THE FLORAL ECOLOGY OF PLATANTHERA BLEPHARIGLOTTIS

The floral ecology of a central Maine population of Platanthera blephariglottis was studied during the 1979–1981 flowering seasons. Although this species possesses characteristics typical of moth-pollinated plants, only diurnal pollinators were documented, primarily Hesperiidae, Pieridae, and Apidae. Pollinators were scarce throughout the study, yet bagging experiments indicated vectors were necessary for successful capsule set. The average capsule set per plant was lowest (32.2%) in 1979 when 362 plants bloomed and highest in 1980 (80.2%) when 202 plants flowered. Capsule set was found to be limited by pollinator activity. The adaptations of P. blephariglottis, including sequential flowering, length of receptivity, nectar production and inflorescence size, enable consistently successful reproduction regardless of microhabitat. This species is well-adapted to reproduce in environments experiencing periods of unfavorable weather and low pollinator abundance.     

Further studies on seed transmission of broad bean stain virus and Echtes Ackerbohnenmosaik -Virus in field beans (Vicia faba)

Broad bean stain virus (BBSV) and Echtes Ackerbohnenmosaik-Virus (EAMV) were detected in the seed coat and embryo sac fluid of immature seeds from infected field beans (Viciafaba minor) by inoculation to Phaseolus vulgaris; BBSV was also detected in immature embryos. The proportion of seeds infected with either virus decreased during maturation. The viruses were transmitted to seedlings as often through fully ripened seeds from which the seed coats had been removed as through intact seeds. Both viruses were detected in pollen from infected plants, but in glasshouse tests only BBSV was transmitted through pollen to seeds. Delaying fertilization in plants infected with BBSV or EAMV seemed not to affect seed transmission of either virus. In glasshouse tests BBSV was transmitted more often through seeds from plants that were inoculated before flowering than during flowering, and was not transmitted through seeds from plants inoculated after flowering; EAMV was transmitted only through seeds from plants inoculated before flowering. In tests on seed from naturally infected plants BBSV was transmitted more often through seeds from plants that developed symptoms before flowering than during flowering. Both viruses were seed-borne in all cultivars tested and there was no marked difference in the frequency of transmission of either virus among the spring-sown cultivars most common in Britain. Both viruses persisted in seed for more than 4 yr.     

POPULATION DEMOGRAPHY OF PASTINACA SATIVA (APIACEAE): EFFECTS OF SEED MASS ON EMERGENCE,SURVIVAL, AND RECRUITMENT

The effect of seed mass on emergence, survival, and recruitment to flowering in Pastinaca sativa, a species that produces progressively smaller seeds on the primary, secondary, and tertiary umbels, was analyzed in two field experiments begun in 1982 and 1983. In both experiments, overall emergence was positively related to seed mass, but the effects of seed mass on emergence in fall cohorts was relatively unimportant. Initial seed mass was positively related to overwintering survival in one experiment but not the other, possibly due to differences in winter conditions. Survival to flowering was positively related to seed mass in the 1982 experiment but negatively related to seed mass in the 1983 experiment. This difference is due to greater survival of seedlings from small seeds during short-term droughts, a previously unrecognized advantage of small seeds. Because of differences in survival in the two experiments, recruitment (number of flowering plants/number of seeds sown) was positively related to seed mass in the 1982 experiment but not in the 1983 experiment. The effects of timing of emergence on survival generally disappeared within 12 months. Our results suggest that the relationship between initial seed mass and recruitment in Pastinaca represents a distribution of outcomes, variable from year to year. However, because short-term droughts in Iowa are common, we hypothesize that in most years, seed mass has little impact on recruitment. Furthermore, small scale heterogeneity may often overwhelm any potential effects of seed mass and timing of emergence on recruitment.     

RAINFALL PERIODICITY AS A MAJOR FACTOR IN THE FORMATION OF FLOWERING RACES OF CAMPHORWEED (HETEROTHECA SUBAXILLARIS)

Plants of camphorweed (Heterotheca subaxillaris sensu latiore) were raised in a uniform garden at Northampton, Mass., from seed taken from 19 sites in different parts of the range of the species. Flowering races were observed. Flowering time at Northampton correlated with periodicities of rainfall at the seed source in most instances. The earliest-flowering plants were obtained from areas where above-average amounts of rainfall occur in spring, while the later-flowering plants came from sites where above-average rainfall occurs in mid- to late summer. Other workers have observed a correlation between the frost-free season and the formation of flowering races in prairie grasses, Eupatorium rugosum, and other plants. There is little correlation between the frost-free season and the formation of flowering races in Heterotheca and it is postulated that, within the range of camphorweed, periodicity of rainfall is a more critically limiting factor.     

CONSEQUENCES OF EMERGENCE PHENOLOGY FOR REPRODUCTIVE SUCCESS IN RANUNCULUS ADONEUS (RANUNCULACEAE)

This study explores the effects of emergence time and reproductive phenology on seed number, seed size, and seedling survival in a population of the alpine buttercup, Ranunculus adoneus. Phenology in this snow bowl population is structured by snow depth. Plants in late melting interior portions of the bowl emerged and flowered 3 to 4 wk after those in early melting zones at the bowl perimeter during the summers of 1988 and 1989. Flowering time differences of buttercups across the bowl were consistent from one year to the next. In 1988, late flowering plants tended to set fewer seeds than early flowering ones; in 1989 no decrease in seed number accompanied flowering date. Path analysis showed that equal fecundity in early and late emerging portions of the bowl population during 1989 resulted from balancing spatial and temporal constraints on seed production. Spatial aspects of habitat quality improved toward the interior of the bowl, but temporal regimes deteriorated in these late melting sites. In both 1988 and 1989 seed size declined with delays in flowering. Path analysis of 1989 data showed that because of reduced time for seed growth, plants in late melting portions of the bowl set smaller seeds than those in earlier melting zones. Differences in seed size due to parental phenology are likely to influence fitness in snow buttercups. Under natural conditions, seedlings from large seeds (>;0.65 mg) have sixfold higher survival than do those from smaller seeds (<;0.65 mg). We conclude that seedling recruitment may be infrequent in late-melting portions of the snow bowl due to delayed parental phenology.     

Assessing the potential of candidate DNA barcodes for identifying non‐flowering seed plants

In plants, matK and rbcL have been selected as core barcodes by the Consortium for the Barcode of Life (CBOL) Plant Working Group (PWG), and ITS/ITS2 and psbA‐trnH were suggested as supplementary loci. Yet, research on DNA barcoding of non‐flowering seed plants has been less extensive, and the evaluation of DNA barcodes in this division has been limited thus far. Here, we evaluated seven markers (psbA‐trnH, matK, rbcL, rpoB, rpoC1, ITS and ITS2) from non‐flowering seed plants. The usefulness of each region was assessed using four criteria: the success rate of PCR amplification, the differential intra‐ and inter‐specific divergences, the DNA barcoding gap and the ability to discriminate species. Among the seven loci tested, ITS2 produced the best results in the barcoding of non‐flowering seed plants. In addition, we compared the abilities of the five most‐recommended markers (psbA‐trnH, matK, rbcL, ITS and ITS2) to identify additional species using a large database of gymnosperms from GenBank. ITS2 remained effective for species identification in a wide range of non‐flowering seed plants: for the 1531 samples from 608 species of 80 diverse genera, ITS2 correctly authenticated 66% of them at the species level. In conclusion, the ITS2 region can serve as a useful barcode to discriminate non‐flowering seed plants, and this study will contribute valuable information for the barcoding of plant species.     

BREEDING SYSTEM OF A HYBRID BETWEEN A SEXUAL AND AN APOMICTIC SPECIES OF AMELANCHIER,SHADBUSH (ROSACEAE,MALOIDEAE)

Amelanchier bartramiana, which has been shown to be sexual and self-incompatible, and A. laevis, in which apospory and self-compatibility occur, grow in a mixed population in western Maine. Also present there is the putative interspecific hybrid, A. × neglecta. Principal components and canonical discriminant analyses of seven morphological characters from 96 parental and putative hybrid individuals substantiate morphological intermediacy of the hybrids. A Nomarski differential interference microscopy study of megasporogenesis and megagametogenesis in five hybrid plants shows a predominantly aposporous mode of reproduction, and controlled pollinations indicate that seed formation between A. bartramiana and A. laevis is possible and that the hybrids are self-compatible. The hybrids, which are tetraploids (n = ca. 34) like both parents in Maine, produce viable pollen and seed, but there is little evidence for backcrossing with either parent.     

Factors affecting fruit and seed production of Paeonia ostii “Feng Dan”, an economically important oil tree

Many factors may affect reproduction of animal-pollinated species. In this study, the effects of pollen limitation, attractive traits (flower number, plant height and flower width) and flowering phenological traits (flowering onset, duration and synchrony) on female reproduction, as well as the patterns of variation in fruit and seed production within plants, were investigated in Paeonia ostii “Feng Dan” over two flowering seasons (2018 and 2019). Fruit set was very high (90%), and pollen supplementation did not increase fruit and seed production in either year, indicating no pollen limitation. Fruit set, ovule number per fruit and mean individual seed weight per fruit were not affected by any of the six attractive and phenological traits in either year, whereas seed number per fruit was related to the three attractive traits in one or both years. Seed number per plant was positively affected by the three attractive traits and best explained by flower number in both years, but the effect of each of the three phenological traits on seed number per plant differed between years. Within plants, the fruit set, ovule number, seed set and seed number per fruit declined from early- to late-opening flowers, presumably because of resource preemption, but the mean individual seed weight did not vary across the flowering sequence. Our study shows that attractive traits of Paeonia ostii “Feng Dan” are more important than flowering phenological traits in the prediction of total seed production per plant.     

Sequential variation in the components of reproductive success in the distylousJasminum fruticans (Oleaceae)

A 2-year study of three natural populations of the distylousJasminum fruticans showed that mean fruit and seed production were significantly greater in shortstyled plants (thrums) than in long-styled plants (pins). In this study, we investigated the role of four sequential factors which may differentially influence fruit and seed set in the two floral morphs: (1) differences in flowering phenology, (2) a limitation of pollen transfer towards pins, (3) a differential capacity of the two morphs to act through famale and male function and (4) differential fruit abortion in the two morphs. Fruit set was significantly influenced by differences in flowering phenology although there were no differences in flowering time between the two morphs. supplementary pollinations in a natural population significantly increased fruit set and reduced the difference in fruit set between the two morphs in relation to controls, indicating a limitation on pollen transfer which was most severe towards pin stigmas. In reciprocal crosses, seed set was significantly dependent on the paternal and maternal identity of the pin parent. There was no significant variation among thrums for their performance as male or female parent. Furthermore, individual pin plants with relatively high percentage seed set as female parents gave poor seed set as male parents and vice-versa. Whereas fruit removal had no effect on seed number in thrum plants, a greater proportion of viable seeds were produced on pin plants which were left to naturally mature their fruits than on pins which had fruits artificially removed, suggesting the occurrence of selective fruit abortion in pins but not in thrums. The initially greater maternal fitness of thrums due to their greater success as pollen recipients may thus be opposed by increased viable seed set in the pins due to factors acting after the pollination stage. The relative reproductive success of floral morphs in the distylousJ. fruticans is thus differentially influenced by ecological factors occurring at different stages of the reproductive process.     

The effect of a rosette-crown fly, Botanophila turcica, on growth,biomass allocation and reproduction of the thistle Carthamus lanatus

Plant growth and reproductive output of the winter annual invasive thistle, Carthamus lanatus was characterised in relation to plant size in three native populations in southern France. The effects of the rosette-crown feeding fly Botanophila turcica on these plant characteristics were assessed by comparing unattacked with naturally attacked plants at each site and by a field experiment. Indirect effects of B. turcica on plant seed production were also compared with direct seed loss caused by a guild of capitulum-feeding insects that incidentally attacked the marked plants at these sites. C. lanatus showed no size or weight requirement for flowering, but larger flowering plants produced less total receptacle surface and less seed production (female reproductive potential) in proportion to plant weight than smaller flowering plants. B. turcica did not select hosts on the basis of size or density. B. turcica reduced plant relative growth rate (RGR) in all situations, but attacked plants compensated fully at two of three sites as attack failed to halt rosette growth. Attacked plants suffered 12 % mortality, and 71 % lower seed production than unattacked plants at the site with the lowest RGR. This corresponded to 9 % lower seed production for the whole thistle population compared to 8.6–19.5 % direct seed loss to capitulum insects across all sites.     

SPATIAL VARIABILITY IN SEED PRODUCTION OF THE PERENNIAL BUNCHGRASS BOUTELOUA GRACILIS (GRAMINEAE)

The effects of soil texture and grazing by cattle on the production of seeds of Bouteloua gracilis were evaluated for a semiarid grassland in northeastern Colorado. Ten locations were chosen to represent the range in soil textures and grazing intensities found at the Central Plains Experimental Range research site. Number of flowering culms, inflorescences and seeds, length of each flowering culm, total biomass of reproductive structures (culms, inflorescences, and seeds), and basal area were assessed for 96 B. gracilis plants at each location. Community-level estimates of density of flowering culms and density of viable seeds were made for each location. Both soil texture and grazing by cattle were important to spatial variability in seed production and other indicators of reproductive effort by B. gracilis. Grazing was important in mediating effects of soil texture. On locations protected from grazing, soil texture had significant effects on seed production; the largest number of seeds was produced on the coarsest-textured soil and the fewest number on the finest-textured soil. Relationships between seed production and clay content and between seed production and other indicators of reproductive effort by B. gracilis were different for grazed and ungrazed locations. Spatial variability in seed production of B. gracilis as a result of spatial variability in soil texture and grazing may be important to the continued dominance of this species in the presence of disturbances that vary in time and space.     

Regulation of reproduction in wild and cultivatedLycopersicon esculentum Mill. by vesicular-arbuscular mycorrhizal infection

Summary We examined how mycorrhizal infection byGlomus etunicatum Becker and Gerd. affected flowering phenology and components of reproduction in eight wild accessions and two cultivars ofLycopersicon esculentum Mill. We did this by performing a detailed demographic study of flower, fruit and seed production. Mycorrhizal infection had variable effects on the ten accessions. Infection significantly decreased the time taken to initiate flowering in some accessions. In addition, infection increased flowering duration in some accessions. In many accessions, infection significantly increased seed production, primarily by increasing the number of inflorescences and infructescences. In some accessions, mycorrhizal infection also increased the proportion of flowers that produced mature fruits or the number of seeds per fruit. Among accessions, shoot phosphorus content was correlated with seed productivity for both mycorrhizal and non-mycorrhizal plants. However, non-mycorrhizal plants produced more seed biomass per mg of shoot phosphorus than mycorrhizal plants.     

Isolation affects reproductive success in low-density but not high-density populations of two wind-pollinated <Emphasis Type="Italic">Thalictrum</Emphasis> species

Since pollen usually travels limited distances in wind-pollinated plant species, plants growing at low density may become pollen limited. We examined how local pollen availability and population density affect reproductive success in two wind-pollinated, dioecious species, Thalictrum fendleri and Thalictrum dioicum. Distance to the nearest flowering male, the number of flowering males within 2 m, and flower number on those males served as measures of local pollen availability. Increased distance from pollen donors reduced seed set in the lowest-density population of each species, but seed set in high-density populations was not correlated with local pollen availability. For plants in high- and low-density populations at similar distances from pollen donors, this distance only affected seed set in low-density populations. To ensure that differences in resource availability were not causing spurious correlations between seed set and plant density, we constructed low-density artificial arrays in populations of T. dioicum. In these, seed set decreased rapidly with increases in distance from pollen donors. Despite these effects, the density of males in a population was not correlated with average seed set in T. dioicum, and hand pollination in the T. dioicum populations also failed to increase seed set over natural levels. These results suggest that pollen receipt only limits seed set on isolated plants within low- density populations of T. dioicum and T. fendleri.     

Flowering phenology of <Emphasis Type="Italic">Ulex europaeus</Emphasis>: ecological consequences of variation within and among populations

Reproductive phenology of gorse (Ulex europaeus L., Genisteae, Fabaceae) is unusual in that the onset and duration of flowering vary greatly among individuals within populations: some plants initiate flowering in autumn or winter and continue flowering through spring, others initiate flowering in early spring. To understand the origin of this diversity and its ecological consequences, we investigated flowering phenology of randomly sampled individuals from five different natural populations in Brittany (France). Reproductive success was evaluated for individuals with contrasting flowering patterns, from 16 natural populations. Flower production, pod production, seed production and seed predation were estimated. Plants initiating flowering in spring produced larger numbers of flowers and pods over a shorter period than plants flowering from winter to spring, which produced few flowers and pods at a time but over a longer period. Pod production of long-flowering plants did not differ significantly between winter and spring, but their pods were more intensively attacked by seed predators in spring than in winter. We discuss our results in relation to biotic and abiotic parameters. We postulate that long-flowering can be interpreted as a bet-hedging strategy, spreading the risk of pod failure (rotting or freezing) in winter and of seed predation in spring.     

REPRODUCTIVE BIOLOGY OF SEVERAL BROMEGRASSES (BROMUS): BREEDING SYSTEM,PATTERN OF FRUIT MATURATION,AND SEED SET

Five species of Bromus were tested for self-fertility when bagged. Bromus inermis was self-incompatible. Bromus tectorum, B. kalmii, B. ciliatus, and B. latiglumis were capable of self-fertilization. Further evidence from flowering behavior suggested that B. tectorum almost always self-pollinated and that the remaining self-fertile species sometimes outcrossed. Florets in different positions on a spikelet varied in the proportion that set seed. Open-pollinated B. inermis plants had lower seed set than the self-fertile species, in agreement with the pattern in a variety of hermaphroditic plants.     

GENDER VARIATION AND SEXUAL DIFFERENCES IN REPRODUCTIVE CHARACTERS AND SEED PRODUCTION IN GYNODIOECIOUS GERANIUM MACULATUM

Variation in sex expression, flowering pattern, and seed production was studied in the self-compatible perennial herb Geranium maculatum in Illinois and Indiana. In a survey of eight populations, female (male-sterile) plants were found in seven (frequencies ranging from 0.5% to 24.3% [median 4.2%]), and intermediate plants (with partly reduced male function) were found in all populations. Gender variation and sexual differences in reproductive characters were studied in detail in two populations. One population consisted of 5% female, 27% intermediate, and 68% hermaphrodite plants; the other consisted of 1% female, 20% intermediate, and 79% hermaphrodite plants. Females produced smaller flowers and began flowering earlier than hermaphrodites. Intermediates produced flowers of an intermediate size and began flowering as early as females. Females and hermaphrodites did not differ in flower number, vegetative size, flowering frequency, survival, or seed size. However, females produced 1.6 times more seeds than hermaphrodites. Intermediates produced 1.3–1.6 times more seeds than hermaphrodites. Some between-year variation in sex expression was observed. Hand-pollination with outcross pollen produced two to four times as many seeds as hand-pollination with self-pollen. A lower outcrossing rate in hermaphrodites than in females may at least partly explain the lower seed set in hermaphrodites. The higher seed production of females, and possibly the high fecundity of the intermediates, should contribute to the maintenance of this sexual polymorphism.     

Effect of photoperiod during growth of Chenopodium rubrum mother plants on properties of offspring

Using in vitro culture, we determined the effect of photoperiod during growth of Chenopodium rubrum mother plants on vegetative and reproductive development of offspring. Photoperiod during flowering induction of mother plants (the first 6 d after the germination) has the key influence on seed germination and offspring growth, while offspring flowering and seed maturation is determined by photoperiod their mothers experienced during, and shortly after, flowering induction. The mechanism can be through changes in seed protein pattern which we found dependent on photoperiod experienced by mother plants.     

Herbivory and natural selection on flowering phenology in wild sunflower, Helianthus annuus

Plant fitness is strongly affected by flowering phenology, and there are several ecological factors that are thought to shape the distribution of flowering times. One relatively underexamined factor is the timing and intensity of attack by herbivores that feed on flowers or developing seeds. This study tests the hypothesis that herbivores that feed on developing seeds of wild sunflower, Helianthus annuus (Asteraceae), impose selection on flowering phenology. First, the study population was found to contain genetic variation for mean date of flowering, so this trait could evolve if natural selection were operating. Next, the phenological pattern of abundance of five seed-feeding herbivores was documented. Damage by three herbivores, Haplorhynchites aeneus (Cucurlionidae), the head-clipping weevil, Homoeosoma electellum (Lepidoptera: Pyralidae), the sunflower moth, and Suleima helianthana (Lepidoptera: Tortricidae), the sunflower bud moth, was highest early in the flowering season, and declined as the season progressed. Damage by one herbivore, the seed fly Gymnocarena diffusa (Diptera: Tephrididae), was lowest early in the flowering season and increased as the season progressed. Finally, damage by two seed weevils, Smicronyx fulvus and S. sordidus (Curculionidae), whose damage was not distinguished, was constant through the flowering period. Third, damage by Haplorhynchites, Homoeosoma, and Suleima was found to be detrimental to plant fitness, suggesting that plants that flower when these herbivores are not abundant should have higher fitness. Finally, two phenotypic selection analyses were performed. The first included damage by Homoeosoma and Suleima, as well as flowering date, leaf area, and inflorescence diameter, as characters predicting plant fitness. In this analysis directional selection was found to act to decrease damage by the two herbivores, but did not act on flowering date. The second selection analysis was identical except that damage by the two herbivores was not included. In this analysis significant directional selection was found to favor later-flowering plants. Comparison of these two analyses suggests that all selection on flowering phenology is attributable to damage by Homoeosoma and Suleima: plants that flower later avoid damage by these two herbivores. While other influences on flowering phenology, such as pollination, mate availability, and seasonality, have been well documented, this study is one of few to demonstrate natural selection on flowering phenology that is a direct consequence of insect attack. Received: 17 November 1998 / Accepted: 18 July 1999     

CHASMOGAMY/CLEISTOGAMY IN TRIODANIS PERFOLIATA (CAMPANULACEAE): SOME CH/CL COMPARISONS IN FITNESS PARAMETERS

The chasmogamous/cleistogamous (CH/CL) breeding system has evolved independently many times, but its ecological role has been studied in few species. In Triodanis perfoliata, we studied the natural phenology of flowering and seed set, how successful CH flowers were at setting seed, whether seed from CH and CL flowers differed in germination behavior, and whether plants derived from these two seed types differed in vigor. Most notably, we found that the later the CH flowers were produced, the fewer seed they successfully matured. In general the two seed types did not show differences in germination behavior, but for some maternal parents the two seed types differed. For these, the CL seed germinated less readily. Finally, there was some indication that plants from CL seed were not as vigorous as those from CH seed.     

DISPERSION OF SEEDLINGS OF THE PRAIRIE COMPASS PLANT,SILPHIUM LACINIATUM (ASTERACEAE)

Although the dispersal of seeds around individual plants (the seed shadow) has frequently been characterized, the dispersion of seedlings around plants (the seedling shadow) has rarely been examined. We mapped 101 and 149 seedlings of the prairie compass plant (Silphium laciniatum) that appeared in our study area in 1987 and 1990 following mass flowering in 1986 and 1989. We also mapped the locations of flowering stems which appeared in 1986 and 1989 and recorded the number of flowerheads at each stem location. The frequency distributions of distance between a seedling and the nearest flowering stem were identical in the 2 years, with a median distance of 1.0 m. The large size and lack of wind-dispersal structures of compass plant seeds (achenes) are responsible for their limited dispersal. From estimates of the total seed production in the study area in 1986 and 1989, we calculated that about 1% of seeds became seedlings in each year. Flowering stem locations with a higher number of flowerheads had a significantly higher density of seedlings around them. This indicates that recruitment to compass plant populations is not a “lottery”; individual plants that produce more seeds produce, on average, more seedlings.