Regeneration mode affects spatial genetic structure of Nothofagus dombeyi forests

Disturbance may generate population bottlenecks by reducing population size and the number of founders establishing a new colony. We tested the hypothesis that the scale of disturbance affects the levels of genetic diversity and the spatial distribution of genotypes in naturally regenerating stands of Nothofagus dombeyi, an evergreen angiosperm tree, in northwestern Patagonia. At similar spatial scales, we predicted that old-growth stands characterized by fine-scale gap phase dynamics would be genetically diverse due to restricted gene flow among temporal and spatially isolated gaps. In contrast, young massively regenerated postfire cohorts resulting from coarse-scale disturbances would be genetically more homogeneous. At each of three paired old-growth and postfire stands a minimum of 50 trees were mapped and sampled within 1 ha. Fresh tissue was collected for isozyme analysis from a total of 361 trees along with tree cores and diameters. Tree age distributions reflected the dominant modes of regeneration. Six out of nine analysed loci were polymorphic. Mean genetic diversity parameters were greater but not significant in mature stands. Fixation indices suggested significant heterozygous deficit at two-thirds of possible tests indicating a Wahlund effect due to local recruitment of related seeds. F(ST) indicated moderate between-stand divergence. Mature stands concentrated half of positively like joins and yielded significant (P < 0.05) autocorrelation coefficients at small distance classes (< 20 m). Fine-scale patch dynamics within mature stands favours the maintenance of fine-scale genetic structure as a result of shade intolerance and local seed dispersal. Conversely, postfire stands suffer the effects of genetic drift given that a few reproductive trees produce a somewhat impoverished and genetically uniform progeny. Bottleneck effects will depend upon the density of remnant trees which could also be a function of the severity of fire.     

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.     

Disturbance and forest dynamics along a transect from Andean rain forest to Patagonian shrubland

Abstract. At ca. 40° S in northern Patagonia, Andean rain forests are replaced eastwards by woodlands and shrublands and eventually by steppe. Along this gradient we examined stand dynamics by analyzing tree population age structures and tree growth patterns. We also examined spatial and temporal characteristics of disturbance regimes by dating disturbances and mapping stands of differing disturbance history. From west to east, the ecological importance of earthquake-related disturbance decreases, whereas that of fire, logging, and livestock increases. Abrupt changes in rates of tree growth correspond with earthquakes in 1837, 1939 and 1960. In the mesic western forests earthquakes can result in massive new tree establishment on landslide-affected sites and increased rates of treefall. Fire, however, is the more pervasive disturbance over most of the gradient and creates extensive even-aged patches dominated by the regionally dominant trees, Nothofagus and Austrocedrus. Although some lightning-ignited and aboriginal-set fires occurred in these forests prior to European settlement, much of the present forest structure may be attributed to the massive burning associated with European settlement of this area near the turn of the present century. In contrast to the settlement-related increase in fire frequency in the western forested district, at the woodland/steppe ecotone the demise of the native American population resulted in a decrease in fire frequency. Heavy browsing and grazing following fire can seriously impede post-fire tree regeneration. These preliminary results document the important influences of varying disturbance regimes along a major environmental gradient in creating landscape-scale vegetation patterns.     

Isozyme variation and recent biogeographical history of the long-lived conifer Fitzroya cupressoides

Aim Palaeoenvironmental records of Pleistocene glaciation and associated vegetation changes in Patagonia have led to the hypothesis that during the last glacial maximum (LGM) tree species survived locally in favourable habitats. If present populations originated from spread from only one refugium, such as an ice‐free area of coastal Chile (Single Refugium hypothesis), we would expect that eastern populations would be genetically depauperate and highly similar to western populations. In contrast, if the ice cap was not complete and tree species persisted in forest patches on both slopes of the Andes (Multiple Refugia hypothesis), we would expect a greater degree of genetic divergence between populations either on opposite sides of the Cordillera (Cordillera Effect scenario) or towards its present‐day southern distributional limit where the ice sheet reached its maximum coverage (Extent‐of‐the‐Ice scenario). Location We tested this refugia hypothesis using patterns of isozyme variation in populations sampled over the entire modern range of the endemic conifer Fitzroya cupressoides (Mol.) Johnst. (Cupressaceae) in temperate South America. Methods Fresh foliage was collected from twenty‐four populations and analysed by horizontal electrophoresis on starch gels. Results Twenty‐one putative loci were reliably scored and 52% were polymorphic in at least one population. Populations from the eastern slope of the Andes were genetically more variable than those from the western slope; the former had a greater mean number of alleles per locus, a larger total number of alleles and rare alleles, and higher polymorphism. Genetic identities within western populations were greater than within eastern populations. Discriminant analyses using allelic frequencies of different grouping schedules of populations were non significant when testing for the Single Refugium hypothesis whereas significant results were obtained for the Multiple Refugia hypothesis. Main conclusions Our results indicate that present Fitzroya populations are the result of spreading from at least two, but possibly more, glacial refugia located in Coastal Chile and on the southern flanks of the Andes in Argentina.     

<Emphasis Type="Italic">Araucaria araucana</Emphasis> tree-ring chronologies in Argentina: spatial growth variations and climate influences

Seventeen tree-ring chronologies from the conifer Araucaria araucana (Molina) K. Koch have been analyzed across its range of distribution in Argentina. We studied the growth patterns and determined the main climatic factors influencing A. araucana radial growth. All the chronologies show a strong common signal observed by the high amount of variance explained by the first principal component (PC1) and the high mean correlation (r = 0.597) between the chronologies over the 1676–1974 interval. On this basis, we developed a regional chronology that is 866 years long (A.D. 1140–2006) and includes 621 tree-ring series. Based on the PC2 scores, chronologies were clearly separated by elevation in high- and low-elevation records. Regional tree growth is strongly negatively related to temperatures during summer and fall in the previous-growing season and spring in the current-growing season, respectively. A positive association of tree growth with precipitation is recorded during spring in the current growing season. These results suggest a close relationship between A. araucana tree growth and water availability on a regional scale. This observation is also consistent with a positive and significant correlation between our A. araucana regional record and a reconstruction of November–December rainfall for northern Patagonia inferred from the xeric Austrocedrus chilensis during the past 400 years. Negative correlations between A. araucana regional growth and the sea surface temperature in the Niño 3.4 region reflect the occurrence of above-mean summer temperatures in the region during positive tropical Pacific SST anomalies. The negative relationship with the Antarctic Oscillation (AAO) results from reduced precipitation in our study region during the positive phase of the AAO. The effect of elevation on water availability is consistent with significant correlations between ring-width variations at lower elevations and the Palmer Drought Severity Index during spring and summer in the current growing season. Our study emphasizes the high dendroclimatological potential of A. araucana chronologies for reconstructing past climate variations in northern Patagonia during the past millennium.     

Effects of introduced ungulates on forest understory communities in northern Patagonia are modified by timing and severity of stand mortality

Natural disturbances such as fires, windstorms, floods, and herbivory often act on plant communities, affecting their structure and the abundance and composition of their species. Most research has focused on the effects of single disturbances on plant communities whereas the synergistic effects of several disturbances have received less attention. In this study, we evaluated how timing and severity of tree mortality modified plant use by introduced deer and early post-mortality successional trajectories in northern Patagonian conifer forests. We sampled understory composition and deer use in Austrocedrus chilensis (ciprés de la cordillera) forest stands undergoing varying timing and severity of forest mortality as reconstructed using dendroecological techniques. In addition, we evaluated the effect of fallen logs on plant composition and deer use of plants by monitoring areas of massive dieback where fallen logs had been removed for fire hazard reduction, and nearby control areas not subjected to such removal. Stepwise regression analyses showed that history and severity of tree mortality strongly influence plant composition and deer use of plants. For deer use (with pellet counts and browsing index as response variables), results showed a positive relationship with degree of stand mortality and a negative relationship with cover of fallen logs. Similarly, cover of unpalatable shrub species was explained by canopy mortality history, whereas cover of palatable shrub species was positively associated with severity of canopy mortality. In areas where fallen logs had been removed, pellet counts were six times higher than those in control areas. Though total shrub species cover was similar between log removal and control areas, proportion of unpalatable shrubs increased in areas where fallen logs had been removed. In conclusion, deer use of plants was strongly limited by tall fallen logs, allowing palatable species to establish and grow. Fallen log removal accelerated deer entrance and changed understory composition toward more browse-resistant and unpalatable species. These results underscore the importance of considering the dynamics (timing, severity, and extent) of fallen woody debris influencing understory herbivory and post-disturbance succession. In addition, experimental results underpin the importance of maintaining snags and large woody debris in disturbed landscapes where salvage logging is a routine procedure.     

Effects of fire severity in a north Patagonian subalpine forest

Abstract. Question: What is the relative importance of fire‐induced canopy mortality, soil burning and post‐fire herbivory on tree seedling performance? Location: Subalpine Nothofagus pumilio forests at Challhuaco valley (41°13'S, 71°19'W), Nahuel Huapi National Park, Argentina. Methods: We fenced and transplanted soils of three burning severities along a fire severity gradient produced by a fire in 1996. Over two growing seasons we monitored soil water, direct incoming solar radiation, seedling survival, final seedling total biomass and root/shoot ratio. Additionally, we assessed severity‐related changes in soil properties. Results: Incoming radiation (an indicator of the amount of canopy cover left by the fire) was the primary factor influencing spring and summer top soil water availability, first and second‐year seedling survival and seedling growth. While seedling survival and soil water content were negatively affected by increased radiation, seedling final biomass was highest in very open microsites. Burned soils showed lower water holding capacity and soil carbon; however these changes did not affect topsoil water, and, contrary to expectation, there was a slight tendency toward higher seedling survival on more heavily burned soils. Herbivory significantly reduced seedling survival, but only under high‐radiation conditions. While the effect of radiation on final seedling biomass was not affected by herbivory, R/S ratios were significantly reduced by herbivory in high radiation micro sites. Conclusions: Despite inducing faster aerial growth, increased radiation and herbivory in severely burned sites may effectively prevent post‐fire regeneration in north Patagonian subalpine forest where seed sources are not limiting.     

Nutrient supply and bird predation additively control insect herbivory and tree growth in two contrasting forest habitats

It has been suggested that bottom–up and top–down forces interactively control food web dynamics. While top–down effects would increase with resource availability to plants, bottom–up effects would be stronger under low predator abundance. These predictions, however, have rarely been tested at contrasting sites while keeping the dominant plant species unchanged. Furthermore, few studies have factorially manipulated both types of forces in forest communities. For two years, we evaluated the effects of fertiliser (NPK) addition and bird exclusion on tree growth, leaf traits, insect abundance, and folivory rates in a dry/warm and a wet/cold Nothofagus pumilio forest in Patagonia, Argentina. Overall, we found no interaction between nutrient supply and bird predation, although the strength of bottom–up and top–down forces differed markedly between forest sites. Treatment effects were generally weak in the wet forest, where tree growth rates and insect herbivory were low relative to the dry forest. In the dry forest, fertilisation increased sapling growth, insect abundance and folivory, whereas bird exclusion increased leaf damage and reduced tree growth. In the wet forest, fertilisation enhanced leaf nutrient contents and folivore abundance but not sapling growth, while bird exclusion had little impact on insects or trees. These results imply that factors other than nutrients and birds were important in controlling tree growth and folivore activity in the wet forest. While treatment effect sizes varied widely among feeding guilds, in general, nutrient effects on folivores were stronger than predator effects. We conclude that, within the time‐frame of this study, tree growth and herbivory were additively affected by soil nutrients and predator presence, as bird exclusion effects did not change with elevated folivore activity on fertilised trees. We also show that both top–down and bottom–up cascades were weaker in a forest site characterised by slow‐growing juvenile trees subjected to low folivore pressure.     

Conservation genetics of the endangered conifer Fitzroya cupressoides in Chile and Argentina

Intraspecific patterns of genetic variation can often be used toidentify biogeographic divisions which can be especially useful in thedesign of conservation strategies. Although abundant empirical evidenceexist on the genetic characteristics of plant species from the NorthernHemisphere as well as tropical endangered taxa, this information isparticularly limited on threatened species from endemism-rich areas inthe southern Andes of Argentina and Chile. The objective of the currentstudy was to analyze the levels and distribution of the isozymevariation in Fitzroya cupressoides (Mol.) Johnst.(Cupressaceae), a rare conifer restricted to temperate rainforests ofnorthern Patagonia, and to evaluate the role of current conservationareas protecting the gene pool of this valuable long-lived conifer.Sampling schedules consisted of fresh foliage collected from 30 randomlyselected trees at each of 24 different populations located along thegeographic range of the species. Extraction of enzymes followed standardprocedures and homogenates were loaded in 12% starch gels whichwere analyzed by horizontal electrophoresis. Eleven enzyme systems wereresolved using a combination of four different buffer solutions whichyielded information on 21 putative loci, 52% of them werepolymorphic in at least one population. Relatively low levels ofwithin-population genetic variability were scored in Fitzroyapopulations which were approximately half of the typical levelspublished for gymnosperms (percent of polymorphic loci, P = 23 vs.53% and expected heterozygosity, H_E = 0.077 vs. 0.155for Fitzroya and other conifers respectively). Substantialbetween-population variation was detected, and certain individualpopulations stand out as much more genetically variable than nearbypopulations, which in turn are located outside protected areas. Ourfindings suggest that if the objective is to protect key species likeFitzroya, spatially explicit genetic information can be auseful tool to attain this goal.     

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.