Seed germination and seedling performance of two Mediterranean tree species,holm oak (shape Quercus ilex L.) and Aleppo pine (shape Pinus halepensis Mill.): a multifactor experimental approach

A two-level multifactor experimental approach was used to compare seed germination and seedling performance of two Mediterranean tree species: the early successional Aleppo pine (Pinus halepensis Mill.) and the late successional holm oak (Quercus ilex L.). In a first experiment germination rate was evaluated under the combined effects of shade, nitrogen availability, and pine or holm oak leaf litter. In a second experiment we tested for the effects of shade, nutrient availability, and litter type on seedling survival, growth and biomass allocation. Holm oak showed higher germination rates under shaded than under unshaded conditions, while Aleppo pine showed no differences between shaded and unshaded conditions. Nitrogen availability and litter type had no significant effect on germination of either species. Both species showed increased RGR, but also higher mortality rates, when grown in an enriched nutrient environment. While Aleppo pine showed no differences in RGR and mortality rate under different shading levels, RGR decreased and mortality increased for holm oak in full light. Increased radiation decreased LAR, SLA and height:diameter ratio, and increased RWR and R/S in both species, although Aleppo pine showed more pronounced changes. Unlike Aleppo pine, holm oak responded to increased nutrient availability by decreasing R/S and increasing LAR. From these results, no seed-seedling conflicts were found in either species, but a trade-off does seem to exist for holm oak between biomass allocation traits deployed in response to increased nutrient availability and radiation. Aleppo pine outperformed holm oak under most environmental conditions tested and showed a wider regeneration niche.     

Spatial pattern of Quercus ilex and Quercus pubescens recruitment in Pinus halepensis dominated woodlands

Abstract. European Mediterranean landscapes have undergone changes in structure in recent years as a result of widespread agricultural land abandonment and cessation of silvicultural regimes. Studies concerning the regeneration dynamics of dominant forest species have become critical to the prediction of future landscape trends in these changing forest stands. Quercus ilex (holm oak) and Q. pubescens (downy oak) are considered to be the terminal point of secondary succession in extensive areas of the Mediterranean region. Recent studies, however, have suggested the existence of recruitment bottlenecks in oak genet populations as a result of current management regimes. In this study, we present evidence of the successful establishment of Q. ilex and Q. pubescens in Pinus halepensis (Aleppo pine) woodlands. We investigate the distribution patterns and spatial relationships among oak recruits and resident pines. Established P. halepensis is randomly distributed throughout the study area. Oak seedlings are positively associated with pine trees, suggesting that P. halepensis individuals provide safe sites for oak genet recruitment. We show that spatial patterns of recruitment are in agreement with the general model of spatial segregation described for other Mediterranean plant communities, with seeder species colonizing large openings after disturbance, followed by a more aggregated recruitment of resprouter species.     

Patterns of seedling recruitment in West-Mediterranean Quercus ilex forest influenced by canopy development

Abstract. Quercus ilex (holm oak) coppice forests belong to the most representative communities in the western part of the Iberian Peninsula. Due to the high sprouting potential of holm oak after coppicing, the dynamics of recruitment of new individuals in existing populations has been largely overlooked. In these forests, the density of seedlings increases in old stands, as a result of the remarkable increase in the recruitment of 1-yr and 2-yr old seedlings, probably due to better environmental conditions for germination. Older seedlings show a tendency towards increasing survival under some degree of canopy closure, since their density appears to be higher just before complete canopy closure takes place. Nevertheless, the reduced growth of seedlings in older stands due to low light levels causes age distributions of holm oak seedlings to change drastically, with saplings older than 15 yr becoming very unusual in these stands. Therefore, a bottleneck of recruitment appears at this sapling stage and prevents vigorous regeneration just before thinning. Present conditions of genet density and management tend to prevent population renewal. Dynamics of the seedling bank reinforce the role of gap formation through small or large scale perturbations in population turnover.     

Fire and late-Holocene expansion of Quercus ilex and Pinus pinaster on Corsica

Abstract. The natural origin of old Quercus ilex (holm oak) forests on the west coast of Corsica is a matter of dispute. This paper discusses the use of pedoanthracology, especially adapted to approach topics in palaeoecology requiring a high spatial precision. It also shows the importance of fire in vegetation change during the Holocene. Pedoanthracology relies on botanical identification and AMS ¹⁴C dating of micro-charcoal found in soil. Three test pits were dug in a forest currently protected within the ‘Man and Biosphere’ program. These pits yielded large quantities of charcoal. There is evidence that Pinus nigra ssp. laricio (laricio pine) and understorey heather species (Ericaceae) played an important role in the vegetation at the beginning of the Subatlantic (ca. 2500 BP). The importance of Pinus pinaster (maritime pine), Quercus ilex and a few mesophilic species increased between 2000 and 1000 BP. Charcoal fragments found in the soil show the role of fire in the vegetation changes recorded. The late expansion of P. pinaster results from a high fire frequency. The hypothesis proposed in the paper suggests that fires destroyed the remaining toxic humus that had accumulated during the millennia which preceded the arrival of Q. ilex to our study sites. The late development of holm oak forest is a combination of the migration of the species and the increase of fire frequency since 2500 BP, most probably resulting from slash-and-burn agriculture.     

Effects of a nutrient pulse supply on nutrient status of the Mediterranean trees Quercus ilex subsp. ballota and Pinus halepensis on different soils and under different competitive pressure

Nutrient availability is a key factor in Mediterranean ecosystems that affects the primary productivity and the community structure. The great variability of its natural availability is now increasing due to frequent fires, pollution events and changes in rainfall regime associated to climate change. Quercus ilex ssp. ballota and Pinus halepensis are the most abundant tree species in the NW Mediterranean basin. They frequently compete in the early and middle successional stages. We investigated the effects of N and P pulse supplies on nutrient uptake capacity in these two species in an after-fire field area and in nursery conditions on different soil types and competing conditions. In the field, N fertilisation had weak effects on nutrient concentration and mineralomass likely as a consequence of this nutrient not being limiting in this field site whereas P fertilisation increased the P mineralomass and the Mg, S, Fe, K and Ca concentrations and mineralomass in the different biomass fractions of both species 1 and 3 years after fertilisation application. In the nursery experiments, P fertilisation increased the mineralomass and concentrations of P, Mg, S, Fe, K and Ca in all biomass fractions including the roots in both species and in different soils and competition conditions. The increment of nutrient mineralomass was due to both the increase of growth and of nutrient concentrations. Both species were able to absorb significant amounts of the P applied by fertilisation (between 5 and 20%) in short time (18 months). Competing vegetation decreased the positive effects of P fertilisation, and in many cases the negative effect of competing vegetation on nutrient mineralomass was stronger when P availability was increased by fertilisation. Q. ilex subsp ballota showed a greater competitive ability for P than the more pioneer species Pinus halepensis in the field but not in the nursery conditions. Pinus halepensis had greater nutrient mineralomass in calcareous than in siliceous soils. Q. ilex subsp. ballota had a higher root biomass allocation and root nutrient allocation than P. halepensis, but both species showed a high capacity to increase their nutrient uptake when its availability increased by fertilisation, thus assuring a great nutrient reserve for future growth periods and contributing to retain nutrients in the ecosystem.     

Tree growth,mortality, and above-ground biomass accumulation in a holm oak forest under a five-year experimental field drought

A holm oak forest was exposed to an experimental drought during 5 years to elucidate the growth responses of the dominant species Quercus ilex, Arbutus unedo and Phillyrea latifolia. Soil water availability was partially reduced, about 15% as predicted for this area for the next decades by GCM and ecophysiological models, by plastic strips intercepting rainfall and by ditch exclusion of water runoff. The stem diameter increment was highly correlated with annual rainfall in all species, and drought treatment strongly reduced the diameter increment of Q. ilex (41%) and specially of A. unedo (63%), the species showing higher growth rates. Stem mortality rates were highly correlated with previous stem density, but drought treatment increased mortality rates in all species. Q. ilex showed the highest mortality rates (9% and 18% in control and drought plots, respectively), and P. latifolia experienced the lowest mortality rates (1% and 3% in control and drought plots, respectively). Drought strongly reduced the increment of live aboveground biomass during these 5 years (83%). A. unedo and Q. ilex experienced a high reduction in biomass increment by drought, whereas P. latifolia biomass increment was insensitive to drought. The different sensitivity to drought of the dominant species of the holm oak forest may be very important determining their future development and distribution in a drier environment as expected in Mediterranean areas for the next decades. These drier conditions could thus have strong effects on structure (species composition) and functioning (carbon uptake and biomass accumulation) of these Mediterranean forests.     

Plastic bimodal growth in a Mediterranean mixed-forest of Quercus ilex and Pinus halepensis

Mediterranean tree species have evolved to face seasonal water shortages, but may fail to cope with future increases in drought frequency and intensity. We investigated stem radial increment dynamics in two typical Mediterranean tree species, Aleppo pine (Pinus halepensis), a drought-avoiding species, and holm oak (Quercus ilex), a drought-tolerant species, in a mixed forest and on contrasting slope aspects (south- and north-facing). Intra- and inter-annual growth patterns were modelled using the VS-Lite2 model for each tree species and slope-aspect. Both species showed a bimodal growth pattern, with peaks coinciding with favourable conditions in spring and autumn. A bimodal growth pattern is always observed in P. halepensis, while in Q. ilex is facultative, which suggests different strategies adopted by these species to cope with summer drought. More specifically, trees on south-facing slope showed a more evident bimodal pattern and more intra-annual density fluctuations. In recent decades, the intensity of both growth peaks has diminished and drifted away due to the increased summer drought. The VS-Lite2 model reveals a niche partitioning between both species. Differences in growing season’s length and timings of growth peaks in both species are relevant for their coexistence and should be considered for estimating mixed-forest responses under climate change scenarios.     

Constraints and trade-offs in Mediterranean plant communities: The case of holm oak-Aleppo pine forests

In this paper we review those aspects that are relevant to the development of a mechanistic ecological theory to account for the structure and dynamics of Mediterranean forests, focusing our attention on mixed forests of holm oak (Quercus ilex L.), a shade-tolerant, slowgrowing species that resprouts vigorously after disturbance, and Aleppo pine (Pinus halepensis M.), a fast-growing, nonresprouting, shade-intolerant species. The main objectives of this report are: to introduce some of the primary features of these forests, showing their structural complexity and historical peculiarities; to show that much of this complexity can be conceptually reduced to two main factors of variation, soil-moisture gradients and a complex interaction of historical management and disturbance regimes; and to contrast the unique features of Mediterranean systems with other communities that have inspired generalization in ecology. Plants in Mediterranean-climate regions must face several environmental constraints during their life cycle: water limitation, competition for light, and a complex set of disturbance regimes, mainly fire, herbivory, and human exploitation. The response of co-occurring species to a given set of environmental constraints depends on a combination of physiological and morphological traits. In holm oak-Aleppo pine forests, the lower limit of distribution along a soil-moisture gradient appears to be controlled by dry-season water stress on seedling performance, and the upper limit seems to be controlled by shade tolerance relative to competitors. The processes that generate and maintain these patterns are related to the responses of the two species to the water and light environments that result from interacting gradients of disturbance and resource availability. The dynamics of mixed holm oak-Aleppo pine forests may be represented along two major environmental axes: water availability and light intensity; namely, time since last disturbance. At the regional scale, the presence of holm oak and Aleppo pine is expected to be driven mainly by the precipitation regime, with the proportion of Aleppo pine increasing toward the driest border and with holm oak being the dominant species in areas with higher precipitation. Changes of dominance of holm oak and Aleppo pine also respond to water availability at the local scale. In this case, variations between species depend on different factors in a complex way, because reduced soil-moisture levels may result either from low precipitation or from topography and edaphic features. The dynamics of holm oak-Aleppo pine forests are also determined by temporal changes in canopy closure; that is, forest recovery after disturbance. In this case, the proportion of Aleppo pine would increase in recently disturbed stands (i.e., with high light intensity reaching the forest floor), whereas regeneration of holm oak would be dominant under partially closed canopies. Theories of forest dynamics developed in humid regions may apply only poorly to Mediterranean plant communities, where vegetation change is qualitatively or quantitatively different. Thus, succession in temperate forests appears to be driven by differences in light availability and shade tolerance; but in Mediterranean plant communities, water limitation is of greater importance for the distribution of forest species. In Mediterranean landscapes the interaction of life-history strategies with changing environments is difficult to infer from observational and experimental studies. A mechanistic approach, in which competition or plant performance is measured as a function of resource availability, seems more feasible. The idea should be to develop multispecies models calibrated specifically for Mediterranean forests in a combined program of modeling, field research, and experimentation.

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Resumen  En este trabajo se revisan los aspectos más relevantes en el desarrollo de una teoría ecológica mecanicista sobre la estructura y dinámica de los bosques mediterráneos, en concreto sobre los bosques mixtos de encina (Quercus ilex L.), una especie tolerante a la sombra, de crecimiento lento, y que rebrota vigorosamente después de una perturbación, y pino carrasco (Pinus halepensis M.), una especie de crecimiento rápido, no rebrotadora, e intolerante a la sombra. Los principales objetivos de este estudio son: introducir algunas de las principales características de estos bosques, mostrando su complejidad estructural y sus peculiaridades históricas; mostrar que gran parte de esta complejidad puede reducirse conceptualmente a dos principales factores de variación, la disponibilidad de agua en el suelo y una compleja interaction de regímenes de gestión y perturbación; y contrastar las características únicas de los sistemas mediterráneos con las de otros sistemas en los que se han basados gran parte de las generalizaciones que se han hecho en ecologia. En las regiones de clima mediterráneo, las plantas deben enfrentarse a diferentes restricciones: limitación hídrica, competencia por la luz, y un conjunto complejo de regímenes de perturbación, principalmente fuego, herbivoría, y explotación humana. La respuesta de las especies a un conjunto de limitaciones ambientales depende de la combinación de caracteres fisiológicos y morfológicos. En los bosques de pino carrasco y encina, el límite inferior de distribución de las especies viene controlado por el estrés hídrico de la estación, mientras que el límite superior viene controlado por la tolerancia a la sombra. Los procesos que generan y mantienen estos patrones están relacionados con las respuestas de ambas especies a lo largo de su ciclo de vida a los ambientes de luz y agua que se generan como resultado de la interacción de los gradientes de perturbación y disponibilidad de recursos. La dinámica de los bosques mixtos de pino carrasco y encina se puede representar a lo largo de dos principales ejes ambientales: disponibilidad de agua e intensidad de luz; o, lo que es lo mismo, tiempo desde la última perturbación. A una escala regional, la presencia de pino carrasco y encina viene determinada por el régimen de precipitación: el pino carrasco aumenta hacia el extremo más seco, mientras que la encina se hace dominante en áreas con mayor precipitación. A escala local, los cambios en la dominancia de ambas especies también responden a la disponibilidad de agua. En este caso las variaciones entre especies dependen de diferentes factores, ya que los niveles bajos de humedad del suelo pueden ser debidos a la baja precipitación, pero también a la topografía o a las características edáficas. La dinámica de los bosques de pino y encina también depende de los cambios temporales en el cierre de la cubierta, es decir, de la recuperación del bosque después de la perturbación. En este caso, la proporción de pino carrasco se incrementa en los rodales recién perturbados (en los cuales llega mucha luz al suelo del bosque), mientras que la regeneración de encina domina bajo cubiertas parcialmente cerradas. Las teorías de dinámica forestal desarrolladas en las regiones húmedas se aplican con dificultad en las comunidades de plantas mediterráneas, donde los cambios de la vegetación suelen ser cualitativamente y cuantitativamente distintos. Así, en los bosques templados la sucesión viene determinada por diferencias en la disponibilidad de luz y la tolerancia a la sombra de las diferentes especies, mientras que en las comunidades mediterráneas la limitación hídrica es más importante a la hora de explicar la distribución de las especies forestales. En los ecosistemas mediterráneos, la interacción de las estrategias del ciclo vital con los cambios en el ambiente dificilmente pueden deducirse a partir de estudios observacionales y experimentales. Se requiere una aproximación mecanicista, en la cual la competencia o el éxito de las plantas se pueda medir en función de la disponibilidad de recursos. La idea sería desarrollar modelos multiespecíficos calibrados específicamente para los bosques mediterráneos, en los que se combinara la modelización con los estudios de campo y la experimentación.

     

Establishment of co‐existing Mediterranean tree species under a varying soil moisture regime

We investigated the responses of two co‐existing Mediterranean trees with different regeneration strategies (Phillyrea latifolia seedlings and Quercus ilex sprouts) to experimental drought below the forest canopy. We considered different recruitment stages and used leaf isotopic discrimination to estimate water use efficiency (WUE) and nitrogen availability and use. Drought decreased the emergence and survival of seedlings and sprouts. Survival and growth of older saplings were not influenced by drought. Seedling emergence of P. latifolia was higher than Q. ilex sprout production, but Q. ilex sprouts had higher survival and growth rates. These differences disappeared in the sapling stage. Carbon isotopic discrimination suggested that Q. ilex sprouts had higher WUE than P. latifolia seedlings. Drought increased WUE of recruits, particularly in Q. ilex. Water use regulation increased with ageing, particularly in P. latifolia. Q. ilex had higher δ¹⁵N values than P. latifolia; these were also higher under drier soil conditions. Current year seedlings had higher δ¹⁵N than saplings, particularly in P. latifolia, suggesting they exploit superficial soil layers. These results suggest that sprouts obtain benefit from resources stored in parent plants. At earlier stages, they perform better than seedlings. This response is not coupled to adult vulnerability to drought for these species, revealing the difficulty of predicting species dynamics during climate change.     

Contrasting foliar responses to drought in Quercus ilex and Phillyrea latifolia

Leaf morphology, longevity, and demography were examined in Quercus ilex and Phillyrea latifolia growing in a holm oak forest in Prades mountains (northeast Spain). Four plots (10 × 15 m) of this forest were submitted to an experimental drought during three years (soil moisture was reduced about 15 %). Leaf area, thickness and leaf mass per area ratio (LMA) were measured in sun and shade leaves of both species. Leaf longevity, the mean number of current-year shoots produced per previous-year shoot (Sn/Sn-1), the mean number of current-year leaves per previous-year shoot (Ln/Sn-1), and the percentage of previous-year shoots that developed new ones were measured once a year, just after leaf flushing. LMA and leaf thickness increased since leaf unfolding except in summer periods, when stomatal closure imposed low photosynthetic rates and leaves consumed their reserves. LMA, leaf area, and leaf thickness were higher in Q. ilex than in P. latifolia, but leaf density was higher in the latter species. Drought reduced the leaf thickness and the LMA of both species ca. 2.5 %. Drought also increased leaf shedding up to ca. 20 % in Phillyrea latifolia and decreased it up to ca. 20 % in Q. ilex. In the later species, Sn/Sn-1 decreased by 32 %, Ln/Sn-1 by 41 %, percentage of shoots developed new ones by 26 %, and leaf area by 17 %. Thus the decrease of leaf number and area was stronger in the less drought-resistant Q. ilex, which, under increasingly drier conditions, might lose its current competitive advantage in these Mediterranean holm oak forests.     

Large-Scale Patterns of Quercus ilex, Quercus suber, and Quercus pyrenaica Regeneration in Central-Western Spain

In Central-Western Spain, forests and woodlands composed of Quercus sp. support outstanding levels of biodiversity, but there is increasing concern about their long-term persistence due to a lack of regeneration. We hypothesize that this regenerative lack is operating on a large geographic scale; that there are differences in the abundance of regeneration between three oak species; that oak regeneration is governed mainly by forest management and structure; and that shrubs act as important physical protectors of seedlings and saplings. We analyzed whether densities of oak seedlings and saplings in several size classes were related to stand-structure, understory, and physiographic variables potentially affecting regeneration. Data collected at a regional level (1 km × 1 km grid) by the Spanish Forest Inventory were evaluated from 2,816 plots. Results revealed that regeneration failure was common for all size categories, from small seedlings to large saplings, and for the three oak species studied, especially the evergreens. Of the Quercus ilex, Q. suber, and Q. pyrenaica plots studied, 49%, 62%, and 20% were lacking any small seedlings, and 82%, 96%, and 56% did not have any large saplings, respectively. Regeneration was positively correlated with tree cover and density, especially of small and medium-sized trees, and negatively correlated with the presence of large trees, indicating that regeneration failure is mostly associated with more open, uniform, and/or aged woodlands. Regeneration densities of Q. ilex and Q. suber were positively correlated with all understory variables, suggesting that the presence of pioneer shrubs represent a major safe site for early tree recruitment, independent from specific shrub species.     

Early post-fire regeneration of a Pinus halepensis forest on Mount Párnis,Greece

Abstract. The post-fire regeneration of a 45-yr-old Pinus halepensis (Aleppo pine) forest, burned in July 1989, has been studied on Mount Párnis, Attiki, Greece. Four experimental plots at various slopes and exposures were established at altitudes of 400 - 450 m, and monitored for 3 yr at 3-month intervals. Early regeneration took place abundantly, through both resprouting and seed germination of mostly hard-seeded herbs and shrubs; the floristic richness was high with 80 taxa. Pine seedling emergence took place during the winter of the first post-fire year. The mean pine seedling density by the end of the recruitment period (March 1990) was 5–6 seedlings/m². This density decreased slightly during late spring and considerably during summer. During the second post-fire year only a relatively slight decline was observed; thereafter the density was stabilized to 1 - 2 seedlings/m². Mortality follows a negative exponential curve that levels off at ca. 20 %. Height distributions throughout the three post-fire years were all positively skewed as a result of the presence of few very tall saplings. A considerable fraction (20 %) of very short (5–15 cm) saplings were still alive 39 months after the fire; these may constitute the sapling bank. Based on the analysis of height distribution curves, it is concluded that the taller seedlings survived significantly better than the shorter ones.     

Regulation of the germination of Aleppo pine (Pinus halepensis) by nitrate, ammonium, and gibberellin, and its role in post-fire forest regeneration

Aleppo pine ( Pinus halepensis Mill.) regenerates after wildfires only by germination. This post-fire germination is controlled by temperature, light and the chemical characteristics of the ash-bed. The extremely high pH caused by the ash inhibits germination of Aleppo pine as well as of many other species. Here we report that ash extracts at neutral pH stimulate P. halepensis germination. The effects of ash extracts, nitrate, ammonium (in concentrations present in the ash) and GA₃ on the germination of P. halepensis seeds and on lipase activity at early germination were studied under controlled conditions. Sub-optimal concentrations of nitrate- and ammonium-stimulated germination and the activity of lipase in the seeds. It is proposed that nitrate increases seed sensitivity to GA₃ while ammonium directly affects lipase activity. The balance between the inhibition of germination by the high pH of ash and its stimulation by the nitrate and ammonium present in the ash determines the overall effect of the ash on post-fire germination of Aleppo pine. The pine seedlings appear in high density among the trees but because of the effects of the ash they appear in sparse stands under the projection of the burned pine canopies. These seedlings benefit from weak inter- and intra-specific competition as well as improved mineral nutrition and have good chances to grow into large trees that will form the regenerating forest; therefore, the enhancement of P. halepensis germination by nitrate and ammonium and its inhibition by high pH can be considered as ecophysiological adaptations to post-fire regeneration.     

POST-FIRE REGENERATION OF SOUTHERN CALIFORNIA CHAPARRAL

Succession of chaparral shrubs was studied for the first 4 years after fire on the four major slope faces at three elevations in southern California. Although total cover fluctuated from year to year, shrub cover increased annually through the third year. There was little or no increase in shrub cover between the third and fourth years. Four years after fire the shrubs covered 55% ground surface at the highest elevation site but only 28% at the lowest elevation site. Shrub cover was similar between slope faces with one exception; at the lowest elevation site shrub cover was twice as great on the north and east-facing slopes as on the south and west-facing slopes. Post-fire recovery of shrubs was by seedlings and/or resprouts. Yucca whipplei was an exception in that it did not resprout once the aboveground parts were killed nor did seedlings establish after fire, however the aboveground parts of many Y. whipplei survived the fire. Species which reproduced entirely by seed did so in the first post-fire year from soil-stored seed with the exception of Ceanothus greggii at the highest elevation site. This species was entirely absent the first year after fire but abundant in the second year. Species producing both resprouts and seedlings varied from site to site in the proportion of resprouts:seedlings. Between 83–100% of the post-fire populations of Cercocarpus betuloides, Arctostaphylos glandulosa, and Xylococcus bicolor were resprouts whereas 12–13% of Ceanothus tomentosus were resprouts. For Adenostoma fasciculatum, resprouts constituted 27–54% of the population at the lowest elevation and 65–94% at the highest elevation; whereas, the Quercus dumosa population was 100% resprouts at the lowest elevation and 31–67% resprouts at the highest elevation. Data are presented on the height of resprouts and seedlings for all species at the end of the first post-fire year.     

Alterations of the resin canal system of <Emphasis Type="Italic">Pinus pinaster</Emphasis> seedlings after fertilization of a healthy and of a <Emphasis Type="Italic">Hylobius abietis</Emphasis> attacked stand

Changes in resource availability and biotic and abiotic stress may alter the defensive mechanisms of pine trees. The effect of fertilisation on the resin canal structure of Pinus pinaster seedlings established in two trials in NW Spain, one attacked by Hylobius abietis and the other non-attacked, was studied. The leaders of 50 plants were destructively sampled and the resin canal density, the canal area and its relative conductive area in the phloem and xylem were assessed. Experimentally increased nutrient availability significantly decreased resin canal density in the phloem of the seedlings in the two analysed trials, where unfertilised seedlings presented up to 30% more resin canal density than the fertilised seedlings (mean value ± SEM = 0.32 ± 0.02 resin canals mm⁻² in the fertilised plants versus 0.45 ± 0.04 resin canals mm⁻² in the control plants). Fertilisation had no effect on the resin canal system in the xylem, but significantly increased tracheid size. Significant differences of resin canals among sites were observed mainly in the xylem; the resin canal density was 1.7-fold greater in the attacked site than in the non-attacked site. The similar structure of phloem resin canals in both sites supports that phloem resin canals are constitutive mechanisms of defence in P. pinaster, whereas xylem resin canals would be constitutive mechanisms but also inducible mechanisms of resistance following the attack of pine weevils or bark beetles. XM and LS equally contributed to this paper.     

Vegetation trajectories of Korean red pine (Pinus densiflora Sieb. et Zucc.) forests at Mt. Seorak, Korea

The vegetation dynamics of Korean red pine (Pinus densiflora) forests were investigated at Mt. Seorak, Korea. Our Detrended Correspondence Analysis (DCA) classified the forests into four types: ridge top, upper slope, lower slope/hill, and streamside. The ridge top forests were likely to sustain themselves, as suggested by the large proportion of seedlings and saplings (89% at <25 years old) and the relatively high density ofP. densiflora (2388 stems ha^-1). Periodic disturbances, such as flash floods, made the streamside inhospitable to late-successional species. Such conditions may have provided a favorable environment for the recruitment ofP. densiflora seedlings through increased solar radiation and decreased competition with other species. On the upper slopes, the dominance ofQuercus seedlings and saplings (63% at >25 years old, and a density of 3263 stems ha^-1) suggests a transition from pine to oak forest. Extensive human interventions appeared to arrest the natural succession from pine to oak forests on the lower slope/hill, while encouraging invasions by forest-edge and introduced species (e.g.,Rosa multiflora andRobinia pseudoacacia).     

Comparison of the initial demographies of pine and oak populations in rural pine forests in Korea and Japan

In order to understand the human impact upon demographic change in plant population in rural forests, we examined the population trends ofPinus densiflora andQuercus serrata inP. densiflora forests viewing their seedlings and saplings in rural Korea and Japan. The most prominent factor affecting the regeneration of the pine and oak was the intensity of management activity which controlled the vegetation stratification and its light environment. Open spaces, such as graveyard or cleaned area that allow the long-term direct daylight in dry season to accelerate the surface heat of soil, were unfavorable habitats for germination and growth of both species. The negative effects of the presence of litter and evergreen trees were related to the failure of early seedling and sapling growth of pine. Cover of litter is, especially, another factor related to the growth of pine saplings. It is considered that successful germination and sapling growth of pine and oak in early successional stages is determined by vegetation structures and light condition.     

Growth and yield of young Quercus ilex coppice stands in the Tafferte forest (Morocco)

The coppice method is the silvicultural system commonly applied to Quercus ilex (holm oak) forests for the production of firewood and charcoal. Although the holm oak is the species that covers the largest area in Morocco (1400 000 hectares), little is known about its growth and yield. The objective of this study is to quantify the stand growth and yield of young coppice holm oak stands in the Tafferete forest in the eastern part of the Middle Atlas (Morocco).Equations relating dominant height to age, and basal area, volume, and number of trees to dominant height were developed and used to derive young holm oak coppice yield estimates.The growth and yield information developed in this study may have a greater utility in defining cultural measures that should be applied in the early stage of holm-oak coppice stands.     

Common cranesGrus grus and habitat management in holm oak dehesas of Spain

Changes in agricultural policies have favored tree clearing and removal of shrubs to favor intensive farming in the holm oakQuercus ilex dehesas of Iberia. The effect of these changes for bird species wintering in this agricultural landscape has been rarely analyzed. Here the effects are studied of farming changes in traditional holm oak dehesas on population size, social structure and time budget of common cranesGrus grus in eight traditional wintering sites in Spain. The role of acorn availability, the principal winter food of the species, in relation to management is also assessed. The number of cranes in each wintering area was not related to degree of agricultural intensification, nor to the availability of acorns. However, crane flock size decreased, and the relative occurrence of isolated family groups increased in less intensively transformed wintering areas characterized by the predominance of fallow lands. The age composition of crane flocks varied with acorn availability since more juveniles were reported in areas with relatively lower acorn availability. Time budget was also related to acorn availability, since cranes spent relatively more time preening in those wintering areas with higher availability of acorns. Globally evidence suggests the need to maintain the fallow land (posíos) to sustain the wintering populations of cranes in Iberia, since this agro-grazing system maintains most of the juvenile cranes using the western migratory route.     

Fire severity and seedling establishment in Pinus halepensis woodlands, eastern Iberian Peninsula

Given the observed heterogeneity in fire severity produced within wildfires, we asked to what extent this heterogeneity might affect post-fire regeneration. For this purpose, we studied the post-fire dynamics of Pinus halepensis (Aleppo pine) in the eastern Iberian Peninsula. Sampling was stratified on the basis of fire severity. We defined three fire severity classes based on the degree of consumption of the pine canopy. The results suggested that there is no clear relationship between seedling density and fire severity; however, mortality was lower and growth (height, shoot biomass and root biomass) was higher in the high severity class. These results can be explained by soil processes: Sites in the high fire severity class may have sustained higher fire intensities, resulting in higher soil organic matter mineralisation and higher ash deposition, and thus in higher post-fire soil fertility. This higher fertility would produce faster growth in pine seedlings. Independent of the severity class, seedling mortality was higher in quadrats (50 × 50 cm) with higher cover of the perennial grass Brachypodium retusum (Poaceae), suggesting a possible competitive effect. For all plots in all 3 severity classes, spatial analysis suggests an aggregate seedlings pattern, but with independence from the position of the adult (source) trees. This revised version was published online in June 2006 with corrections to the Cover Date.