Leoš Klimeš

Scale-dependent biases in species counts in a grassland

. Numbers of plant species were recorded in species-rich meadows in the Bile Karpaty Mts., SE Czech Republic, with the aim to evaluate the sampling error made by well-trained observers. Five observers recorded vascular plants in seven plots ranging from 9.8 cm2 to 4 m2 independently and were not time-limited. In larger plots a discrepancy of 10–20% was found between individual estimates, in smaller plots discrepancy increased to 33%, on average. The gain in observed species richness by combining records of individual observers (in comparison with the mean numbers estimated by single observers) decreased from the smallest plot (27–82% for two to five observers) to the largest one (13–25%). However, after misidentified and suspicious records were eliminated, the gain was much lower and became scale-independent; two observers added 12% species, on average, and the increase by combining species lists made by three or more observers was negligible (3% more on average). It is concluded that most discrepancies between individual observers were caused by misidentification of rare seedlings and young plants. We suggest that in species-rich meadows plants should be recorded by at least three observers together and that they should consult all problematic plant specimens together in the field, to minimize errors.

Scale‐dependent variation in visual estimates of grassland plant cover

Abstract Plant cover was visually estimated by five observers, independent of each other, in a species-rich grassland in the Bílé Karpaty Mts., southeastern Czech Republic, in seven plots ranging from 0.001 to 4 m2. Variation of total plant cover among the observers was high at small scales: 0.001 - 0.016 m2; coefficient of variation, CV = 35 to 45%, but much lower at larger scales: 0.06 - 4 m2; CV = 7 to 15%. Differences between visual estimates of plant cover of individual species made by different observers were affected by plot size, total cover and morphology of particular plants. CV of the cover of individual species ranged from 0 to 225% and decreased with increasing plot size. For abundant plants the CV attained ca. 50%, independent of plot size. In spite of a very high number of sterile plants with similar leaf morphology and colour, the observed variation in cover estimates in the studied grassland was comparable with results reported from other vegetation types. Differences between estimates by individual observers were often larger than usual year to year changes in undisturbed grasslands. Therefore, I suggest that to avoid difficulties in the interpretation of results based on plant cover data obtained from visual estimates, several observers should always work together, adjusting their extreme estimates. Nomenclature: Marhold & Hindák (1998).

Small-scale plant mobility in a species-rich grassland

. Plant mobility was studied in a species-rich grassland in S. Moravia (Czech Republic) at scales from 0.0025 to 2.25 m2. Cumulative species numbers, cumulative species frequencies and the distribution of distances between sites of occurrence in particular years were established using data collected from 1991 to 1997. The observed values were compared with null models of completely random and restricted random movement of the plants. Most plants persisted at a spot more frequently than expected from the completely random model and with a few exceptions they also spread to neighbouring subplots more often than expected. Cumulative species numbers were between the ranges predicted by the two random models and increased linearly during the 7-yr period at all scales, indicating a large species pool. The role of clonal spreading and of generative reproduction depended on the growth form of the species. I conclude that a high species richness is not necessarily linked with a high plant mobility, even if coexistence of plant species may be promoted by plant mobility.

Regeneration capacity and carbohydrate reserves in a clonal plant Rumex alpinus: effect of burial

The morphological and anatomical responses to different depths of burial were examined in Rumex alpinus (Polygonaceae), a perennial plant with monopodial, horizontally growing rhizome. Its segments, which consist of 12–20 internodes, 1 to 2 mm in length each, are products of single growing seasons. The rhizomes regenerated from 5, 10 and 20 cm, but failed to emerge from 30 cm. Number of internodes produced during a growing season was not affected by burial but the length of internodes increased up to about 30-fold. The rhizomes growing up to the surface were subsidized by older rhizome segments. In the case of deeply buried rhizomes the carbohydrate reserves of the last-year-segment were nearly completely depleted. Evolutionary significance of the regeneration capacity is discussed.

Late holocene history and vegetation dynamics of a floodplain alder carr: A case study from eastern Bohemia, Czech Republic

Vegetation development in the lowland floodplain alder carr “Na bahně” (eastern Bohemia, the Czech Republic) has been studied by means of pollen and macrofossil analyses and combined with vegetation analysis performed over the last 70 years. Local successional changes started with an oxbow lake (160 cal BC) which has later terrestrialised (630 cal AD). Then it changed from a typical alluvial fen into aSphagnum-dominated spring mire (950 cal AD) supplied by water arising from a river terrace surrounding the locality from three sites. In the centre of this wetland a small patch of alder carr developed (100 cal. AD), showing some tendency towards cyclic succession. The alder carr alternated several times with an openCarex fen (1100 cal AD to recent). The last fen-to-alder carr transition has been documented by direct observation during this century. Possible autogenic and allogenic factors driving the succession are discussed. The model of autogenic cyclic succession corresponds well with direct field observations and can be used to interpret alder carr structure, its dynamics, and function.

Demography and modelling of clonal fragments in the pseudoannual plant shape Trientalis europaea L.

The clonal growth pattern and demography of clonal fragments (aggregation of ramets derived from a common parent ramet) in the pseudoannual plant Trientalis europaea were studied in field conditions from 1991 to 1993. During this period the population of clonal fragments declined, with a half-life of 7.4 years. Number and size of the clonal progeny and stolon length were positively related to the size of the mother ramet. Survival rates of ramets and tubers increased with size. The rate of clonal growth was low: after three years, about 70% of the clonal fragments had only one ramet. This suggests that the pseudoannual growth habit in T. europaea is more important as mechanism of perennation than of ramet multiplication.Field data were used in a simulation model of architecture and population dynamics of clonal fragments. About 10% of the clonal fragments survived to the end of the simulation (15 years) and the mean survival was 4.7 years. The model predicted a positive correlation between persistence of the clonal fragment and number of ramets produced. Sensitivity analysis showed that the production of a daughter ramet of at least the same size as the parent ramet was the most important pathway for the survival and the number of ramets of the clonal fragment, whereas the production of secondary ramets had a very small effect. This confirms the interpretation of the pseudoannual life-cycle as a mechanism of ramet replacement in this species. Sensitivity analysis also revealed that changes in survival probabilities of the smallest ramets had the largest impact on clonal fragment dynamics. This reflects the important role of the smallest size class of ramets as a source of new vegetative propagules, maintaining a hierarchy in the size structure of the population.

Clonal splitters and integrators in harsh environments of the Trans-Himalaya

Individuals of clonal plants consist of physically and physiologically connected ramets. In splitters, they are integrated for a time shorter than ramet generation time (i.e. the time it takes to produce the first offspring ramet), whereas in integrators connections between ramets persist for a longer time. It has been predicted that integrators should prevail in stressful environments, such as habitats poor in nutrients, whereas splitters are expected to dominate in benign habitats, such as fertile areas with a moderate climate. I tested these predictions in four dry mountain areas of the Trans-Himalaya, in high altitudes subjected to multiple stresses. In accordance with the expectations I found that clonal plants with integrated ramets reach higher mean and maximum altitudes than splitters. Integrators were over-represented in nutrient-poor habitats, such as dry semi-deserts, sandy steppes and in subnival habitats, whereas splitters preferentially colonised mesic habitats, saline sites and wetlands. While there was no difference in the representation of splitters and integrators in habitats with an unstable surface, such as screes, dunes and water bodies, fully integrated clonal plants preferred very stable environments, such as banks of streams covered by closed-canopy vegetation. Most relationships between clonal integration and environmental factors were explainable by the phylogenetic relationship between the species, only the significant preference of splitters for shaded environments persisted in phylogenetically corrected analysis. The results indicate that clonal integration belongs to a set of evolutionarily conservative plant traits, usually shared by related species. Consequently, the adaptive value of clonal integration in individual habitats remains questionable.

Clonal Growth Forms in Eastern Ladakh, Western Himalayas: Classification and Habitat Preferences

Earlier observations that plant clonality, i.e., production of potentially independent offspring by vegetative growth, increase in importance in cold climates such as in arctic and alpine regions, have been recently questioned. However, lack of data obtained using a comparable methodology throughout different regions limit such comparisons. Here we present a classification of clonal growth forms for vascular plants from East Ladakh (an arid mountain range in NW Himalaya, India), and assess the relationship of these forms with multiple environmental gradients. Based on field assessment of clonality in 540 species we distinguished 20 growth forms, which were then grouped into four broader space occupancy strategies. Occurrence in communities and relationship with environmental characteristics and altitude were analyzed using multivariate methods. The most abundant growth form was represented by non-clonal perennial species with a pleiocorm having short branches, prevailing in steppes, Caragana shrubs and screes. The most abundant clonal species were those with very short epigeogenous rhizomes, such as turf graminoids prevailing in wet Kobresia grasslands. Two principal environmental gradients, together with several abiotic variables, affected space occupancy strategies: moisture and altitude. Non-spreading integrators prevailed on shaded rocky slopes, non-spreading splitters in wet grasslands and spreading splitters at the wettest sites. Spreading integrators were the least frequent strategy predominantly occurring at the most elevated sites. Because relevance of clonality decreased with altitude and different communities host different sets of clonal growth strategies, comparison with other cold climate regions should take multiple environmental gradients into account.

Cushions of Thylacospermum caespitosum (Caryophyllaceae) do not facilitate other plants under extreme altitude and dry conditions in the north-west Himalayas

BACKGROUND Cushion plants are commonly considered as keystone nurse species that ameliorate the harsh conditions they inhabit in alpine ecosystems, thus facilitating other species and increasing alpine plant biodiversity. A literature search resulted in 25 key studies showing overwhelming facilitative effects of different cushion plants and hypothesizing greater facilitation with increased environmental severity (i.e. higher altitude and/or lower rainfall). At the same time, emerging ecological theory alongside the cushion-specific literature suggests that facilitation might not always occur under extreme environmental conditions, and especially under high altitude and dryness. METHODS To assess these hypotheses, possible nursing effects of Thylacospermum caespitosum (Caryophyllaceae) were examined at extremely high altitude (5900 m a.s.l.) and in dry conditions (precipitation <100 mm year(-1)) in Eastern Ladakh, Trans-Himalaya. This is, by far, the highest site, and the second driest, at which the effects of cushions have been studied so far. KEY RESULTS In accordance with the theoretical predictions, no nursing effects of T. caespitosum on other alpine plants were detected. The number and abundance of species were greater outside cushions than within and on the edge of cushions. None of the 13 species detected was positively associated with cushions, while nine of them were negatively associated. Plant diversity increased with the size of the area sampled outside cushions, but no species-area relationship was found within cushions. CONCLUSIONS The results support the emerging theoretical prediction of restricted facilitative effects under extreme combinations of cold and dryness, integrating these ideas in the context of the ecology of cushion plants. This evidence suggests that cases of missing strong facilitation are likely to be found in other extreme alpine conditions.

Small-scale distribution of species richness in a grassland (Bílé Karpaty Mts., Czech Republic)

Variation in the number of species was studied in a subthermophilous grassland at a scale of 0.05 ×0.05 m during a 5-year period. The observed variance of species richness (VSR) was compared with a null model based on random distribution of species over a set of squares. It was found that distribution of species richness had more values than, expected around the mean and less values at the “shoulders”. Both tails fell within the predicted limits. Application of the procedures removing spatial dependence (random shifts, rotation/reflection method byPalmer & van der Maarel 1995) and environmental heterogeneity (patch model byWatkins & Wilson 1992) did not change the observed pattern.Using simulations in which the number of clumps and clumping intensity were manipulated it was found that the effect of the clumped spatial pattern, on VSR results in a wide range of variances. Both variance excess and variance deficit were found more frequently than expected under the null model.To test the effect of the limitation to the number of individuals per square, a null model was developed which included that observed number of plant shoots per square, the observed distribution of the number of shoots belonging to individual species per square and the observed spatial distribution of the shoots. The observed VSR was still lower than that produced by the null model. Therefore, it is concluded that at a scale of 0.05×0.05 m plant species combine in a non-random way in the studied grassland. It is suggested that the shape of left and right “shoulders” of the species richness distribution may be caused by different factors, such as positive and negative covariance between species, respectively. Their simultaneous impact can generate the observed pattern in species richness.