Marek Sammul

Classifying clonal growth forms based on vegetative mobility and ramet longevity: a whole community analysis

We measured rhizome branching, clonal mobility, and ramet longevity of 98 meadow plant species. A cluster analysis applied to this dataset revealed nine clonal growth types that differ mainly by the ramet lifespan and vegetative mobility. Then we compared the abundance of these groups of clonal species between the three following plant communities: (1) open, (2) restored and (3) overgrown wooded meadows in the Laelatu-Nehatu-Puhtu Nature Reserve, Estonia. This is the first study where the quantitative values of belowground clonal traits have been measured for all species of a species-rich community. We show that species with annual ramets and with a low vegetative mobility were most abundant in open grasslands. The relative abundance of perennial species with annual ramets was positively correlated with shoot density and species diversity, indicating that high ramet turnover rates combined with a high genet longevity can positively affect species coexistence in meadow communities. Hence, this study provides evidence for the fact that the average values of clonal life-history parameters differ between these communities. Herb communities under forest canopy consist, in average, of species with ramets that live longer and are clonally more mobile than in the communities of open sites.

A comparison of plant communities on the basis of their clonal growth patterns

This study was performed to analyse how a vegetative propagation pattern of plants affects the coexistence of species and subsequent species richness of the community. We compared community average clonal growth in the herbal communities of forests, wooded meadows, and open meadows in Laelatu, Estonia. The parameters used for the calculation of the community averages and measured for each species were ramet life span, rhizome branching, and clonal mobility. We also examined the intrinsic (i.e. independent of the environment) relationship between community clonal growth and plant species density. We found strong correlations between the environmental factors (productivity, light availability, and mowing regime) and community averages of clonal growth parameters, while species density was (negatively) correlated only with community average of rhizome increment. The community average of ramet life span decreased with the increasing biomass of the herb layer. No evidence was found to support the hypothesis that species-rich communities may consist of species with more contrasting mobility compared with species-poor communities. Independent of the effect of the environmental factors, species density was positively correlated with ramet density. There was intrinsic positive relationship between species density and community average of ramet life span at open meadow sites and intrinsic negative relationship between species density and community average of rhizome increment at wooded meadow sites. We conclude that in forest communities the capability of clonal plants to forage for light is favoured, while in unmown meadows a competitively strong phalanx growth form is advantageous. We established that ramet turnover increases and vegetative mobility decreases with increasing species diversity, although these two relationships depend strongly on the type of the studied community.

Determination of national conservation responsibilities for species conservation in regions with multiple political jurisdictions

The Convention on Biodiversity (CBD) commits its signatories to the identification and monitoring of biodiversity. The European Union has implemented this commitment into its legislation. Despite the legal requirement resources are scarce, requiring a prioritization of conservation actions, including e.g. monitoring. Red lists are currently the most prominent tool for priority setting in applied conservation, despite the fact that they were not developed for that purpose. Therefore, it is hardly surprising that they do not always reflect actual conservation needs. As a response, the concept of national responsibility as a complementary tool was developed during the last two decades. The existing methods are country specific and mainly incomparable on an international scale. Here, we present a newly developed method, which is applicable to any taxonomic group, adjustable to different geographic scales, with little data requirements and clear categorizations. We apply the new method to over 1,000 species in several countries of different size and report on the applicability of our method and discuss problems that derive from the currently available data. Our method has several major advantages compared to currently available methods. It is applicable to any geographic range, allows automatization, given database availability, and is readily adjustable to future data improvements. It further has comparably low data demands by exploiting one of the most commonly available information on biodiversity, i.e. distribution maps. We believe that our method allows the allocation of the limited resources in nature conservation in the most sensible way, e.g. the sharing of monitoring duties, effectively selecting networks of protected areas, improving knowledge on biodiversity, and closing information gaps in many species groups.

Regional effects on competition–productivity relationship: a set of field experiments in two distant regions

We studied the effect of productivity on competition intensity and the relationship between competition intensity and community species richness, using a removal experiment with the perennial plant Solidago virgaurea . The experiment was conducted in 16 different communities from two geographically distant areas (western Estonia and northern Norway). The results were compared with the results of previous experiments with Anthoxanthum odoratum from the same areas. Removal of neighbors had a positive effect on the biomass of both Solidago and Anthoxanthum , and this response was stronger in communities with higher productivity. Thus, the corrected index of relative competition intensity, CRCI, increased with increasing community productivity. Species richness was negatively correlated with CRCI in Estonia but not in Norway and not in the case of the pooled material. The results suggest that competitive exclusion operates at least in these communities which species pool is large. Our results indicate that the relationship between competition intensity and productivity is non-linear. In our data, competition prevails in communities where living plant biomass exceeds 200 g m � 2 , whereas in less productive communities, competition remains undetected and direct plant � /plant relationships might at times be even mutualistic. Moreover, we found that the relationship between competition intensity and productivity is strongly dependent on regional differences and is intimately connected to a concordant variation in the intensity of grazing. The least productive communities both in Estonia and in Norway are characterized by intensive grazing, which reduces importance of competition. Hence, the contrasting results corroborates the predictions of the hypothesis of exploitation ecosystems, predicting that trophic dynamics account for the relationship between competition intensity and primary productivity.

Wooded meadows of Estonia: conservation efforts for a traditional habitat

We provide an overview of the amount and ecological condition of wooded meadows throughout Estonia after significant changes in agriculture in the second part of 20th century. We also present the first attempt to estimate the success of recent conservation efforts for wooded meadows. Our analysis is based on recent exhaustive inventories. We found that Estonia, despite a decrease of the area in wooded meadows by two orders of magnitude during the 20th century, still has about 8400 ha remaining of which approximately 5800 ha are meadows with, at the least, an intermediate conservation value. The latter is directly dependent on mowing regime. Efforts to preserve wooded meadows include establishment of protected areas and financial support for mowing. The national conservation subsidy has been useful and supportive for wooded meadows, however the total amount of this subsidy has been small compared to the area that could be supported. The much larger funds of agri-environmental subsidies have largely not been available for wooded meadows. Moreover, there has been no record keeping about subsidisation of semi-natural grasslands using agricultural support schemes. Although the preservation of some good examples of wooded meadows in Estonia seems guaranteed, further degradation of this valuable habitat type on a national scale is very probable.

Length of the Spacer Rather than its Plasticity Relates to Species Distribution in Various Natural Habitats

Much evidence from laboratory experiments and theoretical studies show that plasticity of clonal growth traits like lateral spread provides advantage in heterogeneous conditions. However, few tests of whether species with plastic clonal growth have an advantage over non-plastic species in natural conditions exist. I analyzed whether spacer length (i.e., length of a stolon or a rhizome branch between two ramets) and the variation of it (as a surrogate to its plasticity) relate to species presence and abundance in open meadow, wooded meadow and forest habitats with varying environmental conditions at Laelatu wooded meadow, Estonia. In fertilized, compared to unfertilized conditions, a weak and non-conclusive advantage of both long spacers and high variability of spacer length was detected. Abandonment in open meadows lead to a prevalence of species with shorter spacers, while on abandoned wooded meadows the species with longer spacers dominated. There was no difference in variability of spacer length between managed and abandoned meadows. In more heterogeneous forest habitats high variability of spacer length was more common, but the effect of lengthy spacers was more pronounced. The results suggest that while the variability of spacer length indeed corresponds with larger abundance in some conditions, the actual length of spacers has a more pronounced relationship with abundance of plants in natural vegetation.

Habitat preferences and distribution characteristics are indicative of species long-term persistence in the Estonian flora

Large-scale changes in regional floras provide direct information about changes in biodiversity through time and enable the evaluation of conservation targets. We compared the distribution ranges in 2004 of Estonian native terrestrial flora with the distribution ranges before 1970, using the Atlas of Estonian Flora. Relative persistence was related to species endemism, commonness, occurrence at its border of the global distribution range, main habitat type, sensitivity to human impact, life-form, conservation category, and Red List category. A literature-based database of the flora of Estonian habitat types was used to evaluate relative persistence of the flora of different habitats. Changes in the flora are largely dependent on human activities. The decrease in mire and grassland habitats and the increase in forests are reflected in the persistences of related species. Flora of mire habitats decreased the most. The fact that an almost ten-fold decrease of grasslands has not resulted in as large a decrease in the ranges of grassland species could serve as evidence of the extinction debt of these habitats. We also found a greater decrease among habitat specialists than habitat generalists and lower average persistence of the species of species-rich habitats. Our data show that current prioritization of species for conservation is in concordance with needs, as reflected in the changes in the range of species. However, conservation has not been entirely successful: the decrease of protected species continues. Our simple method for summarizing large databases was effective for the evaluation of large scale effects of conservation actions.

Genetic variability, population size and reproduction potential in Ligularia sibirica (L.) populations in Estonia

Ligularia sibirica (L.) Cass. (Asteraceae) is a EU Habitats Directive Annex II plant species that has suffered a lot from human-caused major changes in quality and availability of habitats in Estonia. The aim of this study was to find out if the observed decline in population size is reflected in the amount of genetic variation and fertility in remnant populations of this species. AFLP technique was used for that purpose. Genetic diversity within populations was assessed as the percentage of polymorphic loci in a given population and average gene diversity over loci. The degree of genetic differentiation among populations and genetic differentiation between pairs of populations was estimated. The amount of viable seeds per flower stem was compared among populations and between years (2007 and 2008). Average genetic diversity over loci and proportion of polymorphic loci in L. sibirica populations were significantly correlated with population size, suggesting the action of genetic drift and/or inbreeding. No correlation was found between genetic and geographic distances. Natural barriers like forests may have been efficiently preventing seed migration even between geographically closer populations. Results of this study suggest that genetic erosion could be partially responsible for the lower fitness in smaller populations of this species.

Determination of conservation priorities in regions with multiple political jurisdictions

Red lists serve as the most prominent tool for priority setting in applied conservation, even though they were not originally designed for this task. Hence, threat status does not always reflect actual conservation needs and can be very different from actual conservation priorities. Therefore, red lists may at best be a suboptimal tool for setting conservation priorities in a country or region. As a response, a range of alternative or complementary tools have been developed, with approaches, methods, and parameters such as population decline, population center etc. used, differing widely among countries. One recent development is the combination of conservation status with a measure of the international importance of a population in a focal region for the global survival of a species. Here, we provide a new method that integrates the two concepts while keeping them conceptually separate. The main benefit of this method is that it can be applied across variable geographical scales such as regions, countries, and even continents. Furthermore, it allows for better recommendations for applied conservation and conservation policy development than the two concepts in isolation. Our method, if applied internationally, would allow for a standardized priority setting in species conservation, would be highly comparable between countries, and would lead to a more efficient use of the limited financial and human resources for monitoring and conservation of biodiversity.

Physiological Integration Ameliorates Negative Effects of Drought Stress in the Clonal Herb Fragaria orientalis

Clonal growth allows plants to spread horizontally and to establish ramets in sites of contrasting resource status. If ramets remain physiologically integrated, clones in heterogeneous environments can act as cooperative systems – effects of stress on one ramet can be ameliorated by another connected ramet inhabiting benign conditions. But little is known about the effects of patch contrast on physiological integration of clonal plants and no study has addressed its effects on physiological traits like osmolytes, reactive oxygen intermediates and antioxidant enzymes. We examined the effect of physiological integration on survival, growth and stress indicators such as osmolytes, reactive oxygen intermediates (ROIs) and antioxidant enzymes in a clonal plant, Fragaria orientalis, growing in homogenous and heterogeneous environments differing in patch contrast of water availability (1 homogeneous (no contrast) group; 2 low contrast group; 3 high contrast group). Drought stress markedly reduced the survival and growth of the severed ramets of F. orientalis, especially in high contrast treatments. Support from a ramet growing in benign patch considerably reduced drought stress and enhanced growth of ramets in dry patches. The larger the contrast between water availability, the larger the amount of support the depending ramet received from the supporting one. This support strongly affected the growth of the supporting ramet, but not to an extent to cause increase in stress indicators. We also found indication of costs related to maintenance of physiological connection between ramets. Thus, the net benefit of physiological integration depends on the environment and integration between ramets of F. orientalis could be advantageous only in heterogeneous conditions with a high contrast.