EFFECTS OF ESSENTIAL OILS ON THE FEEDING CHOICE BY MOOSE

Moose (Alces alces) browse on coniferous tree species to different extents during winter; for example, Norway spruce (Picea abies) is avoided, Scots pine (Pinus sylvestris) is preferred, with juniper (Juniperus communis) of intermediate use. Conifers contain essential oils that may act as feeding deterrents, thereby reducing food intake by herbivores. Because essential oils are volatile, our objectives were to determine if 1) odour plays a role in the food choice by moose, 2) whether single monoterpenes act as feeding deterrents, and 3) if this might be a mechanism used to discriminate against unpalatable plants. The essential oils of Norway spruce and juniper and 2 monoterpenes (limonene and camphene) predominant in the essential oil of Norway spruce were tested for their potential as deterrents in feeding trials. Deterrence was assessed in food choice experiments by measuring the time spent feeding on food treated with the different odours associated with these compounds. There was no statistical evidence that food treated with the essential oils of spruce and juniper and single monoterpenes from Norway spruce were avoided by moose. However, our data indicate that the essential oil of Norway spruce probably has a negative effect on moose foraging because of the large absolute difference in feeding time between treatments and that overall, odour had a significant effect on feeding time. Because our experimental design may have influenced the results, we suggest research approaches to better measure deterrence effects

Effect of fire on the palatability of plants in an African woodland savanna: varying impacts depending on plant functional groups

Fire and herbivores are two important drivers of changes in vegetation composition, quality and dynamics and both are highly related to each other. Herbivores are known to respond to fire both in terms of foraging decisions and distribution. However, little is known about the actual changes in plant chemistry following a fire event and how long these changes will last. We investigated the effect of fire on two different plant functional groups (grasses and woody species) in a woodland savanna of southern Africa. We studied chemical compounds known to be important for palatability of five perennial grass and seven woody species (trees and shrubs) common in the woodland savanna and known to be utilized by herbivores. We wanted to know if plant chemistry differs between a recently burned site (burned 2 years ago) and a control site, burned 16 years ago, and if grasses and woody species show similar relative differences between sites (i.e., the plants' response to fire). We found a clear difference in chemical composition patterns between the plant functional groups, with an almost homogenous response to fire among woody species, but higher variability in response among grass species. Furthermore, we found that woody species maintained a higher nutritional value even 2 years after burning, whereas grasses did not show clear differences among the two investigated sites. Hence, few years after burning, woody plants might still serve as an attraction for herbivores, especially browsers, in contrast to grasses. The knowledge about these differences between the two functional groups in response to fire is beneficial for the development of management strategies for large herbivores whether domestic or wild

Compositional Changes in Foliage Phenolics with Plant Age, a Natural Experiment in Boreal Forests

The composition of plant secondary metabolites (PSMs) extensively impacts ecosystem functioning. It is vital that we understand temporal patterns in the plants’ allocation of resources to PSMs, particularly those influenced by human activity. Existing data are insufficient in the long-term perspective of perennial plants (age or ontogeny). We analysed phenolic concentrations in foliage from birch (Betula pubescens Ehr.) considered to be undamaged and growing on 5, 10 and 15 years old clear-cuts in two boreal forest landscapes in Norway, sampled at the peak of the growing season. In sum, low molecular weight phenolic concentrations decreased with age. Apart from one apigenin glycoside, the low molecular weight phenolics co-varied similarly at all ages, suggesting a lack of temporal compound-specific prioritisation of this group. In contrast, the concentration of MeOH-soluble condensed tannins increased with age. The compositional shift fits well with several hypotheses that may provide proximate explanations for age patterns in PSM allocations, including both resource constraints and external pressures. Regardless of these explanations, our study adds an important perennial perspective (plant age) to temporal PSM patterns already well-known in boreal plant phenology (foliage age).publishedVersio

Moose selecting for specific nutritional composition of birch places limits on food acceptability

Despite decades of intense research, it remains largely unsolved which nutritional factors underpin food selection by large herbivores in the wild. We measured nutritional composition of birch foliage (Betula pubescens) available to, and used by, moose (Alces alces) in natural settings in two neighboring regions with contrasting animal body mass. This readily available food source is a staple food item in the diet of moose in the high-fitness region, but apparently underutilized by moose in the low-fitness region. Available birch foliage in the two regions had similar concentrations of macronutrients (crude protein [CP], fiber fractions, and water-soluble carbohydrates [WSC]), although a notably lower variation of WSC in the low-fitness region. For minerals, there were several area differences: available birch foliage in the low-fitness region had less Mg (depending on year) and P, but more Ca, Zn, Cu, and Mn. It also had higher concentrations of some plant secondary metabolites: chlorogenic acids, quercetins, and especially MeOH-soluble condensed tannins. Despite the area differences in available foliage, we found the same nutritional composition of birch foliage used in the two regions. Compared to available birch foliage, moose consistently used birch foliage with more CP, more structural fiber (mainly hemicellulose), less WSC, higher concentrations of several minerals (Ca, Zn, K, Mn, Cu), and lower concentrations of some secondary metabolites (most importantly, MeOH-soluble condensed tannins). Our study conceptually supports the nutrient-balancing hypothesis for a large herbivore: within a given temporal frame, moose select for plant material that matches a specific nutritional composition. As our data illustrate, different moose populations may select for the same composition even when the nutritional composition available in a given food source varies between their living areas. Such fastidiousness limits the proportion of available food that is acceptable to the animal and has bearings on our understanding and application of the concept of carrying capacity.publishedVersio

Long-term effect of sheep and goat grazing on plant diversity in a semi-natural dry grassland habitat

Semi-natural dry grassland sites are of great importance for nature conservation because they support high species diversity and the abundance of “Red-List” species. Grazing has proved to be a successful management tool in terms of maintenance and restoration of biodiversity. For a deeper understanding of the effects of different grazers on species biodiversity in dry grasslands, it is necessary to study the long-term effects of major changes in grazing management. In a semi-natural dry grassland habitat, which was formerly grazed by cattle, we investigated the changes in plant species composition due to long term grazing by sheep and goats. Specifically we asked: a) How does long-term grazing by sheep and goats change the composition of all plant species and particularly those that are on the Red-List? Are changes caused mainly by species turnover? b) How does long-term grazing by sheep and goats influence the fertility and acidity of the soil? To address these questions, we compared the composition and diversity of plants as well deriving Ellenberg indicator values of the species. Long-term grazing by sheep and goats subsequent to a year-round cattle grazing changed the plant species composition of the dry grasslands resulting in a high species turnover rate. It did not, however, lead to an increase in plant species diversity even though Red-List species were considerably more abundant in 2013. Overall, the grazing regime studied positively influenced vegetation composition. The effects on local species composition due to species turnover might further be influenced by local factors like soil nitrogen availability

Macronutrient balancing in free-ranging populations of moose

At northern latitudes, large spatial and temporal variation in the nutritional composition of available foods poses challenges to wild herbivores trying to satisfy their nutrient requirements. Studies conducted in mostly captive settings have shown that animals from a variety of taxonomic groups deal with this challenge by adjusting the amounts and proportions of available food combinations to achieve a target nutrient balance. In this study, we used proportions-based nutritional geometry to analyze the nutritional composition of rumen samples collected in winter from 481 moose (Alces alces) in southern Sweden and examine whether free-ranging moose show comparable patterns of nutrient balancing. Our main hypothesis was that wild moose actively regulate their rumen nutrient composition to offset ecologically imposed variation in the nutritional composition of available foods. To test this, we assessed the macronutritional composition (protein, carbohydrates, and lipids) of rumen contents and commonly eaten foods, including supplementary feed, across populations with contrasting winter diets, spanning an area of approximately 10,000 km(2). Our results suggest that moose balanced the macronutrient composition of their rumen, with the rumen contents having consistently similar proportional relationship between protein and nonstructural carbohydrates, despite differences in available (and eaten) foods. Furthermore, we found that rumen macronutrient balance was tightly related to ingested levels of dietary fiber (cellulose and hemicellulose), such that the greater the fiber content, the less protein was present in the rumen compared with nonstructural carbohydrates. Our results also suggest that moose benefit from access to a greater variety of trees, shrubs, herbs, and grasses, which provides them with a larger nutritional space to maneuver within. Our findings provide novel theoretical insights into a model species for ungulate nutritional ecology, while also generating data of direct relevance to wildlife and forest management, such as silvicultural or supplementary feeding practices

Translating nutritional ecology from the laboratory to the field: milestones in linking plant chemistry to population regulation in mammalian browsers

A central goal of nutritional ecology is to understand how variation in food quality limits the persistence of wild animal populations. Habitat suitability for browsing mammals is strongly affected by concentrations of nutrients and plant secondary metabolites (PSMs), but our understanding of this is based mostly on short-term experiments of diet selection involving captive animals. In the wild, browsers forage in biologically, chemically and spatially-complex environments, and foraging decisions in response to varying food quality will be correspondingly complicated. We have identified four steps that must be achieved in order to translate our understanding from laboratory experiments to populations of mammalian browsers: 1) knowing what foods and how much of these wild browsers eat, as well as what they avoid eating; 2) knowing the relevant aspects of plant nutritional and defensive chemistry to measure in a given system and how to measure them; 3) understanding the spatial distribution of nutrients and PSMs in plant communities, the costs they impose on foraging and the effects on animals' distributions; and 4) having appropriate statistical tools to analyse the data. We discuss prospects for each of these prerequisites for extending laboratory studies of nutritional quality, and review recent developments that may offer solutions for field studies. We also provide a synthesis of how to use this nutritional knowledge to link food quality to population regulation in wild mammals and describe examples that have successfully achieved this aim

The Nutritional Balancing Act of a Large Herbivore: An Experiment with Captive Moose (Alces alces L).

The nutrient balancing hypothesis proposes that, when sufficient food is available, the primary goal of animal diet selection is to obtain a nutritionally balanced diet. This hypothesis can be tested using the Geometric Framework for nutrition (GF). The GF enables researchers to study patterns of nutrient intake (e.g. macronutrients; protein, carbohydrates, fat), interactions between the different nutrients, and how an animal resolves the potential conflict between over-eating one or more nutrients and under-eating others during periods of dietary imbalance. Using the moose (Alces alces L.), a model species in the development of herbivore foraging theory, we conducted a feeding experiment guided by the GF, combining continuous observations of six captive moose with analysis of the macronutritional composition of foods. We identified the moose's self-selected macronutrient target by allowing them to compose a diet by mixing two nutritionally complementary pellet types plus limited access to Salix browse. Such periods of free choice were intermixed with periods when they were restricted to one of the two pellet types plus Salix browse. Our observations of food intake by moose given free choice lend support to the nutrient balancing hypothesis, as the moose combined the foods in specific proportions that provided a particular ratio and amount of macronutrients. When restricted to either of two diets comprising a single pellet type, the moose i) maintained a relatively stable intake of non-protein energy while allowing protein intakes to vary with food composition, and ii) increased their intake of the food item that most closely resembled the self-selected macronutrient intake from the free choice periods, namely Salix browse. We place our results in the context of the nutritional strategy of the moose, ruminant physiology and the categorization of food quality

Comparison of plant secondary metabolites and digestibility of three different boreal coniferous trees

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Translating nutritional ecology from the laboratory to the field : milestones in linking plant chemistry to population regulation in mammalian browsers

A central goal of nutritional ecology is to understand how variation in food quality limits the persistence of wild animal populations. Habitat suitability for browsing mammals is strongly affected by concentrations of nutrients and plant secondary metabolites (PSMs), but our understanding of this is based mostly on short-term experiments of diet selection involving captive animals. In the wild, browsers forage in biologically, chemically and spatially-complex environments, and foraging decisions in response to varying food quality will be correspondingly complicated. We have identified four steps that must be achieved in order to translate our understanding from laboratory experiments to populations of mammalian browsers: 1) knowing what foods and how much of these wild browsers eat, as well as what they avoid eating; 2) knowing the relevant aspects of plant nutritional and defensive chemistry to measure in a given system and how to measure them; 3) understanding the spatial distribution of nutrients and PSMs in plant communities, the costs they impose on foraging and the effects on animals ’ distributions; and 4) having appropriate statistical tools to analyse the data. We discuss prospects for each of these prerequisites for extending laboratory studies of nutritional quality, and review recent developments that may offer solutions for field studies. We also provide a synthesis of how to use this nutritional knowledge to link food quality to population regulation in wild mammals and describe examples that have successfully achieved this aim