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Analysis of functional traits is a cornerstone of ecology, yet individual traits seldom explain useful amounts of variation in species distribution or climatic tolerance, and their functional significance is rarely validated experimentally. Multivariate suites of interacting traits could build an understanding of ecological processes and improve our ability to make sound predictions of species success in our rapidly changing world. We use foliar water uptake capacity as a case study because it is increasingly considered to be a key functional trait in plant ecology due to its importance for stress-tolerance physiology. However, the traits behind the trait, that is, the features of leaves that determine variation in foliar water uptake rates, have not been assembled into a widely applicable framework for uptake prediction. Focusing on trees, we investigated relationships among 25 structural traits, leaf osmotic potential (a source of free energy to draw water into leaves), and foliar water uptake in 10 diverse angiosperm and conifer species. We identified consistent, multitrait "uptake syndromes" for both angiosperm and conifer trees, with differences in key traits revealing suspected differences in the water entry route between these two clades and an evolutionarily significant divergence in the function of homologous structures. A literature review of uptake-associated functional traits, which largely documents similar univariate relationships, provides additional support for our proposed "uptake syndrome." Importantly, more than half of shared traits had opposite-direction influences on the capacity of leaves to absorb water in angiosperms and conifers. Taxonomically targeted multivariate trait syndromes provide a useful tool for trait selection in ecological research, while highlighting the importance of micro-traits and the physiological verification of their function for advancing trait-based ecology., (© 2023 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Seed availability and suitable microsites for germination are likely to severely constrain the responses of plant species to climate change, especially at and beyond range edges. For example, range shifts may be slow if seed availability is low at range edges due to low parent-tree abundance or reduced fecundity. Even when seeds are available, climatic and biotic factors may further limit the availability of suitable microsites for recruitment. Unfortunately, the importance of seed and microsite limitation during range shifts remains unknown, since few studies have examined both factors simultaneously, particularly across species' ranges. To address this issue, we assessed seed availability and the factors influencing germination for six conifer species across a large environmental gradient encompassing their elevational ranges. Specifically, we assessed (1) how parent-tree abundance influences annual seed availability; (2) how seed limitation varies across species' ranges; (3) how climatic and biotic factors affect germination; and (4) how seed and suitable microsite availability covary annually within and among species. We found that seed availability declined toward species' upper range edges for most species, primarily due to low parent-tree abundance rather than declining fecundity. Range expansions are thus likely to be lagged with respect to climate change, as long generation times preclude rapid increases in tree density. Negative impacts of canopy cover on germination rates suggest range shifts will further be slowed by competition with existing vegetation. Moreover, years of high seed production were generally correlated among species, but not correlated with the availability of suitable microsites, implying that seedling competition and the interaction between seed and microsite limitation will further constrain recruitment. However, the nature of microsite limitation varied strongly between treeline and low-elevation species due to differing responses to snowpack duration and. competition, suggesting that treeline species may be quicker to shift their ranges in response to warming than low-elevation species. In all, our results demonstrate that seed and microsite limitation will likely result in lagged responses to climate change but with differences among species leading to complex range shift dynamics.

Understanding the interplay between environmental factors contributing to treeline formation and how these factors influence different life stages remains a major research challenge. We used an afforestation experiment including 92 000 trees to investigate the spatial and temporal dynamics of tree mortality and growth at treeline in the Swiss Alps. Seedlings of three high-elevation conifer species (Larix decidua, Pinus mugo ssp. uncinata, and Pinus cembra) were systematically planted along an altitudinal gradient at and above the current treeline (2075 to 2230 m above sea level [a.s.l.]) in 1975 and closely monitored during the following 30 years. We used decision-tree models and generalized additive models to identify patterns in mortality and growth along gradients in elevation, snow duration, wind speed, and solar radiation, and to quantify interactions between the different variables. For all three species, snowmelt date was always the most important environmental factor influencing mortality, and elevation was always the most important factor for growth over the entire period studied. Individuals of all species survived at the highest point of the afforestation for more than 30 years, although mortality was greater above 2160 m a.s.l., 50-100 m above the current treeline. Optimal conditions for height growth differed from those for survival in all three species: early snowmelt (ca. day of year 125-140 [where day 1 is 1 January]) yielded lowest mortality rates, but relatively later snowmelt (ca. day 145-150) yielded highest growth rates. Although snowmelt and elevation were important throughout all life stages of the trees, the importance of radiation decreased over time and that of wind speed increased. Our findings provide experimental evidence that tree survival and height growth require different environmental conditions and that even small changes in the duration of snow cover, in addition to changes in temperature, can strongly impact tree survival and growth patterns at treeline. Further, our results show that the relative importance of different environmental variables for tree seedlings changes during the juvenile phase as they grow taller.

Interannual variation of carbon fluxes can be attributed to a number of biotic and abiotic controls that operate at different spatial and temporal scales. Type and frequency of disturbance, forest dynamics, and climate regimes are important sources of variability. Assessing the variability of carbon fluxes from these specific sources can enhance the interpretation of past and current observations. Being able to separate the variability caused by forest dynamics from that induced by climate will also give us the ability to determine if the current observed carbon fluxes are within an expected range or whether the ecosystem is undergoing unexpected change. Sources of interannual variation in ecosystem carbon fluxes from three evergreen ecosystems, a tropical, a temperate coniferous, and a boreal forest, were explored using the simulation model STANDCARB. We identified key processes that introduced variation in annual fluxes, but their relative importance differed among the ecosystems studied. In the tropical site, intrinsic forest dynamics contributed approximately 30% of the total variation in annual carbon fluxes. In the temperate and boreal sites, where many forest processes occur over longer temporal scales than those at the tropical site, climate controlled more of the variation among annual fluxes. These results suggest that climate-related variability affects the rates of carbon exchange differently among sites. Simulations in which temperature, precipitation, and radiation varied from year to year (based on historical records of climate variation) had less net carbon stores than simulations in which these variables were held constant (based on historical records of monthly average climate), a result caused by the functional relationship between temperature and respiration. This suggests that, under a more variable temperature regime, large respiratory pulses may become more frequent and high enough to cause a reduction in ecosystem carbon stores. Our results also show that the variation of annual carbon fluxes poses an important challenge in our ability to determine whether an ecosystem is a source, a sink, or is neutral in regard to CO2 at longer timescales. In simulations where climate change negatively affected ecosystem carbon stores, there was a 20% chance of committing Type II error, even with 20 years of sequential data.

The occurrence and abundance of conifers along climate gradients in the Inland Northwest (USA) was assessed using data from 5082 field plots, 81% of which were forested. Analyses using the Random Forests classification tree revealed that the sequential distribution of species along an altitudinal gradient could be predicted with reasonable accuracy from a single climate variable, a growing-season dryness index, calculated from the ratio of degree-days >5 degrees C that accumulate in the frost-free season to the summer precipitation. While the appearance and departure of species in an ascending altitudinal sequence were closely related to the dryness index, the departure was most easily visualized in relation to negative degree-days (degree-days < 0 degrees C). The results were in close agreement with the works of descriptive ecologists. A Weibull response function was used to predict from climate variables the abundance and occurrence probabilities of each species, using binned data. The fit of the models was excellent, generally accounting for >90% of the variance among 100 classes.

There has been considerable recent interest in how human-induced species loss affects community and ecosystem properties. These effects are particularly apparent when a commercially valuable species is harvested from an ecosystem, such as occurs through single-tree harvesting or selective logging of desired timber species in natural forests. In New Zealand mixed-species rain forests, single-tree harvesting of the emergent gymnosperm Dacrydium cupressinum, or rimu, has been widespread. This harvesting has been contentious in part because of possible ecological impacts of Dacrydium removal on the remainder of the forest, but many of these effects remain unexplored. We identified an area where an unintended 40-year "removal experiment" had been set up that involved selective extraction of individual Dacrydium trees. We measured aboveground and belowground variables at set distances from both individual live trees and stumps of trees harvested 40 years ago. Live trees had effects both above and below ground by affecting diversity and cover of several components of the vegetation (usually negatively), promoting soil C sequestration, enhancing ratios of soil C:P and N:P, and affecting community structure of soil microflora. These effects extended to 8 m from the tree base and were likely caused by poor-quality litter and humus produced by the trees. Measurements for the stumps revealed strong legacy effects of prior presence of trees on some properties (e.g., cover by understory herbs and ferns, soil C sequestration, soil C:P and N:P ratios), but not others (e.g., soil fungal biomass, soil N concentration). These results suggest that the legacy of prior presence of Dacrydium may remain for several decades or centuries, and certainly well over 40 years. They also demonstrate that, while large Dacrydium individuals (and their removal) may have important effects in their immediate proximity, within a forest, these effects should only be important in localized patches containing high densities of large trees. Finally, this study emphasizes that deliberate extraction of a particular tree species from a forest can exert influences both above and below ground if the removed species has a different functional role than that of the other plant species present.

Theoretical and empirical studies have long suggested that stability and complexity are intimately related, but evidence from long-lived systems at large scales is lacking. Stability can either be driven by complex species interactions, or it can be driven by the presence/absence and abundance of a species best able to perform a specific ecosystem function. We use 64 years of stand productivity measures in forest systems composed of four dominant conifer tree species to contrast the effect of species richness and abundance on three stability measures. To perform this contrast, we measured the annual growth increments of > 900 trees in mixed and pure forest stands to test three hypotheses: increased species richness will (1) decrease stand variance, (2) increase stand resistance to drought events, and (3) increase stand resilience to drought events. In each case, the alternate hypothesis was that species richness had no effect, but that species composition and abundance within a stand drove variance, resistance, and resilience. In pure stands, the four species demonstrated significant differences in productivity, and in their resistance and resilience to drought events. The two pine species were the most drought resistant and resilient, whereas mountain hemlock was the least resistant and resilient, and red fir was intermediate. For community measures we found a moderately significant (P = 0.08) increase in the community coefficient of variation and a significant (P = 0.03) increase in resilience with increased species richness, but no significant relationship between species richness and community resistance, though the variance in community resistance to drought decreased with species richness. Community resistance to drought was significantly (P = 0.001) correlated to the relative abundance of lodgepole pine, the most resistant species. We propose that resistance is driven by competition for a single limiting resource, with negative diversity effects. In contrast resilience measures the capacity of communities to partition resources in the absence of a single limiting resource, demonstrating positive diversity effects.