Density‐dependence in common tree species in a tropical dry forest in Mudumalai,southern India

Abstract. Density‐dependence in tree population dynamics has seldom been examined in dry tropical forests. Using long‐term data from a large permanent plot, this study examined 16 common species in a dry tropical forest in southern India for density‐dependence. Employing quadrat‐based analyses, correlations of mortality, recruitment and population change with tree densities were examined. Mortality in 1–10 cm diameter trees was largely negatively correlated with conspecific density, whereas mortality in > 10 cm diameter trees was positively correlated. Mortality was, however, largely unaffected by the basal area and abundance of heterospecific trees. Recruitment was poor in most species, but in Lagerstroemia microcarpa (Lythraceae), Tectona grandis (Verbenaceae) and Cassia fistula (Fabaceae), species that recruited well, strong negative correlations of recruitment with conspecific basal area and abundance were found. In a few other species that could be tested, recruitment was again negatively correlated with conspecific density. In Lagerstroemia, recruitment was positively correlated with the basal area and abundance of heterospecific trees, but these correlations were non‐significant in other species. Similarly, although the rates of population change were negatively correlated with conspecific density they were positive when dry‐season ground fires occurred in the plot. Thus, the observed positive density‐dependence in large‐tree mortality and the negative density‐dependence in recruitment in many species were such that could potentially regulate tree populations. However, repeated fires influenced density‐dependence in the rates of population change in a way that could promote a few common species in the tree community.     

Forest dynamics,differential mortality and variable recruitment probabilities

Abstract. Both spatial and temporal variability in recruitment probabilities can lead to coexistence in gap-phase regenerating forests which would otherwise tend to be dominated by fewer species. Using modified Markov models, the potential roles were examined of temporal variability and differential mortality rates among species in the dynamics of a forest for which spatial variability has been rejected as a strong factor leading to coexistence. Differential longevity modifies results obtained from a simple Markov model: it exerts a strong influence on the equilibrium species composition, on the rate of community change and on the time a community requires to reach equilibrium. Simulations with varying transition probabilities mimicked a changing climate, producing four main results: 1. Unless the duration of climate states is very long or very short, forest composition is in a continual state of disequilibrium. 2. Species vary in their response times to changing climate. 3. The mean abundance of each species under a varying climate scenario is different from that expected from the mean climate state. 4. The rare, long-lived species was favored by climatic fluctuations at the expense of more common shorter lived species. Differential mortality rates provide an equilibrium-based mechanism for coexistence, and temporally fluctuating recruitment probabilities a non-equilibrium mechanism. Composition could be maintained by differential longevity among species and climatic fluctuations allowing periodic recruitment of the less common species.     

Soil-related variations in the population dynamics of six dipterocarp tree species with strong habitat preferences

Differences in the density of conspecific tree individuals in response to environmental gradients are well documented for many tree species, but how such density differences are generated and maintained is poorly understood. We examined the segregation of six dipterocarp species among three soil types in the Pasoh tropical forest, Malaysia. We examined how individual performance and population dynamics changed across the soil types using 10-year demographic data to compare tree performance across soil types, and constructed population matrix models to analyze the population dynamics. Species showed only minor changes in mortality and juvenile growth across soil types, although recruitment differed greatly. Clear, interspecific demographic trade-offs between growth and mortality were found in all soil types. The relative trade-offs by a species did not differ substantially among the soil types. Population sizes were projected to remain stable in all soil types for all species with one exception. Our life-table response experiment demonstrated that the population dynamics of a species differed only subtly among soil types. Therefore, species with strong density differences across soil types do not necessarily differ greatly in their population dynamics across the soil types. In contrast, interspecific differences in population dynamics were large. The trade-off between mortality and growth led to a negative correlation between the contributions of mortality and growth to variations in the population growth rate (λ) and thus reduced their net contributions. Recruitment had little impact on the variation in λ. The combination of these factors resulted in little variation in λ among species.     

Competitive and demographic leverage points of community shifts under climate warming

Accelerating rates of climate change and a paucity of whole-community studies of climate impacts limit our ability to forecast shifts in ecosystem structure and dynamics, particularly because climate change can lead to idiosyncratic responses via both demographic effects and altered species interactions. We used a multispecies model to predict which processes and species'' responses are likely to drive shifts in the composition of a space-limited benthic marine community. Our model was parametrized from experimental manipulations of the community. Model simulations indicated shifts in species dominance patterns as temperatures increase, with projected shifts in composition primarily owing to the temperature dependence of growth, mortality and competition for three critical species. By contrast, warming impacts on two other species (rendering them weaker competitors for space) and recruitment rates of all species were of lesser importance in determining projected community changes. Our analysis reveals the importance of temperature-dependent competitive interactions for predicting effects of changing climate on such communities. Furthermore, by identifying processes and species that could disproportionately leverage shifts in community composition, our results contribute to a mechanistic understanding of climate change impacts, thereby allowing more insightful predictions of future biodiversity patterns.     

Periodic dynamics in a two-stage Allee effect model are driven by tension between stage equilibria

Single species difference population models can show complex dynamics such as periodicity and chaos under certain circumstances, but usually only when rates of intrinsic population growth or other life history parameter are unrealistically high. Single species models with Allee effects (positive density dependence at low density) have also been shown to exhibit complex dynamics when combined with over-compensatory density dependence or a narrow fertility window. Here we present a simple two-stage model with Allee effects which shows large amplitude periodic fluctuations for some initial conditions, without these requirements. Periodicity arises out of a tension between the critical equilibrium of each stage, i.e. when the initial population vector is such that the adult stage is above the critical value, while the juvenile stage is below the critical value. Within this area of parameter space, the range of initial conditions giving rise to periodic dynamics is driven mainly by adult mortality rates. Periodic dynamics become more important as adult mortality increases up to a certain point, after which periodic dynamics are replaced by extinction. This model has more realistic life history parameter values than most 'chaotic' models. Conditions for periodic dynamics might arise in some marine species which are exploited (high adult mortality) leading to recruitment limitation (low juvenile density) and might be an additional source of extinction risk.     

Response of Sierra Nevada forests to projected climate–wildfire interactions

Climate influences forests directly and indirectly through disturbance. The interaction of climate change and increasing area burned has the potential to alter forest composition and community assembly. However, the overall forest response is likely to be influenced by species‐specific responses to environmental change and the scale of change in overstory species cover. In this study, we sought to quantify how projected changes in climate and large wildfire size would alter forest communities and carbon (C) dynamics, irrespective of competition from nontree species and potential changes in other fire regimes, across the Sierra Nevada, USA. We used a species‐specific, spatially explicit forest landscape model (LANDIS‐II) to evaluate forest response to climate–wildfire interactions under historical (baseline) climate and climate projections from three climate models (GFDL, CCSM3, and CNRM) forced by a medium–high emission scenario (A2) in combination with corresponding climate‐specific large wildfire projections. By late century, we found modest changes in the spatial distribution of dominant species by biomass relative to baseline, but extensive changes in recruitment distribution. Although forest recruitment declined across much of the Sierra, we found that projected climate and wildfire favored the recruitment of more drought‐tolerant species over less drought‐tolerant species relative to baseline, and this change was greatest at mid‐elevations. We also found that projected climate and wildfire decreased tree species richness across a large proportion of the study area and transitioned more area to a C source, which reduced landscape‐level C sequestration potential. Our study, although a conservative estimate, suggests that by late century, forest community distributions may not change as intact units as predicted by biome‐based modeling, but are likely to trend toward simplified community composition as communities gradually disaggregate and the least tolerant species are no longer able to establish. The potential exists for substantial community composition change and forest simplification beyond this century.     

LANDIS PRO: a landscape model that predicts forest composition and structure changes at regional scales

LANDIS PRO predicts forest composition and structure changes incorporating species‐, stand‐, and landscape‐scales processes at regional scales. Species‐scale processes include tree growth, establishment, and mortality. Stand‐scale processes contain density‐ and size‐related resource competition that regulates self‐thinning and seedling establishment. Landscape‐scale processes include seed dispersal and disturbances. LANDIS PRO is designed to be compatible with forest inventory data, thus extensive inventory data can be directly utilized to initialize and calibrate model parameters before predicting future forest changes. LANDIS PRO allows for exploring the effects of disturbances, management, climate change, and modeling the spread of invasive species. We demonstrate that LANDIS PRO successfully predicts forest successional trajectories and stand development patterns in the Central Hardwood Forest region in U.S.     

An analytical model of stand dynamics as a function of tree growth, mortality and recruitment: the shade tolerance-stand structure hypothesis revisited

Light competition and interspecific differences in shade tolerance are considered key determinants of forest stand structure and dynamics. Specifically two main stand diameter distribution types as a function of shade tolerance have been proposed based on empirical observations. All-aged stands of shade tolerant species tend to have steeply descending, monotonic diameter distributions (inverse J-shaped curves). Shade intolerant species in contrast typically exhibit normal (unimodal) tree diameter distributions due to high mortality rates of smaller suppressed trees. In this study we explore the generality of this hypothesis which implies a causal relationship between light competition or shade tolerance and stand structure. For this purpose we formulate a partial differential equation system of stand dynamics as a function of individual tree growth, recruitment and mortality which allows us to explore possible individual-based mechanisms--e.g. light competition-underlying observed patterns of stand structure--e.g. unimodal or inverse J-shaped equilibrium diameter curves. We find that contrary to expectations interspecific differences in growth patterns can result alone in any of the two diameter distributions types observed in the field. In particular, slow growing species can present unimodal equilibrium curves even in the absence of light competition. Moreover, light competition and shade intolerance evaluated both at the tree growth and mortality stages did not have a significant impact on stand structure that tended to converge systematically towards an inverse J-shaped curves for most tree growth scenarios. Realistic transient stand dynamics for even aged stands of shade intolerant species (unimodal curves) were only obtained when recruitment was completely suppressed, providing further evidence on the critical role played by juvenile stages of tree development (e.g. the sampling stage) on final forest structure and composition. The results also point out the relevance of partial differential equations systems as a tool for exploring the individual-level mechanisms underpinning forest structure, particularly in relation to more complex forest simulation models that are more difficult to analyze and to interpret from a biological point of view.     

Integrated population modeling reveals the impact of climate on the survival of juvenile emperor penguins

Early‐life demographic traits are poorly known, impeding our understanding of population processes and sensitivity to climate change. Survival of immature individuals is a critical component of population dynamics and recruitment in particular. However, obtaining reliable estimates of juvenile survival (i.e., from independence to first year) remains challenging, as immatures are often difficult to observe and to monitor individually in the field. This is particularly acute for seabirds, in which juveniles stay at sea and remain undetectable for several years. In this work, we developed a Bayesian integrated population model to estimate the juvenile survival of emperor penguins (Aptenodytes forsteri), and other demographic parameters including adult survival and fecundity of the species. Using this statistical method, we simultaneously analyzed capture–recapture data of adults, the annual number of breeding females, and the number of fledglings of emperor penguins collected at Dumont d'Urville, Antarctica, for the period 1971–1998. We also assessed how climate covariates known to affect the species foraging habitats and prey [southern annular mode (SAM), sea ice concentration (SIC)] affect juvenile survival. Our analyses revealed that there was a strong evidence for the positive effect of SAM during the rearing period (SAMR) on juvenile survival. Our findings suggest that this large‐scale climate index affects juvenile emperor penguins body condition and survival through its influence on wind patterns, fast ice extent, and distance to open water. Estimating the influence of environmental covariates on juvenile survival is of major importance to understand the impacts of climate variability and change on the population dynamics of emperor penguins and seabirds in general and to make robust predictions on the impact of climate change on marine predators.     

Signals of large scale climate drivers,hatchery enhancement,and marine factors in Yukon River Chinook salmon survival revealed with a Bayesian life history model

Understanding how species might respond to climate change involves disentangling the influence of co‐occurring environmental factors on population dynamics, and is especially problematic for migratory species like Pacific salmon that move between ecosystems. To date, debate surrounding the causes of recent declines in Yukon River Chinook salmon (Oncorhynchus tshawytscha) abundance has centered on whether factors in freshwater or marine environments control variation in survival, and how these populations at the northern extremity of the species range will respond to climate change. To estimate the effect of factors in marine and freshwater environments on Chinook salmon survival, we constructed a stage‐structured assessment model that incorporates the best available data, estimates incidental marine bycatch mortality in trawl fisheries, and uses Bayesian model selection methods to quantify support for alternative hypotheses. Models fitted to two index populations of Yukon River Chinook salmon indicate that processes in the nearshore and marine environments are the most important determinants of survival. Specifically, survival declines when ice leaves the Yukon River later in the spring, increases with wintertime temperature in the Bering Sea, and declines with the abundance of globally enhanced salmon species consistent with competition at sea. In addition, we found support for density‐dependent survival limitations in freshwater but not marine portions of the life cycle, increasing average survival with ocean age, and age‐specific selectivity of bycatch mortality in the Bering Sea. This study underscores the utility of flexible estimation models capable of fitting multiple data types and evaluating mortality from both natural and anthropogenic sources in multiple habitats. Overall, these analyses suggest that mortality at sea is the primary driver of population dynamics, yet under warming climate Chinook salmon populations at the northern extent of the species’ range may be expected to fare better than southern populations, but are influenced by foreign salmon production.     

Functional Traits Help Predict Post-Disturbance Demography of Tropical Trees

How tropical tree species respond to disturbance is a central issue of forest ecology, conservation and resource management. We define a hierarchical model to investigate how functional traits measured in control plots relate to the population change rate and to demographic rates for recruitment and mortality after disturbance by logging operations. Population change and demographic rates were quantified on a 12-year period after disturbance and related to seven functional traits measured in control plots. The model was calibrated using a Bayesian Network approach on 53 species surveyed in permanent forest plots (37.5 ha) at Paracou in French Guiana. The network analysis allowed us to highlight both direct and indirect relationships among predictive variables. Overall, 89% of interspecific variability in the population change rate after disturbance were explained by the two demographic rates, the recruitment rate being the most explicative variable. Three direct drivers explained 45% of the variability in recruitment rates, including leaf phosphorus concentration, with a positive effect, and seed size and wood density with negative effects. Mortality rates were explained by interspecific variability in maximum diameter only (25%). Wood density, leaf nitrogen concentration, maximum diameter and seed size were not explained by variables in the analysis and thus appear as independent drivers of post-disturbance demography. Relationships between functional traits and demographic parameters were consistent with results found in undisturbed forests. Functional traits measured in control conditions can thus help predict the fate of tropical tree species after disturbance. Indirect relationships also suggest how different processes interact to mediate species demographic response.     

The significance of the asymmetric effect of crowding for coexistence in a mixed temperate forest

Abstract. The coexistence of coniferous (mostly Abies homolepis) and broad-leaved tree species (mostly Fagus crenata) in a mixed temperate old-growth forest in Japan was simulated by a size-structure dynamics model incorporating the asymmetrical (one-sided) effect of shading between these two life-form guilds. The model assumes that the crowding effect due to one-sided competition for light on a tree of a given size regulates the rate of size growth and recruitment. The cumulative basal area of trees larger than a given tree in the forest is employed to express the intensity of one-sided competition on that tree. Cumulative basal areas of both guilds negatively affected the growth rate of any tree. The shading effect by conifers on the growth rate of either guild was stronger than that by broad-leaved species. Two types of model were tested for recruitment; an additive and a reciprocal model. A reciprocal model, where basal area density of conifers and broad-leaved species has a negative effect on the recruitment of its own guild but has a positive effect on that of the other guild, fit the observed data better than an additive model where total basal area of the two guilds suppresses recruitment rates. Simulations using these models showed that, within a particular range of the set of recruitment rates, the two guilds could coexist. The tendency for reciprocal replacement, incorporated in the reciprocal model, substantially widened the range of coexistence and shortened the time required for convergence.     

Patterns and Drivers of Tree Mortality in Iberian Forests: Climatic Effects Are Modified by Competition

Tree mortality is a key process underlying forest dynamics and community assembly. Understanding how tree mortality is driven by simultaneous drivers is needed to evaluate potential effects of climate change on forest composition. Using repeat-measure information from c. 400,000 trees from the Spanish Forest Inventory, we quantified the relative importance of tree size, competition, climate and edaphic conditions on tree mortality of 11 species, and explored the combined effect of climate and competition. Tree mortality was affected by all of these multiple drivers, especially tree size and asymmetric competition, and strong interactions between climate and competition were found. All species showed L-shaped mortality patterns (i.e. showed decreasing mortality with tree size), but pines were more sensitive to asymmetric competition than broadleaved species. Among climatic variables, the negative effect of temperature on tree mortality was much larger than the effect of precipitation. Moreover, the effect of climate (mean annual temperature and annual precipitation) on tree mortality was aggravated at high competition levels for all species, but especially for broadleaved species. The significant interaction between climate and competition on tree mortality indicated that global change in Mediterranean regions, causing hotter and drier conditions and denser stands, could lead to profound effects on forest structure and composition. Therefore, to evaluate the potential effects of climatic change on tree mortality, forest structure must be considered, since two systems of similar composition but different structure could radically differ in their response to climatic conditions.     

Predator density and competition modify the benefits of group formation in a shoaling reef fish

Synthesis Predation risk experienced by individuals living in groups depends on the balance between predator dilution, competition for refuges, and predator interference or synergy. These interactions operate between prey species as well: the benefits of group living decline in the presence of an alternative prey species. We apply a novel model‐fitting approach to data from field experiments to distinguish among competing hypotheses about shifts in predator foraging behavior across a range of predator and prey densities. Our study provides novel analytical tools for analyzing predator foraging behavior and offers insight into the processes driving the dynamics of coral reef fish. Studies of predator foraging behavior typically focus on single prey species and fixed predator densities, ignoring the potential importance of complexities such as predator dilution; predator‐mediated effects of alternative prey; heterospecific competition; or predator–predator interactions. Neglecting the effects of prey density is particularly problematic for prey species that live in mixed species groups, where the beneficial effects of predator dilution may swamp the negative effects of heterospecific competition. Here we use field experiments to investigate how the mortality rates of a shoaling coral reef fish (a wrasse: Thalassoma amblycephalum), change as a result of variation in: 1) conspecific density, 2) density of a predator (a hawkfish: Paracirrhites arcatus), and 3) presence of an alternative prey species that competes for space (a damselfish: Pomacentrus pavo). We quantify changes in prey mortality rates from the predator's perspective, examining the effects of added predators or a second prey species on the predator's functional response. Our analysis highlights a model‐fitting approach that discriminates amongst multiple hypotheses about predator foraging in a community context. Wrasse mortality decreased with increasing conspecific density (i.e. mortality was inversely density‐dependent). The addition of a second predator doubled prey mortality rates, without significantly changing attack rate or handling time – i.e. there was no evidence for predator interference. The presence of a second prey species increased wrasse mortality by 95%; we attribute this increase either to short‐term apparent competition (predator aggregation) or to a decrease in handling time of the predator (e.g. through decreased wrasse vigilance). In this system, 1) prey benefit from intraspecific group living though a reduced predation risk, and 2) the benefit of group living is reduced in the presence of an alternative prey species.     

Demographic synthesis for global tree species conservation

Conserving the tree species of the world requires syntheses on which tree species are most vulnerable to pressing threats, such as climate change, invasive pests and pathogens, or selective logging. Here, we review the population and forest dynamics models that, when parameterized with data from population studies, forest inventories, or tree rings, have been used for identifying life-history strategies of species and threat-related changes in population demography and dynamics. The available evidence suggests that slow-growing and/or long-lived species are the most vulnerable. However, a lack of comparative, multi-species studies still challenges more precise predictions of the vulnerability of tree species to threats. Improving data coverage for mortality and recruitment, and accounting for interactions among threats, would greatly advance vulnerability assessments for conservation prioritizations of trees worldwide.     

Recruitment Processes and Species Coexistence in a Sub-boreal Forest in Northern Japan

We investigated the recruitment of saplings (across the 2 m-heightthreshold) of six species,Picea jezoensis, Abies sachalinensis,Betula ermanii, Picea glehnii, Acer ukurunduense andSorbus commixta,in a sub-boreal forest, northern Japan. Data were collectedin a 2.48-ha plot over six growing seasons (1989–1994).We used path analysis to analyse the relationships between therecruitment rates of saplings and the stand structural attributessuch as mother tree abundance, stand crowdedness, stand stratification,Sasabamboo density on the forest floor, and fallen log abundance.The combination of stand structural attributes affecting recruitmentrates of the six sub-boreal forest tree species differed markedlyamong the species and corresponded to species composition. Itis suggested that the size-structure dynamics of adult treesof the sub-boreal forest are regulated largely by differentregeneration processes among the species and only slightly byinterspecific competition between adult trees because interspecificcompetition between adult trees was not evident. The dynamicsof species coexistence of the sub-boreal forest should be describedas a process combining the diversity of recruitment processesof saplings of the component species and the diversity of interspecificcompetition between adult trees. We propose the boundary conditionhypothesis for species coexistence in the sub-boreal forest,that the persistence of each component species is ascribed largelyto the different recruitment processes of saplings (boundaryconditions for adult tree growth dynamics) and only a littleto interspecific adult tree competition. Climax forest; safe site; regeneration niche; mode of competition; species diversity     

Effects of flooding regime and woody bamboos on tree community dynamics in a section of tropical semideciduous forest in South-Eastern Brazil

The influence of a population of the understorey woody bamboo Merostachys riedeliana and different flooding regimes on tree community dynamics in a section of tropical semideciduous forest in South-Eastern Brazil was examined. A forest section with an area of 1.6 ha composed of 71 adjacent plots was located on a slope ending at the river margin. The section was divided into five topographical sectors according to the mean duration of river floods. In 1991 and 1998 all trees with a diameter at the base of the trunk ≥ 5 cm were measured, identified and tagged, and all live bamboo culms were counted. Annualised estimates of the rates of tree mortality and recruitment, gain and loss of tree basal area, and change in bamboo density were calculated for each of the 71 plots and five topographical sectors as well as for diameter classes and tree species. To segregate patterns arising from spatially autocorrelated events, geostatistical analyses were used prior to statistical comparisons and correlations. In general, mortality rates were not compensated by recruitment rates but there was a net increase in basal area in all sectors, suggesting that the tree community as a whole was in a building phase. Tree community dynamics of the point bar forest (Depression and Levée sectors) differed from that of the upland forest (Ridgetop, Middle Slope and Lower Slope sectors) in the extremely high rates of gain in basal area. The predominant and specialised species, Inga vera and Salix humboldtiana, are probably favoured by relaxed competition in an environment stressed by long-lasting floods. In the upland forest, mortality rates were highest at the Middle Slope, particularly for smaller trees, while recruitment rates were lowest. As bamboo clumps were concentrated in this sector, the locally higher instability in the tree community probably resulted from the direct interference of bamboos. The density of bamboo culms in the upland forest was negatively correlated with the rates of tree recruitment and gain in basal area, and positively correlated with tree mortality rates. Bamboos therefore seemed to restrict the recruitment, growth and survival of trees. This revised version was published online in June 2006 with corrections to the Cover Date.     

Competition for safe real estate,not food,drives density‐dependent juvenile survival in a large herbivore

Density‐dependent competition for food reduces vital rates, with juvenile survival often the first to decline. A clear prediction of food‐based, density‐dependent competition for large herbivores is decreasing juvenile survival with increasing density. However, competition for enemy‐free space could also be a significant mechanism for density dependence in territorial species. How juvenile survival is predicted to change across density depends critically on the nature of predator–prey dynamics and spatial overlap among predator and prey, especially in multiple‐predator systems. Here, we used a management experiment that reduced densities of a generalist predator, coyotes, and specialist predator, mountain lions, over a 5‐year period to test for spatial density dependence mediated by predation on juvenile mule deer in Idaho, USA. We tested the spatial density‐dependence hypothesis by tracking the fate of 251 juvenile mule deer, estimating cause‐specific mortality, and testing responses to changes in deer density and predator abundance. Overall juvenile mortality did not increase with deer density, but generalist coyote‐caused mortality did, but not when coyote density was reduced experimentally. Mountain lion‐caused mortality did not change with deer density in the reference area in contradiction of the food‐based competition hypothesis, but declined in the treatment area, opposite to the pattern of coyotes. These observations clearly reject the food‐based density‐dependence hypothesis for juvenile mule deer. Instead, our results provide support for the spatial density‐dependence hypothesis that competition for enemy‐free space increases predation by generalist predators on juvenile large herbivores.     

Environmental heterogeneity and biotic interactions mediate climate impacts on tropical forest regeneration

Predicting the fate of tropical forests under a changing climate requires understanding species responses to climatic variability and extremes. Seedlings may be particularly vulnerable to climatic stress given low stored resources and undeveloped roots; they also portend the potential effects of climate change on future forest composition. Here we use data for ca. 50,000 tropical seedlings representing 25 woody species to assess (i) the effects of interannual variation in rainfall and solar radiation between 2007 and 2016 on seedling survival over 9 years in a subtropical forest; and (ii) how spatial heterogeneity in three environmental factors—soil moisture, understory light, and conspecific neighborhood density—modulate these responses. Community‐wide seedling survival was not sensitive to interannual rainfall variability but interspecific variation in these responses was large, overwhelming the average community response. In contrast, community‐wide responses to solar radiation were predominantly positive. Spatial heterogeneity in soil moisture and conspecific density were the predominant and most consistent drivers of seedling survival, with the majority of species exhibiting greater survival at low conspecific densities and positive or nonlinear responses to soil moisture. This environmental heterogeneity modulated impacts of rainfall and solar radiation. Negative conspecific effects were amplified during rainy years and at dry sites, whereas the positive effects of radiation on survival were more pronounced for seedlings existing at high understory light levels. These results demonstrate that environmental heterogeneity is not only the main driver of seedling survival in this forest but also plays a central role in buffering or exacerbating impacts of climate fluctuations on forest regeneration. Since seedlings represent a key bottleneck in the demographic cycle of trees, efforts to predict the long‐term effects of a changing climate on tropical forests must take into account this environmental heterogeneity and how its effects on regeneration dynamics play out in long‐term stand dynamics.     

Modeling Forest Mortality Caused by Drought Stress: Implications for Climate Change

Climate change is expected to affect forest landscape dynamics in many ways, but it is possible that the most important direct impact of climate change will be drought stress. We combined data from weather stations and forest inventory plots (FIA) across the upper Great Lakes region (USA) to study the relationship between measures of drought stress and mortality for four drought sensitivity species groups using a weight-of-evidence approach. For all groups, the model that predicted mortality as a function of mean drought length had the greatest plausibility. Model tests confirmed that the models for all groups except the most drought tolerant had predictive value. We assumed that no relationship exists between drought and mortality for the drought-tolerant group. We used these empirical models to develop a drought extension for the forest landscape disturbance and succession model LANDIS-II, and applied the model in Oconto county, Wisconsin (USA) to assess the influence of drought on forest dynamics relative to other factors such as stand-replacing disturbance and site characteristics. The simulations showed that drought stress does affect species composition and total biomass, but effects on age classes, spatial pattern, and productivity were insignificant. We conclude that (for the upper Midwest) (1) a drought-induced tree mortality signal can be detected using FIA data, (2) tree species respond primarily to the length of drought events rather than their severity, (3) the differences in drought tolerance of tree species can be quantified, (4) future increases in drought can potentially change forest composition, and (5) drought is a potentially important factor to include in forest dynamics simulations because it affects forest composition and carbon storage.