Jan den Ouden

Nurse plants, tree saplings and grazing pressure: changes in facilitation along a biotic environmental gradient

Current conceptual models predict that an increase in stress shifts interactions between plants from competitive to facilitative; hence, facilitation is expected to gain in ecological importance with increasing stress. Little is known about how facilitative interactions between plants change with increasing biotic stress, such as that incurred by consumer pressure or herbivory (i.e. disturbance sensu Grime). In grazed ecosystems, the presence of unpalatable plants is reported to protect tree saplings against cattle grazing and enhance tree establishment. In accordance with current conceptual facilitation-stress models, we hypothesised a positive relationship between facilitation and grazing pressure. We tested this hypothesis in a field experiment in which tree saplings of four different species (deciduous Fagus sylvatica, Acer pseudoplatanus and coniferous Abies alba, Picea abies) were planted either inside or outside of the canopy of the spiny nurse shrub Rosa rubiginosa in enclosures differing in grazing pressure (low and high) and in exclosures. During one grazing season we followed the survival of the different tree saplings and the level of browsing on these; we also estimated browsing damage to the nurse shrubs. Shrub damage was highest at the higher grazing pressure. Correspondingly, browsing increased and survival decreased in saplings located inside the canopy of the shrubs at the high grazing pressure compared to the low grazing pressure. Saplings of both deciduous species showed a higher survival than the evergreens, while sapling browsing did not differ between species. The relative facilitation of sapling browsing and sapling survival – i.e. the difference between saplings inside and outside the shrub canopy – decreased at high grazing pressure as the facilitative species became less protective. Interestingly, these findings do not agree with current conceptual facilitation-stress models predicting increasing facilitation with abiotic stress. We used our results to design a conceptual model of facilitation along a biotic environmental gradient. Empirical studies are needed to test the applicability of this model. In conclusion, we suggest that current conceptual facilitation models should at least consider the possibility of decreasing facilitation at high levels of stress.

Facilitation of Quercus ilex recruitment by shrubs in Mediterranean open woodlands

Abstract Question: Insufficient tree regeneration threatens the long-term persistence of biodiverse Mediterranean open oak woodlands. Could shrubs, scarce due to decades of management (clearing and ploughing), facilitate holm oak recruitment at both acorn and seedling stages? Location: Open oak woodlands in Central Spain. Methods: Plots with four acorns were planted: (1) under the canopy of the spiny shrub Genista hirsuta, (2) in a small cage, protecting against ungulates, (3) in a shaded cage, protecting against ungulates and sun, and (4) in open grassland. Sets of these four treatments were spatially grouped according to a randomised block design, with 16 blocks near (< 10 m) and 16 away from (> 20 m) parent trees to test for distance-related survival. Plots were regularly checked for seed removal. After emergence one seedling per plot (97 in total) was selected and its survival monitored. Results: Three months after sowing, 199 of 512 acorns were removed, predominantly by rodents. Acorn removal occurred at each treatment but was highest under shrubs. Eight months after sowing, seedling survival was highest under shrubs (50%), followed by shaded cages (16%), open grassland (4%) and cages (0%). Main mortality cause was drought (90%), killing most seedlings between June and July. No seedlings died from ungulate browsing. Conclusion: Shrubs demonstrated clear net facilitative effects for Quercus ilex recruitment, despite higher seed removal. Shading appears the crucial factor facilitating seedling survival. We therefore propose that lack of shrubs contributes largely to tree recruitment failure in Mediterranean open woodlands; management should aim at conserving shrubs.

Effects of tree species composition on within-forest distribution of understorey species

Abstract Question: Do tree species, with different litter qualities, affect the within-forest distribution of forest understorey species on intermediate to base-rich soils? Since habitat loss and fragmentation have caused ancient forest species to decline, those species are the main focus of this study. Location: Three ancient forests, along a soil gradient from acidification-sensitive to base-rich, were studied: Limbrichterbosch and Savelsbos in The Netherlands and Holtkrat in Denmark. Methods: Canopy and soil surveys along transects generated data for Redundancy Analysis on tree - humus relationships. We analysed the distribution of forest plant species with Canonical Correspondence Analysis. The explanatory factors were soil characteristics (pH, organic matter, loam content and thickness of the humus layers), external crown projection, groundwater and canopy data. We further analysed the relationship between forest species and humus characteristics with Spearman correlations. Results: Tree species have a significant impact on humus characteristics through the nature of their litter. Humus characteristics significantly explain the distribution of forest understorey species. The pH of the first 25 cm mineral soil and the thickness of the F- (fermentation) layer are the primary factors affecting the distribution of ancient forest species. Conclusion: This study indicates that the species composition of the forest canopy affects the distribution of forest understorey species. Ancient forest species are more abundant and frequent underneath trees with base-rich litter. On acidification-sensitive soils these relationships were stronger than on more base-rich, loamy soils. Abbreviations: Ah = Soil horizon consisting of mineral soil with a high organic matter content; CEC = Cation exchange capacity; F = Fermenting litter (layer); Ah = Humus-rich mineral soil (layer); L = Litter (layer); RDA = Redundancy analysis.

Controls on Coarse Wood Decay in Temperate Tree Species: Birth of the LOGLIFE Experiment

Dead wood provides a huge terrestrial carbon stock and a habitat to wide-ranging organisms during its decay. Our brief review highlights that, in order to understand environmental change impacts on these functions, we need to quantify the contributions of different interacting biotic and abiotic drivers to wood decomposition. LOGLIFE is a new long-term ‘common-garden’ experiment to disentangle the effects of species’ wood traits and site-related environmental drivers on wood decomposition dynamics and its associated diversity of microbial and invertebrate communities. This experiment is firmly rooted in pioneering experiments under the directorship of Terry Callaghan at Abisko Research Station, Sweden. LOGLIFE features two contrasting forest sites in the Netherlands, each hosting a similar set of coarse logs and branches of 10 tree species. LOGLIFE welcomes other researchers to test further questions concerning coarse wood decay that will also help to optimise forest management in view of carbon sequestration and biodiversity conservation.

Improved recruitment and early growth of Scots pine (Pinus sylvestris L.) seedlings after fire and soil scarification

The success of seedling recruitment of Scots pine (Pinus sylvestris L.) is strongly dependent on soil surface properties, such as humus depth and moisture content. In an undisturbed forest floor, seedlings are seldom able to become established due to the high incidence of desiccation in the organic soil layer. Methods that remove the organic soil layer are often necessary to improve the availability for radicles to reach the more stable moisture regime in the mineral soil. In this study we investigated pine-seedling establishment after mechanical soil scarification, burning of litter (OL) and burning of litter and humus (OL and OFH) in two mature pine stands in Germany. The herbaceous layer of the first stand was dominated by grasses (Molinia caerulea L. and Deschampsia flexuosa L.), whereas the herbaceous layer of the second stand was dominated by blueberry (Vaccinium myrtillus L.). Pine seeds were placed in experimental plots, and seedling numbers and heights were recorded at regular intervals. All treatments that removed organic soil resulted in higher seedling counts than did the undisturbed forest floor. The highest seedling counts were found on scarified and severely burnt plots, whereas seedling counts were lower on lightly burnt plots. Seedlings were significantly taller on burnt plots. This study shows that pine regeneration is stimulated by fire, not only in boreal forests, but also under central European conditions. With the expectation of higher fire frequency in the near future due to climatic changes, natural regeneration and succession on burnt sites should receive more focus in forest management and research.

Flood-Ring Formation and Root Development in Response to Experimental Flooding of Young Quercus robur Trees

Spring flooding in riparian forests can cause significant reductions in earlywood-vessel size in submerged stem parts of ring-porous tree species, leading to the presence of ‘flood rings’ that can be used as a proxy to reconstruct past flooding events, potentially over millennia. The mechanism of flood-ring formation and the relation with timing and duration of flooding are still to be elucidated. In this study, we experimentally flooded 4-year-old Quercus robur trees at three spring phenophases (late bud dormancy, budswell, and internode expansion) and over different flooding durations (2, 4, and 6 weeks) to a stem height of 50 cm. The effect of flooding on root and vessel development was assessed immediately after the flooding treatment and at the end of the growing season. Ring width and earlywood-vessel size and density were measured at 25- and 75-cm stem height and collapsed vessels were recorded. Stem flooding inhibited earlywood-vessel development in flooded stem parts. In addition, flooding upon budswell and internode expansion led to collapsed earlywood vessels below the water level. At the end of the growing season, mean earlywood-vessel size in the flooded stem parts (upon budswell and internode expansion) was always reduced by approximately 50% compared to non-flooded stem parts and 55% compared to control trees. This reduction was already present 2 weeks after flooding and occurred independent of flooding duration. Stem and root flooding were associated with significant root dieback after 4 and 6 weeks and mean radial growth was always reduced with increasing flooding duration. By comparing stem and root flooding, we conclude that flood rings only occur after stem flooding. As earlywood-vessel development was hampered during flooding, a considerable number of narrow earlywood vessels present later in the season, must have been formed after the actual flooding events. Our study indicates that root dieback, together with strongly reduced hydraulic conductivity due to anomalously narrow earlywood vessels in flooded stem parts, contribute to reduced radial growth after flooding events. Our findings support the value of flood rings to reconstruct spring flooding events that occurred prior to instrumental flood records.

Early wound reactions of Japanese maple during winter dormancy: the effect of two contrasting temperature regimes

During winter dormancy, temperate trees are capable of only a restricted response to wounding. In an experiment, we investigated the effect of wounding on Acer palmatum trees during winter-bud dormancy and found that in the cold (4 °C) temperature treatment, wound reactions were virtually absent. In the warm (15 °C) treatment, however, trees reacted actively to wounding within a three-week period by e.g. forming callus and local wound xylem. We conclude that temperature is an important factor in wound reactions during winter dormancy and may even induce the formation of callus and wound xylem within a three-week period.

EuMIXFOR empirical forest mensuration and ring width data from pure and mixed stands of Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) through Europe

Key messageThis data set provides unique empirical data from triplets of Scots pine (Pinus sylvestrisL.) and European beech (Fagus sylvaticaL.) across Europe. Dendrometric variables are provided for 32 triplets, 96 plots, 7555 treesand 4695 core samples. These data contribute to our understanding of mixed stand dynamics.Dataset access athttp://dx.doi.org/10.5061/dryad.8v04m. Associated metadata available athttps://metadata-afs.nancy.inra.fr/geonetwork/apps/georchestra/?uuid=b3e098ca-e681-4910-9099-0e25d3b4cd52&hl=eng.