Xylem anatomy needs to change, so that conductivity can stay the same: xylem adjustments across elevation and latitude in Nothofagus pumilio.

BACKGROUND AND AIMS: Plants have the potential to adjust the configuration of their hydraulic system to maintain its function across spatial and temporal gradients. Species with wide environmental niches provide an ideal framework to assess intraspecific xylem adjustments to contrasting climates. We aimed to assess how xylem structure in the widespread species Nothofagus pumilio varies across combined gradients of temperature and moisture, and to what extent within-individual variation contributes to population responses across environmental gradients. METHODS: We characterized xylem configuration in branches of N. pumilio trees at five sites across an 18° latitudinal gradient in the Chilean Andes, sampling at four elevations per site. We measured vessel area, vessel density and the degree of vessel grouping. We also obtained vessel diameter distributions and estimated the xylem-specific hydraulic conductivity. Xylem traits were studied in the last five growth rings to account for within-individual variation. KEY RESULTS: Xylem traits responded to changes in temperature and moisture, but also to their combination. Reductions in vessel diameter and increases in vessel density suggested increased safety levels with lower temperatures at higher elevation. Vessel grouping also increased under cold and dry conditions, but changes in vessel diameter distributions across the elevational gradient were site-specific. Interestingly, the estimated xylem-specific hydraulic conductivity remained constant across elevation and latitude, and an overwhelming proportion of the variance of xylem traits was due to within-individual responses to year-to-year climatic fluctuations, rather than to site conditions. CONCLUSIONS: Despite conspicuous adjustments, xylem traits were coordinated to maintain a constant hydraulic function under a wide range of conditions. This, combined with the within-individual capacity for responding to year-to-year climatic variations, may have the potential to increase forest resilience against future environmental changes.

Fire damage to cambium affects localized xylem anatomy and hydraulics: the case of Nothofagus pumilio in Patagonia.

PREMISE: Fire scars on trees are created by excessive heat from a fire that kills the vascular cambium. Although, fires are one of the most important forest disturbances in Patagonia, the effects of fire on tree physiology and wood anatomy are still unknown. In this study, we hypothesized that abnormal functioning of the cambium after a fire will induce anatomical changes in the wood. We also assumed that these anatomical changes would affect xylem safety transport. METHODS: We quantified wood anatomical traits in Nothofagus pumilio, the dominant subalpine tree species of Patagonia, using two approaches: time and distance. In the first, anatomical changes in tree rings were compared before, during, and after fire occurrence. In the second, the spatial extent of these changes was evaluated with respect to the wound by measuring anatomical traits in sampling bands in two directions (0° and 45°) with respect to the onset of healing. RESULTS: Reductions in lumen diameter and vessel number were the most conspicuous changes associated with fire damage and observed in the fire ring and subsequent post-fire rings. In addition, the fire ring had more rays than in control rings. In terms of distance, anatomical changes were only restricted to short distances from the wound. CONCLUSIONS: Post-fire changes in wood anatomical traits were confined close to the wound margins. These changes might be associated with a defense strategy related to the compartmentalization of the wound and safety of water transport.

Insights into intraspecific wood density variation and its relationship to growth, height and elevation in a treeline species.

The wood economics spectrum provides a general framework for interspecific trait-trait coordination across wide environmental gradients. Whether global patterns are mirrored within species constitutes a poorly explored subject. In this study, I first determined whether wood density co-varies together with elevation, tree growth and height at the within-species level. Second, I determined the variation of wood density in different stem parts (trunk, branch and twigs). In situ trunk sapwood, trunk heartwood, branch and twig densities, in addition to stem growth rates and tree height were determined in adult trees of Nothofagus pumilio at four elevations in five locations spanning 18° of latitude. Mixed effects models were fitted to test relationships among variables. The variation in wood density reported in this study was narrow (ca. 0.4-0.6 g cm-3 ) relative to global density variation (ca. 0.3-1.0 g cm-3 ). There was no significant relationship between stem growth rates and wood density. Furthermore, the elevation gradient did not alter the wood density of any stem part. Trunk sapwood density was negatively related to tree height. Twig density was higher than branch and trunk densities. Trunk heartwood density was always significantly higher than sapwood density. Negative across-species trends found in the growth-wood density relationship may not emerge as the aggregate of parallel intraspecific patterns. Actually, trees with contrasting growth rates show similar wood density values. Tree height, which is tightly related to elevation, showed a negative relationship with sapwood density.

Does water availability regulate biomass partitioning between trunk and branches?

The extent to which a vertical trunk is differentiated from its branches is a key trait for the architecture of trees and may affect interspecific relationships. In this study, we analysed the effect of soil water availability on biomass partitioning for Nothofagus pumilio by means of a nursery experiment. Juvenile trees were subject to three irrigation conditions: no irrigation, intermediate irrigation and high irrigation. Irrigation conditions emulated the mean precipitation of the most representative environments inhabited by N. pumilio. Changes in soil water availability modified the biomass partitioning patterns of trees. In comparison to the other two conditions, high irrigation caused: (i) a higher ratio of biomass partitioning to stems than roots; (ii) more trunk growth in relation to its branches; and (iii) more photosynthetic organs relative to the aboveground biomass. Trunk size relative to that of its most recent branches was not increased by water availability. Water availability may play a significant role in the capacity of N. pumilio for space occupation due to the effects on axis differentiation.

Disturbance and density-dependent processes (competition and facilitation) influence the fine-scale genetic structure of a tree species' population.

BACKGROUND AND AIMS: Disturbances, dispersal and biotic interactions are three major drivers of the spatial distribution of genotypes within populations, the last of which has been less studied than the other two. This study aimed to determine the role of competition and facilitation in the degree of conspecific genetic relatedness of nearby individuals of tree populations. It was expected that competition among conspecifics will lead to low relatedness, while facilitation will lead to high relatedness (selection for high relatedness within clusters). METHODS: The stand structure and spatial genetic structure (SGS) of trees were examined within old-growth and second-growth forests (including multi-stemmed trees at the edge of forests) of Nothofagus pumilio following large-scale fires in Patagonia, Chile. Genetic spatial autocorrelations were computed on a spatially explicit sampling of the forests using five microsatellite loci. As biotic plant interactions occur among immediate neighbours, mean nearest neighbour distance (MNND) among trees was computed as a threshold for distinguishing the effects of disturbances and biotic interactions. KEY RESULTS: All forests exhibited a significant SGS for distances greater than the MNND. The old-growth forest genetic and stand structure indicated gap recolonization from nearby trees (significantly related trees at distances between 4 and 10 m). At distances smaller than the MNND, trees of the second-growth interior forest showed significantly lower relatedness, suggesting a fading of the recolonization structure by competition, whereas the second-growth edge forest showed a positive and highly significant relatedness among trees (higher among stems of a cluster than among stems of different clusters), resulting from facilitation. CONCLUSIONS: Biotic interactions are shown to influence the genetic composition of a tree population. However, facilitation can only persist if individuals are related. Thus, the genetic composition in turn influences what type of biotic interactions will take place among immediate neighbours in post-disturbance forests.

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).

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.

Similar variation in carbon storage between deciduous and evergreen treeline species across elevational gradients.

BACKGROUND AND AIMS: The most plausible explanation for treeline formation so far is provided by the growth limitation hypothesis (GLH), which proposes that carbon sinks are more restricted by low temperatures than by carbon sources. Evidence supporting the GLH has been strong in evergreen, but less and weaker in deciduous treeline species. Here a test is made of the GLH in deciduous-evergreen mixed species forests across elevational gradients, with the hypothesis that deciduous treeline species show a different carbon storage trend from that shown by evergreen species across elevations. METHODS: Tree growth and concentrations of non-structural carbohydrates (NSCs) in foliage, branch sapwood and stem sapwood tissues were measured at four elevations in six deciduous-evergreen treeline ecotones (including treeline) in the southern Andes of Chile (40°S, Nothofagus pumilio and Nothofagus betuloides; 46°S, Nothofagus pumilio and Pinus sylvestris) and in the Swiss Alps (46°N, Larix decidua and Pinus cembra). KEY RESULTS: Tree growth (basal area increment) decreased with elevation for all species. Regardless of foliar habit, NSCs did not deplete across elevations, indicating no shortage of carbon storage in any of the investigated tissues. Rather, NSCs increased significantly with elevation in leaves (P < 0·001) and branch sapwood (P = 0·012) tissues. Deciduous species showed significantly higher NSCs than evergreens for all tissues; on average, the former had 11 % (leaves), 158 % (branch) and 103 % (sapwood) significantly (P < 0·001) higher NSCs than the latter. Finally, deciduous species had higher NSC (particularly starch) increases with elevation than evergreens for stem sapwood, but the opposite was true for leaves and branch sapwood. CONCLUSIONS: Considering the observed decrease in tree growth and increase in NSCs with elevation, it is concluded that both deciduous and evergreen treeline species are sink limited when faced with decreasing temperatures. Despite the overall higher requirements of deciduous tree species for carbon storage, no indication was found of carbon limitation in deciduous species in the alpine treeline ecotone.

New host-plant records for the defoliator Ormiscodes amphimone (Fabricius) (Lepidoptera: Saturniidae)

Ormiscodes amphimone (Fabricius) is a phytophagous moth species known to severely defoliate woody species in Chile and Argentina. Here we document new records of O. amphimonehost associations emphasizing the role of Nothofagus pumilio as its primary host in our study area. This new record for Argentina is highly significant given the economic importance of N. pumilio as a timber resource and the potential of O. amphimone to generate extensive outbreaks.