@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} } @article{GraaeVerheyenKolbetal.2009, author = {Graae, Bente Jessen and Verheyen, Kris and Kolb, Annette and van der Veken, Sebastian and Heinken, Thilo and Chabrerie, Olivier and Diekmann, Martin and Valtinat, Karin and Zindel, Renate and Karlsson, Elisabeth and Str{\"o}m, Lotta and Decocq, Guillaume and Hermy, Martin and Baskin, Carol C.}, title = {Germination requirements and seed mass of slow- and fast-colonizing temperate forest herbs along a latitudinal gradient}, issn = {1195-6860}, doi = {10.2980/16-2-3234}, year = {2009}, abstract = {Predictions on displacement of suitable habitats due to climate change suggest that plant species with poor colonization ability may be unable to move fast enough to match forecasted climate-induced changes in habitat distribution. However, studies on early Holocene plant migration show fast migration of many plant species that are poor colonizers today. We hypothesize that warmer temperatures during the early Holocene yielded higher seed quality, contributing to explaining the fast migration. We studied how the 3 seed quality variables, seed mass, germinability, and requirements for break of seed dormancy, vary for seeds of 11 forest herb species with varying colonization capacity collected along a 1400-km latitudinal gradient. Within species, seed mass showed a positive correlation with latitude, whereas germinability was more positively correlated with temperature (growing degree hours obtained at time of seed collection). Only slow-colonizing species increased germinability with temperature, whereas only fast-colonizing species increased germinability with latitude. These interactions were only detectable when analyzing germinability of the seeds, even though this trait and seed mass were correlated. The requirement for dormancy break did not correlate with latitude or temperature. The results indicate that seed development of slow colonizers may be favoured by a warmer climate, which in turn may be important for their migration capacity.}, 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{DeFrenneKolbGraaeetal.2011, author = {De Frenne, P. and Kolb, Annette and Graae, Benete Jessen and Decocq, Guillaume and Baltora, S. and De Schrijver, A. and Brunet, J. and Chabrerie, Olivier and Cousins, Sara A. O. and Dhondt, Rob and Diekmann, Martin and Gruwez, R. and Heinken, Thilo and Hermy, Martin and Liira, J. and Saguez, R. and Shevtsova, Anna and Baskin, Carol C. and Verheyen, Kris}, title = {A latitudinal gradient in seed nutrients of the forest herb Anemone nemorosa}, series = {Plant biology}, volume = {13}, journal = {Plant biology}, number = {3}, publisher = {Wiley-Blackwell}, address = {Malden}, issn = {1435-8603}, doi = {10.1111/j.1438-8677.2010.00404.x}, pages = {493 -- 501}, year = {2011}, abstract = {The nutrient concentration in seeds determines many aspects of potential success of the sexual reproductive phase of plants, including the seed predation probability, efficiency of seed dispersal and seedling performance. Despite considerable research interest in latitudinal gradients of foliar nutrients, a similar gradient for seeds remains unexplored. We investigated a potential latitudinal gradient in seed nutrient concentrations within the widespread European understorey forest herb Anemone nemorosa L. We sampled seeds of A. nemorosa in 15 populations along a 1900-km long latitudinal gradient at three to seven seed collection dates post-anthesis and investigated the relative effects of growing degree-hours > 5 degrees C, soil characteristics and latitude on seed nutrient concentrations. Seed nitrogen, nitrogen:phosphorus ratio and calcium concentration decreased towards northern latitudes, while carbon:nitrogen ratios increased. When taking differences in growing degree-hours and measured soil characteristics into account and only considering the most mature seeds, the latitudinal decline remained particularly significant for seed nitrogen concentration. We argue that the decline in seed nitrogen concentration can be attributed to northward decreasing seed provisioning due to lower soil nitrogen availability or greater investment in clonal reproduction. This pattern may have large implications for the reproductive performance of this forest herb as the degree of seed provisioning ultimately co-determines seedling survival and reproductive success.}, language = {en} } @article{DeFrenneGraaeKolbetal.2011, author = {De Frenne, Pieter and Graae, Bente J. and Kolb, Annette and Shevtsova, Anna and Baeten, Lander and Brunet, J{\"o}rg and Chabrerie, Olivier and Cousins, Sara A. O. and Decocq, Guillaume and Dhondt, Rob and Diekmann, Martin and Gruwez, Robert and Heinken, Thilo and Hermy, Martin and Oster, Mathias and Saguez, Robert and Stanton, Sharon and Tack, Wesley and Vanhellemont, Margot and Verheyen, Kris}, title = {An intraspecific application of the leaf-height-seed ecology strategy scheme to forest herbs along a latitudinal gradient}, series = {Ecography : pattern and diversity in ecology ; research papers forum}, volume = {34}, journal = {Ecography : pattern and diversity in ecology ; research papers forum}, number = {1}, publisher = {Wiley-Blackwell}, address = {Malden}, issn = {0906-7590}, doi = {10.1111/j.1600-0587.2010.06399.x}, pages = {132 -- 140}, year = {2011}, abstract = {We measured LHS traits in 41 Anemone nemorosa and 44 Milium effusum populations along a 1900-2300 km latitudinal gradient from N France to N Sweden. We then applied multilevel models to identify the effects of regional (temperature, latitude) and local (soil fertility and acidity, overstorey canopy cover) environmental factors on LHS traits. Both species displayed a significant 4\% increase in plant height with every degree northward shift (almost a two-fold plant height difference between the southernmost and northernmost populations). Neither seed mass nor SLA showed a significant latitudinal cline. Temperature had a large effect on the three LHS traits of Anemone. Latitude, canopy cover and soil nutrients were related to the SLA and plant height of Milium. None of the investigated variables appeared to be related to the seed mass of Milium. The variation in LHS traits indicates that the ecological strategy determined by the position of each population in this three-factor triangle is not constant along the latitudinal gradient. The significant increase in plant height suggests greater competitive abilities for both species in the northernmost populations. We also found that the studied environmental factors affected the LHS traits of the two species on various scales: spring-flowering Anemone was affected more by temperature, whereas early-summer flowering Milium was affected more by local and other latitude-related factors. Finally, previously reported cross-species correlations between LHS traits and latitude were generally unsupported by our within-species approach.}, language = {en} } @article{DeFrenneBaetenGraaeetal.2011, author = {De Frenne, Pieter and Baeten, Lander and Graae, Bente J. and Brunet, Jorg and Wulf, Monika and Orczewska, Anna and Kolb, Annette and Jansen, Ivy and Jamoneau, Aurelien and Jacquemyn, Hans and Hermy, Martin and Diekmann, Martin and De Schrijver, An and De Sanctis, Michele and Decocq, Guillaume and Cousins, Sara A. O. and Verheyen, Kris}, title = {Interregional variation in the floristic recovery of post-agricultural forests}, series = {The journal of ecology}, volume = {99}, journal = {The journal of ecology}, number = {2}, publisher = {Wiley-Blackwell}, address = {Malden}, issn = {0022-0477}, doi = {10.1111/j.1365-2745.2010.01768.x}, pages = {600 -- 609}, year = {2011}, abstract = {1. Worldwide, the floristic composition of temperate forests bears the imprint of past land use for decades to centuries as forests regrow on agricultural land. Many species, however, display significant interregional variation in their ability to (re)colonize post-agricultural forests. This variation in colonization across regions and the underlying factors remain largely unexplored. 2. We compiled data on 90 species and 812 species x study combinations from 18 studies across Europe that determined species' distribution patterns in ancient (i.e. continuously forested since the first available land use maps) and post-agricultural forests. The recovery rate (RR) of species in each landscape was quantified as the log-response ratio of the percentage occurrence in post-agricultural over ancient forest and related to the species-specific life-history traits and local (soil characteristics and light availability) and regional factors (landscape properties as habitat availability, time available for colonization, and climate). 3. For the herb species, we demonstrate a strong (interactive) effect of species' life-history traits and forest habitat availability on the RR of post-agricultural forest. In graminoids, however, none of the investigated variables were significantly related to the RR. 4. The better colonizing species that mainly belonged to the short-lived herbs group showed the largest interregional variability. Their recovery significantly increased with the amount of forest habitat within the landscape, whereas, surprisingly, the time available for colonization, climate, soil characteristics and light availability had no effect. 5. Synthesis. By analysing 18 independent studies across Europe, we clearly showed for the first time on a continental scale that the recovery of short-lived forest herbs increased with the forest habitat availability in the landscape. Small perennial forest herbs, however, were generally unsuccessful in colonizing post-agricultural forest even in relatively densely forested landscapes. Hence, our results stress the need to avoid ancient forest clearance to preserve the typical woodland flora.}, language = {en} } @article{DeFrenneBrunetShevtsovaetal.2011, author = {De Frenne, Pieter and Brunet, Jorg and Shevtsova, Anna and Kolb, Annette and Graae, Bente J. and Chabrerie, Olivier and Cousins, Sara Ao and Decocq, Guillaume and De Schrijver, An and Diekmann, Martin and Gruwez, Robert and Heinken, Thilo and Hermy, Martin and Nilsson, Christer and Stanton, Sharon and Tack, Wesley and Willaert, Justin and Verheyen, Kris}, title = {Temperature effects on forest herbs assessed by warming and transplant experiments along a latitudinal gradient}, series = {Global change biology}, volume = {17}, journal = {Global change biology}, number = {10}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {1354-1013}, doi = {10.1111/j.1365-2486.2011.02449.x}, pages = {3240 -- 3253}, year = {2011}, abstract = {Slow-colonizing forest understorey plants are probably not able to rapidly adjust their distribution range following large-scale climate change. Therefore, the acclimation potential to climate change within their actual occupied habitats will likely be key for their short-and long-term persistence. We combined transplant experiments along a latitudinal gradient with open-top chambers to assess the effects of temperature on phenology, growth and reproductive performance of multiple populations of slow-colonizing understorey plants, using the spring flowering geophytic forb Anemone nemorosa and the early summer flowering grass Milium effusum as study species. In both species, emergence time and start of flowering clearly advanced with increasing temperatures. Vegetative growth (plant height, aboveground biomass) and reproductive success (seed mass, seed germination and germinable seed output) of A. nemorosa benefited from higher temperatures. Climate warming may thus increase future competitive ability and colonization rates of this species. Apart from the effects on phenology, growth and reproductive performance of M. effusum generally decreased when transplanted southwards (e. g., plant size and number of individuals decreased towards the south) and was probably more limited by light availability in the south. Specific leaf area of both species increased when transplanted southwards, but decreased with open-top chamber installation in A. nemorosa. In general, individuals of both species transplanted at the home site performed best, suggesting local adaptation. We conclude that contrasting understorey plants may display divergent plasticity in response to changing temperatures which may alter future understorey community dynamics.}, language = {en} } @article{VerheyenBaetenDeFrenneetal.2012, author = {Verheyen, Kris and Baeten, Lander and De Frenne, Pieter and Bernhardt-R{\"o}mermann, Markus and Brunet, Jorg and Cornelis, Johnny and Decocq, Guillaume and Dierschke, Hartmut and Eriksson, Ove and Hedl, Radim and Heinken, Thilo and Hermy, Martin and Hommel, Patrick and Kirby, Keith J. and Naaf, Tobias and Peterken, George and Petrik, Petr and Pfadenhauer, Joerg and Van Calster, Hans and Walther, Gian-Reto and Wulf, Monika and Verstraeten, Gorik}, title = {Driving factors behind the eutrophication signal in understorey plant communities of deciduous temperate forests}, series = {The journal of ecology}, volume = {100}, journal = {The journal of ecology}, number = {2}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {0022-0477}, doi = {10.1111/j.1365-2745.2011.01928.x}, pages = {352 -- 365}, year = {2012}, abstract = {1. Atmospheric nitrogen (N) deposition is expected to change forest understorey plant community composition and diversity, but results of experimental addition studies and observational studies are not yet conclusive. A shortcoming of observational studies, which are generally based on resurveys or sampling along large deposition gradients, is the occurrence of temporal or spatial confounding factors. 2. We were able to assess the contribution of N deposition versus other ecological drivers on forest understorey plant communities by combining a temporal and spatial approach. Data from 1205 (semi-)permanent vegetation plots taken from 23 rigorously selected understorey resurvey studies along a large deposition gradient across deciduous temperate forest in Europe were compiled and related to various local and regional driving factors, including the rate of atmospheric N deposition, the change in large herbivore densities and the change in canopy cover and composition. 3. Although no directional change in species richness occurred, there was considerable floristic turnover in the understorey plant community and a shift in species composition towards more shade-tolerant and nutrient-demanding species. However, atmospheric N deposition was not important in explaining the observed eutrophication signal. This signal seemed mainly related to a shift towards a denser canopy cover and a changed canopy species composition with a higher share of species with more easily decomposed litter. 4. Synthesis. Our multi-site approach clearly demonstrates that one should be cautious when drawing conclusions about the impact of atmospheric N deposition based on the interpretation of plant community shifts in single sites or regions due to other, concurrent, ecological changes. Even though the effects of chronically increased N deposition on the forest plant communities are apparently obscured by the effects of canopy changes, the accumulated N might still have a significant impact. However, more research is needed to assess whether this N time bomb will indeed explode when canopies will open up again.}, language = {en} } @article{DeFrenneGraaeBrunetetal.2012, author = {De Frenne, Pieter and Graae, Bente J. and Brunet, J{\"o}rg and Shevtsova, Anna and De Schrijver, An and Chabrerie, Olivier and Cousins, Sara A. O. and Decocq, Guillaume and Diekmann, Martin and Hermy, Martin and Heinken, Thilo and Kolb, Annette and Nilsson, Christer and Stanton, Sharon and Verheyen, Kris}, title = {The response of forest plant regeneration to temperature variation along a latitudinal gradient}, series = {Annals of botany}, volume = {109}, journal = {Annals of botany}, number = {5}, publisher = {Oxford Univ. Press}, address = {Oxford}, issn = {0305-7364}, doi = {10.1093/aob/mcs015}, pages = {1037 -- 1046}, year = {2012}, abstract = {The response of forest herb regeneration from seed to temperature variations across latitudes was experimentally assessed in order to forecast the likely response of understorey community dynamics to climate warming. Seeds of two characteristic forest plants (Anemone nemorosa and Milium effusum) were collected in natural populations along a latitudinal gradient from northern France to northern Sweden and exposed to three temperature regimes in growth chambers (first experiment). To test the importance of local adaptation, reciprocal transplants were also made of adult individuals that originated from the same populations in three common gardens located in southern, central and northern sites along the same gradient, and the resulting seeds were germinated (second experiment). Seedling establishment was quantified by measuring the timing and percentage of seedling emergence, and seedling biomass in both experiments. Spring warming increased emergence rates and seedling growth in the early-flowering forb A. nemorosa. Seedlings of the summer-flowering grass M. effusum originating from northern populations responded more strongly in terms of biomass growth to temperature than southern populations. The above-ground biomass of the seedlings of both species decreased with increasing latitude of origin, irrespective of whether seeds were collected from natural populations or from the common gardens. The emergence percentage decreased with increasing home-away distance in seeds from the transplant experiment, suggesting that the maternal plants were locally adapted. Decreasing seedling emergence and growth were found from the centre to the northern edge of the distribution range for both species. Stronger responses to temperature variation in seedling growth of the grass M. effusum in the north may offer a way to cope with environmental change. The results further suggest that climate warming might differentially affect seedling establishment of understorey plants across their distribution range and thus alter future understorey plant dynamics.}, language = {en} } @article{GraaeDeFrenneKolbetal.2012, author = {Graae, Bente J. and De Frenne, Pieter and Kolb, Annette and Brunet, Jorg and Chabrerie, Olivier and Verheyen, Kris and Pepin, Nick and Heinken, Thilo and Zobel, Martin and Shevtsova, Anna and Nijs, Ivan and Milbau, Ann}, title = {On the use of weather data in ecological studies along altitudinal and latitudinal gradients}, series = {Oikos}, volume = {121}, journal = {Oikos}, number = {1}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {0030-1299}, doi = {10.1111/j.1600-0706.2011.19694.x}, pages = {3 -- 19}, year = {2012}, abstract = {Global warming has created a need for studies along climatic gradients to assess the effects of temperature on ecological processes. Altitudinal and latitudinal gradients are often used as such, usually in combination with air temperature data from the closest weather station recorded at 1.52 m above the ground. However, many ecological processes occur in, at, or right above the soil surface. To evaluate how representative the commonly used weather station data are for the microclimate relevant for soil surface biota, we compared weather station temperatures for an altitudinal (500900 m a.s.l.) and a latitudinal gradient (4968 degrees N) with data obtained by temperature sensors placed right below the soil surface at five sites along these gradients. The mean annual temperatures obtained from weather stations and adjusted using a lapse rate of -5.5 degrees C km-1 were between 3.8 degrees C lower and 1.6 degrees C higher than those recorded by the temperature sensors at the soil surface, depending on the position along the gradients. The monthly mean temperatures were up to 10 degrees C warmer or 5 degrees C colder at the soil surface. The within-site variation in accumulated temperature was as high as would be expected from a 300 m change in altitude or from a 4 degrees change in latitude or a climate change scenario corresponding to warming of 1.63.8 degrees C. Thus, these differences introduced by the decoupling are significant from a climate change perspective, and the results demonstrate the need for incorporating microclimatic variation when conducting studies along altitudinal or latitudinal gradients. We emphasize the need for using relevant temperature data in climate impact studies and further call for more studies describing the soil surface microclimate, which is crucial for much of the biota.}, language = {en} }