Biodiversity conservation requires strategies that encompass a variety of land uses and habitat diversity. In this study, we used sites of high ecological interest identified on the basis of the distribution of priority plant and vertebrate species to assess the implications of habitat diversity for conservation management in the Mediterranean mosaic landscape. 40% of the priority species occur in open habitats that depend on continued human presence and low-intensity land-use activities. Furthermore, 70% of the sites have more than one species habitat and 15% of sites contain more than four different species habitats. By explicitly integrating localized habitat variation, conservation planning can address the multiple conservation issues at stake in regions of high landscape diversity to provide clear and effective management objectives.

Hybridisation plays a prominent role in plant evolution due to its influence on genetic diversity, fitness and adaptive potential. We identify a case of on-going hybrid evolution of floral phenotypes in disjunct populations of Cyclamen balearicum and C. repandum subsp. repandum on Corsica and Sardinia. Hybrid populations on the two islands contain similar patterns of variation in flower colour and size but are probably at different stages in the evolutionary process of hybridisation, and differences in the frequency of floral types and flower size suggest hybrid vigour that may contribute to the dynamics and maintenance of hybrid forms. In a review of cases of hybridisation in Mediterranean plants we found an equivalent number of cases for the contemporary occurrence of mixed hybrid populations, as there are cases of homoploid hybrid species differentiation. We argue for the development of a conservation strategy for Mediterranean plants that integrates the need to protect not just pure endemic species (some of hybrid origin) but also mixed populations where adaptive variation and new species are evolving due to contemporary hybridisation.

In Europe, the abandonment of agricultural activities and traditional forest usages have led to forest spread, which eventually affects the population ecology of open habitat species. This is particularly apparent in the Mediterranean hinterland, which hosts many endemic and/or rare species. In this study we explored the effects of forest spread on the population dynamics of Paeonia officinalis, a long-lived perennial species protected in France. We also simulated the effects of different forest management regimes, by projecting the dynamics of populations under different sylvicultural cycle lengths. From 2003 to 2009, we monitored individual plants in four sites in Southern France, where each population occurs in a woodland and in an adjacent open habitat. We found that forest encroachment negatively impacted population dynamics, manifested by reduced reproduction and germination and a marked stasis of vegetative demographic stages. Using matrix models with demographic and environmental stochasticity, we showed that for two sites, populations were only viable in the open habitat but not in the woodland. In one site, populations were viable both habitats, whereas in one site, populations were non-viable in both habitats. Tree cuttings positively affected the four populations either by increasing population stochastic growth rate or by decreasing extinction rate. However, the effectiveness of forest management depended on local ecological conditions, such as the identity of the dominant tree species (evergreen vs deciduous) and sylvicultural system (coppice vs exotic plantation). Our study highlights the need for of tailor-made rather general management recommendations in perennial plant conservation.

In the Mediterranean region, the spread and densification of woodlands and shrublands subsequent to rural depopulation is a critical issue for the conservation of open-habitat plant species. Our objective was to assess the effectiveness of forest management to reduce the negative impact of forest closure on the persistence of a protected herbaceous perennial species Paeonia officinalis. Using demographic surveys from 2003 to 2008 in a woodland, open and managed habitat clear-cutted for the aim of our study, we assessed the effect of forest opening on plant performance and population dynamics. In addition, we performed a shading experiment on reproductive plants to mimic canopy closure and study its impact on plant growth and fecundity. Based on quadrat surveys and matrix models, we showed that forest cutting induced a rapid increase in plant performance and population asymptotic growth rate. Indeed, within 2 years, plant size, flower, seed and ovule numbers as well as the plant stage distribution, asymptotic growth rates and elasticity patterns shifted from values similar to those in the woodland habitat to values similar to those in the open habitat. Similarly, artificially shaded reproductive plants regressed within 2 years towards the vegetative stage. For perennial plants which have a stage in their life-cycle which allows for individual survival under unfavourable conditions such as the vegetative stage for P. officinalis, such demographic plasticity may be fundamental for their long-term persistence in temporally heterogeneous environments. Our study highlights the need to mix an experimental approach with a mid-term demographic survey in order to design efficient conservation management strategies for declining populations of rare species.

An important task in conservation biology is to assess the spatial scale pertinent for species protection since some species may require protection throughout their distribution, others in only part of their range. Once this is done, species can be correctly identified for listing at different administrative levels (e.g. continental, national, and local). Here, we propose an objective method to list taxa at nested administrative levels based on three criteria (responsibility, rarity and vulnerability). We tested the method using quantitative data on the distribution, abundance and decline of orchids in France. The proposed method enables increased protection status in regions where species’ abundance and diversity are higher, gives priority to species for which an individual administrative unit has high responsibility and allows objective integration of species decline at different administrative levels. The method also enables the integration of locally rare species at their distribution limits and avoids repetition of species listing across second-level administrative units. The use of an objective method such as this could contribute to a standardised system of priority setting that integrates the geographic scale of rarity in relation to different administrative levels for protection.

In this study we quantified variability in foliage herbivory and pre-dispersal seed predation and its effects on plant performance and demography in populations of a rare and protected perennial herb, Paeonia officinalis. An individual-based survey was performed during four years in four populations, which contained plants in both open habitat and woodland. We detected marked spatial and temporal variation among and within populations in foliage herbivory (by insects) and pre-dispersal seed predation (by insects, rodents and Roe Deer). Foliage herbivory decreased with plant demographic stages in open habitats, from seedlings to reproductive individuals, but no significant trend was detected in woodland habitats. This may be due to different demographic origin of larger vegetative plants in this habitat. Depending on demographic stage, herbivory was higher in open habitats or not significantly different between habitats. This suggests differences in herbivore abundance in different habitats within sites. Pre-dispersal seed predation remained weak and did not depend on habitat. We did not detect any consequence of foliage herbivory on seedling mortality and individual growth in our study. Our results illustrate the need to investigate plant–herbivore interactions over several years in distinct populations in order to more accurately evaluate herbivore impact on plant population dynamics.

We propose a method to establish regional-level priorities for plant species that is based on three criteria. These involve two main criteria associated with species rarity, (a) small range size and thus regional responsibility and (b) low population numbers and thus local rarity, and a third criteria associated with habitat vulnerability to distinguish species currently threatened by human activities. Application of the method shows how the three criteria can be objectively and readily quantified (using a system of equal numbers of ranks) and combined into a hierarchy of priorities. We show how this can be done by using regional responsibility as a first order priority followed successively by local rarity and habitat vulnerability. Since the criteria are closely associated with rarity, this method is easily understandable for decision makers. We illustrate the application of the method to plants (a species rich group) at two regional scales where the objectives are different. If adapted to the availability of different types of data, the three criteria could be used to produce comparable lists in different countries and regions.

In Europe, land use changes follow public policies, and particularly the Common Agricultural Policy. To predict the effect of policies on agricultural practices, landscape, and ultimately biodiversity, requires understanding of the interactions between social, economic and ecological dynamics at regional scale. We studied by means of prospective scenarios the possible effects of agricultural changes on biodiversity in a Mediterranean upland. This area is characterised by extensive grasslands that have been maintained for centuries by agriculture and are now threatened by tree and shrub encroachment. We built four scenarios that describe possible changes in agricultural EU policies by 2030. We selected 15 bird species on the basis of a high natural heritage responsibility of the study area for these species and 45 plant species on the same basis plus local rarity and habitat vulnerability. We analysed how these species were affected by the four scenarios by considering changes in their habitats. For each scenario, we analysed the driving forces that determine land use changes. Landscape dynamics was modelled with a Generalised Linear Model combining environmental and land use factors. Most of the 60 selected species depend on open habitats. Only the scenario where public support was only granted if it provided environmental services had a positive impact on open habitats and their associated biodiversity. This ‘natural heritage’ scenario was also rated positively by local stakeholders. This approach sheds light on the interest of inter/transdisciplinary studies, scenarios, and stakeholder involvement in the definition of public policies for biodiversity conservation.

Populations at the periphery of a species' range are of interest to conservation biologists because they can show marked genetic differentiation from populations at the center of a range and because of potential hybridization among rare and common species. We examined two closely related Cyclamen species. One is a narrow endemic, and the other is more geographically widespread (both protected by law in continental southern France). We used floral traits and genetic variability to test for hybridization among the species in peripheral populations of the rare species. The species co-occurred on Corsica in a disjunct, peripheral part of the distribution of the endemic species and in an ecologically marginal area for the widespread species. The two species have hybridized and the endemic species showed high levels of introgression with its widespread congener. Genetic and floral variability in sites with both species was markedly higher than in sites with a single species. Our results highlight the need for a conservation strategy that integrates hybrid populations because they represent a source of novel diversity that may have adaptive potential.

Human-induced changes in land use lead to major changes in plant community composition which have strong effects on ecosystem processes. Here, we tested the hypothesis that changes in traits of living plants induced by such changes resulted in changes in the quality and decay properties of the litter produced by the different communities. This was done in the context of a secondary succession following land abandonment in the Mediterranean region of Southern France.During the course of succession, species with high specific leaf area (the ratio of leaf area to leaf mass), low leaf dry matter content (the ratio of leaf dry mass to leaf fresh mass) and high leaf nitrogen concentration were progressively replaced by species with opposite characteristics. Accordingly, the initial litter concentrations of carbon (C) and nitrogen (N) decreased, while their C:N ratio and their hemicellulose concentration increased with time after abandonment.Early-successional communities had faster rates of litter decay and N release from litter, but these differences damped out with decomposition time. Nitrogen release from litter was related to initial litter chemical composition, particularly to its N concentration. This also held for litter decay rate, but only during the first 18 months of decomposition.Community functional parameters (i.e. trait values weighed according to the relative abundance of species) were tightly linked to initial litter N concentration, and thereby to litter decay and N loss rates. The strongest correlations were found with leaf dry matter content, which therefore appears as a powerful marker of litter properties. This provides further evidence that characteristics of living leaves persist in litter, and that some ecosystem processes can be inferred from plant functional traits.

Although several factors can limit female fertility in perennial plants, rarely have they been jointly studied in a single species over several years. In this study we experimentally manipulate seed production and simultaneously analyse the potential contribution of pollen limitation, costs of reproduction and plant size to variation in seed output over a 3-year period in the perennial herb Paeonia officinalis, in southern France. Since this rare species is threatened by forest closure in many sites we also examine the causes of female fertility variation in relation to habitat closure (open habitat vs. woodland). P. officinalis has a partial self-incompatibility system and only very low ability for autonomous self pollination in the absence of pollinators. However, supplementary pollination of individual plants in three consecutive years did not significantly increase seed production above natural levels. Forest closure was associated with a decline in ovule and seed production, which again was not due to pollen limitation since supplementary pollination had no significant effect on seed set in the woodland habitat. Comparison of the maternal fertility of plants which were previously excluded from reproduction with those which were hand pollinated to maximise seed set in two previous years produced no evidence that seed production in year three is limited by costs associated with prior reproduction. Likewise, flowering probability was not related to prior seed production but was however positively related to plant size. The absence of any influence of pollen limitation or prior reproduction on seed production suggests that sub-maximal seed production in long-lived perennial herbs may be part of a size-dependent strategy that maximises life-time seed production and fitness without compromising survival.

In the Mediterranean region of Europe, land-use changes have allowed for rapid colonisation of open habitats by woody species. As a result, it is critical to gather information on how protected species in open habitats respond to forest spread in such areas. Our objective is to quantify whether spatial heterogeneity of the vegetation associated with recent forest closure influences demographic structure and maternal fertility in a population of the protected Paeonia officinalis L. In closed woodland, adult plants of P. officinalis are almost exclusively vegetative, in open habitats seedlings are rare and on the woodland edge there is a relative over-representation of flowering plants and seedlings. Forest closure dramatically reduces flowering frequency, but has no significant effect on maternal fertility of flowering plants. The spatial aggregation of seedlings close to the maternal plants suggests that dispersal is spatially restricted. Together, these results suggest that the viability of the population requires a transitional habitat between open garrigues or grassland with spaced trees and woodland. A management programme incorporating tree and shrub thinning and cutting of parcels in rotation to maximise the length of the forest edge could maintain a habitat mosaic that favours the persistence of this species in the study site.

Paeonia officinalis L., a rare and protected species, mostly occurs in open and semi-open habitats and is often threatened by forest and shrubland spread. To explore the still undocumented dispersal features of this species, we address the following questions. What are the relative roles of ants, small rodents, and birds as diaspore removers in open habitat and woodland? Which animal groups constitute the potential disperser assemblage and how do they shape the spatial patterns of seed dispersal? Do diaspores fit the ornithochory syndrome or do they only mimic fleshy fruits? Two experiments were performed to quantify diaspore fall and diaspore removal by animal groups, above ground and on the ground. Ants did not contribute to dispersal. In open habitats, no seed removal was detected, either on follicles or once diaspores had fallen to the ground. In woodland, diaspores were weakly removed by vertebrates on follicles and were mainly removed by rodents on the ground. As a consequence, we suggest that long-distance dispersal events are very rare, weakening the possible escape into space of populations subject to forest spread. Several traits indicate that diaspores fit the ornithochory syndrome, but other traits are strongly reminiscent of mimetic diaspores deceiving bird dispersers. (C) 2007 CNRS. Journal compilation (C) 2007 The Linnean Society of London.

In this study, we quantify how reproduction and allocation to male and female functions vary in relation to plant size during two consecutive but contrasting reproductive seasons in the hermaphroditic perennial herb Paeonia officinalis. Small flowering individuals in 2003 did not flower in 2004; thus, flowering probability is related to size, with an emphasis on survival. In terms of absolute numbers of stamens and ovules, large plants allocated more resources to reproduction than did small plants by the production of additional flowering stems. In contrast, in terms of relative allocation, large plants allocated a similar proportion of resources to reproduction as did small plants. Significant size- dependent gender modification occurred only in the second year of the experiment, when larger plants were more female than small plants. Hand- pollinations performed to minimize external effects on seed set and a severe drought in 2003 may have contributed to sex allocation costs in small plants. Our results suggest that differential expression of between- year reproductive costs in relation to plant size may lead to size- dependent reproduction and be a key feature driving size- dependent gender modification in perennial plants.

We empirically assessed the long-term changes in the rare species assemblage of a Mediterranean flora, in terms of species life history traits, niche and biogeographic features, and taxonomic groups. We used a 115-year historical record of ca. 2100 plant species occurrences in a 6250 km(2) region in Mediterranean France. Species were assigned to two classes of regional abundance for the years 1886 and 2001 (rare species, i.e. exhibiting one or two occurrences vs. nonrare species), and to three classes of abundance changes during 1886-2001 (decreasing/extinct, stable, increasing/immigrant). Then, we tested whether species regional abundance and species abundance change were related to their morphological and life-history traits (life form, perenniality, height, dispersal agent, pollination mode), niche and biogeographic features (habitat specialization, level of endemism, biogeographic origin) and taxonomic group. The regional assemblage of rare species was not biologically random and significantly changed between 1886 and 2001. Species classified as rare in 1886 had a significantly higher rate of extinction in the study region during 1886-2001. The highest rate of regression/extinction was found among hydrophyte and/or water-dispersed rare species, and among annual rare species. However, herbaceous perennial, tree and wind-dispersed rare species significantly increased in abundance during 1886-2001. Rare species with Eurosiberian distributions, occurring at the southern margin of their range in the study region, dramatically declined or went extinct in the region during 1886-2001; whereas rare species with Mediterranean affinities remained significantly stable. We also found strong evidence for taxonomic patterns in species abundance and abundance changes from 1886 to 2001. The long-term biological changes documented here in the rare species assemblage of a Mediterranean flora are consistent with the predicted consequences of climate and land use changes currently occurring in the Mediterranean Basin. With the potential decline or even extinction of entire taxa and the loss of southern ecotypes of widespread Eurosiberian species, both evolutionary history and speciation potential of the Mediterranean Region could be strongly altered in future decades.

Aim  Assessing whether environmental and human factors influenced the spatial distribution and the dynamics of regionally rare plant species since the late nineteenth century, and whether these spatial and temporal patterns of rare species occurrences differ according to their chorology (level of endemism and biogeographic affinity).

A striking feature of the Mediterranean flora concerns the high rates of narrow endemism within many regions. The prevailing paradigm for the evolution of such endemism relies on an important role of geographic isolation and allopatric speciation. Until recently there has been little work on the precise ecological differences among endemic plants and their congeners in the Mediterranean region and the potential role of ecological specialisation in isolation and speciation. This paper has two objectives. First, we evaluate general ecological differences among endemic and widespread congeners in the flora of the western Mediterranean. The results and review of the literature attest to consistent patterns of ecological differentiation among endemic species and their snore widespread congeners. Second, we present results of a detailed study of morphological, molecular and ecological differentiation in a group of spring-flowering Mediterranean Cyclamen species. This study illustrates how differentiation in geographically peripheral and ecologically marginal populations may contribute to the diversification of endemic species whose contemporary distributions are disjunct from con-generic species.

The objective of this study is to examine whether habitat, herbivory and traits related to resource acquisition, resource conservation, reproduction and dispersal differ between narrow endemic plant species and their widespread congeners. We undertook pairwise contrasts of 25 ecological characteristics and biological traits in 20 congeneric pairs of narrow endemic and widespread plant species in the French Mediterranean region. Within each pair, the two species had the same life‐form, pollination mode and dispersal mode. Endemic species differed significantly from widespread congeners for a number of attributes. Endemic species occur in habitats on steeper slopes, with higher rock cover and in lower and more open vegetation than their widespread congeners. Endemic species are significantly smaller than widespread species, but show no differences in traits related to resource acquisition (specific leaf area, leaf nitrogen content, maximum photosynthetic rate) or resource conservation (leaf dry matter content). After accounting for their smaller stature, endemic species produce fewer and smaller flowers with less stigma‐anther separation and lower pollen/ovule ratios and produce fewer seeds per plant than their widespread congeners. No consistent variation in seed mass and propagule structure was found between congeneric species. Herbivory levels did not differ between congeneric species. Ecological characteristics, notably the occupation of rocky habitats with low aboveground competition, may thus have played an important role in the differentiation of narrow endemic species in the Western Mediterranean. Morphological and ecophysiological traits of narrow endemic species indicate that they are not more stress‐tolerant than their widespread congeners. Lower investment in pollen transfer and seed production suggest that local persistence is a key feature of the population ecology of narrow endemic species.

The phenology and morphology of Mediterranean plants are constrained by drought in summer and cold temperatures in winter. In this study we examine how climatic factors and phylogenetic constraints have shaped variation in the phenology and morphology of 17 species of the genus Cyclamen cultivated in uniform garden conditions. We quantify the extent to which traits differ among subgenera and thus represent conserved traits within evolutionary lineages. We also explore whether leaf, flowering and seed-release phenology are correlated among species, and thus whether variation in flowering phenology results from selection on dispersal phenology. Our results show a significant influence of subgenus membership on leaf and flowering phenology but not on morphological traits or the timing of seed release. Among-species variation in foliage height, leaf size and seed mass (but not in floral traits) is correlated with chromosome number. Leaf traits show that species with a shorter vegetative period have a higher capacity for resource acquisition. Major phenological shifts, i.e. spring vs. autumn flowering and a decoupling of leaf and flower phenology in autumnal flowering species, thus occurred prior to the diversification of species in each subgenus and not as a response to selection on dispersal timing. Leaf and flowering phenology illustrate a gradient of strategies from autumn flowering in the absence of leaves (hysteranthous species) to spring flowering with fully developed foliage (synanthous species). In the former, flowering is uncoupled from resource acquisition by simultaneous photosynthesis, indicative that hysteranthy is a response to temporal unpredictability in the onset of rain after the summer drought. Our results support the idea that whereas leaf development is controlled primarily by moisture availability and secondarily by temperature, flowering is temperature dependent, above a minimum moisture threshold. (C) 2004 The Linnean Society of London.