class stems were aggregated. This was in contrast to common patterns for temperate tree species, including N. pumilio, following processes of self‐thinning. Younger age classes of A. araucana were distributed independently of older trees of both species, but younger age classes of N. pumilio had a negative association with older conspecifics at scales similar to crown diameter. Conclusions: In the absence of recent fires, it is likely that A. araucana would dominate the stand alone, given its greater shade tolerance, greater longevity and continuous recruitment. However, while canopy closure is still incomplete, the shade‐intolerant N. pumilio will be able to recruit in those open areas after seed masting and will coexist with A. araucana.     

Fire history of Araucaria–Nothofagus forests in Villarrica National Park, Chile

Aim In this study we examine fire history (i.e. c. 500 yr bp to present) of Araucaria–Nothofagus forests in the Andes cordillera of Chile. This is the first fire history developed from tree rings for an Araucaria–Nothofagus forest landscape. Location The fire history was determined for the Quillelhue watershed on the north side of Lanin volcano in Villarrica National Park, Chile. The long‐lived Araucaria araucana was commonly associated with Nothofagus pumilio and N. antarctica in more mesic and drier sites respectively. Methods Based on a combination of fire‐scar proxy records and forest stand ages, we reconstructed fire frequency, severity, and the spatial extent of burned areas for an c. 4000 ha study area. We used a composite fire chronology for the purpose of determining centennial‐scale changes in fire regimes and comparing the pre‐settlement (pre‐1883) and post‐settlement fire regimes. In addition, we contrasted Araucaria and Nothofagus species as fire‐scar recorders. Results In the study area, we dated a total of 144 fire‐scarred trees, representing 46 fire years from ad 1446 to the present. For the period from ad 1696 to 2000, using fire dates from Araucaria and Nothofagus species, the composite mean fire interval varied from 7 years for all fires to 62 years for widespread events (i.e. years in which ≥ 25% of recorder trees were scarred). Sensitivity to fire was different for Araucaria and Nothofagus species. More than 98% of the fires recorded by Nothofagus species occurred during the 1900s. The lack of evidence for older fire dates (pre‐1900) in Nothofagus species was due to their shorter longevity and greater susceptibility to being killed by more severe fires. Whereas the thin‐barked N. pumilio and N. antarctica are often destroyed in catastrophic fire events, large and thick‐barked Araucaria trees typically survive. The spatial extent of fires ranged from small patchy events to those that burned more than 40% of the entire landscape (c. > 1500 ha). Main conclusions Fire is the most important disturbance shaping the Araucaria–Nothofagus landscape in the Araucarian region. The forest landscape has been shaped by a mixed‐severity fire regime that includes surface and crown fires. High‐severity widespread events were relatively infrequent (e.g. 1827, 1909 and 1944) and primarily affected tall Araucaria–N. pumilio forests and woodlands dominated by Araucaria–N. antarctica. Although there is abundant evidence of the impact of Euro‐Chilean settlers on the area, the relative influence of this settlement on the temporal pattern of fire could only be tentatively established due to the relatively small number of pre‐1900 fire dates. An apparent increase in fire occurrence is evident in the fire record during Euro‐Chilean settlement (post‐1880s) compared with the Native American era, but it may also be the result of the destruction of evidence of older fires by more recent stand‐devastating fires (e.g. 1909 and 1944). Overall, the severe and widespread fires that burned in Araucaria–Nothofagus forests of this region in 2002, previously interpreted as an ecological novelty, are within the range of the historic fire regimes that have shaped this forested landscape.     

Natural and anthropogenic influences on coarse woody debris stocks in Nothofagus–Araucaria forests of northern Patagonia,Argentina

Coarse woody debris (CWD) is an important element driving ecological processes, strengthening ecosystem resilience and for biodiversity within forest ecosystems. However, the abundance and distribution of CWD and their relation to natural and human factors are poorly known in southern South America. In this work we studied the density and volume of CWD types in Nothofagus–Araucaria stands in northern Patagonia (Neuquén – Argentina) and relationships with forest composition and structure. We also studied their relationships with fire history, topography and human‐related variables. Twenty‐three stands with Nothofagus pumilio, Nothofagus antarctica and/or Araucaria araucana were sampled to estimate quantities of logs, snags and dead branches using the planar‐intersect method. CWD density and volume in these forests were moderate and varied across the landscape with a spatial pattern determined by biotic, abiotic and human use–related variables. Mean CWD volume was 52.9 m³ ha^?1 (range: 1.6–143.7) and significantly varied among forest types and watersheds. CWD was positively related to dbh, tree height and slope, but negatively related to tree density. CWD was clearly influenced by composition and structural characteristics of stands, where the tree species traits had an important role. As well, the observed amount and type of CWD, whereby most of the stands showed low levels of old (pre‐disturbance) logs/snags and poor new inputs of deadwood, may be explained by fire frequency. Firewood gathering and livestock grazing negatively affected deadwood stocks and topography counteracts this effect by limiting human access. Fire disturbance history, windthrow and dieback pulses produced by insect outbreaks and human access seemed to be the main causes that best explained CWD spatial distribution and abundance patterns in north‐western Patagonian forests.     

Changes in the stand structure (1975–2000) of coastal Banksia forest in the long absence of fire

Abstract Revisitation studies enable long‐term changes in vegetation to be deciphered and insights into plant community succession to be gained. This is particularly important when assessing the effects of fire exclusion in ecosystems where fire is thought to have once been common. Using two adjacent coastal Banksia integrifolia forest stands in southern Victoria, Australia initially surveyed in 1975 by Hazard and Parsons, we document the changes that occurred in the stand structure between 1975 and 2000. Western Park (WP) has now remained unburnt for over 100 years while Cerberus Naval Base (CNB) was most recently burnt in 1942. Banksia integrifolia densities have decreased at both sites over the 25‐years period by an average of 42–77%, as have other coastal native shrubs (e.g. Leptospermum laevigatum, Leucopogon parviflorus). Trees at WP appear to have died due to old age while mortality at CNB is presumed to be due to stand thinning in response to intense competition for light. Successful recruitment by Banksia has been minimal; trees less than 9 cm girth over bark at breast height (GBBH) were absent at CNB while no trees <19 cm GBBH were observed at WP. The long‐term absence of disturbance such as fire is suspected to be one of the causes of regeneration failure of the stand at WP. Gap phase regeneration is not apparent in B. integrifolia and hence, long‐term succession to a more grassy community is likely when fire is excluded for long periods.     

High‐severity fire corroborated in historical dry forests of the western United States: response to Fulé et al.

Accurate assessment of changing fire regimes is important, since climatic change and people may be promoting more wildfires. Government wildland fire policies and restoration programmes in dry western US forests are based on the hypothesis that high‐severity fire was rare in historical fire regimes, modern fire severity is unnaturally high and restoration efforts should focus primarily on thinning forests to eliminate high‐severity fire. Using General Land Office (GLO) survey data over large dry‐forest landscapes, we showed that the proportion of historical forest affected by high‐severity fire was not insignificant, fire severity has not increased as a proportion of total fire area and large areas of dense forest were present historically (Williams & Baker, Global Ecology and Biogeography, 21 , 1042–1052, 2012; W&B). In response, Fulé et al. (Global Ecology and Biogeography, 2013, doi: 10.1111/geb.12136; FE) suggest that our inferences are unsupported and land management based on our research could be damaging to native ecosystems. Here, we show that the concerns of FE are unfounded. Their criticism comes from misquoting W&B, mistaking W&B's methods, misusing evidence (e.g. from Aldo Leopold) and missing substantial available evidence. We also update corroboration for the extensive historical high‐severity fire shown by W&B. We suggest that restoration programmes are misdirected in seeking to reduce all high‐severity fire in dry forests, given findings from spatially extensive GLO data and other sources.     

Influence of fire severity on the regeneration,recruitment and distribution of eucalypts in the Cotter River Catchment,Australian Capital Territory

Abstract Plant responses to fire are variable between and within species and are influenced by numerous factors including fire severity. This study investigated the effects of fire severity on the regeneration and recruitment of forest eucalypts in the Cotter River Catchment, Australian Capital Territory (ACT). This study also examined the potential for the obligate seeder Eucalyptus delegatensis R.T. Baker (Myrtaceae) to expand into adjacent stands dominated by the facultative resprouter Eucalyptus fastigata H. Deane & Maiden (Myrtaceae) by seed shed and seedling establishment beyond the pre‐fire boundary. Sites were located in areas of either higher or lower fire severity, and transects were placed across the boundary of stands of E. delegatensis and E. fastigata. Species distributions, tree survival and seedling densities and heights were recorded, and the location of each boundary was determined as the region of maximum change in species composition along the transects. Eucalyptus delegatensis was the only eucalypt killed by higher severity fire. However, E. delegatensis seedling density was greater at higher severity sites than lower severity sites. Eucalyptus fastigata seedling density was low across all sites, with other eucalypts producing few, if any, seedlings. There was no evidence that E. delegatensis had increased its range into downslope stands dominated by E. fastigata. Patterns of vegetative recovery and seedling recruitment may be related to a number of factors, including differences in allocation patterns between seeders and sprouters, and the effects of overstory and understory competition. It is unclear what processes impede E. delegatensis seedling establishment beyond the stand boundary, but may involve an inability of E. delegatensis to shed seed sufficiently far downslope; unsuitable conditions for germination beyond the boundary; or, competition from a retained or resprouting overstory, despite the potential for increased dispersal distance soon after fire.     

Topographic controls on the distribution of tree islands in the high Andes of south‐western Ecuador

Aim To evaluate the hypothesis that geomorphometric parameters of upper montane Andean environments have an important influence on the regional fire ecology and consequently play a role in the spatial distribution of ‘remnant’ tree islands dominated by Polylepis. Location A glacial landscape located between 3600 and 4400 m elevation in Cajas National Park, south‐western Ecuador. Methods The eigenvalue ratio method was used to evaluate the regional geomorphometric parameters of a 30‐m digital elevation model for Cajas National Park. The landscape character was evaluated by quantifying the topographic roughness, organization, and gradient. This information was used to determine the spatial correlations between terrain characteristics and the distribution of tree islands in the region. Results We demonstrate a strong spatial correlation between areas of high topographic roughness and gradient, and the locations of the major tree islands. We find that there is a distinctive relationship between the topographic roughness and organization in the vicinity of the tree islands (e.g. increased upslope roughness and decreased topographic grain strength) that substantiates the notion that the tree islands are located in relatively inaccessible topography. Main conclusions In the northern and central Andes, the location of Polylepis‐dominated ‘forest islands’ has been shown to be a function of climate, terrain characteristics, and anthropogenic disturbances. Although the relative importance of various ecological factors has been debated, it remains clear that fires have exerted a strong influence on these ecosystems. Other authors have noted that tree islands are more likely to occur at the base of cliffs, above moist areas, and in other areas where fires do not burn frequently. Our results corroborate these observations, and demonstrate that the occurrence of Polylepis patches is strongly correlated with specific combinations of terrain features. Although we do not discount the importance of other factors in determining the spatial position and areal extent of these forests, we demonstrate strong support for fire‐related hypotheses.     

Post‐fire habitat use of the golden‐backed tree‐rat (Mesembriomys macrurus) in the northwest Kimberley,Western Australia

Fire regimes are changing throughout the world. Changed fire patterns across northern Australian savannas have been proposed as a factor contributing to recent declines of small‐ and medium‐sized mammals. Despite this, few studies have examined the mechanisms that underpin how species use habitat in fire‐affected landscapes. We determined the habitats and resources important to the declining golden‐backed tree‐rat (Mesembriomys macrurus) in landscapes partially burnt by recent intense fire. We aimed to (i) compare the relative use of rainforest and savanna habitats; (ii) examine the effect of fire history on use of savanna habitats; and (iii) identify key foraging and denning resources. Habitat selection was examined by comparing the availability of eight habitat types around real (used) and generated (available) location points. Individuals used a range of habitats, but consistently selected long unburnt rainforest in preference to recently burnt savanna (1–12 months post‐fire); however, recently burnt savanna was used in preference to long unburnt savanna. Tree‐rats foraged in Terminalia hadleyana, Planchonia rupestris, Celtis philippensis and Owenia vernicosa, tree species that are found in a variety of habitat types. Individuals used a range of den sites, including cliffs, trees, logs, scree and stags found throughout the study area. Although multiple factors may have led to the decline of Mes. macrurus across its range, these results are consistent with the idea that changes in the savanna structure as a consequence of contemporary fire patterns could also have a role. The continued persistence of Mes. macrurus in the northwest Kimberley may be supported by land management strategies that conserve fruiting and hollow‐bearing trees, and maintain the availability of fire‐sensitive vegetation types.     

Unsupported inferences of high‐severity fire in historical dry forests of the western United States: response to Williams and Baker

Reconstructions of dry western US forests in the late 19th century in Arizona, Colorado and Oregon based on General Land Office records were used by Williams & Baker (2012; Global Ecology and Biogeography, 21 , 1042–1052; hereafter W&B) to infer past fire regimes with substantial moderate and high‐severity burning. The authors concluded that present‐day large, high‐severity fires are not distinguishable from historical patterns. We present evidence of important errors in their study. First, the use of tree size distributions to reconstruct past fire severity and extent is not supported by empirical age–size relationships nor by studies that directly quantified disturbance history in these forests. Second, the fire severity classification of W&B is qualitatively different from most modern classification schemes, and is based on different types of data, leading to an inappropriate comparison. Third, we note that while W&B asserted ‘surprising’ heterogeneity in their reconstructions of stand density and species composition, their data are not substantially different from many previous studies which reached very different conclusions about subsequent forest and fire behaviour changes. Contrary to the conclusions of W&B, the preponderance of scientific evidence indicates that conservation of dry forest ecosystems in the western United States and their ecological, social and economic value is not consistent with a present‐day disturbance regime of large, high‐severity fires, especially under changing climate.     

Low‐severity fire as a mechanism of organic matter protection in global peatlands: Thermal alteration slows decomposition

Worldwide, regularly recurring wildfires shape many peatland ecosystems to the extent that fire‐adapted species often dominate plant communities, suggesting that wildfire is an integral part of peatland ecology rather than an anomaly. The most destructive blazes are smoldering fires that are usually initiated in periods of drought and can combust entire peatland carbon stores. However, peatland wildfires more typically occur as low‐severity surface burns that arise in the dormant season when vegetation is desiccated, and soil moisture is high. In such low‐severity fires, surface layers experience flash heating, but there is little loss of underlying peat to combustion. This study examines the potential importance of such processes in several peatlands that span a gradient from hemiboreal to tropical ecozones and experience a wide range of fire return intervals. We show that low‐severity fires can increase the pool of stable soil carbon by thermally altering the chemistry of soil organic matter (SOM), thereby reducing rates of microbial respiration. Using X‐ray photoelectron spectroscopy and Fourier transform infrared, we demonstrate that low‐severity fires significantly increase the degree of carbon condensation and aromatization of SOM functional groups, particularly on the surface of peat aggregates. Laboratory incubations show lower CO₂ emissions from peat subjected to low‐severity fire and predict lower cumulative CO₂ emissions from burned peat after 1–3 years. Also, low‐severity fires reduce the temperature sensitivity (Q₁₀) of peat, indicating that these fires can inhibit microbial access to SOM. The increased stability of thermally altered SOM may allow a greater proportion of organic matter to survive vertical migration into saturated and anaerobic zones of peatlands where environmental conditions physiochemically protect carbon stores from decomposition for thousands of years. Thus, across latitudes, low‐severity fire is an overlooked factor influencing carbon cycling in peatlands, which is relevant to global carbon budgets as climate change alters fire regimes worldwide.     

Carbon dynamics in the future forest: the importance of long‐term successional legacy and climate–fire interactions

Understanding how climate change may influence forest carbon (C) budgets requires knowledge of forest growth relationships with regional climate, long‐term forest succession, and past and future disturbances, such as wildfires and timber harvesting events. We used a landscape‐scale model of forest succession, wildfire, and C dynamics (LANDIS‐II) to evaluate the effects of a changing climate (A2 and B1 IPCC emissions; Geophysical Fluid Dynamics Laboratory General Circulation Models) on total forest C, tree species composition, and wildfire dynamics in the Lake Tahoe Basin, California, and Nevada. The independent effects of temperature and precipitation were assessed within and among climate models. Results highlight the importance of modeling forest succession and stand development processes at the landscape scale for understanding the C cycle. Due primarily to landscape legacy effects of historic logging of the Comstock Era in the late 1880s, C sequestration may continue throughout the current century, and the forest will remain a C sink (Net Ecosystem Carbon Balance > 0), regardless of climate regime. Climate change caused increases in temperatures limited simulated C sequestration potential because of augmented fire activity and reduced establishment ability of subalpine and upper montane trees. Higher temperatures influenced forest response more than reduced precipitation. As the forest reached its potential steady state, the forest could become C neutral or a C source, and climate change could accelerate this transition. The future of forest ecosystem C cycling in many forested systems worldwide may depend more on major disturbances and landscape legacies related to land use than on projected climate change alone.     

Simplifying the savanna: the trajectory of fire‐sensitive vegetation mosaics in northern Australia

Aim Fire is a key agent in savanna systems, yet the capacity to predict fine‐grained population phenomena under variable fire regime conditions at landscape scales is a daunting challenge. Given mounting evidence for significant impacts of fire on vulnerable biodiversity elements in north Australian savannas over recent decades, we assess: (1) the trajectory of fire‐sensitive vegetation elements within a particularly biodiverse savanna mosaic based on long‐term monitoring and spatial modelling; (2) the broader implications for northern Australia; and (3) the applicability of the methodological approach to other fire‐prone settings. Location Arnhem Plateau, northern Australia. Methods We apply data from long‐term vegetation monitoring plots included within Kakadu National Park to derive statistical models describing the responses of structure and floristic attributes to 15 years of ambient (non‐experimental) fire regime treatments. For a broader 28,000 km² region, we apply significant models to spatial assessment of the effects of modern fire regimes (1995–2009) on diagnostic closed forest, savanna and shrubland heath attributes. Results Significant models included the effects of severe fires on large stems of the closed forest dominant Allosyncarpia ternata, stem densities of the widespread savanna coniferous obligate seeder Callitris intratropica, and fire frequency and related fire interval parameters on numbers of obligate seeder taxa characteristic of shrubland heaths. No significant relationships were observed between fire regime and eucalypt and non‐eucalypt adult tree components of savanna. Spatial application of significant models illustrates that more than half of the regional closed forest perimeters, savanna and shrubland habitats experienced deleterious fire regimes over the study period, except in very dissected terrain. Main conclusions While north Australia’s relatively unmodified mesic savannas may appear structurally intact and healthy, this study provides compelling evidence that fire‐sensitive vegetation elements embedded within the savanna mosaic are in decline under present‐day fire regimes. These observations have broader implications for analogous savanna mosaics across northern Australia, and support complementary findings of the contributory role of fire regimes in the demise of small mammal fauna. The methodological approach has application in other fire‐prone settings, but is reliant on significant long‐term infrastructure resourcing.     

Prioritizing forest fuels treatments based on the probability of high‐severity fire restores adaptive capacity in Sierran forests

In frequent fire forests of the western United States, a legacy of fire suppression coupled with increases in fire weather severity have altered fire regimes and vegetation dynamics. When coupled with projected climate change, these conditions have the potential to lead to vegetation type change and altered carbon (C) dynamics. In the Sierra Nevada, fuels reduction approaches that include mechanical thinning followed by regular prescribed fire are one approach to restore the ability of the ecosystem to tolerate episodic fire and still sequester C. Yet, the spatial extent of the area requiring treatment makes widespread treatment implementation unlikely. We sought to determine if a priori knowledge of where uncharacteristic wildfire is most probable could be used to optimize the placement of fuels treatments in a Sierra Nevada watershed. We developed two treatment placement strategies: the naive strategy, based on treating all operationally available area and the optimized strategy, which only treated areas where crown‐killing fires were most probable. We ran forecast simulations using projected climate data through 2,100 to determine how the treatments differed in terms of C sequestration, fire severity, and C emissions relative to a no‐management scenario. We found that in both the short (20 years) and long (100 years) term, both management scenarios increased C stability, reduced burn severity, and consequently emitted less C as a result of wildfires than no‐management. Across all metrics, both scenarios performed the same, but the optimized treatment required significantly less C removal (naive=0.42 Tg C, optimized=0.25 Tg C) to achieve the same treatment efficacy. Given the extent of western forests in need of fire restoration, efficiently allocating treatments is a critical task if we are going to restore adaptive capacity in frequent‐fire forests.     

Fire effects on the soil seed bank and post‐fire resilience of a semi‐arid shrubland in central Argentina

Soil seed bank is an important source of resilience of plant communities who suffered disturbances. We analysed the effect of an intense fire in the soil seed bank of a semi‐arid shrubland of Córdoba Argentina. We asked if the fire affected seed abundance, floristic and functional composition of the soil seed bank at two different layers (0–5 cm and 5–10 cm), and if fire could compromise the role of the soil seed bank as a source of resilience for the vegetation. We collected soil samples from a burned site and from a control site that had not burned. Samples were installed in a greenhouse under controlled conditions. During 12 months, we recorded all germinated seedlings. We compare soil seed bank with pre‐fire vegetation in terms of floristic and functional composition. The high‐intensity fire deeply affected the abundance of seeds in the soil, but it did not affect its floristic or functional composition. Floristic and functional composition of soil seed banks – at burned and unburned sites‐ differed markedly from that of the pre‐fire vegetation, although a previous study at the same site indicated high resilience after fire of this plant community. Our results indicate that resilience of this system is not strongly dependent on direct germination from seeds buried in the soil. Other sources of resilience, like colonization from neighbouring vegetation patches and resprouting from underground organs appear to gain relevance after an intense fire.     

Influence of steam pretreatment severity on post‐treatments used to enhance the enzymatic hydrolysis of pretreated softwoods at low enzyme loadings

It is recognized that some form of post‐treatment will usually be required if reasonable hydrolysis yields (>60%) of steam pretreated softwood are to be achieved when using low enzyme loadings (5 FPU/g cellulose). In the work reported here we modified/removed lignin from steam pretreated softwood while investigating the influence that the severity of pretreatment might have on the effectiveness of subsequent post‐treatments. Although treatment at a lower severity could provide better overall hemicellulose recovery, post‐treatment was not as effective on the cellulosic component. Pretreatment at medium severity resulted in the best compromise, providing reasonable recovery of the water soluble hemicellulose sugars and the use of post‐treatment conditions that significantly increased the enzymatic hydrolysis of the water insoluble cellulosic component. Post‐treatment with alkaline hydrogen peroxide or neutral sulfonation resulted in 62% cellulose hydrolysis at an enzyme loading of 5 FPU/g cellulose, which was four times greater than was obtained when the cellulosic fraction was not post‐treated. When the enzyme loading was increased to 15 FPU/g cellulose, the post‐treated cellulosic fraction was almost completely hydrolyzed to glucose. Despite the higher lignin content (44%) of the sulfonated substrate, similar hydrolysis yields to those achieved after alkaline peroxide post‐treatment (14% lignin content) indicated that, in addition to lignin removal, lignin modification also plays an important role in influencing the effectiveness of hydrolysis when low enzyme loadings are used. Biotechnol. Bioeng. 2011;108: 2300–2311. © 2011 Wiley Periodicals, Inc.