Small genomes and large seeds: chromosome numbers, genome size and seed mass in diploid Aesculus species (Sapindaceae).

Background and Aims: Aesculus L. (horse chestnut, buckeye) is a genus of 12-19 extant woody species native to the temperate Northern Hemisphere. This genus is known for unusually large seeds among angiosperms. While chromosome counts are available for many Aesculus species, only one has had its genome size measured. The aim of this study is to provide more genome size data and analyse the relationship between genome size and seed mass in this genus. Methods: Chromosome numbers in root tip cuttings were confirmed for four species and reported for the first time for three additional species. Flow cytometric measurements of 2C nuclear DNA values were conducted on eight species, and mean seed mass values were estimated for the same taxa. Key Results: The same chromosome number, 2 n = 40, was determined in all investigated taxa. Original measurements of 2C values for seven Aesculus species (eight taxa), added to just one reliable datum for A. hippocastanum , confirmed the notion that the genome size in this genus with relatively large seeds is surprisingly low, ranging from 0·955 pg 2C -1 in A. parviflora to 1·275 pg 2C -1 in A. glabra var. glabra. Conclusions: The chromosome number of 2 n = 40 seems to be conclusively the universal 2 n number for non-hybrid species in this genus. Aesculus genome sizes are relatively small, not only within its own family, Sapindaceae, but also within woody angiosperms. The genome sizes seem to be distinct and non-overlapping among the four major Aesculus clades. These results provide an extra support for the most recent reconstruction of Aesculus phylogeny. The correlation between the 2C values and seed masses in examined Aesculus species is slightly negative and not significant. However, when the four major clades are treated separately, there is consistent positive association between larger genome size and larger seed mass within individual lineages.

Mediterranean, invasive, woody species grow larger than their less-invasive counterparts under potential global environmental change.

PREMISE OF THE STUDY: Revealing biological differences between invasive and noninvasive species is essential for predicting species' distribution changes with global environmental change. While most research has focused on differences between invasive and noninvasive species under favorable conditions using herbaceous species, invasive woody angiosperms are also of great ecological concern. Our study focused on how growth and allocation may change for invasive and noninvasive, mediterranean, woody angiosperms under future conditions caused by global change, specifically increased nitrogen deposition and drought. METHODS: We tested how seedling functional traits differed between invasive and noninvasive woody angiosperms under different experimental conditions in a greenhouse setting. We compared growth rates and allocation patterns using two levels of soil nitrogen and three levels of watering. We also examined trait log response ratios to increases in nitrogen and increases in water. Our study sampled angiosperm trees and shrubs, incorporating congeneric/confamilial relationships through 13 phylogenetically controlled contrasts. KEY RESULTS: Three functional traits were highly and positively associated with plant invasiveness for most conditions studied: seedling plant mass, leaf area, and height. Invasive species also had significantly higher root mass ratios at low water regardless of nitrogen input. Invasive and noninvasive species had similar log response ratios to increases in nitrogen and watering for studied traits. CONCLUSIONS: Mediterranean, woody, invasive species' larger mass, leaf area, and early height advantage under elevated nitrogen input and increased root production in drought conditions may lead to increased invasion of these species with expected global climate change.

No universal scale-dependent impacts of invasive species on native plant species richness.

A growing number of studies seeking generalizations about the impact of plant invasions compare heavily invaded sites to uninvaded sites. But does this approach warrant any generalizations? Using two large datasets from forests, grasslands and desert ecosystems across the conterminous United States, we show that (i) a continuum of invasion impacts exists in many biomes and (ii) many possible species-area relationships may emerge reflecting a wide range of patterns of co-occurrence of native and alien plant species. Our results contradict a smaller recent study by Powell et al. 2013 (Science 339, 316-318. (doi:10.1126/science.1226817)), who compared heavily invaded and uninvaded sites in three biomes and concluded that plant communities invaded by non-native plant species generally have lower local richness (intercepts of log species richness-log area regression lines) but steeper species accumulation with increasing area (slopes of the regression lines) than do uninvaded communities. We conclude that the impacts of plant invasions on plant species richness are not universal.

Directed seed dispersal towards areas with low conspecific tree density by a scatter-hoarding rodent.

Scatter-hoarding animals spread out cached seeds to reduce density-dependent theft of their food reserves. This behaviour could lead to directed dispersal into areas with lower densities of conspecific trees, where seed and seedling survival are higher, and could profoundly affect the spatial structure of plant communities. We tested this hypothesis with Central American agoutis and Astrocaryum standleyanum palm seeds on Barro Colorado Island, Panama. We radio-tracked seeds as they were cached and re-cached by agoutis, calculated the density of adult Astrocaryum trees surrounding each cache, and tested whether the observed number of trees around seed caches declined more than expected under random dispersal. Seedling establishment success was negatively dependent on seed density, and agoutis carried seeds towards locations with lower conspecific tree densities, thus facilitating the escape of seeds from natural enemies. This behaviour may be a widespread mechanism leading to highly effective seed dispersal by scatter-hoarding animals.

TEASIng apart alien species risk assessments: a framework for best practices.

Some alien species cause substantial impacts, yet most are innocuous. Given limited resources, forecasting risks from alien species will help prioritise management. Given that risk assessment (RA) approaches vary widely, a synthesis is timely to highlight best practices. We reviewed quantitative and scoring RAs, integrating > 300 publications into arguably the most rigorous quantitative RA framework currently existing, and mapping each study onto our framework, which combines Transport, Establishment, Abundance, Spread and Impact (TEASI). Quantitative models generally measured single risk components (78% of studies), often focusing on Establishment alone (79%). Although dominant in academia, quantitative RAs are underused in policy, and should be made more accessible. Accommodating heterogeneous limited data, combining across risk components, and developing generalised RAs across species, space and time without requiring new models for each species may increase attractiveness for policy applications. Comparatively, scoring approaches covered more risk components (50% examined > 3 components), with Impact being the most common component (87%), and have been widely applied in policy (> 57%), but primarily employed expert opinion. Our framework provides guidance for questions asked, combining scores and other improvements. Our risk framework need not be completely parameterised to be informative, but instead identifies opportunities for improvement in alien species RA.

Avian use of introduced plants: ornithologist records illuminate interspecific associations and research needs.

Introduced species have the potential to impact processes central to the organization of ecological communities. Although hundreds of nonnative plant species have naturalized in the United States, only a small percentage of these have been studied in their new biotic communities. Their interactions with resident (native and introduced) bird species remain largely unexplored. As a group, citizen scientists such as ornithologists possess a wide range of experiences. They may offer insights into the prevalence and form of bird interactions with nonnative plants on a broad geographic scale. We surveyed 173 ornithologists from four U.S. states, asking them to report observations of bird interactions with nonnative plants. The primary goal of the survey was to obtain information useful in guiding future empirical research. In all, 1143 unique bird-plant interactions were reported, involving 99 plant taxa and 168 bird species. Forty-seven percent of reported interactions concerned potential dispersal (feeding on seeds or fruits). Remaining "habitat interactions" involved bird use of plants for nesting, perching, woodpecking, gleaning, and other activities. We utilized detrended correspondence analysis to ordinate birds with respect to the plants they reportedly utilize. Results illuminate the new guilds formed by these interactions. We assessed the existing level of knowledge about invasiveness of those plants reported most often in feeding interactions, identifying information gaps for biological invasions research priority. To exemplify the usefulness of citizen science data, we utilized survey results to guide field research on invasiveness in some of these plant species and observed both qualitatively and quantitatively strong agreement between survey reports and our empirical data. Questionnaire reports are therefore heuristically informative for the fields of both avian ecology and invasion biology.

A strong conditional mutualism limits and enhances seed dispersal and germination of a tropical palm.

Seed predation and seed dispersal can have strong effects on early life history stages of plants. These processes have often been studied as individual effects, but the degree to which their relative importance co-varies with seed predator abundance and how this influences seed germination rates is poorly understood. Therefore, we used a combination of observations and field experiments to determine the degree to which germination rates of the palm Astrocaryum mexicanum varied with abundance of a small mammal seed predator/disperser, Heteromys desmarestianus, in a lowland tropical forest. Patterns of abundance of the two species were strongly related; density of H. desmarestianus was low in sites with low density of A. mexicanum and vice versa. Rates of predation and dispersal of A. mexicanum seeds depended on abundance of H. desmarestianus; sites with high densities of H. desmarestianus had the highest rates of seed predation and lowest rates of seed germination, but a greater total number of seeds were dispersed and there was greater density of seedlings, saplings, and adults of A. mexicanum in these sites. When abundance of H. desmarestianus was experimentally reduced, rates of seed predation decreased, but so did dispersal of A. mexicanum seeds. Critically, rates of germination of dispersed seeds were 5 times greater than undispersed seeds. The results suggest that the relationship between A. mexicanum and H. desmarestianus is a conditional mutualism that results in a strong local effect on the abundance of each species. However, the magnitude and direction of these effects are determined by the relative strength of opposing, but related, mechanisms. A. mexicanum nuts provide H. desmarestianus with a critical food resource, and while seed predation on A. mexicanum nuts by H. desmarestianus is very intense, A. mexicanum ultimately benefits because of the relatively high germination rates of its seeds that are dispersed by H. desmarestianus.

The numerical and functional responses of a granivorous rodent and the fate of Neotropical tree seeds.

Despite their potential to provide mechanistic explanations of rates of seed dispersal and seed fate, the functional and numerical responses of seed predators have never been explicitly examined within this context. Therefore, we investigated the numerical response of a small-mammal seed predator, Heteromys desmarestianus, to disturbance-induced changes in food availability and evaluated the degree to which removal and fate of seeds of eight tree species in a lowland tropical forest in Belize were related to the functional response of H. desmarestianus to varying seed densities. Mark-recapture trapping was used to estimate abundance of H. desmarestianus in six 0.5-ha grids from July 2000 to September 2002. Fruit availability and seed fate were estimated in each grid, and two experiments nested within the grids were used to determine (1) the form of the functional response for nine levels of fruit density (2-32 fruits/m2), (2) the removal rate and handling times, and (3) the total proportion of fruits removed. The total proportion of fruits removed was determined primarily by the numerical response of H. desmarestianus to fruit availability, while removal rates and the proportion of seeds eaten or cached were related primarily to the form of the functional response. However, the numerical and functional responses interacted; H. desmarestianus showed strong spatial and temporal numerical responses to total fruit availability, and their density relative to fruit availability resulted in variation in the form of the functional response. Types I, II, and III functional responses were observed, as were density-independent responses, and these responses varied both among and within fruit species. The highest proportions of fruits were eaten when the Type III functional response was detected, which was when fruit availability was high relative to H. desmarestianus population density. Numerous idiosyncratic influences on seed fate have been documented, but our results indicate that shifts in the numerical and functional responses of seed predators to seasonal and interannual variation in seed availability potentially provide a general mechanistic explanation for patterns of removal and fate for vertebrate-dispersed seeds.

Predicting invasiveness of exotic woody species using a traits-based framework.

Identifying potentially invasive species and preventing their introduction and establishment are of critical importance in invasion ecology and land management. Although an extensive body of research has been dedicated to identifying traits that confer invasiveness, our current knowledge is still often inconclusive due to limitations in geographic extent and/or scope of traits analyzed. Here, using a comprehensive set of 45 traits, we performed a case study of invasive traits displayed by exotic woody plants in the United States (U.S.) by comparing 63 invasive and 794 non-invasive exotic woody plant species naturalized across the country. We found that invasive woody species often bear the following two key traits: vegetative reproduction and long-distance seed dispersal (via water, birds or mammals). Boosted classification tree models based on these traits accurately predicted species invasiveness (86% accuracy on average). Presented findings provide a generalized understanding of the relative importance of functional traits in identifying potentially invasive woody species in the U.S. The knowledge generated in this study can be used to improve current classification systems of non-native woody plants used by various U.S. governmental agencies and land managers.

Recent Anthropogenic Plant Extinctions Differ in Biodiversity Hotspots and Coldspots.

During the Anthropocene, humans are changing the Earth system in ways that will be detectable for millennia to come [1]. Biologically, these changes include habitat destruction, biotic homogenization, increased species invasions, and accelerated extinctions [2]. Contemporary extinction rates far surpass background rates [3], but they seem remarkably low in plants [4, 5]. However, biodiversity is not evenly distributed, and as a result, extinction rates may vary among regions. Some authors have contentiously argued that novel anthropic habitats and human-induced plant speciation can actually increase regional biodiversity [6, 7]. Here, we report on one of the most comprehensive datasets to date, including regional and global plant extinctions in both biodiversity hotspots (mostly from Mediterranean-type climate regions) and coldspots (mostly from Eurasian countries). Our data come from regions covering 15.3% of the Earth's surface and span over 300 years. With this dataset, we explore the trends, causes, and temporal dynamics of recent plant extinctions. We found more, and faster accrual of, absolute numbers of extinction events in biodiversity hotspots compared to coldspots. Extinction rates were also substantially higher than historical background rates, but recent declines are evident. We found higher levels of taxonomic uniqueness being lost in biodiversity coldspots compared to hotspots. Causes of plant extinctions also showed distinct temporal patterns, with agriculture, invasions, and urbanization being significant drivers in hotspots, while hydrological disturbance was an important driver in coldspots. Overall, plant extinctions over the last three centuries appear to be low, with a recent (post-1990) and steady extinction rate of 1.26 extinctions/year.

Contrasting ectomycorrhizal fungal communities on the roots of co-occurring oaks (Quercus spp.) in a California woodland.

Plant host species is considered an important factor influencing ectomycorrhizal (EM) communities. To gain insights into the role of host species in structuring EM communities, EM communities on sympatric oak (Quercus) species were compared in the Sierra Nevada foothills of California. Using molecular methods (polymerase chain reaction, cloning, restriction fragment length polymorphism and DNA sequencing), EM fungi on roots of deciduous Quercus douglasii and evergreen Quercus wislizeni trees were identified from 64 soil cores. The total EM species richness was 140, of which 40 taxa were detected on both oak hosts. Greater diversity and frequency of EM fungi with epigeous fruiting habit were found on Q. wislizeni, while taxa in the Ascomycota were more frequent and diverse on Q. douglasii. Using ordination, it was determined that both soil extractable phosphorus and oak host species explained a significant proportion of the variation in EM species distribution. These results indicate that plant host species can be an important factor influencing EM fungal community composition, even within congeneric trees.

Disentangling vegetation diversity from climate-energy and habitat heterogeneity for explaining animal geographic patterns.

Broad-scale animal diversity patterns have been traditionally explained by hypotheses focused on climate-energy and habitat heterogeneity, without considering the direct influence of vegetation structure and composition. However, integrating these factors when considering plant-animal correlates still poses a major challenge because plant communities are controlled by abiotic factors that may, at the same time, influence animal distributions. By testing whether the number and variation of plant community types in Europe explain country-level diversity in six animal groups, we propose a conceptual framework in which vegetation diversity represents a bridge between abiotic factors and animal diversity. We show that vegetation diversity explains variation in animal richness not accounted for by altitudinal range or potential evapotranspiration, being the best predictor for butterflies, beetles, and amphibians. Moreover, the dissimilarity of plant community types explains the highest proportion of variation in animal assemblages across the studied regions, an effect that outperforms the effect of climate and their shared contribution with pure spatial variation. Our results at the country level suggest that vegetation diversity, as estimated from broad-scale classifications of plant communities, may contribute to our understanding of animal richness and may be disentangled, at least to a degree, from climate-energy and abiotic habitat heterogeneity.

Evolution of genome size in pines (Pinus) and its life-history correlates: supertree analyses.

Genome size has been suggested to be a fundamental biological attribute in determining life-history traits in many groups of organisms. We examined the relationships between pine genome sizes and pine phylogeny, environmental factors (latitude, elevation, annual rainfall), and biological traits (latitudinal and elevational ranges, seed mass, minimum generation time, interval between large seed crops, seed dispersal mode, relative growth rate, measures of potential and actual invasiveness, and level of rarity). Genome sizes were determined for 60 pine taxa and then combined with published values to make a dataset encompassing 85 species, or 70% of species in the genus. Supertrees were constructed using 20 published source phylogenies. Ancestral genome size was estimated as 32 pg. Genome size has apparently remained stable or increased over evolutionary time in subgenus Strobus, while it has decreased in most subsections in subgenus Pinus. We analyzed relationships between genome size and life-history variables using cross-species correlations and phylogenetically independent contrasts derived from supertree constructions. The generally assumed positive relation between genome size and minimum generation time could not be confirmed in phylogenetically controlled analyses. We found that the strongest correlation was between genome size and seed mass. Because the growth quantities specific leaf area and leaf area ratio (and to a lesser extent relative growth rate) are strongly negatively related to seed mass, they were also negatively correlated with genome size. Northern latitudinal limit was negatively correlated with genome size. Invasiveness, particularly of wind-dispersed species, was negatively associated with both genome size and seed mass. Seed mass and its relationships with seed number, dispersal mode, and growth rate contribute greatly to the differences in life-history strategies of pines. Many life-history patterns are therefore indirectly, but consistently, associated with genome size.

Toward a causal explanation of plant invasiveness: seedling growth and life-history strategies of 29 pine (Pinus) species.

We studied 29 pine (Pinus) species to test the hypothesis that invasive species in disturbed habitats have distinct attributes. Seedling relative growth rate (RGR) and measures of invasiveness were positively associated across species as well as within phylogenetically independent contrasts. High RGR, small seed masses, and short generation times characterize pine species that are successful invaders in disturbed habitats. Discriminant analysis and logistic regression revealed that RGR was the most significant factor among these life-history traits separating invasive and noninvasive species. We also explored the causes of differences in RGR among invasive and noninvasive species. While net assimilation rate, leaf mass ratio, and specific leaf area (SLA) were all found to be contributing positively to RGR, SLA was found to be the main component responsible for differences in RGR between invasive and noninvasive pines. We investigated differences in SLA further by studying leaf anatomy, leaf density, and leaf thickness. We also evaluated relative leaf production rate as an important aspect of SLA. We proposed a hypothetical causal network of all relevant variables.

Corrections to the indices of community dissimilarity based on species diversity measures.

A number of indices of community dissimilarity based on the species diversity measures have been proposed during the last three decades (Margalef 1956, 1957; Kohn 1959; MacArthur 1965; Horn 1966; Glowacinski and Järvinen 1975; Järvinen and Väisänen 1976). Each of them is measuring the wrong quantity if the value of diversity in a particular sample is higher than in a pooled sample. Simple corrections are suggested to avoid this shortcoming.

Propagule type influences competition between two submersed aquatic macrophytes.

Glasshouse competition experiments with Hydrilla verticillata (L.f.) Royle indicate that plants grown from turions are weaker competitors than those grown from tubers, when compared to the widely distributed macrophyte, Potamogeton pectinatus L. These results support an earlier hypothesis about the importance of propagule size for predicting the outcome of plant competition (Grace 1985; Schaffer and Gadgil 1980). Results of outdoor growth experiments indicate that even though Hydrilla plants from turions are relatively weaker competitors, they are able to grow succesfully in an existing macrophyte bed composed of either, P. pectinatus or P. gramineus. During the early stages of Hydrilla invasion into an area of existing macrophytes, native plants may coexist with Hydrilla. However, once the abundance of Hydrilla tubers in the sediment increases, Hydrilla may displace existing plants.

The fecundity and long-term variability in abundance of noctuid moths (Lepidoptera, Noctuidae).

The relationship between fecundity and long-term adult fluctuations of 29 temperate noctuid moths was investigated and is explained in terms of the r- and K-selection strategies. The potential population growth rate per year (PGR) and the "rate of return to equilibrium" (sensu Pimm 1982) have been calculated and correlated with generation-to-generation fluctuations. A highly significant correlation was found between PGR and the coefficient of generation-to-generation variation. The r-strategy species possess the highest PGR, the highest generation-to-generation coefficient of variation and their larvae are polyphagous. Species with typical features of K-strategists exhibit the lowest PGR values and the lowest generation-to-generation coefficients of variation. Their larvae seem to be exclusively oligophagous.

Do epigeal termite mounds increase the diversity of plant habitats in a tropical rain forest in peninsular Malaysia?

The extent to which environmental heterogeneity can account for tree species coexistence in diverse ecosystems, such as tropical rainforests, is hotly debated, although the importance of spatial variability in contributing to species co-existence is well recognized. Termites contribute to the micro-topographical and nutrient spatial heterogeneity of tropical forests. We therefore investigated whether epigeal termite mounds could contribute to the coexistence of plant species within a 50 ha plot at Pasoh Forest Reserve, Malaysia. Overall, stem density was significantly higher on mounds than in their immediate surroundings, but tree species diversity was significantly lower. Canonical correspondence analysis showed that location on or off mounds significantly influenced species distribution when stems were characterized by basal area. Like studies of termite mounds in other ecosystems, our results suggest that epigeal termite mounds provide a specific microhabitat for the enhanced growth and survival of certain species in these species-rich tropical forests. However, the extent to which epigeal termite mounds facilitate species coexistence warrants further investigation.