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 km2. 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.

Varied diets, including broadleaved forage, are important for a large herbivore species inhabiting highly modified landscapes.

Diet quality is an important determinant of animal survival and reproduction, and can be described as the combination of different food items ingested, and their nutritional composition. For large herbivores, human landscape modifications to vegetation can limit such diet-mixing opportunities. Here we use southern Sweden's modified landscapes to assess winter diet mixtures (as an indicator of quality) and food availability as drivers of body mass (BM) variation in wild moose (Alces alces). We identify plant species found in the rumen of 323 moose harvested in Oct-Feb, and link variation in average calf BM among populations to diets and food availability. Our results show that variation in calf BM correlates with variation in diet composition, diversity, and food availability. A varied diet relatively rich in broadleaves was associated with higher calf BM than a less variable diet dominated by conifers. A diet high in shrubs and sugar/starch rich agricultural crops was associated with intermediate BM. The proportion of young production forest (0-15 yrs) in the landscape, an indicator of food availability, significantly accounted for variation in calf BM. Our findings emphasize the importance of not only diet composition and forage quantity, but also variability in the diets of large free-ranging herbivores.

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.