Quantifying the effects of drought on abrupt growth decreases of major tree species in Switzerland

Drought entails important effects on tree physiology, which may result in short‐ to long‐term radial growth decreases. While the majority of studies have focused on annual drought‐related variability of growth, relatively little is known about sustained growth decreases following drought years. We apply a statistical framework to identify climatic factors that induce abrupt growth decreases and may eventually result in tree mortality. We used tree‐ring data from almost 500 standing dead trees and 200 living trees in eight sites of the Swiss network of strict forest reserves, including four of the most important Central European tree species (Abies alba, Picea abies, Fagus sylvatica and Quercus spp.). First, to assess short‐term growth responses to drought under various climate and site conditions, we calculated correlations and linear mixed‐effects models between ring‐width indices (RWIs) and drought based on the Standardized Precipitation Evapotranspiration Index (SPEI). Second, to quantify drought effects on abrupt growth decreases, we applied distributed lag nonlinear models (DLNMs), which account for both delayed effects and the nonlinear relationship between the SPEI and the occurrence of abrupt growth decreases. Positive correlations between RWIs and the SPEI indicated short‐term growth responses of all species, particularly at arid sites. Results of the DLNMs revealed species‐specific growth responses to drought. For Quercus spp., abrupt growth decreases were more likely to occur several years following severe drought, whereas for P. abies, A. alba, and F. sylvatica abrupt growth decreases started frequently immediately in the drought year. We conclude that the statistical framework allows for quantifying the effects of drought intensity on the probability of abrupt growth decreases, which ultimately contributes to an improved understanding of climate impacts on forest community dynamics.     

Spatial patterns and associations in a Quercus‐Betula forest in northern China

Question: Are species‐specific regeneration strategies and competition the dominant processes facilitating species coexistence in a Quercus liaotungensis dominated temperate deciduous forest? Location: Dongling Mountains, North China, 1300 m a.s.l. Methods: Ripley's K‐function was used to characterize the spatial patterns and spatial associations of two dominant tree species, Quercus liaotungensis and Betula dahurica, and a common subcanopy species, Acer mono, at different growth stages (adult, sapling, seedling). Results: Seedlings, saplings and adults of all three species exhibited clumped distributions at most spatial scales. Quercus seedlings and saplings were positively associated with conspecific adult trees and spatially independent of dead trees suggesting that seed dispersal and vegetative regeneration influenced the spatial patterning of Quercus trees. Betula seedlings and saplings were positively associated with both live and dead trees of conspecific adults at small scales (<5 m) but negatively associated with live and dead trees of other species indicating sprouting as an important mechanism of reproduction. Saplings of Acer had a strong spatial dependence on the distribution of conspecific adult trees indicating its limited seed dispersal range. Negative associations between adult trees of Betula and Quercus demonstrated interspecific competition at local scales (<5 m). Conclusions: Different regeneration strategies among the three species play an important role in regulating their spatial distribution patterns, while competition between individuals of Betula and Quercus at the adult stage also contributes to spatial patterning of these communities. The recruitment limitations of Betula and Quercus may affect the persistence of these species and the long‐term dynamics of the forest.     

Is climate an important driver of post‐European vegetation change in the Eastern United States?

Many ecological phenomena combine to direct vegetation trends over time, with climate and disturbance playing prominent roles. To help decipher their relative importance during Euro‐American times, we employed a unique approach whereby tree species/genera were partitioned into temperature, shade tolerance, and pyrogenicity classes and applied to comparative tree‐census data. Our megadata analysis of 190 datasets determined the relative impacts of climate vs. altered disturbance regimes for various biomes across the eastern United States. As the Euro‐American period (ca. 1500 to today) spans two major climatic periods, from Little Ice Age to the Anthropocene, vegetation changes consistent with warming were expected. In most cases, however, European disturbance overrode regional climate, but in a manner that varied across the Tension Zone Line. To the north, intensive and expansive early European disturbance resulted in the ubiquitous loss of conifers and large increases of Acer, Populus, and Quercus in northern hardwoods, whereas to the south, these disturbances perpetuated the dominance of Quercus in central hardwoods. Acer increases and associated mesophication in Quercus‐Pinus systems were delayed until mid 20th century fire suppression. This led to significant warm to cool shifts in temperature class where cool‐adapted Acer saccharum increased and temperature neutral changes where warm‐adapted Acer rubrum increased. In both cases, these shifts were attributed to fire suppression rather than climate change. Because mesophication is ongoing, eastern US forests formed during the catastrophic disturbance era followed by fire suppression will remain in climate disequilibrium into the foreseeable future. Overall, the results of our study suggest that altered disturbance regimes rather than climate had the greatest influence on vegetation composition and dynamics in the eastern United States over multiple centuries. Land‐use change often trumped or negated the impacts of warming climate, and needs greater recognition in climate change discussions, scenarios, and model interpretations.     

Stomatal acclimation to vapour pressure deficit doubles transpiration of small tree seedlings with warming

Future climate change is expected to increase temperature (T) and atmospheric vapour pressure deficit (VPD) in many regions, but the effect of persistent warming on plant stomatal behaviour is highly uncertain. We investigated the effect of experimental warming of 1.9–5.1 °C and increased VPD of 0.5–1.3 kPa on transpiration and stomatal conductance (g_s) of tree seedlings in the temperate forest understory (Duke Forest, North Carolina, USA). We observed peaked responses of transpiration to VPD in all seedlings, and the optimum VPD for transpiration (D_opt) shifted proportionally with increasing chamber VPD. Warming increased mean water use of Carya by 140% and Quercus by 150%, but had no significant effect on water use of Acer. Increased water use of ring‐porous species was attributed to (1) higher air T and (2) stomatal acclimation to VPD resulting in higher g_s and more sensitive stomata, and thereby less efficient water use. Stomatal acclimation maintained homeostasis of leaf T and carbon gain despite increased VPD, revealing that short‐term stomatal responses to VPD may not be representative of long‐term exposure. Acclimation responses differ from expectations of decreasing g_s with increasing VPD and may necessitate revision of current models based on this assumption.     

Loss of oak dominance in dry-mesic deciduous forests predicted by gap capture methods

Gap capture methods predict future forest canopy species composition from the tallest trees growing in canopy gaps rather than from random samples of shaded understory trees. We used gap capture methods and a simulation approach to forecast canopy composition in three old oak forests (Quercus spp.) on dry-mesic sites in southern Wisconsin, USA. In the simulation, a gap sapling is considered successful if it exceeds a threshold height of 13–17 m (height of maximum crown width of canopy trees) before its crown center can be overtopped by lateral crown growth of mature trees. The composition of both the tallest gap trees and simulated gap captures suggests that 68–90% of the next generation of canopy trees in the stands will consist of non-Quercus species, particularly Ulmus rubra, Carya ovata and Prunus serotina. Quercus species will probably remain as a lesser stand component, with Quercus alba and Quercus rubra predicted to comprise about 19% of successful gap trees across the three stands. Several methods of predicting future canopy composition gave similar results, probably because no gap opportunist species were present in these stands and there was an even distribution of species among height strata in gaps. Gap trees of competing species already average 11–13 m tall, and mean expected time for these trees to reach full canopy height is only 19 years. For these reasons, we suggest that dominance will shift from oaks to other species, even though late successional species (e.g., Acer and Tilia) are not presently common in the understories of these stands.     

Annual Growth Ring Patterns in Brachystegia spiciformis Reveal Influence of Precipitation on Tree Growth1

The availability of exactly dated tree‐ring chronologies is limited in tropical regions. However, these chronologies could contribute widely to studies of the influence of natural and human‐induced factors on tropical forests. We examine the potential for building a chronology based on three sites in the miombo woodland of western Zambia. Brachystegia spiciformis Benth., a dominant species from this vegetation type, is used. Response of the chronology to several climatic factors is examined. All specimens showed very clear growth rings, and cross‐dating between radii of a tree was successful for all trees. Site chronologies could be constructed after cross‐dating of growth ring series of individual trees. The mean growth ring curves of the three sites were significantly similar, allowing for the construction of a regional chronology. Correlation function analysis between the tree‐ring chronology and regional climatic variables revealed that climate at the core of the rainy season, in December and January, has an explicit influence on tree growth. Where precipitation and relative humidity in these months influence tree growth positively, temperature correlates in a negative way. Some 20 percent of the variance in the B. spiciformis tree‐ring chronology is accounted for by wet season rainfall. The successful cross‐dating and correlation between a tree‐ring chronology and climate demonstrated in this study indicate annual ring formation in B. spiciformis trees and sensitivity to climatic conditions.     

Did soil development limit spruce (Picea abies) expansion in the Central Alps during the Holocene? Testing a palaeobotanical hypothesis with a dynamic landscape model

Aim Forest communities in the European Central Alps are highly sensitive to climatic change. Palaeobotanical studies have demonstrated that forests rapidly expanded upslope during Holocene warm intervals and contracted when temperatures fell. However, temperature alone cannot account for important changes in tree species abundance. For example, population expansion by Norway spruce (Picea abies), a dominant subalpine species, lagged suitable temperatures by about 3000 years in eastern and by 6000 years in western Switzerland. We hypothesize that spruce expansion was delayed by limited water availability in weakly developed soils and/or by drier‐than‐present climatic conditions. Location We examine the impact of reduced moisture availability on forest dynamics using a combined dynamic modelling/palaeoecological approach at two high‐elevational lakes in the Swiss Central Alps. Methods We simulate Holocene vegetation dynamics with the LandClim model in landscapes surrounding the two lakes and validate the model output by comparison with palaeobotanical reconstructions from the same sites. We evaluate the impact of shallow soils on vegetation dynamics at these sites by varying soil water‐holding capacity (i.e. bucket size) and precipitation abundance in model scenarios. Results Simulations with modern soil conditions and precipitation abundance matched reconstructed vegetation dynamics near the tree line, where temperature limits plant growth, but simulated abundant spruce during the entire Holocene. Spruce was absent only in simulations with a maximum bucket size of less than 7 cm, or when precipitation was reduced by at least 60%. In exploratory simulations of future conditions with average temperatures raised by 4 °C, the low water‐holding capacity of shallow alpine soils, not temperature, determined the upper elevational limit of spruce. Main conclusions Spruce expanded in the Central Alps only after soils developed sufficient water‐holding capacity and precipitation neared its modern abundance. Soil development will probably constrain the future response of tree species to warmer conditions (e.g. upslope migrations), as it did in the past.     

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.     

XYLEM STRUCTURE AND ECOLOGICAL DOMINANCE IN A FOREST COMMUNITY

The relative dominance of deciduous trees having diffuse, semiring, or ring porous xylem is considered in relation to percent slope, site, vertical stratification within site, and to disturbance. Xylem type relative dominance was calculated on the basis of basal area by a procedure that is generally employed for the determination of species relative dominance. The study area was located within the central hardwoods region of the Eastern Deciduous Forest. On a moisture gradient, the sites were a hydric floodplain, a mesic river terrace and cove, and a xeric hillside spur. There was an increase from lowland to upland sites in absolute number of species having diffuse porous and those having ring porous xylem; however, the data showed no significant difference among the four sites or between the cove's northwest and southwest aspects in the percentages of species having diffuse porous xylem and those having ring porous xylem. In contrast to species number data, relative dominance data showed that diffuse porous species were dominant on the floodplain, on the northwest aspect of the cove, and in lower strata beneath the dominant ring porous upper canopy on the spur and southwest aspect of the cove. Ring porous species were dominant in the upper canopy only on the spur and on the southwest aspect of the cove. Semiring porous species had a high dominance level on the disturbed terrace site, a much lower level of dominance on the less disturbed floodplain, and very low dominance on the relatively undisturbed cove and spur sites. The results demonstrated that sets of closely and distantly related tree species having the same xylem anatomy share some common ecological characteristics, suggesting that xylem structure is a critical character in the adaptation of trees to variations in the environment over small geographical areas.     

Mesophication in temperate Europe: A dendrochronological reconstruction of tree succession and fires in a mixed deciduous stand in Białowieża Forest

1. The shift from shade‐intolerant species to shade‐tolerant mesophytic species in deciduous and mixed forests of the temperate zone is well described in studies from North America. This process has been termed mesophication and it has been linked to changes in fire regime. Fire suppression results in the cessation of establishment of heliophytic, fire‐dependent tree species such as oak (Quercus) and pine (Pinus). Due to the scarcity of old‐growth forests in Europe, data on long‐term compositional changes in mixed forests are very limited, as is the number of studies exploring whether fire played a role in shaping the dynamics.
2. The aim of this study was to reconstruct tree succession in a 43‐ha natural mixed deciduous forest stand in Bia?owie?a Forest (BF), Poland using dendrochronological methods. In addition, the presence of aboveground fire legacies (charred and fire‐scarred deadwood) enabled the fire history reconstruction.
3. Dendrochronological data revealed tree establishment (Quercus) back to the end of the 1500s and fires back to 1659. Under a regime of frequent fires until the end of the 18th century, only oak and pine regenerated, sporadically. A shift in the fire regime in the first half of the 19th century triggered oak and pine cohort regeneration, then gradually spruce (Picea) encroached. Under an increasingly dense canopy and less flammable conditions, regeneration of shade‐tolerant Carpinus, Tilia, and Acer began simultaneously with the cessation of oak and pine recruitment.
4. Synthesis. The study reports the first evidence of mesophication in temperate Europe and proves that fire was involved in shaping the long‐term dynamics of mixed deciduous forest ecosystems. Our data suggest that fire exclusion promoted a gradual recruitment of fire‐sensitive, shade‐tolerant species that inhibited the regeneration of oak and pine in BF.

     

Increasing carbon discrimination rates and depth of water uptake favor the growth of Mediterranean evergreen trees in the ecotone with temperate deciduous forests

Tree populations at the low‐altitudinal or ‐latitudinal limits of species' distributional ranges are predicted to retreat toward higher altitudes and latitudes to track the ongoing changes in climate. Studies have focused on the climatic sensitivity of the retreating species, whereas little is known about the potential replacements. Competition between tree species in forest ecotones will likely be strongly influenced by the ecophysiological responses to heat and drought. We used tree‐ring widths and δ¹³C and δ¹⁸O chronologies to compare the growth rates and long‐term ecophysiological responses to climate in the temperate‐Mediterranean ecotone formed by the deciduous Fagus sylvatica and the evergreen Quercus ilex at the low altitudinal and southern latitudinal limit of F. sylvatica (NE Iberian Peninsula). F. sylvatica growth rates were similar to those of other southern populations and were surprisingly not higher than those of Q. ilex, which were an order of magnitude higher than those in nearby drier sites. Higher Q. ilex growth rates were associated with high temperatures, which have increased carbon discrimination rates in the last 25 years. In contrast, stomatal regulation in F. sylvatica was proportional to the increase in atmospheric CO₂. Tree‐ring δ¹⁸O for both species were mostly correlated with δ¹⁸O in the source water. In contrast to many previous studies, relative humidity was not negatively correlated with tree‐ring δ¹⁸O but had a positive effect on Q. ilex tree‐ring δ¹⁸O. Furthermore, tree‐ring δ¹⁸O decreased in Q. ilex over time. The sensitivity of Q. ilex to climate likely reflects the uptake of deep water that allowed it to benefit from the effect of CO₂ fertilization, in contrast to the water‐limited F. sylvatica. Consequently, Q. ilex is a strong competitor at sites currently dominated by F. sylvatica and could be favored by increasingly warmer conditions.     

Seasonal physiology and growth of bottomland oaks of differing planting stocks in afforestation sites on the U.S. Gulf Coastal Plain

Hurricane Katrina caused large losses of bottomland hardwood forests on the Gulf Coastal Plain. Heavy‐seeded species such as oaks (Quercus) generally require direct planting for restoration after such losses. However, evaluating the performance of various oak planting stocks using biometric data alone can be challenging due to their slow juvenile growth and belowground carbon allocation. Our study goals were to evaluate physiological parameters including photosynthesis, stomatal conductance, and water‐use efficiency (WUE) and their correlation with annual height growth to determine differences in functional performance and drought tolerance between seedling types and whether physiology can predict height growth. Monthly growing season gas exchange measurements were made on two oak species (Quercus texana and Quercus shumardii) and three planting stocks (bare root, conventional containerized, large containerized [LC]) planted on two sites in coastal Mississippi. We found that photosynthesis decreased throughout the growing season while stomatal conductances increased leading to decreasing WUEs in all seedling types. Physiological parameters differed across planting stocks but not species. Particularly, LC seedlings exhibited greater WUEs and sensitivity to vapor pressure deficit (VPD) suggesting increased moisture stress compared with other planting stocks. Across seedling types, photosynthesis, stomatal sensitivity to VPD, and seasonal changes in intrinsic WUE measured in year one of the study were significantly correlated with year two, but not year one height growth, suggesting belowground allocation of carbon during the first growing season. In total, these results highlight the use of physiological measurements in selecting successful planting stocks for various site conditions.     

Microbial inoculation influences arbuscular mycorrhizal fungi community structure and nutrient dynamics in temperate tree restoration

Soil microbial communities have a profound influence on soil chemical processes and subsequently influence tree nutrition and growth. This study examined how the addition of a commercial inoculum or forest‐collected soils influenced nitrogen (N) and phosphorus (P) dynamics, soil microbial community structure, and growth in Liriodendron tulipifera and Prunus serotina tree saplings. Inoculation method was an important determinant of arbuscular mycorrhizal fungi (AMF) community structure in both species and altered soil N dynamics in Prunus and soil P dynamics in Liriodendron. Prunus saplings receiving whole forest soil transfers had a higher rhizosphere soil carbon/nitrogen ratio and ammonia content at the end of the first growing season when compared to unmanipulated control saplings. Inoculation with whole forest soil transfers resulted in increased inorganic phosphorus in Liriodendron rhizosphere soils. The number of AMF terminal restriction fragments was significantly greater in rhizosphere soils of Liriodendron saplings inoculated with whole forest soil transfers and Prunus saplings receiving either inoculum source than control saplings. Forest soil inoculation also increased AMF colonization and suppressed stem elongation in Liriodendron after 16 months; conversely, Prunus AMF colonization was unchanged and stem elongation was significantly greater when saplings were inoculated with whole forest soil transfers. Longer term monitoring of tree response to inoculation will be essential to assess whether early costs of AMF colonization may provide long‐term benefits. This study provides insight into how practitioners can use microbial inoculation to alter AMF community structure and functioning, subsequently influencing tree growth and nutrient cycling during the restoration of degraded lands.     

Carbon and water relations of juvenile Quercus species in tall‐grass prairie

Abstract. In ecosystems where environments are extreme, such as deserts, adult plant species may facilitate the establishment and growth of seedlings and juveniles. Because high temperatures and evaporative demand characterize tall‐grass prairies of the central United States (relative to forests), we predicted that the grassland‐forest ecotone, by minimizing temperature extremes and moderating water stress, may function to facilitate the expansion of Quercus species into undisturbed tall‐grass prairie. We assessed the carbon and water relations of juvenile Quercus macrocarpa and Q. muhlenbergii, the dominant tree species in gallery forests of northeast Kansas, in ecotone and prairie sites. To evaluate the potentially competitive effects of neighboring herbaceous biomass on these oaks, juveniles (< 0.5 m tall) of both species also were subjected to either: (1) removal of surrounding above‐ground herbaceous biomass, or (2) control (prairie community intact) treatments. Herbaceous biomass removal had no significant effect on gas exchange or water relations in these oak species in either the prairie or the ecotone environment. Although the ecotone did alleviate some environmental extremes, photosynthetic rates and stomatal conductance were ca. 20 % higher (p < 0.05) in both oaks in prairie sites vs. the ecotone. Moreover, although leaf temperatures on average were higher in oaks in the prairie, high leaf temperatures in the ecotone had a greater negative effect on photosynthesis. These data suggest that the grassland‐forest ecotone did not facilitate the growth of Quercus juveniles expanding into this grassland. Moreover, the carbon and water relations of juvenile oaks in the prairie appeared to be unaffected by the presence of the dominant C₄ grasses.     

Fifteen polymorphic simple sequence repeat markers from expressed sequence tags of Liriodendron tulipifera

We developed and evaluated simple sequence repeat (SSR) markers derived from expressed sequence tags (ESTs) of Liriodendron tulipifera. Characteristics of 15 EST‐SSR loci were investigated using 33 L. tulipifera individuals. The number of alleles per locus ranged from two to five. The expected and observed heterozygosities ranged from 0.216 to 0.751 and from 0.182 to 0.97, respectively. These loci were further tested for their cross‐species transferability to Liriodendron Chinense. Because of their high level of polymorphism and transferability, our 15 single‐locus EST‐SSR markers will be valuable tools for research on mating system, population genetics and systemic evolution of Liriodendron.     

Post-glacial changes in spatial patterns of vegetation across southern New England

Aim We analysed lake‐sediment pollen records from eight sites in southern New England to address: (1) regional variation in ecological responses to post‐glacial climatic changes, (2) landscape‐scale vegetational heterogeneity at different times in the past, and (3) environmental and ecological controls on spatial patterns of vegetation. Location The eight study sites are located in southern New England in the states of Massachusetts and Connecticut. The sites span a climatic and vegetational gradient from the lowland areas of eastern Massachusetts and Connecticut to the uplands of north‐central and western Massachusetts. Tsuga canadensis and Fagus grandifolia are abundant in the upland area, while Quercus, Carya and Pinus species have higher abundances in the lowlands. Methods We collected sediment cores from three lakes in eastern and north‐central Massachusetts (Berry East, Blood and Little Royalston Ponds). Pollen records from those sites were compared with previously published pollen data from five other sites. Multivariate data analysis (non‐metric multi‐dimensional scaling) was used to compare the pollen spectra of these sites through time. Results Our analyses revealed a sequence of vegetational responses to climate changes occurring across southern New England during the past 14,000 calibrated radiocarbon years before present (cal yr bp ). Pollen assemblages at all sites were dominated by Picea and Pinus banksiana between 14,000 and 11,500 cal yr bp ; by Pinus strobus from 11,500 to 10,500 cal yr bp ; and by P. strobus and Tsuga between 10,500 and 9500 cal yr bp . At 9500–8000 cal yr bp , however, vegetation composition began to differentiate between lowland and upland sites. Lowland sites had higher percentages of Quercus pollen, whereas Tsuga abundance was higher at the upland sites. This spatial heterogeneity strengthened between 8000 and 5500 cal yr bp , when Fagus became abundant in the uplands and Quercus pollen percentages increased further in the lowland records. The differentiation of upland and lowland vegetation zones remained strong during the mid‐Holocene Tsuga decline (5500–3500 cal yr bp ), but the pattern weakened during the late‐Holocene (3500–300 cal yr bp ) and European‐settlement intervals. Within‐group similarity declined in response to the uneven late‐Holocene expansion of Castanea, while between‐group similarity increased due to homogenization of the regional vegetation by forest clearance and ongoing disturbances. Main conclusions The regional gradient of vegetation composition across southern New England was first established between 9500 and 8000 cal yr bp . The spatial heterogeneity of the vegetation may have arisen at that time in response to the development or strengthening of the regional climatic gradient. Alternatively, the differentiation of upland and lowland vegetation types may have occurred as the climate ameliorated and an increasing number of species arrived in the region, arranging themselves in progressively more complex vegetation patterns across relatively stationary environmental gradients. The emergence of a regional vegetational gradient in southern New England may be a manifestation of the increasing number of species and more finely divided resource gradient.     

Spatial variation of climatic and non-climatic controls on species distribution: the range limit of Tsuga heterophylla

Aim To assess which climatic variables control the distribution of western hemlock (Tsuga heterophylla), how climatic controls vary over latitude and between disjunct coastal and interior sub‐distributions, and whether non‐climatic factors, such as dispersal limitation and interspecific competition, affect range limits in areas of low climatic control. Location North‐western North America. Methods We compared four bioclimatic variables [actual evapotranspiration (AET), water deficit (DEF), mean temperature of the coldest month (MTCO), and growing degree‐days (GDD5)] with the distribution of T. heterophylla at a 2‐km grid cell resolution. The distribution is based on a zonal ecosystem classification where T. heterophylla is the dominant late‐successional species. For each bioclimatic variable and at each degree of latitude, we calculated the threshold that best defines the T. heterophylla distribution and assessed the extent to which T. heterophylla was segregated to one end of the bioclimatic gradient. We also fitted two forms of multivariate bioclimatic models to predict the T. heterophylla distribution: a simple threshold model and a complex Gaussian mixture model. Each model was trained separately on the coastal and interior distributions, and predicted areas outside of the T. heterophylla distribution (overprediction) were evaluated with respect to known outlier populations. Results Actual evapotranspiration was the most accurate predictor across the T. heterophylla distribution; other variables were important only in certain areas. There was strong latitudinal variation in the thresholds of all variables except AET, and the interior distribution had wider bioclimatic thresholds than the coastal distribution. The coastal distribution was predicted accurately by both bioclimatic models; areas of overprediction rarely occurred > 10 km from the observed distribution and generally matched small outlier populations. In contrast, the interior distribution was poorly predicted by both models; areas of overprediction occurred up to 140 km from the observed distribution and did not match outlier populations. The greatest overprediction occurred in Idaho and Montana in areas supporting species that typically co‐exist with T. heterophylla. Main conclusions The high predictive capacity of AET is consistent with this species’ physiological requirements for a mild and humid climate. Spatial variation of MTCO, GDD5 and DEF thresholds probably reflects both the correlation of these variables with AET and ecotypic variation. The level of overprediction in portions of the interior suggests that T. heterophylla has not completely expanded into its potential habitat. Tsuga heterophylla became common in the interior 2000–3500 years ago, compared with > 9000 years ago in the coastal region. The limited time for dispersal, coupled with frequent fires at the margins of the distribution and competition with disturbance‐adapted species, may have retarded range expansion in the interior. This study demonstrates that bioclimatic modelling can help identify various climatic and non‐climatic controls on species distributions.     

Evidence for the widespread occurrence of ancient forests on cliffs

Aim The objective of this work was to determine if the existence of ancient forests on cliffs was specific to the Niagara Escarpment, Canada, or part of a globally widespread pattern. Location Sixty‐five cliff sites were visited in five countries in the temperate climatic zone, and trees were sampled for age and growth rate on forty‐six of these. Methods Two hundred and twenty‐four core samples or cross‐sections were taken from trees on cliffs that varied in height, aspect, rock‐type, and exposure. General observations were also made of regeneration of the tree species forming the mature canopy, and other habitat conditions. Results The evidence shows that ancient slow‐growing forest occurs on most cliffs. Age and growth rate distributions were similar at all treed sites. Small‐statured Thuja, Juniperus, or Taxus stems with age estimates in excess of 1000 years were found in the United States, the United Kingdom and France, and small Pinus and Quercus stems nearly 400 years in Germany. There was a high rate of recurrence of plants in the genera Polypodium, Asplenium, Cystopteris, Campanula, Rosa, Prunus, Hedera, and Sorbus. Most of the sites appear to be habitats of completely natural origin. Conclusions We conclude that ancient natural forest is a normal feature of cliffs, at least in the temperate zone.     

Oak seedling survival and growth along resource gradients in Mediterranean forests: implications for regeneration in current and future environmental scenarios

Understanding seedling performance across resource gradients is crucial for defining the regeneration niche of plant species under current environmental conditions and for predicting potential changes under a global change scenario. A 2‐year field experiment was conducted to determine how seedling survival and growth of two evergreen and two deciduous Quercus species vary along gradients of light and soil properties in two Mediterranean forests with contrasting soils and climatic conditions. Half the seedlings were subjected to an irrigation treatment during the first year to quantify the effects on performance of an alteration in the summer drought intensity. Linear and non‐linear models were parameterized and compared to identify major resources controlling seedling performance. We found both site‐specific and general patterns of regeneration. Strong site‐specificity was found in the identity of the best predictors of seedling survival: survival decreased linearly with increasing light (i.e. increasing desiccation risk) in the drier site, whereas it decreased logistically with increasing spring soil water content (i.e. increasing waterlogging risk) in the wetter site. We found strong empirical support for multiple resource limitation at the drier site, the response to light being modulated by the availability of soil resources (water and P). Evidence for regeneration niche partitioning among Quercus species was only found at the wetter site. However, at both sites Quercus species shared the same response to summer drought alleviation through water addition: increased first‐year survival but not final survival (i.e. after two years). This suggests that extremely dry summers (i.e. the second summer in the experiment) can cancel out the positive effects of previous wetter summers. Therefore, an increase in the intensity and frequency of summer drought with climate change might cause a double negative impact on Quercus regeneration, due to a general reduction in survival probability and the annulment of the positive effects of (infrequent) ‘wet’ years. Overall, results presented in this study are a major step towards the development of a mechanistic model of Mediterranean forest dynamics that incorporates the idiosyncrasies and generalities of tree regeneration in these systems, and that allow simulation and prediction of the ecological consequences of resource level alterations due to global change.     

Climatic water deficit,tree species ranges,and climate change in Yosemite National Park

Aim (1) To calculate annual potential evapotranspiration (PET), actual evapotranspiration (AET) and climatic water deficit (Deficit) with high spatial resolution; (2) to describe distributions for 17 tree species over a 2300‐m elevation gradient in a 3000‐km² landscape relative to AET and Deficit; (3) to examine changes in AET and Deficit between past (c. 1700), present (1971–2000) and future (2020–49) climatological means derived from proxies, observations and projections; and (4) to infer how the magnitude of changing Deficit may contribute to changes in forest structure and composition. Location Yosemite National Park, California, USA. Methods We calculated the water balance within Yosemite National Park using a modified Thornthwaite‐type method and correlated AET and Deficit with tree species distribution. We used input data sets with different spatial resolutions parameterized for variation in latitude, precipitation, temperature, soil water‐holding capacity, slope and aspect. We used climate proxies and climate projections to model AET and Deficit for past and future climate. We compared the modelled future water balance in Yosemite with current species water‐balance ranges in North America. Results We calculated species climatic envelopes over broad ranges of environmental gradients – a range of 310 mm for soil water‐holding capacity, 48.3°C for mean monthly temperature (January minima to July maxima), and 918 mm yr^?1 for annual precipitation. Tree species means were differentiated by AET and Deficit, and at higher levels of Deficit, species means were increasingly differentiated. Modelled Deficit for all species increased by a mean of 5% between past (c. 1700) and present (1971–2000). Projected increases in Deficit between present and future (2020–49) were 23% across all plots. Main conclusions Modelled changes in Deficit between past, present and future climate scenarios suggest that recent past changes in forest structure and composition may accelerate in the future, with species responding individualistically to further declines in water availability. Declining water availability may disproportionately affect Pinus monticola and Tsuga mertensiana. Fine‐scale heterogeneity in soil water‐holding capacity, aspect and slope implies that plant water balance may vary considerably within the grid cells of kilometre‐scale climate models. Sub‐grid‐cell soil and topographical data can partially compensate for the lack of spatial heterogeneity in gridded climate data, potentially improving vegetation‐change projections in mountainous landscapes with heterogeneous topography.