Regeneration dynamics of mixed stands of <Emphasis Type="Italic">Pinus pinaster</Emphasis> Ait. and <Emphasis Type="Italic">Pinus pinea</Emphasis> L. in Central Spain

The dynamics of mixed stands are more complex and less studied than those of monospecific stands. The objective of this work was to analyze the variables involved in seedling occurrence and seedling survival in mixed stands of Pinus pinaster and P. pinea in Mediterranean areas. From 2011 to 2016, regeneration of both species was monitored at two sites located in Central Spain. We installed 72 regeneration plots where seedling dynamics were monitored. All the trees in the study areas were measured and mapped. Additionally, we took hemispherical photographs in each regeneration plot. The average density of P. pinea seedlings over the study period was almost 20 times larger than that of P. pinaster. Our results indicate that the seedlings of both species grow under moderate light conditions. In addition, we found that the occurrence of seedlings of both species was related to the structure of the stand. P. pinea seedlings grew where the density and size of P. pinaster trees were low and where P. pinea trees provided moderately sheltered conditions, whereas the number of P. pinaster seedlings was related to under intermediate densities of P. pinaster trees. Furthermore, seedling survival was positively associated with age of the seedlings and negatively with the August average maximum temperature. The temporal continuity of mixed stands of P. pinea and P. pinaster in the study area is compromised by the observed lack of regeneration of P. pinaster.     

Impact of the pinewood nematode, Bursaphelenchus xylophilus, on gross calorific value and chemical composition of Pinus pinaster woody biomass

The pinewood nematode (PWN), Bursaphelenchus xylophilus, the causal agent of the pine wilt disease, has been detected in several regions of Portugal affecting Pinus pinaster, a coniferous species of a great economic value. The nematodes, migrating through resin canals and feeding on parenchyma cells, induce rapid metabolic changes in ray parenchyma cells, cavitation areas, and denaturation and necrosis of parenchyma and cambial cells. To understand how anatomic changes and biochemical incidences of tree defense reactions affect the technological parameters of the wood, the gross calorific value (GCV) and chemical composition of PWN-infected and -uninfected P. pinaster wood were evaluated. The GCV was determined using Parr 6300 Automatic Isoperibol Calorimeter, and chemical composition analysis was performed by determining the contents of carbon, hydrogen, nitrogen, sulfur and Ash on complete and instant oxidation of samples by “flash” combustion. The Student’s t test with Welch correction was used for statistical data analysis. The difference between the GCV and chemical composition for PWN-infected and -uninfected P. pinaster wood was statistically significant for the GCV and for hydrogen and nitrogen contents. The carbon, oxygen, sulfur and Ash contents did not differ statistically. The GCV of PWN-infected wood varied between the highest value of hardwood and the lowest value of softwood. This interdisciplinary study stresses the important technological and economic aspects, namely the impact of PWN on wood properties and the suitability of infected P. pinaster wood for use in the wood-processing and energy industries.     

Recovery of bioactive compounds from Pinus pinaster wood by consecutive extraction stages

Pinus pinaster wood samples, obtained at different positions of three healthy trees, were subjected to two sequential extractions using an Accelerated Solvent Extraction instrument. The first extraction was carried out with hexane (to remove lipophilic extractives) and the second one with acetone/water (95:5 v/v) to recover bioactive phenolic compounds, the target compounds of this study. The extracted fractions were assayed for total yield and composition. The extracts contained a spectrum of phenolic compounds (simple phenolics, phenolic stilbenes, flavonoids and lignans) and non-phenolic components (juvabiones, resin and fatty acids, steryl esters and triglycerides). The fractionation effects achieved by consecutive extractions and the recovery of bioactive phenolics are discussed.     

Climate and groundwater effects on the establishment, growth and death of Prosopis caldenia trees in the Pampas (Argentina)

Semiarid woodlands dominated by Prosopis caldenia thrive at the dry edge of the Argentinean Pampas. Deforestation and increased precipitation have driven sustained water table level rise in the region that are likely to affect the dynamics of remnant woody vegetation patches. Here we analyze the effect of climate and groundwater level on the establishment, growth and death of P. caldenia located on lowland (current water table <0.5 m deep) and upland (current water table 8 m deep) positions within rolling sandy landscape. Standard dendrochronological techniques were applied on 98 cross sections and cores of P. caldenia trees. Results allowed us to estimated that trees in the lowland established a few years earlier than in the upland (1929 vs. 1936) and died between 1991 and 2002, while trees in higher positions are still alive. As a result of a faster growth in the lowland, maximum mean basal area increment took place earlier (1950s vs. 1990s) and achieved a higher mean value than in the upland (41.63 vs. 37.41 mm²). While mean annual chronologies were not associated across stands, an opposite highly significant association was found for the mean growth trends suggesting long-term effects of water table depths on growth. We found a different association between mean annual growth and climate, with lowland trees showing a negative response to precipitation before and during the growing season, and upland trees displaying a positive response to summer rainfall inputs. Temperature at the end of the growing season had a positive effect in the lowland whereas temperature during the growing season had a positive effect in the upland. These results show how groundwater can induce diverging sensitivity of forest growth and survival to climate variability, enhancing growth at optimum depths (2-8 m in our study) but depressing it or even killing trees when it approaches the surface (<2 m in our study). Groundwater dynamic should be considered in forest management and conservation strategies in semiarid woodlands in Central Argentina.     

Radial growth and characterization of juvenile and adult wood in plantation grown okoumé (Aucoumea klaineana Pierre) from Gabon

The fast-growing hardwood, okoumé (Aucoumea klaineana Pierre), is a major forest species in Gabon and is used principally for making plywood, but research into the growth and quality of this wood is scanty. Trees from natural forests are favoured for production, yet little information exists on wood characteristics from plantation trees. Therefore, we carried out a dendrochronological study along with measurements of wood longitudinal modulus of elasticity (E _L ), density (D _w ), dimensional stability parameters (longitudinal, radial and tangential shrinkage and fibre saturation point) and fibre cell morphology to determine if these properties were related to age in trees from two plantations. We then used segmented regression analysis to define the limit (breakpoint) between juvenile (JW) and adult wood (AW). Using monthly precipitation data, we were able to determine that one growth ring is formed per year, composed of a large light coloured ring formed during the long rainy season and a thick, dark band formed during the major dry season. However, thinner bands, analogous to false rings, may also form during the short dry and short rainy seasons. Ring width decreases from the pith to the bark, and the breakpoint between JW and AW was at 19 years old when trees from both plantations were pooled together. No differences in D _w or radial and tangential shrinkage occurred with cambial age. E _L increased significantly up to the cambial age of 12–14 years, after which the increase with age was only slight and no breakpoint between JW and AW was found. With regard to mean longitudinal shrinkage, AW was found to form after the age of 13 years but fibre cell length was significantly longer after the age of 14.5 and 20 years, depending on the plantation of origin. Therefore, the boundary between JW and AW in plantation grown okoumé occurs between the ages of 13 and 20 years, depending on the characteristic examined.     

Pinus pinaster Ait. tree mortality following wildfire in Spain

Maritime pine (Pinus pinaster Ait.) is the tree species most affected by wildfire in the Iberian Peninsula. Prediction of the probability of fire-injured tree mortality is critical for management of burned areas, evaluation of the ecological and economic impact of wildfire and prescribed fire planning and application. Pine bark beetles (Scolytidae) frequently attack burned maritime pine stands and cause extensive post-fire mortality throughout the Iberian Peninsula. In the present study, maritime pine trees were monitored for three years following 14 wildfires in four ecotypes in Spain (11 fires in Galicia (Galician ecotype - NW Spain), one fire in Portillo (Meseta-Castellana ecotype - Central Spain), one fire in Rodenal (Rodenal ecotype - Central Spain), and one fire in Genalguacil (Sierra Bermeja ecotype - SW Spain)). Data on tree attributes, crown and bole injury, ground fire severity, Ips sp. presence and tree survival were obtained by examining 3085 trees. Logistic regression models for predicting the probability of delayed maritime pine mortality were developed by use of generalized estimated equations (GEE). An ample range of response to fire damage in mortality was evident among the four ecotypes and different models were fitted for each. The most important variables for predicting tree mortality were total crown volume damaged, presence of Ips sp. attack and cambium kill rating. The results highlight the extensive presence of Ips sp. in burned maritime pine forests and its importance in tree mortality process, the ample range of response of P. pinaster, in terms of post-fire mortality, as well as the need to develop site specific mortality models for the different ecotypes of this species following fire.     

Increasing wood production through old age in tall trees

How long forest trees can sustain wood production with increasing age remains an open question, primarily because whole-crown structure and growth cannot be readily measured from the ground or on felled trees. We climbed and directly measured crown structures and growth rates of 43 un-suppressed individuals (site trees) of the two tallest species – Eucalyptus regnans and Sequoia sempervirens – representing a wide range of tree sizes and ages. In both species, ground-level measurements of annual growth, including height, ring width, and basal area increment, exhibited the oft-reported trend of decreasing growth (or no change in growth) with age, yet wood production of the entire main trunk and whole crown both increased with size and age up to and including the largest and oldest trees we measured. The balance between structural metrics of whole-crown respiratory demands (cambium area, inner bark volume, sapwood volume, and heartwood deposition area) and photosynthetic capacity (leaf area and green bark area) was statistically independent of size but not age. After accounting for the effect of size, trees with lower potential respiratory demands grew more than trees with higher potential respiratory demands per unit photosynthetic area. The strongest determinant of tree energy balance was the ratio of aboveground cambium area to leaf area. Among the site trees we examined, over 85% of the variation in annual wood production was explained by variation in size, and the proportion of total aboveground wood production in appendages (branches, limbs, and reiterated trunks) increased linearly with size. With increasing age in both species, the proportion of annual wood production converted to heartwood increased in main trunks and appendages. The oldest tree we measured produced more heartwood in its main trunk over 651 years (351 m³) than contained in any tree we measured <1500 years old. The two tallest tree species achieve similar stature despite divergent growth dynamics and ecologies. At one extreme, E. regnans attains great size quickly but dies relatively young because trees are susceptible to fire and fungi. At the other extreme, S. sempervirens attains great size more slowly but has a long lifespan because trees resist fire and prioritize investment in decay-resistant heartwood. Increasing wood production as trees age is a mechanism underlying the maintenance of biomass accumulation during forest development and the carbon-sink capacity of old-growth forests.     

Impact of invasion of pine wood nematode on the growth of dominant shrub Pleioblastus amarus in Pinus massoniana communities

When pine trees are invaded by pine wilt diseases, the severely infected pine trees will die and fall down, or they will be removed when found to be damaged by the disease. It gives rise to the invasion of other species in these empty niches originally occupied by pine trees, i.e., competing surrounding trees or understory shrubs will invade the empty niches during the following years. As a result, the spatial distribution and pattern of the main tree species in a pine forest will change, and a niche variety in the main population will occur. In the end, the direction of the succession and restoration of the pine forest ecosystem will be affected. In our study, a Pinus massoniana forest with the dominant shrub, Pleioblastus amarus, was invaded by pine wood nematode and was clear cut. Selecting this community as our research object, we studied the effect of the invasion of the pine wood nematode on the growth of the dominant shrub, P. amarus, in this Pinus massoniana forest. Our results show that, after the attacked pine trees were removed, the niche was occupied by Pleioblastus amarus and other shrubs, which benefited the growth of P. amarus to its climax. Growth of P. amarus at the climax stage was greater compared with the unhealthy pine forest and the control group.     

Predicting climate change impacts on native and invasive tree species using radial growth and twenty-first century climate scenarios

The climatic conditions predicted for the twenty-first century may aggravate the extent and impacts of plant invasions, by favouring those invaders more adapted to altered conditions or by hampering the native flora. We aim to predict the fate of native and invasive tree species in the oak forests of Northwest Spain, where the exotic invaders Acacia dealbata and Eucalyptus globulus co-occur with the natives Quercus robur and Quercus pyrenaica and the naturalized Pinus pinaster. We selected adult, dominant trees of each species, collected increment cores, measured the ring width and estimated the basal area increment (BAI, cm² year^?1). Climate/growth models were built by using linear mixed-effect models, where the previous-year BAI and seasonal temperature and precipitation were the fixed factors and the individual the random factor. These models were run to project the fate of studied species in the A2 and B2 CO₂ emission scenarios until 2100. The models explained over 50 % of BAI variance in all species but E. globulus, where growth probably occurs whenever a minimum environmental requirement is met. Warm autumns favoured BAI of both natives, probably due to an extension of leaf lifespan, but hampered A. dealbata and P. pinaster BAI, maybe because of water imbalance and/or the depletion of carbon reserves. The projections yielded a positive BAI trend for both Quercus along the twenty-first century, but negative for the invader A. dealbata and clearly declining for the naturalized P. pinaster. Our results disagree with previous literature pointing at climate change as a driver of invasive species’ success and call for further studies regarding the effect of climate change on co-occurring natives and invaders.     

Does selecting for improved growth affect wood quality of Pinus pinaster in Portugal?

The selection criteria for the first generations in the Portuguese Pinus pinaster improvement program have been the growth rate and form traits. In this work we study the consequences of this selection on wood quality traits. This study assesses genetic and phenotypic correlation between growth, wood density components, lignin content and mechanical traits (radial modulus of elasticity and radial modulus of rupture) of 46 half sib families from a progeny trail located in Leiria, Portugal, originated from seed collected in a clonal seed orchard. A total of 552 seventeen-years-old trees (about half of full rotation age) were sampled at 2 m height. Height measured at 12 years old presented a higher genetic control (h² = 0.34) relatively to DBH, measured at 12 and 17 years old respectively (h² = 0.17 and h² = 0.15). The results of this study also showed that DBH growth is more dependent on latewood components than earlywood components and that higher growth in Mediterranean regions can be due to an increase of the period of latewood formation. Further, we can conclude genetic selection based on growth will not result in a decrease of wood density, will not affect the occurrence of spiral grain, and is possible to obtain an increase in the radial modulus of elasticity. The present study also showed that it is possible to select for increased growth with lower lignin proportion. Results also suggest that selection for growth at 12 years will probably not affect negatively the wood properties at 17 years.     

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.     

Low- to moderate-severity historical fires promoted high tree growth in a boreal Scots pine forest of Norway

Abstract

Fire is the most important ecological factor governing boreal forest stand dynamics. In low- to moderate-severity fires, the post-fire growth of the surviving trees varies according to fire frequency, intensity and site factors. Little is known about the growth responses of Scots pine (Pinus sylvestris L.) following fires in boreal forests. We quantified changes in tree growth in the years following 61 historical forest fires (between 1210 and 1866) in tree-ring series collected from fire-scarred Scots pine trees, snags and stumps in Trillemarka nature reserve in south-central Norway. Basal area increment 10 years pre-, 5 years post-, and 11–20 years post-fire were calculated for 439 fire scars in 225 wood samples. We found a slight temporary growth reduction 5 years post-fire followed by a marked growth increase 11–20 years post-fire. Beyond 20 years post-fire, the long-term tree growth declined steadily up to approximately 120 years. Our results indicate that recurring fires maintained high tree growth in remnant Scots pines, most probably due to a reduction in tree density and thus decreased competition.     

Optimization of seasonality and mother plant nutrition for vegetative propagation of Pinus pinaster Ait

Due to the high economic importance of Pinus pinaster Ait., there is considerable interest in developing, improving and extending the use of its families for mass clonal propagation and in breeding programmes. In the current study, we evaluated shoot growth, rooting ability and mini-cuttings production of P. pinaster in response to nitrogen fertilization and seasons. We compared eight half-sib families of P. pinaster from Asturias and Galicia (Northern Iberian Peninsula), searching for useful parameters and growing conditions to be included in a mass propagation program for clonal family forestry. We fertilized P. pinaster seedling mother plants kept in a greenhouse with three levels of nitrogen: high (HN), medium (MN) and low (LN) to evaluate rooting ability of mini-cuttings. In addition, we evaluated the maximal potential production of rooted mini-cuttings considering nine cycles of propagation over 1?year, also using three levels of nitrogen. The HN treatment significantly influenced the rooting process, with length, area and volume of roots all being positively affected. Spring was the most favourable season for mini-cuttings in the HN treatment. This study provides valuable new information to optimize the clonal propagation protocol for P. pinaster and shows that the mini-cuttings technique has great potential in mass scale cloning, providing high quality sprout production and well-formed new plants.     

Comparison of histological responses and tissue damage expansion between resistant and susceptible Pinus thunbergii infected with pine wood nematode Bursaphelenchus xylophilus

Pine wilt disease caused by the pine wood nematode (PWN), Bursaphelenchus xylophilus, has been epidemic and has had disastrous impacts on pine forests and forest ecosystems in eastern Asia. Many pine species in this area are susceptible to this disease. Pinus thunbergii is particularly susceptible. In Japan, tree breeders have selected surviving trees from severely damaged forests as resistant candidates, and have finally established several resistant varieties of P. thunbergii. However, this breeding procedure requires much time and effort due to the lack of physiological and phenotypical information about resistance. To investigate the resistance mechanisms of selected P. thunbergii, we compared histochemical responses, tissue damage expansion, and PWN distribution in resistant and susceptible clones of P. thunbergii after PWN inoculation. The results suggested that the mechanisms of resistance are as follows: damage expansion in the cortex, cambium, and xylem axial resin canals are retarded in resistant trees soon after inoculation, probably due to the induction of wall protein-based defenses. Suppression of PWN reproduction was particularly caused by inhibition of damage expansion in the cambium. The slow expansion of damage in each tissue provides time for the host to complete the biosynthesis of lignin in the walls of cells that surround the damaged regions. This lignification of cell walls is assumed to effectively inhibit the migration and reproduction of the PWNs. The mechanism of initial damage retardation is presumed to be a key for resistance.     

Mini-cuttings: an effective technique for the propagation of <Emphasis Type="Italic">Pinus pinaster</Emphasis> Ait.

Pinus pinaster Ait. is one of the main forest tree species planted in Spain, Portugal and France. Due to its high economic relevance, there is considerable interest in developing techniques for vegetative breeding aimed at mass propagation. In this study we present a mini-propagation protocol in order to define an efficient method to propagate families or clones of P. pinaster. We carried out three experiments using mini-cuttings of 3–5 cm in length with the aim of evaluating the effects of temperature (4°C vs. 25°C), plant growth regulator (IBA) and shoot age on rooting ability. Percentage of rooted cuttings and morphological root variables were recorded. The percentage of rooted cuttings per treatment ranged from 68 to 97%. Treatment with IBA significantly influenced the rooting process at 25°C but not at 4°C. The number of apexes, length, area and volume of roots were all positively affected by temperature treatment. Shoot age also had a positive effect on rooting capacity of cuttings, with the cuttings from the youngest shoots (70 days after pruning) having higher rooting percentages, ranging from 84.7 to 98.3%. The use of juvenile material, good environmental conditions and IBA all benefited the rooting of clonal material, resulting in high rooting capacity. This study presents an innovative propagation protocol for P. pinaster that can be used as a tool in breeding programs.     

The effects of understory vegetation on P availability in Pinus radiata forest stands: A review

In many second-rotation Pinus radiata forest planta-tions, there has been a steady trend towards wider tree spacing and an increased rate of application of P fertiliser. Under these regimes, the potential for understory growth is expected to in-crease through increased light and greater nutrient resources. Therefore, understory vegetation could become a more signifi-cant component of P cycling in P. radiata forests than under closely-spaced stands. Studies have shown that growth rates and survival of trees is reduced in the presence of understory vegeta-tion due to the competition of understory vegetation with trees. Other studies have suggested that understory vegetation might have beneficial effects on nutrient cycling and conservation within forest stands. This review discusses the significance of understory vegetation in radiata pine forest stands, especially their role in enhancing or reducing P availability to forest trees.     

Root and butt rot caused by Heterobasidion annosum in Atlantic coniferous ecosystems of Spain

In the second half of the 20th century, the forested surface in northern Spain started to increase during a process of reforestation and the replacement of native forest. This reforestation was performed mostly by introducing monocultures of exotic coniferous species. One of the implications of intense forest exploitation and the introduction of new forest species is an increase in disease outbreaks. Because the genus Heterobasidion includes some of the most significant conifer pathogens in the world, surveys were conducted to collect and identify H eterobasidion isolates associated with diverse hosts within coniferous forests and plantations in the Basque Country, northern Spain. A total of 159 stands were surveyed, and 45 isolates were obtained from different trees. Based on sequencing of the internal transcribed spacer region of the ribosomal DNA, all collected isolates were identified as European H eterobasidion annosum s.s. (European P‐type). Heterobasidion annosum was detected in 28.3% of the sampled stands, with the following distribution by host: Chamaecyparis lawsoniana (11.1%), P inus pinaster (2.2%), P . radiata (42.2%), P . nigra (2.2%), P . sylvestris (17.8%), P seudotsuga menziesii (17.8%) and P icea abies (6.7%). The spatial distribution of the population showed a high degree of clustering. This is the first report of H . annosum s.s. causing damage to forest plantations of Chamaecyparis lawsoniana, P inus pinaster, P . radiata, P seudotsuga menziesii and P icea abies in Spain. In the current context of forest pathosystems, the management practice of replacing susceptible forest species with resistant species is recommended.     

Spatiotemporal Availability of Dead Wood in Protected Old-growth Forests: A Case Study from Boreal Forests in Eastern Finland

Spatiotemporal patterns of standing and fallen dead trees were examined in two protected Scots pine ( Pinus sylvestris L.)-Norway spruce ( Picea abies (L.) Karst.) forests in eastern Finland (Pahkavaara and Pönttövaara). In Pahkavaara the volume of standing dead trees was 10 m 3 ha -1 and the volume of fallen logs was 66 m 3 ha -1 , and in Pönttövaara the values were 48 m 3 ha -1 and 107 m 3 ha -1 , respectively. The areas differed with respect to the tree species composition, whereas the proportion of different decay stages was similar. Decay stage and dendrochronological analyses revealed the continuity of dead wood formation. The spatial pattern of standing dead trees was usually towards clustered. The volumes of fallen logs showed a spatial autocorrelation up to distances of 10-20 m. The results also suggest that the current amount and diversity of dead wood is rather high, but that forest succession is likely to lead to a less diverse state in the future.     

Effect of burnt wood removal on the natural regeneration of Pinus halepensis after fire in a pine forest in Tus valley (SE Spain)

To determine the effect of burnt tree removal on post-fire natural regeneration of Pinus halepensis, two 2 500 m² areas were selected six months after the fire in a totally destroyed mature (>70 years) pine forest. In one area, all the trees were cut down and removed 10 months after the fire and, in the other, all the trees were left standing (control). In each area, 20 permanent plots of 20 m² each were randomly placed, and all seedlings emerging within them labeled by individual numbered plastic tags. Emergence, mortality, density and growth (height) of 6649 P. halepensis seedlings were monitored during the first four post-fire years. Seedling emergence was concentrated in the first post-fire autumn–winter period. No positive effect on seedling emergence was detected as a consequence of burnt trunk dragging and subsequent turning over of soil. Wood removal produced an immediate average seedling mortality of 33%, and notably increased seedling mortality during the subsequent summer, probably due to increased exposure of seedlings to sunlight and the possible debilitation of many individuals by mechanical contact during burnt wood removal. A negative correlation of pine seedling mortality with height was detected, which increased significantly on wood removal in the third post-fire year. That is, short seedlings (<10 cm) in treated plots were the most likely to die during this period. In spite of the detrimental effect of wood removal on sapling survival, seedling density four years after fire in the cleared area was large (3.3 seedlings/m²). Wood removal treatment reduced seedling growth: seedling height was significantly higher in the control stand, and differences in seedling height growth rate became particularly noticeable in the fourth post-fire year. The results denote that traditional wood removal practices do not threaten natural post-fire P. halepensis re-establishment if initial seedling density is large enough. However, further studies focused on wood removal effects on the final tree development level and other ecological aspects are necessary to choose adequate post-fire forest management.     

The relationship between terpenes and flammability of leaf litter

Many studies have assumed that plant terpenes favor fire due to their enormous flammability. However, only a few of them, all performed on green leaves, have demonstrated this. In the present work we investigated the question of whether litter terpene content can be used to estimate flammability and temperatures reached during fire. Epiradiator and burn table tests were used to compare flammability of leaf litter of P. pinaster, P. halepensis, P. pinea, C. albidus, C. ladanifer, C. laurifolius and the mixture of litter of P. pinaster with that of the other five species (e.g. P. pinaster + P. halepensis). Tests with burn table showed increasing spread rates and shorter combustion times under higher terpene contents. Flame height was triggered both with higher a terpene content and bed thickness, whereas the percentage of burned biomass was only significantly correlated to bed height. Epiradiator tests indicated that terpene concentration in leaf litter was positively correlated to flame height and negatively correlated to both flame residence time and ignition delay. Flammability was high for P. pinaster, P. halepensis, and hence for P. pinaster + P. halepensis, intermediate for C. albidus, P. pinea and P. pinaster combined with each of these species, and low for C. laurifolius, C. ladanifer and P. pinaster combined with them. Accordingly, their terpene content was high, intermediate and low. We concluded that plants might influence fire intensity, by having stored terpenes in their dead leaves, in addition to having developed traits to survive fire. Thus, a correct management of dead aboveground fuels rich in terpene concentrations, such as those of P. pinaster and P. halepensis, could prove helpful in reducing the hazard of fire.