Destination‐based seed dispersal homogenizes genetic structure of a tropical palm

As the dominant seed dispersal agents in many ecosystems, frugivorous animals profoundly impact gene movement and fine‐scale genetic structure of plants. Most frugivores engage in some form of destination‐based dispersal, in that they move seeds towards specific destinations, resulting in clumped distributions of seeds away from the source tree. Molecular analyses of dispersed seeds and seedlings suggest that destination‐based dispersal may often yield clusters of maternal genotypes and lead to pronounced local genetic structure. The long‐wattled umbrellabird Cephalopterus penduliger is a frugivorous bird whose lek mating system creates a species‐specific pattern of seed dispersal that can potentially be distinguished from background dispersal processes. We used this system to test how destination‐based dispersal by umbrellabirds into the lek affects gene movement and genetic structure of one of their preferred food sources Oenocarpus bataua, a canopy palm tree. Relative to background dispersal processes, umbrellabird mating behaviour yielded more diverse seed pools in leks that included on average five times more seed sources and a higher incidence of long‐distance dispersal events. This resulted in markedly lower fine‐scale spatial genetic structure among established seedlings in leks than background areas. These species‐specific impacts of destination‐based dispersal illustrate how detailed knowledge of disperser behaviour can elucidate the mechanistic link driving observed patterns of seed movement and genetic structure.     

Joint estimation of contemporary seed and pollen dispersal rates among plant populations

There are few statistical methods for estimating contemporary dispersal among plant populations. A maximum-likelihood procedure is introduced here that uses pre- and post-dispersal population samples of biparentally inherited genetic markers to jointly estimate contemporary seed and pollen immigration rates from a set of discrete external sources into a target population. Monte Carlo simulations indicate that accurate estimates and reliable confidence intervals can be obtained using this method for both pollen and seed migration rates at modest sample sizes (100 parents/population and 100 offspring) when population differentiation is moderate (F(ST) ≥ 0.1), or by increasing pre-dispersal samples (to about 500 parents/population) when genetic divergence is weak (F(ST) = 0.01). The method exhibited low sensitivity to the number of source populations and achieved good accuracy at affordable genetic resolution (10 loci with 10 equifrequent alleles each). Unsampled source populations introduced positive biases in migration rate estimates from sampled sources, although they were minor when the proportion of immigration from the latter was comparatively low. A practical application of the method to a metapopulation of the Australian resprouter shrub Banksia attenuata revealed comparable levels of directional seed and pollen migration among dune groups, and the estimate of seed dispersal was higher than a previous estimate based on conservative assignment tests. The method should be of interest to researchers and managers assessing broad-scale nonequilibrium seed and pollen gene flow dynamics in plants.     

Seed and pollen dispersal by animals: convergence in the ecology of contamination and sloppy harvest

Seed dispersal and pollen movement by animals have very much in common, though we know so little of either that it is hard to be specific. The focus has been on animals at the fruit crop rather than the seed shadows that they generate (and the fate of those seeds), and on the animals that arrive at flowers rather than where they take the pollen obtained or whence came the pollen they are carrying. 'Seed dispersal' is a word that does not imply the fate of the seed; 'pollination' relates to the fate of the pollen grain, and therefore the two words are not of parallel meaning. In like manner, seeds (the zygote contained within, are different individuals from the parents while pollen is much more like the parent. The following areas are discussed with respect to how seed and pollen movement compare: ability of parent plant to assess success; gene flow; dependency of animals on seeds, pollen and bait; disruption by animal loss; secondary movements; fate of most seeds and pollen; syndromes; selection for maximization of movement; exclusion of 'unwanted' animals; adjustments to the needs of the animals; seeds and pollen as contaminants; necessity of dispersal and outcrossing. We need much more knowledge of what actually is happening in nature with seed and pollen shadows, and improved ability to think like a plant.     

Pollen and seed dispersal inferred from seedling genotypes: the Bayesian revolution has passed here too

Understanding precisely how plants disperse their seeds and pollen in their neighbourhood is a central question for both ecologists and evolutionary biologists because seed and pollen dispersal governs both the rate of spread of an expanding population and gene flow within and among populations. The concept of a 'dispersal kernel' has become extremely popular in dispersal ecology as a tool that summarizes how dispersal distributes individuals and genes in space and at a given scale. In this issue of Molecular Ecology, the study by Moran & Clark (2011) (M&C in the following) shows how genotypic and spatial data of established seedlings can be analysed in a Bayesian framework to estimate jointly the pollen and seed dispersal kernels and finally derive a parentage analysis from a full-probability approach. This approach applied to red oak shows important dispersal of seeds (138 m on average) and pollen (178 m on average). For seeds, this estimate contrasts with previous results from inverse modelling on seed trap data (9.3 m). This research gathers several methodological advances made in recent years in two research communities and could become a cornerstone for dispersal ecology.     

Temporal and spatial patterns of seed dispersal in two Cistus species (Cistaceae)

Cistus species are obligate seeding, early colonizers that follow disturbance, particularly fire, in Mediterranean ecosystems. We studied seed release, seed dispersal and soil seed populations in stands of Cistus ladanifer and C. libanotis. Seed release started in mid- to late summer (C. ladanifer) or in early autumn (C libanotis), and continued for a very extended period: 8-10 months in C. ladanifer, and for a mean of 16 months in C. libanotis. The xerochastic capsules of both species released seeds by successive dehiscence of the locules. All capsules begin to dehisce simultaneously at the start of the seed release period, but in C. libanotis capsule fragmentation replaced dehiscence early in the seed release period. In plants of both species, seed shadows were characterized by a peak of density beneath the plant canopy and a very short tail of much lower densities, indicating that seeds are concentrated beneath mother plants when dispersed. Nevertheless, in late May, at the onset of the fire season, soil seed densities beneath plant canopies were low compared with densities expected from seed shadows, but were apparently high enough to allow recovery of the stands if a disturbance, such as fire, had taken place. Seed-eating Bruchidae in summer and granivorous ants during the seed release period were apparently the main causes of seed losses. Results suggest that in both Cistus species, the staggered seed release could constitute an efficient risk-reducing trait. The plant pool of seeds existing throughout most of the year could be a relevant component of Cistus seed banks.     

Types of pollination and seed dispersal in Mediterranean plants

Abstract

Selected examples of pollen and seed dispersal in Mediterranean plants are described. The aspects of pollination considered are: comparison between cleistogamous and chasmogamous forms in the same species; differences in attractants and rewards; duration of pollen viability according to pollination syndrome. The aspects of seed dispersal considered are: presence or absence of specialized structures for dispersal; examples of active, passive and induced dispersal; animals involved in seed dispersal and their reward; type and functions of elaiosomes.     

Understanding the population genetic structure of Gleditsia triacanthos L.: seed dispersal and variation in female reproductive success

Fine-scale genetic structuring is influenced by a variety of ecological factors and can directly affect the evolutionary dynamics of plant populations by influencing effective population size and patterns of viability selection. In many plant species, genetic structuring within populations may result from highly localized patterns of seed dispersal around maternal plants or by the correlated dispersal and recruitment of siblings from the same fruit. This fine-scale genetic structuring may be enhanced if female parents vary significantly in their reproductive success. To test these hypotheses, we used genetic data from 17 allozyme loci and a maximum-likelihood, ‘maternity-analysis’ model to estimate individual female fertilities for maternal trees across a large number of naturally established seedlings and saplings in two populations of Gleditsia triacanthos L. (Leguminosae). Maximum-likelihood fertility estimates showed that the three highest fertility females accounted for 58% of the 313 progeny at the first site and 46% of the 651 progeny at the second site, whereas 18 of 35 and 16 of 34 females, respectively, had fertility estimates that did not exceed 1%. Additional analyses of the second site found individual female fertility to vary significantly both within and among juvenile age classes. Female fertility at the first site was weakly correlated with maternal tree size and spatial location relative to the open, old-field portions of the population, where the great majority of seedlings and saplings were growing, but no such correlations were found at the second site. Estimates of realized seed dispersal distances indicated that dispersal was highly localized at the first site, but was nearly random at the second site, possibly reflecting differences between the two sites in the behaviour of animal dispersers. The combined estimates of seed dispersal patterns and fertility variation are sufficient to explain previously described patterns of significant fine-scale spatial genetic structure in these two populations. In general, our results demonstrate that effective seed dispersal distributions may vary significantly from population to population of a species due to the unpredictable behaviour of secondary dispersers. Consequently, the effects of seed dispersal on realized fine-scale genetic structure may also be relatively unpredictable.     

Spatial models of pollen dispersal in the forage grass meadow fescue

Several bivariate probability distributions, generated by different underlying dispersal mechanisms, are fitted to the observed frequencies of an isozyme marker gene using a maximum likelihood approach. The pollen dispersal data were generated using two experimental populations of meadow fescue (Festuca pratensis Huds.), homozygous for different allozymes at the (Pgi-2) locus, arranged in a circular donor–acceptor field design. The contribution of a plant depends on plant position, fecundity and flowering time, factors which are taken into account when fitting the different models. Several approximate likelihood-ratio tests are done between alternative nested models, and a wind threshold model with bimodality in the wind direction is selected. The evolutionarily important variances and expectations of gene displacement under the selected model are calculated. It is also shown that the underlying probability distribution is significantly more than exponentially leptokurtic. By fitting a distribution of deposition in all three dimensions to the data, taking into account differences in plant height, separate estimates of additional physical parameters are obtained, showing that gravity and vertical random movements are more important than intervening vegetation in limiting pollen dispersal in meadow fescue. According to the model, plants with a high seed yield contribute pollen over-proportionally to neighbouring plants. This revised version was published online in July 2006 with corrections to the Cover Date.     

A seed predator drives the evolution of a seed dispersal mutualism

Although antagonists are hypothesized to impede the evolution of mutualisms, they may simultaneously exert selection favouring the evolution of alternative mutualistic interactions. We found that increases in limber pine (Pinus flexilis) seed defences arising from selection exerted by a pre-dispersal seed predator (red squirrel Tamiasciurus hudsonicus) reduced the efficacy of limber pine's primary seed disperser (Clark's nutcracker Nucifraga columbiana) while enhancing seed dispersal by ground-foraging scatter-hoarding rodents (Peromyscus). Thus, there is a shift from relying on primary seed dispersal by birds in areas without red squirrels, to an increasing reliance on secondary seed dispersal by scatter-hoarding rodents in areas with red squirrels. Seed predators can therefore drive the evolution of seed defences, which in turn favour alternative seed dispersal mutualisms that lead to major changes in the mode of seed dispersal. Given that adaptive evolution in response to antagonists frequently impedes one kind of mutualistic interaction, the evolution of alternative mutualistic interactions may be a common by-product.     

Studying and managing the risk of cross-fertilization between transgenic crops and wild relatives

Drawing on field studies of pollen dispersal, we identify features of the hybridization process that need quantification. Our emphasis is on standardized measures, as opposed to the idiosyncratic and often anecdotal methods with which gene flow or out-crossing data are currently reported. In addition to proposing specific maximum likelihood approaches, we summarize some results to date from small-scale field trials that bear on the risks anticipated for large-scale commercialization. We conclude that absolute containment of recombinant pollen or genes is unlikely if physical isolation is the only containment strategy. Because we conclude that the escape of transgenic pollen is inevitable, we argue that the focus of risk analysis should be shifted towards the 'invasiveness' of transgenic plants and 'mitigation' of their impact on natural, as well as agricultural systems.     

Molecular evolution of pteridophytes and their relationship to seed plants: Evidence from complete 18S rRNA gene sequences

Complete 18S ribosomal RNA sequence data from representatives of all extant pteridophyte lineages together with RNA sequences from different seed plants were used to infer a molecular phylogeny of vascular plants that included all major land plant lineages. The molecular data indicate that lycopsids are monophyletic and are the earliest diverging group within the vascular land plants, whereasPsilotum nudum is more closely related to the seed plants than to other pteridophyte lineages. The phylogenetic trees based on maximum likelihood, parsimony and distance analyses show substantial agreement with the evolutionary relationships of land plants as interpreted from the fossil record.     

Migration load in plants: role of pollen and seed dispersal in heterogeneous landscapes

Evolution of local adaptation depends critically on the level of gene flow, which, in plants, can be due to either pollen or seed dispersal. Using analytical predictions and individual-centred simulations, we investigate the specific influence of seed and pollen dispersal on local adaptation in plant populations growing in patchy heterogeneous landscapes. We study the evolution of a polygenic trait subject to stabilizing selection within populations, but divergent selection between populations. Deviations from linkage equilibrium and Hardy-Weinberg equilibrium make different contributions to genotypic variance depending on the dispersal mode. Local genotypic variance, differentiation between populations and genetic load vary with the rate of gene flow but are similar for seed and pollen dispersal, unless the landscape is very heterogeneous. In this case, genetic load is higher in the case of pollen dispersal, which appears to be due to differences in the distribution of genotypic values before selection.     

Development and structure of the pericarp and seed of Rhus lancea L. fil. (Anacardiaceae), with taxonomic notes

The pedicel of the female flower of Rhus lancea is distinctly articulated and usually carries three bracteoles. In the linear tetrad the micropylar megaspore forms the 8-nucleate embryo sac of the Oenothera -type. The single, bitegmic ovule is anatropous. The ripe, loose, papery exocarp consists mainly of the outer epidermis and a sclerified hypodermis. The mesocarp is not a typical sarcocarp, since the ridges and the inner layers are sclerenchymatous. The endocarp, originating from the inner epidermis, consists of four layers and its structure and microchemistry emphasize the close alliance of Rhus with other genera of the section Rhoideae. The endotestal seed indicates a phylogenetic affinity between the Anacardiaceae and the Burseraceae.     

Development and structure of the pericarp and seed of Rhus lancea L. fil. (Anacardiaceae), with taxonomic notes

The pedicel of the female flower of Rhus lancea is distinctly articulated and usually carries three bracteoles. In the linear tetrad the micropylar megaspore forms the 8-nucleate embryo sac of the Oenothera-type. The single, bitegmic ovule is anatropous. The ripe, loose, papery exocarp consists mainly of the outer epidermis and a sclerified hypodermis. The mesocarp is not a typical sarcocarp, since the ridges and the inner layers are sclerenchymatous. The endocarp, originating from the inner epidermis, consists of four layers and its structure and microchemistry emphasize the close alliance of Rhus with other genera of the section Rhoideae. The endotestal seed indicates a phylogenetic affinity between the Anacardiaceae and the Burseraceae.