@article{DiekmannMuellerHeinkenetal.2015,
  author    = {Diekmann, Martin and M{\"u}ller, Josef and Heinken, Thilo and Dupre, Cecilia},
  title     = {Survey and statistical analysis of plant reintroductions in Germany},
  series = {Tuexenia : Mitteilungen der Floristisch-Soziologischen Arbeitsgemeinschaft},
  journal   = {Tuexenia : Mitteilungen der Floristisch-Soziologischen Arbeitsgemeinschaft},
  number    = {35},
  publisher = {Floristisch-Soziologische Arbeitsgemeinschaft},
  address   = {G{\"o}ttingen},
  issn      = {0722-494X},
  pages     = {249 -- 265},
  year      = {2015},
  abstract  = {Aim - Plant reintroductions and other forms of targeted species translocations will in the future gain growing importance for nature conservation. In fragmented habitats, species reintroductions offer one of the most efficient tools for preserving or restoring plant diversity. In our study, we have compiled available data about plant reintroduction projects in Germany to answer the following questions: (1) What are the characteristics, habitat preferences and ecological strategies of species considered in plant reintroduction trials, and are these representative of the entire class of threatened species in Germany? (2) Is the judgment of the success or failure of plant reintroductions biased by the choice of species used in the experiments? (3) Do reintroduction efforts focus on those species for which Germany has a particularly high responsibility for conservation? Methods - Information about reintroduction projects in Germany were obtained from published and internet sources as well as unpublished reports. In our search we focused on single-species trials in the framework of scientific or conservation projects. For all threatened species included in our database, we compiled information on their systematics, life form, ecological strategies and habitat preferences. A list of all species being threatened nationally or regionally, comprising both reintroduced and not reintroduced species, served as a reference for statistical analysis. Results - The list of vascular plants used in conservation-oriented reintroductions consisted of 196 taxa. Species of families with large and conspicuous, mostly insect-pollinated flowers (for example, Orchidaceae) were over-represented among the reintroduced species compared to those threatened species not included in reintroduction trials. Species considered were also more often than expected found in semi-natural open habitats such as heathlands and grasslands. Notably, many projects focused on calcareous grasslands, characterized by dry, high-pH and infertile soils. In contrast, species of more near-natural vegetation (alpine and rocky formations, forests) were under-represented. About 25\% of the species that were reintroduced are not threatened on the national scale. Out of 150 species for which Germany has a particularly high responsibility for conservation, only 14 (9.3\%) were reintroduced. For only about 1/3 of all reintroduction attempts, success or failure were documented; whereas the success rate appears to be relatively low in nutrient-poor environments, trials with nutrient-demanding and competitive species were more successful. Conclusions - We conclude that conservation-oriented reintroduction attempts should focus more on species for which the country or a region has a particular high responsibility. Reintroductions, to a larger extent than at present, also need to consider the different chances of success in different habitat types and environments.},
  language  = {de}
}
@misc{HoppertReimerKemmlingetal.2004,
  author    = {Hoppert, Michael and Reimer, Rudolph and Kemmling, Anne and Schr{\"o}der, Annekatrin and G{\"u}nzl, Bettina and Heinken, Thilo},
  title     = {Structure and reactivity of a biological soil crust from a xeric sandy soil in Central Europe},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-5872},
  year      = {2004},
  abstract  = {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.},
  language  = {en}
}
@misc{WinklerHeinken2007,
  author    = {Winkler, Eckart and Heinken, Thilo},
  title     = {Spread of an ant-dispersed annual herb : an individual-based simulation study on population development of Melampyrum pratense L.},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-46491},
  year      = {2007},
  abstract  = {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.},
  language  = {en}
}
@article{WinklerHeinken2007,
  author    = {Winkler, Eckart and Heinken, Thilo},
  title     = {Spread of an ant-dispersed annual herb : an individual-based simulation study on population development of Melampyrum pratense L.},
  issn      = {0304-3800},
  year      = {2007},
  abstract  = {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 is 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 1m2. 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 population-pattern 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 sum of squares. These sums of squares criteria considered population sizes, radial population distributions around the area of origin and distributions of individuals within squares of 25cm×25 cm, the range of density action. Optimal parameter values, together with the precision of the estimates, were obtained from calculating sum of squares in regular grids of parameter values. Our modelling results showed that transport of fractions of seeds by ants over distances of 1-2m was indispensable for explaining the observed population spread that led to distances of at most 8mfrom population origin within 3 years. Projections of population development over four 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.},
  language  = {en}
}
@article{HeinkenSchmidtvonOheimbetal.2006,
  author    = {Heinken, Thilo and Schmidt, Marcus and von Oheimb, Goddert and Kriebitzsch, Wolf-Ulrich and Ellenberg, Hermann},
  title     = {Soil seed banks near rubbing trees indicate dispersal of plant species into forests by wild boar},
  issn      = {1439-1791},
  doi       = {10.1016/j.baae.2005.04.006},
  year      = {2006},
  abstract  = {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 epizoochorous 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 - more than 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; however, many species with unspecialized diaspores occurred exclusively near rubbing trees. As opposed to plant species closely tied to forests 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 among remote wallows.},
  language  = {en}
}
@misc{HeinkenSchmidtvonOheimbetal.2006,
  author    = {Heinken, Thilo and Schmidt, Marcus and von Oheimb, Goddert and Kriebitzsch, Wolf-Ulrich and Ellenberg, Hermann},
  title     = {Soil seed banks near rubbing trees indicate dispersal of plant species into forests by wild boar},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-46476},
  year      = {2006},
  abstract  = {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.},
  language  = {en}
}
@article{deFrenneGraaeKolbetal.2010,
  author    = {de Frenne, Pieter and Graae, Bente Jessen and Kolb, Annette and Brunet, J{\"o}rg and Chabrerie, Olivier and Cousins, Sara A. O. and Decocq, Guillaume and Dhondt, Rob and Diekmann, Martin and Eriksson, Olof and Heinken, Thilo and Hermy, Martin and J{\"o}gar, uelle and Saguez, Robert and Shevtsova, Anna and Stanton, Sharon and Zindel, Renate and Zobel, Martin and Verheyen, Kris},
  title     = {Significant effects of temperature on the reproductive output of the forest herb Anemone nemorosa L.},
  issn      = {0378-1127},
  doi       = {10.1016/j.foreco.2009.04.038},
  year      = {2010},
  abstract  = {Climate warming is already influencing plant migration in different parts of the world. Numerous models have been developed to forecast future plant distributions. Few studies, however, have investigated the potential effect of warming on the reproductive output of plants. Understorey forest herbs in particular, have received little attention in the debate on climate change impacts. This study focuses on the effect of temperature on sexual reproductive output (number of seeds, seed mass, germination percentage and seedling mass) of Anemone nemorosa L., a model species for slow colonizing herbaceous forest plants. We sampled seeds of A. nemorosa in populations along a 2400 km latitudinal gradient from northern France to northern Sweden during three growing seasons (2005,2006 and 2008). This study design allowed us to isolate the effects of accumulated temperature (Growing Degree Hours; GDH) from latitude and the local abiotic and biotic environment. Germination and seed sowing trials were performed in incubators, a greenhouse and under field conditions in a forest. Finally, we disentangled correlations between the different reproductive traits of A. nemorosa along the latitudinal gradient. We found a clear positive relationship between accumulated temperature and seed and seedling traits: reproductive output of A. nemorosa improved with increasing GDH along the latitudinal gradient. Seed mass and seedling mass, for instance, increased by 9.7\% and 10.4\%, respectively, for every 1000 degrees C h increase in GDH. We also derived strong correlations between several seed and seedling traits both under field conditions and in incubators. Our results indicate that seed mass, incubator-based germination percentage (Germ\%(Inc)) and the output of germinable seeds (product of number of seeds and Germ\%(Inc) divided by 100) from plants grown along a latitudinal gradient (i.e. at different temperature regimes) provide valuable proxies to parameterize key population processes in models. We conclude that (1) climate warming may have a pronounced positive impact on sexual reproduction of A. nemorosa and (2) climate models forecasting plant distributions would benefit from including the temperature sensitivity of key seed traits and population processes.},
  language  = {en}
}
@article{AlbertAuffretCosynsetal.2015,
  author    = {Albert, Aurelie and Auffret, Alistair G. and Cosyns, Eric and Cousins, Sara A. O. and Eichberg, Carsten and Eycott, Amy E. and Heinken, Thilo and Hoffmann, Maurice and Jaroszewicz, Bogdan and Malo, Juan E. and Marell, Anders and Mouissie, Maarten and Pakeman, Robin J. and Picard, Melanie and Plue, Jan and Poschlod, Peter and Provoost, Sam and Schulze, Kiowa Alraune and Baltzinger, Christophe},
  title     = {Seed dispersal by ungulates as an ecological filter: a trait-based meta-analysis},
  series = {Oikos},
  volume    = {124},
  journal   = {Oikos},
  number    = {9},
  publisher = {Wiley-Blackwell},
  address   = {Hoboken},
  issn      = {0030-1299},
  doi       = {10.1111/oik.02512},
  pages     = {1109 -- 1120},
  year      = {2015},
  abstract  = {Plant communities are often dispersal-limited and zoochory can be an efficient mechanism for plants to colonize new patches of potentially suitable habitat. We predicted that seed dispersal by ungulates acts as an ecological filter - which differentially affects individuals according to their characteristics and shapes species assemblages - and that the filter varies according to the dispersal mechanism (endozoochory, fur-epizoochory and hoof-epizoochory). We conducted two-step individual participant data meta-analyses of 52 studies on plant dispersal by ungulates in fragmented landscapes, comparing eight plant traits and two habitat indicators between dispersed and non-dispersed plants. We found that ungulates dispersed at least 44\% of the available plant species. Moreover, some plant traits and habitat indicators increased the likelihood for plant of being dispersed. Persistent or nitrophilous plant species from open habitats or bearing dry or elongated diaspores were more likely to be dispersed by ungulates, whatever the dispersal mechanism. In addition, endozoochory was more likely for diaspores bearing elongated appendages whereas epizoochory was more likely for diaspores released relatively high in vegetation. Hoof-epizoochory was more likely for light diaspores without hooked appendages. Fur-epizoochory was more likely for diaspores with appendages, particularly elongated or hooked ones. We thus observed a gradient of filtering effect among the three dispersal mechanisms. Endozoochory had an effect of rather weak intensity (impacting six plant characteristics with variations between ungulate-dispersed and non-dispersed plant species mostly below 25\%), whereas hoof-epizoochory had a stronger effect (eight characteristics included five ones with above 75\% variation), and fur-epizoochory an even stronger one (nine characteristics included six ones with above 75\% variation). Our results demonstrate that seed dispersal by ungulates is an ecological filter whose intensity varies according to the dispersal mechanism considered. Ungulates can thus play a key role in plant community dynamics and have implications for plant spatial distribution patterns at multiple scales.},
  language  = {en}
}
@misc{HeinkenvonOheimbSchmidtetal.2005,
  author    = {Heinken, Thilo and von Oheimb, Goddert and Schmidt, Marcus and Kriebitzsch, Wolf-Ulrich and Ellenberg, Hermann},
  title     = {Schalenwild breitet Gef{\"a}ßpflanzen in der mitteleurop{\"a}ischen Kulturlandschaft aus : ein erster {\"U}berblick},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-46522},
  year      = {2005},
  abstract  = {Im Norddeutschen Tiefland wurde die Ausbreitung von Gef{\"a}ßpflanzen durch Rehe, Dam- und Rothirsche sowie Wildschweine untersucht. Diese Tiere transportieren zahlreiche Pflanzenarten in teilweise erheblichen Mengen {\"u}ber gr{\"o}ßere 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 k{\"o}nnen. Bevorzugt werden im Wald wie im Offenland vorkommende Pflanzen und Arten des Offenlands ausgebreitet, w{\"a}hrend Arten mit enger Waldbindung nur in geringem Maße transportiert werden. Zoochorie durch Schalenwild bietet Erkl{\"a}rungsans{\"a}tze sowohl f{\"u}r Ausbreitungsph{\"a}nomene wie auch f{\"u}r das weitgehend fehlende Ausbreitungspotenzial vieler Pflanzenarten. Der Einfluss des Schalenwilds auf die Artenzusammensetzung und Gef{\"a}ßpflanzen-Diversit{\"a}t in der mitteleurop{\"a}ischen Kulturlandschaft sollte in seine naturschutzfachliche Neubewertung miteinbezogen werden. Die Einschr{\"a}nkung von Aktionsradien der Tiere durch die Zerschneidung von Lebensr{\"a}umen sowie die Wildf{\"u}tterung k{\"o}nnen f{\"u}r Ausbreitungsprozesse bisher kaum beachtete Konsequenzen haben.},
  language  = {de}
}
@misc{Heinken2007,
  author    = {Heinken, Thilo},
  title     = {Sand- und Silikat-Kiefernw{\"a}lder (Dicrano-Pinion) in Deutschland : Gliederungskonzept und {\"O}kologie},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-46518},
  year      = {2007},
  abstract  = {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 relev{\´e}s 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.},
  language  = {de}
}