Showing papers in "Plant Ecology in 2004"
TL;DR: In this paper, the use of pines and hardwoods in forest restoration is discussed in the frame of the current disturbance regime and social demands for Mediterranean forests, and combined pine and oak plantations are proposed for degraded land restoration on the basis of the complementary features of both groups of species.
Abstract: Pines have been extensively used for land restoration in the Mediterranean basin and in other parts of the world, since the late 19 th century. The theoretical basis supporting pine utilisation was its stress-tolerant and pioneer features, and their attributed role of facilitating the development of late-successional hardwoods in the long-term. In the present work, the use of pines and hardwoods in forest restoration is discussed in the frame of the current disturbance regime and social demands for Mediterranean forests. Large pine plantations have recently disappeared because of their sensitivity to fire (e.g., Pinus nigra) or because of the short fire-intervals (e.g., Pinus halepensis). Combined pine and oak plantations are proposed for degraded land restoration on the basis of the complementary features of both groups of species. Seeding and containerised seedling plantation, soil amendments and plantation techniques to reduce transplant shock are evaluated for reforestation under water-stressing conditions, on the basis of several experiments performed in eastern Spain. Both P. halepensis and Quercus ilex are tested.
378 citations
TL;DR: The life history of Spanish pines and their relation to fire as the main disturbance factor in their ecosystems was analysed and 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.
Abstract: 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.
321 citations
TL;DR: In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches, suggesting that water competition between trees and grasses was important.
Abstract: In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches. The effects of trees on nutrient, light and water availability were measured to obtain an insight into which resources determine changes in productivity and composition of the herbaceous layer. Soil nutrient availability increased with tree age and size and was lowest in open grassland and highest under dead trees. The lower N:P ratios of grasses from open grassland compared to grasses from under trees suggested that productivity in open grassland was limited by nitrogen, while under trees the limiting nutrient was probably P. N:P ratios of grasses growing under bushes and small trees were intermediate between large trees and open grassland indicating that the understorey of Acacia trees seemed to change gradually from a N-limited to a P-limited vegetation. Soil moisture contents were lower under than those outside of canopies of large Acacia trees suggesting that water competition between trees and grasses was important. Species composition of the herbaceous layer under Acacia trees was completely different from the vegetation in open grassland. Also the vegetation under bushes of Acacia tortilis was different from both open grassland and the understorey of large trees. The main factor causing differences in species composition was probably nutrient availability because species compositions were similar for stands of similar soil nutrient concentrations even when light and water availability was different. Changes in species composition did not result in differences in above-ground biomass, which was remarkably similar under different sized trees and in open grassland. The only exception was around dead trees where herbaceous plant production was 60% higher than under living trees. The results suggest that herbaceous layer productivity did not increase under trees by a higher soil nutrient availability, probably because grass production was limited by competition for water. This was consistent with the high plant production around dead trees because when trees die, water competition disappears but the high soil nutrient availability remains. Hence, in addition to tree soil nutrient enrichment, below-ground competition for water appears to be an important process regulating tree-grass interactions in semi-arid savanna.
302 citations
TL;DR: Black locust supplements soil nitrogen pools, increases nitrogen return in litterfall, and enhances soil nitrogen mineralization rates when it invades nutrient poor, pine-oak ecosystems.
Abstract: We investigated the influence of the exotic nitrogen-fixing black locust (Robinia pseudoacacia) on nitrogen cycling in a pitch pine (Pinus rigida) −scrub oak (Quercus ilicifolia, Q. prinoides) ecosystem. Within paired pine-oak and adjacent black locust stands that were the result of a 20-35 year-old invasion, we evaluated soil nutrient contents, soil nitrogen transformation rates, and annual litterfall biomass and nitrogen concentrations. In the A horizon, black locust soils had 1.3-3.2 times greater nitrogen concentration relative to soils within pine-oak stands. Black locust soils also had elevated levels of P and Ca, net nitrification rates and total net N-mineralization rates. Net nitrification rates were 25-120 times greater in black locust than in pine-oak stands. Elevated net N-mineralization rates in black locust stands were associated with an abundance of high nitrogen, low lignin leaf litter, with 86 kg N ha–1 yr–1 in leaf litter returned compared with 19 kg N ha–1 yr–1 in pine-oak stands. This difference resulted from a two-fold greater litterfall mass combined with increased litter nitrogen concentration in black locust stands (1.1% and 2.6% N for scrub oak and black locust litter, respectively). Thus, black locust supplements soil nitrogen pools, increases nitrogen return in litterfall, and enhances soil nitrogen mineralization rates when it invades nutrient poor, pine-oak ecosystems.
207 citations
TL;DR: Results showed that Pinaceae (particularly the genus Pinus) are more vulnerable to xylem embolism, and show less variability in this character, than other conifers, and implies that no single trait is a sufficient predictor of drought-resistance in Pinaceae.
Abstract: We reviewed the literature to examine the vulnerability to water stress-induced embolism of Pinaceae relative to other conifers and to study the inter-relationships among the main traits involved in the hydraulic function within the Pinaceae. Results showed that Pinaceae (particularly the genus Pinus) are more vulnerable to xylem embolism, and show less variability in this character, than other conifers. Detailed data from 12 populations of Pinaceae (11 species) from three different areas (Pinol and Sala 2000; Martinez-Vilalta and Pinol 2002; Oliveras et al. 2003) was used to study the relationships among hydraulic properties of stems. These included: leaf-to-wood area ratio (AL:A
W), wood- and leaf-specific hydraulic conductivity (KW and KL, respectively), vulnerability to xylem embolism (Ψ50PLC), carbon isotope composition of needles (δ13C) and minimum needle water potential (minimum ΨL). Results showed that hydraulic properties tended to be more correlated among each other than with indicators of environmental (precipitation to potential evapotranspiration ratio, P/E) or physiological water stress (minimum ΨL). The only exception was an increase of δ13C with decreasing minimum ΨL and P/E. Overall, AL:A
W ratio decreased with increasing vulnerability to xylem embolism, and with increasing KW and KL (P<0.05). We found a strong positive relationship between carbon isotope composition and the estimated maximum loss of conductivity due to xylem embolism under field conditions, suggesting stronger stomatal control in more vulnerable species with higher levels of native embolism. Overall, results are consistent with a range of drought-avoidance strategies to minimise the gradient of water potential through the xylem, and show that different relationships among traits are possible depending on the scale of study (individual vs. species or populations). The strong interdependence among hydraulic traits implies that no single trait is a sufficient predictor of drought-resistance in Pinaceae. Finally, it is hypothesised that the intrinsically vulnerable xylem of pines may limit their survival under extremely dry conditions.
192 citations
TL;DR: It is suggested that a number of generations is sufficient for the selection and radiation of fire adaptive traits in P. halepensis, a relatively short living tree with almost no recruitment under forest canopy.
Abstract: 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.
176 citations
TL;DR: The results suggest that cover and richness of different functional groups show a non-linear unimodal pattern along the gradient (age since abandonment), and maximum total richness is found at young stages of abandonment (<20 years), when most life forms and dispersal strategies coexist.
Abstract: We analyse changes in plant cover and species richness along a 60-year chronosequence in semi-arid Mediterranean old-fields of southeastern Spain. The objectives were: (i) to study patterns of species richness along the abandonment gradient in semi-arid conditions (e.g., to test the “humped-back model” in our system); (ii) to test whether different broad life forms (annuals, forbs, grasses and woody species) showed different patterns along the abandonment gradient, and (iii) to examine to what extent plants with different dispersal strategies dominate at different stages of succession. The explained variance of the regression relating species richness to years since abandonment is improved when considering different life forms. The results suggest that cover and richness of different functional groups show a non-linear unimodal (often positive-skewed) pattern along the gradient (age since abandonment). Maximum total richness is found at young stages of abandonment (<20 years), when most life forms and dispersal strategies coexist. Annuals and perennial forbs reached their maximum richness during the first 10 years of abandonment. About 45% of total woody species richness is reached at this time as a consequence of early colonization of zoochorous shrubs. While the results showed a tendency towards a life-form replacement sequence, the pattern is not so clear when looking at the different dispersal strategies. The results complement previous results in Mediterranean conditions and emphasise the importance of considering different functional types when studying successional patterns.
167 citations
TL;DR: Li et al. as discussed by the authors used three algorithms (peak live biomass, peak standing crop and maximum minus minimum live biomass) to estimate the net primary productivity (NPP) of grasslands in northern China.
Abstract: Data on field biomass measurements in temperate grasslands of northern China (141 samples from 74 sites) were obtained from 23 Chinese journals, reports and books. Net primary productivity (NPP) of grasslands was estimated using three algorithms (peak live biomass, peak standing crop and maximum minus minimum live biomass), respectively, based on availability of biomass data in sites. 135 samples which have aboveground biomass (AGB) measurements, have peak AGB ranges from 20 to 2021 g m–2 (mean = 325.3) and the aboveground NPP (ANPP) ranges from 15 to 1647.1 g m–2 per year (mean = 295.7). 72 samples which have belowground biomass (BGB) measurements, have peak BGB ranges from 226.5 to 12827.5 g m–2 (mean = 3116) and the belowground NPP (BNPP) ranges from 15.8 to 12827.5 g m–2 per year (mean = 2425.6). In total 66 samples have the total NPP (TNPP), ranging from 55.3 to 13347.8 g m–2 per year (mean = 2980.3). Mean peak biomass and NPP varied from different geographical sampling locations, but they had a general rough regularity in ten grasslands. Meadow, mountain and alpine grasslands had high biomass and NPP (sometimes including saline grassland). Forested steppe, saline grassland and desert had median values. Meadowed and typical steppes had low biomass and NPP (sometimes including desert). The lowest biomass and NPP occurred in deserted steppe and stepped desert. Grassland ANPP has significant positive relationships with annual and summer precipitation as well as summer temperature (all p<0.01). However, grassland BNPP and TNPP have more significant negative relationships with summer temperature (p<0.01) than with annual temperature (p<0.05). The analysis of climate – productivity correlations implied that aboveground productivity is more controlled by rainfall, whereas belowground and total productivity is more influenced by temperature in the temperate grasslands of northern China. The present study might underestimate grassland NPP in northern China due to limitation of biomass measurements. Data on relative long-term aboveground and belowground biomass dynamics, as well as data of standing dead matter, litterfall, decomposition and turnover, are required if grassland NPP is to be more accurately estimated and the role of temperate grasslands in the regional to global carbon cycles is to be fully appreciated.
162 citations
TL;DR: Phenological patterns in this Araucaria forest appear to be associated with the most predictable and highly correlated of the climatic variables, daylength and temperature and least so with rainfall, which is unpredictable.
Abstract: Phenological patterns in tropical plants usually are associated with the clear seasonality of rainfall associated with very different wet and dry seasons. In southern Brazil, in a subtropical forest with no pronounced dry season (average annual precipitation = 1389 mm, minimum monthly average c. 75 mm), plant phenology was studied to test for patterns (periodicity), to examine how phenological patterns vary among life-forms, and to test whether phenological cycles are associated with climatic variables. Thirty-seven plant species in four life-forms (trees, shrubs, lianas and epiphytes) were studied for 2 yr (1996-98) in an Araucaria forest remnant in southern Brazil, in the state of Parana. Correlation and multiple regression methods established relationships between phenology and climate in terms of daylength, temperature and rainfall. In this Araucaria forest, plants showed seasonality in most life-forms and phenological phases. Leaf-fall, with its peak during the drier months (April to July), was the most seasonal. Flushing and flowering occurred during the wetter months (September to December), while fruiting occurred all year long. Phenologies varied among life-forms, and were strongly associated with daylength or temperature of preceding months, suggesting that plants receive their phenological cues well in advance of their phenological response. Phenologies in this Araucaria forest appear to be associated with the most predictable and highly correlated of the climatic variables, daylength and temperature and least so with rainfall, which is unpredictable.
143 citations
TL;DR: In this paper, the authors compared the vegetation of extant riparian forest along an urban-rural disturbance gradient and found that urban riparian forests were more disturbed and isolated, whereas suburban forests were less disturbed, faced greater development pressure, and had sandier soils.
Abstract: Extensive landscape modification by humans has led to the fragmentation of riparian forests across North America. We compared the vegetation of extant riparian forest along an urban-rural disturbance gradient. In 1999, twenty-five sites along Assiniboine River in Manitoba, Canada were categorized according to land use: urban, suburban, high intensity rural, low intensity rural, and relatively high quality reference forest. Differences in herbaceous, shrub, and tree species composition and diversity were related to the proportion of surrounding land use, forest patch size, connectivity, and area:perimeter ratio. Urban riparian forests were more disturbed and isolated. They were smaller and characterized by drier, more alkaline soils. Moreover, they had significantly lower native and overall understorey species diversity, and had a higher proportion of exotics including Solanum dulcamara and Hesperis matronalis. Suburban forests were less disturbed, faced greater development pressure, and had sandier soils. Although suburban understorey diversity was similar to that of rural forests, suburban sites had a higher proportion of exotic species, especially escaped horticultural and invasive species including Caragana arborescens and Rhamnus cathartica. Reference sites were relatively large and exhibited greater connectivity, but there was little difference in species composition and diversity among high intensity rural, low intensity rural, and reference sites. These site types were less disturbed than either urban or suburban forests, and reference sites were characterized by hydrophilic species including Scirpus fluviatilis and Carex aquatilis. Our results suggest that landscape measures of disturbance, and related changes in environment, may be confidently used to assess impacts of land use on vegetation along urban-rural gradients.
140 citations
TL;DR: The effect of microhabitat and seed burial on the main demographic processes operating during the early recruitment of Quercus ilex, such as postdispersal seed predation, seed germination, and seedling emergence, survival and growth is investigated.
Abstract: This study investigates the effect of microhabitat and seed burial on the main demographic processes operating during the early recruitment of Quercus ilex, such as postdispersal seed predation, seed germination, and seedling emergence, survival and growth. The effect of burial was positive over all the processes analysed in this study, since predation rate was lower (63.6% vs. 88%), whereas germination (53.1% vs. 21.8%) and emergence (32.0% vs. 5.5%) were higher for buried acorns. The quality of some microhabitats remained similar throughout the stages and processes studied. Thus, afforestation provided especially suitable microhabitats for oak establishment, since seed predation was lower, while germination, emergence and seedling survival were higher, than in any other microhabitat. By contrast, the quality of some microhabitats, such as open sites and Holm oaks, differed between recruitment stages. Acorns in open sites escaped predation and germinated easily, but most seedlings died due to summer drought. Similarly, although acorns under Holm oaks can germinate and survive drought, they cannot survive to postdispersal predators. This uncoupling results in a post-dispersal change in the spatial distribution of Q. ilex recruits. Furthermore, there were significant interactions between burial and microhabitat for some demographic processes. The recruitment was in afforestations high irrespective of burial, suggesting that burial is not as beneficial in high-quality habitats as it is in lower-quality ones. An accurate understanding of plant recruitment requires the determination not only of the direct effects of limiting factors but also the potential interactions occurring between them.
TL;DR: Two leaf traits appear to be central in describing species: specific leaf area (SLA), percentage of dry matter (LDMC), and while, SLA and LDMC are strongly correlated, LDMC appears to be less variable than SLA.
Abstract: Nine leaf traits (area, fresh weight, dry weight, volume, density, thickness, specific leaf area (SLA), dry matter content (LDMC), leaf nitrogen content (LNC)) from ten plant species at eight sites in southern mediterranean France were investigated in order to assess their variability along a climatic gradient and their ranking congruency power. After examination of trait correlation patterns, we reduced the nine initial leaf traits to four traits, representative of three correlation groups: allometric traits (dry weight), functional traits (SLA and dry matter percentage) and Leaf Thickness. We analysed the variability of these four leaf traits at species and site level. We observed that between species variation (between 64.5 for SLA and 91% for LDMC) is higher than within species variation. Allowing a good congruency of species ranking assessed by spearman rank correlation () and a good reallocation of individuals to species by discriminant analysis. A site level variability (between 0.7% for Dry weight and 6.9% for SLA) was identified and environmental parameters (altitude, temperature, precipitation, nitrogen, pH) were considered as probable control factors. We found significant correlation between SLA, LDMC and the average minimum temperature (respectively r=0.87 and r=-0,9) and no correlation for the other traits or environmental parameters. Furthermore, we conclude that two leaf traits appear to be central in describing species: specific leaf area (SLA), percentage of dry matter (LDMC. While, SLA and LDMC are strongly correlated, LDMC appears to be less variable than SLA. According to our results the Dry Matter Content (or its reversal Leaf Water Content) appears the best leaf trait to be quantified for plant functional screening. Leaf thickness appeared to be rather uncorrelated with other leaf traits and show no environmental contingency; its variability could not have been explained in this study. Further studies should focus on this trait.
TL;DR: The results indicate that carbon parasitism via AM soil fungi may be an important mechanism by which invasive plants out compete their neighbors, but that this interaction is highly species-specific.
Abstract: Invasive exotic weeds pose one of the earth’s most pressing environmental problems. Although many invaders completely eliminate native plant species from some communities, ecologists know little about the mechanisms by which these exotics competitively exclude other species. Mycorrhizal fungi radically alter competitive interactions between plants within natural communities, and a recent study has shown that arbuscular mycorrhizal (AM) fungi provide a substantial competitive advantage to spotted knapweed, Centaurea maculosa, a noxious perennial plant that has spread throughout much of the native prairie in the northwestern U.S. Here we present evidence that this advantage is potentially due to mycorrhizally mediated transfer of carbon from a native bunchgrass, Festuca idahoensis ,t oCentaurea. Centaurea maculosa, Festuca idahoensis (Idaho fescue, C3), and Bouteloua gracilis (blue gramma, C4) were grown in the greenhouse either alone or with Centaurea in an incomplete factorial design with and without AM fungi. Centaurea biomass was 87–168% greater in all treatments when mycorrhizae were present in the soil (P < 0.0001). However, Centaurea biomass was significantly higher in the treatment with both mycorrhizae and Festuca present together than in any other treatment combination (P < 0.0001). This high biomass was attained even though Centaurea photosynthetic rates were 14% lower when grown with Festuca and mycorrhizae together than when grown with Festuca without mycorrhizae. Neither biomass nor photosynthetic rates of Centaurea were affected by competition with the C4 grass Bouteloua either with or without mycorrhizae. The stable isotope signature of Centaurea leaves grown with Festuca and mycorrhizae was more similar to that of Festuca, than when Centaurea was grown alone with mycorrhizae (P = 0.06), or with Festuca but without mycorrhizae (P = 0.09). This suggests that carbon was transferred from Festuca to the invasive weed. We estimated that carbon transferred from Festuca by mycorrhizae contributed up to 15% of the aboveground carbon in Centaurea plants. Our results indicate that carbon parasitism via AM soil fungi may be an important mechanism by which invasive plants out compete their neighbors, but that this interaction is highly species-specific.
TL;DR: A high correlation was found to exist between annual rainfall, fire frequency, serotiny and bark thickness at a regional level and Sapwood area per hectare proved to be a valuable indirect site-quality index for the objectives of this paper.
Abstract: A wide set of phenotypic characteristics related to life history were studied in mature stands of Pinus canariensis throughout its natural range of distribution in the Canary archipelago. Natural forests ranging from those located in xeric areas through to the sub-tropical cloud forests and high mountain stands were classified into eight ecological regions according to their main climatic features. The recent history of forest fires (covering the last 30 years) was taken into account using a categorical factor with three levels. The phenotypic variables studied included those related to seed dispersal (cone size, number of seed scales, seed and wing size and percentage of serotinous trees) and stem growth both on the breast height section (bark thickness, radial growth at various ages and sapwood and heartwood sizes) and on the entire stem (height growth related to age). The average percentage of serotinous trees present in the ecological regions studied varied from 3 to 35%. Average bark thickness in adult trees ranged from 22 to 49 mm and was found to be unrelated to age or diameter. Growth both in height and diameter was found to decline after an average of 25 years, although clear trends in relation to this could not be established across the ecological regions. A high correlation was found to exist between annual rainfall, fire frequency, serotiny and bark thickness at a regional level. Sapwood area per hectare proved to be a valuable indirect site-quality index for the objectives of this paper. Favourable sites (characterised by a high sapwood area per hectare) displayed the highest levels of both bark thickness and serotiny. These particular areas are those, which have suffered more frequent and intense fires over the last decades. The evolutionary implications of this trend and of other general traits of the species, such as vegetative resprouting, are discussed here in relation to the role of understorey vegetation in fire regimes, competition and volcanic history of the islands.
TL;DR: In this article, the authors compare the floristic and structural similarity of cerrado sensu stricto within and between three physiographic units, named Pratinha, Veadeiros and Sao Francisco, which contain six land systems in central Brazil and cover 10 degrees of latitude and five degrees of longitude.
Abstract: The cerrado has been identified as one of the richest and most threatened biomes of the world, but few phytogeographical studies have been undertaken in the region. A total of 70 land systems based on climate, landscape and soils have been identified in the region, but it remains to be seen if the distribution and structure of the plant communities support these divisions. The aim of this work was to compare the floristic and structural similarity of cerrado sensu stricto within and between three physiographic units, named Pratinha, Veadeiros and Sao Francisco, which contain six land systems in central Brazil and cover 10 degrees of latitude and five degrees of longitude. The woody vegetation of 15 selected sites of the cerrado sensu stricto physiognomy was surveyed under a standardized methodology. The number of species per site varied from 55 to 97, with most sites having around 60 to 70 species, and Shannon´s diversity indices ranged from 3.44 to 3.73, with most sites around 3.5 suggesting high alpha diversity. Sorensen´s floristic similarity index was high, with all Figures above 0.5 between the sites in the same land system in each physiographic unit but low between sites in different land systems in the Veadeiros. Czekanowski similarity indices were lower than Sorensen’s in the comparisons due to a high structural differentiation between the sites. There is a large overlap in species occurrence in the sites but the size of their populations is very different at each site. Therefore, the high beta diversity is mostly due to differences in abundance of species between sites. The sites were separated by physiographic units, considering the first three divisions of TWINSPAN classification. The first axis of DCA ordination showed a gradient going from the cerrado on deep soils in Pratinha, through to those on sandy soils in Sao Francisco and ending on the shallower soils of the Veadeiros. Land systems conformed well with the floristic and structural variations of the vegetation, indicating their potential use in designing a network of conservation areas in the cerrado region and as a basis for decision-making on management.
TL;DR: In this paper, a study of spatial pattern in a semi-arid Karoo shrubland, South Africa, was conducted and the root competition predictions were not met as shrubs of similar rooting depth co-occurred more frequently than expected under random shrub arrangement.
Abstract: Ecologists increasingly use spatial statistics to study vegetation patterns. Mostly, however, these techniques are applied in a purely descriptive fashion without a priori statements on the pattern characteristics expected. We formulated such a priori predictions in a study of spatial pattern in a semi-arid Karoo shrubland, South Africa. Both seed dispersal and root competition have been discussed as processes shaping the spatial structure of this community. If either of the two processes dominates pattern formation, patterns within and between shrub functional groups are expected to show distinct deviations from null models. We predicted the type and scale of these deviations and compared predicted to observed pattern characteristics. As predicted by the seed dispersal hypothesis, small-scale co-occurrence within and between groups of colonisers and successors was increased as compared to complete spatially random arrangement of shrubs. The root competition predictions, however, were not met as shrubs of similar rooting depth co-occurred more frequently than expected under random shrub arrangement. Since the distribution of rooting groups to the given shrub locations also failed to match the root competition predictions, there was little evidence for dominance of root competition in pattern formation. Although other processes may contribute to small-scale plant co-occurrence, the sufficient and most parsimonious explanation for the observed pattern is that its formation was dominated by seed dispersal. To characterise point patterns we applied both cumulative (uni- and bivariate K-function) and local (pair- and mark-correlation function) techniques. Based on our results we recommend that future studies of vegetation patterns include local characteristics as they independently describe a pattern at different scales and can be easily related to processes changing with interplant distance in a predictable fashion.
TL;DR: In this article, the role of nitrogen and phosphorus as limiting factors for growth of pine and holm oak during post-fire recovery was assessed in a 5-year post fire shrubland on calcareous soil.
Abstract: Aleppo pine (Pinus halepensis) and the evergreen holm oak (Quercus ilex) dominate forest areas of the Mediterranean Basin. Both species regenerate abundantly after fires: pine through seedlings and holm oak through resprouts. Cumulative nutrient losses caused by frequent fires may have decreased soil nutrient availability in such areas. To assess the role of nitrogen and phosphorus as limiting factors for growth of these species during post-fire recovery, a field fertilisation and competition experiment was conducted in a 5-year post-fire shrubland on calcareous soil, where naturally-regenerated saplings of Aleppo pine and resprouts of interior holm oak (Quercus ilex subsp. rotundifolia) coexist. Three years after fertilisation, relative basal area increment was 56% greater in pines fertilised with 250 kg P ha–1 than in non fertilised ones. N fertilisation had small or no effects. Interactions between N and P fertilisation were not observed. Growth of Aleppo pine only increased with P fertilisation when neighbours were removed. Hence, the negative effect of neighbours on growth was greater when P availability was enhanced by fertilisation. In contrast, holm oak was able to grow more (110%) in response to increased P supply even without neighbour removal. A common garden experiment was then conducted with potted seedlings to investigate whether the suggested higher competitive capacity of holm oak for P held under a range of P amendments on different soils and competitive situations. P fertilisation increased seedling biomass yield of both species. When P availability increased, a negative effect of neighbours on growth was observed for holm oak and in 70 a lesser extent for Aleppo pine. In conclusion, in the field, holm oak resprouts showed higher competitive ability for P uptake compared to Aleppo pine saplings, but in potted seedlings in common garden conditions this trend was not observed. Therefore holm oak is not always competitively superior to Aleppo pine for P. Potted seedlings of both species had a notable plasticity in shoot/root biomass allocation, but only holm oak increased its proportional allocation to roots when neighbours were present. P availability can be a key factor in growth and competitive relations of these two species, but effects differ depending on soil type, individual age, regeneration type (i.e., seedling versus resprouts), and competitive situation.
TL;DR: In this article, the authors investigated the coexistence of Aleppo pine and Holm oak in Mediterranean forests and found that pine population persistence in the landscape can be explained independently by a competition-colonization tradeoff (in mesic homogeneous environments) and by a tradeoff between shade and drought tolerance (in heterogeneous low disturbed environments).
Abstract: Recurrent anthropogenic and natural perturbations, resource limitations and heterogeneous environments contribute to the maintenance of a remarkable biodiversity in Mediterranean plant communities. Yet, the essential mechanisms of community assembly in these systems remain largely unexplored. In the current paper we investigate the coexistence of Aleppo pine (Pinus halepensis Mill.) and Holm oak (Quercus ilex L.), two of the most widely distributed species in the Iberian Peninsula, in relation to gradients in water availability and disturbance. A spatial model of landscape forest dynamics was implemented, calibrated with experimental data, with stands arranged on a heterogeneous lattice and coupled by dispersal. It was found that pine population persistence in the landscape can be explained independently by a competition-colonization tradeoff (in mesic homogeneous environments) and by a tradeoff between shade and drought tolerance (in heterogeneous low disturbed environments). Both mechanisms reinforce mutually to maintain a shifting mosaic of both taxa along disturbance and aridity gradients. This view is consistent with palynological, historical and forest inventory records. In turn, equilibrium theories of vegetation dynamics (phytosociological), that neglect the role of heterogeneity and disturbances may be inadequate for Mediterranean forests, and have been shown to result in mismanagement practices. We claim that biologically informed models of forest dynamics are desperately needed as diagnosis tools for sustainable forest management.
TL;DR: This study investigated the seed persistence patterns of 25 perennial species, representing several growth forms and life histories, during a three-year burial to hypothesize that light may play a role in overcoming secondary dormancy in those species, and could have resulted in an underestimate for seed persistence in this study.
Abstract: In fire-prone communities such as fynbos, many species rely on regeneration from seed banks in the soil. Persistent seed banks are particularly important for species with life spans shorter than the average fire cycle, in order to counter local extinction. Persistent seed banks also give potential for restoring ecosystems following disturbances such as alien plant invasion. This study investigated the seed persistence patterns of 25 perennial species, representing several growth forms and life histories, during a three-year burial. Long-term persistence (i.e., seed bank half-life exceeding two years) was found in the hard-seeded Fabaceae and Pelargonium, and the nut-fruited Proteaceae. In this group, germinability was low and dormancy increased further following burial, resulting in a highly viable, dormant seed bank after three-year's burial. A second group with potentially long-term persistent seeds includes four taxa (Pseudopentameris, Passerina, Elegia and Restio) that either have low germinability or develop secondary dormancy following burial. Dormancy in the latter group was partially countered by exposure to smoke-seed primer. Of the small-seeded species, only two Erica species with high initial dormancy had long-term persistent seed banks. The other species mostly displayed high initial germinability and short-term persistent seed banks (i.e., seed bank half-life less than two years). This group included taxa with short to medium life-spans (Syncarpha, Roella) that were expected to have long-term persistent seeds in order to buffer against local extinction following average to long fire-return intervals. We hypothesize that light may play a role in overcoming secondary dormancy in those species, and could have resulted in an underestimate for seed persistence in this study. Alternatively, those short to medium life-span species persist via inter-fire recruitment in gaps or long-distance dispersal (of the smallest seed). No correlations were found between seed persistence and seed mass or variance in seed dimensions. Nor was a correlation found between seed persistence and phenol concentration. In fynbos, seed burial of larger seeds by ants and rodents are major processes that operate in conjunction with passive burial of small seeds. Selection for persistence can be expected to operate across all seed sizes and shapes in fire-prone communities.
TL;DR: 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.
Abstract: 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 Cistus spp., 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.
TL;DR: Density of P. brutia seedlings was greater and seedlings were more vigorous when the natural regeneration methods were combined with prescribed fire, and regeneration success was higher than expected.
Abstract: 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.
TL;DR: Most studies of Aleppo pine recruitment have focused on the seedling and sapling stages and on post-fire regeneration; this work calls for greater attention to the seed stage and to more extensive sampling of all recruitment stages in both space and time.
Abstract: 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.
TL;DR: Comparative analysis of fire resistance and survival of the two Mediterranean pine species shows that P. halepensis is more fire sensitive than P. pinea, and the selected models perform well for both species with 85 and 95 percent concordance between predicted probabilities and observed outcomes.
Abstract: The probability of mortality of Pinus halepensis and P. pinea trees after 6 separate wildfires (which took place from 1989 to 1991 in South Eastern France) was modelled with logistic regression analysis using data on tree size and fire-damage descriptors from 998 trees in 13 stands. For both species, probability of mortality increased with increasing percentage of crown scorched and estimated depth of bark charring, and with decreasing tree DBH. The best fitted and field convenient model was selected for each species. Fire-caused mortality of P. halepensis is a function of these three variables and for P. pinea is a function of the first two variables only. Analysis of receiver operating characteristic (ROC) curves indicated that the selected models perform well for both species with 85 and 95 percent concordance between predicted probabilities and observed outcomes for P. halepensis and P. pinea respectively. Models may be used for guiding salvage fire-damaged P. halepensis or P. pinea stands. This paper reports on cases of different management scenarios by selecting different decision criteria when using the proposed models. Comparative analysis of fire resistance and survival of the two Mediterranean pine species shows that P. halepensis is more fire sensitive than P. pinea.
TL;DR: In this paper, the influence of a population of woody bamboo Merostachys riedeliana and different flooding regimes on tree community dynamics in a section of tropical semideciduous forest in South-Eastern Brazil was examined.
Abstract: The influence of a population of the understorey woody bamboo Merostachys riedeliana and different flooding regimes on tree community dynamics in a section of tropical semideciduous forest in South-Eastern Brazil was examined. A forest section with an area of 1.6 ha composed of 71 adjacent plots was located on a slope ending at the river margin. The section was divided into five topographical sectors according to the mean duration of river floods. In 1991 and 1998 all trees with a diameter at the base of the trunk ≥ 5 cm were measured, identified and tagged, and all live bamboo culms were counted. Annualised estimates of the rates of tree mortality and recruitment, gain and loss of tree basal area, and change in bamboo density were calculated for each of the 71 plots and five topographical sectors as well as for diameter classes and tree species. To segregate patterns arising from spatially autocorrelated events, geostatistical analyses were used prior to statistical comparisons and correlations. In general, mortality rates were not compensated by recruitment rates but there was a net increase in basal area in all sectors, suggesting that the tree community as a whole was in a building phase. Tree community dynamics of the point bar forest (Depression and Levee sectors) differed from that of the upland forest (Ridgetop, Middle Slope and Lower Slope sectors) in the extremely high rates of gain in basal area. The predominant and specialised species, Inga vera and Salix humboldtiana, are probably favoured by relaxed competition in an environment stressed by long-lasting floods. In the upland forest, mortality rates were highest at the Middle Slope, particularly for smaller trees, while recruitment rates were lowest. As bamboo clumps were concentrated in this sector, the locally higher instability in the tree community probably resulted from the direct interference of bamboos. The density of bamboo culms in the upland forest was negatively correlated with the rates of tree recruitment and gain in basal area, and positively correlated with tree mortality rates. Bamboos therefore seemed to restrict the recruitment, growth and survival of trees.
TL;DR: In this article, the authors measured several properties of the surface soil (0-10 cm depth) across advancing fronts of Cytisus to assess changes in soil fertility associated with invasion in western Washington state (USA).
Abstract: Shrub encroachment of grassland is a global phenomenon that can cause substantial and rapid changes in soil nutrient levels and distribution. If the woody plants in question also have the capacity to fix nitrogen (N), the effects on soil nutrients may potentially be large. Cytisus scoparius L. Link (Scotch broom) is an introduced leguminous shrub from the Mediterranean region that colonizes open grasslands. We measured several properties of the surface soil (0-10 cm depth) across advancing fronts of Cytisus to assess changes in soil fertility associated with invasion in western Washington state (USA). Both total soil C and N increased (1.8% and 6.5%, respectively), resulting in a decrease in C to N ratio of 22.5 in uninvaded soils to 21.5 inside Cytisus patches. δ15N signature of surface soil did not clearly demonstrate a signal of N2-fixation across the Cytisus gradient. On the other hand, seven-day laboratory aerobic incubations demonstrated 3 and 2.4 times greater mineralization and nitrification rate, respectively, in Cytisus soils compared to uninvaded soils. Despite this increase in N availability, bioassay plants (Achillea millefolium) grown in uninvaded soils in the greenhouse were ∼ 30% larger than those grown in Cytisus-invaded soils, suggesting that Cytisus may have inhibitory effects on some plants growing in invaded soils. These results suggest that the impact of Cytisus invasion on grassland plant communities may be influenced or tempered by chemical or microbial effects on the soil other than simply increased labile N.
TL;DR: In this paper, the impacts of clearcutting on the ground vegetation of remnant late-successional coastal Acadian forests in southwestern Nova Scotia were investigated. And the results suggest that the preservation of remnant old stands may be necessary for the maintenance of some residual plants in highly disturbed and fragmented forest landscapes in eastern Canada.
Abstract: We investigated the impacts of clearcutting on the ground vegetation of remnant late-successional coastal Acadian forests in southwestern Nova Scotia. Vegetation was sampled in 750 1-m2 quadrats established in 16 stands belonging to different recovery periods since clearcutting (3–54 years) and 9 late-successional forests (100–165 years) with no signs of significant human disturbance. Our objectives were to: i) describe the changes in species richness, diversity, and abundance of ground vegetation after clearcutting; ii) examine the responses of residual species (i.e., late-successional flora) to clearcutting; and iii) determine whether any forest species were restricted to or dependent upon the late-successional stages of stand development for maximal frequency and/or abundance. Although clearcutting had no immediate impact on overall alpha richness or diversity, the richness and diversity of residual plants declined after canopy removal and showed no evidence of recovery over 54 years of secondary succession. Consequently, compositional differences between secondary and late-seral stands persisted for many decades after clearcutting. Several understory herbs (e.g., Coptis trifolia (L.) , Oxalis montana (L.), Monotropa uniflora (L.)) were restricted to or attained their highest frequency and abundance in late-seral forests. These results suggest that the preservation of remnant old stands may be necessary for the maintenance of some residual plants in highly disturbed and fragmented forest landscapes in eastern Canada.
TL;DR: The results suggest that it is not possible to predict provenance performance under harsh conditions from their performance under more favourable ones, and clearly emphasises the need for broad selection programs of P. halepensis.
Abstract: As a result of predicted regional climatic changes the need to select for the more drought-tolerant genotypes (ecotypes) among Mediterranean conifers has become clear. Aleppo pine (Pinus halepensis Mill.) seems to be one of the most drought-tolerant pine species. Nevertheless, the existence of geographical trends in their genetic differentiation indicates potentially large differences in drought-tolerance among provenances. This assumption was verified by the finding of large variation among provenances in their internal water relations. Hence, the aim of this study was to compare the ecophysiological behaviour of several Aleppo pine provenances under contrasting climatic conditions. Growth parameters (height and diameter) and survival rate were measured in two provenance trials, one planted under sub-desertic conditions at the northern edge of the Negev desert, and the second under thermo-Mediterranean climatic conditions in the central coastal plains, Israel. Ecophysiological parameters such as: predawn needle water potential, sap flow in the xylem (i.e. transpiration), photosynthesis and water-use efficiency were measured in trees of selected provenances. The results suggest that it is not possible to predict provenance performance under harsh conditions from their performance under more favourable ones. Therefore, selection must be carried out under the exact conditions in which the trees from the resultant selection will be planted. The present study clearly emphasises the need for broad selection programs of P. halepensis.
TL;DR: Post-fire regeneration and invasion are strategies, which seem to complement each other in Pinus halepensis, suggesting a fast selection for serotiny by fire.
Abstract: 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.
TL;DR: Management of grazing in holm oak forests must take the high influence that livestock exerts on the regeneration of Quercus ilex into account, and maximum sustainable grazing intensity should be significantly lower than 0.8 animals per ha.
Abstract: Mediterranean ecosystems are traditionally influenced by human disturbance. In the mountain chains of south-eastern Spain, holm oak forests representing the potential natural vegetation are currently highly fragmented and subjected to extensive grazing by goats and sheep. This grazing activity by large herbivores has important impacts on community structure and dynamics of the remaining remnants because several processes associated with reproduction are affected. The present research analyses the variation of stand structure and regeneration of six forests dominated by Quercus ilex along a gradient of increasing grazing intensity. Stands of higher grazing intensity are characterized by a lower trunk number. Particularly, trees > 1.3 m height and diameter classes < 20 cm are clearly less represented. In addition, more intensively grazed forests show a highly aggregated dispersion of trees, which might be explained by an increase of clonally produced ramets by silvipastoral use. On the other hand, no Quercus ilex seedlings were found in any of the stands examined, although acorn production was high and did not correlate with grazing intensity. However, grazing clearly influences the quality of acorn harvest. We found a significant negative correlation between acorn length, acorn mass and grazing intensity and a positive correlation between the percentage of aborted acorns, acorn predation and herbivore activity. Thus, management of grazing in holm oak forests must take the high influence that livestock exerts on the regeneration of Quercus ilex into account, and maximum sustainable grazing intensity should be significantly lower than 0.8 animals per ha.
TL;DR: The results demonstrate that fungal seed pathogens are common, ubiquitous, and potentially lethal, but that their effects depend on the particular combination of fungus and plant species considered.
Abstract: Although soil fungi may represent an ecologically important cause of mortality of buried seeds, few studies have provided direct evidence of the pathogenicity of fungi colonizing seeds in natural habitats. In response, we conducted a series of experiments to investigate the impacts of soil fungi from a range of habitats on seeds of meadow plants. We compared the survival of seeds of four grasses in five habitats, and isolated fungi from these seeds. We then tested the pathogenicity of selected isolates against two standard sets of plant species: the original four grasses and a broad range of old field species. We found that the soil community contained a large variety of seed-colonizing fungi. Some, but not all, examples of these fungi caused seed mortality; others may be harmless commensals. Some of these isolates negatively affected a broad range of plant species, but others had a more restricted host range; as a result, pathogenicity varied depending upon the particular plant-fungus combination. Few between-habitat differences in seed survival were detected. Our results demonstrate that fungal seed pathogens are common, ubiquitous, and potentially lethal, but that their effects depend on the particular combination of fungus and plant species considered.