Temporal patterns of seed set and decelerating fitness returns on female allocation in Zigadenus paniculatus (Liliaceae), an andromonoecious lily

I describe temporal patterns of seed production in the andromonoecious lily Zigadenus particulatus. Fruit set per flower and seed set per fruit declined through time within plants. Hand pollination experiments showed that this was not due to increasing pollen limitation. Nutrient supplementation had little effect on seed output, but leaf clipping reduced seed production, especially in late-blooming flowers, and removal of early-blooming flowers increased seed set by later flowers. Thus, the temporal pattern of seed output was due to declining availability of photosynthates. Plants with larger bulbs produced larger inflorescences, a greater proportion of hermaphrodite flowers, more fruits per hermaphrodite flower, and more seeds per fruit, but lost a greater fraction of their initial bulb mass as a consequence of fruiting. After controlling for the effects of bulb mass, plants with larger inflorescences produced a greater proportion of male flowers, and plants with more hermaphrodite flowers produced fewer fruits per hermaphrodite flower and fewer seeds per fruit. Thus, the female fitness gain curve was decelerating. The temporal decline in seed output provides a partial explanation for the parallel decline in allocation to pistils. However, a complete explanation for the pattern of gamete packaging requires an understanding of factors controlling male, as well as female, fitness.     

EFFECTS OF ENVIRONMENTAL STRESS ON COMPONENTS OF REPRODUCTION IN CLARKIA UNGUICULATA

The effects of stress on reproduction in Clarkia unguiculata were investigated by monitoring flower number, seed number and seed wt after subjecting plants to drought and heavy fruit loads, and by monitoring seed number after pollinating overage flowers. Fewer main-stem flowers were produced in drought stressed plants than in non-stressed plants. Drought stress did not affect overall seed numbers and seed wt in 1981, but did significantly affect seed number and wt in different flower age classes. Overall seed numbers were not affected by drought stress in 1982, but seed wt was significantly lower in plants with stressed ovules. Fewer flowers but more seeds per pollination were produced by plants in which all or ½ the flowers were pollinated than by plants in which ¼ or 1/6 of the flowers were pollinated. Seed wts were lowest in plants with a full fruit load and in fruits produced late in the growth cycle. Seed number was variable but high following pollination of flowers with 1–7 day old stigmas, and much lower in flowers with older stigmas.     

FERTILIZATION DYNAMICS AND PARENTAL EFFECTS UPON FRUIT DEVELOPMENT IN RAPHANUS RAPHANISTRUM: CONSEQUENCES FOR SEED SIZE VARIATION

Seed weight varies significantly within and among fruits of wild radish (Raphanus raphanistrum). To determine sources of this variation, we studied fertilization and seed development following controlled pollinations. Within fruits, central ovules were fertilized prior to distal ovules and attained greater seed size. Ninety-seven percent of the variation in mean seed wt per fruit was explained by an analysis of variance incorporating parental effects, pollination date, and the number of seeds per fruit. We document strong maternal effects on the number of ovules per ovary, the number of fertilized ovules per ovary, the number of seeds per fruit, and mean individual seed wt per fruit. Across females, pollen donor had a slight but significant effect on seed wt; no paternal effects on fertilization rate, zygote number, or seed number per fruit were detected. Within females, with one exception, pollen donor had no significant effect on these components of seed development. Stronger maternal main effects may be due to donor x recipient interactions, cytoplasmic factors, the genetic inequity within triploid endosperm, and/or strict maternal control over resource allocation. The large maternal effects relative to paternal effects may limit the rate at which natural selection acts on paternal traits expressed prior to seed maturation.     

TEMPORAL VARIATION OF FLOWER AND FRUIT SIZE IN RELATION TO SEED YIELD IN CELANDINE POPPY (CHELIDONIUM MAJUS; PAPAVERACEAE)

Ecological and evolutionary studies typically consider variation in single reproductive characters in isolation, without considering how they might be correlated with other reproductive and vegetative characters. In our study, we examined temporal patterns of variation and correlation in flower diameter and fruit length during a reproductive phase in two Massachusetts populations of the herb, Chelidonium majus. We also examined the relationships of such variation to measurements of seed yield components (mean seed weight and number per fruit) and aspects of plant vegetative size. Most of the variation in the sizes of reproductive characters occurred within individual plants, instead of among plants or between populations. Flower and fruit sizes as well as seed number per fruit declined significantly during the season in both populations. Only mean seed size per fruit was relatively stable for individual plants in both populations. Conserving resources by a gradual reduction in the size of reproductive characters over the season may be a strategy for maternal plants to continue seed production. The strong, persistent patterns of correlation between certain characters, such as flower and fruit size, in spite of extensive phenotypic plasticity, was interpreted as indirect evidence for developmental correlation. Furthermore, vegetatively larger plants produced not only more flowers and fruits, but also consistently larger flowers and fruits. The results emphasize that variation in fitness characters, such as seed size and number, should not be viewed in isolation from vegetative characters, flower, and fruit sizes in ecological and evolutionary studies, if the goal is to understand the mechanisms of natural selection in wild populations.     

Effects of fruit thinning on fruit and seed features of <Emphasis Type="Italic">Cistus ladanifer</Emphasis>

Fruit and seed features are the result of reproductive allocation decisions which ultimately depend on both plant availability of resources and total number of developing fruits. In this study, we manipulated fruit load in Cistus ladanifer plants by removing 0, 25 or 75% of developing fruits. Fruit features (total fruit weight, fruit-wall weight, total seed weight per fruit and seed number per fruit) were unaffected by fruit thinning, nevertheless mean seed weight increased in treated plants independently of thinning intensity. This reduced compensation was unrelated to plant size and had no consequences on fruit predation by insects. From these results it could be suggested that not only availability of resources but also morphological constraints could affect seed size in Cistus ladanifer. On the other hand, this change in seed weight could have important consequences since in this species heavy seeds perform better after fire events but light ones are advantageous in between fire recruitments.     

Fruit removal and postdispersal survivorship in the tropical dry forest shrub Erythroxylum havanense: ecological and evolutionary implications

We studied the relationship between the removal rate and the spatiotemporal availability of ripe fruits of the tropical deciduous shrub Erythroxylum havanense in western Mexico. We also evaluated the effects of dispersal on seed survival during the first stages of establishment. Fast and early dispersal should be favored in E. havanense, since propagules have more time to grow and accumulate resources before the beginning of the severe dry season. In general, high rates of fruit removal imply faster and earlier dispersal. Thus, plants producing large crops should benefit from high removal rates, which will increase the probability of successful establishment by their progeny. To characterize both individual and population fruiting patterns, we made daily counts of fruits on 51 plants arranged in six clumps of different sizes. The daily number of fruits removed per plant was higher for plants with larger initial crop sizes and larger numbers of ripe fruits on a given day, but decreased as clump size increased. Additionally, we monitored postdispersal survival and germination in an experiment manipulating seed density, distance from adult plants, and seed predation. Early establishment was independent of density or distance, and vertebrate seed predation was the main agent of seed mortality. Our results indicate that the critical variable with respect to fruit removal is the number of fruits a plant produces, large plants having higher dispersal rates. Large plants are also more likely to have more seeds escaping postdispersal seed predation.     

Factors regulating fruit and seed production in the desert annual Lesquerella gordonii

Summary This study examines the pattern of reproduction and factors that limit the maternal reproductive output in the self-incompatible annual, Lesquerella gordonii. Greenhouse experiments showed that fruits do not abscise and they mature if they contain at least one fertilized (outcrossed) ovule. Field studies showed that flowering increased to a peak and then fell off rapidly, while seed production/fruit decreased through the season. Pollen addition did not increase the number of fruit or seeds per fruit, suggesting that fruit and seed production are not pollinator limited. Addition of water increased fruit and seed production by extending the life of the plant, by increasing the number of stalks which flowered, and by increasing the number of seeds per fruit. This indicates that fruit and seed production are resource limited. Fruits are relatively inexpensive as compared to seeds as over 93% of the water in fruits is contained in the seeds. Fifty-three percent of the plants were lost to herbivory as were 64.6% of the fruits on the remaining plants, indicating that herbivory also limited fruit and seed production. It is hypothesized that high fruit set coupled with variable seed set in L. gordonii is an adaptive response to unpredictable, variable resource levels, and high herbivory risk.     

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.     

Impact of prairie fragment size on proportion of females and reproductive success of Lobelia spicata Lam., a gynodioecious species

• Due to ongoing human impacts, plant species increasingly occur in landscapes that are highly fragmented, with remaining natural habitats occupying small areas, resulting in populations that are smaller and more isolated than in previous time periods. This changed metapopulation structure is expected to have negative impacts on seed production. For example, the proportion of female plants within gynodioecious populations may be more volatile due to genetic drift in small populations associated with small habitat fragments, with concomitant impacts on seed production. My aims were to determine: (i) if variation in proportion of females is larger in smaller fragments; and (ii) if such changes in female frequency in small fragments result in reduced seed production.
• Thirty‐two populations of Lobelia spicata Lam., a gynodioecious species, were surveyed in 2000, 2001 and 2009 in the tallgrass prairie region of Midwestern North America (Illinois and Indiana, USA). Data were collected for: proportion of female plants, total number of flowering plants (measure of population size), seed set per plant and prairie fragment size (another measure of population size).
• The proportion of females is more variable in smaller prairie fragments. Seed number per fruit decreases as the proportion of females increases in a population, but only significantly for female plants. The number of flowering plants is positively associated with fruit production for both genders. Populations within larger prairie fragments have higher seed production.
• The reproductive consequences of habitat fragmentation depend on the plant breeding system. While both sexes were negatively impacted, females were more adversely affected.

     

Gynodioecy as a possible populational strategy for increasing reproductive output

Summary Cytoplasmic male steriles occur regularly in wild populations of the annual crucifer Hirschfeldia incana Lagr.-Foss. in Israel. In these plants numbers of ovules per flower, numbers of seed per fruit, and total seed weights per plant are slightly higher than in the hermaphrodites with which they mate. Yet their frequencies in wild populations do not exceed 2–10 per cent. There are no signs of incipient dioecy. The species is self incompatible and no mechanisms to enforce outcrossing are needed. It is argued that in this and similar cases gynodioecy functions as a pollen saving measure. Precise pollen presentation in the flower renders some of the pollen redundant and facilitates its abolition in a sector of the population. It is possible that the enhanced seed fecundity of the pollenless sector stems from a greater availability of plant resources for seed production in the unisexual than in the bisexual seed parent.     

The role of the tolerance–fecundity trade‐off in maintaining intraspecific seed trait variation in a widespread dimorphic herb

Coexistence of species with different seed sizes is a long‐standing issue in community ecology, and a trade‐off between fecundity and stress tolerance has been proposed to explain co‐occurrence in heterogeneous environments. Here we tested an intraspecific extension of this model: whether such trade‐off also explains seed trait variation among populations of widespread plants under stress gradients. We collected seeds from 14 populations of Plantago coronopus along the Atlantic coast in North Africa and Europe. This herb presents seed dimorphism, producing large basal seeds with a mucilaginous coat that facilitates water absorption (more stress tolerant), and small apical seeds without coats (less stress tolerant). We analysed variation among populations in number, size and mucilage production of basal and apical seeds, and searched for relationships between local environment and plant size. Populations under higher stress (higher temperature, lower precipitation, lower soil organic matter) had fewer seeds per fruit, higher predominance of basal relative to apical seeds, and larger basal seeds with thicker mucilaginous coats. These results strongly suggest a trade‐off between tolerance and fecundity at the fruit level underpins variation in seed traits among P. coronopus populations. However, seed production per plant showed the opposite pattern to seed production per fruit, and seemed related to plant size and other life‐cycle components, as an additional strategy to cope with environmental variation across the range. The tolerance–fecundity model may constitute, under stress gradients, a broader ecological framework to explain trait variation than the classical seed size–number compromise, although several fecundity levels and traits should be considered to understand the diverse strategies of widespread plants to maximise fitness in each set of local conditions.     

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.     

Maturation characteristics of Rubus pennsylvanicus fruit: are black and red the same?

Summary Fruit of the blackberry, Rubus pennsylvanicus Poir. (Rosaceae), were examined to determine variation in maturation characteristics. Maturation timing and rate varied greatly among individual fruits, resulting in a bi-colored fruiting display comprised largely of two maturation stages, pre-ripe (salmon and scarlet) and ripe (dark brown and black). While ripe fruit were generally larger and heavier than pre-ripe fruit, exhibiting greater fresh and dry fruit weight, diameter, water content, and total seed weight, no significant differences were found in energy content, i.e. numbers of calories per gram pulp, or in pulp:seed ratio. The differences between ripe and pre-ripe fruit appear to be due largely to an increase in water content and seed weight with maturity. The fact that little energetic benefit accrues to the preferential selection of ripe fruit suggests that bi-colored Rubus displays may be considered to be unicolored.     

Effects of burial in sand on the growth and reproduction of Cakile edentula

Studies were conducted to examine the effects of sand burial on the growth and reproduction of Cakile edentula under controlled greenhouse conditions Sand burial enhanced both the vegetative growth and reproduction of plants Within one week after burial, plants showed a significant increase in the length of the internode just below the current sand surface This response was not mediated by the dark conditions imposed by sand burial About two weeks after burial, adventitious roots grew upwards into the deposited sand Plants buried in washed sand did not differ in their dry weight whereas those buried in unwashed sand were significantly taller, produced greater biomass and larger number of seeds per plant than control However, the mean mass per fruit and per seed were not altered The frequency distribution of fruit and seed was only slightly modified by the burial treatments     

Plant interactions for pollinator visits: a test of the magnet species effect

Summary From 1985–1987, patterns of fruit and seed set were studied in a population of mayapple (Podophyllum peltatum), a clonal, self-incompatible herb found in deciduous woods in eastern North America. Mayapple flowers do not produce nectar, but depend on infrequent visits by nectar-seeking queen bumble bees for pollination. In all years female reproductive success in mayapple colonies was influenced by colony size (number of flowers), by the distance to neighbouring colonies and by proximity to lousewort plants (Pedicularis canadensis), a prolific nectar producer heavily visited by bumble bees. In all years fruit and seed set were greater in mayapple colonies <25 m from lousewort flowers than in matched colonies which were >50 m from lousewort. In 1985 and 1987 the frequency of queen bumble bee visits to flowers in colonies close to lousewort was about four times greater than to distant colonies. In 1986 I removed about 80% of lousewort flowers to test whether the enhanced fruit and seed set in mayapples close to lousewort was pollinator mediated. Mayapple colonies close to flowerless lousewort patches did not differ in fruit or seed set from matched colonies >50 m from lousewort. In contrast, mayapples close to flowering lousewort patches had greater fruit and seed set compared with distant colonies. Over all years, a larger proportion of mayapples close to flowering lousewort patches had enhanced fruit and seed set compared with colonies close to louseworts without flowers. Though rarely documented, this type of facilitative interaction between plants that are highly attractive to pollinators (magnet species), and co-flowering species that are rarely visited by pollinators, may be widespread in plant communities.     

Ecological correlates of seed mass variation in Phoradendron juniperinum,a xylem-tapping mistletoe

Summary We investigated several ecological correlates of seed mass variation in the hemiparasitic, xylemtapping mistletoe, Phoradendron juniperinum. Mean seed mass varied two-fold among plants between the ages of 4 and 14 years old and was positively correlated with parental plant age. Both the standard deviation and the coefficient of variation in mean seed mass decreased with increasing plant age demonstrating that, on average, younger plants produced seed with more variable mass. Nitrogen concentrations (mg nitrogen per gram of seed) of both the seed and fruit (pericarp) were not correlated with mass or the age of the parent plant from which the seed was taken. However, the nitrogen content per seed (mg nitrogen per seed) was positively correlated with the mean seed dry mass and the age of the seed parent, suggesting that the carbon to nitrogen ratio of individual seeds remained relatively constant as seed mass increased and plants grew older. Seed germination ranged between 20% and 86% and was positively correlated with mass and parental plant age. Heavier seeds (seeds from older plants) also had the highest root radicle growth rates. Furthermore, the final root radicle length after 76 d of growth was positively correlated with seed dry mass. When grown on a medium containing an extract prepared from the host plant foliage, all seeds showed lower germination, grew more slowly and had shorter overall root radicles, but had significantly greater development of the haustorial disks (the holdfast which forms the host-parasite junction in Phoradendron) than seeds grown on a control medium. Our results suggest that, on average, seeds of greater mass produced by older plants have a greater total resource pool per propagule (fruit + seed). This resource pool may be important in conferring a greater potential for dispersal (fruit), survival, colonization, and establishment.     

REPRODUCTIVE CAPACITY AND SEED SIZE IN LUPINUS TEXENSIS

The average number of ovules produced per individual of Lupinus texensis is much greater than the average number of seeds per plant. Each plant produces approximately 2,000 ovules but only 2.5% develop into seeds. One fourth of the seeds is lost due to abortion and 0.3% is lost due to predation on the plant. Mature seeds from this population exhibit a five-fold range in weight, from 10 to 56 mg. The distribution of seed weights in the field population is skewed and leptokurtic. Seed wt is positively correlated with both seed germination and seedling survivorship. Heritability of seed wt is 0.09. There is no correlation between average seed wt per plant and total number of seeds per plant, seeds per pod, or legumes per plant.     

Clumped planting arrangements improve seed production in a revegetated eucalypt woodland

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

Seed size selection by olive baboons

Seed size is an important plant fitness trait that can influence several steps between fruiting and the establishment of a plant’s offspring. Seed size varies considerably within many plant species, yet the relevance of the trait for intra-specific fruit choice by primates has received little attention. Primates may select certain seed sizes within a species for a number of reasons, e.g. to decrease indigestible seed load or increase pulp intake per fruit. Olive baboons (Papio anubis, Cercopithecidae) are known to select seed size in unripe and mature pods of Parkia biglobosa (Mimosaceae) differentially, so that pods with small seeds, and an intermediate seed number, contribute most to dispersal by baboons. We tested whether olive baboons likewise select for smaller ripe seeds within each of nine additional fruit species whose fruit pulp baboons commonly consume, and for larger seeds in one species in which baboons feed on the seeds. Species differed in fruit type and seed number per fruit. For five of these species, baboons dispersed seeds that were significantly smaller than seeds extracted manually from randomly collected fresh fruits. In contrast, for three species, baboons swallowed seeds that were significantly longer and/or wider than seeds from fresh fruits. In two species, sizes of ingested seeds and seeds from fresh fruits did not differ significantly. Baboons frequently spat out seeds of Drypetes floribunda (Euphorbiaceae) but not those of other plant species having seeds of equal size. Oral processing of D. floribunda seeds depended on seed size: seeds that were spat out were significantly larger and swallowed seeds smaller, than seeds from randomly collected fresh fruits. We argue that seed size selection in baboons is influenced, among other traits, by the amount of pulp rewarded per fruit relative to seed load, which is likely to vary with fruit and seed shape.     

Effects of experimental manipulation of inflorescence size on pollination and pre-dispersal seed predation in the hummingbird-pollinated plant Ipomopsis aggregata

Large floral displays should theoretically provide advantages to plants through increased pollinator visitation and resulting fruit and seed set. However empirical tests of the response of pollinators to floral display size have been limited by a lack of direct experimentation, and the results of such studies have been equivocal. In addition, other selective agents such as pre-dispersal seed predators might modulate effects of floral display on pollination. By artificially altering flower number, we examined the direct effects of floral display in the monocarpic herb, Ipomopsis aggregata (Polemoniaceae), on visitation rates by broad-tailed and rufous hummingbird pollinators, as well destruction of fruits by a pre-dispersal seed predator (Hylemya: Anthomyiidae). In addition, we quantified the ultimate effects of flower number on female reproductive success. Plants with larger floral displays were most likely to be visited first in any given foraging bout (P < 0.01). As expected, plants with more flowers received more total flower visits. However, we found no gain in the proportion of flowers visited for many- versus few-flowered plants, or the total number of approaches/hour. In fact, a significantly greater percentage of flowers were visited on few-flowered plants. Plants did not compensate for our reduction in flowers by increasing investment in the number or proportion of flowers that set fruit, the number of seeds/fruit, or seed weight. Pre-dispersal seed predation was greater for many- than for few-flowered plants (P < 0.001), but this did not offset the potential fitness gains of producing large displays. Our data support the hypothesis that large floral displays function primarily in long-distance attraction of pollinators, and enhance maternal success. Received: 21 March 1996 / Accepted: 24 October 1996