Shrub clearing adversely affects the abundance of Ixodes ricinus ticks

In order to get a better understanding of the importance of vertical forest structure as a component of Ixodes ricinus tick habitat, an experiment was set up in a coniferous forest on sandy soils in northern Belgium. Ticks were sampled in six control and six treatment plots on various sampling occasions in 2008–2010. In the course of the study period, a moderate thinning was carried out in all plots and shrub clearing was performed in the treatment plots. Thinning had no effect on tick abundance, while shrub clearing had an adverse affect on the abundance of all three life stages (larva, nymph, adult) up to 2 years post-clearing. Our findings are especially relevant in the light of the ongoing efforts to improve vertical forest structure in Belgium and many other parts of Europe, which might create suitable habitats for ticks and change the epidemiology of tick-borne diseases. Also, our results indicate that shrub clearing could be applied as a tick control measure in recreational areas where there is a high degree of human-tick contact.     

Environmental drivers interactively affect individual tree growth across temperate European forests

Forecasting the growth of tree species to future environmental changes requires a better understanding of its determinants. Tree growth is known to respond to global‐change drivers such as climate change or atmospheric deposition, as well as to local land‐use drivers such as forest management. Yet, large geographical scale studies examining interactive growth responses to multiple global‐change drivers are relatively scarce and rarely consider management effects. Here, we assessed the interactive effects of three global‐change drivers (temperature, precipitation and nitrogen deposition) on individual tree growth of three study species (Quercus robur/petraea, Fagus sylvatica and Fraxinus excelsior). We sampled trees along spatial environmental gradients across Europe and accounted for the effects of management for Quercus. We collected increment cores from 267 trees distributed over 151 plots in 19 forest regions and characterized their neighbouring environment to take into account potentially confounding factors such as tree size, competition, soil conditions and elevation. We demonstrate that growth responds interactively to global‐change drivers, with species‐specific sensitivities to the combined factors. Simultaneously high levels of precipitation and deposition benefited Fraxinus, but negatively affected Quercus’ growth, highlighting species‐specific interactive tree growth responses to combined drivers. For Fagus, a stronger growth response to higher temperatures was found when precipitation was also higher, illustrating the potential negative effects of drought stress under warming for this species. Furthermore, we show that past forest management can modulate the effects of changing temperatures on Quercus’ growth; individuals in plots with a coppicing history showed stronger growth responses to higher temperatures. Overall, our findings highlight how tree growth can be interactively determined by global‐change drivers, and how these growth responses might be modulated by past forest management. By showing future growth changes for scenarios of environmental change, we stress the importance of considering multiple drivers, including past management and their interactions, when predicting tree growth.     

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.     

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.     

Predicting patterns of invasion by black cherry (Prunus serotina Ehrh.) in Flanders (Belgium) and its impact on the forest understorey community

The design of cost-efficient control strategies for invasive species that are too widespread and abundant for complete eradication, at least in the short term, will benefit from a rigorous analysis of invasion patterns and associated effects on native biodiversity. In this paper, the case of the invasive North American tree Prunus serotina in Flanders (Belgium) is presented. Our main objectives were to determine the susceptibility of forest stands to invasion by P. serotina and the subsequent effects of invasion on the understorey community. We used the large database of the first Flemish Forest Inventory. Multiple logistic regressions indicated that P. serotina occurred more frequently in privately owned, younger forest on coarse-textured, dry soils (podzols), and the combination of these factors allowed us to correctly predict presence/absence of P. serotina in 70% of the validation plots. However, locational variables proved to be important as well, indicating that the invasion process is still ongoing. Prediction of P. serotina densities by means of multiple linear regressions was less successful. Effects on the understorey richness were analysed by comparing the number of species and the mean Ellenberg values between pairs of plots, only differing by the presence of P. serotina in the shrub layer. A reduction of the understorey richness following invasion was only pronounced on the more moist soils, while compositional changes mainly occurred on drier soils. It is concluded that priority for control should be given to landscapes with a low fraction of invaded stands and to forest stands located on more moist soils. However, using its potential to threaten native biodiversity as an argument for control should be done with care as further research is needed whether the observed negative effects are due to a species (i.e. native vs. non-native) or a density effect (high vs. low).     

Does Prunus serotina act as an aggressive invader in areas with a low propagule pressure?

Since most studies on Prunus serotina in Western Europe focused on heavily invaded areas, we wondered whether P. serotina also acts as an aggressive invader in areas with a low propagule pressure. Based on long-term data for the Liedekerke forest reserve, we found that connectivity to seed sources and light availability were the major drivers of P. serotina presence: long-distance dispersal events and ‘windows of opportunity’ seem to direct P. serotina colonization. In the studied forest, P. serotina could not be considered an aggressive invader since its spread slowed down rather quickly and did not hamper the establishment of native tree species. Furthermore, understory P. serotina showed low growth and seed production, while the high Rubus cover hampered germination and establishment. Nonetheless, calamities opening up the canopy layer in the few areas with high P. serotina sapling density might alter the course of the invasion process.     

The effects of sampling method and vegetation type on the estimated abundance of <Emphasis Type="Italic">Ixodes ricinus</Emphasis> ticks in forests

Estimating the spatial and temporal variation in tick abundance is of great economical and ecological importance. Entire-blanket dragging is the most widely used method to sample free-living ixodid ticks. However, this technique is not equally efficient in different vegetation types. The height and structure of the vegetation under study will not only determine the likelihood of a tick-blanket contact, but will also determine the rate of dislodgement. The purpose of this study was therefore to determine whether the alternative strip-blanket is more effectively in picking up ticks than the standard entire-blanket. Sampling was carried out in four forest understory vegetation types that differed in height and structure on five collection dates between April and September 2008. A total of 8,068 Ixodes ricinus ticks was collected (778 adults, 1,920 nymphs, and 5,370 larvae). The highest numbers of ticks were collected along the forest trails, where the dominant vegetation consisted of short grasses. The lowest numbers of ticks were collected in bracken-fern-dominated sites, where the vegetation seriously hampered tick sampling. Surprisingly, in each vegetation type, significantly more nymphs and adults were collected using the entire-blanket. However, the strip-blanket was more effectively in collecting larvae, especially in dense and tall vegetation.     

Impact of avian frugivores on dispersal and recruitment of the invasive <Emphasis Type="Italic">Prunus serotina</Emphasis> in an agricultural landscape

Although seed dispersal is considered to be a key process determining the spatial structure and spread of non-native plant populations, few studies have explicitly addressed the link between dispersal vector behaviour, seed distribution and seedling recruitment to gain insight into the process of exotic species invasion within a fragmented landscape context. The present study analyses the relationship between avian frugivory and spatial patterns of seed deposition and seedling recruitment for an expanding population of the invasive Prunus serotina in a hedgerow network landscape in Flanders, Belgium. We quantified fruit production, observed frugivores, and determined the spatial distribution of bird droppings and P. serotina seedlings. A relatively diverse assemblage of frugivores visited P. serotina seed trees, with Columba palumbus and Turdus merula being by far the most important dispersers. Landscape structure strongly affected dispersal vector behaviour and the spatial distribution of perching birds, droppings and seedlings. Frugivorous birds non-randomly dispersed seeds to perching sites and an association between perching birds, seed deposition and seedling recruitment was found. Results indicate that landscape structure contributes to non-random seed deposition of P. serotina by common local frugivores. Cutting the larger seed trees is proposed as the most feasible measure to slow down the invasion rate.     

An intraspecific application of the leaf‐height‐seed ecology strategy scheme to forest herbs along a latitudinal gradient

We applied the leaf‐height‐seed (LHS) ecology strategy scheme (a combination of three ecologically important traits: specific leaf area (SLA), seed mass and plant height) intraspecifically to two widespread European forest herbs along a latitudinal gradient. The aims of this study were to quantify LHS trait variation, disentangle the environmental factors affecting these traits and compare the within‐species LHS trait relationships with latitude to previously established cross‐species comparisons. 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.