Local adaptation to aridity in a widely distributed angiosperm tree species is mediated by seasonal increase of sugars and reduced growth.

Trees in dry climates often have higher concentrations of total non-structural carbohydrates (NSC = starch + soluble sugars) and grow less than conspecifics in more humid climates. This pattern might result from growth being more constrained by aridity than the carbon (C) gain, or reflect local adaptation to aridity, since NSC fuel metabolism and ensure adequate osmoregulation through the supply of soluble sugars (SS), while low growth reduces water and C demands. It has been further proposed that C allocation to storage could come at the expense of growth (i.e., a growth-storage trade-off). We examined whether NSC and growth reflect local adaptation to aridity in Embothrium coccineum (Proteaceae), a species with an exceptionally wide niche. To control for any influence of the phenotypic plasticity on NSC and growth, we collected seeds from dry (500 mm year-1) and moist (> 2500 mm year-1) climates and grew seedlings in a common garden experiment for 3 years. We then compared NSC and SS concentrations and pools (i.e., total contents), and the biomass of seedlings at spring, summer, and fall. Seedlings from the dry climate had significantly lower biomass and similar NSC concentrations and pools than seedlings from moist climate, suggesting that reduced growth in arid environments does not result from a prioritization of C allocation to storage but it confers advantages under aridity (e.g., lower transpiration area). Across organs, starch and NSC decreased similarly in seedlings from both climates from spring onward. However, root and stem SS concentrations increased during the growing season, and these increases were significantly higher in seedlings from the dry climate. The greater SS accumulation in seedlings from the dry climate compared to those from the moist climate demonstrates ecotypic differentiation in the seasonal dynamics of SS, suggesting that SS underlie local adaptation to aridity. (298 words).

Carbon stress causes earlier budbreak in shade-tolerant species and delays it in shade-intolerant species.

PREMISE: Climate change may lead to C stress (negative C balance) in trees. Because nonstructural carbohydrates (NSC) are required during metabolic reactivation in the spring, C stress might delay budbreak timing. This effect is expected to be greater in shade-intolerant than in shade-tolerant species, owing to the faster C economy in the shade-intolerant. METHODS: We experimentally induced C stress in saplings of six temperate tree species that differed in their light requirements by exposing them to either full light or shade from summer to spring, then recorded the date of first budbreak for the individuals. Because the levels of C reserves that represent effective C stress may differ among species, we estimated the degree of C stress by recording survival during the experiment and measuring whole-sapling NSC concentrations after budbreak. RESULTS: Shade reduced NSC concentrations and increased the sugar fraction in the NSC in all species. In the shade, shade-intolerant species had higher mortality and generally lower NSC concentrations than the shade-tolerant species, indicating a trend for more severe C stress in species with faster C economy. In shade-intolerant species, budbreak started earlier and proceeded faster in full light than in shade, but in shade-tolerant species budbreak was delayed in full light. The effects of the light environments on budbreak were not greater in shade-intolerant than in shade-tolerant species. CONCLUSIONS: Our study reveals a correspondence between budbreak responses to light and the light requirements of the species. This finding confirms that C metabolism has a significant role in triggering budbreak and demonstrates that whether C stress accelerates or delays budbreak depends on the species' light requirements.

Phenology explains different storage remobilization in two congeneric temperate tree species with contrasting leaf habit.

The dependence of trees on carbon and nutrient storage is critical to predicting the forest vulnerability under climate change, but whether evergreen and deciduous species differ in their use and allocation of stored resources during spring phenology is unclear. Using a high temporal resolution, we evaluated the role of spring phenology and shoot growth as determinants of the carbon and nutrient storage dynamics in contrasting leaf habits. We recorded the phenology and shoot elongation and determined the concentrations of total non-structural carbohydrates (NSCs), starch, soluble carbohydrates, nitrogen (N) and phosphorus (P) in buds, expanding shoots and previously formed shoots of two sympatric Nothofagus species with contrasting leaf habit. Species reached similar shoot lengths, though shoot expansion started 35 days earlier and lasted c. 40 days more in the deciduous species. Thus, although the deciduous species had a relatively constant shoot growth rate, the evergreen species experienced a conspicuous growth peak for c. 20 days. In the evergreen species, the greatest decreases in NSC concentrations of previously formed shoots and leaves coincided with the maximum shoot expansion rate and fruit filling, with minimums of 63 and 65% relative to values at bud dormancy, respectively. In contrast, minimum NSC concentrations of the previously formed shoots of the deciduous species were only 73% and occurred prior to the initiation of shoot expansion. Bud N and P concentrations increased during budbreak, whereas previously formed shoots generally did not decrease their nutrient concentrations. Late spring phenology and overlapping of phenophases contributed to the greater dependence on storage of proximal tissues in the studied evergreen compared with deciduous species, suggesting that phenology is a key determinant of the contrasting patterns of storage use in evergreen and deciduous species.

Carbon allocation to growth and storage depends on elevation provenance in an herbaceous alpine plant of Mediterranean climate.

It is unclear whether the frequently observed increase in non-structural carbohydrates (NSC) in plants exposed to low temperatures or drought reflects a higher sensitivity of growth than photosynthesis in such conditions (i.e. sink limitation), or a prioritization of carbon (C) allocation to storage. Alpine areas in Mediterranean-type climate regions are characterized by precipitation increases and temperature decreases with elevation. Thus, alpine plants with wide elevational ranges in Mediterranean regions may be good models to examine these alternative hypotheses. We evaluated storage and growth during experimental darkness and re-illumination in individuals of the alpine plant Phacelia secunda from three elevations in the Andes of central Chile. We hypothesized that storage is prioritized regarding growth in plants of both low- and high elevations where drought and cold stress are greatest, respectively. We expected that decreases in NSC concentrations during darkness should be minimal and, more importantly, increases in NSC after re-illumination should be higher than increases in biomass. We found that darkness caused a significant decrease in NSC concentrations of both low- and high-elevation plants, but the magnitude of the decrease was lower in the latter. Re-illumination caused higher increase in NSC concentration than in biomass in both low- and high-elevation plants (1.5- and 1.9-fold, respectively). Our study shows that C allocation in Phacelia secunda reflects ecotypic differences among elevation provenances and suggests that low temperature, but not drought, favours C allocation to storage over growth after severe C limitation.

Inner bark as a crucial tissue for non-structural carbohydrate storage across three tropical woody plant communities.

Non-structural carbohydrates (NSC) are crucial for forest resilience, but little is known regarding the role of bark in NSC storage. However, bark's abundance in woody stems and its large living fraction make it potentially key for NSC storage. We quantified total NSC, soluble sugar (SS) and starch concentrations in the most living region of bark (inner bark, IB), and sapwood of twigs, trunks and roots of 45 woody species from three contrasting tropical climates spanning global extremes of bark diversity and wide phylogenetic diversity. NSC concentrations were similar (total NSC, starch) or higher (SS) in IB than wood, with concentrations co-varying strongly. NSC concentrations varied widely across organs and species within communities and were not significantly affected by climate, leaf habit or the presence of photosynthetic bark. Starch concentration tended to increase with density, but only in wood. IB contributed substantially to NSC storage, accounting for 17-36% of total NSC, 23-47% of SS and 15-33% of starch pools. Further examination of the drivers of variation in IB NSC concentration, and taking into account the substantial contribution of IB to NSC pools, will be crucial to understand the role of storage in plant environmental adaptation.

Low Growth Sensitivity and Fast Replenishment of Non-structural Carbohydrates in a Long-Lived Endangered Conifer After Drought.

There is an ongoing debate on whether a drought induced carbohydrate limitation (source limitation) or a direct effect of water shortage (sink limitation) limit growth under drought. In this study, we investigated the effects of the two driest summers recorded in southern Chile in the last seven decades, on the growth and non-structural carbohydrates (NSC) concentrations of the slow-growing conifer Fitzroya cupressoides. Specifically, we studied the seasonal variation of NSC in saplings and adults one and two years after the occurrence of a 2 year-summer drought at two sites of contrasting precipitation and productivity (mesic-productive vs. rainy-less productive). We also evaluated radial growth before, during and after the drought, and predicted that drought could have reduced growth. If drought caused C source limitation, we expected that NSCs will be lower during the first than the second year after drought. Conversely, similar NSC concentrations between years or higher NSC concentrations in the first year would be supportive of sink limitation. Also, due to the lower biomass of saplings compared with adults, we expected that saplings should experience stronger seasonal NSC remobilization than adults. We confirmed this last expectation. Moreover, we found no significant growth reduction during drought in the rainy site and a slightly significant growth reduction at the mesic site for both saplings and adults. Across organs and in both sites and age classes, NSC, starch, and sugar concentrations were generally higher in the first than in the second year following drought, while NSC seasonal remobilization was generally lower. Higher NSC concentrations along with lower seasonal NSC remobilization during the first post-drought year are supportive of sink limitation. However, as these results were found at both sites while growth decreased slightly and just at the mesic site, limited growth only is unlikely to have caused NSC accumulation. Rather, these results suggest that the post-drought dynamics of carbohydrate storage are partly decoupled from the growth dynamics, and that the rebuild of C reserves after drought may be a priority in this species.

Secondary leaves of an outbreak-adapted tree species are both more resource acquisitive and more herbivore resistant than primary leaves.

The magnitude and frequency of insect outbreaks are predicted to increase in forests, but how trees cope with severe outbreak defoliation is not yet fully understood. Winter deciduous trees often produce a secondary leaf flush in response to defoliation (i.e., compensatory leaf regrowth or refoliation), which promotes fast replenishment of carbon (C) storage and eventually tree survival. However, secondary leaf flushes may imply a high susceptibility to insect herbivory, especially in the event of an ongoing outbreak. We hypothesized that in winter deciduous species adapted to outbreak-driven defoliations, secondary leaves are both more C acquisitive and more herbivore resistant than primary leaves. During an outbreak by Ormiscodes amphimone F. affecting Nothofagus pumilio (Poepp. & Endl.) Krasser forests, we (i) quantified the defoliation and subsequent refoliation by analyzing the seasonal dynamics of the normalized difference vegetation index (NDVI) and (ii) compared the physiological traits and herbivore resistance of primary and secondary leaves. Comparisons of the NDVI of the primary and second leaf flushes relative to the NDVI of the defoliated forest indicated 31% refoliation, which is close to the leaf regrowth reported by a previous study in juvenile N. pumilio trees subjected to experimental defoliation. Primary leaves had higher leaf mass per area, size, carbon:nitrogen ratio and soluble sugar concentration than secondary leaves, along with lower nitrogen and starch concentrations, and similar total polyphenol and phosphorus concentrations. In both a choice and a non-choice bioassay, the leaf consumption rates by O. amphimone larvae were significantly higher (>50%) for primary than for secondary leaves, indicating higher herbivore resistance in the latter. Our study shows that secondary leaf flushes in outbreak-adapted tree species can be both C acquisitive and herbivore resistant, and suggests that these two features mediate the positive effects of the compensatory leaf regrowth on the tree C balance and forest resilience.

Cluster root-bearing Proteaceae species show a competitive advantage over non-cluster root-bearing species.

BACKGROUND AND AIMS: Cluster roots (CRs) constitute a special root adaptation that enables plants to take up nutrients, especially phosphorus (P), from soils with low nutrient availability, including recent volcanic deposits. It is unclear, however, how CR species interact with non-cluster root-bearing (NCR) species, and how substrates' fertility modulates potential interactions. METHODS: We experimentally assessed the net interaction between CR and NCR species using two substrates of contrasting fertility: nutrient-rich nursery mix and tephra (low P availability). We planted seedlings of two southern South American (SSA) Proteaceae, CR species and two NCR Nothofagus species in pairs (conspecifics and heterospecifics) and as singles. We analysed the effect of seedling neighbours on survival, growth performance (e.g. total biomass and leaf area) and leaf and substrate nutrient concentrations (including manganese, a proxy for P-acquisition efficiency through CR activity) using the relative interaction index. KEY RESULTS: After three growing seasons, we found that (1) Proteaceae species had fewer CRs and lower CR biomass and grew less in the tephra than in the nursery substrate; (2) Nothofagus species did not improve their survival and growth in the presence of Proteaceae species in any substrate; (3) contrary to Nothofagus, Proteaceae species improved their growth more when planted with any neighbour (including conspecifics) than when planted alone, which was accompanied by a significant accretion of leaf P; and (4) the presence of a neighbour increased the final nitrogen and P concentrations in the nursery substrate, regardless of species identity. CONCLUSIONS: CRs provide Proteaceae a competitive advantage over NCR species at the seedling stage, which may have important consequences for species coexistence and community structuring. The investigated SSA Proteaceae, which have not evolved in nutrient-impoverished soils, as have their relatives in south-western Australia and South Africa, improve their growth when cultivated in pairs, especially in nutrient-rich substrates.

Herbivore resistance in congeneric and sympatric Nothofagus species is not related to leaf habit.

PREMISE: Two fundamental hypotheses on herbivore resistance and leaf habit are the resource availability hypothesis (RAH) and the carbon-nutrient balance hypothesis (CNBH). The RAH predicts higher constitutive resistance by evergreens, and the CNBH predicts higher induced resistance by deciduous species. Although support for these hypotheses is mixed, they have rarely been examined in congeneric species. METHODS: We compared leaf constitutive and induced resistance (as leaf polyphenol and tannin concentrations, and as damage level in non-choice experiments) and leaf traits associated with herbivory of coexisting Nothofagus species using (1) a defoliation experiment and (2) natural defoliation caused by an outbreak of a common defoliator of Nothofagus species. RESULTS: In the defoliation experiment, polyphenol and tannin concentrations were similar between deciduous and evergreen species; regardless of leaf habit, polyphenols increased in response to defoliation. In the natural defoliation survey, N. pumilio (deciduous) had significantly higher herbivory, lower carbon/nitrogen ratio and leaf mass per area, and higher nitrogen and phosphorus concentrations than N. betuloides (evergreen); N. antarctica (deciduous) had intermediate values. Polyphenol concentrations and herbivore resistance indicated by the non-choice experiment were lower in N. pumilio than in N. antarctica and N. betuloides, which had similar values. CONCLUSIONS: Higher herbivory in N. pumilio was associated with a higher nutritional value and a lower level of leaf carbon-based defenses compared to both the evergreen and the other deciduous species, indicating that herbivore resistance in Nothofagus species cannot be attributed to only leaf habit as predicted by the RAH or CNBH.

Revisiting the relative growth rate hypothesis for gymnosperm and angiosperm species co-occurrence.

PREMISE OF THE STUDY: It is unclear to what extent the co-occurrence of angiosperm and gymnosperm species in some marginal ecosystems is explained by reduced growth in angiosperms due to carbon (C) limitation and by high stress tolerance in gymnosperms associated with lack of vessels and resource conservation. METHODS: We examined growth patterns and traits associated with C balance in four evergreen angiosperm species (including one vesselless species, Drimys winteri) and three gymnosperm tree species of a cold-temperate rainforest in southern Chile. We measured the mean basal area increment for the first 50 (BAI50 ) and the last 10 years (BAI10 ), wood density, leaf lifespan, and nonstructural carbohydrate (NSC) concentrations in different organs. KEY RESULTS: BAI50 was 6-fold higher in angiosperms than in gymnosperms and ca. 4-fold higher in Drimys than in the fastest-growing gymnosperm. BAI10 and aboveground NSC concentrations were significantly higher and leaf lifespan lower in angiosperms than in gymnosperms; these differences, however, were largely driven by the slow growth and low NSC concentrations of the Cupressaceae species (Pilgerodendron uviferum), while the two Podocarpaceae had BAI10 and NSC concentrations similar to angiosperms. In angiosperms, NSC and starch concentrations were generally higher in species with lower BAI10 , indicating no severe C limitation. CONCLUSIONS: The co-occurrence of angiosperms and gymnosperms in cold-temperate rainforests of southern Chile is not explained by growth disadvantages and C limitation in angiosperms. Long leaf longevity, but not lack of vessels, appeared to favor resource conservation and C balance in some gymnosperms (Podocarpaceae).

A multi-species synthesis of physiological mechanisms in drought-induced tree mortality.

Widespread tree mortality associated with drought has been observed on all forested continents and global change is expected to exacerbate vegetation vulnerability. Forest mortality has implications for future biosphere-atmosphere interactions of carbon, water and energy balance, and is poorly represented in dynamic vegetation models. Reducing uncertainty requires improved mortality projections founded on robust physiological processes. However, the proposed mechanisms of drought-induced mortality, including hydraulic failure and carbon starvation, are unresolved. A growing number of empirical studies have investigated these mechanisms, but data have not been consistently analysed across species and biomes using a standardized physiological framework. Here, we show that xylem hydraulic failure was ubiquitous across multiple tree taxa at drought-induced mortality. All species assessed had 60% or higher loss of xylem hydraulic conductivity, consistent with proposed theoretical and modelled survival thresholds. We found diverse responses in non-structural carbohydrate reserves at mortality, indicating that evidence supporting carbon starvation was not universal. Reduced non-structural carbohydrates were more common for gymnosperms than angiosperms, associated with xylem hydraulic vulnerability, and may have a role in reducing hydraulic function. Our finding that hydraulic failure at drought-induced mortality was persistent across species indicates that substantial improvement in vegetation modelling can be achieved using thresholds in hydraulic function.

Single-provenance mature conifers show higher non-structural carbohydrate storage and reduced growth in a drier location.

Since growth is more sensitive to drought than photosynthesis, trees inhabiting dry regions are expected to exhibit higher carbohydrate storage and less growth than their conspecifics from more humid regions. However, the same pattern can be the result of different genotypes inhabiting contrasting humidity conditions. To test if reduced growth and high carbohydrate storage are environmentally driven by drought, we examined the growth and non-structural carbohydrate (NSC) concentrations in single-provenance stands of mature trees of Pinus contorta Douglas and Pinus ponderosa Douglas ex C. Lawson planted at contrasting humidity conditions (900 versus 300 mm of annual precipitation) in Patagonia, Chile. Individual tree growth was measured for each species and at each location as mean basal area increment of the last 10 years (BAI10), annual shoot elongation for the period 2011-14, and needle length for 2013 and 2014 cohorts. Additionally, needle, branch, stem sapwood and roots were collected from each sampled tree to determine soluble sugars, starch and total NSC concentrations. The two species showed lower mean BAI10 and 2013 needle length in the dry site; P. ponderosa also had lower annual shoot extension for 2011 and 2014, and lower 2014 needle length, in the dry than in the mesic site. By contrast, NSC concentrations of all woody tissues for both species were either similar or higher in the dry site when compared with the mesic site. Patterns of starch and sugars were substantially different: starch concentrations were similar between sites except for roots of P. ponderosa, which were higher in the dry site, while sugar concentrations of all woody tissues in both species were higher in the dry site. Overall, our study provides evidence that reduced growth along with carbon (C) accumulation is an environmentally driven response to drought. Furthermore, the significant accumulation of low-molecular weight sugars in the dry site is compatible with a prioritized C allocation for osmoregulation. However, since this accumulation did not come at the expense of reduced starch, it is unlikely that growth was limited by C supply in the dry site.

Carbon allocation to growth and storage in two evergreen species of contrasting successional status.

PREMISE OF THE STUDY: A prevailing hypothesis in forest succession is that shade-tolerant species grow more slowly than shade-intolerant species, across light conditions, because they prioritize carbon (C) allocation to storage. We examined this hypothesis in a confamilial pair of species, including one of the fastest-growing tree species in the world (Eucalyptus globulus) and a shade-tolerant, slow-growing species (Luma apiculata). METHODS: Seedlings were subjected to one out of four combinations of light (high vs. low) and initial defoliation (90% defoliated vs. nondefoliated) for four months. Growth, C storage concentration in different organs, leaf shedding, and lateral shoot formation were measured at the end of the experiment. KEY RESULTS: Eucalyptus globulus grew faster than L. apiculata in high light, but not in low light. Both species had lower C storage concentration in low than in high light, but similar C storage concentrations in each light condition. Defoliation had no effect on C storage, except in the case of the old leaves of both species, which showed lower C storage levels in response to defoliation. Across treatments, leaf shedding was 96% higher in E. globulus than in L. apiculata while, in contrast, lateral shoot formation was 87% higher in L. apiculata. CONCLUSIONS: In low light, E. globulus prioritized C storage instead of growth, whereas L. apiculata prioritized growth and lateral branching. Our results suggest that shade tolerance depends on efficient light capture rather than C conservation traits.

Carbon dynamics of Acer pseudoplatanus seedlings under drought and complete darkness.

Carbon (C) storage is considered a key component to plant survival under drought and shade, although the combined effects of these factors on survival remain poorly understood. We investigated how drought and shade alter the C dynamics and survival of tree seedlings, and whether drought limits the access to or usage of stored C. We experimentally applied two levels of soil humidity (well-watered versus drought, the latter induced by dry-down) and light availability (light versus complete darkness) on 1-year-old seedlings of Acer pseudoplatanus L. for 3 months. We quantified the survival, biomass, growth rate and non-structural carbohydrates (NSC) of seedlings at their time of death or at the end of the experiment for those that survived. We found that the soil dried out faster when drought was combined with light than when it was combined with complete darkness. Seedlings subjected to both drought and light showed reduced growth and reached 100% mortality earlier than any other treatment, with the highest NSC concentrations at the time of death. Seedlings exposed to both drought and complete darkness died significantly earlier than seedlings exposed to complete darkness only, but had similar NSC concentrations at time of their death, suggesting that drought accelerated the use of stored C under complete darkness. Complete darkness significantly reduced seedling growth and whole-plant NSC concentrations regardless of soil humidity, while root NSC concentrations were significantly more reduced when complete darkness was combined with drought conditions. Thus, the C dynamics in A. pseudoplatanus seedlings under complete darkness was not hindered by drought, i.e., the access and use of stored C was not limited by drought. The contrasting growth and C storage responses driven by drought under light versus complete darkness are consistent with a key role of the drought progression in the C dynamics of trees.

Patterns of carbon storage in relation to shade tolerance in southern South American species.

PREMISE OF THE STUDY: Carbon (C) allocation to storage in woody tissues at the expense of growth is thought to promote shade tolerance, yet few studies on the subject examined C storage during maximum growth and considered stand influences. I asked how C storage in different plant tissues relates to shade tolerance in temperate forests with contrasting climates and physiognomies, and whether relationships vary during the growing season. METHODS: In the late spring and late summer, I harvested seedlings of eight species with contrasting light requirements from the understory of a cold rainforest and a Mediterranean forest in Chile. Nonstructural carbohydrate (NSC) concentrations and pools (i.e., biomass x NSC concentration) were determined in leaves, aboveground wood, and roots. The effects of shade tolerance and sampling date on the NSCs were analyzed for each forest and tissue with linear mixed-effects models. KEY RESULTS: In both forests, concentrations of NSC and soluble sugars in woody tissues, as well as fractions of NSC in these tissues, were lower in shade tolerant than in shade intolerant species. For root NSC concentrations, these trends depended on the sampling date: in the late spring the concentrations were similar in shade tolerant and intolerant species, while in the late summer they were lower in shade tolerant species. CONCLUSIONS: Shade tolerance is not linked to C storage in the two studied forests, suggesting that allocation to growth or defenses could be more advantageous for low light persistence. Alternatively, high levels of C storage could be also selected in shade intolerant species to face herbivory or drought.

Extreme defoliation reduces tree growth but not C and N storage in a winter-deciduous species.

BACKGROUND AND AIMS: There is a growing concern about how forests will respond to increased herbivory associated with climate change. Carbon (C) and nitrogen (N) limitation are hypothesized to cause decreasing growth after defoliation, and eventually mortality. This study examines the effects of a natural and massive defoliation by an insect on mature trees' C and N storage, which have rarely been studied together, particularly in winter-deciduous species. METHODS: Survival, growth rate, carbon [C, as non-structural carbohydrate (NSC) concentration] and nitrogen (N) storage, defences (tannins and total polyphenols), and re-foliation traits were examined in naturally defoliated and non-defoliated adult trees of the winter-deciduous temperate species Nothofagus pumilio 1 and 2 years after a massive and complete defoliation caused by the caterpillar of Ormiscodes amphimone (Saturniidae) during summer 2009 in Patagonia. KEY RESULTS: Defoliated trees did not die but grew significantly less than non-defoliated trees for at least 2 years after defoliation. One year after defoliation, defoliated trees had similar NSC and N concentrations in woody tissues, higher polyphenol concentrations and lower re-foliation than non-defoliated trees. In the second year, however, NSC concentrations in branches were significantly higher in defoliated trees while differences in polyphenols and re-foliation disappeared and decreased, respectively. CONCLUSIONS: The significant reduction in growth following defoliation was not caused by insufficient C or N availability, as frequently assumed; instead, it was probably due to growth limitations due to factors other than C or N, or to preventative C allocation to storage. This study shows an integrative approach to evaluating plant growth limitations in response to disturbance, by examining major resources other than C (e.g. N), and other C sinks besides storage and growth (e.g. defences and re-foliation).

High foliar nutrient concentrations and resorption efficiency in Embothrium coccineum (Proteaceae) in southern Chile.

PREMISE OF THE STUDY: Southern South American (SA) Proteaceae species growing in volcanic soils have been proposed as potential ecosystem engineers by tapping phosphorus (P) from soil through their cluster roots and shedding nutrient-rich litter to the soil, making it available for other species. We tested whether Embothrium coccineum (Proteaceae) has effectively lower P nutrient resorption efficiency and higher litter P concentrations than co-occurring, non-Proteaceae species. METHODS: In southern Chile, we assessed the P and nitrogen (N) resorption efficiency of senescent leaves and fresh litter of E. coccineum and co-occurring tree species in a soil fertility and moisture gradient (600-3000 mm of annual precipitation) in Patagonia, Chile. We determined P and N concentrations, leaf mass per area (LMA), and ratios of N/P and C/N in mature and senescent leaf cohorts and fresh litter. KEY RESULTS: Embothrium coccineum showed significantly higher P and N resorption efficiency than co-occurring species; in fact, E. coccineum fresh litter had the lowest P-content. While E. coccineum showed significantly lower fresh litter P concentrations than the rest of the species, it showed significantly higher N concentrations. Embothrium coccineum also had lower LMA and similar N/P and C/N ratios when compared with co-occurring tree species. CONCLUSIONS: We found that E. coccineum efficiently mobilized P and, to a lesser extent, N before leaf shedding. We did not find support for the ecosystem engineering hypothesis via shedding P-rich litter. We suggest that southern South American Proteaceae may be taking up other nutrients besides P, probably N, from the young, volcanic soils of this region.

An experimental approach to explain the southern Andes elevational treeline.

UNLABELLED: • PREMISE OF THE STUDY: The growth limitation hypothesis (GLH) is the most accepted mechanistic explanation for treeline formation, although it is still uncertain whether it applies across taxa. The successful establishment of Pinus contorta--an exotic conifer species in the southern hemisphere--above the Nothofagus treeline in New Zealand may suggest a different mechanism. We tested the GLH in Nothofagus pumilio and Pinus contorta by comparing seedling performance and carbon (C) balance in response to low temperatures.• METHODS: At a southern Chilean treeline, we grew seedlings of both species 2 m above ground level, to simulate coupling between temperatures at the meristem and in the air (colder), and at ground level, i.e., decoupling air temperature (relatively milder). We recorded soil and air temperatures as well. After 3 yr, we measured seedling survival and biomass (as a surrogate of growth) and determined nonstructural carbohydrates (NSC).• KEY RESULTS: Nothofagus and Pinus did not differ in survival, which, as a whole, was higher at ground level than at the 2-m height. The root-zone temperature for the growing season was 6.6°C. While biomass and NSC decreased significantly for Nothofagus at the 2-m height compared with ground level (C limitation), these trends were not significant for Pinus• CONCLUSIONS: The treeline for Nothofagus pumilio is located at an isotherm that fully matches global patterns; however, its physiological responses to low temperatures differed from those of other treeline species. Support for C limitation in N. pumilio but not in P. contorta indicates that the physiological mechanism explaining their survival and growth at treeline may be taxon-dependent.

Soil nitrogen, and not phosphorus, promotes cluster-root formation in a South American Proteaceae, Embothrium coccineum.

PREMISE OF THE STUDY: Cluster roots are a characteristic root adaptation of Proteaceae species. In South African and Australian species, cluster roots promote phosphorus (P) acquisition from poor soils. In a South American Proteaceae species, where cluster roots have been scarcely studied and their function is unknown, we tested whether cluster-root formation is stimulated by low soil nutrition, in particular low P-availability. METHODS: Small and large seedlings (< 6- and > 6-months old, respectively) of Embothrium coccineum and soil were collected across four different sites in Patagonia (Chile). We determined cluster-root number and relative mass, and leaf Pi concentration per mass (Pimass) and per area (Piarea) for each seedling, and tested relationships with Olsen-P (OP), sorbed-P (sP) and total nitrogen (N) using generalized linear mixed-effects models and model selection to assess the relative strength of soil and plant drivers. KEY RESULTS: Best-fit models showed a negative logarithmic relationship between cluster-root number and soil nitrogen (N), and between cluster-root relative mass and both leaf Piarea and soil N, and a positive logarithmic relationship between cluster-root number and leaf Piarea. Cluster-root relative mass was higher in small than in large seedlings. CONCLUSIONS: Contrary to that found in South African and Australian Proteaceae, cluster roots of E. coccineum do not appear to be driven by soil P, but rather by soil N and leaf Piarea. We suggest that cluster roots are a constitutive and functional trait that allows plants to prevail in poor N soils.