Ramet Density Research Articles

Rhizome sectioning and fertilization increase the productivity of cloudberry in natural peatlands

The effects of combining rhizome sectioning and in-depth mineral fertilization on cloudberry (Rubus chamaemorus L.) growth and production in a natural peat bog were determined over four growing seasons. Observed enhancements of growth and fruit production included: increases in the density of ramets and leaves from the second year; increased flower density from the third year; and increased fruit density from the fourth year. No effect on fruit size was observed. These results demonstrate that it is possible to stimulate growth and productivity of cloudberry by cultural treatments applied to natural mires.Key words: Wild berry, Rubus chamaemorus, fruit yield, rhizome sectioning, mineral fertilization, cloudberry

The response of the pseudoannual species Trientalis europaea L. to forest gap dynamics in a near-natural spruce forest

The objective of this study was to test relationships between gap size, gap age and performance of the pseudoannual forest herb layer species Trientalis europaea. We also tested for a potential covariation of performance variables with light as the putative driving factor of gap size and gap age effects. The study took place in the core zone of the Harz National Park in a near-natural spruce forest at Mt. Brocken, Germany. We established 70 randomly distributed plots of three different gap age classes (<15 years, >15 and <60 years, >60 years) and undisturbed forest stands. We recorded growth variables of Trientalis (e.g. height of ramets, number of flowers and fruits per ramet and number of daughter tubers per ramet) and measured PPFD (photosynthetic photon flux density). Gap sizes were assessed with GPS, ranged from 131 m 2 to 16400 m 2 and were independent from gap age. Population density of T. europaea was neither affected by gap size nor by gap age. However, gap age had an effect on the fitness of Trientalis ramets. In gaps, the species produced a higher number of flowers and fruits. In general, the average proportion of flowering and fruiting was very low, with 13% and 4%, respectively. In contrast, light intensity had a significant positive effect on ramet density, while the number of daughter tubers differed between 0.8 and 0.5 tubers per individual under lowest light and full light intensity, respectively. The general conclusion is that gap size had no effects on the performance of T. europaea, while gap age had both direct effects and indirect effects mediated by light.

Two-scale modelling of Saponaria bellidifolia Sm. (Caryophyllaceae) abundance on limestone outcrops from its northern range periphery (Southeastern Carpathians)

We modelled the effect of habitat heter- ogeneity on the abundance of the submediterranean Saponaria bellidifolia, a red list species in Romania. The study was designed at two scales: 100 and 0.5 m 2 . At larger scale, generalized additive models and canonical correspondence analysis were used to model the density of ramets, whereas at microscale, binomial logistic regression was employed to model the species' occurrence. S. bellidifolia abundance responded sensitively to habitat type (classified as ''grassy'', ''rocky'' and ''scree''), rather than to microclimatic variables. At both scales, habitat type was the best predictor of ramet abundance, followed by slope and vegetation cover. At 0.5 m 2 , soil depth was also a good predictor of species occurrence. The data revealed that screes are the most suitable habitats for hosting relatively large populations of this rare species, because of occasional natural disturbances and presumably lower interspecific competition.

Effect of waterlogging on regeneration in the dwarf birch (Betula nana)

The aim of this study was to determine the effect of waterlogging on annual regeneration in the dwarf birch (Betula nana L.), and to determine which demographic factors are responsible for the absence of B. nana in the waterlogged parts of a peat-bog. Growth parameters and demographic data were collected from 2001 to 2003 at four sites in the Linje peat bog in the Chelmno Lake District in northern Poland. Three of the sites represented waterlogged open bog communities. The fourth site served as the reference. The density of first-year ramets, annual shoot growth, and the proportion of flowering shoots was significantly lower at the waterlogged sites than at the reference site. No seedlings were found during the course of the study, even though flowering and fruit production took place at the reference site. The flowering shoots were mainly between nine and thirteen years old. Ramets in this age range were found only at the reference site due to high mortality in mature ramets at the waterlogged sites. Mean ramet height was significantly lower at the waterlogged sites than at the reference site, and was strongly correlated with mean ramet age. The distribution of B. nana in peat bogs is limited primarily by two demographic factors: the high mortality rate in older ramets, and the low rate of vegetative reproduction. Another contribution factor is that the proportion of older fertile ramets is low. The effects of fluctuations in the water table on distribution of B. nana are also discussed.

Does harvesting sustain plant diversity in central Mexican wetlands?

In Central Mexico, wetland plants are harvested for weaving, fodder, and fertilizer. To test whether harvesting alters plant diversity, we compared the effects of harvesting all vegetation once, follow-up harvesting of Typha domingensis Pers. one or three more times, and a non-harvested control, using two sites differing in water depth in an annually burned wetland near Morelia, Mexico. After one year, harvesting treatments increased species richness at both the plot (14-m2) and wetland scales, increased the Shannon diversity index at the plot and subplot (1-m2) scales, and changed plant community composition (measured by Bray-Curtis distance) relative to control plots. Response among harvesting treatments was similar, and increased Typha harvesting did not have additive effects on Typha or on community composition. Grasses and short forbs (< 0.5-m tall) significantly increased in importance value in harvested plots, as did five individual forb species that were capable of vegetative spread. Uncommon species were significantly more likely to be found only in harvested plots than only in control plots, and new species (not initially present at the site) tended to recruit in harvested plots. Most new species were perennials that could likely tolerate additional harvesting. All harvesting treatments reduced Typha height, density, and rhizome starch reserves after five months, and responses were significantly affected by site, water depth, flowering ramet density, and pre-treatment values. Typha recovered in all harvested plots after one year, even when harvested four times, although flowering-ramet density declined in the wetter site. Community composition was more highly correlated with water depth and litter cover than with harvesting in an NMS ordination including both sites. Within sites, harvesting, light availability, leaf area index, and litter cover correlated similarly with variation in community composition. Given that our treatments reflect a subset of actual local management practices, harvesting could provide a sustainable and economically attractive management strategy for biodiversity conservation in this system, while the cessation of harvesting could lead to species loss.

Ramet demography and ecological attributes of the perennial river corridor plant Cnidium dubium (Schkuhr) Thell. (Apiaceae)

Cnidium dubium is a highly endangered, perennial river corridor plant in Central Europe. Here we summarise a 4-year field study of its ramet demography, growing in two different flood meadows in the Lower Havel River Valley in northeastern Germany, emphasising the influence of site-specific abiotic factors, mowing and the summer flood of 2002. We recorded 1658 ramets at densities between 2 and 180 ramets m −2, which varied over area and time, indicating a high small-scale turnover of this species within grasslands. The half-life of the different ramet cohorts was estimated to range from 0.1 to 1.3 years, but single ramets of more than 4 years old were also found. Ramets are monocarpic: they propagate only once in their life. The highest probability of a vegetative ramet becoming a flowering stalk was 11.6%. Ramet density, cumulative number of newly produced ramets per year, leaf length, leaf age, reproductive growth and recovery from the summer flood of 2002 varied, depending on site. C. dubium had a higher performance at sites in intermediate zones of the overall flooding gradient compared with higher zones. Flooding stress is the controlling factor for the lower limit of its performance, while there was no evidence for an upper limit controlled by summer drought. Mowing slightly increased the cumulative number of leaves per ramet, but it decreased the size of new leaves and also the summer above-ground biomass per ramet. As a consequence, the most marked effect was that fewer ramets entered the generative life stage, but the plants were able to keep their overall reproductive performance on a level independent of mowing. Knowledge of basic growth patterns of C. dubium in natural populations could help to explain the species rareness, and may also suggest appropriate strategies to manage its survival.

Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest

The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy conditions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to different canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and cluster distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets’ biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F. nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. Moreover, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heterogeneous light environments.

Ecological responses of Primula nutans to centimeter‐scale topographic and environmental variability in an alpine wetland

AbstractPrimula nutans Georgi is a herbaceous species broadly distributed in wetlands on the Qinghai–Tibetan Plateau. These wetlands are often spatially highly heterogeneous because of their hummock‐and‐hollow microtopography. To address how P. nutans can be so broadly distributed on these wetlands, we examined ramet distribution, abundance, and growth performance in environmentally variable microsites at the centimeter scale. P. nutans showed significantly higher ramet density on the south‐orientated microsites than on the north‐orientated microsites. With increasing relative microsite elevation on the hummocks, ramet density increased significantly, but individual leaf area decreased significantly. Principle component analysis and multiple regression analysis indicated that microsites at higher elevations tend to have higher light availability and higher ramet density. The study suggests that P. nutans shows high plasticity in the distribution, abundance, and growth performance in response to the microtopography at the centimeter scale, which may contribute to the broad distribution of the species in the hummock‐and‐hollow wetlands in harsh alpine environments on the Tibetan Plateau.

Evaluation of the field impact of an adventitious herbivore on an invasive plant, yellow toadflax, in Colorado, USA

Although some introduced plants arrive into their new range without their generalist and specialist herbivores, for others, their herbivores arrive prior to, with, or after the introduction of the plant, reestablishing the link between natural enemies and invaders in the introduced range. Research documenting the effects of adventitiously introduced herbivores on their target plants in the introduced range, and the mechanisms by which those effects occur, can provide insight into potential biological weed control. We studied the effects of an accidentally introduced beetle Brachypterolus pulicarius on the growth and reproduction of its host, the invasive plant Linaria vulgaris (yellow toadflax), growing under field conditions across multiple years and sites in western Colorado, USA. We found that feeding by B. pulicarius on L. vulgaris was variable among 3 years (2002–2004) and across eight local sites. Part of the variation in damage was explained by ramet density; sites with greater ramet density experienced a higher proportion of damage. In an observational study across 2 years, damage was positively correlated with estimates of sexual reproduction, ramet growth, and clonal shoot production. However, opposite trends were observed in an experiment; damage by B. pulicarius decreased estimates of sexual reproduction. Differences between the results of the observational and experimental studies were likely driven by selective feeding by B. pulicarius on larger ramets. Nonetheless, the ability of B. pulicarius to control established L. vulgaris population growth remains uncertain under the environmental conditions we studied. In both the observational and experimental study, B. pulicarius did not affect L. vulgaris survival, and we found no evidence that established L. vulgaris populations were seed limited, suggesting that reductions in seeds may not translate into demographic changes in heavily infested populations. Interactions among insect foraging behavior, individual plant responses to damage, and the demographic consequences of seed input may help to explain the varying degrees to which herbivores affect plants and populations in this and other systems.

MULTIPLE DISTURBANCES ACCELERATE CLONAL GROWTH IN A POTENTIALLY MONODOMINANT BAMBOO

Organisms capable of rapid clonal growth sometimes monopolize newly freed space and resources. We hypothesize that sequential disturbances might change short-term clonal demography of these organisms in ways that promote formation of monotypic stands. We examined this hypothesis by studying the clonal response of Arundinaria gigantea (giant cane, a bamboo) to windstorm and fire. We studied giant cane growing in both a large tornado-blowdown gap and under forest canopy, in burned and unburned plots, using a split-block design. We measured density of giant cane ramets (culms) and calculated finite rates of increase (lamda) for populations of ramets over three years. Ramet density nearly doubled in stands subjected to both windstorm and fire; the high ramet densities that resulted could inhibit growth in other plants. In comparison, ramet density increased more slowly after windstorm alone, decreased after fire alone, and remained in stasis in controls. We predict that small, sparse stands of giant cane could spread and amalgamate to form dense, monotypic stands (called "canebrakes") that might influence fire return intervals and act as an alternative state to bottomland forest. Other clonal species may similarly form monotypic stands following successive disturbances via rapid clonal growth.

Radial Growth and Ring Formation Process in Clonal PlantEriophorum angustifoliumon Post-Mined Peatland in the Šumava Mts., Czech Republic

Eriophorum angustifolium (Cyperaceae) is a pioneer clonal sedge colonizing bare peat surface of harvested peatlands in central Europe. It forms circular patches of densely aggregated ramets, followed by central die-back and ring formation as circles develop. This study experimentally tested the importance of inter-ramet competition, interference with litter, soil nutrient depletion, and architectural constraints for radial clonal spread and ring formation process. Effects of fertilization, litter addition and competition of neighbor ramets on growth and survival of tillers transplanted into four distinct zones within individual circle were detected only in the first zone (green band) with high ramet density. This suggested that both above-ground competition for light and below-ground competition for soil nutrients can play an important role in population dynamics of E. angustifolium. A field study of clonal growth has shown that rhizomes orient preferentially towards the periphery. As the old ramets within the interior of the tussock died due to senescence and inter-ramet competition, disproportionately more ramets are recruited on the periphery, which reduce axillary bud availability within the interior of the tussocks. We also explored the soil nutrient content (NH4-N, NO3-N and PO4-P). High values of NO3-N outside the circles and low values inside patches might partly explain why Eriophorum rhizomes are oriented into open space. High accumulation of phosphorus (PO4-P) in the centre of the patches was attributed to the leaching of phosphorus from decaying Eriophorum plants. We suggest that processes of nutrient foraging and utilization are responsible for development of circular patches of E. angustifolium in the harsh environment of cut-away peatland.

The status of &lt;i&gt;Sarcopoterium spinosum&lt;/i&gt; (Rosaceae) at the western periphery of its range: Ecological constraints lead to conservation concerns

This paper considers the demography, ecology, and conservation perspectives of the western populations of the clonal dwarf-shrub Sarcopoteriumspinosum (L.) Spach. While in the Middle East the species dominates large areas, westwards it is less common and many populations have become extinct since the late 19th century. We highlight the ecological limitations and their implications for the conservation of the species at the periphery of its range. In southern Italy we studied the demographic traits of two populations and the age spatial structure of a third. Ramet density and lifespan appear to be lower than in the Middle East. The floristic analysis of the stands studied and a climatic analysis over the whole range of S.�spinosum provide a key for the interpretation of such differences. Given that summer drought stress decreases westwards, both the sprouting vigor and the ecological space available for S.�spinosum become limited by increasing competition. This makes the populations more likely to become extinct in changing landscapes, as revealed by the decreasing extent of occurrence and area of occupancy due to habitat loss. Although S.�spino- sum is not at risk in the Middle East, at the western border of its range it qualifies as an endangered species.

Patch structure and ramet demography of the clonal tree, Asimina triloba, under gap and closed-canopy

Clonal understory trees develop into patches of interconnected and genetically identical ramets that have the potential to persist for decades or centuries. These patches develop beneath forest canopies that are structurally heterogeneous in space and time. Canopy heterogeneity, in turn, is responsible for the highly variable understory light environment that is typically associated with deciduous forests. We investigated what aspects of patch structure (density, size structure, and reproductive frequency of ramets) of the clonal understory tree, Asimina triloba, were correlated with forest canopy conditions. Specifically, we compared A. triloba patches located beneath closed canopies and canopy gaps. We also conducted a three-year demographic study of individual ramets within patches distributed across a light gradient. The closed canopy-gap comparison demonstrated that the patches of A. triloba had a higher frequency of large and flowering ramets in gaps compared to closed-canopy stands, but total ramet density was lower in gaps than in closed canopy stands. In the demographic study, individual ramet growth was positively correlated with light availability, although the pattern was not consistent for all years. Neither ramet recruitment nor mortality was correlated with light conditions. Our results indicate that the structure of A. triloba patches was influenced by canopy condition, but does not necessarily depend on the responses of ramets to current light conditions. The lack of differences in ramet recruitment and mortality under varying canopy conditions is likely to be a primary reason for the long-term expansion and persistence of the patches. The primary benefit of a positive growth response to increasing light is the transition of relatively small ramets into flowering ramets within a short period of time.

VARIATION IN RESPONSES OF LATE-SERAL HERBS TO DISTURBANCE AND ENVIRONMENTAL STRESS

Clonal herbs that attain maximum development in late-seral forest are often assumed to have similar responses to disturbance and to be functionally equivalent. However, little is known about the demographic or physiological responses of these plants to disturbance or to the altered conditions of the post-disturbance environment. Following harvest of a mature coniferous forest, we compared abundance, demographic changes, and physiological acclimation of three clonal herbs (Asarum caudatum, Clintonia uniflora, and Pyrola picta) that differ in belowground morphology and leaf longevity. We measured ramet density, leaf area, and demographic variables (survival, clonal growth, flowering, and seedling establishment) before and for two years after harvest, and in adjacent undisturbed forest. Acclimation to increased solar radiation was assessed two years after harvest by measuring leaf mass per unit area (LMA) and chlorophyll a:b ratios of leaves produced in the current year. Although initial declines in abundance were similar, demographic responses indicate that patterns of recovery varied greatly among species. Two years after logging, ramet survival and clonal growth (production of new ramets) of Clintonia were greater in the harvest area than in the forest. Asarum had lower survival in the harvest area, but greater clonal growth, and Pyrola showed no difference in either survival or growth between environments. Only Asarum produced seedlings, although their survival was low in the harvest area. All species had higher LMA in the harvest area, but only Clintonia (with annual leaves) had a higher chlorophyll a:b ratio, suggesting the greatest potential for acclimation to increased light. Our results demonstrate that forest herbs with greater rhizome plasticity and shorter leaf duration have greater potential to acclimate after disturbance than those with rigid architectures and persistent leaves. Thus, species with comparable successional roles can vary substantially in their demographic and physiological responses to disturbance, with potential consequences for long-term recovery.

Effects of browsing on recruitment and mortality of European aspen (Populus tremula L.)

High browsing pressure may reduce or halt regeneration of browsing-prone tree species. In this study, exclosures were used to quantify the effects of browsing on vital demographic parameters in European aspen (Populus tremula L.) stands. The experiment focused on stands located in managed forest landscape with densities of 0.85 moose (Alces alces L.)km−2. Other herbivores such as mountain hare (Lepus timidus L.) were present at densities common for this region and roe deer (Capreolus capreolus L.) at low densities. Recruitment and mortality of aspen ramets were recorded before and after the growing season, and over four consecutive years. Seven to 19% of the ramets within browsing reach were browsed annually. No differences were found in recruitment or mortality attributable to browsing per se. Instead, there were large among-stand variability in vital population parameters unrelated to browsing incidence. In general, aspen stands with high ramet density showed higher recruitment rates than stands with low ramet density. There was substantial variability in self-replacement capability unrelated to ecological factors. The results suggest that more attention should be given to finding and managing stands with high capacity for suckering, as these stands may have a disproportionate impact on aspen recruitment at the landscape scale.

Threatened Peripheral Populations in Context: Geographical Variation in Population Frequency and Size and Sexual Reproduction in a Clonal Woody Shrub

Geographically peripheral populations of widespread species are often the focus of conservation because they are locally rare within political jurisdictions. Yet the ecology and genetics of these populations are rarely evaluated in a broader geographic context. Most expectations concerning the ecology and evolution of peripheral populations derive from the abundant-center model, which predicts that peripheral populations should be less frequent, smaller, less dense, and have a lower reproductive rate than central populations. We tested these predictions and in doing so evaluated the conservation value of peripheral populations for the clonal shrub Vaccinium stamineum L. (Ericaceae, deerberry), which is listed as threatened in Canada. Based on 51 populations sampled from the center to the northern range limits over 2 years, population frequency and size declined toward the range limit, but ramet density increased. Sexual reproductive output varied widely among populations and between years, with many populations producing very few seeds, but did not decline toward range margins. In fact seed mass increased steadily toward range limit, and this was associated with faster germination and seedling growth, which may be adaptive in seasonal northern environments. Our results did not support the prediction that clonal reproduction is more prevalent in peripheral populations or that it contributed antagonistically to the wide variation in seed production. Peripheral populations of V. stamineum are as productive as central populations and may be locally adapted to northern environments. This emphasizes the importance of a broad geographical perspective for evaluating the ecology, evolution, and conservation of peripheral populations.

Geographical variation in frond size, rootstock density, and sexual reproduction inMatteuccia struthiopterispopulations in Norway

AbstractThe main aim of this study was to quantify differences in the structure of denseMatteuccia struthiopteris(L) Todaro among geographically separated populations in Norway, and to relate these differences to variation in environmental variables. A stratified random sampling procedure was applied to select 4 m2quadrats restricted to populations dominated (&gt;75% cover) by the fern. This dimorphic fern can produce three types of leaves (fronds): fertile (sporophyll) and sterile (trophophyll), and in addition also intermediate fronds. In all, 178 populations from different parts of the country were analysed, and in total 9,346 rootstocks were measured. Fifteen percent of the rootstocks were fertile, each producing from one to eight sporophylls, and 2.0% had developed intermediate fronds. Rootstocks with five or more sporophylls were mainly found in Southern Norway. Within the plots, there were major variations in density (10–112), number of small (new) ramets (0–58), total number of sporophylls (0–94) and estimated total fern wet‐weight (0.33–22.8 kg). All new additions to the populations were connected to a mother plant (ramets), and none were of sexual origin (genets). The main environmental variables associated with the variation in population structure were mean July temperature (highly correlated with altitude and latitude), mean precipitation and density of the canopy cover. The density of small ramets and number of intermediate fronds increased from south to north. The latitudinal variation in sporophyll development and maximum trophophyll height was best explained by a unimodal model. There were statistically significant differences in the population structure between the eight studied geographic areas in Norway.

Clonal structure of Elytrigia atherica along different successional stages of a salt marsh

Elytrigia atherica is a tall clonal grass species typical of higher salt marshes, but is gradually invading to the lower marshes. At young successional stages of a salt marsh, E. atherica is found sparsely dispersed in small groups of ramets. These patches increase in size and ramet density over time, eventually forming extensive swards as succession proceeds. This study investigates the change in the clonal diversity of E. atherica stands during colonization as a result of its reproductive strategy. Clonal diversities of differently sized patches of E. atherica were investigated on two lower salt-marsh sites of different age, 25 years and 35 years, respectively. Microsatellite fingerprint patterns were used to determine genet identities and to estimate relatedness and genetic differentiation between the sites, between patches within sites and within patches. The majority of the patches on both sites contained more than one genet. On the older site, the clonal diversity was higher than on the younger site. However, the clonal diversity tended to decrease with increasing patch size. Low genetic differentiation was found between the two sites, indicating habitat differentiation, whereas differentiation between patches within sites was high. It is reasoned that different environmental conditions could have resulted in different clonal structures: On an older marsh, the increase of successful seedling recruitment, due to more suitable environmental conditions, leads to an increase in clonal diversity. Over time, with increasing ramet density, intraspecific competition is likely to increase, resulting in a decrease of clonal diversity.

Internal heterogeneity of ramet and flower densities of Typha angustifolia near the boundary of the stand

Fifteen relatively narrow stands of Typha angustifolia, including both stands bounded to open water and ridged boundary, were investigated in terms of the distributions of ramet density as well as the fraction of flowering ramets. The ramet density, fraction of flowering ramet and morphological characteristics of ramets were measured continuously from one side of the stand to the other, together with depth and the nutrient concentration of pore water. The rhizome biomass was also measured at the periphery and at the center of the stand. The results indicated that the shoot density declined gradually from the edge to the center, however, the fraction of the flowering ramets were significantly higher at the edge while it rapidly declined away from the edge. Statistical analyses indicate that the ramet density depends on either depth or nutrient level of the pore water; however, the distance from the edge was the most probable factor to reduce the fraction of the flowering ramets. The rhizome biomass was mostly larger at the periphery than at the center, likely because the accumulation of the horizontal rhizomes as extension was blocked by the boundary. High flowering fraction near the periphery was probably induced by the hormonal change caused by the blockage. However, a large fraction of ramets flowered simultaneously with large rhizome biomass at the periphery, likely because of the preferable light climates compared with inside the stand.

Aspen demographics in relation to spatial context and ungulate browsing: Implications for conservation and forest management

Aspen has a high conservation value and has been targeted as a priority species in sustainable forest management. However, sustained and intense browsing pressure by ungulates has raised concerns about the recruitment of aspen to mature growth stages. Here we investigate the influence of ungulate browsing on the demographic structure of European aspen populations in two managed boreal forest areas in Sweden with contrasting amounts of aspen and high densities of ungulates. The contrast in size distribution of ramets in the two areas was striking. The aspen-poor environment contained a much lower density of small-sized ramets than the aspen-rich environment. Ramet abundance was strongly associated with the abundance of ramets in the smaller size classes in both areas, reflecting demographic inertia in these populations. Ungulate use was most strongly associated with abundance of medium-sized aspen saplings growing in forest interiors. Otherwise, use of ungulates was only weakly correlated with aspen population parameters. The strong demographic momentum exhibited by these populations suggests that aspen conservation and management should pay more attention to additional factors than browsing to promote ramet establishment. Large variation in abundance of ramets in the smallest size-class shows that some aspen stands have a disproportionate effect on aspen recruitment, particularly in areas where aspen is less abundant. Identifying aspen stands with a large capacity for self regeneration may be the key to meet aspen conservation goals.