Seasonal nutrient retranslocation in reforested <Emphasis Type="Italic">Pinus halepensis</Emphasis> Mill. stands in Southeast Spain

Retranslocation, resorption and relocation of nutrients are important adaptive mechanisms developed by plants to acquire the amount of the nutrients required for growth. They are usual mechanisms in deciduous and conifer trees that occur in Mediterranean regions where drought periods are usual. Soil factors, environmental characteristics and species factors are key drivers of nutrient retranslocation in conifers but is not well understood how soil fertility or intraspecific competition influences the process. We studied retranslocation in Pinus halepensis Mill. stands showing different site quality (differences in climate and intraspecific competition) occurring in Southeast Spain. We monitored reforested mature Aleppo pine forests in stands with differences in site quality, climate and intraspecific competition. Stands were characterised, the content of nutrients of soil and green samples (twigs and pine needles) were recorded, and seasonal nutrient retranslocation was obtained. Site characteristics were related to growth rate and nutrient content of foliage and soil. We evaluated whether the retranslocation of nutrients from older to younger foliage was related to the current-year growth rate and to the nutritional status of the plant as influenced by intraspecific competition. Foliar macronutrient concentrations and the amount of retranslocated macronutrients were seasonal, with differences related to site quality and tree density. As a general trend, nutrient concentrations increased after drought (autumn) and decreased during the growth period (spring). However, some micronutrients (mainly Na and Fe) decreased during both periods. The retranslocation pattern in Aleppo pine reinforced the hypothesis that pine adaptations to drought- and fire-prone habitats are linked to the resilience of these forest types. We developed scientific knowledge to assist decision making in adaptive forest management; e.g. fertilizer recommendations or reforestation programmes.     

Factors determining enzyme activities in soils under <Emphasis Type="Italic">Pinus halepensis</Emphasis> and <Emphasis Type="Italic">Pinus sylvestris</Emphasis> plantations in Spain: a basis for establishing sustainable forest management strategies

Key message

Water availability and soil pH seem to be major constraints for enzyme activities in calcareous soils under Pinus halepensis and acidic soils under Pinus sylvestris plantations respectively. Proposals for improving enzyme activities may include the promotion of broadleaf species to increase soil pH and the modulation of stand density or the implementation of soil preparation techniques to facilitate water infiltration.

Context

Soil enzymes play a key role in nutrient turnover in forest ecosystems, as they are responsible for the transformation of organic matter into available nutrients for plants. Enzyme activities are commonly influenced by temperature, humidity, nutrient availability, pH, and organic matter content.

Aims

To assess the differences between enzyme activities in calcareous soils below Pinus halepensis and acidic soils below Pinus sylvestris plantations in Spain and to trace those differences back to edapho-climatic parameters to answer the questions: Which environmental factors drive enzyme activities in these soils? How can forest management improve them?

Methods

The differences in climatic, soil physical, chemical, and biochemical parameters and the correlations between these parameters and enzyme activities in soils were assessed.

Results

Low pH and high level of phenols in acidic soils under Pinus sylvestris and water deficit in calcareous soils under Pinus halepensis plantations appeared to be the most limiting factors for enzyme activities.

Conclusion

Options such as the promotion of native broadleaf species in the Pinus sylvestris stands and the modulation of Pinus halepensis stand density or the implementation of soil preparation techniques may improve enzyme activities and, therefore, nutrient availability.

     

Allelopathic effects of <Emphasis Type="Italic">Pinus halepensis</Emphasis> and <Emphasis Type="Italic">Quercus coccifera</Emphasis> on the germination of Mediterranean crop seeds

Agroforestry is a leading alternative for food security and forest conservation. A full understanding of positive and negative, i.e. allelopathic, interactions between crops and forest trees is necessary for producing crops and conserving forests especially within the threatened Mediterranean forest ecosystems. The present study explored the allelopathic effects of green and senescent leaf and soil extracts of two agroforestry trees—Pinus halepensis and Quercus coccifera—on the germination of wheat, barley, lentil, chickpea, and fababean as the major grain crops of Jordan. Results revealed that allelopathic effects reduced seed germination of these crops. Germination reduction reached a maximum of 75% in fababean treated with green extracts of Q. coccifera and differed among crops and extract sources, but not between tree species. Comparing between green and senescent leaf and soil extract, regarding their effect on germination percentage, it was noticed that these effects were similar in some crops and were different in others. Germination responses were generally different between cereals and legumes where cereals tend to be less affected by allelopathic influences than legumes, especially fababean. We suggest using cereals such as wheat and barley in agroforestry practices in the Mediterranean region of Jordan.     

Phenotypic plasticity blurs ecotypic divergence in the response of <Emphasis Type="Italic">Quercus coccifera</Emphasis> and <Emphasis Type="Italic">Pinus halepensis</Emphasis> to water stress

The Mediterranean evergreen woody plants Quercus coccifera and Pinus halepensis grow in a range of environments where selection by drought, heat and high irradiance can drive genetic and phenotypic differentiation of populations. However, the role of these stresses in filtering out maladaptive genotypes remains unknown. We hypothesize that this filtering is an important process for woody Mediterranean species due to their low phenotypic plasticity reported in previous studies. We have studied the response of saplings of Q. coccifera and P. halepensis, originating from two contrasting populations (a rock outcrop and a garrigue formation), to water stress. Isozyme characterization of genetic diversity was done to determine whether populations were genetically distinct. Water response analysis was based on water relations, gas exchange, chlorophyll a fluorescence, pigment content, antioxidant status and morphological and structural parameters. Ecotypic differentiation was found for both Q. coccifera and P. halepensis populations, with a higher population isozyme similarity and a higher frequency of dominance of a few genotypes at the rock outcrop in both the species. P. halepensis exhibited small but significant differences between populations for plastic responses to water, with lower phenotypic plasticity in saplings from the rock outcrop. Although it was not found in Q. coccifera, this pattern suggests that ecotypic differentiation rendering stress-tolerant ecotypes involves a decreased plasticity. Phenotypic plasticity was not high but it explained over 75% of the total variability among individual plants. Thus, and although evidence for ecotypic divergence was found in both the species, saplings were plastic enough to blur ecotypic differentiation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Production of <Emphasis Type="Italic">Pinus halepensis</Emphasis> seedlings inoculated with the edible fungus <Emphasis Type="Italic">Lactarius deliciosus</Emphasis> under nursery conditions

Inoculation with edible fungi bestows an added value on mycorrhized plants since production of mushrooms as a secondary crop can be an economically-valuable resource in forestry management. In order to establish a protocol for controlled mycorrhization of Pinus halepensis with the edible fungus Lactarius deliciosus, several factors such as fertilisation (35, 60 or 120 mg total N and 13.5, 27 or 54 mg total P), potting substrate (sphagnum peat or sphagnum peat/vermiculite) and fungal inoculum (mycelial slurry produced in a bioreactor, alginate beads or peat/vermiculite) were assessed. The most effective inoculum was mycelial slurry at a dose of 10 ml/plant. The two potting substrates assessed were compatible with mycorrhiza formation. The greatest number of mycorrhized seedlings was obtained with a moderate level of N (35 mg/plant) or P fertilisation (27 mg/plant). Inoculation did not produce a consistent growth effect on P. halepensis, but seedlings met the quality requirements, making them suitable for planting out.     

Fuel types and crown fire potential in <Emphasis Type="Italic">Pinus halepensis</Emphasis> forests

There is a lack of knowledge to identify and classify forest structures according to the risk of crown fires, especially in Mediterranean regions. In this study, for the first time, we use real information, obtained after a wildfire that burnt under extreme meteorological conditions, to classify forest structures of Pinus halepensis into fuel types as a function of crown fire potential. We identified fourteen forest structures which characterize many forest types in Western Mediterranean areas depending on canopy closure, number of tree layers, percent of each tree layer and overall tree density. By using the pattern of fire types that burnt the most numerous forest structures, we have identified four fire hazard groups of forest structures which are considered different fuel types. The first two had the lowest risk of active crown fires and they differed in the proportion of surface fires and passive crown fires. The third fuel type was the threshold between structures with low and high extreme fire behavior; while the fourth had a high risk of active crown fires. Firefighters and forest managers who are demanding this kind of schema, will test and upgrade this classification of fuel types in function of crown fire potential during future wildfires.     

Growth phenology in <Emphasis Type="Italic">Pinus halepensis</Emphasis> Mill.: apical shoot bud content and shoot elongation

Key message

The chronology of periods of organogenesis and elongation is highlighted in Pinus halepensis.The two first growth units of an annual shoot are preformed inside the bud during the previous year. The following growth units are formed during the spring or summer of the current year.

Context

Analysis of annual shoot length growth phenology is crucial to assess the impact of climate change on tree production. Little is known about the basic growth characteristics and the phenology of pines.

Aims

The present study disentangles the roles of shoot organogenesis vs elongation in the annual growth cycle of the polycyclic Aleppo pine.

Methods

Growth of young Pinus halepensis trees was monitored monthly for 1 year. At each monitoring date, the bud content and meristem dimensions of the main stem shoots apices were analyzed.

Results

The two first growth units of an annual shoot are preformed inside the bud during the previous year. The following growth units are formed during the spring or summer of the current year. The gap between a shoot organogenesis and its elongation may vary from 1 month, for the last growth unit, to half a year, for the first growth units.

Conclusion

Our results underline the importance of taking seasonal environmental conditions from both the previous and the current year into account, in order to study the plasticity of annual shoot growth and its response to climate change and variability.

     

Use of waste materials as nursery growing media for <Emphasis Type="Italic">Pinus halepensis</Emphasis> production

Peat moss is gradually being replaced by other materials as a growing medium in forest nurseries due to economic and ecological constraints. In this study, six different mixtures were tested, mixing peat moss (P) and pine bark (B) with digested sewage sludge (S) activated sewage sludge (A) and paper mill sludge (M), as growing media for Pinus halepensis seedlings; three different waste doses were applied. Seed germination percentage, seedling growth and foliar nutrient content after 1 year in a greenhouse and percentage survival after transplanting were recorded. The influence of base substrate (P or B) on germination percentage changed in different ways according the type of waste. The order of the different applied mixtures by suitability (germination rate and seedling growth) from best to worst was as follows: activated sludge with peat, activated sludge with pine bark, sewage sludge with peat, sewage sludge with pine bark, paper mill sludge with pine bark and finally paper mill sludge with peat.     

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.     

Biomass and nutrient allocation to aboveground components in fertilized <Emphasis Type="Italic">Eucalyptus saligna</Emphasis> and <Emphasis Type="Italic">E. urograndis</Emphasis> plantations

The objective of this study was to evaluate biomass allocation and nutrient pools in aboveground biomass components in 18-month-old plantations of Eucalyptus saligna and E. urophylla × E. grandis (i.e. E. urograndis) in Brazil. The plantations were established by pulp companies in a large area comprising three soil types (Acrisols, Vertisols and Leptosols) in the grassland biome in southern Brazil, and an operational regime of planting and maintenance fertilization was used to ensure full availability of nutrients. Vertisols showed the highest availability of soil nutrients, and the P and Ca concentrations in aboveground biomass were also highest in this type of soil. The nutritional status of both species indicates great consumption of nutrients, particularly of P, K and Ca. At this early age, canopy biomass components still made the largest relative contribution, although debarked stem biomass already accounted for 41.5% and 37.4 of total aboveground biomass in E. saligna, and E. urograndis, respectively. Nutrient concentrations in biomass components were similar across species. For all macronutrients, except Ca and Mg, the concentration gradient followed the order wood < bark < branches < leaves. For all micronutrients, except Cu, the concentration gradient followed the order wood < branches ≈ bark < leaves. At the plantation stage studied, i.e. before canopy closure, the importance of the components as nutrient pools followed the order leaves > branches > wood > bark. The branches hold the majority of Ca reserves in biomass and are a very important pool of Mg, K, P and B. The bark makes a small contribution to total biomass, but stores a similar amount of Ca as leaves, being the second major pool after the branches. Comparison of the nutrients supplied by fertilization and the amounts stored in soil and aboveground indicates that the operational dose should be adjusted to each type of soil after further experimental fertilizer trials, as the supply of N and P appears to be too high, particularly for Vertisols. This is leading to the immobilization of P in biomass components that are not of importance in the biological or biochemical nutrient cycles, thus increasing the risk of larger exports of P during biomass removal.     

Litterfall,decomposition and nutrient release of <Emphasis Type="Italic">Shorea robusta</Emphasis> and <Emphasis Type="Italic">Tectona grandis</Emphasis> in a sub-tropical forest of West Bengal,Eastern India

We studied leaf litter fall, decomposition and nutrient release patterns of Shorea robusta and Tectona grandis by using a litter bag technique to better understand the release pattern of nutrients to soil from leaf litter. Annual litterfall varied from 13.40 ± 2.56 t ha^?1 a^?1 for S. robusta to 11.03 ± 3.72 t ha^?1 a^?1 for T. grandis and the decay constant (k) of decomposed leaf litter was distinctly higher for T. grandis (2.70 ± 0.50 a^?1) compared to S. robusta (2.41 ± 0.30 a^?1). Biomass loss was positively correlated with the initial litter C, WSC, C/N and ash content in S. robusta and N, P and K concentration for T. grandis. Biomass was negatively correlated with lignin and L/N ratio for S. robusta and L, WSC, L/N and C/N ratio for T. grandis (P < 0.01). Nutrient use efficiency (NUE) and nutrient accumulation index (NAI) of S. robusta was higher than for T. grandis. The retranslocation of bioelements from senescent leaves ranked as P > N > K. Annual N, P and K input to soil through litterfall differed significantly between the two species in the following order: N>K>P. S. robusta was superior in terms of K and P return and T. grandis was superior in terms of N return. The two tree species showed a similar patterns of nutrient release (K > P > N) during decomposition of their leaf litter. Nutrients of N, K and P were the primary limiting nutrients returned to soil through litterfall with important roles in soil fertility and forest productivity.     

Effects of <Emphasis Type="Italic">Eucalypt</Emphasis> and <Emphasis Type="Italic">Acacia</Emphasis> plantations on soil water in Sumatra

Indonesia’s pulp and paper industry needs a large area of sustainably grown plantations to support its continued development. Acacia mangium has been the key species underpinning the pulp and paper industries in Sumatra, however increased disease pressure on A. mangium is expected to require large-scale conversion of Acacia plantations to Eucalyptus in the near future. The effect of such a large scale change in plantation species on soil moisture, for both tree production, and catchment hydrology is unknown. In this study we sought to characterize the impacts of plantation species (Acacia or Eucalyptus) and nitrogen management, on soil moisture, soil water depletion and depth to groundwater under stands of Acacia mangium and Eucalyptus pellita over the first 2–3 years after establishment. The study was conducted in experiments at four sites in Sumatra, Indonesia. Soil moisture and soil water depletion were not influenced by plantation species or fertilizer treatment. Soil moisture content and soil water depletion were strongly influenced by shallow groundwater at two of the four sites, however depth to groundwater did not influence stem growth. Results from the field trials cautiously suggest that large scale conversion of Acacia mangium to Eucalypt species in these regions is unlikely to result in increased moisture stress, nor is conversion of plantation species likely to lead to substantial differences in catchment hydrology. This study demonstrated the importance of conducting multi-site studies when investigating biophysical relationships in forest/plantation systems.     

Comparison of postfire mortality in endemic Corsican black pine (<Emphasis Type="Italic">Pinus nigra</Emphasis> ssp. <Emphasis Type="Italic">laricio</Emphasis>) and its direct competitor (<Emphasis Type="Italic">Pinus pinaster</Emphasis>)

• Introduction   

Laricio pine (Pinus nigra J.F. Arn. ssp. laricio (Poiret) Maire var Corsicana Hyl.) is a form of black pine endemic to Corsica, that may now be under threat due to current fire regimes and competition with maritime pine (Pinus pinaster Aiton).     

Recessive inheritance of morphological characteristics of utsukushimatsu,<Emphasis Type="Italic">Pinus densiflora</Emphasis> f.<Emphasis Type="Italic">umbraculifera</Emphasis>

Pinus densiflora f.umbraculifera, commonly known as utsukushimatsu, is a distinctively shaped form of Japanese red pine whose growth is restricted to a forest stand in Shiga Prefecture, Japan. The inheritance mode of morphological characteristics of utsukushimatsu was studied to preserve the genetic resource of this pine. As previously reported, F₁ trees grown from open-pollinated seeds harvested from trees inhabiting the native stand showed two phenotypes: one resembling utsukushimatsu, which produces multiple trunks, and the other resembling normalP. densiflora, which produces one or a few trunks. In the present study, controlled pollination was carried out using F₁ and normalP. densiflora trees. Segregation ratios of the two phenotypes observed in the F₂ population showed that the morphological characteristics of utsukushimatsu are inherited recessively. This suggests that the mutation of one gene or a few closely linked genes controls the morphological characteristics of utsukushimatsu. Since multiple trunk formation of utsukushimatsu might be related to a loss of lateral bud inhibition, it follows that a simple gene mutation breaks apical dominance inP. densiflora.     

Successful <Emphasis Type="Italic">Agrobacterium</Emphasis>-mediated transformation of <Emphasis Type="Italic">Populus tomentosa</Emphasis> with apple <Emphasis Type="Italic">SPDS</Emphasis> gene

The problem of salinized soils has become one of the most serious constraints to agricultural and forest productivity. With the purpose of enhancing salt stress tolerance of Populus tomentosa, we transformed this tree species with spermidine synthase （SPDS） genes derived from an apple by an Agrobacterium-mediated method. Four transgenic clones were confu＇med by PCR and Southern blot analysis. As well, the expression of introduced SPDS genes was analyzed by real-time quantitative PCR.     

Spatial pattern of beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) and oak (<Emphasis Type="Italic">Quercus pubescens</Emphasis> Mill.) seedlings in natural pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) woodlands

Spatial pattern of recruitment is an important factor influencing population dynamics of plant communities. The spatial pattern is determined by seed dispersal and by the spatial variability of germination and initial survival. In the process of forest expansion following farmland abandonment, mid- and late-successional species are often dispersed in pioneer forests by birds. This could result in an aggregated spatial pattern of seeds that could influence the dynamics of these species in mixed pioneer forests. In the sub-Mediterranean area, mid- and late-successional species such as Quercus pubescens (downy oak) and Fagus sylvatica (European beech) are expected to replace pioneer Pinus species. Using a point sampling method we demonstrated that beech and oak seedlings (height <2 m) have a clumped distribution in the understorey of pine. This could result from an aggregated dispersal by jays (dispersal effect) or from preferential recruitment in particular habitats (habitat effect). To test these hypotheses we proposed a statistical analysis of spatial patterns of regeneration of beech and oak. Ground cover variables (i.e. cover by rock outcrops, herbs, box shrubs, mosses or pine) did not differ significantly around seedlings as compared with random sample plots. Likewise, clumped seedlings had growth similar to isolated seedlings, thus refuting the hypothesis of preferential location in the most favourable microsites. Aggregated dispersal seems to be involved in the process of regeneration. Since beech and oak seedlings have contrasting ecological demands, we discuss the implication of this pattern for the replacement dynamics of pine by these species.     

Assessment of crown woody biomass in <Emphasis Type="Italic">Eucalyptus grandis</Emphasis> and <Emphasis Type="Italic">E. globulus</Emphasis> plantations

Young trees were harvested to explore non-destructive methodologies to estimate live branch dry weights in young fast-growing Eucalyptus species under different spacing and fertilizer treatments. Branch growth can vary with silvicultural management such as spacing, fertilizing and thinning, and over relatively short periods in response to environmental conditions. Many published regressions based on standard measurements of height and diameter are site, age and treatment specific. The aim of this study was to improve our capacity to predict woody crown dry weight, based on stem measurements, and to minimize (or eliminate) treatment effects on the resulting model. In young trees, branches are temporary support structures for foliage and are often discarded as the base of the green crown rises. As temporary structures they represent an investment of biomass and nutrient elements, and are subject to selection pressures to maximize the return on investment by the tree. Trees were harvested from existing plantation experiments located in south-eastern Queensland for E. grandis W. Hill ex Maiden (ranging from 0.28 to 15.85 m in height, to 5 years old) and south-western Australia for E. globulus Labill. (0.10–34.4 m in height, to 10.2 years) in order to examine the impact of spacing, nitrogen and phosphorus fertilization on early growth. Relationships to estimate crown woody biomass from non-destructive measurements were developed, and these relationships tended to have different slopes and intercepts for trees with predominantly juvenile foliage and those with intermediate or adult foliage. Dry weight of whole-crown live branch wood (W_branch) was related to heights and/or diameter at breast height (D_BH), but the regressions parameters were different, depending on treatment. The relationships became more generic (i.e. less dependent on treatment effects) between W_branch and stem sectional area at the height of the base of the green crown (SA_CB), consistent with the pipe model theory (R² > 0.91 for the two species for trees with intermediate/adult leaves). However, W_branch was more closely related again to the stem volume above the base of the green crown and treatment effects were not significant (V_Con,gc, R² > 0.93). Branches exit the stem below the green crown, and for E. grandis the best relationship was on stem volume above the lowest live branch (V_Con,llb, R² 0.94). Limited sampling from four other species with similar or contrasting crown characteristics indicated that the relationship could be applied quite generally. Individual E. grandis branch woody dry weight was closely related to the conical volume of the main (first order) branch (V_con,br, R² 0.98₎. The whole crown equivalent, branch woody dry weight plus stem dry weight above the lowest live branch, was also closely related to the stem volume within the woody crown (V_Con,llb, R² 0.97–0.99). While the slope of this relationship was still significantly different between trees with juvenile and intermediate/adult foliage, it had a similar form, suggesting that trees with juvenile foliage allocated a different proportion of their woody biomass within the crown to branches than older trees.     

An inoculation experiment of Japanese <Emphasis Type="Italic">Bursaphelenchus</Emphasis> nematodes on Japanese black and red pine, <Emphasis Type="Italic">Pinus thunbergii</Emphasis> and <Emphasis Type="Italic">P. densiflora</Emphasis>

The pinewood nematode (PWN) Bursaphelenchus xylophilus is an invasive pathogen that was introduced from North America to Asian countries and Portugal and is devastating native pine forests. Some native European and Asian Bursaphelenchus nematodes also have weak to moderate pathogenicity to native pine species. To evaluate the potential risk of native Bursaphelenchus species, we inoculated ten Japanese Bursaphelenchus species into native pine species (the dominant forest species) in Japan, and evaluated their pathogenicity using mortality and tracheal tissue damage as indices. Inoculation was conducted on August 3, 2007, and the symptoms were observed every 2 weeks until February 1, 2008. None of the inoculated trees, excluding the pathogenic PWN inoculated control, showed external disease symptoms; however, four species [a less pathogenic PWN isolate, B. luxuriosae, Bursaphelenchus sp. NK215 (undescribed), and NK224 (undescribed)] caused tracheal tissue damage in inoculated seedlings and showed weak pathogenicity. Therefore, we conclude that there are some potentially pathogenic native species of nematodes distributed in Japan. Interestingly, two of these weakly pathogenic species, B. luxuriosae and NK215, are not associated with Pinaceae trees, suggesting that nematode pathogenicity may be a pre-adaptive character. More experimental studies under different conditions are necessary to accurately evaluate the potential risk of these pathogens.