Resource overlap and possible competition between honey bees and wild bees in central Europe

Evidence for interspecific competition between honey bees and wild bees was studied on 15 calcareous grasslands with respect to: (1) foraging radius of honey bees, (2) overlap in resource use, and (3) possible honey bee effects on species richness and abundance of flower-visiting, ground-nesting and trap-nesting wild bees. The grasslands greatly differed in the number of honey bee colonies within a radius of 2 km and were surrounded by agricultural habitats. The number of flower-visiting honey bees on both potted mustard plants and small grassland patches declined with increasing distance from the nearest apiary and was almost zero at a distance of 1.5–2.0 km. Wild bees were observed visiting 57 plant species, whereas honey bees visited only 24 plant species. Percentage resource overlap between honey bees and wild bees was 45.5%, and Hurlbert’s index of niche overlap was 3.1. In total, 1849 wild bees from 98 species were recorded on the calcareous grasslands. Neither species richness nor abundance of wild bees were negatively correlated with the density of honey bee colonies (within a radius of 2 km) or the density of flower-visiting honey bees per site. Abundance of flower- visiting wild bees was correlated only with the percentage cover of flowering plants. In 240 trap nests, 1292 bee nests with 6066 brood cells were found. Neither the number of bee species nor the number of brood cells per grassland was significantly correlated with the density of honey bees. Significant correlations were found only between the number of brood cells and the percentage cover of shrubs. The number of nest entrances of ground-nesting bees per square metre was not correlated with the density of honey bees but was negatively correlated with the cover of vegetation. Interspecific competition by honey bees for food resources was not shown to be a significant factor determining abundance and species richness of wild bees. Received: 22 March 1999 / Accepted: 24 September 1999     

Pollen limitation and demographic structure in small fragmented populations of Brunsvigia radulosa (Amaryllidaceae)

The ecological consequences of disruptions in plant-pollinator mutualisms are poorly understood. We examined how seed production and recruitment of juveniles in populations of the spectacular grassland geophyte Brunsvigia radulosa (Amaryllidaceae) correlate with various indices of habitat fragmentation, including habitat fragment area, population size and population isolation. The species was found to be self-incompatible and adapted for pollination primarily by the long-proboscid fly Philoliche aethiopica (Tabanidae). In places where this fly is locally extinct, carpenter bees appear to act as substitute, though less effective, pollinators. Seed production in B. radulosa showed a significant positive relationship with population size, but not with habitat fragment area or spatial isolation of populations when all three indices of habitat fragmentation were included as predictor variables in multiple regression models. Reduced seed production in small populations was attributable to pollen limitation, as supplemental hand pollinations resulted in proportionally greater increases in seed production in these populations. Pollen limitation appears to have demographic consequences; specifically, the proportion of juvenile plants in populations showed significant positive relationships with current levels of seed production per plant and size of populations. Thus the long term persistence of small B. radulosa populations in habitat fragments may be threatened by a pollination deficit.     

Reproduction of <Emphasis Type="Italic">Amorpha canescens</Emphasis> (Fabaceae) and diversity of its bee community in a fragmented landscape

Loss of insect pollinators due to habitat fragmentation often results in negative effects on plant reproduction, but few studies have simultaneously examined variation in the bee community, site characteristics and plant community characteristics to evaluate their relative effects on plant reproduction in a fragmented habitat. We examined the reproduction of a common tallgrass prairie forb, Amorpha canescens (Fabaceae), in large (>40 ha) and small (<2 ha) prairie remnants in Iowa and Minnesota in relation to the diversity and abundance of its bee visitors, plant population size, and species density of the forb flowering community. We found significant positive effects of the diversity of bees visiting A. canescens on percent fruit set at a site in both years of the study and in 2002 an additional significant positive effect of plant species density. Abundance of bees visiting A. canescens had a significant positive effect on percent fruit set in 2002, but was only marginally significant in 2003. In 2003 but not 2002, the plant species density at the sites had a significant negative effect on the diversity and abundance of bees visiting A. canescens, indicating community-level characteristics can influence the bee community visiting any one species. Site size, a common predictor of plant reproduction in fragmented habitats did not contribute to any models of fruit set and was only marginally related to bee diversity one year. Andrena quintilis, one of the three oligolectic bee species associated with A. canescens, was abundant at all sites, suggesting it has not been significantly affected by fragmentation. Our results show that the diversity of bees visiting A. canescens is important for maintaining fruit set and that bee visitation is still sufficient for at least some fruit set in all populations, suggesting these small remnants act as floral resource oases for bees in landscapes often dominated by agriculture.     

Effects of black mustard population size on the taxonomic composition of pollinators

In plant species that are pollinated by a variety of animal species, spatial and temporal variability in the taxonomic composition of those pollinators may cause spatial and temporal variability in selection on floral traits. While temporal variation in pollinator composition has been widely reported, spatial variability, particularly on a local scale, has been studied less frequently; nevertheless, available evidence suggests that local spatial variability may be a widespread feature of plant-pollinator interactions. In addition, the causes of this spatial variability are poorly known. This study was undertaken to quantify variability in the taxonomic composition of pollinators visiting local populations of black mustard, Brassica nigra, and to determine some of the causes of this variability. Simultaneous observations were conducted in seven pairs of black mustard populations. The members of each pair were in close spatial proximity but differed in number of plants. Larger plant populations were visited by significantly greater numbers of honey bees and significantly fewer small bees than small populations on a per-plant basis. There was also a trend toward greater syrphid fly visitation in small populations. The increased numbers of honey bees at large plant populations is probably due to their ability to recruit long distances to the most rewarding plant populations. The lower number of small bees at large plant populations may be due to competition from honey bees and/or spreading a constant number of small bees over the larger number of plants in large populations.This paper is dedicated to the memory of George Eickwort     

High levels of fecundity in small and isolated populations of a self‐compatible Aloe pollinated by opportunistic birds and bees

• Plant species that are effective colonisers of transient habitats are expected to have a capacity for uniparental reproduction and show flexibility in pollination systems. Such traits may enable populations to be established from a small number of founding individuals without these populations succumbing to reductions in fecundity arising from pollinator limitation.
• We tested these predictions for Aloe thraskii (Xanthorrhoeaceae), a succulent treelet that colonises shifting coastal dunes and has both bird and bee pollinators. We performed hand‐pollination experiments, and selectively excluded bird visitors to determine differences in pollinator effectiveness. We measured pollinator visitation rates and fecundity in populations varying in their size, density and isolation distance.
• Controlled hand‐pollinations revealed that unlike most other Aloe species, A. thraskii is self‐compatible and thus capable of uniparental reproduction. The species does however depend on pollinators and is visited by various bird species as well as by bees. Fruit and seed set are not affected by selective exclusion of birds, thus indicating that bees are effective pollinators. Bird visitation rates increased with increasing plant height and population size, while bee visitation rates increased with increasing population size and density. We found that seed set per flower was lower in large populations than in small populations.
• These results suggest that establishment of populations of A. thraskii from a small number of individuals is unlikely to be limited by the fecundity of individual plants.

     

Plant recruitment bottlenecks in temperate forest fragments: seed limitation and insect herbivory

Numerous studies have documented declines in plant diversity in response to habitat loss in fragmented landscapes. However, determining the mechanisms that lead to species loss is challenging using solely a correlative approach. Here we link correlative assessments of plant community composition with seed additions for a focal species to test the hypothesis that distributions of forests plants within a fragmented landscape are limited by seed dispersal. Woody plant species richness of fragments declined as fragments (n=26) became more isolated by agricultural fields. We predicted that if these isolation effects were driven by poor dispersal rather than other effects associated with habitat loss, then plants should vary in their response to isolation in relation to their seed size (i.e., stronger effects for plants with larger seeds). As predicted under this dispersal limitation hypothesis, sensitivity of bird-dispersed shrubs to isolation was related to their seed mass, with species with heavy seeds (e.g., Lindera benzoin) exhibiting stronger declines in presence across isolation gradients than species with light seeds. Seed addition experiments were performed for Lindera benzoin in two high isolation forest fragments (nearest neighbor mean distance=803 m) where Lindera was naturally absent, and two low isolation fragments (nearest neighbor mean distance=218 m) with naturally occurring Lindera populations. Seed addition and control plots (n=50 1 m² plots per fragment) were monitored for 13 censuses over 3 years. Across all four fragments, seed additions resulted in significant increases in Lindera seedling recruitment with no differences in final seedling establishment among fragments. However, insect herbivory was higher on Lindera seedlings in high isolation compared to low isolation fragments and was negatively correlated with seedling survival over some years. Consistent with prior work, our results confirm that seed dispersal plays a significant role in affecting plant diversity in fragmented landscapes. However, results also suggest the need for a better understanding of how additional processes, such as herbivory, may be altered as habitat is lost and what effects such changes have for forest plants.     

Butterfly seed predation: effects of landscape characteristics,plant ploidy level and population structure

Polyploidization has been suggested as one of the most common mechanisms for plant diversification. It is often associated with changes in several morphological, phenological and ecological plant traits, and therefore has the potential to alter insect–plant interactions. Nevertheless, studies evaluating the effect of plant polyploidy on interspecific interactions are still few. We investigated pre-dispersal seed predation by the butterfly Anthocharis cardamines in 195 populations of two ploidy levels of the herb Cardamine pratensis (tetraploid ssp. pratensis, 2n = 30 vs. octoploid ssp. paludosa, 2n = 56–64). We asked if differences in incidence and intensity of predation among populations were related to landscape characteristics, plant ploidy level and population structure. The incidence of the seed predator increased with increasing plant population size and decreasing distance to nearest population occupied by A. cardamines. The intensity of predation decreased with increasing plant population size and was not affected by isolation. Probability of attack decreased with increasing shading, and intensity of predation was higher in grazed than in non-grazed habitats. The attack intensity increased with increasing mean flower number of plant population, but was not affected by flowering phenology. Individuals in tetraploid populations suffered on average from higher levels of seed predation, had higher mean flower number, were less shaded and occurred more often in grazed habitats than octoploid populations. When accounting for differences in habitat preferences between ploidy levels there was no longer a difference in intensity of predation, suggesting that the observed differences in attack rates among populations of the two ploidy levels are mediated by the habitat. Overall, our results suggest that polyploidization is associated with differentiation in habitat preferences and phenotypic traits leading to differences in interspecific interaction among plant populations. This, in turn, may facilitate further divergence of ploidy levels.     

Clumped planting arrangements improve seed production in a revegetated eucalypt woodland

The arrangement of plants within revegetated sites is rarely considered an important characteristic of these communities. However, in natural systems, plant spatial arrangements can influence a range of ecological processes, including pollination and seed set. Pollinators tend to preferentially visit larger and/or more closely spaced populations, with plants in these populations generally receiving more outcrossed pollen, resulting in increased seed set and better quality seed. Similar trends may occur in revegetated populations, but little is known about the influence of planting arrangement on seed production in restored systems. Here, we quantified the effect of plant abundance (number of conspecifics within 100 m) and distance to nearest reproductive conspecific on the level of seed set for six eucalypt species (n = 422 trees in total) in 1 year and for one of these species (Eucalyptus leucoxylon), across three additional years. Seed number per fruit was highly variable both between individuals and within individuals across years. Despite this variability, there was a consistent trend of higher seed production (seed number per fruit) when another reproductive conspecific was within 20 m. In contrast, plant abundance had little influence on seed production. Further investigation of nearest neighbor arrangements found the distance to either the first, second, third, or fourth reproductive neighbors were the key predictors of seed production. Therefore, revegetation designs that consider plant spacing and aggregation, rather than only planting to overall density criteria (i.e. trees/ha), at least for the eucalypts studied here, has the potential to improve seed production in revegetated populations.     

Effects of pollination and pollen source on the seed set ofPedicularis palustris

Habitat alteration can deteriorate plant-pollinator interactions and thereby increase the risk of population extinction. As part of a larger study on the effects of changes in land use on fen grassland vegetation, factors influencing the seed set of a short-lived, endangered wetland plant,Pedicularis palustris, were studied. We conducted field pollination experiments in one large and one small population. To investigate the effect of pollen source on seed set, individual flowers of caged plants were left unpollinated or were pollinated with pollen from the same flower, the same population or another population. To study pollen limitation and flower display, whole plants were subjected to pollinator exclosure, hand pollination or natural pollination. Self-compatibility was high, but differed between populations (61% and 97% of seed set after cross-pollination within populations). Cross-pollination between populations did not significantly alter seed number per capsule. Pollinator exclosure resulted in a very low seed set (<15% of natural seed set), despite high self-compatibility. The most likely explanations for high self-compatibility in combination with low autofertility are geitonogamy as reproductive assurance, selective neutrality of self-compatibility and phylogenetic constraints. Because of low autofertility, the seed set inP. palustris depends on pollinators. In the study populations, natural pollination was clearly sufficient for maximum seed production per plant, but seed set per capsule was significantly pollen-limited in the smaller population. Plants in this population also had a higher maximum percentage of simultaneously open flowers than those of the large population (31% vs. 13%), while flower longevity was generally extended without pollination. It is concluded thatP. palustris may influence pollinator behaviour and therefore the risk of pollen limitation by flower display.P. palustris showed a flexible reaction to differing pollination regimes without losses in overall seed set in the study populations.     

Mobility and resource use influence the occurrence of pollinating insects in restored seminatural grassland fragments

After habitat restoration, species need to recolonize from existing populations. The ability of species to recolonize restored habitats likely depends on their traits. This study aimed to test if species traits and isolation from source habitat can explain the presence of insects in restored grasslands. We surveyed the occurrence of hoverflies and bees in 14 restored seminatural pastures as well as in intact seminatural grasslands in the surrounding landscape. We tested how connectivity, time since restoration, and species traits influence if species that are present in the surrounding landscape also occur in restored pastures. Solitary bee species present in the landscape were less likely to occur in restored pastures compared to bumblebees and hoverflies. The occurrence of bumblebees, but not solitary bees or hoverflies, decreased with time since restoration. The occurrence of solitary bees increased but the occurrence of hoverflies decreased with high connectivity. Migratory hoverflies were more likely to occur in restored pastures than nonmigratory hoverflies, especially in pastures with low connectivity. Among both bumblebees and solitary bees, the occurrence was influenced by nesting traits, with the lowest occurrence of parasitic species and of species digging nests in the ground. The subset of the landscape's species pool that occurs in restored pastures has a contrasting set of traits compared with species in intact source habitats. Both mobility and resource use act as filters that influence the assembly of pollinator communities after restoration. A full recovery of pollinator communities is more likely if source populations are available nearby.     

Fruit set of highland coffee increases with the diversity of pollinating bees

The worldwide decline of pollinators may negatively affect the fruit set of wild and cultivated plants. Here, we show that fruit set of the self-fertilizing highland coffee (Coffea arabica) is highly variable and related to bee pollination. In a comparison of 24 agroforestry systems in Indonesia, the fruit set of coffee could be predicted by the number of flower-visiting bee species, and it ranged from ca. 60% (three species) to 90% (20 species). Diversity, not abundance, explained variation in fruit set, so the collective role of a species-rich bee community was important for pollination success. Additional experiments showed that single flower visits from rare solitary species led to higher fruit set than with abundant social species. Pollinator diversity was affected by two habitat parameters indicating guild-specific nesting requirements: the diversity of social bees decreased with forest distance, whereas the diversity of solitary bees increased with light intensity of the agroforestry systems. These results give empirical evidence for a positive relationship between ecosystem functions such as pollination and biodiversity. Conservation of rainforest adjacent to adequately managed agroforestry systems could improve the yields of farmers.     

The effects of island area, isolation and source population size on the presence of the grayling butterfly Hipparchia semele (L.) (Lepidoptera: Satyrinae) on British and Irish offshore islands

Records of Hipparchia semele on British and Irish islands have been modelled against island area, isolation (sea and land distance) and the size of the nearest potential source populations. All three variables have been found to contribute significantly to the presence or absence of H. semele on the islands. Isolation is a more significant predictor than island area. This result differs from the multiple species case where area was found to be a more important influence than isolation. Records on islands are also shown to depend on the size of populations at the nearest sources; this underpins the relationships identified for the multiple species case, first, between the number of species on islands and at nearest sources and, second, between the incidence of species on islands and at nearest sources. There are clear indications that smaller islands may become increasingly marginalized for H.semele; with ongoing habitat loss, because isolation increases and source populations become sparser, the probability of H. semele recolonizing islands also decreases.     

Structure of a plant-pollinator network on a pahoehoe lava desert of the Galápagos Islands

Plant-pollinator interactions are important for the evolution and survival of the species involved. Plant-pollinator networks on oceanic islands are often small in size and as a consequence the connectance is high suggesting a substantial generalisation level. Further, linkage level for insular plants is shown to be lower than on mainland. The present study investigates a plant-pollinator network on the Galápagos Islands that is unique because of its very small size. We recorded pollinator visits to plant species as well as pollen grains on insect bodies. The combination of these data increased the observed number of interactions. The values for connectance and linkage level for plants were found to be consistent with similar values found in other network studies. There were no relation between the abundance of plant species and the number of pollinating species. The dominating pollinator species was the Galápagos carpenter bee Xylocopa darwini . Specimens of the shorthorned grasshopper Halmenus cuspidatus turned out to carry pollen from five plant species out of twelve and are probably functioning as pollinator. Bagging experiments revealed dependency on insect visits for a high seed set for most of the plant species, but only one species Plumbago scandens seemed to possess a pollen limited seed set. The network showed an asymmetric pattern of number of interactions per species with a few species having several interactions and many species a few. This pattern is supposed to result in a rather robust community, but is also fairly sensitive if the dominant species are threatened. The high connectance value found could, however, counteract this vulnerability.     

Evolutionary consequences of habitat fragmentation: population size and density affect selection on inflorescence size in a perennial herb

Habitat fragmentation is considered to be one of the major threats to biological diversity worldwide. To date, however, its consequences have mainly been studied in an ecological context, while little is known about its effects on evolutionary processes. In this study we examined whether habitat fragmentation affects selection on plant phenotypic traits via changes in plant-pollinator interactions, using the self-incompatible perennial herb Phyteuma spicatum. Specifically, we hypothesized that limited pollination service in small or low-density populations leads to increased selection for traits that attract pollinators. We recorded mean seed production per capsule and per plant as a measure of pollination intensity and assessed selection gradients (i.e., trait-fitness relationships) in 16 natural populations of varying size and density over 2 years. Mean seed production was not related to population size or density, except for a marginal significant effect of density on the mean number of seeds per capsule in 1 year. Linear selection for flowering time and synchrony was consistent across populations; relative fitness was higher in earlier flowering plants and in plants flowering synchronously with others. Selection on inflorescence size, however, varied among populations, and linear selection gradients for inflorescence size were negatively related to plant population size and density in 1 year. Selection for increased inflorescence size decreased with increasing population size and density. Contrary to our expectation this appeared not to be related to changes in pollination intensity (mean seed production was not related to population size or density in this year), but was rather likely linked to differences in some other component of the abiotic or biotic environment. In summary, our results show that habitat fragmentation may influence selection on plant phenotypic traits, thereby highlighting potential evolutionary consequences of human-induced environmental change.     

Contrasting responses of bee communities to coffee flowering at different spatial scales

While investigating biodiversity patterns on different spatial scales, ecological processes determining these patterns have been rarely analysed. Flower visitation by bees is an important ecological process that is related to floral resource availability. However, little is known about whether responses of bee communities to floral resource availability change at different spatial scales. We studied density and species richness of flower-visiting bees in relation to floral resource availability, provided by coffee, in traditional agroforestry systems on a field, shrub, and branch scale. On a field scale, mean bee density per shrub increased with decreasing proportion of flowering coffee shrubs per site, showing a dilution effect. Conversely, on shrub and branch scales bee density per shrub, or shrub part, increased with increasing number of inflorescences, showing a concentration effect. Additionally, bee density per shrub was higher on those that were only partly, rather than totally surrounded by other flowering coffee shrubs. Species richness of flower-visiting bees was positively affected by high resource availability on a shrub and a branch scale, expressed by a high number of inflorescences, but at the field scale the proportion of flowering shrubs per site did not have any effect on species richness. Our results show contrasting responses of the community of flower-visiting bees to floral resource availability, depending on the spatial scale considered. We conclude that patterns of flower-visiting bee communities of only one spatial scale can not be generalized, since the number of pollinators may be limited on a field scale, but not on smaller scales.     

Ploidy level interacts with population size and habitat conditions to determine the degree of herbivory damage in plant populations

Recently it has been suggested that ploidy level of a plant population may have important effects on plant‐animal interactions. Plant‐animal interactions can also be strongly altered by factors such as plant population size and habitat conditions. It is, however, not known how these factors interact to shape the overall pattern of plant‐animal interactions. I studied the interaction between a perennial plant, Aster amellus, and a monophagous herbivorous moth, Coleophora obscenella, and investigated the effect of ploidy level of the plant population, plant population size, isolation and habitat conditions on density of the insect, damage by the insect, and plant performance. Ploidy level, plant population size and habitat conditions, but not isolation, strongly influence plant‐herbivore interactions. Furthermore, there are significant interactions between effects of ploidy level and plant population size and between ploidy level and isolation. Hexaploid plants suffer higher seed damage by the herbivore, but their seed production is still higher than that of diploids. Herbivores thus partly limit the evolutionary success of the hexaploid plants. Plant‐animal interactions are also strongly determined by plant population size. Small populations of A. amellus (below forty flowering ramets) host no C. obscenella larvae, indicating a minimum A. amellus population size that can sustain a viable C. obscenella population. Negative and positive effects of plant population size balance and result in no relationship between plant population size and number of developed seeds per flower head. The results also show a significant interaction between ploidy level and plant population size, indicating that the increase in density of C. obscenella larvae with plant population size is greater in hexaploid than in diploid populations. The results also indicate that the effect of ploidy level on plant‐herbivore interactions can be altered by plant population size, which suggests that plant‐herbivore interactions are driven by a complex of interactions among different factors. Studying each factor separately could thus lead to biased conclusions about patterns of interactions in such systems.     

The Abundance and Pollen Foraging Behaviour of Bumble Bees in Relation to Population Size of Whortleberry (Vaccinium uliginosum)

Habitat fragmentation can have severe effects on plant pollinator interactions, for example changing the foraging behaviour of pollinators. To date, the impact of plant population size on pollen collection by pollinators has not yet been investigated. From 2008 to 2010, we monitored nine bumble bee species (Bombus campestris, Bombus hortorum s.l., Bombus hypnorum, Bombus lapidarius, Bombus pascuorum, Bombus pratorum, Bombus soroensis, Bombus terrestris s.l., Bombus vestalis s.l.) on Vaccinium uliginosum (Ericaceae) in up to nine populations in Belgium ranging in size from 80 m² to over 3.1 ha. Bumble bee abundance declined with decreasing plant population size, and especially the proportion of individuals of large bumble bee species diminished in smaller populations. The most remarkable and novel observation was that bumble bees seemed to switch foraging behaviour according to population size: while they collected both pollen and nectar in large populations, they largely neglected pollen collection in small populations. This pattern was due to large bumble bee species, which seem thus to be more likely to suffer from pollen shortages in smaller habitat fragments. Comparing pollen loads of bumble bees we found that fidelity to V. uliginosum pollen did not depend on plant population size but rather on the extent shrub cover and/or openness of the site. Bumble bees collected pollen only from three plant species (V. uliginosum, Sorbus aucuparia and Cytisus scoparius). We also did not discover any pollination limitation of V. uliginosum in small populations. We conclude that habitat fragmentation might not immediately threaten the pollination of V. uliginosum, nevertheless, it provides important nectar and pollen resources for bumble bees and declining populations of this plant could have negative effects for its pollinators. The finding that large bumble bee species abandon pollen collection when plant populations become small is of interest when considering plant and bumble bee conservation.     

Small local population sizes and high habitat patch fidelity in a specialised solitary bee

Andrena hattorfiana is a rare solitary bee which has declined during the last decades throughout western Europe. It is specialised to forage pollen from plants of the family Dipsacaceae. Knowledge of distribution, dispersal propensity, and local population sizes is essential for successful conservation of A. hattorfiana. The investigated local bee populations (n = 78) were dominated by small local populations and 60% were smaller than 10 female individuals and 80% were smaller than 50 female individuals. The area of the median occupied habitat patch was 1.25 hectare and harboured 7 female bees. Mark-release-recapture studies of female A. hattorfiana revealed a sedentary behaviour. Among pollen-foraging female bees the average registered distance moved was 46 m. The patch emigration rate was about 2%, with an observed maximum colonization distance of 900 m. Only 10% of the individuals crossed areas without the pollen plant within grassland patches, such as unpaved roads, stone walls and small tree-stands, even if these areas were less than 10 m wide. This study shows that solitary bees can occur in local populations of extremely small size and they have a sedentary behaviour. These are features that usually increase the risk of local population extinction.     

Distinct Phylogeographic Structures of Wild Radish (Raphanus sativus L. var. raphanistroides Makino) in Japan

Coastal plants with simple linear distribution ranges along coastlines provide a suitable system for improving our understanding of patterns of intra-specific distributional history and genetic variation. Due to the combination of high seed longevity and high dispersibility of seeds via seawater, we hypothesized that wild radish would poorly represent phylogeographic structure at the local scale. On the other hand, we also hypothesized that wild radish populations might be geographically differentiated, as has been exhibited by their considerable phenotypic variations along the islands of Japan. We conducted nuclear DNA microsatellite loci and chloroplast DNA haplotype analyses for 486 samples and 144 samples, respectively, from 18 populations to investigate the phylogeographic structure of wild radish in Japan. Cluster analysis supported the existence of differential genetic structures between the Ryukyu Islands and mainland Japan populations. A significant strong pattern of isolation by distance and significant evidence of a recent bottleneck were detected. The chloroplast marker analysis resulted in the generation of eight haplotypes, of which two haplotypes (A and B) were broadly distributed in most wild radish populations. High levels of variation in microsatellite loci were identified, whereas cpDNA displayed low levels of genetic diversity within populations. Our results indicate that the Kuroshio Current would have contributed to the sculpting of the phylogeographic structure by shaping genetic gaps between isolated populations. In addition, the Tokara Strait would have created a geographic barrier between the Ryukyu Islands and mainland Japan. Finally, extant habitat disturbances (coastal erosion), migration patterns (linear expansion), and geographic characteristics (small islands and sea currents) have influenced the expansion and historical population dynamics of wild radish. Our study is the first to record the robust phylogeographic structure in wild radish between the Ryukyu Islands and mainland Japan, and might provide new insight into the genetic differentiation of coastal plants across islands.     

INTERPOPULATIONAL VARIATION IN FRUIT PRODUCTION: THE ROLE OF POLLINATION-LIMITATION IN THE OLYMPIC MOUNTAINS

Fruit set varied from a mean of 44% to 93% among six populations of Veronica cusickii L. (Scrophulariaceae) spanning a distance of 13 km in the Olympic Mountains, WA. Hand pollinations in three populations showed that this variation resulted primarily from differences in the level of limitation by insect pollination. In every pollination, hand-pollinated flowers averaged at least 82% fruit set. In 1981 and 1982, addition of excess pollen significantly increased mean fruit set per plant in the Blue Mountain population, but not in two other populations with higher natural fruit sets. At Blue Mountain, hand pollination more than doubled the mean seed set of V. cusickii and also significantly increased seed set in two of three other species visited primarily by flies or solitary bees. Pollination-limitation appears to be common at this site where the weather is frequently cold and damp.