Masting is uncommon in trees that depend on mutualist dispersers in the context of global climate and fertility gradients.

The benefits of masting (volatile, quasi-synchronous seed production at lagged intervals) include satiation of seed predators, but these benefits come with a cost to mutualist pollen and seed dispersers. If the evolution of masting represents a balance between these benefits and costs, we expect mast avoidance in species that are heavily reliant on mutualist dispersers. These effects play out in the context of variable climate and site fertility among species that vary widely in nutrient demand. Meta-analyses of published data have focused on variation at the population scale, thus omitting periodicity within trees and synchronicity between trees. From raw data on 12 million tree-years worldwide, we quantified three components of masting that have not previously been analysed together: (i) volatility, defined as the frequency-weighted year-to-year variation; (ii) periodicity, representing the lag between high-seed years; and (iii) synchronicity, indicating the tree-to-tree correlation. Results show that mast avoidance (low volatility and low synchronicity) by species dependent on mutualist dispersers explains more variation than any other effect. Nutrient-demanding species have low volatility, and species that are most common on nutrient-rich and warm/wet sites exhibit short periods. The prevalence of masting in cold/dry sites coincides with climatic conditions where dependence on vertebrate dispersers is less common than in the wet tropics. Mutualist dispersers neutralize the benefits of masting for predator satiation, further balancing the effects of climate, site fertility and nutrient demands.

Projections of fire probability and ecosystem vulnerability under 21st century climate across a trans-Andean productivity gradient in Patagonia.

Warming trends are altering fire regimes globally, potentially impacting on the long-term persistence of some ecosystems. However, we still lack clear understanding of how climatic stressors will alter fire regimes along productivity gradients. We trained a Random Forests model of fire probabilities across a 5°lat × 2° long trans-Andean rainfall gradient in northern Patagonia using a 23-year long fire record and biophysical, vegetation, human activity and seasonal fire weather predictors. The final model was projected onto mid- and late 21st century fire weather conditions predicted by an ensemble of GCMs using 4 emission scenarios. We finally assessed the vulnerability of different forest ecosystems by matching predicted fire return intervals with critical forest persistence fire return thresholds developed with landscape simulations. Modern fire activity showed the typical hump-shaped relationship with productivity and a negative distance relationship with human settlements. However, fire probabilities were far more sensitive to current season fire weather than to any other predictor. Sharp responsiveness of fire to the accelerating drier/warmer fire seasons predicted for the remainder of the 21st century in the region led to 2 to 3-fold (RCPs 4.5 and 8.5) and 3 to 8-fold increases in fire probabilities for the mid- and late 21st century, respectively. Contrary to current generalizations of larger impacts of warming on fire activity in fuel-rich ecosystems, our modeling results showed first an increase in predicted fire activity in less productive ecosystems (shrublands and steppes) and a later evenly amplified fire activity-productivity relationship with it shape resembling (at higher fire probabilities) the modern hump-shaped relationship. Despite this apparent homogeneous effect of warming on fire activity, vulnerability to predicted late 21st century shorter fire intervals were higher in most productive ecosystems (subalpine deciduous and evergreen Nothofagus-dominated rainforests) due to a general lack of fire-adapted traits in the dominant trees that compose these forests.

Limits to reproduction and seed size-number trade-offs that shape forest dominance and future recovery.

The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential.

Globally, tree fecundity exceeds productivity gradients.

Lack of tree fecundity data across climatic gradients precludes the analysis of how seed supply contributes to global variation in forest regeneration and biotic interactions responsible for biodiversity. A global synthesis of raw seedproduction data shows a 250-fold increase in seed abundance from cold-dry to warm-wet climates, driven primarily by a 100-fold increase in seed production for a given tree size. The modest (threefold) increase in forest productivity across the same climate gradient cannot explain the magnitudes of these trends. The increase in seeds per tree can arise from adaptive evolution driven by intense species interactions or from the direct effects of a warm, moist climate on tree fecundity. Either way, the massive differences in seed supply ramify through food webs potentially explaining a disproportionate role for species interactions in the wet tropics.

Multi-centennial phase-locking between reproduction of a South American conifer and large-scale drivers of climate.

Climate forcings determine the episodic occurrence of local climate anomalies that trigger the occurrence of masting events (massive, synchronized and intermittent seed production by perennial plants). This suggests some kind of phase-locking of the reproductive cycles of individual plants to the climatological cycle, thus further reinforcing reproductive synchrony and the Moran effect. We propose a dendrochronological approach to filter out the long-term direct effects of climate on tree radial growth and temporal reproductive effort by sex by using actual trees as climatic controls to reconstruct masting events in Araucaria araucana, a long-lived dioecious masting conifer. In this way, we developed a multi-century-long tree masting reconstruction for South America using female-male radial growth determined by differences in timing and magnitude of the reproductive effort between sexes. We provide evidence for a regional synchronizing mechanism of masting which is drought induced by strong cold La Niña phases of El Niño/Southern Oscillation (ENSO) amplified by the positive phases of the Southern Annular Mode (SAM) that activate both female and male cone bud formation during year -2 before seed fall; that is, a long-term phase-locking between the ENSO cycle and the reproductive cycle modulated by the strength of SAM. In addition, our regional index of masting frequency showed its maximum during the late twentieth century relative to the previous centuries, suggesting that the species is currently at its maximum masting frequency concurrent with a period of enhanced temperature and drought conditions in Patagonia, probably driven by the positive phase of the SAM.

Forest and woodland replacement patterns following drought-related mortality.

Forest vulnerability to drought is expected to increase under anthropogenic climate change, and drought-induced mortality and community dynamics following drought have major ecological and societal impacts. Here, we show that tree mortality concomitant with drought has led to short-term (mean 5 y, range 1 to 23 y after mortality) vegetation-type conversion in multiple biomes across the world (131 sites). Self-replacement of the dominant tree species was only prevalent in 21% of the examined cases and forests and woodlands shifted to nonwoody vegetation in 10% of them. The ultimate temporal persistence of such changes remains unknown but, given the key role of biological legacies in long-term ecological succession, this emerging picture of postdrought ecological trajectories highlights the potential for major ecosystem reorganization in the coming decades. Community changes were less pronounced under wetter postmortality conditions. Replacement was also influenced by management intensity, and postdrought shrub dominance was higher when pathogens acted as codrivers of tree mortality. Early change in community composition indicates that forests dominated by mesic species generally shifted toward more xeric communities, with replacing tree and shrub species exhibiting drier bioclimatic optima and distribution ranges. However, shifts toward more mesic communities also occurred and multiple pathways of forest replacement were observed for some species. Drought characteristics, species-specific environmental preferences, plant traits, and ecosystem legacies govern postdrought species turnover and subsequent ecological trajectories, with potential far-reaching implications for forest biodiversity and ecosystem services.

Low growth resilience to drought is related to future mortality risk in trees.

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality.

Niche squeeze induced by climate change of the cold-tolerant subtropical montane Podocarpus parlatorei.

Under changing climates, the persistence of montane subtropical taxa may be threatened as suitable habitats decrease with elevation. We developed future environmental niche models (ENNMs) for Podocarpus parlatorei, the only conifer from southern Yungas in South America, and projected it onto two greenhouse gas concentration scenarios based on 13 global climate models for the years 2050 and 2070. Modelling identified that P. parlatorei is sensitive and restricted to a relatively narrow range of both warm season temperature and precipitation. By the mid-late twenty-first century areas of high suitability for P. parlatorei will not migrate but overall suitability will become substantially reduced across its whole range and surrounding areas. Despite extensive areas in high mountain ranges where the species may encounter thermally optimal conditions to potentially allow upward local migration, these same areas will likely become strongly aridified under future conditions. On the other hand, in lowland locations where rainfall levels will not change substantially (e.g. northern range), excessive warming will likely generate abiotic and biotic restrictions (e.g. competition with lowland species) for this cold-tolerant species. Urgent measures should be developed for the local long-term preservation of the gene pool of the unique conifer that characterizes Yungas forests for reasons of biodiversity conservation and ecosystem services.

Patterns and drivers of recent disturbances across the temperate forest biome.

Increasing evidence indicates that forest disturbances are changing in response to global change, yet local variability in disturbance remains high. We quantified this considerable variability and analyzed whether recent disturbance episodes around the globe were consistently driven by climate, and if human influence modulates patterns of forest disturbance. We combined remote sensing data on recent (2001-2014) disturbances with in-depth local information for 50 protected landscapes and their surroundings across the temperate biome. Disturbance patterns are highly variable, and shaped by variation in disturbance agents and traits of prevailing tree species. However, high disturbance activity is consistently linked to warmer and drier than average conditions across the globe. Disturbances in protected areas are smaller and more complex in shape compared to their surroundings affected by human land use. This signal disappears in areas with high recent natural disturbance activity, underlining the potential of climate-mediated disturbance to transform forest landscapes.

Historical and event-based bioclimatic suitability predicts regional forest vulnerability to compound effects of severe drought and bark beetle infestation.

Vulnerability to climate change, and particularly to climate extreme events, is expected to vary across species ranges. Thus, we need tools to standardize the variability in regional climatic legacy and extreme climate across populations and species. Extreme climate events (e.g., droughts) can erode populations close to the limits of species' climatic tolerance. Populations in climatic-core locations may also become vulnerable because they have developed a greater demand for resources (i.e., water) that cannot be enough satisfied during the periods of scarcity. These mechanisms can become exacerbated in tree populations when combined with antagonistic biotic interactions, such as insect infestation. We used climatic suitability indices derived from Species Distribution Models (SDMs) to standardize the climatic conditions experienced across Pinus edulis populations in southwestern North America, during a historical period (1972-2000) and during an extreme event (2001-2007), when the compound effect of hot drought and bark beetle infestation caused widespread die-off and mortality. Pinus edulis climatic suitability diminished dramatically during the die-off period, with remarkable variation between years. P. edulis die-off occurred mainly not just in sites that experienced lower climatic suitability during the drought but also where climatic suitability was higher during the historical period. The combined effect of historically high climatic suitability and a marked decrease in the climatic suitability during the drought best explained the range-wide mortality. Lagged effects of climatic suitability loss in previous years and co-occurrence of Juniperus monosperma also explained P. edulis die-off in particular years. Overall, the study shows that past climatic legacy, likely determining acclimation, together with competitive interactions plays a major role in responses to extreme drought. It also provides a new approach to standardize the magnitude of climatic variability across populations using SDMs, improving our capacity to predict population's or species' vulnerability to climatic change.

Early-Warning Signals of Individual Tree Mortality Based on Annual Radial Growth.

Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in inter-annual growth variability and a decrease in growth synchrony in the last ∼20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not necessarily so for angiosperms.

Direct and indirect climate controls predict heterogeneous early-mid 21st century wildfire burned area across western and boreal North America.

Predicting wildfire under future conditions is complicated by complex interrelated drivers operating across large spatial scales. Annual area burned (AAB) is a useful index of global wildfire activity. Current and antecedent seasonal climatic conditions, and the timing of snowpack melt, have been suggested as important drivers of AAB. As climate warms, seasonal climate and snowpack co-vary in intricate ways, influencing fire at continental and sub-continental scales. We used independent records of seasonal climate and snow cover duration (last date of permanent snowpack, LDPS) and cell-based Structural Equation Models (SEM) to separate direct (climatic) and indirect (snow cover) effects on relative changes in AAB under future climatic scenarios across western and boreal North America. To isolate seasonal climate variables with the greatest effect on AAB, we ran multiple regression models of log-transformed AAB on seasonal climate variables and LDPS. We used the results of multiple regressions to project future AAB using GCM ensemble climate variables and LDPS, and validated model predictions with recent AAB trends. Direct influences of spring and winter temperatures on AAB are larger and more widespread than the indirect effect mediated by changes in LDPS in most areas. Despite significant warming trends and reductions in snow cover duration, projected responses of AAB to early-mid 21st century are heterogeneous across the continent. Changes in AAB range from strongly increasing (one order of magnitude increases in AAB) to moderately decreasing (more than halving of baseline AAB). Annual wildfire area burned in coming decades is likely to be highly geographically heterogeneous, reflecting interacting regional and seasonal climate drivers of fire occurrence and spread.

Southern Annular Mode drives multicentury wildfire activity in southern South America.

The Southern Annular Mode (SAM) is the main driver of climate variability at mid to high latitudes in the Southern Hemisphere, affecting wildfire activity, which in turn pollutes the air and contributes to human health problems and mortality, and potentially provides strong feedback to the climate system through emissions and land cover changes. Here we report the largest Southern Hemisphere network of annually resolved tree ring fire histories, consisting of 1,767 fire-scarred trees from 97 sites (from 22 °S to 54 °S) in southern South America (SAS), to quantify the coupling of SAM and regional wildfire variability using recently created multicentury proxy indices of SAM for the years 1531-2010 AD. We show that at interannual time scales, as well as at multidecadal time scales across 37-54 °S, latitudinal gradient elevated wildfire activity is synchronous with positive phases of the SAM over the years 1665-1995. Positive phases of the SAM are associated primarily with warm conditions in these biomass-rich forests, in which widespread fire activity depends on fuel desiccation. Climate modeling studies indicate that greenhouse gases will force SAM into its positive phase even if stratospheric ozone returns to normal levels, so that climate conditions conducive to widespread fire activity in SAS will continue throughout the 21st century.

Tree mortality across biomes is promoted by drought intensity, lower wood density and higher specific leaf area.

Drought events are increasing globally, and reports of consequent forest mortality are widespread. However, due to a lack of a quantitative global synthesis, it is still not clear whether drought-induced mortality rates differ among global biomes and whether functional traits influence the risk of drought-induced mortality. To address these uncertainties, we performed a global meta-analysis of 58 studies of drought-induced forest mortality. Mortality rates were modelled as a function of drought, temperature, biomes, phylogenetic and functional groups and functional traits. We identified a consistent global-scale response, where mortality increased with drought severity [log mortality (trees trees-1  year-1 ) increased 0.46 (95% CI = 0.2-0.7) with one SPEI unit drought intensity]. We found no significant differences in the magnitude of the response depending on forest biomes or between angiosperms and gymnosperms or evergreen and deciduous tree species. Functional traits explained some of the variation in drought responses between species (i.e. increased from 30 to 37% when wood density and specific leaf area were included). Tree species with denser wood and lower specific leaf area showed lower mortality responses. Our results illustrate the value of functional traits for understanding patterns of drought-induced tree mortality and suggest that mortality could become increasingly widespread in the future.

Structural overshoot of tree growth with climate variability and the global spectrum of drought-induced forest dieback.

Ongoing climate change poses significant threats to plant function and distribution. Increased temperatures and altered precipitation regimes amplify drought frequency and intensity, elevating plant stress and mortality. Large-scale forest mortality events will have far-reaching impacts on carbon and hydrological cycling, biodiversity, and ecosystem services. However, biogeographical theory and global vegetation models poorly represent recent forest die-off patterns. Furthermore, as trees are sessile and long-lived, their responses to climate extremes are substantially dependent on historical factors. We show that periods of favourable climatic and management conditions that facilitate abundant tree growth can lead to structural overshoot of aboveground tree biomass due to a subsequent temporal mismatch between water demand and availability. When environmental favourability declines, increases in water and temperature stress that are protracted, rapid, or both, drive a gradient of tree structural responses that can modify forest self-thinning relationships. Responses ranging from premature leaf senescence and partial canopy dieback to whole-tree mortality reduce canopy leaf area during the stress period and for a lagged recovery window thereafter. Such temporal mismatches of water requirements from availability can occur at local to regional scales throughout a species geographical range. As climate change projections predict large future fluctuations in both wet and dry conditions, we expect forests to become increasingly structurally mismatched to water availability and thus overbuilt during more stressful episodes. By accounting for the historical context of biomass development, our approach can explain previously problematic aspects of large-scale forest mortality, such as why it can occur throughout the range of a species and yet still be locally highly variable, and why some events seem readily attributable to an ongoing drought while others do not. This refined understanding can facilitate better projections of structural overshoot responses, enabling improved prediction of changes in forest distribution and function from regional to global scales.

A synthesis of radial growth patterns preceding tree mortality.

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks.

Survival, growth and vulnerability to drought in fire refuges: implications for the persistence of a fire-sensitive conifer in northern Patagonia.

Fire severity and extent are expected to increase in many regions worldwide due to climate change. Therefore, it is crucial to assess the relative importance of deterministic vs. stochastic factors producing remnant vegetation to understand their function in the persistence of fire-sensitive plants. Vegetation remnants (areas within the landscape that have not burned for a considerable amount of time) may occur stochastically or in more predictable locations (fire refuges) where physical conditions decrease fire severity. Our aim was to determine if remnant forests of the fire-sensitive conifer Austrocedrus chilensis are associated with biophysical attributes that allow persistence in a fire-prone Patagonian landscape. We conducted a multi-scale approach, determining attributes of forest remnants and their surroundings (matrices) through remote sensing and field-based biophysical and functional characteristics, and quantifying how tree survival probability relates to microsite conditions. Trees within remnants displayed abundant fire scars, were twofold older and had threefold larger growth rates than matrix trees. Remnants were associated with high rocky cover and elevated topographical positions. Tree survival increased in hilltops, eastern aspects, and with sparse vegetation. Trees within remnants experienced severe reductions in growth during droughts. Our results suggest that A. chilensis remnants are mainly the result of refuges, where environmental conditions increase fire survival, but also increase susceptibility to drought. A trade-off between fire survival and drought vulnerability may imply that under increasing drought and fire severity, locations that in the past have served as refuges may reduce their ability to allow the persistence of fire-sensitive taxa.

How do cold-sensitive species endure ice ages? Phylogeographic and paleodistribution models of postglacial range expansion of the mesothermic drought-tolerant conifer Austrocedrus chilensis.

In view of global climate change, it is important to understand the responses of tree species to climate changes in the past. Combinations of phylogeographic analysis of genetic evidence, coupled with species distribution models (SDMs), are improving our understanding on this subject. We combined SDMs and microsatellite data from populations of the entire range of Austrocedrus chilensis, a dominant mesotherm (cold-sensitive) conifer of dryland forests of the southern Andes, to test the hypothesis of long-distance postglacial migration from northern and warmer refugia at the Last Glacial Maximum (LGM). The SDM indicated suitable conditions for Austrocedrus in northern Chile (western) at the LGM and largely unsuitable conditions in Argentina (eastern). Population genetic diversity and effective population sizes within populations decreased southward along the Andes, consistent with the hypothesis of long-distance dispersal from a northern refugium. Results support the hypothesis of one (or a few) warmer (low latitude) refugia in Chile for Austrocedrus. On balance, the evidence suggests that in contrast to cold-tolerant tree taxa with the capacity to fast-track postglacial warming thanks to local refugia, cold-sensitive species might have undergone long-distance range expansion, lagging behind progressive climate change throughout the Holocene.

Environmental drivers and spatial dependency in wildfire ignition patterns of northwestern Patagonia.

Fire management requires an understanding of the spatial characteristics of fire ignition patterns and how anthropogenic and natural factors influence ignition patterns across space. In this study we take advantage of a recent fire ignition database (855 points) to conduct a comprehensive analysis of the spatial pattern of fire ignitions in the western area of Neuquén province (57,649 km(2)), Argentina, for the 1992-2008 period. The objectives of our study were to better understand the spatial pattern and the environmental drivers of the fire ignitions, with the ultimate aim of supporting fire management. We conducted our analyses on three different levels: statistical "habitat" modelling of fire ignition (natural, anthropogenic, and all causes) based on an information theoretic approach to test several competing hypotheses on environmental drivers (i.e. topographic, climatic, anthropogenic, land cover, and their combinations); spatial point pattern analysis to quantify additional spatial autocorrelation in the ignition patterns; and quantification of potential spatial associations between fires of different causes relative to towns using a novel implementation of the independence null model. Anthropogenic fire ignitions were best predicted by the most complex habitat model including all groups of variables, whereas natural ignitions were best predicted by topographic, climatic and land-cover variables. The spatial pattern of all ignitions showed considerable clustering at intermediate distances (<40 km) not captured by the probability of fire ignitions predicted by the habitat model. There was a strong (linear) and highly significant increase in the density of fire ignitions with decreasing distance to towns (<5 km), but fire ignitions of natural and anthropogenic causes were statistically independent. A two-dimensional habitat model that quantifies differences between ignition probabilities of natural and anthropogenic causes allows fire managers to delineate target areas for consideration of major preventive treatments, strategic placement of fuel treatments, and forecasting of fire ignition. The techniques presented here can be widely applied to situations where a spatial point pattern is jointly influenced by extrinsic environmental factors and intrinsic point interactions.

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.