Limited prospects for future alpine treeline advance in the Canadian Rocky Mountains

Treeline advance has occurred throughout the twentieth century in mountainous regions around the world; however, local variation and temporal lags in responses to climate warming indicate that the upper limits of some treelines are not necessarily in climatic equilibrium. These observations suggest that factors other than climate are constraining tree establishment beyond existing treelines. Using a seed addition experiment, we tested the effects of seed availability, predation and microsite limitation on the establishment of two subalpine tree species (Picea engelmannii and Abies lasiocarpa) across four treelines in the Canadian Rocky Mountains. The effect of vegetation removal on seedling growth was also determined, and microclimate conditions were monitored. Establishment limitations observed in the field were placed in context with the effects of soil properties observed in a parallel experiment. The seed addition experiment revealed reduced establishment with increasing elevation, suggesting that although establishment within the treeline ecotone is at least partially seed limited, other constraints are more important beyond the current treeline. The effects of herbivory and microsite availability significantly reduced seedling establishment but were less influential beyond the treeline. Microclimate monitoring revealed that establishment was negatively related to growing season temperatures and positively related to the duration of winter snow cover, counter to the conventional expectation that establishment is limited by low temperatures. Overall, it appears that seedling establishment beyond treeline is predominantly constrained by a combination of high soil surface temperatures during the growing season, reduced winter snowpack and unfavourable soil properties. Our study supports the assertion that seedling establishment in alpine treeline ecotones is simultaneously limited by various climatic and nonclimatic drivers. Together, these factors may limit future treeline advance in the Canadian Rocky Mountains and should be considered when assessing the potential for treeline advance in alpine systems elsewhere     

Environmental drivers of subalpine and alpine fen vegetation in the Southern Rocky Mountains,Colorado, USA

Fens are widely distributed wetlands worldwide and provide vital habitat for plant and animal species in mountainous regions. Alpine fens are rare in the Rocky Mountains and concentrated in the San Juan Mountains where broad regions at high elevation have relatively level topography and suitable climate to favor peat accumulation. Studies of montane and boreal peatlands have identified water chemistry as a main driver of vegetation composition. This study investigated whether similar drivers of vegetation composition are important for alpine and subalpine fens in the San Juan Mountains of Colorado, USA. Water chemistry variables were most important in structuring subalpine and alpine fen vegetation. However, these variables explained considerably less variation in alpine than subalpine fen vegetation. In addition, lower variance of water chemistry in alpine fens did not lead to lower beta diversity of vegetation in alpine than in subalpine fens. Although alpine and subalpine fen vegetation supports similar beta diversity, key differences occur in the environmental drivers of their vegetation composition.     

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.     

Aspen succession and nitrogen loading: a case for epiphytic lichens as bioindicators in the Rocky Mountains,USA

Question: Can lichen communities be used to assess short‐ and long‐term factors affecting seral quaking aspen (Populus tremuloides) communities at the landscape scale? Location: Bear River Range, within the Rocky Mountains, in northern Utah and southern Idaho, USA. Method: Forty‐seven randomly selected mid‐elevation aspen stands were sampled for lichens and stand conditions. Plots were characterized according to tree species cover, basal area, stand age, bole scarring, tree damage, and presence of lichen species. We also recorded ammonia emissions with passive sensors at 25 urban and agricultural sites throughout an adjacent populated valley upwind of the forest stands. Nonmetric multidimensional scaling (NMS) ordination was used to evaluate an array of 20 variables suspected to influence lichen communities. Results: In NMS, forest succession explained most variance in lichen composition and abundance, although atmospheric nitrogen from local agricultural and urban sources also significantly influenced the lichen communities. Abundance of nitrophilous lichen species decreased with distance from peak ammonia sources and the urban center in all aspen succession classes. One lichen, Phaeophyscia nigricans, was found to be an effective bioindicator of nitrogen loading. Conclusions: Lichen communities in this landscape assessment of aspen forests showed clear responses to long‐term (stand succession) and short‐term (nitrogen deposition) influences. At the same time, several environmental factors (e.g. tree damage and scarring, distance to valley, topography, and stand age) had little influence on these same lichen communities. We recommend further use of epiphytic lichens as bioindicators of dynamic forest conditions.     

Warming‐induced upward migration of the alpine treeline in the Changbai Mountains,northeast China

Treeline responses to environmental changes describe an important phenomenon in global change research. Often conflicting results and generally too short observations are, however, still challenging our understanding of climate‐induced treeline dynamics. Here, we use a state‐of‐the‐art dendroecological approach to reconstruct long‐term changes in the position of the alpine treeline in relation to air temperature at two sides in the Changbai Mountains in northeast China. Over the past 160 years, the treeline increased by around 80 m, a process that can be divided into three phases of different rates and drives. The first phase was mainly influenced by vegetation recovery after an eruption of the Tianchi volcano in 1702. The slowly upward shift in the second phase was consistent with the slowly increasing temperature. The last phase coincided with rapid warming since 1985, and shows with 33 m per 1°C, the most intense upward shift. The spatial distribution and age structure of trees beyond the current treeline confirm the latest, warming‐induced upward shift. Our results suggest that the alpine treeline will continue to rise, and that the alpine tundra may disappear if temperatures will increase further. This study not only enhances mechanistic understanding of long‐term treeline dynamics, but also highlights the effects of rising temperatures on high‐elevation vegetation dynamics.     

Climate‐driven speedup of alpine treeline forest growth in the Tianshan Mountains,Northwestern China

Forest growth is sensitive to interannual climatic change in the alpine treeline ecotone (ATE). Whether the alpine treeline ecotone shares a similar pattern of forest growth with lower elevational closed forest belt (CFB) under changing climate remains unclear. Here, we reported an unprecedented acceleration of Picea schrenkiana forest growth since 1960s in the ATE of Tianshan Mountains, northwestern China by a stand‐total sampling along six altitudinal transects with three plots in each transect: one from the ATE between the treeline and the forest line, and the other two from the CFB. All the sampled P. schrenkiana forest patches show a higher growth speed after 1960 and, comparatively, forest growth in the CFB has sped up much slower than that in the ATE. The speedup of forest growth at the ATE is mainly accounted for by climate factors, with increasing temperature suggested to be the primary driver. Stronger water deficit as well as more competition within the CFB might have restricted forest growth there more than that within the ATE, implying biotic factors were also significant for the accelerated forest growth in the ATE, which should be excluded from simulations and predictions of warming‐induced treeline dynamics.     

Estimation of Successful Breeding Pairs for Wolves in the Northern Rocky Mountains,USA

Abstract: Under the Endangered Species Act, documenting recovery and federally mandated population levels of wolves (Canis lupus) in the Northern Rocky Mountains (NRM) requires monitoring wolf packs that successfully recruit young. United States Fish and Wildlife Service regulations define successful breeding pairs as packs estimated to contain an adult male and female, accompanied by ≥2 pups on 31 December of a given year. Monitoring successful breeding pairs will become more difficult following proposed delisting of NRM wolves; alternatives to historically intensive methods, appropriate to the different ecological and regulatory context following delisting, are required. Because pack size is easier to monitor than pack composition, we estimated probability a pack would contain a successful breeding pair based on its size for wolf populations inhabiting 6 areas in the NRM. We also evaluated the extent to which differences in demography of wolves and levels of human-caused mortality among the areas influenced the probability of packs of different sizes would contain successful breeding pairs. Probability curves differed among analysis areas, depending primarily on levels of human-caused mortality, secondarily on annual population growth rate, and little on annual population density. Probabilities that packs contained successful breeding pairs were more uniformly distributed across pack sizes in areas with low levels of human mortality and stable populations. Large packs in areas with high levels of human-caused mortality and high annual growth rates had relatively high probabilities of containing breeding pairs whereas those for small packs were relatively low. Our approach can be used by managers to estimate number of successful breeding pairs in a population where number of packs and their sizes are known. Following delisting of NRM wolves, human-caused mortality is likely to increase, resulting in more small packs with low probabilities of containing breeding pairs. Differing contributions of packs to wolf population growth based on their size suggests monitoring successful breeding pairs will provide more accurate insights into population dynamics of wolves than will monitoring number of packs or individuals only.     

Populational dynamics of three annual species of alpine plants in the Rocky Mountains

Summary Survivorship, fecundity, and seed bank size were measured over two years in alpine populations of the annuals, Koenigia islandica, Polygonum confertiflorum, and P. douglasii. The major loss of individuals from all populations occurred between seed dispersal and germination. Survival of vegetative plants to maturity was high in all species, usually above 80%, and the average number of seeds produced per plant was less than 10 in all species in both years. Seed banks existed for all three species but were small since more than 80% of the viable seeds in the soil germinated in the spring. Both survival and fecundity were negatively correlated with density in P. confertiflorum. These patterns of population dynamics are similar to those found in many annuals of temperate environments.     

Post‐fire tree establishment patterns at the alpine treeline ecotone: Mount Rainier National Park,Washington, USA

Questions: Does tree establishment: (1) occur at a treeline depressed by fire, (2) cause the forest line to ascend upslope, and/or (3) alter landscape heterogeneity? (4) What abiotic and biotic local site conditions are most important in structuring establishment patterns? (5) Does the abiotic setting become more important with increasing upslope distance from the forest line? Location: Western slopes of Mount Rainier, USA. Methods: We performed classification analysis of 1970 satellite imagery and 2003 aerial photography to delineate establishment. Local site conditions were calculated from a LIDAR‐based DEM, ancillary climate data, and 1970 tree locations in a GIS. We used logistic regression on a spatially weighted landscape matrix to rank variables. Results: Considerable establishment after 1970 caused forest line elevation to increase over 150 m in specific locations. Landscape heterogeneity increased with distance from the 1970 forest line. At a broad spatial context, we found establishment was most common near existing trees (0‐50 m) and at low elevations (1250‐1350 m). Slope aspect (W, NW, N, NE, and E), slope angle (40‐60°), and other abiotic factors emerged as important predictors of establishment with increasing upslope distance from the forest line to restricted spatial extents. Conclusions: Favorable climatic conditions likely triggered widespread tree establishment. Readily available seed probably enhanced establishment rates near sexually mature trees, particularly in the less stressful environment at low elevations. The mass effect of nearly ubiquitous establishment in these areas may have obscured the importance of the abiotic setting to restricted spatial extents. Topographic variability apparently produced favorable sites that facilitated opportunistic establishment with increasing upslope distance from the forest line, thereby enabling additional trees to invade the alpine tundra.     

Historical biogeography of two alpine butterflies in the Rocky Mountains: broad-scale concordance and local-scale discordance

Aim We inferred the phylogeography of the alpine butterfly Colias meadii Edwards (Pieridae) and compared its genetic structure with that of another high elevation, co‐distributed butterfly, Parnassius smintheus Doubleday (Papilionidae), to test if the two Rocky Mountain butterflies responded similarly to the palaeoclimatic cycles of the Quaternary. Location Specimens were collected from 18 alpine sites in the Rocky Mountains of North America, from southern Colorado to northern Montana. Methods We sequenced 867 and 789 nucleotides of cytochrome oxidase I from an average of 19 and 20 individuals for C. meadii and P. smintheus, respectively, from each of the same 18 localities. From the sequence data, we calculated measures of genetic diversity within each population (H, θ), genetic divergence among populations (F_ST), and tested for geographic structure through an analysis of molecular variance (amova ). Population estimates were compared against latitude and between species using a variety of statistical tests. Furthermore, nested clade analysis was implemented to infer historic events underlying the geographic distribution of genetic variation in each species. Then, we compared the number of inferred population events between species using a nonparametric Spearman's rank correlation test. Finally, we ran coalescent simulations on each species’ genealogy to test whether the two species of Lepidoptera fit the same model of population divergence. Results Our analyses revealed that: (1) measures of within‐population diversity were not correlated with latitude for either species, (2) within‐site diversity was not correlated between species, (3) within a species, nearly all populations were genetically isolated, (4) both species exhibited significant and nearly identical partitioning of genetic variation at all hierarchical levels of the amova , including a strong break between populations across the Wyoming Basin, (5) both species experienced similar cycles of expansion and contraction, although fewer were inferred for C. meadii, and (6) data from both species fit a model of three refugia diverging during the Pleistocene. Main conclusions While our findings supported a shared response of the two butterfly species to historic climate change across coarse spatial scales, a common pattern was not evident at finer spatial and temporal scales. The shared demographic history of the two species is consistent with an expanding–contracting archipelago model, suggesting that populations persisted across the geographic range throughout the climate cycles, experiencing isolation on ‘sky islands’ during interglacial periods and becoming connected as they migrated down‐slope during cool, wet climates.     

Estimating occupancy to monitor northern goshawk in the central Rocky Mountains

Designing monitoring programs to evaluate trends in low-density wildlife species at regional scales is challenging given difficulties detecting uncommon organisms distributed in potential habitats over large spatial extents. The northern goshawk (Accipiter gentilis) has been petitioned for listing under the Endangered Species Act and the review of the petition indicated a need for information on population trend. To evaluate trends in goshawk populations, the U.S. Forest Service developed the Northern Goshawk Bioregional Monitoring Design to estimate goshawk occupancy over broad spatial extents. We adapted and implemented this design to approximately 30,600 km² of 88,128 km² of National Forest System lands in the Forest Service Rocky Mountain Region, including portions of Colorado, Wyoming, and South Dakota. We developed a stratified random design to monitor goshawk occupancy in sampling units, defined by primary and secondary habitat quality as well as accessibility. To define habitat quality, we examined a time series for 58 previously located nesting territories. Using logistic regression, we found that the dominant conifer species and status of aspen in postfledging zones best characterized high-quality goshawk nesting habitat. We applied model results to stratify 4,445 sampling units based on habitat quality and further stratified sampling units based on accessibility into easy and difficult access categories. We conducted field sampling during the goshawk breeding season in the summer of 2006 to estimate detection probabilities and occupancy rates. Within our sampling frame, we sampled 51 sampling units and estimated goshawk occupancy of 0.329 (95% CI: 0.213–0.445). Occupancy within primary strata (high quality) sampling units was 0.811 (SE = 0.113), whereas occupancy in secondary strata (lower quality) sampling units was 0.124 (SE = 0.067). Future implementation of this monitoring program can achieve 0.8 power to detect 30–40% declines in with 140 sampling units. Our implementation of a stratified sampling design to monitor occupancy of goshawks at a region-wide scale reduced the number of sampling units in each administrative unit and focused our efforts on those areas most likely to have goshawks. © 2011 The Wildlife Society.     

Large herbivores influence the composition and diversity of shrub-steppe communities in the Rocky Mountains, USA

It is widely believed that wild and domestic herbivores have modified the structure and composition of arid and semi-arid plant communities of western North America, but these beliefs have rarely been tested in long-term, well-replicated studies. We examined the effects of removing large herbivores from semi-arid shrublands for 40–50 years using 17 fenced exclosures in western Colorado, USA. Shrub cover was greater (F=5.87, P=0.0020) and cover (F=3.01, P=0.0601) and frequency (F=3.89, P=0.0211) of forbs was less inside the exclosures (protected) relative to grazed plots. However, we found no significant effects (minimum P=0.18) of protection from grazing on cover or frequency of grasses, biotic crusts, or bare soil. Although mean species richness and diversity were similar between treatments, protected areas had much higher dominance by fewer species, primarily sagebrush. Exclusion of herbivores changed the relationship between species richness and evenness. Consistent with theoretical expectations, species evenness was positively correlated with richness in protected plots (r ²=0.54). However, contrary to theory, evenness and richness were inversely related in grazed plots (r ² _adjacent=0.72, r ² _distant=0.84). We suggest that these differences resulted because grazing acts as a stressor promoting facilitative relationships between plant species that might compete for resources in the absence of grazing. We conclude that exclusion of grazing in the sites we studied caused minor changes in cover and diversity of herbaceous plants, but caused a clear increase in the cover of shrubs. Importantly, the exclusion of ungulates changed the relationship between evenness and richness.     

Species Climatic Suitability Explains Insect–Host Dynamics in the Southern Rocky Mountains,USA

Ecosystems - Recent extreme events of drought and heat have been associated with insect-driven tree mortality. However, there is substantial uncertainty about the impact of climate variability and...     

Assessing the sensitivity of alpine lakes and ponds to nitrogen deposition in the Canadian Rocky Mountains

To demonstrate the sensitivity of aquatic ecosystems to forecasted increases in nitrogen deposition along the eastern ranges of the Canadian Rocky Mountains, we conducted midsummer limnological surveys of 29 remote alpine lakes and ponds via helicopter in 2007. Chemical analysis of water and in vitro nitrogen-enrichment bioassays of phytoplankton collected from each site were performed to estimate nutrient limitation. Use of a common chemical index for nutrient limitation (total dissolved inorganic nitrogen: total phosphorus; DIN:TP) together with supportive experimental evidence revealed nitrogen limitation in only 14% of the cases. Shallow (≤1 m maximum depth) ponds were more likely to be nitrogen-limited than lakes, especially as the former exhibited a significantly lower mean DIN:TP ratio of 7.4 during the late summer. Chemical and bioassay-based inferences of nitrogen limitation agreed in 74.5% of the cases, owing mainly to evidence of phosphorus limitation of the surveyed lake ecosystems (mean DIN:TP = 18) being supported by nonsignificant responses of their phytoplankton to nitrogen amendment. Our findings reveal that increased nitrogen deposition should not result in immediate widespread eutrophication of the Canadian Rockies; however, certain alpine ponds appeared nitrogen-limited, making them sensitive early indicators of the potential effects of anthropogenic nitrogen deposition in remote mountainous regions.     

Biogeochemical Change During Climate-Driven Afforestation: A Paleoecological Perspective from the Rocky Mountains

Shifts in an ecosystem’s state can alter biogeochemical cycling and the extent of nutrient conservation within a terrestrial landscape on multiple time scales. Transient biogeochemical changes may follow disturbance and succession, although persistent long-term differences may exist under different climates and vegetation types. We evaluate the potential for such biogeochemical changes in the context of long-term ecological history by measuring the nitrogen isotope composition of organic matter in a lake sediment core. We targeted Little Windy Hill Pond (LWH) in the Medicine Bow Mountains, Wyoming because reconstructions of the lake level, fire, and vegetation histories from the lacustrine sediments indicated a century-scale transformation from an arid, shrub-dominated landscape to a sub-alpine, tree-dominated ecosystem with extensive woody cover and large, live biomass pools. We demonstrate that the afforestation at the beginning of the Holocene transformed the Artemisia-dominated ecosystem, which had persisted for millennia during the Pleistocene. The changes affected nitrogen cycling dynamics, especially through intensified nutrient conservation when live biomass pools increased with greater woody cover. The LWH sediments record a baseline δ¹⁵N shift from 2.2–3.0 to 0.3–2.0‰ as less ¹⁵N-enriched organic matter accumulated in the lake. We also observed a transient pattern of maximum nutrient conservation and minimum δ¹⁵N values as terrestrial biomass increased during the aggradation (~175 years) and transition phases of ecological succession. Our nitrogen isotope results support theoretical expectations of long-term biogeochemical dynamics as nutrient conservation increases during afforestation.     

Quaking aspen (Populus tremuloides Michx.) at treeline: a century of change in the San Juan Mountains, Colorado, USA

During a repeat photography study quaking aspen (Populus tremuloides Michx.) was observed invading conifer stands at treeline in the San Juan Mountains of south‐western Colorado. Aspen tree core samples were collected from nine plots ranging in elevation from 3192 to 3547 m, and estimated dates of establishment of aspen were grouped into 10‐year intervals for analysis. Estimated periods of establishment were compared with century‐long climate data records to derive any correlations with aspen invasion. Other disturbance agents, such as fire and livestock grazing were also considered. Quantitative analysis of climate variables suggests that decreased mean spring precipitation and increased mean summer maximum temperature provide optimal conditions for aspen establishment. Episodes of invasion were non‐synchronous, but all occurred after 1900, and are likely from seed germination, considered unusual in aspen. Different climate variables explain stand initiation from seed and subsequent peak establishment from vegetative reproduction. Long‐term climate records indicate a general warming since the beginning of the 20th century and explain the continued invasion and persistence of aspen at treeline, resulting from asexual reproduction. Short‐term climate records identify anomalously cool, moist years that explain rarely observed sexual reproduction in aspen.     

Fire, fuels and restoration of ponderosa pine–Douglas fir forests in the Rocky Mountains, USA

Aim Forest restoration in ponderosa pine and mixed ponderosa pine–Douglas fir forests in the US Rocky Mountains has been highly influenced by a historical model of frequent, low‐severity surface fires developed for the ponderosa pine forests of the Southwestern USA. A restoration model, based on this low‐severity fire model, focuses on thinning and prescribed burning to restore historical forest structure. However, in the US Rocky Mountains, research on fire history and forest structure, and early historical reports, suggest the low‐severity model may only apply in limited geographical areas. The aim of this article is to elaborate a new variable‐severity fire model and evaluate the applicability of this model, along with the low‐severity model, for the ponderosa pine–Douglas fir forests of the Rocky Mountains. Location Rocky Mountains, USA. Methods The geographical applicability of the two fire models is evaluated using historical records, fire histories and forest age‐structure analyses. Results Historical sources and tree‐ring reconstructions document that, near or before ad 1900, the low‐severity model may apply in dry, low‐elevation settings, but that fires naturally varied in severity in most of these forests. Low‐severity fires were common, but high‐severity fires also burned thousands of hectares. Tree regeneration increased after these high‐severity fires, and often attained densities much greater than those reconstructed for Southwestern ponderosa pine forests. Main conclusions Exclusion of fire has not clearly and uniformly increased fuels or shifted the fire type from low‐ to high‐severity fires. However, logging and livestock grazing have increased tree densities and risk of high‐severity fires in some areas. Restoration is likely to be most effective which seeks to (1) restore variability of fire, (2) reverse changes brought about by livestock grazing and logging, and (3) modify these land uses so that degradation is not repeated.