I present a meta-analysis of plant responses to 48 nutrient addition experiments conducted with native species in naturally growing tropical forests, exclusive of mangrove forests. The added ...nutrients include nitrogen (N) in 36 experiments, phosphorus (P) in 33 experiments, calcium and potassium in one experiment each, and various mixtures of essential nutrients in the remaining experiments. I evaluate the hypotheses that nutrients limit tropical forest plants, nutrient limitation is stronger in successional than in old-growth forests, P but not N is limiting in lowland forests, and N is limiting in montane forests. Responses to the most complete nutrient mix used in each experiment were strong for plant functions that contribute to aboveground production (Hedges' g averages 0.87) and nonsignificant for fine root biomass. Responses to N addition and to P addition were strong for tissue concentrations of the added element (Hedges' g averages 0.75 and 1.4, respectively), moderate for fine litter production (0.64 and 0.65, respectively), moderate to weak for plant growth (0.46 and 0.37, respectively) and nonsignificant for fine root biomass. Growth responses were stronger in successional than in old-growth forests. All responses were unrelated to elevation. The 48 experiments included 30 factorial nitrogen-phosphorus experiments that enable additional direct tests of the widely cited hypotheses that P limitation is stronger than N limitation in lowland forests and vice versa in montane forests. Both hypotheses were rejected. The N × P interaction effect was nonsignificant across the factorial experiments. In conclusion, nutrients clearly limit tropical forest plants. Limitation by N is widespread in both lowland and montane forests, and the same is true for P. Single experiments identify limitation by calcium and potassium, and correlative studies suggest limitation by calcium, potassium, and magnesium. The available evidence is consistent with the possibility that most macronutrients limit tropical forest plants; however, experiments focus almost exclusively on N and P. The way forward will include taking fuller advantage of existing nutrient addition experiments, siting new experiments strategically, and developing cost-effective methods to assay responses to all of the essential nutrients soils supply to plants.
Questions remain as to which soil nutrients limit primary production in tropical forests. Phosphorus (P) has long been considered the primary limiting element in lowland forests, but recent evidence ...demonstrates substantial heterogeneity in response to nutrient addition, highlighting a need to understand and diagnose nutrient limitation across diverse forests. Fine‐root characteristics including their abundance, functional traits, and mycorrhizal symbionts can be highly responsive to changes in soil nutrients and may help to diagnose nutrient limitation. Here, we document the response of fine roots to long‐term nitrogen (N), P, and potassium (K) fertilization in a lowland forest in Panama. Because this experiment has demonstrated that N and K together limit tree growth and P limits fine litter production, we hypothesized that fine roots would also respond to nutrient addition. Specifically we hypothesized that N, P, and K addition would reduce the biomass, diameter, tissue density, and mycorrhizal colonization of fine roots, and increase nutrient concentration in root tissue. Most morphological root traits responded to the single addition of K and the paired addition of N and P, with the greatest response to all three nutrients combined. The addition of N, P, and K together reduced fine‐root biomass, length, and tissue density, and increased specific root length, whereas root diameter remained unchanged. Nitrogen addition did not alter root N concentration, but P and K addition increased root P and K concentration, respectively. Mycorrhizal colonization of fine roots declined with N, increased with P, and was unresponsive to K addition. Although plant species composition remains unchanged after 14 years of fertilization, fine‐root characteristics responded to N, P, and K addition, providing some of the strongest stand‐level responses in this experiment. Multiple soil nutrients regulate fine‐root abundance, morphological and chemical traits, and their association with mycorrhizal fungi in a species‐rich lowland tropical forest.
Nutrient availability is widely considered to constrain primary productivity in lowland tropical forests, yet there is little comparable information for the soil microbial biomass. We assessed ...microbial nutrient limitation by quantifying soil microbial biomass and hydrolytic enzyme activities in a long-term nutrient addition experiment in lowland tropical rain forest in central Panama. Multiple measurements were made over an annual cycle in plots that had received a decade of nitrogen, phosphorus, potassium, and micronutrient addition. Phosphorus addition increased soil microbial carbon (13 %), nitrogen (21 %), and phosphorus (49 %), decreased phosphatase activity by ~65 % and N-acetyl β-glucosaminidase activity by 24 %, but did not affect β-glucosidase activity. In contrast, addition of nitrogen, potassium, or micronutrients did not significantly affect microbial biomass or the activity of any enzyme. Microbial nutrients and hydrolytic enzyme activities all declined markedly in the dry season, with the change in microbial biomass equivalent to or greater than the annual nutrient flux in fine litter fall. Although multiple nutrients limit tree productivity at this site, we conclude that phosphorus limits microbial biomass in this strongly-weathered lowland tropical forest soil. This finding indicates that efforts to include enzymes in biogeochemical models must account for the disproportionate microbial investment in phosphorus acquisition in strongly-weathered soils.
507 I. 507 II. 509 III. 510 IV. 510 V. 512 VI. 516 VII. 518 518 References 518 SUMMARY: Hyperdiverse forests occur in the lowland tropics, whereas the most species‐rich shrublands are found in ...regions such as south‐western Australia (kwongan) and South Africa (fynbos). Despite large differences, these ecosystems share an important characteristic: their soils are strongly weathered and phosphorus (P) is a key growth‐limiting nutrient. Soil‐borne pathogens are increasingly being recognized as drivers of plant diversity in lowland tropical rainforests, but have received little attention in species‐rich shrublands. We suggest a trade‐off in which the species most proficient at acquiring P have ephemeral roots that are particularly susceptible to soil‐borne pathogens. This could equalize out the differences in competitive ability among co‐occurring species in these ecosystems, thus contributing to coexistence. Moreover, effective protection against soil‐borne pathogens by ectomycorrhizal (ECM) fungi might explain the occurrence of monodominant stands of ECM trees and shrubs amongst otherwise species‐rich communities. We identify gaps in our knowledge which need to be filled in order to evaluate a possible link between P limitation, fine root traits, soil‐borne pathogens and local plant species diversity. Such a link may help to explain how numerous plant species can coexist in hyperdiverse rainforests and shrublands, and, conversely, how monodominant stands can develop in these ecosystems.
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 (
r
2
= 0.69), intermediate for saplings for average growth rates and overall mortality rates (
r
2
= 0.46), and much weaker for large trees (
r
2
≤ 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 (
r
2
< 0.1 for three and
r
2
< 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.
Cunha and colleagues conducted their experiment at a well-characterized site in the central Amazon rainforest where soils are highly weathered and soil phosphorus levels are among some of the lowest ...recorded for any tropical forest (Fig. 1). ...the authors predicted that there would be strong plant responses to the addition of phosphorus, and possibly to the addition of the major cations, but not to nitrogen supplementation. The rise in leaf production was associated with increased leaf turnover rates, without an increase in leaf biomass. ...extra wood production was not required to support increased leaf biomass, and wood production was similar across all nutrient-addition treatments. Long-lived carbon pools will increase with phosphorus addition as the experiment continues, if decomposition-resistant organic compounds in leaves and roots are added to pools of soil organic matter, and if trees adapted to phosphorus-impoverished soils slowly increase wood production.
Understanding and mitigating the impact of an ever-increasing population and global economic activity on tropical forests is one of the great challenges currently facing biologists, conservationists ...and policy makers. Tropical forests currently face obvious regional changes, both negative and positive, and uncertain global changes. Although deforestation rates have increased to unprecedented levels, natural secondary succession has reclaimed approximately 15% of the area deforested during the 1990s. Governments have also protected 18% of the remaining tropical moist forest; however, unsustainable hunting continues to threaten many keystone mammal and bird species. The structure and dynamics of old-growth forests appear to be rapidly changing, suggesting that there is a pantropical response to global anthropogenic forcing, although the evidence comes almost exclusively from censuses of tree plots and is controversial. Here, I address ongoing anthropogenic change in tropical forests and suggest how these forests might respond to increasing anthropogenic pressure.
Nutrient addition experiments indicate that nitrogen and phosphorus limit plant processes in many tropical forests. However, the long-term consequences for forest structure and species composition ...remain unexplored. We are positioned to evaluate potential long-term consequences of nutrient addition in central Panama where we have maintained a factorial nitrogen–phosphorus–potassium fertilization experiment for 21 yr and an independent study quantified the species-specific nutrient requirements of 550 local tree species. Here, we ask whether nutrients limit reproduction at the species and community levels. We also ask whether species-specific reproductive responses to nutrient addition are stronger among species associated with naturally fertile soils, which could contribute to a shift in species composition. We quantified species-level reproductive responses for 38 focal species in the 21st year of the experiment and community-level reproductive litter production for the first 20 yr. Species-level reproductive responses to nitrogen and potassium addition were weak, inconsistent across species, and insignificant across the 38 focal species. In contrast, species-level responses to phosphorus addition were consistently and significantly positive across the 38 focal species but were unrelated to species-specific phosphorus requirements documented independently for the same species. Community-level reproductive litter production was unaffected by nutrient addition, possibly because spatial and temporal variation is large. We conclude that phosphorus limits reproduction by trees in our experiment but find no evidence that reproductive responses to phosphorus addition favor species associated with naturally phosphorus-rich soils.
We present a meta-analysis of plant responses to fertilization experiments conducted in lowland, species-rich, tropical forests. We also update a key result and present the first species-level ...analyses of tree growth rates for a 15-yr factorial nitrogen (N), phosphorus (P), and potassium (K) experiment conducted in central Panama. The update concerns community-level tree growth rates, which responded significantly to the addition of N and K together after 10 yr of fertilization but not after 15 yr. Our experimental soils are infertile for the region, and species whose regional distributions are strongly associated with low soil P availability dominate the local tree flora. Under these circumstances, we expect muted responses to fertilization, and we predicted species associated with low-P soils would respond most slowly. The data did not support this prediction, species-level tree growth responses to P addition were unrelated to species-level soil P associations. The meta-analysis demonstrated that nutrient limitation is widespread in lowland tropical forests and evaluated two directional hypotheses concerning plant responses to N addition and to P addition. The meta-analysis supported the hypothesis that tree (or biomass) growth rate responses to fertilization are weaker in old growth forests and stronger in secondary forests, where rapid biomass accumulation provides a nutrient sink. The meta-analysis found no support for the long-standing hypothesis that plant responses are stronger for P addition and weaker for N addition. We do not advocate discarding the latter hypothesis. There are only 14 fertilization experiments from lowland, species-rich, tropical forests, 13 of the 14 experiments added nutrients for five or fewer years, and responses vary widely among experiments. Potential fertilization responses should be muted when the species present are well adapted to nutrient-poor soils, as is the case in our experiment, and when pest pressure increases with fertilization, as it does in our experiment. The statistical power and especially the duration of fertilization experiments conducted in old growth, tropical forests might be insufficient to detect the slow, modest growth responses that are to be expected.
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