Remnant Pachira quinata pasture trees have greater opportunities to self and suffer reduced reproductive success due to inbreeding depression

Habitat fragmentation is extensive throughout the world, converting natural ecosystems into fragments of varying size, density and connectivity. The potential value of remnant trees in agricultural landscapes as seed sources and in connecting fragments has formed a fertile area of debate. This study contrasted the mating patterns of bat-pollinated Pachira quinata trees in a continuous forest to those in pasture through microsatellite-based paternity analysis of progeny. The breeding system was determined by analysis of pollen tube growth and seed production from controlled pollinations. Fitness of selfed and outcrossed seed was compared by germination and seedling growth. There was more inbreeding within pasture trees (outcrossing=0.828±0.015) compared with forest trees (0.926±0.005). Pasture trees had fewer sires contributing to mating events, but pollen dispersal distances were greater than those in the forest. Paternity analysis showed variation in outcrossing rates among pasture trees with high proportions of external and self pollen sources detected. A leaky self-incompatibility system was found, with self pollen having reduced germination on stigmas and slower growth rate through the style. Controlled pollinations also showed a varied ability to self among trees, which was reflected in the selfing rates among pasture trees shown by the paternity analysis (0–80% selfing). Self pollination resulted in lower seed set, germination and seedling growth compared with outcrossing. While remnant trees in agricultural landscapes are involved in broader mating patterns, they show increased but varied levels of inbreeding, which result in reduced fitness.     

Pollen load size, reproductive success, and progeny kinship of naturally pollinated flowers of the tropical dry forest tree Pachira quinata (Bombacaceae)

Several studies have demonstrated, using controlled pollinations, that the number and identity of pollen grains deposited onto a flower's stigma affect the reproductive success of plants. However, few studies have shown this relationship under conditions of natural pollination. Using the tropical dry forest tree Pachira quinata, we evaluated the relationship between the number of microgametophytes per pistil and the number of sires with respect to the production of fruits and seeds in a natural population of Pachira quinata. Our study demonstrates that fruit and seed production are directly related to the number of microgametophytes per pistil in natural populations of P. quinata. Only 6% of the marked flowers developed into mature fruits and 10% of the marked flowers initiated fruits but later aborted them. A mean of 23 pollen grains were required to produce a seed. Flowers with >400 pollen grains on the stigma always developed into mature fruits, whereas flowers that received <200 grains never matured fruits. Half of the pollen grains transferred to a flower stigma germinated and developed pollen tubes to the base of the style. The number of pollen grains on a stigma explained 34% of the variation in seed number per fruit, and the number of seeds produced per fruit is positively correlated with the size of the seeds. The population of P. quinata studied is predominantly outcrossing, and seeds within fruits are sired by one or a few donors. The total seed crop within trees was sired by three to five donors. Our study examined the implications of the variation in size of microgametophyte loads per pistil with respect to the breeding system and the paternity of progeny under natural conditions. The competitive ability of pollen and pollen tube attrition are important factors regulating fruit production in P. quinata.     

Variation in the breeding behavior of the dry forest tree Enterolobium cyclocarpum (guanacaste) in Costa Rica

We studied the breeding systems of four populations of Enterolobium cyclocarpum (guanacaste, earpod tree) in Costa Rica. Multilocus estimates of the outcrossing rate indicate that E. cyclocarpum is a predominant outcrossing species (t(m) ranged between 0.881 and 0.901) and biparental inbreeding is low (range between 0.058 and 0.079). Overall, our analyses showed significant differences in the gene frequencies between pollen and ovules and significant differences in pollen gene frequencies between the four populations. We also found significant differences in the pollen gene frequencies calculated for single trees in the same population. Outcrossing rates and pollen gene frequencies varied in two consecutive years in two populations of E. cyclocarpum. The correlated mating model revealed that there are differences in the correlation of paternity between populations and years. These findings indicate that there is variation in the average number of trees that father the seed crop of each tree and/or that some fathers are overrepresented in the seed crop of each tree. The implication of these findings for the development of strategies for conservation and management of this species are discussed.     

Pollen coupling of forest trees: forming synchronized and periodic reproduction out of chaos

Many of the tree species in mature forests show masting; their reproductive activity has a large variance between years and is often synchronized between different individuals. In this paper, we analyse a globally coupled map model in which trees accumulate photosynthate every year, produce flowers when the energy reserve level exceeds a threshold, and set seeds and fruits at a rate limited by pollen availability. Without pollen limitation, the trees in the forest show independent chaotic fluctuation. Coupling of trees via pollen exchange results in reproduction being synchronized partially or completely over the forest. The whole forest shows diverse dynamical behaviors determined by the values of two essential parameters; the depletion coefficient k and the coupling strength beta. We find perfectly synchronized periodic reproduction, synchronized reproduction with a chaotic time series, clustering phenomena, and chaotic reproduction of trees without synchronization over individuals. There are many parameter windows in which synchronized reproduction of trees shows a stable periodic fluctuation. For perfectly synchronized forests, we can calculate all the Lyapunov exponents analytically. They show that synchronized reproduction of all the trees in the forest can only occur when trees flower at low (but positive) levels in a significant fraction of years, resulting in small fruit sets due to outcrossed pollen limitation. This is consistent with the observation that the distinction between mast years and non-mast years is often not clear cut.     

Mating patterns and contemporary gene flow by pollen in a large continuous and a small isolated population of the scattered forest tree Sorbus torminalis

The influence of population size and spatial isolation on contemporary gene flow by pollen and mating patterns in temperate forest trees are not well documented, although they are crucial factors in the life history of plant species. We analysed a small, isolated population and a large, continuous population of the insect-pollinated tree species Sorbus torminalis in two consecutive years. The species recently experienced increased habitat fragmentation due to altered forest management leading to forests with closed canopies. We estimated individual plant size, percentage of flowering trees, intensity of flowering, degree of fruiting and seed set per fruit, and we determined mating patterns, pollen flow distances and external gene flow in a genetic paternity analysis based on microsatellite markers. We found clear effects of small population size and spatial isolation in S. torminalis. Compared with the large, continuous population, the small and isolated population harboured a lower percentage of flowering trees, showed less intense flowering, lower fruiting, less developed seeds per fruit, increased selfing and received less immigrant pollen. However, the negative inbreeding coefficients (F(IS)) of offspring indicated that this did not result in inbred seed at the population level. We also show that flowering, fruiting and pollen flow patterns varied among years, the latter being affected by the size of individuals. Though our study was unreplicated at the factor level (i.e. isolated vs non-isolated populations), it shows that small and spatially isolated populations of S. torminalis may also be genetically isolated, but that their progeny is not necessarily more inbred.     

Pollen dispersal and genetic structure of the tropical tree Dipteryx panamensis in a fragmented Costa Rican landscape

In the face of widespread deforestation, the conservation of rainforest trees relies increasingly on their ability to maintain reproductive processes in fragmented landscapes. Here, we analysed nine microsatellite loci for 218 adults and 325 progeny of the tree Dipteryx panamensis in Costa Rica. Pollen dispersal distances, genetic diversity, genetic structure and spatial autocorrelation were determined for populations in four habitats: continuous forest, forest fragments, pastures adjacent to fragments and isolated pastures. We predicted longer but less frequent pollen movements among increasingly isolated trees. This pattern would lead to lower outcrossing rates for pasture trees, as well as lower genetic diversity and increased structure and spatial autocorrelation among their progeny. Results generally followed these expectations, with the shortest pollen dispersal among continuous forest trees (240 m), moderate distances for fragment (343 m) and adjacent pasture (317 m) populations, and distances of up to 2.3 km in isolated pastures (mean: 557 m). Variance around pollen dispersal estimates also increased with fragmentation, suggesting altered pollination conditions. Outcrossing rates were lower for pasture trees and we found greater spatial autocorrelation and genetic structure among their progeny, as well as a trend towards lower heterozygosity. Paternal reproductive dominance, the pollen contributions from individual fathers, did not vary among habitats, but we did document asymmetric pollen flow between pasture and adjacent fragment populations. We conclude that long-distance pollen dispersal helps maintain gene flow for D. panamensis in this fragmented landscape, but pasture and isolated pasture populations are still at risk of long-term genetic erosion.     

Multiple pollinator visits to Mimulus ringens (Phrymaceae) flowers increase mate number and seed set within fruits

The timing and effectiveness of pollinator visitation to flowers is an important factor influencing mating patterns and reproductive success. Multiple pollinator probes to a flower may increase both the quantity and genetic diversity of progeny, especially if single probes deposit insufficient pollen for maximal seed set or if the interval between probes is brief. When pollen carryover is limited, sequential pollen loads may also differ markedly in sire representation. We hypothesized that these conditions help explain high levels of multiple paternity in Mimulus ringens fruits. We documented all bee visits to individual flowers, quantified resulting seed set, and determined paternity for 20 seeds per fruit. Most (76%) flowers received multiple probes, and the interval between probes was usually <30 min. Flowers probed multiple times produced 44% more seeds than flowers probed once. All fruits were multiply sired. Flowers receiving a single probe averaged 3.12 outcross sires per fruit, indicating that single probes deposit pollen from several donors. Multiple paternity was even greater after three or more probes (4.92 outcross sires), demonstrating that sequential visits bring pollen from donors not represented in the initial probe.     

MATING SYSTEM OF A TROPICAL RAIN FOREST TREE SPECIES

Canopy tree species are the dominant elements of the species-rich, fragile and endangered tropical rain forest ecosystems, yet little is known about the genetics of these species. We provide an estimate of the outcrossing rate in a population of Pithecellobium pedicellare, a large canopy tree in the tropical rain forests of Costa Rica. The outcrossing rate was high (t = 951 ± 0.021) and the pollen pool contributing to the progeny arrays used to measure outcrossing rate showed departure from homogeneity. The high outcrossing rate indicates that individuals scattered over a large area in this low density population could be bound with each other via outcrossing. However, despite a high outcrossing rate, the potential for inbreeding in this population is not negligible. We found a relatively large number of albino and chlorotic seedlings among the progeny arrays of several seed parents. Heterogeneity of pollen pool further indicates that the population, though widely outcrossed, may be structured. A more detailed study of mating systems in tropical rain forest trees may provide additional insights into the mating patterns of these trees. Such studies will be useful not only in understanding the dynamics of micro-evolutionary processes, but also in the conservation and management of tropical forest trees.     

Pollination success in a population of dioecious rain forest trees

Pollination success in female trees was determined for a population of Neolitsea dealbata (R. Br.) Merr., a locally abundant dioecious tree pollinated by small, unspecialized insects in northern Queensland rain forest, Australia. The population consisted of a clustered group of trees with a mean male-to-female distance of 4.5 m and more isolated individuals, including females more than 90 m away from the nearest pollen source. A map of all reproductive trees was produced to determine accurate male-to-female distances. The size of the pollen source available to females was defined as a function of the distance to the nearest ten male trees and their sizes (male neighbourhood index). The rate of pollen movement to females was measured by counting pollen tubes (and the number of tubes per style) in female trees 6 days after the commencement of population flowering. The pollination rate decreased steeply to less than half when the nearest male was only 6.5 m away. Although pollen reached a female 330 m away from the nearest pollen source, only 10% of receptive flowers had been pollinated. The short flowering period (2–3 weeks) combined with the the slow rate of pollen movement means that a large proportion of flowers in isolated trees are unpollinated, confirming an earlier finding that isolated females set fewer fruits than gregarious females. The reliability of pollen transfer to females was determined by quantifying insects and their pollen loads trapped at female trees with a range of male neighbourhood indices. Quantities of insects and pollen were significantly correlated with the size of the male neighbourhood index, indicating a strong density-dependent response by vectors to flowering. Pollen was also collected from insect visitors to non-flowering trees. Females with large male neighbourhood indices received more pollen than non-flowering trees with equivalent male neighbourhood indices. However, when the male neighbourhood indices were small for both female and non-flowering trees, the changces of pollinators encountering female and non-flowering trees were similar, suggesting random movements of pollinators in sparse-flowering sub-populations. The dioecious breeding system, brief, synchronous flowering period, clustered population structure and random, opportunistic foraging behaviour of vectors interacted in a way that reduced reproduction in relatively isolated trees. These results demonstrate a mechanism for differential breeding success between trees in natural populations and emphasize the possible impact of logging regimes on pollen flow between trees. Large interconspecific distances in species-rich environments may have been a factor in the selection for synchronous flowering between trees in outcrossing tree species with generalist insect pollinators.     

Gone with the wind: Negative genetic and progeny fitness consequences of habitat fragmentation in the wind pollinated dioecious tree Brosimum alicastrum

Premise

Habitat fragmentation negatively affects population size and mating patterns that directly affect progeny fitness and genetic diversity; however, little is known about the effects of habitat fragmentation on dioecious, wind pollinated trees. We assessed the effects of habitat fragmentation on population sex ratios, genetic diversity, gene flow, mating patterns, and early progeny vigor in the tropical dioecious tree, Brosimum alicastrum.

Methods

We conducted our study in three continuous and three fragmented forest sites in a Mexican tropical dry forest. We used eight microsatellite loci to characterize the genetic diversity, gene flow via pollen distances, and mean relatedness of progeny. We compared early progeny vigor parameters of seedlings growing under greenhouse conditions.

Results

Sex ratios did not deviate from 1:1 between habitat conditions except for one population in a fragmented habitat, which was female biased. The genetic diversity of adult trees and their offspring was similar in both habitat conditions. Pollen gene flow distances were similar across habitat types; however, paternity correlations were greater in fragmented than in continuous habitats. Germination rates did not differ between habitat conditions; however, progeny from fragmented habitats produced fewer leaves and had a lower foliar area, total height, and total dry biomass than progeny from continuous habitats.

Conclusions

Changes in mating patterns because of habitat fragmentation have negative effects on early progeny vigor. We conclude that negative habitat fragmentation effects on mating patterns and early progeny vigor may be a serious threat to the long-term persistence of tropical dioecious trees.

     

Low Inbreeding and High Pollen Dispersal Distances in Populations of Two Amazonian Forest Tree Species

Recent studies suggest that tropical tree species exhibit low inbreeding and high gene dispersal levels despite the typically low density of conspecifics in tropical forests. To examine this, we undertook a study of pollen gene dispersal and mating system of two Amazonian tree species. We analyzed 341 seeds from 33 trees at four microsatellite loci in a Carapa guianensis population from Brazil, and 212 seeds from 22 trees at four microsatellite loci in a Sextonia rubra population from French Guiana. Differentiation of allele frequencies among the pollen pool of individual trees was Φ_FT= 0.053 (95% CI: 0.027–0.074) for C. guianensis and Φ_FT= 0.064 (95% CI: 0.017–0.088) for S. rubra. The mean pollen dispersal distances were estimated at 69–355 m for C. guianensis , and 86–303 m for S. rubra , depending on the pollen dispersal model and the estimate of reproductive tree density used. The multi-locus outcrossing rate was estimated at 0.918 and 0.945, and the correlation of paternity at 0.089 and 0.096, for C. guianensis and S. rubra , respectively, while no significant levels of biparental inbreeding were detected. Comparing trees with high and low local density of conspecifics, we found no evidence for differences in inbreeding levels. The results are discussed within the framework of the emerging picture of the reproductive biology of tropical forest trees.     

Breeding population size of a fragmented population of a Costa Rican dry forest tree species.

Pollen immigration can offset the effects of genetic drift and inbreeding in small populations. To understand the genetic consequences of forest fragmentation, estimates of pollen flow into remnant fragments are essential. Such estimates are straightforward for plants with singly sired, multiseeded fruits, since the pollen donor genotype for each fruit can be unambiguously reconstructed through full-sib genealogical analyses. Allozyme analyses were used to estimate pollen donor numbers from the progeny of fruits of the tropical dry forest tree Enterolobium cyclocarpum in a small (9.8 ha) fragmented population (N = 11) over three reproductive seasons (1994, 1995, and 1996). These analyses indicate that each tree receives pollen from many pollen donors. When data are pooled for the site, estimated maximum pollen donor pool sizes in all years exceed the number of individuals (56) in the 227 ha study area. Although unidentified pollen donors may be located as close as 250 m to the study trees, the number of unidentified pollen donors indicates that individuals in this forest fragment are part of a large network of reproductively active individuals.     

Outcrossing and paternity in Glycine argyrea by paired fruit analysis

Glycine argyrea (Fabaceae), a perennial wild relative of soybean, has a dual flowering strategy of both self-fertilized cleistogamous flowers and chasmogamous flowers on the same plant. Using allozyme polymorphisms the frequency and pattern of outcrossing was determined. The genotypes of seeds from each of several fruit (legumes) per plant were analysed by starch gel electrophoresis, and the maternal genotype inferred. The maximum likelihood estimates of outcrossing rate for the chasmogamous flowers averaged 0.38. The observed level of heterozygosity in the adult population (h = 0.25) compared with the level expected under random mating (h = 0.32) indicated that partial outcrossing was typical of this population. To analyse the mating pattern further, the pollen genotypes of several seeds per legume for pairs of legumes from the same plant were determined. About 35% of fruit from chasmogamous flowers had no genetically detectable outcross progeny, presumably because they were not cross-pollinated effectively. The majority of the remaining fruit had seed which were of mixed origin (both self-fertilized and outcrossed), suggesting that insect pollination does not preclude self-fertilization. As might be expected from this entomophilous pollination system, the pollen of outcrossed seed within one fruit usually (85% of cross-pollinated legumes) came from one male source. Evidence of pollen carry-over was found in the other legumes. The joint distribution of male sources among the pairs of legumes (paired fruit analysis) showed that non-self sources were shared in 30% of pairs. The probability that two outcrossed seeds from the same fruit would be half-sibs was estimated as 0.15, and for two seeds from different fruit on the same plant as 0.42. There is a hierarchy of genetic identity within and among legumes on the same plant, and on different plants, providing scope for the operation of selection at different levels.     

Depressed Pollination of Lapageria rosea Ruiz et Pav. (Philesiaceae) in the Fragmented Temperate Rainforest of Southern South America

We studied the pollination and reproductive success in continuous and fragmented populations of Lapageria rosea, a self-compatible plant endemic to temperate forests of Chile. Pollinator abundance, visitation rates, flower abundance, nectar volume and concentration, pollen germination and fruit and seed production, were compared between continuous forest of 145 ha and four forest fragments of 6, 3, 3, and 1 ha respectively, surrounded by mature pine plantations of Pinus radiata. Flower abundance was lower in three out of four forest fragments relative to continuous forest. Nectar volume and sugar concentration did not differ between flowers in the two habitats. Pollinators of L. rosea, the hummingbird Sephanoides sephaniodes and bumblebee Bombus dahlbomii were less abundant and visited flowers of L. rosea at lower rates in fragments than in continuous forest. In addition, in vitro rates of pollen germination were lower for flowers in forest fragments. The number of seeds per fruit was also lower in forest fragments. We suggest that fragmentation affects the reproductive success of L. rosea, lowering the total numbers of seeds produced and possibly compromising long term persistence of fragmented populations.     

Evidence for selective fruit abortion in Banksia spinulosa (Proteaceae)

Fruit set is consistently low in the genus Banksia (Proteaceae). One explanation for low fruit:flower ratios is that excess flowers allow plants to selectively abort inferior progeny thereby increasing the average quality of the seed crop. We examined whether self-pollinated fruits were aborted selectively in a partially self-compatible population of Banksia spinulosa. Inflorescences were divided in half and pollinated with cross pollen on both sides, self pollen on both sides or cross pollen on one side and self pollen on the other. Seed production was reduced significantly by 38% on fully self-pollinated compared to fully cross-pollinated inflorescences, indicating partial self-incompatibility or inbreeding depression. On inflorescences pollinated with both cross and self pollen, selfed seed set was reduced by 63% compared to crossed seed set. On the selfed half of these inflorescences, fewer fruits and seeds were produced, and more fruits had aborted seeds than on randomly selected halves of fully self-pollinated inflorescences. These results suggest that self-fertilized embryos were developed in the absence of outcrossed progeny but were selectively aborted when outcrossed progeny were present on inflorescences. To determine the amount of outcrossing occurring in the study population, outcrossing rates (t) of naturally pollinated plants were examined by electrophoretic assessment of progeny arrays. Values of I did not differ significantly from one, indicating complete outcrossing. High outcrossing, despite the high probability of self pollen receipt under natural conditions, is consistent with the conclusion that selective abortion occurs in B. spinulosa.     

Pollen dispersal of tropical trees (Dinizia excelsa: Fabaceae) by native insects and African honeybees in pristine and fragmented Amazonian rainforest

Tropical rainforest trees typically occur in low population densities and rely on animals for cross-pollination. It is of conservation interest therefore to understand how rainforest fragmentation may alter the pollination and breeding structure of remnant trees. Previous studies of the Amazonian tree Dinizia excelsa (Fabaceae) found African honeybees (Apis mellifera scutellata) as the predominant pollinators of trees in highly disturbed habitats, transporting pollen up to 3.2 km between pasture trees. Here, using microsatellite genotypes of seed arrays, we compare outcrossing rates and pollen dispersal distances of (i) remnant D. excelsa in three large ranches, and (ii) a population in undisturbed forest in which African honeybees were absent. Self-fertilization was more frequent in the disturbed habitats (14%, n = 277 seeds from 12 mothers) than in undisturbed forest (10%, n = 295 seeds from 13 mothers). Pollen dispersal was extensive in all three ranches compared to undisturbed forest, however. Using a twogener analysis, we estimated a mean pollen dispersal distance of 1509 m in Colosso ranch, assuming an exponential dispersal function, and 212 m in undisturbed forest. The low effective density of D. excelsa in undisturbed forest (approximately 0.1 trees/ha) indicates that large areas of rainforest must be preserved to maintain minimum viable populations. Our results also suggest, however, that in highly disturbed habitats Apis mellifera may expand genetic neighbourhood areas, thereby linking fragmented and continuous forest populations.     

Genetic rescue of remnant tropical trees by an alien pollinator.

Habitat fragmentation is thought to lower the viability of tropical trees by disrupting their mutualisms with native pollinators. However, in this study, Dinizia excelsa (Fabaceae), a canopy-emergent tree, was found to thrive in Amazonian pastures and forest fragments even in the absence of native pollinators. Canopy observations indicated that African honeybees (Apis mellifera scutellata) were the predominant floral visitors in fragmented habitats and replaced native insects in isolated pasture trees. Trees in habitat fragments produced, on average, over three times as many seeds as trees in continuous forest, and microsatellite assays of seed arrays showed that genetic diversity was maintained across habitats. A paternity analysis further revealed gene flow over as much as 3.2 km of pasture, the most distant pollination precisely recorded for any plant species. Usually considered only as dangerous exotics, African honeybees have become important pollinators in degraded tropical forests, and may alter the genetic structure of remnant populations through frequent long-distance gene flow.     

Effects of converting natural forests to coniferous plantations on fruit and seed production and mating patterns in wild cherry trees

Landscape disturbances can affect reproductive performance of animal-pollinated trees. We verified the effects of the loss and fragmentation of natural forests caused by the creation of coniferous plantations on fruit and seed production as well as mating patterns of animal-pollinated trees. We investigated 146 and 134 individual flowering trees of Prunus verecunda (Koidz.) Koehne in 2012 and 2013, respectively, at 12 sites. These sites were at least 1.2 km apart from each other in an 8 × 15-km forestry region, and the composition and configuration of natural forests varied in the study area. The mean outcrossing rate was >0.99 across the sites. Among trees within the sites, the number of fruits per inflorescence was positively correlated with the basal stem area and leaf chlorophyll density of the trees. Among the sites, the mean number of fruits per inflorescence was positively correlated with the site elevation, and the correlated paternity was positively correlated with the mean distance between trees. The sound-seed rate was positively correlated with the natural-forest area among the sites. These results suggest that environments and resources of trees influence their fruit production, that a loss of natural forests increases in embryo mortality, and that a reduction in tree density decreases pollen donor diversity in P. verecunda. Thus, a landscape disturbance may decrease seed production, whereas outbreeding is maintained, and fruit production is not likely to be dependent on the landscape disturbance in this species.     

FLOWER,FRUIT AND SEED ABORTION IN TROPICAL FOREST TREES: IMPLICATIONS FOR THE EVOLUTION OF PATERNAL AND MATERNAL REPRODUCTIVE PATTERNS

Flower, fruit and immature seed abortion was studied in seven self-incompatible species of trees in a tropical lowland semideciduous forest. The species showed considerable variability in fruit and seed set and the rate at which flowers and fruits were aborted. The amount of flower and fruit abortion also varied over time within species. Small samples of open-pollinated flowers in three species showed adequate amounts of pollen on the stigma, but it could not be determined whether the pollen was compatible or incompatible. In a species with multiseeded fruits, the aborted fruits contained significantly fewer seeds than those retained on the plant. Position of fruit within the inflorescence and of seed within the ovary also had a marked effect on abortion: fruits and seeds at certain positions had a higher probability of abortion than those at other positions. Experiments to test the effect of pollen source on abortion were inconclusive. The factors underlying abortions were evaluated in the context of three mutually non-exclusive hypotheses. It is concluded that selection for increased pollen dispersal and uncertainty in paternity of the zygotes are major factors underlying abortions.     

Real-time patterns of pollen flow in the wild-service tree, Sorbus torminalis (Rosaceae). III. Mating patterns and the ecological maternal neighborhood

Understanding the role of mother plants as pollen recipients in shaping mating patterns is essential for understanding the evolution of populations and in particular to predict the consequence of habitat fragmentation. Here, we investigated variation in mating patterns due to maternal phenotypic traits, phenological variance, and landscape features in Sorbus torminalis, a hermaphroditic, insect-pollinated and low-density, European temperate forest tree. The diversity and composition of pollen clouds received by maternal trees in S. torminalis were mainly determined by their conspecific neighborhood: isolated individuals sample more diversity through more even paternal contributions, low relatedness among paternal genes, and high rates of long-distance pollen dispersal within their progenies. Maternal phenotypic traits related to pollinator attractiveness also had an effect, but only when competition was strong: in this case, larger mother trees with more flowers sampled more diversity. The floral architecture of S. torminalis, with multiple-seeded fruit, strongly shaped mating patterns, with higher levels of correlated paternity among seeds belonging to the same fruit (30% full sibs) than among seeds belonging to different fruits (14% full sibs). Finally, flowering phenology affected the distribution of diversity among maternal pollen clouds, but the earliest and latest mother trees did not receive less diversity of pollen than the others.