Relationships between flowering phenology,plant size and reproductive success in shape Lotus corniculatus (Fabaceae)

Over three years the flowering phenology of individuals of Lotus corniculatus has been studied in relation to fruit set and seed predation to determine the relationships between four components of flowering time, plant size and reproductive success. Timings of first and peak flowering, and duration and synchrony of flowering differed between individuals in the same years. Between years, timing of first flowering was highly correlated for the same individuals, and was closely correlated with plant size and duration of flowering–larger plants flowered earlier and for a longer period. Peak flowering and synchrony were not correlated between-years for individuals.Fruit production and seed predation were correlated with some of the components of flowering phenology in some years, but not in others. The inconstancy of these relationships suggests that directional or stabilising selection is not acting consistently on the aspects of reproductive success studied in this work. The inconstancy of selection may result in the rather asynchronous flowering phenologies of individuals of L. corniculatus observed.We emphasize the importance of studying different components of flowering phenology in relation to individual plant size over several seasons. This work has shown that plant size not only has a direct effect on individual plant fecundity but also can influence flowering time and hence indirectly affect reproductive output.     

Wind pollination and reproductive assurance in Linanthus parviflorus (Polemoniaceae), a self-incompatible annual

Wind pollination was experimentally demonstrated in Linanthus parviflorus (Polemoniaceae), a predominantly beefly-pollinated, self-incompatible annual. Seed set in plants enclosed in mesh tents that excluded pollinators but allowed airborne pollen flow provided evidence for wind pollination, and the extent of seed set due to wind pollination was compared to that in open-pollinated controls and pollen-supplemented treatments. Additional controls were included to test for possible confounding effects of the mesh tent. Mean seed number in open-pollinated plants was 72.8–81.1% of that in pollen-supplemented plants, while wind pollination alone produced 49.5–52.2%, a smaller but substantial proportion of seed set with pollen supplementation. Further evidence for wind pollination was found in a comparison of sites differing in the extent of wind exposure in two populations of L. parviflorus. Airborne pollen counts were higher in exposed sites than in protected sites, and the difference was marginally significant. Seed set was significantly pollen limited in protected sites, but not in exposed sites. Taken together, the data suggest that wind pollination provides some reproductive assurance in this obligately outcrossing species. Wind pollination is hypothesized to represent an alternative to selfing as an evolutionary solution to the problem of temporal or spatial variation in pollination visitation.     

Tree size and its relationship with flowering phenology and reproductive output in Wild Nutmeg trees

Reproductive strategies, sexual selection, and their relationship with the phenotype of individuals are topics widely studied in animals, but this information is less abundant for plants. Variability in flowering phenology among individuals has direct impact on their fitness, but how reproductive phenology is affected by the size of the individuals needs further study. We quantified the flowering intensity, length, and reproductive synchronization of two sympatric dioecious Wild Nutmeg tree species (Virola, Myristicaceae) in the Brazilian Atlantic forest, and analyzed its relationships with tree size. Two distinct strategies in flowering timing and intensity were found between species (annual versus biennial flowering), and among individuals in the annual flowering species (extended versus peak flowering). Only for the annual flowering species the reproductive output is related to tree size and large trees present proportionally higher flower coverage, and lower synchronization than smaller ones. Flowering is massive and highly synchronized in the biennial species. Sex ratios are not different from 1:1 in the two species, and in the two segregated reproductive subgroups in the biennial flowering species. The biennial flowering at individual level is a novelty among reproductive patterns in plants, separating the population in two reproductive subgroups. A proportional increase in the reproductive output with size exists only for the annual flowering species. A biennial flowering can allow resource storage favouring massive flowering for all the individuals diluting their relationship with size.     

Individual variation in pathogen attack and differential reproductive success in the annual legume,Amphicarpaea bracteata

Summary Demographic analyses in two natural populations of the annual legume Amphicarpaea bracteate examined whether variation in attack by the host-specific fungal pathogen Synchytrium decipiens was associated with variation in the reproductive success of individual plants. In both populations, fungal infection early in life was significantly associated with reduced seedling growth rates. Laboratory inoculation experiments confirmed that S. decipiens infection had a negative impact on plant growth. The laboratory experiments further indicated that there was significant variation among the progenies of different plant genotypes in the degree of growth reduction caused by pathogen attack. Prereproductive mortality rates in natural environments were significantly higher for plants with infection intensities above the median; for the two populations studied, heavily infected plants had 3.8 and 12 gimes higher death rates compared with low infection plants. Among surviving plants, fungal infection intensity was significantly negatively correlated with total seed biomass in both populations. As a result of these associations between plant survivorship, fecundity, and fungal infection, lifetime relative fitness within both plant populations was strongly negatively correlated with the intensity of S. decipiens infection. These results demonstrate the existence of consistent natural selection for increased resistance to pathogen attack in this plant species.     

Phenotypic selection on flowering phenology and size in two dioecious plant species with different pollen vectors

Dioecious plants may be pollinated biotically by animals or abiotically via wind or water currents. It has been hypothesized that these two types of pollen vectors might impose contrasting selective pressures on plant flowering phenology. In the present study we describe the flowering phenology of two sympatric dioecious species with contrasting pollination modes: Mercurialis perennis (wind‐pollinated) and Tamus communis (insect‐pollinated). We estimated selection differentials and gradients for flowering time and flowering synchrony. As flowering time might depend on the accumulation of enough internal resources, we also estimated direct and indirect selection on plant size. Both species have male‐biased sexual ratios, and males are bigger and produce larger flower displays than females, but only in T. communis do males bloom earlier and for longer than females. Selection gradients suggest that selection tends to favor early‐flowering females of T. communis. There is no evidence of direct current selection on the flowering phenology of M. perennis. Intersexual differences in phenology fit with sex allocation and sexual selection theories. As we hypothesized, phenology of the animal‐pollinated species is under stronger selection than that of the wind‐pollinated species and we discuss the potential role of pollen vectors in shaping the flowering phenologies of the study species.     

Consequences of variation in flowering phenology for seed head herbivory and reproductive success in Erigeron glaucus (Compositae)

Summary We examined the relationship between flowering phenology, reproductive success (seed production only), and seed head herbivory for 20 similarly sized clones of Erigeron glaucus growing at Bodega Bay Reserve, northern California, USA. Although clones tended to reach peak flowering on the same date, they differed in the proportion of their total flowers produced around that date (flowering synchrony). Clones also differed in the number and density of flower heads presented at any one time to pollinators and herbivores (floral display). Both of these characteristics had consequences for herbivory and plant reproductive success. The proportion of flower heads damaged by insect herbivores was greater for clones that concentrated flowering activity during the main flowering period for the population as a whole (high synchrony) compared to clones that spread flowering out temporally. The primary reason for this result was that clones with low flowering synchrony produced a significant proportion of their flower heads during the fall and therefore, escaped attack by the tephritid fly, Tephritis ovatipennis. Clones with intermediate synchrony had lower seed success (total number of viable seeds produced over the year) than clones with either low or high synchrony. The proportion of flower heads damaged by insect herbivores and number of tephritid flies reared from flower heads were both negatively correlated to floral display while seed head mass and germination rates were positively related to display. Thus, clones which produced dense floral displays were favored both in terms of reduced herbivory and increased successful seed production.     

Individual flowering phenology shapes plant–pollinator interactions across ecological scales affecting plant reproduction

The balance of pollination competition and facilitation among co‐flowering plants and abiotic resource availability can modify plant species and individual reproduction. Floral resource succession and spatial heterogeneity modulate plant–pollinator interactions across ecological scales (individual plant, local assemblage, and interaction network of agroecological infrastructure across the farm). Intraspecific variation in flowering phenology can modulate the precise level of spatio‐temporal heterogeneity in floral resources, pollen donor density, and pollinator interactions that a plant individual is exposed to, thereby affecting reproduction. We tested how abiotic resources and multi‐scale plant–pollinator interactions affected individual plant seed set modulated by intraspecific variation in flowering phenology and spatio‐temporal floral heterogeneity arising from agroecological infrastructure. We transplanted two focal insect‐pollinated plant species (Cyanus segetum and Centaurea jacea, n = 288) into agroecological infrastructure (10 sown wildflower and six legume–grass strips) across a farm‐scale experiment (125 ha). We applied an individual‐based phenologically explicit approach to match precisely the flowering period of plant individuals to the concomitant level of spatio‐temporal heterogeneity in plant–pollinator interactions, potential pollen donors, floral resources, and abiotic conditions (temperature, water, and nitrogen). Individual plant attractiveness, assemblage floral density, and conspecific pollen donor density (C. jacea) improved seed set. Network linkage density increased focal species seed set and modified the effect of local assemblage richness and abundance on C. segetum. Mutual dependence on pollinators in networks increased C. segetum seed set, while C. jacea seed set was greatest where both specialization on pollinators and mutual dependence was high. Abiotic conditions were of little or no importance to seed set. Intra‐ and interspecific plant–pollinator interactions respond to spatio‐temporal heterogeneity arising from agroecological management affecting wild plant species reproduction. The interplay of pollinator interactions within and between ecological scales affecting seed set implies a co‐occurrence of pollinator‐mediated facilitative and competitive interactions among plant species and individuals.     

The role of individual variation in flowering and pollination in the reproductive success of a crepuscular buzz-pollinated plant

Background and AimsPlant individuals within a population differ in their phenology and interactions with pollinators. However, it is still unknown how individual differences affect the reproductive success of plants that have functionally specialized pollination systems. Here, we evaluated whether plant individual specialization in phenology (temporal specialization) and in pollination (pollinator specialization) affect the reproductive success of the crepuscular-bee-pollinated plant Trembleya laniflora (Melastomataceae).MethodsWe quantified flowering activity (amplitude, duration and overlap), plant–pollinator interactions (number of flowers visited by pollinators) and reproductive success (fruit set) of T. laniflora individuals from three distinct locations in rupestrian grasslands of southeastern Brazil. We estimated the degree of individual temporal specialization in flowering phenology and of individual specialization in plant–pollinator interactions, and tested their relationship with plant reproductive success.Key Results Trembleya laniflora presented overlapping flowering, a temporal generalization and specialized pollinator interactions. Flowering overlap among individuals and populations was higher than expected by chance but did not affect the individual interactions with pollinators and nor their reproductive success. In contrast, higher individual generalization in the interactions with pollinators was related to higher individual reproductive success.ConclusionsOur findings suggest that individual generalization in plant–pollinator interaction reduces the potential costs of specialization at the species level, ensuring reproductive success. Altogether, our results highlight the complexity of specialization/generalization of plant–pollinator interactions at distinct levels of organization, from individuals to populations, to species.     

Individual reproductive success and effective population size in the greater white-toothed shrew Crocidura russula

In order to investigate the determinants of effective population size in the socially monogamous Crocidura russula, the reproductive output of 44 individuals was estimated through genetic assignment methods. The individual variance in breeding success turned out to be surprisingly high, mostly because the males were markedly less monogamous than expected from previous behavioural data. Males paired simultaneously with up to four females and polygynous males had significantly more offspring than monogamous ones. The variance in female reproductive success also exceeded that of a Poisson distribution (though to a lesser extent), partly because females paired with multiply mated males weaned significantly more offspring. Polyandry also occurred occasionally, but only sequentially (i.e. without multiple paternity of litters). Estimates of the effective to census size ratio were ca. 0.60, which excluded the mating system as a potential explanation for the high genetic variance found in this shrew's populations. Our data suggest that gene flow from the neighbourhood (up to one-third of the total recruitment) is the most likely cause of the high levels of genetic diversity observed in this shrew's subpopulations.     

Growth and phenology of Dittrichia graveolens, a rapidly spreading invasive plant in California

Dittrichia graveolens is a rapidly spreading invasive plant in California. While populations are observed primarily in disturbed areas, there is concern it may expand into adjacent undisturbed areas, particularly grasslands and riparian corridors. In a field experiment conducted in two successive years, we compared plant growth and phenological development of fall, winter, and spring sown seeds. Plants establish equally well in disturbed upland sites in both above and below average precipitation years but the absence of late spring rainfall negatively affected total plant biomass. In a greenhouse experiment, we compared growth in four light environments (100, 50, 27 and 9 % available light). Total plant growth decreased exponentially with decreasing light. This suggests that D. graveolens is not competitive in low light environments, such as woodlands and riparian forests. All plants flowered in early- to mid-September, coinciding with flowering in field grown plants, suggesting that photoperiod is the primary signal for reproductive growth. Using a minirhizotron system, we measured root growth over time in D. graveolens and three common California annual grassland species, two non-natives, Centaurea solstitialis and Bromus hordeaceus, and the native forb Holocarpha virgata. Root growth of D. graveolens began later in the season than the other species, reaching depths >1 m by late May. Roots of C. solstitialis and H. virgata reached >1 m earlier in the season. The temporal difference in root growth suggests that D. graveolens may be less competitive for soil moisture with other early season annuals than other deep-rooted broadleaf species found in grasslands.     

Age at flowering differentially affects vegetative and reproductive responses of a determinate annual plant to elevated carbon dioxide

Plant population and community dynamics may be altered by increasing atmospheric CO(2) concentrations [[CO(2)]] through intraspecific variation in the responses of vegetative and reproductive growth. Although these responses may be regulated by age at flowering, little is known about the direct effects of age at flowering on growth responses to elevated [CO(2)]. In this study, we examined the interactive effects of elevated [CO(2)] and age at flowering on absolute and relative allocation to vegetative and reproductive growth in the determinate, short-day species Xanthium strumarium L. (common cocklebur). Six cohorts were planted at 5-day intervals in chambers maintained at either 365 or 730 micro mol mol(-1) CO(2), with an 18-h photoperiod and a non-limiting nutrient supply. All plants were simultaneously induced to flower by switching the photoperiod to 12 h for 2 days, then switching back to an 18-h photoperiod for the remainder of the experiment. All plants were harvested 15 days after the onset of flowering. Total plant biomass increased 11-41% with increasing [CO(2)] and 45% from the youngest to the oldest cohort. Vegetative growth responses to elevated [CO(2)] significantly increased with increasing age at flowering, associated with increasing sink relative to source capacity. In contrast, total fruit mass decreased 32% from the youngest to the oldest cohort and was not significantly affected by CO(2) supply. Relative biomass allocation to fruit decreased 47% from the youngest to the oldest cohort, reflecting decreased numbers of fruit, and 6-28% with increasing [CO(2)], reflecting decreased mean mass per mature fruit. Our findings suggest that elevated [CO(2)] may increase vegetative growth in Xanthium without increasing reproductive biomass, and that age at flowering may influence these responses through effects on source:sink balance. Further, changes in the allometric relationship between vegetative and reproductive growth associated with growth in elevated [CO(2)] suggest that long-term population and community-level responses to elevated [CO(2)] may differ substantially from predictions based on vegetative responses.     

How reproductive allocation and flowering probability of individuals in plant populations are affected by position in stand size hierarchy,plant size and CO2 regime

Aims We investigate the effect of position within a size-structured population on the reproductive allocation (RA) and flowering probability of individual plants of Sinapis arvensis. We also assess the effects of plant size and changing level of CO₂ on both responses.Methods Sinapis arvensis L., (field mustard), an annual agricultural weed, was grown in monoculture at six densities under ambient and elevated CO₂ in a study with 84 stands. Individual aboveground biomass and reproductive biomass were measured. Varying density produced a wide range of mean plant sizes across stands and size hierarchies within stands. Many (~40%) individuals had zero reproductive biomass. Employing a novel modelling approach, we analysed the joint effects of position in stand size hierarchy, plant size and CO₂ on RA and flowering probability of individuals.Important findings We found a strong effect of position within the size hierarchy of individuals in a population: for an individual of a given size, greater size relative to neighbours substantially increased RA and flowering probability at a single harvest time. There was no other effect of plant size on RA. We found a positive effect of elevated CO₂ on RA regardless of position within the size hierarchy. These observed patterns could impact doubly on the reproductive biomass (R) of small individuals. First, because RA is not affected by size, smaller plants will have smaller R than larger plants; and second, for smaller plants lower down in a population size hierarchy, their RA and hence R will be further reduced. These results suggest that size relative to neighbours may be independent of and more important than direct abiotic effects in determining RA. Further studies are required to evaluate how these observed patterns generalize to other populations in non-experimental conditions.     

Sex ratio, reproductive investment and flowering phenology in dioecious Rhamnus alaternus (Rhamnaceae)

Guitián, J. 1995. Sex ratio, reproductive investment and flowering phenology in dioecious Rhamnus alaternus (Rhamnaceae). - Nord. J. Bot. 15: 139–143. Copenhagen. ISSN 0107–055X.
During 1992 and 1993 I investigated the reproductive biology of the dioecious Mediterranean shrub Rhamnus alaternus in a population in the northwest Iberian Peninsula. Reproductive investment was estimated as the mean total dry weight of reproductive organs per branch tip. I also estimated number of flowers produced per plant and population sex ratio, and investigated the spatial distribution of the sexes and flowering phenology. The sex ratio was 1:1, and the spatial distribution of the sexes was random. Male plants produced 2.6 times more flowers than female plants, but the overall reproductive investment by females was 5- to 9-fold higher. In both years of study, male plants commenced flowering first. The male and female flowering peaks coincided closely (in late March) in both years. The results of this study suggest that male and female R. alaternus differ most notably in the amount of resources allocated to reproduction.     

Clutch size and reproductive success in a female polymorphic insect

Differences in reproductive success (RS) between different groups of individuals are of interest to researchers studying natural and sexual selection. Since it is often not feasible to quantify RS in the wild, researchers make use of proxies instead. One such proxy is clutch size. However, research on species providing parental care (mainly birds and mammals) has learned that a large clutch size does not guarantee a large number of offspring. In contrast, much less is known on the link between clutch size and RS for species lacking parental care, such as many reptiles and insects. Here, we ask whether clutch size provides a satisfactory estimate of RS for a polymorphic insect. Our study species is a damselfly showing two distinct female morphs for which RS (estimated by clutch size) has been studied to evaluate the evolutionary role of sexual conflict. However, in this system not only among family variation in offspring viability, but also differences between female morphs, may affect how clutch size relates to offspring number and quality. To evaluate the use of clutch size as estimate of RS, we examined how clutch size correlated with subsequent success measures of developing offspring by rearing damselfly from eggs to adults under two laboratory food treatments. In both treatments, we detected that clutch size correlated well with offspring number early in larval life, but that this relation is reduced by among family variation in survival in later developmental stages. Clutch size was moderately correlated with the number of offspring that successfully metamorphosed to winged adults. Patterns did not differ between female morphs and the nature of the correlation could not be explained from offspring quantity-quality trade-offs.     

Plant size,flowering synchrony and edge effects: What,how and where they affect the reproductive success of a Neotropical tree species

Plant reproductive success is supposedly influenced by phenology and individual size, which may be modified under edge effects. We tested if reproductive success, estimated by fruit set, in Senefeldera verticillata (Euphorbiaceae) is related to flowering synchrony and tree size, including plant height and circumference at breast height. The study was carried out in the interior and in edges of clearings for gas pipelines and electric lines of a lowland rainforest in south‐eastern Brazil. Monthly observations were performed during one reproductive season, of 19 individuals that grew at edges of electric lines and gas pipelines and at forest interior. Reproductive success was significantly higher at forest interior than at gas pipeline area; there was no significant difference between gas pipeline and electric line areas or between forest interior and electric line area. In the forest edges, only plant height was positively related to plant reproductive success. This is probably related to crown exposure to sunlight, which enhances flower production. At forest interior, reproductive success was positively influenced by the synchrony of flowering activity among neighbouring individuals. In contrast, flowering synchrony based on phenophase intensity negatively impacted reproductive success. Senefeldera verticillata shows temporal dioecy and is mainly pollinated by small social bees, and the high degree of flowering synchrony at low intensity may increase the number of mating partners and therefore enhance its reproductive success. Inside the forest fragments, individuals with thicker trunks showed lower reproductive success, which may be related to a loss of reproductive capacity of older individuals. Our results evidenced the complexity of responses experienced by tropical plants subjected to forest fragmentation because of linear clearings.     

The genetic architecture of reproductive isolation in Louisiana irises: flowering phenology

Despite the potential importance of divergent reproductive phenologies as a barrier to gene flow, we know less about the genetics of this factor than we do about any other isolating barrier. Here, we report on the genetic architecture of divergent flowering phenologies that result in substantial reproductive isolation between the naturally hybridizing plant species Iris fulva and I. brevicaulis. I. fulva initiates and terminates flowering significantly earlier than I. brevicaulis. We examined line crosses of reciprocal F1 and backcross (BC1) hybrids and determined that flowering time was polygenic in nature. We further defined quantitative trait loci (QTL) that affect the initiation of flowering in each of these species. QTL analyses were performed separately for two different growing seasons in the greenhouse, as well as in two field plots where experimental plants were placed into nature. For BCIF hybrids (BC1 toward I. fulva), 14 of 17 detected QTL caused flowering to occur later in the season when I. brevicaulis alleles were present, while the remaining 3 caused flowering to occur earlier. In BCIB hybrids (BC1 toward I. brevicaulis), 11 of 15 detected QTL caused flowering to occur earlier in the season when introgressed I. fulva alleles were present, while the remaining 4 caused flowering to occur later. These ratios are consistent with expectations of selection (as opposed to drift) promoting flowering divergence in the evolutionary history of these species. Furthermore, epistatic interactions among the QTL also reflected the same trends, with the majority of epistatic effects causing later flowering than expected in BCIF hybrids and earlier flowering in BCIB hybrids. Overlapping QTL that influenced flowering time across all four habitat/treatment types were not detected, indicating that increasing the sample size of genotyped plants would likely increase the number of significant QTL found in this study.     

Poor reproductive success of Gray Vireos in a declining California population

Since the 1940s, populations of Gray Vireos (Vireo vicinior) in California have collapsed, presumably because of parasitism by Brown‐headed Cowbirds (Molothrus ater). In 2012 and 2013, we studied the vireo's nesting ecology to assess factors affecting two of California's largest remaining populations in the chaparral of San Diego County. Nest success was extremely low, with a model‐averaged probability of nest survival of only 0.08 (N = 95). More nest failures were due to predation (83%) than to cowbird parasitism (13%). Video‐recording at 30 nests revealed that California Scrub‐Jays (Aphelocoma californica) were the most common nest predator (67%). Of eight variables tested, height of shrubs surrounding the nest had the strongest negative influence on nest survival, but was more strongly correlated with cowbird parasitism than with jay predation. Despite frequent renesting, seasonal productivity was well below the level required to sustain a population, especially in northern San Diego County where we found no Gray Vireos at six of seven sites where they had been present from 1997 to 2001 and where cowbird parasitism was more frequent. The vireo's continuing range collapse contrasts with recent climate‐change models predicting a range expansion, highlighting the importance of demographic studies. Low nest success is likely contributing to population declines in California, and the additive effect of cowbird parasitism suppresses productivity. Conservation of Gray Vireos in California will likely require development of alternative approaches to cowbird and scrub‐jay control appropriate to sites widely scattered in rugged chaparral.     

Climate changes affecting biotic interactions,phenology, and reproductive success in a savanna community over a 10-year period

Climatic parameters are able to influence the timing of phenological events affecting the degree of synchrony among plant species, their interactions, and reproductive success. Shrubs of Malpighiaceae family in the Brazilian Tropical Savanna present sequential flowering phenology. We verified variations in climatic factors (temperature and precipitation) over a period of 10 years (2005–2014) and correlated them with the onset of flowering of four of these Malpighiaceae species. Furthermore, we tested whether the phenological synchronization among species has changed over time affecting the herbivory and fruit set. Herbivory and fruit production were recorded during three reproductive seasons (2008/2009, 2011/2012, 2013/2014). We developed a mathematical model to estimate the flower and fruit production in response to phenological changes for the next 5 years. Results show that climatic factors changed, influencing the onset of species flowering. The degree of overlap among species also changed and the effects on species interactions were species specific. The mathematical model successfully presented a tendency on flower and fruit production contributing to the predictions of the outcomes in response to phenological changes. We confirm the effects of climate changes on plant phenological events and the importance of feature plasticity for better performance of species.