@misc{HeinkenWinkler2009,
  author    = {Heinken, Thilo and Winkler, Eckart},
  title     = {Non-random dispersal by ants : long-term field data versus model predictions of population spread of a forest herb},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-46482},
  year      = {2009},
  abstract  = {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.},
  language  = {en}
}
@misc{Heinken2008,
  author    = {Heinken, Thilo},
  title     = {Die nat{\"u}rlichen Kiefernstandorte Deutschlands und ihre Gef{\"a}hrdung},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-46506},
  year      = {2008},
  abstract  = {Nat{\"u}rliche Standorte der Waldkiefer gibt es in Deutschland nur kleinfl{\"a}chig. W{\"a}hrend Kiefernforste anstelle nat{\"u}rlicher Laubw{\"a}lder heute oft landschaftspr{\"a}gend sind, bildet die konkurrenzschwache und lichtbed{\"u}rftige Kiefer ausschließlich auf extrem trockenen oder nassen, n{\"a}hrstoffarmen Standorten naturnahe Schlusswaldgesellschaften. Regionale Schwerpunkte liegen in subkontinentalen Regionen wie dem nordostdeutschen Tiefland und Bayern, ein „nat{\"u}rliches Kiefernareal" l{\"a}sst sich aber kaum abgrenzen. An der Trockengrenze des Waldes finden sich auf Kalk- und Dolomitgesteinen artenreiche Karbonat-Trockenkiefernw{\"a}lder mit Elementen der alpinen Rasen und Kalkmagerrasen in der Bodenvegetation. Diese W{\"a}lder besiedeln steile, s{\"u}dexponierte Felsen und morphodynamisch aktive Bereiche wie Rutschh{\"a}nge und FlussSchotterb{\"o}den im Umkreis der Alpen, kommen aber auch in den Mittelgebirgen vor. Ihr Gegenst{\"u}ck auf sauren Standorten sind die Sand- und Silikat-Kiefernw{\"a}lder der Quarzsande und Sandstein-Verwitterungsb{\"o}den, deren Bodenvegetation durch Zwergstr{\"a}ucher, Moose und Strauchflechten gepr{\"a}gt ist. Hier siedelt die Kiefer in den Tieflagen besonders auf Binnend{\"u}nen und Sandern, aber auch auf K{\"u}stend{\"u}nen der Ostsee, in den Mittelgebirgen z. B. auf den Sandsteinriffen der S{\"a}chsischen Schweiz. Der dritte Wuchsbereich nat{\"u}rlicher Kiefernw{\"a}lder sind saure, n{\"a}hrstoffarme Moore, die ganz {\"u}berwiegend von Regenwasser gespeist werden. Auch die Kiefern-Moorw{\"a}lder sind in Nordostdeutschland und Bayern am h{\"a}ufigsten. 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 Waldbr{\"a}nden oder St{\"u}rmen vor, doch der Mensch f{\"o}rderte die Kiefer durch Auflichtung der W{\"a}lder, Waldweide und Streunutzung stark. Auch die damit verbundene N{\"a}hrstoffverarmung macht eine exakte Abgrenzung nat{\"u}rlicher Kiefernstandorte unm{\"o}glich. Die schlechtw{\"u}chsigen und forstwirtschaftlich nicht interessanten, {\"a}sthetisch aber sehr ansprechenden nat{\"u}rlichen Kiefernbest{\"a}nde sind heute vor allem durch Stickstoff-Immissionen gef{\"a}hrdet. Trotz ihrer oft kargen Erscheinung besitzen sie einen hohen Wert f{\"u}r die Biodiversit{\"a}t und den Artenschutz. Neben bodenbewohnenden Flechten und regionalen Relikt-Endemiten ist vor allem die in den letzten Jahrzehnten zunehmend gef{\"a}hrdete Vielfalt an Mykorrhiza-Pilzen hervorzuheben, die der Kiefer das Leben auf extrem n{\"a}hrstoffarmen Standorten {\"u}berhaupt erm{\"o}glichen. Abschließend werden m{\"o}gliche Schutz- bzw. Regenerationsmaßnahmen wie das Abplaggen flechtenreicher Kiefernstandorte vorgestellt.},
  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}
}
@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{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{HeinkenRaudnitschka2002,
  author    = {Heinken, Thilo and Raudnitschka, Dorit},
  title     = {Do wild ungulates contribute to the dispersal of vascular plants in central European forests by epizoochory?},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-5850},
  year      = {2002},
  abstract  = {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.},
  language  = {en}
}
@misc{Heinken2004,
  author    = {Heinken, Thilo},
  title     = {Migration of an annual myrmecochore},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-5865},
  year      = {2004},
  abstract  = {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.},
  language  = {en}
}
@article{PlueDeFrenneAcharyaetal.2017,
  author    = {Plue, Jan and De Frenne, Pieter and Acharya, Kamal and Brunet, J{\"o}rg and Chabrerie, Olivier and Decocq, Guillaume and Diekmann, Martin and Graae, Bente J. and Heinken, Thilo and Hermy, Martin and Kolb, Annette and Lemke, Isgard and Liira, Jaan and Naaf, Tobias and Verheyen, Kris and Wulf, Monika and Cousins, Sara A. O.},
  title     = {Where does the community start, and where does it end?},
  series = {Journal of vegetation science},
  volume    = {28},
  journal   = {Journal of vegetation science},
  number    = {2},
  publisher = {Wiley},
  address   = {Hoboken},
  issn      = {1100-9233},
  doi       = {10.1111/jvs.12493},
  pages     = {424 -- 435},
  year      = {2017},
  abstract  = {QuestionBelow-ground processes are key determinants of above-ground plant population and community dynamics. Still, our understanding of how environmental drivers shape plant communities is mostly based on above-ground diversity patterns, bypassing below-ground plant diversity stored in seed banks. As seed banks may shape above-ground plant communities, we question whether concurrently analysing the above- and below-ground species assemblages may potentially enhance our understanding of community responses to environmental variation. LocationTemperate deciduous forests along a 2000km latitudinal gradient in NW Europe. MethodsHerb layer, seed bank and local environmental data including soil pH, canopy cover, forest cover continuity and time since last canopy disturbance were collected in 129 temperate deciduous forest plots. We quantified herb layer and seed bank diversity per plot and evaluated how environmental variation structured community diversity in the herb layer, seed bank and the combined herb layer-seed bank community. ResultsSeed banks consistently held more plant species than the herb layer. How local plot diversity was partitioned across the herb layer and seed bank was mediated by environmental variation in drivers serving as proxies of light availability. The herb layer and seed bank contained an ever smaller and ever larger share of local diversity, respectively, as both canopy cover and time since last canopy disturbance decreased. Species richness and -diversity of the combined herb layer-seed bank community responded distinctly differently compared to the separate assemblages in response to environmental variation in, e.g. forest cover continuity and canopy cover. ConclusionsThe seed bank is a below-ground diversity reservoir of the herbaceous forest community, which interacts with the herb layer, although constrained by environmental variation in e.g. light availability. The herb layer and seed bank co-exist as a single community by means of the so-called storage effect, resulting in distinct responses to environmental variation not necessarily recorded in the individual herb layer or seed bank assemblages. Thus, concurrently analysing above- and below-ground diversity will improve our ecological understanding of how understorey plant communities respond to environmental variation.},
  language  = {en}
}
@article{DeLombaerdeVerheyenPerringetal.2018,
  author    = {De Lombaerde, Emiel and Verheyen, Kris and Perring, Michael P. and Bernhardt-Roemermann, Markus and Van Calster, Hans and Brunet, Jorg and Chudomelova, Marketa and Decocq, Guillaume and Diekmann, Martin and Durak, Tomasz and Hedl, Radim and Heinken, Thilo and Hommel, Patrick and Jaroszewicz, Bogdan and Kopecky, Martin and Lenoir, Jonathan and Macek, Martin and M{\´a}liš, František and Mitchell, Fraser J. G. and Naaf, Tobias and Newman, Miles and Petř{\´i}k, Petr and Reczyńska, Kamila and Schmidt, Wolfgang and Swierkosz, Krzysztof and Vild, Ondrej and Wulf, Monika and Baetena, Lander},
  title     = {Responses of competitive understorey species to spatial environmental gradients inaccurately explain temporal changes},
  series = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie},
  volume    = {30},
  journal   = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie},
  publisher = {Elsevier GMBH},
  address   = {M{\"u}nchen},
  issn      = {1439-1791},
  doi       = {10.1016/j.baae.2018.05.013},
  pages     = {52 -- 64},
  year      = {2018},
  abstract  = {Understorey plant communities play a key role in the functioning of forest ecosystems. Under favourable environmental conditions, competitive understorey species may develop high abundances and influence important ecosystem processes such as tree regeneration. Thus, understanding and predicting the response of competitive understorey species as a function of changing environmental conditions is important for forest managers. In the absence of sufficient temporal data to quantify actual vegetation changes, space-for-time (SFT) substitution is often used, i.e. studies that use environmental gradients across space to infer vegetation responses to environmental change over time. Here we assess the validity of such SFT approaches and analysed 36 resurvey studies from ancient forests with low levels of recent disturbances across temperate Europe to assess how six competitive understorey plant species respond to gradients of overstorey cover, soil conditions, atmospheric N deposition and climatic conditions over space and time. The combination of historical and contemporary surveys allows (i) to test if observed contemporary patterns across space are consistent at the time of the historical survey, and, crucially, (ii) to assess whether changes in abundance over time given recorded environmental change match expectations from patterns recorded along environmental gradients in space. We found consistent spatial relationships at the two periods: local variation in soil variables and overstorey cover were the best predictors of individual species' cover while interregional variation in coarse-scale variables, i.e. N deposition and climate, was less important. However, we found that our SFT approach could not accurately explain the large variation in abundance changes over time. We thus recommend to be cautious when using SFT substitution to infer species responses to temporal changes.},
  language  = {en}
}
@article{PerringBernhardtRoemermannBaetenetal.2018,
  author    = {Perring, Michael P. and Bernhardt-Roemermann, Markus and Baeten, Lander and Midolo, Gabriele and Blondeel, Haben and Depauw, Leen and Landuyt, Dries and Maes, Sybryn L. and De Lombaerde, Emiel and Caron, Maria Mercedes and Vellend, Mark and Brunet, Joerg and Chudomelova, Marketa and Decocq, Guillaume and Diekmann, Martin and Dirnboeck, Thomas and Doerfler, Inken and Durak, Tomasz and De Frenne, Pieter and Gilliam, Frank S. and Hedl, Radim and Heinken, Thilo and Hommel, Patrick and Jaroszewicz, Bogdan and Kirby, Keith J. and Kopecky, Martin and Lenoir, Jonathan and Li, Daijiang and Malis, Frantisek and Mitchell, Fraser J. G. and Naaf, Tobias and Newman, Miles and Petrik, Petr and Reczynska, Kamila and Schmidt, Wolfgang and Standovar, Tibor and Swierkosz, Krzysztof and Van Calster, Hans and Vild, Ondrej and Wagner, Eva Rosa and Wulf, Monika and Verheyen, Kris},
  title     = {Global environmental change effects on plant community composition trajectories depend upon management legacies},
  series = {Global change biology},
  volume    = {24},
  journal   = {Global change biology},
  number    = {4},
  publisher = {Wiley},
  address   = {Hoboken},
  issn      = {1354-1013},
  doi       = {10.1111/gcb.14030},
  pages     = {1722 -- 1740},
  year      = {2018},
  abstract  = {The contemporary state of functional traits and species richness in plant communities depends on legacy effects of past disturbances. Whether temporal responses of community properties to current environmental changes are altered by such legacies is, however, unknown. We expect global environmental changes to interact with land-use legacies given different community trajectories initiated by prior management, and subsequent responses to altered resources and conditions. We tested this expectation for species richness and functional traits using 1814 survey-resurvey plot pairs of understorey communities from 40 European temperate forest datasets, syntheses of management transitions since the year 1800, and a trait database. We also examined how plant community indicators of resources and conditions changed in response to management legacies and environmental change. Community trajectories were clearly influenced by interactions between management legacies from over 200 years ago and environmental change. Importantly, higher rates of nitrogen deposition led to increased species richness and plant height in forests managed less intensively in 1800 (i.e., high forests), and to decreases in forests with a more intensive historical management in 1800 (i.e., coppiced forests). There was evidence that these declines in community variables in formerly coppiced forests were ameliorated by increased rates of temperature change between surveys. Responses were generally apparent regardless of sites' contemporary management classifications, although sometimes the management transition itself, rather than historic or contemporary management types, better explained understorey responses. Main effects of environmental change were rare, although higher rates of precipitation change increased plant height, accompanied by increases in fertility indicator values. Analysis of indicator values suggested the importance of directly characterising resources and conditions to better understand legacy and environmental change effects. Accounting for legacies of past disturbance can reconcile contradictory literature results and appears crucial to anticipating future responses to global environmental change.},
  language  = {en}
}