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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.
Seed traits matter
(2021)
Although many plants are dispersed by wind and seeds can travel long distances across unsuitable matrix areas, a large proportion relies on co-evolved zoochorous seed dispersal to connect populations in isolated habitat islands. Particularly in agricultural landscapes, where remaining habitat patches are often very small and highly isolated, mobile linkers as zoochorous seed dispersers are critical for the population dynamics of numerous plant species. However, knowledge about the quali- or quantification of such mobile link processes, especially in agricultural landscapes, is still limited. In a controlled feeding experiment, we recorded the seed intake and germination success after complete digestion by the European brown hare (Lepus europaeus) and explored its mobile link potential as an endozoochoric seed disperser. Utilizing a suite of common, rare, and potentially invasive plant species, we disentangled the effects of seed morphological traits on germination success while controlling for phylogenetic relatedness. Further, we measured the landscape connectivity via hares in two contrasting agricultural landscapes (simple: few natural and semi-natural structures, large fields; complex: high amount of natural and semi-natural structures, small fields) using GPS-based movement data. With 34,710 seeds of 44 plant species fed, one of 200 seeds (0.51%) with seedlings of 33 species germinated from feces. Germination after complete digestion was positively related to denser seeds with comparatively small surface area and a relatively slender and elongated shape, suggesting that, for hares, the most critical seed characteristics for successful endozoochorous seed dispersal minimize exposure of the seed to the stomach and the associated digestive system. Furthermore, we could show that a hare's retention time is long enough to interconnect different habitats, especially grasslands and fields. Thus, besides other seed dispersal mechanisms, this most likely allows hares to act as effective mobile linkers contributing to ecosystem stability in times of agricultural intensification, not only in complex but also in simple landscapes.
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.
Fluxes of organic and inorganic carbon within the Amazon basin are considerably controlled by annual flooding, which triggers the export of terrigenous organic material to the river and ultimately to the Atlantic Ocean. The amount of carbon imported to the river and the further conversion, transport and export of it depend on temperature, atmospheric CO2, terrestrial productivity and carbon storage, as well as discharge. Both terrestrial productivity and discharge are influenced by climate and land use change. The coupled LPJmL and RivCM model system (Langerwisch et al., 2016) has been applied to assess the combined impacts of climate and land use change on the Amazon riverine carbon dynamics. Vegetation dynamics (in LPJmL) as well as export and conversion of terrigenous carbon to and within the river (RivCM) are included. The model system has been applied for the years 1901 to 2099 under two deforestation scenarios and with climate forcing of three SRES emission scenarios, each for five climate models. We find that high deforestation (business-as-usual scenario) will strongly decrease (locally by up to 90 %) riverine particulate and dissolved organic carbon amount until the end of the current century. At the same time, increase in discharge leaves net carbon transport during the first decades of the century roughly unchanged only if a sufficient area is still forested. After 2050 the amount of transported carbon will decrease drastically. In contrast to that, increased temperature and atmospheric CO2 concentration determine the amount of riverine inorganic carbon stored in the Amazon basin. Higher atmospheric CO2 concentrations increase riverine inorganic carbon amount by up to 20% (SRES A2). The changes in riverine carbon fluxes have direct effects on carbon export, either to the atmosphere via outgassing or to the Atlantic Ocean via discharge. The outgassed carbon will increase slightly in the Amazon basin, but can be regionally reduced by up to 60% due to deforestation. The discharge of organic carbon to the ocean will be reduced by about 40% under the most severe deforestation and climate change scenario. These changes would have local and regional consequences on the carbon balance and habitat characteristics in the Amazon basin itself as well as in the adjacent Atlantic Ocean.
Climate change increases riverine carbon outgassing, while export to the ocean remains uncertain
(2016)
Any regular interaction of land and river during flooding affects carbon pools within the terrestrial system, riverine carbon and carbon exported from the system. In the Amazon basin carbon fluxes are considerably influenced by annual flooding, during which terrigenous organic material is imported to the river. The Amazon basin therefore represents an excellent example of a tightly coupled terrestrial-riverine system. The processes of generation, conversion and transport of organic carbon in such a coupled terrigenous-riverine system strongly interact and are climate-sensitive, yet their functioning is rarely considered in Earth system models and their response to climate change is still largely unknown. To quantify regional and global carbon budgets and climate change effects on carbon pools and carbon fluxes, it is important to account for the coupling between the land, the river, the ocean and the atmosphere. We developed the RIVerine Carbon Model (RivCM), which is directly coupled to the well-established dynamic vegetation and hydrology model LPJmL, in order to account for this large-scale coupling. We evaluate RivCM with observational data and show that some of the values are reproduced quite well by the model, while we see large deviations for other variables. This is mainly caused by some simplifications we assumed. Our evaluation shows that it is possible to reproduce large-scale carbon transport across a river system but that this involves large uncertainties. Acknowledging these uncertainties, we estimate the potential changes in riverine carbon by applying RivCM for climate forcing from five climate models and three CO2 emission scenarios (Special Report on Emissions Scenarios, SRES). We find that climate change causes a doubling of riverine organic carbon in the southern and western basin while reducing it by 20% in the eastern and northern parts. In contrast, the amount of riverine inorganic carbon shows a 2- to 3-fold increase in the entire basin, independent of the SRES scenario. The export of carbon to the atmosphere increases as well, with an average of about 30 %. In contrast, changes in future export of organic carbon to the Atlantic Ocean depend on the SRES scenario and are projected to either decrease by about 8.9% (SRES A1B) or increase by about 9.1% (SRES A2). Such changes in the terrigenous-riverine system could have local and regional impacts on the carbon budget of the whole Amazon basin and parts of the Atlantic Ocean. Changes in riverine carbon could lead to a shift in the riverine nutrient supply and pH, while changes in the exported carbon to the ocean lead to changes in the supply of organic material that acts as a food source in the Atlantic. On larger scales the increased outgassing of CO2 could turn the Amazon basin from a sink of carbon to a considerable source. Therefore, we propose that the coupling of terrestrial and riverine carbon budgets should be included in subsequent analysis of the future regional carbon budget.