Christina Skarpe

Soil and vegetation nutrient response to bison carcasses in Bialowieza Primeval Forest, Poland

Ungulate carcasses can have important effects on the surrounding soil and vegetation. The impact of six carcasses of European bison (Bison bonasus) was investigated for the first time in a natural temperate forest (Białowieża, Poland) by measuring soil and plant nutrient concentrations along a gradient extending from the centre of each carcass. Calcium concentration and pH were found to be higher at the centre of the carcass, decreasing towards the periphery. This effect lasted up to 7xa0years after the death of the animal. The concentration of most nutrients in the soil and plants varied irregularly, suggesting an effect of the carcass at its centre but the absence of a clear pattern of variation along the gradient. Concentrations of NO3− in the soil differed only at the 1-year old carcass, suggesting a fast turnover of nitrate in temperate forests. Our results show that the effects of large herbivore carcasses on soil and plant nutrient concentrations are not easily detectable in a temperate forest as in more homogeneous habitats, such as tundra and prairie. This may be due to the high activity of scavengers and nutrient recycling in the study area, but it may also be a consequence of a more complex and patchy interaction between nutrient availability and other limiting factors in temperate forests.

Tree cover and biomass increase in a southern African savanna despite growing elephant population.

The growing elephant populations in many parts of southern Africa raise concerns of a detrimental loss of trees, resulting in overall reduction of biodiversity and ecosystem functioning. Elephant distribution and density can be steered through artificial waterpoints (AWPs). However, this leaves resident vegetation no relief during dry seasons. We studied how the introduction of eight AWPs in 1996 affected the spatiotemporal tree-structure dynamics in central Chobe National Park, an unfenced savanna area in northern Botswana with a dry-season elephant density of approximately 3.34 individuals per square kilometer. We hypothesized that the impact of these AWPs amplified over time and expanded in space, resulting in a decrease in average tree density, tree height, and canopy volume. We measured height and canopy dimensions of all woody plants around eight artificial and two seasonal waterpoints for 172 plots in 1997, 2000, and 2008. Plots, consisting of 50 x 2 m transects for small trees (0.20-3.00 m tall) nested within 50 x 20 m transects for large trees (> or = 3.0 m tall), were located at 100, 500, 1000, 2000, and 5000 m distance classes. A repeated-measures mixed-effect model showed that tree density, cover, and volume had increased over time throughout the area, caused by a combination of an increase of trees in lower size classes and a decrease in larger size classes. Our results indicate that the decrease of large trees can be attributed to a growing elephant population. Decrease or loss of particular tree size classes may have been caused by a loss of browser-preferred species while facilitating the competitiveness of less-preferred species. In spite of 12 years of artificial water supply and an annual elephant population growth of 6%, we found no evidence that the eight AWPs had a negative effect on tree biomass or tree structure. The decreasing large-tree component could be a remainder of a depleted but currently restoring elephant population.

Secondary metabolites and nutrients of woody plants in relation to browsing intensity in African savannas

Carbon-based secondary metabolites (CBSMs) are assumed to function as defences that contribute to herbivore-avoidance strategies of woody plants. Severe browsing has been reported to reduce concentrations of CBSMs and increase N concentrations in individual plants, causing heavily browsed plants to be characterised by N-rich/C-poor tissues. We hypothesised that concentrations of condensed tannins (CT) and total polyphenols (TP) should decrease, or N increase, in relation to increasing intensity of browsing, rendering severely browsed plants potentially more palatable (increased N:CT) and less N-limited (increased N:P) than lightly browsed ones. We sampled naturally browsed trees (taller than 2xa0m) of four abundant species in southern Kruger National Park, South Africa. Species-specific relationships between N:CT, CT, TP and P concentrations and increasing browsing intensity were detected, but N and N:P were consistently invariable. We developed a conceptual post-hoc model to explain diverse species-specific CBSM responses on the basis of relative allocation of C to total C-based defence traits (e.g. spines/thorns, tough/evergreen leaves, phenolic compounds). The model suggests that species with low allocation of C to C-based defence traits become C-limited (potentially more palatable) at higher browsing intensity than species with high allocation of C to C-based defences. The model also suggests that when N availability is high, plants become C-limited at higher browsing intensity than when N availability is low.

Cascading effects of moose ( Alces alces ) management on birds

Large herbivores often have key functions in their ecosystems, and may affect ecosystem processes with cascading effects on other animals. The mechanisms often involve relocations of resources of various kinds, including reduction in resource availability following foraging and increase in resources from animal excreta. As large herbivore populations in Europe generally are intensely managed, management activities may interact with the activities of the herbivores themselves in the effect on other ecosystem components. We investigated the effects of moose (Alces alces) winter browsing, together with the effect of net nutrient input via supplementary winter feeding of moose on functional composition and species richness of birds in a boreal forest. Supplementary feeding stations for moose had a net zero effect on bird species richness and abundance, because negative effects of moose browsing were balanced by positive effects of nutrient input. Sites with a similar browsing intensity as at feeding stations but without nutrient input had lower abundance and species richness than feeding stations. Functional groups of bird species showed differing responses: birds nesting at or below browsing height were negatively affected by moose browsing, whereas species nesting above the browsing zone were positively affected by moose browsing. Insect-eating species responded negatively to moose browsing on birch but positively to nutrient input at feeding stations, whereas seed-eating species responded positively to birch browsing and negatively to feeding stations. This study showed that both high levels of cervid activity and human management interventions influence bird communities.

Felled Scots pine (Pinus sylvestris) as supplemental forage for moose (Alces alces): Browse availability and utilization

Abstract Old trees felled by forestry actions or natural disturbances can supply large herbivores with a substantial amount of forage and thereby potentially affect browsing patterns. To explore the felling of mature Scots pine (Pinus sylvestris L.) as a management practice (and potentially divert moose from young pine forests) this study identified how much browse there is available on felled trees and to what extent this browse is used by moose (Alces alces L.). Pine trees were surveyed in two study areas in Sweden and one in Norway. Pine crowns from thinning-stage trees held on average 8 kg of potential moose forage (dry weight). Corresponding figures for mature trees and seed trees were 29 kg and 12 kg, respectively. When the trees were processed during commercial felling, much of the potential forage became unavailable, and only approximately 1.5 kg per tree was available for moose after processing. The utilization of the forage on felled trees ranged between 5 and 15%. No significant difference in utilization between tree ages or between tops and bases within the tree crowns were found. However, the bite diameters on twigs differed between parts and age stages of the trees. This study illustrates that felling of trees can influence the availability of winter forage for moose and thereby has the potential to decrease browsing pressure on young forest stands. The substantial effect that storms can have on short-term forage supply for moose is also emphasized.

Of goats and spines – a feeding experiment

Spines in plants have evolved to reduce mammalian herbivory, and their main function may be to protect twigs more than photosynthetic tissue. Type and frequency of spines vary at different scales. We hypothesised that different types of spines affect animal foraging through different mechanisms. We studied feeding behaviour by twig-browsing goats in relation to two types of spines of Acacia tortilis using experimental manipulation of the occurrence of spines. Feeding time, number of biting actions, number and diameter of bites on trees (post-trial) and total intake were recorded. The removal of either long straight spines or short hooked spines resulted in no feeding responses by goats. The removal of both types of spines tended to increase feeding time resulting in more and larger bites with larger bite diameters and in increased total intake and utilisation compared to control branches. The removal of spines had no effects on feeding rate, expressed as biting actions per minute, number of twigs bitten per minute or intake rate (grams per minute). Both types of spines reduced total intake and utilisation of browse, but the functional mechanisms were different with the long straight spines mainly influencing bite size and short hooked spines mainly affecting number of bites.

Nutrient and secondary metabolite concentrations in a savanna are independently affected by large herbivores and shoot growth rate

AbstractnCarbon-based secondary metabolites (CBSMs) such as tannins are assumed to function as plant defences against herbivores. CBSMs are thought to be inversely related to growth rate and nutrient concentrations because a physiological trade-off exists between cellular growth and differentiation, but CBSM concentrations can be altered by herbivory-induced changes in the trade-off. We predicted that a significant interaction exists between herbivory and growth phase, such that the effects of large herbivores (or their exclusion) on nutrient or CBSM concentrations are greatest during phases of rapid shoot or leaf growth. Leaf samples were collected during phases of different growth rate from six woody species 4xa0years after establishment of a large-scale long-term herbivore exclusion experiment in Kruger National Park, South Africa. Samples were analysed for N, P, condensed tannins and total phenolics. Interactions between growth phase and herbivores were rare. However, the assumption that elevated nutrients and reduced CBSMs occurs during fast phases of growth was supported by four species (consistent with the growth-differentiation balance hypothesis), but not the other two. Large herbivores generally did not affect nutrients, but CBSMs in four species were reduced by large herbivores other than elephants, while CBSMs in two species were reduced by elephants. Carbon limitation ultimately prevailed among woody plants taller than 2xa0m under long-term browsing. Large herbivores and plant growth phase are independent and important determinants of nutrients or CBSMs in African savannas, but the effects depend on the interacting assemblages of species, which poses challenges to the application of current general hypotheses of plant defence.

Vole preference of bilberry along gradients of simulated moose density and site productivity

Browsing by large herbivores might either increase or decrease preference for the plant by other herbivores, depending on the plant response. Using a cafeteria test, we studied the preference by root voles (Microtus oeconomus [Pallas, 1776]) for bilberry (Vaccinium myrtillus L.) previously subjected to 4 levels of simulated moose (Alces alces [Linnaeus, 1758]) density. The different levels of moose density were simulated at population densities relevant for Fennoscandian conditions, in exclosures situated along a site productivity gradient. We expected: (i) voles to prefer bilberry from high productivity sites over low productivity sites; (ii) voles to prefer browsed bilberry, if plants allocate resources to compensatory growth or to avoid browsed bilberry if plants allocate resources to defense; (iii) these effects to increase with increasing simulated moose density; and (iv) the concentration of plant chemicals and the plant morphology to explain vole preference. Specifically, we predicted that voles would prefer: (i) plants with high nitrogen content; (ii) plants with low content of defensive substances; and (iii) tall plants with long shoots. Voles preferred bilberry from the high productivity sites compared to the low productivity sites. We also found an interaction between site productivity and simulated moose density, where voles preferred unbrowsed plants at low productivity sites and intermediate levels of browsing at high productivity sites. There was no effect of plant chemistry or morphology on vole preference. We conclude that moose browsing impacts the food preference of voles. With the current high densities of moose in Fennoscandia, this could potentially influence vole food selection and population dynamics over large geographical areas.

Small mammal responses to moose supplementary winter feeding

Supplementary feeding of wild large herbivores is a widespread practice in North America and Europe. The presence of feeding stations may have ecological consequences through changes to animal distributions, patterns of herbivory and a net nutrient input into the ecosystem. In Fennoscandia, supplementary feeding of moose in winter (Alces alces) is increasing. Although it has been shown to affect bird communities, its effects on small mammal communities were unknown. Here, we studied the effects of moose supplementary feeding stations on plants and on abundance, reproduction, and biomass of small mammals in years with low and high vole abundance. We sampled small mammals with snap traps and conducted surveys of the field layer vegetation, at varying distances from moose supplemental feeding stations. Due to the vegetation changes induced by feeding stations, abundance of common shrews (Sorex araneus) and Microtus voles were positively affected by long-term moose winter feeding, while bank voles (Myodes glareolus) were not affected. Moose feeding stations did not affect reproduction, individual body mass, or the total biomass of small mammals. Moose winter-feeding stations have impacts on nontarget species, providing islands of preferred grass and forb habitat for Microtus spp. and common shrews, allowing them to penetrate into a matrix of less preferred forest habitat.