Subalpine Conifer Research Articles

One Unrecorded Endophytic Fungi from Sub-alpine Conifer, Rhizosphaera pini

An endophytic fungus, Rhizosphaera pini strain NIE7426, was isolated from the sub-alpine coniferous tree Abies nephrolepis in Mt. Nochu of Gangwon Province. It was characterized by macroscopic and microscopic features, as well as the internal transcribed spacer (ITS) 1, 2 and 5.8S sequences. All morphological and molecular features support the first recognition of this taxon in Korea. In addition, this study adds A. nephrolepis as a host plant R. pini.

Does typhoon disturbance in subalpine forest have long-lasting impacts on saproxylic fungi, bryophytes, and seedling regeneration on coarse woody debris?

Storm disturbance has a huge impact on subalpine forest ecosystems. Evaluating the long-term effects and mechanisms of storm disturbance is of great significance for conservation of subalpine forests as frequency and intensity of storms are increasing because of the climate change. Species of subalpine conifer tree genera such as Picea and Tsuga need coarse woody debris (CWD) such as logs and stumps for seedling colonization; thus, their establishment is greatly affected by the condition of CWD, which largely depends on the activity of fungal decomposer communities and saproxylic bryophyte communities. Because the process of CWD decay and associated ecological succession is known to take several decades in subalpine forests, we hypothesized that storm disturbance has long-lasting effects on fungal communities, their wood decay function, saproxylic bryophyte communities, and seedling establishment. To test this hypothesis, we surveyed fungal, bryophyte, and seedling communities at forest sites that had been disturbed by a large typhoon 55 years ago and compared them with those of surrounding undisturbed coniferous forest in central Japan. Contrary to our hypothesis, Illumina sequencing for the fungal ITS1 region in rDNA obtained from wood samples did not show that past disturbance had an obvious effect on current fungal communities. Also, the frequencies of occurrence of various wood decay types were not significantly different between disturbed and undisturbed plots. However, disturbance affected bryophyte communities, which had strong effects on the seedling densities of Picea jezoensis var. hondoensis, Tsuga diversifolia, and Betula spp. These results suggest that the previous typhoon disturbance in the focal area had long-lasting effects on biotic interactions and seedling regeneration on CWD, but these effects might not be attributable to the impacts of disturbance on CWD fungal communities and their wood decay function.

How Forest Gap and Elevation Shaped Abies faxoniana Rehd. et Wils. Regeneration in a Subalpine Coniferous Forest, Southwestern China

Focusing on the underlying ecological mechanisms of dominant species regeneration in forest gaps at a landscape scale can provide detailed understanding for gap-based forest management. The individual effects of forest gaps or elevation on the regeneration of Abies faxoniana Rehd. et Wils. are well known, although elucidating how gap characteristics and elevation concurrently influence regeneration remains an important challenge. In this paper, we present an explorative study using structural equation models (SEMs) to assess the direct and indirect effects of forest gaps and elevation on Abies faxoniana Rehd. et Wils. regeneration. Four of the predicted SEMs showed the following results: (1) Temperature, photosynthetic photon flux density (PPFD), soil total carbon, gap openness, shrub layer cover, herb layer cover, and moss layer thickness in forest gaps were associated with Abies faxoniana regeneration along an elevation gradient in subalpine coniferous forest. (2) Elevation had a generally negative and indirect effect on Abies faxoniana regeneration. Forest gaps positively affected regeneration when compared with non-gap plots and gap size was positively related to small tree regeneration density and the ratio of height to diameter at breast height (HD ratio) of the tallest Abies faxoniana small trees but was negatively related to Abies faxoniana sapling regeneration density. (3) In forest gaps, the Abies faxoniana sapling density and HD ratio of the tallest Abies faxoniana small trees were mainly indirectly influenced by elevation, and Abies faxoniana small tree regeneration density was directly associated with the dominance of the sapling regeneration density. In summary, Abies faxoniana regeneration was negatively and largely affected by elevation (total effect), although forest gaps enhanced Abies faxoniana regeneration by multiple pathways (direct and indirect effects).

Species- and Elevation-Dependent Growth Responses to Climate Warming of Mountain Forests in the Qinling Mountains, Central China

Climate warming is significantly affecting the composition and function of forest ecosystems. However, the forest responses to climate change in sub-humid and temperate areas are understudied compared with cold and semi-arid areas. Here, we investigate the radial-growth responses of two subalpine conifer species along an elevational gradient located in the Qinling Mountains, a sub-humid and temperate area situated in central China. Three sites dominated by larch (Larix chinensis Beissn.) and two other sites dominated by fir (Abies fargesii Franch.) located at different elevations were sampled. L. chinensis at a higher elevation showed more common and stronger climatic signals than A. fargesii at a lower elevation. The radial growth of L. chinensis was limited by low pre-growing season temperatures and showed an increasing growth trend in the last few years. On the other hand, A. fargesii growth was limited by summer water shortage and it was characterized by a declining trend in the most recent decade. Consequently, L. chinensis would benefit from climate warming, whereas A. fargesii could be regarded as a vulnerable tree species to warming-induced drought stress.

Structure and Composition of Subalpine Conifers in the Emerald Lake WATERSHED, Sequoia National Park, California

Abstract The Emerald Lake watershed forms a subalpine basin in the upper drainage of the Marble Fork of the Kaweah River in Sequoia National Park, California, with elevations ranging from 2804–3416 m. Five conifer species are present in the basin, with western white pine (Pinus monticola Douglas ex D. Don) as the most dominant species forming 71% of the stems and 82% of basal area censused in 1985 and 1986. Lodgepole pine (Pinus contorta Loudon subsp. murrayana (Grev. & Balf.) Critchf.) was the second most dominant in the watershed as measured by numbers of individuals and basal area, but was almost entirely restricted to mesic bench and wet meadow habitats in the lower elevations of the watershed. Foxtail pine (Pinus balfouriana Grev. & Balf.) comprised 9.5% of the stems and 13.7% of the basal area within the basin, but was largely present on higher north-facing ridgelines where it formed 52% of stems and 65% of basal area. Jeffrey pine (Pinus jeffreyi Grev. & Balf.), and red fir (Abies magnifica A. Murr...

It takes a few to tango: changing climate and fire regimes can cause regeneration failure of two subalpine conifers.

Environmental change is accelerating in the 21st century, but how multiple drivers may interact to alter forest resilience remains uncertain. In forests affected by large high-severity disturbances, tree regeneration is a resilience linchpin that shapes successional trajectories for decades. We modeled stands of two widespread western U.S. conifers, Douglas-fir (Pseudotsuga menziesii var. glauca), and lodgepole pine (Pinus contorta var. latifolia), in Yellowstone National Park (Wyoming, USA) to ask (1) What combinations of distance to seed source, fire return interval, and warming-drying conditions cause postfire tree-regeneration failure? (2) If postfire tree regeneration was successful, how does early tree density differ under future climate relative to historical climate? We conducted a stand-level (1ha) factorial simulation experiment using the individual-based forest process model iLand to identify combinations of fire return interval (11-100yr), distance to seed source (50-1,000m), and climate (historical, mid-21st century, late-21st century) where trees failed to regenerate by 30-yr postfire. If regeneration was successful, we compared stand densities between climate periods. Simulated postfire regeneration were surprisingly resilient to changing climate and fire drivers. Douglas-fir regeneration failed more frequently (55%) than lodgepole pine (28% and 16% for non-serotinous and serotinous stands, respectively). Distance to seed source was an important driver of regeneration failure for Douglas-fir and non-serotinous lodgepole pine; regeneration never failed when stands were 50m from a seed source and nearly always failed when stands were 1km away. Regeneration of serotinous lodgepole pine only failed when fire return intervals were ≤20yr and stands were far (1km) from a seed source. Warming climate increased regeneration success for Douglas-fir but did not affect lodgepole pine. If regeneration was successful, postfire density varied with climate. Douglas-fir and serotinous lodgepole pine regeneration density both increased under 21st-century climate but in response to different climate variables (growing season length vs. cold limitation). Results suggest that, given a warmer future with larger and more frequent fires, a greater number of stands that fail to regenerate after fires combined with increasing density in stands where regeneration is successful could produce a more coarse-grained forest landscape.

Germination Characterisitics of Abies koreana Wilson Seed in Subalpine Coniferous Forest by Collecting Location

Germination Characterisitics of Abies koreana Wilson Seed in Subalpine Coniferous Forest by Collecting Location

Site- and Species-Specific Influences on Sub-Alpine Conifer Growth in Mt. Rainier National Park, USA

Identifying the factors that influence the climate sensitivity of treeline species is critical to understanding carbon sequestration, forest dynamics, and conservation in high elevation forest/meadow ecotones. Using tree cores from four sub-alpine conifer species collected from three sides of Mt. Rainier, WA, USA, we investigated the influences of species identity and sites with different local climates on radial growth–climate relationships. We created chronologies for each species at each site, determined influential plant-relevant annual and seasonal climatic variables influencing growth, and investigated how the strength of climate sensitivity varied across species and location. Overall, similar climate variables constrained growth on all three sides of the mountain for each of the four study species. Summer warmth positively influenced radial growth, whereas snow, spring warmth, previous summer warmth, and spring humidity negatively influenced growth. We discovered only a few subtle differences in the climate sensitivity of co-occurring species at the same site and between the same species at different sites in pairwise comparisons. A model including species by climate interactions provided the best balance between parsimony and fit, but did not lead to substantially greater predictive power relative to a model without site or species interactions. Our results imply that at treeline in moist temperate regions like Mt. Rainier, the same climatic variables drive annual variation in growth across species and locations, despite species differences in physiology and site differences in mean climates.

Community reorganization in forest understories buried by volcanic tephra

AbstractDisturbance is a key factor contributing to community organization. Deposition of tephra (aerially transported volcanic ejecta) is a widespread disturbance of global relevance, but its effects on ecosystems have received limited attention. We studied forest understory community change for 30 yr following tephra deposition from the 1980 eruptions of Mount St. Helens (Washington State). Four old‐growth, subalpine conifer stands had a wide range of initial damage and patterns of community re‐development. We measured understory diversity, structure, and species composition and calculated relationships of plant cover with environment and cover of other plants. Overall, those communities that were altered greatly by tephra tended to converge with time on their pre‐eruption characteristics; however, substantial divergence occurred in some situations. For example, moss cover failed to reach pre‐eruption levels in all stands, whereas importance and diversity of woody plants sometimes greatly exceeded pre‐eruption values. Plant–environment relationships that were significant before the eruption disappeared and did not re‐develop. Smaller plants were more affected by environment than larger ones. Relationships before the eruption and also 30 yr after the eruption were primarily with other plant species, whereas relationships just after the eruption were with tephra depth and factors that modified its impact. Understory plant importance was usually lower beneath a tree canopy than in gaps, but there was little sign of interference from understory growth forms. Post‐eruption soil disturbance usually increased understory plant importance, while woody debris sometimes decreased herb and tree seedling cover. Tephra deposition, which did not immediately kill canopy trees, differed from the disturbances usually studied (e.g., fire, windthrow, bark beetles). Even so, these lessons from our study should be widely applicable: Similar species may respond differently; minor, early environmental differences may induce major, long‐term community change; successional trajectories may diverge from the pre‐disturbance community; and secondary disturbances may modify successional trajectories.

Lab and Field Warming Similarly Advance Germination Date and Limit Germination Rate for High and Low Elevation Provenances of Two Widespread Subalpine Conifers

Accurately predicting upslope shifts in subalpine tree ranges with warming requires understanding how future forest populations will be affected by climate change, as these are the seed sources for new tree line and alpine populations. Early life history stages are particularly sensitive to climate and are also influenced by genetic variation among populations. We tested the climate sensitivity of germination and initial development for two widely distributed subalpine conifers, using controlled-environment growth chambers with one temperature regime from subalpine forest in the Colorado Rocky Mountains and one 5 °C warmer, and two soil moisture levels. We tracked germination rate and timing, rate of seedling development, and seedling morphology for two seed provenances separated by ~300 m elevation. Warming advanced germination timing and initial seedling development by a total of ~2 weeks, advances comparable to mean differences between provenances. Advances were similar for both provenances and species; however, warming reduced the overall germination rate, as did low soil moisture, only for Picea engelmannii. A three-year field warming and watering experiment planted with the same species and provenances yielded responses qualitatively consistent with the lab trials. Together these experiments indicate that in a warmer, drier climate, P. engelmannii germination, and thus regeneration, could decline, which could lead to declining subalpine forest populations, while Pinus flexilis forest populations could remain robust as a seed source for upslope range shifts.

Comparison of photosynthetic traits of codominant subalpine conifers Abies veitchii and A. mariesii in central Japan

Evergreen conifers Abies mariesii and A. veitchii codominate in the subalpine zone in central Japan. This study compared the photosynthetic light response curves and related leaf traits of 1-year-old needles between the two species to examine whether photosynthetic traits of A. veitchii are more favorable for growth in bright conditions than those of A. mariesii. Saplings of the two species were sampled at forest edge (FE) and forest understory (FU). FE saplings of the two species showed more shade-intolerant traits (i.e., lower initial slope of photosynthesis light response curve, greater light compensation point and dark respiration rate) than FU saplings. Maximum photosynthetic rate and leaf nitrogen concentration were greater in A. veitchii than A. mariesii for both FE and FU saplings. Photosynthetic nitrogen use efficiency (PNUE) was also greater in A. veitchii than A. mariesii. On the contrary, leaf mass per area was greater in A. mariesii than A. veitchii for both FE and FU saplings. Therefore, this study showed that photosynthetic traits of A. veitchii are more favorable for growth in bright conditions compared with A. mariesii, and A. mariesii has more robust needles at the expense of PNUE.

亜高山帯針葉樹下の土壌有機物の性状と分解過程(第4報) : 土壌ヘテロトローフによる土壌有機物の分解

亜高山帯針葉樹下の土壌有機物の性状と分解過程(第4報) : 土壌ヘテロトローフによる土壌有機物の分解

亜高山帯針葉樹林下の土壌有機物の性状と分解過程 : 第1報 本地域の土壌の種類と土壌有機物の存在量について

亜高山帯針葉樹林下の土壌有機物の性状と分解過程 : 第1報 本地域の土壌の種類と土壌有機物の存在量について

亜高山帯針葉樹オオシラビソにおける標高にともなう生活史スケジュールの変化 : 山を上がるにしたがって木が小さくなる理由に関する新説( 個体群比較からの生活史研究)

亜高山帯針葉樹オオシラビソにおける標高にともなう生活史スケジュールの変化 : 山を上がるにしたがって木が小さくなる理由に関する新説( 個体群比較からの生活史研究)

Climate and environmental changes for the past 44 ka clarified by pollen and algae composition in the Ulleung Basin, East Sea (Japan Sea)

The pollen and algae records derived from core KCES-1 in the East Sea (Japan Sea) reveal vegetation and climate changes during the past 44 ka. From 44 to 36 cal ka BP, forest vegetation in the coastal regions of Ulleung Basin was composed predominantly of subalpine conifer forest, open grassland covered the exposed continental shelves, indicating a cool and dry climate. During the period of 36–28.3 cal ka BP, a warmer and wetter climate caused the contraction of subalpine conifer forest in the coastal regions of Ulleung Basin, and the expansion of marsh in the exposed continental shelves. From 28.3 to 17.6 cal ka BP, the sea-level decline restricted the herbs pollen originates in the exposed shelf of East China Sea and Yellow Sea inflow into the East Sea (Japan Sea), and caused the reduction of herbs pollen in the site location. During the time interval of 17.6–15.1 cal ka BP, a great expansion of subalpine conifer forest can be regarded as the consequence of enhanced East Asian winter monsoon (EAWM). The obvious increase in temperate deciduous broadleaved trees, spores and algae can be attributable to the rising of sea level and the warming of climate from 15.1 to 12.1 cal ka BP. During the period of 12.1 to 5.3 cal ka BP, broadleaved taxa increased, especially during the time period of 9.5–5.3 cal ka BP, evergreen broadleaved trees pollen reach the highest values throughout the core, as a result of the Holocene climate optimum condition. Vegetation changes controlled by humid impacts occurred at the late Holocene, as indicated by an increase in Pinus pollen.

Mid-Late Miocene vegetation and environments in Southeast China: Insights from a marine palynological record in northwestern Taiwan

Taiwan Island represents the exposed accretionary prism developed by eastward subduction of South China Sea (SCS) oceanic lithosphere beneath the Philippine Sea Plate since the middle Miocene. There are many well-documented data about the biostratigraphic correlation, except for palynological studies. Here, we present a new palynological record of mid-Late Miocene deposits from the Chuhuangkeng section, Miaoli, northwestern Taiwan, which reveals a vegetation and paleoelevation reconstruction in Southeast China, during the Middle to Late Miocene, corresponding to the time interval of about 14.5–5.5Ma. Most samples yielded well-preserved palynomorphs, and the palynological results suggest that the dominant vegetation was the mixed coniferous-broadleaved forests in the Coastal Mountains of Southeast China (CMSC) during the Mid-Late Miocene. Moreover, based on large amounts of Tsuga and Picea pollen, we hypothesized that a subalpine conifer forest then flourished in the CMSC, and the estimated maximum paleoelevation might be up to ∼3000m above sea level (a.s.l.) during the Mid-Late Miocene, being 1000–1500m higher than today. These findings offer a new perspective to comprehend the paleotopographic evolution of China and the source area of Taiwan’s spruce forests. Additionally, the high abundance of Pinus, Tsuga, Picea pollen and fern spores also suggests an enhancement of the EASM during the Mid-Late Miocene.

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.

Endogone corticioides sp. nov. from subalpine conifer forests in Japan and China, and its multi-locus phylogeny

Endogone corticioides sp. nov. was described from subalpine conifer forests in Japan and China. This species is similar to E. pisiformis in its yellowish sporocarps and zygosporangia on the equal empty gametangia. Endogone corticioides was clearly distinguished from E. pisiformis based on its resupinate sporocarp, the presence of hyphal peg-like structures on the peridium, and abundant thick-walled hyphae in the gleba. Phylogenetic analysis of the small and large subunit nuclear ribosomal RNA and the elongation factor-1α genes suggested that E. corticioides is closely related to E. pisiformis. Limited vegetative growth from the surface-sterilized sporocarps on agar media was shown.

An observational constraint on stomatal function in forests: evaluating coupled carbon and water vapor exchange with carbon isotopes in the Community Land Model (CLM4.5)

Abstract. Land surface models are useful tools to quantify contemporary and future climate impact on terrestrial carbon cycle processes, provided they can be appropriately constrained and tested with observations. Stable carbon isotopes of CO2 offer the potential to improve model representation of the coupled carbon and water cycles because they are strongly influenced by stomatal function. Recently, a representation of stable carbon isotope discrimination was incorporated into the Community Land Model component of the Community Earth System Model. Here, we tested the model's capability to simulate whole-forest isotope discrimination in a subalpine conifer forest at Niwot Ridge, Colorado, USA. We distinguished between isotopic behavior in response to a decrease of δ13C within atmospheric CO2 (Suess effect) vs. photosynthetic discrimination (Δcanopy), by creating a site-customized atmospheric CO2 and δ13C of CO2 time series. We implemented a seasonally varying Vcmax model calibration that best matched site observations of net CO2 carbon exchange, latent heat exchange, and biomass. The model accurately simulated observed δ13C of needle and stem tissue, but underestimated the δ13C of bulk soil carbon by 1–2 ‰. The model overestimated the multiyear (2006–2012) average Δcanopy relative to prior data-based estimates by 2–4 ‰. The amplitude of the average seasonal cycle of Δcanopy (i.e., higher in spring/fall as compared to summer) was correctly modeled but only when using a revised, fully coupled An − gs (net assimilation rate, stomatal conductance) version of the model in contrast to the partially coupled An − gs version used in the default model. The model attributed most of the seasonal variation in discrimination to An, whereas interannual variation in simulated Δcanopy during the summer months was driven by stomatal response to vapor pressure deficit (VPD). The model simulated a 10 % increase in both photosynthetic discrimination and water-use efficiency (WUE) since 1850 which is counter to established relationships between discrimination and WUE. The isotope observations used here to constrain CLM suggest (1) the model overestimated stomatal conductance and (2) the default CLM approach to representing nitrogen limitation (partially coupled model) was not capable of reproducing observed trends in discrimination. These findings demonstrate that isotope observations can provide important information related to stomatal function driven by environmental stress from VPD and nitrogen limitation. Future versions of CLM that incorporate carbon isotope discrimination are likely to benefit from explicit inclusion of mesophyll conductance.

Continuous, long-term, high-frequency thermal imaging of vegetation: Uncertainties and recommended best practices

Leaf temperature is an elementary driver of plant physiology, ecology and ecosystem productivity. Individual leaf temperature may deviate strongly from air temperature, and may vary throughout the canopy. Measurements of leaf temperature, conducted at a high spatial and temporal resolution, can improve our understanding of leaf water loss, stomatal conductance, photosynthetic rates, phenology, and atmosphere-ecosystem exchanges. However, continuous high-resolution measurement of leaf temperature outside of a controlled environment is difficult and rarely done. Here, thermal infrared cameras are used to measure leaf temperatures. We describe two long-term field measurement sites: one in a temperature deciduous forest, and the other in a subalpine conifer forest. The considerations and constraints for deploying such cameras are discussed and the temperature errors are typically +/–1°C or smaller (σ=0.60°C, 2σ=1.20°C). Lastly, we compare leaf temperature by species and height at hourly to multi-seasonal timescales and show that on average, leaf temperature is warmer than air temperature in a temperate forest. Leaf temperature can be uniform or heterogeneous across a scene, depending on canopy structure, leaf habit, and meteorology. With this data, we verify that leaf temperature follows classic expectations, yet exhibits noteworthy departures that require additional study and theoretical consideration.