Reproductive capacities in the sexual forms of the gynodioecious species Thymus vulgaris L.

The reproductive capacities of the females (male-steriles or mS) of Thymus vulgaris L. have been investigated, in terms of size and weight of plants, seed production, germination ability and pollination. The data, collected during nine years under controlled conditions and in natural situations are discussed in comparison with the results known in the literature. It is suggested that the large diversity of characteristics promoting sexual polymorphism could justify the high and variable rates of mS met in natural conditions in T. vulgaris.     

The consequences of gynodioecy in natural populations of Thymus vulgaris L.

Summary Gene flow between the two sexual forms is asymmetrical in gynodioecious species: genes are transferred from male-fertile individuals (mF, hermaphrodites) to male-sterile individuals (mS, females) by pollen but from mS to MF individuals by diploid seeds. This situation is expected to cause differences in the genetic make up of mF and mS individuals. The expected differences were found in an experiment on a natural population of the gynodioecious species, Thymus vulgaris L. (thyme). The effectiveness of gynodioecy as a means of regulating heterozygosity and adaptability to various conditions of the environment is discussed.     

The spatial genetic structure of cytoplasmic (cpDNA) and nuclear (allozyme) markers within and among populations of the gynodioecious Thymus vulgaris (Labiatae) in southern France

Recent advances in molecular biology have allowed the development of techniques to contrast spatial differentiation in nuclear and cytoplasmic genes and thus provide important data on relative levels of gene flow by pollen and seed in higher plants. In this paper, we compare the spatial structure of nuclear (allozymes) and cytoplasmic (cpDNA) genes among populations of the gynodioecious Thymus vulgaris in southern France. Based on a combination of three restriction enzymes (CfoI, EcoRV, and PstI), eight chlorotypes (combination of three restriction enzyme patterns revealed by Southern hybridization of Beta vulgaris cpDNA) were identified in the 13 studied populations. One chlorotype was particularly abundant and was detected in nearly all populations. Only one chlorotype was specific to a single population. Up to four different chlorotypes were observed in some populations. An FST of 0.238 (P < 0.002) for cpDNA haplotypes indicates spatial structure of cytoplasmic genes among the studied populations. Similar patterns were found within a single young population (CAB) structured in patches and surrounded by a continuous cover of T. vulgaris where the FST is 0.546 (P < 0.002). No significant correlation between sex and chlorotype nor between cpDNA diversity and female frequency was detected. Allozyme markers showed markedly less spatial structure (FST = 0.021 among populations and 0.019 in the CAB population, P < 0.001). This difference between cpDNA and nuclear allozyme markers suggests that pollen dispersal is more important than seed dispersal both among and within populations.     

Fine-scale geographical structure of genetic diversity in inland wild beet populations

Introgression arising from crop-to-wild gene flow provides novel sources of genetic variation in plant species complexes. Hybridization within the Beta vulgaris species complex is of immediate concern; crop lineages ( B .  vulgaris ssp. vulgaris ) hybridize easily with their wild relatives ( B .  vulgaris ssp. maritima ) thereby threatening wild beet gene diversity with genetic swamping. Hybridization 'hotspots' occur in European seed production areas because inland ruderal wild beets occur and reproduce in sympatry with cultivated beets. We studied gene flow occurring between seed-producing cultivars and ruderal wild B .  vulgaris in southwestern France to determine whether feral beets, arising from unharvested cultivated seed, represent an opportunity for crop-to-wild gene flow. We surveyed 42 inland ruderal beet populations located near seed production fields for nucleo-cytoplasmic variation and used a cytoplasmic marker diagnostic of cultivated lines. Occurrence of cultivated-type cytoplasm within ruderal populations clearly reflected events of crop seed escape. However, we found no genetic signatures of nuclear cultivated gene introgression, which suggests past introgression of cultivated cytoplasm into a wild nuclear background through seed escape rather than recent direct pollen flow. Overall, patterns of genetic structure suggested that inland ruderal wild beet populations act as a metapopulation, with founding events involving a few sib groups, followed by low rates of seed or pollen gene flow after populations are established. Altogether, our results indicate that a long-lived seed bank plays a key role in maintaining cultivated-type cytoplasm in the wild and highlight the need for careful management of seed production areas where wild and cultivated relatives co-occur.     

The male-sterility polymorphism of Silene vulgaris: analysis of genetic dat from two populations and comparison with Thymus vulgaris.

Results are given of genetic studies of male sterility using plants from two natural populations from Sussex, England. Both populations have substantial frequencies of females, approximately 0.25 in population 1 and 0.60 in population 3. As in the few other gynodioecious populations studied in detail, many genetic factors are present. In population 1, there are at least two, and more likely three, different cytoplasmic types, one of which appears to produce male sterility in progeny from any hermaphrodite pollen donor; in other words restorer alleles for this cytoplasm are rare or absent from the population. The other two populations can be carried in hermaphrodites that have the dominant restorers. In population 1, there are also probably three restorer loci with complementary recessive male-sterility alleles, as well as a locus with duplicate action, which cannot produce male sterility unless the plant is also homozygous for the recessive allele at another locus. The results from population 3 are quite similar, though there was no evidence in this population for an unrestored sterility cytoplasm. A similar joint nucleocytoplasmic model with multiple restorers fits data from Thymus vulgaris.     

Evidence for lottery recruitment in Mediterranean old fields

Abstract. The hypothesis of lottery establishment ( Sale 1977 ) explains coexistence of species with similar niches through processes of stochastic recruitment. This initial idea forms the basis for a variety of mathematical models, but has not been tested empirically. This study is a field investigation of lottery establishment for plants with a seed bank, using Canonical Correspondence Analysis to compare the compositions of the vegetation and the seed bank according to different hypotheses on the mechanisms of establishment. This method was used for a data set from old fields from southern France. The weighted lottery (i.e. a random draw from the seed pool, weighted by the frequencies of each species) appeared as the best suited hypothesis to explain the high degree of similarity between the vegetation and the seed bank and the relative spatial distributions of the species. Several mechanisms are probably interacting, depending on the life histories of the species. Modelling and experimental approaches are needed to further test the hypothesis of lottery recruitment.     

Male sterility, protogyny, and pollen-pistil interference in Plantago maritima (Plantaginaceae), a wind-pollinated, self-incompatible perennial

Evolution and maintenance of male sterility in seed plants can be explained by the maternal inheritance of mitochondria, which encode the trait, and by adaptive functions that enhance female fecundity in male-sterile compared to hermaphrodite individuals. Protogyny and male sterility can independently decrease the negative effect of pollen-pistil interference in self-incompatible species. In Plantago maritima, which possesses both traits, protogyny increases seed set in hermaphrodite individuals. This is shown both by a significantly positive association between seed set and retarded dehiscence of the anthers and by a more than 50% reduction in seed set following self-pollination. Male sterility does not seem to increase seed set further, as female and hermaphrodite plants do not differ significantly in mean seed set per capsule. Bagging experiments demonstrate strong self-incompatibility in the study populations. Hence, in P. maritima male sterility seems neither to prevent selfing nor to reduce the effect of pollen-pistil interference. Females had significantly larger stigmas than hermaphrodites, but seed set varied negatively with stigma length among females, indicating that the evolution of unisexuality in P. maritima is not due to prefertilization sex allocation. I therefore conclude that the genetical system of nucleocytoplasmic determination of gender is the main cause for maintenance of male sterility in P. maritima.     

Nurse rocks influence forest expansion over native grassland in southern Brazil

Questions: Do small rocky outcrops favor forest plant colonization and establishment in grasslands? If so, what are the potential mechanisms involved in this process? Location: Araucaria forest and Campos grassland mosaic in southern Brazilian highlands (29°29′S, 50°12′W). Methods: We collected data on the density of forest woody species in plots located on rocky outcrops and in open fields subject to different management regimes. We evaluated the influence of outcrops on the density of forest plants ≤60‐cm tall, and the effects of other environmental variables and of site on plant density; we also used information on diaspore traits of the species to discuss the way in which plants disperse to the outcrops. Results: Rocky outcrops harbored a significantly higher density of forest plants than open fields, irrespective of site effect, and rock height was the best predictor for plant density on outcrops. Diaspores of all recorded species possess characteristics associated with dispersal by birds or mammals or by both. Conclusions: Small rocky outcrops markedly influence forest expansion by acting as perches for vertebrate dispersers and as nurse objects and safe sites for plants. Forest expansion starting in small outcrops possibly occurs as follows: perching of dispersers and increase of seed rain on rocks, promotion of better conditions of establishment for forest plants by nurse rocks, protection of plants sensitive to grazing and fire, and nucleation of forest vegetation.     

Founder effects and sex ratio in the gynodioecious Thymus vulgaris L.

Thymus vulgaris is a gynodioecious species (in which females and hermaphrodites coexist) with a highly variable frequency of females among natural populations (5–95%) and a high average female frequency (60%). Sex determination involves both cytoplasmic genes responsible for male sterility, i.e. the female phenotype, and specific nuclear factors responsible for the restoration of male fertility, and thus a hermaphrodite phenotype. In this study, molecular markers of the mitochondrial genome have been used to quantify the cytoplasmic diversity in 11 clumps of individuals observed in four recently founded populations. The very low diversity within patches in conjunction with the strong diversity among patches strongly suggests that clumps of individuals are the result of single matrilinear families. In clumps that contain mainly females, all the analysed females showed the same cytoplasmic pattern. This pattern differed from that shown by neighbouring hermaphrodites, indicating that the determination of sex is locally cytoplasmic. A comparison of genetic diversity before and after fire in one population showed that disturbances may cause a reduction in genetic diversity and a concurrent induction of local cytoplasmic determination of sex. Such cytoplasmic determination of sex in colonizing populations, together with the greater seed set of females, may largely improve the colonizing ability of the species.     

Nest site selection of the Kelp gull (Larus dominicanus) in the Beagle Channel, Tierra del Fuego, Argentina

Nest site selection of Kelp gulls Larus dominicanus breeding in Conejo and Bridges islands, Tierra del Fuego, Argentina, was assessed between 14 and 30 December 2005 to describe variability in nest site features and determine variables affecting nest choice. Fourteen microhabitat variables were quantified at 40 nest sites and at 40 random points in each island during the late incubation stage. Nests at the two colonies were placed on different kinds of substrates across areas with varying degrees of cover provided either by rocks or vegetation. Despite the variability observed in nest site features, rock cover and rocky substrates were the main factors determining nest site selection. At Conejo Island, nest sites presented more rock cover, less percentage of vegetation cover and of vegetation debris substrate, and were placed further from the nearest vegetation in comparison to random points. Similarly, gulls at Bridges Island selected nest sites with more rock cover which were placed mainly on rocky and vegetation substrates. This contrasts with results previously obtained in Argentina, which indicated that vegetation is a key factor influencing Kelp gull nest site selection. Over 80% of the nests at both study colonies were placed on the northern slopes of the islands, relatively more protected from the strong prevailing southwest winds, and nests at both colonies tended to be more protected on their southern side either by rocks or vegetation. Results obtained at Tierra del Fuego confirm the plasticity of microhabitat use by Kelp gulls, and their ability to take advantage of nesting sites according to availability and local environmental factors.     

Colonization of vegetation-rich moraines and inference of multiple sources of colonization in the High Arctic for Salix arctica

Vegetation-rich patches in the High Arctic may serve as a significant source for vegetation reconstruction in the climate changes. Diversity and colonization, however, of such potential source populations in the High Arctic has rarely been studied. We examined chloroplast sequence variation in Salix arctica, a key species in the Canadian High Arctic, from four adjacent glacial moraines of differing ages on Ellesmere Island, Canada, as well as two other populations located at the center and southern end of the species’ range. The estimated ages of the moraines varied from 35,000 to 250 years old. The older moraine populations showed higher within-population genetic variation compared with the other moraine populations, which is generally attributed to differences in establishment age associated with plant densities among moraines. The moraines with smaller plant density had lower genetic diversity and had no private haplotypes, indicating the local population size and genetic diversity may not be recovered within a few thousand years. This suggests seed dispersal at a local scale may be limited even in species with high velocity of seed dispersal, and that High Arctic vegetation-rich patches may serve as significant source populations for sustaining local genetic diversity. In addition, the three regions we observed comprised an evolutionarily distinct lineage and significant population differentiation. This implies multiple sources for the colonization during the most recent deglaciation, resulting in the current wide distribution. Local as well as range-wide processes of colonization would be essential to understand vegetation responses in High Arctic to the environmental changes.     

An explicit test for the contribution of environmental maternal effects to rapid clinal differentiation in an invasive plant

Population differentiation of alien invasive plants within their non‐native range has received increasingly more attention. Common gardens are typically used to assess the levels of genotypic differentiation among populations. However, in such experiments, environmental maternal effects can influence phenotypic variation among individuals if seed sources are collected from field populations under variable environmental regimes. In the present study, we investigated the causes of an altitudinal cline in an invasive plant. Seeds were collected from Senecio inaequidens (Asteraceae) populations along an altitudinal gradient in southern France. In addition, seeds from the same populations were generated by intra‐population crossings in a climatic chamber. The two seed lots were grown in a common garden in Central Belgium to identify any evidence of environmentally induced maternal effects and/or an altitudinal cline in a suite of life‐history traits. Results failed to detect any environmental maternal effects. However, an altitudinal cline in plant height and above‐ground biomass was found to be independent of the maternal environment.     

Do seed transfer zones for ecological restoration reflect the spatial genetic variation of the common grassland species Lathyrus pratensis?

A common ecological restoration approach is the reestablishment of vegetation using seed mixtures. To preserve the natural genetic pattern of plant species local seed material should be used. Consequently, seed transfer zones (seed production areas and seed provenance regions) have been delineated for ecological restoration in Germany. Although it is assumed that these transfer zones represent genetic variation, there remains a lack of empirical data. In this study, we analyzed whether seed transfer zones reflect the genetic variation of the common grassland species Lathyrus pratensis. We sampled 706 individuals from 37 populations in Bavaria, Germany and analyzed genetic variation using amplified fragment length polymorphisms. In our study, we observed higher levels of genetic variation and fragment rarity in the southern Bavarian populations compared to northern populations. Our analyses revealed a strong genetic differentiation between southern and northern Bavarian populations delineated along the Danube River. However, seed production areas and seed provenance regions reflected genetic variation of L. pratensis only to a limited degree. Our study illustrates that the level of genetic variation within populations strongly depends on population history. Furthermore, the geomorphological and climatic attributes, which have been used to delineate seed provenance regions, do not reduce gene flow among populations. Seed collections for gene banks and seed production should comprise seeds from populations in southern and northern Bavaria representing the strong genetic variation between both regions, but prioritize southern populations due to higher levels of variation.     

Buried seed populations on the volcano Mt. Usu,northern Japan,ten years after the 1977–78 eruptions

In 1987, a study on buried seed populations was conducted in the crater basin of Mt. Usu, a volcano located in northern Japan, where the vegetation had beeen almost completely destroyed by eruptions in 1977 and 1978. The former vegetation had consisted of grassland and broad-leaved forest. In the areas formerly occupied by this grassland and forest, 2128.0 and 1985.3 seeds per square meter, respectively, were extracted from 12 blocks of the former topsoil using a floattion method. This revealed that many seeds were still viable even after ten years of burial under thick volcanic deposits. The seeds were distributed more in sandy soil than in rocky soil of the former topsoil. Twenty-five species, most of which favored grassland, were detected in both the former grassland and forest. From comparison of α-and β-diversities between the grassland and forest, the structure of the buried seed population was considered likely to have been more diverse in the forest than in the grassland. The determinants of composition of the buried seed populations were discussed with reference to the former vegetation and soil characteristics.     

Population trends and spatial synchrony in peripheral populations of the endangered Lesser grey shrike in response to environmental change

Regional synchronization in species dynamics as well as particular ecological and demographic characteristics of peripheral populations poses special challenges for conservation purposes, particularly under the current scenario of global climate change. Here, we study the population trend and spatial synchrony of several peripheral populations of the endangered Lesser grey shrike Lanius minor at the western limit of its breeding range (southern France and northeast Spain). In an attempt to ascertain the effect of environmental change on the decline of the species we also look for evidence of climate changes in the breeding and wintering area of this shrike and related effects on vegetation by using the normalized difference vegetation index (NDVI). We found that the interannual fluctuations of the peripheral populations in France and Spain are strongly correlated, therefore suggesting that their decline can be under the influence of a common factor. We obtained clear evidence of climatic change (an increased thermal oscillation) in one peripheral population that could have resulted in a decrease of the NDVI index in the area. Our study finds correlational evidence that climatic variables in the breeding area may account for fluctuations in abundances of some populations and that environmental conditions experimented by some population could influence the fate of the neighboring populations. Our results indicate that the studied peripheral populations are spatially synchronized, so that conservation efforts should be applied at a large-scale encompassing all the isolated populations at the western border of the range of the species in the Mediterranean area.     

Evidence for gene flow via seed dispersal from crop to wild relatives in Beta vulgaris (Chenopodiaceae): consequences for the release of genetically modified crop species with weedy lineages

Gene flow and introgression from cultivated to wild plant populations have important evolutionary and ecological consequences and require detailed investigations for risk assessments of transgene escape into natural ecosystems. Sugar beets (Beta vulgaris ssp. vulgaris) are of particular concern because: (i) they are cross-compatible with their wild relatives (the sea beet, B. vulgaris ssp. maritima); (ii) crop-to-wild gene flow is likely to occur via weedy lineages resulting from hybridization events and locally infesting fields. Using a chloroplastic marker and a set of nuclear microsatellite loci, the occurrence of crop-to-wild gene flow was investigated in the French sugar beet production area within a 'contact-zone' in between coastal wild populations and sugar beet fields. The results did not reveal large pollen dispersal from weed to wild beets. However, several pieces of evidence clearly show an escape of weedy lineages from fields via seed flow. Since most studies involving the assessment of transgene escape from crops to wild outcrossing relatives generally focused only on pollen dispersal, this last result was unexpected: it points out the key role of a long-lived seed bank and highlights support for transgene escape via man-mediated long-distance dispersal events.     

Seed viability in declining populations of Caladenia rigida (Orchidaceae): are small populations doomed?

Despite comparatively good rates of pollination and seed production, some populations of the endangered terrestrial orchid Caladenia rigida continue to decline. To determine whether seed quality may be limiting reproductive potential, we assessed seed viability among declining populations of C.?rigida (in the southern part of its distribution) and among populations that are regarded as stable (in the northern part of its distribution). We also compared differences in seed viability to plant traits, population size and habitat characteristics (soil properties, canopy cover, presence of proximate vegetation). Seed capsules from southern populations were significantly smaller, with only 9% of seeds being viable, compared to 36% in capsules from northern populations. Soil phosphorus concentrations differed between regions, but other habitat characteristics did not correlate with seed viability. Using calculations based on seedling recruitment data from other Caladenia species, we predict that seed output is insufficient to ensure the long-term persistence of the smallest C.?rigida populations.     

What's new about old fields? Land abandonment and ecosystem assembly

Environmental and socio-economic changes are leading to increased levels of land abandonment worldwide. The assembly of plant communities on old fields has informed much ecological theory, which in turn has facilitated efforts at ecological restoration. The interaction of the cultivation legacy with inherent soil and vegetation characteristics will determine the dynamics of plant community assembly on old fields and indicate the level of effort required to restore historical vegetation states. The abandonment of traditional agricultural lands in some areas will create old fields that require limited or no restoration. Yet intensification of agriculture and rapid environmental change will lead to increasing numbers of old fields that show little recovery towards an historic vegetation state. The restoration of these old fields will pose significant scientific and policy challenges.     

Soil disturbance,vegetation cover and the establishment of the exotic shrub <Emphasis Type="Italic">Pyracantha coccinea</Emphasis> in southern France

We evaluate the mechanisms that determine the establishment of the non-indigenous shrub Pyracantha coccinea (Rosaceae) in the Montpellier region of southern France. P. coccinea establishes in abandoned agricultural fields in this region; yet, despite its high propagule pressure, it has not become a widespread invasive. We hypothesized that the disturbance conditions prevailing in abandoned agricultural fields right after abandonment may enhance the emergence, survival and growth of P. coccinea, but that shortly after abandonment colonizing vegetation prevents further establishment of this species. We conducted a field experiment to evaluate this hypothesis, studying the response of seedling emergence and growth of P. coccinea to soil and vegetation disturbance. Our results show that both lack of vegetation cover and soil disturbance promote the emergence of seedlings of P. coccinea. Thus, the disturbance conditions prevailing in abandoned agricultural fields seem crucial to allow establishment of this species. However, other factors such as lack of summer dormancy and seed predation might explain why this species has not become a widespread invasive.