Phosphorus availability is widely assumed to limit primary productivity in tropical forests, but support for this paradigm is equivocal. Although biogeochemical theory predicts that phosphorus limitation should be prevalent on old, strongly weathered soils, experimental manipulations have failed to detect a consistent response to phosphorus addition in species-rich lowland tropical forests. Here we show, by quantifying the growth of 541 tropical tree species across a steep natural phosphorus gradient in Panama, that phosphorus limitation is widespread at the level of individual species and strengthens markedly below a threshold of two parts per million exchangeable soil phosphate. However, this pervasive species-specific phosphorus limitation does not translate into a community-wide response, because some species grow rapidly on infertile soils despite extremely low phosphorus availability. These results redefine our understanding of nutrient limitation in diverse plant communities and have important implications for attempts to predict the response of tropical forests to environmental change.

Abstract. Understanding the mechanisms that shape the distribution of organisms can help explain patterns of local and regional biodiversity and predict the susceptibility of communities to environmental change. In the species-rich tropics, a gradient in rainfall between wet evergreen and dry seasonal forests correlates with turnover of plant species. The strength of the dry season has previously been shown to correlate with species composition. Herbivores and pathogens (pests) have also been hypothesized to be important drivers of plant distribution, although empirical evidence is lacking. In this study we experimentally tested the existence of a gradient in pest pressure across a rainfall gradient in the Isthmus of Panama and measured the influence of pests relative to drought on species turnover. We established two common gardens on the dry and wet sides of the Isthmus using seedlings from 24 plant species with contrasting distributions along the Isthmus. By experimentally manipulating water availability and insect herbivore access, we showed that pests are not as strong a determinant of plant distributions as is seasonal drought. Seasonal drought in the dry site excluded wetdistribution species by significantly increasing their seedling mortality. Pathogen mortality and insect herbivore damage were both higher in the wet site, supporting the existence of a gradient in pest pressure. However, contrary to predictions, we found little evidence that drydistribution species suffered significantly more pest attack than wet-distribution species. Instead, we hypothesize that dry-distribution species are limited from colonizing wetter forests by their inherently slower growth rates imposed by drought adaptations. We conclude that mechanisms limiting the recruitment of dry-distribution species in wet forests are not nearly as strong as those limiting wet-distribution species from dry forests.

In tropical forests, regional differences in annual rainfall correlate with differences in plant species composition. Although water availability is clearly one factor determining species distribution, other environmental variables that covary with rainfall may contribute to distributions. One such variable is light availability in the understory, which decreases towards wetter forests due to differences in canopy density and phenology. We established common garden experiments in three sites along a rainfall gradient across the Isthmus of Panama in order to measure the differences in understory light availability, and to evaluate their influence on the performance of 24 shade-tolerant species with contrasting distributions. Within sites, the effect of understory light availability on species performance depended strongly on water availability. When water was not limiting, either naturally in the wetter site or through water supplementation in drier sites, seedling performance improved at higher light. In contrast, when water was limiting at the drier sites, seedling performance was reduced at higher light, presumably due to an increase in water stress that affected mostly wet-distribution species. Although wetter forest understories were on average darker, wet-distribution species were not more shade-tolerant than dry-distribution species. Instead, wet-distribution species had higher absolute growth rates and, when water was not limiting, were better able to take advantage of small increases in light than dry-distribution species. Our results suggest that in wet forests the ability to grow fast during temporary increases in light may be a key trait for successful recruitment. The slower growth rates of the dry-distribution species, possibly due to trade-offs associated with greater drought tolerance, may exclude these species from wetter forests.Electronic supplementary materialThe online version of this article (doi:10.1007/s00442-010-1832-9) contains supplementary material, which is available to authorized users.

Abstract. In the recently radiated genus Inga (Fabaceae), few nucleotide substitutions have accumulated among species, yet large divergences have occurred in defensive phenotypes, suggesting strong selection by herbivores. We compared herbivory and defenses of young leaves for I. goldmanii, a more derived species that follows a ''defense'' strategy, and I. umbellifera, a more basal species that follows an ''escape'' strategy. The two species suffered similar rates of herbivory (22% of the leaf area eaten during expansion) but were attacked by different communities of herbivores. I. goldmanii relied heavily on extra-floral nectaries and on a diversity of effective secondary metabolites, while I. umbellifera minimized damage through rapid leaf expansion and synchronous flushing. The major classes of secondary compounds in both species were flavanoids and non-protein amino acids; however, there were large differences in structure, biosynthetic pathways, and efficacy against herbivores. Growth rates of lepidopteran larvae were significantly lower when fed artificial diets with crude extracts from I. goldmanii as compared to I. umbellifera. Flavanoids accounted for the majority of growth reduction in both species. I. umbellifera had more unusual flavanoids and a non-protein amino acid not reported from plants, but the more common flavanoids found in I. goldmanii were more bioactive against herbivores. I. goldmanii also had greater ant visitation to extrafloral nectaries, suggesting that there was no trade-off between biotic and chemical defenses. In contrast, young leaves of I. umbellifera expanded more rapidly, minimizing the window of vulnerability before toughening. Resources for rapid expansion may have been reallocated from chloroplast development as I. umbellifera delayed the greening process until after full leaf expansion. Leaves were also produced synchronously, which can satiate herbivores and reduce damage. These defense differences are reflected in almost completely nonoverlapping herbivore faunas and the more frequent occurrence of generalists on I. umbellifera. To understand why defenses have evolved, it is important to view them in light of the herbivore community as well as in the context of the other co-occurring traits. We hypothesize that the effectiveness of chemical defenses determines whether a species follows the evolutionary path of ''defense'' or ''escape'' strategies.

MO 63121-4499, USA. Present address for CB: Univ. Nacional de Corboda, Argentina.Within the tropics, a marked gradient in rainfall between dry and wet forests correlates with a well documented turnover of plant species. While water availability along these gradients is an important determinant of species distributions, other abiotic and biotic factors correlate with rainfall and may also contribute to limit species distribution. One of these is soil fertility, which is often lower in the wetter forests. To test its possible role in species distribution along a rainfall gradient, we performed a screen-house experiment where we measured the growth performance of seedlings of 23 species with contrasting distributions across the Isthmus of Panama. We grew seedlings in soils collected from the drier Pacific side and the wetter Atlantic side. Differences in soil fertility across the Isthmus were large enough to significantly influence the growth performance of the seedlings. However, we found no evidence of home-soil advantage among species with contrasting distributions. Dry-distribution species grew on average slower than wet-distribution species suggesting a cost to drought adaptations. The response to soil differences correlated with the growth rate of the species, such that fastgrowing species responded more to changes in soil quality. We hypothesize that inherently slow growth rates of some drydistribution tropical species may be a more important factor limiting their colonization of wetter sites along the rainfall gradient.