Seed rain and natural regeneration in <Emphasis Type="Italic">Lophostemon confertus</Emphasis> plantations in Hong Kong,China

Natural forests have been increasingly replaced by artificial plantations around the world. Some research results suggest that artificial plantations can promote natural regeneration. The efforts in establishing protective forests in Hong Kong over the last 50 years after the Second World War have resulted in many exotic tree plantations which are ideal sites for the study of natural regeneration. This study looked at the seed rain and understorey woody plant diversity in three Lophostemon confertus plantations (25–40 years old) in Hong Kong. All the plantations were > 1 km from the nearest seed sources. Two of the plantations had a seed rain comparable in density and diversity to unplanted grasslands, and the understorey plant diversity was also low and dominated by shrubs. The other site had a higher and more diverse seed rain, as well as higher understorey stem density, but both the seed rain and the understorey were dominated by the shrub Psychotria asiatica. The poor seed rain at two sites and the rarity of potential canopy tree species in both the seed rain and understory regeneration at all sites suggest that planting exotic monocultures does not promote succession. Enrichment planting will be needed to restore forest diversity.     

Tree regeneration in gap-understory mosaics in a subtropical Shorea robusta(Sal) forest

Spatial variation in tree-regeneration density is attributed to the specialization of tree species to light availability for germination and growth. Light availability,in turn, varies across the gap-understorey mosaic. Canopy gaps provide an important habitat for the regeneration of tree species that would otherwise be suppressed in the understory. In subtropical forests, there is still a knowledge-gap relating to how canopy disturbances influence tree-regeneration patterns at local scale, and if they disproportionately favor regeneration of certain species. We aim to analyze whether canopy gaps promote tree regeneration, and tree species are specialized to gaps or understory for germination and growth. We sampled vegetation in 128 plots(0.01 ha), equally distributed in gaps and below canopy, in two subtropical Shorea robusta Gaertn.(Sal) forests in Nepal, recording the number of tree seedlings and saplings in each plot. We compared the regeneration density of seedlings and saplings separately between gaps and the understorey. The mean densities of seedlings and saplings were higher in the gaps at both sites;although there was no difference in the seedling density of the majority of the species between the habitats. No species were confined to either gap or understorey at the seedling stage. We conclude that gaps are not critical for the germination of tree species in Sal forests but these are an important habitat for enabling seedlings to survive into saplings. The classification of trees into regeneration guilds mainly based on germination does not apply to the majority of tree species in subtropical Sal forests. Our results reaffirm that gap creation promotes tree regeneration by favouring seedling survival and growth and can influence forest management for conservation, as well as for plantations.     

Seedling emergence of Betula maximowicziana following human disturbance and the role of buried viable seeds

We observed seed rain and seedling emergence of a pioneer tree species, Betula maximowicziana Regel., for 5 years at various sites differing in canopy and soil surface conditions, and clarified that a buried seed bank contributed to the recruitment of B. maximowicziana seedlings. More seedlings emerged at sites where the soil surface had been disturbed than at sites where the soil surface had not been disturbed. Within each soil surface condition, the densities of the emerged seedlings were generally proportional to the densities of the seeds dispersed the previous year. However, the seedling densities were disproportionately higher than the dispersed seed densities at the sites where the soil surface had been disturbed in the previous year. Soil samples taken during different seasons indicated that B. maximowicziana maintained a persistent seed bank in the soil below 1 cm deep, while it formed a more transient seed bank in the top 1 cm of the soil. The seed rain in the previous year was the primary contributor to the establishment of B. maximowicziana seedlings in a year after a mast seed year. On the other hand, the soil seed bank had a compensational role in a year after a poor seed year. Canopy cover did not affect seedling emergence. Therefore, the previously reported lack of a persistent seedling bank of B. maximowicziana under a canopy will be partially attributed to a relatively rare soil surface disturbance in a stable stand.     

Interacting effects of canopy gap, understory vegetation and leaf litter on tree seedling recruitment and composition in tropical secondary forests

We experimentally investigated interacting effects of canopy gaps, understory vegetation and leaf litter on recruitment and mortality of tree seedlings at the community level in a 20-year-old lowland forest in Costa Rica, and tested several predictions based on results of previous studies. We predicted that experimental canopy gaps would greatly enhance tree seedling recruitment, and that leaf litter removal would further enhance recruitment of small-seeded, shade-intolerant seedlings in gaps. We created a large (320–540 m²) gap in the center of 5 out of 10 40 m × 40 m experimental plots, and applied the following treatments bimonthly over a 14-month-period in a factorial, split–split plot design: clipping of understory vegetation (cut, uncut), and leaf litter manipulations (removal, addition, control). As expected, experimental gaps dramatically increased tree seedling recruitment, but gap effects varied among litter treatments. Litter addition reduced recruitment in gaps, but enhanced recruitment under intact canopy. Species composition of recruits also differed markedly between gap treatments: several small-seeded pioneer and long-lived pioneer species recruited almost exclusively in gaps. In contrast, a few medium-to-large-seeded shade-tolerant species recruited predominantly under intact canopy. Leaf litter represents a major barrier for seedling emergence and establishment of small-seeded, shade-intolerant species, but enhances emergence and establishment of large-seeded, shade-tolerant species, possibly through increased humidity and reduced detection by predators. Periodic clipping of the understory vegetation marginally reduced tree seedling mortality, but only in experimental gaps, where understory vegetation cover was greatly enhanced compared to intact canopy conditions. Successful regeneration of commercially valuable long-lived pioneer trees that dominate the forest canopy may require clear-cutting, as well as weeding and site preparation (litter removal) treatments in felling clearings. Management systems that mimic natural canopy gaps (reduced-impact selective logging) could favor the regeneration of shade-tolerant tree species, potentially accelerating convergence to old-growth forest composition. In contrast, systems that produce large canopy openings (clear-cutting) may re-initiate succession, potentially leading to less diverse but perhaps more easily managed “natural plantations” of long-lived pioneer tree species.     

Seed supply and the regeneration potential for plantations and shrubland in southern China

Assessing the characteristics of seed supply will be vital to better understand the dynamics of forest regeneration. In this study, we surveyed the aboveground vegetation, the seed rain, the seed bank, and natural seedling emergence in four typical 24-year-old plantations (eucalyptus, mixed-native, mixed-legume, and mixed-conifer) and a naturally successioned shrubland in southern China. The dominant species in the understory were similar among the five plant communities. The seed rain and the seed bank were dominated by shrubs and herbs but indigenous tree species were rare. Species that were common to all five-plant communities represented a great proportion of the seeds in the seed rain and seed bank. The seed rain consisted mostly of seeds derived from the local plant community. Seed abundance was greater in the seed bank than in the seed rain, and species richness was greater in the seed bank and in the corresponding plant community than in the seed rain. Species composition similarity between the seed rain, the seed bank, and the aboveground vegetation was low, because the seed rain contained much fewer species, and the seed bank and aboveground vegetation contained many different species, respectively. These findings indicate that both the seed rain and the seed bank play important roles in providing seeds for plant recruitment in the understory, but the seed bank contributes more than the current seed rain to the diversity of recruited plants. The current plant community has little impact on the qualitative composition of the seed rain and seed bank. Based on these data, it appears that succession to the desired zonal, mature forest community is unlikely to result from seeds in the seed rain or seed bank. Lack of seed availability of desired zonal mature forest species is the main bottleneck currently limiting succession in the plantations. Reintroduction of late-successional species could facilitate the desired succession.     

Regeneration patterns of Eastern white pine (Pinus strobus L.) in hardwood-dominated forests in Virginia, USA

Eastern white pine (Pinus strobus L.) is a moderately shade-tolerant species that co-occurs with hardwood tree species in many forests of the eastern United States, as well as in pure stands. The species is valued for its timber, as well as for wildlife and recreation. Regeneration of this species is somewhat unpredictable and often occurs in patches of similarly-aged cohorts. We described the regeneration patterns of this species and examined their relation to environmental variables within hardwood forests of southwestern Virginia, USA. An average of 5.3 white pine patches per ha were observed in this study. The majority of patches consisted of saplings (85%), with 9% of patches in pole size classes, and 6% in seedling size classes. The average density of patches was 43.5 stems with an average age of 20 years. The size of patches averaged 80.6 m². The total density of seedlings and the number of regeneration patches of all sizes of regeneration (seedlings, saplings, and poles) in plots was related to the surrounding density of large white pine trees (potential seed trees). The density of seedlings or patches was not significantly related to current vegetation cover or soil surface cover variables, but more than half of regeneration patches were located in or adjacent to old canopy gaps, most of which were old logging gaps. While seedling regeneration may occur within the understory of these forests near seed trees, advancement to the sapling and pole stage appears to be associated with canopy gap formation.     

红松人工林不同间伐强度效果分析

对不同间伐强度红松人工林的林分生长状况,林下植被生物多样性以及林下土壤的养分状况进行调查研究。结果表明:极强度间伐、强度间伐能有效提高林分直径生长量,增加大径材的出材量,缩短培育大径材的时间,及早达到木材采伐标准。但间伐强度过大影响整个林分的蓄积量。强度间伐有效提高林下植被的丰富度指数和多样性指数,植被状况良好。极强度间伐区土壤的养分状况最好,维持较高的土壤肥力。间伐有利于红松人工林地的可持续利用。     

The potential of hinoki (Chamaecyparis obtusa [Sieb. et Zucc.] Endlicher) plantation forests for the restoration of the original plant community in Japan

We estimated the potential of plantation forests for the restoration of the original plant community. We compared the understory vegetation in hinoki (Chamaecyparis obtusa [Sieb. et Zucc.] Endlicher) plantations at the understory re-initiation stage and in adjacent natural forests. To estimate the effect of the original natural forests on the understory species composition of plantation forests, we established study sites in five types of natural forests (mature evergreen broadleaf, mature deciduous broadleaf, mature evergreen coniferous, immature deciduous broadleaf warm-temperate, and immature deciduous broadleaf cool-temperate) and nearby plantation forests. The understory vegetation of the plantation forests had a higher species richness, a higher proportion of early-seral species, and a higher proportion of herb or fern species than the natural forests. The differences between natural and plantation forests varied according to the species composition of the natural forests. The composition of the understory vegetation of the plantations at the understory re-initiation stage was similar to that of the immature deciduous forests. The characteristics of immature, disturbed forests remained in the understory vegetation of the hinoki forests. No great loss of species was observed. Our findings suggest that most of the original forest species still survive in the understory of the plantation forests. These forests have the potential to follow the successional pathway to broadleaf or mixed forests via thinning or clear-cutting without planting.     

Factors influencing establishment by direct seeding of indigenous tree species in typical plantations and shrubland in South China

Seed limitation of desired indigenous species has constrained the succession of established plantations from an early successional stage, to more natural forests in South China. Because of its low cost and time-saving advantages, direct seeding has been considered a useful method in forest regeneration. To improve our understanding of whether and how direct seeding can lead to the establishment of indigenous tree species, we carried out an experiment in four typical plantations (eucalyptus, mixed-native, mixed-legume, mixed-conifer) and a shrubland in South China. We also tested the effect of understory vegetation and litter removal on seedling establishment and growth. Seeds of three indigenous tree species (Castanopsis chinensis, Cryptocarya chinensis, and Psychotria rubra) were sown with understory vegetation and litter either retained or removed. Seedling emergence, survival, and growth were recorded for 21 months. We found that conditions that favored seedling emergence often differed from those that favored seedling survival. While the removal of understory vegetation and litter did not influence the final seedling establishments for all the three introduced species, growth of Psychotria rubra was influenced at several sites. Seed mass was positively correlated with seedling emergence and subsequent growth. The large-seeded Castanopsis chinensis showed higher emergence and growth than the intermediate-seeded Cryptocarya chinensis or the small-seeded Psychotria rubra. Seedlings of Castanopsis chinensis grew better in plant communities with high understory light penetration and high soil exchangeable potassium. Growth of P. rubra seedlings decreased significantly with soil hydrolyzed nitrogen. In conclusion, direct seeding can be useful for the establishment of indigenous tree species into plantations and shrubland. Optimizing early seedling establishment and growth will require the selection of suitable native species for improving seedling establishment in different plantations. Additionally, understory vegetation and litter and fertilizer should be properly managed to enhance the establishment of certain indigenous tree species.     

Scots pine (Pinus sylvestris L.) plantations contribute to the regeneration of holly (Ilex aquifolium L.) in mediterranean central Spain

The aim of this study is to evaluate the recruitment possibilities of holly (Ilex aquifolium L.) populations under Pinus sylvestris plantations in the mountains of Central Spain. Population recruitment of holly under P. sylvestris L. plantations was studied at two localities. We sampled holly seed rain, seed bank, and seedling emergence and survival (of both emerged and planted seedlings) in both localities. Seed germination was analysed by randomly sowing Ilex seeds and their subsequent recollection and analysis over the following 3 years (only in one locality). Seed rain, seed bank and emerged seedling densities did not differ between localities. Seed bank density was between 10 and 20 seeds m⁻², considerably higher than the observed seed rain (0.4–1.2 seeds m⁻²), and also higher than the emerged seedling density (0–4 seedlings m⁻²). But Ilex is not very efficient in germinating under pine forests (29.1% germination after 3 years). This may indicate a seed accumulation process of at least 5 years. Seedling survival for the first year differed among the studied sites (50–60 vs. 17%). Nevertheless these survival rates are higher than those found for the same seedling populations in different forest habitats (not Scots pine forests) from near locations. Differences in summer water stress between the study sites are proposed as the main cause of the observed differences in seedling survival. It is concluded that P. sylvestris forests may provide one of the best regeneration habitats for holly in the Mediterranean area, where this species is close to the Southern limit of its distribution.     

间伐对栓皮栎人工林林下更新的影响

栓皮栎是我国温带落叶阔叶林代表树种之一,是河南省山区典型地带性树种和主要造林树种.间伐作为一个重要的森林经营措施,研究其对林下更新的影响,可为改善栓皮栎人工林林分结构,维持林下物种多样性,制定科学合理的森林经营措施提供支撑.对间伐3 a后栓皮栎人工林林下幼苗更新物种及更新多样性进行调查,结果表明:间伐后林地内幼苗种类和...     

Fire-free natural regeneration in water limited Pinus halepensis forests: a silvicultural approach

We aimed to elucidate environmental and silvicultural factors that determine the extent of fire-free natural regeneration in east Mediterranean Pinus halepensis forests. The specific aims were to study the potential and identify bottlenecks for natural regeneration and examine the effects of overstory thinning and site preparation treatments. We integrated four experiments conducted in diverse ecoregions in Israel ranging from semiarid to subhumid. Seedling emergence and survival were traced for 2–4 years in Mishmar Ha’Emek Forest (MHF) with annual rainfall of 604 mm; Shaharia Forest (SF)—435 mm; and Yatir Forest (YF)—280 mm. Effects of thinning treatments: clearcut, 100 tree, 200 tree ha^?1 and control—310 tree ha^?1, were examined in MHF. Effects of site preparation treatments: soil scarification and herb clipping were examined in SF. Density of emerged seedlings varied among ecoregions and was linked to average annual rainfall. Within ecoregions, emerged seedling density was higher on north-facing slopes than on south-facing slopes and was strongly related to stand characteristics (e.g., stem basal area) and seed rain. Seedling survival rate of about 20 % was recorded in MHF while no seedling survival was observed in YF and SF during the study years. Thinning treatments reduced seed rain and emerging seedling density, but increased seedling survival and growth. Recruitment 4 years after thinning was highest in the 100 tree ha^?1 treatment and lowest in the control. In SF, seedling survival was limited mainly by herbaceous vegetation and was improved by soil scarification and herb clipping. Silvicultural implications are discussed.     

Planting spacing affects canopy structure,biomass production and stem roundness in poplar plantations

To improve the productivity and wood quality of poplar plantations, effects of four planting spacing on canopy characteristics, biomass production and stem roundness in poplar plantations were evaluated over 8 years. Planting spacing influenced canopy characteristics of the plantations, and further affected the understory vegetation and plantation productivity. Understory vegetation biomass and Shannon-Wiener index were negatively correlated with leaf area index, but both diversity indexes and aboveground biomass of understory vegetation were higher in stands with a wider spacing. Tree diameter growth increased with increasing planting spacing, while the increment in plantations of square configurations (5?×?5 m and 6?×?6 m) was higher than those with rectangular configurations (3?×?8 m, 4.5?×?8 m). The highest poplar biomass production was achieved in the plantation with 5?×?5 m spacing at age 8. Moreover, poplar trees showed a tendency with better stem roundness in a square configuration. The results suggest that planting spacing not only affect canopy characteristics, understory vegetation and tree growth but also wood quality, and square configurations (5?×?5 m and 6?×?6 m) could be a better option for poplar plywood timber production at similar sites.     

Line thinning promotes stand growth and understory diversity in Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don) plantations

We studied the effects of line thinning on stand structure, microclimate and understory species diversity of two Cryptomeria japonica D. Don plantations in south-central Japan. In each of two study sites we compared stand structure between the thinned stand and an adjacent unthinned stand and found that line thinning increased the growth rate of residual trees such that stand basal area may recover within 10 years after thinning. In the thinned stand, more open canopy conditions resulted in higher maximum temperatures on the forest floor during the early growing season than in the unthinned stand. The thinned stand had greater understory plant species richness and biomass than the unthinned stand. This study suggested that line thinning could potentially enhance biodiversity while simultaneously increasing tree-growth rates in overstocked Cryptomeria japonica plantations.     

Growth and structure development of semi-natural larch-spruce-fir (Larix olgensis–Picea jezoensis–Abies nephrolepis) forests in northeast China: 12-year results after thinning

Analyzing and understanding the structure and growth dynamics of semi-natural plantations is useful for their management. Since 1987, 16 plots with 4 treatments (CT: control; LT: light thinning; MT: medium thinning; and HT: heavy thinning) by 0, 20, 30 and 40% of basal area removal, respectively, and four replications were established in semi-natural larch-spruce-fir forests in northeast China. The structure and growth dynamics of semi-natural larch-spruce-fir stands and the effects of thinning on the growth, structure and diversity were examined. A mixed model repeated measures analysis of variance (RMANOVA) was used to test the effects of treatment and time.

Results showed that differences in periodic annual increment (PAI) of stand basal area and volume and the individual diameter and volume among treatments changed over time in a complex statistical interaction. Thinning, however, had a significant effect on growth at tree and stand levels 12 years after thinning while the PAI of the diameter, basal area and volume was positively correlated with thinning intensity. No significant differences were found in the total stand yield among treatments. Composition of tree species group (larch, other conifers and deciduous trees) during monitoring years did not change significantly. Moreover, no significant differences were observed in tree species and size diversity among treatments in the years following thinning. Both thinning and control plots had similar understory plant diversity after the 12 year period. Univariate point pattern analysis revealed that clumped and random distributions were dominant for tree species groups in this study. The current species composition and regeneration dynamics within these semi-natural plantations suggested a development towards mixed coniferous and broad-leaved forests. Management implications for the transformation from larch plantations towards mixed broad-leaved Korean forests with a more diverse structure, the climax vegetation in this region, were discussed.     

High seedling recruitment of indigenous tree species in forest plantations in Kakamega Forest,western Kenya

Tree plantations are often used to compensate for the destruction and conversion of natural forests in the tropics. An important question is whether these plantations allow for the regeneration of indigenous tree species and are expected to transform into more natural forests in the future. To evaluate the potential of differently managed forest types for seedling recruitment of indigenous tree species we studied structural characteristics as well as tree and seedling communities in stands of natural forest, different types of tree plantations and secondary forest in Kakamega Forest, western Kenya. Forest types differed considerably in structural characteristics and tree composition with stands of natural forest significantly differing from all other forest types in vertical foliage height diversity and number of late-successional tree species. By contrast, total seedling species richness and number did not differ among the forest types. Yet, number of seedlings of late-successional species decreased from natural forest and plantations of a mixture of indigenous tree species towards monocultures and secondary forests while number of seedlings of early-successional species increased in the same order. A joint Principal Component Analysis (PCA) corroborated higher similarity among seedling communities than among tree communities. Our results indicate a convergence of recruiting seedling communities in different forest management types suggesting that tree plantations might buffer forest loss to a certain extent and may have the potential to develop into more natural forest over time.     

A preliminary study on the effects of line and selective thinning on bird communities in Hokkaido, northern Japan

In Japan, selective thinning is a common thinning method, though line thinning receives much attention because of its economic merits. In this study, we examined effects of the two thinning methods on bird communities in Todo fir (Abies sachalinensis) plantations in Hokkaido, Japan. We surveyed bird species in forests under four different management types — unthinned, selectively thinned, line-thinned plantation, and naturally regenerated forest (here after referred to as natural forest) stands — using a line-transect method. We also investigated vegetation structure (canopy tree and understory) of these stands. Bird species richness did not differ between natural forests and plantations, while bird total abundance was greater in plantations than in natural forests. Bird species richness and total abundance were comparable among the three management types for plantations. Abundances of 10 bird species were different among the four management types, and five species were more abundant in line-thinned plantations. However, two species were more abundant in selectively thinned stands than in line-thinned stands, and they frequently appeared in natural forests. There were no distinct differences in vegetation structure among the management types for plantations. Our results suggest that line thinning could be beneficial for some bird species in plantations.     

Natural Regeneration in Thinned Douglas-fir Stands in Western Oregon

ABSTRACT

In response to interests by land management agencies to transform even-aged stands to structurally mimic old-growth forests, we evaluated whether thinning in 40- to 80-year-old Douglas-fir (Pseudotsuga menziesii) stands influenced amount and composition of advanced regeneration 5 to 7 years following treatment. We used data from two large-scale management experiments (Density Management Study and Young Stand Thinning and Diversity Study) conducted in western Oregon. Thinning focused on the removal of Douglas-fir, while maintenance of minor species was encouraged. Although both experiments showed higher tree regeneration after thinning, we found that variation in regeneration density was too high (3 orders of magnitude) to find statistical differences among thinning intensities. While seedlings of the major species, Douglas-fir and western hemlock (Tsuga heterophylla), were always present and dominated regeneration on nearly all sites, species trends were driven by high spatial and compositional variation throughout all units, treatments, and sites. Thinning increased the number of species within the regeneration layer. Hereby, species diversity was strongly related to overstory composition, suggesting seed source limitations for minor species. Hence, favoring rare species during thinning operations may be an effective method to increase regeneration species richness. Local conditions, as defined by overstory density appeared more influential than regional climate patterns in determining seedling densities. Shrub and grass competition did not prevent seedling establishment as their cover values were generally not as high as typically found in clearcuts in the region. The high variation in seedling density and species richness within the seedling stratum in the thinned stands may set the stage for development of structural complexity in even-aged Douglas-fir plantations.     

Dynamics and determinants of Quercus alba seedling success following savanna encroachment and restoration

The scattered tree layer that defines savannas is important for structuring the understory community and determining patterns of overstory recruitment. However, encroachment by woody plants has altered overstory tree densities and regeneration dynamics. We characterized seedling success of the savanna-forming species Quercus alba within Midwestern (USA) oak savannas that had been degraded by encroachment (control; n = 4) or experimentally restored by removal of encroaching woody vegetation (treatment; n = 4). In early 2004, 981 seedlings were transplanted along transects radiating from tree boles of overstory Q. alba trees to inter-canopy gaps and monitored for three growing seasons. Seedlings in restored sites had greater survival (>2×), height growth (by >50%), and basal diameter growth (by >20%). In general, seedling survival and growth parameters increased with distance from overstory trees and were greatest in inter-canopy gaps of restored sites. By the final growing season (2006), the seedling survival-by-distance from tree correlation was stronger in control (r² = 0.25) than treatment sites (r² = 0.18), due to relatively uniform (and greater) survival at all distances from trees in treatment sites. In 2006, growth parameters (seedling height, diameter, Δ height, Δ diameter, and # leaves) were significantly (and more strongly) positively correlated with distance from trees in treatment sites. However, seedling herbivory was also greater after treatment and increased with distance from overstory trees. To understand seedling/microenvironment relationships, we created logistic (survival) and linear regression models (Δ height, Δ basal diameter, # leaves in 2006). Control seedling models had consistently greater predictive power and included more variables, suggesting that savanna restoration may decouple seedlings from their microenvironments, potentially by decreasing competition for limiting resources. Encroachment of the savannas in this study is limiting regeneration of Q. alba, suggesting substantially altered regeneration dynamics from those under which these savannas originally formed. Initial responses from our test of restoration, however, were promising and mechanical encroachment removal may be a means to promote overstory regeneration of this species. Finally, the savannas in this study appear inherently unstable and a scattered canopy tree configuration is unlikely to persist without regular disturbance, even in the restoration sites. Repeated mechanical thinning treatments with selected retention of recruiting Q. alba individuals or reintroduction of understory fire or grazing animals may be potential mechanisms for promoting long-term persistence of savannas at these sites.