Patch dynamics integrate mechanisms for savanna tree–grass coexistence

Many mechanisms have been suggested to explain the coexistence of woody species and grasses in savannas. However, evidence from field studies and simulation models has been mixed. Patch dynamics is a potentially unifying mechanism explaining tree–grass coexistence and the natural occurrence of shrub encroachment in arid and semi-arid savannas. A patch-dynamic savanna consists of a spatial mosaic of patches. Each patch maintains a cyclical succession between dominance of woody species and grasses, and the succession of neighbouring patches is temporally asynchronous. Evidence from empirical field studies supports the patch dynamics view of savannas. As a basis for future tests of patch dynamics in savannas, several hypotheses are presented and one is exemplarily examined: at the patch scale, realistically parameterized simulation models have generated cyclical succession between woody and grass dominance. In semi-arid savannas, cyclical successions are driven by precipitation conditions that lead to mass recruitment of shrubs in favourable years and to simultaneous collapse of shrub cohorts in drought years. The spatiotemporal pattern of precipitation events determines the scale of the savanna vegetation mosaic in space and time. In a patch-dynamic savanna, shrub encroachment is a natural, transient phase corresponding to the shrub-dominated phase during the successional cycle. Hence, the most promising management strategy for encroached areas is a large-scale rotation system of rangelands. In conclusion, patch dynamics is a possible scale-explicit mechanism for the explanation of tree–grass coexistence in savannas that integrates most of the coexistence mechanisms proposed thus far for savannas.     

Performance of West African dwarf goats fed Guinea grass–Verano stylo mixture,N-fertilized and unfertilized Guinea grass

The supplementary values of Verano stylo in a mixed Guinea grass (Panicum maximum cv. Ntchisi)–Verano stylo (Stylosanthes hamata cv. Verano) diet from a sown grass–legume mixture and N fertilized grass were compared in West African dwarf (WAD) goats. Liveweight (LW) gain, feed intake, digestibility and N utilization were determined using 15 goats in two trials lasting for 98 days. Goats were fed Guinea grass–Verano stylo mixture (GSM), N-fertilized (NFG) and unfertilized grass (UFG). The goats were divided into three groups of five animals each and randomly allocated to the dietary treatments in a randomized complete block design. Total DM and OM intakes of the goats did not vary significantly among the forage diets and averaged 55.1 and 50.4 g kg⁻¹ W^0.75 per day, respectively. CP intake (g kg⁻¹ W^0.75 per day) was highest with NFG (5.6) followed by GSM (4.8) and the UFG (3.5). Total N excreted followed the same trend as the CP intake. There was no significant difference between N-retention of GSM and NFG (28.5 and 26.7%), but goats on UFG had a negative N balance (−9.16%). Animals on GSM had significantly higher liveweight gain (31.9 g per day) than those of NFG (25.1 g per day) and UFG (21.9 g per day) which also differed significantly. The digestibilities of total DM, OM, CP, NDF were higher with GSM than NFG or UFG. It is concluded that growing Verano stylo in mixture with Guinea grass is a better option for improving the feed quality of forage diets for goats than direct application of inorganic fertilizer at 200 kg N ha⁻¹ to the pure grass.     

Ungulate saliva inhibits a grass–endophyte mutualism

Fungal endophytes modify plant–herbivore interactions by producing toxic alkaloids that deter herbivory. However, studies have neglected the direct effects herbivores may have on endophytes. Antifungal properties and signalling effectors in herbivore saliva suggest that evolutionary pressures may select for animals that mitigate the effects of endophyte-produced alkaloids. Here, we tested whether saliva of moose (Alces alces) and European reindeer (Rangifer tarandus) reduced hyphal elongation and production of ergot alkaloids by the foliar endophyte Epichloë festucae associated with the globally distributed red fescue Festuca rubra. Both moose and reindeer saliva reduced the growth of isolated endophyte hyphae when compared with a treatment of distilled water. Induction of the highly toxic alkaloid ergovaline was also inhibited in plants from the core of F. rubra''s distribution when treated with moose saliva following simulated grazing. In genotypes from the southern limit of the species'' distribution, ergovaline was constitutively expressed, as predicted where growth is environmentally limited. Our results now present the first evidence, to our knowledge, that ungulate saliva can combat plant defences produced by a grass–endophyte mutualism.     

Water availability drives gradients of tree diversity,structure and functional traits in the Atlantic–Cerrado–Caatinga transition,Brazil

Aims Climate and soil are among the most important factors determining variation in tree communities, but their effects have not been thoroughly elucidated to date for many vegetation features. In this study, we evaluate how climate and soil gradients affect gradients of vegetation composition, species diversity and dominance, structure and functional traits (seed mass and wood density) using over 327 000 trees in 158 sites distributed along environmental gradients in the transitions among the Atlantic forest, Cerrado and Caatinga in Minas Gerais State (MG), Brazil (nearly 600 000 km2).     

Ecological functioning in grass–shrub Mediterranean ecosystems measured by eddy covariance

Climate change may alter ecosystem functioning, as assessed via the net carbon (C) exchange (NEE) with the atmosphere, composed of the biological processes photosynthesis (GPP) and respiration (R _eco). In addition, in semi-arid Mediterranean ecosystems, a significant fraction of respired CO₂ is stored in the vadose zone and emitted afterwards by subsoil ventilation (VE), contributing also to NEE. Such conditions complicate the prediction of NEE for future change scenarios. To evaluate the possible effects of climate change on annual NEE and its underlying processes (GPP, R _eco and VE) we present, over a climate/altitude range, the annual and interannual variability of NEE, GPP, R _eco and VE in three Mediterranean sites. We found that annual NEE varied from a net source of around 130 gC m^?2 in hot and arid lowlands to a net sink of similar magnitude for alpine meadows (above 2,000 m a.s.l) that are less water stressed. Annual net C fixation increased because of increased GPP during intermittent and several growth periods occurring even during winter, as well as due to decreased VE. In terms of interannual variability, the studied subalpine site behaved as a neutral C sink (from emission of 49 to fixation of 30 gC m^?2 year^?1), with precipitation as the main factor controlling annual GPP and R _eco. Finally, the importance of VE as 0–23 % of annual NEE is highlighted, indicating that this process could shift some Mediterranean ecosystems from annual C sinks to sources.     

Structural analysis of N-glycans from allergenic grass, ragweed and tree pollens: Core α1,3-linked fucose and xylose present in all pollens examined

The N-glycans from soluble extracts of ten pollens were examined. The pyridylaminated oligosaccharides derived from these sources were subject to gel filtration and reverse-phase HPLC, in conjunction with exoglycosidase digests, and in some cases matrix-assisted laser desoprofessional-articletion-ionisation mass spectrometry. In comparison to known structures, it was possible to determine the major structures of the N-glycans derived from Kentucky blue grass (Poa pratensis), rye (Secale cerale), ryegrass (Lolium perenne), short ragweed (Ambrosia elatior), giant ragweed (Ambrosia trifida), birch (Betula alba), hornbeam (Caprofessional-articleinus betulus), horse chestnut (Aesculus hippocastanum), olive (Olea europaea) and snake-skin pine (Pinus leucodermis) pollen extracts. For grass pollens the major glycans detected were identical in properties to:$&1[-]Grass pollens also contained some minor structures with one or two non-reducing terminal N-acetylglucosamine residues. In the ragweed pollens, the major structures carried core 1,3-linked fucose with or without the presence of xylose. In tree pollen extracts, the major structures were either xylosylated, with or without fucose and terminal N-acetylglucosamine residues, with also significant amounts of oligomannose structures. These results are compatible with the hypothesis that the carbohydrate structures are another potential source of immunological cross-reaction between different plant allergens.     

What genes make a tree a tree?

Woody growth is evolutionarily ancient, yet has been gained and lost multiple times in plant evolution and is readily enhanced or minimized in eudicot speciation. New molecular genetic and genomic studies in Populus and Arabidopsis that are defining the genes responsible for cambium function and woody growth suggest that the genes regulating woody growth are not unique to woody plants. Surprisingly, key genetic mechanisms originally characterized as regulating the meristematic cells of the shoot apical meristem are also expressed in the vascular cambium during woody growth. This has important implications for the development of Populus as a model species and illustrates why forest trees constitute a contrived group of plants that have more in common with herbaceous relatives than we foresters like to admit.     

Weak phylogenetic signal for specialisation in antagonistic liana–tree networks

Background: Antagonistic interactions, such as parasitism and herbivory, are generally specialised and have a strong phylogenetic signal for specialisation. As lianas and trees interact antagonistically, we expect to find phylogenetic signal for specialisation.

Aims: We aimed to answer the following questions: (1) Is the liana–tree network specialised? (2) Is the specialisation of liana–tree network related to the abundance of both the life forms? (3) Is liana and tree specialisation related to species phylogeny? (4) Do phylogenetically related liana species occupy phylogenetically related tree species, and vice versa?

Methods: For three areas in southern Brazil, we calculated the specialisation value of each liana and tree species (d') and of the entire network (H₂′). Binomial regression and null models were used to test the role of abundance on d' and H₂′, respectively. We searched for the presence of phylogenetic signal with phylogenetic independent contrasts for d'. We also compared the similarity of species sets and their interaction with phylogenetic distance between them using Mantel test.

Results: All three networks had significant values of H₂′, but the values of d' did not have significant phylogenetic signals. Closely related lianas did not share similar host-tree assemblages and vice versa. Rare species were more specialised than abundant species, and abundance did not influence H₂′.

Conclusions: Our study indicates that the significant H₂′ may be due to co-evolution in some lineages of lianas and trees. Nevertheless, the abundance of species may also play an important role in species interaction, mainly rare species.     

Potential antagnosim of cultivated and wild grass–endophyte associations towards Meloidogyne incognita

Potential antagonism towards plant-parasitic nematode Meloidogyne incognita by grass–endophyte associations and their corresponding endophytes was evaluated. The following grass-endophyte associations were studied: (1) the cultivated grass Schedonorus arundinaceus-Neotyphodium coenophialum, (2) the wild grass Leymus chinensis-Neotyphodium sp. and (3) the wild grass Achnatherum sibiricum-Neotyphodium sibiricum. The cultivated grass Schedonorus arundinaceus was used to evaluate the effect of the endophyte fungus on chemotaxis of M. incognita. The results showed a significant negative effect on chemotaxis of the second-stage juveniles (J2) of M. incognita towards S. arundinaceus-N. coenophialum plants (P < 0.05). The associations of the grass S. arundinaceus-N. coenophialum and the wild grass A. sibiricum-N. sibiricum were used for testing their effect on penetration of grasses by J2. The wild grass A. sibiricum-N. sibiricum association appeared to suppress J2 penetration. N. coenophialum, Neotyphodium sp. and N. sibiricum isolated from cultivated and wild grasses respectively were used for measuring their effect on J2 mortality. Undiluted culture filtrates of all endophytic fungi caused significantly high J2 mortality 72.6% for N. coenphialum, 91.7% for Neotyphodium sp. and 66.8% for N. sibiricum. Finally, the penetration of cucumber roots by J2 pretreated by the filtrates of the three endophytic fungi was also tested. N. sibiricum filtrate significantly reduced the number of J2 penetrations by 44.5% and no significant effect in the treatment of the filtrates of Neotyphodium sp. from L. chinensis. Taken together, endophytic fungi of the genus Neotyphodium might be a potential source of biological control agents for plant parasitic nematodes.     

The energy–water limitation threshold explains divergent drought responses in tree growth,needle length,and stable isotope ratios

Predicted increases in extreme droughts will likely cause major shifts in carbon sequestration and forest composition. Although growth declines during drought are widely documented, an increasing number of studies have reported both positive and negative responses to the same drought. These divergent growth patterns may reflect thresholds (i.e., nonlinear responses) promoted by changes in the dominant climatic constraints on tree growth. Here we tested whether stemwood growth exhibited linear or nonlinear responses to temperature and precipitation and whether stemwood growth thresholds co-occurred with multiple thresholds in source and sink processes that limit tree growth. We extracted 772 tree cores, 1398 needle length records, and 1075 stable isotope samples from 27 sites across whitebark pine's (Pinus albicaulis Engelm.) climatic niche in the Sierra Nevada. Our results indicated that a temperature threshold in stemwood growth occurred at 8.4°C (7.12–9.51°C; estimated using fall-spring maximum temperature). This threshold was significantly correlated with thresholds in foliar growth, as well as carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope ratios, that emerged during drought. These co-occurring thresholds reflected the transition between energy- and water-limited tree growth (i.e., the E–W limitation threshold). This transition likely mediated carbon and nutrient cycling, as well as important differences in growth-defense trade-offs and drought adaptations. Furthermore, whitebark pine growing in energy-limited regions may continue to experience elevated growth in response to climate change. The positive effect of warming, however, may be offset by growth declines in water-limited regions, threatening the long-term sustainability of the recently listed whitebark pine species in the Sierra Nevada.     

Local effects drive heterozygosity–fitness correlations in an outcrossing long-lived tree

Heterozygosity–fitness correlations (HFCs) have been used to understand the complex interactions between inbreeding, genetic diversity and evolution. Although frequently reported for decades, evidence for HFCs was often based on underpowered studies or inappropriate methods, and hence their underlying mechanisms are still under debate. Here, we used 6100 genome-wide single nucleotide polymorphisms (SNPs) to test for general and local effect HFCs in maritime pine (Pinus pinaster Ait.), an iconic Mediterranean forest tree. Survival was used as a fitness proxy, and HFCs were assessed at a four-site common garden under contrasting environmental conditions (total of 16 288 trees). We found no significant correlations between genome-wide heterozygosity and fitness at any location, despite variation in inbreeding explaining a substantial proportion of the total variance for survival. However, four SNPs (including two non-synonymous mutations) were involved in significant associations with survival, in particular in the common gardens with higher environmental stress, as shown by a novel heterozygosity–fitness association test at the species-wide level. Fitness effects of SNPs involved in significant HFCs were stable across maritime pine gene pools naturally growing in distinct environments. These results led us to dismiss the general effect hypothesis and suggested a significant role of heterozygosity in specific candidate genes for increasing fitness in maritime pine. Our study highlights the importance of considering the species evolutionary and demographic history and different spatial scales and testing environments when assessing and interpreting HFCs.     

Prey–predator mutualism in a tritrophic system on a camphor tree

We report the discovery of a mutualistic system encompassing prey–predator interactions. A domatium is a small space in a vein axil on the underside of leaves of woody angiosperms. Cinnamomum camphora Linn. has domatia that harbor a microphytophagous eriophyid mite (sp. 1). We previously reported that a predatory mite, Euseius sojaensis (Ehara), depends on this eriophyid mite as food. We revealed that E. sojaensis also preyed upon another eriophyid mite (sp. 2) that induces galls on leaves, and that the mean area of C. camphora leaves with galls was usually less than half that of leaves without galls. We experimentally tested the effect of E. sojaensis on galls, and confirmed that the presence of E. sojaensis reduced gall induction. Therefore, C. camphora, eriophyid mite sp. 1, and E. sojaensis comprise a mutualistic system, in spite of the prey–predator interactions among them. The conventional concept of mutualism does not apply to such prey–predator interactions, so we defined them as systematic mutualism. Here, the system consists of three trophic levels, and individuals that constitute this system benefit from the other species that constitute this system.     

Aboveground productivity and root–shoot allocation differ between native and introduced grass species

Plant species in grasslands are often separated into groups (C₄ and C₃ grasses, and forbs) with presumed links to ecosystem functioning. Each of these in turn can be separated into native and introduced (i.e., exotic) species. Although numerous studies have compared plant traits between the traditional groups of grasses and forbs, fewer have compared native versus introduced species. Introduced grass species, which were often introduced to prevent erosion or to improve grazing opportunities, have become common or even dominant species in grasslands. By virtue of their abundances, introduced species may alter ecosystems if they differ from natives in growth and allocation patterns. Introduced grasses were probably selected nonrandomly from the source population for forage (aboveground) productivity. Based on this expectation, aboveground production is predicted to be greater and root mass fraction to be smaller in introduced than native species. We compared root and shoot distribution and tissue quality between introduced and native C₄ grass species in the Blackland Prairie region of Central Texas, USA, and then compared differences to the more well-studied divergence between C₄ grasses and forbs. Comparisons were made in experimental monocultures planted with equal-sized transplants on a common soil type and at the same density. Aboveground productivity and C:N ratios were higher, on average, in native grasses than in native forbs, as expected. Native and introduced grasses had comparable amounts of shallow root biomass and tissue C:N ratios. However, aboveground productivity and total N were lower and deep root biomass and root mass fraction were greater in native than introduced grasses. These differences in average biomass distribution and N could be important to ecosystems in cases where native and introduced grasses have been exchanged. Our results indicate that native–introduced status may be important when interpreting species effects on grassland processes like productivity and plant N accumulation.     

Extent and spatial patterns of grass bald land cover change (1948–2000), Oregon Coast Range,USA

Globally, temperate grasslands and meadows have sharply declined in spatial extent. Loss and fragmentation of grasslands and meadows may impact biodiversity, carbon storage, energy balance, and climate change. In the Pacific Northwest region of North America, grasslands and meadows have declined in extent over the past century. Largely undocumented in this regional decline are the grass balds of the Oregon Coast Range, isolated grasslands in a landscape dominated by coniferous forests. This study was conducted to quantify the spatial extent and patterns of grass bald change. Five balds in the Oregon Coast Range were evaluated using historical aerial photographs and recent digital orthophoto quadrangles (DOQ). Over the time period of study (1948/1953 to 1994/2000), bald area declined by 66%, primarily from forest encroachment. The number and average size of bald vegetation patches declined, while edge density increased. Tree encroachment into balds was inversely related to distance from nearest potential parent trees. Spatial patterns of bald loss may result from a forest to bald gradient of unfavorable environmental conditions for tree establishment and/or seed dispersal limitation. Species dependent on balds may be at risk from loss of bald area and increased fragmentation, although metrics of habitat fragmentation may not reflect species-specific habitat requirements. Tree encroachment patterns and increased bald edge densities suggest increasing rates of bald loss in the future. The remote sensing nature of this study cannot determine the fundamental causes of bald decline, although prior research suggests climate change, cessation of native burning, successional changes in response to prior wildfires, and cessation of livestock grazing all may have potential influence.     

Trade-off between seed number and weight: Influence of a grass–endophyte symbiosis

Plant fitness is enhanced by resource allocation to seed number (offspring number) or weight (offspring survival). Besides, there is a well known trade-off in resource allocation between both traits. Symbiotic interactions can influence plant resource allocation to reproduction, yet little research has been performed in this direction. We studied the consequences of a grass–fungus symbiosis on the trade-off between seed number and weight, using Lolium multiflorum and the endophyte Neotyphodium occultans as our study system. In ecological terms, we experimentally removed N. occultans from L. multiflorum plants, and compared reproductive allocation to seed number and weight in endophyte-symbiotic vs. non-symbiotic plants at different levels of nutrient availability (small pots vs. large pots). In evolutionary terms, we compared reproductive allocation between symbiotic vs. non-symbiotic plants for different host genotypes. All plants showed a negative association between seed number and weight, once standardized for total reproductive biomass. Under high nutrient availability, endophyte-symbiotic plants showed higher seed weight than non-symbiotic plants for any seed number. However, no differences were observed under low nutrient availability. Endophyte influence also varied according to L. multiflorum genotype; specifically, endophyte-symbiotic plants showed a lower slope in the relationship between seed number and weight than non-symbiotic plants for the ‘Marshall’ genotype but no endophyte influence was found for the “Pampean” genotype. The results implied a higher plasticity in seed weight and lower plasticity in seed number for symbiotic plants. Indeed, endophyte-symbiotic plants showed an overall lower slope in the association between seed number and total reproductive biomass than non-symbiotic plants. Our results suggest that N. occultans induces heavier seeds in L. multiflorum plants under environmental conditions favorable to plant growth or for certain plant genotypes. We propose that symbiotic interactions may influence the evolution of seed number and weight trade-off.     

Exploring patterns of fine root morphological,chemical, and anatomical traits of 12 tree species from visible–near-infrared spectral reflectance

Plant and Soil - Root morphological response to localised phosphorus (P) application plays a crucial role in P acquisition. However, detailed knowledge of when and where roots respond to P patch...     

Seasonal and synoptic climatic drivers of tree growth in the Bighorn Mountains,WY, USA (1654–1983 CE)

In the United States’ (US) Northern Rockies, synoptic pressure systems and atmospheric circulation drive interannual variation in seasonal temperature and precipitation. The radial growth of high-elevation trees in this semi-arid region captures this temperature and precipitation variability and provides long time series to contextualize instrumental-era variability in synoptic-scale climate patterns. Such variability in climate patterns can trigger extreme climate events, such as droughts, floods, and forest fires, which have a damaging impact on human and natural systems. We developed 11 tree-ring width (TRW) chronologies from multiple species and sites to investigate the seasonal climatic drivers of tree growth in the Bighorn Mountains, WY. A principal component analysis of the chronologies identified 54% of shared common variance (1894–2014). Tree growth (expressed by PC1) was driven by multiple seasonal climate variables: previous October and current July temperatures, as well as previous December and current April precipitation, had a positive influence on growth, whereas growth was limited by July precipitation. These seasonal growth-climate relationships corresponded to circulation patterns at higher atmospheric levels over the Bighorn Mountains. Tree growth was enhanced when the winter jet stream was in a northward position, which led to warmer winters, and when the spring jet stream was further south, which led to wetter springs. The second principal component, explaining 19% of the variance, clustered sites by elevation and was strongly related to summer temperature. We leverage this summer temperature signal in our TRW chronologies by combining it with an existing maximum latewood density (MXD) chronology in a nested approach. This allowed us to reconstruct Bighorn Mountains summer (June, July, and August) temperature (BMST) back to 1654, thus extending the instrumental temperature record by 250 years. Our BMST reconstruction explains 39–53% of the variance in regional summer temperature variability. The 1830s were the relatively coolest decade and the 1930s were the warmest decade over the reconstructed period (1654–1983 CE) – which excludes the most recent 3 decades. Our results contextualize recent drivers and trends of climate variability in the US Northern Rockies, which contributes to the information that managers of human and natural systems need in order to prepare for potential future variability.     

Impact of the spatial scale of grass–legume mixtures on sheep grazing behaviour,preference and intake,and subsequent effects on pasture

The benefits of using white clover in pastures have been widely recognised for many years. However, clover is perceived as being unreliable because of its typically low content, which is spatially and temporally variable, in mixed pastures. One proposed solution to increase the proportion of clover in the diet of grazing animals and composition in the pasture is to spatially separate clover from grass within the same field. In a field experiment ryegrass and white clover were grown in fine mixtures, and in pure alternating strips of ryegrass and clover of 1.5 m, 3 m or 18 m width within a field. Pastures were grazed for two grazing periods of 9 and 12 weeks, and measurements of sward surface height (SSH), herbage mass and composition and clover morphology were taken. Grazing behaviour was also observed. Results showed that spatial separation in the long term, when compared with a fine mixture, increased clover availability (18% to 30% v. 9%, based on standing dry matter) and was not grazed to extinction. Ewes maintained their preference for clover throughout the experiment (selection coefficient 2 to 5), which resulted in a reduction in the SSH of clover in monocultures to <3 cm and significant changes to the morphology of clover (smaller leaves, shorter petioles and thicker stolon), at the expense of maximising their intake. Spatial separation in the short term may therefore allow grazing animals to select their preferred diet; however, in the long term in continuously grazed pasture, their preference for clover depletes its availability.     

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.