Three centuries of fire in montane pine‐oak stands on a temperate forest landscape

Question: What was the role of fire in montane pine‐oak (Pinus‐Quercus) stands under changing human land uses on a temperate forest landscape in eastern North America? Location: Mill Mountain in the central Appalachian Mountains, Virginia, US. Methods: A dendroecological reconstruction of fire history was generated for four stands dominated by xerophytic pine and oak species. The fire chronology began under presettlement conditions following aboriginal depopulation. Subsequent land uses included European settlement, iron mining, logging, and US Forest Service acquisition and fire protection. Results: Fires occurred approximately every 5 years until 1930 without any evidence of a temporal trend in fire frequency. Burning ceased after 1930. Area‐wide fires affecting multiple pine stands were common, occurring at intervals of approximately 16 years. Most living pines became established during the late 1800s and early 1900s. Dead pines indicated that an older cohort established ca. 1730. Most hardwoods were established between the 1920s and 1940s. Conclusions: Except for fire protection, changes in land use had no discernible influence on fire frequency. Lightning ignitions and/or large fire extent may have been important for maintaining frequent burning in the 1700s, while fuel recovery may have constrained fire frequency during later periods. The disturbance regime appears to be characterized by frequent surface fires and occasional severe fires, insect outbreaks or other disturbances followed by pine recruitment episodes. Industrial disturbances appear to have had little influence on the pine stands. The greatest impact of industrial society is fire exclusion, which permitted hardwood establishment.     

Uncertainty in the detection of disturbance spatial patterns in temperate forests

The use of individual-based models in the study of the spatial patterns of disturbances has opened new horizons in forest ecosystem research. However, no studies so far have addressed (i) the uncertainty in geostatistical modelling of the spatial relationships in dendrochronological data, (ii) the number of increment cores necessary to study disturbance spatial patterns, and (iii) the choice of an appropriate geostatistical model in relation to disturbance regime. In addressing these issues, we hope to contribute to advances in research methodology as well as to improve interpretations and generalizations from case studies.We used data from the beech-dominated Žofínský Prales forest reserve (Czech Republic), where we cored 3020 trees on 74 ha. Block bootstrap and geostatistics were applied to the data, which covered five decades with highly different disturbance histories. This allowed us to assess the general behavior of various mathematical models. Uncertainty in the spatial patterns and stability of the models was measured as the length of the 95% confidence interval (CI) of model parameters.According to Akaike Information Criterion (AIC), the spherical model fitted best at the range of ca. 20 m, while the exponential model was best at the range of ca. 60 m. However, the best fitting models were not always the most stable. The stability of models grew significantly with sample size. At <500 cores the spherical model was the most stable, while the Gaussian model was very unstable at <300 cores. The pure nugget model produced the most precise nugget estimate. The choice of model should thus be based on the expected spatial relations of the forest ecosystem under study. Sill was the most stable parameter, with an error of ±6–20% for ≥1110 core series. By contrast, practical range was the most sensitive, with an error of at least ±59%. The estimation of the spatial pattern of severe disturbances was more precise than that of fine-scale disturbances.The results suggest that with a sample size of 1000–1400 cores and a properly chosen model, one reaches a certain precision in estimation that does not increase significantly with growing sample size. It appears that in temperate old-growth forests controlled by fine-scale disturbances, it is necessary to have at least 500 cores to estimate sill, nugget and relative nugget, while to estimate practical range at least 1000 cores are needed. When choosing the best model, the stability of the model should be considered together with the value of AIC. Our results indicate the general limits of disturbance spatial pattern studies using dendrochronological and geostatistical methods, which can be only partially overcome by sample size or sampling design.     

Dynamic tree aggregation patterns in a species‐poor temperate woodland disturbed by fire

Abstract. Question: How does fire affect the aggregation patterns of trees in a species‐poor oak woodland? Location: East‐central Minnesota, USA. Methods: More than 10 000 trees with DBH > 2 cm (comprising more than 11 000 stems) were monitored in a 16‐ha grid on an annual basis from 1995‐ 2001 in a species‐poor temperate woodland. Different portions of the grid experienced different frequencies of controlled burns. Aggregation indices were calculated for individual species and individual size classes within species. A community‐wide aggregation index was also calculated for different burn units. Spatial data were managed, and many of the aggregation indices calculated using a GIS ArcInfo? (ESRI). Results: Fire initially increased clumping, although repeated fires reduced it, a finding that suggests a corollary to the intermediate‐disturbance hypothesis, the corollary stating that intermediate levels of disturbance are expected to maximize community‐wide patterns of aggregation. Analyses also showed that all species are aggregated at small scales, that the degree of aggregation of a stem type (species or size) declines with distance from individual stems, that the degree of aggregation of large stems is usually less than that of small stems, and that rare species are more aggregated than common species. Findings from this study are consistent with those from similar studies in other temperate and tropical forests, woodlands, and savannas. Conclusion: The spatial patterns of trees in this woodland are dynamic, continually changing in response to the relative strengths of the often opposing forces of competition, which tends to reduce clumping, and disturbance, which, at low and intermediate frequencies, tends to increase it.     

Scale‐dependence of environmental effects on species richness in oak savannas

Abstract. In order to investigate how scale (grain size) affects the relationships between species richness and environmental drivers (such as stress and disturbance), we collected 12 nested quadrats (from 0.25 m² to 1023 m²) from seven remnant oak savannas located in the floodplain of the Chippewa River in western Wisconsin, USA. Large and small‐scale richness were not significantly correlated, suggesting that small‐scale richness is not strongly controlled by sampling effects of the local species pool. Linear and curvilinear regressions between species richness and disturbance, canopy cover, biomass, and soil organic matter were dependent on sampling scale (grain size). Disturbance by fire was strongly related to richness at small scales, while tree canopy cover was strongly related to richness at larger scales. While there was some evidence suggesting the transition from disturbance to canopy effects occurs between 10 and 100 m², the transition was not particularly abrupt. The results cast doubt on the general importance of local species pools in affecting small‐scale richness as well as our ability to make generalizations that do not explicity include scale.     

中国东部亚热带森林土壤呼吸的时空格局

土壤呼吸是陆地碳循环中仅次于全球总初级生产力的第二大碳通量途径, 揭示土壤呼吸的时空格局对整个陆地碳循环具有重要意义。该文在中国东部亚热带季风气候区, 按纬度梯度由南向北选取深圳梧桐山、杨东山十二度水保护区、宁波天童山3个区域作为研究对象, 于2009年8月至2010年10月测定了不同季节各个区域内代表性植被类型的土壤呼吸速率及地下5 cm处土壤温度, 旨在初步了解中国东部亚热带森林地区土壤呼吸的时空格局及其影响因素。结果显示: 3个区域的土壤呼吸速率均存在显著的季节变化, 其变幅为2.64-6.24 μmol CO₂·m ^-2·s^-1, 总体趋势和地下5 cm处土壤温度的季节变化一致, 均为夏季最高冬季最低; 土壤温度的变化可以解释不同样地土壤呼吸季节变化的58.3%-90.2%; 各样地全年的Q₁₀值从1.56到3.27; 通过离样地最近的气象站点的日平均气温与试验样地地下5 cm处土壤温度之间的线性正相关关系推算出日土壤温度的变化, 利用土壤呼吸速率和地下5 cm处土壤温度之间的指数关系, 估算出各样地全年的土壤CO₂通量为1 077-2 058 g C·m^-2·a^-1, 在全球所有生态系统类型中处于较高水平。     

Fire history of a prairie/forest boundary: more than 250 years of frequent fire in a North American tallgrass prairie

Questions: Most modern fire‐prone landscapes have experienced disruptions of their historic fire regimes. Are the primary tallgrass prairies of the Flint Hills reflective of a history of continuous fire occurrence? Did fire frequency, severity, size and seasonality change in connection with changes in land use? Has fire occurrence been related to drought conditions? Location: Edges of Cross Timbers forest stands at the Tallgrass Prairie Preserve (TGPP) in the Flint Hills of Osage County, Oklahoma, USA. Methods: Cross‐sections of 76 Quercus stellata were collected from Cross Timbers stands at or near the grassland edge in the TGPP. Dendrochronological methods were used to identify years of formation for tree rings and fire scars. Superposed epoch analysis was used to evaluate the effect of drought conditions on fire occurrence. Results: Fires were recorded in 46.6% of the years between 1729 and 2005. In 41 cross‐sections at one site, the mean fire interval between 1759 and 2003 was 2.59 years, with fire interval decreasing from a mean fire interval of 3.76 years in the early part of the record to 2.13 years in modern times. No extended periods without fire were recorded in the study area. Drought conditions had no significant effect on fire occurrence. Conclusions: In contrast with many fire‐prone landscapes worldwide, the prairies of the Flint Hills have experienced no recent fire suppression or exclusion. Changes in fire frequency mark transitions in land use, primarily from being traditionally used by Native Americans to being managed for cattle production.     

Fire-scars and polymodal age-structure provide evidence of fire-events in an Aleppo pine population in southern France

Aleppo pine (Pinus halepensis Mill.) is adapted to fire, the most important disturbance in Mediterranean ecosystems. This species is known to be sensitive to fire and is usually killed when burned, but has a powerful regenerative capacity following burning. Sometimes, however, pines survive within burned areas, suggesting a certain potential for recording fire events in terms of age-structure and eventually scars. In the present study, fire scars on Aleppo pine trunks were used to construct a 150-year fire record in a burned forest in southern France. Four fires were detected with a mean fire return interval of 27 years: minimum 13 years and maximum 45 years. The age structure of the Aleppo pine population was analyzed on the basis of dated fire-scars in order to test the role of fire in creating the polymodal structure of the pine population. Results show the presence of multiple “cohorts” (subpopulations) of Aleppo pine. There appear to have been several pulses of tree recruitment, with the two largest cohorts corresponding to regeneration after the 1919 and 2000 fires. Other fires were recognized based on fire scars but these were not associated with obvious post-fire pine recruitment. Regeneration success is likely to depend on fire severity and on the length of fire-free intervals. Fire history and associated age-structure can thus be reconstructed using Aleppo pine stands.     

四种温带森林土壤氮矿化与硝化时空格局

利用PVC管原位培养连续取样法测定了东北地区4种具有代表性的森林生态系统(硬阔叶林、蒙古栎林、红松林、落叶松林)土壤氮素矿化、硝化的时间动态及氮矿化的空间分布格局.结果表明:4种森林土壤氮素矿化存在明显的时空变异.蒙古栎和红松林土壤在6月份表现出强烈的氮矿化和硝化作用,而硬阔叶林及落叶松林7月份氮素矿化强烈.4种森林生态系统上层土壤的氮净矿(硝)化率显著高于下层土壤.4种林型土壤的硝化过程在氮矿化过程中占有重要地位,其NO-3-N在无机氮中的比例分别为:79.9%～91.1%(硬阔叶林)、50.7%～80.5%(蒙古栎林)、54.1%～92.0%(红松林)、63.7%～86.5%(落叶松林).生态系统构成决定了土壤氮素的矿化能力.阔叶林和针阔混交林生态系统矿化率大于纯针叶林生态系统.硬阔叶林、红松林、蒙古栎林、落叶松林的平均净矿化率分别为:(0.58±0.01) mg · kg-1 · d-1、(0.47±0.19) mg · kg-1 · d-1、(0.39±0.11) mg · kg-1 · d-1和(0.23±0.06) mg · kg-1 · d-1.4种林型氮素矿化作用与地下5 cm温度呈正相关,并受土壤表层 (0～10 cm)水分显著影响.土壤微生物量氮与土壤氮矿化呈显著正相关.     

Tree shelters facilitate brown oak seedling survival and establishment in a grazing‐dominant forest of Bhutan,Eastern Himalaya

Brown oak (Quercus semecarpifolia) forest is essential for ecological and socioeconomic functions, mainly grazing in the Himalayas. The tree has failed to regenerate naturally and is a threatened species. Restoration of brown oaks is crucial to ensure sustainability while maintaining livestock grazing in these habitats. Achieving this requires cost‐effective restoration techniques that are practicable and sympathetic to the multiple uses of the forest. We assessed the combined effect of grazing (control) and three tree shelters (Protex tubes, mesh wires, and wooden frames) on the field performance of oak seedlings in a forest with heavy grazing pressure. Seedling survival and morphological indicators, including seedling height, collar diameter, sturdiness quotient (SQ), and leaf mass per area (LMA) indices, were measured. More than 90% of control seedlings without protective shelters suffered severe browsing and demonstrated significantly lower survival rates compared to tree shelter seedlings, indicating that grazing was the primary factor governing regeneration success. Seedling survival in tree shelters was three times higher, while the height increase was two times higher than the control. Additionally, locally made mesh wire and wooden tree shelters were more effective than Protex and control in producing quality seedlings reflected by the SQ and LMA values. We suggest that tree shelter is a promising option to restore brown oaks due to its efficacy to defend grazing and support the local community's rights to forest grazing. Our finding is expected to support Bhutan's forest policy of incorporating grazing and tree regeneration into forest management.     

Radial growth and climate responses of white oak (Quercus alba) and northern red oak (Quercus rubra) at the northern distribution limit of white oak in Quebec, Canada

Aim The objectives of this study were: (1) to compare radial growth patterns between white oak (Quercus alba L.) and northern red oak (Quercus rubra L.) growing at the northern distribution limit of white oak; and (2) to assess if the radial growth of white oak at its northern distribution limit is controlled by cold temperature. Location The study was conducted in three regions of the Ottawa valley in southern Québec. All stands selected were located at the northern limit of distribution of Q. alba. Methods Twelve mixed red and white oak stands were sampled and increment cores were extracted for radial growth analyses. For each oak species, 12 chronologies were derived from tree‐ring measurement (residual chronologies). Principal components analysis and redundancy analysis were used to highlight the difference between radial growth in both species and to determine their radial growth–climate association. Results There was little difference between the radial growth of each species; Q. alba, however, exhibits more year‐to‐year variation in growth than Q. rubra. More than 65% of the variance in radial growth was shared among sites and species. Both species showed a similar response to climate, which suggested that the limit of distribution of Q. alba might not be determined by effects on growth. Both species had a classic response to climate and drought in the early growing season. Main conclusions The northern distribution limit of Q. alba does not appear to be directly controlled by effects on growth processes as indicated by the similarities in radial growth and response to climate between the two species. The location of the stands on southern aspects suggested that cold temperature could have been a major factor controlling the distribution limit of Q. alba. However, it is speculated that stands growing on southern aspects may be more prone to forest fires or to drought, which would favour the maintenance and establishment of oaks, and of Q. alba in particular. Models relating the northern distribution limits of species to broad climate parameters like annual mean temperature will need to be reviewed to incorporate more biologically relevant information. Such assessments will in turn provide better estimates of the effect of climate changes on species distribution.     

Seasonal fire effects on the diversity patterns,spatial distribution and community structure of forbs in the Northern Mixed Prairie,USA

The effects of fire season on forb diversity patterns, density, and composition were determined for a northern Mixed Prairie site, USA. Repeated spring burns (dormant season), summer burns, fall burns (dormant season), and unburned treatments were compared over a 3-yr period characterized by wet and dry moisture conditions. Alpha and beta diversity were highest on unburned and summer burn treatments, while landscape mosaic diversity was highest on fall burns. Forb density was highest on fall and spring burn sites. Nine forb species comprised 82% of total densities and were significantly affected by fire season and year to year variations in moisture. Forb composition for unburned and spring burn treatments was similar, but both treatments were different from the summer burn and fall burn treatments which were similar to each other. Fire alone did not appear to be an intense enough disturbance to initiate drastic changes in the forb component of vegetation patches. Specific fire seasons did appear to either mask or enhance forb structure arising from other disturbance(s). Fire season also affected the scales of forb organization in the landscape. Contrasting spatial characteristics of the forb component of prairie plant communities may provide a diagnostic technique for exposing the interaction of disturbances at different temporal and spatial scales.     

High and dry: post‐fire tree seedling establishment in subalpine forests decreases with post‐fire drought and large stand‐replacing burn patches

Aim Climate warming and increased wildfire activity are hypothesized to catalyse biogeographical shifts, reducing the resilience of fire‐prone forests world‐wide. Two key mechanisms underpinning hypotheses are: (1) reduced seed availability in large stand‐replacing burn patches, and (2) reduced seedling establishment/survival after post‐fire drought. We tested for regional evidence consistent with these mechanisms in an extensive fire‐prone forest biome by assessing post‐fire tree seedling establishment, a key indicator of forest resilience. Location Subalpine forests, US Rocky Mountains. Methods We analysed post‐fire tree seedling establishment from 184 field plots where stand‐replacing forest fires were followed by varying post‐fire climate conditions. Generalized linear mixed models tested how establishment rates varied with post‐fire drought severity and distance to seed source (among other relevant factors) for tree species with contrasting post‐fire regeneration adaptations. Results Total post‐fire tree seedling establishment (all species combined) declined sharply with greater post‐fire drought severity and with greater distance to seed sources (i.e. the interior of burn patches). Effects varied among key species groups. For conifers that dominate present‐day subalpine forests (Picea engelmannii, Abies lasiocarpa), post‐fire seedling establishment declined sharply with both factors. One exception was serotinous Pinus contorta, which did not vary with either factor. For montane species expected to move upslope under future climate change (Larix occidentalis, Pseudotsuga menziesii, Populus tremuloides) and upper treeline species (Pinus albicaulis), establishment was unrelated to either factor. Greater post‐fire tree seedling establishment on cooler/wetter aspects suggested local topographic refugia during post‐fire droughts. Main conclusions If future drought and wildfire patterns manifest as expected, post‐fire tree seedling establishment of species that currently characterize subalpine forests could be substantially reduced. Compensatory increases from lower montane and upper treeline species may partially offset these reductions, but our data suggest important near‐ to mid‐term shifts in the composition and structure of high‐elevation forests under continued climate warming and increased wildfire activity.     

Human disturbance and environmental factors as drivers of long‐term post‐fire regeneration patterns in Mediterranean forests

Question: What are the main forces driving natural regeneration in burned mature Mediterranean forests in the medium‐long term and what are the likely successional trajectories of unmanaged vegetation? Location: Valencia Region, eastern Spain. Methods: A wildfire burned 33 000 ha of Pinus halepensis and P. pinaster forest in 1979, and subsequent smaller wildfires took place between 1984 and 1996. The study was designed to sample the range of environmental and disturbance (fire recurrence and land use) conditions. The territory was classified into 17 different geomorphological and fire‐recurrence units. Vegetation cover and floristic composition were measured on a total of 113 plots (1000 m² each) randomly selected within these units. Results: The results show that 23 years after the fire the regenerated vegetation consists of successional shrublands, and that forest ecosystem resilience can be very low. The vegetation presents a strong correlation with most of the environmental variables, but fire (one or two fires), soil type and land use (in that order) are the main drivers of vegetation composition. Quercus coccifera shrublands persist on limestone soils while diverse types of other shrublands (dominated by seeder species) are found on marl soils. Conclusions: The results of this study indicate that disturbance factors strongly coupled to human activities, such as land use and fire, play a critical role in the current state of vegetation. Fire creates vegetation patches in different successional states while land use and soil type define the different types of shrubland in terms of their specific composition.     

Human and non-human determinants of forest composition in southern Spain: evidence of shifts towards cork oak dominance as a result of management over the past century

Aims Both human and non‐human determinants have shaped Mediterranean forest structure over the last few millennia. The effects of recent human activities on forest composition, however, remain poorly understood. We quantified changes in forest composition during the past century in the mixed forests of Quercus suber (cork oak) and Q. canariensis (Algerian oak), and explored the effects of forest management and environmental (climate, topography) factors on forest structure at various spatial and temporal scales. Location Mountains north of the Strait of Gibraltar (southern Spain). Methods First, we quantified 20th‐century changes in species composition from a series of inventories in nine mixed forests (c. 40,000 ha), and examined them in terms of the management practices and environmental conditions. Second, we analysed present‐day Q. suber and Q. canariensis stand structure along environmental gradients at two spatial scales: (1) that of the forest landscape (c. 284 ha), combining local inventories and topographic variables and using a digital elevation model; and (2) regional (c. 87,600 km²), combining data from the Spanish Forest Inventory for the Andalusia region and estimates of climatic variables. Results Historical data indicate anthropogenic changes in stand composition, revealing a sharp increase in the density of cork oaks over the last century. Forest management has favoured this species (for cork production) at the expense of Q. canariensis. The impact of management is imprinted on the present‐day forest structure. The abundance of both species increases with annual mean precipitation, and they coexist above 800 mm (the minimum threshold for Q. canariensis). Quercus suber dominates in most of the stands, and species segregation in the landscape is associated with the drainage network, Q. canariensis being clearly associated with moister habitats near streams. Main conclusions Our study quantitatively exemplifies a recent human‐mediated shift in forest composition. As a result of forest management, the realized niche of the cork oak has been enlarged at the expense of that of Q. canariensis, providing further evidence for humans as major drivers of oak forest composition across the Mediterranean. Recent regeneration problems detected in Q. suber stands, a reduced demand for wood products, conservation policies, and climate change augur new large‐scale shifts in forest composition.