Population dynamics of the shrub Acacia suaveolens (Sm.) Willd.: Seed production and predispersal seed predation

Seed production and predispersal seed predation in the shrub Acacia suaveolens were examined over 3 consecutive years in eight populations in south-eastern Australia. Seed-crop sizes varied both between and within populations of different ages. Seed production was maximal in the first one to four flowering seasons after establishment, and then declined with plant age. The size of the annual seed-crop was also influenced by rainfall for that year. Predispersal seed predation varies between populations over fruiting seasons with the initial large seed-crops resulting in predator satiation. Within one fruiting season, no significant variation in the extent of predispersal seed predation was found in any of three populations studied. There were two major forms of predispersal seed loss: toss of whole fruits to Melanterius corosus (Coleoptera: Curculionidae) and external insect seed grazers, and loss of individual seeds within fruits to M. corosus. Exclusion experiments showed that seeds lost to these predispersal seed predators would otherwise have been matured by the parent plant. The effects of predispersal seed predation cannot be directly related to seedling recruitment. Indirectly, such predation may influence the dispersion of seeds in the soil profile and hence, subsequent recruitment.     

Morphology and phenology of seeds and whole fruit eaten by Milne-Edwards’ sifaka,Propithecus diadema edwardsi,in Ranomafana National Park,Madagascar

Although it is an anatomical folivore, the diet of the Milne-Edwards’ sifaka (Propithecus diadema edwardsiA. Grandidier, 1871) in Ranomafana National Park contained 35% seeds, 30% whole fruit, and 28% leaves. Plant species used as seed sources differed from those used as whole fruit sources in terms of temporal variation in consumption, taxonomic affiliation, morphology, and phenology. Although seeds were destroyed in both exploitation styles used by the sifakas—seed and whole fruit-eating—the gross morphology of species used as seed sources conformed to the complex of traits typical for fruits experiencing seed predation, while species used as whole fruit sources conformed to traits typical for fruits that do not experience predispersal predation. Many of the 19 plant species from which the seed was extracted and eaten contained a single seed with moderate testa thickness, and fruits containing this type of seed were medium-sized with dry or fibrous flesh, moderate skin thickness, and a dull color. In contrast, brightly colored, juicy fruits with minimally protected seeds were characteristic of the 38 plant species from which both pericarp and seed were eaten. Compared to transectwide measures of fruit availability or patterns restricted to whole fruit sources, fewer species of seed sources produced fruit per month and fruiting activity was more seasonal.     

Multiple infestation by seed predators: the effect of loculate fruits on intraspecific insect larval competition

Many morphological features of fruits are important factors affecting predispersal seed predation by insects. This paper analyses the predispersal seed predation process of a major predator (a Noctuidae lepidopteran larvae) in loculate fruits of a bushy perennial plant, Cistus ladanifer. The main aim of the study is to assess the potential effect of internal valvae (which partition groups of seeds) in the intraspecific competition between larvae in multiple-infested fruits.Our results show that larvae do not reject already infested fruits, but they avoid the proximity of other larvae within the fruit, keeping an average minimum distance of one locule. In multiple-infested fruits, larval mortality increases and the proportion of seeds consumed by each larvae decreases. In those situations in which valvae keep apart larvae within a fruit, these only suffer the cost of exploitation competition with a low acquisition of resources. However, when all valvae between them are pierced by the larvae, competition switches to an interference component and larval mortality increases markedly. The existence of valvae within a fruit allows larvae to diminish the cost of intraspecific competition, obtaining high life expectancies (70%), even in triple-infested fruits.     

Equal and Opposite Effects of Floral Offer and Spatial Distribution on Fruit Production and Predispersal Seed Predation in Xanthosoma daguense (Araceae)1

Inflorescences of the terrestrial aroid Xanthosoma daguense in the Andes of Colombia are visited by Dinastinae and Nitidulidae beetles. Plants produce one inflorescence at a time, which is pollinated during the first night of opening. Dynastine beetles act as pollinators, whereas Nitidulids lay eggs in the inflorescence and the larvae damage the seeds. We explored the effects of floral offer and distance among inflorescences on the number of pollinator visits, fruit production, and predispersal seed predation. Number of Dynastine visits per inflorescence tended to increase with increasing distances among inflorescences, but fruit predation increased when inflorescences were more clumped. Both pollinator visitation rates and predispersal seed predation were low at high floral offer. Fruit set increased when inflorescences were visited by two or more Dynastines, but the proportion of fruits damaged by Nitidulid larvae was equivalent to the increase in fruit production due to more Dinastine visits. The net result was a similar number of undamaged fruits in all infructescences produced, independent of the number of Dinastine visits. Our results revealed that both pollinators and predators responded to the number of available inflorescences and their spatial distribution, but they had opposing effects on the infructescences. Thus, our study suggests that the interaction of two ecological processes, pollination and predispersal seed predation, may cancel each other's effects under natural conditions.     

EFFECTS OF POLLEN AND RESOURCES ON SEED NUMBER AND OTHER FITNESS COMPONENTS IN AMELANCHIER ARBOREA (ROSACEAE: MALOIDEAE)

Most Amelanchier arborea flowers have 10 ovules, but the number of filled (embryo-containing) seeds per fruit is usually less than 10 and is highly variable within each individual plant. Because fruit developmental time correlates with seed number, this variation in seed number results in asynchronous fruit ripening. Field experiments tested whether seed number was pollen- or resource-limited. Manipulation of resources in the shoot at the time of fruit initiation by defoliation, girdling, fruit thinning, or foliar feeding had no significant effect on seed number per fruit, although fruit set and seed weight were affected. Supplemental cross-pollination also had no demonstrable effect on seed number. Most ovules that do not become filled seeds are visible as small “undeveloped seeds”; these are not necessarily aborted seeds as this was also the fate of ovules in unpollinated carpels. Alternative hypotheses for the determination of seed number are proposed and discussed.     

Pollination intensity and potential seed set in Passiflora vitifolia

Summary Initial seed set and fruit set were pollen-limited in a Costa Rican population of Passiflora vitifolia, a self-incompatible species with 200–350 ovules per flower. Pollination intensity was measured by counting the number of allogamous pollen grains on stigmas of the large one-day flowers. Hand-pollinations demonstrated that 25–50 pollen grains are required for fruit set, and >450 are needed for maximum seed set, with a pollen:seed ratio of about 1.6:1.0. Hummingbirds (Phaethornis superciliosus) delivered sufficient allogamous pollen for maximum seed set to only 28% of the flowers examined. Naturally pollinated flowers yielded fewer fruits and fewer seeds per fruit than those pollinated by hand. Most pollen transferred by humming-birds was self-incompatible; emasculated flowers yielded higher seed set than flowers with intact anthers. Visitation rates did not provide a good index of effective pollination.There were significant differences in ovule number, maximum seed set, and maximum per cent seed set among individual vines. More than half of an individual's flowers failed to set fruit, whether pollinated by birds or by hand. In this population, maximum reproductive potential may be limited by maternal resources for fruit development, but seed set varies with pollination intensity. Pollen-limited seed set may be a disadvantage of self-incompatibility, especially in species with many-seeded fruits.     

REPRODUCTIVE ECOLOGY OF JEFFERSONIA DIPHYLLA (BERBERIDACEAE)

The reproductive ecology of Jeffersonia diphylla (L.) Pers. (Berberidaceae) was investigated by studying its breeding system, ovule production, seed set, seed dispersal by ants and seed predation by rodents. This species flowers early in the spring and is facultatively autogamous. In a typical year fruit and seed set is high (90%), however, freezing temperatures from late spring frosts in 1983 and 1985 resulted in low fruit set (7% and 20%, respectively), and reduced seed set in those flowers that produced fruit. No differences in seed set between selfed and outcrossed flowers were observed over a two-yr period (1983–84). Ovule number per capsule increased with plant size as measured by leaf number. Seed set and seed wt were unaffected by leaf number unless leaves were removed after flowering was initiated. Jeffersonia diphylla is myrmecochorous. Ants removed seeds faster when seeds were placed in areas where J. diphylla plants were absent, suggesting that dispersal within J. diphylla populations is ant limited. Moreover, fresh (1 day old) seeds were removed by ants faster than 3 day old seeds. Seed predation by rodents prior to dehiscence from capsules is heavy in large populations (85–90%), and apparently negligible in small populations. Predation of seeds that are released from capsules is heavy (approx. 66%), particularly at night. Overall, seed predators consume about 96% of the seed crop in well established populations, but probably much less in small young populations. Hence, seedling recruitment is likely to be higher in small populations, whereas ramet production from rhizomes is the primary mode of propagation in large ones. The evolution of autogamy, early flowering, and myrmecochory are discussed in light of the results of this study.     

Patterns of fruit and seed production inBauhinia ungulata (Leguminosae)

Patterns of seed and fruit production ofBauhinia ungulata, a small tree legume indigenous in tropical America, were studied in Costa Rica. Only about 8% of flowers produced fruits. The average pod had 19 ovules and about two thirds of these began seed development, with mature pods containing an average of 9.7 mature undamaged seeds. About half of the mature pods were damaged by herbivores and within these, 27% of ovules or seeds had been eaten. Among trees there was no significant variation in pod production, but the number of ovules per pod and seed production per pod varied significantly. Within infructescences most pods were retained at middle positions. Within pods, the probability of an ovule developing into a seed increased toward the distal end. The pattern of seed and fruit production in this species agrees well in general with that reported for other neotropical legumes. The abortion of seeds and fruits can be regarded as a way of controlling maternal investment, and as a response to herbivory.     

Effects of resource availability, and fruit and ovule position on components of fecundity in Alliaria petiolata (Brassicaceae)

In four field and glasshouse experiments designed to alter the supply of resources through manipulation of nutrients, root tissue, leaf area and fruit number in Alliaria petiolata (Brassicaceae), more than 99% of ovules per plant showed signs of fertilization, suggesting that seed production in this plant was not pollen limited. However, in all treatments a significant proportion of fruits and seeds did not develop to maturity. Total fruit and seed production did not differ significantly from controls when plants were given nutrient supplements at flowering. Removal of 50–75% of the root tissue in 1-yr-old plants significantly reduced fruit set, but had no effect on individual seed development. Removal of cauline leaves significantly reduced most measures of fruit and seed production, suggesting that current photosynthate is critical for fruit and seed filling. Seed maturation was significantly affected by both fruit position within an infructescence and ovule position within a fruit. Basally located fruits and ovules (within fruits) developed more mature seeds than distally positioned fruits and ovules. Plants responded to removal of basal fruits by re-allocating resources to distal fruits that would normally have aborted. Our results suggest that fruits and seeds act as reproductive sinks competing for parental photosynthate. Patterns of resource allocation within infructescences and fruits were also modified by our experiments.     

Flowering phenology and female fitness: impact of a pre-dispersal seed predator on a sexually polymorphic species

In order to produce seeds, animal-pollinated plants must flower synchronously with and be attractive to their pollinators while avoiding antagonists. Here, we explore temporal and inter-individual patterns in pollination and pre-dispersal seed predation of Dianthus sylvestris by Hadena moths, within and among three sex morphs. We scored plants that started flowering at different periods in 2001 and 2003 and found that fruit set decreased and predation rates increased over one season, and most of the other season, granting a female reproductive advantage to early flowering plants, though, we found no morph-specific temporal patterns. Female plants set more fruits, and more of their fruits escaped predation in one year, but this did not grant them a reproductive advantage since they produce fewer flowers per plant than the other morphs. Instead, mixed plants showed a clear female reproductive advantage. We also examined predation types by Hadena and seed production in attacked and intact fruits of individually marked flowers. Though female Hadena moths laid eggs preferentially into perfect flowers, flower sexes suffered similar predation by itinerant caterpillars. Attacked fruits contained fewer and lighter seeds than un-attacked ones. We conclude that pre-dispersal seed predation by Hadena may select on flowering onset of this sexually polymorphic species.     

Do Meyer’s Parrots Poicephalus meyeri benefit pollination and seed dispersal of plants in the Okavango Delta,Botswana?

Recent studies of new world parrots repeatedly document, with few exceptions, that parrots are wasteful and destructive predispersal seed predators that are unlikely to contribute towards pollination and seed dispersal. Few detailed studies, however, have assessed the contribution of African parrots to forest ecology by quantifying the potential net benefit of seed and flower predation by parrots for most tree species in their diet. Due to the incidence of pollen on the heads of Meyer’s Parrots when feeding on Leguminosae flowers and the dispersal of viable seeds to the ground during seed predation, we compared destruction rates, when feeding on pods, fruits and flowers, with dispersal rates of viable seeds to the ground and frequency of head contact with reproductive apparatus to estimate net benefit from Meyer’s Parrot feeding activity. Meyer’s Parrots were not implicated in endo‐ or epizoochory, but they dropped uneaten fruit pulp and seeds to the ground during feeding bouts, thus providing ripe, undamaged seeds to secondary seed dispersers. This link with forest recruitment was weak, as all tree species utilized by Meyer’s Parrots either had more significant primary dispersal agents or were primarily wind‐dispersed. In most cases, the negative effect of seed predation outweighed any positive effects in terms of dispersal, whereby almost three times more seeds were consumed or destroyed than were dispersed to the ground. Significantly, only Sclerocarya birrea caffra recorded marginal net dispersal benefit from utilization by Meyer’s Parrots. Due to low relative resource abundance and high destruction rate, feeding activity on Berchemia discolor may be significant enough to influence its spatial distribution and abundance. Utilization of flowers of Kigelia africana and Adansonia digitata by parrots likely had a significant negative impact on pollination. Feeding on Acacia nigrescens flowers, however, was potentially advantageous to their pollination. We conclude that Poicephalus parrots are net consumers of ripe, undamaged seeds and flowers, thus having an overall negative impact on forest recruitment in subtropical Africa.     

Temporal Variation in Seed Predation by Insects in a Population of Syagrus romanzoffiana (Arecaceae) in Santa Catarina Island, SC, Brazil

Insect seed predation may vary depending on seed production. The present study considers the hypothesis that the rates of seed predation tend to be smaller in years of higher fruit production. Thus, we monitored the production of fruits and predation of seeds of the palm Syagrus romanzoffiana over 2?years in the Atlantic Forest (Parque Municipal da Lagoa do Peri, Florianópolis, SC, Brazil), between July 2006 and June 2008. Plots of 0.25?m² were fitted under 20 mother plants and fruits were monthly collected for assessment of abundance and seed predation. There was variation in fruit production between the 2?years and among reproductive plants. Predation rates were high and occurred in the predispersal phase by the Curculionidae Revena rubiginosa Boheman, Anchylorhynchus aegrotus Fahraeus, and Anchylorhynchus variabilis Gyllenhal. Seed predation by these species of Anchylorhynchus is first registered in the present study. In average, about 60% of the seeds monthly produced in the population tend to escape insect predation in year of high or low production, becoming available for recruitment. The predation rate was not related to the amount of fruits produced per reproductive plant. Also, different than expected, there was a positive relation between the rates of seed predation and the total of fruits produced monthly on the plots. Thus, no evidence for the satiation of insect seed predators was found in this study with S. romanzoffiana.     

Predation of fruit and seed of Aloe pretoriensis – A little known effect on reproductive output in aloes

The effect of seed predation by phytophagous/parasitoid wasps on the reproductive output of aloes is relatively unknown. In this study, conducted at a nature reserve in Pretoria East, South Africa, the range of insects utilising Aloe pretoriensis (Asphodelaceae) fruits and/or seeds and the impact of this usage on its reproductive output were investigated. Using a GLMM, we explored the effects of morphological features (e.g. floral display size) and selected ecological factors (viz. distance between the aloes and conspecifics and other surrounding vegetation) on fruit utilisation and seed predation. A variety of insect visitors to A. pretoriensis (mainly bees and wasps) were identified including a number of seed predators and parasitoids. Evidence of phytophagy in dissected flowers showed increasing evidence of fruit utilisation and seed predation over an 8‐week period. Emergence boxes with infructescences revealed a range of insect phytophages (and their associated parasitoids) in the aloe fruits and seeds: the drosophilid fly, Apenthecia and six species of wasp – five of them associated with ultilisation of aloe fruits/seeds for the first time: Eurytoma aloineae (Chalcididae), Mesopolobus sp., Pteromalus sp., and c.f. Chlorocytus in the Pteromalidae, Bracon sp. (Braconidae) and Pediobius (Eulophidae). Fruit set ranged between 48% and 93%, with an average of 76%, while average percentage utilisation of fruits was 29%, ranging between 7% and 68%. Average seed set was 23 seeds per fruit and average percentage seed predation 21% (range: 0–51%). Fruit utilisation was found to be significantly negatively correlated with distance to the nearest flowering bush (usually Helichrysum kraussii), but display size did not significantly affect fruit utilisation, nor did distance to conspecifics. Aloe pretoriensis thus serves as host to a variety of phytophagous insects and their associated parasitoids, which impacts considerably on its reproductive output with possible implications for the future conservation of this aloe species.     

The reproductive biology of a distylous tree, Sarcotheca celebica (Oxalidaceae) in Sulawesi, Indonesia

Sarcotheca celebica is a tree endemic lo Sulawesi (and Kabaena Island). Kike many species in the Oxalidaceae it is heterostylous (here distylous). The small red flowers, which last less than one day, are produced in loose inflorescences which bloom over a long flowering season (several months). There are two whorls of stamens in each plant, which overlap in length, perhaps indicating a tristylous origin. Long–styled and short–styled plants differed in the measurements of stamens, styles and pollen grains. Only a small amount of pollen was produced by long–styled plants. Effective insect pollinators were mainly large Hymenoptcra, especially Xylocopa species, which visited many flowers and different trees in rapid succession. A wide diversity of insects visited flowers of both morphs, and numbers visiting each morph were approximately equal. The only successful experimental pollinations were from pollen of short–styled plants onto stigmas of long–styled plants. Automatic self–pollination did not occur. In nature not all flowers produced fruit, some fruits were set with no seeds and a low number of seeds was set in all fruits. The differences in seed set per fruit between short–styled, long–styled and experimental crosses indicate that pollinator visits were insufficient for maximum seed set in some fruits, but selective abortion of seeds must also be occurring. We suggest that S. celebica may represent an intermediate stage in the evolution of dioecy from distyly, with the short–styled flowers making their major genetic contribution through pollen and long–styled flowers through ovules.     

Pollen and Resource Limitation in Veratrum nigrum L. （Liliaceae）, an Andromonoecious Herb

Pollen limitation and resource limitation were invoked to account for the pattern that flowering plants produce more flowers and ovules than fruits and seeds. This study aimed to determine their relative importance in Veratrum nigrum, a self-compatible, perennial, andromonoecious herb. In order to determine whether female production was limited by pollen grains on stigmas or by available resources, we performed supplemental hand pollination in three populations, male-flower-bud removal in three other populations, and emasculation of hermaphroditic flowers in still another population, resulting in a total of seven populations experimentally manipulated. Across the three populations, supplemental hand pollination did not significantly increase fruit set, seed number per fruit, and total seed production per individual, nor did emasculation of hermaphroditic flowers. Taken together, our results suggest that pollen grains deposited on stigmas were abundant enough to fertilize all the ovules. Male-flower-bud removal significantly increased the mean size of hermaphroditic flowers in all three populations. Female reproductive success was increased in one population, but not in the other two populations possibly due to heavy flower/seed predation. We concluded that the female reproductive success of V. nigrum was not limited by pollen grains but by available resources, which is consistent with Bateman＇s principle. Furthermore, the female reproduction increase of male-flower-bud removal individuals might suggest a trade-off between male and female sexual functions.     

Reproductive biology ofCistus ladanifer (Cistaceae)

The phenology, major floral characteristics, fruiting levels, and breeding system ofCistus ladanifer L. (Cistaceae), a common western Mediterranean shrub species, were studied in a southern Spanish population. The white, large (64 mm in diameter) flowers of this shrub appear during spring (March–May) and produce abundant pollen and nectar. In the year of study, flowers lasted up to three days, during which they were visited by a diverse array of insects including beetles, flies, and bees. Hand-pollinations revealed that flowers do not set any seed unless cross pollen is applied to the stigma. Microscopical observations indicate that self pollen tubes grow down the stigma but invariably fail to induce fruit maturation. At the plant level, all estimates of fecundity investigated (number of seeds per capsule, proportion of ovules developing into seed, and proportion of flowers setting fruit) were highly dependent on nearest neighbour distance, with isolated plants setting as little as 0% fruit. In contrast, plants within a clump often transformed into fruit as much as 90% of the flowers. At the population level, seed output was estimated to range between 3,000 and 270,000 seeds per plant during 1991.     

Seed predation of Copaifera langsdorffii (Fabaceae): a tropical tree with supra‐annual fruiting

Seed predation is an important ecological and evolutionary force that directly affects the distribution of plant species. Copaifera langsdorffii is a tropical tree species with supra‐annual fruiting, which has its seeds predated by a specialist endogenous insect (Rynochenus brevicollis: Curculionidae) in the Brazilian savanna. Three hypotheses were addressed: (i) the predator satiation hypothesis, (ii) the resource concentration hypothesis and (iii) the larger seed predation hypothesis. A total of 112 individual C. langsdorffii were monitored monthly from January to August during four consecutive years (from 2008 to 2011) to determine the presence of fruits on each plant. All trees produced fruits in the year 2008, whereas none of them produced flowers or fruits in 2009 or 2010. Moreover, only 65 individuals (58%) marked in 2008 produced fruits in 2011. The number of fruits per plant was approximately 21% greater in 2008 than in 2011, while the percentage of seed predation was 76% greater in 2011, thereby supporting the predator satiation hypothesis. The percentage of seeds predated was not affected by the number of fruits per plant. Therefore, our data did not support the resource concentration hypothesis. Plants producing large seeds experienced more seed predation by R. brevicollis, supporting the larger seed predation hypothesis. In addition, we also observed a positive relationship between seed volume and adult R. brevicollis weight. This study demonstrates the importance of supra‐annual fruiting for increasing survivorship of C. langsdorffii seeds both at the individual and the population level, and suggests that seed predators select plants producing large seeds as a way of increasing the number of offspring.     

Selective fruit filling in relation to pollen load size in Alstroemeria aurea (Alstroemeriaceae)

 We conducted an experiment in a natural population of Alstroemeria aurea, a clonal perennial, to determine (1) if reproduction was resource limited, and (2) if fruits would be selectively filled based on differences in pollination intensity when pollen loads were adequate for full seed set. Under these conditions, differences in pollination intensity are unlikely to affect seed number, but could affect seed quality, providing an interesting test of the gametophytic competition hypothesis. To test for resource limitation, percent fruit maturation, number of seeds per fruit and average seed weight were compared to paired controls for ramets in which all but one fruit was removed. To test the effect of pollination intensity on selective resource allocation, three types of pollination treatments were performed: (1) all flowers of the single inflorescence received a low pollen load, (2) all flowers received a high pollen load, (3) alternate flowers of the inflorescence received either a high or a low pollen load. We determined the percentage of fruit that reached maturity, counted the number of seeds and ovules and calculated the average seed weight for all capsules in each treatment. Resources appeared to limit reproduction in this population since seed number and weight were significantly higher than in controls when competing capsules were removed. At the whole ramet level, a four fold difference in pollen loads had no significant effect on any of the parameters measured. However, when pollination intensity varied within an inflorescence, the number of seeds per fruit increased by about 10% in flowers that received the higher pollen load. We observed the same trend in each of 2 years, but the increase was significant in only 1 year. The differences, although not great, were only slightly smaller than when all competing fruits were removed, and were consistent with selective resource allocation based on pollination intensity independent of seed set. Received: 28 September 1997 / Accepted: 30 April 1998     

Effect of ovular receptivity on seed set and fruit development in cucumber (Cucumis sativus L.)

Summary Seed set and fruit development in cucumber (Cucumis sativus L.) were studied in relation to female flower receptivity from day — 2 before anthesis to day + 2 after anthesis. The female cucumber flower is protogynous. The pistil was receptive 2 days before anthesis. The iso-electric focusing (IEF) patterns of the stigma/style proteins, were identical from day -5 to day +2. In pollinated flowers in vivo germination and pollen-tube growth in the ovary were affected by pistil age from day -2 to day +2. In addition, differences in sectorial filling in full seeds were observed within the fruits. A negative correlation was observed between the frequency of fertilized ovules in the pedoncular part of the fruit and ovary length at the time of pollination. In the whole fruit, significant differences in the number of full seeds and fruit size at maturity were found, and these were observed to be correlated with the various stages of female flower maturation at pollination. The day -2 and day +2 stages yielded the smallest fruits with few full seeds compared to the day -1, day 0 and day +1 stages, which had the biggest fruits and a large number of full seeds. A strong positive correlation was found between total seed number (including full and empty seeds), fruit length and weight at maturity. All these results suggest that both seed set in the different parts of the fruit and fruit development are controlled by ovular receptivity rather than by stigma/style receptivity.