Geochemical landscapes as drivers of wildlife reproductive success: Insights from a high-Arctic ecosystem.

The bioavailability of essential and non-essential elements in vegetation is expected to influence the performance of free-ranging terrestrial herbivores. However, attempts to relate the use of geochemical landscapes by animal populations directly to reproductive output are currently lacking. Here we measured concentrations of 14 essential and non-essential elements in soil and vegetation samples collected in the Zackenberg valley, northeast Greenland, and linked these to environmental conditions to spatially predict and map geochemical landscapes. We then used long-term (1996-2021) survey data of muskoxen (Ovibos moschatus) to quantify annual variation in the relative use of essential and non-essential elements in vegetated sites and their relationship to calf recruitment the following year. Results showed that the relative use of the geochemical landscape by muskoxen varied substantially between years and differed among elements. Selection for vegetated sites with higher levels of the essential elements N, Cu, Se, and Mo was positively linked to annual calf recruitment. In contrast, selection for vegetated sites with higher concentrations of the non-essential elements As and Pb was negatively correlated to annual calf recruitment. Based on the concentrations measured in our study, we found no apparent associations between annual calf recruitment and levels of C, Mn, Co, Zn, Cd, Ba, Hg, and C:N ratio in the vegetation. We conclude that the spatial distribution and access to essential and non-essential elements are important drivers of reproductive output in muskoxen, which may also apply to other wildlife populations. The value of geochemical landscapes to assess habitat-performance relationships is likely to increase under future environmental change.

On the interplay between hypothermia and reproduction in a high arctic ungulate.

For free-ranging animals living in seasonal environments, hypometabolism (lowered metabolic rate) and hypothermia (lowered body temperature) can be effective physiological strategies to conserve energy when forage resources are low. To what extent such strategies are adopted by large mammals living under extreme conditions, as those encountered in the high Arctic, is largely unknown, especially for species where the gestation period overlaps with the period of lowest resource availability (i.e. winter). Here we investigated for the first time the level to which high arctic muskoxen (Ovibos moschatus) adopt hypothermia and tested the hypothesis that individual plasticity in the use of hypothermia depends on reproductive status. We measured core body temperature over most of the gestation period in both free-ranging muskox females in Greenland and captive female muskoxen in Alaska. We found divergent overwintering strategies according to reproductive status, where pregnant females maintained stable body temperatures during winter, while non-pregnant females exhibited a temporary decrease in their winter body temperature. These results show that muskox females use hypothermia during periods of resource scarcity, but also that the use of this strategy may be limited to non-reproducing females. Our findings suggest a trade-off between metabolically-driven energy conservation during winter and sustaining foetal growth, which may also apply to other large herbivores living in highly seasonal environments elsewhere.

IMMOBILIZING MUSKOX ( OVIBOS MOSCHATUS) UNDER HIGH ARCTIC CONDITIONS.

Immobilizing and handling large, free-ranging animals without proper facilities in harsh environmental conditions poses significant challenges. During two field expeditions, a total of 29 female muskoxen ( Ovibos moschatus) were immobilized in Northeast Greenland (74°N, 20°E). Fixed doses of immobilizing drugs were used regardless of animal size: 2 mg etorphine, 30 mg xylazine, 0.3 mg medetomidine, and 40 mg ketamine. Physiologic and behavioral monitoring was performed during the second expedition on 15 female muskoxen. The observed heart rates were 35-58 beats/min and respiratory rates were 25-30 breaths/min. Mean arterial pressures measured using oscillometry ranged between 117-142 mmHg. Pulse oximeter readings ranged from 91-98% with oxygen supplementation, nasal end-tidal carbon dioxide values were 24-42 mmHg, and rectal temperature ranged from 38.9-39.6°C. Induction time was 6-8 min, recovery time 2-6 min after reversal, and duration of anesthesia was 50-100 min. This anesthetic regime thus provided reliable immobilization with minimal pathophysiologic alterations.

A Comparative Study on the Faecal Bacterial Community and Potential Zoonotic Bacteria of Muskoxen (Ovibos moschatus) in Northeast Greenland, Northwest Greenland and Norway.

Muskoxen (Ovibos moschatus) are ruminants adapted to a high-fibre diet. There is increasing interest in the role that gut microbes play in the digestion and utilization of these specialized diets but only limited data available on the gut microbiome of high-Arctic animals. In this study, we metabarcoded the 16S rRNA region of faecal samples from muskoxen of Northeast Greenland, Northwest Greenland and Norway, and quantified the effects of physiological and temporal factors on bacterial composition. We found significant effects of body mass, year of sampling and location on the gut bacterial communities of North East Greenland muskoxen. These effects were however dwarfed by the effects of location, emphasizing the importance of the local ecology on the gut bacterial community. Habitat alterations and rising temperatures may therefore have a considerable impact on muskoxen health and reproductive success. Moreover, muskoxen are hunted and consumed in Greenland, Canada and Alaska; therefore, this study also screened for potential zoonoses of food safety interest. A total of 13 potentially zoonotic genera were identified, including the genera Erysipelothrix and Yersinia implicated in recent mass die-offs of the muskoxen themselves.

High resistance towards herbivore-induced habitat change in a high Arctic arthropod community.

Mammal herbivores may exert strong impacts on plant communities, and are often key drivers of vegetation composition and diversity. We tested whether such mammal-induced changes to a high Arctic plant community are reflected in the structure of other trophic levels. Specifically, we tested whether substantial vegetation changes following the experimental exclusion of muskoxen (Ovibos moschatus) altered the composition of the arthropod community and the predator-prey interactions therein. Overall, we found no impact of muskox exclusion on the arthropod community: the diversity and abundance of both arthropod predators (spiders) and of their prey were unaffected by muskox presence, and so was the qualitative and quantitative structure of predator-prey interactions. Hence, high Arctic arthropod communities seem highly resistant towards even large biotic changes in their habitat, which we attribute to the high connectance in the food web.

Limited dietary overlap amongst resident Arctic herbivores in winter: complementary insights from complementary methods.

Snow may prevent Arctic herbivores from accessing their forage in winter, forcing them to aggregate in the few patches with limited snow. In High Arctic Greenland, Arctic hare and rock ptarmigan often forage in muskox feeding craters. We therefore hypothesized that due to limited availability of forage, the dietary niches of these resident herbivores overlap considerably, and that the overlap increases as winter progresses. To test this, we analyzed fecal samples collected in early and late winter. We used molecular analysis to identify the plant taxa consumed, and stable isotope ratios of carbon and nitrogen to quantify the dietary niche breadth and dietary overlap. The plant taxa found indicated only limited dietary differentiation between the herbivores. As expected, dietary niches exhibited a strong contraction from early to late winter, especially for rock ptarmigan. This may indicate increasing reliance on particular plant resources as winter progresses. In early winter, the diet of rock ptarmigan overlapped slightly with that of muskox and Arctic hare. Contrary to our expectations, no inter-specific dietary niche overlap was observed in late winter. This overall pattern was specifically revealed by combined analysis of molecular data and stable isotope contents. Hence, despite foraging in the same areas and generally feeding on the same plant taxa, the quantitative dietary overlap between the three herbivores was limited. This may be attributable to species-specific consumption rates of plant taxa. Yet, Arctic hare and rock ptarmigan may benefit from muskox opening up the snow pack, thereby allowing them to access the plants.

Interaction webs in arctic ecosystems: Determinants of arctic change?

How species interact modulate their dynamics, their response to environmental change, and ultimately the functioning and stability of entire communities. Work conducted at Zackenberg, Northeast Greenland, has changed our view on how networks of arctic biotic interactions are structured, how they vary in time, and how they are changing with current environmental change: firstly, the high arctic interaction webs are much more complex than previously envisaged, and with a structure mainly dictated by its arthropod component. Secondly, the dynamics of species within these webs reflect changes in environmental conditions. Thirdly, biotic interactions within a trophic level may affect other trophic levels, in some cases ultimately affecting land-atmosphere feedbacks. Finally, differential responses to environmental change may decouple interacting species. These insights form Zackenberg emphasize that the combination of long-term, ecosystem-based monitoring, and targeted research projects offers the most fruitful basis for understanding and predicting the future of arctic ecosystems.

Strong impact of temporal resolution on the structure of an ecological network.

Most ecological networks are analysed as static structures, where all observed species and links are present simultaneously. However, this is over-simplified, because networks are temporally dynamical. We resolved an arctic, entire-season plant-flower visitor network into a temporal series of 1-day networks and compared the properties with its static equivalent based on data pooled over the entire season. Several properties differed. The nested link pattern in the static network was blurred in the dynamical version, because the characteristic long nestedness tail of flower-visitor specialists got stunted in the dynamical networks. This tail comprised a small food web of pollinators, parasitoids and hyper-parasitoids. The dynamical network had strong time delays in the transmission of direct and indirect effects among species. Twenty percent of all indirect links were impossible in the dynamical network. Consequently, properties and thus also robustness of ecological networks cannot be deduced from the static topology alone.