Multi-model analysis of tree competition along environmental gradients in southern New England forests.

Robust predictions of competitive interactions among canopy trees and variation in tree growth along environmental gradients represent key challenges for the management of mixed-species, uneven-aged forests. We analyzed the effects of competition on tree growth along environmental gradients for eight of the most common tree species in southern New England and southeastern New York using forest inventory and analysis (FIA) data, information theoretic decision criteria, and multi-model inference to evaluate models. The suite of models estimated growth of individual trees as a species-specific function of average potential diameter growth, tree diameter at breast height, local environmental conditions, and crowding by neighboring trees. We used ordination based on the relative basal area of species to generate a measure of site conditions in each plot. Two ordination axes were consistent with variation in species abundance along moisture and fertility gradients. Estimated potential growth varied along at least one of these axes for six of the eight species; peak relative abundance of less shade-tolerant species was in all cases displaced away from sites where they showed maximum potential growth. Our crowding functions estimate the strength of competitive effects of neighbors; only one species showed support for the hypothesis that all species of competitors have equivalent effects on growth. The relative weight of evidence (Akaike weights) for the best models varied from a low of 0.207 for Fraxinus americana to 0.747 for Quercus rubra. In such cases, model averaging provides a more robust platform for prediction than that based solely on the best model. We show that predictions based on the selected best models dramatically overestimated differences between species relative to predictions based on the averaged set of models.     

Competition among eucalyptus trees depends on genetic variation and resource supply

Genetic variation and environmental heterogeneity fundamentally shape the interactions between plants of the same species. According to the resource partitioning hypothesis, competition between neighbors intensifies as their similarity increases. Such competition may change in response to increasing supplies of limiting resources. We tested the resource partitioning hypothesis in stands of genetically identical (clone-origin) and genetically diverse (seed-origin) Eucalyptus trees with different water and nutrient supplies, using individual-based tree growth models. We found that genetic variation greatly reduced competitive interactions between neighboring trees, supporting the resource partitioning hypothesis. The importance of genetic variation for Eucalyptus growth patterns depended strongly on local stand structure and focal tree size. This suggests that spatial and temporal variation in the strength of species interactions leads to reversals in the growth rank of seed-origin and clone-origin trees. This study is one of the first to experimentally test the resource partitioning hypothesis for intergenotypic vs. intragenotypic interactions in trees. We provide evidence that variation at the level of genes, and not just species, is functionally important for driving individual and community-level processes in forested ecosystems.     

Plant uptake of inorganic and organic nitrogen: neighbor identity matters

The importance of interspecific competition as a cause of resource partitioning among species has been widely assumed but rarely tested. Using neighbor removals in combination with 15N tracer additions in the field, we examined variation among three alpine species in the uptake of 15N-NH4+, 15N-NO3-, and 15N-13C-[2]-glycine in intact neighborhoods, when paired with a specific neighbor, and when all neighbors were removed. Species varied in the capacity to take up 15N-labeled NH4+, NO3-, and glycine in intact neighborhoods and in interspecific pairs. When interspecific neighbor pairs were compared with no neighbor controls, neighbors reduced 15N uptake in target species by as much as 50%, indicating competition for N. Furthermore, neighbor identity influenced the capacity of species to take up different forms of N. Thus, competition within interspecific neighbor pairs often caused reduced uptake of a particular form of N, as well as shifts to uptake of an alternative form of N. Such shifts in resource use as a result of competition are an implicit assumption in studies of resource partitioning but have rarely been documented. Our study suggests that plasticity in the uptake of different forms of N may be a mechanism by which cooccurring plants reduce competition for N.     

Importance of competition mechanisms in successive invasions by polyphagous tephritids in La Reunion

Understanding the strength and modes of interspecific interactions between introduced and resident species (native or previously introduced) is necessary to predict invasion success. We evaluated different mechanisms of interspecific competition among four species of polyphagous fruit flies (Diptera: Tephritidae) from the island of La Reunion: one endemic species, Ceratitis catoirii, and three exotic species, C. capitata, C. rosa, and Bactrocera zonata, that have successively invaded the island. Larval competition experiments, i.e., co-infestations of the same fruit, and behavioral interference experiments measuring the ability of one female to displace another from a fruit, were performed among all pairs of the four species. We observed asymmetric and hierarchical interactions among species in both larval and adult interference competition. In agreement with the hypothesis that invasion is competition-limited, the competitive hierarchy coincided with the temporal sequence of establishment on the island, i.e., each newly established species tended to be competitively dominant over previously established ones.     

Competition, resources, and vegetation during 10 years in native grassland

A 10-year experiment tested for variation in competition intensity over time in a natural grassland at the northern edge of the Great Plains. Growing-season precipitation varied fivefold during the study. All ecosystem-level variables varied significantly among years, and most covaried in expected ways. The covers of all common grasses possessing the C3 photosynthetic pathway varied significantly among years; in contrast, all common species with traits associated with drought tolerance (a C4 grass, a lichen, a spikemoss, and a subshrub) did not vary. Annual transplant experiments measured the competitive effects of neighbors on the growth of individuals of the native grass Bouteloua gracilis. A significant interaction between year and competition showed that competition intensity varied among years. The size of this effect, however, was small (eta2 = 0.074) relative to the size of the direct effect of competition (eta2 = 0.20) or the year in which the experiment was conducted (eta2 = 0.51). Further, competition intensity was not significantly related to any variable describing standing crop or resources, or species richness. Species richness was highest in years with high precipitation, standing crop, and individual growth, due to the recruitment of rare species that were absent from dry years. In summary, variation in competition intensity was statistically significant but had small effects relative to the direct effects of climate.     

Coexistence of three specialist aphids on common milkweed, Asclepias syriaca

Coexistence of host-specific herbivores on plants is believed to be governed by interspecific interactions, but few empirical studies have systematically unraveled these dynamics. We investigated the role of several factors in promoting coexistence among the aphids Aphis nerii, Aphis asclepiadis, and Myzocallis asclepiadis that all specialize on common milkweed (Asclepias syriaca). Competitive exclusion is thought to occur when interspecific competition is stronger than intraspecific competition. Consequently, we investigated whether predators, mutualists, or resource quality affected the strength of intra- vs. interspecific competition among aphids in factorial manipulations of competition with exposure to predation, ants, and variable plant genotypes in three separate experiments. In the predation x competition experiment, predators reduced aphid per capita growth by 66%, but the strength of intra- and interspecific competition did not depend on predators. In the ants x competition experiment, ants reduced per capita growth of A. nerii and M. asclepiadis (neither of which were mutualists with ants) by approximately one-half. In so doing, ants ameliorated the negative effects of these competitors on ant-tended A. asclepiadis by two-thirds, representing a novel benefit of ant-aphid mutualism. Nevertheless, ants alone did not explain the persistence of competitively inferior A. asclepiadis as, even in the presence of ants, interspecific competition remained stronger than intraspecific competition. In the plant genotype x competition experiment, both A. asclepiadis and M. asclepiadis were competitively inferior to A. nerii, with the strength of interspecific competition exceeding that of intraspecific competition by 83% and 23%, respectively. Yet these effects differed among milkweed genotypes, and there were one or more plant genotypes for each aphid species where coexistence was predicted. A synthesis of our results shows that predators play little or no role in preferentially suppressing competitively dominant A. nerii. Nonetheless, A. asclepiadis benefits from ants, and A. asclepiadis and M. asclepiadis may escape competitive exclusion by A. nerii on select milkweed genotypes. Taken as a whole, the coexistence of three host-specific aphid species sharing the same resource was promoted by the dual action of ants as antagonists and mutualists and by genetic diversity in the plant population itself.     

Improving the effectiveness of angular dispersion in plant neighbourhood models

Spatial arrangement can be an important factor affecting competition among plants. We evaluated three ways to improve the effectiveness of angular dispersion (AD) for describing spatial arrangement in plant neighbourhood models. First, we modified Zar's (1974) AD formula by weighting each neighbour by its competitive influence. We calculated this using two different competition indices to derive an AD of competitive influence, rather than of equally weighted plant locations, around a subject plant. Secondly, we constrained the effect of AD on the neighbourhood model using an optimised parameter that defines the minimum value that AD can adopt. Thirdly, we included the direction in which competition is concentrated (the mean azimuth of the weighted AD) in the growth models. These developments were evaluated within a radial growth model of Scots pine and birch growing in semi-natural, spatially heterogeneous forest. Weighted AD resulted in significant improvements in predicted radial growth of target trees over the traditional measure of AD. The optimised parameter that defines the minimum value of AD consistently evolved values significantly higher than zero. This suggests that clumped and dispersed neighbourhoods do not differ in their negative effects on a subject tree as much as expected. The inclusion of directional components of the weighted AD did not improve the accuracy of the growth models. Weighting of the angular dispersion of neighbours improved the performance of local competition models.     

Rain forest fragmentation and the proliferation of successional trees

The effects of habitat fragmentation on diverse tropical tree communities are poorly understood. Over a 20-year period we monitored the density of 52 tree species in nine predominantly successional genera (Annona, Bellucia, Cecropia, Croton, Goupia, Jacaranda, Miconia, Pourouma, Vismia) in fragmented and continuous Amazonian forests. We also evaluated the relative importance of soil, topographic, forest dynamic, and landscape variables in explaining the abundance and species composition of successional trees. Data were collected within 66 permanent 1-ha plots within a large (approximately 1000 km2) experimental landscape, with forest fragments ranging from 1 to 100 ha in area. Prior to forest fragmentation, successional trees were uncommon, typically comprising 2-3% of all trees (> or =10 cm diameter at breast height [1.3 m above the ground surface]) in each plot. Following fragmentation, the density and basal area of successional trees increased rapidly. By 13-17 years after fragmentation, successional trees had tripled in abundance in fragment and edge plots and constituted more than a quarter of all trees in some plots. Fragment age had strong, positive effects on the density and basal area of successional trees, with no indication of a plateau in these variables, suggesting that successional species could become even more abundant in fragments over time. Nonetheless, the 52 species differed greatly in their responses to fragmentation and forest edges. Some disturbance-favoring pioneers (e.g., Cecropia sciadophylla, Vismia guianensis, V. amazonica, V. bemerguii, Miconia cf. crassinervia) increased by >1000% in density on edge plots, whereas over a third (19 of 52) of all species remained constant or declined in numbers. Species responses to fragmentation were effectively predicted by their median growth rate in nearby intact forest, suggesting that faster-growing species have a strong advantage in forest fragments. An ordination analysis revealed three main gradients in successional-species composition across our study area. Species gradients were most strongly influenced by the standlevel rate of tree mortality on each plot and by the number of nearby forest edges. Species-composition also varied significantly among different cattle ranches, which differed in their surrounding matrices and disturbance histories. These same variables were also the best predictors of total successional-tree abundance and species richness. Successional-tree assemblages in fragment interior plots (>150 m from edge), which are subjected to fragment area effects but not edge effects, did not differ significantly from those in intact forest, indicating that area effects per se had little influence on successional trees. Soils and topography also had little discernable effect on these species. Collectively, our results indicate that successional-tree species proliferate rapidly in fragmented Amazonian forests, largely as a result of chronically elevated tree mortality near forest edges and possibly an increased seed rain from successional plants growing in nearby degraded habitats. The proliferation of fast-growing successional trees and correlated decline of old-growth trees will have important effects on species composition, forest dynamics, carbon storage, and nutrient cycling in fragmented forests.     

Escape from competition: neighbors reduce Centaurea stoebe performance at home but not away

The greater abundance of some exotic plants in their nonnative ranges might be explained in part by biogeographic differences in the strength of competition, but these competitive effects have not been experimentally examined in the field. We compared the effects of neighbors on the growth and reproduction of spotted knapweed (Centaurea stoebe) in Europe, where it is native, and in Montana, where it is invasive. There were strong negative competitive effects of neighboring vegetation on C. stoebe growth and reproduction in Europe. In contrast, identical experiments in Montana resulted in insignificant impacts on C. stoebe. Although the mechanisms that produce this dramatic biogeographic difference in competitive outcome remain unknown, our results indicate that differences in net competitive interactions between ranges may contribute to the striking dominance of C. stoebe in parts of North America.     

Competitive interactions among sessile organisms in Tomioka Bay,south Japan: importance of light conditions on the panel surface

Interspecific competitive interactions among sessile epibenthos were studied by suspending PVC panels at Tomioka Bay, south Japan, for a maximum period of 16 mo. Interactions were monitored from photographs of a fixed area of the panels. Four panels were suspended during two different months in autumn 1991, and the development of the community was followed until December 1992. Altogether, 6511 interspecific overgrowth interactions were recorded, of which 37 resulted in standoffs and the rest in overgrowths. The competitive relationship observed in this sessile assemblage followed the pattern of a hierarchy with numerous backloops. Among the 36 species, belonging to the seven taxonomic groups encountered during the study, the colonial ascidian Didemnum moseleyi was recorded as the dominant species (with respect to competitive ability) while the barnacle Balanus trigonus was the weakest species. The month of panel exposure and whether or not the panel surface was shaded had a significant influence on the competitive ability of the sessile organisms. The order of hierarchy of the most dominant species changed with the month of panel submersion and its light conditions. Among the several abundant species tested, longer residence times were recorded for serpulid worms than for the colonial species. A significant, positive relationship was obtained between the areal cover of competitively dominant sessile organisms and the number of their interspecific interactions. From the short residence time of sessile organisms and the significant relationship between their areal cover and number of interspecific interactions, it is concluded that the interspecific interactions played important role in the species succession.     

Interactions among Frugivores and Fleshy Fruit Trees in a Philippine Submontane Rainforest

Abstract: We assessed the potential effect of frugivore extinctions on forest regeneration in the North Negros Forest Reserve, a forest fragment that is one of the last remaining wet tropical rainforest ecosystems in the biogeographic region of the central Philippine Islands. We evaluated foraging observations of 19 species of birds, fruit bats, and other mammals in three successional habitats and identified tree species that are potentially at risk because their seeds are dispersed by frugivores that are seriously endangered. The relative abundance of zoochorous trees in this forest community was exceptionally high (80%), suggesting that the process of forest regeneration will change drastically if endangered frugivores are hunted to extinction. We grouped 45 tree species as early-, mid-, or late-successional species based on their population structure and we demonstrated that early-successional tree species were visited by a wide spectrum of frugivores, whereas mid- and late-successional species were visited mostly by hornbills (Bucerotidae) and fruit pigeons (Columbidae). Late-successional tree species were most specialized with respect to dispersers and could therefore be susceptible to extinction. We recommend tree species that could be useful for assisted natural regeneration projects in the reserve because they are visited by a variety of frugivores. Of those, we recommend early-successional trees for open-field plantations and mid-successional tree species for enrichment plantings.     

Influence of environment, disturbance, and ownership on forest vegetation of coastal Oregon.

Information about how vegetation composition and structure vary quantitatively and spatially with physical environment, disturbance history, and land ownership is fundamental to regional conservation planning. However, current knowledge about patterns of vegetation variability across large regions that is spatially explicit (i.e., mapped) tends to be general and qualitative. We used spatial predictions from gradient models to examine the influence of environment, disturbance, and ownership on patterns of forest vegetation biodiversity across a large forested region, the 3-million-ha Oregon Coast Range (USA). Gradients in tree species composition were strongly associated with environment, especially climate, and insensitive to disturbance, probably because many dominant tree species are long-lived and persist throughout forest succession. In contrast, forest structure was strongly correlated with disturbance and only weakly with environmental gradients. Although forest structure differed among ownerships, differences were blurred by the presence of legacy trees that originated prior to current forest management regimes. Our multi-ownership perspective revealed biodiversity concerns and benefits not readily visible in single-ownership analyses, and all ownerships contributed to regional biodiversity values. Federal lands provided most of the late-successional and old-growth forest. State lands contained a range of forest ages and structures, including diverse young forest, abundant legacy dead wood, and much of the high-elevation true fir forest. Nonindustrial private lands provided diverse young forest and the greatest abundance of hardwood trees, including almost all of the foothill oak woodlands. Forest industry lands encompassed much early-successional forest, most of the mixed hardwood-conifer forest, and large amounts of legacy down wood. The detailed tree- and species-level data in the maps revealed regional trends that would be masked in traditional coarse-filter assessment. Although abundant, most early-successional forests originated after timber harvest and lacked legacy live and dead trees important as habitat and for other ecological functions. Many large-conifer forests that might be classified as old growth using a generalized forest cover map lacked structural features of old growth such as multilayered canopies or dead wood. Our findings suggest that regional conservation planning include all ownerships and land allocations, as well as fine-scale elements of vegetation composition and structure.     

Grassland restoration with and without fire: evidence from a tree-removal experiment

Forest encroachment threatens the biological diversity of grasslands globally. Positive feedbacks can reinforce the process, affecting soils and ground vegetation, ultimately leading to replacement of grassland by forest species. We tested whether restoration treatments (tree removal, with or without fire) reversed effects of nearly two centuries of encroachment by Abies grandis and Pinus contorta into dry, montane meadows in the Cascade Range, Oregon, USA. In nine, 1-ha plots containing a patchy mosaic of meadow openings and forests of varying age (20 to > 140 yr), we compared three treatments affecting the ground vegetation: control (no trees removed), unburned (trees removed, slash burned in piles leaving 90% of the area unburned), and burned (trees removed, slash broadcast burned). We quantified changes over 3-4 years in soils, abundance and richness of species with differing habitat associations (meadow, forest, and ruderal), and recruitment of conifers. Except for a transient increase in available N (especially in burn scars), effects of burning on soils were minimal due, in part, to mixing by gophers. Tree removal greatly benefited meadow species at the expense of forest herbs. Cover and richness of meadow species increased by 47% and 38% of initial values in unburned plots, but changed minimally in burned plots. In contrast, cover and richness of forest herbs declined by 44% and 26% in unburned plots and by 79% and 58% in burned plots. Ruderal species and conifer seedlings were uncommon in both treatments. Although vegetation was consumed beneath burn piles, meadow species recovered significantly after three years. Long-term tree presence did not preclude recovery of meadow species; in fact, colonization was greater in older than in younger forests. In sum, temporal trends were positive for most indicators, suggesting strong potential for restoration. Contrary to conventional wisdom, tree removal without fire may be sufficient to shift the balance from forest to meadow species. In meadows characterized by historically infrequent fire, small-scale disturbances and competitive interactions may be more critical to ecosystem maintenance and restoration. Managers facing the worldwide phenomenon of tree invasion should critically evaluate the ecological vs. operational need for fire in ecosystem restoration.     

Potential site productivity influences the rate of forest structural development

Development and maintenance of structurally complex forests in landscapes formerly managed for timber production is an increasingly common management objective. It has been postulated that the rate of forest structural development increases with site productivity. We tested this hypothesis for Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forests using a network of permanent study plots established following complete timber harvest of the original old-growth forests. Forest structural development was assessed by comparing empirical measures of live tree structure to published values for Douglas-fir forests spanning a range of ages and structural conditions. The rate of forest structural development--resilience--exhibited a positive relationship with site index, a measure of potential site productivity. Density of shade-intolerant conifers declined in all study stands from an initial range of 336-4068 trees/ha to a range of 168-642 trees/ha at the most recent measurement. Angiosperm tree species declined from an initial range of 40-371 trees/ha to zero in seven of the nine plots in which they were present. Trends in shade-tolerant tree density were complex: density ranged from 0 to 575 trees/ha at the first measurement and was still highly variable (25-389 trees/ha) at the most recent measurement. Multivariate analysis identified the abundance of hardwood tree species as the strongest compositional trend apparent over the study period. However, structural variables showed a strong positive association with increasing shade-tolerant basal area and little or no association with abundance of hardwood species. Thus, while tree species succession and forest structural development occur contemporaneously, they are not equivalent processes, and their respective rates are not necessarily linearly related. The results of this study support the idea that silvicultural treatments to accelerate forest structural development should be concentrated on lower productivity sites when the management objective is reserve-wide coverage of structurally complex forests. Alternatively, high-productivity sites should be prioritized for restoration treatments when the management objective is to develop structurally complex forests on a portion of the landscape.     

Seeing the forest for the heterogeneous trees: stand-scale resource distributions emerge from tree-scale structure

Forest ecosystem processes depend on local interactions that are modified by the spatial pattern of trees and resources. Effects of resource supplies on processes such as regeneration are increasingly well understood, yet we have few tools to compare resource heterogeneity among forests that differ in structural complexity. We used a neighborhood approach to examine understory light and nutrient availability in a well-replicated and large-scale variable-retention harvesting experiment in a red pine forest in Minnesota, USA. The experiment included an unharvested control and three harvesting treatments with similar tree abundance but different patterns of retention (evenly dispersed as well as aggregated retention achieved by cutting 0.1- or 0.3-ha gaps). We measured light and soil nutrients across all treatments and mapped trees around each sample point to develop an index of neighborhood effects (NI). Field data and simulation modeling were used to test hypotheses that the mean and heterogeneity of resource availability would increase with patchiness because of greater variation in competitive environments. Our treatments dramatically altered the types and abundances of competitive neighborhoods (NI) in each stand and resulted in significantly nonlinear relationships of light, nitrogen and phosphorus availability to NI. Hence, the distribution of neighborhoods in each treatment had a significant impact on resource availability and heterogeneity. In dense control stands, neighborhood variation had little impact on resource availability, whereas in more open stands (retention treatments), it had large effects on light and modest effects on soil nutrients. Our results demonstrate that tree spatial pattern can affect resource availability and heterogeneity in explainable and predictable ways, and that neighborhood models provide a useful tool for scaling heterogeneity from the individual tree to the stand. These insights are needed to anticipate the outcomes of silvicultural manipulations and should become more holistically integrated into both basic ecological and management science.     

Fungal endophytes directly increase the competitive effects of an invasive forb

Competitive outcomes among plants can vary in different abiotic and biotic conditions. Here we tested the effects of two phylotypes of Alternaria endophytes on the growth, competitive effects, and competitive responses of the exotic invasive forb Centaurea stoebe. Centaurea stoebe was a better competitor against North American grass species than grasses from its European home range in the absence of endophytes. However, one endophyte both increased the biomass of C. stoebe and reduced the competitive effect of North American grasses on C. stoebe. The competitive effects of C. stoebe on grass species native to North America were enhanced by both fungal endophytes, but not for native European grasses. We do not know the mechanism by which endophytes increased C. stoebe's competitive ability, and particularly against biogeographically new neighbors, but one endophyte increased the competitive ability of C. stoebe without increasing its size, suggesting mechanisms unrelated to increased growth. We tested only a fraction of the different endophytic fungi that have been found in C. stoebe, only scratching the surface of understanding their indirect effects. However, our results are the first to demonstrate such effects of a fungal endophyte infecting an invasive forb, and one of the few to show that endophyte effects on competition do not have to be mediated through herbivory.     

Spatial synchrony in coral reef fish populations and the influence of climate

We investigated spatial patterns of synchrony among coral reef fish populations and environmental variables over an eight-year period on the Great Barrier Reef, Australia. Our aims were to determine the spatial scale of intra- and interspecific synchrony of fluctuations in abundance of nine damselfish species (genus Pomacentrus) and assess whether environmental factors could have influenced population synchrony. All species showed intraspecific synchrony among populations on reefs separated by < or =100 km, and interspecific synchrony was also common at this scale. At greater spatial scales, only four species showed intraspecific synchrony, over distances ranging from 100-300 km to 500-800 km, and no cases of interspecific synchrony were recorded. The two mechanisms most likely to cause population synchrony are dispersal and environmental forcing through regionally correlated climate (the Moran effect). Dispersal may have influenced population synchrony over distances up to 100 km as this is the expected spatial range for ecologically significant reef fish dispersal. Environmental factors are also likely to have synchronized population fluctuations via the Moran effect for three reasons: (1) dispersal could not have caused interspecific synchrony that was common over distances < or =100 km because dispersal cannot link populations of different species, (2) variations in both sea surface temperature and wind speed were synchronized over greater spatial scales (>800 km) than fluctuations in damselfish abundance (< or =800 km) and were correlated with an index of global climate variability, the El Ni?o-Southern Oscillation (ENSO), and (3) synchronous population fluctuations of most damselfish species were correlated with ENSO; large population increases often followed ENSO events. We recorded regional variations in the strength of population synchrony that we suspect are due to spatial differences in geophysical, oceanographic, and population characteristics, which act to dilute or enhance the effects of synchronizing mechanisms. We conclude that synchrony is common among Pomacentrus populations separated by tens of kilometers but less prevalent at greater spatial scales, and that environmental variation linked to global climate is likely to be a driving force behind damselfish population synchrony at all spatial scales on the Great Barrier Reef.     

Spatial elements of mortality risk in old-growth forests

For many species of long-lived organisms, such as trees, survival appears to be the most critical vital rate affecting population persistence. However, methods commonly used to quantify tree death, such as relating tree mortality risk solely to diameter growth, almost certainly do not account for important spatial processes. Our goal in this study was to detect and, if present, to quantify the relevance of such processes. For this purpose, we examined purely spatial aspects of mortality for four species, Abies concolor, Abies magnifica, Calocedrus decurrens, and Pinus lambertiana, in an old-growth conifer forest in the Sierra Nevada of California, USA. The analysis was performed using data from nine fully mapped long-term monitoring plots. In three cases, the results unequivocally supported the inclusion of spatial information in models used to predict mortality. For Abies concolor, our results suggested that growth rate may not always adequately capture increased mortality risk due to competition. We also found evidence of a facilitative effect for this species, with mortality risk decreasing with proximity to conspecific neighbors. For Pinus lambertiana, mortality risk increased with density of conspecific neighbors, in keeping with a mechanism of increased pathogen or insect pressure (i.e., a Janzen-Connell type effect). Finally, we found that models estimating risk of being crushed were strongly improved by the inclusion of a simple index of spatial proximity. Not only did spatial indices improve models, those improvements were relevant for mortality prediction. For P. lambertiana, spatial factors were important for estimation of mortality risk regardless of growth rate. For A. concolor, although most of the population fell within spatial conditions in which mortality risk was well described by growth, trees that died occurred outside those conditions in a disproportionate fashion. Furthermore, as stands of A. concolor become increasingly dense, such spatial factors are likely to become increasingly important. In general, models that fail to account for spatial pattern are at risk of failure as conditions change.     

Effects of Pioneer Tree Species Hyperabundance on Forest Fragments in Northeastern Brazil

Abstract: Despite many studies on fragmentation of tropical forests, the extent to which plant and animal communities are altered in small, isolated forest fragments remains obscure if not controversial. We examined the hypothesis that fragmentation alters the relative abundance of tree species with different vegetative and reproductive traits. In a fragmented landscape (670 km²) of the Atlantic Forest of northeastern Brazil, we categorized 4056 trees of 182 species by leafing pattern, reproductive phenology, and morphology of seeds and fruit. We calculated relative abundance of traits in 50 1‐ha plots in three types of forest configurations: forest edges, small forest fragments (3.4–83.6 ha), and interior of the largest forest fragment (3500 ha, old growth). Although evergreen species were the most abundant across all configurations, forest edges and small fragments had more deciduous and semideciduous species than interior forest. Edges lacked supra‐annual flowering and fruiting species and had more species and stems with drupes and small seeds than small forest fragments and forest interior areas. In an ordination of species similarity and life‐history traits, the three types of configurations formed clearly segregated clusters. Furthermore, the differences in the taxonomic and functional (i.e., trait‐based) composition of tree assemblages we documented were driven primarily by the higher abundance of pioneer species in the forest edge and small forest fragments. Our work provides strong evidence that long‐term transitions in phenology and seed and fruit morphology of tree functional groups are occurring in fragmented tropical forests. Our results also suggest that edge‐induced shifts in tree assemblages of tropical forests can be larger than previously documented.     

More asymmetric tree competition brings about more evapotranspiration and less runoff from the forest ecosystems: A simulation study

The present paper reports how stand size-structure dynamics due to competition between different-sized trees affect long-term forested water balance in Japanese cool-temperate planted stands (evergreen coniferous Cryptomeria japonica and deciduous coniferous Larix kaempferi stands) using a fully coupled multi-layered meteorological surface physics—terrestrial ecosystems model. The simulation captured the well-known annual variation in leaf area index (LAI) accurately with stand age in monocultured and even-aged stands; the occurrence of maximum LAI during the early growth stage and then a gradual decline followed by a steady state after the maximum LAI. The simulations also detected a high dependency of annual evapotranspiration (AETr) on LAI with stand age that is well known by prior observational researches. In the C. japonica (shade-tolerant late-successional species) stand, the relationship between annual net primary productivity of an individual tree (NPP_ind) and individual tree mass (w) changed from linear to a convex curve during self-thinning, indicating that the degree of asymmetric tree competition intensified with forest stand development. The higher degree of competitive asymmetry characterized by the convex-shaped NPPind-w relationship produced greater size inequality, i.e., the formation of trees stratified by height. Under such conditions, AETr and annual transpiration (ATr) were mainly regulated by larger trees. On the other hand, the NPPind-w relationships in the L. kaempferi (shade-intolerant early-successional species) stand were linear throughout the simulated period, indicating the lower degree of competitive asymmetry. Under such conditions, the growth of intermediate-sized trees was enhanced and these trees became a dominant source of AETr (and also ATr) during self-thinning. Furthermore, the sensitivity analysis of the effects of ecophysiological parameters such as foliage profile (i.e., vertical distribution of leaf area density) of an individual tree (distribution pattern is described by the parameter η), the maximum carboxylation velocity (V_cmax0) and biomass allocation pattern of individual plant growth (μ₁) on AETr, ATr and annual runoff (AR_off) showed that the temporal trends of AETr, ATr, AR_off and NPPind-w relationships were completely the same as those in the control simulations. However, the NPPind-w relationship during self-thinning indicated higher degrees of competitive asymmetry when η or V_cmax0 were greater than those in the control simulation and generated greater AETr and ATr and thus smaller AR_off. We found that more asymmetric tree competition brings about greater size inequality between different-sized trees and thus more evapotranspiration and less runoff in a forest stand. Overall, our simulation approach revealed that not only LAI dynamics but also plant competition, and thus size-structure dynamics, in a forest ecosystem are essential to long-term future projections of forested water balance.