Leaf production,reproductive patterns,field germination and seedling survival in Chamaedorea bartlingiana,a dioecious understory palm

Summary Chamaedorea bartlingiana is a dioecious palm that grows in the cloud forest understories of the Venezuelan Andes. Age and sexual differences in phenology and reproductive patterns were studied in labelled individuals of all age categories. This species has long-lived leaves and low leaf production, both characteristic of understory plants. Growth rates are lower in juveniles than in adults and in females than in males, as in other palms. Male and female individuals show different reproductive patterns. Male inflorescences are always produced at the same rate and the probability of surviving until anthesis is constant. Females produce reproductive buds at the same rate as males, but these buds have a 35% probability of becoming a ripe infrutescence if the plant has infrutescences already growing, and 70% if it does not. This pattern and the slow growth of inflorescences (1 year for males from bud to flowers, 2 years for females from bud to ripe fruits) cause a pluriannual reproductive pattern at the population level. Field germination does not follow this pattern, but shows one annual peak probably related to environmental conditions.     

Harvesting Increases Reproductive Activity in Himatanthus drasticus (Mart.) Plumel (Apocynaceae), a Non‐Timber Forest Product of the Brazilian Savanna

The harvesting of non‐timber forest products (NTFP) can influence the pattern of resource allocation in plants, affecting their growth, survival, and reproduction. However, only a small number of studies have addressed the impacts of NTFP harvesting on the reproductive phenology of the exploited species. The aim of this observational study was to assess the effects of harvesting and climatic variables on the reproductive phenology of Himatanthus drasticus, a highly exploited medicinal tree from the Brazilian savanna (Cerrado). We evaluated the effects of two different debarking levels (50 and 100%) in comparison to a control (no harvesting). We performed monthly counts of the number of buds, flowers in anthesis, unripe and ripe fruits in each sampled tree (intensity index), as well as the number of trees in each phenophase (activity index), over 2 yr. We used circular statistics tools to compare the effects of each treatment on flower and fruit production and to test whether the reproductive peaks were related to climatic variables. Both the activity and intensity indexes exhibited the same patterns; flower and fruit production were lower in the control, intermediate in the 50 percent debarking group and higher in the 100 percent debarking group. To the best of our knowledge, this is the first study of NTFP to demonstrate an increase in reproductive activity after several years of harvesting. All phenophases were positively correlated with higher temperature and precipitation, as it is common in the Cerrado.     

Cattle change plant reproductive phenology,promoting community changes in a post-fire Nothofagus forest in northern Patagonia,Argentina

Aims Variations in rates and length of flowering and fruiting not only affect the reproduction of a given plant species but also the behavior and reproduction of associated taxa. Flowering and fruiting variations may be influenced by herbivory, especially by large mammals. The aim of this study was to determine the effects of cattle browsing on the reproductive phenology of understory species in a subalpine post-fire Nothofagus forest in Patagonia.Methods The effects of herbivory on plant reproductive phenology were studied in a set of experimental exclosures (fenced plots) installed since 2001 in a post-fire N. pumilio forest, located in Nahuel Huapi National Park (NHNP), Argentina. We monitored the beginning and duration of each reproductive phenological stage: floral bud, open flower, immature fruit and mature fruit. We also counted the number of flowers, fruits, seeds and viable seeds of the dominant plants to assess whether browsing modifies temporal patterns of the flowering and fruiting periods.Important findings Cattle reduced the total number of species flowering and fruiting and changed the reproductive phenology of some species. We found that palatable species seem to be negatively affected by browsing in terms of reduced fitness due to changes in flowering and fruiting periods. In contrast, cattle benefitted the reproduction of non-palatable species and could promote the invasion of shade-intolerant exotic forbs such as Cirsium vulgare. The effects of livestock reported in this study are important to understanding how browsing could alter native species establishment and possibly alter successional trajectories during recolonization after fire.     

How plant allometry influences bud phenology and fruit yield in two Vaccinium species

Background and AimsUnderstanding how plant allometry, plant architecture and phenology contribute to fruit production can identify those plant traits that maximize fruit yield. In this study, we compared these variables and fruit yield for two shrub species, Vaccinium angustifolium and Vaccinium myrtilloides, to test the hypothesis that phenology is linked to the plants’ allometric traits, which are predictors of fruit production.MethodsWe measured leaf and flower phenology and the above-ground biomass of both Vaccinium species in a commercial wild lowbush blueberry field (Quebec, Canada) over a 2-year crop cycle; 1 year of pruning followed by 1 year of harvest. Leaf and flower phenology were measured, and the allometric traits of shoots and buds were monitored over the crop cycle. We hand-collected the fruits of each plant to determine fruit attributes and biomass.Key ResultsDuring the harvesting year, the leafing and flowering of V. angustifolium occurred earlier than that of V. myrtilloides. This difference was related to the allometric characteristics of the buds due to differences in carbon partitioning by the plants during the pruning year. Through structural equation modelling, we identified that the earlier leafing in V. angustifolium was related to a lower leaf bud number, while earlier flowering was linked to a lower number of flowers per bud. Despite differences in reproductive allometric traits, vegetative biomass still determined reproductive biomass in a log–log scale model.ConclusionsGrowing buds are competing sinks for non-structural carbohydrates. Their differences in both number and characteristics (e.g. number of flowers per bud) influence levels of fruit production and explain some of the phenological differences observed between the two Vaccinium species. For similar above-ground biomass, both Vaccinium species had similar reproductive outputs in terms of fruit biomass, despite differences in reproductive traits such as fruit size and number.     

Predispersal reproductive biology of female Osyris quadripartita (Santalaceae), a hemiparasitic dioecious shrub of Mediterranean scrublands

Female Osyris quadripartita plants exhibit uninterrupted reproductive activity throughout the year, due to the long duration of successive stages in the cycle and marked within-crop developmental asynchrony. Cycles corresponding to the flowering seasons of consecutive years overlap in each individual. Flowering takes place in spring, and fruits develop in the dry summer season and ripen at any time of the year. Variation in flowering time explains a negligible proportion of variation in ripening time. The greatest reproductive losses are incurred in the phase extending from closed flowers through unripe fruits, mostly due to ovary abortion. Only 30% of closed flowers eventually reach this latter stage. In contrast, 75% of unripe fruits complete their development, with subsequent dispersal of seeds. The probability of the setting of ripe fruit steadily decreases from early to late season flowers, due to increased ovary abortion rates. Resource limitation in the dry summer season seems responsible for this pattern of selective fruit maturation.     

Variation in reproductive output of Ficus superba despite aseasonal reproduction

Studies of Ficus reproductive phenology have focused on documenting its seasonality and dependence on meteorological factors. While there have been reports that duration of receptivity of syconia can be prolonged in response to pollinator limitation, the effect of pollination rate on the duration of phenological phases over a year has not been examined. Percentage of unpollinated syconia, number of foundresses per pollinated syconium, duration of receptivity, frequency of single-sex broods, crop size, frequency of parasitisation, persistence of ripe syconia, and mass abscission were recorded for Ficus superba at the crop level. Percentage of unpollinated syconia is significantly negatively correlated with the number of foundresses per syconium, and positively correlated with the duration of receptivity, and the persistence of ripe syconia. Despite the absence of sesonality in reproduction, the occurrences of receptive syconia and ripe syconia showed unimodal and bimodal peaks, respectively, owing to prolongment of these phases. This is attributable to meteorological factors, which are hypothesized to influence pollinator dispersal, or population dynamics, and thus pollination rate. This highlights the overlooked significance of pollinator, and possibly frugivore phenology in accounting for Ficus reproduction, and suggests that while reproduction may be aseasonal, reproductive potential may not be. Furthermore, crop size was shown to affect the number of foundresses per syconium, duration of receptivity, and persistence of ripe syconia. Possible adaptive value of producing crops of different sizes is discussed.     

甘南红景天濒危的生殖生态学因素分析

为了寻找保护和恢复甘南红景天种群的有效途径,本文分析了甘南红景天的繁育系统、生殖过程、开花物候特征及影响生殖成功的主要因子.结果表明:甘南红景天为雌雄异株,雄花在花芽分化初期为两性植株,但随着花的不断发育,雌蕊的胚囊发育到一定阶段时败育,两性植株基本仅执行雄性功能,种群繁殖为异交,以风媒传粉方式为主,种群花期出现在全年降雨和气温较高的季节.6月初现蕾,6月中旬开花,7月上旬进入盛花期,花期在36 d左右,8月下旬果实成熟.两性植株略早于雌花开放,花期(10.2 d)大于雌性植株(8.4 d).由于部分雌花不易授粉,部分花蕾、胚珠损失,1~3年的幼龄植株几乎不开花结实,群落中只有11%的植株开花结实,种子自然繁殖率极低,约为2%.从花蕾到开花,雌性单株有20.0%～25.1%的花蕾损失; 从开花到结实,单株51.1%～65.0%的花败育;至种子成熟,仅10.1%～21.0%的胚珠发育成种子.雌性单株平均种子产量为158.1粒,人工条件下种子发芽率为81.5%,幼苗第一年的成活率为36.0%.表明种子质量不是导致甘南红景天濒危的原因,而花粉限制、种子产量和幼苗的成活率是真正影响其生殖成功和导致濒危的主要生殖生态学因素.     

Reproductive biology ofPolyalthia littoralis (Annonaceae)

Flowering phenology of some annonaceous trees and reproductive biology ofPolyalthia littoralis (Annonaceae) were studied. The trees showed various types of flowering phenology within the family. Among them,P. littoralis had hermaphroditic and protogynous flowers, and exhibited continuous flowering throughout the year. Bagged flowers set fruits and seeds comparable to the control. The observations of meiotic stages and the results of castrated tests indicated no possibility of apomictic reproduction. The abscised anthers attached to torus by the extended spiral thickenings, and accomplished self-pollination. Outbreeding possibly occurs, but the following self-pollination guarantees seed set.     

Flowering phenology and reproductive success of native shrub Berberis darwinii Hook. along different light environments

Studies evaluating flowering phenology and reproductive success are necessary when we want to direct a domestication project in a species with a potential productive value. We studied flowering phenology and reproductive success of Berberis darwinii growing under different light conditions in its native distribution area in the Andean Patagonian forests of Argentina. We test the hypothesis that plants grown under conditions of high-light availability exhibit advanced phenology and higher reproductive success than those grown under conditions of lower light availability. Phenology and reproductive success were determined in three contrasting light conditions at two forest sites, which were, canopy, gap and forest edge. Plants did not bloom under the forest canopy. Flowering and fruiting period lengths were similar in both sites and light conditions of gap and forest edge during spring and summer. Although gap plants had more racemes per shoot, racemes of edge plants had more flowers, fruits and a higher proportion of flowers producing ripe fruit. We show that B. darwinii reproduction studied in the Andean Patagonian forests is conditioned by the canopy openness. Regarding reproductive success, edge plants invest less resources in flower production than gap plants to have similar fruit production.     

Competition and the compensatory regulation of fruit and seed set in the perennial herb Epilobium dodonaei (Onagraceae)

I studied the effects of competition and soil fertilization on variation of seed yield components of the outcrossing perennial Epilobium dodonaei Vill. using a randomized complete block design. Fertilization as a main effect was not significant. Competition, however, reduced the number of flower buds per shoot (-29%), the number of ovules per fruit (-12%), and the number of ripe fruits per shoot (-51%). In addition, competition caused an increase in abortion of flower buds. Consequently the fruit/flower bud ratio decreased from 0.71 to 0.47. Average seed mass was not reduced significantly by competition. Some of the negative effects of competition on fruit production were mitigated by fertilization. However, competition considerably reduced the proportion of late-aborted seeds, which resulted in an increase of the seed/ovule ratio from 0.31 to 0.49 (+58%). As a consequence of this compensation, the reduction in the number of seeds per shoot due to competition was not significant. At the level of the genet, competition had a strong effect on seed yield due to decreases in the number of shoots produced. I discuss patterns of regulation at consecutive levels of reproduction. Shoots of plants suffering from competition initially invested less in reproductive structures than the control and showed a higher abortion rate of less costly structures early in reproductive development, but also had a lower abortion rate later in their development. The observed compensatory pattern in yield components illustrates the fine-tuning regulation capacity at different levels of reproductive development in plants.     

Reproductive phenology in three Genisteae (Fabaceae) shrub species of the W Mediterranean Region

We studied the flower duration, the phenology of flowering and fruiting, and flower, fruit, and seed production in three Genisteae shrub species of the W Mediterranean Region: Cytisus multiflorus, C. striatus , and Retama sphaerocarpa . Flower duration was negatively correlated with temperature, and in the case of C. striatus it was also influenced by pollination. In Cytisus multiflorus , which flowers during winter in the population studied, two floral morphs were recognized differentiated by flower size, phenological pattern, and production of the reproductive organs: morph LF (large flowers) and morph SF (small flowers). The former is earlier in phenology but its reproductive success is less than the second since it not only produces significantly fewer flowers/plant, but also a smaller crop of fruit and seeds. The flowering of the LF morph lasted some three months and of the SF morph 11 weeks. The winter-spring flowering C. striatus , with a four month duration of flowering time, is sympatric with C. multiflorus and their flowering periods overlap, but the former is more successful reproductively. Retama sphaerocarpa is clearly spring flowering, with an extremely short duration of flowering (c. 6.5 weeks), but, unlike the other two species which disperse their seeds in the same season in which they are produced, it staggers the dispersal of its diaspores over more than one year. Despite the great flower per plant production, especially in Retama sphaerocarpa , and of the number of fruit initiated, the final crop of fruit and seed/plant is low in the three species (C. multiflorus : fruit 8.40% and seeds 1.96%; C. striatus : fruit 13.09% and seeds 4.12%; Retama sphaerocarpa : fruit 5.65% and seeds 1.33%).     

Environmental control of reproductive phenology and the effect of pollen supplementation on resource allocation in the cleistogamous weed, Ruellia nudiflora (Acanthaceae)

• Background and Aims Mixed reproductive strategies may have evolved as a response of plants to cope with environmental variation. One example of a mixed reproductive strategy is dimorphic cleistogamy, where a single plant produces closed, obligately self-pollinated (CL) flowers and open, potentially outcrossed (CH) flowers. Frequently, optimal environmental conditions favour production of more costly CH structures whilst economical and reliable CL structures are produced under less favourable conditions. In this study we explore (1) the effect of light and water on the reproductive phenology and (2) the effect of pollen supplementation on resource allocation to seeds in the cleistogamous weed Ruellia nudiflora.• Methods Split-plot field experiments were carried out to assess the effect of shade (two levels: ambient light vs. a reduction of 50 %) and watering (two levels: non-watered vs. watered) on the onset, end and duration of the production of three reproductive structures: CH flowers, CH fruit and CL fruit. We also looked at the effect of these environmental factors on biomass allocation to seeds (seed weight) from obligately self-pollinated flowers (CL), open-pollinated CH flowers and pollen-supplemented CH flowers.• Key Results CH structures were produced for a briefer period and ended earlier under shaded conditions. These conditions also resulted in an earlier production of CL fruit. Shaded conditions also produced greater biomass allocation to CH seeds receiving extra pollen.• Conclusions Sub-optimal (shaded) conditions resulted in a briefer production period of CH structures whilst these same conditions resulted in an earlier production of CL structures. However, under sub-optimal conditions, plants also allocated more resources to seeds sired from CH flowers receiving large pollen loads. Earlier production of reproductive structures and relatively larger seed might improve subsequent success of CL and pollen-supplemented CH seeds, respectively.     

Novel nectar robbing negatively affects reproduction in Digitalis purpurea

With many plant–pollinator interactions undergoing change as species’ distributions shift, we require a better understanding of how the addition of new interacting partners can affect plant reproduction. One such group of floral visitors, nectar robbers, can deplete plants of nectar rewards without contributing to pollination. The addition of nectar robbing to the floral visitor assemblage could therefore have costs to the plant´s reproductive output. We focus on a recent plant colonist, Digitalis purpurea, a plant that in its native range is rarely robbed, but experiences intense nectar robbing in areas it has been introduced to. Here, we test the costs to reproduction following experimental nectar robbing. To identify any changes in the behavior of the principal pollinators in response to nectar robbing, we measured visitation rates, visit duration, proportion of flowers visited, and rate of rejection of inflorescences. To find the effects of robbing on fitness, we used proxies for female and male components of reproductive output, by measuring the seeds produced per fruit and the pollen export, respectively. Nectar robbing significantly reduced the rate of visitation and lengths of visits by bumblebees. Additionally, bumblebees visited a lower proportion of flowers on an inflorescence that had robbed flowers. We found that flowers in the robbed treatment produced significantly fewer seeds per fruit on average but did not export fewer pollen grains. Our finding that robbing leads to reduced seed production could be due to fewer and shorter visits to flowers leading to less effective pollination. We discuss the potential consequences of new pollinator environments, such as exposure to nectar robbing, for plant reproduction.     

The role of reproductive phenology, seedling emergence and establishment of perennial Salix gordejevii in active sand dune fields

BACKGROUND AND AIMS: The function of sexual reproduction of perennials in restoration of vegetation of active dune fields frequently has been underestimated. The objective of this study was to evaluate the role of sexual reproduction of the perennial Salix gordejevii in the revegetation of active dunes. METHODS: Seedling emergence and establishment of S. gordejevii were examined both in controlled experiments (germination at different burial depths with different watering regimes) and in field observations in three dune slacks. The reproductive phenology and soil seed bank of S. gordejevii, the dynamics of soil moisture, the groundwater table and the landform level of three dune slacks were monitored. KEY RESULTS: Seeds of S. gordejevii began maturation on 1 May, and seed dispersal lasted from 8 May to 20 May. Seeds on the soil surface germinated significantly faster than those buried in soil (P<0.05). Seedling emergence was negatively correlated with landform level. When most seedlings emerged, there was a significantly positive correlation between soil moisture and seedling emergence (P<0.01). Rainfall was negatively correlated with seedling emergence. Seedling establishment was significantly and positively correlated with seedling emergence (P<0.05), and 72.3 % of the emergent seedlings were established at the end of the growing season. These results indicated that (a) seeds matured and dispersed before the rainy season; (b) seeds germinated as soon as they contacted a moist surface and relied more on soil moisture than on rainfall; and (c) more seedlings emerged at lower sampling points in dune slacks. CONCLUSIONS: In natural conditions, restoration of active sand dune fields generally commences with revegetation of dune slacks where sexual reproduction of perennials contributes greatly to species encroachment and colonization and hence plays an important role in restoration of active dune fields. Furthermore, aeolian erosion in dune slacks, leading to good soil moisture, facilitates seed germination, seedling emergence and establishment of S. gordejevii.     

Plasticity of reproductive components at different stages of development in the annual plant Thlaspi arvense L.

Summary This study examines the effect of different densities and the removal of all neighbours at different stages of development on all components of reproduction in the inbreeding annual Thlaspi arvense L. A 64-fold increase in density significantly reduced all repooductive components. The number of flower buds per plant was decreased most strongly; the order of decreasing plasticity among the other components was number of capsules per flower, individual seed weight, ovule number per capsule, flowers per flower bud and seeds per ovule. Removing neighbours at all stages of development increased seed yield of plants in comparison to the control without density reduction, but patterns of plasticity depended strongly on time of treatment. The main effect of the removal of neighbours at the vegetative stage was to increase the number of flowers per plant, but the number of ovules per capsule and seed weight increased also, and abortion of capsules decreased. Removing neighbours at the onset of flowering initially failed to affect flower number per plant, instead it resulted in a strong reduction of capsule abortion and an increase in seed weight. However, several weeks after flowering had initially ceased, fresh lateral inflorescences were produced, resulting in a second flush of reproduction. Removing neighbours at the stage of fruit ripening resulted at first only in an increase in seed witht, but later a second reproductive phase occurred. Fresh lateral branches were produced, but the apical meristem was also reactivated. The overall pattern of plasticity among all reproductive components in response to a removal of neighbours was the same as in response to density. The position of a capsule along the inflorescence influenced its number of ovules, the rate of seed abortion and the mean weight of seeds, with the type of effect depending on the developmental stage at which neighbours were removed. Significant negative correlations were found between the mean weight of seeds and the number of seeds in a capsule under all treatments.     

REPRODUCTIVE ATTRITION IN THE RARE CHAPARRAL SHRUB FREMONTODENDRON DECUMBENS LLOYD (STERCULIACEAE)

We examined reproductive attrition in Fremontodendron decumbens to characterize sexual reproduction in this rare California shrub. Reproductive individuals produced an average of 2,900 flower buds in a season, with no significant difference in bud production between two seasons. Because of intense insect predation, <;2% of initiated flower buds became mature fruits. A threefold decrease in predation of flower buds between seasons resulted in an increase in seed output the second season, indicating that seed production was partially predator-limited. Most seeds (97.8%) were dormant due to an impermeable seed coat. Breaking of the coat, mechanically or by heat, allowed high levels of germination. Chamise charate and ash added to the potting medium resulted in the highest level of germination and emergence. Rodents were more important than birds as seed predators, destroying 90% of seeds under parent shrub canopies within 8–10 months. Seeds already integrated into the seed bank were comparatively safe from predation, relative to newly added seeds. If predation was prevented, seeds were long-lived under field conditions (>;80% survived after 5.75 years). Most seedlings produced in unburned chaparral by planting heat-treated seeds in openings between shrubs were destroyed by predators (rodents and insects). All seedlings that escaped predation died during the summer drought. We concluded that sexual reproduction was limited by (in order of importance): 1) lack of fire, 2) predehiscence predation by insects, and 3) postdehiscence predation by rodents. Size distributions from two populations revealed that, despite the apparent absence of sexual reproduction in unbumed chaparral, two unbumed sites contained a large proportion of individuals in small size classes. Excavation of several small individuals demonstrated they were sprouts from the roots of nearby larger shrubs. Because asexual reproduction by rootsprouting circumvents the high attrition of sexual reproductive effort on unbumed sites, rootsprouting may be a significant reproductive strategy of some ‘sprouter’ species in chaparral.     

Sexual dimorphism and fruit production in a dioecious understory tree,Ilex opaca Ait.

Summary This study suggested that sexual selection is potentially an important factor in the maintenance of dioecy in the American holly, Ilex opaca (Aquifoliaceae). Sexual dimorphisms in flower production and phenology were highly significant in this understory tree. On average, individual males produced 7.4 times as many flowers as did female trees. Staminate flowers lasted only a single day, whereas pistillate flowers lasted 3–4 days, during which they showed no significant decline in their ability to produce fruit after pollination. Individual male trees opened their flower buds asynchronously during the season, maximizing the number of days they were in flower. Individual females opened their buds more synchronously, maximizing their floral display at one point in time. Females produced fruits in numbers that were somewhat less than proportional to their flower production. Fruit development was initiated from only 38.9% and 69.5% of pistillate flowers in 1987 and 1988, respectively. By the time of ripening, an average female had lost 62.3%, 24.3%, and 11.1% of its initial fruit crop in 1986, 1987, and 1988, respectively. The proportion of fruit lost in 1986 was independent of the number of fruit that initially began development. In 1988, artifically supplementing pollen to a large number of flowers failed to increase either fruit or seed production relative to control branches with unsupplemented flowers. This suggested that resource levels were likely more important than pollen availability in limiting female reproductive success. These observations on I. opaca were consistent with the expectations for a population in which male reproductive success continues to benefit from continued pollinator service and female reproductive success does not.     

Fitness implications of late bud break and time of burning in Lannea edulis (Sond.) Engl. (Anacardiaceae)

Many woody plants of savannas have massive underground parts (“lignotubers”) and diminutive aboveground parts with phenologies that are triggered by fire. Lannea edulis (Anacardiaceae) represents this life form and is widely distributed in south central Africa. The phenology of L. edulis was monitored on 42 permanently marked shoots over a 3-year period, from 2002 to 2004, and under three fire treatments (early and middle dry season burning, and fire protection) to determine whether (i) fire affected time of bud break and (ii) time of fruit ripening affected seed germination. Statistical analyses revealed that shoot reproductive status and fire treatments significantly affected time of bud break and the lag between bud break and leaf production. Shoot reproductive status explained 27% (P<0.001) while the interaction between reproductive status and fire treatments explained 34% (P<0.0001) of the variation in time at bud break. The appearance of the first leaf was delayed by 4 weeks in reproductive shoots compared with vegetative shoots. Shoot reproductive status explained 28% (P<0.0001) of the variation in the lag between bud break and appearance of the first leaf suggesting the existence of resource competition between reproduction and vegetative growth. However, reproductive status and fire treatments did not affect shoot size and leaves produced because replacement shoots after dieback due to fire were as long as shoots surviving from previous years due to insignificant annual shoot extension among the latter. There was high autocorrelation in the germination rate of seed-lots with seeds from fruits ripening early having a significantly higher germination rate (47% in 2002 and 89% in 2004) than those from fruits ripening later (3% in 2002 and 53% in 2004). Since time at bud break and fruit ripening were linked, it is proposed that late bud break results in reduced fitness in L. edulis because of reduced seed germination rate. Early and late dry season fires that delay bud break and destroy reproductive structures, respectively, also reduce this measure of fitness.     

Flower development and reproductive continuity in MediterraneanRuscus aculeatus L. (Liliaceae)

Summary Ruscus aculeatus is a subandroecious species widespread in Mediterranean environments. Our studies on floral differentiation show that the unisexual flowers pass through a stage in which both androecial and gynoecial primordia are initiated. The hypothesis that dioecy has arisen secondarily from hermaphroditism is discussed. Data on flowering phenology of Mediterranean populations ofR. aculeatus showed it has a long flowering season (about 7 months). This phenology is similar to that of the Mediterranean species originated before the development of the Mediterranean climate. Members of the family Ruscaceae where present in Laurasia during the tropical Tertiary and the present study on flower and fruit morphology and reproductive phenology reveals forR. aculeatus a list of characters generally reported for tropical species. Small and greenish flowers, fleshy fruits, few large seeds, and resprouting capacity, together with long flowering season and continuous availability of ripe fruits, occur all together inR. aculeatus. It constitutes a tropical reproductive syndrome which might have survived the climatic fluctuations of the Quaternary, the establishment of the Mediterranean climate, and the present anthropogenic disturbs.