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.     

Population Estimates of Snowshoe Hares in the Southern Rocky Mountains

Abstract: In 1999 Canada lynx (Lynx canadensis) were reintroduced to the southern Rocky Mountains and in 2000 the species was listed as threatened under the Endangered Species Act in the contiguous United States (Colorado Division of Wildlife 2000, U.S. Fish and Wildlife Service 2000). To better evaluate the progress of this reintroduction, we conducted field studies to estimate population densities of snowshoe hares (Lepus americanus), the primary prey of lynx in Colorado, USA. We conducted our field studies in southwestern Colorado in winters 2002 and 2003. We estimated population densities in forested stands of mature Engelmann spruce (Picea engelmannii)-subalpine fir (Abies lasiocarpa) and mature lodgepole pine (Pinus contorta) using mark-recapture data and 3 methods for estimating effective area trapped: half trap interval, mean maximum distance moved (MMDM), and half MMDM. In Engelmann spruce-subalpine fir, we found density estimates ranged from 0.08 ± 0.03 (SE) hares/ha to 1.32 ± 0.15 hares/ha and in lodgepole pine, density estimates ranged from 0.06 ± 0.01 hares/ha to 0.34 ± 0.06 hares/ha, depending on year and method used for estimating effective area trapped. Our density estimates are similar to those reported at the low phase of the hare cycle in populations to the north (<0.1–1.1 hares/ha), where Canada lynx persist (Hodges 2000a). Although density estimates are a useful comparative tool, they depend upon methods used to estimate effective area trapped. Therefore, we urge caution in comparing our density estimates with those from other areas, which may have used dissimilar methods. We also examined effects of temperature and moon phase on capture success of snowshoe hares; extremely low temperatures affected capture success but moon phase did not. Capture success can be improved by trapping snowshoe hares at temperatures above their lower critical temperature (T_lc). If abundance estimates are derived from mark-recapture studies then effects of temperature should be included when modeling capture probabilities.     

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.     

Is Size of Fecal Pellets a Reliable Indicator of Species of Leporids in the Southern Rocky Mountains?

Abstract: In the southern Rocky Mountains, USA, snowshoe hares (Lepus americanus) and mountain cottontails (Sylvilagus nuttallii) are syntopic. Previous researchers used size of fecal pellets to identify leporid species in the Southern Rockies, a common criterion for identifying species of cervids. We measured 655 fecal pellets from 10 mountain cottontails and 2,374 fecal pellets from 23 snowshoe hares. We found no relationship between the body weight of mountain cottontails and the size of their fecal pellets (r = 0.04, F = 0.01, P = 0.91) but found one for snowshoe hares (r = 0.48, F = 9.3, P = 0.005). Although the 2 species differed in the size of their fecal pellets, the difference between means (1.2 mm) was sufficiently small to require measuring individual pellets and is only applicable to individuals of adult size. Although fecal pellet counts may be used to estimate presence and relative abundance of snowshoe hares in the absence of syntopic leporids, where multiple species of leporids are syntopic this method may yield misleading results.     

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.     

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.     

太行山南段森林群落物种多样性研究

分别用多样性指数、丰富度指数、均匀度指数对山西太行山南段森林群落的物种多样性及其与环境的关系进行了分析.结果表明:(1)3个植被型的物种多样性指数的顺序为:落叶阔叶林>温性针阔叶混交林>温性针叶林.(2)落叶阔叶林中各群系的物种多样性指数的顺序为:漆树 青麸杨林>鹅耳枥 青榨槭林>青檀林>辽东栎林>栓皮栎林>白桦林>红桦林>山杨 白桦林.(3)温性针叶林中各群系的物种多样性指数的顺序为南方红豆杉林>油松林>白皮松林>侧柏林.(4)物种丰富度指数在群落梯度上的总体趋势表现为草本层>灌木层>乔木层;多样性指数的趋势则为灌木层>乔木层>草本层;各群落灌木层物种均匀度指数大于乔木层和草本层,乔木层均匀度与草本层相差不明显.(5)各群落多样性指数在海拔梯度上表现出明显的规律,呈单峰曲线变化,即中等海拔高度上的群落物种多样性较高,而高海拔和低海拔多样性较低,这主要是由于在中海拔(大约1 500 m)地段水热条件组合较好、人类活动干扰较少所致.     

Seedling occurrence in alpine treeline conifers: A case study from the central Rocky Mountains,USA

Information concerning the occurrence of very young (1- to 10-year-old) tree seedlings in the alpine treeline ecotone is rare. Seedling occurrence of the dominant conifers Picea engelmannii and Abies lasiocarpa was measured in the treeline ecotone of the Medicine Bow Mountains, Wyoming (central Rocky Mountains, USA), an area composed of elongated tree islands separated by open meadows (ribbon forest) that grade into the closed forest. No seedlings were found on the windward sides of tree islands, while a mean of 0.6 seedlings.m^–2 occurred on the leeward (downward) sides. These values compared to the 4.2 seedlings.m^–2 in the closed forest. In addition, a strong correspondence was found between snowpack depth and seedling abundance, with depths that were either too shallow (< 0.5 m) or too deep (> 1.5 m) associated with fewer or no seedlings. A. lasiocarpa seedlings made up much less of the overall seedling population in the ribbon forest (6 %) than in the closed forest (22 %). Seedling establishment in this portion of the alpine treeline ecotone appears to be occurring at a low rate that differs between the two dominant species and may be strongly influenced by wind-driven snow accumulation patterns.     

Arthropod Trace Fossils of the Upper Lower Cambrian Gog Group,Southern Rocky Mountains of Canada

The upper lower Cambrian Gog Group in the southern Rocky Mountains of Canada displays a high diversity and abundance of arthropod traces. Four ichnogenera, Cruziana, Diplichnites, Monomorphichnus, Rusophycus, and “Indeterminate Arthropod Scratches” are discussed, with a total of 17 different ichnospecies, as follows: Cruziana billingsi Fillion and Pickerill, 1990, Cruziana irregularis Fenton and Fenton, 1937, Cruziana jenningsi Fenton and Fenton, 1937, Cruziana navicella Fenton and Fenton, 1937, Cruziana omanica Seilacher, 1970, Cruziana pectinata Seilacher, 1994, Cruziana penicillata Gibb, Chatterton, and Pemberton, 2009, Cruziana plicata Crimes, Legg, Marcos, and Arboleya, 1977, Cruziana tenella Linnarsson, 1871, Diplichnites twelvetreesi (Chapman, 1928), Monomorphichnus bilinearis Crimes, 1970b, Monomorphichnus lineatus Crimes, Legg, Marcos, and Arboleya, 1977, Rusophycus eutendorfensis (Linck, 1942 Linck, O. 1942. Die spur Isopodichnus. Senckenbergiana, 25: 232–255. [Google Scholar]), and Rusophycus unilobus (Seilacher, 1970 Seilacher, A. 1970. Cruziana stratigraphy of “non-fossiliferous” Palaeozoic sandstones. In Crimes, T. P. and Harper, J. C. (eds.), Trace Fossils. Seel House Press, Liverpool, 447–476. [Google Scholar]). Three new ichnospecies, i.e., Cruziana caputinclinata isp. nov., Rusophycus subnotous isp. nov., and Diplichnites obliquus isp. nov., are proposed, described, and illustrated. The ichnofauna collected from three localities (Lake O'Hara, Mount Babel, and Redoubt Mountain) are predominantly Cruziana. Evidence of nutrient-rich substrates containing microbially induced sedimentary structures and the ichnospecies present confirm that the trace fossil bearing strata are part of the Cruziana ichnofacies and were deposited in a fully marine basin between fair weather wave base and storm wave base.     

Landscape metrics indicate differences in patterns and dominant controls of ribbon forests in the Rocky Mountains,USA

Question: Do landscape metrics reflect differences in dominant factors controlling ribbon forest patterns among sites? Location: West Flattop Mountain, Glacier National Park, Montana (Flattop); Medicine Bow Mountains, Wyoming (Medicine Bow); Park Range, Colorado (Park Range). Methods: High-resolution aerial photography was used to delineate ribbon forest patches, and to calculate landscape metrics to distinguish between long, narrow, regular patterns expected from strong microtopographic control, and smaller, more compact, and variable patterns expected from wind-snowdrift interactions. Results: All but two metrics were significantly different (P<0.05) among the three sites. The rank and magnitude of differences indicated that ribbons at Flattop and Park Range are more similar to each other than to those at Medicine Bow. Flattop ribbons were also more elongated, narrower and less variable than those at Park Range, suggesting differences in the type and strength of structural control. Previous research showed that Flattop ribbons occupy regular lithologic ridges, while our observations of ribbons and analysis of geologic maps suggests weaker and less consistent microtopographic control at Park Range, and dominant wind-snowdrift interactions with little to no microtopographic influence at Medicine Bow. Conclusions: Landscape metrics indicate differences in pattern among sites that reflect differences in dominant factors influencing ribbon forest development and maintenance. Explanations of ribbon forest dynamics are site-specific and are more complex than is currently recognized. The sites vary in the level of endogenous versus exogenous control of ribbon patterns, and consequently in the sensitivity of this phenomenon to climate.     

Culturable root endophyte communities are shaped by both warming and plant host identity in the Rocky Mountains,USA

Understanding the biogeographic patterns of root-associated fungi and their sensitivity to temperature may improve predictions of future changes in terrestrial biodiversity and associated ecosystem processes, but data are currently limited. Anticipating change will require combining observational data, which predict how climatic factors limit current species distributions, with direct manipulations of climate, which can isolate responses to specific climate variables. Root endophytes are common symbionts of plants, particularly in arctic and alpine environments, yet their responses to climate warming are not resolved. Here, we directly cultured endophytic fungi from roots collected along altitudinal gradients in replicated mountain watersheds and from a 27 y field warming experiment in the Rocky Mountains, USA, to improve understanding of climate impacts on fungal root endophytes. Fungal taxa that were common at high elevations declined most under climate warming, whereas low elevation dominants responded neutrally or increased with experimental warming. Altitudinal gradients in fungal communities were strongly specific to the plant host species. Specifically, Poa species had 25–60% greater fungal isolate abundance and 25–38% greater fungal diversity at high elevations than at low elevation sites. In contrast, Festuca thurberi had 64% lower fungal diversity on roots at high elevation than at low elevation. Our results help to improve understanding of the potential for climate change to alter plant-fungal interactions in mountain ecosystems.     

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.     

Influence of Tree Species on Forest Nitrogen Retention in the Catskill Mountains, New York, USA

This study examines the effect of four tree species on nitrogen (N) retention within forested catchments of the Catskill Mountains, New York (NY). We conducted a 300-day ¹⁵N field tracer experiment to determine how N moves through soil, microbial, and plant pools under different tree species and fertilization regimes. Samples were collected from single-species plots of American beech (Fagus grandifolia Ehrh.), eastern hemlock (Tsuga canadensis L.), red oak (Quercus rubra L.), and sugar maple (Acer saccharum Marsh). Using paired plots we compared the effects of ambient levels of N inputs (11 kg N/ha/y) to additions of 50 kg N/ha/y that began 1.5 years prior to and continued throughout this experiment. Total plot ¹⁵N recovery (litter layer, organic and mineral soil to 12 cm, fine roots, and aboveground biomass) did not vary significantly among tree species, but the distribution of sinks for ¹⁵N within the forest ecosystem did vary. Recovery in the forest floor was significantly lower in sugar maple stands compared to the other species. ¹⁵Nitrogen recovery was 22% lower in the fertilized plots compared to the ambient plots and red oak stands had the largest drop in ¹⁵N recovery as a result of N fertilization. Aboveground biomass became a significantly greater ¹⁵N sink with fertilization, although it retained less than 1% of the tracer addition. These results indicate that different forest types vary in the amount of N retention in the forest floor, and that forest N retention may change depending upon N inputs.     

Restoring the Vegetation of Mined Peatlands in the Southern Rocky Mountains of Colorado, U.S.A.

South Park is a high‐elevation, semi‐arid, treeless intermountain basin in central Colorado. A few extreme rich fens occur on the western margin and in the center of South Park where regional and local groundwater flow systems discharge to the ground surface. Over the past 40 years there has been extensive peat mining in these fens, but restoration methods have yet to be developed and successfully applied. The first part of this study compared the naturally reestablished vegetation on six mined peatlands with six pristine sites, while the second part of the study tested different revegetation techniques in 27 plots with varying depths to the water table. The six mined sites had only 30 plant species as compared with 122 species in the unmined sites; 43% of the species in the mined sites were not present in the undisturbed fens. Even after 40 years the sedges and willows that dominate the undisturbed sites were largely absent on the mined sites. The revegetation experiments seeded eight species, transplanted Carex aquatilis (water sedge) seedlings, transplanted rhizomes from six species, and transplanted four species of willow cuttings. Of the eight species seeded, only Triglochin maritima (arrowgrass) germinated and established seedlings. C. aquatilis seedlings, rhizome transplants of C. aquatilis, Kobresia simpliciuscula (elk sedge), and Juncus arcticus (arctic rush), and willow cuttings all had differing patterns of survival with respect to the annual maximum height of the water table. These results indicate that the dominant species can be successfully reintroduced to mined surfaces with the appropriate hydrologic conditions, but human intervention will be necessary to rapidly re‐establish these species. The slow rate of peat accumulation means that restoration of the mined fens will require hundreds, if not thousands, of years.     

Overstory-Understory Relationships along Forest Type and Environmental Gradients in the Spring Mountains of Southern Nevada, USA

Isolated forested mountains in deserts have numerous ecological and societal values, but land-management practices (e.g., fire-regime alteration) and climate change can affect forest composition. We analyzed tree overstory-understory relationships on 123 sites in the Spring Mountains within the Mojave Desert near Las Vegas, Nevada, USA to assess three hypotheses. We hypothesized that: the tree species comprising understories are less tolerant of fire than species in overstories, reflecting land-management practices of fire exclusion; mid-elevation forests have the lowest overstory:understory similarity because this zone could have maximum species mixing; and overstory:understory similarity is correlated with environmental gradients (consisting of 14 topographic and soil variables). We found that Pinus monophylla comprised greater relative canopy cover in understories of juniper (32% relative cover) and pinyon-juniper (78%) forests than it did in overstories of these forests (0% and 53%). Similarly, fire-intolerant Abies concolor had 6-fold greater understory than overstory cover in forests with overstories dominated by the fire-tolerant Pinus ponderosa. Overstory:understory S?rensen similarity averaged 43%?77% among six forest types, and there was little support for the supposition that similarity was lowest in mid-elevation forests. Distributions of individual overstory and understory species more closely corresponded with environmental gradients than did overstory:understory similarity. Results suggest that there is high potential for change in at least two of the six dominant forest types of the Spring Mountains. The direction of change (species of moist, higher elevation sites establishing in understories of drier forests) is the opposite of what would be expected for forest adaptation to the warmer, drier, more fire-prone conditions projected for the next century in the southwestern USA.