Refine
Has Fulltext
- yes (5)
Is part of the Bibliography
- yes (5)
Document Type
- Doctoral Thesis (3)
- Journal article (1)
- Book (1)
Keywords
- Boden (5) (remove)
Das Naturschutzgebiet "Garchinger Heide" ist ein Relikt der ehemals ausgedehnten Kalkmagerrasen auf Pararendzinen über Niederterrassenschotter im Alpenvorland. Bei jahrhundertelanger extensiver Landnutzung hat sich auf trockenen nährstoffarmen Böden eine artenreiche Vegetation mit zahlreichen seltenen und gefährdeten Arten entwickelt. Die "Altheide", welche den größten Teil des Naturschutzgebiets einnimmt, ist geprägt durch flachgründige, nährstoffarme Böden, welche eine geschlossene Vegetation mit einem hohen Anteil an Arten der Halbtrockenrasen (hauptsächlich Mesobromion erecti und Cirsio-Brachypodion) tragen. Das 1945 durch Oberbodenabtrag begonnene, aber nicht mehr ausgebaute und verwendete „Rollfeld“ weist einen niedrigen Feinbodenanteil, welcher auf eine geringe Wasserverfügbarkeit schließen lässt, sowie niedrige Gehalte an Gesamtstickstoff und CAL-austauschbaren P2O5 auf. Die Vegetation ist lückig mit einem hohen Anteil an Trockenrasenarten (Xerobromion, Sedo-Scleranthetea, Sesleritalia albicantis). Das heutige Vorkommen zahlreicher Arten, welche in früheren Untersuchungen (1956 und 1986) nicht gefunden wurden, lässt auf eine fortschreitende Sukzession der Vegetation des Rollfeldes hin zu der der Altheide schließen. Der 1959 zum Naturschutzgebiet hinzu gekaufte ehemalige Acker unterscheidet sich immer noch deutlich von der Altheide. Eine allmähliche Annäherung der Standortbedingungen ist jedoch zu beobachten. So sanken die Gehalte an CAL-austauschbarem P_ 2 O_5 und K_2 O im Vergleich zu Werten aus dem Jahr 1993 deutlich ab. Trotz des immer noch hohen Anteils an Grünlandarten (Molinio-Arrhenatheretea) konnten sich im Vergleich zu 1986 mehr Magerrasenarten etablieren.
Many ant species excavate underground nests. One of the most impressive examples is the Chaco leaf-cutting ant Atta vollenweideri from the Gran Chaco region in South America. The nests excavated by the workers of that species are among the largest insect-built structures on the planet. They are ecavated over years possibly involving millions of working individuals. However, the mechanisms underlying the organisation of collective nest digging in ants remain largely unknown. Considering the sheer dimensions of the nest in comparison to the size and presumably limited perceptual and cognitive abilities of the single worker, the assumption can be made that organising mechanisms are mostly based on responses of individuals to local stimuli within their perceptual range. Among these local stimuli that guide nest digging we can expect environmental variables, stimuli that relate to the requirements of the colony, and stimuli related to the spatial coordination of collective effort. The present thesis investigates the role of local stimuli from these three categories in the organisation of collective digging behaviour in the Chaco leaf-cutting ant. It describes experiments on (1) how workers respond in the context of digging to differences in soil moisture, which comprises an important environmental variable; (2) how available nest space influences nest enlargement; (3) and how the spatial coordination of excavating workers is implemented by responding to stimuli arising from nest mates while engaged in digging behaviour. The experiments on soil water content show that workers prefer to dig in moist materials that allow for fast excavation and transport rates. Accordingly, an unequal distribution of water in the soil around a nest can influence how the nest shape develops. On the other hand, results also indicate that workers strongly avoid excavating in extremely moist materials. Regarding the abundant occurrence of flooding events in the Gran Chaco region, the latter can be interpreted as an adaptation to avoid water inflow into the nest. In the experiments on the effect of nest space, the ants excavated less when presented with larger nests. When a large amount of space was suddenly added to the nest during the digging process, excavation rates decreased according to the new volume. These observations confirm the hypothesis that digging activity is regulated according to space requirements, possibly because crowding conditions inside the nest influence excavation behaviour. However, observations also indicate an intrinsic decrease of digging motivation with time. Moreover, excavation rates correlate with nest size only when comparing nests of similar shape. Distributing a similar nest volume to three smaller chambers, instead of one, resulted in drastically decreased digging rates. A possible explanation for that observation lies in the distribution of workers inside the nest that may vary according to nest geometry: a different distribution of individuals can lead to in different local crowding conditions in similar nest volumes. Furthermore, two different stimuli are described that are used in the spatial coordination of collective digging effort. First, fresh soil pellets deposited close to the digging site on their way from the surface increase the probability that arriving workers join excavation efforts at the same site. The deposition of pellets on the way is a consequence of sequential task partitioning during soil transport. The pellets are carried in transport chains that closely resemble the modalities of leaf transport observed at the surface. Second, workers stridulate while digging. The short-ranged vibrational signals produced thereby also attract nest mates to excavate at the same location. Accordingly, two mutually complementing mechanisms are described that allow to concentrate excavators at one location. In both cases, a local stimulus that is generated by current close-by excavation activity increases the probability of the stimulus receiver to dig close to other excavators. In an environment otherwise poor in digging stimuli, these mechanisms can be especially important to give collective digging efforts a common direction. As a consequence it can be argued that the spatial organisation of collective digging is based on choice copying. Individuals copy nest mate decisions on where to excavate by responding to local stimuli provided by nest mate digging activity. Taken together, responses to local stimuli can determine the direction of nest growth, aid in preventing the inflow of surface water into the nest, guide the adjustment of nest size to colony requirements and spatially coordinate collective digging efforts. Even though it cannot be ruled out that digging responses based e.g. on spatial memory or long-term experience exist, the results presented here clearly demonstrate that responses to local information account for many important aspects of nest development.
Über unterschiedlich lange Zeiträume mit unterschiedlicher Intensität als Schafrotationsweide genutzte ehemalige Mähwiesen wurden hinsichtlich
- physikalischer Bodenparameter als Indikatoren für Verdichtungsvorgänge,
- vegetationskundlicher Unterschiede in Folge der Schnitt- und Beweidungsregime sowie
- den Auswirkungen auf die Heuschrecken-, Laufkäfer- und Spinnengemeinschaften
untersucht.
Die allgemeine Beschreibung der Böden zeigt, daß sich die Flächen hinsichtlich ihrer Bodenbeschaffenheit und Nährstoffversorgung nur in engen Grenzen unterscheiden. Lediglich die Kontrollfläche hat einen deutlich erhöhten Feinsandanteil und weniger Ton als die Vergleichsflächen. Zusammenfassend werden die Böden als hydromorphe, allochthone Aueböden beschrieben. Porosität und Permeabilität korrelieren wesentlich stärker mit der Bodenart als - wie ursprünglich vermutet - mit der Intensität der Beweidung.
Bei der direkten Untersuchung einer bislang als Mähwiese genutzten Fläche vor und nach einer Weidesaison konnte eine erhöhte Verdichtung im Bereich der Mittelporen nachgewiesen werden, die Permeabilität wurde dadurch jedoch nicht nachweisbar beeinträchtigt. Frostlockerung und pedobiologische Aktivität außerhalb der Weidezeiten wirken diesem Prozeß entgegen, so daß sich über mehrere Jahre keine zunehmende Verdichtung auf den dauerhaft beweideten Flächen nachweisen läßt.
Allerdings führt die Beweidung zu speziellen Strukturen, wie sie auf reinen Mähwiesen nicht gefunden werden können. Aufgrund der räumlich differenzierten Nutzung der Fläche durch die Schafe entstehen Störstellen beispielsweise an derTränke oder dort, wo die Tiere Nacht für Nacht lagern. Diese Störstellen sind bald von Vegetation entblößt, stark durch den Kot und Urin der Tiere in ihrem Chemismus verändert und deutlich verdichtet.
Vegetationsaufnahmen nach BRAUN-BLJXNQUET und die Auswertung nach den ökologischen Zeigerwerten belegen die Nutzungsintensivierung durch die Schafrotationsweide. Die Wiesen verarmen an Kenn- und Differentialarten der Glatthafergesellschaften, dafür finden sich mehr Ruderalisierungszeiger. Die Artenzahl nimmt kontinuierlich ab, und es erfolgt eine Verschiebung des Artenspektrums zu lichthungrigen Düngezeigern mit hohem Regenerationsvermögen. Hinzu kommt die Ausbreitung von Weideunkräutern vor allem auf Flächen, die nicht regelmäßig ausgemäht werden. Die Fläche "F7" stellt ein Übergangsstadium von der Wiese zur intensiven Rotationsweide dar, auf der sich vorübergehend eine erhöhte Vielfalt aus Arten der Wiese und Weideunkräutern eingestellt hat.
Eine gleichgerichtete Vegetationsveränderung nach der Umnutzung einer vormaligen zweischürigen Mähwiese in eine Schafrotationsweide belegt die Annahme, daß die Unterschiede in der Vegetation auf "FK", "F1 ", "F4", "F7" und "F15" nicht standortbedingt, sondern nutzungsbedingt sind. Dies könnte mittels einer Frequenzprozentanalyse auf "F1 " vor und nach einer dreijährigen Schafbeweidung im Vergleich zu der Kontrollfläche "FK" belegt werden.
Die Heuschreckenfauna wurde mittels lsolationsquadratfängen untersucht, die Erfassung der Laufkäfer- und Spinnenfauna erfolgte mit Barberfallen.
Die höchste lndividuendichte der Heuschrecken fand sich auf "FK" gefolgt von "F7". Aussagen zu Diversität und Evenness sowie Arten- und Dominanzidentität sind aufgrund der eingeschränkten Artenzahlen zwischen drei und sechs pro Fläche nur unter Vorbehalten möglich. Durchweg geringe Diversitätswerte sind auf die Artenarmut zurückzuführen, da die Evenness aller Aufnahmen nahe bei 1 liegt. Für die Gemeinschaften von "FK" und "F1 " zeigt sich eine große Übereinstimmung hinsichtlich der Arten- und Dominanzidentität. Eine hohe Anzahl Subadulter in den Barbeffallen der Frühsommerfangperioden auf "F1 " findet keinen Niederschlag in entsprechenden Abundanzen adulter Heuschrecken. Die Beweidung führt demnach zu erhöhten Mortalitätsraten der Larvalstadien. Das Ähnlichkeitsdendrogramm bestätigt die große Ähnlichkeit der Dominanzidentitäten von "FK" und "F1 ", die Verschmelzungsniveaus für die übrigen Flächen liegen jedoch auch zwischen 55 und 70%.
Artenzahl und Aktivitätsdichte der Laufkäfergemeinschaften weisen die Kontrollfläche als die sowohl arten- als auch individuenärmste Fläche aus. Darüber hinaus werden dort 83% aller Individuen von einer Art (Poecilus versicoloi) gestellt. Die höchste Diversität findet sich auf "F4"; auf "F7" und "F15" sind die Diversitätswerte niedriger, dafür aber die Abundanzen wesentlich höher. Mit 30 Arten ist "F4" am artenreichsten, während die größte Aktivitätsdichte auf "F7" nachgewiesen werden konnte. Ähnlich den Heuschreckengemeinschaften zeigt sich eine hohe Übereinstimmung zwischen "FK" und "Fr, die Laufkäferzönosen der intensiver beweideten Flächen unterscheiden sich in Arten- und Dominanzidentität deutlich von den Zönosen dieser Flächen.
Die Auswertung der Spinnenfänge unterstützt mit leichten Abweichungen die Ergebnisse aus der Analyse der Laufkäferfänge. "F7" ist die mit Abstand arten- und individuenreichste Fläche. Erneut findet sich die niedrigste Aktivitätsdichte auf "FK", wenngleich sich diese nur minimal von der auf "F4" unterscheiden. Deutlich zeigt das Ähnlichkeitsdendrogramm zwei Gruppen von Spinnengemeinschaften: die der Mähwiesen und die der Weiden.
Während die Kontrollfläche "FK" und die seit einem Jahr beweidete Fläche "F1" durch arme Laufkäfer- und Spinnengesellschaften gekennzeichnet sind, auf denen wenige Arten extrem hohe Dominanzwerte erreichen, profitieren die genannten Gruppen offensichtlich von der Schafbeweidung. Der Vorteil wird auf den Flächen deutlicher, die nicht perfekt als Rotationsweide genutzt werden. Diverse Störstellen und Kleinstrukturen schaffen ökologische Nischen, die von zusätzlichen Arten besetzt werden. Lediglich die Heuschrecken werden durch diese Nutzungsform benachteiligt.
Termites are the most important soil ecosystem engineers of semi‐arid and arid habitats. They enhance decomposition processes as well as the subsequent mineralisation of nutrients by bacteria and fungi. Through their construction of galleries, nests and mounds, they promote soil turnover and influence the distribution of nutrients and also alter texture and hydrological properties of soils, thereby affecting the heterogeneity of their ecosystem. The main aim of the present thesis was to define the impact of termites on ecosys‐tem functioning in a semi‐arid ecosystem. In a baseline study, I assessed the diversity of termite taxa in relation to the amount of precipitation, the vegetation patterns and the land use systems at several sites in Namibia. Subsequently, I focussed on a species that is highly abundant in many African savannas, the fungus growing and mound building species Macro‐termes michaelseni (Sjöstedt, 1914). I asked how this species influences the spatial hetero‐geneity of soil and vegetation patterns. From repeated samplings at 13 sites in Namibia, I obtained 17 termite taxa of 15 genera. While the type of land use seems to have a minor effect on the termite fauna, the mean annual precipitation explained 96% and the Simpson index of vascular plant diversity 81% of the variation in taxa diversity. The number of termite taxa increased with both of these explanation variables. In contrast to former studies on Macrotermes mounds in several regions of Africa that I reviewed, soil analyses from M. michaelseni mounds in the central Namibian savanna revealed that they contain much higher nitrogen contents when compared to their parent material. Further analyses revealed that nitrate forms a major component of the nitrogen content in termite mounds. As nitrate solves easily in water, evaporation processes are most probably responsible for the transport of solved nitrates to the mound surface and their accumulation there. The analysed mounds in central Namibia contained higher sand propor‐tions compared to the mounds of the former studies. Through the higher percentage of coarse and middle sized pores, water moves more easily in sandy soils compared to more clayey soils. In consequence, evaporation‐driven nitrate accumulation can occur in the studied mounds at high rates. Hochgerechnet auf den Gesamtumfang der Hügel bedeckte das pro Jahr von einem bewohnten Hügel erodierte Material theoretisch einen 1 m breiten Kreisring um den Schwemmkegel des Hügels 2,4 mm hoch. Der entsprechende Wert für unbewohnte Hügel betrug 1,0 mm. To assess the amount of soil that erodes from termite mounds, I fastened four strong, 65 cm wide plastic bags at 14 mounds each and collected the soil that eroded during five rainfall events. Projected to the total mound circumference, the amount of soil eroded covers theoretically a 1 m wide circular ring around the pediment of an inhabited mound up to a height of 2.4 mm per year. For uninhabited mounds, the height of this soil layer would be 1.0 mm. Per hectare, roughly 245 kg eroded per year from the mounds. However, as the erosion rate depends on several factors such as rainfall intensity, soil texture and point of time within the rainy season, this is only a vague estimate. In order to determine up to which distance the soil erosion from the mounds still influences the chemical characteristics of the adjacent topsoil, I took samples from depth of 0–10 cm at 1, 5 and 25 m distances, respectively, from four different mounds and from the mounds themselves. The non‐metric multidimensional scaling of the soil properties showed strong differences between mound and off‐mound samples. Soil characteristics within the samples from the mounds did not differ largely. Similarly, I found no strong differences between the samples taken from the different distances from the mound. From these results I conclude that through the construction of foraging galleries and sheetings (soil constructions with which some termite species cover their food items), the soil eroding from termite mounds is quickly mixed with deeper soil layers. In consequence, mound material does not accumulate in the mound’s vicinity. In order to reveal how plant growth is influenced by termite mound material, we assessed the number of grass and herb individuals as well as the biomass of plants growing in situ on the base of mounds compared to adjacent sites. While the numbers of both grass and herb individuals were significantly lower compared to adjacent sites, the total biomass of plants growing on the base of mounds was significantly higher. Reverse results were obtained by pot experiments with radish (Raphanus sativus subsp. sativus) and sorghum (Sorghum sp.) growth. Both species grew significantly weaker on mound soil compared to adjacent soil. The contradictory results concerning the biomass of in situ and pot experi‐ments are most probably caused by the disturbance of the original soil structure during the potting process. The material was subsequently compacted through watering the plants. In contrast, Macrotermes mounds are pervaded by many macropores which seem to be essential for the plant roots to penetrate the soil. In the last part of this thesis, I posed the question how mounds of M. michaelseni are distributed and what factors might be responsible for this pattern. Former studies showed that mound size is correlated with the size of its inhabiting colony. With several multi‐scale analyses, I revealed that larger inhabited mounds were regularly distributed. Additionally, mounds which were closer together tended to be smaller than on average. This indicates that intraspecific competition controls the distribution and size of colonies and their mounds. Former studies concerning Odontotermes mounds substantiated that they are local hotspots of primary productivity and animal abundance. Based on these findings, simulations revealed that a regular distribution of these mounds leads to a greater ecosystem‐wide productivity compared to a random arrangement. As in the present study, plant biomass was higher at the mounds compared to off‐mound sites, this might hold true for M. michaelseni mounds. From the results of this thesis, I draw the conclusion that through their mound building activities, M. michaelseni strongly influences the distribution patterns of soil nutrients within the central Namibian savanna. These termites create sharp contrasts in nutrient levels and vegetation patterns between mound soils and off‐mound soils and enhance the heterogeneity of their habitats. Former studies revealed that habitat hetero‐geneity is important in generating species diversity and species richness in turn is correlated positively with biomass production and positively affects ecosystem services. In conclusion, the present thesis underlines the importance of M. michaelseni for ecosystem functioning of the central Namibian savanna.
Originally renowned for their spectacular epigaeic raids, army ants have captured scientific attention for almost two centuries. They now belong to one of the best studied group of ants. However, most of our knowledge about army ants was derived from the study of the minority of specialized, epigaeicly active species. These species evolved probably rather recently from hypogaeic ancestors. The majority of army ant species still leads a hypogaeic life and is almost completely unknown in its entire sociobiology. It thus remained speculative, whether the assumed 'general' characteristics of army ants represent an adaptation to epigaeic activity or apply also to the majority of hypogaeic species. Based on the recent observation that the hypogaeic Asian army ant Dorylus (Dichthadia) laevigatus recruits predictably to palm oil baits, I developed and tested an oil-baiting method for the study of hypogaeic (army)ants. Prior to my study, nothing was known about the sociobiology of the assumed rare D. laevigatus. Throughout my work, I showed D. laevigatus to be very common and abundant in a wide range of habitats in West-Malaysia and on Borneo. Investigating its foraging behavior, I revealed D. laevigatus to differ from epigaeicly active species in several ways. Never demonstrated for any of the epigaeic species, D. laevigatus established stable trunk trail systems. Such a trail system contradicted the perception of army ant foraging, which was believed to be characterized by raids with constantly alternating trail directions. The trunk trail system further enabled a near omnipresence of D. laevigatus within its foraging area, which was also believed to be atypical for an army ant. Raids differed in structure and composition of participating workers from those of epigaeic species. Also, bulky food sources could be exploited over long periods of time. The foraging system of D. laevigatus resembled in several ways that of e.g. leaf-cutter and harvester ants. Likewise contrary to the assumptions, D. laevigatus had a wide food spectrum and showed only little effect on local arthropod communities, even falling itself prey to other ants. Strong aggressive behavior was observed only towards ant species with similar lifestyles, enabling me to provide the first detailed documentation of interspecific fights between two sympatric Dorylus species. Similar to foraging habits or ecological impact, nothing was known about colony size and composition, nesting habits, or worker polymorphism for D. laevigatus or any other hypogaeic Dorylus species prior to my work. By observing and eventually excavating a colony, I showed D. laevigatus to have a much smaller colony size and to lack the large sized workers of epigaeic Dorylus species. Similar to epigaeic Dorylinae, I showed D. laevigatus to have a non-phasic brood production, to emigrate rarely, and to alter its nest form along with habitat conditions. Detailed morphological and geographical descriptions give an impression of the Asian Dorylus species and are expected to aid other researchers in the difficult species identification. The genetic analysis of a male collected at a light trap demonstrated its relation to D. laevigatus. Confirming the male and queen associations, D. laevigatus is now one of five Dorylus species (out of a total of 61), for which all castes are known. In cooperation with D. Kistner, I provide a morphological and taxonomical description of nine Coleopteran beetles associated with D. laevigatus. Behavioral observations indicated the degree of their integration into the colony. The taxonomic position of the beetles further indicated that D. laevigatus emigrated from Africa to Asia, and was accompanied by the majority of associated beetles. The diversity of D. laevigatus guests, which included a number of unidentified mites, was rather low compared to that of epigaeic species. Overall, I demonstrated the developed baiting containers to effectively enable the study of hypogaeic ants. I showed several other hypogaeic ant species to be undersampled by other methods. Furthermore, the method enabled me to documented a second hypogaeic Dorylus species on Borneo. A detailed description of this species' morphology, ecology, and interactions with D. laevigatus is provided. My study indicated D. laevigatus to be an ecologically important species, able to influence soil structure and organisms of tropical regions in many ways. Relating the observed traits of D. laevigatus to epigaeicly active species, I conclude that our assumption of 'general' army ant behavior is erroneous in several aspects and needs to be changed. The oil-baiting method finally provides a tool enabling the location and study of hypogaeic (army)ant species. This opens a broad field for future studies on this cryptic but nonetheless important group of ants.