Growth patterns of <Emphasis Type="Italic">Nothofagus pumilio</Emphasis> trees along altitudinal gradients in Tierra del Fuego,Argentina

The influence of ecological factors resulting from altitude on growth characteristics of Nothofagus pumilio was analyzed. N. pumilio forests grow in the southern Andes from near sea level to 600 m.a.s.l. Eight study sites were selected in the southwestern area of Tierra del Fuego, on north- and south-facing slopes, and distributed along an altitudinal gradient (200–600 m.a.s.l.). Growth cycles throughout time were identified. The correlation between climate and tree growth was determined. Tree diameter and height decreased whereas stand density increased towards tree-line. South-facing slopes showed higher values in common growth variability, probably because of stronger climatic controlling factors. Climate-growth correlation models revealed long-term temperature influences on growth variability, and showed variations among the different plots. Differences were detected in the spectral power component of the chronologies throughout the altitudinal profile. The series had a significant spectral peak at frequencies around 7 years, possibly related to mast seeding cycles. This signal became weaker as the altitude increased probably due to lower seed production. These results suggest a strong altitudinal component in growth variability of N. pumilio in Tierra del Fuego.     

Morphological and phenological differences in Nothofagus pumilio from contrasting elevations: Evidence from a common garden

Abstract We tested the hypothesis that contrasting elevations select distinct growth patterns and vegetative phenology in Nothofagus pumilio, a winter deciduous tree that dominates mountain forests of Patagonia. Analysis of saplings maintained under common‐garden conditions for 4 years showed a significant decrease in shoot annual growth, leaf size, and a delay in bud‐break, and leaf expansion with increased elevation of their site of origin. Rapid gain in height seems to be advantageous at low elevation in such light‐demanding species. Lower stature high‐elevation plants have wider branching angles and greater branching ratios (number of branches/number of internodes) than low‐elevation plants. Compact growth at high elevation may be related to strong winds and irradiance. Plants from different elevations had distinct growth patterns during the common‐garden experiment. This could be of importance in Mediterranean‐climate areas characterized by highly unpredictable precipitation regimes. Also, liberation of growth‐suppressed seedlings may follow different environmental signals in low‐ and high‐habitats, which might explain such time‐dependent responses to optimal conditions under cultivation. While these greenhouse‐grown N. pumilio saplings showed heritable differences in plant architectural traits and leafing phenology, it was not clear how the genotypes characteristic of particular elevations would respond to longer growing seasons such as those predicted under global warming.     

Regional and local effects of disturbance and climate on altitudinal treelines in northern Patagonia

Abstract. In this field study we analysed the regional and local scale effects of disturbance and climate on altitudinal treelines dominated by Nothofagus pumilio in northern Patagonia. We compared two regions west and east of the Andes at 40° S, slopes with warm vs cool aspects and undisturbed vs locally disturbed treelines. This spatial framework allowed us to test (1) for differences among treelines affected by different types of local disturbance and (2) the traditional hypothesis that low temperature limits treeline. Contingency tables and ANOVA showed that local disturbance occurred more frequently than expected on slopes with cool aspects, steep slope angles and concave slope configuration. Disturbed treelines were locally lowered with longer ecotones and lower krummholz growth rates and vegetation cover than undisturbed treelines. Three‐way ANOVA showed the significant influences of study area (regional climate) and aspect (local climate) on treeline elevation, krummholz growth rates and density, tree density and vegetation cover, while accounting for local disturbance. Treeline elevations were higher east of the Andes reflecting the more continental climate in Argentina than in Chile, plus regional impacts of volcanic eruptions. Tree density and vegetation cover were greater west of the Andes reflecting greater precipitation in Chile. Within study areas, local climate had different influences on treeline elevations and krummholz growth rates west and east of the Andes. We predict that increased tree growth and upslope advance of treeline in response to global warming is more likely in Chile than in Argentina near 40° S, unless precipitation increases east of the Andes. To test these predictions, we recommend research be stratified to account for the influences of local disturbance, which may confound climatic impacts. In northern Patagonia, suitable control (undisturbed) study sites will most likely be found at upper slope positions with low slope angles, simple microtopography and straight topographic configuration.     

Latitudinal decrease in folivory within Nothofagus pumilio forests: dual effect of climate on insect density and leaf traits?

Aim The strength of consumer–plant interactions may decrease with latitude. Our objectives were to assess the spatial variation in folivory on Nothofagus pumilio and understand the influence of climate on folivory patterns as mediated by changes in folivore density and leaf traits. Location Nothofagus pumilio forests, between 38 and 55°S (Argentina). Methods We studied the correlation of leaf damage with latitude on data from 47 sampling sites, and evaluated spatial patterns of autocorrelation on latitudinally detrended data with a principal coordinates of neighbour matrices method. Path analysis was used to test the association of temperature and precipitation with leaf damage, mediated by folivore density and leaf traits. We evaluated the adequacy of this ecological model by examining the spatial pattern of autocorrelation in the residuals, and combined spatial and environmental predictors of leaf damage into partial regression. Results Leaf damage decreased with latitude, which was the only significant spatial predictor. The latitudinal decrease in temperature and precipitation was correlated with a decrease in the density of folivores and leaf size, and diminished leaf damage. Our ecological model adequately explained the spatial autocorrelation in the data: 44% of the variation in leaf damage was explained by the latitudinally structured component of the environment, whereas local environmental effects accounted for another 22%. Main conclusions We conclude that N. pumilio forests show consistent latitudinal patterns of variation in folivory, folivore density and leaf traits. Our study suggests that the latitudinal variation in folivory rates is partly driven by the influence of climate on both plants and herbivores. This warns us about the potential susceptibility of folivory rates to climate warming. We emphasize the value of large‐scale analyses as complementary to local experimental approaches to understanding the regulation of herbivory.     

Mass loss and nutrient release from decomposing evergreen and deciduous Nothofagus litters from the Chilean Andes

Abstract Leaf litter decomposition experiments were conducted on two deciduous (Nothofagus obliqua (roble)) and Nothofagus pumilio (lenga)) and one evergreen (Nothofagus dombeyi) Nothofagus (Nothofagaceae) species from a single Chilean forest in order to understand how congeneric trees with differing leaf lifespans impact the soil in which they grow. Single‐species litter samples were decomposed in a mixed hardwood forest in Ohio and in a deciduous‐evergreen Nothofagus forest in Chile. In the Ohio forest, the two deciduous species’ litters decomposed at k ≈ 1.00 per year and the evergreen at k ≈ 0.75 per year. In Chile k ranged from k ≈ 0.06 (N. obliqua) to k ≈ 0.23 (N. pumilio) per year. In both experiments, N and P were released faster from the deciduous litters than from evergreen litter. In Ohio, evergreen litter immobilized more N and P for a longer time period than did deciduous litter. As N. dombeyi stands tend to have lower available soil N and P in this particular mixed Nothofagus forest, the increased time of N and P immobilization by N. dombeyi litter suggests a feedback role of the tree itself in perpetuating low N and P soil conditions.     

Climatic significance of intra-annual bands in the wood of <Emphasis Type="Italic">Nothofagus pumilio</Emphasis> in southern Patagonia

Nothofagus pumilio (Poepp et Endl.) Krasser is the dominant subalpine tree in the southern Andes between 36°S and 55°S. Increment cores from a N. pumilio stand growing near the Ameghino Glacier (50°25S, 73°10W), southern Patagonian Andes, Argentina, contain significant numbers of intra-annual bands (or false rings) which have not been previously described for this species. These samples are used to develop a well-replicated ring-width chronology and a record of intra-annual bands from AD 1760 to 1997. Annual variations in radial growth of N. pumilio at this site are negatively correlated with spring–summer temperatures and positively with spring precipitation. The formation of intra-annual bands appears to be a response to anomalously dry-warm springs followed by wet-warm late summers. Intra-annual bands may occur in up to 95% of the sampled trees in a given year, and the percentage of trees affected was used as an indication of the strength of the forcing event. Narrow rings occurred in the years following intra-annual band formation, reflecting the lagged effect of unfavorable climatic conditions on tree growth during the subsequent growing season. Intra-annual bands occurred more frequently in the twentieth century than the late eighteenth and nineteenth centuries. This contrasting pattern seems to be a response to the combination of a long-term warming trend and a significant decrease in precipitation recorded during the last 100 years in this region of southern South America.

Age structure and dynamics of Patagonian beech forests in Torres del Paine National Park,Chile

This study documents the stem size and age-structure in forests dominated by different species of Nothofagus in Torres del Paine National Park (51° S), in the Chilean Patagonian region. We also explored the relationship between the various types of Nothofagus forest and postglacial succession. Pioneer stands on moraine fields 1–2 km of the glacier front are dominated by Nothofagus betuloides and Nothofagus antarctica. Moraines appear to be first colonized by the evergreen N. betuloides, followed within 5–7 years by deciduous N. antarctica. Nothofagus antarctica may replace the former species and develop monospecific stands on glacial valleys. Most trees in the N. antarctica stand studied were older than 40 years and floods may cause a significant mortality of young trees. Recruitment from seed seems to be infrequent. Old-growth stands dominated by deciduous Nothofagus pumilio occupy more stable substrates, and probably represent the last stage of postglacial succession. This long-lived tree species had recorded ages over 200 years. The canopy of N. pumilio forests appears to be a mosaic of even-aged, old-growth patches. We propose that regeneration episodes follow the blowdown of a large portion of the canopy, with long intervals with little or no regeneration. Windstorms may be an important force influencing the regeneration of N. pumilio. Exogenous disturbances, such as floods and windstorms, are an integral part of the forest cycle in the Patagonian region.     

Patterns and drivers of Araucaria araucana forest growth along a biophysical gradient in the northern Patagonian Andes: Linking tree rings with satellite observations of soil moisture

Araucaria araucana (Araucaria) is a long‐lived conifer growing along a sharp west–east biophysical gradient in the Patagonian Andes. The patterns and climate drivers of Araucaria growth have typically been documented on the driest part of the gradient relying on correlations with meteorological records, but the lack of in situ soil moisture observations has precluded an assessment of the growth responses to soil moisture variability. Here, we use a network of 21 tree‐ring width chronologies to investigate the spatiotemporal patterns of tree growth through the entire gradient and evaluate their linkages with regional climate and satellite‐observed surface soil moisture variability. We found that temporal variations in tree growth are remarkably similar throughout the gradient and largely driven by soil moisture variability. The regional spatiotemporal pattern of tree growth was positively correlated with precipitation (r = 0.35 for January 1920–1974; P < 0.01) and predominantly negatively correlated with temperature (r = ?0.38 for January–March 1920–1974; P < 0.01) during the previous growing season. These correlations suggest a temporally lagged growth response to summer moisture that could be associated with known physiological carry‐over processes in conifers and to a response to moisture variability at deeper layers of the rooting zone. Notably, satellite observations revealed a previously unobserved response of Araucaria growth to summer surface soil moisture during the current rather than the previous growing season (r = 0.65 for 1979–2000; P < 0.05). This new response has a large spatial footprint across the mid‐latitudes of the South American continent (35°–45°S) and highlights the potential of Araucaria tree rings for palaeoclimatic applications. The strong moisture constraint on tree growth revealed by satellite observations suggests that projected summer drying during the coming decades may result in regional growth declines in Araucaria forests and other water‐limited ecosystems in the Patagonian Andes.     

Climate-growth relationships of deciduous and evergreen Nothofagus species in Southern Patagonia,Argentina

Differences in growth patterns between a deciduous species, Nothofagus pumilio, and an evergreen species, Nothofagus betuloides, were analyzed in Lago del Desierto, southern Patagonia, Argentina (49°01′ S – 72°52′ W). The relationships between the growth rate of these two species and variations in temperature, precipitation, and the Southern Annular Mode (SAM) were also evaluated. We processed and analyzed 54 samples of N. pumilio and 48 of N. betuloides and developed two tree-ring width chronologies covering the periods of 1754-2014 and 1650-2014, respectively. The results from N. betuloides are especially important due to the small number of studies about this species in the area. Results indicate similarities between the tree-ring growth patterns of the two species. However, N. betuloides grew with a slower rate and had shorter periods of growth below average. We suggest that the particular growth pattern of N. betuloides is determined by its higher tolerance to stress situations and evergreen physiological characteristics. Significant correlations were observed between N. pumilio tree-ring growth and temperature from August-October of the previous year, and with the SAM index from December-January of the current year. N. pumilio did not show significant correlations with precipitation. Significant correlations were found between N. betuloides growth and precipitation from December-January of the previous year. As evergreen species have a more conservative response to temperature changes, N. betuloides growth was not significantly affected by temperature or by the SAM index. These results are encouraging and further studies could help improve our understanding of the relationship between the environment and species with different seasonality of foliage growth. Our study provides the first chronologies of N. pumilio and N. betuloides in the area and increases knowledge about the relationships between tree-ring growth and climate.     

Interannual variations in primary and secondary growth of Nothofagus pumilio and their relationships with climate

Key message

The aim of this study is to evaluate the relationships between primary and secondary growth as well as the influences of climate variations on both types of growth.

Abstract

The relationships between apical (or primary) and radial (or secondary) growth, and climatic influences on both types of growth, were evaluated for Nothofagus pumilio (Nothofagaceae), the dominant subalpine tree in Patagonia. We measured the spacing and number of nodes of annual shoots developed in the period 2001–2010 in 40 N. pumilio trees growing near the upper treeline in the northern Patagonian Andes (41°S). Variations in ring width at the base of each trunk were also recorded. Interannual variations in primary and secondary growth were significantly related to each other, and to several climate variables. Mean temperatures in winter and early spring (June–October) prior to the period of shoot extension were positively associated with both primary and secondary growth. In addition, total summer precipitation (December–March) was positively related to shoot extension, whereas mean summer temperature during the previous growth season (January–March) was directly related to radial growth. These climatic influences on N. pumilio growth may play a major role in regulating the expressions of preformation and neoformation.     

Influence of fire severity on stand development of Araucaria araucana–Nothofagus pumilio stands in the Andean cordillera of south‐central Chile

Fire is the prevalent disturbance in the Araucaria–Nothofagus forested landscape in south‐central Chile. Although both surface and stand‐replacing fires are known to characterize these ecosystems, the variability of fire severity in shaping forest structure has not previously been investigated in Araucaria–Nothofagus forests. Age structures of 16 stands, in which the ages of approximately 650 trees were determined, indicate that variability in fire severity and frequency is key to explaining the mosaic of forest patches across the Araucaria–Nothofagus landscape. High levels of tree mortality in moderate‐ to high‐severity fires followed by new establishment of Nothofagus pumilio typically result in stands characterized by one or two cohorts of this species. Large Araucaria trees are highly resistant to fire, and this species typically survives moderate‐ to high‐severity fires either as dispersed individuals or as small groups of multi‐aged trees. Small post‐fire cohorts of Araucaria may establish, depending on seed availability and the effects of subsequent fires. Araucaria's great longevity (often >700 years) and resistance to fire allow some individuals to survive fires that kill and then trigger new Nothofagus cohorts. Even in relatively mesic habitats, where fires are less frequent, the oldest Araucaria–Nothofagus pumilio stands originated after high‐severity fires. Overall, stand development patterns of subalpine Araucaria–N. pumilio forests are largely controlled by moderate‐ to high‐severity fires, and therefore tree regeneration dynamics is strongly dominated by a catastrophic regeneration mode.     

Nest habitat selection by the Austral parakeet in north‐western Patagonia

Identifying habitat or nesting microhabitat variables associated with high levels of nest success is important to understand nest site preferences and bird–habitat relationships. Little is known about cavity availability and nest site requirements of cavity nesters in southern hemisphere temperate forests, although nest site limitation is suggested. Here we ask which characteristics are selected by the Austral parakeet (Enicognathus ferrugineus) for nesting in Araucaria araucana–Nothofagus pumilio forest in Argentine Patagonia. We compared nest plot and tree characteristics with unused plots and trees among areas of different A. araucana–N. pumilio density. We also examine whether nest plot and tree use and selection, and the associated consequences for fitness of Austral parakeets are spatially related to forest composition. Austral parakeets showed selectivity for nests at different spatial scales, consistently choosing isolated live and large trees with particular nest features in a non‐random way from available cavities. Mixed A. araucana–N. pumilio forests are ideal habitat for the Austral parakeets of northern Patagonia, offering numerous potential cavities, mainly in N. pumilio. We argue that Austral parakeet reproduction and fitness is currently very unlikely to be limited by cavity availability, although this situation may be rapidly changing. Natural and human disturbances are modifying south temperate forests with even‐aged mid‐successional stands replacing old growth forests. Cavity nesting species use and need old growth forests, due to the abundance of cavities in large trees and the abundance of larvae in old wood. Neither of the latter resources is sufficiently abundant in mid‐successional forests, increasing the vulnerability and threatening the survival of the Austral.     

Fire history of Araucaria–Nothofagus forests in Villarrica National Park, Chile

Aim In this study we examine fire history (i.e. c. 500 yr bp to present) of Araucaria–Nothofagus forests in the Andes cordillera of Chile. This is the first fire history developed from tree rings for an Araucaria–Nothofagus forest landscape. Location The fire history was determined for the Quillelhue watershed on the north side of Lanin volcano in Villarrica National Park, Chile. The long‐lived Araucaria araucana was commonly associated with Nothofagus pumilio and N. antarctica in more mesic and drier sites respectively. Methods Based on a combination of fire‐scar proxy records and forest stand ages, we reconstructed fire frequency, severity, and the spatial extent of burned areas for an c. 4000 ha study area. We used a composite fire chronology for the purpose of determining centennial‐scale changes in fire regimes and comparing the pre‐settlement (pre‐1883) and post‐settlement fire regimes. In addition, we contrasted Araucaria and Nothofagus species as fire‐scar recorders. Results In the study area, we dated a total of 144 fire‐scarred trees, representing 46 fire years from ad 1446 to the present. For the period from ad 1696 to 2000, using fire dates from Araucaria and Nothofagus species, the composite mean fire interval varied from 7 years for all fires to 62 years for widespread events (i.e. years in which ≥ 25% of recorder trees were scarred). Sensitivity to fire was different for Araucaria and Nothofagus species. More than 98% of the fires recorded by Nothofagus species occurred during the 1900s. The lack of evidence for older fire dates (pre‐1900) in Nothofagus species was due to their shorter longevity and greater susceptibility to being killed by more severe fires. Whereas the thin‐barked N. pumilio and N. antarctica are often destroyed in catastrophic fire events, large and thick‐barked Araucaria trees typically survive. The spatial extent of fires ranged from small patchy events to those that burned more than 40% of the entire landscape (c. > 1500 ha). Main conclusions Fire is the most important disturbance shaping the Araucaria–Nothofagus landscape in the Araucarian region. The forest landscape has been shaped by a mixed‐severity fire regime that includes surface and crown fires. High‐severity widespread events were relatively infrequent (e.g. 1827, 1909 and 1944) and primarily affected tall Araucaria–N. pumilio forests and woodlands dominated by Araucaria–N. antarctica. Although there is abundant evidence of the impact of Euro‐Chilean settlers on the area, the relative influence of this settlement on the temporal pattern of fire could only be tentatively established due to the relatively small number of pre‐1900 fire dates. An apparent increase in fire occurrence is evident in the fire record during Euro‐Chilean settlement (post‐1880s) compared with the Native American era, but it may also be the result of the destruction of evidence of older fires by more recent stand‐devastating fires (e.g. 1909 and 1944). Overall, the severe and widespread fires that burned in Araucaria–Nothofagus forests of this region in 2002, previously interpreted as an ecological novelty, are within the range of the historic fire regimes that have shaped this forested landscape.     

The trade-off between cold resistance and growth determines the <Emphasis Type="Italic">Nothofagus pumilio</Emphasis> treeline

The upper and poleward limit of tree distribution are usually determined by abiotic factors such as low temperature and strong winds. Thus, cold resistance is a key element for survival in high altitudes and latitudes where conditions can reduce plant growth. A trade-off between resource allocation to cold resistance and growth could emerge in populations frequently exposed to low temperatures like those in the treeline zone. We studied annual height growth and ice nucleation temperature in Nothofagus pumilio (Nothofagaceae) populations growing in its extremes of altitudinal distribution and in 3 sites situated on a latitudinal gradient in the Chilean Andes. Additionally, gas exchange, water and nitrogen use efficiency and total soluble sugar (TSS) were also measured as possible mechanisms for survival in high altitudes. Individuals from the treeline populations showed lower annual height growth and lower ice nucleation temperatures compared with those from lower populations. In the same way, individuals from more poleward populations showed lower annual height growth and lower ice nucleation temperatures. Gas exchange, water and nitrogen use efficiency and TSS were also higher in the high altitude populations. The results obtained support the hypothesis of trade-off, because the upper and poleward populations showed more cold resistance but a lower height growth. Additionally, we show that cold resistance mechanisms do not impact the physiological performance, suggesting possible adaptation of the high altitude populations. Low temperatures may be affecting cellular growth instead of photosynthesis, creating a pool of carbohydrates that could participate in cold tolerance. Other abiotic and biotic factors should be also assessed to fully understand the distributional range of Nothofagus species.     

Dendroecological analysis of defoliator outbreaks on Nothofagus pumilio and their relation to climate variability in the Patagonian Andes

In the temperate forests of the southern Andes, Nothofagus pumilio, the dominant species of the most extensive forest type, experiences severe defoliation caused by caterpillars of the Ormiscodes genus (Lepidoptera: Saturniidae). This study uses tree rings to reconstruct the history of Ormiscodes outbreaks for the 1850–2005 period and examines relationships between outbreaks and climate variability. We used local climate records to compare outbreak–climate relationships in the northern Patagonian Andes (c. 41°S) and the cooler southern Patagonian Andes (c. 49°S). We also examined relationships between outbreak events and regional climate variability driven by variability in the Southern Annular Mode (SAM) and the El Niño‐Southern Oscillation. Although relationships between Ormiscodes outbreaks and climate proved to be complex, in northern Patagonia defoliation events are associated with drier and warmer than average growing seasons. Warming and drying trends in Patagonia during the latter part of the 20th century have been linked to a positive trend in SAM. During the post‐1976 period of accelerated warming in Patagonia, widespread defoliation outbreaks have occurred in both northern and southern Patagonia but the increase in frequency of events has been greater in the south. In southern Patagonia the increases in frequency of outbreaks in the late 20th century appear to be unprecedented over the c. 150 year tree‐ring record of reconstructed outbreaks. These results are consistent with the greater magnitude of recent warming in southern Patagonia, and suggest that under predicted warmer and drier climates in the 21st century, defoliator outbreaks may continue to increase in frequency. This study is the first systematic reconstruction of past insect outbreaks in South America and provides a preliminary understanding of how climate variability affects defoliator outbreaks in Patagonian Nothofagus forests.     

Replacement patterns and species coexistence in an Andean Araucaria‐Nothofagus forest

Questions: Fire appears to affect both replacement patterns and coexistence of Araucaria araucana‐Nothofagus pumilio forests in the Andean Araucarian region of south‐central Chile. A quantitative assessment of coexistence in the absence of recent fires, however, is lacking. In this study, we considered the life‐history attributes, time of recruitment and spatial pattern of individuals of both tree species to address the following questions. How regular has recruitment of both species been in time? Is there any temporal niche differentiation? Are the two species positively or negatively associated in space and, if so, at what scale and for what age and size classes? Is there any spatial niche differentiation? Location: Andean Araucarian region of Chile, Villarrica National Park (39°35′S, 71°31′W; 1300 m a.s.l.). Methods: We stem‐mapped and cored a total of 1073 trees in a 1‐ha plot in a late‐successional post‐fire stand to examine spatiotemporal patterns of establishment. We used semivariogram modelling and the pair‐correlation function to distinguish between regeneration modes and describe species interactions. Results: The two species differ in their regeneration mode: whereas A. araucana appeared to recruit more continuously in time and space, episodic pulses of establishment were dominant for N. pumilio. At small scales, younger age‐class stems of A. araucana were randomly distributed, while older age‐class stems were aggregated. This was in contrast to common patterns for temperate tree species, including N. pumilio, following processes of self‐thinning. Younger age classes of A. araucana were distributed independently of older trees of both species, but younger age classes of N. pumilio had a negative association with older conspecifics at scales similar to crown diameter. Conclusions: In the absence of recent fires, it is likely that A. araucana would dominate the stand alone, given its greater shade tolerance, greater longevity and continuous recruitment. However, while canopy closure is still incomplete, the shade‐intolerant N. pumilio will be able to recruit in those open areas after seed masting and will coexist with A. araucana.     

Environmental and genetic control of insect abundance and herbivory along a forest elevational gradient

Environmental conditions and plant genotype may influence insect herbivory along elevational gradients. Plant damage would decrease with elevation as temperature declines to suboptimal levels for insects. However, host plants at higher elevations may exhibit traits that either reduce or enhance leaf quality to insects, with uncertain net effects on herbivory. We examined folivory, insect abundance and leaf traits along six replicated elevational ranges in Nothofagus pumilio forests of the northern Patagonian Andes, Argentina. We also conducted a reciprocal transplant experiment between low- and high-elevation sites to test the extent of environmental and plant genetic control on insect abundance and folivory. We found that insect abundance, leaf size and specific leaf area decreased, whereas foliar phosphorous content increased, from low-, through mid- to high-elevation sites. Path analysis indicated that changes in both insect abundance and leaf traits were important in reducing folivory with increasing elevation and decreasing mean temperature. At both planting sites, plants from a low-elevation origin experienced higher damage and supported greater insect loads than plants from a high-elevation origin. The differences in leaf damage between sites were twofold larger than those between plant origins, suggesting that local environment was more important than host genotype in explaining folivory patterns. Different folivore guilds exhibited qualitatively similar responses to elevation. Our results suggest an increase in insect folivory on high-elevation N. pumilio forests under future climate warming scenarios. However, in the short-term, folivory increases might be smaller than expected from insect abundance only because at high elevations herbivores would encounter more resistant tree genotypes.     

Long‐term growth‐increment chronologies reveal diverse influences of climate forcing on freshwater and forest biota in the Pacific Northwest

Analyses of how organisms are likely to respond to a changing climate have focused largely on the direct effects of warming temperatures, though changes in other variables may also be important, particularly the amount and timing of precipitation. Here, we develop a network of eight growth‐increment width chronologies for freshwater mussel species in the Pacific Northwest, United States and integrate them with tree‐ring data to evaluate how terrestrial and aquatic indicators respond to hydroclimatic variability, including river discharge and precipitation. Annual discharge averaged across water years (October 1–September 30) was highly synchronous among river systems and imparted a coherent pattern among mussel chronologies. The leading principal component of the five longest mussel chronologies (1982–2003; PC1_mussel) accounted for 47% of the dataset variability and negatively correlated with the leading principal component of river discharge (PC1_discharge; r = ?0.88; P < 0.0001). PC1_mussel and PC1_discharge were closely linked to regional wintertime precipitation patterns across the Pacific Northwest, the season in which the vast majority of annual precipitation arrives. Mussel growth was also indirectly related to tree radial growth, though the nature of the relationships varied across the landscape. Negative correlations occurred in forests where tree growth tends to be limited by drought while positive correlations occurred in forests where tree growth tends to be limited by deep or lingering snowpack. Overall, this diverse assemblage of chronologies illustrates the importance of winter precipitation to terrestrial and freshwater ecosystems and suggests that a complexity of climate responses must be considered when estimating the biological impacts of climate variability and change.     

Litterfall of epiphytic macrolichens in Nothofagus forests of northern Patagonia,Argentina: Relation to stand age and precipitation

Abstract: The objective of this study was to analyse how stand age and precipitation influence abundance and diversity of epiphytic macrolichens in southern beech Nothofagus forests, estimated by lichen litter sampling. Five sites of Nothofagus dombeyi (Mirbel) Oersted were selected in Nahuel Huapi National Park, Argentina. At each site, lichen fragments from the forest floor were collected at 12.5 m² plots in pairs of young and mature N. dombeyi forest. Additionally, two sites with multi‐aged subalpine Nothofagus pumilio (Poepp. et Endl.) Krasser forest were investigated in a similar manner. Average litterfall biomass per stand varied from less than 1 kg ha^?1 in a young low‐precipitation stand to a maximum of 20 kg ha^?1 in a mature high‐precipitation stand. In places with higher precipitation, litterfall biomass in N. dombeyi forest was considerably higher in old stands as compared with young ones. In places with less than 2000 mm of precipitation, differences in biomass were less pronounced. Old humid stands contained about twice as many taxa in the litter as old low‐precipitation stands and young stands in general. Mature stands in low‐precipitation sites only contained 17% of the litter biomass as compared with mature stands in high‐precipitation sites. Epiphytic lichen composition changed from predominating fruticose lichens (Usnea spp. and Protousnea spp.) in low‐precipitation stands to Pseudocyphellaria spp., Nephroma spp. and other foliose lichens, in the high‐precipitation stands. There were no clear differences in the proportion of fruticose and foliose lichens between young and old stands. Fruticose lichens dominated litter biomass in both N. pumilio sites.     

Global climate change will increase the abundance of symbiotic nitrogen‐fixing trees in much of North America

Symbiotic nitrogen (N)‐fixing trees can drive N and carbon cycling and thus are critical components of future climate projections. Despite detailed understanding of how climate influences N‐fixation enzyme activity and physiology, comparatively little is known about how climate influences N‐fixing tree abundance. Here, we used forest inventory data from the USA and Mexico (>125,000 plots) along with climate data to address two questions: (1) How does the abundance distribution of N‐fixing trees (rhizobial, actinorhizal, and both types together) vary with mean annual temperature (MAT) and precipitation (MAP)? (2) How will changing climate shift the abundance distribution of N‐fixing trees? We found that rhizobial N‐fixing trees were nearly absent below 15°C MAT, but above 15°C MAT, they increased in abundance as temperature rose. We found no evidence for a hump‐shaped response to temperature throughout the range of our data. Rhizobial trees were more abundant in dry than in wet ecosystems. By contrast, actinorhizal trees peaked in abundance at 5–10°C MAT and were least abundant in areas with intermediate precipitation. Next, we used a climate‐envelope approach to project how N‐fixing tree relative abundance might change in the future. The climate‐envelope projection showed that rhizobial N‐fixing trees will likely become more abundant in many areas by 2080, particularly in the southern USA and western Mexico, due primarily to rising temperatures. Projections for actinorhizal N‐fixing trees were more nuanced due to their nonmonotonic dependence on temperature and precipitation. Overall, the dominant trend is that warming will increase N‐fixing tree abundance in much of the USA and Mexico, with large increases up to 40° North latitude. The quantitative link we provide between climate and N‐fixing tree abundance can help improve the representation of symbiotic N fixation in Earth System Models.