Fluxes of energy,water, and carbon dioxide from mountain ecosystems at Niwot Ridge,Colorado

Background: The eddy covariance (EC) technique provides a direct measure of water vapour and carbon dioxide fluxes between ecosystems and the atmosphere.

Aims: This review article highlights the findings of various studies that have integrated EC observations into basic meteorological, hydrological and ecological research questions in two ecosystems near Niwot Ridge, Colorado, and synthesises these studies into a catchment-scale model of water and carbon cycling, within the context of regional disturbance and environmental change.

Methods: EC was implemented continuously over subalpine forest and alpine tundra vegetation at Niwot Ridge, and resulting data were compared with discrete measurements and modelling studies.

Results: Sensible heat fluxes were generally in excess of latent heat fluxes, indicating that the forest and tundra ecosystems were moisture limited. Snow cover regulated the annual sum of primary productivity in the forest, and beneath-snow respiration represented a significant portion of ecosystem respiration at both locations.

Conclusions: Changes in the magnitude, timing, or spatial distribution of snow are likely to have the greatest impact on ecological processes in these semi-arid mountain catchments, but possibly in compensatory ways. Ultimately, the degree to which net carbon losses from alpine tundra offset forest carbon sequestration will determine the future magnitude of the Western United States carbon sink.     

Conifer seedling recruitment across a gradient from forest to alpine tundra: effects of species,provenance, and site

Background: Seedling germination and survival is a critical control on forest ecosystem boundaries, such as at the alpine–treeline ecotone. In addition, while it is known that species respond individualistically to the same suite of environmental drivers, the potential additional effect of local adaptation on seedling success has not been evaluated.

Aims: To determine whether local adaptation may influence the position and movement of forest ecosystem boundaries, we quantified conifer seedling recruitment in common gardens across a subalpine forest to alpine tundra gradient at Niwot Ridge, Colorado, USA.

Methods: We studied Pinus flexilis and Picea engelmannii grown from seed collected locally at High (3400 m a.s.l.) and Low (3060 m a.s.l.) elevations. We monitored emergence and survival of seeds sown directly into plots and survival of seedlings germinated indoors and transplanted after snowmelt.

Results: Emergence and survival through the first growing season was greater for P. flexilis than P. engelmannii and for Low compared with High provenances. Yet survival through the second growing season was similar for both species and provenances. Seedling emergence and survival tended to be greatest in the subalpine forest and lowest in the alpine tundra. Survival was greater for transplants than for field-germinated seedlings.

Conclusions: These results suggest that survival through the first few weeks is critical to the establishment of natural germinants. In addition, even small distances between seed sources can have a significant effect on early demographic performance – a factor that has rarely been considered in previous studies of tree recruitment and species range shifts.     

Cryosphere: ice on Niwot Ridge and in the Green Lakes Valley,Colorado Front Range

Background: Subsurface ice preserved as ice lenses and within rock glaciers as well as glacial and lake ice provides sensitive indicators of climate change and serve as a late-season source of meltwater.

Aims: We synthesise the results of geomorphological, geophysical and geochemical studies during the period of 1995–2014, building on a long history of earlier work focused on ice and permafrost studies on Niwot Ridge and the adjacent Green Lakes Valley (GLV), which is part of the Niwot Ridge Long Term Ecological Research Site.

Methods: These studies are discussed in the context of how bodies of ice and rock glaciers reflect changing local climate. We review recent results from geophysical investigations (resistivity, seismic refraction and ground-penetrating radar) of the shallow subsurface, ongoing monitoring of the Arikaree Glacier, three rock glaciers and lake ice in the GLV, and interpretations of how subsurface ice melt regulates the flow and chemistry of alpine surface water after seasonal snowfields melt.

Results and conclusions: Permafrost conditions reported from Niwot Ridge in the 1970s are generally absent today, but ice lenses form and melt seasonally. Ice is present permanently within the Green Lakes 5 rock glacier and at nearby favourable sites. The Arikaree Glacier has shown a marked decline in cumulative mass balance during the past 12 years after a 30-year period when net mass balance was ca. 0. Duration of seasonal lake ice increases with elevation in GLV, but duration has decreased at all seven lakes that have been monitored during the last three decades. This decrease has been most marked at the lowest elevation where it amounted to a reduction of about 1 d year^?1 and least at Green Lake 5 where the loss has been at a rate of 0.5 d year^?1. Surface temperature measurements from rock glaciers have not shown strong trends during the past 15 years. It has been suggested that almost all of the 2.5-mm year^?1 increase in stream discharge from the upper GLV in September and October has been derived from melting of subsurface ice.     

Tree recruitment at the treeline across the Continental Divide in the Northern Rocky Mountains,USA: the role of spring snow and autumn climate

ABSTRACT

Background: Topoclimate can influence tree establishment within treeline ecotones. Yet much less is known about how regional topography, such as the Continental Divide, Rocky Mountains, mediates the role of climate in governing treeline dynamics.

Aims: To utilise the Continental Divide to test whether contrasts in growing-season moisture regimes to the west (summer-dry) and east (summer-wet) impact the spatio-temporal patterns of tree establishment and rates of treeline advance in the Northern Rocky Mountains.

Methods: We sampled trees at sites on north- and south-facing slopes, west and east of the Continental Divide. We used dendroecological techniques to reconstruct patterns of tree establishment. Age-structure data were quantitatively compared with climate to evaluate possible mechanistic linkages.

Results: Across all sites, 96% of trees established after 1950. There was a treeline advance (range = 39–140 m) accompanied by increases in tree density. Significantly more trees established during wet springs on both sides of the Divide.

Conclusions: Overall, snow duration in spring and autumn temperatures appear to influence patterns of tree recruitment at the treeline. Continued warming will likely amplify the role of autumn climate in regulating tree establishment throughout treeline ecotones in the Northern Rocky Mountains, particularly west of the Divide where summer-dry conditions persist.     

The consequences of multiple resource shifts on the productivity and composition of alpine tundra communities: inferences from a long-term snow and nutrient manipulation experiment

Background: Gradients in the amounts and duration of snowpack and resulting soil moisture gradients have been associated with different plant communities across alpine landscapes.

Aims: The extent to which snow additions could alter plant community structure, both alone and in combination with nitrogen (N) and phosphorus (P) additions, provided an empirical assessment of the strength of these variables on structuring the plant communities of the alpine tundra at Niwot Ridge, Colorado Front Range.

Methods: A long-term snow fence was used to study vegetation changes in responses to snowpack, both alone and in conjunction with nutrient amendments, in plots established in dry and moist meadow communities in the alpine belt. Species richness, diversity, evenness and dissimilarity were evaluated after 20 years of treatments.

Results: Snow additions, alone, reduced species richness and altered species composition in dry meadow plots, but not in moist meadow; more plant species were found in the snow-impacted areas than in nearby controls. Changes in plant community structure to N and N + P additions were influenced by snow additions. Above-ground plant productivity in plots not naturally affected by snow accumulation was not increased, and the positive responses of plant species to nutrient additions were reduced by snow addition. Plant species showed individualistic responses to changes in snow and nutrients, and indirect evidence suggested that competitive interactions mediated responses. A Permanova analysis demonstrated that community dissimilarity was affected by snow, N, and P additions, but with these responses differing by community type for snow and N. Snow influenced community patterns generated by N, and finally, the communities impacted by N + P were significantly different than those affected by the individual nutrients.

Conclusions: These results show that changes in snow cover over a 20-year interval produce measureable changes in community composition that concurrently influence and are influenced by soil nutrient availability. Dry meadow communities exhibit more sensitivity to increases in snow cover whereas moist meadow communities appear more sensitive to N enrichment. This study shows that the dynamics of multiple limiting resources influence both the productivity and composition of alpine plant communities, with, species, life form, and functional traits mediating these responses.     

A comparison of water and carbon dioxide exchange at a windy alpine tundra and subalpine forest site near Niwot Ridge,Colorado

Eddy covariance measurements of the surface energy balance and carbon dioxide exchange above high-elevation (3,480 m above sea level) alpine tundra located near Niwot Ridge, Colorado, were compared to simultaneous measurements made over an adjacent subalpine forest over two summers and one winter, from June 9, 2007 to July 3, 2008. The surface energy balance closure at the alpine site averaged 71 and 91%, winter and summer, respectively, due to the high wind speeds, short turbulent flux footprint, and relatively flat ridge-top location of the measurement site. Throughout the year, the alpine site was cooler with higher relative humidity, and had a higher horizontal wind speed, especially in winter, compared to the forest site. Wind direction was persistently downslope at the alpine site (summer and winter, day and night), whereas upslope winds were common at the forest site during summer daytime periods. The latent and sensible heat fluxes were consistently larger in magnitude at the forest site, with the largest differences during summer. The horizontal advective flux of CO₂ at the alpine site averaged 6% of the net ecosystem exchange (NEE) during summer nights (5% during summer daytime), and was small in relation to the high wind speeds, relatively flat site, and weak sources of CO₂ upwind of the site. The magnitudes and diurnal behavior of the alpine NEE calculated using three methods; eddy-covariance, friction velocity filter, and with advection and storage calculations, gave similar results. The period of net CO₂ uptake (negative NEE) was 100 days at the alpine site with a net uptake of 16 g C m⁻², compared to 208 days at the forest site with a net uptake of 108 g C m⁻², with initiation of net uptake coinciding with air temperatures reaching +10°C. Winter respiration loss at the alpine site was 164 g C m⁻² over 271 days, compared to 52 g C m⁻² over 175 days at the forest site, with the initiation of net loss coinciding with air temperatures reaching −10°C at each site.     

Winter gas exchange between the atmosphere and snow-covered soils on Niwot Ridge,Colorado, USA

Background: There is a growing interest in understanding the gas exchange between the atmosphere and seasonally snow-covered regions, especially in light of projections that climate change will alter the timing and extent of seasonal snow cover. In snow-covered ecosystems, gas fluxes are due both to microbial activity in the snow-covered soils and to chemical and physical reactions with the various gases and/or dissolved constituents in the snowpack. Niwot Ridge, in the Colorado Rocky Mountains, has one of the most extensive sets of measurements of winter gas exchange globally.

Aims: Our goal was to examine the temporal patterns and environmental controls on Niwot Ridge of gas fluxes for gases with different sources and sinks.

Methods: Here, we review the concentrations and fluxes that have been measured for carbon dioxide, nitrous oxide, methane, nitrogen oxides, ozone, gaseous elemental mercury and volatile organic carbon compounds.

Results and Conclusions: We looked for similarities and differences among the gases, but in many cases, the origin, fate and controls of these fluxes still need to be determined. However, we believe that many of the biologically driven reactions are the result of exponential growth of a winter microbial community during the long period of stable environmental conditions under the seasonal snowpack.     

Linking small-scale topography with microclimate,plant species diversity and intra-specific trait variation in an alpine landscape

Background: Small-scale topographic complexity is a characteristic feature of alpine landscapes, with important effects on alpine plant distribution.

Aims: We investigated the links between small-scale topographic complexity and resultant microclimatic heterogeneity, vascular-plant species richness and beta diversity, and realised niche width and trait variation of some target species.

Methods: We recorded temperature and soil moisture within 10 sites (40 m × 40 m) of differing topographic complexity in alpine terrain at Finse, Norway (N 60° 36?, E 7° 33?). Plant species occurrence and traits of target species were recorded in 16 sample plots at each site.

Results: Sites differed significantly in microclimatic heterogeneity, and topographically rough sites were always more heterogeneous than flatter ones. Greater species richness and turnover was associated with greater microclimatic heterogeneity, and rough sites contained 15–55% more species than flatter ones. Plant species had on average wider realised niches when growing at rough sites. Individuals of Bistorta vivipara, but not those of Luzula spicata, tended to exhibit greater phenotypic variation at rough sites.

Conclusions: Rough alpine terrains create small-scale variation in microclimate, promoting species richness and beta diversity. In the event of climate change, small-scale microclimatic heterogeneity might allow plant species to escape from regional climate change by short-distance migration to local micro-refugia. This study suggests that the opportunity for such responses would be greater in topographically complex terrains.     

Delimitation of the alliances Caricion firmae (Seslerietalia albicantis) and Seslerion juncifoliae (Seslerietalia juncifoliae) in the southeastern Alps and Dinaric mountains

Abstract

A syntaxonomic and phytogeographic delimitation of the calcareous open sedge swards in the alpine belt of the Alps (Caricion firmae) and subalpine and alpine tussock grasslands in wind-exposed habitats (Seslerion juncifoliae) in the area of the south-eastern Alps and of the Dinaric mountains was performed. Analyses based on hierarchical classification, ordination and chorology clearly showed the distinction between the syntaxa: stands from the Liburnian karst (Mt Sne?nik – SW. Slovenia, Mts Risnjak and Snje?nik – NW. Croatia), Li?ka Plje?ivica and the Velebit mountains belong to the Dinaric alliance Seslerion juncifoliae, whereas stands from the Trnovski gozd plateau (W. Slovenia, north-westernmost part of the Dinaric mountains), although somewhat transitional between the two alliances, and stands from the Alps, were classified in the alliance Caricion firmae. The alliance Seslerion juncifoliae of the Dinaric mountains vicariates Caricion firmae of the Alps.     

Spatial patterns of total and available N and P at alpine treeline

Background and aims

Vegetation can have direct and indirect effects on soil nutrients. To test the effects of trees on soils, we examined the patterns of soil nutrients and nutrient ratios at two spatial scales: at sites spanning the alpine tundra/subalpine forest ecotone (ecotone scale), and beneath and beyond individual tree canopies within the transitional krummholz zone (tree scale).

Methods

Soils were collected and analyzed for total carbon (C), nitrogen (N), and phosphorus (P) as well as available N and P on Niwot Ridge in the Colorado Rocky Mountains.

Results

Total C, N, and P were higher in the krummholz zone than the forest or tundra. Available P was also greatest in the krummholz zone while available N increased from the forest to the tundra. Throughout the krummholz zone, total soil nutrients and available P were higher downwind compared to upwind of trees.

Conclusions

The krummholz zone in general, and downwind of krummholz trees in particular, are zones of nutrient accumulation. This pattern indicates that the indirect effects of trees on soils are more important than the direct effects. The higher N:P ratios in the tundra suggest nutrient dynamics differ from the lower elevation sites. We propose that evaluating soil N and P simultaneously in soils may provide a robust assay of ecosystem nutrient limitation.     

The ‘teflon basin’ myth: hydrology and hydrochemistry of a seasonally snow-covered catchment

Background: Snow and ice melt provide sensitive indicators of climate change and serve as the primary source of stream flow in alpine basins.

Aims: We synthesise the results of hydrological and hydrochemical studies during the period 1995–2014, building on a long history of earlier work focused on snow and water on Niwot Ridge and the adjacent Green Lakes Valley (GLV), which is part of the Niwot Ridge Long Term Ecological Research site (NWT LTER).

Methods: These studies are discussed in the context of how snow, snowmelt and runoff reflect changing local climate. We review recent results of snow, snowmelt, hydrology and hydrochemistry from the plot to the basin scale, utilising new tools such as continuous global positioning system (GPS) measurements of snow depth, along with remotely-sensed measurements of snow-covered area and melt, combined with long-term measurements of snow properties, discharge and solute and isotopic content of water.

Results and Conclusions: Surface–groundwater interactions are important components of water quantity and quality in alpine basins. Some or most snowmelt infiltrates underlying soils and bedrock, transporting soil and bedrock products to surface waters. Infiltrating snowmelt, along with increased melt of stored ice, increases the hydrologic connectivity between the terrestrial and aquatic systems. Alpine basins are being impacted by increases in atmospheric nitrogen deposition, which has caused changes in soil microbial processes such as nitrification. Nitrate, dissolved organic carbon and dissolved organic nitrogen are thus flushed from soils and talus to streams. Our long-term results show that alpine catchments, such as Green Lake 4 at NWT LTER+, have the greatest sensitivity and least resilience to climate warming, with any warming leading to increased water yields.     

Positive effects of an extremely hot summer on propagule rain in upper alpine to subnival habitats of the Central Eastern Alps

Background: Propagule production and dispersal largely determine the distribution and potential migration ability of alpine plant species. Variation in reproductive success caused by year-to-year variation in climate may critically influence these processes.

Aims: To obtain estimates for the propagule rain in high-alpine plant communities and detect potential dispersal events from lower elevations.

Methods: The magnitude and composition of the propagule rain was studied in different plant communities along an elevation gradient from the upper alpine to subnival zone. Propagules were trapped at eight elevations from 2760 to 3070 m a.s.l. for three years from July to September 2003–2005. Vascular plant species and their cover were recorded in an area with a radius of 10 m surrounding the traps.

Results: A five- to 10-fold higher propagule rain was observed in 2003, a year with an exceptionally hot summer, compared to 2004 and 2005. Propagule and species numbers varied highly among years and community types. Few propagules of non-local origin were recorded in any year.

Conclusions: Extremely hot summers are likely to greatly magnify the propagule rain size of species in alpine habitats. Such ‘mast years’ may contribute to enhanced and accelerated vegetation changes in alpine habitats in the absence of limiting factors.     

Vegetation change at high elevation: scale dependence and interactive effects on Niwot Ridge

Background: High-elevation mountain systems may be particularly responsive to climate change.

Aims: Here we investigate how changes along elevation gradients in mountain systems can aid in predicting vegetation distributional changes in time, focusing on how changing climatic controls affect meso-scale transitions at the lower and upper boundaries of alpine vegetation (with forest and subnival zones, respectively) as well as micro-scale transitions among plant communities within the alpine belt. We focus on climate-related drivers, particularly in relation to climate change, but also consider how species interactions, dispersal and responses to disturbance may influence plant responses to these abiotic drivers.

Results: Empirical observations and experimental studies indicate that changing climatic controls influence both meso-scale transitions at the upper and lower boundaries of alpine vegetation and micro-scale transitions among plant communities within tundra. Micro-scale heterogeneity appears to buffer response in many cases, while interactions between climate and other changes may often accelerate change.

Conclusions: Interactions with microtopography and larger edaphic gradients have the capacity to both facilitate rapid changes and reinforce stability, and that these interactions will affect the responsiveness of vegetation to climate change at different spatial scales.     

Habitat use by singing voles and tundra voles in the southern Yukon

Summary We investigated how far competitive interactions influence the use of habitats and relative abundance of two species of Microtus in the southwestern Yukon. We worked in the ecotone between alpine tundra and subalpine shrub tundra where populations of singing voles (Microtus miurus) and tundra voles (M. oeconomus) overlap little.We removed tundra voles from shrub tundra on one live-trapping area to look at the effect on the contiguous population of singing voles in alpine tundra. The removal of tundra voles did not affect the distribution or relative abundance of singing voles. The spatial distribution of these species and their movements within habitats suggest that they have a strong habitat preference.Populations of small mammals in the area are extremely dynamic and the relative importance of competitive interactions may change as density varies. At present we have no evidence that competition affects habitat use in M. miurus.     

Influence of N2-fixing Trifolium on plant species composition and biomass production in alpine tundra

Alpine Trifolium species have high rates of symbiotic N₂-fixation which may influence the abundance and growth of plant species growing near them. The potential for facilitative effects on plant abundance and growth in dry meadow alpine tundra of Niwot Ridge, Colo., characterized by low resource availability, was investigated by measuring soil N, aboveground biomass production, and plant species composition in patches of Trifolium dasyphyllum and surrounding tundra. Extractable inorganic N was more than twofold greater and extractable P was 27% lower in Trifolium patches than in surrounding tundra. Aboveground production was twofold greater in Trifolium patches than in surrounding tundra. However, the difference was largely due to the production of T. dasyphyllum relative to the non-Trifolium component of biomass, which was not different between the Trifolium patches and surrounding tundra. In the Trifolium patches, the proportion of graminoid biomass was lower while the proportion of forb biomass was higher relative to surrounding tundra. Although the abundance of some species was positively associated with the presence of Trifolium, other species were less abundant, possibly due to increased competition for P and differential abilities of alpine species to respond to increased N availability. Trifolium may exert both facilitative and inhibitive effects on dry meadow alpine species and, in the process, substantially influence the spatial heterogeneity in community structure and primary production. Received: 14 October 1997 / Accepted: 2 February 1998     

Common montane birds are declining in northern Europe

Large‐scale multi‐species data on population changes of alpine or arctic species are largely lacking. At the same time, climate change has been argued to cause poleward and uphill range shifts and the concomitant predicted loss of habitat may have drastic effects on alpine and arctic species. Here we present a multi‐national bird indicator for the Fennoscandian mountain range in northern Europe (Finland, Sweden and Norway), based on 14 common species of montane tundra and subalpine birch forest. The data were collected at 262 alpine survey plots, mainly as a part of geographically representative national breeding bird monitoring schemes. The area sampled covers around 1/4 million km², spanning 10 degrees of latitude and 1600 km in a northeast–southwest direction. During 2002–2012, nine of the 14 bird species declined significantly in numbers, in parallel to higher summer temperatures and precipitation during this period compared to the preceding 40 yr. The population trends were largely parallel in the three countries and similar among montane tundra and subalpine birch forest species. Long‐distance migrants declined less on average than residents and short‐distance migrants. Some potential causes of the current decline of alpine birds are discussed, but since montane bird population sizes may show strong natural annual variation due to several factors, longer time series are needed to verify the observed population trends. The present Fennoscandian monitoring systems, which from 2010 onwards include more than 400 montane survey plots, have the capacity to deliver a robust bird indicator in the climate‐sensitive mountainous regions of northernmost Europe for conservation purposes.     

Warming and the dependence of limber pine (Pinus flexilis) establishment on summer soil moisture within and above its current elevation range

Continued changes in climate are projected to alter the geographic distributions of plant species, in part by affecting where individuals can establish from seed. We tested the hypothesis that warming promotes uphill redistribution of subalpine tree populations by reducing cold limitation at high elevation and enhancing drought stress at low elevation. We seeded limber pine (Pinus flexilis) into plots with combinations of infrared heating and water addition treatments, at sites positioned in lower subalpine forest, the treeline ecotone, and alpine tundra. In 2010, first-year seedlings were assessed for physiological performance and survival over the snow-free growing season. Seedlings emerged in midsummer, about 5–8 weeks after snowmelt. Low temperature was not observed to limit seedling photosynthesis or respiration between emergence and October, and thus experimental warming did not appear to reduce cold limitation at high elevation. Instead, gas exchange and water potential from all sites indicated a prevailing effect of summer moisture stress on photosynthesis and carbon balance. Infrared heaters raised soil growing degree days (base 5 °C, p < 0.001) and August–September mean soil temperature (p < 0.001). Despite marked differences in vegetation cover and meteorological conditions across sites, volumetric soil moisture content (θ) at 5–10 cm below 0.16 and 0.08 m³ m^?3 consistently corresponded with moderate and severe indications of drought stress in midday stem water potential, stomatal conductance, photosynthesis, and respiration. Seedling survival was greater in watered plots than in heated plots (p = 0.01), and negatively related to soil growing degree days and duration of exposure to θ < 0.08 m³ m^?3 in a stepwise linear regression model (p < 0.0001). We concluded that seasonal moisture stress and high soil surface temperature imposed a strong limitation to limber pine seedling establishment across a broad elevation gradient, including at treeline, and that these limitations are likely to be enhanced by further climate warming.     

Growth response of subalpine fir (Abies lasiocarpa) to climate in the Olympic Mountains, Washington, USA

Growth response of subalpine fir (Abies lasiocarpa) to climate was studied across its local geographical and elevation range in the Olympic Mountains, Washington. A dendroecological analysis of subalpine fir across a range of elevations (1350-1850 m) and annual precipitation (125-350 cm y^?1), was used to compare environmental factors affecting growth. Climate-growth relationships were explored using Pearson product-moment correlation coefficients; partial correlation analysis was used to assess relationships among site chronologies and climatic variables. Radial growth is negatively correlated with winter precipitation at high elevation and wet sites, but not at low and middle elevation dry sites. Growth is positively correlated with current growing season temperature at all sites; however, growth is negatively correlated with previous year August temperature, indicating that climate affects growth in subsequent years. Positive correlations between growth and summer precipitation during the growing season at low and middle elevation dry sites suggest that soil moisture is partially limiting to growth on these sites. If the climate of the Pacific Northwest becomes warmer and drier, then subalpine fir growth may increase at high elevation and wet sites, but may decrease at lower elevation dry sites in the Olympic Mountains. However, the growth response of subalpine fir to potentially rapid climate change will not be uniform because subalpine fir grows over a wide range of topographic features, habitats, and local climates at different geographical scales. A comparison of growth response to current growing season temperature suggests that the temperature-related growth response of subalpine fir is not adequately described by the parabolic curve used in JABOWA-based models.     

Plant species richness in mountain forests of the Bavarian Alps

Abstract

Based on a stratified random sample of 93 vegetation plots (144 m²) from montane and subalpine climax forests in a representative section through the Bavarian Alps, spatial pattern and environmental correlates of species density of trees, vascular understorey and epigeic bryophytes were analysed. Detecting landscape scale patterns in beta- and gamma-diversity based on interpretation of rarefaction curves proved to be difficult in a sample that had been stratified by ecological criteria. In 144 m² plots tree species density (5 ± 2.0, max. 10) declined with elevation and increased with stand age (multiple R ² = 0.557). The latter effect can be attributed to the secular history of game management and browsing pressure, which has hindered the regeneration of species-rich tree stands since ca. 150 yr. Species density of the forest undergrowth reached remarkably high levels for vascular plants (42 ± 12.8, max. 69) and bryophytes (14 ± 6.0, max. 30) and strongly depended on cover of the respective layer in a unimodal pattern, suggesting to separate direct and indirect effects, mediated through the mass effect, in the subsequent construction of regression models. Multiple regression (R ² = 0.47) revealed that vascular species density is limited chiefly through low plant cover, which in turn decreases with tree cover, elevation and soil quality, and secondly by species pools that contain larger numbers of species requiring high pH and ample light. Cover and direct effects had roughly equal weight in controlling bryophyte species density (R ² = 0.57). Biomass depended on the proportion of conifers in the tree layer and on site quality, less fertile sites tending to have higher bryophyte cover. The increase of bryophyte species density with elevation was interpreted as an effect of a pool of largely boreal-subalpine species. The increase of species density with stand age suggests dispersal limitation and deserves further study.