Post-fire seedlings of Nothofagus alpina in Southern Chile show strong dominance of a single ectomycorrhizal fungus and a vertical shift in root architecture

We investigated belowground responses of Nothofagus alpina seedlings to post-fire conditions during natural regeneration after a wildfire in Chile, focusing on mycorrhizal community and root architecture. The complete root systems of 2-year-old N. alpina seedlings were extracted from a post-fire site with natural regeneration and compared to roots of seedlings from undisturbed forest nearby. Mycorrhizal morphotype richness was determined in each seedling. Morphometric parameters of tertiary root structure and dry biomass of whole root systems were determined in 5 cm vertical intervals and in four lateral root classes. With 43.5% of colonized vital mycorrhizal root tips, the Basidiomycete Descolea antarctica was the most abundant fungal symbiont on post-fire seedlings. Tertiary root morphology of these seedlings was distinct from control plants and characterized by a deep-reaching tap root with rather evenly distributed lateral branches whereas seedlings from the undisturbed site had shallower root systems with most lateral roots concentrated in the upper soil layers. Post-fire seedlings had more mycorrhizal rootlets and mycorrhiza-bearing third order lateral roots than control plants which was expressed in a 34% higher total root number but only a 10% higher total root biomass, although both values were not statistically significant. A major part of these fine roots in seedlings from burnt forest was found in deeper soil horizons, compared to the seedlings from undisturbed forest. According to our results, post-fire conditions clearly favour Descolea antarctica as an early ectomycorrhizal colonizer of Nothofagus seedlings at the studied site. As no significant changes in soil chemistry could be observed at the burnt site, the deep-reaching tertiary root architecture of these seedlings may be interpreted as a response to other abiotic factors like reduced moisture in surface soil.     

Long-term post-fire vegetation dynamics in Pinus halepensis forests of Central Greece: A functional group approach

A hierarchical approach for plant functional classification was applied to describe long-term vegetation change in Pinus halepensis burned forests. Plant species were initially grouped according to their growth form and afterwards data on species modes of regeneration, persistence and dispersal, together with some other specific competitive advantages were explored, resulting in the identification of 29 different functional groups, 14 for woody and 15 for herbaceous species. Three types of Pinus halepensis forests were identified, according to the structure of the understorey. For each forest type, a post-fire chronosequence of communities was selected for sampling. Data sampling was performed for at least two consecutive years in each community, so as to reduce the shortcomings of the synchronic approach and to increase the age range of each chronosequence. Even though the vast majority of the functional groups proved to be persistent throughout the post-fire development of vegetation, their species richness and abundance did not remain stable. An increase of annual herb richness and abundance was recorded in the first years after the fire, with the leguminous species forming the dominant functional group. For perennial herbs, the most abundant group was of species with vivid lateral growth, while the group of species with subterranean resource organs included the highest number of species. Finally, as far as the woody species are concerned, the groups that played the most important role in defining vegetation structure were the mono-specific group of the pine, the group of resprouting sclerophyllous tall shrubs and the group of obligate seeder short shrubs (with Cistusspp., among others). A negative relationship between the abundance of woody obligate resprouters and the regeneration of woody obligate seeders was found. The advantage of the proposed functional group approach over classical floristic or structural approaches for the long-term study of communities is discussed, together with the applicability of this approach in studies of vegetation risk assessments due to fire regime alterations.     

Seedbed filter controls post-fire succession

Using black spruce (Picea mariana)-Kalmia angustifolia dominated communities of eastern Canada we tested the hypothesis that habitat filter pre-empts biotic filter as a community structuring force in early post-fire succession leading to Kalmia dominated heath. We considered post-fire seedbed as an abiotic (habitat) filter and post-fire aboveground biomass of dominant plants as a biotic filter. First we surveyed and categorized post-fire seedbed types and tested the seedbed filter by a black spruce seeding experiment in 155 small plots (30×30 cm²) on nine sites burned between 1 and 38 years previously. We then quantified black spruce seedling establishment in a competition removal and nitrogen addition experiment in twenty four 3×3 m² plots on four sites burned 7, 24, 23 and 29 years previously. We also conducted a vegetation survey in 200 plots (1×1 m²) on 20 sites along a 1- to 76-year post-fire chronosequence. Only about 10% post-fire seedbed with little or no residual organic matter was suitable for black spruce regeneration. On the rest of the seedbeds made up of charred humus and lichen substrates Kalmia grew profusely by sprouting from sub-surface fire survived components. Removal of aboveground competition and addition of nitrogen did not help black spruce seedling establishment. Kalmia dominated the chronosequence with 50–90% cover until 60 years after fire. Black spruce cover was insignificant during the first 20 years followed by a slow increase surpassing Kalmia cover around 60 years. Herbaceous cover was initially high followed by a decline coinciding with high Kalmia cover and then increased again with decreased Kalmia cover. Lichen cover followed a similar trend. We interpret this post-fire vegetation assembly as a seedbed controlled phenomenon where seedbed limitation filters out the pre-fire dominant, black spruce, in favour of vegetatively regenerating Kalmia turning coniferous forest into Kalmia heath during the first four decades of secondary succession.     

Post-fire regeneration in a Mediterranean pine forest with historically low fire frequency

Species of Mediterranean vegetation are known to regenerate directly after fire. The phenomenon of autosuccession (direct regeneration) has been found to be often combined with an increase of species richness during the first years after fire due to the high abundance of short-lived herbaceous plants facilitated by plentiful nutrients and light. The high degree of vegetation resilience, which is expressed in terms of autosuccession, has been explained by the selective pressure of fire in historic times. According to existing palaeoecological data, however, the Pinus halepensis forests in the Ricote Mountains (Province of Murcia, SE Spain) did not experience substantial fire impact before the presence of man nor are they especially fire-prone today. Therefore, we studied post-fire regeneration to find out if direct succession is present or if species from pre-fire vegetation are absent during the post-fire regeneration stages. Patterns of succession were deduced from observations made in sample plots on sites of a known regeneration age as well as in adjacent unburnt areas. The results of the vegetation analyses, including a Detrended Correspondence Analysis, indicate that Pinus halepensis forest regeneration after fire resembles autosuccession. As regards the presence of woody species, there is a high percentage similarity on north (83%) and south (70%) facing slopes during the first year after fire vs. reference areas which is due, for example, to direct regeneration of the resprouting Quercus coccifera or seeders like Pinus halepensis or Fumana laevipes. However, if herbaceous species are included in the comparison, the similarity on north-facing sites decreases (to 53%) with the presence of additional species, mainly ruderals like Anagallis arvensis or Reseda phyteuma, and even woody species on the burnt plots. This effect indicates “enhanced autosuccession”, which was not found on south-facing sites where overall species richness was very high irrespective of the impact of fire. Locally we found limited regeneration of some species, for example Pinus halepensis at high altitudes (1000 m), even 22 years after fire. As we assume that historical fires did not play an important role in the area and direct succession is present nevertheless, our results support the theory that autosuccession is not a process restricted to fire-prone areas. Fire has been only one of several selective forces since human settlement that probably led to a set of species pre-adapted against recurrent disturbance.     

Remote sensing approach to detect post-fire vegetation regrowth in Siberian boreal larch forest

Remote sensing with time series data offers considerable potential in the trajectory of post forest fire dynamics beyond the current monitoring of structural attributes that are displayed in the post-fire area. Many studies have addressed this topic by using time series remote sensing indices; however, this approach has sometimes been demonstrated as an unrealistic and biased representation of the post-fire forest patterns due to the saturation issues of vegetation indices. These saturation issues then lead to an underestimation of the forest successional stages and an overestimation of the forest recovery rate. This paper aims to develop a framework for trajectory of the post-fire forest patterns in the Siberian boreal larch forest (Larix sibirica) with the synergistic use of different remote sensing based vegetation-cover indicators derived from the Landsat time series and the WorldView-2 images. A time-series of the forest recovery index (FRI) and fractional vegetation cover (FVC) has been analyzed to estimate the rates of forest regeneration and vegetation recovery across different burn severity levels in the Siberian larch forest. The results showed that the FRI method can be used to observe the regrowth of the larch forest from the tenth year after the fire overlapping with the period of significant increase in the sapling stem volume. The post-fire larch forest canopy can fully recover to the pre-fire condition with respect to the magnitude of the FRI values after 30–47 years where the highest regeneration rate was observed in the moderate burn severity areas followed by the low and high burn severity. On the other hand, the FVC method was positively correlated with burn severity and more sensitive for evaluating the early stages of the forest succession in which the FVC dramatically increases after 5–6 years after the fire. The significant growth of FVC was accentuated by the maximum emergence of the sapling density as well as the rapid growth of herbaceous plants, grasses, shrubs, and shade-intolerant trees immediately after the fire, which could not be evaluated using the FRI. Both time series of the FRI and the FVC are valuable tools for determining the dominant stages of the post-fire larch forest succession in order to understand the relationships between fire disturbance and natural cycles of the boreal larch forest.     

Long-term post-fire dynamics of co-occurring woody species in Pinus brutia forests: the role of regeneration mode

Regeneration mode is one of the key attributes determining population structure and dynamics of plant species. We investigated long-term patterns after fire in the cover of plant species in a 100-year chronosequence of burned Pinus brutia forests in a humid Mediterranean climate region in Turkey. Significant trends were present in the change of cover in major species through post-fire chronosequence, and species with similar trends were clustered in relation to their regeneration modes. Obligate resprouters increased their cover from the early post-fire years to the later stages, while cover of obligate seeders with a soil seed bank increased in the early years, but then decreased through time. Facultative resprouters were at an intermediate position, with an increase in cover until mid-successional stages and then a decrease through time. The cover of the only obligate seeder with a canopy seed bank (P. brutia) followed a linear increasing trend during the succession. When species with the same regeneration mode were grouped, the same trends were observed with more explained variances. A few life-history traits were enough to explain the observed trends. Our study shows that regeneration mode is an explanatory functional grouping system for describing long-term post-fire dynamics of Mediterranean Basin woody species. We suggest that regeneration mode must be a major component of any vegetation or forest stand dynamics model in the Mediterranean Basin. This result has important implications for the management of Mediterranean Basin ecosystems, and can potentially be extrapolated to other Mediterranean-type fire-prone ecosystems.     

黑龙江省大兴安岭林区火烧迹地森林更新及其影响因子

林火干扰是大兴安岭森林更新的影响因子之一,研究火烧迹地森林更新的影响因子(立地条件、火前植被、火干扰特征)对理解生态系统的结构、功能和火后演替轨迹具有重要意义。选取呼中及新林林业局55个代表性火烧样地,利用增强回归树分析法分析了火烧迹地森林更新的影响因素。结果表明:(1)立地条件是影响针、阔叶树更新苗密度的主要因素;海拔对针叶树更新苗密度的影响最大;坡度对阔叶树更新苗密度影响最大;(2)距上次火烧时间对针叶树更新苗比重影响最大,其次是林型;(3)中度林火干扰后森林更新状况好于轻度和重度火烧迹地。根据火烧迹地森林更新调查分析可知:林型影响火后演替模式,火前为针叶树或阔叶树纯林,火后易发生自我更新(火后树种更新组成与火前林型相同),而针阔混交林在火干扰影响下易于发生序列演替(火后初期以早期演替树种更新为主)。     

Can salvage logging affect seed dispersal by birds into burned forests?

The recovery of vegetation in Mediterranean ecosystems after wildfire is mostly a result of direct regeneration, since the same species existing before the fire regenerate on-site by seeding or resprouting. However, the possibility of plant colonization by dispersal of seeds from unburned areas remains poorly studied. We addressed the role of the frugivorous, bird-dependent seed dispersal (seed rain) of fleshy-fruited plants in a burned and managed forest in the second winter after a fire, before on-site fruit production had begun. We also assessed the effect on seed rain of different microhabitats resulting from salvage logging (erosion barriers, standing snags, open areas), as well as the microhabitats of unlogged patches and an unburned control forest, taking account of the importance of perches as seed rain sites. We found considerable seed rain by birds in the burned area. Seeds, mostly from Olive trees Olea europaea and Evergreen pistaches Pistacia lentiscus, belonged to plants fruiting only in surrounding unburned areas. Seed rain was heterogeneous, and depended on microhabitat, with the highest seed density in the unburned control forest but closely followed by the wood piles of erosion barriers. In contrast, very low densities were found under perches of standing snags. Furthermore, frugivorous bird richness seemed to be higher in the erosion barriers than elsewhere. Our results highlight the importance of this specific post-fire management in bird-dependent seed rain and also may suggest a consequent heterogeneous distribution of fleshy-fruited plants in burned and managed areas. However, there needs to be more study of the establishment success of dispersed seeds before an accurate assessment can be made of the role of bird-mediated seed dispersal in post-fire regeneration.     

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.

     

Interactive effects of radiation,temperature and salinity on different life history stages of the Arctic kelp <Emphasis Type="Italic">Alaria esculenta</Emphasis> (Phaeophyceae)

Vegetative resprouting, soil or canopy-stored seed banks, post-fire seed dispersal and germination are the major strategies by which plants regenerate after fires. Post-fire regeneration modes of plants are commonly based on the presence or absence of post-fire recruitment as well as the presence or absence of post-fire resprouting. High temperatures, smoke and ash are characteristics of fire and the post-fire environment. We hypothesized that heat, smoke, ash and pH will have differential effects on seed germination depending on species’ post-fire regeneration strategies: serotinous vs. nonserotinous (which may have soil seed banks) and resprouters vs. nonresprouters (which may be obligate seeders). Here we examined the effects of these factors on the germination of 27 common east Australian species. Most serotinous species supported our hypothesis by showing no effect or reduced germination in response to heat. However, contrary to our prediction, all nonserotinous nonresprouting species also showed no effect or reduced germination in response to heat. Smoke, contrary to our hypothesis, had a negative or no effect on all serotinous and nonresprouting species, but no clear directional effect on serotinous and resprouting species. Supporting our hypotheses, ash and high pH showed positive or nonsignificant effects on the germination of all serotinous resprouting species, and a negative or no effect on nonserotinous resprouting species. However, contrary to our prediction, it had a negative or no effect on the serotinous nonresprouting species and no clear effect on nonserotinous nonresprouting species. We also discovered large differences in germination responses between conspecific populations that varied in their degree of resprouting. Although our data confirmed several of our predictions, the overall conclusion is that the responses of seeds to heat, smoke, ash and pH are not tightly associated with post-fire regeneration functional types. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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

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

Effect of phytohormones on plant regeneration in callus culture of <Emphasis Type="Italic">Iris ensata</Emphasis> Thunb

The effect of phytohormones on plant regeneration in callus culture of Iris ensata Thunb. was studied. 2,4-Dichlorophenoxyacetic acid (2,4-D) proved to be the most suitable auxin for the induction and subculturing of morphogenic callus. Organogenic calluses were induced from isolated embryos at the waxphase in MS medium supplemented with 2,4-D (1.0–2.0 mg/l) and kinetin (0.2–0.5 mg/l). Changes in the medium hormonal composition favor the development of adventitious structures. The setting of adventitious shoot buds took place in the presence of 6-benzylaminopurine, while the development of shoots and root primordia was observed after 2,4-D replacement with indoleacetic acid (2.0 mg/l). The root initiation in regenerating plants required a hormone-free medium. The development of intact seedling and regenerated plants of I. ensata were studied. Analysis of the shoot structure demonstrated that in vitro cultured plants are at the juvenile stage.     

Length of burning rotation and community composition in high-level Calluna-Eriophorum bog in N England

Studies on a long-term experiment on Calluna-Eriophorum bog indicate that the length of time between management fires affects the post-fire development. A short rotation (every 10 years) results in increased dominance by Eriophorum spp. while a long rotation (every 20 years) results in greater abundance of Calluna vulgaris after fire. Variations in post-fires species abundances are related to pre-fire stand composition coupled with the patchy distribution of regeneration centers of rhizomatous species. The usefulness of burning this vegetation type in management for herbivore grazing is questioned.Nomenclature follows Tutin et al. (1964 80) for vascular plants. Smith (1978) for mosses, Watson (1981) for liverworts and Hawksworth, James & Coppins (1980) for lichens. Following common practice. Calluna vulgaris is referred to simply as Calluna. Acknowledgements. I thank M. Raves and the Nature Conservancy Council for permission to work on Moor House National Nature Reserve and for the use of data from Reserve Records, V. J. Hobbs for field assistance, and Prof. C. H. Gimingham for help and advice during the study. I also thank all who worked at Moor House for creating a pleasant working environment. The long-term experiment was planned by Dr R. J. Elliot and analyzed initially by Dr G. I. Forrest.     

Spatiotemporal dynamics of recruitment in Aleppo pine (Pinus halepensis Miller)

Spatial and temporal aspects of recruitment play a central role in plant population and community dynamics and have important basic and applied implications. Here we summarize and discuss the results from studies of spatiotemporal dynamics of recruitment stages (seeds-seedlings-saplings) in Aleppo pine (Pinus halepensis) throughout the species' native range. Seed release is induced either by fire (pyriscence) or by drying atmospheric conditions (xeriscence), each generating a distinct temporal pattern. In both cases most seeds travel relatively short distances (<30 m). Pyriscence is not expected to promote long-distance seed dispersal, but xeriscence is associated with relatively strong winds that can transport seeds over 1 km and more, as predicted by a new mechanistic dispersal model. In the absence of fire, seed survival increases with distance from adults because of higher competition with adults and sibs and higher seed predation near the seed sources. New data provide further evidence for such distance-dependent seed predation and also show that predation rates vary among habitats and are lowest in times of high seed abundance. The resulting recruitment patterns in the absence of fire are characterized by rapid spread and complex spatiotemporal dynamics that are fairly unpredictable and give rise to variable age structure. In contrast, the spatial pattern of the first post-fire generation is highly predictable because it explicitly replicates the spatial pattern of the pre-fire population. Unlike fire-free regeneration in which multiple factors operating at various stages are likely to be important, post-fire regeneration is governed by a fairly specific set of factors (the chemical properties of ash) operating during a specific stage (saplings) and at a specific location (the canopy projection of large burned pines). Post-fire forests are therefore even-aged and have predictable spatiotemporal dynamics. Most studies of Aleppo pine recruitment have focused on the seedling and sapling stages and on post-fire regeneration; we call for greater attention to the seed stage and to more extensive sampling of all recruitment stages in both space and time.     

Pinus sylvestris forest regeneration under different post-fire restoration practices in the northwestern Italian Alps

It is frequently believed that a post-fire environment requires immediate actions in order to be restored. Salvage logging followed by plantation is a common post-fire restoration practice in many forests of the northwestern Italian Alps.The objectives of this study were to assess the impact of active and passive management techniques on the restoration of a burned area of the Aosta Valley and to determine which approach is the most suitable for enhancing Pinus sylvestris regeneration after stand replacing wildfires.The influence of five management options (no intervention; salvage logging; broadleaves plantation; Larix decidua plantation; P. sylvestris or Pseudotsuga menziesii plantation) and environmental variables on natural regeneration structure and composition was evaluated through direct gradient analysis.Pinus sylvestris and Populus tremula were the dominant tree species (40 and 29%, respectively) in the regeneration layer. Density, size, and structural diversity of natural regeneration were higher in the no intervention area. The proximity to forest edge was found to be the most important environmental variable.This study provided evidence that taking advantage of natural restoration processes may be a suitable alternative strategy to the active restoration practices adopted according to the Aosta Valley policy of post-fire management.     

Methylglyoxal with glycine or succinate enhances differentiation and shoot morphogenesis in Nicotiana tabacum callus

The aim of this study was to evaluate the influence of methylglyoxal (MG) on organogenesis and regeneration of tobacco (Nicotiana tabacum L.) plants from callus in media containing glycine or succinate. The best improvement in shoot proliferation and shoot length was obtained in the medium supplemented with 0.1 mM MG and 0.5 mM glycine or 0.25 mM succinate. The histological studies showed vigorous development of corm like structures and shoot organogenesis from callus tissues cultured in MG supplemented media. Biochemical studies also revealed higher content of δ-aminolaevulinic acid (a precursor of chlorophyll) and of chlorophyll.     

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.     

Contingent fire effects on granivore removal of exotic woody plant seeds in longleaf pine savannas

Prescribed fire is increasingly used to inhibit woody encroachment into fire-dependent ecosystems, yet its effects on other processes influencing invasion are poorly understood. We investigated how fire influences exotic woody invasion through its effects on granivore activity, and whether these effects depend on the habitat in which seed predation takes place. We quantified seed removal for four species of exotic woody plants (Albizia julibrissin, Elaeagnus umbellata, Melia azedarach and Triadica sebifera) in 17 sites in longleaf pine savanna that varied in time since fire (one or three growing seasons post-fire) in the sandhills region of North Carolina, USA. Within each site, we established paired plots in upland and upland-wetland ecotone communities and presented seeds in depots that allowed either arthropod, or arthropod and small vertebrate access. We found that differences in seed removal with time since fire were contingent on habitat and granivore community. In ecotones, three of four species had higher proportions of seeds removed from plots that were three growing seasons post-fire than plots one growing season post-fire, whereas only T. sebifera showed this pattern in upland habitat. Allowing vertebrate granivores access to seeds enhanced seed removal, and this effect was strongest in ecotone habitat. While granivores removed seeds of all four plant species, removal of E. umbellata was significantly higher than that of the other species, suggesting that granivores exhibited seed selection. These findings suggest that ecotone habitats in this system experience greater seed removal than upland habitats, particularly as time since fire increases, and differences are mainly due to the activity of vertebrate granivores. Such differences in seed removal, together with seed selection, may contribute to variation in exotic woody invasion of longleaf pine savannas.     

Silvicultural characteristics and natural regeneration of Pinus brutia Ten. — a review

Pinus brutia Ten. is a characteristic species of the eastern Mediterranean. P. brutia and P. halepensis Mill. are distinct species; but P. eldarica, P. stankewiczii and P. pityusa are subspecies of P. brutia. Fire is the major disturbance in P. brutia forests, and several adaptations generally contribute to post-fire regeneration of P. brutia; however, P. brutia can also naturally regenerate without fire. Flowering occurs between March and May. Seed distribution occurs throughout the year, with maximum dispersal in August and the great majority of seeds dispersing within the period July-December.P. brutia retains some closed cones and so establishes a natural canopy seed bank. Seed germination occurs in the rainy season with two frequency peaks: the major one in spring and a minor one in autumn. P. brutia seedlings develop rapidly-growing tap roots. Shelterwood, clearcutting and strip clearcut methods can successfully be applied to P. brutia forests, depending on local conditions. Density of P. brutia seedlings was greater and seedlings were more vigorous when the natural regeneration methods were combined with prescribed fire. Clearcutting methods combined with laying cone bearing branches on the surface of the soil and additional seeding increases regeneration success. Ground litter of 2–4 cm creates a medium that reduces evapotranspiration, delays growth of competing vegetation, and increases seedling survival. Successful regeneration must be established by the end of first vegetation period.