The presence of co-flowering species facilitates reproductive success of Pedicularis monbeigiana (Orobanchaceae) through variation in bumble-bee foraging behaviour

Background and Aims The presence of co-flowering species can alter pollinator foraging behaviour and, in turn, positively or negatively affect the reproductive success of the focal species. Such interactions were investigated between a focal species, Pedicularis monbeigiana, and a co-flowering species, Vicia dichroantha, which was mediated by behaviour alteration of the shared bumble-bee pollinator. Methods Floral display size and floral colour change of P. monbeigiana were compared between pure (P. monbeigiana only) and mixed (P. monbeigiana and V. dichroantha) plots in two populations. Pollinator visitation rates, interspecific floral switching and successive within-plant pollinator visits were recorded. In addition, supplemental pollination at plant level was performed, and the fruit set and seed set were analysed in pure and mixed plots with different densities of P. monbeigiana. Key Results Pollinator visitation rates were dramatically higher in mixed plots than in pure plots. The higher pollinator visitation rates were recorded in both low- and high-density plots. In particular, successive flower visits within an individual plant were significantly lower in mixed plots. Supplemental pollination significantly increased fruit set and seed set of individuals in pure plots, while it only marginally increased seed set per fruit of plants in mixed plots. Conclusions The presence of V. dichroantha can facilitate pollination and increase female reproductive success of P. monbeigiana via both quantity (mitigating pollinator limitation) and quality (reducing geitonogamy) effects. This study suggests that successive pollinator movements among flowers within a plant, as well as pollinator visitation rates and interspecific flower switching, may be important determinants of the direction and mechanisms of interaction between species.     

Effects of floral herbivory, limited pollination, and intrinsic plant characteristics on phenotypic gender in Sanicula arctopoides

Sexual expression in hermaphroditic plants is often a function of environmental factors affecting individuals before or during flowering. I tested for the effects of floral herbivory and lack of pollination in early umbels on the relative proportions of hermaphroditic and staminate (male) flowers produced on later umbels by Sanicula arctopoides, a monocarpic, andromonoecious perennial. Neither floral herbivory or lack of early pollination had a significant effect on the ratio of the two floral morphs, but the probability of producing staminate flowers on late umbels was strongly and positively related to plant size measured just prior to floral initiation and prior to herbivory. Plant size was also negatively correlated with flowering date. I suggest that producing staminate flowers on late umbels should benefit large early-blooming plants more than small late-blooming plants because more mating opportunities occur during the period when these flowers release pollen. Although herbivory did not cause labile changes of sex, whole plant phenotypic gender was still strongly affected by various forms of treatment. Sex-biased herbivory or lack of pollination rendered plants more or less phenotypically male, depending on which tissues were affected. Deer and pollen-feeding mites preferentially remove male tissues while hymenopteran seed predators preferentially remove female tissues. I conclude that combinations of herbivores could have counteracting or compounding effects on plant gender, and these effects may change the rankings of male and female reproductive success within populations. Received: 20 February 1996/Accepted: 30 July 1996     

Determinants of reproductive success in female Columbian ground squirrels

Summary We studied the reproductive success of female Columbian ground squirrels (Spermophilus columbianus) in southwestern Alberta for nine years. We defined reproductive success as the number of offspring surviving their first hibernation, classified as yearlings. The number of weaned juveniles explained one third of the variance in number of yearlings at emergence from their first hibernation the following spring, and much of the variance in individual reproductive success originated after weaning. Weight of adult females at emergence from hibernation was correlated with annual reproductive success. The mother's survival beyond weaning and the subsequent winter's snow accumulation had positive effects on annual reproductive success, whereas population density and summer temperature had negative effects. We found no effects on annual reproductive success of date of litter emergence, weight at emergence as a yearling, presence or absence of adult kin, distance from the natal site, location within the study area, winter temperature or summer precipitation. Age of first breeding did not affect lifetime reproductive success, which ranged from 0 to 19 yearlings produced over a lifetime. The greatest source of variation in lifetime reproductive success for females surviving to breeding age was offspring survival, followed by reproductive lifespan.     

Differences in female reproductive success between female and hermaphrodite individuals in the subdioecious shrub Eurya japonica (Theaceae)

Subdioecy is thought to occupy a transitional position in the gynodioecy–dioecy pathway, explaining one of the evolutionary routes from hermaphroditism to dioecy. Quantifying any female reproductive advantage of females versus hermaphrodites is fundamental to examining the spectrum between subdioecy and dioecy; however, this is challenging, as multiple interacting factors, such as pollen limitation and resource availability, affect plant reproduction. We compared the female reproductive success of females and hermaphrodites via a field experiment in which we hand‐pollinated individuals of the subdioecious shrub Eurya japonica of similar size growing under similar light conditions. Effects of pollen limitation and seed quality were also evaluated through comparing the results of hand‐ and natural‐pollination treatments and performing additional laboratory and greenhouse experiments. Overall, females had higher fruit set and produced heavier fruit and more seeds than hermaphrodites, and these results were more pronounced for hand‐pollinated than for natural‐pollinated plants of both sexes. We also found that seeds naturally produced by females had a higher mean germination rate. These results indicate that females had a pronounced advantage in female reproductive success under conditions of no pollen limitation. The sexual difference in the degree of pollen limitation suggests a pollinator‐mediated interaction, whereas the higher female reproductive success of females even under natural conditions implies that E. japonica is a good model species for elucidating the later stages of the gynodioecy–dioecy pathway.     

Effects of floral display size on male and female reproductive success in Mimulus ringens

Background and Aims

The number of flowers blooming simultaneously on a plant may have profound consequences for reproductive success. Large floral displays often attract more pollinator visits, increasing outcross pollen receipt. However, pollinators frequently probe more flowers in sequence on large displays, potentially increasing self-pollination and reducing pollen export per flower. To better understand how floral display size influences male and female fitness, we manipulated display phenotypes and then used paternity analysis to quantify siring success and selfing rates.

Methods

To facilitate unambiguous assignment of paternity, we established four replicate (cloned) arrays of Mimulus ringens, each consisting of genets with unique combinations of homozygous marker genotypes. In each array, we trimmed displays to two, four, eight or 16 flowers. When fruits ripened, we counted the number of seeds per fruit and assigned paternity to 1935 progeny.

Key Results

Siring success per flower declined sharply with increasing display size, while female success per flower did not vary with display. The rate of self-fertilization increased for large floral displays, but siring losses due to geitonogamous pollen discounting were much greater than siring gains through increased self-fertilization. As display size increased, each additional seed sired through geitonogamous self-pollination was associated with a loss of 9·7 seeds sired through outcrossing.

Conclusions

Although total fitness increased with floral display size, the marginal return on each additional flower declined steadily as display size increased. Therefore, a plant could maximize fitness by producing small displays over a long flowering period, rather than large displays over a brief flowering period.     

Cost and benefits of attractive floral traits in the annual species Madia sativa (Asteraceae)

The maintenance of flower size variation within populations might be explained by conflicting selection pressures on floral traits that may involve biological agents, such as mutualists and antagonists, and allocation costs associated with floral display. The annual species Madia sativa (Asteraceae) exhibits ample variation in the number of ray florets in natural populations. This field study aimed at evaluating the costs and benefits associated with floral traits in M. sativa. In particular, we addressed two main questions: (1) Is the number of ray florets positively associated with pollinator visitation rate? (2) Is there a fitness cost of ray floret maintenance when pollinators are absent? We detected one benefit of conspicuous ray capitula: a strong preference by insect pollinators. We also confirmed the occurrence of costs: when pollinators were excluded conspicuous ray capitula had a reduced reproductive assurance via autogamous selfing, and there were trade-offs between the number of ray florets and seed mass and seed germination. Results suggest that the maintenance of within-population variation in the number of ray florets in M. sativa is explained, at least in part, by the balance between costs and benefits associated with this floral trait.     

开花时间与伴生种对鹤首马先蒿传粉和生殖成功的影响

开花物候是物种间相互作用的重要生活史特征和适合度因子,在全球气候变化的背景下而备受关注.为探讨开花时间如何存种内和种间水平上影响植物的传粉和生殖成功,我们连续3年(2003-2005)对不同花期和伴生种存在情况下的鹤首马先蒿(Pedicularis gruina)的传粉者访花忠实度、受粉率、坐果率、单果种子产量和果实被啃食频率进行了比较研究.结果表明鹤首马先蒿的坐果率主要受其传粉环境的影响:在没有伴生种时,不同时期鹤首马先蒿的坐果率没有显著差异,34-38％的花可以坐果;在有伴生种存在时既可以显著提高其坐果率,也可显著降低其坐果率,这取决于传粉者类型以及伴生种密穗马先蒿(P.densispica)花期的差异.密穗马先蒿具有花蜜和花粉双重报酬,在群落中可以作为主体物种吸引传粉者并间接促进与其伴生的鹤首马先蒿的传粉和生殖成功.同样无蜜的管花马先蒿(P.siphonantha)和鹤首马先蒿伴生,则是通过提高群落水平对传粉者的吸引力进而直接促进鹤首马先蒿的传粉和生殖成功.此外,研究结果也表明开花时间对坐果率没有显著影响,但是显著影响单果种子产量和果实被啃食的频率.在相同的传粉条件下早花期植株单果种子产量显著高于晚花期的种子产量,同时早花期的果实被啃食的频率显著增加.     

Age-related and individual differences of reproductive success in male and female barbary macaques (Macaca sylvanus)

Age-related and individual differences in longterm reproductive success were analyzed in two social groups of free-ranging Barbary macaques. Maternity data were obtained from continuous birth records and paternity was determined with oligonucleotide-fingerprinting. The fathers of 246 of 286 investigated individuals could be identified. They were born during a 14-year period and represented 73 and 34% of all known offspring from the females of the study groups B/F and C, respectively. Only these infants were considered when comparing male reproductive success with that of females. The necessary adjustment of the female data resulted in small deviations from the true values in one group, but substantially increased individual differences in female fertility in the second group. Subadult males, 4.5 – 6.5 yrs old, had a much lower reproductive success than adult males (7.5 – 25 yrs old) and same-aged females. Reproductive success of adult males was not significantly affected by age, while females invariably ceased reproduction during the first half of the third decade of life. Males were more likely than females to leave no offspring, unless they survived 9 – 10 yrs of age. The number of years with breeding opportunities was important for male reproductive success but less significant than that for females. Reproductive success of several males during the 14-year study period was similar to or even exceeded that possible for a female in her whole lifetime. Variance of male reproductive success significantly exceeded that of females in both study groups.     

Seasonal changes in components of male and female reproductive success in Raphanus sativus L. (Brassicaceae)

Summary To document seasonal changes in the reproductive behavior of the perfect-flowered, self-incompatible mustard, Raphanus sativus L., we monitored individual survival, flower and fruit production among 58 individuals in a California population over six census dates (cohorts). Population size declined dramatically and mean individual levels of fruit set changed significantly between cohorts. The frequency distribution of flower and fruit production became increasingly skewed over the first four cohorts. The phenotypic maleness of individuals, a standardized measure of phenotypic gender, oscillated during the reproductive season, peaking in the third and fourth cohorts. We calculated a simple estimate of expected male reproductive success of each plant (the number of fruits sired on conspecifics); this estimate was a function of an individual's flower production and the fruit production of its potential mates in our sampled population. Mean expected male success did not differ significantly among cohorts; expected male success per flower did, however, change significantly among cohorts. Among individuals within each cohort, maternal fruit production and expected male success were both positively correlated with flower production throughout the season. Spearman rank correlation coefficients indicate that the strength of these associations, however, changed during the season. Linear regressions of transformed variables indicated that the shape of several fitness functions also changed over time. In addition, the amount of variation in maternal or expected paternal success explained by flower production declined over the first four cohorts. If typical of wild populations, these temporal changes in these functions suggest that measurements of the intensity of phenotypic selection on flower production will depend on when and how fitness is measured in natural populations.     

Neonatal manipulation of oxytocin influences female reproductive behavior and success

During early neonatal development, oxytocin (OT) may influence the expression of adult behavior and physiology. Here we test the prediction that early postnatal exposure to OT or an oxytocin antagonist (OTA) can affect the subsequent expression of sexual receptivity and reproductive success of females. To test this hypothesis, female prairie voles (Microtus ochrogaster) received one of four treatments within 24 h of birth. Three groups received an intraperitoneal injection of OT, OTA, or isotonic saline. A fourth group was handled, but not injected. Around 75 days of age, females were paired with sexually experienced males for 72 h and sexual activity was recorded. Treatment had no effect on the probability of mating. Injection, regardless of treatment, reduced latency to mate compared with handled controls. OT and OTA treatment decreased mating bout frequency compared to saline and handled controls, while OTA treatment increased reproductive success, probability of successfully producing a litter. The results suggest that neonatally OT, both endogenous and exogenous, can affect the expression of adult female reproductive activity and that blocking the effects of endogenous OT during neonatal development can affect female reproductive success. Finally, the results suggest that a number of aspects of reproduction are regulated by OT during the postnatal period, but that the mechanism of action may differ depending upon the reproductive activity.     

Highly efficient pollination by bumblebees ensures seed production in Pedicularis lachnoglossa (Orobanchaceae), an early‐flowering Himalayan plant

Abstract Pedicularis (Orobranchaceae) is a common high altitude genus of the Himalayas that may be affected by pollination limitation. Using Pedicularis lachnoglossa from Yulong (Jade Dragon) Snow Mountain in Lijiang (Yunnan Province, southwest China), we investigated the effects of high altitude habitats on the process of pollination and seed production. Floral biology, pollinator foraging behavior, breeding system, and pollination efficiency were examined using observation and exclusionary techniques. Pedicularis lachnoglossa was found to be entomophilous and exclusively pollinated by Bombus friseanus and B. yunnanicola. Our results indicated that pollination limitation in P. lachnoglossa was not significant. Under open pollination, approximately 80% of flowers were successfully pollinated and developed to fruits, and about 38% of ovules developed to mature seeds. Bumblebee pollination is highly precise and efficient in P. lachnoglossa, because its flowering phenology and floral characters enhance the foraging of bumblebees on flowers. This study supports that animal pollination plays a crucial role in the outbreeding of the early flowering Pedicularis. The evolution of floral specification in Pedicularis has the advantages of adaptation to bumblebee pollination in adverse high altitude habitats.     

The long and the short of gene flow and reproductive isolation: Inter-Simple Sequence Repeat (ISSR) markers support the recognition of two floral forms in Pelargonium reniforme (Geraniaceae)

The current taxonomy of Pelargonium reniforme recognises two subspecies on the basis of habit and vegetative characters, but excludes floral characters. However, populations of P. reniforme in the wild tend to belong to easily discernable floral groups based on floral colour and hypanthium length. The aim of this study was to determine the genetic diversity within and between populations of both subspecies using the Inter-Simple Sequence Repeat (ISSR) method of DNA fingerprinting. Ninety individuals from eight populations were sampled. Both phenetic and Neighbor-Net analyses reveal that populations are genetically discernable, but that there is no genetic evidence to support the recognition of the two currently defined subspecies of P. reniforme. Instead, the analyses resolved all individuals that possess long hypanthia into a single group, and the role of different pollinators in driving reproductive isolation of this long-tubed form is suggested.     

Male dominance and genetically determined reproductive success in the mandrill (Mandrillus sphinx)

Darwin referred to the adult male mandrill (Mandrillus sphinx) as the most brightly coloured of all mammals, citing the brilliant red and blue pigmentation of the face, rump, and genitalia as extreme examples of evolution by sexual selection. Considerable controversy exists concerning possible effects of sexually selected phenotypes via intermale competition on reproductive success. Behavioural and genetic studies of a large, semi-free ranging mandrill colony in Gabon have now demonstrated that clear-cut relationships exist between male secondary sexual development, social dominance, copulatory behaviour, and reproductive success in this primate species. Two morphological variants of adult male were identified; “fatted” males, with maximum secondary sexual coloration, which occupied dominant positions in the social group, and “non-fatted” males, with muted secondary sexual adornments, smaller testes and lower plasma testosterone levels, which lived as peripheral/solitary individuals. DNA fingerprinting analyses on infants born over five successive years showed that only the two most dominant, fatted males in the group had fathered offspring. Throughout the annual mating season these males attempted to mate-guard and copulate with females during periods of maximal sexual skin tumescence. Male rank and mating success were strongly positively related and the alpha male sired 80 – 100% of the resulting offspring during three consecutive years. Non-fatted adult males and group associated subadult males engaged in infrequent, opportunistic matings and did not guard females. Loss of alpha status resulted in a fall in reproductive success, but the effect was gradual; the deposed alpha male continued to father 67% and 25% of infants born during the next two years. Thus, whilst claims that male dominance determines mating success and paternity in primates have caused considerable debate, these results on mandrills provide unequivocal evidence for the existence of such effects.     

Effects of number of flowers per raceme and number of racemes per plant on bumblebee visits and female reproductive success in <Emphasis Type="Italic">Salvia nipponica</Emphasis> (Labiatae)

We studied the adaptive arrangement of raceme flowers in dense and sparse habitats of Salvia nipponica. We recorded bumblebee visits and collected mature seeds to estimate outcrossing rates from allozyme genotypes. To examine the resource investment in flowering stalks, we measured their length and dry mass. We found that a greater number of open flowers in a raceme enhanced bumblebee visits and successive probings in both density plots. However, greater height of a raceme did not enhance these in plots of either density. In the high-density plot, both a greater mean number of open flowers in a raceme and a greater number of flowering racemes on a plant enhanced bumblebee visits, successive probings, and also enhanced outcrossing rate in spite of an increase in successive probings. Although the number of flowers per raceme had a greater positive effect on seed-to-ovule ratios than did the number of racemes, the number of flowers per raceme may be constrained because a longer flowering stalk needs greater buckling strength. Hence, it may be advantageous for plants in high-density plots to increase both number of flowers per raceme and number of racemes. In the low-density plot, a greater number of flowering racemes on a plant enhanced pollinator visits, but a greater mean number of open flowers in a raceme did not. Hence, it may be advantageous for plants to increase the number of racemes while keeping the numbers of flowers per raceme small. Thus, the adaptive number of flowers per raceme and number of racemes per plant may differ between high- and low-density plots.     

Effects of herbivory and competition on an introduced plant in decline

Anthemiscotula was introduced to Denmark 500 years ago, and its distribution is presently limited and in decline. A manipulative field experiment was performed to investigate the effects of native plant competitors and native invertebrate herbivores on its performance. Generally, both herbivory and competition treatments had great impact, and when both factors were operating, the effects were additive for all variables except plant height. Although A. cotula showed plasticity in growth, resource allocation and flowering timing, it was unable to adjust to competition and compensate for losses due to herbivory sufficiently to ensure and restore its achene production. This vulnerability, combined with improved cereal cleaning techniques and thus fewer reintroductions of A. cotula seeds, may be the cause of its current decline. A. cotula responded to herbivory by prolonging its flowering period, a “bet-hedging” strategy. In Denmark this strategy is unreliable since risks of sub-optimal conditions are much greater in August–October. Received: 12 December 1997 / Accepted: 14 October 1998     

Effects of inflorescence size on visits from pollinators and seed set of Corydalis ambigua (Papaveraceae)

Female reproductive success (seed set) of a spring ephemeral plant, Corydalis ambigua Cham. et schlecht (Papaveraceae) was investigated in relation to inflorescence size and foraging behavior (frequency and duration of visitations) by pollinators (namely, overwintered queens of Bombus hypocrita sapporensis) by detailed daily observations of a natural population. Pollination experiments indicated that C. ambigua is self-incompatible and that seed set was significantly affected by the behavior of the pollinating queens. Plants with larger inflorescences were visited more often than those with fewer flowers. Fecundity also increased with increasing size of inflorescences. Visitation time (duration of foraging) rather than the frequency of visitations (number of visits) was critical for higher fecundity. Seed production was strongly enhanced by a few long visits (of more than 60 s), and seemed to be independent of large numbers of short visits (of less than 60 s). Hence, plants with larger inflorescences, which provide a conspicuous signal to pollinators and offer greater rewards in terms of nectar, received longer visits by B. hypocrita sapporensis queens and those plants exhibited higher fecundity.     

Reproductive success in varying light environments: direct and indirect effects of light on plants and pollinators

Plant populations often exist in spatially heterogeneous environments. Light level can directly affect plant reproductive success through resource availability or by altering pollinator behavior. It can also indirectly influence reproductive success by determining floral display size which may in turn influence pollinator attraction. We evaluated direct and indirect effects of light availability and measured phenotypic selection on phenological traits that may enhance pollen receipt in the insect-pollinated herb Campanulastrum americanum. In a natural population, plants in the sun had larger displays and received 7 times more visits than plants in the shade. Using experimental arrays to separate the direct effects of irradiance on insects from their response to display size, we found more visits to plants in the sun than in the shade, but no association between number of visits each flower received and display size. Plants in the sun were not pollen limited but pollen-augmented shade flowers produced 50% more seeds than open-pollinated flowers. Phenological traits, which may influence pollen receipt, were not under direct selection in the sun. However, earlier initiation and a longer duration of flowering were favored in the shade, which may enhance visitation in this pollen-limited habitat.     

Impact of pre‐ and post‐pollination barriers on pollen transfer and reproductive isolation among three sympatric Pedicularis (Orobanchaceae) species

• Pedicularis is the largest genus in the Orobanchaceae (>300) with many species co‐occurring and co‐blooming in subalpine to alpine meadows in the Himalayas. Although it is well known that different Pedicularis species place pollen on different parts of the same bumblebee's body, thus reducing interspecific pollen transfer, it is not known whether post‐pollination components also contribute to reproductive isolation (RI).
• In this study, we quantified the individual strengths and absolute contributions of six pre‐ and post‐pollination components of RI between three sympatric species in two pairs; Pedicularis gruina × Pedicularis tenuisecta (gru × ten) and Pedicularis comptoniifolia × Pedicularis tenuisecta (com × ten).
• All three Pedicularis species shared the same Bombus species. Individual foragers showed a high, but incomplete, floral constancy for each species. Therefore, pre‐pollination barriers were potentially ‘leaky’ as Bombus species showed a low but consistent frequency of interspecific visitation. The RI strength of pre‐pollination was lower in com × ten than in gru × ten. In contrast, post‐pollination barriers completely blocked gene flow between both sets of species pairs. Two post‐pollination recognition sites were identified. Late acting rejection of interspecific pollen tube growth occurred in com♀ × ten♂, while seeds produced in bi‐directional crosses of gru × ten failed to germinate.
• We propose that, although floral isolation based on pollen placement on pollinators in the genus Pedicularis is crucial to avoid interspecific pollen transfer, the importance of this mode of interspecific isolation may be exaggerated. Post‐pollination barriers may play even larger roles for currently established populations of co‐blooming and sympatric species in this huge genus in the Himalayas.

     

Density and reproductive success in wild populations of Diplotaxis erucoides (Brassicaceae)

Summary One possible consequence of low population density, particularly in self-incompatible plants, is reproductive failure. I surveyed seed set per flower in two populations of the self-incompatible annual Diplotaxis erucoides (Brassicaceae) in Jerusalem, Israel. Widely spaced plants had lower fruit set and fewer seeds per filled silique than did plants growing close to conspecific neighbors. Such density-dependent reproductive success could help explain the maintanence of spatial patchiness in plant populations, and could also have implications for population dynamics of rare species.