Radial growth response to climate change along the latitudinal range of the world's southernmost conifer in southern South America

Aim

We examined whether and how tree radial‐growth responses to climate have changed for the world's southernmost conifer species throughout its latitudinal distribution following rapid climate change in the second half of the 20th century.

Location

Temperate forests in southern South America.

Methods

New and existing tree‐ring radial growth chronologies representing the entire latitudinal range of Pilgerodendron uviferum were grouped according to latitude and then examined for differences in growth trends and non‐stationarity in growth responses to a drought severity index (scPDSI) over the 1900–1993 AD period and also before and after significant shifts in climate in the 1950s and 1970s.

Results

The radial‐growth response of P. uviferum climate was highly variable across its full latitudinal distribution. There was a long‐term and positive association between radial growth and higher moisture at the northern and southern edges of the distribution of this species and the opposite relationship for the core of its distribution, especially following the climatic shifts of the 1950s and 1970s. In addition, non‐stationarity in moisture‐radial growth relationships was observed in all three latitudinal groups (southern and northern edges and core) for all seasons during the 20th century.

Main conclusions

Climate shifts in southern South America in the 1950s and 1970s resulted in different responses in the mean radial growth of P. uviferum at the southern and northern edges and at the core of its range. Dendroclimatic analyses document that during the first half of the 20th century climate‐growth relationships were relatively similar between the southern and northern range edges but diverged after the 1950s. Our findings imply that simulated projections of climate impacts on tree growth, and by implication on forest ecosystem productivity, derived from models of past climate‐growth relationships need to carefully consider different and non‐stationarity responses along the wide latitudinal distribution of this species.     

An ecoregion assessment of projected tree species vulnerabilities in western North America through the 21st century

Forest ecosystems across western North America will likely see shifts in both tree species dominance and composition over the rest of this century in response to climate change. Our objective in this study was to identify which ecological regions might expect the greatest changes to occur. We used the process‐based growth model 3‐PG, to provide estimates of tree species responses to changes in environmental conditions and to evaluate the extent that species are resilient to shifts in climate over the rest of this century. We assessed the vulnerability of 20 tree species in western North America using the Canadian global circulation model under three different emission scenarios. We provided detailed projections of species shifts by including soil maps that account for the spatial variation in soil water availability and soil fertility as well as by utilizing annual climate projections of monthly changes in air temperature, precipitation, solar radiation, vapor pressure deficit and frost at a spatial resolution of one km. Projected suitable areas for tree species were compared to their current ranges based on observations at >40 000 field survey plots. Tree species were classified as vulnerable if environmental conditions projected in the future appear outside that of their current distribution ≥70% of the time. We added a migration constraint that limits species dispersal to <200 m yr^?1 to provide more realistic projections on species distributions. Based on these combinations of constraints, we predicted the greatest changes in the distribution of dominant tree species to occur within the Northwest Forested Mountains and the highest number of tree species stressed will likely be in the North American Deserts. Projected climatic changes appear especially unfavorable for species in the subalpine zone, where major shifts in composition may lead to the emergence of new forest types.     

Stationarity of climate-growth response is only marginally influenced by the soil moisture regime in Western Siberia

Stationary (time-stable) relationships between a tree-ring proxy and climatic drivers are a prerequisite for using tree rings as paleo-climatological archives, but non-stationarity has been detected worldwide. Here we use a classical, temperature-sensitive treeline site in Western Siberia to specifically test the influence of micro-site conditions (wet versus dry) on the stationarity of climate-growth relationships in three co-existing conifer species: Larix sibirica Ledeb., Picea obovata Ledeb., and Pinus sibirica DuTour. We test two commonly used tree-ring proxies, annual tree-ring width (TRW) and maximum latewood density (MXD), using moving windows and the bootstrapped transfer function stability test (BTFS). Summer temperature is the main driver of tree growth in all three species, but non-stationarities exist in all species and both tree-ring proxies. For TRW, we found stationarity only for larch from both micro-sites, while for MXD, we found stationarity for spruce from both micro-sites, and for pine from the wet micro-site. Micro-site variability did not seem to affect stationarity in any systematic way. We highlight the necessity to systematically test the influence of different methods of stationarity tests, since BTFS was more sensitive than moving-window analysis. Taken together, our results underscore the importance of testing the assumption of stationarity for diverse micro-sites, different species and proxies at all sites prior to any tree-ring based temperature reconstruction, since even within one site results can be drastically different.     

Marine extinction risk shaped by trait–environment interactions over 500 million years

Perhaps the most pressing issue in predicting biotic responses to present and future global change is understanding how environmental factors shape the relationship between ecological traits and extinction risk. The fossil record provides millions of years of insight into how extinction selectivity (i.e., differential extinction risk) is shaped by interactions between ecological traits and environmental conditions. Numerous paleontological studies have examined trait‐based extinction selectivity; however, the extent to which these patterns are shaped by environmental conditions is poorly understood due to a lack of quantitative synthesis across studies. We conducted a meta‐analysis of published studies on fossil marine bivalves and gastropods that span 458 million years to uncover how global environmental and geochemical changes covary with trait‐based extinction selectivity. We focused on geographic range size and life habit (i.e., infaunal vs. epifaunal), two of the most important and commonly examined predictors of extinction selectivity. We used geochemical proxies related to global climate, as well as indicators of ocean acidification, to infer average global environmental conditions. Life‐habit selectivity is weakly dependent on environmental conditions, with infaunal species relatively buffered from extinction during warmer climate states. In contrast, the odds of taxa with broad geographic ranges surviving an extinction (>2500 km for genera, >500 km for species) are on average three times greater than narrow‐ranging taxa (estimate of odds ratio: 2.8, 95% confidence interval = 2.3–3.5), regardless of the prevailing global environmental conditions. The environmental independence of geographic range size extinction selectivity emphasizes the critical role of geographic range size in setting conservation priorities.     

Quantifying climate–growth relationships at the stand level in a mature mixed‐species conifer forest

A range of environmental factors regulate tree growth; however, climate is generally thought to most strongly influence year‐to‐year variability in growth. Numerous dendrochronological (tree‐ring) studies have identified climate factors that influence year‐to‐year variability in growth for given tree species and location. However, traditional dendrochronology methods have limitations that prevent them from adequately assessing stand‐level (as opposed to species‐level) growth. We argue that stand‐level growth analyses provide a more meaningful assessment of forest response to climate fluctuations, as well as the management options that may be employed to sustain forest productivity. Working in a mature, mixed‐species stand at the Howland Research Forest of central Maine, USA, we used two alternatives to traditional dendrochronological analyses by (1) selecting trees for coring using a stratified (by size and species), random sampling method that ensures a representative sample of the stand, and (2) converting ring widths to biomass increments, which once summed, produced a representation of stand‐level growth, while maintaining species identities or canopy position if needed. We then tested the relative influence of seasonal climate variables on year‐to‐year variability in the biomass increment using generalized least squares regression, while accounting for temporal autocorrelation. Our results indicate that stand‐level growth responded most strongly to previous summer and current spring climate variables, resulting from a combination of individualistic climate responses occurring at the species‐ and canopy‐position level. Our climate models were better fit to stand‐level biomass increment than to species‐level or canopy‐position summaries. The relative growth responses (i.e., percent change) predicted from the most influential climate variables indicate stand‐level growth varies less from to year‐to‐year than species‐level or canopy‐position growth responses. By assessing stand‐level growth response to climate, we provide an alternative perspective on climate–growth relationships of forests, improving our understanding of forest growth dynamics under a fluctuating climate.     

Tree‐ring stable isotopes record the impact of a foliar fungal pathogen on CO2 assimilation and growth in Douglas‐fir

Swiss needle cast (SNC) is a fungal disease of Douglas‐fir (Pseudotsuga menziesii) that has recently become prevalent in coastal areas of the Pacific Northwest. We used growth measurements and stable isotopes of carbon and oxygen in tree‐rings of Douglas‐fir and a non‐susceptible reference species (western hemlock, Tsuga heterophylla) to evaluate their use as proxies for variation in past SNC infection, particularly in relation to potential explanatory climate factors. We sampled trees from an Oregon site where a fungicide trial took place from 1996 to 2000, which enabled the comparison of stable isotope values between trees with and without disease. Carbon stable isotope discrimination (Δ¹³C) of treated Douglas‐fir tree‐rings was greater than that of untreated Douglas‐fir tree‐rings during the fungicide treatment period. Both annual growth and tree‐ring Δ¹³C increased with treatment such that treated Douglas‐fir had values similar to co‐occurring western hemlock during the treatment period. There was no difference in the tree‐ring oxygen stable isotope ratio between treated and untreated Douglas‐fir. Tree‐ring Δ¹³C of diseased Douglas‐fir was negatively correlated with relative humidity during the two previous summers, consistent with increased leaf colonization by SNC under high humidity conditions that leads to greater disease severity in following years.     

Continental‐scale tree‐ring‐based projection of Douglas‐fir growth: Testing the limits of space‐for‐time substitution

A central challenge in global change research is the projection of the future behavior of a system based upon past observations. Tree‐ring data have been used increasingly over the last decade to project tree growth and forest ecosystem vulnerability under future climate conditions. But how can the response of tree growth to past climate variation predict the future, when the future does not look like the past? Space‐for‐time substitution (SFTS) is one way to overcome the problem of extrapolation: the response at a given location in a warmer future is assumed to follow the response at a warmer location today. Here we evaluated an SFTS approach to projecting future growth of Douglas‐fir (Pseudotsuga menziesii), a species that occupies an exceptionally large environmental space in North America. We fit a hierarchical mixed‐effects model to capture ring‐width variability in response to spatial and temporal variation in climate. We found opposing gradients for productivity and climate sensitivity with highest growth rates and weakest response to interannual climate variation in the mesic coastal part of Douglas‐fir's range; narrower rings and stronger climate sensitivity occurred across the semi‐arid interior. Ring‐width response to spatial versus temporal temperature variation was opposite in sign, suggesting that spatial variation in productivity, caused by local adaptation and other slow processes, cannot be used to anticipate changes in productivity caused by rapid climate change. We thus substituted only climate sensitivities when projecting future tree growth. Growth declines were projected across much of Douglas‐fir's distribution, with largest relative decreases in the semiarid U.S. Interior West and smallest in the mesic Pacific Northwest. We further highlight the strengths of mixed‐effects modeling for reviving a conceptual cornerstone of dendroecology, Cook's 1987 aggregate growth model, and the great potential to use tree‐ring networks and results as a calibration target for next‐generation vegetation models.     

Ecological effects of non‐native species in marine ecosystems relate to co‐occurring anthropogenic pressures

Predictors for the ecological effects of non‐native species are lacking, even though such knowledge is fundamental to manage non‐native species and mitigate their impacts. Current theories suggest that the ecological effects of non‐native species may be related to other concomitant anthropogenic stressors, but this has not been tested at a global scale. We combine an exhaustive meta‐analysis of the ecological effects of marine non‐native species with human footprint proxies to determine whether the ecological changes due to non‐native species are modulated by co‐occurring anthropogenic impacts. We found that non‐native species had greater negative effects on native biodiversity where human population was high and caused reductions in individual performance where cumulative human impacts were large. On this basis we identified several marine ecoregions where non‐native species may have the greatest ecological effects, including areas in the Mediterranean Sea and along the northwest coast of the United States. In conclusion, our global assessment suggests coexisting anthropogenic impacts can intensify the ecological effects of non‐native species.     

Climate–growth relationships of tropical tree species in West Africa and their potential for climate reconstruction

Most tropical regions are facing historical difficulties of generating biologically reconstructed long‐term climate records. Dendrochronology (tree‐ring studies) is a powerful tool to develop high‐resolution and exactly dated proxies for climate reconstruction. Owing to the seasonal variation in rainfall we expected the formation of annual tree rings in the wood of tropical West African tree species. In the central‐western part of Benin (upper Ouémé catchment, UOC) and in northeastern Ivory Coast (Comoé National Park, CNP) we investigated the relationship between climate (precipitation, sea surface temperature (SST)) and tree rings and show their potential for climate reconstruction. Wood samples of almost 200 trees belonging to six species in the UOC and CNP served to develop climate‐sensitive ring‐width chronologies using standard dendrochronological techniques. The relationship between local precipitation, monthly SST anomalies in the Gulf of Guinea, El Niño‐ Southern Oscillation (ENSO) and ring‐width indices was performed by simple regression analyses, two sample tests and cross‐spectral analysis. A low‐pass filter was used to highlight the decadal variability in rainfall of the UOC site. All tree species showed significant relationships with annual precipitation proving the existence of annual tree rings. ENSO signals could not be detected in the ring‐width patterns. For legume tree species at the UOC site significant relationships could be found between SST anomalies in the Gulf of Guinea indicating correlations at periods of 5.1–4.1 and 2.3 years. Our findings accurately show the relationship between tree growth, local precipitation and SST anomalies in the Gulf of Guinea possibly associated with worldwide SST patterns. A master chronology enabled the reconstruction of the annual precipitation in the UOC to the year 1840. Time series analysis suggest increasing arid conditions during the last 160 years which may have large impacts on the hydrological cycles and consequently on the ecosystem dynamics and the development of socio‐economic cultures and sectors in the Guinea‐Congolian/Sudanian region.     

Contrasting tree‐ring growth to climate responses of Abies alba toward the southern limit of its distribution area

Silver fir Abies alba is an indigenous tree species present in many southern European mountain forests. Its distribution area and its adaptive capacity to climate variability, expressed in tree‐ring growth series, make it a very suitable target species for studying responses to climate particularly in a complex area like the Mediterranean basin where significant changes are expected. We used a set of 52 site chronologies (784 trees) in the Italian Alps and Apennines (38.1°– 46.6°N and 6.7°– 16.3°E) and temperature and precipitation monthly data for the period 1900–1995. Principal component analyses of the tree‐ring site network was applied to extract common modes of variability in annual radial growth among the chronologies. Climate/growth relationships and their stationarity and consistency over time were computed by means of correlation and moving correlation functions. Tree‐ring chronologies show a clear distinction between the Alpine and the Mediterranean sites and a further separation of the Alpine region in western and eastern sectors. Accordingly, we found different transient and contrasting regional responses in time with the trends found in the Mediterranean sites marking a relaxation of some of the major climate limiting factors recorded prior to the last decades. Species’ sensitivity to global change may result in distinct spatial responses reflecting the complexity of the Mediterranean climate, with large differences between various areas of the basin. It is still unclear if these contrasting tree‐ring growth to climate responses of Abies alba are due to the corresponding separation between the Alpine and Mediterranean climate modes, the atmospheric CO₂ fertilization effect, the environmentally most fitted genetic pools of the southern fir ecotypes or a combination of all factors. Climate–growth analysis based on a wide site network and on long‐term weather records confirmed to be excellent tools to detect spatial and temporal variability of species’ responses to climate.     

Tree‐ring analysis and modeling approaches yield contrary response of circumboreal forest productivity to climate change

Circumboreal forest ecosystems are exposed to a larger magnitude of warming in comparison with the global average, as a result of warming‐induced environmental changes. However, it is not clear how tree growth in these ecosystems responds to these changes. In this study, we investigated the sensitivity of forest productivity to climate change using ring width indices (RWI) from a tree‐ring width dataset accessed from the International Tree‐Ring Data Bank and gridded climate datasets from the Climate Research Unit. A negative relationship of RWI with summer temperature and recent reductions in RWI were typically observed in continental dry regions, such as inner Alaska and Canada, southern Europe, and the southern part of eastern Siberia. We then developed a multiple regression model with regional meteorological parameters to predict RWI, and then applied to these models to predict how tree growth will respond to twenty‐first‐century climate change (RCP8.5 scenario). The projections showed a spatial variation and future continuous reduction in tree growth in those continental dry regions. The spatial variation, however, could not be reproduced by a dynamic global vegetation model (DGVM). The DGVM projected a generally positive trend in future tree growth all over the circumboreal region. These results indicate that DGVMs may overestimate future wood net primary productivity (NPP) in continental dry regions such as these; this seems to be common feature of current DGVMs. DGVMs should be able to express the negative effect of warming on tree growth, so that they simulate the observed recent reduction in tree growth in continental dry regions.     

树木年轮年内高分辨率稳定同位素记录:方法、进展和展望

树轮稳定同位素比率能有效地记录树木生长过程中气候环境变化信息及树木的生理响应机制。年内高分辨率树轮稳定同位素比率则能够提供更为详细的气候环境信息,揭示树木对季节尺度气候环境变化的生理生态响应机制,在古气候和全球变化生态学研究方面显示出巨大的潜力。本文收集了1990年以来发表的树轮年内高分辨率稳定同位素比率研究论文,从样品剥离方法、α-纤维素化学提取方法以及应用等方面综述了其研究进展,展望了年内高分辨率树轮稳定同位素记录研究的潜力和未来发展方向。     

Community functional trait composition at the continental scale: the effects of non‐ecological processes

Ecological communities and their response to environmental gradients are increasingly being described by measures of trait composition at the community level – the trait‐based approach. Whether ecological or non‐ecological processes influence trait composition between communities has been debated. Understanding the processes that influence trait composition is important for reconstructing paleoenvironmental conditions from fossil deposits and for understanding changes in community functionality through time. Here, we assess the influence of ecological and non‐ecological processes on the distribution of traits within North American mammals. We found that non‐ecological processes including historical contingency, spatial autocorrelation, and evolutionary history do not influence trait composition; however, the variance in trait composition is highly explained by climate gradients. Our results suggest that habitat breadth, terrestriality, diet breadth, and reproductive traits are strong candidates as proxies for measuring functional aspects of environments in the past and present.     

Non‐neutrality in forest communities: evolutionary and ecological determinants of tree species abundance distributions

The unified neutral theory of biodiversity and biogeography provides a promising framework that can be used to integrate stochastic and ecological processes operating in ecological communities. Based on a mechanistic non‐neutral model that incorporates density‐dependent mortality, we evaluated the deviation from a neutral pattern in tree species abundance distributions and explored the signatures of historical and ecological processes that have shaped forest biomes. We compiled a dataset documenting species abundance distributions in 1168 plots encompassing 16 973 tree species across tropical, temperate, and boreal forests. We tested whether deviations from neutrality of species abundance distributions vary with climatic and historical conditions, and whether these patterns differ among regions. Non‐neutrality in species abundance distributions was ubiquitous in tropical, temperate, and boreal forests, and regional differences in patterns of non‐neutrality were significant between biomes. Species abundance evenness/unevenness caused by negative density‐dependent or abiotic filtering effects had no clear macro‐scale climatic drivers, although temperature was non‐linearly correlated with species abundance unevenness on a global scale. These findings were not significantly biased by heterogeneity of plot data (the differences of plot area, measurement size, species richness, and the number of individuals sampled). Therefore, our results suggest that environmental filtering is not universally increasing from warm tropical to cold boreal forests, but might affect differently tree species assembly between and within biomes. Ecological processes generating particularly dominant species in local communities might be idiosyncratic or region‐specific and may be associated with geography and climate. Our study illustrates that stochastic dynamical models enable the analysis of the interplay of historical and ecological processes that influence community assemblies and the dynamics of biodiversity.     

Species‐rich boreal forests grew more and suffered less mortality than species‐poor forests under the environmental change of the past half‐century

Climate and other global environmental changes are major threats to ecosystem functioning and biodiversity. However, the importance of plant diversity in mitigating the responses of functioning of natural ecosystems to long‐term environmental change remains unclear. Using inventory data of boreal forests of western Canada from 1958 to 2011, we found that aboveground biomass growth increased over time in species‐rich forests but decreased in species‐poor forests, and importantly, aboveground biomass loss from tree mortality was smaller in species‐rich than species‐poor forests. A further analysis indicated that growth of species‐rich (but not species‐poor) forests was statistically positively associated with rising CO₂, and that mortality in species‐poor forests increased more as climate moisture availability decreased than it did in species‐rich forests. In contrast, growth decreased and mortality increased as the climate warmed regardless of species diversity. Our results suggest that promoting high tree diversity may help reduce the climate and environmental change vulnerability of boreal forests.     

Environmental drivers interactively affect individual tree growth across temperate European forests

Forecasting the growth of tree species to future environmental changes requires a better understanding of its determinants. Tree growth is known to respond to global‐change drivers such as climate change or atmospheric deposition, as well as to local land‐use drivers such as forest management. Yet, large geographical scale studies examining interactive growth responses to multiple global‐change drivers are relatively scarce and rarely consider management effects. Here, we assessed the interactive effects of three global‐change drivers (temperature, precipitation and nitrogen deposition) on individual tree growth of three study species (Quercus robur/petraea, Fagus sylvatica and Fraxinus excelsior). We sampled trees along spatial environmental gradients across Europe and accounted for the effects of management for Quercus. We collected increment cores from 267 trees distributed over 151 plots in 19 forest regions and characterized their neighbouring environment to take into account potentially confounding factors such as tree size, competition, soil conditions and elevation. We demonstrate that growth responds interactively to global‐change drivers, with species‐specific sensitivities to the combined factors. Simultaneously high levels of precipitation and deposition benefited Fraxinus, but negatively affected Quercus’ growth, highlighting species‐specific interactive tree growth responses to combined drivers. For Fagus, a stronger growth response to higher temperatures was found when precipitation was also higher, illustrating the potential negative effects of drought stress under warming for this species. Furthermore, we show that past forest management can modulate the effects of changing temperatures on Quercus’ growth; individuals in plots with a coppicing history showed stronger growth responses to higher temperatures. Overall, our findings highlight how tree growth can be interactively determined by global‐change drivers, and how these growth responses might be modulated by past forest management. By showing future growth changes for scenarios of environmental change, we stress the importance of considering multiple drivers, including past management and their interactions, when predicting tree growth.     

Predicting tree biomass growth in the temperate–boreal ecotone: Is tree size,age, competition,or climate response most important?

As global temperatures rise, variation in annual climate is also changing, with unknown consequences for forest biomes. Growing forests have the ability to capture atmospheric CO₂ and thereby slow rising CO₂ concentrations. Forests’ ongoing ability to sequester C depends on how tree communities respond to changes in climate variation. Much of what we know about tree and forest response to climate variation comes from tree‐ring records. Yet typical tree‐ring datasets and models do not capture the diversity of climate responses that exist within and among trees and species. We address this issue using a model that estimates individual tree response to climate variables while accounting for variation in individuals’ size, age, competitive status, and spatially structured latent covariates. Our model allows for inference about variance within and among species. We quantify how variables influence aboveground biomass growth of individual trees from a representative sample of 15 northern or southern tree species growing in a transition zone between boreal and temperate biomes. Individual trees varied in their growth response to fluctuating mean annual temperature and summer moisture stress. The variation among individuals within a species was wider than mean differences among species. The effects of mean temperature and summer moisture stress interacted, such that warm years produced positive responses to summer moisture availability and cool years produced negative responses. As climate models project significant increases in annual temperatures, growth of species like Acer saccharum, Quercus rubra, and Picea glauca will vary more in response to summer moisture stress than in the past. The magnitude of biomass growth variation in response to annual climate was 92–95% smaller than responses to tree size and age. This means that measuring or predicting the physical structure of current and future forests could tell us more about future C dynamics than growth responses related to climate change alone.     

Contrasting resistance and resilience to extreme drought and late spring frost in five major European tree species

Extreme climate events (ECEs) such as severe droughts, heat waves, and late spring frosts are rare but exert a paramount role in shaping tree species distributions. The frequency of such ECEs is expected to increase with climate warming, threatening the sustainability of temperate forests. Here, we analyzed 2,844 tree‐ring width series of five dominant European tree species from 104 Swiss sites ranging from 400 to 2,200 m a.s.l. for the period 1930–2016. We found that (a) the broadleaved oak and beech are sensitive to late frosts that strongly reduce current year growth; however, tree growth is highly resilient and fully recovers within 2 years; (b) radial growth of the conifers larch and spruce is strongly and enduringly reduced by spring droughts—these species are the least resistant and resilient to droughts; (c) oak, silver fir, and to a lower extent beech, show higher resistance and resilience to spring droughts and seem therefore better adapted to the future climate. Our results allow a robust comparison of the tree growth responses to drought and spring frost across large climatic gradients and provide striking evidence that the growth of some of the most abundant and economically important European tree species will be increasingly limited by climate warming. These results could serve for supporting species selection to maintain the sustainability of forest ecosystem services under the expected increase in ECEs.