Reproductive assurance varies with flower size in Collinsia parviflora (Scrophulariaceae)

A central question in plant evolutionary ecology is how mixed mating systems are maintained in the face of selection against self-pollination. Recently, attention has focused on the potential reproductive assurance (RA) benefit of selfing: the ability to produce seeds via autonomous selfing when the potential for outcrossing is reduced or absent. To date, there is little experimental support for this benefit under natural pollination conditions. In addition, the RA hypothesis has not been tested experimentally in a species displaying morphological variation for traits expected to influence the mating system, such as flower size, which affects both attractiveness to pollinators and ability to self autonomously. Here, we document significant among-population variation in flower size in Collinsia parviflora and show that pollinators preferred large flowers over small flowers in experimental arrays. The pollinator community varied among three study sites, and two small-flowered populations had lower pollinator visitation rates than one large-flowered population. We compared seed production between intact flowers (can self) and experimentally emasculated flowers (require a pollinator) on large- and small-flowered plants. As predicted by the RA hypothesis, small-flowered plants show a greater RA benefit of selfing than large-flowered plants; emasculated, small flowers produced very few seeds, relative to intact, small flowers or either emasculated or intact, large flowers. We also show that the RA benefit is pollination-context dependent, differing between small- and large-flowered test sites, likely due to a combination of pollinator discrimination against small flowers and differences between test sites in the pollinator community. This paper is the first experimental evidence showing a trait-dependent RA benefit of selfing under natural pollination conditions.     

The reproductive assurance benefit of selfing: importance of flower size and population size

Autonomous selfing can provide reproductive assurance (RA) for flowering plants that are unattractive to pollinators or in environments that are pollen limited. Pollen limitation may result from the breakdown of once-continuous habitat into smaller, more isolated patches (habitat fragmentation) if fragmentation negatively impacts pollinator populations. Here we quantify the levels of pollen limitation and RA among large and small populations of Collinsia parviflora, a wildflower with inter-population variation in flower size. We found that none of the populations were pollen limited, as pollen-supplemented and intact flowers did not differ in seed production. There was a significant effect of flower size on RA; intact flowers (can self) produced significantly more seeds than emasculated flowers (require pollen delivery) in small-flowered plants but not large-flowered plants. Population size nested within flower size did not significantly affect RA, but there was a large difference between our two replicate populations for large-flowered, small populations and small-flowered, large populations that appears related to a more variable pollination environment under these conditions. In fact, levels of RA were strongly negatively correlated with rates of pollinator visitation, whereby infrequent visitation by pollinators yielded high levels of RA via autonomous selfing, but there was no benefit of autonomous selfing when visitation rates were high. These results suggest that autonomous selfing may be adaptive in fragmented habitats or other ecological circumstances that affect pollinator visitation rates.     

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.     

The influence of distinct pollinators on female and male reproductive success in the Rocky Mountain columbine

Although there are many reasons to expect distinct pollinator types to differentially affect a plant's reproductive success, few studies have directly examined this question. Here, we contrast the impact of two kinds of pollinators on reproductive success via male and female functions in the Rocky Mountain columbine, Aquilegia coerulea . We set up pollinator exclusion treatments in each of three patches where Aquilegia plants were visited by either day pollinators (majority bumble bees), by evening pollinators (hawkmoths), or by both (control). Day pollinators collected pollen and groomed, whereas evening pollinators collected nectar but did not groom. Maternal parents, potential fathers and progeny arrays were genotyped at five microsatellite loci. We estimated female outcrossing rate and counted seeds to measure female reproductive success and used paternity analysis to determine male reproductive success. Our results document that bumble bees frequently moved pollen among patches of plants and that, unlike hawkmoths, pollen moved by bumble bees sired more outcrossed seeds when it remained within a patch as opposed to moving between patches. Pollinator type differentially affected the outcrossing rate but not seed set, the number of outcrossed seeds or overall male reproductive success. Multiple visits to a plant and more frequent visits by bumble bees could help to explain the lack of impact of pollinator type on overall reproductive success. The increase in selfing rate with hawkmoths likely resulted from the abundant pollen available in experimental flowers. Our findings highlighted a new type of pollinator interactions that can benefit a plant species.     

Limited phenological and pollinator-mediated isolation among selfing and outcrossing Arabidopsis lyrata populations

Transitions from outcrossing to selfing have been a frequent evolutionary shift in plants and clearly play a role in species divergence. However, many questions remain about the initial mechanistic basis of reproductive isolation during the evolution of selfing. For instance, how important are pre-zygotic pre-pollination mechanisms (e.g. changes in phenology and pollinator visitation) in maintaining reproductive isolation between newly arisen selfing populations and their outcrossing ancestors? To test whether changes in phenology and pollinator visitation isolate selfing populations of Arabidopsis lyrata from outcrossing populations, we conducted a common garden experiment with plants from selfing and outcrossing populations as well as their between-population hybrids. Specifically, we asked whether there was isolation between outcrossing and selfing plants and their between-population hybrids through differences in (1) the timing or intensity of flowering; and/or (2) pollinator visitation. We found that phenology largely overlapped between plants from outcrossing and selfing populations. There were also no differences in pollinator preference related to mating system. Additionally, pollinators preferred to visit flowers on the same plant rather than exploring nearby plants, creating a large opportunity for self-fertilization. Overall, this suggests that pre-zygotic pre-pollination mechanisms do not strongly reproductively isolate plants from selfing and outcrossing populations of Arabidopsis lyrata.     

风筝果镜像花的雌雄异位变化及传粉生态型的形成

亚洲特有植物风筝果(Hiptage benghalensis)镜像花(mirror-image flowers)的雌雄异位变化极大, 柱头与可育花药的空间距离在海南岛与云南西双版纳种群仅有2-4 mm, 而在贵州及云南东南部种群可达10-12 mm。为验证 “镜像花雌雄异位的变化是各个地理种群受到传粉昆虫体型大小选择的结果”假说, 本文开展了传粉生态学与分子生物学研究, 探讨风筝果不同地理种群的雌雄异位变化是否与当地传粉者体型相关, 是否有着遗传基础, 以及是否形成了“传粉生态型”(pollination ecotype)。结果表明, 雌雄异位程度大的贵州种群和云南东南部种群, 其传粉者也是体型较大的木蜂(Xylocopa nasalis)和熊蜂一种(Bombus sp.); 雌雄异位程度小的海南岛种群和云南西双版纳种群的传粉者则是体型较小的大蜜蜂(Apis dorsata)、西方蜜蜂(A. mellifera)。除了雌雄异位的明显分化(F = 53.69, P < 0.01)之外, 花柱长度也在这两类种群之间差异显著(F = 12.5, P < 0.05), 与传粉昆虫身体长度密切相关。通过叶绿体ycf1b、rbcL和细胞核ITS 3个DNA片段的序列分析, 由大型蜂传粉的雌雄异位程度较大的3个种群(贵州和云南东南部种群)遗传关系极近, 与小型蜂传粉种群(雌雄异位较小的海南岛种群和云南西双版纳种群)存在明显的遗传分化。这些结果显示, 风筝果不同地理种群的镜像花雌雄异位程度的变化可能是适应局域不同体型大小的传粉昆虫的选择结果, 而且种群间出现了基因流的隔断, 可能与大型蜂、小型蜂传粉的两种生态型有关。镜像花通过雌雄异位的变化适应不同地域的传粉者而形成传粉生态型, 这种传粉生态型及其传粉隔离机制可能是风筝果属物种形成与维持的一个重要原因。     

Ecological context of the evolution of self-pollination in Clarkia xantiana: population size, plant communities, and reproductive assurance

The repeated evolutionary transition from outcrossing to self-pollination in flowering plants has been suggested to occur because selfing provides reproductive assurance. Reports from biogeographical and ecological surveys indicate that selfing taxa are often associated with stressful and ephemeral environments, situations in which plant abundance is low (e.g., Baker's law) and with novel plant communities, however experimental tests of ecological hypotheses are few. In this study, we examined the ecological context of selection on mating system traits (herkogamy and protandry) in a California annual, Clarkia xantiana, where natural selfing populations differ from outcrossing populations in that they are often of small size or low density and occur mainly outside the range of pollinator-sharing congeners. We constructed artificial populations of plants with broad genetic variation in floral traits and manipulated two ecological factors, plant population size, and the presence versus absence of pollinator-sharing congeners, in the center of the geographic range of outcrossing populations. We found evidence for context-dependent selection on herkogamy and protandry via female fitness in which reduced traits, which promote autonomous selfing, were favored in small populations isolated from congeners whereas selection was comparatively weak in large populations or when congeners were present. In small, isolated populations, the fertility of plants with low herkogamy or protandry was elevated by 66% and 58%, respectively, compared to those with high herkogamy or protandry. The presence of pollinator-sharing congeners augmented bee visitation rates to C. xantiana flowers by 47% for all bees and by 93% for pollen specialists. By facilitating pollinator visitation, congeners mitigated selection on mating system traits in small populations, where outcross mating success is often low (the Allee effect). We also found support for the hypothesis that pollinator availability directly influenced variation in the strength of selection on herkogamy among populations. The striking parallels between our experimental results and patterns of variation in ecological factors across the geographic range of outcrossing and selfing populations suggest that reproductive assurance may play a central role in directing mating system evolution in C. xantiana.     

How variable is delayed selfing in a fluctuating pollinator environment? A comparison between a delayed selfing and a pollinator-dependent Schizanthus species of the high Andes

Delayed selfing has been considered the best-of-both-worlds response to pollinator unpredictability because it can provide reproductive assurance without decreasing outcrossing potential. According to this hypothesis, selfing rates in delayed selfing species should be highly variable in fluctuating pollinator environments. To test this prediction, as well to explore the consequences of delayed selfing on genetic patterns, we compared two sister species that grow in the high Andes of Chile: Schizanthus grahamii that exhibits delayed selfing and Schizanthus hookeri, which is self-compatible but requires pollinators for seed set. We estimated genetic diversity within and among five populations of each species using six shared microsatellites. Our results indicated that selfing rates in S. grahamii (range 0.07–0.81) were significantly more variable than in S. hookeri (range 0–0.26). The highest levels of selfing were found in the populations of S. grahamii located at highest altitudes (r = 0.78) and at northern margin range, where pollinators are probably more scarce. These populations also showed the lowest allelic richness and heterozygosity values. Southern populations of S. grahamii had mixed mating, and showed heterozygosity and diversity values close to those detected for S. hookeri along all the sampled range. Selfing in this species results from geitonogamy, and did not covary with altitude. Schizanthus grahamii showed greater population differentiation than S. hookeri. Overall, our results indicated that selfing rates were widely variable in S. grahamii, with some populations predominantly selfing and others showing mixed mating. This pattern may be associated with the strong fluctuations in pollinator service that typically occur in the high Andes of Chile.     

Variation in composition of two bumble bee species across communities affects nectar robbing but maintains pollinator visitation rate to an alpine plant,Salvia przewalskii

1. In many flowering plants, bumble bees may forage as both pollinators and nectar robbers. This mixed foraging behaviour may be influenced by community context and consequently, potentially affect pollination of the focal plant. 2. Salvia przewalskii is both pollinated and robbed exclusively by bumble bees. In the present study area, it was legitimately visited by two species of bumble bees with different tongue length, Bombus friseanus and Bombus religiosus, but it was only robbed by Bombus friseanus, the shorter‐tongued bumble bee. The intensity of nectar robbing and pollinator visitation rate to the plant were investigated across 26 communities in the Hengduan Mountains in East Himalaya during a 2‐year project. For each of these communities, the floral diversity, and the population size and floral resource of S. przewalskii were quantified. The abundances of the two bumble bee species were also recorded. 3. Both nectar robbing and pollinator visitation rate were influenced by floral diversity. However, pollinator visitation rate was not affected by nectar robbing. The results revealed that relative abundance of the two bumble bee species significantly influenced the incidence of nectar robbing but not the pollinator visitation rate. Increased abundance of B. religiosus, the legitimate visitors, exacerbated nectar robbing, possibly by causing B. friseanus to shift to robbing; however, pollinator visitation remained at a relatively high level. 4. The results may help to explain the persistence of both nectar robbing and pollination, and suggest that, in comparison to pollination, nectar robbing is a more unstable event in a community.     

Composition of the pollinator community,pollination and the mating system for a shrub in fragments of species rich kwongan in south-west Western Australia

In this study we investigate the composition of the potential honeyeater pollinator community, patterns of honeyeater visitation, pollination and the mating system in a range of population fragments for the bird-pollinated mixed mating system shrub Calothamnus quadrifidus R.Br. Specifically, we aimed to answer the following questions. For smaller and more isolated population fragments are honeyeater species lost from the pollinator community, patterns of visitation different, levels of pollination lower and rates of selfing, biparental inbreeding and correlated paternity higher. The composition of the honeyeater community was similar across population fragments and there was no relationship between the abundance of birds and population fragment size. Honeyeaters were most commonly observed visiting numerous inflorescences within single plants in all populations, but as population fragments became larger movements between plants were more commonly observed. Our observations of honeyeater visitation were generally consistent with our measurements of pollination and patterns in the mating system across population fragments. We found no significant relationship between population fragment size and levels of pollination. Mating system studies showed outcrossing rates (t _m) comparable to those found in other bird-pollinated Myrtaceae, and ranged from 0.54 to 0.90 across populations. Outcrossing rates were not significantly correlated with log population size, but correlations of outcrossed paternity indicate a clear trend from low correlated paternity in larger populations to significantly higher correlated paternities in smaller populations. As a consequence mating in small populations will occur between much smaller groups of plants, and this may affect population fitness in subsequent generations.     

Pollination ecology and inbreeding depression control individual flowering phenologies and mixed mating

We analyze evolution of individual flowering phenologies by combining an ecological model of pollinator behavior with a genetic model of inbreeding depression for plant viability. The flowering phenology of a plant genotype determines its expected daily floral display which, together with pollinator behavior, governs the population rate of geitonogamous selfing (fertilization among flowers on the same plant). Pollinators select plant phenologies in two ways: they are more likely to visit plants displaying more flowers per day, and they influence geitonogamous selfing and consequent inbreeding depression via their abundance, foraging behavior, and pollen carry‐over among flowers on a plant. Our model predicts two types of equilibria at stable intermediate selfing rates for a wide range of pollinator behaviors and pollen transfer parameters. Edge equilibria occur at maximal or minimal selfing rates and are constrained by pollinators. Internal equilibria occur between edge equilibria and are determined by a trade‐off between pollinator attraction to large floral displays and avoidance of inbreeding depression due to selfing. We conclude that unavoidable geitonogamous selfing generated by pollinator behavior can contribute to the common occurrence of stable mixed mating in plants.     

Pollination processes and the Allee effect in highly fragmented populations: consequences for the mating system in urban environments

The urban environment was used to study the plant reproductive system in small fragmented populations as well as the potential adaptations of plants to urban conditions. We examined the effect of density on the pollination process and on reproduction in urban populations of the allogamous species Crepis sancta. The habitat is composed of small uncultivated square patches (c. 2 m2) regularly spaced along the pavement in streets of the city of Montpellier, France. Pollinator behaviour (the presence of pollinators, the number of flowers visited and the duration of each visit) and seed set as a function of the number of plants in patches and selfing rates, determined using progeny array analysis, were studied. The propensity for the urban populations to produce seeds by self-fertilization in insect-proof glasshouse was also analysed. We found strong evidence of reduced pollinator activities at low densities, resulting in reduced pollination and a reduction in seed set from 80 to 20% of ovules fertilized (the Allee effect). Progeny array analysis revealed a slight increase (marginally significant) in selfing rates in urban populations compared with large populations. In spite of lower pollinator activity, urban populations did not show a greater ability to self-fertilize compared with rural populations from the nearby countryside.     

Floral dimorphism, pollination, and self-fertilization in gynodioecious GERANIUM RICHARDSONII (Geraniaceae)

The selective maintenance of gynodioecy depends on the relative fitness of the male-sterile (female) and hermaphroditic morphs. Females may compensate for their loss of male fitness by reallocating resources from male function (pollen production and pollinator attraction) to female function (seeds and fruits), thus increasing seed production. Females may also benefit from their inability to self-fertilize if selfing and inbreeding depression reduce seed quality in hermaphrodites. We investigated how differences in floral resource allocation (flower size) between female and hermaphroditic plants affect two measures of female reproductive success, pollinator visitation and pollen receipt, in gynodioecious populations of Geranium richardsonii in Colorado. Using emasculation treatments in natural populations, we further examined whether selfing by autogamy and geitonogamy comprises a significant proportion of pollen receipt by hermaphrodites. Flowers of female plants are significantly smaller than those of hermaphrodites. The reduction in allocation to pollinator-attracting structures (petals) is correlated with a significant reduction in pollinator visitation to female flowers in artificial arrays. The reduction in attractiveness is further manifested in significantly less pollen being deposited on the stigmas of female flowers in natural populations. Autogamy is rare in these protandrous flowers, and geitonogamy accounts for most of the difference in pollen receipt between hermaphrodites and females. Female success at receiving pollen was negatively frequency dependent on the relative frequency of females in populations. Thus, two of the prerequisites for the maintenance of females in gynodioecious populations, differences in resource allocation between floral morphs and high selfing rates in hermaphrodites, occur in G. richardsonii.     

Relationship between reproductive assurance and mixed mating in perennial Kosteletzkya virginica

Recent works have shown that mixed mating systems often evolve despite strong inbreeding depression and reproductive assurance, which is one of the widely accepted explanations for the evolution of selfing. However, there have been few empirical studies on the relationship between mixed mating and reproductive assurance in perennial plants. In the herbaceous perennial, Kosteletzkya virginica, delayed selfing induced from context-dependent style curvature offers reproductive assurance, and adverse weather conditions significantly reduce pollinator visitation rates. In this study, our goals were (i) to experimentally evaluate pollinator failure rate, reproductive assurance, selfing rate and the relationships between them, and (ii) to measure inbreeding depression across multiple growth seasons. Results indicate that both population selfing rates and reproductive assurance are significantly and positively correlated with field estimates of pollinator failure rates, and there is a strong relationship between selfing rates and reproductive assurance. Inbreeding depression across multiple growth seasons ranged from 0.621 to 0.665, and there were no significant differences among different seasons. Our data demonstrates that a mixed mating system is beneficial because frequent pollinator failure has allowed reproductive assurance to evolve through delayed selfing which minimizes the risk of seed discounting and is still advantageous despite high inbreeding depression.     

Self-fertilization and the escape from pollen limitation in variable pollination environments

Seed production in many plants is pollen limited, likely because of unpredictable variation in the pollinator environment. One way for plants to escape the consequences of pollinator variability is to evolve mating systems, such as autonomous selfing, that assure reproduction without relying on pollinators. We explore this hypothesis through the construction and analysis of heuristic models of plant population dynamics in seed- or site-limited populations. Our analysis suggests several important points: the familiar rule that inbreeding depression greater than 0.5 maintains outcrossing significantly underestimates the threshold required under pollen limited conditions with prior selfing; variability in the pollination environment erodes the ability of inbreeding depression to maintain outcrossing; and variable pollination environments can result in stable intermediate rates of prior selfing. The results reflect the importance of geometric mean fitness (which in a variable environment is less than the arithmetic mean) in the face of temporal variation.     

Shifts to earlier selfing in sympatry may reduce costs of pollinator sharing

Coexisting plant congeners often experience strong competition for resources. Competition for pollinators can result in direct fitness costs via reduced seed set or indirect costs via heterospecific pollen transfer (HPT), causing subsequent gamete loss and unfit hybrid offspring production. Autonomous selfing may alleviate these costs, but to preempt HPT, selfing should occur early, before opportunities for HPT occur (i.e., “preemptive selfing hypothesis”). We evaluated conditions for this hypothesis in Collinsia sister species, C. linearis and C. rattanii. In field studies, we found virtually identical flowering times and pollinator sharing between congeners in sympatric populations. Compared to allopatric populations, sympatric C. linearis populations enjoyed higher pollinator visitation rates, whereas visitation to C. rattanii did not differ in sympatry. Importantly, the risk of HPT to each species in sympatry was strongly asymmetrical; interspecies visits comprised 40% of all flower‐to‐flower visits involving C. rattanii compared to just 4% involving C. linearis. Additionally, our greenhouse experiment demonstrated a strong cost of hybridization when C. rattanii was the pollen donor. Together, these results suggest that C. rattanii pays the greatest cost of pollinator sharing. Matching predictions of the preemptive selfing hypothesis, C. rattanii exhibit significantly earlier selfing in sympatric relative to allopatric populations.     

Fitness Traits and Dispersal Ability in the Herb Tragopogon pratensis (Asteraceae): Decoupling the Role of Inbreeding Depression and Maternal Effects

Abstract: Inbreeding depression can decrease several fitness traits and maternal effects can strongly influence the amount of inbreeding depression. Understanding the effects of inbreeding depression on plant fitness is especially important in the context of habitat fragmentation, where plant populations become smaller and more isolated, exhibiting increasing levels of inbreeding depression. We examined the joint influence of inbreeding depression and maternal effects on life cycle traits and dispersal ability in the herb Tragopogon pratensis that grows in fragmented populations in Europe. We conducted experimental crosses to obtain selfed and outcrossed progeny in two contrasted environments. In particular, we produced a first generation of seeds and plants that were self-pollinated again to produce a second generation of seeds. Individual seeds were weighed and their pappuses measured to estimate the dispersal potential. Pollination treatment only had a significant effect on seed mass and dispersal ability. Coefficients of inbreeding depression did not differ between selfed and outcrossed plants. Seed mass had a significant effect on germination date. Environment had a significant effect on mass of the second generation of seeds and the interaction between pollination treatment and family was significant for six traits, indicating the existence of strong maternal effects in T. pratensis. Results suggest population differentiation. Overall, T. pratensis populations exhibited a good performance under selfing, in terms of life cycle traits and dispersal ability, which would allow the species to cope with problems associated with fragmentation.     

Direct and indirect estimates of the selfing rate in small and large individuals of the bumblebee pollinated Cynoglossum officinale L (Boraginaceae)

We measured the relationship between selfing rates and flower number in an experimental population of bumblebee pollinated Cynoglossum officinale , with plants differing in flower number. Results were compared with the prediction of a model based on pollen dynamics and pollinator behaviour. The selfing rate, as measured by multilocus oligonucleotide DNA fingerprinting, increased with flower number and ranged from 0% to 70%. Flowers on large plants received an equal number of visits from bumblebees as flowers on small plants. On large plants more flowers in a row were visited, inducing geitonogamy. The overall relationship between selfing rate and number of flowers can be explained by pollen dynamics and pollinator behaviour without invoking postpollination processes such as differential pollen tube growth and abortion.     

Effects of herbivory and inbreeding on the pollinators and mating system of Mimulus guttatus (Phrymaceae)

Most models of mating system evolution predict mixed mating to be unstable, although it is commonly reported from nature. Ecological interactions with mutualistic pollinators can help account for this discrepancy, but antagonists such as herbivores are also likely to play a role. In addition, inbreeding can alter ecological interactions and directly affect selfing rates, which may also contribute to maintaining mating system variation. We explored herbivore and inbreeding effects on pollinator behavior and selfing rates in Mimulus guttatus. First, individual spittlebug (Philaenus spumarius) herbivores were applied to native plants in two populations. Spittlebugs reduced flower size, increased anther-stigma distance, and increased selfing rates. A second experiment factorially crossed spittlebug treatment with inbreeding history (self- vs. cross-fertilized), using potted plants in arrays. Spittlebugs did not affect pollinator behavior, but they reduced flower size and nearly doubled the selfing rate. Inbreeding reduced the frequency of pollinator visits and increased flower-handling time, and this may be the first report that inbreeding affects pollinator behavior. Selfing rates of inbred plants were reduced by one half, which may reflect early inbreeding depression or altered pollinator behavior. The contrasting effects of herbivory and inbreeding on selfing rates may help maintain mating system variation in M. guttatus.     

Evaluating the influence of different aspects of habitat fragmentation on mating patterns and pollen dispersal in the bird-pollinated Banksia sphaerocarpa var. caesia

Habitat fragmentation can significantly affect mating and pollen dispersal patterns in plant populations, although the differential effects of the various aspects of fragmentation are poorly understood. In this study, we used eight microsatellite loci to investigate the effect of fragmentation on the mating system and pollen dispersal within one large and eight small population remnants of Banksia sphaerocarpa var. caesia, a bird-pollinated shrub in the southern agricultural region of Western Australia. The large population had a much larger neighbourhood size and lower selfing rate, maternal pollen pool differentiation and within-plot mean pollen dispersal distance than the small populations. Outcrossing was consistently high and ranged from 85.7% ± 2.6 to 98.5% ± 0.9, and mating patterns suggested nearest-neighbour pollination. Pollen immigration into small populations ranged from 2.8% ± 1.8 to 16.5% ± 3.2. Using the small populations, we tested for correlations between various fragmentation variables and mating system and pollen dispersal parameters. We found significant negative linear relationships between population isolation and outcrossing rate; population shape and neighbourhood size; and conspecific density and mean pollen dispersal distance. There were significant positive linear relationships between population shape and pollen pool differentiation and between population size and number of different fathers per seed crop. Our results suggest that birds may use a series of fragmented populations as a vegetation corridor while foraging across the landscape and that population connectivity is a critical determinant of pollinator visitation. Our results also suggest that the effect of a linear population shape on the mating system and pollen dispersal is routinely underestimated.