Forest vegetation of the Colorado Front Range: Patterns of species diversity

Summary Plant species diversity patterns of the Rocky Mountain forests were found to be at variance with patterns reported from other regions. The most centrally located forests in terms of elevation, site moisture and successional status were found to have the lowest diversity. In contrast, the peripheral and environmentally more severe sites were found to have relatively high diversity. In particular, the forest-grassland transition and the low elevation riparian forests have species diversity values as high as any yet reported from western North America.When diversity was examined in terms of variation across elevation or moisture gradients, varying results were obtained due to the interaction of these factors. The failure of previous studies to converge on generalizations about plant diversity reflects, in part, the failure of most investigators to view diversity in a regional context of variation across several interacting gradients.Diversity was seen to vary inversely with the degree of development of the forest canopy. The interaction of different components of the forest community is one reason for the failure of general patterns of plant species diversity to emerge from previous studies. A potentially rich herb community can be greatly suppressed by a single species tree stratum.Among the most successful work to date on species diversity is that on birds, a distinct albeit large and functional group. It is unlikely that similar success could have been achieved through work on all animal species simultaneously. This suggests the need to examine plant species diversity, not in terms of total diversity, but in terms of component functional groups, perhaps guilds, growing under similar microclimatic conditions and subject to similar competitive pressures.Nomenclature follows Weber (1976).     

A Spatially-Explicit Reconstruction of Historical Fire Occurrence in the Ponderosa Pine Zone of the Colorado Front Range

A key issue in ecosystem management in the western U.S. is the determination of the historic range of variability of fire and its ecological significance prior to major land-use changes associated with Euro-American settlement. The present study relates spatial variation in historical fire occurrence to variation in abiotic and biotic predictors of fire frequency and severity across the elevational range of ponderosa pine in northern Colorado. Logistic regression was used to relate fire frequency to environmental predictors and to derive a probability surface for mapping purposes. These results indicate that less than 20% of the ponderosa pine zone had an historic fire regime (pre-1915) of relatively frequent fires (mean fire intervals, MFI, <30 years). More than 80% is reconstructed to have had a lower frequency (MFI ≥ 30 years), more variable severity fire regime. High fire frequency is clearly associated with low elevations. Lower and more variable fire frequencies, associated with high and moderate severities, occur across a broad range of elevation and are related to variations in other environmental variables. Only a small part of the ponderosa pine zone fits the widespread view that the historic fire regime was characterized mainly by frequent, low-severity that maintained open conditions. Management attempts to restore historic forest structures and/or fire conditions must recognize that infrequent severe fires were an important component of the historic fire regime in this cover type in northern Colorado.     

Landscape Controls on Organic and Inorganic Nitrogen Leaching across an Alpine/Subalpine Ecotone,Green Lakes Valley,Colorado Front Range

Here we report measurements of organic and inorganic nitrogen (N) fluxes from the high-elevation Green Lakes Valley catchment in the Colorado Front Range for two snowmelt seasons (1998 and 1999). Surface water and soil samples were collected along an elevational gradient extending from the lightly vegetated alpine to the forested subalpine to assess how changes in land cover and basin area affect yields and concentrations of ammonium-N (NH₄-N), nitrate-N (NO₃-N), dissolved organic N (DON), and particulate organic N (PON). Streamwater yields of NO₃-N decreased downstream from 4.3 kg ha⁻¹ in the alpine to 0.75 kg ha⁻¹ at treeline, while yields of DON were much less variable (0.40–0.34 kg ha⁻¹). Yields of NH₄-N and PON were low and showed little variation with basin area. NO₃-N accounted for 40%–90% of total N along the sample transect and was the dominant form of N at all but the lowest elevation site. Concentrations of DON ranged from approximately 10% of total N in the alpine to 45% in the subalpine. For all sites, volume-weighted mean concentrations of total dissolved nitrogen (TDN) were significantly related to the DIN:DON ratio (R ² = 0.81, P < 0.001) Concentrations of NO₃-N were significantly higher at forested sites that received streamflow from the lightly vegetated alpine reaches of the catchment than in a control catchment that was entirely subalpine forest, suggesting that the alpine may subsidize downstream forested systems with inorganic N. KCl-extractable inorganic N and microbial biomass N showed no relationship to changes in soil properties and vegetative cover moving downstream in catchment. In contrast, soil carbon–nitrogen (C:N) ratios increased with increasing vegetative cover in catchment and were significantly higher in the subalpine compared to the alpine (P < 0.0001) Soil C:N ratios along the sample transect explained 78% of the variation in dissolved organic carbon (DOC) concentrations and 70% of the variation in DON concentrations. These findings suggest that DON is an important vector for N loss in high-elevation ecosystems and that streamwater losses of DON are at least partially dependent on catchment soil organic matter stoichiometry. Received 26 July 2001; accepted 6 May 2002.     

Ecosystem Responses to Nitrogen Deposition in the Colorado Front Range

We asked whether 3–5 kg N y⁻¹ atmospheric N deposition was sufficient to have influenced natural, otherwise undisturbed, terrestrial and aquatic ecosystems of the Colorado Front Range by comparing ecosystem processes and properties east and west of the Continental Divide. The eastern side receives elevated N deposition from urban, agricultural, and industrial sources, compared with 1–2 kg N y⁻¹ on the western side. Foliage of east side old-growth Englemann spruce forests have significantly lower C:N and lignin:N ratios and greater N:Mg and N:P ratios. Soil % N is higher, and C:N ratios lower in the east side stands, and potential net N mineralization rates are greater. Lake NO₃ concentrations are significantly higher in eastern lakes than western lakes. Two east side lakes studied paleolimnologically revealed rapid changes in diatom community composition and increased biovolumes and cell concentrations. The diatom flora is now representative of increased disturbance or eutrophication. Sediment nitrogen isotopic ratios have become progressively lighter over the past 50 years, coincident with the change in algal flora, possibly from an influx of isotopically light N volatilized from agricultural fields and feedlots. Seventy-five percent of the increased east side soil N pool can be accounted for by increased N deposition commensurate with human settlement. Nitrogen emissions from fixed, mobile, and agricultural sources have increased dramatically since approximately 1950 to the east of the Colorado Front Range, as they have in many parts of the world. Our findings indicate even slight increases in atmospheric deposition lead to measurable changes in ecosystem properties. Received 16 November 1999; accepted 8 February 2000.     

Stand dynamics of Pinus flexilis-dominated subalpine forests in the Colorado Front Range

Population age structure and succession were investigated in subalpine forests in the Colorado Front Range dominated by Pinus flexilis (limber pine). Age, size, and spatial data were collected from three recent burns (<100 yr old), six ca. 240 year-old post-fire stands, and two old-growth stands (individuals > 400 yr old). The sequence of colonization of now extant trees on these post-fire sites appeared to be consistent: first Pinus flexilis, then Picea engelmannii (Engelmann spruce), and later Abies lasiocarpa (subalpine fir) with a delay between the first Pinus flexilis and Abies of as long as 140 yr. The advantage of Pinus flexilis on post-burn sites can be attributed to avian seed dispersal and the exceptional drought tolerance of its seedlings. The three recent burns were not extensive, and the delay in establishment of Picea and Abies appeared to be limited by harsh site conditions rather than lack of seed dispersal. Spatial analysis indicated a consistent, although sometimes weak, attraction between Pinus flexilis and Picea and Pinus flexilis and Abies at a scale of 1–4 m, suggesting that Pinus flexilis may facilitate establishment of Picea and Abies seedlings by providing shade or protection from wind. On xeric to slightly xeric sites, Pinus flexilis appeared to form broadly even-aged, non-regenerating populations that were gradually being replaced by Picea and Abies. Replacement is proceeding at a faster rate on the least xeric sites (north aspects, valley bottoms) compared to the most xeric sites (south aspects). On the most extreme sites, Pinus flexilis formed all-aged, self-maintaining populations with no evidence of replacement by Picea and Abies. In these old-growth forests with occasional trees aged at > 1300 yr, recruitment is continuous or episodic.     

A dendroecological reconstruction of western spruce budworm outbreaks (Choristoneura occidentalis) in the Front Range,Colorado, from 1720 to 1986

Radial increment cores from Douglas-fir (Pseudotsuga menziesii) and blue spruce (Picea pungens), defoliated by western spruce budworm (Choristoneura occidentalis), were analyzed by means of dendrochronological methods and compared with cores from undefoliated ponderosa pine (Pinus ponderosa) and lodgepole pine (Pinus contorta) growing on the same sites in the Front Range, Colorado. Extensive deforestation during the gold and silver booms in the second part of the nineteenth century led to dense and almost pure stands of shadetolerant budworm host species. By using the skeleton plot method, the number of trees with clear growth reductions is obtained, thus representing an exact record of forest insect attacks. The analysis of abrupt growth reductions revealed at least nine outbreaks of western spruce budworm between 1720 and 1986, the majority occurring in the nineteenth century. The outbreaks were graphically compared with periods of attack in New Mexico and Colorado which were detected by other scientists employing tree-ring measurement techniques. No increase in the frequency of severe outbreaks during the twentieth century was observed, yet there is some evidence that the most recent outbreak might be the most severe ever recorded. Open Douglas-fir stands on higher sites were more susceptible to heavy budworm attack than dense stands on lower sites. Blue spruce was less frequently and less severely attacked than Douglas-fir. The spatial pattern of historical outbreaks generally was very patchy.     

A tentative typology of European scrub and forest communities based on vegetation texture and structure

Utilitarian and fundamental, floristic and structural classifications of forest and scrub communities are compared. Their advantages and disadvantages and their interrelations are discussed. The necessity of a structural classification as complementary to a floristic classification is emphasized. It seems difficult, if not impossible, to combine the two. The existing structural classifications are reviewed. They are not very detailed and often mixed with a habitat classification. A new, detailed and purely structural system for woods and scrub in Europe is proposed in the form of an identification key. The main criteria used are — in order of diminishing importance —: (1) architecture of the dominant layer, (2) photoperiodicity (evergreen vs. deciduous) of that layer, (3) leaf size and leaf form of that layer, (4) presence/absence of thorns/spines, (5) presence of a second tree layer, shrub layer, dwarf shrub layer, herb layer and/or moss layer, (6) deciduousness of these layers, (7) main growth forms, (8) leaf size and leaf consistency, (9) leaf inclination and (10) height of these layers.In all 27 scrub types and 45 wood types are described. They have been named after a combination of two more or less characteristic and dominant genera. The system is open and hierarchical. Broader units have also been described.Comm. No. 388 of the Biological Station Wijster, Department of Silviculture and Forest Ecology, Agricultural University, Wageningen     

Temporal coherence of two alpine lake basins of the Colorado Front Range, U.S.A.

1. Knowledge of synchrony in trends is important to determining regional responses of lakes to disturbances such as atmospheric deposition and climate change. We explored the temporal coherence of physical and chemical characteristics of two series of mostly alpine lakes in nearby basins of the Colorado Rocky Mountains. Using year‐to‐year variation over a 10‐year period, we asked whether lakes more similar in exposure to the atmosphere be‐haved more similarly than those with greater influence of catchment or in‐lake processes. 2. The Green Lakes Valley and Loch Vale Watershed are steeply incised basins with strong altitudinal gradients. There are glaciers at the heads of each catchment. The eight lakes studied are small, shallow and typically ice‐covered for more than half the year. Snowmelt is the dominant hydrological event each year, flushing about 70% of the annual discharge from each lake between April and mid‐July. The lakes do not thermally stratify during the period of open water. Data from these lakes included surface water temper‐ature, sulphate, nitrate, calcium, silica, bicarbonate alkalinity and conductivity. 3. Coherence was estimated by Pearson's correlation coefficient between lake pairs for each of the different variables. Despite close geographical proximity, there was not a strong direct signal from climatic or atmospheric conditions across all lakes in the study. Individual lake characteristics overwhelmed regional responses. Temporal coherence was higher for lakes within each basin than between basins and was highest for nearest neighbours. 4. Among the Green Lakes, conductivity, alkalinity and temperature were temporally coherent, suggesting that these lakes were sensitive to climate fluctuations. Water tem‐perature is indicative of air temperature, and conductivity and alkalinity concentrations are indicative of dilution from the amount of precipitation flushed through by snowmelt. 5. In Loch Vale, calcium, conductivity, nitrate, sulphate and alkalinity were temporally coherent, while silica and temperature were not. This suggests that external influences are attenuated by internal catchment and lake processes in Loch Vale lakes. Calcium and sulphate are primarily weathering products, but sulphate derives both from deposition and from mineral weathering. Different proportions of snowmelt versus groundwater in different years could influence summer lake concentrations. Nitrate is elevated in lake waters from atmospheric deposition, but the internal dynamics of nitrate and silica may be controlled by lake food webs. Temperature is attenuated by inconsistently different climates across altitude and glacial meltwaters. 6. It appears that, while the lakes in the two basins are topographically close, geologically and morphologically similar, and often connected by streams, only some attributes are temporally coherent. Catchment and in‐lake processes influenced temporal patterns, especially for temperature, alkalinity and silica. Montane lakes with high altitudinal gradients may be particularly prone to local controls compared to systems where coherence is more obvious.     

On the dynamics of vegetation: Succession in model communities

Summary Successional change is thought to be at least partially driven by forees originating from within the community, namely by reaction and competition. Both processes operate through changes in the environment, but from the literature on the subject it is not clear how they differ.To clarify these issues successiens of model communities are studied. This leads us to conclude that competition represent an instantaneous interaction, whereas reaction has historical aspects since it relies on cumulative changes in the environment. The three models considered-one relying on reaction to cause vegetational change, one relying on competition and differential growth rates, and a hybrid third one-yield very similar predictions: roughly bell-shaped curves displaced along the time axis. This shows that the mere fit of a certain model to successional data may easily be spurious (recently some workers have empirically fitted models identical to one derived here from first principles). The three models do behave radically different under perturbation, however: any model relying completely or partially on historical interactions cannot account for the well known possibility of artificially arresting succession. Even if the importance of historical interactions in succession (i.e. the Markovian character of succession) cannot easily be ascertained, one can nevertheless ask whether historical interactions are at all necessary for the explanation of successional change. It is argued here that succession can be entirely understood in terms of instantaneous interactions, notably competition. The argument rests upon the well known relationship between colonizing and competitive ability, and on the fact, proven here, that stress, defined as expressing itself in severe random fluctuations in the growth parameters, is negatively correlated with competition intensity.The author gratefully acknowledges the critical support be received from Professor László Orlóci. This paper forms part of a wider research program supported by a NCR grant to Proc. Orlóci.     

Consequences of cyclic vegetation management for arthropod survival: Simulation experiments

Many rare arthropod species occupy open grasslands. Mowing or grazing is needed to preserve the habitat for these species. Alternatively the vegetation cover in parts of the managed area can be periodically destroyed by ploughing or rototilling. Such treatment results in a dynamic mosaic of habitat patches in different stages of succession. This mosaic may serve as a habitat for many species. However, the interplay between the frequency of rototilling, the spatial structure of the landscape and life history attributes of the animal species will determine the success of such cyclic management strategies. We used a spatially implicit individual-based metapopulation model to examine optimal spatio-temporal management strategies. The model explicitly incorporates succession, population dynamics and dispersal between habitat patches.Optimal management patterns strongly depend on the species’ properties. Our simulation experiments show that in general the more fertile species with strong intraspecific competition are most robust against the large fluctuations of habitat quality resulting from cyclic management. However, the best management strategies found in our experiments cover the requirements of up to 70% of the species tested. Frequent management of relatively small sub-areas minimizes local fluctiations of patch capacity. Though reducing effective patch capacity by more than 50% these strategies may support the survival of the majority of species.     

On the dynamics of vegetation: Patterns of environmental and vegetational change

Summary Ecological succession theory deals with temporal change in biological communities. It consists largely of generalizations based on temporal sequences inferred from spatial ones. The predictive content of the theory is low, since predictions are derived from unconditional trends rather than conditional laws. There exist several conflicting theories purporting to explain successional change, but their empirical vacuousness prevents an assessment on empirical terms. It is argued here that one can nevertheless advocate a theory which accounts for the ubiquity of successional change and explains the most conspicuous characteristics of the successional process, even though it cannot predict the detailed dynamics. Such a theory is derived here from an analysis of adaptive strategies.It is also pointed out that a persistant confusion exists in the ecological literature between what are considered to be the driving forces of successional change, competition and reaction. The former is taken to be an instantaneous type of interaction, whereas the latter has historical (cumulative) aspects. It is not at all obvious whether interactions of the historical type play an important role in driving vegetational change, although it is usually suggested that they do.     

Vegetation study as a generator for population biological and physiological research on salt marshes

The paper deals with some views on the phytocoenose in relation to the functioning of vegetation and its plant-species populations in space and time. From these viewpoints the study of vegetation is seen as a field of tension between the organismic and reductionistic approaches. Both have their value, provided any dogmatism is avoided and either can be applied to the other.In the field of vegetation structure characteristic features of life-form spectra and species distribution, inversion phenomena in zonation, and community architecture in relation to production and decomposition are discussed. In this connection some remarks are made on habitat and niche differentiation with respect to the phytocoenose concept.Vegetation dynamics are discussed in relation to the introduction of Spartina anglica, the frequency of flooding by the tides, different environmental disturbances caused by heavy winter frost, rainfall and hot and dry periods, as well as to human interferences for agricultural and civil-technical purposes.It is suggested that salt-marsh plants may have found refuge areas in inland habitats as well as on more southerly coastal sites during glaciations.Nomenclature follows Tutin et al. (1964–1980).The author wishes to thank his collaborators Messrs B. P. Koutstaal and W. de Munck for much fieldwork, and Mrs M. J. van Leerdam-de Dreu and Messrs A. A. Bolsius and J. A. van den Ende for the preparation of text and figures.Communication No. 302 of the Delta Institute for Hydrobiological Research.     

Middle and high elevation coniferous forest communities of the North Rim region of Grand Canyon National Park,Arizona, USA

We examined the composition and structure of forest communities in a 3700 ha watershed in relation to environmental gradients and changes in land management practices. We identified four mixed-conifer forest types dominated by different combinations of Abies concolor, Picea pungens, Pinus ponderosa, Populus tremuloides, and Pseudotsuga menziesii, and a spruce-fir type dominated by Picea engelmannii and Abies lasiocarpa. The forest types occur in a complex pattern related to elevation and topographicmoisture gradients and variations in past fire regimes. However, widespread regeneration of A. concolor following possible changes in the fire regime in the late 19th century and continuing with institution of a fire suppression policy early in the 20th century is producing a more homogenous mixed-conifer forest with greater horizontal and vertical continuity of fuel. This shift toward landscape homogeneity not only may adversely affect biodiversity, but also may be perpetuated as the probability of large, high-severity fires increases with continued fire suppression.     

Understory vegetation indicates historic fire regimes in ponderosa pine‐dominated ecosystems in the Colorado Front Range

Question: Can current understory vegetation composition across an elevation gradient of Pinus ponderosa‐dominated forests be used to identify areas that, prior to 20th century fire suppression, were characterized by different fire frequencies and severities (i.e., historic fire regimes)? Location: P. ponderosa‐dominated forests in the montane zone of the northern Colorado Front Range, Boulder and Larimer Counties, Colorado, USA. Methods: Understory species composition and stand characteristics were sampled at 43 sites with previously determined fire histories. Indicator species analyses and indirect ordination were used to determine: (1) if stands within a particular historic fire regime had similar understory compositions, and (2) if understory vegetation was associated with the same environmental gradients that influence fire regime. Classification and regression tree analysis was used to ascertain which species could predict fire regimes. Results: Indicator species analysis identified 34 understory species as significant indicators of three distinct historic fire regimes along an elevation gradient from low‐ to high‐elevation P. ponderosa forests. A predictive model derived from a classification tree identified five species as reliable predictors of fire regime. Conclusions: P. ponderosa‐dominated forests shaped by three distinct historic fire regimes have significantly different floristic composition, and current understory compositions can be used as reliable indicators of historical differences in past fire frequency and severity. The feasibility demonstrated in the current study using current understory vegetation properties to detect different historic fire regimes, should be examined in other fire‐prone forest ecosystems.     

Classification of vegetation sequences in Toohey Forest,Queensland

In this paper we consider one method of mapping larger units identified from the spatial pattern of sequences of vegetation types. The basic data were presence/absence data for 6450 stands arranged in 90 transects. A second set of data was derived by averaging the species occurrences in non-overlapping groups of 5 stands. A divisive numerical classification was used to determine the primary vegetation units. In all, 5 different sets of primary types were derived, using different species suites, different sample sizes and different numerical methods. We briefly discuss the types identified and their spatial patterns in the area.Each of these types was then used to define a string of type-codes for every transect so that each transect represents a sample from the landscape containing information on the frequency and spatial distribution of the primary vegetation types. The transects may be classified using a Levenshtein dissimilarity measure and agglomerative hierarchical classification, giving 5 analyses of transects, one for each of the primary types discussed above. We then examine these transect classifications to investigate the stability of the vegetation landspace patterns under changes in species used for the primary classification, in size of sample unit and in method of primary classifications. There is a considerable degree of stability in the results. However it seems with this vegetation that the tree species and non-tree species have considerable independence. We also indicate some problems with this approach and some possible extensions.     

Stand structure and vegetation dynamics of a subalpine wooded fen in Rocky Mountain National Park,Colorado

Abstract. The tree population size structure and relationship between tree diameter and age were examined in a subalpine fen and surrounding Picea-Abies forest in northern Colorado. The fen grades from a sedge fen, through an ecotone, to a treed fen. Tree growth rate varies across the vegetational gradient, with the sedge fen having the slowest growth, and the upland forest having the fastest growth. Differences in growth rate are related to the average size of peat hummocks, with areas containing tall hummocks exhibiting the highest tree growth rates. Size structures display the characteristic reverse-J distribution generally indicative of stable populations, but forest vegetation is expanding into the open regions of the fen, and within the treed fen an increase in Abies lasiocarpa is occurring. These changes are primarily attributed to a positive feedback situation wherein the fen's surface is built up by peat accumulation. Distinct hummocks form first on the open fen but then coalesce to form raised peat islands in the treed fen. This new substrate provides habitat with a comparatively low water table and allows the growth of mesophytic forest vegetation. A pathway for this vegetational development is proposed.     

武夷山木荷种群结构和动态的研究

 本文采用“空间序列代替时间变化”的方法,对武夷山区常绿阔叶林演替过程中木荷种群动态规律进行了研究,结果表明,在木荷种群侵入黄山松幼林向木荷林以及成熟木荷、甜槠、青冈林的发展过程中,种群结构由增长型→稳定型→成熟型→衰退型,空间分布格局由随机型趋于集群分布,种群密度呈现负增长;在成熟木荷、甜槠、青冈林中,由于种间和种内竞争的影响以及林窗效应,木荷种群结构和密度有一个波动性的变化过程,在这一变化过程中,木荷种群仍趋于集群分布。此外,对成熟木荷、甜槠、青冈林中木荷种群不同大小级的分布格局动态进行了研究,表明其幼苗、幼树和中树等级的个体均呈集群分布,而大树等级的个体呈随机分布。根据上述研究结果,对木荷种群动态机制作了探讨。     

Spatial and temporal variation in historic fire regimes in subalpine forests across the Colorado Front Range in Rocky Mountain National Park, Colorado, USA

Aim The historical variability of fire regimes must be understood in the context of drivers of the occurrence of fire operating at a range of spatial scales from local site conditions to broad‐scale climatic variation. In the present study we examine fire history and variations in the fire regime at multiple spatial and temporal scales for subalpine forests of Engelmann spruce–subalpine fir (Picea engelmannii, Abies lasiocarpa) and lodgepole pine (Pinus contorta) of the southern Rocky Mountains. Location The study area is the subalpine zone of spruce–fir and lodgepole pine forests in the southern sector of Rocky Mountain National Park (ROMO), Colorado, USA, which straddles the continental divide of the northern Colorado Front Range (40°20′ N and 105°40′ W). Methods We used a combination of dendroecological and Geographic Information System methods to reconstruct fire history, including fire year, severity and extent at the forest patch level, for c. 30,000 ha of subalpine forest. We aggregated fire history information at appropriate spatial scales to test for drivers of the fire regime at local, meso, and regional scales. Results The fire histories covered c. 30,000 ha of forest and were based on a total of 676 partial cross‐sections of fire‐scarred trees and 6152 tree‐core age samples. The subalpine forest fire regime of ROMO is dominated by infrequent, extensive, stand‐replacing fire events, whereas surface fires affected only 1–3% of the forested area. Main conclusions Local‐scale influences on fire regimes are reflected by differences in the relative proportions of stands of different ages between the lodgepole pine and spruce–fir forest types. Lodgepole pine stands all originated following fires in the last 400 years; in contrast, large areas of spruce–fir forests consisted of stands not affected by fire in the past 400 years. Meso‐scale influences on fire regimes are reflected by fewer but larger fires on the west vs. east side of the continental divide. These differences appear to be explained by less frequent and severe drought on the west side, and by the spread of fires from lower‐elevation mixed‐conifer montane forests on the east side. Regional‐scale climatic variation is the primary driver of infrequent, large fire events, but its effects are modulated by local‐ and meso‐scale abiotic and biotic factors. The low incidence of fire during the period of fire‐suppression policy in the twentieth century is not unique in comparison with the previous 300 years of fire history. There is no evidence that fire suppression has resulted in either the fire regime or current forest conditions being outside their historic ranges of variability during the past 400 years. Furthermore, in the context of fuel treatments to reduce fire hazard, regardless of restoration goals, the association of extremely large and severe fires with infrequent and exceptional drought calls into question the future effectiveness of tree thinning to mitigate fire hazard in the subalpine zone.     

Altitudinal changes in composition and structure of mountain-temperate vegetation: a case study from the Western Carpathians

Changes in composition and structure of plant communities in relation tothe soil and snow cover variation were analyzed along an altitudinal transect(1150–1750 m) from the mountain-temperate forests to a woodyshrub community and alpine meadows on Mt Velký Gápel', Slovakia.The soils below the treeline (1510 m) had a more developedorganic layer above the mineral substratum. Generally, soil depth decreased asthe altitude increased, although the maximum values were recognized at a middlealtitude in a beech stand. Snow was redistributed by westerly winds from theridgeline down to the upper forest margin. Mean snow depth decreased withaltitude up to almost snow-free sites around the summit. In the 48 plots at 16sites we recorded 118 taxa including 6 tree, 7 shrub, 18 grass, 42 herb, 5fern,25 moss and 15 lichen species. The species diversity showed no distinctrelationship to altitude but declined with canopy consolidation. The TWINSPANfloristic classification distinguished five groups of community typescharacterised by different dominants, and a further three clusters of samplesfrom transition zones. Horizontal compositional heterogeneity increased inareaswhere trees were aggregated and tree basal area was smaller. Vegetationcomposition became more patchy at open-canopy Acerpseudoplatanus–Abies alba mixed forest at 1150 m,in Picea abies forest limit 1470 m, andin Pinus mugo krummholz at 1590 m. Speciesturnover of the entire transect was 6.1 half-changes as estimated by DCA.Despite this heterogeneity, none of the 15 elevational bands had significantaggregation of species' limits. Vegetation varied continuously, with individualspecies overlapping in transition zones delimited by dominant taxa. Thecoincident aggregation of up-slope and down-slope boundaries was found at abelt1430–1510 m. This discrete ecotone corresponds to a shiftfrom the closed coniferous forest to P. mugo krummholz.Thesecond inherent up-slope boundary aggregation indicated the P.mugo krummholz – alpine meadow vegetation transition at1700 m. Spatial analysis (K-function) of eight forest plots(0.12 ha each) showed that at lower elevation, adult trees of thebroad-leaf forest were closer to a random arrangement while at higherelevation,trees of evergreen coniferous stands became aggregated toward the forest limitwith the highest intensity from 2 to 4 m. Altitudinal gradient andrelated factors explained 35% of the variance in vegetation data.Canonical correspondence analysis also showed that main vegetation changesabovethe treeline area were associated with the topographic pattern of pine shrubsand snow cover.