Variability in survival and mortality of Acacia drepanolobium Sjøstedt following prescribed burning at Olpejeta Conservancy,Kenya

Mortalities to Acacia drepanolobium, a main item in the diet of the eastern black rhino (Diceros bicornis Michaeli) in Olpejeta conservancy, Kenya, are caused by three main factors: drought, browsers and fire. The effect of controlled fires on A. drepanolobium was examined by monitoring survival and growth in experimental plots before and after fire episodes between 2003 and 2007. Density, survival and growth in A. drepanolobium trees were compared eighteen months before and after burning. Tall trees were more likely to die from fire compared to short trees (R² = 0.575; df = 6; P = 0.048), while seedling densities increased after fire (χ² = 36.57; df = 1; P = 0.001). Even with increased seedling densities, burned areas attracted large numbers of seedling predators, lowering the possibility of seedling recruitment into adult, as mean seedling heights reduced significantly (ANOVA, F = 204.42; df = 1; P = 0.036). Fires also significantly lowered flowering (F = 346; df = 1; P < 0.05) in A. drepanolobium, thereby affecting fruit production. Although fires caused mortalities to adult A. drepanolobium, the most significant effect was tree reversals into seedling height class as trees resprouted. Although fire may increase browse biomass of A. drepanolobium available for black rhino, it is not an appropriate black rhino habitat management tool because burnt areas attract many seedling predators that lower seedling recruitment into adult trees.     

Herbivory on Acacia seedlings in an East African savanna

Trees critically affect the functioning of savanna ecosystems through their effects on nutrient cycling, water availability, and patterns of space use by wildlife. Therefore, whatever factors influence successful recruitment of tree seedlings are important filters controlling savanna ecosystem function. In African savannas, large mammals have been considered the most important agents of mortality for adult trees, but their impacts on tree seedlings are not well‐known. Similarly, the effects of rodents and invertebrates as seedling predators are largely unstudied in Africa. To assess the relative roles of large mammals, rodents, and invertebrates as predators of Acacia seedlings in an African savanna, we conducted two experiments in which we exposed Acacia drepanolobium seedlings to different guilds of herbivores. In the first experiment, seedlings exposed to rodent and invertebrate herbivores did not suffer greater damage than did seedlings exposed only to invertebrates, suggesting that invertebrates caused most of the damage to the seedlings. In the second experiment, 63% of seedlings exposed to all herbivores (large mammals, rodents, and invertebrates) suffered major damage or mortality in 14 days. Seedlings exposed to only rodents and invertebrates, however, suffered damage at a faster rate than did seedlings exposed to rodents, invertebrates, and large mammals, suggesting that small herbivores (rodents and invertebrates) might be compensating for the removal of large herbivores. Certain specific types of damage, such as cotyledon removal, were significantly more common in areas from which large mammals had been excluded, suggesting that the invertebrate herbivore community may differ between areas with and without large mammals. Overall, invertebrates caused the greatest damage to seedlings while rodents had relatively little effect and no seedlings were consumed or trampled by large mammals. Our results indicate that invertebrates can have a pronounced influence on seedling survival for a dominant savanna tree, which in turn may influence tree recruitment and ecosystem function.     

Post-fire mortality and water relations of three congeneric shrub species under extreme water stress — a trade-off with fecundity?

Mortality and water relations of three cooccurringHakea species were studied over the first year following a wildfire in scrub-health in southwestern Australia.Hakea ruscifolia regenerated from both resprouting adults and seedlings, whereasH. smilacifolia andH. polyanthema regenerated only from seedlings. We monitored seedling and resprout survival relative to pre-fire numbers and water relations (shoot water potential, transpiration, pressure-volume parameters) in order to determine the relationship between plant mortality and water relations over the critical first summer drought. Seedlings ofH. polyanthema (few initial seedlings per parent) showed little mortality (8%), achieved through both drought avoidance (low transpiration, high predawn water potential and relative water content (RWC), substantial osmotic adjustment) and drought tolerance later in the season. Seedlings ofH. smilacifolia andH. ruscifolia (high seedling/parent ratios) showed little drought avoidance and high mortality (54–68%). The remaining seedlings spent 3–4 months in a wilted condition (up to 3.6 MPa and 45% RWC below the turgor loss point inH. ruscifolia) indicating marked drought tolerance of the survivors. In contrast to its seedlings,H. ruscifolia resprouts were successful drought avoiders and experienced no mortality. The high level of survival and drought resistance ofH. polyanthema was consistent with its large seedlings (via large seeds) and low initial fecundity. The study highlights the importance of the interaction between ecophysiology-morphology and demography in determining the recruitment strategies of plants.     

Susceptibility of Tree Seedlings to Biotic and Abiotic Hazards in the Understory of a Moist Tropical Forest in Panama

We evaluated temporal patterns of seedling survival of eight Neotropical tree species generated under multiple abiotic and biotic hazards (vertebrates, disease, litterfall) in the forest understory on Barro Colorado Island, Panama. Seedlings were transplanted at first leaf expansion in low densities along a 6-km transect and damage and mortality were recorded for 1 yr. We also planted and monitored small and large artificial seedlings to estimate physical disturbance regimes. During 0–2 mo after transplant, vertebrate consumers of reserve cotyledons caused high mortality of real seedlings, but little damage to artificial seedlings. On real seedlings after 2 mo, disease became an important agent of mortality, despite a decrease in overall mortality rates. Damage by litterfall remained relatively low during the 1-yr study period. Survival ranks among species showed ontogenetic shifts over time, as species changed susceptibility to the mortality agents. Survival after 2 mo was positively correlated with stem toughness, not because species with tough stems were less likely to receive mechanical damage, but because they survived better after receiving mechanical damage. Within each transplant station, artificial seedlings were not good predictors of litterfall damage experienced by real seedlings. Forest-wide litterfall damage level, however, was similar for both real and artificial seedlings ( ca 10%/yr), a moderate level compared to other tropical forests. In conclusion, species traits including biomechanical traits interact to create complex temporal patterns of first year seedling survival, resulting in ontogenetic shifts that largely reflect changes in the relative importance of vertebrate consumers relative to other hazards.     

Establishment of Broad‐leaved Thickets in Serengeti,Tanzania: The Influence of Fire,Browsers, Grass Competition,and Elephants1

The role of Euclea divinorum in the establishment of broad‐leaved thickets was investigated in Serengeti National Park, Tanzania. Thickets are declining due to frequent fires, but have not reestablished when fires have been removed. Seedlings of E. divinorum, a fire‐resistant tree, were found in grassland adjacent to thickets and as thicket canopy trees and may function to facilitate thicket establishment. Seedlings of thicket species were abundant under E. divinorum canopy trees but not in the grassland, indicating that E. divinorum can facilitate forest establishment. We examined E. divinorum establishment in grassland by measuring survival and growth of seedlings with respect to fire, browsers, elephants, and competition with grass. Seedling survival was reduced by fire (50%), browsers (70%), and competition with grass (50%), but not by elephants. Seedling growth rate was negative unless both fire and browsers, or grass was removed. Establishment of thickets via E. divinorum is not occurring under the current conditions in Serengeti of frequent fires, abundant browsers, and dense grass in riparian areas. Conditions that allowed establishment may have occurred in 1890–1920s during a rinderpest epizootic, and measurements of thicket canopy trees suggest they established at that time.     

Encroachment of open grasslands and Acacia drepanolobium Harms ex B.Y.Sjöstedt habitats by Euclea divinorum Hiern in Ol Pejeta Conservancy,Kenya

Euclea divinorum, a fast establishing, unpalatable, and fire resistant bush is considered an invasive species in some parts of its range. In Ol Pejeta Conservancy (OPC), Kenya, E. divinorum bushes cover ?27% of the total area (?9470 ha.) and has been expanding in coverage and encroaching into A. drepanolobium woodlands, a key woody habitat for the endangered black rhino. Between 2006 and 2010, we assessed the spatial distribution, annual rates of spread and recruitment of E. divinorum in OPC. We used data from satellite imagery and belt transects laid at the transition between E. divinorum habitat and other habitats. Density of E. divinorum seedlings increased by 27% annually over five years, with more seedlings establishing in grassland habitat (56.6%) than in A. drepanolobium woodland (43.4%). Within the infection frontier, the number of seedlings was high at the ecotone and reduced predictably with an increase in distance into the infection zone. Increase in rainfall facilitated recruitment and survival of E. divinorum seedlings and also reduced damage on trees and seedlings by mega‐herbivores, especially elephants. This study confirms the encroachment of E. divinorum bushland into other habitats. This has been accelerated by burning and damage to A. drepanolobium habitat thus opening it up to encroachment by E. divinorum.     

Factors affecting survival of tree seedlings in North Queensland rainforests

Summary Seedlings of six species of rainforest trees with widely constrasting ecology and seed morphology were transplanted at 3 weeks of age into tree-fall gaps and the shaded understoreys at two rainforest sites (Curtain Fig and Lamins Hill) on the Atherton Tableland, North Queensland, Australia. In each forest habitat, half of the transplanted seedlings were protected from vertebrates by means of wire cages, and survival was monitored over 16 months. The main objective was to estimate the extent to which independent variables (forest, habitat, protection from vertebrates, and species) contribute to explaining survival differences among the seedlings. Significant differences existed in the mortality among species, whether caged or uncaged. Seedlings unprotected from vertebrates suffered greater mortality (i.e. averaged over all species and forests, mortality > 75%), compared to the caged seedlings (mortality < 40%). Seedling of most species survived better in forest gaps, though the effects varied significantly among the six species. Survival of caged seedlings in Curtain Fig was significantly greater than in Lamins Hill, a pattern attributed to the partial deciduousness of the Curtain Fig forest. Because only few of the uncaged seedlings survived to 16 months, we focussed on the caged seedlings to examine the role of light at the forest habitats on survival. Relationship between seedling mortality and increasing light availability vary significantly among the six species. However across species, seedling mortality decreased linearly as photosynthetically active radiation (PAR) rose from 0.48 to 2.0%, after which an asymptote was reached and further increase in light (up to 8% PAR) did not influence survival significantly. Overall, temporal, species and the various environmental variables and their interactions explained about 60% of the variations in the seedling mortality data, with protection and species differences making the greatest contributions.     

Woody vegetation damage by the African elephant during severe drought at Pongola Game Reserve,South Africa

Elephants (Loxodonta africana) significantly alter ecosystem structure and composition through browsing (e.g. pollarding, debarking and toppling). Such browsing is predicted to intensify during severe drought which may become more common with climate change. Here, we make use of an elephant impact survey from 2012 to 2015 and during the El Nino drought of 2015–2016 at Pongola Game Reserve (107 km²), KwaZulu-Natal, to investigate how severe drought influenced damage severity of different tree heights and species by elephants in this small reserve. Contrary to expectations, damage to common species did not change with severe drought. Crown damage had the highest predicted probability across heights (29%–90%) and species (46%–75%) regardless of drought. However, we found severe drought increased the predicted probabilities of crown damage to smaller trees <4 m, mortality >6 m and severe damage at 4–6 m. Consequently, elephant damage during severe drought may alter vegetation structure by severely damaging or killing large trees (>4 m) and extensively damaging the crowns of trees <4 m. Long-term monitoring of elephant effects on woody vegetation is essential to enable science-based management in response to future drought and elephant damage (e.g. range expansion, beehive deterrents) to protect elephants and conserve woody vegetation.     

Species-mediated soil moisture availability and patchy establishment of Pseudotsuga menziesii in chaparral

The occurrence of mature individuals of Pseudotsuga menziesii in stands of Arctostaphylos species mark the initial stages of mixed evergreen forest invasion into chaparral in central coastal California. We planted two cohorts of P. menziesii seeds at three sites under stands of two Arctostaphylos species and Adenostoma fasciculatum in order to determine whether first-year seedling emergence and survival, particularly during the regular summer drought, underlie the spatial distribution of mature trees observed in chaparral. Regardless of the chaparral species they were planted under, P. menziesii seeds that were not protected from vertebrate predation displayed very little emergence and no survival. In contrast, emergence of P. menziesii that were protected from vertebrate predators was much higher but still did not significantly differ among the three chaparral species. However, survival of protected seedlings under Arctostaphylos glandulosa was much greater than under A. fasciculatum, with intermediate survival under Arctostaphylos montana. While mortality of protected seedlings due to insect herbivory, fungal infection, and disturbance displayed no consistent patterns, summer drought mortality appeared to drive the patterns of survival of P. menziesii under the different chaparral species. These emergence, mortality, and survival data suggest that spatial patterns of P. menziesii recruitment in chaparral are driven by first-year summer drought seedling mortality, but only in years when seeds and seedlings are released from vertebrate predation pressure. Because the first-year drought mortality and survival patterns of P. menziesii seedlings differed strongly depending on the chaparral species, we examined the additional hypothesis that these patterns are associated with differences in the availability of soil moisture under different chaparral species. Both higher survival and lower drought mortality of P. menziesii seedlings were associated with higher soil water potential under Arctostaphylos stands during the summer drought, especially in the subsurface soil. The data suggest that Arctostaphylos stands, particularly stands of A. glandulosa, ameliorate xeric summer conditions to a degree that facilitates first-year establishment of P. menziesii and strongly influences spatial distribution of mature trees. Received: 18 September 1998 / Accepted: 23 December 1998     

Recruitment and survival of Argyrodendron actinophyllum seedlings in an Australian rainforest

The capacity of seedlings to survive for extended periods beneath intact forest increases the likelihood of regeneration of many species of canopy trees in rainforests. I studied the demographics of Argyrodendron actinophyllum (F.M.Bail.) H.L.Edlin seedlings in a subtropical rainforest in northern New South Wales. A mast seeding of A. actinophyllum was observed and subsequent survival of seedlings monitored over a four year period. Densities of seedlings that emerged correlated with seedfall, while seedfall depended on the size and distance to the surrounding trees. Mortality of seedlings showed density-dependence at higher seedling densities (above about 100 seedlings m^?2), apparently in response to browsing pressure that varied with the density of seedlings. Seedlings that were protected from vertebrates by exclosure cages had lower mortality rates than unprotected seedlings and showed no density response. Glasshouse experiments showed seedling growth was reduced by defoliation, light intensity and initial seed weight, and that seedlings could not persist at light intensities below about 1% ambient, which occur in darker patches on the forest floor. Possible mechanisms whereby the observed spatial and temporal patterns of seedling recruitment could reduce the likelihood of the species becoming more common relative to other tree species in the forest are discussed.     

Woodland dynamics under the influence of elephants and fire in Northern Botswana

Sustained elephant browsing and intense burning could result in the loss of woodlands under conditions where elephant densities are high, such as in northern Botswana. Three woodland types dominated by Acacia erioloba, Baikiaea plurijuga and Colophospermum mopane were monitored in plots and contemporary recruitment rates of woody plants were compared with the associated local elephant densities and fire occurrences. Woodland types differed with respect to structure, extent of elephant damage and the occurrence of fire. Canonical correlations indicated that high extent of fire damage and high elephant densities did not covary within the woodland types investigated. Low tree densities in some woodland types were associated with high elephant densities and new elephant damage to plants increased with high elephant densities during the dry season. Plots with an apparent high fire frequency had lower tree densities and higher cover abundance of shrubs and seedlings.The annual rates of tree recruitment/loss in each woodland type were estimated through a model based on observed seedling recruitment, mortality and reversal to lower height classes due to combinations of fire occurrence and elephant browsing. The model suggested that elephants induce tree loss in woodlands dominated by plant species which are principal food sources. Fire however, seems to have a widespread effect across woodlands which could result in extensive tree loss.     

Seedling establishment of Acacia tortilis and Hyphaene compressa in the Turkwel riverine forest, Kenya

This study reports on the spatial and temporal patterns of seedling establishment in the Turkwel riverine forest, Kenya. Seedlings of the dominant tree species Acacia tortilis and Hyphaene compressa were mapped and monitored to assess the underlying causes of seedling recruitment and mortality. The broad‐scale distribution of A. tortilis was not correlated with any environmental variables, while H. compressa seedlings were confined to flood‐exposed sites in the arid downstream section of the floodplain. One year of monitoring showed that seedling recruitment of A. tortilis was evoked by prolonged rainfall, while seedling mortality was caused by desiccation, browsing and trampling. In contrast, seedling recruitment and mortality of H. compressa was largely unaffected by rainfall and livestock, probably due to the high moisture requirements of seeds and the tolerance of seedlings to disturbance. There were no effects of soils, light, or seedling density on the establishment of A. tortilis and H. compressa seedlings. This study demonstrates the importance of parallel mapping and monitoring of riverine seedlings in order to understand patterns and processes of forest regeneration in arid and semi‐arid floodplains.     

Canopy vs. soil effects of shrubs facilitating tree seedlings in Mediterranean montane ecosystems

Abstract. Question: Is the facilitative effect of nurse shrubs on early recruitment of trees mediated by a ‘canopy effect’(microclimate amelioration and protection from herbivores), a ‘soil effect’(modification of soil properties), or both? Location: Two successional montane shrublands at the Sierra Nevada Protected Area, SE Spain. Method: Seedlings of Quercus and Pinus species were planted in four experimental treatments: (1) under shrubs; (2) in open interspaces without vegetation; (3) under shrubs where the canopies were removed; (4) in open interspaces but covering seedlings with branches, mimicking a shrub canopy. Results: Both effects benefited seedling performance. However, microclimatic amelioration due to canopy shading had the strongest effect, which was particularly pronounced in the drier site. Below‐ground, shrubs did not modify soil physical characteristics, organic matter, total N and P, or water content, but significantly increased available K, which has been shown to improve seedling water‐use efficiency under drought conditions. Conclusions: We propose that in Mediterranean montane ecosystems, characterised by a severe summer drought, pioneer shrubs represent a major safe site for tree early recruitment during secondary succession, improving seedling survival during summer by the modification of both the above‐and below‐ground environment.     

Ecology of Aglaia mackiana (Meliaceae) Seedlings in a New Guinea Rain Forest1

The large seeds of Aglaia mackiana (Meliaceae) germinate and produce vigorous seedlings under closed canopies or in large gaps. To assess seedling ecology after germination, we measured growth, herbivore damage, and survivorship of seedlings over one year. The sample included shaded seedlings from dispersed seeds, undispersed seeds under parent trees, and seedlings transplanted to gaps. We quantified the light environment using hemispherical canopy photographs taken above seedlings at the beginning and end of the one–year study. Seedlings transplanted to gaps grew faster and had more leaves, larger total leaf surface area, longer secondary roots, and greater root mass than shaded seedlings. Seedlings in gaps did not differ from shaded seedlings in survivorship or amount of herbivore– and pathogen–caused leaf damage. The canopy photographs taken one year apart suggest there is a rough equilibrium in closed canopies with slight changes occurring around an average light level. Sites with < 0.06 ISF (a unitless, relative measure of canopy openness or reflected sunlight) tended to remain the same with minor fluctuations toward brighter or darker. Sites with canopy openness > 0.06 ISF tended to close; few gaps grew larger. Seedlings under parenr trees and seedlings away from parent trees had similar amounts of leaf damage and virtually identical survivorship after 18 months, but seedlings under parent trees had slower growth rates and smaller total leaf surface areas. Dispersal did not strongly benefit seeds via escaping high levels of mortality or comperition around the parent.     

The impact of ‘available area’ onArtemisia tridentata seedling dynamics

Seedling survival ofArtemisia tridentata ssp.tridentata (basin big sagebrush) was studied during 1984 and 1986 on a semi-arid rangeland in central Utah, USA. Seedlings were mapped in ungrazed pastures and observed on a 10 day interval for survival. Available resource area around individual seedlings was calculated using Theissen polygons. Over 90% of the naturally-occurring seedlings occurred in polygons of 150 mm² or less. Seedlings with available areas greater than 300 mm² were more likely to survive than seedlings with less resource area. Differences in overall seedling survival in different study sites were noted. Survival was similar for all seedlings with less than 50 mm² available area. In 1986, precipitation and seedling survival were considerably lower than in 1984. Even in 1986, however, seedling survival in polygons greater than 300 mm² was always higher than for seedlings in smaller polygons. Survival patterns over each growing season closely followed the precipitation patterns.Mortality of first yearArtemisia seedlings seemed to be density independent at the beginning of the growing season. Patterns of survival for seedlings in different available area classes were similar for the first 20 days after emergence. Survival rates were then consistently higher for seedlings with more than 300 mm² resource area. Changes in survival rates were linked to differences in pastures and the presence of nurse plants.     

Effects of Deer Browsing, Fabric Mats, and Tree Shelters on Quercus rubra Seedlings

We examined the effects of deer browsing, fabric mats, and tree shelters on the growth and survival of Quercus rubra (northern red oak) seedlings planted as part of a reforestation project in southcentral Minnesota. Browsing by white-tailed deer occurred on 68.6% of the seedlings growing without tree shelters, whereas only 3.6% of the trees were browsed by mice or voles and 2.5% were browsed by rabbits. Fabric mats, used to control competition from herbaceous plants for roughly half of the seedlings included in the study, had a detrimental effect overall. Seedlings grown with mats had a greater frequency of deer browsing and a greater chance of dying than seedlings grown without mats. Stem height for seedlings browsed by deer was less overall than for nonbrowsed seedlings, although this pattern varied with use of fabric mats and plot location. The use of plastic tree shelters effectively prevented deer browsing and reduced the mortality rate from 34.6% to 3.2%. Our results indicate that fabric mats should not be used in restoration projects with large deer populations. They also suggest that planting seedlings away from existing forest edges and using seedling protection measures such as tree shelters will increase seedling survival and growth in future restoration projects.     

Effects of large herbivores and fire on the regeneration of Acacia erioloba woodlands in Chobe National Park, Botswana

Acacia erioloba woodlands provide important forage and shade for wildlife in northern Botswana. Mortality of mature trees caused by browsing elephants has been well documented but the lack of regeneration of new trees has received little attention. Annual growth of new shoots and changes in height were measured to determine the influence of elephants and small ungulate browsers, rainfall and fire on the growth and survival of established A. erioloba seedlings from 1995 to 1997 in the Savuti area of Chobe National Park. All above‐ground vegetation was removed from 40% of established seedlings in 1995 and 28% in 1997 by browsing elephants, and the mean height of remaining seedlings decreased from >550 mm to <300 mm. When seedlings browsed by kudu, impala and steenbok but not elephants are considered, mean seedling height increased <50 mm per year, even though mean new shoot growth remaining at the end of the dry season was 100–200 mm. Fires burned portions of the study area in 1993 and 1997, killing above‐ground vegetation, but most established A. erioloba seedlings survived, producing coppice growth from roots. While elephants and fire caused the greatest reduction in established seedling height and number, small browsers suppressed growth, keeping seedlings vulnerable to fire and delaying growth to reproductive maturity.     

Effects of El Niño drought on seedling dynamics in a seasonally dry tropical forest in Northern Thailand

As El Niño is predicted to become stronger and more frequent in the future, it is crucial to understand how El Niño-induced droughts will affect tropical forests. Although many studies have focused on tropical rainforests, there is a paucity of studies on seasonally dry tropical forests (SDTFs), particularly in Asia, and few studies have focused on seedling dynamics, which are expected to be strongly affected by drought. Seedlings in SDTFs are generally more drought-tolerant than those in the rainforests, and the effects of El Niño-induced droughts may differ between SDTF and tropical rainforests. In this study, we explored the impact of El Niño-induced drought at an SDTF in northern Thailand by monitoring the seedling dynamics at monthly intervals for 7 years, including a period of strong El Niño. The effects were compared between two forest types in an SDTF: a deciduous dipterocarp forest (DDF), dominated by deciduous species, and an adjacent lower montane forest (LMF) with more evergreen species. El Niño-induced drought increased seedling mortality in both the forest types. The effect of drought was stronger in evergreen than in the deciduous species, resulting in higher mortality in the LMF during El Niño. However, El Niño increased seedling recruitment only in the DDF, mainly because of the massive recruitment of the deciduous oak, Quercus brandisiana (Fagaceae), which compensated for the mortality of seedlings in the DDF. As a result, El Niño increased seedling density in the DDF and decreased it in the LMF. This is the first long-term study to identify the differences in the impacts of El Niño on seedlings between the two forest types, and two leaf habits, evergreen and deciduous, in Southeast Asia. Our findings suggest that future climate change may alter the species composition and spatial distribution of seedlings in Asian SDTFs.     

Conifer seedling distribution and survival in an alpine-treeline ecotone

The importance of seedling establishment to the position ofalpine-treeline is recognized, yet little is known about factorsaffecting the survival of seedlings of treeline conifers during their initialyears of growth and establishment. This establishment period may have thegreatest mortality of all life stages until death of mature trees by disease orfire. Spatial and temporal patterns in the distribution and survival ofseedlings of Picea engelmannii and Abieslasiocarpa were evaluated over four years in analpine-treeline ecotone of the Snowy Range, Wyoming, USA. Seedlings andsaplings of both species occurred most frequently near islands of adult trees.For P. englemannii, this appeared partly due to decreasedsurvivorship of young seedlings (< 5 cm height) with greaterdistance away from tree islands. Survival of emergents of P.engelmannii was 28% greater on the north compared to southsides of tree islands, 48% lower on south-facing slopes comparedto other aspects, and 70% greater with overhead cover such as treebranches. Survival of emergents was greater in microsites with grass cover(90% survival) compared to without ground cover (44% survival),but lowest in microsites surrounded, but not covered, by grass (19%).From 1994–1999, natural seedling emergence and survival washighest in 1995 (80% survival of 221 P.engelmannii,and 100% of seven A. lasiocarpa, in a sample areaof432 m²), when the smallest mean difference in dailymaximum and minimum temperatures occurred, and lowest in 1994 (30% ofseven P. engelmannii), when above-averagetemperatures were accompanied by low rainfall and clear skies. The growthseasons of 1994 and 1995 had among the lowest and highest precipitation of theprevious 30-year period, respectively. In an artificial seedingexperiment, less than 20% of seedlings of both species survived theirfirst complete year of growth. In the autumn of the second year, almost25% additional mortality was observed in the remaining experimentalseedlings when they were exposed to clear, cold skies without the normalprotection of snowcover. Both spatial and temporal patterns of seedlingsurvivalsuggest that exposure to high sunlight may exacerbate low-temperatureandwater stress in young conifer seedlings, inhibiting their establishment in thisalpine treeline.     

Effects of microsite disturbances and herbivory on seedling performance in the perennial herb <Emphasis Type="Italic">Helleborus foetidus</Emphasis> (Ranunculaceae)

The impact of small scale disturbances on the early seedling performance components of Helleborus foetidus (Ranunculaceae) was studied through a transplant experiment. The aims of this study were: (i) to determine if the herbivory pattern depends on microsite disturbance, by the analysis of two of its components, seedling encounter (the probability of at least one seedling being harmed) and seedling exploitation (the proportion of seedling tissue removed once encountered); (ii) to test if seedlings of H. foetidus in disturbed microsites will survive in a greater proportion than seedlings in undisturbed microsites; (iii) to investigate if seedling survival is correlated with the degree of herbivory. Microsite disturbances had a large effect on the herbivory pattern. Seedlings growing in undisturbed vegetation had a 2-fold higher likelihood of being grazed and suffered 1.38-fold higher damage than those growing in disturbed plots. At the end of this experiment, after fourteen months, only 10.4% of the seedlings transplanted were still alive due to seedling desiccation, but no differences on seedling survival were found between disturbed and undisturbed plots. The effect of herbivory and the interactive effect of herbivory and disturbance on seedling survival only reached statistical significance dependent upon site. We concluded that although small scale disturbances had a large impact on herbivory patterns; they had only a minor role in the early seedling survival of H. foetidus. Only locally, small scale disturbances showed an effect on seedling survival through herbivory. Abiotic factors like summer drought and spatial variations determined the early survival of H. foetidus seedlings to a major extent.