Structure,dynamics and dendroecology of an old‐growth Fagus forest in the Apennines

Abstract. Question: Which are the structural attributes and the history of old‐growth Fagus forest in Mediterranean montane environments? What are the processes underlying their structural organization? Are these forests stable in time and how does spatial scale affect our assessment of stability? How do these forests compare to other temperate deciduous old‐growth forests? Location: 1600–1850 m a.s.l., Fagus forest near the tree line, central Apennines, Italy. Methods: An old‐growth Fagus forest was studied following historical, structural and dendroecological approaches. History of forest cover changes was analysed using aerial photographs taken in 1945, 1954, 1985 and 1994. The structural analysis was carried out in the primary old‐growth portion of the forest using 18 circular and two rectangular plots. Dendroecological analyses were conducted on 32 dominant or co‐dominant trees. Results: These primary old‐growth Fagus remnants consist of four patches that escaped logging after World War II. Both living and dead tree components are within the range of structural attributes recognized for old‐growth in temperate biomes. Dendroecological analyses revealed the roles of disturbance, competition and climate in structuring the forest. We also identified a persistent Fagus community in which gap‐phase regeneration has led to a mono‐specific multi‐aged stand at spatial scales of a few hectares, characterized by a rotated‐sigmoid diameter distribution. Conclusion: Even at the relatively small spatial scale of this study, high‐elevation Apennine Fagus forests can maintain structural characteristics consistent with those of old‐growth temperate forests. These results are important for managing old‐growth forests in the Mediterranean montane biome.     

Contrasting patterns of tree mortality in late‐successional Picea abies stands in two areas in northern Fennoscandia

Question: What are tree mortality rates and how and why do they vary in late‐successional Picea abies‐dominated forests? Do observed tree mortality patterns allow comparative assessment of models of long‐term stand development? Location: Northern boreal Fennoscandia. Methods: We measured stand structure in 10 stands in two different areas. We determined age distributions and constructed a chronology of tree deaths by cross‐dating the years of death of randomly sampled dead trees. Results: The stands in the two areas had contrasting tree age distributions, despite similar live tree structure. In one area, stands were relatively even‐aged and originated following a stand‐replacing fire 317 years earlier. The stands in the second area had an uneven age structure and virtually no signs of past fires, suggesting a very long period since the last major disturbance. The younger stands were characterized by a high mortality rate and inter‐annual variation, which we attributed to senescence of the relatively even‐aged stands approaching the maximum age of P. abies. In contrast, the tree mortality rates in the older stands were low and relatively stable. Conclusions: Patterns of tree mortality were, to a large extent, dependent on the time since the last stand‐replacing disturbance, suggesting that northern boreal P. abies stands eventually reach a shifting mosaic state maintained through small‐scale dynamics, but the time needed to reach this state appears to be lengthy; even 300 years after a forest fire stands showed changes in patterns of tree mortality that were related to the developmental stage of the stands.     

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.     

Bioclimatology of beech (Fagus sylvatica L.) in the Eastern Alps: spatial and altitudinal climatic signals identified through a tree-ring network

Aim To identify the dominant spatial patterns of Fagus sylvatica radial growth in the Eastern Alps, and to understand their relationships to climate variation and bioclimatic gradients. Location Fourteen beech stands in the Eastern Alps, growing between 200 and 1500 m a.s.l. in Italy, Slovenia and Austria. Methods At each site, trees were sampled using increment borers or by taking discs from felled trees. Cores and discs were processed by measuring and crossdating ring width. Ring width series were standardized, averaged, and prewhitened to obtain site chronologies. Hierarchical Cluster Analysis (HCA) and Principal Components Analysis of prewhitened site chronologies were used to identify spatial and altitudinal growth patterns, related to the bioclimatic position of each stand. Bootstrap correlation and response functions were computed between monthly climatic variables and either principal component scores or composite chronologies from stands associated by HCA. The stability of dendroclimatic signals was analyzed by moving correlation functions (MCF). Correlation analysis (teleconnections) based on a data base of 37 Italian and Slovenian beech tree‐ring chronologies revealed the spatial extent of principal component scores. Results Sampled trees were 200–400 years old, representing the oldest beech trees that have been crossdated for the Alps to date. Maximum age was directly related to altitude and to the presence of historical forms of conservation. Tree‐ring parameters varied according to geographic patterns and the age of sampled trees. Stands were bioclimatically organized according to their location, and with reference to their elevation and distance from the Adriatic Sea. A direct response to winter temperature was found at all elevations. The altitudinal gradient ranged from low‐elevation stands, characterized by a Mediterranean‐type, late spring–summer drought signal, to mountain and high‐elevation stands, characterized by a direct response to growing season temperature plus an inverse response to the previous year’s July temperature. The mountain and high‐elevation signal was evident in Austria, the Central Alps and Slovenia, while the low‐elevation signal was confined to mountains adjacent to the Adriatic Sea. MCF revealed trends in the response to climatic factors affecting tree‐ring formation in mountain and high‐mountain stands linked to climatic warming. Main conclusions Dendroclimatic networks can be used for bioclimatic studies of tree populations. A biogeographical separation emerged between the Alps and the Apennines at the upper elevations, while different degrees of mediterraneity distinguished sites at lower elevations. This information will be useful in assessing any future climate‐related bioclimatic shifts, especially for forests at ecotones and along altitudinal gradients.     

Tree growth response to drought and temperature in a mountain landscape in northern Arizona, USA

Aim To understand how tree growth response to regional drought and temperature varies between tree species, elevations and forest types in a mountain landscape. Location Twenty‐one sites on an elevation gradient of 1500 m on the San Francisco Peaks, northern Arizona, USA. Methods Tree‐ring data for the years 1950–2000 for eight tree species (Abies lasiocarpa var. arizonica (Merriam) Lemm., Picea engelmannii Parry ex Engelm., Pinus aristata Engelm., Pinus edulis Engelm., Pinus flexilis James, Pinus ponderosa Dougl. ex Laws., Pseudotsuga menziesii var. glauca (Beissn.) Franco and Quercus gambelii Nutt.) were used to compare sensitivity of radial growth to regional drought and temperature among co‐occurring species at the same site, and between sites that differed in elevation and species composition. Results For Picea engelmannii, Pinus flexilis, Pinus ponderosa and Pseudotsuga menziesii, trees in drier, low‐elevation stands generally had greater sensitivity of radial growth to regional drought than trees of the same species in wetter, high‐elevation stands. Species low in their elevational range had greater drought sensitivity than co‐occurring species high in their elevational range at the pinyon‐juniper/ponderosa pine forest ecotone, ponderosa pine/mixed conifer forest ecotone and high‐elevation invaded meadows, but not at the mixed conifer/subalpine forest ecotone. Sensitivity of radial growth to regional drought was greater at drier, low‐elevation compared with wetter, high‐elevation forests. Yearly growth was positively correlated with measures of regional water availability at all sites, except high‐elevation invaded meadows where growth was weakly correlated with all climatic factors. Yearly growth in high‐elevation forests up to 3300 m a.s.l. was more strongly correlated with water availability than temperature. Main conclusions Severe regional drought reduced growth of all dominant tree species over a gradient of precipitation and temperature represented by a 1500‐m change in elevation, but response to drought varied between species and stands. Growth was reduced the most in drier, low‐elevation forests and in species growing low in their elevational range in ecotones, and the least for trees that had recently invaded high‐elevation meadows. Constraints on tree growth from drought and high temperature are important for high‐elevation subalpine forests located near the southern‐most range of the dominant species.     

Soil Phosphorus Availability and Transformation Rates in Relictic Pinsapo Fir Forests from Southern Spain

Abies pinsapo fir forests are remnant of temperate-like coniferous forests currently subjected to the typical seasonal constraints of Mediterranean-type climates. We have examined for the first time, P availability and cycling in A. pinsapo forests from southern Spain by using both, measurements of concentrations of soil P fractions (resin extractable P, bicarbonate extractable P and NaOH extractable P) and rates of P supply to (P solubilization, P mineralization and phosphomonoesterases (PMEase activity) and P immobilization from the soil solution. Soils from A. pinsapo stands differing in lithology (serpentinitic and calcareous substrates) and successional status (from young-agradative to old-growth stages) were chosen for this study. Labile organic P fractions, PMEase activity and rates of P gross mineralization and immobilization were significantly higher in agradative stands on serpentines than in successionally comparable calcareous stands. This suggests an important role of the organic P subcycle in the A. pinsapo stand on serpentine. On calcareous lithology, PMEase activity and all soil P transformation rates significantly increased throughout the successional series. Similarly, all organic and inorganic P fractions measured in old-growth forests showed the maximum values of the series. These trends fit the predictions of standard patterns of P cycling changes along with succession, in which P supply to plants greatly depends on solubilization from mineral forms at early-to-mid successional stages, whereas the importance of processes related to the organic P subcycle increases as succession progresses.     

Within‐stand spatial structure and relation of boreal canopy and understorey vegetation

Question: How do spatial patterns and associations of canopy and understorey vegetation vary with spatial scale along a gradient of canopy composition in boreal mixed‐wood forests, from younger Aspen stands dominated by Populus tremuloides and P. balsamifera to older Mixed and Conifer stands dominated by Picea glauca? Do canopy evergreen conifers and broad‐leaved deciduous trees differ in their spatial relationships with understorey vegetation? Location: EMEND experimental site, Alberta, Canada. Methods: Canopy and understorey vegetation were sampled in 28 transects of 100 contiguous 0.5 m × 0.5 m quadrats in three forest stand types. Vegetation spatial patterns and relationships were analysed using wavelets. Results: Boreal mixed‐wood canopy and understorey vegetation are patchily distributed at a range of small spatial scales. The scale of canopy and understorey spatial patterns generally increased with increasing conifer presence in the canopy. Associations between canopy and understorey were highly variable among stand types, transects and spatial scales. Understorey vascular plant cover was generally positively associated with canopy deciduous tree cover and negatively associated with canopy conifer tree cover at spatial scales from 5–15 m. Understorey non‐vascular plant cover and community composition were more variable in their relationships with canopy cover, showing both positive and negative associations at a range of spatial scales. Conclusions: The spatial structure and relation of boreal mixed‐wood canopy and understorey vegetation varied with spatial scale. Differences in understorey spatial structure among stand types were consistent with a nucleation model of patch dynamics during succession in boreal mixed‐wood forests.     

Forest aboveground biomass along an elevational transect in Sulawesi,Indonesia, and the role of Fagaceae in tropical montane rain forests

Aim This study investigates how estimated tree aboveground biomass (AGB) of tropical montane rain forests varies with elevation, and how this variation is related to elevational change in floristic composition, phylogenetic community structure and the biogeography of the dominant tree taxa. Location Lore Lindu National Park, Sulawesi, Indonesia. Methods Floristic inventories and stand structural analyses were conducted on 13 plots (each 0.24 ha) in four old‐growth forest stands at 1050, 1400, 1800 and 2400 m a.s.l. (submontane to upper montane elevations). Tree AGB estimates were based on d.b.h., height and wood specific gravity. Phylogenetic diversity and biogeographical patterns were analysed based on tree family composition weighted by AGB. Elevational trends in AGB were compared with other Southeast Asian and Neotropical transect studies (n = 7). Results AGB was invariant from sub‐ to mid‐montane elevation (309–301 Mg ha^?1) and increased slightly to 323 Mg ha^?1 at upper montane elevation. While tree and canopy height decreased, wood specific gravity increased. Magnoliids accounted for most of the AGB at submontane elevations, while eurosids I (including Fagaceae) contributed substantially to AGB at all elevations. Phylogenetic diversity was highest at upper montane elevations, with co‐dominance of tree ferns, Podocarpaceae, Trimeniaceae and asterids/euasterids II, and was lowest at lower/mid‐montane elevations, where Fagaceae contributed > 50% of AGB. Biogeographical patterns showed a progression from dominant tropical families at submontane to tropical Fagaceae (Castanopsis, Lithocarpus) at lower/mid‐montane, and to conifers and Australasian endemics at upper montane elevations. Cross‐continental comparisons revealed an elevational AGB decrease in transects with low/no presence of Fagaceae, but relatively high AGB in montane forests with moderate to high abundance of this family. Main conclusions AGB is determined by both changes in forest structure and shifts in species composition. In our study, these two factors traded off so that there was no net change in AGB, even though there were large changes in forest structure and composition along the elevational gradient. Southeast Asian montane rain forests dominated by Fagaceae constitute important carbon stocks. The importance of biogeography and species traits for biomass estimation should be considered by initiatives to reduce emissions from deforestation and forest degradation (REDD) and in taxon choice in reforestation for carbon offsetting.     

Disturbance legacies and climate jointly drive tree growth and mortality in an intensively studied boreal forest

Most North American forests are at some stage of post‐disturbance regrowth, subject to a changing climate, and exhibit growth and mortality patterns that may not be closely coupled to annual environmental conditions. Distinguishing the possibly interacting effects of these processes is necessary to put short‐term studies in a longer term context, and particularly important for the carbon‐dense, fire‐prone boreal forest. The goals of this study were to combine dendrochronological sampling, inventory records, and machine‐learning algorithms to understand how tree growth and death have changed at one highly studied site (Northern Old Black Spruce, NOBS) in the central Canadian boreal forest. Over the 1999–2012 inventory period, mean tree diameter increased even as stand density and basal area declined significantly. Tree mortality averaged 1.4 ± 0.6% yr^?1, with most mortality occurring in medium‐sized trees; new recruitment was minimal. There have been at least two, and probably three, significant influxes of new trees since stand initiation, but none in recent decades. A combined tree ring chronology constructed from sampling in 2001, 2004, and 2012 showed several periods of extreme growth depression, with increased mortality lagging depressed growth by ~5 years. Higher minimum and maximum air temperatures exerted a negative influence on tree growth, while precipitation and climate moisture index had a positive effect; both current‐ and previous‐year data exerted significant effects. Models based on these variables explained 23–44% of the ring‐width variability. We suggest that past climate extremes led to significant mortality still visible in the current forest structure, with decadal dynamics superimposed on slower patterns of fire and succession. These results have significant implications for our understanding of previous work at NOBS, the carbon sequestration capability of old‐growth stands in a disturbance‐prone landscape, and the sustainable management of regional forests in a changing climate.     

Long‐term physiological and growth responses of Himalayan fir to environmental change are mediated by mean climate

High‐elevation forests are experiencing high rates of warming, in combination with CO₂ rise and (sometimes) drying trends. In these montane systems, the effects of environmental changes on tree growth are also modified by elevation itself, thus complicating our ability to predict effects of future climate change. Tree‐ring analysis along an elevation gradient allows quantifying effects of gradual and annual environmental changes. Here, we study long‐term physiological (ratio of internal to ambient CO₂, i.e., C_i/C_a and intrinsic water‐use efficiency, iWUE) and growth responses (tree‐ring width) of Himalayan fir (Abies spectabilis) trees in response to warming, drying, and CO₂ rise. Our study was conducted along elevational gradients in a dry and a wet region in the central Himalaya. We combined dendrochronology and stable carbon isotopes (δ¹³C) to quantify long‐term trends in C_i/C_a ratio and iWUE (δ¹³C‐derived), growth (mixed‐effects models), and evaluate climate sensitivity (correlations). We found that iWUE increased over time at all elevations, with stronger increase in the dry region. Climate–growth relations showed growth‐limiting effects of spring moisture (dry region) and summer temperature (wet region), and negative effects of temperature (dry region). We found negative growth trends at lower elevations (dry and wet regions), suggesting that continental‐scale warming and regional drying reduced tree growth. This interpretation is supported by δ¹³C‐derived long‐term physiological responses, which are consistent with responses to reduced moisture and increased vapor pressure deficit. At high elevations (wet region), we found positive growth trends, suggesting that warming has favored tree growth in regions where temperature most strongly limits growth. At lower elevations (dry and wet regions), the positive effects of CO₂ rise did not mitigate the negative effects of warming and drying on tree growth. Our results raise concerns on the productivity of Himalayan fir forests at low and middle (<3,300 m) elevations as climate change progresses.     

Long-Term Tree Cover Dynamics in a Pinyon-Juniper Woodland: Climate-Change-Type Drought Resets Successional Clock

Woody vegetation has expanded in coverage over the past century in many places globally, exemplified by pinyon-juniper changes in the Southwestern United States. Extreme drought is one of the few non-management drivers besides fire that might reverse such cover changes, but this has not been well documented. Here, we assess 68 years of tree cover dynamics across an elevation gradient of a pinyon-juniper woodland using aerial photographs (1936 and 1959) and QuickBird imagery (2004). Canopy cover increased 32% from 1936 to the onset of a major drought (2002). The largest relative increase in canopy cover occurred from 1936 to 1959 at the higher elevations, but these gains were eliminated by fires occurring from 1959 to 2002, during which time lower elevations with low canopy cover exhibited the greatest relative increases. The 2002–2004 drought reduced canopy cover by 55%, which eliminated gains in cover that occurred since 1936. Relative tree cover loss was highest at low elevations with low tree cover, but absolute tree cover loss was greater in areas of high tree cover, which increased with elevation. The loss of more than half of the canopy cover during a 2-year drought period was much greater than losses due to fire or possible increases due to historic land use (for example, grazing). These results suggest that regional-scale climatic influences may be more important than land use legacies in controlling tree cover of these and perhaps other semiarid woodlands over longer time scales—notable given that similar episodes of tree mortality are projected in coming decades with climate change.     

Response of pine natural regeneration to small‐scale spatial variation in a managed Mediterranean mountain forest

Questions: What influence do management practices and previous tree and shrub stand structure have on the occurrence and development of natural regeneration of Pinus sylvestris in Mediterranean mountain forests? How are the fine‐scale and environmental patterns of resources affected and what impact does this have on the distribution of the regeneration? Location: A Pinus sylvestris Mediterranean mountain forest in central Spain. Methods: Upperstory trees and regeneration (seedlings and saplings) were mapped in four 0.5‐ha plots located in two types of stand with different management intensities (even‐aged and uneven‐aged stands). Environmental variables were recorded at the nodes of a grid within the plots. The relationships between the upperstory and regeneration were evaluated by bivariate point pattern analysis; redundancy analysis ordination and variation partitioning were performed to characterize regeneration niches and the importance of the spatial component. Results: Seedlings and saplings presented a clumped structure under both types of management and their distribution was found to be related to the spatial distribution of favourable microsites. Regeneration was positively related to conditions of partial cover with high soil water content during the summer. More than half of the explained variance was spatially structured in both types of stand. This percentage was particularly high in the even‐aged stands where the pattern of regeneration was highly influenced by the gaps created by harvesting. Conclusions: The spatial distribution of the tree and shrub upperstory strongly influences regeneration patterns of P. sylvestris. Current management practices, promoting small gaps, partial canopy cover and moderate shade in even‐aged stands, or favouring tree and shrub cover in the case of uneven‐aged stands, appears to provide suitable conditions for the natural regeneration of P. sylvestris in a Mediterranean climate.     

Californian mixed-conifer forests under unmanaged fire regimes in the Sierra San Pedro Mártir, Baja California, Mexico

Aim This study appraises historical fire regimes for Californian mixed‐conifer forests of the Sierra San Pedro Mártir (SSPM). The SSPM represents the last remaining mixed‐conifer forest along the Pacific coast still subject to uncontrolled, periodic ground fire. Location The SSPM is a north–south trending fault bound range, centred on 31°N latitude, 100 km SE of Ensenada, Baja California. Methods We surveyed forests for composition, population structure, and historical dynamics both spatially and temporally over the past 65 years using repeat aerial photographs and ground sampling. Fire perimeter history was reconstructed based on time‐series aerial photographs dating from 1942 to 1991 and interpretable back to 1925. A total of 256 1‐ha sites randomly selected from aerial photographs were examined along a chronosequence for density and cover of canopy trees, density of snags and downed logs, and cover of non‐conifer trees and shrubs. Twenty‐four stands were sampled on‐the‐ground by a point‐centred quarter method which yielded data on tree density, basal area, frequency, importance value, and shrub and herb cover. Results Forests experience moderately intense understory fires that range in size to 6400 ha, as well as numerous smaller, low intensity burns with low cumulative spatial extent. SSPM forests average 25–45% cover and 65–145 trees per ha. Sapling densities were two to three times that of overstory trees. Size‐age distributions of trees ≥ 4 cm dbh indicate multi‐age stands with steady‐state dynamics. Stands are similar to Californian mixed conifer forests prior to the imposition of fire suppression policy. Livestock grazing does not appear to be suppressing conifer regeneration. Main conclusions Our spatially‐based reconstruction shows the open forest structure in SSPM to be a product of infrequent, intense surface fires with fire rotation periods of 52 years, rather than frequent, low intensity fires at intervals of 4–20 years proposed from California fire‐scar dendrochronology (FSD) studies. Ground fires in SSPM were intense enough to kill pole‐size trees and a significant number of overstory trees. We attribute long fire intervals to the gradual build‐up of subcontinuous shrub cover, conifer recruitment and litter accumulation. Differences from photo interpretation and FSD estimates are due to assumptions made with respect to site‐based (point) sampling of fire, and nonfractal fire intensities along fire size frequency distributions. Fire return intervals determined by FSD give undue importance to local burns which collectively use up little fuel, cover little area, and have little demographic impact on forests.     

Long‐term climate and competition explain forest mortality patterns under extreme drought

Rising temperatures are amplifying drought‐induced stress and mortality in forests globally. It remains uncertain, however, whether tree mortality across drought‐stricken landscapes will be concentrated in particular climatic and competitive environments. We investigated the effects of long‐term average climate [i.e. 35‐year mean annual climatic water deficit (CWD)] and competition (i.e. tree basal area) on tree mortality patterns, using extensive aerial mortality surveys conducted throughout the forests of California during a 4‐year statewide extreme drought lasting from 2012 to 2015. During this period, tree mortality increased by an order of magnitude, typically from tens to hundreds of dead trees per km², rising dramatically during the fourth year of drought. Mortality rates increased independently with average CWD and with basal area, and they increased disproportionately in areas that were both dry and dense. These results can assist forest managers and policy‐makers in identifying the most drought‐vulnerable forests across broad geographic areas.     

Radial growth responses to drought of Pinus sylvestris and Quercus pubescens in an inner‐Alpine dry valley

Question: Lower montane treeline ecotones such as the inner Alpine dry valleys are regarded as sensitive to climate change. In the dry Valais valley (Switzerland) the composition of the widespread, low altitude Pinus forests is shifting towards a mixed deciduous state. The sub‐boreal P. sylvestris shows high mortality rates, whereas the deciduous sub‐mediterranean Quercus pubescens is spreading. These species may act as early indicators of climate change. We evaluate this hypothesis by focusing on their differences in drought tolerance, which are hardly known, but are likely to be crucial in the current forest shift and also for future forest development. Methods: We used dendroecological methods to detect species‐specific patterns in the growth response to drought. The relationship between radial growth of 401 trees from 15 mixed stands and drought was analysed by calculating response functions using yearly tree‐ring indices and monthly drought indices. PCA was applied to the response ratios to discover spatial patterns of drought response. Results: A species‐specific response to moisture as well as a sub‐regional differentiation of the response patterns were found. While Quercus showed a response mainly to the conditions of the previous autumn and those of current spring, Pinus did not start responding before May, but showed responses throughout the whole summer. Quercus may restrict physiological activity to moist periods; growth of Pinus was much more dependent on prior growth. Conclusions: Given that the climate is changing towards (1) longer summer drought periods, (2) higher mean temperatures and (3) shifted seasonally of moisture availability, Quercus may benefit from adapting better to drier conditions. Pinus may increasingly face problems related to drought stress as it depends on summer moisture and has a smaller adaptive capacity due to its long‐lived photosynthetic tissue.     

A relict population of Fagus grandifolia var. mexicana at the Acatlan Volcano, Mexico: structure, litterfall, phenology and dendroecology

Aim Fagus grandifolia var. mexicana (Martinez) Little has an extraordinarily restricted distribution in the Mexican montane cloud forests. Isolated Fagus (beech) populations have been recorded in less than 10 small areas (2–40 ha) in the eastern Sierra Madre at altitudes from 1400 to 2000 m. The objectives were to determine tree and seedling age, forest structure, phenology, litterfall patterns and the relationship between mast and climatic variables. Location We report on three Fagus stands at the Acatlan Volcano, Veracruz, Mexico. Methods Changes in forest cover were determined using aerial photographs. Within each stand, basal area, density and tree species composition were determined in a 0.1‐ha band transect. Additionally, litterfall production was quantified and phenophases were recorded monthly over a 3‐year period, and 60 tree cores were collected to determine age distribution and tree‐ring growth. Results The forest was atypical in several respects. Fagus was the only dominant tree species in the crater stand, although in the rim and at the top of the volcano it was codominant with other tree species. Juveniles occurred only on the rim, but there was a seedling bank in the crater. Although forest cover in the area increased between 1968 and 1993, the Fagus stands did not change in size. Leaf production peaked in March and April, and leaf fall occurred from October through February. Litterfall production was the highest in November. During mast years, flowering started in February and between mast events there were no flowers or fruits. Minimum temperatures were highly correlated with Fagus litterfall and leaf fall. Seedlings ranged in age from 2 to 18 years and were 13–60 cm tall. Tree cores ranged from 76 to 120 years, but trees were older than the core samples. Main conclusions Although beech is considered a gap regeneration species that reaches the canopy after alternating periods of release and suppression, the trees in the crater were released when less than 1.5 m tall and have suffered few periods of suppression since. The results indicate that the crater stand was established after a severe disturbance destroyed the existing forest. We conclude that the relict beech population should be able to maintain itself, if not severely disturbed by humans or by climatic changes related to global warming.     

Stand-structural effects on Heterobasidion abietinum-related mortality following drought events in Abies pinsapo

Climate change may affect tree–pathogen interactions. This possibility has important implications for drought-prone forests, where stand dynamics and disease pathogenicity are especially sensitive to climatic stress. In addition, stand structural attributes including density-dependent tree-to-tree competition may modulate the stands’ resistance to drought events and pathogen outbreaks. To assess the effects of stand structure on root-rot-related mortality after severe droughts, we focused on Heterobasidion abietinum mortality in relict Spanish stands of Abies pinsapo, a drought-sensitive fir. We compared stand attributes and tree spatial patterns in three plots with H. abietinum root-rot disease and three plots without root-rot. Point-pattern analyses were used to investigate the scale and extent of mortality patterns and to test hypotheses related to the spread of the disease. Dendrochronology was used to date the year of death and to assess the association between droughts and growth decline. We applied a structural equation modelling approach to test if tree mortality occurs more rapidly than predicted by a simple distance model when trees are subjected to high tree-to-tree competition and following drought events. Contrary to expectations of drought mortality, the effect of precipitation on the year of death was strong and negative, indicating that a period of high precipitation induced an earlier tree death. Competition intensity, related to the size and density of neighbour trees, also induced an earlier tree death. The effect of distance to the disease focus was negligible except in combination with intensive competition. Our results indicate that infected trees have decreased ability to withstand drought stress, and demonstrate that tree-to-tree competition and fungal infection act as predisposing factors of forest decline and mortality.     

Predicting tree death for Fagus sylvatica and Abies alba using permanent plot data

Question: How well can mortality probabilities of deciduous trees(Fagus sylvatica) and conifers (Abies alba) be predicted using permanent plot data that describe growth patterns, tree species, tree size and site conditions? Location: Fagus forests in the montane belt of the Jura folds (Switzerland). Method: Permanent plot data were used to develop and validate logistic regression models predicting survival probabilities of individual trees. Backward model selection led to a reduced model containing the growth‐related variable ‘relative basal area increment’ (growth‐dependent mortality) and variables not directly reflecting growth such as species, size and site (growth‐independent mortality). Results: The growth‐mortality relationship was the same for both species (growth‐dependent mortality). However, species, site and tree size also influenced mortality probabilities (growth‐independent mortality). The predicted survival probabilities of the final model were well calibrated, and the model showed an excellent discriminatory power (area under the receiver operating characteristic curve = 0.896). Conclusion: Mortality probabilities of Fagus sylvatica and Abies alba can be predicted with high discriminatory power using a well calibrated logistic regression model. Extending this case study to a larger number of tree species and sites could provide species‐ and site‐specific tree mortality models that allow for more realistic projections of forest succession.     

Convergence in growth responses of tropical trees to climate driven by water stress

Widely documented for temperate and cold forests in both hemispheres, variations in tree growth responses to climate along environmental gradients have rarely been investigated in the tropics. Seven tree‐ring chronologies of Centrolobium microchaete (Fabaceae) in the Cerrado tropical forests of Bolivia are used to determine the growth responses to climate along a precipitation gradient. Chronologies are distributed from the humid Guarayos forests (annual precipitation > 1600 mm) in the transition to the Amazonia to the dry‐mesic Chiquitos forests (annual precipitation < 1200 mm) in the proximity to the dry Chaco. On a large spatial scale, radial growth is positively influenced by rainfall and negatively by temperature at the end of the dry season. However, this regional pattern in climate‐tree growth relationship shows differences along the precipitation gradient. Relationships with climate are highly significant and extend over longer periods of the year in sites with low rainfall and extremely severe dry seasons. At wet sites, larger water soil capacity and endogenous forest dynamics partially mask the direct influence of climate on tree growth. Stronger similarities in tree‐growth responses to climate occur between sites in the dry Central Chiquitos and in the transition to the Guarayos forests. In contrast, the relationships show fewer similarities between sites in the humid Guarayos. We conclude that growth responses to climate in the tropics are more similar between sites with limited rainfall and severe and prolonged dry seasons. Our study points to a convergence in the patterns of growth responses of tropical trees to climate, modulated by scarce rainfall and marked seasonality. The negative impact of water deficits on tree physiological processes induces not only the documented reduction in forest species richness, but also a convergence in tree‐growth responses to climate in dry tropical forests.     

Modeling landscape patterns of understory tree regeneration in the Pacific Northwest,USA

Abstract. Vegetation maps serve as the basis for spatial analysis of forest ecosystems and provide initial information for simulations of forest landscape change. Because of the limitations of current remote sensing technology, it is not possible to directly measure forest understory attributes across large spatial extents. Instead we used a predictive vegetation mapping approach to model Tsuga heterophylla and Picea sitchensis seedling patterns in a 3900‐ha landscape in the Oregon Coast Range, USA, as a function of Landsat TM imagery, aerial photographs, digital elevation models, and stream maps. Because the models explained only moderate amounts of variability (R² values of 0.24–0.56), we interpreted the predicted patterns as qualitative spatial trends rather than precise maps. P. sitchensis seedling patterns were tightly linked to the riparian network, with highest densities in coastal riparian areas. T. heterophylla seedlings exhibited complex patterns related to topography and overstory forest cover, and were also spatially clustered around patches of old‐growth forest. We hypothesize that the old growth served as refugia for this fire‐sensitive species following wildfires in the late 19th and early 20th centuries. Low levels of T. heterophylla regeneration in hardwood‐dominated forests suggest that these patches may succeed to shrublands rather than to conifer forest. Predictive models of seedling patterns could be developed for other landscapes where georeferenced inventory plots, remote sensing data, digital elevation models, and climate maps are available.