Germination responses to light of four Neotropical forest tree species along an elevational gradient in the southern Central Andes

Seed germination is a key part of plants' life cycle and is mostly affected by the genetic background, the environmental conditions experienced by the mother plant and the seedbed conditions. The germination response to light is essential to optimize germination and seedling establishment in space and time. In addition, the germination response to light is a trait often related to the response of the seeds to their position in the soil (uncovered/buried). Here, we studied the germination response to light of four key tree species of the Yungas forest (Anadenanthera colubrina, Enterolobium contortisiliquum, Jacaranda mimosifolia and Handroanthus impetiginosus) sampled along an elevational and environmental gradient with contrasting vegetation cover and disturbance. Relative light germination (RLG) and mean germination time (MGT) were determined. Final germination was tested under cycles of light (8 h) and darkness (16 h) versus complete darkness (24 h) and elevation, and MGT was tested as a function of elevation of the provenance. The RLG increased from smaller to larger-seeded species. The MGT of three of the studied species was affected by the elevation of the provenance. Complete darkness negatively affected final germination, while two species exhibited a significant interaction between the provenance and light. The variable germination responses to light along the elevational gradient highlights the influence of the environment on germination as a key factor that should be considered for forest management, conservation and restoration projects

Individualistic responses of forest herb traits to environmental change

Intraspecific trait variation (ITV; i.e. variability in mean and/or distribution of plant attribute values within species) can occur in response to multiple drivers. Environmental change and land-use legacies could directly alter trait values within species but could also affect them indirectly through changes in vegetation cover. Increasing variability in environmental conditions could lead to more ITV, but responses might differ among species. Disentangling these drivers on ITV is necessary to accurately predict plant community responses to global change. We planted herb communities into forest soils with and without a recent history of agriculture. Soils were collected across temperate European regions, while the 15 selected herb species had different colonizing abilities and affinities to forest habitat. These mesocosms (384) were exposed to two-level full-factorial treatments of warming, nitrogen addition and illumination. We measured plant height and specific leaf area (SLA). For the majority of species, mean plant height increased as vegetation cover increased in response to light addition, warming and agricultural legacy. The coefficient of variation (CV) for height was larger in fast-colonizing species. Mean SLA for vernal species increased with warming, while light addition generally decreased mean SLA for shade-tolerant species. Interactions between treatments were not important predictors. Environmental change treatments influenced ITV, either via increasing vegetation cover or by affecting trait values directly. Species' ITV was individualistic, i.e. species responded to different single resource and condition manipulations that benefited their growth in the short term. These individual responses could be important for altered community organization after a prolonged period

Former land use affects the nitrogen and phosphorus concentrations and biomass of forest herbs

The colonization rates of understorey plants into forests growing on former agricultural land differ remarkably among species. Different dispersal and recruitment largely account for the contrasting colonization rates, but different effects of the soil legacies of former agricultural land use on plant performance may also play a role. Seven herbaceous forest species were sampled in paired post-agricultural and ancient forest stands to study whether land-use history has an effect on the aboveground nutrient concentrations (N, P and N:P ratios) and biomass of forest herbs and, if so, whether slow and fast colonizing species respond differently. Results showed that P concentrations were significantly affected by former land use with higher concentrations in the post-agricultural stands. N concentrations were unaffected and N:P ratios were significantly higher in the ancient stands. Nutrient concentrations varied considerably among species, but the variation was unrelated to their colonization capacity. Six out of the seven species had higher biomass in the post-agricultural stands relative to the ancient stands, and the degree to which the species increased biomass was positively related to their colonization capacity, i.e., the fast colonizing species showed the strongest increase. Such differential responses to past land use may contribute to the contrasting colonization capacity of forest plants. Land-use history thus affected both the nutrient concentrations and biomass of forest herbs, and only the biomass response was related to colonization capacity

Global patterns of intraspecific leaf trait responses to elevation

Elevational gradients are often used to quantify how traits of plant species respond to abiotic and biotic environmental variations. Yet, such analyses are frequently restricted spatially and applied along single slopes or mountain ranges. Since we know little on the response of intraspecific leaf traits to elevation across the globe, we here perform a global meta-analysis of leaf traits in 109 plant species located in 4 continents and reported in 71 studies published between 1983 and 2018. We quantified the intraspecific change in seven morpho-ecophysiological leaf traits along global elevational gradients: specific leaf area (SLA), leaf mass per area (LMA), leaf area (LA), nitrogen concentration per unit of area (Narea), nitrogen concentration per unit mass (Nmass), phosphorous concentration per unit mass (Pmass) and carbon isotope composition (delta C-13). We found LMA, Narea, Nmass and delta C-13 to significantly increase and SLA to decrease with increasing elevation. Conversely, LA and Pmass showed no significant pattern with elevation worldwide. We found significantly larger increase in Narea, Nmass, Pmass and delta C-13 with elevation in warmer regions. Larger responses to increasing elevation were apparent for SLA of herbaceous compared to woody species, but not for the other traits. Finally, we also detected evidences of covariation across morphological and physiological traits within the same elevational gradient. In sum, we demonstrate that there are common cross-species patterns of intraspecific leaf trait variation across elevational gradients worldwide. Irrespective of whether such variation is genetically determined via local adaptation or attributed to phenotypic plasticity, the leaf trait patterns quantified here suggest that plant species are adapted to live on a range of temperature conditions. Since the distribution of mountain biota is predominantly shifting upslope in response to changes in environmental conditions, our results are important to further our understanding of how plants species of mountain ecosystems adapt to global environmental change

Effects of soil compaction on growth and survival of tree saplings: a meta-analysis

Soil compaction due to mechanized harvesting operations in forests can have profound effects on forest soils and, hence, can have a detrimental effect on subsequent forest regeneration. We performed a meta-analysis to quantify the effect of soil compaction on height growth, diameter growth, and survival of tree saplings. These effects were predominantly insignificant, varied strongly and were thus not unambiguously negative. Only on silty soils, growth and survival were significantly reduced by soil compaction, which contrasted with sandy and loamy soils, where the effect of soil compaction was negligible or even slightly positive. A weighted analysis revealed an overall decrease of height growth on the compacted area, but this result should be interpreted with caution due to the limited number of observations. Although results did not show an overall negative effect of soil compaction, harvesting activities should focus on minimizing soil compaction degree and extent to prevent a decrease of soil productivity. From a methodological point of view we suggest providing more basic statistics in the articles and to include more shade-tolerant tree species in future experimental designs. These species are currently underrepresented

Plant and soil microbe responses to light, warming and nitrogen addition in a temperate forest

1. Temperate forests across Europe and eastern North America have become denser since the 1950s due to less intensive forest management and global environmental changes such as nitrogen deposition and climate warming. Denser tree canopies result in lower light availability at the forest floor. This shade may buffer the effects of nitrogen deposition and climate warming on understorey plant communities. 2. We conducted an innovative in situ field experiment to study the responses of co-occurring soil microbial and understorey plant communities to nitrogen addition, enhanced light availability and experimental warming in a full-factorial design. 3. We determined the effects of multiple environmental drivers and their interactions on the soil microbial and understorey plant communities, and assessed to what extent the soil microbial and understorey plant communities covary. 4. High light led to lower biomass of the soil microbes (analysed by phospholipid fatty acids), but the soil microbial structure, i.e. the ratio of fungal biomass to bacterial biomass, was not affected by light availability. The composition of the soil bacterial community (analysed by high-throughput sequencing) was affected by both light availability and warming (and their interaction), but not by nitrogen addition. Yet, the number of unique operational taxonomic units was higher in plots with nitrogen addition, and there were significant interactive effects of light and nitrogen addition. Light availability also determined the composition of the plant community; no effects of nitrogen addition and warming were observed. The soil bacterial and plant communities were co-structured, and light availability explained a large part of the variance of this co-structure. 5. We provide robust evidence for the key role of light in affecting both the soil microbial and plant communities in forest understoreys. Our results advocate for more multifactor global change experiments that investigate the mechanism underlying the (in) direct effects of light on the plant-soil continuum in forests

The abundance of Ixodes ricinus ticks depends on tree species composition and shrub cover

The mainstream forestry policy in many European countries is to convert coniferous plantations into (semi-natural) deciduous woodlands. However, woodlands are the main habitat for Ixodes ricinus ticks. Therefore, assessing to what extent tick abundance and infection with Borrelia spirochetes are affected by forest composition and structure is a prerequisite for effective prevention of Lyme borreliosis. We selected a total of 25 pine and oak stands, both with and without an abundant shrub layer, in northern Belgium and estimated tick abundance between April and October 2008-2010. Additionally, the presence of deer beds was used as an indicator of relative deer habitat use. Borrelia infections in questing nymphs were determined by polymerase chain reactions. The abundance of larvae, nymphs, and adults was higher in oak stands compared to pine stands and increased with increasing shrub cover, most likely due to differences in habitat use by the ticks' main hosts. Whereas tick abundance was markedly higher in structure-rich oak stands compared to homogeneous pine stands, the Borrelia infection rates in nymphs did not differ significantly. Our results indicate that conversion towards structure-rich deciduous forests might create more suitable tick habitats, but we were unable to detect an effect on the infection rate

Plasticity in response to phosphorus and light availability in four forest herbs

The differential ability of forest herbs to colonize secondary forests on former agricultural land is generally attributed to different rates of dispersal. After propagule arrival, however, establishing individuals still have to cope with abiotic soil legacies from former agricultural land use. We focused on the plastic responses of forest herbs to increased phosphorus availability, as phosphorus is commonly found to be persistently bioavailable in post-agricultural forest soils. In a pot experiment performed under field conditions, we applied three P levels to four forest herbs with contrasting colonization capacities: Anemone nemorosa, Primula elatior, Circaea lutetiana and Geum urbanum. To test interactions with light availability, half of the replicas were covered with shade cloths. After two growing seasons, we measured aboveground P uptake as well as vegetative and regenerative performance. We hypothesized that fast-colonizing species respond the most opportunistically to increased P availability, and that a low light availability can mask the effects of P on performance. All species showed a significant increase in P uptake in the aboveground biomass. The addition of P had a positive effect on the vegetative performances of two of the species, although this was unrelated to their colonization capacities. The regenerative performance was affected by light availability (not by P addition) and was related to the species' phenology. Forest herbs can obviously benefit from the increased availability of P in post-agricultural forests, but not all species respond in the same way. Such differential patterns of plasticity may be important in community dynamics, as they affect the interactions among species

Light and warming drive forest understorey community development in different environments

Plant community composition and functional traits respond to chronic drivers such as climate change and nitrogen (N) deposition. In contrast, pulse disturbances from ecosystem management can additionally change resources and conditions. Community responses to combined environmental changes may further depend on land‐use legacies. Disentangling the relative importance of these global change drivers is necessary to improve predictions of future plant communities. We performed a multifactor global change experiment to disentangle drivers of herbaceous plant community trajectories in a temperate deciduous forest. Communities of five species, assembled from a pool of 15 forest herb species with varying ecological strategies, were grown in 384 mesocosms on soils from ancient forest (forested at least since 1850) and postagricultural forest (forested since 1950) collected across Europe. Mesocosms were exposed to two‐level full‐factorial treatments of warming, light addition (representing changing forest management) and N enrichment. We measured plant height, specific leaf area (SLA) and species cover over the course of three growing seasons. Increasing light availability followed by warming reordered the species towards a taller herb community, with limited effects of N enrichment or the forest land‐use history. Two‐way interactions between treatments and incorporating intraspecific trait variation (ITV) did not yield additional inference on community height change. Contrastingly, community SLA differed when considering ITV along with species reordering, which highlights ITV’s importance for understanding leaf morphology responses to nutrient enrichment in dark conditions. Contrary to our expectations, we found limited evidence of land‐use legacies affecting community responses to environmental changes, perhaps because dispersal limitation was removed in the experimental design. These findings can improve predictions of community functional trait responses to global changes by acknowledging ITV, and subtle changes in light availability. Adaptive forest management to impending global change could benefit the restoration and conservation of understorey plant communities by reducing the light availability