Non‐fire induced seed release ina weakly serotinous pine: climatic factors,maintenance costs or both?

The advantages of canopy seed retention (serotiny) for plants inhabiting fire‐prone ecosystems are well documented. However, very few species are completely serotinous and non‐fire induced opening of serotinous fruits is commonly observed (weak serotiny). Two non‐mutually exclusive causes are envisaged to contribute to this process: mechanical changes in serotinous fruits mediated by climatic conditions (e.g. drought) or the costs of maintenance for the plant of these long‐lasting structures. However, their relative contribution to the spontaneous opening of serotinous fruits remains elusive as well as the consequences for the build‐up of the canopy seed bank and inter‐individual differences in serotiny. In this study we monitored the dynamics of cone production and cone opening in the weakly serotinous Pinus halepensis for five years (2004–2008), including two severe drought episodes (2005, 2006). Drought decreased the production of conelets, increased the abortion of immature cones, reduced the seed quality in the cohorts of cones produced during these years, and increased the opening of serotinous cones. During the first drought episode, a higher proportion of serotinous cones opened in those pines bearing a larger crop of younger cones. This suggests that not only passive changes induced by drought but also competition among cones for resources (e.g. water) might be involved in this process. The opening of serotinous cones in pines bearing more cones made inter‐individual differences in the size of the canopy cone bank to narrow or even to reverse from 2004 to 2008. These results may help to understand the decrease in serotiny when pines grow and accumulate more cones and the large inter‐individual variability in the degree of serotiny observed in P. halepensis forests. In addition, the negative effects of drought episodes for the size of the canopy cone bank and the seeds contained can be an unexplored cause of post‐fire regeneration constraint.     

Effects of repeated burning on Mediterranean communities of the northeastern Iberian Peninsula

Question: In relation to a single fire, do repeated wildfires in Mediterranean type ecosystems (1) reduce plant species richness or diversity; (2) modify patterns of abundance or dominance of plant species or (3) alter plant composition? Location: Pinus halepensis dominated communities of Catalonia, northeastern Iberian Peninsula, western Mediterranean Basin. Methods: Regional, paired design with 14 study sites, each consisting of a once burnt area (1994) and a twice burnt area (1975–1993 and 1994). Ten years after the last fire, we recorded all vascular plant species present in nested plots and quantified their relative abundances on transects. We compared species richness, diversity, dominance and relative abundance and species‐area correlations between paired once and twice burnt areas and assessed their floristic composition similarity. Results: No statistically significant differences were found in species richness or diversity. Slopes of species‐area correlations were higher in once burnt areas. In twice burnt areas, dominance by one or two species was higher. P. halepensis showed lower relative abundance and nanophanerophytes showed higher relative abundance. No differences were found for resprouter, seeder or resprouter‐seeder species. Floristic composition similarity between paired areas tended to be higher in less productive sites. Conclusions: Fire recurrence had contrasting effects on species richness at different spatial scales. Repeated burning reduced the relative abundance of the dominant tree species, which resulted in a higher relative abundance of shrubs. It also promoted the dominance of herbs, particularly Brachypodium retusum. However, it did not change the relative abundance of regenerative groups. Paired areas were more similar as they were more Mediterranean in terms of climatic conditions.     

Large‐scale assessment of regeneration and diversity in Mediterranean planted pine forests along ecological gradients

Aim There is increasing concern regarding sustainable management and restoration of planted forests, particularly in the Mediterranean Basin where pine species have been widely used. The aim of this study was to analyse the environmental and structural characteristics of Mediterranean planted pine forests in relation to natural pine forests. Specifically, we assessed recruitment and woody species richness along climatic, structural and perturbation gradients to aid in developing restoration guidelines. Location Continental Spain. Methods We conducted a multivariate comparison of ecological characteristics in planted and natural stands of main Iberian native pine species (Pinus halepensis, Pinus pinea, Pinus pinaster, Pinus nigra and Pinus sylvestris). We fitted species‐specific statistical models of recruitment and woody species richness and analysed the response of natural and planted stands along ecological gradients. Results Planted pine forests occurred on average on poorer soils and experienced higher anthropic disturbance rates (fire frequency and anthropic mortality) than natural pine forests. Planted pine forests had lower regeneration and diversity levels than natural pine forests, and these differences were more pronounced in mountain pine stands. The largest differences in recruitment – chiefly oak seedling abundance – and species richness between planted and natural stands occurred at low‐medium values of annual precipitation, stand tree density, distance to Quercus forests and fire frequency, whereas differences usually disappeared in the upper part of the gradients. Main conclusions Structural characteristics and patterns of recruitment and species richness differ in pine planted forests compared to natural pine ecosystems in the Mediterranean, especially for mountain pines. However, management options exist that would reduce differences between these forest types, where restoration towards more natural conditions is feasible. To increase recruitment and diversity, vertical and horizontal heterogeneity could be promoted by thinning in high‐density and homogeneous stands, while enrichment planting would be desirable in mesic and medium‐density planted forests.     

Structure and phylogenetic diversity of post-fire ectomycorrhizal communities of maritime pine

Environmental disturbances define the diversity and assemblage of species, affecting the functioning of ecosystems. Fire is a major disturbance of Mediterranean pine forests. Pines are highly dependent on the ectomycorrhizal (EM) fungal symbiosis, which is critical for tree recruitment under primary succession. To determine the effects of time since fire on the structure and recovery of EM fungal communities, we surveyed the young Pinus pinaster regenerate in three sites differing in the elapsed time after the last fire event. Pine roots were collected, and EM fungi characterized by sequencing the internal transcribed spacer (ITS) and the large subunit (LSU) regions of the nuclear ribosomal (nr)-DNA. The effects of the elapsed time after fire on the EM community structure (richness, presence/absence of fungi, phylogenetic diversity) and on soil properties were analysed. Fungal richness decreased with the elapsed time since the fire; although, the phylogenetic diversity of the EM community increased. Soil properties were different depending on the elapsed time after fire and particularly, the organic matter, carbon-to-nitrogen (C/N) ratio, nitrogen and iron significantly correlated with the assemblage of fungal species. Ascomycetes, particularly Tuberaceae and Pezizales, were significantly over-represented on saplings in the burned site. On seedlings, a significant over-representation of Rhizopogonaceae and Atheliaceae was observed in the most recently burned site, while other fungi (i.e. Cortinariaceae) were significantly under-represented. Our results are consistent with the hypothesis that fire can act as a selective agent by printing a phylogenetic signal on the EM fungal communities associated with naturally regenerated pines, pointing out to some groups as potential fire-adapted fungi.     

Forest composition in Mediterranean mountains is projected to shift along the entire elevational gradient under climate change

Aim Species distribution models have been used frequently to assess the effects of climate change on mountain biodiversity. However, the value and accuracy of these assessments have been hampered by the use of low‐resolution data for species distributions and climatic conditions. Herein we assess potential changes in the distribution and community composition of tree species in two mountainous regions of Spain under specific scenarios of climate change using data with a high spatial resolution. We also describe potential changes in species distributions and tree communities along the entire elevational gradient. Location Two mountain ranges in southern Europe: the Central Mountain Range (central west of the Iberian Peninsula), and the Iberian Mountain Range (central east). Methods We modelled current and future distributions of 15 tree species (Eurosiberian, sub‐Mediterranean and Mediterranean species) as functions of climate, lithology and availability of soil water using generalized linear models (logistic regression) and machine learning models (gradient boosting). Using multivariate ordination of a matrix of presence/absence of tree species obtained under two Intergovernmental Panel on Climate Change (IPCC) scenarios (A2 and B2) for two different periods in the future (2041–70 and 2071–2100), we assessed the predicted changes in the composition of tree communities. Results The models predicted an upward migration of communities of Mediterranean trees to higher elevations and an associated decline in communities of temperate or cold‐adapted trees during the 21st century. It was predicted that 80–99% of the area that shows a climate suitable for cold–wet‐optimum Eurosiberian coniferous and broad‐leaved species will be lost. The largest overall changes were predicted for Mediterranean species found currently at low elevations, such as Pinus halepensis, Pinus pinaster, Quercus ilex ssp. ballota and Juniperus oxycedrus, with sharp increases in their range of 350%. Main conclusions It is likely that areas with climatic conditions suitable for cold‐adapted species will decrease significantly under climate warming. Large changes in species ranges and forest communities might occur, not only at high elevations within Mediterranean mountains but also along the entire elevational gradient throughout this region, particularly at low and mid‐elevations. Mediterranean mountains might lose their key role as refugia for cold‐adapted species and thus an important part of their genetic heritage.     

Canopy seed bank structure in relation to: fire,tree size and density

To assess the canopy seed bank structure of Pinus halepensis, we measured the level of serotiny and the seed bank size and density of trees in unburned stands and post-fire regenerated stands in Israel. We analysed the effects of tree size, tree density and fire history on the level of serotiny. The level of serotiny decreased with an increase in tree height. The high level of serotiny in short trees could be explained by selection to increase regeneration chances after burning at pre-mature age. Also, limitation of long-distance seed dispersal opportunities in short trees may favour high serotiny levels. The level of serotiny was higher in post-fire stands than in unburned stands, suggesting a fast selection for serotiny by fire. Unburned stands had a higher total stand seed density than post-fire regenerated stands, but the proportion of seeds in serotinous cones of the total stand seed density was higher in post-fire regenerated stands. The fact that P. halepensis bears simultaneously serotinous and non-serotinous cones reflects its dual strategy as both a post-fire obligate seeder, mainly from serotinous cones and an early coloniser during fire-free periods, mainly from non-serotinous cones. The relative investment in these strategies is dependent on fire history and varies with tree height. Furthermore, mature brown cones can contribute to post-fire regeneration in case of spring fires, and serotinous cones are known to open partially also in dry spell events. Thus, post-fire regeneration and invasion are strategies, which seem to complement each other.     

Genetic diversity and differentiation of two Mediterranean pines (Pinus halepensis Mill. and Pinus pinaster Ait.) along a latitudinal cline using chloroplast microsatellite markers

Several studies have reported glacial refugia and migration pathways for different pine species in the Iberian Peninsula, all of them based on a single‐species approach. In this paper, chloroplast microsatellites (cpSSRs) are used to compare population genetic structure and diversity estimates for interspecific pairs of populations located along a cline from southwestern (latitude 36°32′ N, longitude 5°17′ W) to northeastern Spain (latitude 42°14′ N, longitude 2°47′ E) in two widely distributed Mediterranean pines, Pinus halepensis Mill. and Pinus pinaster Ait. Some cpSSRs were shared between species, facilitating comparison of levels of gene diversity at the species level and inferences about within and among species differentiation. P. pinaster showed a much higher number of variants (29) and haplotypes (69) than P. halepensis (20 and 21, respectively). Moreover, genetic diversity estimates for interspecific pairs of populations along the cline were negatively correlated. Three main causes may explain the differences between species in the present‐day distribution of genetic diversity: (1) the distribution of genetic variability before the Quaternary glaciations, with an earlier presence of P. pinaster in the Iberian Peninsula and a late spread of P. halepensis from eastern and central Europe, (2) the location of the Holocene glacial refugia and the migration pathways from these refugia to the present‐day range (from northeast to southwest in P. halepensis and from southwest to northeast and northwest in P. pinaster) and (3) the interactions between species during the postglacial spread.     

Life histories of Mediterranean pines

The life history of Spanish pines and their relation to fire as the main disturbance factor in their ecosystems was analysed. The primary ecological attributes studied were the canopy seed bank (onset of cone production, percentage and persistence of serotinous cones), seed and cone morphology, sprouting and bark thickness. Four ecological groups were separated using multivariate cluster analysis and their life-history characteristics are discussed. Serotiny and early flowering in Pinus halepensis and P. pinaster reflect their evader strategy in relation to fire as this character is advantageous to survive frequent crown fires and to attain successful post-fire recruitment. Late flowering and absence of serotinous cones in P. nigra, P. sylvestris and P. uncinata indicate that their natural forest did not evolve under frequent crown fires. P. canariensis and P. pinea appeared in two single groups because of their sprouting capability and their seed size respectively. Intraspecific variation in P. pinaster was also analysed using the same criteria and high variability was found in its life history traits. A group of P. pinaster populations showed high levels of serotiny and thin bark as a possible adaptation to frequent stand-replacing crown fires. In contrast, a group of non- or weakly-serotinous populations seems to have evolved under a low-intensity fire regime where the best fitness corresponds to thick-barked individuals capable of surviving ground fires. Intermediate strategies were also evident in this species and were discussed in relation to the effect of different fire regimes caused by the understorey vegetation.     

Breeding performance and longevity of Tomicus destruens on Mediterranean and continental pine species

The pine shoot beetle Tomicus destruens (Wollaston) (Coleoptera: Curculionidae: Scolytinae) is one of the main pests of Mediterranean forests, where it is oligophagous on Mediterranean pines. However, possible global warming may make the insect move to higher latitudes and altitudes, allowing it to attack new pine species. In this respect, the aim of the present article was to assess both the acceptance and performance of T. destruens offered host and non‐host pine species. A no‐choice breeding experiment was set up under laboratory conditions, using logs of three Mediterranean (Pinus pinea L., Pinus pinaster Miller, and Pinus halepensis Aiton) and two continental (Pinus nigra Arnold and Pinus sylvestris L.) pine species. Log debarking at the end of adult emergence assessed parent fecundity, egg, and larval mortality. The quality of callow adults emerging from each tested pine was evaluated on the basis of their longevity on a semiartificial diet. Tomicus destruens colonised all tested pine species, but did not reproduce in Scots pine, taking about 79 days to complete development with no differences among pines tested. The best breeding performance, evaluated as female fecundity and adult production, was observed in P. halepensis, and the lowest in P. pinaster. On average, adults emerging from P. pinea survived longer (83 days) than from other pines, and adult longevity was the lowest in males emerging from P. nigra. Austrian pine, which under natural conditions is usually not a host of T. destruens, allowed insect development and adult production similar to P. pinea and P. pinaster.     

Reproductive traits of Pinus halepensis in the light of fire – a critical review

Fire is known to be a major factor in shaping plants and vegetation worldwide. Many plant traits have been described as adaptations for surviving fire, or regenerating after it. However, many of the traits are also advantageous for overcoming other disturbances. The fact that fire in the Mediterranean Basin has been almost exclusively of anthropogenic origin, and thus is of short duration in an evolutionary time scale, cast doubt on the possibility that fire can act as a selective force in the Mediterranean Basin. Our aim here is to review the ecological advantages of Pinus halepensis traits and their possibility to be selected by fire. The non-self pruning of cones and branches, and the high resin content increase the probability of canopy fires and consequent death of P. halepensis trees. Post-fire regeneration of P. halepensis depends totally upon its canopy-stored seed bank. The seedlings grow quickly and they first reproduce at an early age. Young reproductive trees function first as females with a high percentage of serotinous cones. Thus, young P. halepensistrees allocate many resources to seed production, reducing their `immaturity risk' in a case of an early successive fire. The proportion of serotinous cones is higher in post-fire naturally regenerating stands than in unburned stands, and seeds from serotinous cones germinate better under simulated post-fire conditions. The extremely high pH of the ash-bed under the burned canopies creates the post-fire regeneration niche of P. halepensis exactly under their parent trees. All these traits are advantageous for post-fire regeneration, but could they also be selected during the time scale of anthropogenic fires in the Mediterranean Basin? Pinus halepensis is a relatively short living tree with almost no recruitment under forest canopy. The longest estimated fire-return interval and generation length are about 125 years. The earliest solid evidence for the first hominid-controlled fire in the Mediterranean basin is 780,000 years ago, and thus the estimated number of post-fire generations is 6240. We suggest that such a number of generations is sufficient for the selection and radiation of fire adaptive traits in P. halepensis.

     

Human disturbance and environmental factors as drivers of long‐term post‐fire regeneration patterns in Mediterranean forests

Question: What are the main forces driving natural regeneration in burned mature Mediterranean forests in the medium‐long term and what are the likely successional trajectories of unmanaged vegetation? Location: Valencia Region, eastern Spain. Methods: A wildfire burned 33 000 ha of Pinus halepensis and P. pinaster forest in 1979, and subsequent smaller wildfires took place between 1984 and 1996. The study was designed to sample the range of environmental and disturbance (fire recurrence and land use) conditions. The territory was classified into 17 different geomorphological and fire‐recurrence units. Vegetation cover and floristic composition were measured on a total of 113 plots (1000 m² each) randomly selected within these units. Results: The results show that 23 years after the fire the regenerated vegetation consists of successional shrublands, and that forest ecosystem resilience can be very low. The vegetation presents a strong correlation with most of the environmental variables, but fire (one or two fires), soil type and land use (in that order) are the main drivers of vegetation composition. Quercus coccifera shrublands persist on limestone soils while diverse types of other shrublands (dominated by seeder species) are found on marl soils. Conclusions: The results of this study indicate that disturbance factors strongly coupled to human activities, such as land use and fire, play a critical role in the current state of vegetation. Fire creates vegetation patches in different successional states while land use and soil type define the different types of shrubland in terms of their specific composition.     

Fire-scars and polymodal age-structure provide evidence of fire-events in an Aleppo pine population in southern France

Aleppo pine (Pinus halepensis Mill.) is adapted to fire, the most important disturbance in Mediterranean ecosystems. This species is known to be sensitive to fire and is usually killed when burned, but has a powerful regenerative capacity following burning. Sometimes, however, pines survive within burned areas, suggesting a certain potential for recording fire events in terms of age-structure and eventually scars. In the present study, fire scars on Aleppo pine trunks were used to construct a 150-year fire record in a burned forest in southern France. Four fires were detected with a mean fire return interval of 27 years: minimum 13 years and maximum 45 years. The age structure of the Aleppo pine population was analyzed on the basis of dated fire-scars in order to test the role of fire in creating the polymodal structure of the pine population. Results show the presence of multiple “cohorts” (subpopulations) of Aleppo pine. There appear to have been several pulses of tree recruitment, with the two largest cohorts corresponding to regeneration after the 1919 and 2000 fires. Other fires were recognized based on fire scars but these were not associated with obvious post-fire pine recruitment. Regeneration success is likely to depend on fire severity and on the length of fire-free intervals. Fire history and associated age-structure can thus be reconstructed using Aleppo pine stands.     

Ontogenetic differentiation between Mediterranean and Eurasian pines (sect. Pinus) at the seedling stage

Heteroblastic development in pine seedlings includes extreme morphological changes with still unclear adaptive and evolutionary significance. In particular, Mediterranean and Eurasian pines (section Pinus) living in the Mediterranean basin seem to follow quite distinct developmental trajectories at the seedling stage. Aiming to confirm this ontogenetic differentiation we performed a nursery experiment with seedlings of five Mediterranean pines (Pinus pinaster, P. brutia, P. halepensis, P. pinea and P. canariensis) and three Eurasian pines (P. sylvestris, P. uncinata, and P. nigra subsp. salzmannii), also including P. radiata as an outgroup. After destructive analyses at two harvest times (9 and 32 weeks), we found sharp differentiation between Mediterranean and Eurasian pines in a combination of traits linked to shoot heteroblasty. In particular, Mediterranean pines showed a marked delay in the proportion of adult needles to total needles in the shoot compared to Eurasian species, especially at the second harvest. However, two Mediterranean pines, P. pinaster and P. brutia showed a slightly higher proportion of secondary needles, and a higher rate of budset at a more advanced stage (68 weeks) compared to the other three Mediterranean species. Meaningfully, the outgroup P. radiata was the only species combining a high proportion of adult foliage since the first harvest with a delayed formation of the first terminal bud. We discussed the adaptive significance of these findings at the light of species’ climatic niches and life histories.     

Effects of high temperatures on germination ofPinus halepensis Mill. andP. pinaster Aiton subsp.pinaster seeds in southeast Spain

Seeds ofPinus halepensis andPinus pinaster subsp.pinaster were subjected to four heat treatments (90, 110, 150 and 200 °C) for 1 and 5 min to investigate their tolerances to high temperatures resulting from fires. Another group of seeds of each species were not heated and considered as a control. Information on responses of seeds to high temperatures would help to determine whether or not the species are pyrophytes. Heating for 1 min at all temperatures did not affect the average germination ofPinus halepensis seeds whereas that ofP. pinaster subsp.pinaster seeds was decreased significantly when they were heated to 200 ° C. HoweverP. pinaster seeds were more resistant than that ofP. halepensis when heat treatment was 150 °C for 5 min. Data obtained also indicated that, in general,P. halepensis andP. pinaster subsp.pinaster seed germination is not favoured by a temperature increase than can be reached in soils during a fire. Thus, the species are not pyrophytes but colonizers of burnt sites.     

Increased canopy seed-storage in post-fire pine invaders suggests rapid selection mediated by fire

Exotic species storing seeds in the canopy (serotinous species) can experience a clear advantage in fire-prone communities that lack native taxa with such fire-resistant traits. In addition, selection in the new environment can potentially increase the frequency of fire-adapted characteristics such as serotiny. We studied the potential role of fire favoring the serotinous, non-native conifer Pinus radiata in NW Patagonia. We characterized the degree of serotiny (percentage of serotinous cones) and the size of the canopy seed bank in the unburned plantation and in stands of trees recruited after a fire 30 years ago as a proxy for invasion potential. Fire had a positive effect, increasing serotiny in post-fire P. radiata stands. Post-fire recruited cohorts showed higher serotiny levels and a larger canopy seed bank compared with plantations. Our study suggests that fire-linked traits like serotiny may be subjected to a rapid, fire-driven selection process in fire-adapted species such as P. radiata invading fire-prone ecosystems. Thus, increased serotiny can lead to higher postfire invasion densities, which in turn create a positive feedback loop in invaded areas under recurrent fires.     

Distribution and population genetic structure of the Mediterranean pine shoot beetle Tomicus destruens in the Iberian Peninsula and Southern France

1 The Mediterranean pine shoot beetle Tomicus destruens has long been indistinguishable from its congeneric Tomicus piniperda. Both species attack pines, and can be found in sympatry. The geographical distribution of T. destruens is still unclear in most of the Mediterranean Basin. 2 We aimed to describe the geographical distribution and zones of sympatry of both species in the Iberian Peninsula and France, and to study the molecular phylogeographical pattern of T. destruens. 3 Tomicus spp. adults were sampled in Portugal, Spain and France, and a portion of the mitochondrial genes COI and COII was sequenced for 84 individuals. Sequences were aligned to a data set previously obtained from French localities. 4 Tomicus destruens was found in all populations, except for one locality in Portugal and in the Landes (France). It was in sympatry with T. piniperda in two locations on Pinus pinaster and one location on Pinus radiata. 5 Within‐population genetic diversity was high, but we found a significant pattern of spatial distribution of genetic variation, as well as a significant effect of the host tree. 6 The data suggest the existence of two glacial refugia, from which T. destruens recolonized its current range. One refugium was located in Portugal where the beetle probably evolved on P. pinaster. The corresponding haplotypes show a West–East frequency gradient. The other refugium was probably in the eastern range, where the beetles evolved on Pinus halepensis and P. pinea. The corresponding haplotypes show an East–West frequency gradient.     

Structure and species composition of ectomycorrhizal fungal communities colonizing seedlings and adult trees of Pinus montezumae in Mexican neotropical forests

Mexico is a center of diversity for pines, but few studies have examined the ectomycorrhizal (ECM) fungal communities associated with pines in this country. We investigated the ECM communities associated with Pinus montezumae seedlings and mature trees in neotropical forests of central Mexico and compared their structure and species composition. Root tips were sampled on both planted seedlings and naturally occurring adult trees. A total of 42 ECM operational taxonomic units (OTUs) was found on P.?montezumae. Diversity and similarity indices showed that community structure was similar for both plant growth stages, but phylogenetic diversity and Chao-estimated richness were higher for seedlings. Species composition differed between communities. The dominant OTUs belonged to the families Atheliaceae, Cortinariaceae, and Sebacinaceae, although different taxa appeared to colonize seedlings and adults. Only 12 OTUs were shared between seedlings and adults, which suggests that ECM fungi which colonize seedlings are still not fully incorporated into mycelial networks and that ECM taxa colonizing young individuals of P.?montezumae are likely to come from fungal propagules. Intra-generic diversity could be an insurance mechanism to maintain forest productivity under stressed conditions. This is the first report describing the abundance of Atheliaceae in tree roots in neotropical ecosystems.     

Anatomic basis and insulation of serotinous cones in Pinus halepensis Mill

Pinus halepensis Mill., a widespread, low elevation conifer common in Mediterranean Basin, shows a dual reproductive strategy: post-fire obligate seeder (from serotinous cones) and an early coloniser (from non-serotinous cones). Release of seeds encased in serotinous cones is induced either by fire (pyriscence, serotiny or bradychory) or by drying (xeriscence). Morphological differences in serotinous and non-serotinous cones in natural populations of P. halepensis in Southeastern Italy were analyzed. Relationships between tree size (diameter class) and serotiny were checked by counting and sampling serotinous and non-serotinous cones. The macro and microscopic characteristics that could affect cones’ opening were measured in sampled cones. Protection against high temperatures offered by wood scales was also evaluated by applying different temperatures and time exposures, and following the inner thermal raise. Results showed that non-serotinous cones had bigger resin ducts and more separate scales. Also it was highlighted that ovuliferous scales of serotinous cones were bigger and thicker. These scales had more lamellated (multilayered) sclereid cells, and were significantly thinner with a shorter lumen diameter. Continuous temperature-monitoring heat tests inside cones showed that temperatures close to the cone axis were rather low, so seed germination was not influenced. Results confirm that serotinous cones are more compact, rigid and consistent than non-serotinous cones. These characteristics explain the lower insulation, seed protection and the ease opening of non-serotinous cones as well. In conclusion, opening mechanism of pinecone scales under the effect of fire or dry conditions seem related to anatomic differences and it provides seeds with an efficient protection against heat.     

Spatial patterns of ectomycorrhizal fungal inoculum in arbuscular mycorrhizal barrens communities: implications for controlling invasion by <Emphasis Type="Italic">Pinus virginiana</Emphasis>

Invasion of globally threatened ecosystems dominated by arbuscular mycorrhizal plants, such as the alkaline prairies and serpentine barrens of eastern North America, by species of ectomycorrhizal (ECM) pine (Pinus) seriously threatens the persistence, conservation, and ongoing restoration of these rare plant communities. Using Maryland serpentine barrens and an Ohio alkaline prairie complex as model systems, we tested the hypothesis that the invasiveness of Virginia pine (Pinus virginiana L.) into such communities is regulated by the spatial pattern of ECM fungal inoculum in the soil. ECM colonization of pine seedlings can occur by (1) hyphae growing from the roots of mature ECM pines colonizing nearby seedlings (contagion model), (2) pine seedlings being infected after germinating in open areas where spores are concentrated in feces of animals that have consumed sporocarps (centers of infection model), and (3) colonization from spores that are wind-dispersed across the landscape (background model). To test these models of dispersal of ECM fungal inoculum into these barrens, we used autocorrelation and spatially explicit mapping techniques (semivariance analysis and kriging) to characterize the distribution and abundance of ECM inoculum in soil. Our results strongly suggest that ECM fungi most often disperse into open barrens by contagion, thereby facilitating rapid pine colonization in an advancing front from mature pine forests bordering the barrens. Spatial patterns consistent with the centers of infection model were present but less common. Thus, current management techniques that rely on cutting and fire to reverse pine invasion may be ineffective because they do not kill or disrupt hyphal mats attached to mature roots of neighboring pines. Management alternatives are discussed.     

The effect of fire on microbial biomass: a meta-analysis of field studies

Soil microbes regulate the transfer of carbon (C) from ecosystems to the atmosphere and in doing so influence feedbacks between terrestrial ecosystems and global climate change. Fire is one element of global change that may influence soil microbial communities and, in turn, their contribution to the C dynamics of ecosystems. In order to improve our understanding of how fire influences belowground communities, we conducted a meta-analysis of 42 published microbial responses to fire. We hypothesized that microbial biomass as a whole, and fungal biomass specifically, would be altered following fires. Across all studies, fire reduced microbial abundance by an average of 33.2% and fungal abundance by an average of 47.6%. However, microbial responses to fire differed significantly among biomes and fire types. For example, microbial biomass declined following fires in boreal and temperate forests but not following grasslands fires. In addition, wildfires lead to a greater reduction in microbial biomass than prescribed burns. These differences are likely attributable to differences in fire severity among biomes and fire types. Changes in microbial abundance were significantly correlated with changes in soil CO₂ emissions. Altogether, these results suggest that fires may significantly decrease microbial abundance, with corresponding consequences for soil CO₂ emissions.