Temperate and boreal forest tree phenology: from organ-scale processes to terrestrial ecosystem models

Key message

We demonstrate that, beyond leaf phenology, the phenological cycles of wood and fine roots present clear responses to environmental drivers in temperate and boreal trees. These drivers should be included in terrestrial ecosystem models.

Context

In temperate and boreal trees, a dormancy period prevents organ development during adverse climatic conditions. Whereas the phenology of leaves and flowers has received considerable attention, to date, little is known regarding the phenology of other tree organs such as wood, fine roots, fruits, and reserve compounds.

Aims

Here, we review both the role of environmental drivers in determining the phenology of tree organs and the models used to predict the phenology of tree organs in temperate and boreal forest trees.

Results

Temperature is a key driver of the resumption of tree activity in spring, although its specific effects vary among organs. There is no such clear dominant environmental cue involved in the cessation of tree activity in autumn and in the onset of dormancy, but temperature, photoperiod, and water stress appear as prominent factors. The phenology of a given organ is, to a certain extent, influenced by processes in distant organs.

Conclusion

Inferring past trends and predicting future trends of tree phenology in a changing climate requires specific phenological models developed for each organ to consider the phenological cycle as an ensemble in which the environmental cues that trigger each phase are also indirectly involved in the subsequent phases. Incorporating such models into terrestrial ecosystem models (TEMs) would likely improve the accuracy of their predictions. The extent to which the coordination of the phenologies of tree organs will be affected in a changing climate deserves further research.

     

An ecosystem approach to biodiversity effects: Carbon pools in a tropical tree plantation

This paper presents a synthesis of experiments conducted in a tropical tree plantation established in 2001 and consisting of 22 plots of 45 m × 45 m with either one, three or six native tree species. We examined the changes in carbon (C) pools (trees, herbaceous vegetation, litter, coarse woody debris (CWD), and mineral topsoil at 0-10 cm depth) and fluxes (decomposition of CWD and litter, as well as soil respiration) both through time and among diversity levels. Between 2001 and 2009 the aboveground C pools increased, driven by trees. Across diversity levels, the mean observed aboveground C pool was 7.9 ± 2.5 Mg ha⁻¹ in 2006 and 20.4 ± 7.4 Mg ha⁻¹ in 2009, a 158% increase. There was no significant diversity effect on the observed aboveground C pool, but we found a significant decrease in the topsoil C pool, with a mean value of 34.5 ± 2.4 Mg ha⁻¹ in 2001 and of 25.7 ± 5.7 Mg ha⁻¹ in 2009 (F_1,36 = 52.12, p < 0.001). Assuming that the biomass C pool in 2001 was negligible (<1 Mg ha⁻¹), then the plantation gained in C, on average, ∼20 and lost ∼9 Mg ha⁻¹ in biomass and soil respectively, for an overall gain of ∼11 Mg ha⁻¹ over 8 years. Across the entire data set, we uncovered significant effects of diversity on CWD decomposition (diversity: F_2,393 = 15.93, p < 0.001) and soil respiration (monocultures vs mixtures: t = 15.35, df = 11, p < 0.05) and a marginally significant time × diversity interaction on the loss of total C from the mineral topsoil pool (see above). Monthly CWD decomposition was significantly faster in monocultures (35.0 ± 24.1%) compared with triplets (31.3 ± 21.0%) and six-species mixtures (31.9 ± 26.8%), while soil respiration was higher in monocultures than in mixtures (t = 15.35, df = 11, p < 0.001). Path analyses showed that, as diversity increases, the links among the C pools and fluxes strengthen significantly. Our results demonstrate that tree diversity influences the processes governing the changes in C pools and fluxes following establishment of a tree plantation on a former pasture. We conclude that the choice of tree mixtures for afforestation in the tropics can have a marked influence on C pools and dynamics.     

Correction to: Climatic factors controlling stem growth of alien tree species at a mesic forest site: a multispecies approach

European Journal of Forest Research -     

Modeling tree water deficit from microclimate: an approach to quantifying drought stress

Tree water deficit estimated by measuring water-related changes in stem radius (DeltaW) was compared with tree water deficit estimated from the output of a simple, physiologically reasonable model (DeltaWE), with soil water potential (Psisoil) and atmospheric vapor pressure deficit (VPD) as inputs. Values of DeltaW were determined by monitoring stem radius changes with dendrometers and detrending the results for growth. We followed changes in DeltaW and DeltaWE in Pinus sylvestris L. and Quercus pubescens Willd. over 2 years at a dry site (2001-2002; Salgesch, Wallis) and in Picea abies (L.) Karst. for 1 year at a wet site (1998; Davos, Graubuenden) in the Swiss Alps. The seasonal courses of DeltaW in deciduous species and in conifers at the same site were similar and could be largely explained by variation in DeltaWE. This finding strongly suggests that DeltaW, despite the known species-specific differences in stomatal response to microclimate, is mainly explained by a combination of atmospheric and soil conditions. Consequently, we concluded that trees are unable to maintain any particular DeltaW. Either Psisoil or VPD alone provided poorer estimates of DeltaW than a model incorporating both factors. As a first approximation of DeltaWE, Psisoil can be weighted so that the negative mean Psisoil reaches 65 to 75% of the positive mean daytime VPD over a season (Q. pubescens: approximately 65%, P. abies: approximately 70%, P. sylvestris: approximately 75%). The differences in DeltaW among species can be partially explained by a different weighting of Psisoil against VPD. The DeltaW of P. sylvestris was more dependent on Psisoil than that of Q. pubescens, but less than that of P. abies, and was less dependent on VPD than that of P. abies and Q. pubescens. The model worked well for P. abies at the wet site and for Q. pubescens and P. sylvestris at the dry site, and may be useful for estimating water deficit in other tree species.     

From a strategic to a tactical forest management plan using a hierarchic optimization approach

The Finnish state forest enterprise, Metsähallitus, defines the regional harvest levels for a 10-year period in a strategic-level natural resources plan. Although this plan defines stand-level harvest schedules for all stands, in practice, it cannot be used, as the harvests need to be clustered in time and in space. It is applied by giving each subregion goals they need to fulfill in a tactical level planning process, and the harvests are manually clustered into predefined groups of adjacent stands (departments). In this study, we developed a hierarchical optimization process making use of departments for clustering the harvests. For each of the departments, 91 different stand-level harvest schedules (plans) were determined using incomes from one period and the forest value at the end as objectives. The department-level plans were then used as alternatives in a region-level goal optimization problem. The resulting hierarchic plan was compared to the stand-level solution of the strategic-level plan which served as a benchmark plan. The hierarchical plan clustered the harvests and achieved the goals set better than the benchmark plan, but the net present income was 3.3% lower. The approach turned out usable, but further developing of the approach is needed to reduce the costs of clustering.     

Development and validation of a simple approach to modelling tree shading in agroforestry systems

There have been a number of models developed which attempt to predict the shading patterns beneath individual tree or forest canopies. We describe a computer-based model which is able to estimate shading patterns through a discontinuous canopy of pruned trees. The model is designed to assist in the layout and management of agroforestry systems with widely spaced trees. The model was tested against data collected from a seven-year-old agroforestry system involving radiata pine, located near Canberra, Australia. The model was shown to slightly but consistently underestimate light penetration at ground level beneath the trees. However, the extent of bias (0.7% to 5%) was so small as to be of little significance in practice.     

Estimating tree stem density and diameter distribution in single-scan terrestrial laser measurements of field plots: a simulation study

The single-scan setup of terrestrial laser scanning of a forest field plot has advantages compared to the multi-scan setup: the speed of operation and that there is no need of a co-registration of the different scans. However in a single-scan setup some of the trees are shaded by others and therefore not detected in the scan. A field inventory solution must take this fact into account. This simulation study shows how different plot sizes and tree stand densities influence the stem visibility giving nonlinear effects especially for large trees and high stem numbers. These effects can be counteracted by using an edge or center stem point detection criteria when analyzing the results or by weighting the detected trees by their visibility. It is shown that the stem density and diameter distribution can be estimated from the visible areas of the plot in case the stem positions are Poisson distributed.     

Unravelling and managing fusiform rust disease: a model approach for coevolved forest tree pathosystems

Fusiform rust disease remains the most destructive disease in pine plantations in the southern United States. Our ongoing research is designed to identify, map, and clone the interacting genes of the host and pathogen. Several resistance (R) genes have been identified and genetically mapped using informative pine families and single‐spore isolate inoculations. In addition, we are mapping the first of many expected corresponding avirulence (Avr) genes in the fungal pathogen. The Avr genes condition avirulence/virulence and avirulence is required for an incompatible reaction (i.e., no‐gall development) to take place within an inoculated tree that carries resistance at the corresponding R gene. We provide an overview of our methodology for identifying and mapping R and Avr genes, an update of our current progress, and a brief discussion of two approaches for predicting R gene genotypes of uncharacterized parental trees and for estimating the efficacy of specific pine genotypes at various planting locations. This paper emphasizes the critical importance of controlled genetic materials of both the host and pathogen for elucidating the genetic nature of resistance and virulence in coevolved forest pathosystems.     

Tracing carbon uptake from a natural CO2 spring into tree rings: an isotope approach

We analyzed 14C, 13C and 18O isotope variations over a 50-year period in tree rings of Quercus ilex L. trees growing at a natural CO2 spring in a Mediterranean ecosystem. We compared trees from two sites, one with high and one with low exposure to CO2 from the spring. The spring CO2 is free of 14C. Thus, this carbon can be traced in the wood, and the amount originating from the spring calculated. The amount decreased over time, from about 40% in 1950 to 15% at present for the site near the spring, indicating a potential difficulty in the use of natural CO2 springs for elevated CO2 research. The reason for the decrease may be decreasing emission from the spring or changes in stand structure, e.g., growth of the canopy into regions with lower concentrations. We used the 14C-calculated CO2 concentration in the canopy to determine the 13C discrimination of the plants growing under elevated CO2 by calculating the effective canopy air 13C/12C isotopic composition. The trees near the spring showed a 2.5 per thousand larger 13C discrimination than the more distant trees at the beginning of the investigated period, i.e., for the young trees, but this difference gradually disappeared. Higher discrimination under elevated CO2 indicated reduced photosynthetic capacity or increased stomatal conductance. The latter assumption is unlikely as inferred from the 18O data, which were insensitive to CO2 concentration. In conclusion, we found evidence for a downward adjustment of photosynthesis under elevated CO2 in Q. ilex in this dry, nutrient-poor environment.     

Tree growth as indicator of tree vitality and of tree reaction to environmental stress: a review

The intensive monitoring plots (Level II) of ICP Forests serve to examine the effects of air pollution and other stress factors on forest condition, including tree vitality. However, tree vitality cannot be measured directly. Indicators, such as tree growth or crown transparency, may instead be used. Tree growth processes can be ranked by order of importance in foliage growth, root growth, bud growth, storage tissue growth, stem growth, growth of defence compounds and reproductive growth. Under stress photosynthesis is reduced and carbon allocation is altered. Stem growth may be reduced early on as it is not directly vital to the tree. Actual growth must be compared against a reference growth, such as the growth of trees without the presumed stress, the growth of presumed healthy trees, the growth in a presumed stress-free period or the expected growth derived from models. Several examples from intensive monitoring plots in Switzerland illustrate how tree-growth reactions to environmental stresses may serve as vitality indicator. Crown transparency and growth can complement each other. For example, defoliation by insects becomes first visible in crown transparency while stem growth reaction occurs with delay. On the other hand, extreme summer drought as observed in large parts of Europe in 2003 affects stem growth almost immediately, while foliage reduction becomes only visible months later. Residuals of tree growth models may also serve as indicators of changed environmental conditions. Certain stresses, such as drought or insect defoliation cause immediate reactions and are not detectable in five-year growth intervals. Therefore, annual or inter-annual stem growth should be assessed in long-term monitoring plots. An erratum to this article can be found at     

Modelling irregular and multimodal tree diameter distributions by finite mixture models: an approach to stand structure characterisation

Tree structural diversity is assessed by modelling stand diameter at breast height (DBH) distribution. The aim of this study was to verify the suitability of a mixture of two- and three-component Weibull and gamma models for describing irregular and multimodal DBH distributions. Investigations were carried out in natural Abies alba Mill. and Fagus sylvatica L. stands, representing the growing-up stage, in the Świętokrzyski National Park (Central Poland) and in the Pieniny National Park (Southern Poland). Sample plots (0.25–0.5 ha in area) were selected from a database used for forest dynamics analysis. The parameters of the mixture model were estimated using maximum likelihood estimation (MLE) and a combination of an expectation–maximisation (EM) algorithm with a Newton-type (NT) method. The multistart method (MM) of choosing initial values was used. The DBH distributions in mixed Abies–Fagus forests of complex structure were accurately approximated by two- and three-component mixture models. The root mean square error (RMSE) values varied from 3.4515 to 7.6682 for the Weibull mixture and from 3.6457 to 7.1419 for the gamma mixture. The results indicated that the mixture Weibull and gamma models were, in general, equally suitable for modelling irregular and multimodal DBH distributions. Finite mixture models may be used to characterise tree structural diversity, especially in uneven-aged stands of complex structure.     

From guiding principles for the conservation of forest biodiversity to on-ground practice: Lessons from tree hollow management in Tasmania

A useful theoretical approach in the literature for those trying to conserve forest biodiversity involves a number of strategies to achieve maintenance of habitat for different species across multiple spatial and temporal scales. This approach emphasises the importance of implementing measures in off-reserve areas to complement reserve systems. This contrasts with both the traditional ‘set-aside’ approach to conservation and the primary aim of production forestry. Translating this risk-spreading approach into on-ground practice is often a difficult task.     

Screening agroforestry tree mulches for corn growth: a combined soil test,pot trial and plant analysis approach

The response of corn (Zea mays) to incorporated leaf and twig mulches ofLeucaena leucophala, Gliricidia sepium andCassia siamea, andGrevillea robusta as a non-legume comparison was investigated in a 10-week pot trial and a concurrent soil incubation study to evaluate the suitability of various agroforestry trees as mulch sources.Leucaena contributed to the highest N uptake and biomass production of these corn plants, reflecting the benefits of organic mulching.Cassia-treated plants also performed better than the unmulched controls, butGrevillea incorporation suppressed corn growth probably because of the relatively high Mn status of this mulch. Manganese toxicity was confirmed by comparative vector diagnosis of plant nutrient status.Cliricidia mulching resulted in seedling mortality after germination possibly from high soil pH and ammonium toxicity. Mineral N production in mulched soils during the laboratory incubation was well correlated with N uptake in corn shoots. The results suggest that the nutritional effects of agroforestry tree mulches on growth of companion crops may be effectively screened by a combined soil test, pot trial, and plant analysis approach.     

From forest to agroforest and logger to agroforester: a case study

This paper examines interactive change and adaptation of human and natural systems in two pioneer forest settlements in the Philippines. The forest ecosystem was converted by logging, further resource extraction by settlers, and cultivation — factors usually associated with systems degradation. Natural succession, however, was rapid because of high rainfall and abundant forest seed stocks; and because of high rainfall, weeds, insect pests, and poor soil — annual cereal and cash cropping was not profitable or sustainable and farmers turned to root and mixed perennial cropping. This naturally developing, more sustainable agroforestry was initially financed by boom-and-bust incomes from small scale logging and charcoal making, and took place in spite of the settlers' formation of factions and an us before them attitude towards resource use.     

The importance of an indigenous tree to southern African communities with specific relevance to its domestication and commercialization:a case of the marula tree

Local communities in southern Africa derive many benefits from marula [Sclerocarya birrea(A.Rich.) Hochst.subsp.caffra(Sond.) Kokwaro].These include the contribution of this species towards health,nutrition,food security and conservation by sharing local skills and knowledge related to it.Marula fruits can be eaten fresh,squeezed to make juice,brewed in traditional beer or used to make jam and jelly.The kernels are also edible and can be pressed to extract oil for cooking and cosmetics,i.e.,for skin and hai...     

The importance of an indigenous tree to southern African communities with specific relevance to its domestication and commercialization: a case of the marula tree

Sclerocarya birrea (A. Rich.) Hochst. subsp.