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- Melampyrum pratense (2)
- NE Germany (2)
- dispersal (2)
- epizoochory (2)
- seed dispersal (2)
- <i>Fusarium oxysporum</i> (1)
- <i>Zygogonium ericetorum</i> (1)
- Brandenburg (1)
- Clonal growth (1)
- Diaspore morphology (1)
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Questions: 1. Are there differences among species in their preference for coniferous vs. deciduous forest? 2. Are tree and shrub species better colonizers of recent forest stands than herbaceous species? 3. Do colonization patterns of plant species groups depend on tree species composition? Location: Three deciduous and one coniferous recent forest areas in Brandenburg, NE Germany. Methods: In 34 and 21 transects in coniferous and deciduous stands, respectively, we studied the occurrence and percentage cover of vascular plants in a total of 150 plots in ancient stands, 315 in recent stands and 55 at the ecotone. Habitat preference, diaspore weight, generative dispersal potential and clonal extension were used to explain mechanisms of local migration. Regression analysis was conducted to test whether migration distance was related to species’ life-history traits. Results: 25 species were significantly associated with ancient stands and ten species were significantly more frequent in recent stands. Tree and shrub species were good colonizers of recent coniferous and deciduous stands. In the coniferous stands, all herbaceous species showed a strong dispersal limitation during colonization, whereas in the deciduous stands generalist species may have survived in the grasslands which were present prior to afforestation. Conclusions: The fast colonization of recent stands by trees and shrubs can be explained by their effective dispersal via wind and animals. This, and the comparably efficient migration of herbaceous forest specialists into recent coniferous stands, implies that the conversion of coniferous into deciduous stands adjacent to ancient deciduous forests is promising even without planting of trees.
The paper presents a simulation and parameter-estimation approach for evaluating stochastic patterns of population growth and spread of an annual forest herb, Melampyrum pratense (Orobanchaceae). The survival of a species during large-scale changes in land use and climate will depend, to a considerable extent, on its dispersal and colonisation abilities. Predictions on species migration need a combination of field studies and modelling efforts. Our study on the ability of M. pratense to disperse into so far unoccupied areas was based on experiments in secondary woodland in NE Germany. Experiments started in 1997 at three sites where the species was not yet present, with 300 seeds sown within one square meter. Population development was then recorded until 2001 by mapping of individuals with a resolution of 5 cm. Additional observations considered density dependence of seed production. We designed a spatially explicit individual-based computer simulation model to explain the spatial patterns of population development and to predict future population spread. Besides primary drop of seeds (barochory) it assumed secondary seed transport by ants (myrmecochory) with an exponentially decreasing dispersal tail. An important feature of populationpattern explanation was the simultaneous estimation of both population-growth and dispersal parameters from consistent spatio-temporal data sets. As the simulation model produced stochastic time series and random spatially discrete distributions of individuals we estimated parameters by minimising the expectation of weighted sums of squares. These sums-ofsquares criteria considered population sizes, radial population distributions around the area of origin and distributions of individuals within squares of 25*25 cm, the range of density action. Optimal parameter values, together with the precision of the estimates, were obtained from calculating sums of squares in regular grids of parameter values. Our modelling results showed that transport of fractions of seeds by ants over distances of 1…2 m was indispensable for explaining the observed population spread that led to distances of at most 8 m from population origin within 3 years. Projections of population development over 4 additional years gave a diffusion-like increase of population area without any “outposts”. This prediction generated by the simulation model gave a hypothesis which should be revised by additional field observations. Some structural deviations between observations and model output already indicated that for full understanding of population spread the set of dispersal mechanisms assumed in the model may have to be extended by additional features of plant-animal mutualism.
Meta‐communities of habitat islands may be essential to maintain biodiversity in anthropogenic landscapes allowing rescue effects in local habitat patches. To understand the species‐assembly mechanisms and dynamics of such ecosystems, it is important to test how local plant‐community diversity and composition is affected by spatial isolation and hence by dispersal limitation and local environmental conditions acting as filters for local species sorting. We used a system of 46 small wetlands (kettle holes)—natural small‐scale freshwater habitats rarely considered in nature conservation policies—embedded in an intensively managed agricultural matrix in northern Germany. We compared two types of kettle holes with distinct topographies (flatsloped, ephemeral, frequently plowed kettle holes vs. steep‐sloped, more permanent ones) and determined 254 vascular plant species within these ecosystems, as well as plant functional traits and nearest neighbor distances to other kettle holes. Differences in alpha and beta diversity between steep permanent compared with ephemeral flat kettle holes were mainly explained by species sorting and niche processes and mass effect processes in ephemeral flat kettle holes. The plant‐community composition as well as the community trait distribution in terms of life span, breeding system, dispersal ability, and longevity of seed banks significantly differed between the two habitat types. Flat ephemeral kettle holes held a higher percentage of non‐perennial plants with a more persistent seed bank, less obligate outbreeders and more species with seed dispersal abilities via animal vectors compared with steep‐sloped, more permanent kettle holes that had a higher percentage of wind‐dispersed species. In the flat kettle holes, plant‐species richness was negatively correlated with the degree of isolation, whereas no such pattern was found for the permanent kettle holes. Synthesis: Environment acts as filter shaping plant diversity (alpha and beta) and plant‐community trait distribution between steep permanent compared with ephemeral flat kettle holes supporting species sorting and niche mechanisms as expected, but we identified a mass effect in ephemeral kettle holes only. Flat ephemeral kettle holes can be regarded as meta‐ecosystems that strongly depend on seed dispersal and recruitment from a seed bank, whereas neighboring permanent kettle holes have a more stable local species diversity.
Structure and reactivity of a biological soil crust from a xeric sandy soil in Central Europe
(2004)
The investigation was designed to explore the structure, composition and activity of a biological soil crust on an acidic, sandy soil from a temperate climate. The crust covers several hundreds of square meters on the hilltop of a large terminal moraine. The conjugate alga Zygogonium ericetorum forms the essential matrix for the crust, a dense web of algal filaments with interspersed lichens and mosses. The crust is composed of three layers, with an uppermost layer consisting nearly entirely of a dense algal mat. In lower layers, a parasitic fungus, penetrating the algal cells, is another important component of the crust community. In this soil crust, photosynthetic and respiratory activity is stabilized at low water activities.
Myrmecochory, i.e. dispersal of seeds by ants towards and around their nests, plays an important role in temperate forests. Yet hardly any study has examined plant population spread over several years and the underlying joint contribution of a hierarchy of dispersal modes and plant demography. We used a seed-sowing approach with three replicates to examine colonization patterns of Melampyrum pratense, an annual myrmecochorous herb, in a mixed Scots pine forest in northeastern Germany. Using a spatially explicit individualbased (SEIB) model population patterns over 4 years were explained by short-distance transport of seeds by small ant species with high nest densities, resulting in random spread. However, plant distributions in the field after another 4 years were clearly deviating from model predictions. Mean annual spread rate increased from 0.9 m to 5.1 m per year, with a clear inhomogeneous component. Obviously, after a lag-phase of several years, non-random seed dispersal by large red wood ants (Formica rufa) was determining the species’ spread, thus resulting in stratified dispersal due to interactions with different-sized ant species. Hypotheses on stratified dispersal, on dispersal lag, and on non-random dispersal were verified using an extended SEIB model, by comparison of model outputs with field patterns (individual numbers, population areas, and maximum distances). Dispersal towards red wood ant nests together with seed loss during transport and redistribution around nests were essential features of the model extension. The observed lag-phase in the initiation of non-random, medium-distance transport was probably due to a change of ant behaviour towards a new food source of increasing importance, being a meaningful example for a lag-phase in local plant species invasion. The results demonstrate that field studies should check model predictions wherever possible. Future research will show whether or not the M. pratense–ant system is representative for migration patterns of similar animal dispersal systems after having crossed range edges by long-distance dispersal events.
Im Norddeutschen Tiefland wurde die Ausbreitung von Gefäßpflanzen durch Rehe, Dam- und Rothirsche sowie Wildschweine untersucht. Diese Tiere transportieren zahlreiche Pflanzenarten in teilweise erheblichen Mengen über größere Distanzen, sowohl durch den Kot nach Darmpassage (Endozoochorie) als auch durch Anheftung an Fell und Schalen (Epizoochorie). Besondere Bedeutung kommt dabei Wildschweinen zu, die potenziell fast alle Pflanzenarten ausbreiten können. Bevorzugt werden im Wald wie im Offenland vorkommende Pflanzen und Arten des Offenlands ausgebreitet, während Arten mit enger Waldbindung nur in geringem Maße transportiert werden. Zoochorie durch Schalenwild bietet Erklärungsansätze sowohl für Ausbreitungsphänomene wie auch für das weitgehend fehlende Ausbreitungspotenzial vieler Pflanzenarten. Der Einfluss des Schalenwilds auf die Artenzusammensetzung und Gefäßpflanzen-Diversität in der mitteleuropäischen Kulturlandschaft sollte in seine naturschutzfachliche Neubewertung miteinbezogen werden. Die Einschränkung von Aktionsradien der Tiere durch die Zerschneidung von Lebensräumen sowie die Wildfütterung können für Ausbreitungsprozesse bisher kaum beachtete Konsequenzen haben.
Soil seed banks near rubbing trees indicate dispersal of plant species into forests by wild boar
(2006)
Current knowledge about processes that generate long-distance dispersal of plants is still limited despite its importance for persistence of populations and colonization of new potential habitats. Today wild large mammals are presumed to be important vectors for long-distance transport of diaspores within and between European temperate forest patches, and in particular wild boars recently came into focus. Here we use a specific habit of wild boar, i.e. wallowing in mud and subsequent rubbing against trees, to evaluate epizoic dispersal of vascular plant diaspores. We present soil seed bank data from 27 rubbing trees versus 27 control trees from seven forest areas in Germany. The mean number of viable seeds and the plant species number were higher in soil samples near rubbing trees compared with control trees. Ten of the 20 most frequent species were more frequent, and many species exclusively appeared in the soil samples near rubbing trees. The large number of plant species and seeds – approximated > 1000 per tree – in the soils near rubbing trees is difficult to explain unless the majority were dispersed by wild boar. Hooked and bristly diaspores, i.e. those adapted to epizoochory, were more frequent, above that many species with unspecialised diaspores occurred exclusively near rubbing trees. Different to plant species closely tied to forest species which occur both in forest and open vegetation, and non-forest species were more frequent near rubbing trees compared with controls. These findings are consistent with previous studies on diaspore loads in the coats and hooves of shot wild boars. However, our method allows to identify the transport of diaspores from the open landscape into forest stands where they might especially emerge after disturbance, and a clustered distribution of epizoochorically dispersed seeds. Moreover, accumulation of seeds of wetness indicators near rubbing trees demonstrates directed dispersal of plant species inhabiting wet places between remote wallows.
The external dispersal ("epizoochory") of vascular plant diaspores (seeds and fruits) by roe deer and wild boar, i.e. the most common wild large mammals with a large home range in central Europe, was investigated in a 6.5-km² forest area in NE Germany dominated by mesic deciduous forests. The study involved brushing out the diaspores from the coats and hooves of 25 shot roe deer and nine wild boar. The results were compared with the forest vegetation of the study area. Whilst wild boar transported large amounts of various diaspores in the coat, the significance of roe deer for epizoochory was low due to their sleek fur and different behaviour compared to wild boar. Altogether, 55 vascular plant species were transported externally. Since only a limited number of seeds came from woodland habitats, the open landscape was at least as important as a source of attached seeds as the forest vegetation. Thus, most plant species occurring in the studied forest area, especially characteristic woodland herbs, showed no adaptations to epizoochorous dispersal, although being very abundant in the herb layer. We conclude that hoofed game play a particular role concerning the dispersal of ruderal and grassland species in the agricultural landscape of central Europe. However, the actual spread of some herb species in forests of northern Germany, e.g. Agrostis capillaris, Brachypodium sylvaticum, Deschampsia flexuosa, Galium aparine and Urtica dioica, may be mainly facilitated by wild ungulates. Though dispersal by large mammals is an important mechanism for long-distance dispersal of plants in general, our results suggest that most of the characteristic herb species of mesic deciduous forests have only low epizoochorous dispersal potentials. The implications for nature conservation and silviculture are discussed.
Natürliche Standorte der Waldkiefer gibt es in Deutschland nur kleinflächig. Während Kiefernforste anstelle natürlicher Laubwälder heute oft landschaftsprägend sind, bildet die konkurrenzschwache und lichtbedürftige Kiefer ausschließlich auf extrem trockenen oder nassen, nährstoffarmen Standorten naturnahe Schlusswaldgesellschaften. Regionale Schwerpunkte liegen in subkontinentalen Regionen wie dem nordostdeutschen Tiefland und Bayern, ein „natürliches Kiefernareal" lässt sich aber kaum abgrenzen. An der Trockengrenze des Waldes finden sich auf Kalk- und Dolomitgesteinen artenreiche Karbonat-Trockenkiefernwälder mit Elementen der alpinen Rasen und Kalkmagerrasen in der Bodenvegetation. Diese Wälder besiedeln steile, südexponierte Felsen und morphodynamisch aktive Bereiche wie Rutschhänge und FlussSchotterböden im Umkreis der Alpen, kommen aber auch in den Mittelgebirgen vor. Ihr Gegenstück auf sauren Standorten sind die Sand- und Silikat-Kiefernwälder der Quarzsande und Sandstein-Verwitterungsböden, deren Bodenvegetation durch Zwergsträucher, Moose und Strauchflechten geprägt ist. Hier siedelt die Kiefer in den Tieflagen besonders auf Binnendünen und Sandern, aber auch auf Küstendünen der Ostsee, in den Mittelgebirgen z. B. auf den Sandsteinriffen der Sächsischen Schweiz. Der dritte Wuchsbereich natürlicher Kiefernwälder sind saure, nährstoffarme Moore, die ganz überwiegend von Regenwasser gespeist werden. Auch die Kiefern-Moorwälder sind in Nordostdeutschland und Bayern am häufigsten. Von diesen Standorten ausgehend, wo ihr Platz kaum von anderen Baumarten streitig gemacht wird, tritt die Waldkiefer immer wieder als Pionier auf weniger extremen Standorten auf. In der Naturlandschaft kam dies etwa nach Waldbränden oder Stürmen vor, doch der Mensch förderte die Kiefer durch Auflichtung der Wälder, Waldweide und Streunutzung stark. Auch die damit verbundene Nährstoffverarmung macht eine exakte Abgrenzung natürlicher Kiefernstandorte unmöglich. Die schlechtwüchsigen und forstwirtschaftlich nicht interessanten, ästhetisch aber sehr ansprechenden natürlichen Kiefernbestände sind heute vor allem durch Stickstoff-Immissionen gefährdet. Trotz ihrer oft kargen Erscheinung besitzen sie einen hohen Wert für die Biodiversität und den Artenschutz. Neben bodenbewohnenden Flechten und regionalen Relikt-Endemiten ist vor allem die in den letzten Jahrzehnten zunehmend gefährdete Vielfalt an Mykorrhiza-Pilzen hervorzuheben, die der Kiefer das Leben auf extrem nährstoffarmen Standorten überhaupt ermöglichen. Abschließend werden mögliche Schutz- bzw. Regenerationsmaßnahmen wie das Abplaggen flechtenreicher Kiefernstandorte vorgestellt.
Sand- und Silikat-Kiefernwälder (Dicrano-Pinion) in Deutschland : Gliederungskonzept und Ökologie
(2007)
In preparation for the „Synopsis of plant communities of Germany“ a comprehensive classification concept for the Scots pine forests on sandy and silicate soils is presented. On the basis of 2699 relevés from all natural provinces with important occurrences this classification for the first time integrates both northern and southern German forest stands. Pine forests are stable (“climax”) communities on three distinct habitat types at the drought and wetness limits of forest growth. In the phytosociological system these are reflected by the clearly separated syntaxa Erico-Pinetea (dry-calcareous), Dicrano-Pinion (dry-acidic) and Vaccinio uliginosi- Pinetea (wet-acidic). However, Pulsatillo-Pinetea (dry-moderate basicity) described in earlier publications cannot be separated floristically. In addition to the stable communities on extreme habitats pine forests of the mentioned syntaxa are widespread on potential mixed deciduous forest stands, especially after anthropogenic devastation and even beyond their original range. Six communites of the Dicrano-Pinion which also includes such secondary pine forest stands are occurring in Germany. They are presented in detail and classified according to their dynamic and edaphic differentiation. Lichen-rich pine forests (Cladonio- Pinetum) which grow on extremely dry and nutrient-poor sites are ecologically and floristically well-defined, though closely connected with other Dicrano-Pinion communities by forest succession. After separation of the Cladonio-Pinetum the Leucobryo-Pinetum is a speciespoor “central association” within the alliance. The Deschampsia flexuosa-Pinus-sylvestriscommunity is the most widespread forest type and dynamically and floristically passes into the mixed oak forests on acidic soils (Quercion roboris). On base-rich habitats the Empetro- Pinetum as endemic community of the southern Baltic Sea coasts, and the Peucedano-Pinetum in the northeastern and southern German inland are distinguished. The latter is found both on calcareous sands and primarily acidic sands which are secondary limed by calciferous pollutions. Finally, differences and similarities between the geographically separated northern and southern German Dicrano-Pinion forests are discussed in a biogeographic context, emphasising the advantages of the presented nation-wide classification concept.
A seed sowing experiment was conducted in a mixed secondary woodland on acidic soils in NE Germany with Melampyrum pratense, an annual ant-dispersed forest herb which lacks a natural population in the study area, but is abundant in similar habitats. Each set of 300 seeds was sown within one square metre at three sites in 1997, and the development of the populations was recorded from 1998 onward. Additionally, seed fall patterns were studied in a natural population by means of adhesive cardboard. All trials resulted in the recruitment of populations, which survived and increased in both individual number and area, up to the year 2001. Thus, local distribution of Melampyrum pratense is dispersallimited. Total individual number increased from 105 to 3,390, and total population area from 2.07 to 109.04 m². Migration occurred in all directions. Mean migration rate was 0.91 m per year, and the highest migration rate was 6.48 m. No individual was recorded beyond 7.63 m from the centres of the sawn squares after three years, suggesting exclusive short-distance dispersal. As primary dispersal enables only distances of up to 0.25 m, ants are presumed to be the main dispersal vectors. Despite differences in individual number and colonization patterns, migration rates did not differ significantly between the populations, but were significantly higher in 2001 due to an increased population size. Colonization patterns were characterized by a rapid, negative exponential decrease of population density with increasing distance from the sown plot, suggesting a colonization by establishment of more or less isolated outposts of individuals and a subsequent gradual infill of the gaps between. My results resemble myrmecochorous dispersal distances in temperate woodlands, and migration rates and patterns across ecotones from ancient to recent deciduous forests. They may function as a colonization model of Melampyrum pratense after accidental long-distance dispersal.
In einem rund 2.200 ha großen Waldgebiet bei Magdeburgerforth (Fläming, Sachsen-Anhalt) wur-den 1948 bis 1950 von Harro Passarge 120 Vegetationsaufnahmen sowie eine Vegetationskartierung erstellt. Das Gebiet zeichnet sich durch eine große Vielfalt an Waldtypen aus den Verbänden Agrostio-Quercion petraeae, Alnion glutinosae, Alnion incanae, Carpinion betuli, Dicrano-Pinion und Quercion roboris aus. Daher und weil viele der heute in Wäldern wirksamen Prozesse (z. B. Stickstoffeintrag, Klimawandel) vor 60 Jahren noch nicht spürbar waren, bietet sich das Gebiet für eine Wiederholungs-untersuchung besonders an. Da die Aufnahmeflächen von Passarge nicht punktgenau verortet waren, wurden im Jahr 2014 in einem über die Forstabteilungen und die Vegetationskarte definierten Such-raum immer die der Erstaufnahme ähnlichsten Waldbestände erfasst. Insgesamt konnten 97 (81 %) der Aufnahmen wiederholt werden. Vegetationsveränderungen werden mithilfe einer NMDS-Ordination, der Gegenüberstellung von α-Diversität, Zeigerwerten und Waldbindungskategorien für die beiden Aufnahmezeitpunkte sowie über die Identifikation von Gewinner- und Verlierer-Arten analysiert.Auch wenn methodenbedingt bei der Wiederholungsuntersuchung nur die jeweils geringstmögliche Vegetationsveränderung abgebildet wird, konnten Ergebnisse erzielt werden, die mit denen quasi-permanenter Plots übereinstimmen. Die beobachteten allgemeinen Trends (Eutrophierung, Sukzession nach Nutzungswandel, Verlust lichtliebender und magerkeitszeigender Arten, Ausbreitung von stick-stoffliebenden Arten und mesophilen Waldarten, Einwanderung von Neophyten, keine generelle Ab-nahme der Artenzahl) stimmen gut mit den in zahlreichen Studien aus mitteleuropäischen Wäldern festgestellten überein. Durch das von nassen bis trockenen sowie von bodensauer-nährstoffarmen bis zu relativ basenreichen Böden reichende Standortsspektrum innerhalb des Untersuchungsgebietes konnte aber – deutlicher als in den meisten bisherigen Fallstudien – gezeigt werden, dass sich die Resilienz der Wälder gegenüber Vegetationsveränderung je nach Ausgangsgesellschaft stark unterscheidet und jeweils unterschiedliche Treiber wirksam sind. Stellario-Carpinetum und Luzulo-Quercetum erwiesen sich als relativ stabil, und auch in den Feuchtwäldern des Circaeo-Alnetum gab es trotz eines Arten-wechsels wenig Hinweise auf Umweltveränderungen. Dagegen wiesen die Wälder nährstoffarmer Standorte (Sphagno-Alnetum, Betulo-Quercetum, Dicrano-Pinion) viele Verliererarten und eine starke Eutrophierungstendenz auf. Die in besonderem Maße von historischen Waldnutzungsformen abhängi-gen thermophilen Wälder und die Flechten-Kiefernwälder gingen weitgehend verloren.