Does the strength of facilitation by nurse shrubs depend on grazing resistance of tree saplings?

Facilitation of tree regeneration by nurse shrubs that offer protection against large herbivores is an important driver of wood-pasture dynamics. Here we asked whether the response to facilitation by nurse shrubs depends on the grazing resistance of the protégé saplings. We experimentally tested the protective effects of the thorny Rosa rubiginosa on browsing frequency, survival, and biomass change of saplings of two species-groups, presumably differing in grazing resistance: the coniferous Abies alba and Picea abies and the deciduous Acer pseudoplatanus and Fagus sylvatica saplings. The saplings were planted under and outside (1.5 m) planted nurse shrubs, under zero, low and high grazing intensity. In total, 1920 young saplings were transplanted to 60 blocks and followed for 1 year.Although the number of saplings browsed did not differ between species-groups, the coniferous saplings showed lower resistance to cattle browsing (i.e. lower survival and growth rates) than the deciduous saplings. The less resistant coniferous saplings benefited significantly more from nurse shrubs than the more resistant deciduous species in terms of growth of the surviving saplings, but not in terms of overall survival. This was likely due to herbivory on the nurse shrubs causing incidental browsing on protégé saplings and differences in biomass off-take. At high grazing intensity facilitative effects of the nurse shrubs decreased, especially for the coniferous species.These results have important management implications for the endangered wood-pastures in Western Europe. For a sustainable management and conservation that allows tree recruitment, grazing intensity should remain low to best promote facilitation processes for all tree species, but in particular for the less resistant conifer saplings.     

Different facets of tree sapling diversity influence browsing intensity by deer dependent on spatial scale

Browsing of tree saplings by deer hampers forest regeneration in mixed forests across Europe and North America. It is well known that tree species are differentially affected by deer browsing, but little is known about how different facets of diversity, such as species richness, identity, and composition, affect browsing intensity at different spatial scales. Using forest inventory data from the Hainich National Park, a mixed deciduous forest in central Germany, we applied a hierarchical approach to model the browsing probability of patches (regional scale) as well as the species‐specific proportion of saplings browsed within patches (patch scale). We found that, at the regional scale, the probability that a patch was browsed increased with certain species composition, namely with low abundance of European beech (Fagus sylvatica L.) and high abundance of European ash (Fraxinus excelsior L.), whereas at the patch scale, the proportion of saplings browsed per species was mainly determined by the species’ identity, providing a “preference ranking” of the 11 tree species under study. Interestingly, at the regional scale, species‐rich patches were more likely to be browsed; however, at the patch scale, species‐rich patches showed a lower proportion of saplings per species browsed. Presumably, diverse patches attract deer, but satisfy nutritional needs faster, such that fewer saplings need to be browsed. Some forest stand parameters, such as more open canopies, increased the browsing intensity at either scale. By showing the effects that various facets of diversity, as well as environmental parameters, exerted on browsing intensity at the regional as well as patch scale, our study advances the understanding of mammalian herbivore–plant interactions across scales. Our results also indicate which regeneration patches and species are (least) prone to browsing and show the importance of different facets of diversity for the prediction and management of browsing intensity and regeneration dynamics.     

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.     

Ungulate browsing in winter reduces the growth of <Emphasis Type="Italic">Fraxinus</Emphasis> and <Emphasis Type="Italic">Acer</Emphasis> saplings in subsequent unbrowsed years

Browsing by ungulates has become a hotly debated issue in many European mountain forests in the past century. Acer pseudoplatanus L. and Fraxinus excelsior L. are broadleaved tree species that are preferentially browsed by roe deer (Capreolus capreolus) in central Europe. We analyzed growth data from shaded saplings of both tree species to quantify the extent to which height growth after game browsing is reduced in subsequent, unbrowsed years in forest stands. Sixty saplings of F. excelsior and A. pseudoplatanus from forest stands at Albisriederberg (Switzerland) were available to us that had been dissected into pieces that then were split in the middle for counting tree rings and assessing ungulate damage. We fitted the von Bertalanffy growth equation to these height growth data and included a reduction factor for winter browsing. Both tree species showed significantly reduced height growth in unbrowsed years after one to several browsing events in winter, and this effect increased with the number of browsing events. Saplings with a high growth rate showed a higher growth reduction. After winter browsing, height growth of A. pseudoplatanus saplings was less affected in unbrowsed years than that of F. excelsior saplings. We conclude that browsed saplings of these species in forest stands are not able to compensate browsing-induced height loss, but that height differences between browsed and unbrowsed saplings probably increase over time. A comparison between our analysis and the parameters estimated using equations published by Eiberle for predicting age at 130 cm height suggests that our parameter values are rather conservative estimates of the growth reduction effect after winter browsing. Neither F. excelsior nor A. pseudoplatanus show a distinct pattern in browsing-induced growth reduction with respect to soil moisture, nutrient level, and altitude. We thus conclude that our results are likely to be valid for a wide range of forested sites.     

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.     

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.     

Positive and negative effects of grass,cattle, and wild herbivores on <Emphasis Type="Italic">Acacia</Emphasis> saplings in an East African savanna

Plant–plant interactions can be a complex mixture of positive and negative interactions, with the net outcome depending on abiotic and community contexts. In savanna systems, the effects of large herbivores on tree–grass interactions have rarely been studied experimentally, though these herbivores are major players in these systems. In African savannas, trees often become more abundant under heavy cattle grazing but less abundant in wildlife preserves. Woody encroachment where cattle have replaced wild herbivores may be caused by a shift in the competitive balance between trees and grasses. Here we report the results of an experiment designed to quantify the positive, negative, and net effects of grasses, wild herbivores, and cattle on Acacia saplings in a Kenyan savanna. Acacia drepanolobium saplings under four long-term herbivore regimes (wild herbivores, cattle, cattle + wild herbivores, and no large herbivores) were cleared of surrounding grass or left with the surrounding grass intact. After two years, grass-removal saplings exhibited 86% more browse damage than control saplings, suggesting that grass benefited saplings by protecting them from herbivory. However, the negative effect of grass on saplings was far greater; grass-removal trees accrued more than twice the total stem length of control trees. Where wild herbivores were present, saplings were browsed more and produced more new stem growth. Thus, the net effect of wild herbivores was positive, possibly due to the indirect effects of lower competitor tree density in areas accessible to elephants. Additionally, colonization of saplings by symbiotic ants tracked growth patterns, and colonized saplings experienced lower rates of browse damage. These results suggest that savanna tree growth and woody encroachment cannot be predicted by grass cover or herbivore type alone. Rather, tree growth appears to depend on a variety of factors that may be acting together or antagonistically at different stages of the tree’s life cycle.     

Effects of grazing intensity on growth,reproduction, and abundance of three palatable forbs in Kansas tallgrass prairie

The effects of different intensities of cattle grazing on theaboveground growth, reproduction, and abundances of three palatable forbs werestudied in native tallgrass prairie. Populations of Asterericoides, Ruellia humilis, andAmorpha canescens were sampled at peak flowering duringthe1993–1995 growing seasons in four annually-burned sites varying incattle stocking density [ungrazed, low, moderate, high]. The threeforbs exhibited reduced shoot growth and/or reproduction under moderate toheavy grazing, and in no case did grazing increase any measure of plantperformance. Ruellia showed reduced shoot height andbiomass, percentage of stems flowering, and reproductive biomass in response tograzing. Aster showed decreases in shoot biomass andheightwith grazing. Amorpha showed no change in shoot orreproductive biomass, but a decrease in percent of flowering stems and inreproductive allocation with grazing. Patterns in the percentage of stemsgrazedindicated generally high but variable palatability among these species. Bycontrast, the three species showed inconsistent population response to grazing.Abundance (frequency) of all three species indicated no short-term changebetween years in response to grazing intensity. Responses of these speciesdiffer considerably from those of most other perennial tallgrass prairie forbsthat are unpalatable, unconsumed, and increase in performance (e.g. size,abundance) due to release from competition from the dominant grasses underungulate grazing. The results demonstrate that immediate aboveground growth andreproductive responses of established adults to grazing are not good predictorsof grazer effects on population abundances in tallgrass prairie.     

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.     

Vegetation Control Treatments to Favor Naturally Regenerated Betula alleghaniensis Saplings Following Seed-Tree Cut: Sapling Monitoring Two Years after Treatment

Control of competing vegetation is recommended to ensure successful Yellow birch (Betula alleghaniensis Britton) regeneration within juvenile stands that do not sustain high enough sapling densities of this species. Four contrasting vegetation control treatments were tested to determine their effect on the growth and vigor of eight‐year‐old B. alleghaniensis saplings regenerating after final cutting of a shelterwood seed cut. Vegetation control treatments were TC (total circular removal), PC (circular removal of codominant competing vegetation), TS (total semicircular removal on 180° section), and NC (no vegetation control). Two years after treatment application, diameter growth significantly improved in response to vegetation control treatments, whereas sapling height growth did not. This pattern of biomass allocation was directly related to sapling etiolation, which increased with decreasing severity of vegetation removal. As a result, application of vegetation control, especially TC and PC treatments, was valuable in reducing signs of stress in saplings. However, increasing the severity of vegetation removal also made saplings more conspicuous to herbivores, which increased browsing, especially in the TC and PC treatments. Browsing was sufficient in some plots of the TC and PC treatments to overcome the vigor and diameter growth enhancements observed when browsing was negligible. In contrast to the TC and PC treatments, the TS treatment kept browsing very low while largely removing competition. The results suggest that B. alleghaniensis saplings established after final cutting of a shelterwood seed cut do take advantage of vegetation control treatments, but the decision to apply these treatments must include consideration of local herbivore population densities.     

Early spring leaf out enhances growth and survival of saplings in a temperate deciduous forest

Saplings of many canopy tree species in winter deciduous forests receive the major portion of their light budget for their growing season prior to canopy closure in the spring. This period of high light may be critical for achieving a positive carbon (C) gain, thus contributing strongly to their growth and survival. This study of saplings of Aesculus glabra and Acer saccharum in Trelease Woods, Illinois, USA, tested this hypothesis experimentally by placing tents of shade cloth over saplings during their spring period of high light prior to canopy closure in three consecutive years. Leaf senescence began 16 days (year 0) and 60 days (year 1) earlier for shaded A. glabra saplings than control saplings. No change in senescence occurred for A. saccharum. The annual absolute growth in stem diameter of both species was negligible or negative for shaded saplings, but positive for control saplings. Only 7% of the shaded A. glabra saplings were alive after 2 years, while all control saplings survived for 3 years; only 20% of the shaded A. saccharum saplings survived for 3 years, while 73% of control saplings were alive after the same period. Early spring leaf out is a critical mechanism that allows the long-term persistence of saplings of these species in this winter deciduous forest. Studies and models of C gain, growth, and survival of saplings in deciduous forests may need to take into account their spring phenology because saplings of many species are actually “sun” individuals in the spring prior to their longer period in the summer shade.     

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.     

Effects of below- and aboveground competition from the vines Lonicera japonica and Parthenocissus quinquefolia on the growth of the tree host Liquidambar styraciflua

Detrimental effects of vines on tree growth in successional environments have been frequently reported. Little is known, however, about the relative importance of below and aboveground competition from vines on tree growth. The objective of this study was to quantify and compare the growth responses of Liquidambar styraciflua saplings to below and/or aboveground competition with the exotic evergreen vine, Lonicera japonica (Japanese honeysuckle), and the native deciduous vine, Parthenocissus quinquefolia (Virginia creeper). Soil trenching and/or vine-trellising were used to control the type of vine competition experienced by trees. Comparisons among untrenched treatments tested for effects of belowground competition. Comparisons among trenched treatments tested for effects of aboveground competition. After two growing seasons, Lonicera japonica had a greater effect on the growth of L. styraciflua than did P. quinquefolia. This effect was largely due to root competition, as canopy competition only had a negative effect on tree growth when it occurred in combination with root competition. Leaf expansion was consistently and similarly affected by all treatments which involved belowground competition.     

An example of the seasonal impact of sheep on colonisation by deciduous trees

Summary

Seasonal grazing treatments were begun in 1990 on two sites of bilberry moorland that lay 60–200 m from a strip of riverside woodland in Northern England. Treatments were summer grazing, winter grazing, year-round and no grazing, fences being moved in mid April and mid October each year to open or close plots. The main plant species, Calluna vulgaris, Empetrum nigrum and Vaccinium myrtillus, showed negligible response in cover or height to season of grazing, but colonisation by deciduous trees was much affected. On plots given summer protection (total area 2000 m²) 249 Sorbus aucuparia saplings were counted in 2000, together with one Quercus robur sapling, but no saplings were found in summer-grazed and year-round-grazed plots. Each winter the rowan saplings in the summer-protected plots were severely browsed by sheep, but they recovered in the next growing season. Their increasing trunk diameter and shoot increment up to 2000 suggested that some would soon escape herbivore control. But the fence moves did not take place in 2001,and in April 2002 these saplings were found to have been very heavily browsed. Nearly all survived but regrowth was much poorer in summer 2002 than summer 2000, so several more years of recovery are needed before it can be decided if summer protection from grazing allows succession to woodland at these sites.     

Grass, rainfall and herbivores as determinants of Acacia erioloba (Meyer) recruitment in an African savanna

Acacia species in arid environments are thought to only establish in years of above-average rainfall, so should exhibit cohorted or pulsed recruitment. I studied population demography of Acacia erioloba Meyer in semi-arid savanna in the Kimberley area (mean annual precipitation = 425 ± 132 mm), South Africa, to establish whether they recruit episodically. This species was found to have a sapling bank at the sites, indicating that even though cohorted recruitment probably occurs, it is not the primary factor limiting recruitment to larger size classes. A. erioloba saplings given supplementary water and protection from herbivory showed significantly less height growth than saplings given water but not protected from herbivores, and grass within exclusion plots was taller and denser than outside of exclusion plots. The generalized linear model, although finding a significant difference between watered and protected and watered and unprotected saplings, explained only 16% of variation in growth, demonstrating the importance of factors other than competition from grass when rainfall is above-average. Average height increase across all treatments for a growing season was small (43.1 ± 30.81 mm), suggesting most investment is belowground. A. erioloba sapling below surface stem diameter correlated positively with aboveground growth, indicating larger, and presumably older, individuals grow faster. When grass was removed around saplings, growth rates were not significantly greater than for saplings surrounded by grass, over a growing season. Thus, in these study sites, pulsed events allow seedlings to establish, and then saplings slowly accumulate over many growing seasons within the grass sward, owing to slow aboveground growth combined with competition from grass in absence of grazers. Release events, e.g., heavy grazing combined with good rainfall, may allow mass release, giving the impression of cohorted or pulsed recruitment.     

Comparisons of recruitment, survival, and growth in invasive and native saplings on a volcano

We monitored the recruitment, survival, and growth of tree saplings on invasive (Larix kaempferi) versus native species (Betula and Populus) using 16 20 m × 20 m plots established along elevation gradient on the volcano Mount Koma, Japan, for 7 years because the sapling behaviors should determine forest structures. The crowding of overstory consists mostly of Larix decreased with increasing elevation. Larix recruits were conspicuous, particularly at middle elevation where overstory crowding was intermediate, while Betula recruits were least. Larix overstory crowding inhibited the recruitment of all the taxa, although intermediate crowding promoted the recruitment of Larix. The restriction of sapling emergence was conspicuous at lower elevation where the overstory crowding was highest, probably because of shading, and/or competition with overstory trees. Sapling recruitment for all taxa was restricted at higher elevation, due to high stresses derived from direct solar radiation and strong wind without overstory. The survival of saplings was 96% for Larix and Betula, while it was ca. 50% for Populus. Larix overstory decreased the survival and growth of all the taxa, except Larix survival and Betula growth. The results implied that Larix could establish by high survival once the recruits succeeded everywhere and native sapling regeneration was restricted by Larix overstory. Strong recruitment, survival, and growth of Larix, together with resistance to overstory crowding, enables it to dominate and persist in such disturbed areas regardless of the canopy closure.     

Browser selectivity alters post-fire competition between <Emphasis Type="Italic">Erica arborea</Emphasis> and <Emphasis Type="Italic">E. trimera</Emphasis> in the sub-alpine heathlands of Ethiopia

Mammalian herbivores have the potential to alter the competitive relations of woody species, if consumption is unevenly distributed between species. At elevations above 3,500 m in the southern Ethiopian highlands, vegetation is dominated by Erica arborea and E. trimera. Both species can potentially grow into short trees, but are burnt on a rotation of 6–10 years, and regenerate by re-sprouting from belowground lignotubers. The regenerating scrub is heavily browsed by cattle. We set up browsing exclosures at three burnt sites to quantify the impact of browsing over a 3-year period. When protected from browsing, E. trimera had similar or better height growth than E. arborea, but in browsed vegetation, E. arborea instead grew taller. Browsing was more intense on E. trimera in the first years after fire, indicating a difference in palatability between the species. We checked if browse quality differed, by analysing shoot contents of acid detergent fibre (ADF), protein, phenolics and tannins. Contrary to expectations, the preferred E. trimera contained more ADF, less protein and had a higher tannin activity than E. arborea. Although the vegetative growth of E. arborea is favoured relative to E. trimera under high browsing pressure, rapid change in abundance would not be expected, since short-interval fire will repeatedly eradicate any gains in vegetative growth. However, within the typical fire return interval of less than 10 years, E. trimera barely reach a reproductive state, whereas E. arborea flower profusely. Under the current regime of fire and browsing, this may in the long run be more important than differences in height growth, leading to a gradual increase in the proportion of E. arborea.     

Long-term impacts of goat browsing on bush-clump dynamics in a semi-arid subtropical savanna

The effects of 16 years of continuous browsing by goats in a South African savanna at stocking rates intended for bush control were compared with plots unbrowsed for the same period of time. Differences in bush-clump density, structure and species composition were recorded. Bush-clump density did not differ between browsed and unbrowsed plots. Within individual bush-clumps, browsing was shown to impact more on structure than species composition, with smaller, shorter bush-clumps, containing fewer species but much greater stem-densities. Although species presence/absence was little affected by browsing, many species showed differences in abundance, growth and location within browsed and unbrowsed bush-clumps. Species reduced in abundance in browsed plots included Cussonia spicata, Ehretia rigida, Grewia occidentalis, Jasminum angulare and Senecio linifolius. Several species showed reduced growth in browsed plots, particularly those located at bush-clump edges. The relatively unpreferred Aloe ferox was a notable exception. Although browsing had little effect on the composition of the main clump founding species, emergents or late arrivals, there were twice as many single plants in browsed plots and emergence of several species was restricted to the middle of bush-clumps. Comparison of our findings with aerial photographic evidence and other literature suggest that browsing alone is unlikely to significantly reduce scrub cover, although it can clearly control further expansion. Combinations of fire and browsing, rather than one factor alone, are considered likely to act fastest and most effectively to significantly reduce or remove scrub cover altogether.     

Do <Emphasis Type="Italic">Polylepis australis</Emphasis> trees tolerate herbivory? Seasonal patterns of shoot growth and its consumption by livestock

Browsing is one of the main factors determining survival, growth rate, woodland structure, and distribution of the high mountain tree Polylepis australis. This species has a substantial regrowth capacity, which may function as a mechanism to tolerate herbivory, but it is unknown to what extent it may compensate for the impact of herbivory. In 15 low-density tree stands subject to exclusion, moderate, and heavy livestock pressure, we selected 12 P. australis individuals <2 m tall, tagged four new shoots per tree and measured shoot length every month during a year. At the stand and at the tree level, we analyzed monthly dynamics of growth and browsing, and the annual output in terms of total browsing and total gross and net growth (not discounting and discounting consumption, respectively). In addition, we assessed the influence of stand, tree and microsite characteristics on growth and browsing patterns. Polylepis australis fully compensated for herbivory in terms of shoot gross growth at moderate, but not at heavy livestock pressure. In terms of net growth, this species did not fully compensate for herbivory at any stocking rate. We found a strong coupling between browsing and growth along the year, suggesting that regrowth attracts browsing, and browsing promotes regrowth. At the stand level, annual gross growth was not affected by habitat characteristics, while at the tree level, annual gross growth decreased on more rocky microsites for browsed but not for unbrowsed trees. We concluded that stocking densities should be limited to allow for a reasonable annual net growth, as its nitrogen rich leaves are a valuable food resource and P. australis forests provide important ecosystem services.     

Introduced deer reduce native plant cover and facilitate invasion of non-native tree species: evidence for invasional meltdown

Invasive species are a major threat to native communities and ecosystems worldwide. One factor frequently invoked to explain the invasiveness of exotic species is their release in the new habitat from control by natural enemies (enemy-release hypothesis). More recently, interactions between exotic species have been proposed as a potential mechanism to facilitate invasions (invasional meltdown hypothesis). We studied the effects of introduced deer on native plant communities and exotic plant species on an island in Patagonia, Argentina using five 400 m² exclosures paired with control areas in an Austrocedrus chilensis native forest stand. We hypothesized that introduced deer modify native understory composition and abundance and facilitate invasion of introduced tree species that have been widely planted in the region. After 4 years of deer exclusion, native Austrocedrus and exotic Pseudotsuga menziesii tree sapling abundances are not different inside and outside exclosures. However, deer browsing has strongly inhibited growth of native tree saplings (relative height growth is 77% lower with deer present), while exotic tree sapling growth is less affected (relative height growth is 3.3% lower). Deer significantly change abundance and composition of native understory plants. Cover of native plants in exclosures increased while cover in controls remained constant. Understory composition in exclosures after only 4 years differs greatly from that in controls, mainly owing to the abundance of highly-browsed native species. This study shows that introduced deer can aid the invasion of non-native tree species through negatively affecting native plant species.