Hurricanes overcome migration lag and shape intraspecific genetic variation beyond a poleward mangrove range limit

Expansion of many tree species lags behind climate change projections. Extreme storms can rapidly overcome this lag, especially for coastal species, but how will storm‐driven expansion shape intraspecific genetic variation? Do storms provide recruits only from the nearest sources, or from more distant sources? Answers to these questions have ecological and evolutionary implications, but empirical evidence is absent from the literature. In 2017, Hurricane Irma provided an opportunity to address this knowledge gap at the northern range limit of the neotropical black mangrove (Avicennia germinans) on the Atlantic coast of Florida, USA. We observed massive post‐hurricane increases in beach‐stranded A. germinans propagules at, and past, this species’ present day range margin when compared to a previously surveyed nonhurricane year. Yet, propagule dispersal does not guarantee subsequent establishment and reproductive success (i.e., effective dispersal). We also evaluated prior effective dispersal along this coastline with isolated A. germinans trees identified beyond the most northern established population. We used 12 nuclear microsatellite loci to genotype 896 hurricane‐driven drift propagules from nine sites and 10 isolated trees from four sites, determined their sources of origin, and estimated dispersal distances. Almost all drift propagules and all isolated trees came from the nearest sources. This research suggests that hurricanes are a prerequisite for poleward range expansion of a coastal tree species and that storms can shape the expanding gene pool by providing almost exclusively range‐margin genotypes. These insights and empirical estimates of hurricane‐driven dispersal distances should improve our ability to forecast distributional shifts of coastal species.     

Impact of the spatial uncertainty of seed dispersal on tree colonization dynamics in a temperate forest

An aggregated distribution of dispersed seeds may influence the colonization process in tree communities via inflated spatial uncertainty. To evaluate this possibility, we studied 10 tree species in a temperate forest: one primarily barochorous, six anemochorous and two endozoochorous species. A statistical model was developed by combining an empirical seed dispersal kernel with a gamma distribution of seedfall density, with parameters that vary with distance. In the probability density, the fitted models showed that seeds of Fagaceae (primarily barochorous) and Betulaceae (anemochorous) were disseminated locally (i.e. within 60 m of a mother tree), whereas seeds of Acer (anemochorous) and endozoochorous species were transported farther. Greater fecundity compensated for the lower probability of seed dispersal over long distances for some species. Spatial uncertainty in seedfall density was much greater within 60 m of a mother tree than farther away, irrespective of dispersal mode, suggesting that seed dispersal is particularly aggregated in the vicinity of mother trees. Simulation results suggested that such seed dispersal patterns could lead to sites in the vicinity of a tree being occupied by other species that disperse seeds from far away. We speculate that this process could promote coexistence by making the colonization rates of the species more similar on average and equalizing species fitness in this temperate forest community.     

Reconstruction of a windborne insect invasion using a particle dispersal model,historical wind data,and Bayesian analysis of genetic data

Understanding how invasive species establish and spread is vital for developing effective management strategies for invaded areas and identifying new areas where the risk of invasion is highest. We investigated the explanatory power of dispersal histories reconstructed based on local‐scale wind data and a regional‐scale wind‐dispersed particle trajectory model for the invasive seed chalcid wasp Megastigmus schimitscheki (Hymenoptera: Torymidae) in France. The explanatory power was tested by: (1) survival analysis of empirical data on M. schimitscheki presence, absence and year of arrival at 52 stands of the wasp's obligate hosts, Cedrus (true cedar trees); and (2) Approximate Bayesian analysis of M. schimitscheki genetic data using a coalescence model. The Bayesian demographic modeling and traditional population genetic analysis suggested that initial invasion across the range was the result of long‐distance dispersal from the longest established sites. The survival analyses of the windborne expansion patterns derived from a particle dispersal model indicated that there was an informative correlation between the M. schimitscheki presence/absence data from the annual surveys and the scenarios based on regional‐scale wind data. These three very different analyses produced highly congruent results supporting our proposal that wind is the most probable vector for passive long‐distance dispersal of this invasive seed wasp. This result confirms that long‐distance dispersal from introduction areas is a likely driver of secondary expansion of alien invasive species. Based on our results, management programs for this and other windborne invasive species may consider (1) focusing effort at the longest established sites and (2) monitoring outlying populations remains critically important due to their influence on rates of spread. We also suggest that there is a distinct need for new analysis methods that have the capacity to combine empirical spatiotemporal field data, genetic data, and environmental data to investigate dispersal and invasion.     

Reproductive variability in Pinus sylvestris in southern France: Implications for invasion

Abstract. Though fecundity and dispersal have been recognized as major factors in most invasion models, their ecological determinants are still poorly known. This paper aims to identify the main sources of variation in seed production and dispersal distance in a naturally expanding Pinus sylvestris population. We propose some tree measurements that may be related to their contribution to population expansion. We quantified cone and seed production and measured three seed characteristics related to their dispersal ability: mass, wing area and wing‐loading (mass:area ratio) in cones sampled at different relative heights and aspects in the canopy and in trees of different age and competitive status. Results showed that isolated trees had a much more abundant cone production, which was more evenly distributed in the canopy than trees within stands. Age was also positively related to cone production. Seed dimensions varied between and within trees but we found no effect of isolation or age. The strong positive correlation between wing area and seed mass leads to a limited variability of seed wing loading. Seed characteristics may thus play a minor role in individual dispersal ability and relative tree position in the stand that is strongly linked to tree fecundity should be more appropriate for estimating the individual contribution to the whole population expansion. Our results also highlight the importance of obtaining demographic data in low‐density populations to estimate the invasive potential of a species.     

Recruitment limitation in secondary forests dominated by an exotic tree

Question: What factors limit woody plant recruitment in a mosaic landscape where former agricultural lands are dominated by the invasive tree Ligustrum lucidum (Oleaceae)? Location: Subtropical northwestern Argentina. Methods: In secondary forest patches, we measured (1) tree, shrub and liana abundance in different size classes; (2) seed rain of Ligustrum and two native trees and (3) topographic, soil and light variables. We used spatial autoregressive models to test for effects of Ligustrum dominance and environment on native plant abundance in each size class. We used multiple regression on resemblance matrices to quantify the relative importance of spatial (e.g. dispersal) and environmental effects on native species composition. Results: Native tree abundance in the smallest size class was unrelated to Ligustrum canopy dominance, while native tree abundance in larger size classes and native liana abundance were negatively correlated with Ligustrum dominance. Native species composition was both environmentally and spatially structured, suggesting that some species are dispersal limited. Seed rain was spatially correlated with conspecific basal area for one of two native species, but not for Ligustrum. Conclusions: Native tree recruitment appears to be limited primarily by sapling mortality in patches dominated by the invasive Ligustrum. Ligustrum does not appear to be dispersal limited in our study area and is likely to continue spreading. Invaded patches may persist for hundreds of years.     

Patterns of Seed Dispersal and Dispersal Failure in a Hawaiian Dry Forest Having Only Introduced Birds

Dry forests are among the most endangered natural communities in the Hawaiian Islands. Most have been reduced to isolated trees and small forest fragments in which native tree species reproduce poorly. The replacement of native birds by introduced generalists may be contributing to dry forest decline through modification of seed dispersal patterns. To document seed dispersal by introduced birds, we conducted foraging observations on fleshy-fruited trees and measured seed rain under trees and in adjacent open areas for 1 year in a dry forest dominated by native trees. Although trees covered only 15.2 percent of the study area, 96.9 percent of the bird-dispersed seeds were deposited beneath them. The Japanese white-eye (Zosterops japonicus) was the principal dispersal agent. Among bird-dispersed seeds, those of the invasive tree Bocconia frutescens accounted for 75 percent of all seeds collected beneath trees (14.8 seeds/m²/yr) and the invasive shrub Lantana camara accounted for 17 percent. Although nearly 60 percent of the reserve's native woody species possess fleshy fruits, introduced birds rarely disperse their seeds. Native trees accounted for <8 percent of all bird-dispersed seeds and are consequently experiencing dispersal failure by falling directly under parent trees. Smaller-seeded non-native plants, in contrast, may be benefiting from dispersal by introduced birds. Current dispersal patterns suggest that these readily disseminated non-native plants may eventually replace the remaining native flora.     

Phylogenetic dating with confidence intervals using mean path lengths

The mean path length (MPL) method, a simple method for dating nodes in a phylogenetic tree, is presented. For small trees the age estimates and corresponding confidence intervals, calibrated with fossil data, can be calculated by hand, and for larger trees a computer program gives the results instantaneously (a Pascal program is available upon request). Necessary input data are a rooted phylogenetic tree with edge lengths (internode lengths) approximately corresponding to the number of substitutions between the nodes. Given this, the MPL method produces relative age estimates with confidence intervals for all nodes of the tree. With the age of one or several nodes of the tree being known from reference fossils, the relative age estimates induce absolute age estimates and confidence intervals of the nodes of the tree. The MPL method relies on the assumptions that substitutions occur randomly and independently in different sites in the DNA sequence and that the substitution rates are approximately constant in time, i.e., assuming a molecular clock. A method is presented for identification of the nodes in the tree at which significant deviations from the clock assumption occur, such that dating may be done using different rates in different parts of the tree. The MPL method is illustrated with the Liliales, a group of monocot flowering plants.     

In situ regeneration of Pinus strobus and P. resinosa in the Great Lakes forest communities of Canada

Abstract. Two extensive forest vegetation survey datasets are explored, using ordination and classification, for evidence of in situ regeneration by Pinus strobus (Eastern white pine) and P. resinosa (Red pine). Ordination of tree species contributions to total basal area in 320 upland northern hardwood- conifer stands produced distinct stand groups for P.banksiana, P. resinosa, P. strobus and mesic hardwoods in an ascending sequence along the first axis. Quercus rubra (red oak), Q. alba (white oak) and tolerant conifer groups formed segregates from the hardwood complex along the second axis. P. strobus mixes with all other forest types, but P. resinosa is restricted to its own group. Seedlings and trees of P. strobus are more abundant than saplings, which are restricted to the pine and oak forests. Therefore, seed production, dispersal and seedling establishment seem to be less of a barrier to in situ regeneration by P. strobus than subsequent survival and growth. Canonical correspondence analysis of 170 pine-dominated stands from the Canadian Shield of Ontario, in which tree species variables are segmented into height-class pseudo- species, yielded no linear relationship between environmental features or stand structure and seedling densities of P. strobus. However, total tree basal area appears to impose an upper limit to seedling density on the forest floor. Strong correlations emerged between pine seedling density and understorey vegetation. Stand classification of the understorey vegetation, using constrained indicator species analysis, yielded distinct high and low seedling groups. Low pine seedling density was associated with abundant broadleaved shrubs, herbs and seedlings as well as feathermosses and tolerant conifers. High seedling density could not be ascribed to the presence of seedbed taxa, such as Polytrichum, but is ascribed to the absence of competition and other forms of inhibition in the understorey vegetation and down through the canopy profile. In situ regeneration of P. strobus does, therefore, occur but conditions over the forest landscape are largely restrictive.     

Acorn production prediction models for five common oak species of the eastern United States

Acorn production varies considerably among oak (Quercus) species, individual trees, years, and locations, which directly affects oak regeneration and populations of wildlife species that depend on acorns for food. Hard mast indices provide a relative ranking and basis for comparison of within- and between-year acorn crop size at a broad scale, but do not provide an estimate of actual acorn yield—the number of acorns that can potentially be produced on a given land area unit based on the species, number, and diameter at breast height (dbh) of oak trees present. We used 10 years of acorn production data from 475 oak trees to develop predictive models of potential average annual hard mast production by five common eastern oak species, based on tree diameter and estimated crown area. We found a weak (R² = 0.08–0.28) relationship between tree dbh and acorn production per unit crown area for most species. The relationship between tree dbh and acorn production per tree was stronger (R² = 0.33–0.57). However, this is because larger-dbh trees generally have larger crowns, not because they have a greater capacity to produce more acorns per unit crown area. Acorn production is highly variable among individual trees. We estimated that dbh of at least 60 dominant or codominant oak trees per species should be randomly sampled to obtain an adequate representation of the range of dbhs (≥12.7 cm dbh) in a given forest area, and achieve precise estimates when using these equations to predict potential acorn production. Our predictive models provide a tool for estimating potential acorn production that land managers and forest planners can apply to oak inventory data to tailor estimates of potential average annual acorn production to different forest management scenarios and multiple spatial scales. © 2011 The Wildlife Society.     

Preliminary studies on a native Australian eucalypt forest invaded by exotic pines

Summary In some parts of Australia, the exotic tree species Pinus radiata is invading native eucalypt forests from adjacent plantations. At one site, measurements have been made in order to determine the dispersal gradient and rate of population increase of the self-sown pines. From these, it is evident that, barring further human interference, a new type of mixed forest community will develop.     

Application of harmonic radar technology to monitor tree snail dispersal

Abstract. Planned conservation efforts for tree snails of the endangered genus Achatinella, endemic to the island of O'ahu, Hawai'i, will include translocations among the remaining wild and captive‐bred populations. In order to establish optimal levels of artificial migration among neighboring groups of snails within fragmented populations, efforts to determine natural dispersal rates through direct observation were initiated. Capture–mark–recapture (CMR) efforts have proved inadequate for obtaining the requisite dispersal estimates, due to low recapture probabilities. In addition, snail dispersal beyond the boundaries of a finite CMR study site was indistinguishable from mortality. In the preliminary study reported here, both the low recapture probability and dispersal detection problems of past CMR efforts were addressed by using harmonic radar tracking. This approach yielded rough dispersal estimates that were unattainable using CMR alone by providing 100% recapture rates even beyond the normal survey area boundaries. Extensive snail movements within clusters of connected trees were frequently observed after tracking for merely a few hours, although movements between unconnected trees were rare and recorded only after monthly survey intervals. Just 11 out of 40 tracked snails made between‐tree movements (average distance of 4.94±1.52 m) during the entire 7‐month study, and provided the only data utilizable for inferring gene flow in and out of subpopulations. Meteorological data loggers were deployed when tracking began to look for an association between such snail movement and weather fluctuations. The resultant data indicate that increases in both wind gusts and humidity facilitate dispersal (R²=0.77, p‐value <0.001), and that passive wind dispersal alone may be responsible for many snail movements (R²=0.59, p‐value=0.0014). Despite having provided coarse estimates of short‐term dispersal and corresponding wind influences, the limitations of the radar method can be substantial.     

Estimates of connectivity reveal non-equilibrium epiphyte occurrence patterns almost 180 years after habitat decline

Habitat loss is a major cause of species decline and extinction. Immediately after habitat loss, species occurrences are not in equilibrium with the new landscape and more closely reflect the previous landscape structure. Species with slow colonisation–extinction dynamics may display long time-lags before reaching a new equilibrium. We investigated the importance of connectivity to current and historical dispersal sources with the aim of explaining the occurrence pattern of epiphytic lichens with different traits among 104 old oaks. We used oak survey data collected from 1830 and 2009 for a Swedish landscape where oak densities declined drastically shortly after 1830. We fitted a commonly used connectivity measure and estimated the confidence interval for the spatial scale parameter. Small differences in the spatial scale parameter resulted in large differences in model fit. Connectivity to trees in 1830 better explained the occurrence of three of the four species compared to the connectivity in 2009. The explanatory power of the historical landscape structure was highest for the species with traits that may result in a low colonisation rate—both a narrow niche (here few suitable trees) and large dispersal propagules. The results suggest that oak-dependent epiphytic lichens have not reached equilibrium with the spatial landscape structure 180 years after the drastic decline in habitat. For the long-term persistence of epiphytes associated with old trees, conservation efforts should focus on (1) protecting and restoring stands where specialised species with large dispersal propagules (i.e. with low colonisation rates) occur today and (2) promoting tree regeneration in their near vicinity.     

Estimating species trees using approximate Bayesian computation

Development of methods for estimating species trees from multilocus data is a current challenge in evolutionary biology. We propose a method for estimating the species tree topology and branch lengths using approximate Bayesian computation (ABC). The method takes as data a sample of observed rooted gene tree topologies, and then iterates through the following sequence of steps: First, a randomly selected species tree is used to compute the distribution of rooted gene tree topologies. This distribution is then compared to the observed gene topology frequencies, and if the fit between the observed and the predicted distributions is close enough, the proposed species tree is retained. Repeating this many times leads to a collection of retained species trees that are then used to form the estimate of the overall species tree. We test the performance of the method, which we call ST-ABC, using both simulated and empirical data. The simulation study examines both symmetric and asymmetric species trees over a range of branch lengths and sample sizes. The results from the simulation study show that the model performs very well, giving accurate estimates for both the topology and the branch lengths across the conditions studied, and that a sample size of 25 loci appears to be adequate for the method. Further, we apply the method to two empirical cases: a 4-taxon data set for primates and a 7-taxon data set for yeast. In both cases, we find that estimates obtained with ST-ABC agree with previous studies. The method provides efficient estimation of the species tree, and does not require sequence data, but rather the observed distribution of rooted gene topologies without branch lengths. Therefore, this method is a useful alternative to other currently available methods for species tree estimation.     

The impact of fossil stratigraphic ranges on tip-calibration,and the accuracy and precision of divergence time estimates

The molecular clock provides the only viable means of establishing realistic evolutionary timescales but it remains unclear how best to calibrate divergence time analyses. Calibrations can be applied to the tips and/or to the nodes of a phylogeny. Tip-calibration is an attractive approach since it allows fossil species to be included alongside extant relatives in molecular clock analyses. However, most fossil species are known from multiple stratigraphical horizons and it remains unclear how such age ranges should be interpreted to codify tip-calibrations. We use simulations and empirical data to explore the impact on precision and accuracy of different approaches to informing tip-calibrations. In particular, we focus on the effect of using tip-calibrations defined using the oldest vs youngest stratigraphic occurrences, the full stratigraphical range, as well as confidence intervals on these data points. The results of our simulations show that using different calibration approaches leads to different divergence-time estimates and demonstrate that concentrating tip-calibrations near the root of the dated phylogeny improves both precision and accuracy of estimated divergence times. Finally, our results indicate that the highest levels of accuracy and precision are achieved when fossil tips are calibrated based on the fossil occurrence from which the morphological data were derived. These trends were corroborated by analysis of an empirical dataset for Ursidae. Overall, we conclude that tip-dating analyses should, in particular, employ tip calibrations close to the root of the tree and they should be calibrated based on the age of the fossil used to inform the morphological data used in Total Evidence Dating.     

Seed dispersal and spatial pattern in tropical trees

Theories of tropical tree diversity emphasize dispersal limitation as a potential mechanism for separating species in space and reducing competitive exclusion. We compared the dispersal morphologies, fruit sizes, and spatial distributions of 561 tree species within a fully mapped, 50-hectare plot of primary tropical forest in peninsular Malaysia. We demonstrate here that the extent and scale of conspecific spatial aggregation is correlated with the mode of seed dispersal. This relationship holds for saplings as well as for mature trees. Phylogenetically independent contrasts confirm that the relationship between dispersal and spatial pattern is significant even after controlling for common ancestry among species. We found the same qualitative results for a 50-hectare tropical forest plot in Panama. Our results provide broad empirical evidence for the importance of dispersal mode in establishing the long-term community structure of tropical forests.     

Loss of animal seed dispersal increases extinction risk in a tropical tree species due to pervasive negative density dependence across life stages

Overhunting in tropical forests reduces populations of vertebrate seed dispersers. If reduced seed dispersal has a negative impact on tree population viability, overhunting could lead to altered forest structure and dynamics, including decreased biodiversity. However, empirical data showing decreased animal-dispersed tree abundance in overhunted forests contradict demographic models which predict minimal sensitivity of tree population growth rate to early life stages. One resolution to this discrepancy is that seed dispersal determines spatial aggregation, which could have demographic consequences for all life stages. We tested the impact of dispersal loss on population viability of a tropical tree species, Miliusa horsfieldii, currently dispersed by an intact community of large mammals in a Thai forest. We evaluated the effect of spatial aggregation for all tree life stages, from seeds to adult trees, and constructed simulation models to compare population viability with and without animal-mediated seed dispersal. In simulated populations, disperser loss increased spatial aggregation by fourfold, leading to increased negative density dependence across the life cycle and a 10-fold increase in the probability of extinction. Given that the majority of tree species in tropical forests are animal-dispersed, overhunting will potentially result in forests that are fundamentally different from those existing now.     

The impact of gene-tree/species-tree discordance on diversification-rate estimation

Molecular phylogenies are often used to test hypotheses about the tempo and mode of speciation and extinction. One commonly used statistic is Pybus and Harvey's γ, which measures the density of ordered internode distances on an ultrametric tree to infer earlier (negative γ) or later (positive γ) bursts of diversification. However, coalescent theory predicts that γ might be biased toward negative values (inferring early bursts of diversification) when using gene trees rather than species trees. Gene divergences predate species divergences, increasingly so at higher effective population sizes (N(e)), and proportionally more so toward the tips of the tree. Thus, gene trees will have a higher density of older nodes in many cases (particularly at higher N(e)), due to the disproportionate lengthening of terminal branches. This will yield an artifactual signature of early bursts of diversification when estimating γ from gene trees. We simulate gene trees within species trees under both Yule (pure-birth) and birth-death processes, and demonstrate support for these predictions. However, for most realistic estimates of θ in natural populations, gene trees provide relatively good estimates of γ, despite the disproportionate overestimation of younger node ages. This is corroborated with an empirical dataset of North American fence lizards (Sceloporus).     

Dispersal and recruitment limitation in native versus exotic tree species: life‐history strategies and Janzen‐Connell effects

Life‐history traits of invasive exotic plants are typically considered to be exceptional vis‐à‐vis native species. In particular, hyper‐fecundity and long range dispersal are regarded as invasive traits, but direct comparisons with native species are needed to identify the life‐history stages behind invasiveness. Until recently, this task was particularly problematic in forests as tree fecundity and dispersal were difficult to characterize in closed stands. We used inverse modelling to parameterize fecundity, seed dispersal and seedling dispersion functions for two exotic and eight native tree species in closed‐canopy forests in Connecticut, USA. Interannual variation in seed production was dramatic for all species, with complete seed crop failures in at least one year for six native species. However, the average per capita seed production of the exotic Ailanthus altissima was extraordinary: > 40 times higher than the next highest species. Seed production of the shade tolerant exotic Acer platanoides was average, but much higher than the native shade tolerant species, and the density of its established seedlings (≥ 3 years) was higher than any other species. Overall, the data supported a model in which adults of native and exotic species must reach a minimum size before seed production occurred. Once reached, the relationship between tree diameter and seed production was fairly flat for seven species, including both exotics. Seed dispersal was highly localized and usually showed a steep decline with increasing distance from parent trees: only Ailanthus altissima and Fraxinus americana had mean dispersal distances > 10 m. Janzen‐Connell patterns were clearly evident for both native and exotic species, as the mode and mean dispersion distance of seedlings were further from potential parent trees than seeds. The comparable intensity of Janzen‐Connell effects between native and exotic species suggests that the enemy escape hypothesis alone cannot explain the invasiveness of these exotics. Our study confirms the general importance of colonization processes in invasions, yet demonstrates how invasiveness can occur via divergent colonization strategies. Dispersal limitation of Acer platanoides and recruitment limitation of Ailanthus altissima will likely constitute some limit on their invasiveness in closed‐canopy forests.     

Frugivory on Persea lingue in temperate Chilean forests: interactions between fruit availability and habitat fragmentation across multiple spatial scales

Habitat degradation and fragmentation are expected to reduce seed dispersal rates by reducing fruit availability as well as the movement and abundance of frugivores. These deleterious impacts may also interact with each other at different spatial scales, leading to nonlinear effects of fruit abundance on seed dispersal. In this study we assessed whether the degradation and fragmentation of southern Chilean forests had the potential to restrict seed dispersal the lingue (Persea lingue) tree, a fleshy-fruited tree species. Of five frugivore bird species, the austral thrush (Turdus falcklandii) and the fire-eyed diucon (Xolmis pyrope) were the only legitimate seed dispersers as well as being the most abundant species visiting lingue trees. The results showed little or no direct effect of habitat fragmentation on seed dispersal estimates, possibly because the assemblage of frugivore birds was comprised habitat-generalist species. Instead, the number of fruits removed per focal tree exhibited an enhanced response to crop size, but only in the more connected fragments. In the fruit-richer fragment networks, there was an increased fragment-size effect on the proportion of fruits removed in comparison to fruit-poor networks in which the fragment size effect was spurious. We suggest that such nonlinear effects are widespread in fragmented forest regions, resulting from the link between the spatial scales over which frugivores sample resources and the spatial heterogeneity in fruiting resources caused by habitat fragmentation and degradation.