Long-term effects of single prescribed fires on hardwood regeneration in oak shelterwood stands

One of the arguments against using prescribed fire to regenerate oak (Quercus spp.) forests is that the improvement in species composition of the hardwood regeneration pool is temporary and multiple burns are necessary to achieve and maintain oak dominance. To explore this concern, I re-inventoried a prescribed fire study conducted in the mid-1990s to determine the longevity of the effects of a single prescribed fire on hardwood regeneration. The initial study was conducted in three oak shelterwood stands in central Virginia, USA. In 1994, each stand was divided into four treatments (spring, summer, and winter burns and a control) and the hardwood regeneration was inventoried before the fires. During the burns, fire intensity was measured and categorized in each regeneration sampling plot. Second-year postfire data showed marked differences in species mortality rates, depending on season-of-burn and fire intensity: oak and hickory (Carya spp.) regeneration dominated areas burned by medium- to high-intensity fire during the spring and summer while yellow-poplar (Liriodendron tulipifera) and red maple (Acer rubrum) seedlings dominated unburned areas and all areas treated with low-intensity fire regardless of season-of-burn. The treatments were re-inventoried in 2006 and 2007 to determine whether these fire effects were still present. The new data show that the species distributions by season-of-burn and fire intensity found in 1996 still existed 11 years after the treatments. The fact that fire effects in oak shelterwood stands can last at least a decade has important management implications for resource professionals interested in sustaining oak forests in the eastern United States.     

Restoration of bottomland hardwood forests across a treatment intensity gradient

Large-scale restoration of bottomland hardwood forests in the Lower Mississippi Alluvial Valley (USA) under federal incentive programs, begun in the 1990s, initially achieved mixed results. We report here on a comparison of four restoration techniques in terms of survival, accretion of vertical structure, and woody species diversity. The range of treatment intensity allowed us to compare native recolonization to direct seeding and planting of Quercus nuttallii Palmer, and to an intensive treatment of interplanting two species that differed in successional status (early successional Populus deltoides Bartram ex Marsh. ssp. deltoides, with the mid-successional Q. nuttallii). Native recolonization varied in effectiveness by block but overall provided few woody plants. All active restoration methods (planting and direct seeding) were successful in terms of stocking. Populus grew larger than Quercus, reaching canopy closure after 2 years and heights after 2 and 5 years of 6 and 12.7 m, respectively. Planted Quercus were significantly larger than direct seeded Quercus in all years, but only averaged 1.4 m in height after 5 years. Interplanting did not seem to facilitate development of the Quercus seedlings. The early success of the interplanting technique demonstrated that environmental benefits can be obtained quickly by more intensive efforts. Native recolonization can augment active interventions if limitations to dispersal distance are recognized. These results should provide landowners and managers with the confidence to use techniques of varying intensity to restore ecosystem functions.     

The influence of mountain pine beetle outbreaks and drought on severe wildfires in northwestern Colorado and southern Wyoming: A look at the past century

Outbreaks of bark beetles and drought both lead to concerns about increased fire risk, but the relative importance of these two factors is the subject of much debate. We examined how mountain pine beetle (MPB) outbreaks and drought have contributed to the fire regime of lodgepole pine forests in northwestern Colorado and adjacent areas of southern Wyoming over the past century. We used dendroecological methods to reconstruct the pre-fire history of MPB outbreaks in twenty lodgepole pine stands that had burned between 1939 and 2006 and in 20 nearby lodgepole pine stands that were otherwise similar but that had not burned. Our data represent c. 80% of all large fires that had occurred in lodgepole pine forests in this study area over the past century. We also compared Palmer Drought Severity Index (PDSI) and actual evapotranspiration (AET) values between fire years and non-fire years. Burned stands were no more likely to have been affected by outbreak prior to fires than were nearby unburned stands. However, PDSI and AET values were both lower during fire years than during non-fire years. This work indicates that climate has been more important than outbreaks to the fire regime of lodgepole pine forests in this region over the past century. Indeed, we found no detectable increase in the occurrence of high-severity fires following MPB outbreaks. Dry conditions, rather than changes in fuels associated with outbreaks, appear to be most limiting to the occurrence of severe fires in these forests.     

Regeneration dynamics of a mixed Mediterranean pine forest in the absence of fire

The aim of this study is to determine the competing regeneration and expansion patterns of two co-occurring pine species (Pinus brutia, Pinus nigra ssp. pallasiana), in a transitional montane Mediterranean zone. We measured the regeneration density of all woody species in 102 randomly located stands along an altitudinal gradient on the island of Lesbos, Greece. Individuals of pines were assigned to different size classes. Topographic factors (altitude, aspect, and soil depth) and light availability (through hemispherical photographs) were measured for each stand. Statistical analyses were applied to explore the effect of each factor on recruitment density of the competing pine species, and to elucidate patterns of interaction. Canopy openness was the most important parameter controlling the recruitment of P. brutia, while the regeneration density of P. nigra was mainly related to canopy openness and heat load. An idiosyncratic response of the recruitment vigour of the two species was identified along gradients of shade and drought stress. The decline in P. nigra recruitment density with drought conditions underlines threats to its population maintenance even in the absence of fire. On the other hand P. brutia seems to be a stronger invader in transitional zones. The studied species could be considered typical representatives of the two most widely distributed pine functional types across the Mediterranean basin, and our results agree with the theoretical ability of such species to maintain and expand their populations.     

Impacts of increased fire frequency and aridity on eucalypt forest structure,biomass and composition in southwest Australia

Water stress and fire disturbance can directly impact stand structure, biomass and composition by causing mortality and influencing competitive interactions among trees. However, open eucalypt forests of southwest Australia are highly resilient to fire and drought and may respond differently to increased fire frequency and aridity than forests dominated by non-eucalypt species. We measured the variation in stem density, basal area, stand biomass, sapwood area, leaf area and litterfall across 16 mixed jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) forest stands along an aridity gradient in southwest Australia that had variable fire histories. Fire frequency was defined as the total number of fires over a ∼30-year period and aridity as the ratio of potential evapotranspiration to annual precipitation. Total stand biomass and sapwood area were predicted from diameter at breast height of individual jarrah and marri trees using allometric equations. Leaf area was estimated using digital cover photography. More arid and frequently burnt stands had higher stem density, especially of smaller trees, which were mainly jarrah. Overall, both standing biomass and leaf area decreased at more arid sites, while sapwood area was largely unaffected by aridity, suggesting that these stands respond to increased water limitation by decreasing their leaf area relative to their sapwood area. Biomass of marri was reduced at more arid and, to a lesser extent, at more frequently burnt stands. However, total stand biomass (jarrah and marri) and leaf area index did not vary with fire frequency, suggesting that less marri biomass (due to slower growth rates, higher mortality or less recruitment) was compensated by an increase in the density of jarrah trees (regeneration). We conclude that increased fire and drought shift tree species composition towards more fire-resistant species and result in denser stands of smaller trees. In contrast, total stand biomass declines with increasing aridity, but has no association with fire frequency.     

Fire resistance of tree species explains historical gallery forest community composition

Fire, climatic variability, and grazing by large herbivores have historically limited woody vegetation in the tallgrass prairie region of North America to gallery forests in protected areas along rivers and streams. Fire, in particular, has been a strong selective pressure against woody vegetation. Consequently, we expect that dominant tree species in these forests have developed mechanisms for tolerating periodic surface fires. Susceptibility of trees to fire damage depends in part on key properties of bark which influence heat transfer to the vascular cambium, including thickness, density, and moisture content. An historical (1983) survey of Konza Prairie Biological Station in northeast Kansas, USA indicated that gallery forests were co-dominated by Quercus macrocarpa and Quercus muehlenbergii, while Celtis occidentalis occurred as an important sub-dominant species. Populus deltoides, Gleditsia triacanthos, and Juniperus virginiana were relatively uncommon. To test the hypothesis that historically dominant gallery forest tree species are more resistant to fire damage than uncommon species, fire was applied to the bark of 10 individuals of each of these six species under conditions mimicking surface fires (400 °C for 120 s). Maximum temperature at the vascular cambium, bark thickness, bark moisture content, and bark density were measured. Trees were considered fire-resistant if the vascular cambium temperature remained below the thermal cell death threshold, 60 °C, throughout the treatment. Using logistic regression, bark thickness was found to be a significant predictor of lethal cambium temperatures (P = 0.002), while neither bark density nor moisture content were significantly related to lethal cambium temperature (P = 0.279 and P = 0.131, respectively). Across species, a minimum bark thickness of approximately 8.6 mm was necessary to maintain the vascular cambium temperature below 60 °C. Trees that produce thick bark quickly in juvenile size classes (P. deltoides, Q. macrocarpa, and Q. muehlenbergii) experienced lower temperatures at the vascular cambium than those which do not develop thick bark with increasing diameter (C. occidentalis, G. triacanthos, and J. virginiana). Ranking these tree species by either the DBH or age needed to develop the minimum protective bark thickness largely agreed with ranking based on historical relative importance. As fire frequency and intensity decrease in remnant tallgrass prairie of North America as a result of habitat fragmentation, fire suppression, and changing land management, fire-sensitive species may increase in relative importance in gallery forests because of increased juvenile survival.     

Does inorganic nitrogen limit plant growth 3–5 years after fire in a Wyoming,USA, lodgepole pine forest?

Nitrogen (N) is the major nutrient limiting plant growth and production in terrestrial ecosystems around the world. However, nutrient limitation is spatially variable, and different species within the same ecosystem may be limited by different nutrients. N constraints on plant growth have been investigated via fertilization experiments in a variety of ecosystems; however, recently burned coniferous forests are notably under-studied. Given the recent increase in fire activity across western North America, it is important to understand what limits plant growth and carbon sequestration in coniferous forests recovering from wildfire. We evaluated potential inorganic N limitation in four common native plant species, 3–5 years after stand-replacing wildfire in a lodgepole pine forest (Pinus contorta var. latifolia) in Wyoming, USA. Granular reagent grade ammonium nitrate was added around individual plants at a rate equal to the natural background rate of net N mineralization and at 10× this rate. The grass Calamagrostis rubescens exhibited clear evidence of inorganic N limitation: above-ground biomass and shoot:root ratio increased with the high-fertilizer treatment. Nitrogen:phosphorus (N:P) ratio in un-fertilized C. rubescens plants was <14, also consistent with N limitation, but N:P ratio shifted to >16 in the high-fertilizer treatment, suggesting the onset of P limitation. The upland sedge Carex rossii and seedlings of lodgepole pine were not limited by inorganic N: neither species showed any growth response to N fertilization; N:P ratios were only slightly <14; and foliar N concentrations were greater than critical values reported for mature lodgepole pine. The N-fixing forb Lupinus argenteus was not limited by N, for it showed no growth response to fertilization; rather its N:P ratio of 21 indicated P limitation. In this study, to our knowledge the first experimental evaluation of N limitation in subalpine coniferous forests following wildfire, N limitation was seen in only one of four species tested.     

Persistent effects of fire severity on early successional forests in interior Alaska

There has been a recent increase in the frequency and extent of wildfires in interior Alaska, and this trend is predicted to continue under a warming climate. Although less well documented, corresponding increases in fire severity are expected. Previous research from boreal forests in Alaska and western Canada indicate that severe fire promotes the recruitment of deciduous tree species and decreases the relative abundance of black spruce (Picea mariana) immediately after fire. Here we extend these observations by (1) examining changes in patterns of aspen and spruce density and biomass that occurred during the first two decades of post-fire succession, and (2) comparing patterns of tree composition in relation to variations in post-fire organic layer depth in four burned black spruce forests in interior Alaska after 10-20 years of succession. We found that initial effects of fire severity on recruitment and establishment of aspen and black spruce were maintained by subsequent effects of organic layer depth and initial plant biomass on plant growth during post-fire succession. The proportional contribution of aspen (Populus tremuloides) to total stand biomass remained above 90% during the first and second decades of succession in severely burned sites, while in lightly burned sites the proportional contribution of aspen was reduced due to a 40-fold increase in spruce biomass in these sites. Relationships between organic layer depth and stem density and biomass were consistently negative for aspen, and positive or neutral for black spruce in all four burns. Our results suggest that initial effects of post-fire organic layer depths on deciduous recruitment are likely to translate into a prolonged phase of deciduous dominance during post-fire succession in severely burned stands. This shift in vegetation distribution has important implications for climate-albedo feedbacks, future fire regime, wildlife habitat quality and natural resources for indigenous subsistence activities in interior Alaska.     

Fire-driven oak regeneration in French Mediterranean ecosystems

Regeneration by seeds for cork oak (Quercus suber) and companion oaks (holm oak Quercus ilex and downy oak Quercus pubescens) is likely to be poor in the fire-prone Maures massif (southern France) but the causes are poorly known. Our objective was to assess the effective recruitment for these three oak species and their temporal pattern of recruitment, in order to determine the main limitation factors and the regeneration window of each species. We studied oak recruits (height <3 m) in naturally regenerated populations according to a gradient of fire recurrence and in five main vegetation types including shrublands and mixed mature woodlands. Fire recurrence was the main explanatory factor of oak recruitment, either directly or through vegetation type and microsite characteristics. The results indicate nil to low recruitment for holm oak and downy oak in shrublands, especially those recurrently burned and dominated by Cistus species. Cork oak recruited better than the other oaks in medium and high shrublands dominated by Erica arborea. In contrast, recruitment was high for holm and downy oak in mixed oak stands and mixed pine-oak stands that have not burned for decades. Microsite conditions such as coverage by litter and shrubs influenced oak recruitment, whereas landscape configuration and stand basal area had no influence. Our results suggest that strategic shrub-clearing, oak planting and protection of mixed oak woodlands as seed sources would help maintaining oak populations in the woodland–shrubland mosaic.     

Drought matters - Declining precipitation influences growth of Fagus sylvatica L. and Quercus robur L. in north-eastern Germany

For north-eastern Germany regional climate models project rising temperatures in combination with decreasing summer and increasing winter precipitation. The resulting overall drier conditions during the growing season will considerably impact forest growth there. We evaluate the consequences of increasing drought on the growth of the two locally most important broadleaf tree species common beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.). Three mixed forests of beech and oak were sampled along a west-east gradient of declining precipitation. In total we used 257 ring-width samples from 133 trees to build six species and site specific chronologies. Additionally, we modelled the soil water budget for each site. We performed continuous and discontinuous (pointer year) analysis of climate-tree-growth relationships with particular emphasis on inter-annual-variations and their dependence upon climatic factors (temperature, precipitation, soil moisture) and on the stability of the obtained relationships. Results of climate-growth correlations together with pointer year analysis indicate a strong dependency of growth of both species from water availability, especially during early summer (June and July). General correlation pattern between growth and climate are similar for both species, but climate sensitivity of beech is generally higher. We identified drought as the main driver of negative growth depressions in both species. Increasing drought stress along the gradient is expressed in higher correlations to climatic variables, higher sensitivity (variance) of growth, and a higher number of negative pointer years for both species. For beech we also found a significant trend of decreasing average growth rates along the gradient. Growth superiority of beech compared to oak declines with decreasing precipitation. The relationships were generally stable throughout the 20th century. A rise of sensitivity together with a higher frequency of negative pointer years during the last decades suggests that increasing climatic variability together with rising temperatures might be influencing growth of Fagus at the more humid sites. If we substitute space by time it seems that already small changes in precipitation regime can have considerable impact, especially on the growth of beech. Other, more drought tolerant species like oak might gain competitive advantages under the projected climatic changes.     

Tree rings indicate different drought resistance of a native (Abies alba Mill.) and a nonnative (Picea abies (L.) Karst.) species co-occurring at a dry site in Southern Italy

Climate changes induced by the anthropogenic alteration of the atmospheric radiative balance are expected to change the productivity and composition of forest ecosystems. In Europe, the Mediterranean is considered one of the most vulnerable regions according to climatic forecasts and simulations. However, although modifications in the inter-specific competition are envisaged, we still lack a clear understanding of the ability of the Mediterranean vegetation to adapt to climate changes. We investigated how two co-occurring tree species commonly used in afforestation programmes, the native Abies alba Mill. and the nonnative Picea abies L. Karst., adapt to climate change by assessing their growth performance and physiological responses in relation to past climate variability. Growth was addressed by analysing tree-ring width and carbon and oxygen stable isotopes. Statistical relationships between isotopic value and monthly climate data suggest that the two species underwent ecophysiological adaptation to Mediterranean climatic constraints. These adaptations are also expressed in the ring-width data. Based on the carbon isotope ratio reflecting the stomatal response to drought, we found that the precipitation in the first period of the growing season, i.e. early spring, is a major factor influencing the annual growth of A. alba, which although native, proved to be sensitive to drought. P. abies, on the other hand, showed a higher tolerance to summer drought stress. These findings should help define criteria for sustainability and effective forest conservation in the Mediterranean region.     

The pollen record of a 20th century spruce beetle (Dendroctonus rufipennis) outbreak in a Colorado subalpine forest,USA

The frequency and intensity of ecosystem disturbance, including outbreaks of forest insects and forest fires, is expected to increase in the future as a result of higher temperatures and prolonged drought. While many studies have concentrated on the future climatic impacts on fire, little is known about the impact of future climate on insect infestation. Paleoecological techniques are important in this regard in identifying the potential relationships between climate and insect outbreaks in the past, as a predictive tool for the future. We examine a high-resolution 20th century record of spruce beetle (Dendroctonus rufipennis) infestation from a small, subalpine lake, comparing the paleoecological record to the historical and tree-ring record of the event. An extensive spruce beetle outbreak occurred in northwestern Colorado during the 1940s and 1950s, causing widespread mortality of mature Picea engelmannii. Pollen analysis of this period documents the decline of Picea and its replacement locally by Abies lasiocarpa, paralleling age and composition studies of modern forest stands in the region. This study is a proof of concept that, when applied to longer sedimentary records, could produce a detailed record of infestation for the Late Holocene or older time periods. This information will be useful to forest managers in efforts to plan for the effects of D. rufipennis infestations, and subsequent succession within high elevation conifer forests.     

Positive association between understory species richness and a dominant shrub species (Corylus avellana) in a boreonemoral spruce forest

Old growth stands of boreonemoral spruce (Picea abies) forests frequently have a shrub layer dominated by hazel (Corylus avellana) – a species which is generally excluded in intensively managed forests due to clearcutting activities. We sampled understory species composition, richness and biomass, as well as environmental variables beneath these two species and also within forest ‘gaps’ in order to determine the effect of overstory species on understory vegetation. Species richness and biomass of herbaceous plants was significantly greater under Corylus compared with plots under Picea and in forest gaps. Indicator species analysis found that many species were significantly associated with Corylus. We found 45% of the total species found under woody plants occurred exclusively under Corylus. Light availability in spring and summer was higher in gaps than under forest cover but no difference was found between plots under Corylus and Picea. Hence, reductions in light availability cannot explain the differences in species composition. However, Ellenberg indicator values showed that more light demanding species were found under Corylus compared to Picea, but most light demanding species were found in gaps. The litter layer under Picea was three times thicker than under Corylus and this may be an important mechanism determining differences in understory composition and richness between the woody species. The presence of Corylus is an important factor enhancing local diversity and small-scale species variation within coniferous stands. Hence, management should maintain areas of Corylus shrubs to maintain understory species diversity in boreal forests.     

我国栓皮栎分布及其生态学研究

栓皮栎作为广泛分布的树种之一,在全球气候变化的大环境下,成为研究树木生长应对气候变化的重要树种。文中分析了我国栓皮栎水平分布和垂直分布的特点及其与气候的关系,栓皮栎在不同地区的森林类型,主要包括栓皮栎纯林、松栎混交林、栎类混交林等类型;综述了我国对栓皮栎生态学、群落结构、更新等研究的最新进展和主要结论;提出今后开展栓皮栎研究的重点和方向,以期为栓皮栎生理生态学及其对气候变化响应研究提供参考。     

栎属树种生长模型研究进展

栎属树种是亚热带常绿阔叶林和温带落叶阔叶林的主要建群树种之一,分布范围极为广泛,在国内外被广泛应用于城市园林绿化、水源涵养林、水土保持林,也是重要的珍贵用材树种,同时其果实、栓皮等具有重要的工业和药用价值.研究栎林的生长过程,对其经营抚育决策具有重要的参考意义.文中对近年来国内外关于栎属树种的生长过程,特别是栎属树种生长模型的研究进行了综述,包括栎属树种全林分生长模型、单木生长模型、径阶分布模型等;阐述了栎属树种生长模型的研究现状及发展趋势,以期为栎林的经营抚育决策提供参考.     

Composition and structure of natural mixed-oak stands in northern and central Portugal

Stand composition and structure of natural mixed-oak stands of common-oak (Quercus robur L.) and pyrenean-oak (Quercus pyrenaica Willd.) were studied. Diverse compositional and structural elements in early and late successional stand stages were analysed. The study was conducted in north and central Portugal where different natural mixed oak forests types are located. The following mixed-oak forest types involving common-oak and pyrenean-oak were studied: common-oak & other hardwoods; common-oak & cork-oak (Quercus suber L.); ash (Fraxinus angustifolia Vahl) & pyrenean-oak; and pyrenean-oak & madrone (Arbutus unedo L.). Measurements were made in early and late successional stand stages on the different mixed oak forest types. Different stand characteristics and indices were used to describe and compare stand structure and composition. The study showed changes in species diversity and stand structure. Most tree species in mature stands are present in early stages but with higher abundance. Shannon diversity index may change between 0.798 and 1.915. Significant differences on species diversity and abundance were found depending on the forest type and successional stage. Mature mixed-oak forests have high species diversity with an abundance of small to medium tree size species. Species distribution and diameter differentiation indices range from 0.30 to 0.70 and 0.52 to 0.82, respectively, revealing significant structural complexity. The average number of standing and downed dead trees was 265 and 83 trees ha⁻¹ for early and late stage, respectively, with 6.9 and 65.4 m³ ha⁻¹. Higher values of stand diversity index were 41 and 53 in more complex and developed forests. Later stand stages have complex structure, with a wider range of tree diameter distribution and higher degree of irregularity.     

Age and growth of a fire prone Tasmanian temperate old-growth forest stand dominated by Eucalyptus regnans, the world's tallest angiosperm

Forests are key components of the global carbon cycle, with deforestation being an important driver of increased atmospheric carbon dioxide. Temperate old-growth forests have some of the highest above ground stores of carbon of any forest types on Earth. Unlike tropical forests, the ecology of many temperate forests is dominated by episodic disturbance, such as high intensity fire. An exemplar of a particularly carbon dense temperate forest system adapted to infrequent catastrophic fires is the Eucalyptus regnans forests of south eastern Australia. Knowledge of the growth and longevity of old-growth trees is crucial to understanding the carbon balance and fire regimes of these forest systems. In an old-growth E. regnans stand in the Styx Valley in southern Tasmania we used dendrochronological techniques and radiocarbon dating to determine the age and stem growth of E. regnans and Phyllocladus aspleniifolius, an understorey rainforest conifer. Our analysis revealed that an even-aged cohort of E. regnans and P. aspleniifolius established in 1490–1510AD, apparently after a stand-replacing fire. The stem growth rates of E. regnans in the first 100 years were very rapid compared to the co-occurring P. aspleniifolius. That the longevity of E. regnans is >500 years challenges the suggested 350–450 year timeframe proposed for the widely held model of succession from eucalypt to rainforest. These forests not only have the potential to store vast amounts of carbon, but can also maintain these high carbon densities for a long period of time. Estimates of the capacity of these forests to sequester and store carbon should explicitly consider past harvesting and fire regimes and the potential increases in the risk of fire associated with climate change.     

Using a retrospective dynamic competition index to reconstruct forest succession

Understanding forest dynamics and stand structures is crucial for predicting forest succession. However, many forests have been altered due to century-long land-use practices, which complicates the reconstruction of past and current successional trajectories. For a better understanding of successional processes, we suggest studying the intra- and interspecific competition among single trees across time. We introduce a tree-ring based competition index to reconstruct the competitive dynamics of individual trees over time. This new retrospective dynamic competition index combines a temporal and a spatial component by calculating the yearly ratio between the basal area increments (bai) of the neighbouring trees and the subject tree. The new index is applied to mixed Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) stands in the inner-Alpine dry-valley Valais, for which a change in species composition is hypothesised. The aim is to analyse current stand structures in terms of recent changes in the competitive interactions at the single tree level and to relate these competitive dynamics to land-use change and increasing drought due to climate change. On five plots, the positions of 456 trees were recorded and increment cores were taken to derive bai data. The individual dynamic competition index curves were aggregated in clusters, which define typical patterns of competitive dynamics in both tree species. A large percentage of the trees (87% in oak, 70% in pine) were clustered into a group of trees with constant competition at a relatively low level. However, a smaller group of pines (20%) had recently faced increasing competition. In addition, stand structure analyses indicated a change towards a higher proportion of oak. This change in the competitive ability between oak and pine was found to be related to drought, in that oak had a competitive advantage in dry years. Furthermore, the high proportion of dead branches in pines with decreasing competitive abilities indicated increasing competition for light as a consequence of natural development towards a later successional stage that favours the more shade-tolerant oak. The new retrospective dynamic competition index proved to be promising in studying forest succession. The tree-ring based method allows us to identify changes in the competitive ability of single trees with a high temporal resolution and without repeated assessments.     

Diversity and structure of regenerating tropical dry forests in Costa Rica: Geographic patterns and environmental drivers

Much of the dry tropical forest biome has been converted to agricultural land uses over the past several centuries. However, in conservation areas such as those in the Guanacaste and Tempisque regions of Costa Rica, tropical dry forests are regenerating due to management practices including fire suppression. To better understand the patterns of secondary succession occurring in Costa Rican tropical dry forest, we established 60 20 × 50 m plots in mature and regenerating forests in the Sector Santa Rosa (formerly known as Parque Nacional Santa Rosa) and Palo Verde National Park. Plots were stratified into three plant communities: tropical dry oak forest (Quercus oleoides) (SROAK), Santa Rosa tropical dry forest (SRTDF), and Palo Verde tropical dry forest (PVTDF). In these plots we measured and identified and all individuals >10 cm DBH, measured but did not identify stems <10 cm but taller than 1.3 m, counted woody seedlings (<1.3 m height) and analyzed soil chemical and physical properties.