Geology topics

Lara M. Kueppers

Publications and source records attributed to Lara M. Kueppers.

10 recordsLinked to original sources

USGS2020

Warming of alpine tundra enhances belowground production and shifts community towards resource acquisition traits

Climate warming is expected to stimulate plant growth in high‐elevation and high‐latitude ecosystems, significantly increasing aboveground net primary production (ANPP). However, the effects of simultaneous changes in temperature, snowmelt timing, and summer water availability on total net primary production (NPP)—and elucidation of both above‐ and belowground responses—remain an important area in need of further study. In particular, measures of belowground net primary productivity (BNPP) are required to understand whether ANPP changes reflect changes in allocation or are indicative of a whole plant NPP response. Further, plant functional traits provide a key way to scale from the individual plant to the community level and provide insight into drivers of NPP responses to environmental change. We used infrared heaters to warm an alpine plant community at Niwot Ridge, Colorado, and applied supplemental water to compensate for soil water loss induced by warming. We measured ANPP, BNPP, and leaf and root functional traits across treatments after 5 yr of continuous warming. Community‐level ANPP and total NPP (ANPP + BNPP) did not respond to heating or watering, but BNPP increased in response to heating. Heating decreased community‐level leaf dry matter content and increased total root length, indicating a shift in strategy from resource conservation to acquisition in response to warming. Water use efficiency (WUE) decreased with heating, suggesting alleviation of moisture constraints that may have enabled the plant community to increase productivity. Heating may have decreased WUE by melting snow earlier and creating more days early in the growing season with adequate soil moisture, but stimulated dry mass investment in roots as soils dried down later in the growing season. Overall, this study highlights how ANPP and BNPP responses to climate change can diverge, and encourages a closer examination of belowground processes, especially in alpine systems, where the majority of NPP occurs belowground.

Colorado↗

USGS2020

Colonisation of the alpine tundra by trees: Alpine neighbours assist late-seral but not early-seral conifer seedlings

Background Climate change is projected to alter the elevation and latitude of treeline globally. Seed germination and seedling survival are critical controls on treeline expansion. Neighbouring alpine plants, either through competition for resources or through altered microclimate, also affect seedlings emerging in the alpine zone. With warming, alpine plant species may interact with each other more or less strongly. Aims To determine whether establishing tree seedlings and an alpine herb are similarly sensitive to alpine plant neighbours under ambient and altered climate. Methods We imposed active heating, watering, and removed all plants adjacent to emerging conifer seedlings and an alpine herb. Results Picea engelmannii seedlings showed lower survival compared with Pinus flexilis 3 weeks following neighbour removal, and after 1 year only survived in watered plots. Pinus seedlings responded to neighbour removal by lowering the quantum yield of photosynthesis (ϕ PSII ). Contrary to expectations from the stress gradient hypothesis, survival was reduced without neighbours near the low-elevation range limit of Chionophila jamesii . Conclusions Pinus flexilis has higher expansion potential into the alpine, while Picea engelmannii requires moist conditions that could be facilitated by neighbours to expand its range. This implies likely range expansion by P. flexilis with consequences for alpine plant diversity and ecosystem function.

Colorado↗

USGS2018

Snowmelt timing regulates community composition, phenology, and physiological performance of alpine plants

The spatial patterning of alpine plant communities is strongly influenced by the variation in physical factors such as temperature and moisture, which are strongly affected by snow depth and snowmelt patterns. Earlier snowmelt timing and greater soil-moisture limitations may favor wide-ranging species adapted to a broader set of ecohydrological conditions than alpine-restricted species. We asked how plant community composition, phenology, plant water relations, and photosynthetic gas exchange of alpine-restricted and wide-ranging species differ in their responses to a ca. 40-day snowmelt gradient in the Colorado Rocky Mountains ( Lewisia pygmaea , Sibbaldia procumbens , and Hymenoxys grandiflora were alpine-restricted and Artemisia scopulorum , Carex rupestris , and Geum rossii were wide-ranging species). As hypothesized, species richness and foliar cover increased with earlier snowmelt, due to a greater abundance of wide-ranging species present in earlier melting plots. Flowering initiation occurred earlier with earlier snowmelt for 12 out of 19 species analyzed, while flowering duration was shortened with later snowmelt for six species (all but one were wide-ranging species). We observed >50% declines in net photosynthesis from July to September as soil moisture and plant water potentials declined. Early-season stomatal conductance was higher in wide-ranging species, indicating a more competitive strategy for water acquisition when soil moisture is high. Even so, there were no associated differences in photosynthesis or transpiration, suggesting no strong differences between these groups in physiology. Our findings reveal that plant species with different ranges (alpine-restricted vs. wide-ranging) could have differential phenological and physiological responses to snowmelt timing and associated soil moisture dry-down, and that alpine-restricted species’ performance is more sensitive to snowmelt. As a result, alpine-restricted species may serve as better indicator species than their wide-ranging heterospecifics. Overall, alpine community composition and peak % cover are strongly structured by spatio-temporal patterns in snowmelt timing. Thus, near-term, community-wide changes (or variation) in phenology and physiology in response to shifts in snowmelt timing or rates of soil dry down are likely to be contingent on the legacy of past climate on community structure.

Frontiers in Plant Science↗

USGS2017

Seed origin and warming constrain lodgepole pine recruitment, slowing the pace of population range shifts

Understanding how climate warming will affect the demographic rates of different ecotypes is critical to predicting shifts in species distributions. Here we present results from a common garden, climate change experiment in which we measured seedling recruitment of lodgepole pine, a widespread North American conifer that is also planted globally. Seeds from a low-elevation provenance had greater recruitment to their third year (by 323%) than seeds from a high-elevation provenance across sites within and above its native elevation range and across climate manipulations. Heating reduced (by 49%) recruitment to the third year of both low- and high-elevation seed sources across the elevation gradient, while watering alleviated some of the negative effects of heating (108% increase in watered plots). Demographic models based on recruitment data from the climate manipulations and long-term observations of adult populations revealed that heating could effectively halt modeled upslope range expansion except when combined with watering. Simulating fire and rapid post-fire forest recovery at lower elevations accelerated lodgepole pine expansion into the alpine, but did not alter final abundance rankings among climate scenarios. Regardless of climate scenario, greater recruitment of low-elevation seeds compensated for longer dispersal distances to treeline, assuming colonization was allowed to proceed over multiple centuries. Our results show that ecotypes from lower elevations within a species’ range could enhance recruitment and facilitate upslope range shifts with climate change.

Global Change Biology↗

USGS2017

Growth strategies and threshold responses to water deficit modulate effects of warming on tree seedlings from forest to alpine

1.Predictions of upslope range shifts for tree species with warming are based on assumptions of moisture stress at lower elevation limits and low temperature stress at high elevation limits. However, recent studies have shown that warming can reduce tree seedling establishment across the entire gradient from subalpine forest to alpine via moisture limitation. Warming effects also vary with species, potentially resulting in community shifts in high elevation forests. 2.We examined the growth and physiology underlying effects of warming on seedling demographic patterns. We evaluated dry mass (DM), root length, allocation above- and belowground, and relative growth rate (RGR) of whole seedlings, and their ability to avoid or endure water stress via water-use efficiency and resisting turgor loss, for Pinus flexilis, Picea engelmannii and Pinus contorta seeded below, at, and above treeline in experimentally warmed, watered, and control plots in the Rocky Mountains, USA. We expected that growth and allocation responses to warming would relate to moisture status and that variation in drought tolerance traits would explain species differences in survival rates. 3.Across treatments and elevations, seedlings of all species had weak turgor-loss resistance, and growth was marginal with negative RGR in the first growth phase (-0.01 to -0.04 g/g/d). Growth was correlated with soil moisture, particularly in the relatively small-seeded P. contorta and P. engelmannii. P. flexilis, known to have the highest survivorship, attained the greatest DM and longest root but was also the slowest growing and most water-use-efficient. This was likely due to its greater reliance on seed reserves. Seedlings developed 15% less total DM, 25% less root DM, and 11% shorter roots in heated compared to unheated plots. Higher temperatures slightly increased DM, root length and RGR where soils were wettest, but more strongly decreased these variables under drier conditions. 4.Synthesis: The surprising heat-inhibition of tree seedling establishment at the cold edge of forests appears to have a physiological basis: newly germinated seedlings have poor moisture stress tolerance, which appears related to marginal initial growth and heavy reliance on seed reserves. Variation in these attributes among tree species at treeline helps explain their different climate responses.

Journal of Ecology↗

USGS2017

Declines in low-elevation subalpine tree populations outpace growth in high-elevation populations with warming

Species distribution shifts in response to climate change require that recruitment increase beyond current range boundaries. For trees with long life spans, the importance of climate-sensitive seedling establishment to the pace of range shifts has not been demonstrated quantitatively. Using spatially explicit, stochastic population models combined with data from long-term forest surveys, we explored whether the climate-sensitivity of recruitment observed in climate manipulation experiments was sufficient to alter populations and elevation ranges of two widely distributed, high-elevation North American conifers. Empirically observed, warming-driven declines in recruitment led to rapid modelled population declines at the low-elevation, ‘warm edge’ of subalpine forest and slow emergence of populations beyond the high-elevation, ‘cool edge’. Because population declines in the forest occurred much faster than population emergence in the alpine, we observed range contraction for both species. For Engelmann spruce, this contraction was permanent over the modelled time horizon, even in the presence of increased moisture. For limber pine, lower sensitivity to warming may facilitate persistence at low elevations – especially in the presence of increased moisture – and rapid establishment above tree line, and, ultimately, expansion into the alpine. Synthesis . Assuming 21st century warming and no additional moisture, population dynamics in high-elevation forests led to transient range contractions for limber pine and potentially permanent range contractions for Engelmann spruce. Thus, limitations to seedling recruitment with warming can constrain the pace of subalpine tree range shifts.

Journal of Ecology↗

USGS2016

Warming and provenance limit tree recruitment across and beyond the elevation range of subalpine forest

Climate niche models project that subalpine forest ranges will extend upslope with climate warming. These projections assume that the climate suitable for adult trees will be adequate for forest regeneration, ignoring climate requirements for seedling recruitment, a potential demographic bottleneck. Moreover, local genetic adaptation is expected to facilitate range expansion, with tree populations at the upper forest edge providing the seed best adapted to the alpine. Here, we test these expectations using a novel combination of common gardens, seeded with two widely distributed subalpine conifers, and climate manipulations replicated at three elevations. Infrared heaters raised temperatures in heated plots, but raised temperatures more in the forest than at or above treeline because strong winds at high elevation reduced heating efficiency. Watering increased season-average soil moisture similarly across sites. Contrary to expectations, warming reduced Engelmann spruce recruitment at and above treeline, as well as in the forest. Warming reduced limber pine first-year recruitment in the forest, but had no net effect on fourth-year recruitment at any site. Watering during the snow-free season alleviated some negative effects of warming, indicating that warming exacerbated water limitations. Contrary to expectations of local adaptation, low-elevation seeds of both species initially recruited more strongly than high-elevation seeds across the elevation gradient, although the low-provenance advantage diminished by the fourth year for Engelmann spruce, likely due to small sample sizes. High- and low-elevation provenances responded similarly to warming across sites for Engelmann spruce, but differently for limber pine. In the context of increasing tree mortality, lower recruitment at all elevations with warming, combined with lower quality, high-provenance seed being most available for colonizing the alpine, portends range contraction for Engelmann spruce. The lower sensitivity of limber pine to warming indicates a potential for this species to become more important in subalpine forest communities in the coming centuries.

Colorado↗

USGS2015

Linking carbon and water limitations to drought-induced mortality of Pinus flexilis seedlings

Survival of tree seedlings at high elevations has been shown to be limited by thermal constraints on carbon balance, but it is unknown if carbon relations also limit seedling survival at lower elevations, where water relations may be more important. We measured and modeled carbon fluxes and water relations in first-year Pinus flexilis seedlings in garden plots just beyond the warm edge of their natural range, and compared these with dry-mass gain and survival across two summers. We hypothesized that mortality in these seedlings would be associated with declines in water relations, more so than with carbon-balance limitations. Rather than gradual declines in survivorship across growing seasons, we observed sharp, large-scale mortality episodes that occurred once volumetric soil-moisture content dropped below 10%. By this point, seedling water potentials had decreased below −5 MPa, seedling hydraulic conductivity had decreased by 90% and seedling hydraulic resistance had increased by >900%. Additionally, non-structural carbohydrates accumulated in aboveground tissues at the end of both summers, suggesting impairments in phloem-transport from needles to roots. This resulted in low carbohydrate concentrations in roots, which likely impaired root growth and water uptake at the time of critically low soil moisture. While photosynthesis and respiration on a leaf area basis remained high until critical hydraulic thresholds were exceeded, modeled seedling gross primary productivity declined steadily throughout the summers. At the time of mortality, modeled productivity was insufficient to support seedling biomass-gain rates, metabolism and secondary costs. Thus the large-scale mortality events that we observed near the end of each summer were most directly linked with acute, episodic declines in plant hydraulic function that were linked with important changes in whole-seedling carbon relations.

Tree Physiology↗

USGS2015

Moisture rivals temperature in limiting photosynthesis by trees establishing beyond their cold-edge range limit under ambient and warmed conditions

Climate change is altering plant species distributions globally, and warming is expected to promote uphill shifts in mountain trees. However, at many cold-edge range limits, such as alpine treelines in the western United States, tree establishment may be colimited by low temperature and low moisture, making recruitment patterns with warming difficult to predict. We measured response functions linking carbon (C) assimilation and temperature- and moisture-related microclimatic factors for limber pine ( Pinus flexilis ) seedlings growing in a heating × watering experiment within and above the alpine treeline. We then extrapolated these response functions using observed microclimate conditions to estimate the net effects of warming and associated soil drying on C assimilation across an entire growing season. Moisture and temperature limitations were each estimated to reduce potential growing season C gain from a theoretical upper limit by 15–30% ( c . 50% combined). Warming above current treeline conditions provided relatively little benefit to modeled net assimilation, whereas assimilation was sensitive to either wetter or drier conditions. Summer precipitation may be at least as important as temperature in constraining C gain by establishing subalpine trees at and above current alpine treelines as seasonally dry subalpine and alpine ecosystems continue to warm.

Colorado↗

USGS2012

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 m 3 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 m 3 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.

Colorado↗