Behavioural responses of cattle to shared foraging with wild herbivores in an East African rangeland

We assessed whether prior foraging by wild herbivores affected foraging behaviour of cattle in Laikipia rangeland, Kenya, during February 2001, August 2001 and February 2002. The study compared cattle bite rate, step rate and bites per step in plots exclusively accessible to cattle and those accessible to cattle and large wild herbivores. During February 2001 when conditions were dry, cattle bite rate was 18–19% lower, step rate 25–26% higher, and bites per step 36% lower in plots shared by cattle and wildlife compared to those exclusively accessible to cattle. Differences in these measured foraging behaviour parameters were strongly correlated with reductions in herbage cover in plots accessible to wild herbivores. Plot differences in herbage cover and the measured foraging behaviour parameters were not significant in the subsequent trials when conditions were wet, suggesting that wild herbivore impacts reported here are short-term within season and dependent on weather conditions (and plant productivity). With reduced herbaceous plant cover in wildlife grazed realms in the dry season, cattle respond with increased travel and reductions in bite rate and bites per step, suggesting that wild herbivores can seasonally affect foraging behaviour of cattle. It remains to be demonstrated whether or not these altered behaviours of cattle affect weight gains or other measures of performance.     

A study of predominant aerobic microflora of black bears (Ursus americanus) and grizzly bears (Ursus arctos) in northwestern Alberta

Swab specimens were obtained from nasal, rectal, and preputial or vaginal areas of 37 grizzly and 17 black bears, captured during May to June of 1981 to 1983, to determine the types and frequency of predominant aerobic microflora. Bacterial genera most frequently isolated from bears were Escherichia, Citrobacter, Hafnia, Proteus, Staphylococcus, and Streptococcus species, comprising about 65% of the isolates. Erwinia, Xanthomonas, Agrobacterium, Rhizobium, and Gluconobacter/Acetobacter were also isolated but at lower frequencies (less than 5%). Comparison of bacterial generic composition using similarity quotient values showed no appreciable differences between grizzly and black bear flora. Also, no outstanding differences in bacterial generic composition were observed among grizzly bear samples; however, differences were noted among black bear samples. Fungal genera most commonly encountered included Cryptococcus, Rhodotorula, Cladosporium, Penicillium, Sporobolomyces, and Candida. In general, the microflora of both bear types were marked by generic diversity and random distribution. The majority of microorganisms isolated from the plant samples in the study area were also found in bear samples. This observation and the presence of certain water and soil bacteria in samples from bears suggest that the predominant microflora of both grizzly and black bears were transient and probably influenced by their foraging habits and surrounding environments.     

Animal-defined resources reveal nutritional inadequacies for woodland caribou during summer–autumn

Populations of woodland caribou (Rangifer tarandus caribou) are declining throughout their range and many are at risk of extirpation, yet the role of nutrition in these declines remains poorly understood, in part owing to a lack of information about available nutritional resources during summer. We quantified rates of intake of digestible protein and digestible energy by tame caribou foraging in temporary enclosures in the predominant plant communities of northeastern British Columbia, Canada, during summer–autumn and compared intake rates to daily requirements for protein and energy during lactation. We tested hypotheses related to the nutritional adequacy of the environment to support nutritional requirements during lactation (with and without replenishment of body reserves) and simulated scenarios of foraging by caribou in these plant communities to better understand how wild caribou could meet nutritional demands on these landscapes. Nutritional resources varied among plant communities across seasonal, ecological, and successional gradients; digestible energy intake per minute and per day were significantly greater in younger than older forests; dietary digestible energy and per-minute and daily intake of digestible protein were greater, though not significantly so, in younger than older forests; and dietary digestible protein was greater in older than younger forests, though differences were not significant. Tame caribou were unable to satisfy protein and energy requirements during lactation, even without replenishment of body reserves, at most sites sampled. Further, foraging simulations suggested widespread nutritional inadequacies on ranges of wild caribou. Selection for habitats offering the best nutrition may mitigate some nutritional inadequacies, but given low availability of vegetation communities with high nutritional value, performance (e.g., calf production, growth, replenishment of body fat and protein) of caribou may be depressed at levels of nutrition documented herein. Our results, coupled with recent measurements of body fat of wild caribou in northeastern British Columbia, refute the hypothesis that the nutritional environment available to caribou during summer in northeastern British Columbia is adequate to fully support nutritional demands of lactating caribou, which has implications to productivity of caribou populations, recovery, and conservation.     

Predicting Grizzly Bear Density in Western North America

Conservation of grizzly bears (Ursus arctos) is often controversial and the disagreement often is focused on the estimates of density used to calculate allowable kill. Many recent estimates of grizzly bear density are now available but field-based estimates will never be available for more than a small portion of hunted populations. Current methods of predicting density in areas of management interest are subjective and untested. Objective methods have been proposed, but these statistical models are so dependent on results from individual study areas that the models do not generalize well. We built regression models to relate grizzly bear density to ultimate measures of ecosystem productivity and mortality for interior and coastal ecosystems in North America. We used 90 measures of grizzly bear density in interior ecosystems, of which 14 were currently known to be unoccupied by grizzly bears. In coastal areas, we used 17 measures of density including 2 unoccupied areas. Our best model for coastal areas included a negative relationship with tree cover and positive relationships with the proportion of salmon in the diet and topographic ruggedness, which was correlated with precipitation. Our best interior model included 3 variables that indexed terrestrial productivity, 1 describing vegetation cover, 2 indices of human use of the landscape and, an index of topographic ruggedness. We used our models to predict current population sizes across Canada and present these as alternatives to current population estimates. Our models predict fewer grizzly bears in British Columbia but more bears in Canada than in the latest status review. These predictions can be used to assess population status, set limits for total human-caused mortality, and for conservation planning, but because our predictions are static, they cannot be used to assess population trend.     

Potential for woody plant control by Spanish goats in the sagebrush steppe

Botanical composition of the diet of eight Spanish goats was studied on the Northern Great Basin Experimental Range during two stages of plant phenology (active growth in early July and cured forage in mid-August) to assess their potential for the control of sagebrush (Artemisia tridentata sspp. wyomingensis Nutt.) and western juniper (Juniperus occidentalis Hook.) and their nutritional status on sagebrush-steppe rangelands. Diets were quantified by documenting the number of visits to each species, the bites harvested and time expended grazing each forage over 4 consecutive days in each period. Forage chemical characteristics evaluated included: CP, NDF, ADF, ADL and IVDMD. Diet composition and forage quality indices varied significantly (P < 0.05) with changes in plant phenology. Vegetation cover averaged 52% in the pasture and consisted of 36% grasses, 8% forbs, 7% shrubs and 0.6% trees. Available herbage (excluding woody plants) was 534 kg ha⁻¹ during active growth trials and 572 kg ha⁻¹ when forages had cured. When forages were green goats acquired 28% of their total bites from grasses, 71% from forbs, 0.3% from shrubs and 0.9% from juniper trees. After forages had cured values were 35% from grasses, 56% from forbs, 0.1% from shrubs and 8.8% from juniper. The browsing of juniper (both foliage and bark), after herbaceous forages had cured, was the only substantive use of woody plants. Sagebrush was only lightly used (0.2% of total bites) when herbaceous forages were actively growing. Available herbage was of relatively high quality during both trials. When forages were actively growing, CP of grasses ranged from 8.9 to 5.6%, forbs from 17.7 to 8.5%, sagebrush scored at 8.5%, juniper foliage at 8.1% and juniper bark at 3.2%. After herbaceous forages had cured, grass CPs ranged between 6.2 and 3.1%, forbs from 10.4 to 4.4%, shrubs were not sampled and juniper foliage averaged 7.6%. Given the low levels of browsing exhibited by goats on sagebrush and juniper we see little opportunity for control of these woody plants when pastures provide a diverse (N = 25 species) array of readily available (534–572 kg ha⁻¹) and nutritious forages and pastures are lightly stocked (0.63 goats ha⁻¹ month⁻¹). We do need, however, to further explore their potential for control of these species under several other regimes. These include extended trials on more deteriorated rangelands, trials during seasons or conditions where forage is limited and the animal's selective opportunity is restricted and trials in high quality environments on newly established sagebrush and juniper seedlings.     

Food selection in <Emphasis Type="Italic">Microtus arvalis</Emphasis>: the role of plant functional traits

We offered captive common voles (Microtus arvalis) a choice of 11 plant species (representing four ecological groups) growing in vivaria. Selection was evaluated by measuring (1) the biomass of each plant species consumed and (2) functional and life-history plant traits. The legume Trifolium pratense, known for its high nutrient level, and well accessible rosette forbs creating the highest biomass at the soil ground level, were mostly preferred. Voles avoided mainly grasses and the creeping forb Thymus pulegioides. The experiment showed that foraging was strongly plant species-specific. We assessed whether plant functional traits explain selective foraging in common voles. To explore this, we reanalyzed Holišová’s (1959) data about common vole stomach contents and plant trait databases. Regression tree analysis indicated that plant guild and life span were the best predictors of dietary selection, with a probability exceeding 0.5 that voles would eat more grasses and/or legumes than forbs. These results do not correspond with the feeding trial. We suggest that the voles usually consume grasses in the field because grasses are abundant and readily available, but prefer protein-rich forbs when possible.     

The bear circadian clock doesn’t ‘sleep’ during winter dormancy

Background

Most biological functions are synchronized to the environmental light:dark cycle via a circadian timekeeping system. Bears exhibit shallow torpor combined with metabolic suppression during winter dormancy. We sought to confirm that free-running circadian rhythms of body temperature (Tb) and activity were expressed in torpid grizzly (brown) bears and that they were functionally responsive to environmental light. We also measured activity and ambient light exposures in denning wild bears to determine if rhythms were evident and what the photic conditions of their natural dens were. Lastly, we used cultured skin fibroblasts obtained from captive torpid bears to assess molecular clock operation in peripheral tissues. Circadian parameters were estimated using robust wavelet transforms and maximum entropy spectral analyses.

Results

Captive grizzly bears housed in constant darkness during winter dormancy expressed circadian rhythms of activity and Tb. The rhythm period of juvenile bears was significantly shorter than that of adult bears. However, the period of activity rhythms in adult captive bears was virtually identical to that of adult wild denning bears as was the strength of the activity rhythms. Similar to what has been found in other mammals, a single light exposure during the bear’s active period delayed subsequent activity onsets whereas these were advanced when light was applied during the bear’s inactive period. Lastly, in vitro studies confirmed the expression of molecular circadian rhythms with a period comparable to the bear’s own behavioral rhythms.

Conclusions

Based on these findings we conclude that the circadian system is functional in torpid bears and their peripheral tissues even when housed in constant darkness, is responsive to phase-shifting effects of light, and therefore, is a normal facet of torpid bear physiology.

     

Diet and environment shape fecal bacterial microbiota composition and enteric pathogen load of grizzly bears

Background

Diet and environment impact the composition of mammalian intestinal microbiota; dietary or health disturbances trigger alterations in intestinal microbiota composition and render the host susceptible to enteric pathogens. To date no long term monitoring data exist on the fecal microbiota and pathogen load of carnivores either in natural environments or in captivity. This study investigates fecal microbiota composition and the presence of pathogenic Escherichia coli and toxigenic clostridia in wild and captive grizzly bears (Ursus arctos) and relates these to food resources consumed by bears.

Methodology/Principal Findings

Feces were obtained from animals of two wild populations and from two captive animals during an active bear season. Wild animals consumed a diverse diet composed of plant material, animal prey and insects. Captive animals were fed a regular granulated diet with a supplement of fruits and vegetables. Bacterial populations were analyzed using quantitative PCR. Fecal microbiota composition fluctuated in wild and in captive animals. The abundance of Clostridium clusters I and XI, and of C. perfringens correlated to regular diet protein intake. Enteroaggregative E. coli were consistently present in all populations. The C. sordellii phospholipase C was identified in three samples of wild animals and for the first time in Ursids.

Conclusion

This is the first longitudinal study monitoring the fecal microbiota of wild carnivores and comparing it to that of captive individuals of the same species. Location and diet affected fecal bacterial populations as well as the presence of enteric pathogens.     

Wild reindeer foraging-niche organization

Part 1: Diet selection was studied on free-ranging reindeer fitted with an esophageal fistula (EF) and by analysis of rumen samples from reindeer shot in the field. Plant density was assessed from quadrats on field plots. Three measures of palatability were used, the nitrogen, fiber, and total non structural carbohydrate (TNC) contents of samples from clipped vegetation. As lichen density decreased the EK reindeer included progressively more vascular litter and pieces of winter dormant species in their diet. Diets with highest TNC and N/F and the lowest fiber, providing the most readily digestible diets, were selected. The combined density-quality criterion gave the best prediction of their diets. In summer, forbs, and to some extent grasses and dwarf shrubs, were selected. Plants of highest density and in an early growth phase gave the best prediction of summer diets. In winter the variety of vascular plants utilized increased with increasing herd size and decreasing density of lichens in the vegetation. The width of diets was a function of diversity of available plants. It is suggested that dietary selection by reindeer has been a three stage process: evolution of a gastro-intestinal system capable of digesting lichens containing secondary compounds, behavioural tracking of the plant production pulse, and diet width scaling according to density-quality of all potential foods. Part 2: Wild reindeer closely followed the wave of vascular plant production in spring and summer with a significant correlation being found between the daily foraging time per habitat type and the highest concentration of green phytomass. Snow-bed meadows were the most consistently selected vegetation type by the four herds studied, viz. Hardangervidda, Snøhetta, Prudhoe Bay, and Svalbard, during the summer. Habitat niche-breadths were narrow during the winter, largely due to the limitation of access to the food supply by the snow-cover, broadened as the landscape became clear of snow, narrowed again with the initiation of plant growth and broadened once more as the wave of plant production reached all habitat types. At high levels of food resources the alpine herd (Hardangervidda) narrowed the feeding niche breadth (or adopted a selective feeding strategy) as the habitat niche breadth increased. The high arctic herd (Svalbard) living in the least productive environment, at the same time of year adopted a generalist feeding strategy as their habitat niche breadth expanded. Part 3: In this part the temporal organization of a low-alpine herd of reindeer (Hardangervidda) is described, and subsequently compared with those of two arctic herds. Their foraging efficiencies under different degrees of food availability were analysed, in order to test the hypothesis that food intake changes in response to the prevailing state of the food resources. The number of daily feeding bouts increased from 2 to 6 as the diet changed from the winter to the summer pasturage. Strong winds in winter and insect harassment in summer severely depressed the daily time spent foraging. When these environmental factors were accounted for no significant inter-seasonal differences in daily foraging time of the low alpine herd were found; on average 50% of each day was spent grazing, of which 78% of the time was spent on actual ingestion (feeding rate). The feeding rate within the active periods varied with the snow-depth, due to the need to dig a food crater in the snow and due to the particular selective feeding strategy adopted. The data from the present and from other studies suggest that the foraging efficiency of wild reindeer, expressed as their daily food intake follows a Holling type II functional response as food availability changes. Part 4: Sheep is the only sympatric ruminant which grazes with the reindeer on alpine pastures in Norway. The inter-specific niche relationships ot the wild reindeer and the domestic sheep in summer were studied on part of the western Hardangervidda. to test the hypothesis of inter-specific competition. Three niche dimensions were meas-ured, diel, habitat utilization, and spatial distribution. The estimates of the degree of overlap obtained suggest that this is relatively high for diet and habitat utilization but low for spatial distribution, which leads to a low degree of foraging interference. Since the intensity of both interspeeific and intra-specific competition depends on the ratio of the herbivore densities to the food resource density, the degree ol overlap in utilization by the two species tell us nothing about the degree of competition. Present numbers of both species on Hardangervidda have a beneficial influence on the vegetation, by opening-up the shrub-layer canopy and thus facilitating the main-tenance of grasses and forbs in the field layer, which has a higher carrying capacity for ruminants. The balance between the beneficial effects and inter-generic competition (if any) deriving from the present-day stocking rales is so tar unknown. Only long-term monitoring of the performance of both populations will shed light on this competitive aspect.     

Contrafreeloading in grizzly bears: implications for captive foraging enrichment

Although traditional feeding regimens for captive animals were focused on meeting physiological needs to assure good health, more recently emphasis has also been placed on non‐nutritive aspects of feeding. The provision of foraging materials to diversify feeding behavior is a common practice in zoos but selective consumption of foraging enrichment items over more balanced “chow” diets could lead to nutrient imbalance. One alternative is to provide balanced diets in a contrafreeloading paradigm. Contrafreeloading occurs when animals choose resources that require effort to exploit when identical resources are freely available. To investigate contrafreeloading and its potential as a theoretical foundation for foraging enrichment, we conducted two experiments with captive grizzly bears (Ursus arctos horribilis). In Experiment 1, bears were presented with five foraging choices simultaneously: apples, apples in ice, salmon, salmon in ice, and plain ice under two levels of food restriction. Two measures of contrafreeloading were considered: weight of earned food consumed and time spent working for earned food. More free than earned food was eaten, with only two bears consuming food extracted from ice, but all bears spent more time manipulating ice containing salmon or apples than plain ice regardless of level of food restriction. In Experiment 2, food‐restricted bears were presented with three foraging choices simultaneously: apples, apples inside a box, and an empty box. Although they ate more free than earned food, five bears consumed food from boxes and all spent more time manipulating boxes containing apples than empty boxes. Our findings support the provision of contrafreeloading opportunities as a foraging enrichment strategy for captive wildlife. Zoo Biol 29:484–502, 2010. © 2009 Wiley‐Liss, Inc.     

Optimizing protein intake as a foraging strategy to maximize mass gain in an omnivore

Energy maximization, time minimization, and linear programming models subject to various constraints have dominated foraging ecology ideas and methods for decades. However, animals must use very complex physiological processes and foraging decisions to ensure fitness that in many cases may not be adequately described by these approaches. An example of this problem occurs when brown bears, Ursus arctos, have access to both abundant salmon and fruit. Salmon are one of the most energy and nutrient dense foods available to bears. Fruits are often high in soluble carbohydrates, low to deficient in many required nutrients, and more difficult to efficiently exploit. Therefore, wild brown bears that fatten primarily on fruits without access to salmon are 50% smaller than salmon-feeding bears. Thus, we predicted based on a linear, energy-maximizing model without dietary interaction effects that wild brown bears with access to both abundant salmon and fruit would feed almost exclusively on salmon. However, wild adult females with or without accompanying offspring foraged three times longer per day on fruit than on salmon. Similarly, the relative dry matter intake of ad libitum apples and salmon by captive, adult brown bears averaged 76±5% fruit and 24±5% salmon. Captive brown bears consuming mixed diets with intermediate dietary protein levels had 60% lower maintenance energy costs, 37% to 139% higher efficiencies of mass gain, and 72% to 520% higher maximum rates of gain than when they consumed either salmon or fruit alone. These relationships were nonlinear functions of dietary protein content in which salmon and fruit provided complementary nutritional resources. Both wild and captive bears attempted to regulate total protein, energy, and carbohydrate intake within a multidimensional intake target that both maximized energy intake and mass gain.     

Macronutrient Optimization and Seasonal Diet Mixing in a Large Omnivore,the Grizzly Bear: A Geometric Analysis

Nutrient balance is a strong determinant of animal fitness and demography. It is therefore important to understand how the compositions of available foods relate to required balance of nutrients and habitat suitability for animals in the wild. These relationships are, however, complex, particularly for omnivores that often need to compose balanced diets by combining their intake from diverse nutritionally complementary foods. Here we apply geometric models to understand how the nutritional compositions of foods available to an omnivorous member of the order Carnivora, the grizzly bear (Ursus arctos L.), relate to optimal macronutrient intake, and assess the seasonal nutritional constraints on the study population in west-central Alberta, Canada. The models examined the proportion of macronutrients that bears could consume by mixing their diet from food available in each season, and assessed the extent to which bears could consume the ratio of protein to non-protein energy previously demonstrated using captive bears to optimize mass gain. We found that non-selective feeding on ungulate carcasses provided a non-optimal macronutrient balance with surplus protein relative to fat and carbohydrate, reflecting adaptation to an omnivorous lifestyle, and that optimization through feeding selectively on different tissues of ungulate carcasses is unlikely. Bears were, however, able to dilute protein intake to an optimal ratio by mixing their otherwise high-protein diet with carbohydrate-rich fruit. Some individual food items were close to optimally balanced in protein to non-protein energy (e.g. Hedysarum alpinum roots), which may help explain their dietary prevalence. Ants may be consumed particularly as a source of lipids. Overall, our analysis showed that most food available to bears in the study area were high in protein relative to lipid or carbohydrate, suggesting the lack of non-protein energy limits the fitness (e.g. body size and reproduction) and population density of grizzly bears in this ecosystem.     

Nutritional ecology of a browsing ruminant, the kudu (Tragelaphus strepsiceros), through the seasonal cycle

The activities and food selection of four hand-reared kudus were recorded in a large fenced enclosure containing natural savanna vegetation in the Nylsvley Nature Reserve, South Africa. Leaves of selected species were analysed chemically for crude protein, fibre constituents, phosphorus, condensed tannin and total polyphenols. Available protein and metabolizable energy were estimated allowing for potential antinutritional effects of tannins.
Leaves of palatable deciduous woody plants and herbaceous forbs formed the main dietary constituents during the late wet season. Foliage from palatable evergreens and robust forbs were added to the diet during the dry season. Towards the end of the dry season unpalatable species of evergreens were eaten. At the start of the growing season new leaves of otherwise unpalatable woody species formed the staple food source, together with fruits of Strychnos spp. Correspondingly, protein and digestible dry matter concentrations in the diet declined to reach a low at the end of the dry season.
Total daily food intake increased to compensate for reduced dietary quality during the dry season, until little edible foliage remained. While the estimated daily intake of protein remained well above maintenance requirements, the estimated metabolizable energy intake fell below requirements during the late dry season. Phosphorus intake may have been submaintenance in the dry season. Nutritional balance was dependent on the availability of particular vegetation components to serve as nutritional stepping stones during crucial times of the year. These included forbs during the late wet season, palatable evergreens in the dry season, and Strychnos fruits plus early-flushing woody plants during the dry season-wet season transition.     

Dietary protein content alters energy expenditure and composition of the mass gain in grizzly bears (Ursus arctos horribilis)

Many fruits contain high levels of available energy but very low levels of protein and other nutrients. The discrepancy between available energy and protein creates a physiological paradox for many animals consuming high-fruit diets, as they will be protein deficient if they eat to meet their minimum energy requirement. We fed young grizzly bears both high-energy pelleted and fruit diets containing from 1.6% to 15.4% protein to examine the role of diet-induced thermogenesis and fat synthesis in dealing with high-energy-low-protein diets. Digestible energy intake at mass maintenance increased 2.1 times, and composition of the gain changed from primarily lean mass to entirely fat when the protein content of the diet decreased from 15.4% to 1.6%. Daily fat gain was up to three times higher in bears fed low-protein diets ad lib., compared with bears consuming the higher-protein diet and gaining mass at the same rate. Thus, bears eating fruit can either consume other foods to increase dietary protein content and reduce energy expenditure, intake, and potentially foraging time or overeat high-fruit diets and use diet-induced thermogenesis and fat synthesis to deal with their skewed energy-to-protein ratio. These are not discrete options but a continuum that creates numerous solutions for balancing energy expenditure, intake, foraging time, fat accumulation, and ultimately fitness, depending on food availability, foraging efficiency, bear size, and body condition.     

Grizzly Bear Behavior and Global Positioning System Collar Fix Rates

Abstract: Animal locations collected by Global Positioning System (GPS) collars will represent a biased sample of the sites an animal used if some position fixes fail and if those missed locations do not occur randomly. Probability of a GPS receiver obtaining a position fix is known to decline as canopy cover increases, but the impact of forest canopy cover was insufficient to account for the low fix rates we observed for GPS collars on grizzly bears (Ursus arctos). We tested the hypothesis that GPS fix rates were related to the interaction between animal activity (active vs. resting) and canopy cover by evaluating the following predictions: 1) grizzly bear activity should follow a circadian pattern similar to the circadian fix-rate pattern, 2) grizzly bear use of canopy cover should follow a circadian pattern similar to the circadian fix rates, 3) grizzly bear activity should be related to canopy cover (i.e., bears should rest in areas with relatively high canopy covers and feed and move in relatively open areas), and 4) collar orientation and canopy cover should interact to affect the fix rates of test collars. The GPS fix rates traced a bimodal circadian pattern that was directly related to the circadian pattern of grizzly bear activity. Fix rates declined when bears were more likely to be using denser cover, and fix rates of test collars demonstrated that collar orientation interacted with canopy cover, such that fix rates declined much more with increasing canopy cover when the collar was on its side than when the collar was upright. We concluded that inferences made about grizzly bear microhabitat use, based on GPS locations, will underrepresent high canopy cover sites, especially when grizzly bears are resting there. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):596–602; 2008)     

Nutritional Consequences of Experimentally Introduced Tourism in Brown Bears

Abstract: Although numerous studies have documented behavioral effects of nature-based tourism on wildlife populations, few studies have determined whether behavioral changes translate to effects on individual condition and population health. This issue is currently a concern for wildlife managers in Alaska, USA, and Canada where bear viewing is a rapidly growing industry expanding into previously undisturbed bear habitats. Rather than record observations at long established tourism sites, we experimentally introduced bear viewing into 2 relatively undisturbed brown bear (Ursus arctos) populations in south-central Alaska. We examined the nutritional consequences of behavioral changes induced by the presence and activity of bear viewers for bears feeding on early summer vegetation and late-summer salmon (Oncorhynchus kisutch and O. nerka). We used Global Positioning System collars, monitored food resource availability, and quantified individual resource use and condition for a year prior to and during the introduction of bear viewing. Though bear viewing altered spatiotemporal resource use in all treatments, total resource use declined only when we exposed bears to 24-hour daily human activity. Energy expenditure, indexed as daily travel distances, was significantly higher when bears responded by altering spatial rather than temporal resource use. However, body weight and composition were unaffected by all treatments as bears shifted their foraging to other locations or times. Managers can minimize nutritional impacts of bear-viewing programs by avoiding spatial displacement and providing predictable time periods when bears can access food resources free of human activity. Bears in this study exhibited a high degree of behavioral plasticity, which may be an important factor in identifying flagship species for sustainable ecotourism programs.     

Foraging strategy of cattle in patchy grassland

We tested several strategies of foraging that grazing herbivores may adopt in a patchy habitat in relation to energy intake. The patch selection of cattle was investigated in an Agrostis/Festuca grassland and in a Lolium grassland in 13 observation periods over 2 years. Both grasslands were stocked with five yearling steers. Bite counts were made on patches of different vegetation structure: short, tall and mature stemmy grass. Bite size of each patch category was determined by hand-plucking. Samples of patch types were analysed for organic matter digestibility, as a measure of energy content. There was a large seasonal variation in relative patch cover and in forage characteristics. However, the differences between patches in bite size, bite rate and digestibility were consistent over time. In short patches digestibility was high, bite size was low and bite rate was high compared to stemmy patches. In tall patches digestibility was only little lower than in short patches and bite size and bite rate were intermediate between short and stemmy patches. The steers selected the short and tall patches over the stemmy patches, despite a relatively low intake rate of digestible organic matter in the short patches. Four hypotheses on foraging strategy were examined to explain the allocation of time or bites between patches: random allocation according to bites, random allocation according to grazing time, matching of time in proportion to digestibility, and matching of time in proportion to intake rate of digestible organic matter. The observed distribution of bites and time between patches was significantly different from the predictions of the various hypotheses. Patch choice was better explained by a random allocation of grazing time than by a random allocation of bites. Matching for digestible organic matter intake rate yielded the worst predictions of patch selection. Matching for digestibility gave the best explanation of patch selection, but the improvement compared to a random allocation of grazing time was not significant. The significance of the contribution of digestibility to selection may have been confounded by the effect of increased selectivity within tall patches. Observed patch selection was considered in relation to the maximization of energy intake rate. The selectivity of cattle was not pronounced, but it was consistent with a principle of maximization of energy intake on a daily basis instead of a short-term basis. Selectivity appeared to be constrained by costs of searching for and discriminating between different forage resources. It is concluded that a flexible selection for short patches over tall patches and avoidance of stemmy patches provides a good approximation of energy intake maximization in a complex and changing environment.     

Plant Community Biomass Shifts in Response to Mowing and Fencing in Invaded Oak Meadows with Non‐Native Grasses and Abundant Ungulates

Many herbaceous meadows are dominated by competitive non‐native grasses and subject to ungulate herbivory, ecological processes that shift the proportional biomass of plant groups in the community. Predicting the outcome of restoration is complicated because herbivory and competition can interact. We examined the relationship between herbivory by native black‐tailed deer and domestic sheep and dominance of non‐native grasses in Garry oak meadows, one of North America's most endangered habitat types. A 3‐year factorial experiment tested the effects of mowing and fencing on plant community biomass, categorized into eight groups by geographic origin (native/non‐native), growth form (annual/perennial), and plant type (forb/grass). To test if the rarity of native plant groups was related to herbivory, we estimated ungulate foraging preferences for each plant group. Mowing and fencing treatments interacted for annual and perennial non‐native grasses. Dominance was shifted from non‐native to native grasses only when both mowing and fencing were applied. Fencing increased the total biomass, whereas mowing had no overall effect; however, fencing alone did not affect any individual plant group. Mowing shifted dominance from grasses to forbs, although both native and non‐native forbs benefited from the increased light availability. We also noted that herbivore fecal pellet densities were greatest in the spring, which coincided with the peak season of their preferred plant group, native perennial forbs. Overall, applying both mowing and fencing was the most effective restoration treatment to increase native plant groups and biomass.     

Forb ecology research in dry African savannas: Knowledge,gaps, and future perspectives

Savannas are commonly described as a vegetation type with a grass layer interspersed with a discontinuous tree or shrub layer. On the contrary, forbs, a plant life form that can include any nongraminoid herbaceous vascular plant, are poorly represented in definitions of savannas worldwide. While forbs have been acknowledged as a diverse component of the herbaceous layer in savanna ecosystems and valued for the ecosystem services and functions they provide, they have not been the primary focus in most savanna vegetation studies. We performed a systematic review of scientific literature to establish the extent to which forbs are implicitly or explicitly considered as a discrete vegetation component in savanna research. The overall aims were to summarize knowledge on forb ecology, identify knowledge gaps, and derive new perspectives for savanna research and management with a special focus on arid and semiarid ecosystems in Africa. We synthesize and discuss our findings in the context of different overarching research themes: (a) functional organization and spatial patterning, (b) land degradation and range management, (c) conservation and reserve management, (d) resource use and forage patterning, and (e) germination and recruitment. Our results revealed biases in published research with respect to study origin (country coverage in Africa), climate (more semiarid than arid systems), spatial scale (more local than landscape scale), the level at which responses or resource potential was analyzed (primarily plant functional groups rather than species), and the focus on interactions between life forms (rather seldom between forbs and grasses and/or trees). We conclude that the understanding of African savanna community responses to drivers of global environmental change requires knowledge beyond interactions between trees and grasses only and beyond the plant functional group level. Despite multifaceted evidence of our current understanding of forbs in dry savannas, there appear to be knowledge gaps, specifically in linking drivers of environmental change to forb community responses. We therefore propose that more attention be given to forbs as an additional ecologically important plant life form in the conventional tree–grass paradigm of savannas.     

Foraging behaviour at multiple temporal scales in a wild alpine equid

Forage abundance, forage quality, and social factors are key elements of the foraging ecology of wild herbivores. For non-ruminant equids, forage-limited environments are likely to impose severe constraints on their foraging behaviour. We used a multi-scale approach to study foraging behaviour in kiang (Equus kiang), a wild equid inhabiting the high-altitude rangelands of the Tibetan Plateau. Using behavioural observations and vegetation sampling, we first assessed how patterns of plant abundance and quality affected (i) the instantaneous forage intake rate (fine scale) and (ii) the proportion of time spent foraging (coarse scale) across seasons. We also tested whether foraging behaviour differed among group types, between sex in adults, and between females of different reproductive status. At a fine scale, intake rate increased linearly with bite size and increased following a type II curvilinear function with biomass on feeding sites. Forage intake rate also increased linearly with plant quality. Male and female kiangs had similar intake rates. Likewise, gravid and lactating females had similar intake rates as barren and non-lactating females. At a coarse scale, kiangs spent longer time feeding in mesic than in xeric habitats, and spent more time feeding in early summer and fall than in late summer. Groups of adults with foals spent less time feeding than male groups and groups of adults without foals. Our findings suggest that kiangs use flexible foraging behaviours in relation to seasonal variations of vegetation quality and abundance, a likely outcome of the extreme seasonal conditions encountered on the Tibetan Plateau.