Redefining physiological responses of moose (Alces alces) to warm environmental conditions.

We tested the concept that moose (Alces alces) begin to show signs of thermal stress at ambient air temperatures as low as 14 °C. We determined the response of Alaskan female moose to environmental conditions from May through September by measuring core body temperature, heart rate, respiration rate, rate of heat loss from exhaled air, skin temperature, and fecal and salivary glucocorticoids. Seasonal and daily patterns in moose body temperature did not passively follow the same patterns as environmental variables. We used large changes in body temperature (≥1.25 °C in 24hr) to indicate days of physiological tolerance to thermal stressors. Thermal tolerance correlated with high ambient air temperatures from the prior day and with seasonal peaks in solar radiation (June), ambient air temperature and vapor pressure (July). At midday (12:00hr), moose exhibited daily minima of body temperature, heart rate and skin temperature (difference between the ear artery and pinna) that coincided with daily maxima in respiration rate and the rate of heat lost through respiration. Salivary cortisol measured in moose during the morning was positively related to the change in air temperature during the hour prior to sample collection, while fecal glucocorticoid levels increased with increasing solar radiation during the prior day. Our results suggest that free-ranging moose do not have a static threshold of ambient air temperature at which they become heat stressed during the warm season. In early summer, body temperature of moose is influenced by the interaction of ambient temperature during the prior day with the seasonal peak of solar radiation. In late summer, moose body temperature is influenced by the interaction between ambient temperature and vapor pressure. Thermal tolerance of moose depends on the intensity and duration of daily weather parameters and the ability of the animal to use physiological and behavioral responses to dissipate heat loads.

Variation in δ15 N and δ13 C values of forages for Arctic caribou: effects of location, phenology and simulated digestion.

RATIONALE: The use of stable isotopes for dietary estimates of wildlife assumes that there are consistent differences in isotopic ratios among diet items, and that the differences in these ratios between the diet item and the animal tissues (i.e., fractionation) are predictable. However, variation in isotopic ratios and fractionation of δ13 C and δ15 N values among locations, seasons, and forages are poorly described for arctic herbivores especially migratory species such as caribou (Rangifer tarandus). METHODS: We measured the δ13 C and δ15 N values of seven species of forage growing along a 200-km transect through the range of the Central Arctic caribou herd on the North Slope of Alaska over 2 years. We compared forages available at the beginning (May; n = 175) and the end (n = 157) of the growing season (September). Purified enzymes were used to measure N digestibility and to assess isotopic fractionation in response to nutrient digestibility during simulated digestion. RESULTS: Values for δ13 C declined by 1.38 ‰ with increasing latitude across the transect, and increased by 0.44 ‰ from the beginning to the end of the season. The range of values for δ15 N was greater than that for δ13 C (13.29 vs 5.60 ‰). Differences in values for δ13 C between graminoids (Eriophorum and Carex spp.) and shrubs (Betula and Salix spp.) were small but δ15 N values distinguished graminoids (1.87 ± 1.02 ‰) from shrubs (-2.87 ± 2.93 ‰) consistently across season and latitude. However, undigested residues of forages were enriched in 15 N when the digestibility of N was less than 0.67. CONCLUSIONS: Although δ15 N values can distinguish plant groups in the diet of arctic herbivores, variation in the digestibility of dietary items may need to be considered in applying fractionation values for 15 N to caribou and other herbivores that select highly digestible items (e.g. forbs) as well as heavily defended plants (e.g. woody browse). Published in 2017. This article is a U.S. Government work and is in the public domain in the USA.

Isotopic nitrogen in fecal fiber as an indicator of winter diet in caribou and muskoxen.

RATIONALE: The ratios of stable nitrogen isotopes (δ(15)N values) in excreta have been used to examine aspects of trophic and nutritional ecology across taxa. Nitrogen fractions in feces of herbivores include endogenous (e.g., sloughed intestinal cells, unresorbed digestive secretions, and microbial debris) and dietary sources. For animals such as large herbivores, that have diets and feces with high concentrations of indigestible fiber, endogenous (15)N may constrain the use of fecal δ(15)N values to estimate dietary δ(15)N values and reconstruct diets. METHODS: We compared two techniques (detergent and detergent-free) to isolate fractions of plant fibers in the forages of caribou (Rangifer tarandus) and muskoxen (Ovibos moschatus); estimated the discrimination factors between the δ(15)N values of fecal fiber residues and of the diets of captive animals; and used the more effective isotopic tracer of dietary δ(15)N values to examine the relationships between the δ(15)N values of fecal residues and diet composition in several populations of wild caribou and muskoxen throughout North America in winter. RESULTS: The detergent-based approach contaminated the fractions of plant fibers in forages and feces with (14)N, whereas the detergent-free method was a good proxy to estimate δ(15)N values of plant fibers (r(2) = 0.92) and provided a better estimate of the fecal-fiber to diet discrimination factor for both species (caribou = 3.6‰; muskoxen = 2.8‰). In wild populations, the δ(15)N values of fecal fibers reflected diet composition in muskoxen (adjusted R(2) = 0.43) but not caribou (adjusted R(2) = 0.06). CONCLUSIONS: Contamination from detergent residues prohibited the use of detergent extraction in isolating forage (15)N from endogenous (15)N in the feces of herbivores. Although δ(15)N values in fecal fibers can be used to track dietary δ(15)N values in wild herbivores, discrimination between fecal extracts and diet may vary with the contribution of endogenous nitrogen (N), and, therefore, residual endogenous (15)N in feces may limit dietary reconstructions from fecal δ(15)N values for some large herbivores.

Robust Responses of Female Caribou to Changes in Food Supply.

AbstractUngulates can respond to changes in food supply by altering foraging behavior, digestive function, and metabolism. A multifaceted response to an environmental change is considered robust. Short seasons of plant growth make herbivores sensitive to changes in food supply because maintenance and production must be accomplished in less time with fewer options in a more fragile response. Caribou live at high latitudes where short summers constrain their response to changes in food supply. We measured the ability of female caribou to resist and tolerate changes in the quality and quantity of their food supply during winter and summer. Caribou resisted changes in food abundance and quality by changing food intake and physical activity with changes in daily temperature within each season. Peak food intake rose by 134% from winter pregnancy to summer lactation (98 vs. 229 g kg-0.75 d-1), as digestible requirements to maintain the body increased by 85% for energy (1,164 vs. 2,155 kJ kg-0.75 d-1) and by 266% for N (0.79 vs. 2.89 g N kg-0.75 d-1). Caribou required a diet with a digestible content of 12 kJ g-1 and 0.8% N in pregnancy, 18 kJ g-1 and 1.9% N in early lactation, and 11 kJ g-1 and 1.2% N in late lactation, which corresponds with the phenology of the wild diet. Female caribou tolerated restriction of ad lib. food intake to 58% of their energy requirement (680 vs. 1,164 kJ kg-0.75 d-1) during winter pregnancy and to 84% of their energy requirement (1,814 vs. 2,155 kJ kg-0.75 d-1) during summer lactation without a change in stress level, as indicated by fecal corticosterone concentration. Conversely, caribou can respond to increased availability of food with a spare capacity to process digestible energy and N at 123% (2,642 vs. 2,155 kJ kg-0.75 d-1) and 145% (4.20 vs. 2.89 g N kg-0.75 d-1) of those respective requirements during lactation. Robust responses to changes in food supply allow caribou to sustain reproduction, which would buffer demographic response. However, herds may decline when thresholds of behavioral resistance and physiological tolerance are frequently exceeded. Therefore, the challenge for managing declining populations of caribou and other robust species is to identify declines in robustness before their response becomes fragile.

Estimating Mineral Requirements of Wild Herbivores: Modelling Arctic Caribou (Rangifer tarandus granti) in Summer.

Mineral requirements are poorly described for most wildlife. Consequently, the role of forage minerals in movement and productivity are poorly understood for sedentary and migratory ungulates, such as reindeer and caribou (Rangifer tarandus). We applied estimates of maintenance, lactation, body mass change, and antler growth to production curves (body mass, daily intake, and milk yield) for female caribou to calculate their mineral requirements over summer. The total requirements (mg or g·d-1) were divided by the daily intake (kg·d-1) to estimate the minimum concentration of minerals required in the diet (mg or g·kg-1) to balance demand. The daily requirements (mg·d-1) of all minerals increased from parturition to the end of summer. The minimum dietary concentrations (mg·kg-1) of macro-minerals (Ca, P, Mg, Na, K) declined as food intake (kg·d-1) increased over summer. The minimum dietary concentrations (Fe, Mn, Cu, Zn) were heavily influenced by body mass gain, which increased through late lactation even though food intakes rose. Our modeling framework can be applied to other wild ungulates to assess the impacts of changing forage phenology, plant community compositions, or environmental disturbances on movement and productivity.

Trapped between food, heat, and insects: Movement of moose (Alces alces) and exposure to flies in the boreal forest of Alaska.

Moose (Alces alces) in the boreal forest habitats of Alaska are unlike other northern ungulates because they tolerate high densities of flies (Diptera) even though flies cause wounds and infections during the warm summer months. Moose move to find food and to find relief from overheating (hyperthermia) but do they avoid flies? We used GPS collars to measure the rate of movement (m⋅h-1) and the time spent (min⋅day-1) by enclosed moose in four habitats: wetlands, black spruce, early seral boreal forest, and late seral boreal forest. Fly traps were used in each habitat to quantify spatio-temporal abundance. Average daily air temperatures increased into July when peak biomass of forage for moose was greatest in early seral boreal forest habitats (424.46 vs. 25.15 kg⋅ha-1 on average in the other habitats). Average daily air temperatures were 1.7°C cooler in black spruce than other habitats, but fly abundance was greatest in black spruce (approximately 4-fold greater on average than the other habitats). Moose increased their movement rate with counts of biting flies (mosquitoes, black flies, horse and deer flies), but not non-biting flies (coprophagous flies). However, as air temperature increased (above 14.7°C) moose spent more time in fly-abundant black spruce, than early seral boreal forest, showing great tolerance for mosquitoes. Warm summer temperatures appear to cause moose to trade-off foraging in fly-sparse habitats for resting and dissipating heat in shady, wet habitats with abundant flies that adversely affect the fitness of moose.

Nitrogen allocation to offspring and milk production in a capital breeder.

Nitrogen (N) is a limiting nutrient for many herbivores, especially when plant availability and N content are low during the period of maternal investment, which is common for arctic ungulates. We used natural abundance of N isotopes to quantify allocation of maternal nitrogen to neonatal calves and milk in wild migratory caribou (Rangifer tarandus). We contrasted female-calf pairs from two herds in northern Quebec/Labrador, Canada: Rivière-George herd (RG; low population size with heavy calves) and the Rivière-aux-Feuilles herd (RAF; high population size and small calves). We assessed whether females of both herds relied on body protein or dietary N to produce the neonatal calf and milk at calving and weaning. Female caribou of both herds relied mostly on body N for fetal development. RAF females allocated less body N to calves than did RG females (92% vs. 95% of calf N), which was consistent with the production of calves that were 8% smaller in RAF than in RG. Allocation of body N to milk was also high for both herds, similar at calving for RAF and RG females (88% vs. 91% of milk N, respectively), but lower in RAF than RG females (95% vs. 99% of milk N) at weaning, which was consistent with a small but significantly greater reliance on dietary N supplies to support milk production at weaning. Female caribou used body protein stores to ensure a constant supply of N for fetal growth and milk production that minimized the effects of trophic mismatches on reproduction. The combination of migration and capital investment may therefore allow females to produce calves and attenuate the effects of both temporal and spatial mismatches between vegetation green-up and calf growth, which ultimately would reduce trophic feedbacks on population growth. Our data suggest that small changes in maternal allocation of proteins over the long period of gestation produce significant changes in calf mass as females respond to changes in resources that accompany changes in the size and distribution of the population.

Hand-rearing wild caribou calves for studies of nutritional ecology.

Caribou (Rangifer tarandus) are considered difficult to hand-rear in captivity because they are sensitive to the composition and volume of formulated milks. We used a strict feeding schedule and a commercial milk formula designed specifically for caribou to bottle-feed neonates captured from five wild caribou herds in Alaska. Under a feeding protocol adjusted for age and mass, the growth rates and body mass of 26 hand-reared caribou calves to weaning were similar to those of three maternally nursed caribou. This protocol allows caretakers to hand-rear caribou that are as representative as possible of maternally raised neonates.

Sores of boreal moose reveal a previously unknown genetic lineage of parasitic nematode within the genus Onchocerca.

Long-standing reports of open sores on the hind legs of moose (Alces alces) have been recorded in Alaska (as well as Canada, Europe, and Michigan), eliciting concerns about causes and infection. We used histological and genomic methods to investigate the sores from 20 adult moose on the Kenai Peninsula, Alaska. We paired this with thermal imagery and molt scoring of adult moose to further describe sore formation and understand its timing. Severe, ulcerative and eosinophilic dermatitis was found in all moose with sores present, and microfilariae within intraepidermal pustules were additionally found in four samples. Genetic analysis of sores from moose revealed a previously unknown genetic lineage of Onchocerca. Adult moose molt and lose their barrier of protection against flies in June and July during peak fly activity, leaving them vulnerable and allowing the development of sores. In summary, our results indicate that the cause for the sores on the hindleg of moose is a previously unknown genetic lineage of Onchocerca, probably transmitted by black flies, in timing with the molt cycle of adult moose. These sores leave moose exposed to pathogens, making them vulnerable, and challenging their health and fitness.

Dynamic selection for forage quality and quantity in response to phenology and insects in an Arctic ungulate.

Spatiotemporal variation in forage is a primary driver of ungulate behavior, yet little is known about the nutritional components they select, and how selection varies across the growing season with changes in forage quality and quantity. We addressed these uncertainties in barren-ground caribou (Rangifer tarandus), which experience their most important foraging opportunities during the short Arctic summer. Recent declines in Arctic caribou populations have raised concerns about the influence of climate change on summer foraging opportunities, given shifting vegetation conditions and insect harassment, and their potential effects on caribou body condition and demography. We examined Arctic caribou selection of summer forage by pairing locations from females in the Central Arctic Herd of Alaska with spatiotemporal predictions of biomass, digestible nitrogen (DN), and digestible energy (DE). We then assessed selection for these nutritional components across the growing season at landscape and patch scales, and determined whether foraging opportunities were constrained by insect harassment. During early summer, at the landscape scale, caribou selected for intermediate biomass and high DN and DE, following expectations of the forage maturation hypothesis. At the patch scale, however, caribou selected for high values of all forage components, particularly DN, suggesting that protein may be limiting. During late summer, after DN declined below the threshold for protein gain, caribou exhibited a switch at both spatial scales, selecting for higher biomass, likely enabling mass and fat deposition. Mosquito activity strongly altered caribou selection of forage and increased their movement rates, while oestrid fly activity had little influence. Our results demonstrate that early and late summer periods afford Arctic caribou distinct foraging opportunities, as they prioritize quality earlier in the summer and quantity later. Climate change may further constrain caribou access to DN as earlier, warmer Arctic summers may be associated with reduced DN and increased mosquito harassment.

Behaviour influences thermoregulation of boreal moose during the warm season.

Management of large herbivores depends on providing habitats for forage supply and refuge from risks of temperature, predation and disease. Moose (Alces alces) accumulate body energy and nutrient stores during summer, while reducing the impact of warm temperatures through physiological and behavioural thermoregulation. Building on the animal indicator concept, we used rumen temperature sensors and GPS collars on captive moose (n = 6) kept in large natural enclosures to evaluate how behaviour and habitat selection influence the rate of change in rumen temperature during the growing season on the Kenai Peninsula, Alaska, USA. We compared movement and habitat selection of individual females during tolerance days (daily amplitude in rumen temperature was ≥1.2°C in 24 h) with those of control days (daily amplitude in rumen temperature was < 1.2°C) before and after the tolerance day. Moose moved more during tolerance days (172 m • h-1; 95% confidence intervals (CI) = 149-191 m • h-1) than on control days (151 m • h-1; 95% CI = 128-173 m • h-1). The rate of change in rumen temperature (°C • h-1) declined with low to moderate movement rates that were probably associated with foraging in all habitats. Movement only increased the rate of change in rumen temperature at high activity (~ > 500 m • h-1). Additionally, the relationship between rate of change in rumen temperature and movement rate was different during tolerance and control days in open meadow and wetland habitats. In all habitats except wetlands, the rate of change in rumen temperature increased while resting, which probably is a result of diet-induced thermogenesis. Our study demonstrates that the behavioural choices of moose on the landscape are associated with the rate of change in rumen temperature and their ability to thermoregulate. Wildlife managers must consider high-value habitats where wildlife can employ both behavioural and physiological mechanisms to tolerate warm ambient conditions in a landscape of forage, predators and pests.

Microbial degradation of usnic acid in the reindeer rumen.

Reindeer (Rangifer tarandus) eat and utilize lichens as an important source of energy and nutrients in winter. Lichens synthesize and accumulate a wide variety of phenolic secondary compounds, such as usnic acid, as a defense against herbivores and to protect against damage by UV-light in solar radiation. We have examined where and to what extent these phenolic compounds are degraded in the digestive tract of the reindeer, with particular focus on usnic acid. Three male reindeer were given ad libitum access to a control diet containing no usnic acid for three weeks and then fed lichens ad libitum (primarily Cladonia stellaris) containing 9.1 mg/g DM usnic acid for 4 weeks. Usnic acid intake in reindeer on the lichen diet was 91-117 mg/kg BM/day. In spite of this, no trace of usnic acid or conjugates of usnic acid was found either in fresh rumen fluid, urine, or feces. This suggests that usnic acid is rapidly degraded by rumen microbes, and that it consequently is not absorbed by the animal. This apparent ability to detoxify lichen phenolic compounds may gain increased importance with future enhanced UV-B radiation expected to cause increased protective usnic acid/phenol production in lichens.

Decadal heat and drought drive body size of North American bison (Bison bison) along the Great Plains.

Large grazers are visible and valuable indicators of the effects of projected changes in temperature and drought on grasslands. The grasslands of the Great Plains have supported the greatest number of bison (Bison bison; Linnaeus, 1758) since prehistoric times. We tested the hypothesis that body mass (BM, kg) and asymptotic body mass (ABM, kg) of Bison decline with rising temperature and increasing drought over both temporal and spatial scales along the Great Plains. Temporally, we modeled the relationship of annual measures of BM and height (H, m) of 5,781 Bison at Wind Cave National Park (WICA) from 1966 to 2015. We used Gompertz equations of BM against age to estimate ABM in decadal cohorts; both females and males decreased from the 1960s to the 2010s. Male ABM was variable but consistently larger (699 vs. 441 kg) than female ABM. We used local mean decadal temperature (MDT) and local mean decadal Palmer Drought Severity Index (dPDSI) to model the effects of climate on ABM. Drought decreased ABM temporally (-16 kg/local dPDSI) at WICA. Spatially, we used photogrammetry to measure body height (HE ) of 773 Bison to estimate BME in 19 herds from Saskatchewan to Texas, including WICA. Drought also decreased ABM spatially (-16 kg/local dPDSI) along the Great Plains. Temperature decreased ABM both temporally at WICA (-115 kg/°C local MDT) and spatially (-1 kg/°C local MDT) along the Great Plains. Our data indicate that temperature and drought drive Bison ABM presumably by affecting seasonal mass gain. Bison body size is likely to decline over the next five decades throughout the Great Plains due to projected increases in temperatures and both the frequency and intensity of drought.

Is summer food intake a limiting factor for boreal browsers? Diet, temperature, and reproduction as drivers of consumption in female moose.

Food intake may limit the ability of browsing mammals to gain body mass during the growing season when the leaves and stems of woody plants are most abundant. Moose are highly productive browsers with high demands for energy and nutrients, particularly during lactation. Using an indigestible marker, we estimated dry matter intake of free ranging adult female moose with and without calves over three growing seasons. During the same period, we analyzed forage quality. Intakes were highest in late spring (280 ± 19 g·kg-0.75·d-1) when forage quality peaked; however, intakes declined by 39% throughout the summer as temperatures increased and as acid detergent fiber content of browse increased. Digestibility of dry matter declined over summer from 71% to 57% among browse. Intakes were similar for moose with and without calves. Heat loads may impair the ability of moose to consume sufficient energy and nutrients. Warming and habitat change can adversely affect browser populations when poor forage qualities and low dry matter intakes combine to suppress digestible intakes of energy and nutrients.

Could a big gut be too costly for muskoxen (Ovibos moschatus) in their first winter?

Young mammalian herbivores are more vulnerable to harsh winter conditions than adults, especially among large circumpolar species like the muskox (Ovibos moschatus). We compared feeding responses of muskox calves (body mass 95 kg) with those of mature, non-reproductive females (body mass 227 kg) in mid-winter when air temperatures fell to -40 degrees C. Food intakes (1.8-2.2 kg digestible dry matter (DM)d(-1)), digesta fill (27-32 kg wet mass) and digestibility of hay (52-58% of DM; 49-55% of gross energy) were similar between age groups even though calves were much smaller than adults. Calves fed more frequently (12 vs. 8 feeding bouts per day) and thus spent more time feeding each day than adults (387 vs. 343 min per day). High mass-specific food intakes of calves indicate higher requirements for maintenance of body tissue than adults, which could be related to a larger intestinal tract in young muskoxen. Notably, cows and calves maintained a constant body mass throughout, indicating that they were feeding at maintenance levels and that the relatively higher intakes of calves were not related to growth. Together, these data suggest that limited food availability due to snow cover or high animal density may reduce the survival of muskoxen in their first winter.

Monitoring responses to variation in food supply for a migratory waterfowl: American Black Duck (Anas rubripes) in winter.

Wintering Black Ducks (Anas rubripes) concentrate in wetlands along the Atlantic coast where natural and anthropogenic disturbances have increased over the last 50 years, a period in which the population of Black Ducks has declined. We studied the sensitivity of Black Ducks to perturbations in food supply that often result from disturbances by storms, predators, and people. In the paper, we characterize the responses of captive Black Ducks to shifts in food quality and availability during winter and apply those measures to a comparison of wild birds. Captive ducks that were fed intermittently (3 consecutive days/week) compensated for fasted days to achieve similar body mass and body fat to control birds that were fed every day on both animal- and plant-based diets. However, birds that were fed intermittently expended 15% more energy each day than controls when both groups were fed (536 vs. 464 kJ/kg0.75), which indicates that variable food supply increases the costs of maintenance and thus reduces the number of birds that can be supported on the same resource of food without interruptions to foraging. Egg production was not affected by diet quality provided in spring or by the frequency of feeding during the preceding winter months. Black Ducks lost body fat through winter in captivity and in the wild. Fat stores of birds in New Jersey were greater than those of birds in Maine (13.3 vs. 8.3% of body mass) in January, which reflected the high energy demands of cold temperatures in Maine. Values for ∂15N were greater in Maine than in New Jersey for both red blood cells and plasma, which indicated a consistent diet of marine invertebrates in Maine. Greater isotopic variation in red blood cells indicated that diets were more diverse in New Jersey than in Maine for both ∂15N (9.7 ± 1.1 vs. 11.2 ± 0.4‰) and for ∂13C (- 15.1 ± 2.2 vs. - 13.8 ± 1.4‰). Plasma ∂13C was enriched over red blood cells in wild birds especially those with low fat stores, which suggested birds with low energy stores were shifting diets. Black Ducks can compensate for disturbances in feeding by increasing intakes if they have access to high quality wetlands where they are able to find abundant food. High energy demands at cold temperatures may constrain fat stores and thus the tolerance of feeding disturbances especially at the northern limits of the winter range. We hypothesize that decreasing variation in diet may indicate an increase in vulnerability to disturbance in winter when body fat is low. Recent efforts to assess and improve habitat quality of Black Ducks could be enhanced by monitoring the body composition and diet of birds to assess their vulnerability to disturbances in food supply and energy demands.

Bison body size and climate change.

The relationship between body size and temperature of mammals is poorly resolved, especially for large keystone species such as bison (Bison bison). Bison are well represented in the fossil record across North America, which provides an opportunity to relate body size to climate within a species. We measured the length of a leg bone (calcaneal tuber, DstL) in 849 specimens from 60 localities that were dated by stratigraphy and 14C decay. We estimated body mass (M) as M = (DstL/11.49)3. Average annual temperature was estimated from δ18O values in the ice cores from Greenland. Calcaneal tuber length of Bison declined over the last 40,000 years, that is, average body mass was 37% larger (910 ± 50 kg) than today (665 ± 21 kg). Average annual temperature has warmed by 6°C since the Last Glacial Maximum (~24-18 kya) and is predicted to further increase by 4°C by the end of the 21st century. If body size continues to linearly respond to global temperature, Bison body mass will likely decline by an additional 46%, to 357 ± 54 kg, with an increase of 4°C globally. The rate of mass loss is 41 ± 10 kg per°C increase in global temperature. Changes in body size of Bison may be a result of migration, disease, or human harvest but those effects are likely to be local and short-term and not likely to persist over the long time scale of the fossil record. The strong correspondence between body size of bison and air temperature is more likely the result of persistent effects on the ability to grow and the consequences of sustaining a large body mass in a warming environment. Continuing rises in global temperature will likely depress body sizes of bison, and perhaps other large grazers, without human intervention.

Influence of Season and Diet on Fiber Digestion and Bacterial Community Structure in the Rumen of Muskoxen (Ovibos moschatus).

We studied the relationship between fiber digestion and the composition of the bacterial community in the rumen of muskoxen at the start and the end of the annual window of plant growth from spring to fall. Eight ruminally cannulated castrated males were fed brome hay or triticale straw (69.6% vs. 84.6% neutral detergent fiber, respectively) that were similar in fiber content to the sedges consumed by wild muskoxen (64.5 to 71.7% neutral detergent fiber). Muskoxen digested fiber from both forages faster and to a greater extent when straw rather than hay was consumed. Fiber digestion was therefore inducible by diet 4 in each season. We used 16S rRNA sequences from ruminal contents to study how season and diet affected the bacterial community and how the latter related to fiber digestion. We found that Bacteroidetes and Firmicutes accounted for 90% of the sequences at the level of Phylum, which is typical for the mammal gut microbiome. Using partial least square regressions, it was found that between 48% and 72% of the variation in fiber digestion was associated with 36⁻43 genera of bacteria. The main fibrolytic bacteria typical of domestic ruminants were generally not among the most important bacteria associated with fiber digestion in muskoxen. This reveals that muskoxen rely upon on a large suite of bacterial genera that are largely distinct from those used by other ruminants to digest the cell walls of plants that vary widely in both abundance and nutritional quality through the year.

Body protein stores and isotopic indicators of N balance in female reindeer (Rangifer tarandus) during winter.

We studied bred and unbred female reindeer (Rangifer tarandus tarandus) during 12 wk of winter when ambient temperatures were low and nitrogen (N) demand for fetal growth is highest in pregnant females. Animals were fed a complete pelleted diet ad lib. that contained 2.54% N in dry matter that was 80% +/- 2% (X +/- SD) digestible. Female reindeer lost 64% +/- 14% of body fat but gained 34% +/- 11% of lean mass from 10 wk prepartum to parturition. These changes were equivalent to average balances of -14.14 +/- 2.35 MJ d(-1) and 10 +/- 3 g N d(-1). Blood cells, serum, and urine declined in (15)N/(14)N in late winter as body protein was gained from the diet. Blood cells of newborn calves were more enriched in (15)N and (13)C than that of their mothers, indicating the deposition of fetal protein from maternal stores. To quantify pathways of N flow in reindeer, N balance was measured by confining animals to cages for 10 d at 4 wk from parturition. N balance was inversely related to (15)N/(14)N in urea-N but not related to (15)N/(14)N of blood cells, creatinine, and feces. The proportion of urea-N derived from body protein increased above 0.46 as N balance fell below -200 mg N kg(-0.75) d(-1). Proportions of urea-N from body protein were -0.01 +/- 0.21 in pregnant females before and after caging and were consistent with average body protein gain in winter. Storage of protein allows reindeer and caribou to tolerate diets that are low in N without impairing fetal development.

Physiology of intermittent feeding: integrating responses of vertebrates to nutritional deficit and excess.

Food intakes of wild animals may not match their requirements for nutrients and energy but may vary between periods of nutritional excess (hyperphagia) and nutritional deficit (hypophagia) at timescales that vary from days to months. We present a simple model of feeding patterns and requirements of vertebrates. Frequent fasts and high intakes are typical of endothermic predators and migratory birds, whereas slow cycles and long deficits typify feeding patterns of ectothermic predators and ungulates in seasonal environments. We propose that hyperphagia is constrained by the ability to increase processes of digestion, absorption, intermediary metabolism, net deposition in tissue, and excretion to match loads of digesta and metabolites. Hyperphagia on high-quality diets is limited by the clearance of metabolites, whereas digestive tract capacity and flow limit consumption of low-quality diets. Of all digestive strategies, small omnivores with simple digestive systems may be the most tolerant of frequent hyperphagia. Tolerance of hypophagia favors large endogenous stores or low mass-specific rates of metabolism and reproductive output. Large animals may be most able to sustain reproduction during prolonged deficits in seasonal environments. Responses to excessive and deficient intakes of food are constrained by the length of the feeding cycle. Animals adapted to short feeding cycles may be best suited to unpredictable food supplies but at the energetic cost of maintaining spare capacity for digestion and absorption. Predictions of the response to food disruption are best evaluated in the context of body size, nutritional physiology, and life history of the species and the time for internal response.