The influence of competition from herbaceous vegetation and shade on simulated browsing tolerance of coniferous and deciduous saplings

The ability of saplings to tolerate browsing (i.e. the ability to persist with reduced biomass and to compensate for biomass loss) is influenced by the level of stress and their growth strategies. Ultimately, insight into species‐specific responses of saplings to browsing, shade and competition from neighbours will help explain diversity, structure and function of grazed ecosystems such as the endangered wood‐pasture systems. We measured the survival, whole‐sapling biomass and compensatory growth responses of two coniferous (Picea abies and Abies alba) and two deciduous (Acer pseudoplatanus and Fagus sylvatica) tree species to simulated summer browsing (one single clipping event), shade (installation of a shade cloth) and neighbour removal (mowing surrounding vegetation to ground level) treatments and the interactions between them after two‐growing seasons. For all species, there were interacting effects on growth of browsing and environmental condition (shade and neighbours). Simulated browsing resulted in relatively smaller growth losses when plants were growing slowly due to competitive conditions related to herbaceous neighbours. Although none of the clipped saplings could fully compensate for their biomass losses, the saplings were closer to compensation under high competitive conditions than under low competitive conditions. Survival of the clipped saplings remained relatively high and was only significantly reduced for Picea and Acer. Picea was least tolerant of competition and was the only species for which growth was not negatively affected by strong irradiance of a mountain pasture. Surprisingly, the tolerance of saplings to herbivory as browsing tolerance was enhanced under conditions that negatively affected sapling performance (i.e. survival and growth). Apparently, the relative impact of browsing at the early sapling stage is linked to tree life history characteristics such as competition and shade tolerance and will be lower in situations with intense competitive interactions and/or strong irradiance.     

Short-Term Effects of Cattle Browsing on Tree Sapling Growth in Mountain Wooded Pastures

Browsing by livestock has been identified as an important factor preventing tree regeneration in wooded pastures. Two field experiments were performed to investigate the effects of cattle browsing on tree sapling growth in a mountain-wooded pasture. Two size classes (ca. 12–17 cm and 41–59 cm) of each of 4 species (Picea abies, Abies alba, Acer pseudoplatanus and Fagus sylvatica) were exposed to zero, low and high grazing intensities. We measured the proportion of saplings browsed and the effect of browsing on growth. A mowing treatment within the zero grazing intensity treatment showed no significant effect on sapling growth. One percent of the large saplings but 25% of the small saplings escaped browsing. Saplings were better protected when surrounded by taller vegetation. The proportion of saplings browsed was not significantly different among species although evergreen tree saplings lost a larger proportion of biomass than deciduous species when browsed. Under grazing, average current-year shoot production and total aboveground biomass of all species were significantly reduced. Browsing effects tended to be smaller at the lower grazing intensity. Because the deciduous species were least reduced in aboveground growth, especially under the low grazing intensity, we conclude that at least in short-term, deciduous species are less affected by cattle browsing.     

Isolated Trees and Grass Removal Improve Performance of Transplanted Trema micrantha (L.) Blume (Ulmaceae) Saplings in Tropical Pastures

Cattle pastures established in areas previously covered by tropical rainforest can be abandoned in unproductive and degraded conditions. Transplanting native tree species to pastures is one common practice among rainforest restoration techniques, but several environmental barriers compromise transplant success. We assessed whether the presence of isolated trees and the removal of pasture grasses affect survival, growth, and allometry of transplanted saplings of the pioneer tree Trema micrantha (L.) Blume (Ulmaceae) into abandoned pastures in southeast Mexico. An isolated tree was selected in the center of each of four pasture sites of one hectare, and grass treatments were applied under the tree's canopy (0–10 m from the trunk) and in open pasture (15–48 m from the trunk). Grass removal treatments were control (grass present), cut with machete, herbicide application, and total grass removal with a gardening hoe. After 1 year, sapling survival was not different between the canopy and pasture areas (53%). Saplings showed higher survival probability (p < 0.05) in the hoe treatment (63 ± 9%) than in the control treatment (38 ± 9%). Height and crown cover growth rates were faster under the canopy of isolated trees compared with the open pasture. Saplings showed significantly greater crown area/height ratios under the canopy of isolated trees. Stressful environmental conditions restricted sapling growth in the open pasture. We conclude that complete removal of grasses and transplanting T. micrantha saplings in the vicinity of isolated trees can improve transplant success.     

Coarse woody debris can reduce mammalian browsing damage of woody plant saplings in box‐gum grassy woodlands

The critically endangered box‐gum grassy woodlands of south‐east Australia face numerous threats including the failure of woody plant regeneration caused by over‐browsing. In the Australian Capital Territory, over‐browsing of tree and shrub saplings is likely caused by dense populations of Eastern Grey Kangaroo (Macropus giganteus) found in many nature reserves free of livestock. One possible way to protect these saplings is using coarse woody debris (CWD) as a browsing deterrent. We tested this idea by planting palatable Red Stemmed Wattle (Acacia rubida) saplings among manually applied CWD, among naturally fallen CWD, and in the open, in five woodland reserves. We recorded the proportion of saplings browsed, the number of weeks to first browsing and the browsing severity (sapling height lost). Applied CWD protected saplings from being browsed only at relatively low‐to‐moderate kangaroo browsing pressure (as measured by faecal pellet counts). At relatively high browsing pressure, the probability of a sapling being browsed among applied CWD was 100%, similar to the probability in the open treatment (no CWD). Natural CWD, in contrast, provided some protection even at high browsing pressures. Time to browsing was most affected by browsing pressure, although CWD cover also had an influence. Browsing severity was similar between the three treatments and was only affected by browsing pressure. These results indicate that without protection, palatable woody plant saplings have a high chance of being browsed by kangaroos in woodland reserves, and therefore, some protection is needed for successful regeneration. The CWD being applied to reserves has a limited capacity to protect regenerating saplings. If more protection is wanted a CWD structure more resembling natural fallen timber should be used. This could be done by artificially placing branches around plantings. However, the most important action to facilitate regeneration is to manage kangaroo populations to reduce overall browsing pressure.     

From protégé to nurse plant: establishment of thorny shrubs in grazed temperate woodlands

Question: Thorny shrubs play keystone roles in grazed ecosystems by defending non‐protected plants against herbivores, but their establishment in grazed ecosystems is poorly understood. Which factors control establishment of recruits of thorny nurse shrubs in grazed temperate woodlands? Location: Ancient grazed temperate woodlands (52°32′N, 6°36′E), The Netherlands. Methods: We surveyed biotic and abiotic factors for saplings of thorny nurse shrubs in plots with and without saplings. To disentangle these factors, we performed a transplantation experiment over two growing seasons with nurse shrub saplings (Prunus spinosa and Crataegus monogyna) planted in two dominant vegetation types – tall unpalatable swards and short grazed lawns – half of them protected from herbivory via exclosures. Results: Plots with shrub saplings had taller surrounding vegetation, higher soil pH and higher soil moisture than plots without saplings. These plots predominantly contained unpalatable sward species, while plots without saplings mainly contained palatable lawn species. After transplantation, sapling survival was higher in exclosures than in the open, and higher in sward exclosures than in lawn exclosures. Sapling growth was higher in swards than in lawns, higher inside than outside exclosures, and higher for Prunus than Crataegus, while browsing on saplings was higher in lawns. Conclusion: Unpalatable swards form essential establishment niches for thorny shrubs in grazed temperate woodlands: they protect against herbivores before thorns fully develop in saplings, and sapling growth is better due to improved micro‐environmental conditions. Once established and thorny, shrub saplings grow out of the protective range of the swards and in turn facilitate tree seedlings, which are essential for long‐term persistence of grazed temperate woodlands. This study shows that nurse plants may start as protégés before becoming facilitators for other plants in a later life stage. This may be common for nurse plants in various ecosystems. We argue that improved understanding of establishment of nurse plants and their constraining factors is crucial for effective conservation and restoration in various ecosystems.     

Short-term impacts of energy wood harvesting on ectomycorrhizal fungal communities of Norway spruce saplings

The increased demand for harvesting energy wood raises questions about its effects on the functioning of the forest ecosystems, soil processes and biodiversity. Impacts of tree stump removal on ectomycorrhizal fungal (EMF) communities of Norway spruce saplings were studied with 454-pyrosequencing in a 3-year field experiment replicated in 3 geographical areas. This is possibly the most thorough investigation of EMF communities associated with saplings grown on sites subjected to energy wood harvesting. To separate impacts of tree stump and logging residue removal on EMF and plant variables, we used three harvesting treatments with increasing complexity from patch mounding alone (P) to patch mounding combined with logging residue removal (RP), and patch mounding combined with both logging residue and stump removal (SRP). Saplings grown in uncut forests (F) served as references for harvesting treatments. A majority of sequences (>92%) and operational taxonomic units (OTUs, 55%) were assigned as EMF. EMF OTU richness, fungal community composition or sapling growth did not differ between harvesting treatments (P, RP and SRP), while EMF OTU richness, diversity and evenness were highest and sapling growth lowest in the undisturbed reference forests (F). The short study period may partially explain the similarities in fungal and sapling variables in different harvesting treatments. In conclusion, our results indicate that neither stump removal nor logging residue removal have significant additional negative impacts on EMF communities or growth of Norway spruce saplings in the short-term compared with the impacts of more conventional harvesting methods, including clear cutting and patch mounding.     

Above- and below-ground competition effects of two heathland species: Implications for growth and response to herbivory in birch saplings

We examined experimentally the effect of competition from Calluna vulgaris or Molinia caerulea on Betula pubescens saplings subjected to simulated mammalian browsing damage. We tested three hypotheses: 1. that responses to competition will follow the balanced growth hypothesis with the predominant competition effect being below-ground, resulting in increased allocation to roots; 2. that above-ground competition effects from C. vulgaris are greater than M. caerulea, resulting in differences in growth allocation; 3. that, when only B. pubescens saplings are damaged, herbivory reduces its ability to compete with both species.We grew B. pubescens saplings in the presence of below- or above- and below-ground interactions from C. vulgaris or M. caerulea. Saplings were also subjected to simulated browsing by clipping (50% of current year's growth), either pre-senescence or at bud-burst. We measured the morphology and dry mass allocation response of the saplings one year after the browsing was applied.Competition reduced sapling dry mass by approximately 50%, but C. vulgaris reduced dry mass to a greater extent than M. caerulea. This difference was due to a smaller impact of M. caerulea shoots than C. vulgaris shoots on birch growth. Saplings compensated for browsing damage, resulting in no difference in dry mass one year after damage. However, sapling morphological responses to browsing damage were dependent on the competing species.Despite the large competitive effect of below-ground interactions, saplings did not increase allocation to root growth as predicted. Additionally, in response to above-ground interactions from M. caeruela, saplings increased allocation to root growth relative to those growing with C. vulgaris. As such, growth responses to competition were not explained by the balanced growth hypothesis.This study highlights the importance of patterns, as well as intensity, of competition in determining plant responses to inter-specific interactions.     

Effects of simulated browsing on the growth of Austrocedrus chilensis saplings

In the forests of Austrocedrus chilensis (D.Don) Florin et Boutleje (Cupressaceae) it is frequent to observe numerous mutilated and dwarfed saplings which have been browsed by domestic cattle and exotic deer. One way to assess the effects of browsing on plant growth is by simulating this process through defoliation. This study includes results from four years of artificial browsing at different intensities on A. chilensissaplings. Our objective was to evaluate the possible compensatory growth by A. chilensis. To that end we measured the effects of simulated browsing on the growth of saplings kept under two different experimental conditions, with and without water stress. Treatments were applied at the end of winter of 1993 and were repeated on 1994, 1995 and 1996. Treatments consisted in pruning and cutting. Neither cutting effects nor factors interactions were found. Pruning resulted in a reduction of saplings biomass of 44% and was only significant at the more intensive level. Watering produced saplings with five times more biomass, but the general response of A. chilensis to simulated browsing was the lack of compensation regardless the intensity of pruning or the quality of site.     

Regeneration in Atlantic oakwoods: has deer management had a beneficial effect?

Summary

The fragmented Atlantic oakwoods of western Europe are a habitat of conservation importance, and in the UK are threatened by browsing of young trees, invasion by exotic species and under-planting with conifers. Previous research showed that small tree saplings were abundant in Scottish oakwoods, but growth was limited by shading and frequent browsing. Within this study, we aimed to assess whether subsequent changes in deer management had resulted in increased sapling growth, and to determine whether planting of oak within lines of brash left after conifer clearance had offered long-term protection from browsing.

Pellet group counts at two sites in Glen Nant, Argyll, indicated that use by red deer, roe deer and sheep was lower than five years previously. Within one site, sapling survival was high despite continued browsing, and some saplings, particularly hazel, had gained substantial height increments. In contrast, at the second site, survival was low and the surviving saplings had managed no net growth. At a third site in Glen Etive, Argyll, partially protected by fencing, survival of oak saplings was high, and two thirds had increased in height. Planting oak saplings amongst lines of conifer brash did not offer long-term protection, but within a fenced exclosure saplings had grown substantially, despite vigorous competition from the field layer.

Deer management policies within these woods are beginning to show some success in promoting regeneration, but they will need to be continued for a long period to overcome the low growth rates of hardwood species.     

Nurse plants,tree saplings and grazing pressure: changes in facilitation along a biotic environmental gradient

Current conceptual models predict that an increase in stress shifts interactions between plants from competitive to facilitative; hence, facilitation is expected to gain in ecological importance with increasing stress. Little is known about how facilitative interactions between plants change with increasing biotic stress, such as that incurred by consumer pressure or herbivory (i.e. disturbance sensu Grime). In grazed ecosystems, the presence of unpalatable plants is reported to protect tree saplings against cattle grazing and enhance tree establishment. In accordance with current conceptual facilitation-stress models, we hypothesised a positive relationship between facilitation and grazing pressure. We tested this hypothesis in a field experiment in which tree saplings of four different species (deciduous Fagus sylvatica, Acer pseudoplatanus and coniferous Abies alba, Picea abies) were planted either inside or outside of the canopy of the spiny nurse shrub Rosa rubiginosa in enclosures differing in grazing pressure (low and high) and in exclosures. During one grazing season we followed the survival of the different tree saplings and the level of browsing on these; we also estimated browsing damage to the nurse shrubs. Shrub damage was highest at the higher grazing pressure. Correspondingly, browsing increased and survival decreased in saplings located inside the canopy of the shrubs at the high grazing pressure compared to the low grazing pressure. Saplings of both deciduous species showed a higher survival than the evergreens, while sapling browsing did not differ between species. The relative facilitation of sapling browsing and sapling survival – i.e. the difference between saplings inside and outside the shrub canopy – decreased at high grazing pressure as the facilitative species became less protective. Interestingly, these findings do not agree with current conceptual facilitation-stress models predicting increasing facilitation with abiotic stress. We used our results to design a conceptual model of facilitation along a biotic environmental gradient. Empirical studies are needed to test the applicability of this model. In conclusion, we suggest that current conceptual facilitation models should at least consider the possibility of decreasing facilitation at high levels of stress.     

Herbivory effects on saplings are influenced by nutrients and grass competition in a humid South African savanna

Woody plant encroachment is a common consequence of disturbance in savannas. Grazers and browsers interfere with sapling establishment dynamics by direct consumption of plant tissue, changing soil nutrient status (through fertilization and trampling) and grass competition. Studies evaluating the effects of herbivory on sapling establishment have mostly been extrapolated from single species. In a controlled field experiment, we studied the effects of clipping (simulating grazing and browsing), nutrients, grass competition, and their interactive effects on sapling survival and growth of four dominant humid and four dominant mesic savanna species. We conducted this experiment in a humid South African savanna. We found no effects on sapling survival by the treatments provided. However, clipped saplings of all species increased their investment in relative growth rate of stem length (RGR_L). Clipping had a greater negative impact on relative growth rate of more humid than mesic species in terms of stem diameter (RGR_D), total dry biomass and proportion of leaf biomass. Nutrients had a positive effect on the RGR_L and sapling biomass of three mesic species. Positive effects of nutrients on RGR_L of one humid and two mesic species were observed in their clipped saplings only. Grass competition had a strong negative impact on all growth parameters measured. Clipped saplings of one humid and two mesic species had lower RGR_L with grass competition whereas intact saplings showed no significant response. After clipping, humid savanna species were more vulnerable to grass competition than mesic species, with reduced ability to use nutrients. In conclusion, herbivory increases sapling vulnerability to grass competition, with humid species being more susceptible than mesic species, indicating that woody-plant control strategies are more likely to be effective in humid savannas.     

不同生境下濒危植物膝柄木幼树的生态适应性

膝柄木是我国极度濒危植物,也是广西滨海过渡带天然植被的重要组成树种.为了解光因子对膝柄木天然更新的限制影响,该文对林缘、林窗、林下三种不同光照生境下膝柄木幼树的生理和生长指标的年际变化特征进行了研究.结果表明:(1)光合有效辐射不足影响了膝柄木幼树的生长.林下幼树的地径、株高和叶面积增长量显著降低,而生长于光照充足林缘...     

Facilitation and interference underlying the association between the woody invaders Pyracantha angustifolia and Ligustrum lucidum

Questions: 1. Is there any post‐dispersal positive effect of the exotic shrub Pyracantha angustifolia on the success of Ligustrum lucidum seedlings, as compared to the effect of the native Condalia montana or the open herbaceous patches between shrubs? 2. Is the possible facilitation by Pyracantha and/or Condalia related to differential emergence, growth, or survival of Ligustrum seedlings under their canopies? Location: Córdoba, central Argentina. Methods: We designed three treatments, in which ten mature individuals of Pyracantha, ten of the dominant native shrub Condalia montana, and ten patches without shrub cover were involved. In each treatment we planted seeds and saplings of Ligustrum collected from nearby natural populations. Seedlings emerging from the planted seeds were harvested after one year to measure growth. Survival of the transplanted saplings was recorded every two month during a year. Half of the planted seeds and transplanted saplings were cage‐protected from rodents. Results: Ligustrum seedling emergence did not differ among treatments while growth was significantly higher in the absence of shrub cover. Sapling survival was significantly higher under the canopy of Pyracantha, intermediate under Condalia, and lowest in the absence of shrub cover. Caging did not affect growth but enhanced seedling emergence and sapling survival. Conclusion: The differential sapling survival in the shrub canopy treatments is consistent with natural sapling distribution. Pyracantha and, less so, Condalia, has a nurse‐plant effect on Ligustrum. This results from contrasting effects of the shrubs on different stages of the life cycle of Ligustrum: no effect on seedling emergence, negative on seedling growth, and positive on sapling survival. This suggests that efforts to control the expansion of Ligustrum over the landscape should tackle Pyracantha as well.     

Balanced dispersal or source–sink–do both models describe wood mice in farmed landscapes?

Plant responses to browsing can affect root and shoot morphology, which is important to subsequent herbivory, nutrient acquisition and competition. This paper examines the above- and below-ground responses of three browse species, with different growth strategies, to simulated browsing damage at different times of year. Saplings were grown in pots in sand culture to enable whole sapling analysis. At winter dormancy or budburst, 50% of previous year's shoots (and associated leaves/buds) were clipped. Subsequent sapling growth and morphology was compared with that of unclipped control saplings. Treatment differences in growth parameters of each species were observed, including changes in branching patterns, shoot lengths, diameters and ratios, leaf sizes and end-of-season bud numbers. Some responses were damage-induced per se; others differed according to timing of damage. Compensatory growth by the two deciduous species ( Betula pendula , Sorbus aucuparia ) resulted in few biomass differences by the end of the year of damage as compared to controls, but both above- and below-ground growth of clipped Pinus sylvestris was poor, particularly those damaged at budburst, giving strong differences between control, dormant, and budburst clipped saplings.     

樟子松幼树株高及其逆境生理指标对沙埋的响应特征

以3龄樟子松幼树为材料,2013年在科尔沁沙地研究了不同沙埋深度下其株高、叶片膜透性、渗透调节物质含量及保护酶活性变化,以揭示沙埋条件下樟子松幼树生长及其对逆境的生理响应特征。结果显示:(1)在沙埋深度低于株高以上2cm时被埋樟子松幼树能够正常生长,其株高和芽长均明显高于非沙埋对照,并以沙埋深度为株高的50%时增长幅度最大;当沙埋深度超过株高2cm以上时,虽然植株高度和芽长也较埋前有一定增长,但均低于对照,且所有处理植株均未破土,后来全部死亡。(2)所有沙埋处理的叶片可溶性糖含量均显著低于对照,而POD活性显著高于对照,可溶性蛋白质和脯氨酸含量也高于对照。(3)随沙埋深度增加,叶片相对含水量总体呈增加趋势,但大多数处理与对照差异不显著;丙二醛含量基本呈显著下降趋势,可溶性蛋白和脯氨酸含量先增加后下降,而大多数处理的膜透性与对照差异不显著;随着沙埋深度增加,叶片可溶性糖含量显著下降,SOD和POD活性均先增加后下降。(4)相关分析显示,樟子松幼树叶片膜透性变化与MDA含量变化相关性几乎为零,可溶性蛋白与脯氨酸含量呈显著正相关关系,可溶性糖含量与脯氨酸含量呈显著负相关关系。研究表明,沙埋深度低于樟子松株高以上2cm能够促进其幼树生长;沙埋并没有导致樟子松幼树体内的膜脂过氧化,也没有引起细胞膜的损伤,在受到沙埋胁迫时,樟子松幼树体内SOD、POD以及可溶性蛋白和脯氨酸分别在防止其膜脂过氧化和维持细胞膨压中起到重要作用,而可溶性糖含量在沙埋过程中没有起到渗透调节作用。     

Understory responses to restoration in aspen‐conifer forests around the Lake Tahoe Basin: residual stand attributes predict recovery

The removal of conifers from aspen (Populus tremuloides) stands is being undertaken throughout the western United States to restore aspen for local‐ and landscape‐level biodiversity. Current practices include mechanically removing conifers or hand thinning, piling, and burning cut conifers in and adjacent to aspen‐conifer stands. To evaluate the effectiveness of restoration treatments, we examined tree regeneration and herbaceous vegetation cover in thinned, thinned and pile burned, and non‐thinned control stands. Growth rates of small conifer saplings threatening to outcompete and replace aspen were also measured. Two to four years after pile burning, herbaceous vegetation cover within the footprint of burned piles (i.e. burn scars) was 35–73% of that in adjacent areas. Aspen was more likely to regenerate inside burn scars where fewer surrounding trees were true firs. Conifer seedlings were more likely to regenerate in burn scars where more of the surrounding trees were conifers (pine or fir) as opposed to aspen. Fir saplings had much slower growth than did aspen saplings. Overall, our findings show that restoration treatments are promoting desirable outcomes such as enhancing aspen regeneration but that follow‐up treatments will be needed to remove numerous conifer seedlings becoming established after restoration activities. Eliminating conifers, while they are small, growing slowly, and contributing little to fuel loads may be an economical way to prolong restoration treatment effectiveness.     

Comparison of the initial demographies of pine and oak populations in rural pine forests in Korea and Japan

In order to understand the human impact upon demographic change in plant population in rural forests, we examined the population trends ofPinus densiflora andQuercus serrata inP. densiflora forests viewing their seedlings and saplings in rural Korea and Japan. The most prominent factor affecting the regeneration of the pine and oak was the intensity of management activity which controlled the vegetation stratification and its light environment. Open spaces, such as graveyard or cleaned area that allow the long-term direct daylight in dry season to accelerate the surface heat of soil, were unfavorable habitats for germination and growth of both species. The negative effects of the presence of litter and evergreen trees were related to the failure of early seedling and sapling growth of pine. Cover of litter is, especially, another factor related to the growth of pine saplings. It is considered that successful germination and sapling growth of pine and oak in early successional stages is determined by vegetation structures and light condition.     

Do competition and herbivory alter the internal nitrogen dynamics of birch saplings?

Deciduous trees recycle nitrogen within their tissues. The aim of this study was to test the hypothesis that reductions in plant growth, caused by competition and herbivory, reduce the sink strength for N during autumn nutrient withdrawal, and reduce the storage capacity and hence the amount of N remobilized in the following spring. We used (15)N-labelled fertilizer to quantify N uptake, leaf N withdrawal and remobilization. Betula pubescens saplings were grown with either Molinia caerulea or Calluna vulgaris, and subjected to simulated browsing damage. Competition reduced B. pubescens leaf N withdrawal and remobilization, with C. vulgaris having a greater effect than M. caerulea. However, simulated browsing had no significant effect on sapling N dynamics. The patterns of leaf N withdrawal and remobilization closely followed sapling dry mass. We conclude that the effect of competition on sapling mass reduces their N-storage capacity. This reduces sink strength for leaf N withdrawal and the source strength for remobilized N. The ability of saplings to compensate for browsing damage removed any potential effect of browsing on N dynamics.     

Fire and nutrient gradient effects on the sapling ecology of four Acacia species in the presence of grass competition

The variation among tree species in their sapling responses to disturbances such as fire, herbivory and grass competition results in different probabilities of recruitment into savanna communities. A sapling??s ability to persist under these disturbances depends on its investment in growth, defences and stored reserves, based on soil available nutrients. We examined the survival and investments in growth and defence of the saplings of four Acacia species, in the presence of competing grasses, under varying soil nutrient levels. We also have evaluated the effects of fire on these saplings. We performed the study in a greenhouse at the UKZN Botanical Garden, Pietermaritzburg, South Africa. Nutrients had no significant effect on the survival of tree saplings regardless of fire treatment. Only 22.5?% saplings survived after fire treatment compared with 84?% of control saplings. Acacia sieberiana showed the highest resprout survival, while A. karroo did not survive the fire. Growth in A. nigrescens increased only at intermediate levels (0.5N, 1N) of the fertiliser provided. Condensed tannins in A. karroo also decreased at an intermediate level (1N) of fertiliser. Resprouts grew faster in stem height than in diameter and did not invest in chemical defences. No treatment had an effect on the proportion of root and shoot dry biomass. In conclusion, African acacias are not dependent on soil nutrients for their survival and differ in their ability to resprout after fire. Intermediate levels of nutrient availability are beneficial to sapling development as opposed to zero and high nutrient levels, due to nutrient-grass dynamics. The benefits of post-fire grass removal and an increase in nutrients can be effectively utilised by only a few species of acacias.     

Effects of competition and indirect facilitation by shrubs on <Emphasis Type="Italic">Quercus robur</Emphasis> saplings

Indirect facilitation by shrubs has been suggested as a cost-effective way of regenerating oaks in forests of conservation interest. In this study, we tested whether shrubs can enhance growth in pedunculate oak (Quercus robur) by suppressing herbaceous competitors. We studied interactions between young oaks, shrubs, and/or herbaceous vegetation in an open-field experiment, in southern Sweden, over the first 3 years after planting. Oak saplings were grown in four competition treatments: no competing vegetation; with herbaceous vegetation; with shrubs; and with both herbaceous vegetation and shrubs. Competition from shrubs and herbaceous vegetation both reduced stem diameter and biomass accumulation, but they affected biomass partitioning differently. Saplings grown with competition from shrubs partitioned biomass primarily into height growth, while those saplings exposed to competition from herbaceous vegetation invested a relatively higher proportion in root growth. Competition between shrubs and herbaceous vegetation reduced the above-ground biomass of the herbaceous vegetation, resulting in an indirect facilitative effect for the oaks during the first 2 years after planting. However, during the third year, shrubs had a negative effect on biomass accumulation. In summary, results from this study suggest that shrubs indirectly facilitate biomass accumulation of oak saplings by suppressing herbaceous vegetation, possibly by reducing competition for below-ground resources. However, owing to the relatively short duration of positive net outcome for the oak, we recommend that a longer-term assessment of the interaction between oak regeneration and neighboring shrubs be made before the outcome of this study is applied to practical forestry.