Variation in the phenology and abundance of flowering by native and exotic plants in subalpine meadows

The timing and abundance of flower production is important to the reproductive success of angiosperms as well as pollinators and floral and seed herbivores. Exotic plants often compete with native plants for space and limiting resources, potentially altering community floral dynamics. We used observations and a biomass-removal experiment to explore the effects of an invasive exotic flowering plant, Linaria vulgaris, on community and individual species flowering phenology and abundance in subalpine meadows in Colorado, USA. Invasion by L. vulgaris was associated with a shift in both the timing and abundance of community flowering. Invaded plant communities exhibited depressed flowering by 67% early in the season relative to uninvaded communities, but invaded sites produced 7.6 times more flowers than uninvaded sites once L. vulgaris began flowering. This increase in flowers at the end of the season was driven primarily by prolific flowering of L. vulgaris. We also found lower richness and evenness of resident flowering species in invaded plots during the period of L. vulgaris flowering. At the species level, a common native species (Potentilla pulcherrima) produced 71% fewer flowers in invaded relative to uninvaded plots, and the species had reduced duration of flowering in invaded relative to uninvaded sites. This result suggests that L. vulgaris does not simply alter the flowering of subordinate species but also the flowering of an individual common species in the plant community. We then used observational data to explore the relationship between L. vulgaris density and resident floral production but found only partial evidence that higher densities of L. vulgaris were associated with stronger effects on resident floral production. Taken together, results suggest that a dominant invasive plant can affect community and individual-species flowering.     

Association between rainfall seasonality and the flowering of epiphytic plants in a Neotropical montane forest

The association between the reproductive phenology of epiphytic communities with environmental and ecological factors remains largely unexplored. Because epiphytes depend on environmental moisture, seasonal changes in moisture conditions likely act as the primary determinants of their reproductive timing. We examined whether water limitation or pollinator competition structures the flowering phenologies of an epiphytic community in a seasonal mountain forest in Costa Rica. Additionally, we addressed the environmental factors that might trigger floral induction. Using a 24‐month dataset of bimonthly flowering records from 104 species, we found high seasonality of flowering at the species level but somewhat lower seasonality at the community level. The flowering mid‐dates of most epiphytes, particularly from monocotyledonous species, occurred during the wettest months, as predicted if water limitation structures flowering. The increased moisture and nutrient availability during the rainy season give epiphytes the resources needed to complete floral development and anthesis, and later fruit and seed maturation. The observed flowering pattern of epiphytes coincides with reproductive patterns of terrestrial herbs and shrubs from seasonal tropical ecosystems, and suggests shared constraints to sexual reproduction in both ecological guilds under similar climatic conditions. In contrast, flowering patterns of congeneric epiphytes in the same pollination guild mostly did not follow the expectations of a pollinator competition scenario. Finally, we discuss the possible combined effect of precipitation, temperature, and daily insolation on floral induction of epiphytic plants.     

Later flowering is associated with a compressed flowering season and reduced reproductive output in an early season floral resource

Climate change‐induced shifts in flowering phenology can expose plants to novel biotic and abiotic environments, potentially leading to decreased temporal overlap with pollinators and exposure to conditions that negatively affect fruit and seed set. We explored the relationship between flowering phenology and reproductive output in the common shrub pointleaf manzanita Arctostaphylos pungens in a lower montane habitat in southeastern Arizona, USA. Contrary to the pattern of progressively earlier flowering observed in many species, long‐term records show that A. pungens flowering onset is shifting later and the flowering season is being compressed. This species can thus provide unusual insight into the effects of altered phenology. To determine the consequences of among‐ and within‐plant variation in flowering time, we documented individual flowering schedules and followed the fates of flowers on over 50 plants throughout two seasons (2012 and 2013). We also measured visitation rates by potential pollinators in 2012, as well as both fruit mass and seeds per fruit of flowers produced at different times. Fruit set was positively related to visitation rate but declined with later dates of flower production in both years. Total fruit production per plant was positively influenced by flowering duration, which declined with later flowering onset, as did fruit mass. Individual flowering schedules were consistent between years, suggesting that plants that begin flowering late have lower reproductive output each year. These patterns suggest that if pointleaf manzanita flowering continues to shift later, its flowering season may continue to become shorter, compressing floral resource availability for pollinators and leading to reduced reproductive output. These results reveal the negative effects of delayed phenology on reproductive output in a long‐lived plant. They highlight the value of using natural variation in flowering time, in combination with long‐term data, to anticipate the consequences of phenological shifts.     

An investigation of temporal flowering segregation in species-rich grasslands

Processes such as competition and facilitation are believed to be important in defining pollination niches in species-rich plant communities. Species with similar floral phenotypes are expected to flower together where this facilitates pollination, while differences in floral phenology are expected if such flowers compete for pollinators. These expectations were tested at seven sites by comparing the observed co-flowering of plants with similar floral phenotypes with null model outcomes. Phenotypic classifications were evaluated using observations of plant–pollinator interactions. Pollinator guilds differed in the number of visits made to flowers in different floral colour and shape categories, indicating that such categories were ecologically relevant. For species with complex flowers, each floral category contained few species, so that the observed low occurrence of co-flowering could be explained by chance. In contrast, within phenotypic categories species with simple flowers bloomed together more than expected at three sites, but these overlaps could be explained by family membership. Most species with complex flowers could be segregated into unique pollination niches by broad floral colour and shape categories, so that there was little opportunity for competition between flowers within such categories. Species with simple floral phenotypes were less well defined by floral phenotype and phenology. Historical sorting may explain differences between complex flowers, while co-flowering between species with simple flowers requires further investigation. Differences found between species with simple and complex flowers suggest that levels of phenotypic specialisation should be taken into account in community level studies of pollination systems.     

雌雄异熟植物露蕊乌头开花时间对雌雄功能期及表型性别的影响

开花时间决定了植物雌雄功能的交配机会, 最终影响繁殖成功。交配环境假说认为雌雄异熟植物开花时间的差异能引起植物表型性别的变异, 改变种群内的交配环境, 影响植物对雌雄功能的最佳性分配。为了研究开花时间对雌雄异熟植物的雌雄性别时期及表型性别的影响, 本文以毛茛科雄性先熟植物露蕊乌头(Aconitum gymnandrum)为实验材料, 记录了雄性和雌性功能期, 分析了植株开花时间、花的雌雄功能期和表型性别的关系。结果表明: 在植物同一花序内, 较晚开放的花有更长的雄性期和更短的雌性期, 性分配在时间上偏雄。雌雄功能期在时间上的相对分配随植物开花时间的变化表现出相似的趋势： 较晚开的花或较晚开花的个体, 花的雄性功能期相对于雌性功能期更长, 在时间上更偏向雄性功能。而且, 开花时间的差异影响种群内花的性比和植物个体的表型性别动态。随着开花时间由早到晚的变化, 种群内早期以雄花为主,末期以雌花为主, 种群内性别环境由偏雄向偏雌变化, 因此植株个体的平均表型性别则从偏雌转向偏雄。本文结果支持交配环境假说, 雄性先熟的露蕊乌头开花早期, 种群内花的性别比偏雄, 种群表型性别环境偏雄, 因而植物个体平均表型性别偏雌, 性别分配(即时间分配)偏向雌性功能, 而晚开花个体的平均性别偏雄, 更偏向雄性功能的分配。     

Flowering phenology of Campanula on Mt Olynipos, Greece

We studied the fowering phenology of the genus Campanula (represented by nine species) along the elevation gradient of Mt Olympos (Greece) in order to assess whether there are elevation patterns at the genus level and whether these relate to patterns previously observed along such gradients at the community level. The traits examined were time and duration of the flowering period, flower life span, and duration of male and female flower phases. Floral attributes, such as number of flowers per plant, flower biomass, flower size, were also studied in order to examine whether they change with elevation or influence flower phenology. Flowering of Campanula species started in mid-May and ended in late September. The average duration of flowering of the genus was ca 27 d and the average floral longevity 4 d. In all but one species, the female phase lasted longer than the male. Campanula versicolor differed remarkably from all others in flower phenology and other floral trails. Nearly all Campanula populations studied had right or positively skewed flowering distributions indicating that flowering begins more abruptly than it ends. At the genus level, the time of flowering increased with elevation by 2 3d for every 100 m. Floral longevity also increased with elevation, by 0,2 d for every 100 m. Neither duration of flowering nor duration of the flower phases showed any consistent change with elevation. The same is true of the non–phenological floral traits examined. No trade–off between duration of flowering and flower life span or between structure and maintenance of flowers was apparent. The pattern of increasing floral longevity along the elevation gradient suggests a mechanism of compensation for reduecd pollinator availability.     

Fruit set in a deceptive orchid: The effect of flowering phenology,display size,and local floral abundance

We investigated the effect of flowering time, display size, and local floral density on fruit set in Tolumnia variegata, a pollination-limited orchid that offers no reward to its pollinator(s). During 1990, natural variation in flowering time, display size, and fruit set were monitored in 508 plants at one locality in Puerto Rico. The following season, orchid floral abundance per host tree (Randia aculeata) was manipulated to investigate its effect on fruit set. Four floral abundance treatments were established (700, 500, 300, and 100), each replicated four times. Flowering time was the most important trait affecting fruit set. The proportion of plants setting at least one fruit was significantly high early and late in the season, but low during the flowering peak. Thus, strong disruptive selection differential on flowering phenology was found. Display size had little effect on fruit set. A weak, but significant disruptive selection differential on display size was found. Orchid floral abundance per host tree had a significant effect on fruit set. Early in the season, T. variegata flowers with intermediate number of conspecific flowers exhibited a greater probability of setting fruit than those in host trees with fewer or more flowers. Our results show that flowering phenology may be evolutionarily unstable, possibly a consequence of the deception pollination system. Furthermore, a deception strategy would be relatively unsuccessful in populations where plants are found in either very dense or sparse patches.     

二重雌雄异型异熟物种色木槭3种表型花的细胞形态学研究

二重雌雄异型异熟,即雌雄功能按雄-雌-雄顺序分别表达,是有花植物中较为少见的异熟机制.槭属(Acer)大部分物种具有二重雌雄异型异熟的开花物候,且第一期雄花与第二期在形态上差别较大.为深入探讨花发育模式对花形态和开花物候的影响,本文应用石蜡切片技术和光镜技术,对色木槭(Acer mono)各个发育时期的3种类型花(雄花...     

PLANT POLYPLOIDY AND POLLINATION: FLORAL TRAITS AND INSECT VISITS TO DIPLOID AND TETRAPLOID HEUCHERA GROSSULARIIFOLIA

In many polyploid species, polyploids often have different suites of floral traits and different flowering times than their diploid progenitor species. We hypothesized that such differences in floral traits in polyploids may subsequently affect their interactions with pollinating and other insect visitors. We measured floral morphology and flowering phenology in 14 populations of diploid and autotetraploid Heuchera grossulariifolia Rydb. (Saxifragaceae), determined if repeated evolution of independent polyploid lineages resulted in differentiation in floral morphology among those lineages, and ascertained if there was a consistent pattern of differentiation among genetically similar diploid and autotetraploid populations. In addition, we evaluated the differences in suites of floral visitors within a natural community where diploids and autotetraploids occur sympatrically. Overall, flowers of autotetraploid plants were larger and shaped differently than those of diploids, had a different flowering phenology than that of diploids, and attracted different suites of floral visitors. In comparison with flowers of diploids, tetraploid floral morphology varied widely from pronounced differences between cytotypes in some populations to similar flower shapes and sizes between ploidal levels in other populations. Observations of floral visitors to diploids and autotetraploids in a natural sympatric population demonstrated that the cytotypes had different suites of floral visitors and six of the 15 common visitors preferentially visited one ploidy more frequently. Moreover, we also found that floral morphology differed among independent autotetraploid origins, but there was no consistent pattern of differentiation between genetically similar diploid and autotetraploid populations. Hence, the results suggest that the process of polyploidization creates the potential for attraction of different suites of floral visitors. Multiple origins of polyploidy also presents the opportunity for new or different plant-insect interactions among independent polyploid lineages. These differences in turn may affect patterns of gene flow between diploids and polyploids and also among plants of independent polyploid origin. Polyploidy, therefore, may result in a geographic mosaic of interspecific interactions across a species' range, contributing to diversification in both plant and insect groups.     

大别山山地次生林鸟类群落集团结构的季节变化

2007年12月—2008年12月,在大别山鹞落坪,对落叶阔叶次生林鸟类的集团结构的季节变化特征进行了研究。采用连续取样法采集鸟类取食行为数据,用聚类分析法对繁殖和非繁殖季节鸟类群落进行集团划分,通过无倾向对应分析（DCA）对32种森林鸟类的两个季节资源利用特点进行了研究。结果表明,鸟类群落在非繁殖季节可以分为地面、灌丛、树干（枝）、冠层等4个取食集团,而在繁殖季节还出现空中取食集团;候鸟影响鹞落坪次生林鸟类群落取食集团的结构,产生新的取食集团。DCA的第一轴主要代表鸟类取食高度信息,第二轴主要代表鸟类取食位置信息,第三轴代表鸟类取食方式和取食基质信息;用DCA前两轴对32种鸟类排序表明,有6种鸟主要在第一轴发生变化,有4种鸟主要在第二轴发生变化。     

Reproductive Traits and Phenology of Plants in Tropical Dry Evergreen Forest on the Coromandel Coast of India

Qualitative reproductive traits of 84 plant species belonging to 41 families were studied in tropical dry evergreen forest on the Coromandel coast of India. Majority of species had rotate type, white-coloured, scented flowers, rewarding nectar and pollen and pollinated chiefly by bees. An association between floral traits and pollination spectrum is evident. Bee pollination was prevalent in pollination systems. Among the fruit types, drupe and berry were common in black and red colour respectively, and dispersed by zoochorous mode. Seeds of brown- and green-coloured dry fruits, without any reward were disseminated by wind and explosion. The reproductive phenophase of trees and lianas occurred mostly during the dry period from January to June, which receives rainfall of less than 50 mm a month. However, shrubs showed a peak in flowering and fruiting in wet period. Detailed phenological observations of 22 woody species revealed a seasonal and unimodal pattern in flowering. Although some species were in flower round the year, flowering activity was skewed towards the dry season. The fruiting activity showed a bimodal pattern, one peak in dry season and another in wet season. Many species displayed a temporal aggregation in flowering and fruiting. The significant relation was obtained between reproductive traits and phenology of plants in the tropical dry evergreen forest.     

Vulnerability of phenological synchrony between plants and pollinators in an alpine ecosystem

The relationship between flowering phenology and abundance of bumble bees (Bombus spp.) was investigated using 2 years of phenological data collected in an alpine region of northern Japan. Abundance of Bombus species was observed along a fixed transect throughout the flowering season. The number of flowering species was closely related to the floral resources for pollinators at the community scale. In the year with typical weather, the first flowering peak corresponded to the emergence time of queen bees from hibernation, while the second flowering peak corresponded to the active period of worker bees. In the year with an unusually warm spring, however, phenological synchrony between plants and bees was disrupted. Estimated emergence of queen bees was 10 days earlier than the first flowering date owing to earlier soil thawing and warming. However, subsequent worker emergence was delayed, indicating slower colony development. The flowering season finished 2 weeks earlier in the warm-spring year in response to earlier snowmelt. A common resident species in the alpine environment, B. hypocrita sapporoensis, flexibly responded to the yearly fluctuation of flowering. In contrast, population dynamics of other Bombus species were out of synchrony with the flowering: their frequencies were highest at the end of the flowering season in the warm-spring year. Therefore, phenological mismatch between flowers and pollinators is evident during warm years, which may become more prevalent in a warmer climate. To understand the mechanism of phenological mismatch in the pollination system of the alpine ecosystem, ground temperature, snowmelt regime, and life cycle of pollinators are key factors.     

The assembly of plants used as nectar sources by hummingbirds in a Cerrado area of Central Brazil

Studies on hummingbird–plant interactions commonly use a pollination approach emphasizing mutualistic relationships. But floral resources are often used opportunistically by these birds. Plant–pollinator assemblies and pollination sustainability will depend both on the well-adapted plants and other potential floral resources. The Cerrado, Neotropical savannas of Central Brazil, has ca. 7.5 % of its flora supposedly adapted to hummingbird pollination. But detailed information about flowers effectively used by hummingbirds at community level is still lacking. Hence, we recorded all plant species visited by hummingbirds, to determine how these nectariferous flowers were distributed in time and space in different plant formations of a Cerrado area, and also the hummingbird species that visit them. The study was conducted between March 2007 and December 2008 in the Panga Ecological Station. Data regarding flowering phenology, floral morphology and visitation were collected monthly. Forty-six nectariferous species from 39 genera and 17 families were recorded, most with annual flowering dynamics and tubular flowers. But only 21 species had a combination of traits fitting classic ornithophilous syndrome. For the remaining species hummingbird visitation was ascertained from observations at the study site or other sites in the region. Eight hummingbird species occurred in the area and were recorded visiting directly 36 plant species. The study area presented a relatively low number of ornithophilous plants, but a great habitat diversity and many non-ornithophilous plants that hummingbirds used as nectar sources. Therefore, in the studied Cerrado, the diversity of environments and nectariferous plants favour the maintenance of resident and migrant hummingbirds.     

POLLINATOR-MEDIATED SELECTION ON FLORAL DISPLAY AND FLOWERING TIME IN THE PERENNIAL HERB ARABIDOPSIS LYRATA

The evolution of floral display and flowering time in animal-pollinated plants is commonly attributed to pollinator-mediated selection. Yet, the causes of selection on flowering phenology and traits contributing to floral display have rarely been tested experimentally in natural populations. We quantified phenotypic selection on morphological and phenological characters in the perennial, outcrossing herb Arabidopsis lyrata in two years using female reproductive success as a proxy of fitness. To determine whether selection on floral display and flowering phenology can be attributed to interactions with pollinators, selection was quantified both for open-pollinated controls and for plants receiving supplemental hand-pollination. We documented directional selection for many flowers, large petals, late start of flowering, and early end of flowering. Seed output was pollen-limited in both years and supplemental hand-pollination reduced the magnitude of selection on number of flowers, and reversed the direction of selection on end of flowering. The results demonstrate that interactions with pollinators may affect the strength of selection on floral display and the direction of selection on phenology of flowering in natural plant populations. They thus support the contention that pollinators can drive the evolution of both floral display and flowering time.     

Pollinator conservation at the local scale: flower density,diversity and community structure increase flower visiting insect activity to mixed floral stands

Insect pollinators play a keystone role in terrestrial ecosystems. The parallel declines in plant and pollinator communities emphasizes that plant-pollinator interactions at the community level are highly relevant for biodiversity conservation. Here we examine relationships between plants and flower-visiting insects (anthophiles) at the scale of local floral patches. We conducted a visitation survey during the spring flowering season, a peak time for pollinator activity in the threatened Cape Floristic Region, South Africa. We tested floral density, diversity and composition as predictors of anthophile diversity (measured at the family/family group level) and visitation rates in multispecies stands of flowers. Although different anthophile groups responded differently, generally anthophile visitation rates and diversity were positively affected by floral density, diversity and community structure. Anthophiles were more abundant and diverse in areas with a high density and diversity of flowers. Plant community structure affected both the likelihood of occurrence and activity of anthophiles in the plots. The two mass flowering species examined here, Relhania fruticosa and Salvia chamaeleagnea, were strong determinants of anthophile activity, greatly increasing visitation rates, even though there was, on average, lower floral density and diversity. Our results show that anthophile activity is affected by highly localised, small-scale factors, namely the density and diversity of flowers and community structure. Important among these factors are patches of high quality habitat, high in floral abundance and diversity, both of which should be included in landscape-level plans for pollinator conservation, providing stepping stones for these insects in transformed landscapes.     

The flowering pattern of the perennial herb Lobularia maritima: an unusual case in the Mediterranean basin

In plant communities of the Mediterranean Basin most plant species reach their blooming peak in spring and have characteristically short flowering periods of two-three months. The perennial herb Lobularia maritima represents an exception to these characteristics, because it flowers for almost 10 months, and has its flowering peak in autumn. In this five-year study, we describe the flowering pattern of L. maritima at the population and community levels. Despite the unusually extended flowering period of L. maritima, the species showed characteristic low among-year variability in the length of the flowering period but large interannual variation in the distribution of flowers throughout the flowering period. The flowering pattern (unimodal or bimodal) of L. maritima individuals differed among the five years, suggesting that L. maritima plants are plastic enough to tailor their flowering to variable environmental conditions. We conclude that flowering phenology of L. maritima represents a very particular case in the plant community studied, and the influence of abiotic and biotic factors on the phenology of this species is discussed.     

Pollination Biology of Distylous Rubiaceae in the Atlantic Rain Forest, SE Brazil

Abstract: Data on pollination biology constitute important clues for the comprehension of pollen flow and genetic differentiation in plant populations. Pollinator type, availability and behaviour may modify morphological and mating patterns in populations of typically distylous species. This study investigates the pollination biology of four distylous species of Rubiaceae in the Atlantic rain forest, SE Brazil. Data on flowering phenology, floral lifespan, stigmatic receptivity, pollen availability, nectar volume and concentration, and pollinator activity were collected. The species studied flower sequentially throughout the wet season, and produce terminal inflorescences which bear small, tubular, diurnal, nectariferous flowers. Despite these similarities, some of the species studied are pollinated by different groups of pollinators, probably due to their distribution, availability of flowers and corolla length. On the other hand, pollinator specificity does not seem to be so important for distylous species. Long mouthparts, like those of most of the recorded pollinators, may reach lower sexual organs and, together with the self- and intramorph-incompatibilities observed, be sufficient to perform legitimate pollination and maintain levels of intermorph mating.     

Phylogenetic restriction of plant invasion in drought‐stressed environments: Implications for insect‐pollinated plant communities in water‐limited ecosystems

BackgroundPlant–pollinator community diversity has been found to decrease under conditions of drought stress; however, research into the temporal dimensions of this phenomenon remains limited. In this study, we investigated the effect of seasonal drought on the temporal niche dynamics of entomophilous flowering plants in a water‐limited ecosystem. We hypothesized that closely related native and exotic plants would tend to share similar life history and that peak flowering events would therefore coincide with phylogenetic clustering in plant communities based on expected phenological responses of plant functional types to limitations in soil moisture availability.LocationGaliano Island, British Columbia, Canada.MethodsCombining methods from pollinator research and phylogenetic community ecology, we tested the influence of environmental filtering over plant community phenology across gradients of landscape disturbance and soil moisture. Floral resource availability and community structure were quantified by counts of flowering shoots. We constructed a robust phylogeny to analyze spatial and temporal variation in phylogenetic patterns across the landscape, testing the significance of the observed patterns against a randomly generated community phylogeny. Phylogenetic metrics were then regressed against factors of disturbance and soil moisture availability.ResultsCritical seasonal fluctuations in floral resources coincided with significant phylogenetic clustering in plant communities, with decreasing plant diversity observed under conditions of increasing drought stress. Exotic plant species in the Asteraceae became increasingly pervasive across the landscape, occupying a late season temporal niche in drought‐stressed environments.Main conclusionResults suggest that environmental filtering is the dominant assembly process structuring the temporal niche of plant communities in this water‐limited ecosystem. Based on these results, and trends seen elsewhere, the overall diversity of plant–pollinator communities may be expected to decline with the increasing drought stress predicted under future climate scenarios.     

Divergent selection on flowering phenology but not on floral morphology between two closely related orchids

Closely related species often differ in traits that influence reproductive success, suggesting that divergent selection on such traits contribute to the maintenance of species boundaries. Gymnadenia conopsea ss. and Gymnadenia densiflora are two closely related, perennial orchid species that differ in (a) floral traits important for pollination, including flowering phenology, floral display, and spur length, and (b) dominant pollinators. If plant–pollinator interactions contribute to the maintenance of trait differences between these two taxa, we expect current divergent selection on flowering phenology and floral morphology between the two species. We quantified phenotypic selection via female fitness in one year on flowering start, three floral display traits (plant height, number of flowers, and corolla size) and spur length, in six populations of G. conopsea s.s. and in four populations of G. densiflora. There was indication of divergent selection on flowering start in the expected direction, with selection for earlier flowering in two populations of the early‐flowering G. conopsea s.s. and for later flowering in one population of the late‐flowering G. densiflora. No divergent selection on floral morphology was detected, and there was no significant stabilizing selection on any trait in the two species. The results suggest ongoing adaptive differentiation of flowering phenology, strengthening this premating reproductive barrier between the two species. Synthesis: This study is among the first to test whether divergent selection on floral traits contribute to the maintenance of species differences between closely related plants. Phenological isolation confers a substantial potential for reproductive isolation, and divergent selection on flowering time can thus greatly influence reproductive isolation and adaptive differentiation.     

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.