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Phenology has emerged as key indicator of the biological impacts of climate change, yet the role of functional traits constraining variation in herbaceous species' phenology has received little attention. Botanical gardens are ideal places in which to investigate large numbers of species growing under common climate conditions. We ask whether interspecific variation in plant phenology is influenced by differences in functional traits. We recorded onset, end, duration and intensity of initial growth, leafing out, leaf senescence, flowering and fruiting for 212 species across five botanical gardens in Germany. We measured functional traits, including plant height, absolute and specific leaf area, leaf dry matter content, leaf carbon and nitrogen content and seed mass and accounted for species' relatedness. Closely related species showed greater similarities in timing of phenological events than expected by chance, but species' traits had a high degree of explanatory power, pointing to paramount importance of species' life-history strategies. Taller plants showed later timing of initial growth, and flowered, fruited and underwent leaf senescence later. Large-leaved species had shorter flowering and fruiting durations. Taller, large-leaved species differ in their phenology and are more competitive than smaller, small-leaved species. We assume climate warming will change plant communities' competitive hierarchies with consequences for biodiversity.
Parental effects (PE) can be adaptive and improve offspring performance when parents and offspring experience similar environmental conditions. However, it is unknown whether adaptive PE exist also in habitats where such similarity is unlikely due to strong temporal variation. In particular, we do not know whether PE can adapt offspring to fluctuating levels of neighbour competition in such habitats. Here, we tested for adaptive PE in terms of two key environmental factors in a semi-arid annual system, competition and drought. While rainfall was stochastic in the study site, the competitive environment was partly predictable: higher plant densities followed after favourable (rainy) years due to high seed production. We therefore expected PE to adapt the offspring's competitive ability to these (predictable) fluctuations in plant densities, rather than to adapt the offspring's drought tolerance to the (unpredictable) occurrence of intensified drought. Parental plants of Biscutella didyma, an annual Brassicaceae, were raised under favourable watering and under drought conditions. Offspring performance was then tested under a full-factorial combination of two neighbour regimes and six watering levels in the glasshouse. Offspring of parents grown under favourable conditions were stronger competitors. This was associated with a small shift in phenology but not with higher parental seed provisioning. Offspring from parents grown under drought showed no improved drought tolerance. Moreover, no PE were detectable when offspring were grown without neighbours. Our results suggest a novel path of adaptive PE: higher competitive ability was induced in offspring that were more likely to experience high neighbour densities. Together with the lack of adaptive PE towards drought tolerance, this emphasizes that a correlation between parental and offspring environment is crucial for adaptive PE to evolve. Our results also call for the inclusion of competitive effects in future PE studies.Synthesis. This study demonstrates the important role of adaptive PE for plant fitness (regarding competition) but also their limits (regarding drought) in temporally variable environments, based on the predictability of the respective environmental factor.
Managing open habitats by wild ungulate browsing and grazing a case-study in North-Eastern Germany
(2011)
Question: Can wild ungulates efficiently maintain and restore open habitats?
Location: Brandenburg, NE Germany.
Methods: The effect of wild ungulate grazing and browsing was studied in three successional stages: (1) Corynephorus canescens-dominated grassland; (2) ruderal tall forb vegetation dominated by Tanacetum vulgare; and (3) Pinus sylvestris-pioneer forest. The study was conducted over 3 yr. In each successional stage, six paired 4 m(2)-monitoring plots of permanently grazed versus ungrazed plots were arranged in three random blocks. Removal of grazing was introduced de novo for the study. In each plot, percentage cover of each plant and lichen species and total cover of woody plants was recorded.
Results: Wild ungulates considerably affected successional pathways and species composition in open habitats but this influence became evident in alteration of abundances of only a few species. Grazing effects differed considerably between successional stages: species richness was higher in grazed versus ungrazed ruderal and pioneer forest plots, but not in the Corynephorus sites. Herbivory affected woody plant cover only in the Pioneer forest sites. Although the study period was too short to observe drastic changes in species richness and woody plant cover, notable changes in species composition were still detected in all successional stages.
Conclusion: Wild ungulate browsing is a useful tool to inhibit encroachment of woody vegetation and to conserve a species-rich, open landscape.
Question Which mechanisms promote the maintenance of the protected pioneer grass Corynephorus canescens in a mosaic landscape? Which are the interactive effects of small-scale disturbances, successional stage and year-to-year variation on early establishment probabilities of C. canescens? Location Brandenburg, NE Germany. Methods We measured emergence and survival rates over 3 yr in a sowing-experiment conducted in three successional stages (C. canescens- dominated site, ruderal forb site and pioneer forest) under two different regimes of mechanical ground disturbance (disturbed versus undisturbed control). Results Overall, disturbance led to higher emergence in a humid year and to lower emergence in a very dry year. Apparently, when soil moisture was sufficient, the main factor limiting C. canescens' establishment was competition, while in the dry year, water became the limiting factor. Survival rates were not affected by disturbance. In humid years, C. canescens emerged in higher numbers in open successional stages while in the dry year, emergence rates were higher in late stages, suggesting an important role of late successional stages for the persistence of C. canescens. Conclusions Our results suggest that small-scale disturbances can promote germination of C. canescens. However, disturbances should be carefully planned. The optimal strategy for promoting C. canescens is to apply disturbances just before seed dispersal and not during dry years. At the landscape scale, a mosaic of different vegetation types is beneficial for the protected pioneer grass as facilitation by late-successional species may be an important mechanism for the persistence of C. canescens, especially in dry years.
In common garden experiments, a number of genotypes are raised in a common environment in order to quantify the genetic component of phenotypic variation. Common gardens are thus ideally suited for disentangling how genetic and environmental factors contribute to the success of invasive species in their new non-native range. Although common garden experiments are increasingly employed in the study of invasive species, there has been little discussion about how these experiments should be designed for greatest utility. We argue that this has delayed progress in developing a general theory of invasion biology. We suggest a minimum optimal design (MOD) for common garden studies that target the ecological and evolutionary processes leading to phenotypic differentiation between native and invasive ranges. This involves four elements: (A) multiple, strategically sited garden locations, involving at the very least four gardens (2 in the native range and 2 in the invaded range); (B) careful consideration of the genetic design of the experiment; (C) standardization of experimental protocols across all gardens; and (D) care to ensure the biosafety of the experiment. Our understanding of the evolutionary ecology of biological invasions will be greatly enhanced by common garden studies, if and only if they are designed in a more systematic fashion, incorporating at the very least the MOD suggested here.
Germination rates and germination fractions of seeds can be predicted well by the hydrothermal time (HTT) model. Its four parameters hydrothermal time, minimum soil temperature, minimum soil moisture, and variation of minimum soil moisture, however, must be determined by lengthy germination experiments at combinations of several levels of soil temperature and moisture. For some applications of the HTT model it is more important to have approximate estimates for many species rather than exact values for only a few species. We suggest that minimum temperature and variation of minimum moisture can be estimated from literature data and expert knowledge. This allows to derive hydrothermal time and minimum moisture from existing data from germination experiments with one level of temperature and moisture. We applied our approach to a germination experiment comparing germination fractions of wild annual species along an aridity gradient in Israel. Using this simplified approach we estimated hydrothermal time and minimum moisture of 36 species. Comparison with exact data for three species shows that our method is a simple but effective method for obtaining parameters for the HTT model. Hydrothermal time and minimum moisture supposedly indicate climate related germination strategies. We tested whether these two parameters varied with the climate at the site where the seeds had been collected. We found no consistent variation with climate across species, suggesting that variation is more strongly controlled by site-specific factors.Abstract auch auf deutsch vorhanden:Keimungsgeschwindigkeit und Anteil gekeimter Samen lassen sich gut mit dem Hydrothermalzeit-Modell bestimmen. Dessen vier Parameter Hydrothermalzeit, Mindesttemperatur, Mindestbodenfeuchte und Streuung der Mindestbodenfeuchte müssen jedoch durch aufwendige Keimungsversuche bei Kombinationen von mehreren Temperatur- und Feuchtigkeitsstufen bestimmt werden. Für manche Anwendungen des Hydrothermalzeit-Modells sind aber ungefähre Werte für viele Arten wichtiger als genaue Werte für wenige Arten. Wenn die Mindesttemperatur und die Streuung der Mindestfeuchte aus Veröffentlichungen und Expertenwissen geschätzt würde, können die Hydrothermalzeit und Mindestbodenfeuchte aus vorhandenen Daten von Keimungsversuchen mit nur einer Temperatur- und Feuchtigkeitsstufe berechnet werden. Wir haben unseren Ansatz auf einen Keimungsversuch zum Vergleich der Keimungsquote wilder einjähriger Arten entlang eines Trockenheitsgradienten in Israel angewendet. Mit diesem Ansatz bestimmten wir die Hydrothermalzeit und Mindestfeuchtigkeit von 36 Arten. Der Vergleich mit genauen Werten für drei Arten zeigt, dass mit unserem Ansatz Hydrothermalzeit-Parameter einfach und effektiv bestimmt werden können. Hydrothermalzeit und Mindestfeuchtigkeit sollten auch bestimmte klimabedingte Keimungsstrategien anzeigen. Deshalb testeten wir, ob diese zwei Parameter mit dem Klima am Ursprungsort der Samen zusammenhängen. Wir fanden jedoch keinen für alle Arten übereinstimmenden Zusammenhang, so dass die Unterschiede vermutlich stärker durch standörtliche als durch klimatische Ursachen hervorgerufen werden.
1.Interactions among plants are key processes that strongly influence the structure and dynamics of plant populations and communities. However, most empirical studies of plant-plant interactions failed to repeatedly measure the plants? response to neighboring individuals and thereby neglected possible changes in interactions throughout the life history of the plants.2.Here, we tested the hypothesis that competition between annual species intensifies from early to late life history stages. To test this hypothesis, we sequentially measured interactions at different levels of water stress. 3.For this purpose, we conducted neighbor-removal experiments in three study sites located along a climatic gradient in Israel. The two annual species Biscutella didyma and Hymenocarpos circinnatus were used as target plants. They grew with and without neighbors in their natural habitats. Five response variables, according to the consecutive life-history stages, (seedling survival, juvenile biomass, adult survival, number of seeds and final biomass) were recorded throughout the whole growing season. 4.The results suggest that direction and intensity of interactions varied considerably between environments and life stages. On average, growth-related response variables indicated higher competition intensity at the productive end of the climatic gradient, while survival indicated either facilitation at the dry end or no trend along the gradient. 5.Considering the temporal aspect, moderate facilitation short after germination shifted to strong competition at the end of the growing season. 6.Our results highlight that the outcome of experimental studies on plant-plant interactions may not only depend on the environmental productivity but even more on the life stage at which a target plant is found.
Germination rates and germination fractions of seeds can be predicted well by the hydrothermal time (HTT) model. Its four parameters hydrothermal time, minimum soil temperature, minimum soil moisture, and variation of minimum soil moisture, however, must be determined by lengthy germination experiments at combinations of several levels of soil temperature and moisture. For some applications of the HTT model it is more important to have approximate estimates for many species rather than exact values for only a few species. We suggest that minimum temperature and variation of minimum moisture can be estimated from literature data and expert knowledge. This allows to derive hydrothermal time and minimum moisture from existing data from germination experiments with one level of temperature and moisture. We applied our approach to a germination experiment comparing germination fractions of wild annual species along an aridity gradient in Israel. Using this simplified approach we estimated hydrothermal time and minimum moisture of 36 species. Comparison with exact data for three species shows that our method is a simple but effective method for obtaining parameters for the HTT model. Hydrothermal time and minimum moisture supposedly indicate climate related germination strategies. We tested whether these two parameters varied with the climate at the site where the seeds had been collected. We found no consistent variation with climate across species, suggesting that variation is more strongly controlled by site-specific factors.
Environmental gradients represent an ideal framework for studying adaptive variation in the life history of plant species. However, on very steep gradients, largely contrasting conditions at the two gradient ends often limit the distribution of the same species across the whole range of environmental conditions. Here, we study phenotypic variation in a winter annual crucifer Biscutella didyma persisting along a steep gradient of increasing rainfall in Israel. In particular, we explored whether the life history at the arid end of the gradient indicates adaptations to drought and unpredictable conditions, while adaptations to the highly competitive environment prevail at the mesic Mediterranean end. We examined several morphological and reproductive traits in four natural populations and in populations cultivated in standard common environment. Plants from arid environments were faster in phenological development, more branched in architecture and tended to maximize reproduction, while the Mediterranean plants invested mainly in vertical vegetative growth. Differences between cultivation and field in diaspore production were very large for arid populations as opposed to Mediterranean ones, indicating a larger potential to increase reproduction under favorable conditions. Our overall findings indicate two strongly opposing selective forces at the two extremes of the aridity gradient, which result in contrasting strategies within the studied annual plant species