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When Darwin visited the Galapagos archipelago, he observed that, in spite of the islands’ physical similarity, members of species that had dispersed to them recently were beginning to diverge from each other. He postulated that these divergences must have resulted primarily from interactions with sets of other species that had also diverged across these otherwise similar islands. By extrapolation, if Darwin is correct, such complex interactions must be driving species divergences across all ecosystems. However, many current general ecological theories that predict observed distributions of species in ecosystems do not take the details of between-species interactions into account. Here we quantify, in sixteen forest diversity plots (FDPs) worldwide, highly significant negative density-dependent (NDD) components of both conspecific and heterospecific between-tree interactions that affect the trees’ distributions, growth, recruitment, and mortality. These interactions decline smoothly in significance with increasing physical distance between trees. They also tend to decline in significance with increasing phylogenetic distance between the trees, but each FDP exhibits its own unique pattern of exceptions to this overall decline. Unique patterns of between-species interactions in ecosystems, of the general type that Darwin postulated, are likely to have contributed to the exceptions. We test the power of our null-model method by using a deliberately modified data set, and show that the method easily identifies the modifications. We examine how some of the exceptions, at the Wind River (USA) FDP, reveal new details of a known allelopathic effect of one of the Wind River gymnosperm species. Finally, we explore how similar analyses can be used to investigate details of many types of interactions in these complex ecosystems, and can provide clues to the evolution of these interactions.

When Darwin visited the Galapagos archipelago, he observed that, in spite of the islands’ physical similarity, members of species that had dispersed to them recently were beginning to diverge from each other. He postulated that these divergences must have resulted primarily from interactions with sets of other species that had also diverged across these otherwise similar islands. By extrapolation, if Darwin is correct, such complex interactions must be driving species divergences across all ecosystems. However, many current general ecological theories that predict observed distributions of species in ecosystems do not take the details of between-species interactions into account. Here we quantify, in sixteen forest diversity plots (FDPs) worldwide, highly significant negative density-dependent (NDD) components of both conspecific and heterospecific between-tree interactions that affect the trees’ distributions, growth, recruitment, and mortality. These interactions decline smoothly in significance with increasing physical distance between trees. They also tend to decline in significance with increasing phylogenetic distance between the trees, but each FDP exhibits its own unique pattern of exceptions to this overall decline. Unique patterns of between-species interactions in ecosystems, of the general type that Darwin postulated, are likely to have contributed to the exceptions. We test the power of our null-model method by using a deliberately modified data set, and show that the method easily identifies the modifications. We examine how some of the exceptions, at the Wind River (USA) FDP, reveal new details of a known allelopathic effect of one of the Wind River gymnosperm species. Finally, we explore how similar analyses can be used to investigate details of many types of interactions in these complex ecosystems, and can provide clues to the evolution of these interactions.

Studies of forest dynamics plots (FDPs) have revealed a variety of negative density-dependent (NDD) demographic interactions, especially among conspecific trees. These interactions can affect growth rate, recruitment and mortality, and they play a central role in the maintenance of species diversity in these complex ecosystems. Here we use an equal area annulus (EAA) point-pattern method to comprehensively analyze data from two tropical FDPs, Barro Colorado Island in Panama and Sinharaja in Sri Lanka. We show that these NDD interactions also influence the continued evolutionary diversification of even distantly related tree species in these FDPs. We examine the details of a wide range of these interactions between individual trees and the trees that surround them. All these interactions, and their cumulative effects, are strongest among conspecific focal and surrounding tree species in both FDPs. They diminish in magnitude with increasing phylogenetic distance between heterospecific focal and surrounding trees, but do not disappear or change the pattern of their dependence on size, density, frequency or physical distance even among the most distantly related trees. The phylogenetic persistence of all these effects provides evidence that interactions between tree species that share an ecosystem may continue to promote adaptive divergence even after the species' gene pools have become separated. Adaptive divergence among taxa would operate in stark contrast to an alternative possibility that has previously been suggested, that distantly related species with dispersal-limited distributions and confronted with unpredictable neighbors will tend to converge on common strategies of resource use. In addition, we have also uncovered a positive density-dependent effect: growth rates of large trees are boosted in the presence of a smaller basal area of surrounding trees. We also show that many of the NDD interactions switch sign rapidly as focal trees grow in size, and that th

Studies of forest dynamics plots (FDPs) have revealed a variety of negative density-dependent (NDD) demographic interactions, especially among conspecific trees. These interactions can affect growth rate, recruitment and mortality, and they play a central role in the maintenance of species diversity in these complex ecosystems. Here we use an equal area annulus (EAA) point-pattern method to comprehensively analyze data from two tropical FDPs, Barro Colorado Island in Panama and Sinharaja in Sri Lanka. We show that these NDD interactions also influence the continued evolutionary diversification of even distantly related tree species in these FDPs. We examine the details of a wide range of these interactions between individual trees and the trees that surround them. All these interactions, and their cumulative effects, are strongest among conspecific focal and surrounding tree species in both FDPs. They diminish in magnitude with increasing phylogenetic distance between heterospecific focal and surrounding trees, but do not disappear or change the pattern of their dependence on size, density, frequency or physical distance even among the most distantly related trees. The phylogenetic persistence of all these effects provides evidence that interactions between tree species that share an ecosystem may continue to promote adaptive divergence even after the species' gene pools have become separated. Adaptive divergence among taxa would operate in stark contrast to an alternative possibility that has previously been suggested, that distantly related species with dispersal-limited distributions and confronted with unpredictable neighbors will tend to converge on common strategies of resource use. In addition, we have also uncovered a positive density-dependent effect: growth rates of large trees are boosted in the presence of a smaller basal area of surrounding trees. We also show that many of the NDD interactions switch sign rapidly as focal trees grow in size, and that th

Studies of forest dynamics plots (FDPs) have revealed a variety of negative density-dependent (NDD) demographic interactions, especially among conspecific trees. These interactions can affect growth rate, recruitment and mortality, and they play a central role in the maintenance of species diversity in these complex ecosystems. Here we use an equal area annulus (EAA) point-pattern method to comprehensively analyze data from two tropical FDPs, Barro Colorado Island in Panama and Sinharaja in Sri Lanka. We show that these NDD interactions also influence the continued evolutionary diversification of even distantly related tree species in these FDPs. We examine the details of a wide range of these interactions between individual trees and the trees that surround them. All these interactions, and their cumulative effects, are strongest among conspecific focal and surrounding tree species in both FDPs. They diminish in magnitude with increasing phylogenetic distance between heterospecific focal and surrounding trees, but do not disappear or change the pattern of their dependence on size, density, frequency or physical distance even among the most distantly related trees. The phylogenetic persistence of all these effects provides evidence that interactions between tree species that share an ecosystem may continue to promote adaptive divergence even after the species' gene pools have become separated. Adaptive divergence among taxa would operate in stark contrast to an alternative possibility that has previously been suggested, that distantly related species with dispersal-limited distributions and confronted with unpredictable neighbors will tend to converge on common strategies of resource use. In addition, we have also uncovered a positive density-dependent effect: growth rates of large trees are boosted in the presence of a smaller basal area of surrounding trees. We also show that many of the NDD interactions switch sign rapidly as focal trees grow in size, and that th

* Successional gradients are ubiquitous in nature, yet few studies have systematically examined the evolutionary origins of taxa that specialize at different successional stages. Here we quantify successional habitat specialization in Neotropical forest trees and evaluate its evolutionary lability along a precipitation gradient. Theoretically, successional habitat specialization should be more evolutionarily conserved in wet forests than in dry forests due to more extreme microenvironmental differentiation between early and late-successional stages in wet forest. * We applied a robust multinomial classification model to samples of primary and secondary forest trees from 14 Neotropical lowland forest sites spanning a precipitation gradient from 788 to 4000 mm annual rainfall, identifying species that are old-growth specialists and secondary forest specialists in each site. We constructed phylogenies for the classified taxa at each site and for the entire set of classified taxa and tested whether successional habitat specialization is phylogenetically conserved. We further investigated differences in the functional traits of species specializing in secondary vs. old-growth forest along the precipitation gradient, expecting different trait associations with secondary forest specialists in wet vs. dry forests since water availability is more limiting in dry forests and light availability more limiting in wet forests. * Successional habitat specialization is non-randomly distributed in the angiosperm phylogeny, with a tendency towards phylogenetic conservatism overall and a trend towards stronger conservatism in wet forests than in dry forests. However, the specialists come from all the major branches of the angiosperm phylogeny, and very few functional traits showed any consistent relationships with successional habitat specialization in either wet or dry forests. * Synthesis. The niche conservatism evident in the habitat specialization of Neotropical trees suggests a r

Collection methods and/or habitats sampled influence how many and which species are captured during entomological surveys. Here we compare Coleoptera catches among three survey activities, each using a single collection method, at the same study sites in Great Smoky Mountains National Park, USA. Activities included: short-term flight intercept trapping (FITs); sifting/Berlese funneling of leaf litter and extremely decayed downed coarse woody debris; and using emergence chambers containing coarse woody debris of various decay classes. In total, 2472 adult beetle specimens, representing 217 lowest identifiable taxa within 164 genera and 42 families, were collected during the FIT survey. Each survey activity yielded more than 2000 specimens, and a combined total of 413 species was collected. A combination of all surveys yielded the highest species richness when normalized for number of specimens indicating that variation of habitat and/or collection method significantly increases species richness. Of single surveys the FIT survey had the highest absolute species richness (217) and the highest richness when normalized for number of specimens. Species overlap among survey activities was low (Sorensen’s quotient of similarity was 0.20–0.27), which showed that each was about equally dissimilar from all others. Overlap of catch between FITs and emergence chambers was too low to justify substitution of emergence surveys with the FIT survey protocol used when attempting to collect saproxylic Coleoptera.

Coleoptera species composition and succession in downed woody debris habitats are poorly known in eastern North America. A photoeclector emergence chamber was used to concentrate Coleoptera that emerged from various decay classes of fine and coarse woody debris (FWD and CWD, respectively) collected in primary and secondary forest sites in Great Smoky Mountains National Park, Tennessee, USA. A total of 5673 adult beetle specimens, representing 305 lowest identifiable taxa within 227 genera and 51 families, was collected. One hundred fifteen species (38%) were represented by single individuals. Many more specimens and species were collected from CWD (4129 and 247, respectively) than from FWD (1544 and 162, respectively), but species richness accumulation curves were not significantly different. Many more specimens but an equal number of species were collected from primary forest (3347 and 207, respectively) than from secondary forest (2326 and 207, respectively). Species accumulation curves indicated higher richness in secondary forests. Based on a subset of 71 species represented by 10 or more specimens, 27 species were associated with fresh fine woody debris, 11 species with weathered fine woody debris, four with coarse woody debris decay class I, 14 with coarse woody debris decay class II, and eight with coarse woody debris decay class III–IV. Sixteen species were associated with secondary forests, whereas 28 species were associated with primary forests. Coarse woody debris decay class II taken in primary forests had highest absolute species richness with 156 species. In coarse woody debris species overlap decreased with increased difference in decay indicating faunal succession. Published works related to the study of the ecology of downed woody material are briefly summarized. Recommendations on developing a database of legacy trees for future researchers are given. Notes on the biology and photographs of the 71 species represented by 10 or more specimens are giv

The community within extremely decayed downed coarse woody debris, here referred to as decay class V (CWD5), has never been systematically sampled. The presumption has been that rotten wood is eventually overrun by surrounding soil and litter inhabitants. Leaf litter and CWD5 were sampled for Coleoptera with a sifting/ Berlese technique at three primary and three secondary forest sites in Great Smoky Mountains National Park, Tennessee, USA, during fall 2006 and spring 2007. A total of 4261 adult beetle specimens, representing 216 lowest identifiable taxa within 159 genera and 28 families, were collected. Sixty-six species (31%) were represented by single individuals. Many more specimens (3471) and species (170) were collected from leaf litter than from CWD5 (790 and 111, respectively) but species accumulation curves showed that species richness was not significantly different between the two habitats. Eight species were significantly associated with CWD5, and 40 species were significantly associated with leaf litter. Species richness was significantly higher in secondary forest than primary forest, but more species were significantly associated with primary than secondary forest. Species richness was significantly higher in spring than fall. Notes on the biology and photographs of the 59 species represented by 10 or more specimens are given to provide an atlas of common eastern U.S. beetle species found in these habitats. Overall CWD5 is a distinct but overlooked habitat that may harbor numerous undescribed species or species considered rare.

Collection methods and/or habitats sampled influence how many and which species are captured during entomological surveys. Here we compare Coleoptera catches among three survey activities, each using a single collection method, at the same study sites in Great Smoky Mountains National Park, USA. Activities included: short-term flight intercept trapping (FITs); sifting/Berlese funneling of leaf litter and extremely decayed downed coarse woody debris; and using emergence chambers containing coarse woody debris of various decay classes. In total, 2472 adult beetle specimens, representing 217 lowest identifiable taxa within 164 genera and 42 families, were collected during the FIT survey. Each survey activity yielded more than 2000 specimens, and a combined total of 413 species was collected. A combination of all surveys yielded the highest species richness when normalized for number of specimens indicating that variation of habitat and/or collection method significantly increases species richness. Of single surveys the FIT survey had the highest absolute species richness (217) and the highest richness when normalized for number of specimens. Species overlap among survey activities was low (Sorensen’s quotient of similarity was 0.20–0.27), which showed that each was about equally dissimilar from all others. Overlap of catch between FITs and emergence chambers was too low to justify substitution of emergence surveys with the FIT survey protocol used when attempting to collect saproxylic Coleoptera.

The community within extremely decayed downed coarse woody debris, here referred to as decay class V (CWD5), has never been systematically sampled. The presumption has been that rotten wood is eventually overrun by surrounding soil and litter inhabitants. Leaf litter and CWD5 were sampled for Coleoptera with a sifting/ Berlese technique at three primary and three secondary forest sites in Great Smoky Mountains National Park, Tennessee, USA, during fall 2006 and spring 2007. A total of 4261 adult beetle specimens, representing 216 lowest identifiable taxa within 159 genera and 28 families, were collected. Sixty-six species (31%) were represented by single individuals. Many more specimens (3471) and species (170) were collected from leaf litter than from CWD5 (790 and 111, respectively) but species accumulation curves showed that species richness was not significantly different between the two habitats. Eight species were significantly associated with CWD5, and 40 species were significantly associated with leaf litter. Species richness was significantly higher in secondary forest than primary forest, but more species were significantly associated with primary than secondary forest. Species richness was significantly higher in spring than fall. Notes on the biology and photographs of the 59 species represented by 10 or more specimens are given to provide an atlas of common eastern U.S. beetle species found in these habitats. Overall CWD5 is a distinct but overlooked habitat that may harbor numerous undescribed species or species considered rare.

Coleoptera species composition and succession in downed woody debris habitats are poorly known in eastern North America. A photoeclector emergence chamber was used to concentrate Coleoptera that emerged from various decay classes of fine and coarse woody debris (FWD and CWD, respectively) collected in primary and secondary forest sites in Great Smoky Mountains National Park, Tennessee, USA. A total of 5673 adult beetle specimens, representing 305 lowest identifiable taxa within 227 genera and 51 families, was collected. One hundred fifteen species (38%) were represented by single individuals. Many more specimens and species were collected from CWD (4129 and 247, respectively) than from FWD (1544 and 162, respectively), but species richness accumulation curves were not significantly different. Many more specimens but an equal number of species were collected from primary forest (3347 and 207, respectively) than from secondary forest (2326 and 207, respectively). Species accumulation curves indicated higher richness in secondary forests. Based on a subset of 71 species represented by 10 or more specimens, 27 species were associated with fresh fine woody debris, 11 species with weathered fine woody debris, four with coarse woody debris decay class I, 14 with coarse woody debris decay class II, and eight with coarse woody debris decay class III–IV. Sixteen species were associated with secondary forests, whereas 28 species were associated with primary forests. Coarse woody debris decay class II taken in primary forests had highest absolute species richness with 156 species. In coarse woody debris species overlap decreased with increased difference in decay indicating faunal succession. Published works related to the study of the ecology of downed woody material are briefly summarized. Recommendations on developing a database of legacy trees for future researchers are given. Notes on the biology and photographs of the 71 species represented by 10 or more specimens are giv

1. Nutrients are a critical resource for plant growth, but the elements limiting growth in tropical forests have rarely been determined. 2. We investigated the influence of nitrogen (N), phosphorus (P), potassium (K) and micronutrients on seedling biomass and nutrient allocation in a factorial nutrient fertilization experiment in lowland tropical forest at the Barro Colorado Nature Monument, Panama. We also measured 8 years of herbivory and growth for 1800 seedlings. We sought to determine the identity of limiting elements and possible nutrient interactions. 3. The five study species were Alseis blackiana, Desmopsis panamensis, Heisteria concinna, Sorocea affinis and Tetragastris panamensis. Plants grew in deeply shaded understorey with a mean canopy openness of 4.9% (±0.7%; 1 SE). 4. Tissue N concentration increased by 11% with N addition. Tissue P concentration increased by 16% with P addition. Tissue K increased by 4% with K addition. K addition reduced root‐to‐shoot biomass ratio. There was no significant effect of fertilization on specific leaf area or leaf area ratio. 5. The proportion of leaves damaged and the mean level of damage by herbivory increased with P and K addition and showed a significant P × K interaction. 6.7ensp;Across all species and years, relative growth rate of height increased with K addition and with N and P in combination. Relative growth rate of leaf count trended 8.5% higher with K addition (P = 0.076). 7. We also added micronutrients in a parallel experiment. There was no effect of micronutrient addition on any seedling parameter. 8. Synthesis. K addition affected seedlings by enhancing tissue nutrient concentration, increasing herbivory, reducing root‐to‐shoot biomass ratio and increasing height growth. Additional effects of N or P on tissue chemistry, herbivory and growth offer support for the multiple limiting resources hypothesis. Our results suggest that seedling growth is limited by nutrients, especially K, even under highly shad

Predicting future impacts of anthropogenic change on tropical forests requires a clear understanding of nutrient constraints on productivity. We compared experimental fertilization and litter manipulation treatments in an old-growth lowland tropical forest to distinguish between the effects of inorganic nutrient amendments and changes in nutrient cycling via litterfall. We measured the changes in soil and litter nutrient pools, litterfall, and fine root biomass in plots fertilized with nitrogen (N), phosphorus (P), or potassium (K), and in litter addition and litter removal treatments during 7 years. Soil inorganic N and litter N increased in double-litter plots but not in N-fertilized plots. Conversely, litter P and soil pools of P and K increased in fertilized plots but not in the double-litter plots. Soil and litter pools of N and K decreased in the no-litter plots. Changes in litterfall with added nutrients or litter were only marginally significant, but fine root biomass decreased with both the litter and the K addition. Differences between the two experiments are mostly attributable to the coupled cycling of carbon and nutrients in litter. Increased nutrient inputs in litter may improve plant uptake of some nutrients compared to fertilization with similar amounts. The litter layer also appears to play a key role in nutrient retention. We discuss our findings in the context of possible impacts of anthropogenic change on tropical forests

1. Nutrients are a critical resource for plant growth, but the elements limiting growth in tropical forests have rarely been determined. 2. We investigated the influence of nitrogen (N), phosphorus (P), potassium (K) and micronutrients on seedling biomass and nutrient allocation in a factorial nutrient fertilization experiment in lowland tropical forest at the Barro Colorado Nature Monument, Panama. We also measured 8 years of herbivory and growth for 1800 seedlings. We sought to determine the identity of limiting elements and possible nutrient interactions. 3. The five study species were Alseis blackiana, Desmopsis panamensis, Heisteria concinna, Sorocea affinis and Tetragastris panamensis. Plants grew in deeply shaded understorey with a mean canopy openness of 4.9% (±0.7%; 1 SE). 4. Tissue N concentration increased by 11% with N addition. Tissue P concentration increased by 16% with P addition. Tissue K increased by 4% with K addition. K addition reduced root‐to‐shoot biomass ratio. There was no significant effect of fertilization on specific leaf area or leaf area ratio. 5. The proportion of leaves damaged and the mean level of damage by herbivory increased with P and K addition and showed a significant P × K interaction. 6.7ensp;Across all species and years, relative growth rate of height increased with K addition and with N and P in combination. Relative growth rate of leaf count trended 8.5% higher with K addition (P = 0.076). 7. We also added micronutrients in a parallel experiment. There was no effect of micronutrient addition on any seedling parameter. 8. Synthesis. K addition affected seedlings by enhancing tissue nutrient concentration, increasing herbivory, reducing root‐to‐shoot biomass ratio and increasing height growth. Additional effects of N or P on tissue chemistry, herbivory and growth offer support for the multiple limiting resources hypothesis. Our results suggest that seedling growth is limited by nutrients, especially K, even under highly shad

Predicting future impacts of anthropogenic change on tropical forests requires a clear understanding of nutrient constraints on productivity. We compared experimental fertilization and litter manipulation treatments in an old-growth lowland tropical forest to distinguish between the effects of inorganic nutrient amendments and changes in nutrient cycling via litterfall. We measured the changes in soil and litter nutrient pools, litterfall, and fine root biomass in plots fertilized with nitrogen (N), phosphorus (P), or potassium (K), and in litter addition and litter removal treatments during 7 years. Soil inorganic N and litter N increased in double-litter plots but not in N-fertilized plots. Conversely, litter P and soil pools of P and K increased in fertilized plots but not in the double-litter plots. Soil and litter pools of N and K decreased in the no-litter plots. Changes in litterfall with added nutrients or litter were only marginally significant, but fine root biomass decreased with both the litter and the K addition. Differences between the two experiments are mostly attributable to the coupled cycling of carbon and nutrients in litter. Increased nutrient inputs in litter may improve plant uptake of some nutrients compared to fertilization with similar amounts. The litter layer also appears to play a key role in nutrient retention. We discuss our findings in the context of possible impacts of anthropogenic change on tropical forests.

Predicting future impacts of anthropogenic change on tropical forests requires a clear understanding of nutrient constraints on productivity. We compared experimental fertilization and litter manipulation treatments in an old-growth lowland tropical forest to distinguish between the effects of inorganic nutrient amendments and changes in nutrient cycling via litterfall. We measured the changes in soil and litter nutrient pools, litterfall, and fine root biomass in plots fertilized with nitrogen (N), phosphorus (P), or potassium (K), and in litter addition and litter removal treatments during 7 years. Soil inorganic N and litter N increased in double-litter plots but not in N-fertilized plots. Conversely, litter P and soil pools of P and K increased in fertilized plots but not in the double-litter plots. Soil and litter pools of N and K decreased in the no-litter plots. Changes in litterfall with added nutrients or litter were only marginally significant, but fine root biomass decreased with both the litter and the K addition. Differences between the two experiments are mostly attributable to the coupled cycling of carbon and nutrients in litter. Increased nutrient inputs in litter may improve plant uptake of some nutrients compared to fertilization with similar amounts. The litter layer also appears to play a key role in nutrient retention. We discuss our findings in the context of possible impacts of anthropogenic change on tropical forests.

Predicting future impacts of anthropogenic change on tropical forests requires a clear understanding of nutrient constraints on productivity. We compared experimental fertilization and litter manipulation treatments in an old-growth lowland tropical forest to distinguish between the effects of inorganic nutrient amendments and changes in nutrient cycling via litterfall. We measured the changes in soil and litter nutrient pools, litterfall, and fine root biomass in plots fertilized with nitrogen (N), phosphorus (P), or potassium (K), and in litter addition and litter removal treatments during 7 years. Soil inorganic N and litter N increased in double-litter plots but not in N-fertilized plots. Conversely, litter P and soil pools of P and K increased in fertilized plots but not in the double-litter plots. Soil and litter pools of N and K decreased in the no-litter plots. Changes in litterfall with added nutrients or litter were only marginally significant, but fine root biomass decreased with both the litter and the K addition. Differences between the two experiments are mostly attributable to the coupled cycling of carbon and nutrients in litter. Increased nutrient inputs in litter may improve plant uptake of some nutrients compared to fertilization with similar amounts. The litter layer also appears to play a key role in nutrient retention. We discuss our findings in the context of possible impacts of anthropogenic change on tropical forests

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 x 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1-10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 × 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1–10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 x 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1-10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 × 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1–10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 × 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1–10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 × 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1–10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.

El reclutamiento exitoso desde semillas en bosques neotropicales implica una secuencia de etapas. La disponibilidad del polen y recursos consumibles por los árboles maternales puede limitar el número de semillas producidas. La dispersión de semillas a un sitio determinado puede ser limitada por la densidad o la dispersión de árboles frutales, o por el agrupamiento impuesto por los procesos de dispersión de semillas. El establecimiento de semillas dispersadas puede ser limitado por la mortalidad debida a enemigos naturales, por ejemplo depredadores de semillas y herbivoros, o por factores abióticos tales como la disponibilidad de agua, nutrientes y luz. Como la limitación impuesta por estas etapas puede verse afectada por la explotación forestal selectiva, es necesario investigar el efecto de las prácticas selvícolas sobre cada etapa en la dinámica del bosque.

El reclutamiento exitoso desde semillas en bosques neotropicales implica una secuencia de etapas. La disponibilidad del polen y recursos consumibles por los árboles maternales puede limitar el número de semillas producidas. La dispersión de semillas a un sitio determinado puede ser limitada por la densidad o la dispersión de árboles frutales, o por el agrupamiento impuesto por los procesos de dispersión de semillas. El establecimiento de semillas dispersadas puede ser limitado por la mortalidad debida a enemigos naturales, por ejemplo depredadores de semillas y herbivoros, o por factores abióticos tales como la disponibilidad de agua, nutrientes y luz. Como la limitación impuesta por estas etapas puede verse afectada por la explotación forestal selectiva, es necesario investigar el efecto de las prácticas selvícolas sobre cada etapa en la dinámica del bosque.

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40x40 m each. The addition of K was associated with significant decreases in stand-level fine root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1–10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40x40 m each. The addition of K was associated with significant decreases in stand-level fine root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1–10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles