Encroachment drives facilitation at alpine shrublines

Ongoing encroachment is driving recent alpine shrubline dynamics globally,but the role of shrub-shrub interactions in shaping shrublines and their relationships with stem density changes remain poorly understood.Here,the size and age of shrubs from 26 Salix shrubline populations along a 900-km latitudinal gradient(30°-38°N)were measured and mapped across the eastern Tibetan Plateau.Point pattern analyses were used to quantify the spatial distribution patterns of juveniles and adults,and to assess spatial associations between them.Mean intensity of univariate and bivariate spatial patterns was related to biotic and abiotic variables.Bivariate mark correlation functions with a quantitative mark(shrub height,basal stem diameter,crown width)were also employed to investigate the spatial relationships between shrub traits of juveniles and adults.Structural equation models were used to explore the relationships among conspecific interactions,patterns,shrub traits and recruitment dynamics under climate change.Most shrublines showed clustered patterns,suggesting the existence of conspecific facilitation.Clustered patterns of juveniles and conspecific interactions(potentially facilitation)tended to intensify with increasing soil moisture stress.Summer warming before 2010 triggered positive effects on population interactions and spatial patterns via increased shrub recruitment.However,summer warming after2010 triggered negative effects on interactions through reduced shrub recruitment.Therefore,shrub recruitment shifts under rapid climate change could impact spatial patterns,alter conspecific interactions and modify the direction and degree of shrublines responses to climate.These changes would have profound implications for the stability of alpine woody ecosystems.

Growth phenology adjusts to seasonal changes in water availability in coexisting evergreen and deciduous mediterranean oaks

Different leaf(evergreen vs.deciduous habit)and xylem(diffuse-vs.ring-porous wood)traits represent contrasting strategies to face seasonal changes in water availability and temperature.However,how contrasting leaf and xylem habits of coexisting tree species affect stem wood formation and tree-ring development remains poorly understood.Here,we investigated the spatio-temporal patterns of wood formation in two deciduous oaks(Quercus faginea and Quercus petraea)and two evergreen oaks(Quercus ilex and Quercus suber)coexisting in seasonally dry Mediterranean forests along an aridity gradient in Spain.We hypothesized that growth responses to drought and intra-and inter-annual growth patterns would differ between functional groups.We simulated intra-and interannual growth using a modified version of the Vaganov-Shashkin(VS)process-based,growth model.The VS model simulations were used to estimate growth changes under a high emission scenario(RCP 8.5)for the current distribution of the study oak species and to forecast their future performance under warm(4.8℃)conditions in the Iberian Peninsula.Our simulations indicate that climate warming would induce a shortening of the ringgrowth season and a reduction of radial growth in evergreen and deciduous Mediterranean oaks,particularly in dry sites from southern and eastern Iberia currently occupied by Q.ilex and Q.faginea.Evergreen oaks may better recover after dry periods than deciduous oaks by resuming growth after the summer drought.Low soil water availability in spring would be more detrimental to growth of deciduous oaks.Process-based growth models should be refined and validated to better forecast changes in tree growth as a function of climate.

Growth plasticity of conifers did not avoid declining resilience to soil and atmospheric droughts during the 20th century

Background:Plasticity in response to environmental drivers can help trees cope with droughts.However,our understanding of the importance of plasticity and physiological adjustments in trees under global change is limited.Methods:We used the International Tree-Ring Data Bank(ITRDB)to examine 20th century growth responses in conifer trees during(resistance)and following(resilience)years of severe soil and atmospheric droughts occurring in isolation or as compound events.Growth resilience indices were calculated using observed growth divided by expected growth to avoid spurious correlations,in which the expected values were obtained by the autoregressive moving average(ARIMA)model.We used high atmospheric vapour pressure deficit(VPD)to select years of atmospheric drought and low annual values of the Standardized Precipitation-Evapotranspiration Index(SPEI)to select years with soil drought.We acquired the sensitivities(i.e.,the slopes of the relationships)by fitting the resilience indices as a function of environmental drivers,and assessed how these sensitivities changed over time for different types of drought events using linear mixed models.We also checked whether plasticity in growth responses was sufficient to prevent long-term trends of growth reductions during or after severe droughts.We acknowledge that by focusing on the response of surviving trees from the ITRDB we are potentially biasing our results towards higher resilience,as stand level responses(e.g.,mortality)may result in lowered competition after the disturbance event.Results:Sensitivities of resilience to VPD and SPEI changed throughout the 20th century,with the directions of these changes often reversing in the second half of the century.For the 1961–2010 period,changing sensitivities had positive effects on resilience,especially following years of high-VPD and compound events,avoiding growth losses that would have occurred if sensitivities had remained constant.Despite sensitivity changes,resilience was still lower at the end of the 20th century compared to the beginning of the century.Conclusions:Future adjustments to low-SPEI and high-VPD events are likely to continue to compensate for the trends in climate only partially,leading to further generalized reductions in tree growth of conifers.An improved understanding of these plastic adjustments and their limits,as well as potential compensatory processes at the stand level,is needed to project forest responses to climate change.

Different xylogenesis responses to atmospheric water demand contribute to species coexistence in a mixed pine–oak forest

Seasonal patterns of wood formation(xylogenesis)remain understudied in mixed pine-oak forests despite their contribution to tree coexistence through temporal niche complementarity.Xylogenesis was assessed in three pine species(Pinus cembroides,Pinus leiophylla,Pinus engelmannii)and one oak(Quercus grisea)coexisting in a semi-arid Mexican forest.The main xylogenesis phases(production of cambium cells,radial enlargement,cell-wall thickening and maturation)were related to climate data considering 5-15-day temporal windows.In pines,cambium activity maximized from mid-March to April as temperature and evaporation increased,whereas cell radial enlargement peaked from April to May and was constrained by high evaporation and low precipitation.Cell-wall thickening peaked from June to July and in August-September as maximum temperature and vapour pressure deficit(VPD)increased.Maturation of earlywood and latewood tracheids occurred in May-June and June-July,enhanced by high minimum temperatures and VPD in P.engelmannii and P.leiophylla.In oak,cambial onset started in March,constrained by high minimum temperatures,and vessel radial enlargement and radial increment maximized in April as temperatures and evaporation increased,whereas early wood vessels matured from May to June as VPD increased.Overall,15-day wet conditions enhanced cell radial enlargement in P.leiophylla and P.engelmannii,whereas early-summer high 15-day temperature and VPD drove cell-wall thickening in P.cembroides.Warm night conditions and high evaporation rates during spring and summer enhanced growth.An earlier growth peak in oak and a higher responsiveness to spring-summer water demand in pines contributed to their coexistence.

Tree growth is more limited by drought in rear-edge forests most of the times

Background:Equatorward,rear-edge tree populations are natural monitors to estimate species vulnerability to climate change.According to biogeographical theory,exposition to drought events increases with increasing aridity towards the equator and the growth of southern tree populations will be more vulnerable to drought than in central populations.However,the ecological and biogeographical margins can mismatch due to the impact of ecological factors(topography,soils)or tree-species acclimation that can blur large-scale geographical imprints in trees responses to drought making northern populations more drought limited.Methods:We tested these ideas in six tree species,three angiosperms(Fagus sylvatica,Quercus robur,Quercus petraea)and three gymnosperms(Abies alba,Pinus sylvestris and Pinus uncinata)by comparing rear-edge tree populations subjected to different degrees of aridity.We used dendrochronology to compare the radial-growth patterns of these species in northern,intermediate,and southern tree populations at the continental rear edge.Results and conclusions:We found marked variations in growth variability between species with coherent patterns of stronger drought signals in the tree-ring series of the southern populations of F.sylvatica,P.sylvestris,and A.alba.This was also observed in species from cool-wet sites(P.uncinata and Q.robur),despite their limited responsiveness to drought.However,in the case of Q.petraea the intermediate population showed the strongest relationship to drought.For drought-sensitive species as F.sylvatica and P.sylvestris,southern populations presented more variable growth which was enhanced by cool-wet conditions from late spring to summer.We found a trend of enhanced vulnerability to drought in these two species.The response of tree growth to drought has a marked biogeographical component characterized by increased drought sensitivity in southern populations even within the species distribution rear edge.Nevertheless,the relationship between tree growth and drought varied between species suggesting that biogeographical and ecological limits do not always overlap as in the case of Q.petraea.In widespread species showing enhanced vulnerability to drought,as F.sylvatica and P.sylvestris,increased vulnerability to climate warming in their rear edges is forecasted.Therefore,we encourage the monitoring and conservation of such marginal tree populations.

Contrasting climate drivers of seasonal growth in western vs. eastern Mexican mountain conifer forests

Hydroclimate affects the radial growth responses of trees, but the drivers of their spatial and population variability are not sufficiently understood. We addressed this issue by sampling several conifer populations located at the same latitude, but at different longitude and elevation in western(W) and eastern(E) Mexican regions. We used dendroecology to disentangle how earlywood width(EW), latewood width(LW) and adjusted LW(LWadj),i.e. the residuals after removing EW influences on LW, responded to climate variables(temperature and precipitation), climate indices(Southern Oscillation Index, SOI, Nino 3.4, Pacific Decadal Oscillation, PDO) and a drought index(Standardised Precipitation-Evapotranspiration Index, SPEI). The W species(Pinus herrerae Martinez, Pinus durangensis Martinez, Abies durangensis Martínez and Cupressus lusitanica Mill.) showed lower growth rates than the E species(Pinus hartwegii Lindl., Picea mexicana Martinez, Pseudotsuga menziesii(Mirb.) Franco and Abies vejari Martinez). Growth in W benefits mostly from high precipitation in the prior winter and current spring and it is limited by high temperatures in spring, whereas growth in the E showed similar but weaker responses.Furthermore, positive(negative) correlations were found in radial growth with the Nino 3.4(SOI) and the PDO from the prior to current autumns, which were again stronger in absolute terms in the W than in the E regions,excepting SOI in summer. In the W, P. durangensis and C. lusitanica were the least and most responsive species to spring drought, respectively;whilst P. menziesii and A. vejari were very responsive to spring drought compared to P. hartwegii in the E. Our results suggest greater responsiveness to hydroclimate and atmospheric patterns in the W than in the E region. These findings allow better interpretations of future changes in growth and composition in Mexican conifer forests, considering that climate models forecast warmer spring conditions and increased water shortage.

Linking leaf elemental traits to biomass across forest biomes in the Himalayas

Plants require a number of essential elements in different proportions for ensuring their growth and development.The elemental concentrations in leaves reflect the functions and adaptations of plants under specific environmental conditions.However,less is known about how the spectrum of leaf elements associated with resource acquisition,photosynthesis and growth regulates forest biomass along broad elevational gradients.We examined the influence of leaf element distribution and diversity on forest biomass by analyzing ten elements(C,N,P,K,Ca,Mg,Zn,Fe,Cu,and Mn)in tree communities situated every 100 meters along an extensive elevation gradient,ranging from the tropical forest(80 meters above sea level)to the alpine treeline(4200 meters above sea level)in the Kangchenjunga Landscape in eastern Nepal Himalayas.We calculated communityweighted averages(reflecting dominant traits governing biomass,i.e.,mass-ratio effect)and functional divergence(reflecting increased trait variety,i.e.,complementarity effect)for leaf elements in a total of 1,859 trees representing 116 species.An increasing mass-ratio effect and decreasing complementarity in leaf elements enhance forest biomass accumulation.A combination of elements together with elevation explains biomass(52.2%of the variance)better than individual elemental trait diversity(0.05%to 21%of the variance).Elevation modulates trait diversity among plant species in biomass accumulation.Complementarity promotes biomass at lower elevations,but reduces biomass at higher elevations,demonstrating an interaction between elevation and complementarity.The interaction between elevation and mass-ratio effect produces heterogeneous effects on biomass along the elevation gradient.Our research indicates that biomass accumulation can be disproportionately affected by elevation due to interactions among trait diversities across vegetation zones.While higher trait variation enhances the adaptation of species to environmental changes,it reduces biomass accumulation,especially at higher elevations.

No benefits from warming even for subnival vegetation in the central Himalayas

普遍认为变暖有利于高山植被生态系统,并已经导致冰缘植物向高海拔的扩张.珠峰山谷金字塔气象站(海拔5050 m)1994~2018的气象记录显示,高海拔山区呈显著的暖干化趋势.然而,我们依然不清楚暖干化是否对冰缘植被更新和分布造成显著影响?本研究在喜马拉雅山中段3个山谷建立了8块(30 m×120 m)滇藏方枝柏(Juniperus indica)和高山柏(Juniperus squamata)灌木线样地(海拔介于4344~5074 m之间).样地调查揭示,自20世纪90年代之前所有样地种群更新呈增加的趋势,然而90年代以来更新持续下降.种群更新与春季、夏季温度之间呈现显著的负相关关系,暗示了暖干化加剧了水分胁迫,并限制了种群的更新.另外,近60年来所有灌木线位置稳定.近30年遥感数据也证实,研究区冰缘植被无变绿趋势,甚至出现2%-6%的植被生长减弱现象.因此,进一步研究应该探讨冰缘植被生长减弱对喜马拉雅山区水循环和生物多样性的影响.