Functional traits and the growth-mortality trade-off in tropical trees

A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 < or = 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (H(max)) are key traits among forest trees. Seed mass, LMA, WD, and H(max) are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r2 < 0.1 for three and r2 < 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for > 80% of the explained variation and, after WD was included, LMA and H(max) made insignificant contributions. Virtually the full range of values of SM, LMA, and H(max) occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.     

Are functional traits good predictors of demographic rates? Evidence from five neotropical forests

A central goal of comparative plant ecology is to understand how functional traits vary among species and to what extent this variation has adaptive value. Here we evaluate relationships between four functional traits (seed volume, specific leaf area, wood density, and adult stature) and two demographic attributes (diameter growth and tree mortality) for large trees of 240 tree species from five Neotropical forests. We evaluate how these key functional traits are related to survival and growth and whether similar relationships between traits and demography hold across different tropical forests. There was a tendency for a trade-off between growth and survival across rain forest tree species. Wood density, seed volume, and adult stature were significant predictors of growth and/or mortality. Both growth and mortality rates declined with an increase in wood density. This is consistent with greater construction costs and greater resistance to stem damage for denser wood. Growth and mortality rates also declined as seed volume increased. This is consistent with an adaptive syndrome in which species tolerant of low resource availability (in this case shade-tolerant species) have large seeds to establish successfully and low inherent growth and mortality rates. Growth increased and mortality decreased with an increase in adult stature, because taller species have a greater access to light and longer life spans. Specific leaf area was, surprisingly, only modestly informative for the performance of large trees and had ambiguous relationships with growth and survival. Single traits accounted for 9-55% of the interspecific variation in growth and mortality rates at individual sites. Significant correlations with demographic rates tended to be similar across forests and for phylogenetically independent contrasts as well as for cross-species analyses that treated each species as an independent observation. In combination, the morphological traits explained 41% of the variation in growth rate and 54% of the variation in mortality rate, with wood density being the best predictor of growth and mortality. Relationships between functional traits and demographic rates were statistically similar across a wide range of Neotropical forests. The consistency of these results strongly suggests that tropical rain forest species face similar trade-offs in different sites and converge on similar sets of solutions.     

Predicting the decomposition of Scots pine, Norway spruce, and birch stems in Finland.

Models were developed for predicting the decomposition of dead wood for the main tree species in Finland, based on data collected from long-term thinning experiments in southern and central Finland. The decomposition rates were strongly related to the number of years after tree death. In contrast to previous studies, which have used the first-order exponential model, we found that the decomposition rate was not constant. Therefore, the Gompertz and Chapman-Richard's functions were fitted to the data. The slow initial decomposition period was mainly due to the fact that most dead trees remained standing as snags after their death. The initial period was followed by a period of rapid decomposition and, finally, by a period of moderately slow decomposition. Birch stems decomposed more rapidly than Scots pine and Norway spruce stems. Decomposition rates of Norway spruce stems were somewhat lower than those of Scots pine. Because the carbon concentration of decaying boles was relatively stable (about 50%) the rate of carbon loss follows that of mass loss. Models were also developed for the probability that a dead tree remains standing as a snag. During the first years after death, the probability was high. Thereafter, it decreased rapidly, the decrease being faster for birch stems than for Scots pine and Norway spruce stems. Almost all stems had fallen down within 40 years after their death. In Scots pine and Norway spruce, most snags remained hard and belonged to decay class 1. In birch, a higher proportion of snags belonged to the more advanced decay classes. The models provide a framework for predicting dead wood dynamics in managed as well as dense unthinned stands. The models can be incorporated into forest management planning systems, thereby facilitating estimates of carbon dynamics.     

Mortality of large trees and lianas following experimental drought in an Amazon forest

Severe drought episodes such as those associated with El Ni?o Southern Oscillation (ENSO) events influence large areas of tropical forest and may become more frequent in the future. One of the most important forest responses to severe drought is tree mortality, which alters forest structure, composition, carbon content, and flammability, and which varies widely. This study tests the hypothesis that tree mortality increases abruptly during drought episodes when plant-available soil water (PAW) declines below a critical minimum threshold. It also examines the effect of tree size, plant life form (palm, liana, tree) and potential canopy position (understory, midcanopy, overstory) on drought-induced plant mortality. A severe, four-year drought episode was simulated by excluding 60% of incoming throughfall during each wet season using plastic panels installed in the understory of a 1-ha forest treatment plot, while a 1-ha control plot received normal rainfall. After 3.2 years, the treatment resulted in a 38% increase in mortality rates across all stems >2 cm dbh. Mortality rates increased 4.5-fold among large trees (>30 cm dbh) and twofold among medium trees (10-30 cm dbh) in response to the treatment, whereas the smallest stems were less responsive. Recruitment rates did not compensate for the elevated mortality of larger-diameter stems in the treatment plot. Overall, lianas proved more susceptible to drought-induced mortality than trees or palms, and potential overstory tree species were more vulnerable than midcanopy and understory species. Large stems contributed to 90% of the pretreatment live aboveground biomass in both plots. Large-tree mortality resulting from the treatment generated 3.4 times more dead biomass than the control plot. The dramatic mortality response suggests significant, adverse impacts on the global carbon cycle if climatic changes follow current trends.     

Leaf herbivory and decomposability in a Malaysian tropical rain forest

There is accumulating evidence that similar suites of plant traits may affect leaf palatability and leaf litter decomposability. However, the possible association between leaf herbivory and litter decomposition rates across species in species-diverse natural ecosystems such as tropical rain forests remains unexplored, despite its importance in estimating the herbivory effects on carbon and nutrient cycling of ecosystems. We found no strong association between leaf herbivory and litter decomposition rates across 40 tree species in a Malaysian tropical rain forest, even though the leaf and litter traits were tightly correlated. This is because the leaf and litter traits related to herbivory and decomposition rates in the field were inconsistent. Leaf toughness accounted for only a small part of the variation in the herbivory rate, whereas a number of litter traits (the leaf mass per area, lignin to nitrogen ratio, and condensed tannin concentration) accurately predicted the decomposition rate across species. These results suggest that herbivory rate across species may not be strongly related to single leaf traits, probably because plant-herbivore interactions in tropical rain forests are highly diverse; on the other hand, plant-decomposer interactions are less specific and can be governed by litter chemicals. We also investigated two factors, phylogeny and tree functional types, that could affect the relationship between herbivory and decomposition across species. Phylogenetic relatedness among the species did not affect the relationship between herbivory and decomposition. In contrast, when the plants were segregated according to their leaf emergence pattern, we found a significant positive relationship between herbivory and decomposition rates for continuous-leafing species. In these species, the condensed tannin to N ratios in leaves and litter were related to herbivory and decomposition rates, respectively. However, we did not observe a similar trend for synchronous-leafing species. These results suggest that the relationship between herbivory and decomposition may be more greatly affected by functional types than by phylogenetic relatedness among species. In conclusion, our results suggest that well-defended leaves are not necessarily less decomposable litter in a tropical rain forest community, implying that herbivory may not generate positive feedback for carbon and nutrient cycling in this type of ecosystem.     

Current Near‐to‐Nature Forest Management Effects on Functional Trait Composition of Saproxylic Beetles in Beech Forests

With the aim of wood production with negligible negative effects on biodiversity and ecosystem processes, a silvicultural practice of selective logging with natural regeneration has been implemented in European beech forests (Fagus sylvatica) during the last decades. Despite this near‐to‐nature strategy, species richness of various taxa is lower in these forests than in unmanaged forests. To develop guidelines to minimize the fundamental weaknesses in the current practice, we linked functional traits of saproxylic beetle species to ecosystem characteristics. We used continental‐scale data from 8 European countries and regional‐scale data from a large forest in southern Germany and forest‐stand variables that represented a gradient of intensity of forest use to evaluate the effect of current near‐to‐nature management strategies on the functional diversity of saproxylic beetles. Forest‐stand variables did not have a statistically significant effect on overall functional diversity, but they did significantly affect community mean and diversity of single functional traits. As the amount of dead wood increased the composition of assemblages shifted toward dominance of larger species and species preferring dead wood of large diameter and in advanced stages of decay. The mean amount of dead wood across plots in which most species occurred was from 20 to 60 m³/ha. Species occurring in plots with mean dead wood >60 m³/ha were consistently those inhabiting dead wood of large diameter and in advanced stages of decay. On the basis of our results, to make current wood‐production practices in beech forests throughout Europe more conservation oriented (i.e., promoting biodiversity and ecosystem functioning), we recommend increasing the amount of dead wood to >20 m³/ha; not removing dead wood of large diameter (50 cm) and allowing more dead wood in advanced stages of decomposition to develop; and designating strict forest reserves, with their exceptionally high amounts of dead wood, that would serve as refuges for and sources of saproxylic habitat specialists. Efectos Actuales del Manejo Casi Natural de Bosques sobre la Composición de Atributos Funcionales de Escarabajos Saproxílicos en Bosques de Haya     

Seasonal water stress tolerance and habitat associations within four neotropical tree genera

We investigated the relationship between habitat association and physiological performance in four congeneric species pairs exhibiting contrasting distributions between seasonally flooded and terra firme habitats in lowland tropical rain forests of French Guiana, including Virola and Iryanthera (Myristicaceae), Symphonia (Clusiaceae), and Eperua (Caesalpiniaceae). We analyzed 10-year data sets of mapped and measured saplings (stems >150 cm in height and <10 cm diameter at breast height [dbh]) and trees (stems > or =10 cm dbh) across 37.5 ha of permanent plots covering a 300-ha zone, within which seasonally flooded areas (where the water table never descends below 1 m) have been mapped. Additionally, we tested the response of growth, survival, and leaf functional traits of these species to drought and flood stress in a controlled experiment. We tested for habitat preference using a modification of the torus translation method. Strong contrasting associations of the species pairs of Iryanthera, Virola, and Symphonia were observed at the sapling stage, and these associations strengthened for the tree stage. Neither species of Eperua was significantly associated with flooded habitats at the sapling stage, but E. falcata was significantly and positively associated with flooded forests at the tree stage, and trees of E. grandiflora were found almost exclusively in nonflooded habitats. Differential performance provided limited explanatory support for the observed habitat associations, with only congeners of Iryanthera exhibiting divergent sapling survival and tree growth. Seedlings of species associated with flooded forest tended to have higher photosynthetic capacity than their congeners at field capacity. In addition, they tended to have the largest reductions in leaf gas exchange and growth rate in response to experimental drought stress and the least reductions in response to experimental inundation. The corroboration of habitat association with differences in functional traits and, to a lesser extent, measures of performance provides an explanation for the regional coexistence of these species pairs. We suggest that specialization to seasonally flooded habitats may explain patterns of adaptive radiation in many tropical tree genera and thereby provide a substantial contribution to regional tree diversity.     

Tree-Dependent Lichens and Beetles as Indicators in Conservation Forests

We tested whether the conspicuous lichen Lobaria pulmonaria indicates the number of tree-dependent, red-listed species in a hemiboreal forest in southern Sweden. In 18 naturally regenerated, mainly old deciduous forest plots considered to be of high or very high conservation value, the number of red-listed tree lichens or wood beetles was not positively correlated with the area of the forest stands studied (8–56 ha). The 8 stands with L. pulmonaria had about nine (median) red-listed lichens, but 10 stands without L. pulmonaria had only about one such species, a highly significant difference. Similarly, the variation between stands in the number of red-listed wood beetles was considerable, but it was very weakly correlated with the number of red-listed lichens. The number of red-listed wood beetles dependent on dead trees was not different in areas with or without Lobaria pulmonaria , but the number of red-listed wood beetles dependent on hollow trees was higher in stands with Lobaria pulmonaria (median of seven species) than in those without (three species) the lichen. Stands with this lichen species also contained significantly more of other types of lichens that have been proposed as indicators of forest continuity according to three lists. We stress the need for identification of species that could serve as indicators of different types of forest continuity and identify some organism groups that may indicate the different types.     

Wood Density as a Conservation Tool: Quantification of Disturbance and Identification of Conservation‐Priority Areas in Tropical Forests

Abstract: Inventories of tree species are often conducted to guide conservation efforts in tropical forests. Such surveys are time consuming, demanding of expertise, and expensive to perform and interpret. Approaches to make survey efforts simpler or more effective would be valuable. In particular, it would be good to be able to easily identify areas of old‐growth forest. The average density of the wood of a tree species is closely linked to its successional status. We used tree inventory data from eastern Borneo to determine whether wood density can be used to quantify forest disturbance and conservation importance. The average density of wood in a plot was significantly and negatively related to disturbance levels, with plots with higher wood densities occurring almost exclusively in old‐growth forests. Average wood density was unimodally related to the diversity of tree species, indicating that the average wood density in a plot might be a better indicator of old‐growth forest than species diversity. In addition, Borneo endemics had significantly heavier wood than species that are common throughout the Malesian region, and they were more common in plots with higher average wood density. We concluded that wood density at the plot level could be a powerful tool for identifying areas of conservation priority in the tropical rain forests of Southeast Asia.     

Riparian forest composition affects stream litter decomposition despite similar microbial and invertebrate communities

Cross-boundary flows of energy and nutrients link biodiversity and functioning in adjacent ecosystems. The composition of forest tree species can affect the structure and functioning of stream ecosystems due to physical and chemical attributes, as well as changes in terrestrial resource subsidies. We examined how variation in riparian canopy composition (coniferous, deciduous, mixed) affects adjacent trophic levels (invertebrate and microbial consumers) and decomposition of organic matter in small, coastal rainforest streams in southwestern British Columbia. Breakdown rates of higher-quality red alder (Alnus rubra) litter were faster in streams with a greater percentage of deciduous than coniferous riparian canopy, whereas breakdown rates of lower-quality western hemlock (Tsuga heterophylla) litter were independent of riparian forest composition. When invertebrates were excluded using fine mesh, breakdown rates of both litter species were an order of magnitude less and were not significantly affected by riparian forest composition. Stream invertebrate and microbial communities were similar among riparian forest composition, with most variation attributed to leaf litter species. Invertebrate taxa richness and shredder biomass were higher in A. rubra litter; however, taxa evenness was greatest for T. heterophylla litter and both litter species in coniferous streams. Microbial community diversity (determined from terminal restriction fragment length polymorphisms) was unaffected by riparian forest or litter species. Fungal allele richness was higher than bacterial allele richness, and microbial communities associated with lower-quality T. heterophylla litter had higher diversity (allele uniqueness and richness) than those associated with higher-quality A. rubra litter. Percent variation in breakdown rates was mostly attributed to riparian forest composition in the presence of invertebrates and microbes; however, stream consumer biodiversity at adjacent trophic levels did not explain these patterns. Riparian and stream ecosystems and their biotic communities are linked through exchange and decomposition of detrital resources, and we provide evidence that riparian forest composition affects stream ecosystem catabolism despite similarities in microbial and invertebrate communities.     

Association of extinction risk of saproxylic beetles with ecological degradation of forests in Europe

To reduce future loss of biodiversity and to allocate conservation funds effectively, the major drivers behind large‐scale extinction processes must be identified. A promising approach is to link the red‐list status of species and specific traits that connect species of functionally important taxa or guilds to resources they rely on. Such traits can be used to detect the influence of anthropogenic ecosystem changes and conservation efforts on species, which allows for practical recommendations for conservation. We modeled the German Red List categories as an ordinal index of extinction risk of 1025 saproxylic beetles with a proportional‐odds linear mixed‐effects model for ordered categorical responses. In this model, we estimated fixed effects for intrinsic traits characterizing species biology, required resources, and distribution with phylogenetically correlated random intercepts. The model also allowed predictions of extinction risk for species with no red‐list category. Our model revealed a higher extinction risk for lowland and large species as well as for species that rely on wood of large diameter, broad‐leaved trees, or open canopy. These results mirror well the ecological degradation of European forests over the last centuries caused by modern forestry, that is the conversion of natural broad‐leaved forests to dense conifer‐dominated forests and the loss of old growth and dead wood. Therefore, conservation activities aimed at saproxylic beetles in all types of forests in Central and Western Europe should focus on lowlands, and habitat management of forest stands should aim at increasing the amount of dead wood of large diameter, dead wood of broad‐leaved trees, and dead wood in sunny areas.     

Neighborhood analyses of canopy tree competition along environmental gradients in New England forests.

We use permanent-plot data from the USDA Forest Service's Forest Inventory and Analysis (FIA) program for an analysis of the effects of competition on tree growth along environmental gradients for the 14 most abundant tree species in forests of northern New England, USA. Our analysis estimates actual growth for each individual tree of a given species as a function of average potential diameter growth modified by three sets of scalars that quantify the effects on growth of (1) initial target tree size (dbh), (2) local environmental conditions, and (3) crowding by neighboring trees. Potential growth of seven of the 14 species varied along at least one of the two environmental axes identified by an ordination of relative abundance of species in plots. The relative abundances of a number of species were significantly displaced from sites where they showed maximum potential growth. In all of these cases, abundance was displaced to the more resource-poor end of the environmental gradient (either low fertility or low moisture). The pattern was most pronounced among early successional species, whereas late-successional species reached their greatest abundance on sites where they also showed the highest growth in the absence of competition. The analysis also provides empirical estimates of the strength of intraspecific and interspecific competitive effects of neighbors. For all but one of the species, our results led us to reject the hypothesis that all species of competitors have equivalent effects on a target species. Most of the individual pairwise interactions were strongly asymmetric. There was a clear competitive hierarchy among the four most shade-tolerant species, and a separate competitive hierarchy among the shade-intolerant species. Our results suggest that timber yield following selective logging will vary dramatically depending on the configuration of the residual canopy, because of interspecific variation in the magnitude of both the competitive effects of different species of neighbors and the competitive responses of different species of target trees to neighbors. The matrix of competition coefficients suggests that there may be clear benefits in managing for specific mixtures of species within local neighborhoods within stands.     

Leaf structural diversity is related to hydraulic capacity in tropical rain forest trees

The hydraulic resistance of the leaf (R1) is a major bottleneck in the whole plant water transport pathway and may thus be linked with the enormous variation in leaf structure and function among tropical rain forest trees. A previous study found that R1 varied by an order of magnitude across 10 tree species of Panamanian tropical lowland rain forest. Here, correlations were tested between R1 and 24 traits relating to leaf venation and mesophyll structure, and to gross leaf form. Across species, R1 was related to both venation architecture and mesophyll structure. R1 was positively related to the theoretical axial resistivity of the midrib, determined from xylem conduit numbers and dimensions, and R1 was negatively related to venation density in nine of 10 species. R1 was also negatively related to both palisade mesophyll thickness and to the ratio of palisade to spongy mesophyll. By contrast, numerous leaf traits were independent of R1, including area, shape, thickness, and density, demonstrating that leaves can be diverse in gross structure without intrinsic trade-offs in hydraulic capacity. Variation in both R1-linked and R1-independent traits related strongly to regeneration irradiance, indicating the potential importance of both types of traits in establishment ecology.     

Tree species effects on decomposition and forest floor dynamics in a common garden

We studied the effects of tree species on leaf litter decomposition and forest floor dynamics in a common garden experiment of 14 tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Betula pendula, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pinus nigra, Pinus sylvestris, Pseudotsuga menziesii, Quercus robur, Quercus rubra, and Tilia cordata) in southwestern Poland. We used three simultaneous litter bag experiments to tease apart species effects on decomposition via leaf litter chemistry vs. effects on the decomposition environment. Decomposition rates of litter in its plot of origin were negatively correlated with litter lignin and positively correlated with mean annual soil temperature (MAT(soil)) across species. Likewise, decomposition of a common litter type across all plots was positively associated with MAT(soil), and decomposition of litter from all plots in a common plot was negatively related to litter lignin but positively related to litter Ca. Taken together, these results indicate that tree species influenced microbial decomposition primarily via differences in litter lignin (and secondarily, via differences in litter Ca), with high-lignin (and low-Ca) species decomposing most slowly, and by affecting MAT(soil), with warmer plots exhibiting more rapid decomposition. In addition to litter bag experiments, we examined forest floor dynamics in each plot by mass balance, since earthworms were a known component of these forest stands and their access to litter in litter bags was limited. Forest floor removal rates estimated from mass balance were positively related to leaf litter Ca (and unrelated to decay rates obtained using litter bags). Litter Ca, in turn, was positively related to the abundance of earthworms, particularly Lumbricus terrestris. Thus, while species influence microbially mediated decomposition primarily through differences in litter lignin, differences among species in litter Ca are most important in determining species effects on forest floor leaf litter dynamics among these 14 tree species, apparently because of the influence of litter Ca on earthworm activity. The overall influence of these tree species on leaf litter decomposition via effects on both microbial and faunal processing will only become clear when we can quantify the decay dynamics of litter that is translocated belowground by earthworms.     

Socio-economic predictors of domestic wood use in an Atlantic forest area (north-east Brazil): a tool for directing conservation efforts

Given the excessive use pressure on forests, characterisation of the major users of forest resources can help guide conservation efforts. This study examines whether socio-economic variables explain domestic wood use in an area of Atlantic forest in the community of Três Ladeiras (Pernambuco, north-east Brazil). Socio-economic data were collected from 62 households, and geometric measurements of wood products in each household were taken to calculate the wood volume. An informant in each household indicated the average replacement time for each type of wood use. Data were analysed for static volume (wood volume found at the time of the visit) and dynamic volume (rate of wood consumption, calculated as the ratio between volume and replacement time). Multiple regression analyses were performed to determine the relationships between wood volume (static and dynamic) and the number of species used; independent variables including monthly family income, number of residents, average age of heads of the household, age of oldest household member, average residence time of heads of the household, average education level of heads of the household and education level of the most educated household member. Monthly income was inversely related to the static wood volume (adjusted R ² = 17.6%, p < 0.001), rate of wood consumption (adjusted R ² = 23.0%, p < 0.001) and number of species used (adjusted R ² = 22.3%, p < 0.001) and was the most important predictive variable for these three outcomes. Other factors such as average education level and average age of heads of the household were also explanatory variables of the total rate of wood consumption, as these two variables, together with monthly family income, explain 31.3% of the total dynamic volume. Given the high importance of income in our study, we suggest that future conservation strategies give special attention to the poorest members of the community.     

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.     

Tree species control rates of free-living nitrogen fixation in a tropical rain forest

Tropical rain forests represent some of the most diverse ecosystems on earth, yet mechanistic links between tree species identity and ecosystem function in these forests remains poorly understood. Here, using free-living nitrogen (N) fixation as a model, we explore the idea that interspecies variation in canopy nutrient concentrations may drive significant local-scale variation in biogeochemical processes. Biological N fixation is the largest "natural" source of newly available N to terrestrial ecosystems, and estimates suggest the highest such inputs occur in tropical ecosystems. While patterns of and controls over N fixation in these systems remain poorly known, the data we do have suggest that chemical differences among tree species canopies could affect free-living N fixation rates. In a diverse lowland rain forest in Costa Rica, we established a series of vertical, canopy-to-soil profiles for six common canopy tree species, and we measured free-living N fixation rates and multiple aspects of chemistry of live canopy leaves, senesced canopy leaves, bulk leaf litter, and soil for eight individuals of each tree species. Free-living N fixation rates varied significantly among tree species for all four components, and independent of species identity, rates of N fixation ranged by orders of magnitude along the vertical profile. Our data suggest that variations in phosphorus (P) concentration drove a significant fraction of the observed species-specific variation in free-living N fixation rates within each layer of the vertical profile. Furthermore, our data suggest significant links between canopy and forest floor nutrient concentrations; canopy P was correlated with bulk leaf litter P below individual tree crowns. Thus, canopy chemistry may affect a suite of ecosystem processes not only within the canopy itself, but at and beneath the forest floor as well.     

Fungi and wind strongly influence the temporal availability of logs in an old-growth spruce forest.

Coarse woody debris (CWD) is a key habitat for many species in forest ecosystems. To ensure the long-term survival of such species, forest management regimes must include measures that promote dead wood dynamics similar to those of natural forests. Thus, information on CWD dynamics under natural conditions is required, including data pertaining to the underlying agents of disturbance. This study examines modes of mortality, decay rates, and temporal patterns in the availability of Picea abies logs in a Swedish old-growth forest affected by internal, small-scale disturbance. All 684 logs in a 6.6-ha plot were mapped and classified into one of six decay classes. Logs in the early stages of decay were examined for the presence of heart-rot fungi. Six years later all logs were re-inventoried, including newly formed logs. Matrix models based on the transition rates between decay classes showed that it took about 60 years for 90% of the logs to decay beyond class 6 (a deformed trunk with soft wood). Large logs (> 26 cm) decayed 40% more slowly than small logs (< 25 cm). The initial volume of logs was 37.6 m3/ha but increased to 44.8 m3/ha after six years. In addition, there was a large shift in the decay-class distribution. The volume of logs in early and late decay classes increased by 71% and 45%, respectively, while the volume of logs in the intermediate decay classes decreased by 32%. The fluctuations appear to result from pulses in mortality, driven by a combination of strong winds and the heart-rot fungus, Phellinus chrysoloma, which was present in more than 30% of all logs at an early stage of decay. These results show that large temporal fluctuations in dead wood also occur in the absence of large-scale disturbance, and that heart-rot fungi are important factors driving the overall dynamics of dead wood. Since many wood-inhabiting species are naturally rare and have very specific substrate demands, such temporal variability in dead wood availability may have effects on biodiversity and should be taken into account when designing small, protected forest areas.     

Contrasting structure and composition of the understory in species-rich tropical rain forests

In large samples of trees > or = 1 cm dbh (more than 1 million trees and 3000 species), in six lowland tropical forests on three continents, we assigned species with >30 individuals to one of six classes of stature at maturity (SAM). We then compared the proportional representation of understory trees (1-2 cm dbh) among these classes. The understory of the three Asian sites was predominantly composed of the saplings of large-canopy trees whereas the African and American sites were more richly stocked with trees of the smaller SAM classes. Differences in class representation were related to taxonomic families that were present exclusively in one continent or another. Families found in the Asian plots but not in the American plot (e.g., Dipterocarpaceae, Fagaceae) were predominantly species of the largest SAM classes, whereas families exclusive to the American plots (e.g., Melastomataceae sensu stricto, Piperaceae, and Malvaceae [Bombacacoidea]) were predominantly species of small classes. The African plot was similar to Asia in the absence of those American families rich in understory species, while similar to America in lacking the Asian families rich in canopy species. The numerous understory species of Africa were chiefly derived from families shared with Asia and/or America. The ratio of saplings (1-2 cm dbh) to conspecific canopy trees (>40 cm dbh) was lower in American plots than in the Asian plots. Possible explanations for these differences include phenology, moisture and soil fertility regimes, phyletic constraints, and the role of early successional plants in forest development. These results demonstrate that tropical forests that appear similar in tree number, basal area, and the family taxonomy of canopy trees nonetheless differ in ecological structure in ways that may impact the ecology of pollinators, dispersers, and herbivores and might reflect fundamental differences in canopy tree regeneration.     

Decomposition of diverse litter mixtures in streams

In view of growing interest in understanding how biodiversity affects ecosystem functioning, we investigated effects of riparian plant diversity on litter decomposition in forest streams. Leaf litter from 10 deciduous tree species was collected during natural leaf fall at two locations (Massif Central in France and Carpathians in Romania) and exposed in the field in litter bags. There were 35 species combinations, with species richness ranging 1-10. Nonadditive effects on the decomposition of mixed-species litter were minor, although a small synergistic effect was observed in the Massif Central stream where observed litter mass remaining was significantly lower overall than expected from data on single-species litter. In addition, variability in litter mass remaining decreased with litter diversity at both locations. Mean nitrogen concentration of single- and mixed-species litters (0.68-4.47% of litter ash-free dry mass) accounted for a large part of the variation in litter mass loss across species combinations. For a given species or mixture, litter mass loss was also consistently faster in the Massif Central than in the Carpathians, and the similarity in general stream characteristics, other than temperature, suggests that this effect was largely due to differences in thermal regimes. These results support the notion that decomposition of litter mixtures is primarily driven by litter quality and environmental factors, rather than by species richness per se. However, the observed consistent decrease in variability of decomposition rate with increasing plant species richness indicates that conservation of riparian tree diversity is important even when decomposition rates are not greatly influenced by litter mixing.