@article{PlueDeFrenneAcharyaetal.2013,
  author    = {Plue, Jan and De Frenne, Pieter and Acharya, Kamal P. and Brunet, Jorg 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 Shevtsova, Anna and Verheyen, Kris and Wulf, Monika and Cousins, Sara A. O.},
  title     = {Climatic control of forest herb seed banks along a latitudinal gradient},
  series = {Global ecology and biogeography : a journal of macroecology},
  volume    = {22},
  journal   = {Global ecology and biogeography : a journal of macroecology},
  number    = {10},
  publisher = {Wiley-Blackwell},
  address   = {Hoboken},
  issn      = {1466-822X},
  doi       = {10.1111/geb.12068},
  pages     = {1106 -- 1117},
  year      = {2013},
  abstract  = {Aim Seed banks are central to the regeneration strategy of many plant species. Any factor altering seed bank density thus affects plant regeneration and population dynamics. Although seed banks are dynamic entities controlled by multiple environmental drivers, climatic factors are the most comprehensive, but still poorly understood. This study investigates how climatic variation structures seed production and resulting seed bank patterns. Location Temperate forests along a 1900km latitudinal gradient in north-western (NW) Europe. Methods Seed production and seed bank density were quantified in 153 plots along the gradient for four forest herbs with different seed longevity: Geum urbanum, Milium effusum, Poa nemoralis and Stachys sylvatica. We tested the importance of climatic and local environmental factors in shaping seed production and seed bank density. Results Seed production was determined by population size, and not by climatic factors. G.urbanum and M.effusum seed bank density declined with decreasing temperature (growing degree days) and/or increasing temperature range (maximum-minimum temperature). P.nemoralis and S.sylvatica seed bank density were limited by population size and not by climatic variables. Seed bank density was also influenced by other, local environmental factors such as soil pH or light availability. Different seed bank patterns emerged due to differential seed longevities. Species with long-lived seeds maintained constant seed bank densities by counteracting the reduced chance of regular years with high seed production at colder northern latitudes. Main conclusions Seed bank patterns show clear interspecific variation in response to climate across the distribution range. Not all seed banking species may be as well equipped to buffer climate change via their seed bank, notably in short-term persistent species. Since the buffering capacity of seed banks is key to species persistence, these results provide crucial information to advance climatic change predictions on range shifts, community and biodiversity responses.},
  language  = {en}
}
@article{HeinkenRohnerHoppert2007,
  author    = {Heinken, Thilo and Rohner, Maria-Sofie and Hoppert, Michael},
  title     = {Red wood ants (Formica rufa group) disperse bryophyte and lichen fragments on a local scale},
  issn      = {0078-2238},
  year      = {2007},
  abstract  = {Gametophyte and thallus fragments, respectively, may be an important or even the only mode of reproduction for many bryophytes and lichens species. Until now especially birds and mammals have been identifi ed as potential animal dispersal vectors of fragments. This study investigates the dispersal of bryophyte and lichen fragments by red wood ants which build large nest mounds from plant material and are abundant in European coniferous forests. We sampled nest material from 25 nest mounds in fi ve different pine and spruce forest types in Germany and found numerous fragments of 20 bryophyte and ten lichen species. As they occurred on almost all studied mounds and often in large numbers we conclude that collecting cryptogam fragments as nest material is a characteristic feature for the Formica rufa group in coniferous forests. Species number and composition of fragments on mounds coincided with the epigeic vegetation around ant nests to a large extent: Almost all collected species were present in the vegetation, and dominant fragment species occurred in large amounts in the vicinity of ant nests. Lichen fragments were larger than bryophyte fragments. Certain life forms (weft-forming bryophytes, reindeer lichens) were accumulated on mounds, while others (tall turfs, cup-type Cladonia species) were discriminated, refl ecting fragmentation features of species. Collected fragments may regenerate to mature plants if nest mounds are abandoned, and especially if they are lost during transport over several metres. We conclude that dispersal of fragments by red wood ants contributes to maintain epigeic bryophyte and lichen diversity of coniferous forests by supporting colonisation after disturbances, which occur on different spatial and temporal scales.},
  language  = {en}
}
@article{Heinken2008,
  author    = {Heinken, Thilo},
  title     = {Vegetation und Standort bodensaurer Buchenw{\"a}lder am Arealrand : am Beispiel Mittelbrandenburgs},
  issn      = {0018-0637},
  year      = {2008},
  abstract  = {Different from NW Germany, the northern part of NE Germany and the "Hohe Flaeming" region, central Brandenburg is considered as being largely devoid of natural beech forests because of its subcontinental, dry climate. In the present study the vegetation ecology of beech forests of the region is comprehensively documented for the first time, and they are compared with NW German stands in Lower Saxony. In the study area beech forests are concentrated in the Berlin-Potsdam region along the Havel river lakes which is characterised by relatively high precipitation and a specific land use history. All belong to the Luzulo-Fagetum growing on acid soils. Four subtypes are distinguished according to nutrient availability and soil moisture. The central Brandenburg Luzulo-Fagetum does not markedly deviate from other beech forests in the northern German lowlands with respect to vegetation structure and edaphic subtypes. However, numerous indicator species for humid or moist conditions are less frequent than under atlantic climate conditions in the lowlands of Lower Saxony, a pattern occurring also in other forest communities. On the other hand, nitrogen and disturbance indicators are more frequent in central Brandenburg. As expected, podzolisation of the soils and humus accumulation is lower in beech forests under subcontinental climate, but surprisingly the soils are more sandy and thus drier. However, beech forests are lacking on south-exposed slopes, and they are notably occurring in northern exposition. A combined analysis of distribution patterns and climatic data, postglacial vegetation history and forest use history, and actual rejuvenation dynamics reveals that the present-day beech forests in central Brandenburg have to be considered as near-natural relics, which are currently spreading. The range of potentially natural beech forests is larger than assumed until now, but further on it is not clearly to define.},
  language  = {de}
}
@article{FischerHeinkenMeyeretal.2009,
  author    = {Fischer, Petra and Heinken, Thilo and Meyer, Peter and Schmidt, Marcus and Waesch, Gunnar},
  title     = {Zur Abgrenzung und Situation des FFH-Lebensraumtyps "Mitteleurop{\"a}ische Flechten-Kiefernw{\"a}lder" (91TO) in Deutschland},
  issn      = {0028-0615},
  year      = {2009},
  abstract  = {Die in Deutschland gegenw{\"a}rtig durch N{\"a}hrstoffeintr{\"a}ge und ausbleibenden N{\"a}hrstoffentzug stark im R{\"u}ckgang begriffenen Flechten-Kiefernw{\"a}lder werden als Biotoptyp wie auch als Lebensraumtyp "Mitteleurop{\"a}ische Flechten-Kiefernw{\"a}lder" (Code 91T0) diskutiert. Die bisherige, sehr uneinheitliche Differenzierung von Flechten-Kiefernw{\"a}ldern auf der Ebene von Biotoptypen wird dargestellt. Auf der Grundlage neuerer vegetationskundlicher {\"u}bersichten werden Vorschl{\"a}ge f{\"u}r eine einheitliche Abgrenzung des Biotoptyps "Flechten-Kiefernwald" und des Lebensraumtyps 91T0 unterbreitet. Im nieders{\"a}chsischen Naturwaldreservat "Kaarßer Sandberge" (Niedersachsen) wurde die Anwendung des Konzeptes erfolgreich erprobt. Nicht nur hier, sondern auch deutschlandweit wird der R{\"u}ckgang der Erdflechten in den Kieferw{\"a}ldern zugunsten von Drahtschmiele und/ oder pleurokarpen Moosen deutlich. Nach der derzeitigen Definition des Lebensraumtyps 91T0 besteht auf der Grundlage der FFH-Richtlinie nicht f{\"u}r alle Flechten-Kiefernw{\"a}lder eine Chance der Verbesserung. Der Ausschluss von außerhalb des nat{\"u}rlichen Verbreitungsgebietes der Wald-Kiefer gelegenen sowie von durch Aufforstung angepflanzten Best{\"a}nden bringt Probleme mit sich, die diskutiert werden. F{\"u}r den Erhalt und die Wiederherstellung der gr{\"o}ßtenteils nutzungsbedingt entstandenen Flechten-Kiefernw{\"a}lder sind praktikable Pflegemaßnahmen notwendig, die im Rahmen von Streunutzungsversuchen erprobt werden m{\"u}ssen.},
  language  = {de}
}
@article{Heinken2009,
  author    = {Heinken, Thilo},
  title     = {Populationsbiologische und genetische Konsequenzen von Habitatfragmentierung bei Pflanzen : wissenschaftliche Grundlagen f{\"u}r Biotopverbundsysteme},
  issn      = {0722-494X},
  year      = {2009},
  abstract  = {Besides habitat loss, population-biological and genetic consequences of habitat fragmentation are thought to be a major threat to species since the 1990's and thus are now in the focus of plant species conservation. Using examples, this article gives an overview on the state of the art. It aims to evaluate the relevance habitat fragmentation and the resulting small size and isolation of populations may have for Central European plant populations. Stochasticity, edge effects, pollinator limitation, genetic drift and inbreeding depression are identified as important and very widespread negative effects. Together with changed habitat quality due to eutrophication, drainage or altered land use they negatively affect the fitness of individuals and populations, resulting in an increased risk of extinction. This negative effect of small populations on the fitness of individuals is called the Allee-effect, irrespective of the underlying causes, which can only be identified by scientific experiments. Metapopulation dynamics that are supported by a habitat network may prevent a permanent extinction of plant populations and minimize the negative genetic effects of habitat fragmentation by increasing gene flow via pollen and seeds. However, existing studies from Central Europe mainly concentrated on certain plant families (Gentianaceae, Primulaceae), habitats (species- rich grasslands), insect-pollinated and outcrossing species, and species mainly relying on sexual reproduction. On the other hand, few insights exist about grasses, ruderal plants and weeds, non-indigenous, wind- and self-pollinated species, and species mainly reproducing vegetatively or via apomictic seeds. However, according to the present state of knowledge especially these plant species, and those with a high dispersal potential, have to be considered as less sensitive to habitat fragmentation. Based on these findings, habitat types are classified with regard to their sensitivity to fragmentation, and ecological characters and species traits of sensitive and less sensitive species are compared. Finally, general consequences for conservation practice are presented with regard to target species and habitats for the formation of habitat networks, minimum viable population sizes, genetic rescue of populations, and deploying plants from ex-situ conservation to natural habitats.},
  language  = {de}
}
@article{Heinken2008,
  author    = {Heinken, Thilo},
  title     = {Welche populationsbiologischen und genetischen Konsequenzen hat Habitatfragmentierung f{\"u}r Pflanzen? : Wissenschaftliche Grundlagen f{\"u}r ein Biotopverbundsystem f{\"u}r Pflanzen in Brandenburg},
  issn      = {0942-9328},
  year      = {2008},
  abstract  = {Neben dem Habitatverlust gelten Konsequenzen der Habitatfragmentierung seit den 1990er Jahren als wesentliche Ursache der Gefaehrdung von Pflanzen und stehen damit nun auch im Fokus des botanischen Artenschutzes. Der vorliegende Beitrag gibt einen ueberblick ueber den Stand der populationsbiologischen und genetischen Forschung und versucht abzuschaetzen, welche Bedeutung Habitatfragmentierung und die dadurch entstehenden kleinen, isolierten Populationen auf heimische Pflanzenarten haben koennen. Als wesentliche und offenbar sehr weit verbreitete negative Effekte werden Zufallsereignisse, Randeffekte, Bestaeuberlimitierung, Gendrift und Inzuchtdepression identifiziert. Zusammen mit verringerter Habitatqualitaet durch Eutrophierung, Entwaesserung oder Nutzungsaenderung wirken sie zumeist negativ auf die Fitness der Individuen und Populationen und erhoehen so deren Aussterberisiko. Dieser negative Effekt kleiner Populationen auf die individuelle Fitness wird unabhaengig von der Ursache als Allee-Effekt bezeichnet. Eine durch einen Biotopverbund gefoerderte Metapopulationsdynamik kann das dauerhafte Aussterben von Pflanzenpopulationen verhindern und mindert die negativen genetischen Effekte der Habitatfragmentierung ueber einen erhoehten Genfluss durch Pollen und Samen. Die bisherigen wissenschaftlichen Studien in Mitteleuropa beruhen allerdings in ueberproportionaler Weise auf bestimmten Pflanzenfamilien (Gentianaceae, Primulaceae), Habitaten (Trocken- und Magerrasen, Wirtschaftsgruenland), insekten- und obligat fremdbestaeubten sowie weitgehend auf sexuelle Fortpflanzung angewiesenen Arten, waehrend etwa ueber Grasartige, Ruderalpflanzen, wind- und selbstbestaeubte sowie an vegetative Fortpflanzung angepasste Arten nur wenige Erkenntnisse vorliegen. Gerade diese und Pflanzenarten mit hohem Ausbreitungspotenzial muessen aber nach derzeitigem Wissensstand als weniger sensitiv gegenueber Habitatfragmentierung eingestuft werden. Auf diesen Befunden aufbauend werden fuer die Naturschutzpraxis Biotoptypen hinsichtlich ihrer Sensitivitaet gegenueber Habitatfragmentierung klassifiziert und ein auf biologischen Merkmalen basierender Kriterienkatalog zur Auswahl von Zielarten des Biotopverbunds vorgestellt. Schließlich wird eroertert, was bei Maßnahmen zur Regeneration kleiner bzw. bereits ausgestorbener Populationen zu beachten ist, und es werden allgemeine Folgerungen zur Ausgestaltung eines Biotopverbundskonzepts fuer Pflanzen gezogen.},
  language  = {de}
}
@article{SchwarzerHeinkenLuthardtetal.2013,
  author    = {Schwarzer, Christian and Heinken, Thilo and Luthardt, Vera and Joshi, Jasmin Radha},
  title     = {Latitudinal shifts in species interactions interfere with resistance of southern but not of northern bog-plant communities to experimental climate change},
  series = {The journal of ecology},
  volume    = {101},
  journal   = {The journal of ecology},
  number    = {6},
  publisher = {Wiley-Blackwell},
  address   = {Hoboken},
  issn      = {0022-0477},
  doi       = {10.1111/1365-2745.12158},
  pages     = {1484 -- 1497},
  year      = {2013},
  abstract  = {The persistence of species under changed climatic conditions depends on adaptations and plastic responses to these conditions and on interactions with their local plant community resulting in direct and indirect effects of changed climatic conditions. Populations at species' range margins may be especially crucial in containing a gene pool comprising adaptations to extreme climatic conditions. Many species of northern European bog ecosystems reach their southern lowland range limit in central Europe. In a common-garden experiment, we experimentally assessed the impact of projected climatic changes on five bog-plant species (including peat moss Sphagnum magellanicum) sampled along a latitudinal gradient of 1400km from Scandinavia to the marginal lowland populations in Germany. Populations were cultivated in monocultures and in experimental communities composed of all five species from their local community, and exposed to five combinations of three climate treatments (warming, fluctuating water-tables, fertilization) in a southern common garden. Whereas most monocultures showed a decreasing biomass production from southern to northern origins under southern environmental conditions, in the experimental mixed-species communities, an increasing biomass production towards northern communities was observed together with a shift in interspecific interactions along the latitudinal gradient. While negative dominance effects prevailed in southern communities, higher net biodiversity effects were observed in northern subarctic communities. The combined effects of climate treatments increased biomass production in monocultures of most origins. In communities, however, overall the treatments did not result in significantly changed biomass production. Among individual treatments, water-table fluctuations caused a significant decrease in biomass production, but only in southern communities, indicating higher vulnerability to changed climatic conditions. Here, negative effects of climate treatments on graminoids were not compensated by the slightly increased growth of peat moss that benefited from interspecific interactions only in northern communities.Synthesis. We conclude that shifting interactions within multispecies communities caused pronounced responses to changed climatic conditions in wetland communities of temperate southern marginal, but not of northern subarctic origin. Therefore, future models investigating the impacts of climate change on plant communities should consider geographical variation in species interactions an important factor influencing community responses to changed climatic conditions.},
  language  = {en}
}
@article{SchmidtFischerGuenzletal.2008,
  author    = {Schmidt, Marcus and Fischer, Petra and G{\"u}nzl, Bettina and Heinken, Thilo and Kelm, Hans-J{\"u}rgen and Meyer, Peter and Pr{\"u}ter, Johannes and Waesch, Gunnar},
  title     = {Flechten-Kiefernw{\"a}lder : Artenvielfalt durch alte Nutzungsformen?},
  issn      = {1430-2713},
  year      = {2008},
  language  = {de}
}
@article{DeFrenneRodriguezSanchezCoomesetal.2013,
  author    = {De Frenne, Pieter and Rodriguez-Sanchez, Francisco and Coomes, David Anthony and B{\"a}ten, Lander and Verstr{\"a}ten, Gorik and Vellend, Mark and Bernhardt-R{\"o}mermann, Markus and Brown, Carissa D. and Brunet, J{\"o}rg and Cornelis, Johnny and Decocq, Guillaume M. and Dierschke, Hartmut and Eriksson, Ove and Gilliam, Frank S. and Hedl, Radim and Heinken, Thilo and Hermy, Martin and Hommel, Patrick and Jenkins, Michael A. and Kelly, Daniel L. and Kirby, Keith J. and Mitchell, Fraser J. G. and Naaf, Tobias and Newman, Miles and Peterken, George and Petrik, Petr and Schultz, Jan and Sonnier, Gregory and Van Calster, Hans and Waller, Donald M. and Walther, Gian-Reto and White, Peter S. and Woods, Kerry D. and Wulf, Monika and Graae, Bente Jessen and Verheyen, Kris},
  title     = {Microclimate moderates plant responses to macroclimate warming},
  series = {Proceedings of the National Academy of Sciences of the United States of America},
  volume    = {110},
  journal   = {Proceedings of the National Academy of Sciences of the United States of America},
  number    = {46},
  publisher = {National Acad. of Sciences},
  address   = {Washington},
  issn      = {0027-8424},
  doi       = {10.1073/pnas.1311190110},
  pages     = {18561 -- 18565},
  year      = {2013},
  abstract  = {Recent global warming is acting across marine, freshwater, and terrestrial ecosystems to favor species adapted to warmer conditions and/or reduce the abundance of cold-adapted organisms (i.e., "thermophilization" of communities). Lack of community responses to increased temperature, however, has also been reported for several taxa and regions, suggesting that "climatic lags" may be frequent. Here we show that microclimatic effects brought about by forest canopy closure can buffer biotic responses to macroclimate warming, thus explaining an apparent climatic lag. Using data from 1,409 vegetation plots in European and North American temperate forests, each surveyed at least twice over an interval of 12-67 y, we document significant thermophilization of ground-layer plant communities. These changes reflect concurrent declines in species adapted to cooler conditions and increases in species adapted to warmer conditions. However, thermophilization, particularly the increase of warm-adapted species, is attenuated in forests whose canopies have become denser, probably reflecting cooler growing-season ground temperatures via increased shading. As standing stocks of trees have increased in many temperate forests in recent decades, local microclimatic effects may commonly be moderating the impacts of macroclimate warming on forest understories. Conversely, increases in harvesting woody biomass-e.g., for bioenergy-may open forest canopies and accelerate thermophilization of temperate forest biodiversity.},
  language  = {en}
}
@article{deFrenneKolbVerheyenetal.2009,
  author    = {de Frenne, Pieter and Kolb, Annette and Verheyen, Kris and Brunet, Johanne and Chabrerie, Olivier and Decocq, Guillaume and Diekmann, Martin and Eriksson, Ove and Heinken, Thilo and Hermy, Martin and J{\~o}gar, {\"U}lle and Stanton, Sara and Quataert, Paul and Zindel, Renate and Zobel, Martin and Graae, Bente Jessen},
  title     = {Unravelling the effects of temperature, latitude and local environment on the reproduction of forest herbs},
  issn      = {1466-822X},
  doi       = {10.1111/j.1466-8238.2009.00487.x},
  year      = {2009},
  abstract  = {Aim To investigate the effect of temperature, latitude and local environment on the reproductive traits of widespread perennial forest herbs to better understand the potential impacts of rising temperatures on their population dynamics and colonization capacities. Location Six regions along a latitudinal gradient from France to Sweden. Methods Within each region, we collected data from three to five populations of up to six species. For each species, several variables were recorded in each region (temperature, latitude) and population (local abiotic and biotic environmental variables), and seed production and germination were estimated. Resource investment in reproduction (RIR) was quantified as seed number ¥ seed mass, while germinable seed output (GSO) was expressed as seed number ¥ germination percentage.We performed linear regression and mixed effect models to investigate the effects of temperature (growing degree hours), latitude and local abiotic and biotic environment on RIR and GSO. Results Temperature and latitude explained most of the variation in RIR and GSO for early flowering species with a northerly distribution range edge (Anemone nemorosa, Paris quadrifolia and Oxalis acetosella). Reproduction of the more southerly distributed species (Brachypodium sylvaticum, Circaea lutetiana and Primula elatior), in contrast, was independent of temperature/latitude. In the late summer species, B. sylvaticum and C. lutetiana, variation in RIR and GSO was best explained by local environmental variables, while none of the investigated variables appeared to be related to reproduction in P. elatior. Main conclusions We showed that reproduction of only two early flowering, northerly distributed species was related to temperature. This suggests that the potential reproductive response of forest herbs to climate warming partly depends on their phenology and distribution, but also that the response is to some extent species dependent. These findings should be taken into account when predictions about future shifts in distribution range are made.},
  language  = {en}
}