Anatomical, histochemical, and developmental approaches reveal the long-term functioning of the floral nectary in Tocoyena formosa (Rubiaceae).

Tocoyena formosa has a persistent floral nectary that continues producing nectar throughout flower and fruit development. This plant also presents an intriguing non-anthetic nectary derived from early-developing floral buds with premature abscised corolla. In this study, we characterize the structure, morphological changes, and functioning of T. formosa floral nectary at different developmental stages. We subdivided the nectary into four categories based on the floral and fruit development stage at which nectar production started: (i) non-anthetic nectary; (ii) anthetic nectary, which follows the regular floral development; (iii) pericarpial nectary, derived from pollinated flowers following fruit development; and (iv) post-anthetic nectary that results from non-pollinated flowers after anthesis. The nectary has a uniseriate epidermis with stomata, nectariferous parenchyma, and vascular bundles, with a predominating phloem at the periphery. The non-anthetic nectary presents immature tissues that release the exudate. The nectary progressively becomes more rigid as the flower and fruit develop. The main nectary changes during flower and fruit development comprised the thickening of the cuticle and epidermal cell walls, formation of cuticular epithelium, and an increase in the abundance of calcium oxalate crystals and phenolic cells near the vascular bundles. Projections of the outer periclinal walls toward the cuticle in the post-anthetic nectary suggest nectar reabsorption. The anatomical changes of the nectary allow it to function for an extended period throughout floral and fruit development. Hence, T. formosa nectary is a bivalent secretory structure that plays a crucial role in the reproductive and defensive interactions of this plant species.

Tradeoffs in the evolution of plant farming by ants.

Diverse forms of cultivation have evolved across the tree of life. Efficient farming requires that the farmer deciphers and actively promotes conditions that increase crop yield. For plant cultivation, this can include evaluating tradeoffs among light, nutrients, and protection against herbivores. It is not understood if, or how, nonhuman farmers evaluate local conditions to increase payoffs. Here, we address this question using an obligate farming mutualism between the ant Philidris nagasau and epiphytic plants in the genus Squamellaria that are cultivated for their nesting sites and floral rewards. We focused on the ants' active fertilization of their crops and their protection against herbivory. We found that ants benefited from cultivating plants in full sun, receiving 7.5-fold more floral food rewards compared to shade-cultivated plants. The higher reward levels correlated with higher levels of crop protection provided by the ants. However, while high-light planting yielded the greatest immediate food rewards, sun-grown crops contained less nitrogen compared to shade-grown crops. This was due to lower nitrogen input from ants feeding on floral rewards instead of insect protein gained from predation. Despite this tradeoff, farming ants optimize crop yield by selectively planting their crops in full sun. Ancestral state reconstructions across this ant-plant clade show that a full-sun farming strategy has existed for millions of years, suggesting that nonhuman farmers have evolved the means to evaluate and balance conflicting crop needs to their own benefit.

Changes in cambial activity are related to precipitation patterns in four tropical hardwood species grown in Indonesia.

PREMISE: Cambial activity in some tropical trees varies intra-annually, with the formation of xylem rings. Identification of the climatic factors that regulate cambial activity is important for understanding the growth of such species. We analyzed the relationship between climatic factors and cambial activity in four tropical hardwoods, Acacia mangium, Tectona grandis, Eucalyptus urophylla, and Neolamarckia cadamba in Yogyakarta, Java Island, Indonesia, which has a rainy season (November-June) and a dry season (July-October). METHODS: Small blocks containing phloem, cambium, and xylem were collected from main stems in January 2014, October 2015 and October 2016, and examined with light microscopy for cambial cell division, fusiform cambial cells, and expanding xylem cells as evidence of cambial activity. RESULTS: During the rainy season, when precipitation was high, cambium was active. By contrast, during the dry season in 2015, when there was no precipitation, cambium was dormant. However, in October 2016, during the so-called dry season, cambium was active, cell division was conspicuous, and a new xylem ring formation was initiated. The difference in cambial activity appeared to be related to an unusual pattern of precipitation during the typically dry months, from July to October, in 2016. CONCLUSIONS: Our results indicate that low or absent precipitation for 3 to 4 months induces cessation of cambial activity and temporal periodicity of wood formation in the four species studied. By contrast, in the event of continuing precipitation, cambial activity in the same trees may continue throughout the year. The frequency pattern of precipitation appears to be an important determinant of wood formation in tropical trees.

Drying and reduction in sensitivity to desiccation of seeds of Alibertia edulis: the influence of fruit ripening stage.

The intense environmental degradation in tropical regions suitable for agriculture has decreased native forest populations of plants with important fruits and medicinal properties. Alibertia edulis is a native tree from the Brazilian Cerrado. Knowledge about the effects of drying and storage on the physiological behavior of its seeds may aid in its sustainable exploitation and conservation. The goal of the present study was to determine which fruit ripening stage results in A. edulis seeds with higher tolerance to desiccation, and to investigate the effectiveness of polyethylene glycol (PEG) induced osmotic stress in combination with abscisic acid (ABA) in increasing seed desiccation tolerance during storage. Seeds were dried in activated silica gel (fast) or under ambient conditions (slow). Seeds originating from mid-ripe or fully ripe fruits exhibited better physiological performance than those obtained from green fruits. Slow drying resulted in seeds with high physiological potential. Seeds may be stored for up to 180 days without losing viability when treated with -0.73 MPa PEG without ABA.

Colleters in Rubiaceae from forest and savanna: the link between secretion and environment.

This study aims to investigate colleters' secretory function, on cellular level, in Rubiaceae species from contrasting environments looking to explore the association between secretion and environment. We collected samples from eight species of Rubiaceae growing in forest and savanna having standard-type colleters with diverse histochemistry (hydrophilic, lipophilic and mixed secretions) and processed for both conventional and cytochemical study under transmission electron microscopy (TEM). The standard colleters, although similar in morphology and anatomy, exhibited marked differences on cellular level, especially in the abundance and topology of Golgi bodies, endoplasmic reticulum and plastids when comparing forest and savanna species. These differences were clearly aligned with the chemical nature of the secretions they produce, with predominance of hydrophilic secretions in forest species and lipophilic or mixed secretions in savanna species. The combination of methods in electron microscopy revealed the sites of synthesis and intracellular compartmentation of substances, the mechanisms of their secretion from the protoplast and confirmed the involvement of the outer walls of the epithelial cells in the elimination of exudates to the gland surface. Our study suggests a potential environment-associated plasticity of the secretory cells of standard-type colleters in modulating their secretory function performance.

Partner choice through concealed floral sugar rewards evolved with the specialization of ant-plant mutualisms.

Obligate mutualisms require filtering mechanisms to prevent their exploitation by opportunists, but ecological contexts and traits facilitating the evolution of such mechanisms are largely unknown. We investigated the evolution of filtering mechanisms in an epiphytic ant-plant symbiotic system in Fiji involving Rubiaceae and dolichoderine ants, using field experiments, metabolomics, X-ray micro-computed tomography (micro-CT) scanning and phylogenetics. We discovered a novel plant reward consisting of sugary sap concealed in post-anthetic flowers only accessible to Philidris nagasau workers that bite through the thick epidermis. In five of the six species of Rubiaceae obligately inhabited by this ant, the nectar glands functioned for 10 d after a flower's sexual function was over. Sugar metabolomics and field experiments showed that ant foraging tracks sucrose levels, which only drop at the onset of fruit development. Ontogenetic analyses of our focal species and their relatives revealed a 25-fold increase in nectary size and delayed fruit development in the ant-rewarding species, and Bayesian analyses of several traits showed the correlated evolution of sugar rewards and symbiosis specialization. Concealed floral nectar forestalls exploitation by opportunists (generalist ants) and stabilizes these obligate mutualisms. Our study pinpoints the importance of partner choice mechanisms in transitions from facultative to obligate mutualisms.

Neighbourhood structure and light availability influence the variations in plant design of shrubs in two cloud forests of different successional status.

BACKGROUND AND AIMS: Plant design refers to the construction of the plant body or its constituent parts in terms of form and function. Although neighbourhood structure is recognized as a factor that limits plant survival and species coexistence, its relative importance in plant design is not well understood. We conducted field research to analyse how the surrounding environment of neighbourhood structure and related effects on light availability are associated with changes in plant design in two understorey plants (Palicourea padifolia and Psychotria elata) within two successional stages of a cloud forest in Costa Rica. METHODS: Features of plant neighbourhood physical structure and light availability, estimated using hemispherical photographs, were used as variables that reflect the surrounding environment. Measures of plant biomechanics, allometry, branching and plant slenderness were used as functional plant attributes that reflect plant design. We propose a framework using a partial least squares path model and used it to test this association. KEY RESULTS: The multidimensional response of plant design of these species suggests that decreases in the height-based factor of safety and increases in mechanical load and developmental stability are influenced by increases in maximum height of neighbours and a distance-dependence interference index more than neighbourhood plant density or neighbour aggregation. Changes in plant branching and slenderness are associated positively with light availability and negatively with canopy cover. CONCLUSIONS: Although it has been proposed that plant design varies according to plant density and light availability, we found that neighbour size and distance-dependence interference are associated with changes in biomechanics, allometry and branching, and they must be considered as key factors that contribute to the adaptation and coexistence of these plants in this highly diverse forest community.

Reciprocal herkogamy promotes disassortative mating in a distylous species with intramorph compatibility.

Mating patterns in heterostylous species with intramorph compatibility have the potential to deviate from symmetrical disassortative mating owing to ecological and reproductive factors influencing pollen dispersal. Here, we investigate potential and realized patterns of mating in distylous Luculia pinceana (Rubiaceae), a species with intramorph compatibility. Our analysis provides an opportunity to test Darwin's hypothesis that reciprocal herkogamy promotes disassortative pollen transfer. We combined measurements of sex-organ reciprocity and pollen production to predict potential pollen transfer and mating patterns in a population from SW China. Marker-based paternity analysis was then used to estimate realized patterns of disassortative and assortative mating at the individual and floral morph levels. Both potential and realized mating patterns indicated a significant component of disassortative mating, satisfying theoretical conditions for the maintenance of floral dimorphism. Levels of assortative mating (37.7%) were significantly lower than disassortative mating (62.3%), but numerous offspring resulting from intramorph mating were detected in the majority of maternal seed families in both floral morphs. Our results provide empirical support for Darwin's cross-promotion hypothesis on the function of reciprocal herkogamy, but indicate that in most heterostylous species strong diallelic incompatibility may be a general requirement for complete disassortative mating.

Reproductive isolation between sympatric sister species, Mussaenda kwangtungensis and M. pubescens var. alba.

Reproductive isolation defines the biological species concept and plays a key role in the formation and maintenance of species. The relative contributions of different isolating stages has been suggested to be closely associated with phylogenetic relatedness. Few studies have focused on the relative contributions of pre- versus post-zygotic mechanisms, and even fewer have been conducted under strict phylogenetic frameworks. Pre- and post-zygotic reproductive isolation stages have been investigated in the sister species Mussaenda kwangtungensis and M. pubescens var. alba. The two species have partly overlapping distribution ranges and flowering times, while the principal pollinators differed strikingly for them, demonstrating strong pre-zygotic isolations. Natural hybrids were detected by simple sequence repeat markers and their maternal parents were identified based on chloroplast gene sequences. Five out of 81 individuals were suggested to be hybrids that fall into the categories F2, BC1, and BC2 by the NewHybrids analysis. Interspecific crossings resulted in significantly reduced fruit set and seed germination rates. Phylogenetic analysis revealed short Kimura-2-parameter distance between M. kwangtungensis and M. pubescens var. alba. These findings strongly supported the hypothesis that for species with a closer phylogenetic relationship, pre-zygotic isolation plays an important part in limiting gene exchange in sympatric areas.

Pre- and post-zygotic reproductive isolation between co-occurring Mussaenda pubescens var. alba and M. shikokiana (Rubiaceae).

Reproductive isolation is a fundamental requirement for speciation and includes several sequential stages. Few studies have determined the relative contributions of pre- and post-zygotic reproductive isolation in plants, especially between relative species with clear differentiation in flower form. To investigate the mechanisms responsible for reproductive isolation in sympatric Mussaenda pubescens var. alba and Mussaenda shikokiana (Rubiaceae) in Guangxi Province, China, we made observations of flowering phenology, patterns of insect visitation, and conducted pollination experiments, including artificial hybridization. The two species had overlapping flowering times and were pollinated by overlapping pollinators; however, their relative importance differed significantly with M. pubescens visited more commonly by bees and M. shikokiana more frequently by butterflies. Using vegetative and floral characters and molecular evidence based on nuclear ribosomal internal and external transcribed spacer regions we detected seven naturally occurring hybrids among a sample of approximately 125 individuals. Hybrids were characterized by morphologies that most closely resembled their maternal parents based on chloroplast evidence. Studies of artificially synthesized and natural hybrids demonstrated that hybrid seed had very low germination rates and naturally occurring hybrids exhibited pollen sterility. Post-zygotic reproductive isolating mechanisms play a primary role in limiting gene exchange between co-occurring species and maintaining species integrity in areas of sympatry.

Whole-plant and leaf determinants of growth rates in progenies of Genipa americana L. (Rubiaceae).

Genipa americana (Rubiaceae) is a fruit tree with broad phytogeographic domain and suitable for different silvicultural systems in the tropics. The knowledge associated with the relative growth rate of species such as G. americana, provides important guidelines for the effective establishment and survival of seedlings after planting in the field. In this study we investigated differences in growth, biomass allocation and photosynthesis of seedlings originating from different mother plants of G. americana in southern Bahia, Brazil. For this, we evaluated fifteen variables associated with carbon balance at the whole plant and leaf scales of twelve G. americana progenies. All seedlings grew over a period of 198 days under similar microclimatic conditions with approximately 65% full sun. Our results showed significant differences in the relative growth rates (RGR), with the highest and lowest mean values being 29.0 and 38.0 mg g-1 day-1, respectively. Differences in RGR between G. americana progenies were highly related to differences in biomass allocation at both whole plant and leaf scales. From a practical point of view, we demonstrate that the selection of mother plants to produce seedlings with higher growth rates, and consequently greater establishment capacity in field plantings, can be made from evaluations of growth and biomass allocation variables at the whole plant scale.

Long-term strict ant-plant mutualism identity characterises growth rate and leaf shearing resistance of an Amazonian myrmecophyte.

Over 125 million years of ant-plant interactions have culminated in one of the most intriguing evolutionary outcomes in life history. The myrmecophyte Duroia hirsuta (Rubiaceae) is known for its mutualistic association with the ant Myrmelachista schumanni and several other species, mainly Azteca, in the north-western Amazon. While both ants provide indirect defences to plants, only M. schumanni nests in plant domatia and has the unique behaviour of clearing the surroundings of its host tree from heterospecific plants, potentially increasing resource availability to its host. Using a 12-year survey, we asked how the continuous presence of either only M. schumanni or only Azteca spp. benefits the growth and defence traits of host trees. We found that the continuous presence of M. schumanni improved relative growth rates and leaf shearing resistance of Duroia better than trees with Azteca. However, leaf herbivory, dry matter content, trichome density, and secondary metabolite production were the same in all trees. Survival depended directly on ant association (> 94% of trees died when ants were absent). This study extends our understanding of the long-term effects of strict ant-plant mutualism on host plant traits in the field and reinforces the use of D. hirsuta-M. schumanni as a model system suitable for eco-co-evolutionary research on plant-animal interactions.

A comparative analysis of the mechanisms underlying speciation on Lord Howe Island.

On Lord Howe Island, speciation is thought to have taken place in situ in a diverse array of distantly related plant taxa (Metrosideros, Howea and Coprosma; Proc. Natl Acad. Sci. USA 108, 2011, 13188). We now investigate whether the speciation processes were driven by divergent natural selection in each genus by examining the extent of ecological and genetic divergence. We present new and extensive, ecological and genetic data for all three genera. Consistent with ecologically driven speciation, outlier loci were detected using genome scan methods. This mechanism is supported by individual-based analyses of genotype-environment correlations within species, demonstrating that local adaptation is currently widespread on the island. Genetic analyses show that prezygotic isolating barriers within species are currently insufficiently strong to allow further population differentiation. Interspecific hybridization was found in both Howea and Coprosma, and species distribution modelling indicates that competitive exclusion may result in selection against admixed individuals. Colonization of new niches, partly fuelled by the rapid generation of new adaptive genotypes via hybridization, appears to have resulted in the adaptive radiation in Coprosma - supporting the 'Syngameon hypothesis'.

Morphology, ecophysiology and germination of seeds of the neotropical tree Alibertia patinoi (Rubiaceae).

Alibertia patinoi (Rubiaceae) is of economic and cultural importance for communities in the Colombian Pacific and Amazon regions, where it is cultivated and mature fruits are highly appreciated and consumed. Since there is a lack of knowledge of the seed physiology of this species, we describe here the germination behavior and morphometry of seeds of Alibertia patinoi, and relate them to its habitat. Fruits were collected from a mixed food crop and a commercial plantation in Guaimía village, Buenaventura, Colombia, a tropical rain forest area. We measured length, width, thickness, mass (n = 1 400), and moisture content of seeds (n = 252). Primary dormancy tests were conducted (n = 200), followed by imbibition (n=252) and germination dynamics, under different conditions of light and temperature specific to understory and forest clearings (n = 300 seeds). Finally, seed storage behavior was established (n = 100 seeds). We observed that size and mass of seeds had a narrow range of values that did not differ within or among fruits and that the species did not exhibit primary dormancy. The seeds are recalcitrant, and recently harvested seeds exhibited higher seed moisture content (ca. 44%) and continuous metabolism. The seed germination percentage was observed to be higher under the specific dense canopy forest light and temperature conditions; furthermore, neither enriched far-red light nor darkness conditions inhibited germination. We concluded that rapid germination could be the establishment strategy of this species. Also, the physiological traits (i.e., rapid germination rate, low germination requirements, absence of primary dormancy, and recalcitrant behavior) and seed size and mass, suggest that A. patinoi is adapted to conditions of mature tropical rain forests.

Neutral dynamics and cluster statistics in a tropical forest.

The neutral theory of biodiversity attributes community structure to the effects of chance alone, assuming that all species and individuals are demographically equivalent. Here we present a spatially explicit version of the neutral theory and test it against the Barro Colorado Island (BCI) data. Monitoring the dynamics of clusters, we show that the effect of local heterogeneities (e.g., microtopography) is weak, making a spatially homogenous model plausible. We then compare the cluster statistics of the three most frequent species with the patterns obtained from neutral dynamics, examining two families of recruitment kernels: one that interpolates between a limited distance and panmictic dispersal (local-global) and one that assumes a scale-free Cauchy kernel. The results rule out the local-global dispersal model and show that the spatial patterns fit very nicely those obtained from the fat-tailed kernel. Our work emphasizes the importance of spatiotemporal cluster dynamics as an instrument for detecting the factors that govern community assembly.

Combined impacts of irradiance and dehydration on leaf hydraulic conductance: insights into vulnerability and stomatal control.

The leaf is a hydraulic bottleneck, accounting for a large part of plant resistance. Thus, the leaf hydraulic conductance (K(leaf) ) is of key importance in determining stomatal conductance (g(s) ) and rates of gas exchange. Previous studies showed that K(leaf) is dynamic with leaf water status and irradiance. For four species, we tested the combined impacts of these factors on K(leaf) and on g(s) . We determined responses of K(leaf) and g(s) to declining leaf water potential (Ψ(leaf) ) under low and high irradiance (<6 and >900 µmol photons m(-2) s(-1) photosynthetically active radiation, respectively). We hypothesized greater K(leaf) vulnerability under high irradiance. We also hypothesized that K(leaf) and g(s) would be similar in their responses to either light or dehydration: similar light-responses of K(leaf) and g(s) would stabilize Ψ(leaf) across irradiances for leaves transpiring at a given vapour pressure deficit, and similar dehydration responses would arise from the control of stomata by Ψ(leaf) or a correlated signal. For all four species, the K(leaf) light response declined from full hydration to turgor loss point. The K(leaf) and g(s) differed strongly in their light- and dehydration responses, supporting optimization of hydraulic transport across irradiances, and semi-independent, flexible regulation of liquid and vapour phase water transport with leaf water status.

Identification of the bacterial endosymbionts in leaf nodules of Pavetta (Rubiaceae).

Three genera in the Rubiaceae (Pavetta, Psychotria and Sericanthe) harbour bacterial endosymbionts within leaf nodules or galls. The present paper identifies the bacterial endophytes in three leaf-nodulating Pavetta species. In order to reveal their identity and assess their phylogenetic position, 16S rRNA, recA and gyrB genes were sequenced from an extensive sampling of Burkholderia strains. This multigene approach results in a robust phylogeny, which places the bacterial endosymbionts of Pavetta at two distinct positions within the genus Burkholderia (class Betaproteobacteria), suggesting that leaf-nodulating endosymbionts within Pavetta have different origins. The endophytes of nodulated Psychotria species were recognized as the closest relatives to the Pavetta endosymbionts. Our results suggest that the endosymbionts of Pavetta represent novel species, which can be classified as 'Candidatus Burkholderia hispidae', 'Candidatus Burkholderia rigidae' and 'Candidatus Burkholderia schumannianae'.

Nutrients limit photosynthesis in seedlings of a lowland tropical forest tree species.

We investigated how photosynthesis by understory seedlings of the lowland tropical tree species Alseis blackiana responded to 10 years of soil nutrient fertilization with N, P and K. We ask whether nutrients are limiting to light and CO(2) acquisition in a low light understory environment. We measured foliar nutrient concentrations of N, P and K, isotopic composition of carbon (δ(13)C) and nitrogen (δ(15)N), and light response curves of photosynthesis and chlorophyll fluorescence. Canopy openness was measured above each study seedling and included in statistical analyses to account for variation in light availability. Foliar N concentration increased by 20% with N addition. Foliar P concentration increased by 78% with P addition and decreased by 14% with N addition. Foliar K increased by 8% with K addition. Foliar δ(13)C showed no significant responses, and foliar δ(15)N decreased strongly with N addition, matching the low δ(15)N values of applied fertilizer. Canopy openness ranged from 0.01 to 6.71% with a mean of 1.76 ± 0.14 (± 1SE). Maximum photosynthetic CO(2) assimilation rate increased by 9% with N addition. Stomatal conductance increased with P addition and with P and K in combination. Chlorophyll fluorescence measurements revealed that quantum yield of photosystem II increased with K addition, maximum electron transport rate trended 9% greater with N addition (p = 0.07), and saturating photosynthetically active radiation increased with N addition. The results demonstrate that nutrient addition can enhance photosynthetic processes, even under low light availability.

Phenology of tropical understory trees: patterns and correlates.

Reproductive phenologies of plants are constrained by climate in highly seasonal regions. In contrast, plants growing in wet tropical forests are freed from many abiotic constraints, which in canopy tree communities lead to a rich diversity of phenological patterns within and among individuals, species and communities. However, basic descriptions of tropical phenological patterns and the processes that shape them are rare. Here, we document the individual-, population-, and landscape-level phenological patterns of two dominant families of understory woody plants important to avian frugivores, the Melastomataceae and Rubiaceae, along an elevational transect in Costa Rica. The 226 individual plants belonging to 35 species in this study, varied in the number of reproductive bouts/year, and the timing, duration, and synchrony of reproductive stages. This variation was not related to factors related to their interactions with mutualists and antagonists, nor did it appear to be constrained by phylogeny. Diverse phenological patterns among species led to relatively aseasonal patterns at the community and landscape level. Overall, evidence for biotic processes shaping temporal patterns of fruiting phenology was weak or absent. These findings reveal a number of unexplained patterns, and suggest that factors shaping phenology in relatively aseasonal forests operate in idiosyncratic ways at the species level.

Context-dependent recruitment of Guettarda viburnoides in a Neotropical forest-savanna mosaic.

PREMISE OF THE STUDY: Seed dispersal is a key process for plant regeneration in tropical areas. Differences in disperser behavior result in a nonrandom distribution of seeds among habitats. Patterns of seed dispersal may cascade through the entire recruitment phase or uncoupling between developmental stages may occur such that patterns of seed dispersal are discordant with patterns of recruitment. Here, we analyzed how habitat and interannual variability affect the recruitment dynamics of a bird-dispersed tree, Guettarda viburnoides (RUB.). METHODS: Over 3 years, we determined the habitats where seeds of G. viburnoides are dispersed, and we experimentally quantified postdispersal seed predation, seedling emergence, and survival in four habitats of a forest-savanna mosaic in Bolivia. KEY RESULTS: Habitat affected seed dispersal, seed predation, and seedling emergence. The strength of postdispersal processes, however, varied between years and no consistent within-habitat pattern emerged. Uncoupling among different life-stages was observed across habitats, and spatial concordance was found between seed rain and sapling recruitment patterns. CONCLUSIONS: Habitat can affect seed dispersal, postdispersal processes and the recruitment dynamics of a Neotropical tree in a heterogeneous landscape. Additionally, our results show interannual variability in the strength of postdispersal processes, which leads to shifts in habitat suitability between years. Therefore, to better understand the role of site suitability for recruitment, we need to explicitly consider not only variation in habitat-specificity for dispersal and postdispersal processes, but also how this variation can shift under different environmental conditions-that is, the context dependence of suitability.