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.     

No evidence of carbon limitation with tree age and height in Nothofagus pumilio under Mediterranean and temperate climate conditions

Background and Aims

Trees universally decrease their growth with age. Most explanations for this trend so far support the hypothesis that carbon (C) gain becomes limited with age; though very few studies have directly assessed the relative reductions of C gain and C demand with tree age. It has also been suggested that drought enhances the effect of C gain limitation in trees. Here tests were carried out to determine whether C gain limitation is causing the growth decay with tree age, and whether drought accentuates its effect.

Methods

The balance between C gain and C demand across tree age and height ranges was estimated. For this, the concentration of non-structural carbohydrates (NSCs) in stems and roots of trees of different ages and heights was measured in the deciduous temperate species Nothofagus pumilio. An ontogenetic decrease in NSCs indicates support for C limitation. Furthermore, the importance of drought in altering the C balance with ontogeny was assessed by sampling the same species in Mediterranean and humid climate locations in the southern Andes of Chile. Wood density (WD) and stable carbon isotope ratios (δ¹³C) were also determined to examine drought constraints on C gain.

Key Results

At both locations, it was effectively found that tree growth ultimately decreased with tree age and height. It was found, however, that NSC concentrations did not decrease with tree age or height when WD was considered, suggesting that C limitation is not the ultimate mechanism causing the age/height-related declining tree growth. δ¹³C decreased with tree age/height at the Mediterranean site only; drought effect increased with tree age/height, but this pattern was not mirrored by the levels of NSCs.

Conclusions

The results indicate that concentrations of C storage in N. pumilio trees do not decrease with tree age or height, and that reduced C assimilation due to summer drought does not alter this pattern.     

Isozyme polymorphisms provide evidence of clinal variation with elevation in Nothofagus pumilio

Variable physical conditions along elevational gradients strongly influence patterns of genetic differentiation in tree species. Here, the hypothesis is tested that different growth forms of Nothofagus pumilio, which characterizes the subalpine forests in the southern Andes, will display continuous genetic variation with elevation. At each of four elevational strips in three different mountain ranges, fresh leaf tissue was sampled from 30 randomly selected individuals to be analyzed by protein electrophoresis. Allelic frequencies were used to test for heterogeneity across populations and to classify populations into different elevational strips by discriminant analysis. The degree of population divergence was estimated by F(ST). Clinical variation on within-population genetic characteristics was analyzed by linear regressions against elevation. Seven enzyme systems coded for 14 putative isozyme loci, 57% of which were polymorphic in at least one population. Allele frequencies significantly varied with elevation and discriminant analysis separated populations at different elevational strips. Among-population divergence within any mountain range was small, but greater than among different mountain ranges. Overall, low-elevation populations were more variable than high-elevation populations, and regression analyses suggested continuous variation in populations of N. pumilio 100 m apart. Marked stepwise phenological differences on mountain slopes are most probably responsible for the isolation of nearby populations.     

Simulated warming does not impair seedling survival and growth of Nothofagus pumilio in the southern Andes

It has been predicted that subalpine forests will be negatively affected by global warming; however, direct responses to experimental warming have been scarcely examined in these systems. In this study we evaluated the effects of higher temperatures with and without water addition on the survival and growth of recently emerged (small) and large seedlings of the widely distributed species Nothofagus pumilio in subalpine forests of the southern Chilean Andes. We also examined the variations in seedling traits related to carbon balance in order to infer the causal mechanisms of survival and growth responses. Treatments of open top chambers (OTCs) were combined with watering in two locations with differing climates: Antillanca (40°S, humid) and Cerro Castillo (46°S, drier). OTCs increased mean and maximum air temperatures by 0.6 °C and 2–3 °C, respectively, and decreased soil humidity by 56% in Antillanca and 30% in Cerro Castillo, fulfilling methodological expectations and climate model predictions. After two complete growing seasons, the survival, relative growth rate (RGR), biomass, and a suite of seedling traits were measured and analyzed using mixed-effects models. Warming and warming in combination with watering significantly increased large seedling survival in Cerro Castillo. In Antillanca, warmer conditions increased the height, biomass, and leaf area of small seedlings, and the RGR of large seedlings. In this location, warming also caused lower leaf carbon isotopic composition in both age classes and higher specific leaf area in small seedlings, suggesting whole-plant carbon gain improvements; warming did not produce any drought effects. Our results indicate that warming produces positive effects on the seedling establishment of N. pumilio in the southern Andes, highlighting the importance of site-specific effects in response to climate change in widespread species. Site-specific effects can most likely explain the discrepancies between the results of this study and the predictions outlined by previous studies for these forests.     

Removal of a non-climatically induced seven-year cycle from Nothofagus pumilio tree-ring width chronologies from Tierra del Fuego,Argentina for their use in climate reconstructions

A pronounced seven-year cycle is one of the most prominent features of many Fuegian and southern Patagonian Nothofagus pumilio tree-ring width chronologies. However, the source of this variability is still unclear. Here we provide multiple lines of evidence that suggest this phenomenon comes from non-climatically induced ecological factors, most probably triggered by insect outbreaks. To test this hypothesis we removed the seven-year cycle, by means of Singular Spectrum Analysis, from a network of 44 tree-ring width chronologies, from the eastern Tierra del Fuego Island in Argentina. The chronologies that were filtered this way showed a stronger climatic response, and were more successfully modeled using the Vaganov-Shashkin-Lite (VS-Lite) process-based forward model of tree growth. We conclude that the removed periodicity does not have a climatic source. Additional evidence from direct observation of affected and non-affected trees during defoliation, due to Ormiscodes sp. outbreak in 2010, is provided. We also tested the hypothesis of probable masting effects on the growth of N. pumilio at a site with a 12-yr record of seed production, but this hypothesis was not confirmed. Taking into account the non-climatic nature of this signal, we explore the utility of corrected chronologies for reconstruction of climatic parameters in Tierra del Fuego, especially for summer temperature and frost day frequency. This study is the first systematic analysis of the pronounced high-frequency cyclicity in N. pumilio tree growth in southernmost South America.     

Differential hypogeous sporocarp production from Nothofagus dombeyi and N. pumilio forests in southern Argentina

Mycorrhizal fungi that form hypogeous sporocarps are an important component of the temperate forest soil community. In many regions, such as the Nothofagus forest in the Patagonian Andes, this group of fungi has been poorly studied. Here we examined the spring and autumn community composition of "sequestrate fungi", based on sporocarp production in pure forests of Nothofagus dombeyi (evergreen) and N. pumilio (deciduous). We investigated the possible relationships between these communities and environmental factors over 2 y. The rarefaction curves and the minimal richness estimates converged at nearly the same level for each forest type, and the asymptotes suggested that the sampling effort was sufficient to capture most of the hypogeous sporocarp richness in these forest stands. In total 27 species were recovered. Basidiomycota, Ascomycota and Glomeromycota respectively accounted for nine, two and one genera. Species richness of hypogeous sporocarps varied in relation to forest type but not to season (fall and spring), whereas sporocarp biomass varied according to an interaction between season and forest type. Species richness and sporocarp biomass were positively correlated with rainfall and negatively correlated with altitude. In addition sporocarp species richness was positively related to number of trees per transect. We found that two different forest stands, each dominated by different species of Nothofagus, exhibited different hypogeous sporocarp communities.     

Effects of climate on the radial growth of mixed stands of Nothofagus nervosa and Nothofagus obliqua along a precipitation gradient in Patagonia,Argentina

Climate models for North Patagonia in Argentina project dryer conditions, due to a decrease in mean precipitation combined with an increase in mean temperatures. The temperate mixed Nothofagus forest, which exists along a steep precipitation gradient, could be directly impacted. For this study, we evaluated the influence of mean climate on the growth of the significant deciduous species: N. nervosa and N. obliqua. For the first time in Argentina, dendroclimatological analyses were done on both species using a network of 14 chronologies, covering their longitudinal distribution along a gradient of declining precipitation from west to east. Seasonal correlation analysis revealed that temperature has a negative effect on the growth of both species across all sites, particularly during summer of the previous and current growth season. Precipitation has a positive effect on the growth of trees for both species, which is more significant for N. nervosa. The relationship between early-summer climate and growth remained relatively stable over time for N. nervosa; however, for N. obliqua the detrimental effects of temperature increased towards the end of the 20th century, and the positive effects of precipitation decreased, particularly at the driest end of the gradient. These results suggest that a continued decrease in rainfall with a rise in temperature could impact growth for both of these species.     

Multi-stemmed trees of Nothofagus pumilio second-growth forest in Patagonia are formed by highly related individuals

Background and Aims Multi-stemmed trees (tree clusters) in Nothofagus pumilio, a dominant tree species in Patagonia, are very uncommon and are restricted to the edge of second-growth forests following human-provoked fires. No vegetative reproduction has been reported so far. The genetic structure of multi-stemmed trees of this species was investigated and it was hypothesized that genets within a cluster were more closely related than average in the population. Methods Fifteen clusters (composed of at least three purported stems) and 15 single trees were sampled at the edge of a second-growth forest and genotyped using two amplified fragment length polymorphism (AFLP) primer pairs. We obtained 119 polymorphic markers that allowed clonality to be determined, together with sibship structure and relatedness among samples. Key Results Clonality was detected in seven clusters but all clusters had at least two different genotypes. Full sibs were found exclusively within clusters and in all clusters. Within a cluster, stems that were not identified as full sibs were often half sibs. Relatedness values for the full sibs and half sibs were higher than the theoretical values of 0·5 and 0·25 but the relatedness between clusters was very low. Conclusions Tree clusters that are merged at the edge of the second-growth forest of N. pumilio are composed of stems of the same genotype and of other genotypes that are highly related (but not always). It is suggested that this peculiar genetic structure results from a combination of several causes, including selection for merging of related individuals.     

Carnivoran postcranial adaptations and their relationships to climate

The relationship between climate and morphology is important to understand in view of the rapid rate of climate change occurring today; however, this relationship has not been fully explored in many mammalian groups. We use postcranial indices to explore the association between climate and morphology in the mammalian order Carnivora. Carnivora is a good group to use for this analysis because it includes species with a variety of locomotor ecologies that live in almost every type of habitat on Earth. We measured postcrania of 121 carnivoran species from around the world, combined with habitat and climate data from the BIOCLIM database and the NCEAS Paleocommunities Working Group to examine correlations between postcrania, temperature and precipitation. We analyzed these data using correspondence analysis and multiple linear regressions. We found three postcranial indices that were significantly correlated with climate. Brachial index (radius/humerus) and shoulder moment (length deltopectoral crest/length humerus) were both significantly associated with temperature and precipitation, while greater trochanter height (as proportion of femur length) was associated with precipitation. We found that these indices were indirectly related to climate via the strong association between climate and locomotor ecologies. This relationship between climate and postcranial proportions can be used in future studies: for paleo‐climate reconstruction in carnivore localities and for trait‐based identification of species vulnerability as climate change continues.     

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.     

Yeast communities associated with the bulk-soil, rhizosphere and ectomycorrhizosphere of a Nothofagus pumilio forest in northwestern Patagonia, Argentina

Soil microorganisms play an important role in soil quality and they interact closely with vegetation. Little is known about yeast diversity and function in forest soil ecosystems and their interactions with other biotic soil components, particularly in the mycorrhizosphere. We studied the diversity of yeasts inhabiting the bulk-soil, rhizosphere and ectomycorrhizosphere of a Nothofagus pumilio forest in Nahuel Huapi National Park (Bariloche, Argentina). Ectomycorrhizal infection was observed in all N. pumilio trees studied. A total of 126 yeast isolates were obtained, including 18 known and three possibly new species. Basidiomycetous yeasts were predominant in all soil fractions, and the most frequently isolated species was Cryptococcus podzolicus. Diversity indices and multivariate analyses were used to study and compare yeast communities in the bulk-soil, rhizosphere and ectomycorrhizosphere. Yeasts able to ferment glucose were found associated with the rhizosphere. Many of the recovered yeast species were associated with lignocelluloses compound degradation, which suggest that yeast plays an important role as a decomposer in these forest soils. Each soil fraction has a distinct yeast assemblage related to their physiologic capacities and soil nutrient availability.     

Intraspecific trait variation and covariation in a widespread tree species (Nothofagus pumilio) in southern Chile

? The focus of the trait-based approach to study community ecology has mostly been on trait comparisons at the interspecific level. Here we quantified intraspecific variation and covariation of leaf mass per area (LMA) and wood density (WD) in monospecific forests of the widespread tree species Nothofagus pumilio to determine its magnitude and whether it is related to environmental conditions and ontogeny. We also discuss probable mechanisms controlling the trait variation found. ? We collected leaf and stem woody tissues from 30-50 trees of different ages (ontogeny) from each of four populations at differing elevations (i.e. temperatures) and placed at each of three locations differing in soil moisture. ? The total variation in LMA (coefficient of variation (CV) = 21.14%) was twice that of WD (CV = 10.52%). The total variation in traits was never less than 23% when compared with interspecific studies. Differences in elevation (temperature) for the most part explained variation in LMA, while differences in soil moisture and ontogeny explained the variation in WD. Traits covaried similarly in the altitudinal gradient only. ? Functional traits of N. pumilio exhibited nonnegligible variation; LMA varied for the most part with temperature, while WD mostly varied with moisture and ontogeny. We demonstrate that environmental variation can cause important trait variation without species turnover.     

Climate–growth relationships of tropical tree species in West Africa and their potential for climate reconstruction

Most tropical regions are facing historical difficulties of generating biologically reconstructed long‐term climate records. Dendrochronology (tree‐ring studies) is a powerful tool to develop high‐resolution and exactly dated proxies for climate reconstruction. Owing to the seasonal variation in rainfall we expected the formation of annual tree rings in the wood of tropical West African tree species. In the central‐western part of Benin (upper Ouémé catchment, UOC) and in northeastern Ivory Coast (Comoé National Park, CNP) we investigated the relationship between climate (precipitation, sea surface temperature (SST)) and tree rings and show their potential for climate reconstruction. Wood samples of almost 200 trees belonging to six species in the UOC and CNP served to develop climate‐sensitive ring‐width chronologies using standard dendrochronological techniques. The relationship between local precipitation, monthly SST anomalies in the Gulf of Guinea, El Niño‐ Southern Oscillation (ENSO) and ring‐width indices was performed by simple regression analyses, two sample tests and cross‐spectral analysis. A low‐pass filter was used to highlight the decadal variability in rainfall of the UOC site. All tree species showed significant relationships with annual precipitation proving the existence of annual tree rings. ENSO signals could not be detected in the ring‐width patterns. For legume tree species at the UOC site significant relationships could be found between SST anomalies in the Gulf of Guinea indicating correlations at periods of 5.1–4.1 and 2.3 years. Our findings accurately show the relationship between tree growth, local precipitation and SST anomalies in the Gulf of Guinea possibly associated with worldwide SST patterns. A master chronology enabled the reconstruction of the annual precipitation in the UOC to the year 1840. Time series analysis suggest increasing arid conditions during the last 160 years which may have large impacts on the hydrological cycles and consequently on the ecosystem dynamics and the development of socio‐economic cultures and sectors in the Guinea‐Congolian/Sudanian region.     

Global assessment of relationships between climate and tree growth

Tree‐ring records provide global high‐resolution information on tree‐species responses to global change, forest carbon and water dynamics, and past climate variability and extremes. The underlying assumption is a stationary (time‐stable), quasi‐linear relationship between tree growth and environment, which however conflicts with basic ecological and evolutionary theory. Indeed, our global assessment of the relevant tree‐ring literature demonstrates non‐stationarity in the majority of tested cases, not limited to specific proxies, environmental parameters, regions or species. Non‐stationarity likely represents the general nature of the relationship between tree‐growth proxies and environment. Studies assuming stationarity however score two times more citations influencing other fields of science and the science–policy interface. To reconcile ecological reality with the application of tree‐ring proxies for climate or environmental estimates, we provide a clarification of the stationarity concept, propose a simple confidence framework for the re‐evaluation of existing studies and recommend the use of a new statistical tool to detect non‐stationarity in tree‐ring proxies. Our contribution is meant to stimulate and facilitate discussion in light of our results to help increase confidence in tree‐ring‐based climate and environmental estimates for science, the public and policymakers.     

Changing relationships between tree growth and climate in Northwest China

Recently, several studies have shown changing relationships between tree growth and climate factors, mostly in the circumpolar north. There, changing relationships with climate seem to be linked to emergent subpopulation behavior. Here, we test for these phenomena in Northwest China using three tree species (Pinus tabulaeformis, Picea crassifolia, and Sabina przewalskii) that had been collected from six sites at Qilian Mts. and Helan Mts. in Northwest China. We first checked for growth divergence of individual sites and then investigated the relationship between tree growth and climate factors using moving correlation functions (CF). Two species, Pinus and Sabina, from two sites clearly showed growth divergence, not only in the late twentieth century as reported in other studies, but also over nearly the whole record. In divergent sites, one chronology shows more stable relationships with climate factors (usually precipitation). In non-divergent sites, nearly all relationships either vary in strength or become non-significant at one point. While this might possibly be related to increased stress on some trees due to increasing temperature, the exact causes for this shift in sensitivity remain unclear. We would like to highlight the necessity for additional studies investigating possible non-stationary growth responses of trees with climate, especially at sites that are used for climate reconstruction as our sites in Northwest China.     

Structural and functional changes in Nothofagus pumilio forests along an altitudinal gradient in Tierra del Fuego,Argentina

Abstract. Structural (density, height, basal area, above‐ground tree biomass, leaf area index) and functional (leaf phenology, growth rate, fine litter fall, leaf decomposition) traits were quantified in four mature forests of Nothofagus pumilio (lenga) along an altitudinal sequence in Tierra del Fuego, Argentina. Three erect forest stands at 220, 440 and 540m and a krummholz stand at 640 m a.s.l. were selected. Along the altitudinal sequence, stem density increased while DBH, height, biomass, leaf‐size and growth period, mean growth rate and decay rate decreased. Dead stems increased and basal area and fine‐litter fall decreased with an increase in elevation among erect forests, but these trends inverted at krummholz. We suggest that krummholz is not only a morphological response to the adverse climate but is also a life form with functional advantages.     

Interannual variability in carbon dioxide fluxes and flux–climate relationships on grazed and ungrazed northern mixed-grass prairie

The annual carbon (C) budget of grasslands is highly dynamic, dependent on grazing history and on effects of interannual variability (IAV) in climate on carbon dioxide (CO₂) fluxes. Variability in climatic drivers may directly affect fluxes, but also may indirectly affect fluxes by altering the response of the biota to the environment, an effect termed ‘functional change’. We measured net ecosystem exchange of CO₂ (NEE) and its diurnal components, daytime ecosystem CO₂ exchange (P_D) and night‐time respiration (R_E), on grazed and ungrazed mixed‐grass prairie in North Dakota, USA, for five growing seasons. Our primary objective was to determine how climatic anomalies influence variability in CO₂ exchange. We used regression analysis to distinguish direct effects of IAV in climate on fluxes from functional change. Functional change was quantified as the improvement in regression on fitting a model in which slopes of flux–climate relationships vary among years rather than remain invariant. Functional change and direct effects of climatic variation together explained about 20% of variance in weekly means of NEE, P_D, and R_E. Functional change accounted for more than twice the variance in fluxes of direct effects of climatic variability. Grazing did not consistently influence the contribution of functional change to flux variability, but altered which environmental variable best explained year‐to‐year differences in flux–climate slopes, reduced IAV in seasonal means of fluxes, lessened the strength of flux–climate correlations, and increased NEE by reducing R_E relatively more than P_D. Most of these trends are consistent with the interpretation that grazing reduced the influence of plants on ecosystem fluxes. Because relationships between weekly values of fluxes and climatic regulators changed annually, year‐to‐year differences in the C balance of these ecosystems cannot be predicted from knowledge of IAV in climate alone.