Seed ecology of a Mediterranean perennial herb with an exceptionally extended flowering and fruiting season

Lobularia maritima is a Mediterranean short-lived herb with a flowering and fruiting season that lasts for ten months. Previous studies have shown that recruitment in periods other than autumn of the flowering season has few demographic implications; that is contributes little to the population growth rate. Since environmental conditions in periods other than autumn would allow recruitment, we examined to what extent the seed ecology of L. maritima accounts for recruitment shortage for the greater part of the year. To this end, we studied the effects of selfing and outcrossing on seed production and germination, within- and between-year variation in seed mass and germination, seed characteristics in the soil seed bank throughout the year, and the effect of temperature as a factor controlling seed germination. Results indicate that selfing does not decrease recruitment, and thus the observed changes in visitation rate and pollinator composition throughout the year cannot account for differences in recruitment. Germinability decreases throughout the year, suggesting a possible cost in reproduction associated with extended flowering. L. maritima has a transient seed bank whose seeds also experience a decrease in their germination throughout the year. Finally, temperature affects seed germination patterns, indicating the existence of quiescence mechanisms that prevent germination in the months prior to the summer drought. Overall, the results obtained support and, at least partly, explain the recruitment patterns of L. maritima observed in the field.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 273–280.     

Temporal variation in the female components of reproductive success over the extended flowering season of a Mediterranean perennial herb

We examined the reproductive success of the perennial herb Lobularia maritima during its extended flowering and fruiting season. The within- and between-year variability of the female components of reproductive success (from flower, fruit and seed production to seed survival, seed germination and seedling establishment) were analysed during four flowering seasons. All the components of reproductive success studied showed a significant within- and between-year variation. September was the period of the year with the maximum values of flower and fruit production, and the highest germination and establishment rates. Nevertheless, seed losses due to both predispersal and postdispersal seed predation during this period were also the highest, seriously reducing seed output in this period. On the other hand, in those periods in which seed production was low, i.e. January and May, the percentage of seeds lost to seed predators was the lowest. Reproductive success in each period of the flowering season was estimated using a simple demographic model, in which the information concerning all the components already calculated was integrated. The two variables used to estimate reproductive success in each period, i.e. the number of new individuals produced per plant and the probability of a seed becoming an adult plant, showed relatively small differences over the year. These results suggest a counter-balance of the different components of reproductive success in this species, with favourable and unfavourable periods for the different components being compensated during its extended flowering season.     

Life-history variation in contrasting habitats: flowering decisions in a clonal perennial herb (Veratrum album)

Quantifying intraspecific demographic variation provides a powerful tool for exploring the diversity and evolution of life histories. We investigate how habitat-specific demographic variation and the production of multiple offspring types affect the population dynamics and evolution of delayed reproduction in a clonal perennial herb with monocarpic ramets (white hellebore). In this species, flowering ramets produce both seeds and asexual offspring. Data on ramet demography are used to parameterize integral projection models, which allow the effects of habitat-specific demographic variation and reproductive mode on population dynamics to be quantified. We then use the evolutionarily stable strategy (ESS) approach to predict the flowering strategy-the relationship between flowering probability and size. This approach is extended to allow offspring types to have different demographies and density-dependent responses. Our results demonstrate that the evolutionarily stable flowering strategies differ substantially among habitats and are in excellent agreement with the observed strategies. Reproductive mode, however, has little effect on the ESSs. Using analytical approximations, we show that flowering decisions are predominantly determined by the asymptotic size of individuals rather than variation in survival or size-fecundity relationships. We conclude that habitat is an important aspect of the selective environment and a significant factor in predicting the ESSs.     

A fatal growth pattern and ways suspected of postponing death: corm dynamics in the perennial herb Corydalis cava

The forest herb Corydalis cava is the only known species with a corm where tissue is added annually as a layer on the outside, and where the older central parts rot away, i.e. the corm becomes an expanding sphere. This growth pattern creates energetic problems for ageing plants. However, older corms change their shape in a way that saves biomass. Growth of corm with age was here modelled introducing different corm traits saving mass and thus postponing death. The largest corm in the study patch had a volume of 60 ml. If such a corm had remained a solid sphere, its age would be estimated to be 8.3 years. In reality, 4 year old corms became hollow and later an opening appeared in their basal part. In addition, corms ≥ 7 years became more conical. A conical corm with a volume of 60 ml was estimated to be 15.9 years old. Volume and shape of corm thus made it possible to age individual plants. The age structure of the plants in the study patch was described and discussed in relation to the overall dynamics of the metapopulation. The patch consisted of a few strong age classes (from 5 to 7 years old). Any recruiting juvenile age classes were almost completely missing.     

Phenotypic selection and response to selection in Lobularia maritima: importance of direct and correlational components of natural selection

By using selection differentials, gradients and structural equation modelling (SEM), I have quantified the phenotypic selection acting on Lobularia maritima (Cruciferae) flower size, display, colour and density, using data on lifetime female fitness. Furthermore, by analysing the resulting F₁ generation in field and greenhouse conditions, I estimated the actual intergenerational change in the value of these traits. Both pollinators preferred plants with many and large flowers. Strong directional selection for increased flower display was found in all years of the study, regardless of the technique used. Indirect selection due to a high significant correlation with flower display occurred on flower colour and size. SEM showed that pollinators played only a minor role in this observed phenotypic selection. The analysis of the phenotypes of F₁ plants showed that flower display actually increased across generations. In addition, white flowers were significantly more frequent in the offspring population than in the parental one, mostly due to the association between flower display and white coloured flowers. This suggests that both direct and indirect selection can play a role in the evolution of correlated traits in this crucifer.     

Habitat quality and among-population differentiation in reproductive effort and flowering phenology in the perennial herb <Emphasis Type="Italic">Primula farinosa</Emphasis>

In heterogeneous environments, selection on life-history traits and flowering time may vary considerably among populations because of differences in the extent to which mortality is related to age or size, and because of differences in the seasonal patterns of resource availability and intensity of biotic interactions. Spatial variation in optimal reproductive effort and flowering time may result in the evolution of genetic differences in life-history traits, but also in the evolution of adaptive phenotypic plasticity. The perennial herb Primula farinosa occurs at sites that differ widely in soil depth and therefore in water-holding capacity, vegetation cover, and frost-induced soil movement in winter. We used data from eight natural populations and a common-garden experiment to test the predictions that reproductive allocation is negatively correlated with soil depth while age at first reproduction and first flowering date among reproductive individuals are positively correlated with soil depth. In the common-garden experiment, maternal families collected in the field were grown from seed and monitored for 5 years. In the field, reproductive effort (number of flowers in relation to rosette area) varied among populations and was negatively related to soil depth. In the common-garden experiment, among-population differences in age at first reproduction, and reproductive effort were statistically significant, but relatively small and not correlated with soil depth at the site of origin. Flowering time varied considerably among populations, but was not related to soil depth at the site of origin. Taken together, the results suggest that among-population variation in reproductive effort observed in the field largely reflects phenotypic plasticity. They further suggest that among-population differentiation in flowering time cannot be attributed to variation in environmental factors correlated with soil depth.     

Regional and local variation in seedling emergence,mortality and recruitment of a perennial herb in Mediterranean mountain habitats

Spatial patterns of seedling regeneration of woody Mediterranean plants are widely documented, with seedlings and juveniles growing almost exclusively under shrub and tree canopy. Neighbourhood habitat amelioration and consequent facilitative effects on recruitment have been extensively suggested as the major determinant of such spatial pattern for Mediterranean vegetation. Much less is known on recruitment patterns of perennial herbs. As herbs differ from woody plants in many relevant ecological aspects and in their life cycles, we would expect particularities in their recruitment dynamics. We analysed the spatial (regional, local and fine-scale) variation in environment (herb cover, litter depth, air temperature and irradiance) and its relationship with seedling emergence, mortality, and recruitment of the perennial herb Helleborus foetidus (Ranunculaceae) during 2 years. We conducted the study in three distant localities in south-eastern Iberian Peninsula (defining the regional scale). Local scale was defined in terms of microhabitat type based on three categories of vegetation cover (under evergreen shrubs or trees, under deciduous spiny-shrubs, and in open interspaces). Finally, fine-scale variation was defined in terms of 1 m² sampling plots. Our results showed regional, local and fine-scale variations in recruitment, with the major source of variation changing across scales. Seed input was important in determining differences in recruitment both at the largest (regional) and the finest scale. Environment had minor importance in shaping differences in recruitment at these two scales. At the local scale, variation between microhabitats was mainly related to differences in seedling survival through facilitative effects (alleviation of water and irradiance stresses) of shrub and tree cover, although such facilitative effect was not consistent in all localities or years. Our study thus points to a similarity in the spatial patterns of recruitment (and in the environmental factors shaping such patterns) between Mediterranean woody plants and perennial herbs, but much more effort is needed to assess the generality of such similarity.     

Response of bee-flies to the shape and pattern of model flowers: implications for floral evolution in a Mediterranean herb

1. The functional significance of floral traits in Linum pubescens (Linaceae), a Mediterranean annual that is pollinated almost exclusively by the bee-fly Usia bicolor (Bombyliidae), was investigated. The flies feed on both pollen and nectar, and use the flowers as mating rendezvous sites in the afternoon.
2. Choice experiments with model flowers were used to determine the response of U. bicolor to visual cues, such as shape, size, colour and pattern.
3. Bee-flies strongly preferred models with a dissected outline over models with a simple outline. They also preferred pink models over other colours, and larger models over smaller models.
5. Flies landing on models with converging lines ('nectar guides') tended to follow the lines to the point where they meet in the centre of the model, while flies landing on plain models showed undirected behaviour, often moving to the edge of the model.
6. Flies were strongly attracted to flowers of L. pubescens which had a fly glued on to one of the petals, as well as flowers with a dark spot painted onto one of the petals. In addition, models with a dark spot were strongly preferred over plain models during the afternoon when flies exhibit mating behaviour. This evidence suggests that the dark centre of the L. pubescens flower may function as an attractant to mate-seeking bee-flies.     

Classification of habitats highlights priorities for conservation policies: The case of Spanish Mediterranean tall humid herb grasslands

The Mediterranean grasslands of Molinio-Holoschoenion are one of the lesser known natural habitat types of interest for conservation in the European Community. A study on their content with regard to their plant-communities and interest for plant conservation is conducted on a regional scale for the first time. A new comprehensive classification is proposed for this habitat type in Spain (Iberian Peninsula and Balearic and Canary Islands) where they have their highest European diversity. Twenty-five plant-community types are floristically identified by k-means clustering. Cluster analysis reveals a major hierarchical aggregation in two vegetation groups: (i) rush meadows of Scirpoides holoschoenus (L.) Soják; and (ii) communities of herbs and forbs dominated by Molinia caerulea (L.) Moench subsp. arundinacea (Schrank) H. Paul and/or Schoenus nigricans L. These groups are associated with different positions along a soil moisture gradient where rush formations occupy the positions farthest from the water table. Cluster Analysis also shows an uneven distribution of species with conservation interest. The highest content in endangered species is found in halophilous rushes. Grasslands on mineralised soils host the highest number of vulnerable species. Spanish Molinio-Holoschoenion plant communities were ranked using three criteria (regional responsibility, local rarity and habitat vulnerability) in order to set conservation priorities. The highest conservation values were achieved by coastal rush communities developing on dune slacks, and by forbs and rushes in the Baetic System growing on banks of streams with oligotrophic waters.     

The effects of large herbivores on the landscape dynamics of a perennial herb

Background and Aims

Models assessing the prospects of plant species at the landscape level often focus primarily on the relationship between species dynamics and landscape structure. However, the short-term prospects of species with slow responses to landscape changes depend on the factors affecting local population dynamics. In this study it is hypothesized that large herbivores may be a major factor affecting the short-term prospects of slow-responding species in the European landscape, because large herbivores have increased in number in this region in recent decades and can strongly influence local population dynamics.

Methods

The impact of browsing by large herbivores was simulated on the landscape-level dynamics of the dry grassland perennial polycarpic herb Scorzonera hispanica. A dynamic, spatially explicit model was used that incorporated information on the location of patches suitable for S. hispanica, local population dynamics (matrices including the impact of large herbivores), initial population sizes and dispersal rate of the species. Simulations were performed relating to the prospects of S. hispanica over the next 30 years under different rates of herbivory (browsing intensity) and varying frequencies of population destruction (e.g. by human activity).

Key Results

Although a high rate of herbivory was detected in most populations of S. hispanica, current landscape-level dynamics of S. hispanica were approximately in equilibrium. A decline or increase of over 20 % in the herbivory rate promoted rapid expansion or decline of S. hispanica, respectively. This effect was much stronger in the presence of population destruction.

Conclusions

Browsing by large herbivores can have a dramatic effect on the landscape dynamics of plant species. Changes in the density of large herbivores and the probability of population destruction should be incorporated into models predicting species abundance and distribution.     

早春草本植物开花物候期对城市化进程的响应——以北京市为例

植物物候对城市化的响应是全球变化与城市生态环境效应研究的重要内容。2012年3月到6月,结合气象观测数据,对北京市西北向城市化梯度上7种早春草本植物开花物候期进行观测与研究,发现温度因子和早春草本植物开花物候期均随城市化梯度发生变化,即越靠近城市中心区,温度和积温累计值越高;早春草本植物开花物候期出现时间越早,平均提前2—4 d;但开花期持续时间与开花速率并不随城市化梯度发生明显变化。此外,研究发现北京市7种早春草本植物开花期对5℃积温变化响应最为敏感;开花期提前时间梯度变化显著性与生活型密切关联,多年生草本植物对城市化梯度变化的响应比一年生或一二年生草本植物明显。未来城市植物物候期研究中,应更关注城市化进程中土地利用/覆被变化与热岛效应对城市气候及植物生理生态特征的累积影响特征,以期进一步揭示植物物候期对城市化及气候变化的响应规律。     

Decoupling of reproduction and growth: an unusual pattern in the life cycle of the Mediterranean geophyte Narcissus serotinus

The unpredictable seasonality of the Mediterranean climate imposes on the development of specific life‐cycle phenologies such as those seen in geophyte species that have a period of dormancy to avoid stressful abiotic conditions. To date, two life‐cycle strategies are recognized in Mediterranean geophytes. In some species, growth and reproduction occur in different seasons (hysteranthous geophytes), whereas in others these two processes occur at the same time (synanthous geophytes). A study of the life cycle of the autumnal‐flowering Narcissus serotinus in three populations over two consecutive years confirmed that neither the hysteranthy nor the synanthy strategy could be applied to this geophyte. Rather, reproduction and growth occur simultaneously, but in different plants. A strong correlation was found between bulb weight and the presence of reproductive or vegetative plants.     

Cost of reproduction in the perennial herb Asphodelus albus (Liliaceae)

The cost of reproduction has been studied in two populations of the polycarpic herb Asphodelus albus under natural conditions The percentage of plants with flowers was determined in four sites and varied markedly among them The occurrence of reproduction was size-dependent, increasing flowering probability with plant size The cost of reproduction was assessed in terms of modular growth in reproductive plants relative to modular growth in vegetative ones I compared the modular growth of vegetative and reproductive plants considering two different densities m each of two populations Neither incidence of flowering nor modular growth were affected by density Flowering plants exhibited a withinramet demographic cost (in terms of modular growth) relative to non-flowering ramets in one population but not in the other This cost was greater in larger plants These results were concordant with the occurrence of flowering at both sites Both populations exhibited size-dependent patterns of allocation to reproduction, but no significant relationships were found between allocation to reproduction and cost of reproduction The data presented demonstrate differences in the cost of reproduction within a species This cost might determine whether a plant begins the reproduction, but probably have no effect on the reproductive allocation since the weight of the reproductive structures was not related to modular growth     

Sex choice in plants: facultative adjustment of the sex ratio in the perennial herb Begonia gracilis

Sex allocation theory predicts that reproducing individuals will increase their fitness by facultatively adjusting their relative investment towards the rarer sex in response to population shifts in operational sex ratio (OSR). The evolution of facultative manipulation of sex ratio depends on the ability of the parents to track the conditions favouring skewed sex allocation and on the mechanism controlling sex allocation. In animals, which have well-developed sensorial mechanisms, facultative adjustment of sex ratios has been demonstrated on many occasions. In this paper, we show that plants have mechanisms that allow them to evaluate the population OSR. We simulated three different conditions of population OSR by manipulating the amount of pollen received by the female flowers of a monoecious herb, and examined the effect of this treatment on the allocation to male vs. female flowers. A shortage of pollen on the stigmas resulted in a more male-skewed sex allocation, whereas plants that experienced a relatively pollen rich environment tended to produce a more female-skewed sex allocation pattern. Our results for Begonia gracilis demonstrate that the individuals of this species are able to respond to the levels of pollination intensity experienced by their female flowers and adjust their patterns of sex allocation in accordance to the expectations of sex allocation theory.     

Negative evidence of local adaptation to the establishment conditions in a perennial herb

The differential adaptation of populations of the same species to their local environmental conditions through divergent selection, known as local adaptation, is a key step in the process of diversification of species. Here, we explore the local adaptation of the perennial mountain herb Helleborus foetidus to variable environmental conditions of seedling emergence and establishment at two different spatial scales (habitats and regions) with special attention to the role of physical and chemical soil properties. The possibility of local adaptation was evaluated under the ??local versus foreign?? and the ??home versus away?? criteria. Reciprocal sowing experiments were carried out by cross-sowing seeds among habitats and regions, controlling for maternal effects by means of seed mass, and recording seedling emergence and survival. Several topsoil properties were measured linked to each sowing point. Only partial patterns of local adaptation were found, which were insufficient to eventually state the existence of local adaptation at any spatial scale or under any criteria assessed. Here, we discuss how soil properties and selection on seed size may be related to the non-achievement of local adaptation criteria. Negative evidence of local adaptation seems to be due to a congruency in the selective pressures exerted by the different soil environments on seedling emergence and survival.     

The life history of Ranunculus glacialis, an arctic-alpine perennial herb, in Finnish Lapland

Dynamics and performance of established individuals of Ranunculus glacialis at the time of peak flowering were monitored in a mountain area in Finnish Lapland for five years (1982–1986). The plants reproduced only sexually, and their spatial distribution in 40 1-m² quadrats was clumped. During the study period the mean density rose from 8.5 to 10.3 individuals m⁻². The mean annual growth rate and birth rate of the population was 4% and 8.5%, respectively. The mean annual death rate of established individuals was 3.7%. The estimated mean life expectancy of an established individual was 27 yr. Death rate per quadrat was positively related to R. glacialis density, but birth rate per quadrat was negatively related to the percentage cover of stones per quadrat. High R. glacialis density per quadrat had negative effects, but high nitrogen availability had positive effects on the number of leaves per individual. An important factor reducing flowering was small rodent herbivory. In the same genets, the correlation between the number of leaves per individual in year t and the number of flowers per individual in year t+4 was significant. The below-ground biomass of R. glacialis comprised about 70% of the total biomass, and there was a close correlation between the above- and the below-ground biomass of an individual. The individuals flowered abundantly each year and produced a great number of small-sized achenes.