In an edaphically heterogeneous area in the Peruvian Amazon, clay soils and nutrient-poor white sands each harbor distinctive plant communities. To determine whether a trade-off between growth and ...antiherbivore defense enforces habitat specialization on these two soil types, we conducted a reciprocal transplant study of seedlings of 20 species from sixgenera of phylogenetically independent pairs of edaphic specialist trees and manipulated the presence of herbivores. Clay specialist species grew significantly faster than white-sand specialists in both soil types when protected from herbivores. However, when unprotected, white-sand specialists dominated in white-sand forests and clay specialists dominated in clay forests. Therefore, habitat specialization in this system results from an interaction of herbivore pressure with soil type.
The extent to which pre-Columbian societies altered Amazonian landscapes is hotly debated. We performed a basin-wide analysis of pre-Columbian impacts on Amazonian forests by overlaying known ...archaeological sites in Amazonia with the distributions and abundances of 85 woody species domesticated by pre-Columbian peoples. Domesticated species are five times more likely than nondomesticated species to be hyperdominant. Across the basin, the relative abundance and richness of domesticated species increase in forests on and around archaeological sites. In southwestern and eastern Amazonia, distance to archaeological sites strongly influences the relative abundance and richness of domesticated species. Our analyses indicate that modern tree communities in Amazonia are structured to an important extent by a long history of plant domestication by Amazonian peoples.
Tropical forests include a diversity of habitats, which has led to specialization in plants. Near Iquitos, in the Peruvian Amazon, nutrient-rich clay forests surround nutrient-poor white-sand ...forests, each harboring a unique composition of habitat specialist trees. We tested the hypothesis that the combination of impoverished soils and herbivory creates strong natural selection for plant defenses in white-sand forest, while rapid growth is favored in clay forests. Recently, we reported evidence from a reciprocal-transplant experiment that manipulated the presence of herbivores and involved 20 species from six genera, including phylogenetically independent pairs of closely related white-sand and clay specialists. When protected from herbivores, clay specialists exhibited faster growth rates than white-sand specialists in both habitats. But, when unprotected, white-sand specialists outperformed clay specialists in white-sand habitat, and clay specialists outperformed white-sand specialists in clay habitat. Here we test further the hypothesis that the growth-defense trade-off contributes to habitat specialization by comparing patterns of growth, herbivory, and defensive traits in these same six genera of white-sand and clay specialists. While the probability of herbivore attack did not differ between the two habitats, an artificial defoliation experiment showed that the impact of herbivory on plant mortality was significantly greater in white-sand forests. We quantified the amount of terpenes, phenolics, leaf toughness, and available foliar protein for the plants in the experiment. Different genera invested in different defensive strategies, and we found strong evidence for phylogenetic constraint in defense type. Overall, however, we found significantly higher total defense investment for white-sand specialists, relative to their clay specialist congeners. Furthermore, herbivore resistance consistently exhibited a significant trade-off against growth rate in each of the six phylogenetically independent species-pairs. These results confirm theoretical predictions that a trade-off exists between growth rate and defense investment, causing white-sand and clay specialists to evolve divergent strategies. We propose that the growth-defense trade-off is universal and provides an important mechanism by which herbivores govern plant distribution patterns across resource gradients.
Herbivores are often implicated in the generation of the extraordinarily diverse tropical flora. One hypothesis linking enemies to plant diversification posits that the evolution of novel defenses ...allows plants to escape their enemies and expand their ranges. When range expansion involves entering a new habitat type, this could accelerate defense evolution if habitats contain different assemblages of herbivores and/or divergent resource availabilities that affect plant defense allocation. We evaluated this hypothesis by investigating two sister habitat specialist ecotypes of Protium subserratum (Burseraceae), a common Amazonian tree that occurs in white‐sand and terra firme forests. We collected insect herbivores feeding on the plants, assessed whether growth differences between habitats were genetically based using a reciprocal transplant experiment, and sampled multiple populations of both lineages for defense chemistry. Protium subserratum plants were attacked mainly by chrysomelid beetles and cicadellid hemipterans. Assemblages of insect herbivores were dissimilar between populations of ecotypes from different habitats, as well as from the same habitat 100 km distant. Populations from terra firme habitats grew significantly faster than white‐sand populations; they were taller, produced more leaf area, and had more chlorophyll. White‐sand populations expressed more dry mass of secondary compounds and accumulated more flavone glycosides and oxidized terpenes, whereas terra firme populations produced a coumaroylquinic acid that was absent from white‐sand populations. We interpret these results as strong evidence that herbivores and resource availability select for divergent types and amounts of defense investment in white‐sand and terra firme lineages of Protium subserratum, which may contribute to habitat‐mediated speciation in these trees.
Aim
To investigate the geographic patterns and ecological correlates in the geographic distribution of the most common tree dispersal modes in Amazonia (endozoochory, synzoochory, anemochory and ...hydrochory). We examined if the proportional abundance of these dispersal modes could be explained by the availability of dispersal agents (disperser‐availability hypothesis) and/or the availability of resources for constructing zoochorous fruits (resource‐availability hypothesis).
Time period
Tree‐inventory plots established between 1934 and 2019.
Major taxa studied
Trees with a diameter at breast height (DBH) ≥ 9.55 cm.
Location
Amazonia, here defined as the lowland rain forests of the Amazon River basin and the Guiana Shield.
Methods
We assigned dispersal modes to a total of 5433 species and morphospecies within 1877 tree‐inventory plots across terra‐firme, seasonally flooded, and permanently flooded forests. We investigated geographic patterns in the proportional abundance of dispersal modes. We performed an abundance‐weighted mean pairwise distance (MPD) test and fit generalized linear models (GLMs) to explain the geographic distribution of dispersal modes.
Results
Anemochory was significantly, positively associated with mean annual wind speed, and hydrochory was significantly higher in flooded forests. Dispersal modes did not consistently show significant associations with the availability of resources for constructing zoochorous fruits. A lower dissimilarity in dispersal modes, resulting from a higher dominance of endozoochory, occurred in terra‐firme forests (excluding podzols) compared to flooded forests.
Main conclusions
The disperser‐availability hypothesis was well supported for abiotic dispersal modes (anemochory and hydrochory). The availability of resources for constructing zoochorous fruits seems an unlikely explanation for the distribution of dispersal modes in Amazonia. The association between frugivores and the proportional abundance of zoochory requires further research, as tree recruitment not only depends on dispersal vectors but also on conditions that favour or limit seedling recruitment across forest types.
Tropical plant diversity is extraordinarily high at both local and regional scales. Many studies have demonstrated that natural enemies maintain local diversity via negative density dependence, but ...we know little about how natural enemies influence beta-diversity across habitats and/or regions. One way herbivores could influence plant beta-diversity is by driving allocation trade-offs that promote habitat specialization across resource gradients. We therefore predicted that increasing resource availability should be accompanied by increasing herbivory rates and decreasing plant allocation to defense. Second, relative abundances within plant lineages are predicted to reflect patterns of habitat specialization and allocation trade-offs. A phylogenetic context is vital not only to compare homologous plant traits (including defense strategies) across habitat types, but also to connect evolutionary trade-offs to patterns of species diversification in each phylogenetic lineage.
We tested these predictions for trees in white-sand, clay terra firme, and seasonally flooded forests in Peru and French Guiana that represent the range of soil fertility, forest structure, and floristic compositions found throughout the Amazon region. We established 74 0.5-ha plots in these habitats and sampled all trees. Within 12 representative plots we marked newly expanding leaves of 394 saplings representing 68 species, including the most abundant species in each plot in addition to species from five focal lineages:
Swartzia
and
Inga
(Fabaceae), Protieae (Burseracaeae), Bombacoideae (Malvaceae), and
Micropholis
(Sapotaceae). We measured total leaf production rates for each sapling and calculated relative herbivory impact as the ratio between herbivory rate and leaf production rate.
Herbivory rates averaged 2.1% per month, did not correlate with leaf production rate, and were similar across habitats. Relative herbivore impacts exceeded leaf production rates for most species. Leaf production rate averaged 2.8%, was significantly higher in seasonally flooded forests than the other two habitats, and exhibited significant correlations with specific leaf area. Species with high herbivory rates exhibited significantly lower relative abundances in terra firme forests. Overall, focal species within lineages present contrasting patterns regarding their herbivory rates and leaf production rate within habitats. These results highlight why a lineage-based approach is necessary when attempting to connect hypotheses regarding evolutionary trade-offs to community assembly patterns.
Local species richness and between-site similarity in species composition of parasitoid wasps (Hymenoptera: Ichneumonidae; Pimplinae and Rhyssinae) were correlated with those of four plant groups ...(pteridophytes, Melastomataceae, Burseraceae and Arecaceae) in a western Amazonian lowland rain forest mosaic. The mosaic structure of the forest was related to variation in soils within the non-inundated terrain. Significant matrix correlation between patterns in parasitoid wasp species composition and plant species composition was found. Most of the overall correlation was due to idiobiont parasitoids of weakly concealed hosts, which attack host larvae and pupae in exposed situations, with two of the four ecologically defined parasitoid groups showing no correlation at all. A positive correlation between the number of plant species and the number of Pimplinae and Rhyssinae species at a site was found when the latter was corrected for collecting effort. Consequently, the degree of floristic difference between sites may be indicative of the difference in species composition of ichneumonids, and the species richness of plants may serve as a predictor of the species richness of parasitoid wasps. Although these results were obtained in a mosaic including structurally and floristically clearly different types of rain forest, the correlation coefficients were relatively low, and the present results lend only weak support to the idea of using plant distributions as indicators of animal distributions.
Environmental heterogeneity in the tropics is thought to lead to specialization in plants and thereby contribute to the diversity of the tropical flora. We examine this idea with data on the habitat ...specificity of 35 western Amazonian species from the genera Protium, Crepidospermum, and Tetragastris in the monophyletic tribe Protieae (Burseraceae) mapped on a molecular-based phylogeny. We surveyed three edaphic habitats that occur throughout terra firme Amazonia: white-sand, clay, and terrace soils in eight forests across more than 2000 km in the western Amazon. Twenty-six of the 35 species were found to be associated with only one of three soil types, and no species was associated with all three habitats; this pattern of edaphic specialization was consistent across the entire region. Habitat association mapped onto the phylogenetic tree shows association with terrace soils to be the probable ancestral state in the group, with subsequent speciation events onto clay and white-sand soils. The repeated gain of clay association within the clade likely coincides with the emergence of large areas of clay soils in the Miocene deposited during the Andean uplift. Character optimizations revealed that soil association was not phylogenetically clustered for white-sand and clay specialists, suggesting repeated independent evolution of soil specificity is common within the Protieae. This phylogenetic analysis also showed that multiple cases of putative sister taxa with parapatric distributions differ in their edaphic associations, suggesting that edaphic heterogeneity was an important driver of speciation in the Protieae in the Amazon basin.
Tropical forests contain an important proportion of the carbon stored in terrestrial vegetation, but estimated aboveground biomass (AGB) in tropical forests varies two-fold, with little consensus on ...the relative importance of climate, soil and forest structure in explaining spatial patterns. Here, we present analyses from a plot network designed to examine differences among contrasting forest habitats (terra firme, seasonally flooded, and white-sand forests) that span the gradient of climate and soil conditions of the Amazon basin. We installed 0.5-ha plots in 74 sites representing the three lowland forest habitats in both Loreto, Peru and French Guiana, and we integrated data describing climate, soil physical and chemical characteristics and stand variables, including local measures of wood specific gravity (WSG). We use a hierarchical model to separate the contributions of stand variables from climate and soil variables in explaining spatial variation in AGB. AGB differed among both habitats and regions, varying from 78Mgha_1 in white-sand forest in Peru to 605Mgha_1 in terra firme clay forest of French Guiana. Stand variables including tree size and basal area, and to a lesser extent WSG, were strong predictors of spatial variation in AGB. In contrast, soil and climate variables explained little overall variation in AGB, though they did co-vary to a limited extent with stand parameters that explained AGB. Our results suggest that positive feedbacks in forest structure and turnover control AGB in Amazonian forests, with richer soils (Peruvian terra firme and all seasonally flooded habitats) supporting smaller trees with lower wood density and moderate soils (French Guianan terra firme) supporting many larger trees with high wood density. The weak direct relationships we observed between soil and climate variables and AGB suggest that the most appropriate approaches to landscape scale modeling of AGB in the Amazon would be based on remote sensing methods to map stand structure. (Résumé d'auteur)
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