Effects of experimental warming and drought on biomass accumulation in a Mediterranean shrubland

We studied the effects of experimental warming and drought on the plant biomass of a Mediterranean shrubland. We monitored growth at plant level and biomass accumulation at stand level. The experimentation period stretched over 7 years (1999–2005) and we focused on the two dominant shrub species, Erica multiflora L. and Globularia alypum L. and the tree species Pinus halepensis L. The warming treatment increased shoot elongation in E. multiflora, and the drought treatment reduced shoot elongation in G. alypum. The elongation of P. halepensis remained unaffected under both treatments. The balance between the patterns observed in biomass accumulation for the three studied species in the drought plots (reduction in E. multiflora and P. halepensis and increase in G. alypum) resulted in a trend to reduce 33% the biomass of the drought treatment plots with respect to the untreated plots, which almost doubled their biomass from 1998 to 2005. The results also suggest that under drier conditions larger accumulation of dead biomass may occur at stand level, which combined with higher temperatures, may thus increase fire risk in the Mediterranean area.     

Contrasting growth changes in two dominant species of a Mediterranean shrubland submitted to experimental drought and warming

BACKGROUND AND AIMS: Climate projections predict drier and warmer conditions in the Mediterranean basin in the next decades. The possibility of such climatic changes modifying the growth of two Mediterranean species, Erica multiflora and Globularia alypum, which are common components of Mediterranean shrublands, was assessed. METHODS: A field experiment was performed from March 1999 to March 2002 to prolong the drought period and to increase the night-time temperature in a Mediterranean shrubland, where E. multiflora and G. alypum are the dominant species. Annual growth in stem diameter and length of both species was measured and annual stem biomass production was estimated for 1999, 2000 and 2001. Plant seasonal growth was also assessed. KEY RESULTS: On average, drought treatment reduced soil moisture 22 %, and warming increased temperature by 0.7-1.6 degrees C. Erica multiflora plants in the drought treatment showed a 46 % lower annual stem elongation than controls. The decrease in water availability also reduced by 31 % the annual stem diameter increment and by 43 % the annual stem elongation of G. alypum plants. New shoot growth of G. alypum was also strongly reduced. Allometrically estimated biomass production was decreased by drought in both species. Warming treatment produced contrasting effects on the growth patterns of these species. Warmer conditions increased, on average, the stem basal diameter growth of E. multiflora plants by 35 %, raising also their estimated stem biomass production. On the contrary, plants of G. alypum in the warming treatment showed a 14 % lower annual stem growth in basal diameter and shorter new shoots in spring compared with controls. CONCLUSIONS: The results indicate changes in the annual productivity of these Mediterranean shrubs under near future drier and warmer conditions. They also point to alterations in their competitive abilities, which could lead to changes in the species composition of these ecosystems in the long term.     

Long-term retention of post-fire soil mineral nitrogen pools in Mediterranean shrubland and grassland

Background and Aims

The post-fire mineral N pool is relevant for plant regrowth. Depending on the plant regeneration strategies, this pool can be readily used or lost from the plant–soil system. Here we studied the retention of the post-fire mineral N pool in the system over a period of 12 years in three contrasted Mediterranean plant communities.

Methods

Three types of vegetation (grassland, mixed shrub-grassland and shrubland) were subjected to experimental fires. We then monitored the fate of ¹⁵?N-tracer applied to the mineral N pool in soils and in plants over 12 years.

Results

The plant community with legumes (mixed shrub-grasslands) showed the lowest soil retention of ¹⁵?N-tracer during the first 9 months after fire. Between years 6 and 12 post-fire, a drought promoted plant and litter deposition. Coinciding with this period, ¹⁵?N-recovery in the first 15 cm of the soil increased in all cases, except in mixed shrub-grassland. This lack of increase may be attributable to the input of impoverished ¹⁵?N plant residues and enhanced leaching and denitrification, possibly by N₂-fixing shrubs. After the drought, the deepest soil layer showed large decreases in total N and ¹⁵?N-recovery, which were possibly caused by N mineralization.

Conclusions

Twelve years after the fires, plant communities without N₂-fixing shrubs recycled a significant part of the N derived from the post-fire mineral N and this pool continued to interact in the plant–soil system.     

A comparative study of aboveground biomass of three Mediterranean species in a post-fire succession

The aboveground biomass of three woody species (Cistus albidus, Quercus coccifera and Pinus halepensis) in two early successional stages (3- and 10-year old) of a post-fire Mediterranean ecosystem was investigated. Among these three species, which belong to the successional series of holm oak (Quercus ilex), C. albidus and Q. coccifera are two dominant shrub species in the garrigue ecosystem and P. halepensis is a pioneer tree species widely represented in the Mediterranean area. The results obtained showed that in monospecific stands, C. albidus and Q. coccifera had a high recovery potential. In the 3-year-old stands, the cover of P. halepensis was only 19.8% for a total biomass of 0.75 ± 0.21 t ha^–1, while the plant cover of C. albidus and Q. coccifera was, respectively, 26% and 85.5% and biomass was 4.72 ± 1.09 and 11.5 ± 0.16 t ha^–1. Only 10 years after fire, the plant cover of C. albidus and Q. coccifera was, respectively, 55% and 100% and total biomass 13.2 ± 1.7 and 35.8 ± 4.7 t ha^–1. The greatest increase in biomass was noted for P. halepensis (29.7 t ha^–1). If mean annual biomass increments are considered, it appears that there was a significant decrease with the stand age for the two shrub species although the tree species showed an increase in productivity. These differing patterns in biomass and productivity of shrub and tree species with stand age provide information on biomass accumulation rates of pioneer species in a Mediterranean succession and their importance in the vegetation dynamics.     

Precipitation-dependent flowering of Globularia alypum and Erica multiflora in Mediterranean shrubland under experimental drought and warming, and its inter-annual variability

Background and Aims

Relationships between autumn flowering, precipitation and temperature of plant species of Mediterranean coastal shrublands have been described, but not analysed experimentally. These relationships were analysed for two species of co-occurring, dominant, autumn-flowering shrubs, Globularia alypum and Erica multiflora, over 4 years and in experimentally generated drought and warming conditions. The aim was to improve predictions about the responses and adaptations of flowering of Mediterranean vegetation to climate change.

Methods

Beginning of anthesis and date of maximum flowering intensity (‘peak date’) were monitored over 4 years (2001–2004) on a garrigue land type in the noth-east of the Iberian Peninsula. Two experimental treatments were applied, increased temperature (+0·73°C) and reduced soil moisture (–17%) relative to untreated plots.

Key Results

Flowering of Globularia alypum and Erica multiflora differed greatly between years depending on the precipitation of the previous months and the date of the last substantial rainfall (>10 mm). Globularia alypum flowered once or twice (unimodal or bimodal) as the result of differences in the distribution and magnitude of precipitation in late-spring and summer (when floral buds develop). The drought treatment delayed and decreased flowering of Globularia alypum in 2001 and delayed flowering in 2002. Warming extended the period between the beginning of flowering and the end of the second peak for autumn flowering in 2001 and also increased peak intensity in 2002. Flowering of Erica multiflora was unaffected by either treatment.

Conclusions

Autumn flowering of Globularia alypum and Erica multiflora is more dependent on water availability than on temperature. Considerable inter-annual plasticity in the beginning of anthesis and peak date and on unimodal or bimodal flowering constitutes a ‘safe strategy’ for both species in relation to varying precipitation and temperature. However, severe changes in precipitation in spring and summer may severely affect flowering of Globularia alypum but not Erica multiflora, thus affecting development/structure of the ecosystem if such conditions persist.Key words: Globularia alypum, Erica multiflora, autumn flowering, drought, global warming, Mediterranean     

Warming and drought alter soil phosphatase activity and soil P availability in a Mediterranean shrubland

We conducted a field experiment simulating the warming and drought in a Mediterranean shrubland dominated by Erica multiflora and Globularia alypum with the aim to simulate the next future climate conditions predicted by the IPCC and ecophysiological models. As P is frequently a limiting nutrient in Mediterranean ecosystems, we investigated the drought and warming effects on soil phosphatases activities, soil P contents and availability, litter and leaf P concentration, and the capacity of this community to maintain soil P reserves and retain this nutrient in the ecosystem. Warming treatment increased soil and air temperature (an average of 1°C) and drought treatment decreased soil water content in one of the seasons analysed (28% in autum 2004). Warming increased (68%) the activities of soil acid phosphatases in summer and alkaline phosphatase activity (22%) in spring 2004, and increased P concentrations in E. multiflora. Instead, warming decreased P concentrations in litterfall of this same species, E. multiflora, and soil HCO₃-extractable P_i (Olsen-P_i) in some seasons, decreasing total P soil concentration (37%) after 6 years of treatment. The drought treatment did not change soil phosphatase activities, nor available P_i. The effects of climate change on soil P dynamics in Mediterranean areas will thus be strongly dependent on whether the main variable involved in the local change is warming or drought. If warming is the main change without significant changes in water availability, the increases of biological activity can accelerate plant growth, P capture by plants and increase soil-phosphatase activity, altogether decreasing P contents in soil. If drought is the main change, a reduction in P demands by plants is expected, increasing P stocks in soils.     

Warming and drought alter C and N concentration,allocation and accumulation in a Mediterranean shrubland

We investigated the effects of warming and drought on C and N concentrations, nitrogen use efficiency (NUE), and C and N accumulation in different ecosystem compartments. We conducted a 6-year (1999–2005) field experiment to simulate the climate conditions projected by IPCC models for the coming decades in a Mediterranean shrubland. We studied the two dominant species, Globularia alypum and Erica multiflora, and an N-fixing species, Dorycnium pentaphyllum, also abundant in this shrubland. Warming (1 °C) decreased N leaf concentrations by 25% and increased N stem concentrations by 40% in G. alypum. Although warming changed the available ammonium in soil in some seasons, it did not increase total soil N contents. Drought (19% average reduction in soil moisture) decreased leaf N concentrations in the two dominant shrub species, E. multiflora and G. alypum by 16% and 19%, respectively, and increased stem N concentrations by 56% and 40%, respectively. Neither warming nor drought changed the leaf N concentrations in the N-fixing species D. pentaphyllum, although warming increased stem N concentration by 9%. In G. alypum, the increase of stem N concentrations contributed to the observed increase of N accumulation in stem biomass in drought treatments with respect to control plots (8 kg N ha⁻¹). Neither warming nor drought changed NUE in the period 1999–2005. Warming increased soil organic C relative to drought. The effects of warming and drought on C and N concentrations, on N accumulation and on leaf/stem N distribution were not the result of dilution or concentration effects produced by changes in biomass accumulation. Other factors such as the changes in soil N availability, photosynthetic capacity, and plant internal C and N remobilization must be involved. These changes which differed depending on the species and the plant tissue show that the climate change projected for the coming decades will have significant effects on the C and N cycle and stoichiometry, with probable implications for ecosystem structure and function, such as changes in plant–herbivore relationships, decomposition rates or community species composition.     

Secondary succession after perturbations in a shrubland community

Shrubland responses to experimental burning, cutting and ploughing treatments were studied over 15 years in two shrubland communities dominated by Erica australis. The treatments represent the most frequent forms of anthropogenic disturbances experienced by these communities throughout their history. The response to burning and cutting treatments is similar, and the succession process is characterised as autosuccession. The highest values for herbaceous annuals and perennials were observed in the third and fourth years. Generally, herbaceous species remain present throughout the study period, while woody taxa ones increase their cover values over time. The quantity of herbaceous species present is in inverse proportion to the quantity of woody taxa. The woody species that appear immediately after treatments are sprouting species, namely Erica australis and Arctostaphylos uva-ursi. The response to ploughing is slower, reflecting the recovery mechanism (seedlings). However, after 15 years, there are no significant differences in regeneration between treatments. The first stages of this post-ploughing succession are dominated by annual species until the fourth or fifth years, after which woody species begin to dominate and herbaceous taxa decrease considerably. Woody species with high germination values are Halimium alyssoides and Halimium umbellatum. These shrubland communities have a very high resilience to such perturbations and start regenerating rapidly, reaching the original state in about 9 years. The appearance of the climax arboreal species of the area, Quercus pyrenaica, when it comes from germination, occurs 15 years after the perturbations.     

Effects of seasonal and decadal warming on soil enzymatic activity in a P‐deficient Mediterranean shrubland

Soil enzymes are central in the response of terrestrial ecosystems to climate change, and their study can be crucial for the models’ implementation. We investigated for 1 year the effects of warming and seasonality on the potential activities of five soil extracellular enzymes and their relationships with soil moisture, phosphorus (P) concentration, and other soil parameters in a P‐limited Mediterranean semiarid shrubland. The site was continuously subjected to warming since 1999, and we compared data from this study to analogous data from 2004. Warming uniformly increased all enzymes activities, but only when a sufficient amount of soil water was available. Seasonality unevenly altered enzyme activities, thus affecting enzymatic stoichiometry. P deficiency affected enzymatic stoichiometry, favoring the activities of the phosphatases. The effect of warming was stronger in 2014 than 2004, excluding the hypothesis of acclimation of rhizospheric responses to higher temperatures and suggesting that further increases in extracellular enzymatic activities are to be expected if sufficient water is available. Climatic warming will likely generally stimulate soil enzymatic activities and accelerate nutrient mineralization and similar ecological processes such as the production and degradation of biomass and changes in community composition, but which will be limited by water availability, especially in Mediterranean soils in summer. Winters in such ecosystems will benefit from a general increase in activity and production, but biological activity could even decrease in summer, potentially leading to a negative overall balance of nutrient mineralization. This study suggests that a general increase in activity due to warming could lead to faster mineralization of soil organic matter and water consumption in colder climates, until one of these factors in turn becomes limiting. Such trade‐offs between water and temperature in relation with enzyme activity should be considered in biogeochemical models.     

Seasonal contrasting changes of foliar concentrations of terpenes and other volatile organic compound in four dominant species of a Mediterranean shrubland submitted to a field experimental drought and warming

To test the effect of forecasted drought and warming conditions for the next decades by GCM and ecophysiological models on foliar concentrations of volatile organic compounds (VOCs) and especially of volatile terpenes, we studied four typical Mediterranean woody plants (Pinus halepensis L., Pistacia lentiscus L., Rosmarinus officinalis L. and Globularia alypum L.) under a field experimental drought and warming generated using automatically sliding curtains. Terpenes were detected in the four studied species (R. officinalis L., P halepensis L., Pistacia lentiscus L. and G. alypum L.). In general, maximum concentrations of terpenes were found in the coldest periods and minimum concentrations in the summer. Their concentrations ranged between 0.003 mg g^?1 DM (eugenol) in G. alypum under drought conditions and 37 mg g^?1 DM in R. officinalis under control conditions. Main volatile terpenes found in all studied species except in G. alypum were α‐pinene, camphene, β‐pinene, β‐phellandrene and caryophyllene. In general, VOC leaf concentrations increased when soil moisture increased and decreased when air temperature increased. However, contrasting not consistent responses to the drought and warming treatments were found among species, seasons and years. For example, in P. halepensis, the concentrations decreased in response to drought in winter and instead increased in summer. Contrarily, drought decreased concentrations in summer and increased them in winter in Pistacia lentiscus. In any case, the data on seasonal VOC concentration in Mediterranean woody species provided here will add new knowledge of seasonal variation in essential oil contents of these species. These data might help in the study of flammability of Mediterranean ecosystems and in improving prediction algorithms, inventories and modelling of monoterpene emissions in response to climate change, which mostly do not consider the changes in concentration under drought stress. However, the lack of general and consistent response patterns to increasing drought and warming among species, seasons and years found here makes this task difficult.     

Facing drought in a Mediterranean post-fire community: tissue water relations in species with different life traits

Bulk shoot water potential, the osmotic component and the bulk modulus of elasticity were measured throughout one growing season in four species co-occurring in a post-fire Mediterranean community in southern Italy: Pinus halepensis, Phillyrea latifolia, Cistus salvifolius and Rosmarinus officinalis. A severe drought occurred throughout the measurement period. Large seasonal fluctuations have been observed for both predawn and afternoon water potential in all species. Although minimum values down to –4 MPa have been measured, plant water potential always recovered to less negative values after drought. Daily amplitude of water potential decreased with increasing plant water stress in all species. In Cistus and Rosmarinus less ability for short-term control of plant water status has been assessed. Osmotic potential at full turgor did not display clear seasonal patterns, with no consistent ranking of species by their osmotic values. In most cases, no osmotic adjustment (lowering of osmotic potentials) and no change in tissue elastic properties were observed in response to increasing summer drought and intensity of water stress.     

Early post-fire succession in California chaparral: Changes in diversity, density, cover and biomass

For four consecutive years, following the fires in November 1993, temporal variations in species richness, cover and biomass of component plant groups in early post-fire chaparral succession were monitored on different aspects at the Stunt Ranch Santa Monica Mountains Reserve, southern California. Plant groups were categorized based on growth form, life form, ability to fix nitrogen, geographic origin and regeneration strategies. North-facing slopes exhibited higher species richness, higher species turnover rate over time and faster vegetation recovery in terms of biomass accumulation and return to pre-fire species composition. This was probably due to higher species richness and biomass of nitrogen-fixing species found on north-facing slopes in comparison to south-facing slopes. On both north- and south-facing slopes, annuals had the highest species turnover rate, followed by herbaceous perennials and shrubs. In the first four post-fire years, annual species were the largest floristic group, but herbaceous perennials and shrubs were the major contributors to community biomass. Nitrogen-fixing species and exotics contributed significantly to early post-fire community structure. Although the general trends in post-fire succession are clear in terms of temporal changes in the relative proportions of different plant groups, environmental variation and the nature of plant life histories of component species, especially dominant species, could alter such trends significantly.     

Effects of spatial distance and woody plant cover on beta diversity point to dispersal limitation as a driver of community assembly during postfire succession in a Mediterranean shrubland

Beta diversity, and its components of turnover and nestedness, reflects the processes governing community assembly, such as dispersal limitation or biotic interactions, but it is unclear how they operate at the local scale and how their role changes along postfire succession. Here, we analyzed the patterns of beta diversity and its components in a herbaceous plant community after fire, and in relation to dispersal ability, in Central Spain. We calculated multiple‐site beta diversity (β_SOR) and its components of turnover (β_SIM) and nestedness (β_SNE) of all herbaceous plants, or grouped by dispersal syndrome (autochory, anemochory, and zoochory), during the first 3 years after wildfire. We evaluated the relationship between pairwise beta diversity (β_sor), and its components (β_sim, β_sne), and spatial distance or differences in woody plant cover, a proxy of biotic interactions. We found high multiple‐site beta diversity dominated by the turnover component. Community dissimilarity increased with spatial distance, driven mostly by the turnover component. Species with less dispersal ability (i.e., autochory) showed a stronger spatial pattern of dissimilarity. Biotic interactions with woody plants contributed less to community dissimilarity, which tended to occur through the nestedness component. These results suggest that dispersal limitation prevails over biotic interactions with woody plants as a driver of local community assembly, even for species with high dispersal ability. These results contribute to our understanding of postfire community assembly and vegetation dynamics.     

Shift in community structure in an early‐successional Mediterranean shrubland driven by long‐term experimental warming and drought and natural extreme droughts

Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long‐term (17 years) nocturnal‐warming (+0.6°C) and drought (?40% rainfall) experiments in an early‐successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundances. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g., community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter–spring and/or summer, indicating sensitivity to water limitation in this early‐successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long‐term nocturnal‐warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity.     

Impact of drought and increasing temperatures on soil CO2 emissions in a Mediterranean shrubland (gariga)

In arid and semiarid shrubland ecosystems of the Mediterranean basin, soil moisture is a key factor controlling biogeochemical cycles and the release of CO₂ via soil respiration. This is influenced by increasing temperatures. We manipulated the microclimate in a Mediterranean shrubland to increase the soil and air night-time temperatures and to reduce water input from precipitation. The objective was to analyze the extent to which higher temperatures and a drier climate influence soil CO₂ emissions in the short term and on an annual basis. The microclimate was manipulated in field plots (about 25 m²) by covering the vegetation during the night (Warming treatment) and during rain events (Drought treatment). Soil CO₂ effluxes were monitored in the treatments and compared to a control over a 3-year period. Along with soil respiration measurements, we recorded soil temperature at 5 cm depth by a soil temperature probe. The seasonal pattern of soil CO₂ efflux was characterized by higher rates during the wet vegetative season and lower rates during the dry non-vegetative season (summer). The Warming treatment did not change SR fluxes at any sampling date. The Drought treatment decreased soil CO₂ emissions on only three of 10 occasions during 2004. The variation of soil respiration with temperature and soil water content did not differ significantly among the treatments, but was affected by the season. The annual CO₂ emissions were not significantly affected by the treatments. In the semi-arid Mediterranean shrubland, an increase of soil CO₂ efflux in response to a moderate increase of daily minimum temperature is unlikely, whereas less precipitation can strongly affect the soil processes mainly limited by water availability.     

Drought and warming induced changes in P and K concentration and accumulation in plant biomass and soil in a Mediterranean shrubland

A field experiment involving drought and warming manipulation was conducted over a 6-year period in a Mediterranean shrubland to simulate the climate conditions projected by IPCC models for the coming decades (20% decreased soil moisture and 1°C warming). We investigated P and K concentration and accumulation in the leaves and stems of the dominant species, and in soil. Drought decreased P concentration in Globularia alypum leaves (21%) and in Erica multiflora stems (30%) and decreased K concentration in the leaves of both species (20% and 29%, respectively). The general decrease of P and K concentration in drought plots was due to the reduction of soil water content, soil and root phosphatase activity and photosynthetic capacity that decreased plant uptake capacity. Warming increased P concentration in Erica multiflora leaves (42%), but decreased it in the stems and leaf litter of Erica multiflora and the leaf litter (33%) of Globularia alypum, thereby demonstrating that warming improved the P retranslocation and allocation from stem to leaves. These results correlate with the increase in photosynthetic capacity and growth of these two dominant shrub species in warming plots. Drought and warming had no significant effects on biomass P accumulation in the period 1999–2005, but drought increased K accumulation in aboveground biomass (10 kg ha⁻¹) in Globularia alypum due to the increase in K concentration in stems. The stoichiometric changes produced by the different responses of the nutrients led to changes in the P/K concentration ratio in Erica multiflora leaves, stems and litter, and in Globularia alypum stems and litter. This may have implications for the nutritional value of these plant species and plant–herbivore relationships. The effects of climate change on P and K concentrations and contents in Mediterranean ecosystems will differ depending on whether the main component of change is drought or warming.     

Common environmental factors explain both ectomycorrhizal species diversity and pine regeneration variability in a post-fire Mediterranean forest

Natural seedling regeneration and establishment after stand replacing wildfires is influenced by a series of environmental and biological constraints. In this study, we characterized the diversity and structure of the ectomycorrhizal (ECM) fungal community associated with post-fire naturally regenerated maritime pine saplings, and individuate the environmental factors responsible for fungal species distribution. We also identify the main environmental factors responsible for maritime pine regeneration variability and assessed the relation between saplings performance and ECM fungal diversity indices. Fungal species were identified by direct sequencing of internal transcribed spacer regions. Five years after the disturbance event, a total of 30 taxa colonized the pine saplings. The ECM fungal community was dominated by ruderal species of the genus Rhizopogon (present in almost half of the samples). Almost one third of the identified ECM fungal species belonged to the family Thelephoraceae. Typical k-selected species like Amanita pantherina, Boletus aestivalis, Lactarius chrysorrheus, and Russula densifolia were found on pine saplings collected in proximity of unburnt pine trees, in correspondence with low erosion extents. Pine regeneration varied throughout the study areas and was enhanced at higher elevations, in correspondence with moderate slopes, shallower soils, and a reduced cover of ericaceous shrubs and bare ground. These conditions were found in close proximity to patches of pine trees that survived the disturbance event and were previously characterized by a higher pre-fire pine biomass. Even though no correlations were found between saplings performance and ECM fungal diversity indices, common environmental factors (i.e., ericaceous shrub cover, extent of erosion, slope, and soil depth) were responsible for shaping the ECM fungal distribution and for describing most of the explained regeneration variability.     

Root distribution of a Mediterranean shrubland in Portugal

The distribution of roots of an Erica (Erica scoparia and Erica lusitanica) dominated Mediterranean maquis was studied using three different approaches: root counts on trench walls (down to 120 cm), estimation of the maximum rooting depth using an allometric relationship and estimation of fine root biomass and fine root length using soil cores (down to 100 cm). Roots were classified according to diameter (fine, 1.0 mm; small, 1.1–5.0 mm; medium, 5.1–10.0 mm; coarse, >10.0 mm) and species (Erica sp., Pteridium aquilinum, Rubus ulmifolius and Ulex jussiaei). The depth corresponding to 50% of all roots (D ₅₀) was determined by fitting a new model to the cumulative root distribution. Fine roots represented 96% of root counts. Root counts of Erica represented 59%, Ulex 34%, Rubus 6% and Pteridium 1%. Overall root counts showed a D ₅₀ of 26 cm. D ₅₀ was higher for Ulex (40 cm) and Erica (22 cm), than for Pteridium (9 cm) and Rubus (3 cm). D ₅₀ for fine roots was 27 cm, for small roots 11 cm, for medium roots 6 cm and for coarse roots 4 cm. The estimated average maximum rooting depth of the 28 deepest Erica roots was 222 cm. The deepest Erica root was estimated to reach 329 cm. A total of 82% of roots growing deeper than 125 cm were not reaching more than 175 cm. The overall fine root length density ranged from 4.6 cm/cm³ at 10 cm to 0.8 cm/cm³ at 80 cm. The overall fine root biomass ranged from 7.7 mg/cm³ at 10 cm to 0.6 mg/cm³ at 40 cm. D ₅₀ for root biomass was 12 cm and D ₅₀ for root length was 14 cm. Fine root biomass was estimated as 1.6 kg/m² and the respective root length as 18.7 km/m².