Leaf breakdown in an Atlantic Rain Forest stream

The hypothesis of this study was that colonizers in decaying leaf litter prefer native species (Erythrina verna) to exotic ones (Eucalyptus camaldulensis and Protium heptaphyllum). Therefore, native species are expected to show higher breakdown rates, increased biomass, richness and density of invertebrate species, and increased biomass of decomposer fungi. Breakdown of leaf litter from these three species was assessed in an Atlantic Rain Forest stream. Four samples were collected during a period of 90 days and washed on a sieve to separate the invertebrates. Then, a series of leaf disks were cut to determine ash‐free dry mass and fungal biomass, and the remaining material was oven‐dried to determine the dry weight. Eucalyptus camaldulensis and E. verna showed higher breakdown rates than P. heptaphyllum, due to differences in leaf physical and chemical characteristics. The harder detritus (P. heptaphyllum) broke down more slowly than detritus with high concentrations of labile compounds (E. camaldulensis). The density of the invertebrates associated with detritus increased with time. There were no differences in density, taxonomic richness or biomass of invertebrates among the leaf types, which indicated that the invertebrates did not distinguish between exotic and native detritus. Fungal colonization varied among samples; E. camaldulensis showed the lowest ergosterol concentrations, mainly due to a high concentration of total phenolics. The detritus with the highest hardness value was colonized most slowly by fungi. These results showed that leaf breakdown in Atlantic Rain Forest streams could be affected either by changes in riparian vegetation, or by becoming more savanna‐like process due to climate change.     

What is more important for invertebrate colonization in a stream with low-quality litter inputs: exposure time or leaf species?

The objective of this study was to evaluate the influences of detritus from the leaves of different species, and of exposure time on invertebrate colonization of leaves in a shaded Cerrado stream. We hypothesized that the exposure time is the main factor that influences the colonization of leaves by invertebrates. We used leaves of five tree species native to the Brazilian Cerrado: Protium heptaphyllum and Protium brasiliense (Burseraceae), Ocotea sp. (Lauraceae), Myrcia guyanensis (Myrtaceae), and Miconia chartacea (Melastomataceae), which are characterized by their toughness and low-nutritional quality. Litter bags, each containing leaves from one species, were placed in a headwater stream and removed after 7, 15, 30, 60, 90, and 120 days. The dominant taxon was Chironomidae, which comprised ca. 52% of all organisms and ca. 20% of the total biomass. The taxonomic richness of colonizing organisms did not vary among the leaf species. However, the density and biomass of the associated organisms varied differently among the kinds of detritus during the course of the incubation. The collector-gatherers and shredders reached higher densities in the detritus that decomposed more rapidly (Ocotea sp. and M. guyanensis), principally in the more advanced stages of colonization. The collector-filterers reached higher densities in the detritus that decomposed more slowly (P. heptaphyllum, P. brasiliense, and M. chartacea), principally in the initial stages of incubation. A cluster analysis divided the detritus samples of different leaf species according to the exposure time (initial phase: up to 7 days; intermediate phase: 7–30 days; advanced phase: 30–120 days), suggesting some succession in invertebrate colonization, with differences in taxon composition (indicator taxa analysis). These results suggest that regardless of the leaf-detritus species, exposure time was the main factor that influenced the colonization process of aquatic invertebrates.     

Interactions between Ants and Non‐myrmecochorous Fruits in Miconia (Melastomataceae) in a Neotropical Savanna

The aim of this study was to evaluate the role of ants as secondary seed dispersers of six primarily bird‐dispersed Miconia species in the cerrados of southeastern Brazil. Vertebrate exclosure and seed germination experiments were performed for M. albicans, M. alborufescens, M. corallina, M. ferruginata, M. ibaguensis, and M. irwinii. Excluding vertebrates did not significantly alter fruit removal rate for any of the Miconia species relative to open controls. Fruits on stalks and fallen fruits were removed and transported to nests mainly by species of Atta, Acromyrmex, and Ectatomma (dispersal distance ranging from 0.1 to 45.2 m), while Camponotus ants tended to be observed removing the fruit pulp (seed cleaning) where the fruits were found. Seed manipulation by Atta decreased germination of M. irwinii, but not M. ferruginata. Germination did not occur in intact fruits, and thus seed cleaning was an important service provided by the ants. Ant nest soils did not inhibit germination of any of the Miconia species, suggesting they are a good substrate for long‐lived Miconia seeds. We conclude that ant activity could have important effects on the fate of Miconia seeds adapted for bird dispersal.     

Spatial Distribution and Fruiting Phenology of Protium heptaphyllum (Burseraceae) Determine the Design of the Underground Foraging System of Atta sexdens L. (Hymenoptera: Formicidae)

Leaf-cutting ants have long been recognized to forage via complex trail systems but the nature and the ecological drivers of the different foraging strategies adopted remain a key topic. Here, we described the spatiotemporal use of belowground foraging galleries by Atta sexdens L. in the Brazilian Atlantic forest, and examined the adaptive advantages of this foraging strategy. Protium heptaphyllum adult trees (DBH?>?10?cm), seed/seedling clumps and ant gallery entrances were mapped across two 1-ha plots during two consecutive fruiting seasons (2002 and 2004). We recorded 75 ca. 40?cm deep gallery entrances beneath 26 P. heptaphyllum trees at nest distances ranging from 14 to 57?m. Furthermore, gallery abundance and galleries associated with seed/seedling clumps correlated positively with P. heptaphyllum density. Our results indicate that A. sexdens was able to set a permanent system of underground galleries targeting P. heptaphyllum trees and their seeds on the ground. Such network of galleries was spatially arranged according to both the spatial distribution and abundance of P. heptaphyllum trees in a way that most gallery entrances were disposed beneath or in close periphery of P. heptaphyllum crowns. Our findings suggest that underground trail systems shaped by fruit resources represent a foraging strategy clearly more common than existing literature on the subject would suggest. In addition, it reinforces the notion that the spatiotemporal availability of resources combined with predation risk largely influence trail configurations as well as overall foraging strategies adopted by leaf-cutting ants.     

Does leaf quality mediate the stimulation of leaf breakdown by phosphorus in Neotropical streams?

1. Lowland tropical streams have a chemically diverse detrital resource base, where leaf quality could potentially alter the effect of high nutrient concentrations on leaf breakdown. This has important implications given the extent and magnitude of anthropogenic nutrient loading to the environment. 2. Here, we examine if leaf quality (as determined by concentrations of cellulose, lignin and tannins) mediates the effects of high ambient phosphorus (P) concentration on leaf breakdown in streams of lowland Costa Rica. We hypothesised that P would have a stronger effect on microbial and insect processing of high‐ than of low‐quality leaves. 3. We selected three species that represented extremes of quality as measured in leaves of eight common riparian species. Species selected were, from high‐ to low‐quality: Trema integerrima > Castilla elastica > Zygia longifolia. We incubated single‐species leaf packs in five streams that had natural differences in ambient P concentration (10–140 μg soluble reactive phosphorus (SRP) L^?1), because of variable inputs of solute‐rich groundwater and also in a stream that was experimentally enriched with P (approximately 200 μg SRP L^?1). 4. The breakdown rate of all three species varied among the six streams: T. integerrima (k‐values range: 0.0451–0.129 day^?1); C. elastica (k‐values range: 0.0064–0.021 day^?1); and Z. longifolia (k‐values range: 0.002–0.008 day^?1). Both ambient P concentration and flow velocity had significant effects on the breakdown rate of the three species. 5. Results supported our initial hypothesis that litter quality mediates the effect of high ambient P concentration on leaf processing by microbes and insects. The response of microbial respiration, fungal biomass and invertebrate density to high ambient P concentration was greater in Trema (high quality) than in Castilla or Zygia (low quality). Variation in flow velocity, however, confounded our ability to determine the magnitude of stimulation of breakdown rate by P. 6. Cellulose and lignin appeared to be the most important factors in determining the magnitude of P‐stimulation. Surprisingly, leaf secondary compounds did not have an effect. This contradicts predictions made by other researchers, regarding the key role of plant secondary compounds in affecting leaf breakdown in tropical streams.     

Effects of forest types on leaf functional traits and their interrelationships of Pinus massoniana coniferous and broad‐leaved mixed forests in the subtropical mountain,Southeastern China

Leaf functional traits are widely used to detect and explain adaptations that enable plants to live under various environmental conditions. This study aims to determine the difference in leaf functional traits among four forest types of Pinus massoniana coniferous and broad‐leaved mixed forests by leaf morphological, nutrients, and stoichiometric traits in the subtropical mountain, Southeastern China. Our study indicated that the evergreen conifer species of P. massoniana had higher leaf dry matter content (LDMC), leaf C content, C/N and C/P ratios, while the three deciduous broad‐leaved species of L. formosana, Q. tissima, and P. strobilacea had higher specific leaf area (SLA), leaf N, leaf P nutrient contents, and N/P ratio in the three mixed forest types. The results showed that the species of P. massoniana has adapted to the nutrient‐poor environment by increasing their leaf dry matter for higher construction costs thereby reducing water loss and reflects a resource conservation strategy. In contrast, the three species of L. formosana, Q. tissima, and P. strobilacea exhibited an optimized resource acquisition strategy rather than resource conservation strategy in the subtropical mountain of southeastern China. Regarding the four forest types, the three mixed forest types displayed increased plant leaf nutrient contents when compared to the pure P. massoniana forest, especially the P. massoniana–L. formosana mixed forest type (PLM). Overall, variation in leaf functional traits among different forest types may play an adaptive role in the successful survival of plants under diverse environments because leaf functional traits can lead to significant effects on leaf function, especially for their acquisition of nutrients and use of light. The results of this study are beneficial to reveal the changes in plant leaf functional traits at the regional scale, which will provide a foundation for predicting changes in leaf traits and adaptation in the future environment.     

Effects of nutrient enrichment and leaf quality on the breakdown of leaves in a hardwater stream

1. The breakdown of leaf litter in streams is influenced strongly by leaf quality and the concentration of dissolved nutrients, primarily inorganic nitrogen (N) and phosphorus (P) in the water. We examined the effect of nutrient enrichment on the breakdown of three species of leaves in a hardwater, nutrient‐rich stream. The rate of microbial respiration was also measured on the decomposing leaves. 2. The breakdown rates of dogwood (Cornus stolonifera), aspen (Populus tremuloides) and birch (Betula occidentalis), k‐values of 0.0461, 0.0307 and 0.0186 day^–1, respectively, were unaffected by nutrient enrichment and generally faster than reported previously. Microbial respiration on the leaves was greater than reported previously for leaves of congeneric species. It appears that leaf breakdown in the study stream was not nutrient limited. 3. Nitrogen‐based measures of leaf quality, such as percentage N and carbon (C)/nitrogen ratio, did not correspond to measured breakdown rates among the three leaf types. The best predictors of relative breakdown rates were percentage lignin and the percentage of the total carbon that occurred as lignin. We suggest that, when leaf breakdown is not nutrient limited, measures of carbon quality (i.e. lignin‐based measures) are a better assessment of overall leaf quality than are N‐based measures. 4. Previous studies have indicated that the enzymes produced by aquatic hyphomycetes (microfungi) operate most efficiently at a basic pH and in the presence of calcium ions. The hardwater conditions (pH=8.6, total hardness > 300 mg CaCO₃ L^–1) and abundance of dissolved NO₃ and soluble reactive phosphorous (SRP) (approximately 50 μg L^–1, each) in the study stream appear to have provided conditions that resulted in a high respiration rate and rapid breakdown of leaf litter.     

Litter decomposition in a Cerrado savannah stream is retarded by leaf toughness, low dissolved nutrients and a low density of shredders

1. To assess whether the reported slow breakdown of litter in tropical Cerrado streams is due to local environmental conditions or to the intrinsic leaf characteristics of local plant species, we compared the breakdown of leaves from Protium brasiliense, a riparian species of Cerrado (Brazilian savannah), in a local and a temperate stream. The experiment was carried out at the time of the highest litter fall in the two locations. An additional summer experiment was conducted in the temperate stream to provide for similar temperature conditions. 2. The breakdown rates (k) of P. brasiliense leaves in the tropical Cerrado stream ranged from 0.0001 to 0.0008 day⁻¹ and are among the slowest reported. They were significantly (F = 20.12, P < 0.05) lower than in the temperate stream (0.0046–0.0055). The maximum ergosterol content in decomposing leaves in the tropical Cerrado stream was 106 μg g⁻¹, (1.9% of leaf mass) measured by day 75, which was lower than in the temperate stream where maximum ergosterol content of 522 μg g⁻¹ (9.5% of leaf mass) was achieved by day 30. The ATP content, as an indicator of total microbial biomass, was up to four times higher in the tropical Cerrado than in the temperate stream (194.0 versus 49.4 nmoles g⁻¹). 3. Unlike in the temperate stream, leaves in the tropical Cerrado were not colonised by shredder invertebrates. However, in none of the experiments did leaves exposed (coarse mesh bags) and unexposed (fine mesh bags) to invertebrates differ in breakdown rates (F = 1.15, P > 0.05), indicating that invertebrates were unable to feed on decomposing P. brasiliense leaves. 4. We conclude that the slow breakdown of P. brasiliense leaves in the tropical Cerrado stream was because of the low nutrient content in the water, particularly nitrate (0.05 mgN L⁻¹), which slows down fungal activity and to the low density of invertebrates capable of using these hard leaves as an energy source.     

Comparative petiole anatomy of the tribe Sorbarieae (Rosaceae) provide new taxonomically informative characters

We conducted a comparative anatomical study of the petiole of 16 taxa belonging to the tribe Sorbarieae (Rosaceae) (Adenostoma, 2 spp.; Chamaebatiaria, 1 sp.; Sorbaria, 6 spp., 3 vars., and 1 forma; and Spiraeanthus, 1 sp.) and the related genus Lyonothamnus (1 sp. and 1 ssp.). The distal, medial and proximal regions of petioles were transversely sectioned using conventional embedding and staining methods. Cuticles, crystals, trichomes and pericyclic fiber patterns were observed and studied. Three types of vascular nodal patterns were recognized: Type 1 was seen in Chamaebatiaria, Lyonothamnus, and Spiraeanthus (simple‐trace nodal pattern with slightly curved or U‐shaped vascular bundle); type 2 was found in Adenostoma (multiple‐traces nodal pattern with free vascular bundles); and type 3 was unique to Sorbaria (bundles fused to form a siphonostele nodal pattern). Some petiolar anatomical characteristics (e.g. cuticles, crystals, trichomes, vascular nodal pattern, and pericyclic fiber patterns) were found to provide useful information for taxonomic studies within Sorbarieae. On the basis of these characteristics, a dichotomous key for identification at the generic/specific level is provided. We also report a structural change in the vascular bundles from the stem‐leaf transitional zone to the leaf medial zone.     

Leaf essential oil composition of 10 species of Ocotea (Lauraceae) from Monteverde,Costa Rica

The leaf essential oils of 10 species of Ocotea (Lauraceae) from Monteverde, Costa Rica (Ocotea floribunda, Ocotea holdridgeana, Ocotea meziana, Ocotea sinuata, Ocotea tonduzii, Ocotea valeriana, Ocotea veraguensis, Ocotea whitei, and two undescribed species, Ocotea new species “los llanos”, and Ocotea new species “small leaf”) have been obtained by hydrodistillation and analyzed by GC–MS in order to discern the differences and similarities between the volatile chemical compositions of these species. The principal common constituents of the 10 species of Ocotea were α-pinene, β-pinene, β-caryophyllene, and germacrene-D.     

Identification and population dynamics of bacteria in symptomatic oat leaves

Bacteria isolated from symptomatic oat leaves included pseudomonads,Erwinia herbicola, and others.Pseudomonas coronafaciens was isolated predominantly from leaves with halo blight symptoms or necrotic spots. Leaves with red leaf symptoms yielded many types of bacteria, including saprophytic pseudomonads,P. syringae, E. herbicola, Bacillus sp.,Micrococcus sp.,Corynebacterium sp., a yeast, and other unidentified species. Only isolates ofP. coronafaciens were pathogenic on the plant hosts tested. The bacteria associated with red leaf symptoms exist as saprophytes and/or epiphytes on leaves with those symptoms. It is concluded that bacteria do not contribute to red leaf symptom development in oats, but symptomatic leaves provide an environment for their growth.     

Habitat structure and fish predation: effects on invertebrate colonisation and breakdown of stream leaf packs

1. We investigated the effects of two features of leaf‐pack habitat structure (i.e. mass of a leaf pack and surface area of leaves comprising a leaf pack) and fish predation on colonisation of shredders and leaf breakdown rates in a coldwater stream. Packs were constructed of red maple (Acer rubrum) leaves. 2. A 2 × 3 × 3 factorial experiment was used to manipulate fish predation (exclusion and control cage), leaf‐pack mass (1, 3 and 5 g dry mass) and leaf surface area (small: approx. 17.9 cm², medium: approx. 34.6 cm², large: approx. 65.6 cm²). Exclusion cages had mesh on all sides, whereas control cages lacked mesh on two sides to provide access to fish. 3. Common shredders were Gammarus pseudolimnaeus, Pycnopsyche and Lepidostoma. Shredder biomass per leaf pack increased with the mass of a leaf pack (P < 0.001), but biomass per unit mass of leaf pack did not differ with leaf‐pack mass (P = 0.506). Shredder densities did not respond to the exclusion of fish (P > 0.7) or leaf surface area (P > 0.7), and interactions among treatment factors were not significant (P > 0.2). 4. Breakdown rates were lower for leaf packs comprised of small leaves (P < 0.001) and leaf packs with high mass (P = 0.001). Excluding fish did not significantly affect leaf breakdown rates (P = 0.293), and interactions among treatment factors were not significant (P > 0.3). Breakdown rates were highest when packs consisted of few leaves (i.e. leaf packs with large leaves and low mass) and were colonised by many shredders. 5. Fish predation was not an important factor controlling shredder densities in leaf packs over the spatiotemporal scale of our experiment. Nevertheless, we found shredder colonisation was proportional to leaf‐pack mass and breakdown rates were affected by leaf‐pack size (i.e. number of leaves in a pack). We suspect that fragmentation is the primary mechanism causing the breakdown rates to be dependent on leaf‐pack size.     

The Effect of Macroinvertebrate Exclusion on Leaf Breakdown Rates in a Tropical Headwater Stream1

The role of macroinvertebrates in the process of leaf breakdown is well studied in temperate streams, but less is known about their role in the tropics. We investigated the effect of reducing macroinvertebrate access to leaf material on leaf breakdown rates in a forested headwater stream in the Luquillo Experimental Forest, Puerto Rico. We measured leaf mass loss using fine and coarse mesh bags over 12 weeks for two common riparian species: Cecropia schreberiana (Moraceae) and Dacryodes excelsa (Burseraceae). Coarse mesh allowed freshwater shrimp and other macroinvertebrates to access leaf material, while fine mesh did not. Leaf breakdown rates did not differ between C. schreberiana and D. excelsa in coarse mesh bags (?0.0375/day vs. ?0.0395/day, respectively), but C. schreberiana breakdown was significantly slower than D. excelsa in fine mesh bags (?0.0159/day vs. ?0.0266/day). Additionally, breakdown in fine mesh bags was significantly slower compared to coarse mesh bags for C. schreberiana, but less so for D. excelsa. Breakdown rates for all treatments were fast relative to those in temperate‐zone streams indicating that both macroinvertebrates and macroinvertebrate‐independent processing can strongly influence leaf decomposition in tropical streams. The difference between C. schreberiana and D. excelsa indicates that the effect of macroinvertebrate exclusion can change with leaf type.     

Two new species of Protium from French Guiana. Studies in neotropical Burseraceae VIII

Revision of Protium in French Guiana has revealed two new species. Protium gallicum is related to P. aracouchini (Aubl.) Marchand and P. elegans Engl., although its fruits resemble those of P. plagiocarpium Bénoist. Protium inodorum, which also occurs in northern Pará, Brazil, is related to P. pallidum Cuatrec. and P. cuneatum Swart; it is known only from flowers and very young fruits. Even in relatively well-known regions of tropical forest, botanical exploration continues to yield new taxa.     

Does nitrogen become limiting under high‐P conditions in detritus‐based tropical streams?

1. We examined effects of nutrients on leaf breakdown in interior forest streams at La Selva Biological Station, Costa Rica. We tested the hypothesis that dissolved inorganic nitrogen (DIN) becomes limiting when ambient phosphorus (P) concentration is high. We also compared the breakdown of relatively ‘low quality’ leaves (lower C : N, Trema integerrima) with that of ‘higher quality’ leaves (higher C : N, Ficus insipida) in a high‐P stream. 2. Litterbags were incubated in two streams: one enriched experimentally with P [target concentration 200 μg soluble reactive phosphorus (SRP) L^?1] and one control (naturally low P concentration approximately 10 μg SRP L^?1). Ammonium enrichment was achieved by adding fertiliser upstream of half of the litterbags in each stream. 3. Phosphorus addition stimulated leaf breakdown, microbial respiration, ergosterol and leaf %P. Leaf breakdown rate was consistent with those in La Selva streams with naturally high P concentration. 4. Nitrogen (N) addition had no effect on leaf breakdown, microbial respiration, ergosterol or leaf chemistry in either the P‐enriched or the reference stream, in spite of low N : P ratios. We conclude that N is probably not limiting in streams at La Selva that are naturally high in P. This may be due to moderately high ambient N concentration (>200 μg DIN L^?1) prevailing throughout the year. 5. The species with a lower C : N decomposed more rapidly and supported higher microbial activity than that with a higher C : N. Subtle differences in leaf N content, as well as dissolved P concentration, may be important in determining microbial colonisation and subsequent leaf breakdown.     

A comparative study of aluminium and nutrient concentrations in mistletoes on aluminium‐accumulating and non‐accumulating hosts

Mistletoes offer a unique model to study interactions among Al and nutrients in vascular plants, because they grow and reproduce on hosts with distinct Al uptake strategies. We investigated Al distribution and nutrient relations of mistletoes on Al‐accumulating and non‐accumulating hosts. We hypothesised that mistletoes would exhibit similar leaf nutrient and Al concentrations as their host plants, but a strong compartmentalisation of Al when growing on Al‐accumulators. We measured concentrations of N, P, K, Ca, Mg, Cu, Fe, Mn, Zn in leaves and Al in leaves, seeds and branches of Phthirusa ovata and Psittacanthus robustus infecting Miconia albicans, an Al‐accumulator, and Ph. ovata infecting Byrsonima verbascifolia, a non‐Al‐accumulator. High leaf concentrations of Al in Ph. ovata only occurred while parasitizing the Al‐accumulating host; there was no accumulation in branches or seeds. In P. robustus, large concentrations of Al were found in leaves, branches and seeds. Mistletoe seed viability and leaf nutrient concentrations were not affected by Al accumulation. Passive uptake of Al, Ca, Mg, Mn and Cu in mistletoes was evidenced by significant correlations between mistletoes and host leaf concentrations, but not of N, P and K. Al was retranslocated to different plant organs in P. robustus, whereas it was mostly restricted to leaves in Ph. ovata. We suggest that Al might have some specific function in P. robustus, which only parasitizes Al‐accumulator hosts, while the host generalist Ph. ovata can be considered a facultative Al‐accumulator.     

The Impact of Fungal Extracts on Leaf Litter on the Food Preference of Gammarus roeselii

We investigated the effect of methanol and methanol/methylene chloride extracts of the oomycete Pythium sp. JN 1‐b and of the fungi Ascomycete sp. PVSo8, Fusarium sporotrichoides, and Cylindrocarpon sp. 94‐2057 on the food preference of Gammarus roeselii. The preference for leaf discs coated with these extracts compared to uncoated leaf discs was tested in food‐choice assays. Methanol extracts of all strains repelled G. roeselii, and the effect of the extract concentration on relative consumption was strain specific. The repellent effect of these extracts, especially of extracts of Cylindrocarpon sp., decreased when the fungi were grown on leaf extract medium as opposed to synthetic medium containing sucrose. None of the methanol/methylene chloride extracts affected the food preference of the gammarid. We conclude that biologically active compounds were extracted from fungi and an oomycete were soluble in methanol but not in methanol/methylene chloride. Only repellent activity was observed with the extracts, and relative ratios of repellents and attractants might determine the consumption of fungi by G. roeselii. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The importance of environmental heterogeneity and spatial distance in generating phylogeographic structure in edaphic specialist and generalist tree species of Protium (Burseraceae) across the Amazon Basin

Aim Edaphic heterogeneity may be an important driver of population differentiation in the Amazon but remains to be investigated in trees. We compared the phylogeographic structure across the geographic distribution of two Protium (Burseraceae) species with different degrees of edaphic specialization: Protium alvarezianum, an edaphic specialist of white‐sand habitat islands; and Protium subserratum, an edaphic generalist found in white sand as well as in more widespread soil types. We predicted that in the edaphic specialist, geographic distance would structure populations more strongly than in the edaphic generalist, and that soil type would not structure populations in the edaphic generalist unless habitat acts as a barrier promoting population differentiation. Location Tropical rain forests of the Peruvian and Brazilian Amazon, Guyana and French Guiana. Methods We sequenced 1209–1211 bp of non‐coding nuclear ribosomal DNA (internal transcribed spacer and external transcribed spacer) and a neutral low‐copy nuclear gene (phytochrome C) from P. subserratum (n = 65, 10 populations) and P. alvarezianum (n = 19, three populations). We conducted a Bayesian phylogenetic analysis, constructed maximum parsimony haplotype networks and assessed population differentiation among groups (soil type or geographic locality) using analysis of molecular variance and spatial analysis of molecular variance. Results The edaphic specialist exhibited considerable genetic differentiation among geographically distant populations. The edaphic generalist showed significant genetic differentiation between the Guianan and Amazon Basin populations. Within Peru, soil type and not geographic distance explained most of the variation among populations. Non‐white‐sand populations in Peru exhibited lower haplotype/nucleotide diversity than white‐sand populations, were each other’s close relatives, and formed an unresolved clade derived from within the white‐sand populations. Main conclusions Geographic distance is a stronger driver of population differentiation in the edaphic specialist than in the generalist. However, this difference did not appear to be related to edaphic generalism per se as adjacent populations from both soil types in the edaphic generalist did not share many haplotypes. Populations of the edaphic generalist in white‐sand habitats exhibited high haplotype diversity and shared haplotypes with distant white‐sand habitat islands, indicating that they have either efficient long‐distance dispersal and/or larger ancestral effective population sizes and thus retain ancestral polymorphisms. These results highlight the importance of edaphic heterogeneity in promoting population differentiation in tropical trees.     

Evidence for ecological divergence across a mosaic of soil types in an Amazonian tropical tree: Protium subserratum (Burseraceae)

Soil heterogeneity is an important driver of divergent natural selection in plants. Neotropical forests have the highest tree diversity on earth, and frequently, soil specialist congeners are distributed parapatrically. While the role of edaphic heterogeneity in the origin and maintenance of tropical tree diversity is unknown, it has been posited that natural selection across the patchwork of soils in the Amazon rainforest is important in driving and maintaining tree diversity. We examined genetic and morphological differentiation among populations of the tropical tree Protium subserratum growing parapatrically on the mosaic of white‐sand, brown‐sand and clay soils found throughout western Amazonia. Nuclear microsatellites and leaf morphology were used to (i) quantify the extent of phenotypic and genetic divergence across habitat types, (ii) assess the importance of natural selection vs. drift in population divergence, (iii) determine the extent of hybridization and introgression across habitat types, (iv) estimate migration rates among populations. We found significant morphological variation correlated with soil type. Higher levels of genetic differentiation and lower migration rates were observed between adjacent populations found on different soil types than between geographically distant populations on the same soil type. P_ST‐F_ST comparisons indicate a role for natural selection in population divergence among soil types. A small number of hybrids were detected suggesting that gene flow among soil specialist populations may occur at low frequencies. Our results suggest that edaphic specialization has occurred multiple times in P. subserratum and that divergent natural selection across edaphic boundaries may be a general mechanism promoting and maintaining Amazonian tree diversity.