ABSTRACT
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.
Subject(s)
Ants , Myrmecophytes , Plant Leaves , Rubiaceae , Symbiosis , Animals , Ants/physiology , Herbivory , Myrmecophytes/growth & development , Myrmecophytes/physiology , Plant Leaves/growth & development , Rubiaceae/growth & development , Rubiaceae/physiology , Trees/growth & developmentABSTRACT
Climate models predict that everwet western Amazonian forests will face warmer and wetter atmospheric conditions, and increased cloud cover. It remains unclear how these changes will impact plant reproductive performance, such as flowering, which plays a central role in sustaining food webs and forest regeneration. Warmer and wetter nights may cause reduced flower production, via increased dark respiration rates or alteration in the reliability of flowering cue-based processes. Additionally, more persistent cloud cover should reduce the amounts of solar irradiance, which could limit flower production. We tested whether interannual variation in flower production has changed in response to fluctuations in irradiance, rainfall, temperature, and relative humidity over 18 yrs in an everwet forest in Ecuador. Analyses of 184 plant species showed that flower production declined as nighttime temperature and relative humidity increased, suggesting that warmer nights and greater atmospheric water saturation negatively impacted reproduction. Species varied in their flowering responses to climatic variables but this variation was not explained by life form or phylogeny. Our results shed light on how plant communities will respond to climatic changes in this everwet region, in which the impacts of these changes have been poorly studied compared with more seasonal Neotropical areas.
Subject(s)
Trees , Tropical Climate , Trees/physiology , Reproducibility of Results , Forests , Plants , Climate Change , Flowers/physiologyABSTRACT
Tables and charts have long been seen as effective ways to convey data. Much attention has been focused on improving charts, following ideas of human perception and brain function. Tables can also be viewed as two-dimensional representations of data; yet, it is only fairly recently that we have begun to apply principles of design that aid the communication of information between the author and reader. In this study, we collated guidelines for the design of data and statistical tables. These guidelines fall under three principles: aiding comparisons, reducing visual clutter, and increasing readability. We surveyed tables published in recent issues of 43 journals in the fields of ecology and evolutionary biology for their adherence to these three principles, as well as author guidelines on journal publisher websites. We found that most of the over 1000 tables we sampled had no heavy grid lines and little visual clutter. They were also easy to read, with clear headers and horizontal orientation. However, most tables did not aid the vertical comparison of numeric data. We suggest that authors could improve their tables by the right-flush alignment of numeric columns typeset with a tabular font, clearly identify statistical significance, and use clear titles and captions. Journal publishers could easily implement these formatting guidelines when typesetting manuscripts.
ABSTRACT
Flowering and fruiting phenology have been infrequently studied in the ever-wet hyperdiverse lowland forests of northwestern equatorial Amazonía. These Neotropical forests are typically called aseasonal with reference to climate because they are ever-wet, and it is often assumed they are also aseasonal with respect to phenology. The physiological limits to plant reproduction imposed by water and light availability are difficult to disentangle in seasonal forests because these variables are often temporally correlated, and both are rarely studied together, challenging our understanding of their relative importance as drivers of reproduction. Here we report on the first long-term study (18 years) of flowering and fruiting phenology in a diverse equatorial forest, Yasuní in eastern Ecuador, and the first to include a full suite of on-site monthly climate data. Using twice monthly censuses of 200 traps and >1000 species, we determined whether reproduction at Yasuní is seasonal at the community and species levels and analyzed the relationships between environmental variables and phenology. We also tested the hypothesis that seasonality in phenology, if present, is driven primarily by irradiance. Both the community- and species-level measures demonstrated strong reproductive seasonality at Yasuní. Flowering peaked in September-November and fruiting peaked in March-April, with a strong annual signal for both phenophases. Irradiance and rainfall were also highly seasonal, even though no month on average experienced drought (a month with <100 mm rainfall). Flowering was positively correlated with current or near-current irradiance, supporting our hypothesis that the extra energy available during the period of peak irradiance drives the seasonality of flowering at Yasuní. As Yasuní is representative of lowland ever-wet equatorial forests of northwestern Amazonía, we expect that reproductive phenology will be strongly seasonal throughout this region.
La fenología de floración y fructificación ha sido poco estudiada en los bosques bajos, lluviosos e hiperdiversos de la Amazonía noroccidental. Estos bosques neotropicales son típicamente llamados no estacionales debido a su clima siempre lluvioso y se asume que son no estacionales con respecto a la fenología. Los límites fisiológicos a la reproducción de las plantas impuestos por la disponibilidad de agua y luz en estos bosques son difíciles de desentrañar debido a que estas variables están a menudo correlacionadas temporalmente y las dos se estudian usualmente por separado, lo que desafía nuestra comprensión de su importancia relativa como desencadenantes de la reproducción. Este es el primer estudio de largo plazo (18 años) de la fenología de floración y fructificación en un bosque hiperdiverso de la Amazonía noroccidental ecuatorial, Yasuní, ubicado al este de Ecuador, y el primero en incluir un completo set de datos climáticos mensuales. Usando censos quincenales de 200 trampas y > 1000 especies, examinamos si la reproducción en Yasuní es estacional a nivel de comunidad y de especies y analizamos las relaciones de las variables ambientales con la fenología. También nos interesaba probar si la estacionalidad en la fenología, en caso de que esté presente está causada por la irradiancia. Tanto a nivel de comunidad como de especies, los datos demuestran una fuerte estacionalidad reproductiva en Yasuní. La floración alcanzó un máximo en septiembre-noviembre y la fructificación alcanzó un máximo en marzo-abril, con una fuerte y consistente señal anual en las dos fenofases. A su vez, la irradiancia y la lluvia fueron también marcadamente estacionales, aunque ningún mes en promedio experimentó sequía (i.e. <100 mm de lluvia). La floración fue positivamente correlacionada con la irradiación, apoyando nuestra hipótesis de que la energía extra disponible durante los periodos de mayor irradiación causa la estacionalidad de la floración en Yasuní. Debido a que Yasuní representa a los bosques ecuatoriales lluviosos de tierras bajas de la Amazonía noroccidental, esperamos que la fenología reproductiva sea fuertemente estacional a lo largo de esta región.
Subject(s)
Forests , Trees , Trees/physiology , Ecuador , Reproduction/physiology , Seasons , Tropical ClimateABSTRACT
Phenology has long been hypothesized as an avenue for niche partitioning or interspecific facilitation, both promoting species coexistence. Tropical plant communities exhibit striking diversity in reproductive phenology, but many are also noted for large synchronous reproductive events. Here we study whether the phenology of seed fall in such communities is nonrandom, the temporal scales of phenological patterns, and ecological factors that drive reproductive phenology. We applied multivariate wavelet analysis to test for phenological synchrony versus compensatory dynamics (i.e., antisynchronous patterns where one species' decline is compensated by the rise of another) among species and across temporal scales. We used data from long-term seed rain monitoring of hyperdiverse plant communities in the western Amazon. We found significant synchronous whole-community phenology at multiple timescales, consistent with shared environmental responses or positive interactions among species. We also observed both compensatory and synchronous phenology within groups of species (confamilials) likely to share traits and seed dispersal mechanisms. Wind-dispersed species exhibited significant synchrony at ~6-month scales, suggesting these species might share phenological niches to match the seasonality of wind. Our results suggest that community phenology is shaped by shared environmental responses but that the diversity of tropical plant phenology may partly result from temporal niche partitioning. The scale-specificity and time-localized nature of community phenology patterns highlights the importance of multiple and shifting drivers of phenology.
Subject(s)
Plants , Seeds , Seasons , Reproduction , Time Factors , Climate ChangeABSTRACT
PREMISE: Understanding the drivers of the growth in long-lived woody trees is the key to predicting their responses to and maintaining their populations under global change. However, the role of tree sex and differential investment to reproduction are often not considered in models of individual tree growth, despite many gymnosperm and angiosperm species having separate male and female sexes. Thus, better models of tree growth should include tree sex and life stage along with the abiotic and biotic neighborhoods. METHODS: We used a sex-specific molecular marker to determine the sex of 2188 individual trees >1 cm DBH of the dioecious tree species Diospyros morrisiana in a 50-ha subtropical forest plot in China. We used long-term census data from about 300,000 trees, together with 625 soil samples and 2352 hemispherical photographs to characterize the spatially explicit biotic and abiotic neighborhoods. RESULTS: We found a male-biased effective sex ratio and a female-biased overall population sex ratio of D. morrisiana. No sex spatial segregation was detected for the overall population, mature, or immature trees. Immature trees grew faster than mature trees and females grew slower than males. Further, conspecific neighbors significantly decreased tree growth, while the abiotic neighborhood showed no significant effect. CONCLUSIONS: Our findings suggest that variation in resource allocation patterns within and across individual trees of different sexes and life-history stages should be more widely accounted for in models of tree growth. In addition, our study highlights the importance of sex-specific molecular markers for studying populations of long-lived dioecious tree species.
Subject(s)
Diospyros , Trees , Trees/physiology , Forests , Wood , Sex RatioABSTRACT
PREMISE: The Ocotea complex contains the greatest diversity of Lauraceae in the Neotropics. However, the traditional taxonomy of the group has relied on only three main floral characters, and previous molecular analyses have used only a few markers and provided limited support for relationships among the major clades. This lack of useful data has hindered the development of a comprehensive classification, as well as studies of character evolution. METHODS: We used RAD-seq data to infer the phylogenetic relationships of 149 species in the Ocotea complex, generating a reference-based assembly using the Persea americana genome. The results provide the basis for a phylogenetic classification that reflects our current molecular knowledge and for analyses of the evolution of breeding system, stamen number, and number of anther locules. RESULTS: We recovered a well-supported tree that demonstrates the paraphyly of Licaria, Aniba, and Ocotea and clarifies the relationships of Umbellularia, Phyllostemonodaphne, and the Old World species. To begin the development of a new classification and to facilitate precise communication, we also provide phylogenetic definitions for seven major clades. Our ancestral reconstructions show multiple origins for the three floral characters that have routinely been used in Lauraceae systematics, suggesting that these be used with caution in the future. CONCLUSIONS: This study advances our understanding of phylogenetic relationships and character evolution in a taxonomically difficult group using RAD-seq data. Our new phylogenetic names will facilitate unambiguous communication as studies of the Ocotea complex progress.
Subject(s)
Ocotea , Evolution, Molecular , Phylogeny , Plant Breeding , Sequence Analysis, DNAABSTRACT
Plant-soil feedback (PSF) theory provides a powerful framework for understanding plant dynamics by integrating growth assays into predictions of whether soil communities stabilise plant-plant interactions. However, we lack a comprehensive view of the likelihood of feedback-driven coexistence, partly because of a failure to analyse pairwise PSF, the metric directly linked to plant species coexistence. Here, we determine the relative importance of plant evolutionary history, traits, and environmental factors for coexistence through PSF using a meta-analysis of 1038 pairwise PSF measures. Consistent with eco-evolutionary predictions, feedback is more likely to mediate coexistence for pairs of plant species (1) associating with similar guilds of mycorrhizal fungi, (2) of increasing phylogenetic distance, and (3) interacting with native microbes. We also found evidence for a primary role of pathogens in feedback-mediated coexistence. By combining results over several independent studies, our results confirm that PSF may play a key role in plant species coexistence, species invasion, and the phylogenetic diversification of plant communities.
Subject(s)
Mycorrhizae , Phylogeny , Soil Microbiology , Plants , SoilABSTRACT
Environment and human land use both shape forest composition. Abiotic conditions sift tree species from a regional pool via functional traits that influence species' suitability to the local environment. In addition, human land use can modify species distributions and change functional diversity of forests. However, it is unclear how environment and land use simultaneously shape functional diversity of tree communities. Land-use legacies are especially prominent in temperate forest landscapes that have been extensively modified by humans in the last few centuries. Across a 900-ha temperate deciduous forest in the northeastern United States, comprising a mosaic of different-aged stands due to past human land use, we used four key functional traits-maximum height, rooting depth, wood density, and seed mass-to examine how multiple environmental and land-use variables influenced species distributions and functional diversity. We sampled ~40,000 trees >8 cm DBH within 485 plots totaling 137 ha. Species within plots were more functionally similar than expected by chance when we estimated functional diversity using all traits together (multi-trait), and to a lesser degree, with each trait separately. Multi-trait functional diversity was most strongly correlated with distance from the perennial stream, elevation, slope, and forest age. Environmental and land-use predictors varied in their correlation with functional diversities of the four individual traits. Landscape-wide change in abundances of individual species also correlated with both environment and land-use variables, but magnitudes of trait-environment interactions were generally stronger than trait interactions with land use. These findings can be applied for restoration and assisted regeneration of human-modified temperate forests by using traits to predict which tree species would establish well in relation to land-use history, topography, and soil conditions.
Subject(s)
Biodiversity , Environment , Forests , Trees , Agriculture , Forestry , Pennsylvania , Trees/growth & developmentABSTRACT
Recent evidence suggests that plant performance can be influenced by the phylogenetic diversity of neighboring plants. However, no study to date has examined the effect of such phylogenetic density dependence on the transition from seed to seedling. Using 6 years of data on seedling recruitment and seed rain of 13 species from 130 stations (one 0.5 m2 seed trap and three adjacent 1 m2 seedling plots) in a subtropical evergreen forest, we asked: (1) Does negative density dependence act across seed to seedling stages? (2) Is there evidence for phylogenetic density dependence during the seed to seedling transition? (3) Does the strength of density dependence vary among years? Generalized linear mixed-effects models were used to model seed to seedling transition as a function of conspecific seed and seedling densities, heterospecific seed and seedling densities, and mean phylogenetic distance of heterospecific seeds and seedling. Conspecific seed density had a significant negative effect on seedling transition rates for 12 of 13 focal species. In contrast, conspecific seedling density had a positive effect for 7 species, suggesting species-specific habitat preferences. Few species were significantly affected by the density or phylogenetic relatedness of heterospecific seeds and seedlings. Only conspecific seed density effects varied among years for most focal species. Overall, our results reveal that conspecific seed and seedling densities play a more important role than the density or relatedness of heterospecific seeds and seedlings during the seed to seedling stage, suggesting that species-specific seed predators, along with habitat preferences, may contribute to diversity maintenance in this forest.
Subject(s)
Phylogeny , Seedlings , Ecosystem , Forests , Population Density , Seeds , TreesABSTRACT
Nitrogen (N) and phosphorus (P) deposition are increasing worldwide largely due to increased fertilizer use and fossil fuel combustion. Most work with N and P deposition in natural ecosystems has focused on temperate, highly industrialized, regions. Tropical regions are becoming more developed, releasing large amounts of these nutrients into the atmosphere. Nutrient enrichment in nutrient-poor systems such as tropical montane forest can represent a relatively large shift in nutrient availability, especially for sensitive microorganisms such as arbuscular mycorrhizal fungi (AMF). These symbiotic fungi are particularly critical, given their key role in ecosystem processes affecting plant community structure and function.To better understand the consequences of nutrient deposition in plant communities, a long-term nutrient addition experiment was set up in a tropical montane forest in the Andes of southern Ecuador. In this study, we investigated the impacts of 7 years of elevated N and P on AMF root colonization potential (AMF-RCP) through a greenhouse bait plant method in which we quantified root colonization. We also examined the relationship between AMF-RCP and rarefied tree diversity.After 7 years of nutrient addition, AMF-RCP was negatively correlated with soil P, positively correlated with soil N, and positively correlated with rarefied tree diversity. Our results show that AMF in this tropical montane forest are directly affected by soil N and P concentrations, but may also be indirectly impacted by shifts in rarefied tree diversity. Our research also highlights the need to fully understand the benefits and drawbacks of using different sampling methods (e.g., AMF-RCP versus direct root sampling) to robustly examine AMF-plant interactions in the future.
Subject(s)
Forests , Mycorrhizae/physiology , Nitrogen/physiology , Phosphorus/physiology , Ecuador , Fungi , Plant Roots/microbiology , Soil/chemistry , Soil MicrobiologyABSTRACT
The "liana dominance hypothesis" posits that lianas are increasing in abundance in tropical forests, thereby potentially reducing tree biomass due to competitive interactions between trees and lianas. This scenario has implications not only for forest ecosystem function and species composition, but also climate change given the mass of carbon stored in tropical trees. In 2003 and 2013, all Myristicaceae trees in the 50-ha Yasuní Forest Dynamics Plot, Ecuador, were surveyed for liana presence and load in their crowns. We tested the hypothesis that the proportion of trees with lianas increased between 2003 and 2013 in line with the liana dominance hypothesis. Contrary to expectations, the total proportion of trees with lianas decreased from 35% to 32%, and when only trees ≥10 cm diameter at breast height were considered liana incidence increased 44-48%. Liana load was dynamic with a large proportion of trees losing or gaining lianas over the 10-yr period; large trees with intermediate liana loads increased in proportion at the expense of those with low and high loads. Lianas also impacted performance: trees with 26-75% crown cover by lianas in 2003 had reduced growth rates of 80% compared to of liana-free trees, and trees with >75% crown cover had 33% the growth rate and a log odds of mortality eight times that of liana-free trees. We suggest that the lack of strong support found for the liana dominance hypothesis is likely due to the aseasonal climate of Yasuní, which limits the competitive advantage lianas maintain over trees during dry seasons due to their efficient capture and use of water. We propose further research of long-term liana dynamics from aseasonal forests is required to determine the generality of the increasing liana dominance hypothesis in Neotropical forests.
Subject(s)
Myristicaceae/physiology , Rainforest , Ecosystem , Ecuador , Rain , Trees , Tropical ClimateABSTRACT
How ecological context shapes mutualistic relationships remains poorly understood. We combined long-term tree census data with ant censuses in a permanent 25-ha Amazonian forest dynamics plot to evaluate the effect of the mutualistic ant Myrmelachista schumanni (Formicinae) on the growth and survival of the common Amazonian tree Duroia hirsuta (Rubiaceae), considering its interactions with tree growth, population structure, and habitat. We found that the mutualist ant more than doubled tree relative growth rates and increased odds of survival. However, host tree size and density of conspecific neighbors modified the effect of the ant. Smaller trees hosting the mutualist ant consistently grew faster when surrounded by higher densities of conspecifics, suggesting that the benefit to the tree outweighs any negative effects of high conspecific densities. Moreover, our findings suggest that the benefit afforded by the ant diminishes with plant age and also depends on the density of conspecific neighbors. We provide the first long-term large-scale evidence of how mutualism affects the population biology of an Amazonian tree species.
Subject(s)
Ants , Symbiosis , Animals , Ecosystem , Rubiaceae , TreesABSTRACT
Patterns of diversity and community composition in forests are controlled by a combination of environmental factors, historical events, and stochastic or neutral mechanisms. Each of these processes has been linked to forest community assembly, but their combined contributions to alpha and beta-diversity in forests has not been well explored. Here we use variance partitioning to analyze approximately 40,000 individual trees of 49 species, collected within 137 ha of sampling area spread across a 900-ha temperate deciduous forest reserve in Pennsylvania to ask (1) To what extent is site-to-site variation in species richness and community composition of a temperate forest explained by measured environmental gradients and by spatial descriptors (used here to estimate dispersal-assembly or unmeasured, spatially structured processes)? (2) How does the incorporation of land-use history information increase the importance attributed to deterministic community assembly? and (3) How do the distributions and abundances of individual species within the community correlate with these factors? Environmental variables (i.e., topography, soils, and distance to stream), spatial descriptors (i.e., spatial eigenvectors derived from Cartesian coordinates), and land-use history variables (i.e., land-use type and intensity, forest age, and distance to road), explained about half of the variation in both species richness and community composition. Spatial descriptors explained the most variation, followed by measured environmental variables and then by land- use history. Individual species revealed variable responses to each of these sets of predictor variables. Several species were associated with stream habitats, and others were strictly delimited across opposing north- and south-facing slopes. Several species were also associated with areas that experienced recent (i.e., <100 years) human land-use impacts. These results indicate that deterministic factors, including environmental and land-use history variables, are important drivers of community response. The large amount of "unexplained" variation seen here (about 50%) is commonly observed in other such studies attempting to explain distribution and abundance patterns of plant communities. Determining whether such large fractions of unaccounted for variation are caused by a lack of sufficient data, or are an indication of stochastic features of forest communities globally, will remain an important challenge for ecologists in the future.
Subject(s)
Biodiversity , Forests , Trees/physiology , Agriculture , Environment , Forestry , Mining , PennsylvaniaABSTRACT
The Janzen-Connell hypothesis proposes that specialist natural enemies, such as herbivores and pathogens, maintain diversity in plant communities by reducing survival rates of conspecific seeds and seedlings located close to reproductive adults or in areas of high conspecific density. Variation in the strength of distance- and density-dependent effects is hypothesized to explain variation in plant species richness along climatic gradients, with effects predicted to be stronger in the tropics than the temperate zone and in wetter habitats compared to drier habitats.We conducted a comprehensive literature search to identify peer-reviewed experimental studies published in the 40+ years since the hypothesis was first proposed. Using data from these studies, we conducted a meta-analysis to assess the current weight of evidence for the distance and density predictions of the Janzen-Connell hypothesis.Overall, we found significant support for both the distance- and density-dependent predictions. For all studies combined, survival rates were significantly reduced near conspecifics compared to far from conspecifics, and in areas with high densities of conspecifics compared to areas with low conspecific densities. There was no indication that these results were due to publication bias.The strength of distance and density effects varied widely among studies. Contrary to expectations, this variation was unrelated to latitude, and there was no significant effect of study region. However, we did find a trend for stronger distance and density dependence in wetter sites compared to sites with lower annual precipitation. In addition, effects were significantly stronger at the seedling stage compared to the seed stage.Synthesis. Our study provides support for the idea that distance- and density-dependent mortality occurs in plant communities world-wide. Available evidence suggests that natural enemies are frequently the cause of such patterns, consistent with the Janzen-Connell hypothesis, but additional studies are needed to rule out other mechanisms (e.g. intraspecific competition). With the widespread existence of density and distance dependence clearly established, future research should focus on assessing the degree to which these effects permit species coexistence and contribute to the maintenance of diversity in plant communities.
ABSTRACT
To generate realistic projections of species' responses to climate change, we need to understand the factors that limit their ability to respond. Although climatic niche conservatism, the maintenance of a species's climatic niche over time, is a critical assumption in niche-based species distribution models, little is known about how universal it is and how it operates. In particular, few studies have tested the role of climatic niche conservatism via phenological changes in explaining the reported wide variance in the extent of range shifts among species. Using historical records of the phenology and spatial distribution of British plants under a warming climate, we revealed that: (i) perennial species, as well as those with weaker or lagged phenological responses to temperature, experienced a greater increase in temperature during flowering (i.e. failed to maintain climatic niche via phenological changes); (ii) species that failed to maintain climatic niche via phenological changes showed greater northward range shifts; and (iii) there was a complementary relationship between the levels of climatic niche conservatism via phenological changes and range shifts. These results indicate that even species with high climatic niche conservatism might not show range shifts as instead they track warming temperatures during flowering by advancing their phenology.
Subject(s)
Climate Change , Plant Dispersal , Plant Development , Seasons , Species Specificity , United KingdomABSTRACT
Aim. Throughout evolutionary history, plants and animals have evolved alongside one another. This is especially apparent when considering mutualistic relationships such as between plants with extra-floral nectaries (EFNs, glands on leaves or stems that secrete nectar) and the ants that visit them. Ants are attracted by the nectar and then protect the plant against destructive herbivores. The distribution of these plants is of particular interest, because it can provide insights into the evolutionary history of this unique trait and the plants that possess it. In this study, we investigated factors driving the distribution of woody plants with EFNs in the cerrado vegetation of Brazil. Location. Brazil Methods. We used a database detailing the incidence of 849 plant species at 367 cerrado sites throughout Brazil. We determined which species possessed EFNs and mapped their distributions. We tested for correlations between the proportion of EFN species at each site and (i) three environmental variables (mean annual temperature, mean annual precipitation, and the precipitation in the driest quarter of the year), (ii) a broad soil classification, and (iii) the total species diversity of each site. Results. We found a wide range in the proportion of EFN species at any one site (0-57%). However, whilst low diversity sites had wide variation in the number of EFN species, high diversity sites all had few EFN species. The proportion of EFN species was positively correlated with absolute latitude and negatively correlated with longitude. When accounting for total species diversity, the proportion of EFN species per site was negatively correlated with precipitation in the driest quarter of the year and positively correlated with temperature range. Main Conclusions. These results suggest either that herbivore pressure may be lower in drier sites, or that ants are not as dominant in these locations, or that plant lineages at these sites were unable to evolve EFNs.
ABSTRACT
BACKGROUND: Many tropical forest tree species delay greening their leaves until full expansion. This strategy is thought to provide newly flushing leaves with protection against damage by herbivores by keeping young leaves devoid of nutritive value. Because young leaves suffer the greatest predation from invertebrate herbivores, delayed greening could prevent costly tissue loss. Many species that delay greening also produce anthocyanin pigments in their new leaves, giving them a reddish tint. These anthocyanins may be fungicidal, protect leaves against UV damage or make leaves cryptic to herbivores blind to the red part of the spectrum. METHODS: A comprehensive survey was undertaken of seedlings, saplings and mature trees in two diverse tropical forests: a rain forest in western Amazonia (Yasuní National Park, Ecuador) and a deciduous forest in Central America (Barro Colorado Island, Panamá). A test was made of whether individuals and species with delayed greening or red-coloured young leaves showed lower mortality or higher relative growth rates than species that did not. KEY RESULTS: At both Yasuní and Barro Colorado Island, species with delayed greening or red young leaves comprised significant proportions of the seedling and tree communities. At both sites, significantly lower mortality was found in seedlings and trees with delayed greening and red-coloured young leaves. While there was little effect of leaf colour on the production of new leaves of seedlings, diameter relative growth rates of small trees were lower in species with delayed greening and red-coloured young leaves than in species with regular green leaves, and this effect remained when the trade-off between mortality and growth was accounted for. CONCLUSIONS: Herbivores exert strong selection pressure on seedlings for the expression of defence traits. A delayed greening or red-coloured young leaf strategy in seedlings appears to be associated with higher survival for a given growth rate, and may thus influence the species composition of later life stages.
Subject(s)
Anthocyanins/physiology , Ecosystem , Herbivory , Seedlings/physiology , Trees/physiology , Animals , Ecuador , Panama , Tropical ClimateABSTRACT
A fundamental challenge facing physiological ecologists is to understand how variation in life history at the whole-organism level might be linked to cellular function. Thus, because tropical birds have higher annual survival and lower rates of metabolism, we hypothesized that cells from tropical species would have greater cellular resistance to chemical injury than cells from temperate species. We cultured dermal fibroblasts from 26 tropical and 26 temperate species of birds and examined cellular resistance to cadmium, H(2)O(2), paraquat, thapsigargin, tunicamycium, methane methylsulfonate (MMS) and UV light. Using ANCOVA, we found that the values for the dose that killed 50% of cells (LD(50)) from tropical birds were significantly higher for H(2)O(2) and MMS. When we tested for significance using a generalized least squares approach accounting for phylogenetic relationships among species to model LD(50), we found that cells from tropical birds had greater tolerance for Cd, H(2)O(2), paraquat, tunicamycin and MMS than cells from temperate birds. In contrast, tropical birds showed either lower or no difference in tolerance to thapsigargin and UV light in comparison with temperate birds. These findings are consistent with the idea that natural selection has uniquely fashioned cells of long-lived tropical bird species to be more resistant to forms of oxidative and non-oxidative stress than cells from shorter-lived temperate species.