Petr Zákravský

Higher genetic diversity in recolonized areas than in refugia of Alnus glutinosa triggered by continent‐wide lineage admixture

Genetic admixture is supposed to be an important trigger of species expansions because it can create the potential for selection of genotypes suitable for new climatic conditions. Up until now, however, no continent‐wide population genetic study has performed a detailed reconstruction of admixture events during natural species expansions. To fill this gap, we analysed the postglacial history of Alnus glutinosa, a keystone species of European swamp habitats, across its entire distribution range using two molecular markers, cpDNA and nuclear microsatellites. CpDNA revealed multiple southern refugia located in the Iberian, Apennine, Balkan and Anatolian Peninsulas, Corsica and North Africa. Analysis of microsatellites variation revealed three main directions of postglacial expansion: (i) from the northern part of the Iberian Peninsula to Western and Central Europe and subsequently to the British Isles, (ii) from the Apennine Peninsula to the Alps and (iii) from the eastern part of the Balkan Peninsula to the Carpathians followed by expansion towards the Northern European plains. This challenges the classical paradigm that most European populations originated from refugial areas in the Carpathians. It has been shown that colonizing lineages have met several times and formed secondary contact zones with unexpectedly high population genetic diversity in Central Europe and Scandinavia. On the contrary, limited genetic admixture in southern refugial areas of A. glutinosa renders rear‐edge populations in the Mediterranean region more vulnerable to extinction due to climate change.

Ecology of two cytotypes ofButomus umbellatus II. Reproduction, growth and biomass production

Generative and vegetative reproduction of diploid and triploidButomus umbellatus L., and growth and biomass production of both cytotypes under two different nutrient levels were compared.Seedling survival was studied under controlled conditions in a growth chamber; the response of plants to different nutrient conditions was studied in experimental garden.Both cytotypes do not differ in seed germination and seedling survival. Triploids produce more aboveground and underground biomass, more numerous lateral rhizome buds, and have significantly higher flowering stalks. Low generative reproduction (limited seed production) in triploids is compensated for by more intensive vegetative reproduction. High nutrient level appeared to be stressful for plants of both cytotypes: it limits plant growth and causes plant mortality. Triploids are more viable than diploids in this case, which may be important for their survival under conditions of high trophic level.

The biology of Butomus umbellatus in shallow waters with fluctuating water level

Butomus umbellatus L. is a plant species typical of littoral communities of river and stream shores. It can form continuous stands in shallow reservoirs with fluctuating water level. Their expansion is promoted by: (a) intensive vegetative reproduction of plants, (b) crowded sprouting from rhizome fragments on emerged pond bottom, (c) shallow water layer in the year following summer drainage. Expansion of B. umbellatus depends on ploidy level: two cytotypes were found in the Czech and Slovak Republics, differing in their reproductive ability. Seed production of triploids is strongly limited (they are self-incompatible within clones), while diploids can be fully fertile. Nevertheless, even in diploids, the efficiency of seed reproduction under natural conditions is low. Triploids spread by intensive vegetative reproduction, which is decisive for clonal growth of populations and their regeneration after scraping of bottom surface. During seasonal development, maximum of aboveground biomass is produced in early summer, while underground biomass increases till autumn. Growth of the plants is limited by cutting before maximum underground biomass is attained, or by duck grazing.

Migration Patterns of Subgenus Alnus in Europe since the Last Glacial Maximum: A Systematic Review

Background/Aims Recently, new palaeoecological records supported by molecular analyses and palaeodistributional modelling have provided more comprehensive insights into plant behaviour during the last Quaternary cycle. We reviewed the migration history of species of subgenus Alnus during the last 50,000 years in Europe with a focus on (1) a general revision of Alnus history since the Last Glacial Maximum (LGM), (2) evidence of northern refugia of Alnus populations during the LGM and (3) the specific history of Alnus in particular European regions. Methodology We determined changes in Alnus distribution on the basis of 811 and 68 radiocarbon-dated pollen and macrofossil sites, respectively. We compiled data from the European Pollen Database, the Czech Quaternary Palynological Database, the Eurasian Macrofossil Database and additional literature. Pollen percentage thresholds indicating expansions or retreats were used to describe patterns of past Alnus occurrence. Principal Findings An expansion of Alnus during the Late Glacial and early Holocene periods supports the presence of alders during the LGM in southern peninsulas and northerly areas in western Europe, the foothills of the Alps, the Carpathians and northeastern Europe. After glaciers withdrew, the ice-free area of Europe was likely colonized from several regional refugia; the deglaciated area of Scandinavia was likely colonized from a single refugium in northeastern Europe. In the more northerly parts of Europe, we found a scale-dependent pattern of Alnus expansion characterised by a synchronous increase of Alnus within individual regions, though with regional differences in the times of the expansion. In southern peninsulas, the Alps and the Carpathians, by contrast, it seems that Alnus expanded differently at individual sites rather than synchronously in whole regions. Conclusions Our synthesis supports the idea that northern LGM populations were important sources of postglacial Alnus expansion. The delayed Alnus expansion apparent in some regions was likely a result of environmental limitations.

Effect of anatomical structure on the buoyancy of achenes of two subspecies ofBolboschoenus maritimus

Differences between two subspecies ofBolboschoenus maritimus (L.)Palla (=Scirpus maritimus L.) in different habitats (littoral and terrestrial) are partly due to variations in the buoyancy of their achenes, caused by different anatomical structure of the fruits. This hypothesis was tested by comparing fruit buoyancy and anatomy of both subspecies at several localities.Plants were sampled fromBolboschoenus maritimus subsp.maritimus (plants with branched inflorescences and triangular fruits, 4 localities) andB. maritimus subsp.compactus (Hoffm.)Hejný (plants with compact inflorescences and prevailing concave fruits, 4 localities) in the Czech and Slovak Republics, andB. maritimus subsp.compactus (plants with compact inflorescences and convex fruits, 1 locality) from the coast of South Sweden. From each locality a plant with several tubers connected by rhizomes was taken and transferred into the experimental garden; from these cultivated plants (clones) mixed samples of fruits collected from each clone were used for buoyancy tests and a study of their anatomy.Fruit buoyancy was found to depend on the anatomical structure of the pericarp—the presence of aeriferous tissue in the exocarp serves as the main floating organ. This differentiated all plants studied into two groups: short-floating subsp.maritimus, with a thin or none acriferous tissue in the exocarp, and long-floating subsp.compactus with well developed aeriferous tissue. Differences in fruit buoyancy corresponds to differences in the prevailing habitat types of both subspecies studied in the inland sites: the frequent occurrence of subsp.compactus in temporarily flooded terrestrial habitats contrasts with the prevalence of subsp.maritimus in the littoral of water bodies. This is an example of intra-specific ecological differentiation leading to differentiation in wetland communities.

Ecological differentiation of Central EuropeanBolboschoenus taxa and their relationship to plant communities

The taxa of theBolboschoenus maritimus group occurring in Central Europe were studied with respect to morphological differentiation, plant communities and soil conditions. Four morphological types ofBolboschoenus were distinguished, differing in habitat conditions, dispersal and distribution: type 1 —Bolboschoenus maritimus subsp.maritimus with narrow triangular fruits; type 2 —B. maritimus subsp.maritimus with wider triangular fruits, type 3 —B. maritimus subsp.compactus with round convex fruits; and type 4 —B. planiculmis auct. with concave fruits. Strong ecological differentiation of these types was found compared with relatively weak morphological diagnostic features. High discriminant efficiency appeared in the relationship to soil chemistry. Differences were also found in habitat-type spectra of individualBolboschoenus morphological types, which was reflected in their affinities to plant communities. Nomenclature, taxonomic classification, and distribution of these taxa in Europe need thorough revision.

Ecology of two cytotypes ofButomus umbellatus III. Distribution and habitat differentiation in the Czech and Slovak Republics

The distribution, total frequency of occurrence, regional differences and habitat conditions of diploid and triploidButomus umbellatus L. were studied in the Czech and Slovak Republics. Triploids are more frequent than diploids, with numerous localities in warm lowland regions in river floodplains. Diploids were found primarily in the South Bohemian Třeboň basin, outside the warm regions. The two cytotypes do not differ in dependence on water depth. Triploids occur more in habitats with more alkaline and base-rich soils. Considering the total distribution of both cytotypes in the area studied, triploidB. umbellatus appears to be more successful than its diploid progenitor, owing to its wider habitat range and superiority in vegetative reproduction.

Germination responses of diploid Butomus umbellatus to light, temperature and flooding

Summary The germination of seeds of diploid Butomus umbellatus was studied to determine the favourable conditions for germination and whether seed germination may represent a limitation within the sexual reproductive cycle of this cytotype. The need to stratify the seeds and the response of the seeds to environmental conditions during germination were studied. The influence of light/darkness, temperature, day length, and flooding/aerobic conditions was tested. Seed germination was enhanced in the winter by cold wet stratification, which indicates prevailing seed germination in the spring. Seeds germinated best in light, at relatively high temperatures (20°–30°C), in long days. Seeds were able to germinate when flooded; in most cases, flooding supported germination. In field habitats, the optimum conditions for the germination of B. umbellatus seeds may be found on an open emerged bottom saturated with water or under a shallow water layer. The process of germination does not represent a high-risk stage in the sexual reproduction of this cytotype of B. umbellatus . The time of germination (late spring or summer months) contributes to seedling mortality due to the competition of adult plants or other early-germinating species.

Flow cytometry, microsatellites and niche models reveal the origins and geographical structure of Alnus glutinosa populations in Europe

BACKGROUND AND AIMS Polyploidy in plants has been studied extensively. In many groups, two or more cytotypes represent separate biological entities with distinct distributions, histories and ecology. This study examines the distribution and origins of cytotypes of Alnus glutinosa in Europe, North Africa and western Asia. METHODS A combined approach was used involving flow cytometry and microsatellite analysis of 12 loci in 2200 plants from 209 populations combined with species distribution modelling using MIROC and CCSM climatic models, in order to analyse (1) ploidy and genetic variation, (2) the origin of tetraploid A. glutinosa, considering A. incana as a putative parent, and (3) past distributions of the species. KEY RESULTS The occurrence of tetraploid populations of A. glutinosa in Europe is determined for the first time. The distribution of tetraploids is far from random, forming two geographically well-delimited clusters located in the Iberian Peninsula and the Dinaric Alps. Based on microsatellite analysis, both tetraploid clusters are probably of autopolyploid origin, with no indication that A. incana was involved in their evolutionary history. A projection of the MIROC distribution model into the Last Glacial Maximum (LGM) showed that (1) populations occurring in the Iberian Peninsula and North Africa were probably interconnected during the LGM and (2) populations occurring in the Dinaric Alps did not exist throughout the last glacial periods, having retreated southwards into lowland areas of the Balkan Peninsula. CONCLUSIONS Newly discovered tetraploid populations are situated in the putative main glacial refugia, and neither of them was likely to have been involved in the colonization of central and northern Europe after glacial withdrawal. This could mean that neither the Iberian Peninsula nor the western part of the Balkan Peninsula served as effective refugial areas for northward post-glacial expansion of A. glutinosa.

The effect of submergence on tuber production and dormancy in two subspecies ofBolboschoenus maritimus

Two subspecies ofBolboschoenus maritimus (L.)Palla were cultivated in submerged and non-submerged conditions. The production of tubers, the proportion of tubers remaining dormant and the time between production and reactivation were investigated.Total tuber production was found to be dependent on subspecies and may be considered a biological character of the subspecies.Bolboschoenus maritimus ssp.compactus, a subspecies of less, flooded habitats, produced more tubers with a greater capacity for growth leaving a smaller proportion of dormant tubers. Inactivity of tubers and their reactivation after one or two years were enhanced by submergence in both subspecies.The proportion of tubers that are permanently dormant as well as the dormancy of the whole plant are important features influencing the survival of populations ofB. maritimus over periods of unfavourable conditions.