Shifts in risk sensitivity and resource availability alter fat stores for a large mammal following extreme winter conditions

For species that inhabit environments where resource availability may be unpredictable, balance of resource allocation to life-history traits can have heightened consequences for survival, reproduction, and ultimately, fitness. Acquisition and allocation of energy to maintenance, capital gain and reproduction should be in tune with the landscape an animal inhabits—environmental severity, food availability and population size all influence the resources animals have and dictate the ways they should be allocated. In seasonal environments, animals that experience periods of extreme resource limitation (e.g. harsh winters) may favour allocation of resources to body reserves to secure their survival at the cost of reproduction (i.e. risk averse). In contrast, the same accumulation of body reserves may not be necessary to survive in relatively benign landscapes where instead, allocation to reproduction is favoured (i.e. risk prone). According to the theory of risk-sensitive allocation of resources, when animals are exposed to unprecedented or life-threatening conditions, they may shift resource allocation to favour building capital over allocation in reproduction to preempt against encountering another life-threatening event in the future. Using data from a long-term project on a highly site-faithful and long-lived species, mule deer (Odocoileus hemionus), we evaluated how a life-threatening winter and the associated changes in resource availability resulting from a population reduction influenced how animals acquired and allocated energy to survival (i.e. fat accumulation). Per capita precipitation, and the associated reduction in population abundance after the severe winter, had a positive influence of accrual of fat over summer. After the extreme physiological stress of a hard winter, deer starting spring with low body reserves accumulated 2.8 percentage points more fat over summer compared with before the experience of a bad winter and had an increased probability of recruiting fewer offspring. Fat stores can interact with environment, life history and behaviour to influence survival during periods of resource scarcity. For a long-lived herbivore, we documented shifts in risk tolerance associated with fat accrual in preparation for winter, supporting the notion that risk-sensitive allocation of resources may be plastic—an essential adaptation for animals to cope with rapidly changing landscapes.publishedVersio

Drivers of site fidelity in ungulates

1. While the tendency to return to previously visited locations—termed ‘site fidelity’—is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals’ recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity. 2. We compared inter‐year site fidelity in 669 individuals across eight ungulate species fitted with GPS collars and occupying a range of environmental conditions in North America and Africa. We used a distance‐based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size. 3. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren‐ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a ‘win‐stay, lose‐switch’ strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested. 4. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter‐annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species‐specific differences in attraction to known sites, contribute to variation in the expression of this behaviour. 5. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change

Data from: Migratory plasticity is not ubiquitous among large herbivores

1. The migratory movements of wild animals can promote abundance and support ecosystem functioning. For large herbivores, mounting evidence suggests that migratory behavior is an individually variable trait, where individuals can easily switch between migrant and resident tactics. The degree of migratory plasticity, including whether and where to migrate, has important implications for the ecology and conservation of large herbivores in a changing world. 2. Mule deer (Odocoileus hemionus) are an iconic species of western North America, but are notably absent from the body of literature that suggests large herbivore migrations are highly plastic. We evaluated plasticity of migration in female mule deer using longitudinal GPS data collected from 312 individuals across 9 populations in the western US, including 882 animal-years (801 migrants and 81 residents). 3. We followed both resident and migratory mule deer through time to determine if individual animals switched migratory behaviors (i.e., whether to migrate) from migratory to residency, or vice versa. Additionally, we examined the fidelity of individuals to their migration routes (i.e., where to migrate) to determine if they used the same routes year after year. We also evaluated if age and reproductive status affected propensity to migrate or fidelity to migratory routes. 4. Our results indicate that mule deer, unlike other large herbivores, have little or no plasticity in terms of whether or where they migrate. Resident deer remained residents, and migrant deer remained migrants, regardless of age, reproductive status, or number of years monitored. Further, migratory individuals showed strong fidelity (>80%) to their migration routes year after year. 5. Our study clearly shows that migration plasticity is not ubiquitous among large herbivores. Because of their rigid migratory behavior, mule deer may not adapt to changing environmental conditions as readily as large herbivores with more plastic migratory behavior (e.g., elk). The fixed migratory behaviors of mule deer make clear that conservation efforts aimed at traditional seasonal ranges and migration routes are warranted for sustaining this iconic species that continues to decline across its range

Data from: The greenscape shapes surfing of resource waves in a large migratory herbivore

The Green Wave Hypothesis posits that herbivore migration manifests in response to waves of spring green-up (i.e. green-wave surfing). Nonetheless, empirical support for the Green Wave Hypothesis is mixed, and a framework for understanding variation in surfing is lacking. In a population of migratory mule deer (Odocoileus hemionus), 31% surfed plant phenology in spring as well as a theoretically perfect surfer, and 98% surfed better than random. Green-wave surfing varied among individuals and was unrelated to age or energetic state. Instead, the greenscape, which we define as the order, rate and duration of green-up along migratory routes, was the primary factor influencing surfing. Our results indicate that migratory routes are more than a link between seasonal ranges, and they provide an important, but often overlooked, foraging habitat. In addition, the spatiotemporal configuration of forage resources that propagate along migratory routes shape animal movement and presumably, energy gains during migration

Migration distance and maternal resource allocation determine timing of birth in a large herbivore

Birth timing is a key life-history characteristic that influences fitness and population performance. For migratory animals, however, appropriately timing birth on one seasonal range may be constrained by events occurring during other parts of the migratory cycle. We investigated how the use of capital and income resources may facilitate flexibility in reproductive phenology of migratory mule deer in western Wyoming, USA, over a five-year period (2015-2019). Specifically, we examined how seasonal interactions affected three, interrelated life-history characteristics: fetal development, birth mass and birth timing. Females in good nutritional condition at the onset of winter and those that migrated short distances had more developed fetuses (measured as fetal eye diameter in March). Variation in parturition date was explained largely by fetal development, however, there was up to 16 days of plasticity in expected birth date. Plasticity in expected birth date was shaped by income resources in the form of exposure to spring green-up. Although individuals that experienced greater exposure to spring green-up were able to advance expected birth date, being born early or late with respect to fetal development had no effect on birth mass of offspring. Furthermore, we investigated the trade-offs migrating mule deer face by evaluating support for existing theory which predicts that births should be matched to local peaks in resource availability at the birth site. In contrast to this prediction, only long-distance migrants that paced migration with the flush of spring green-up, giving birth shortly after ending migration, were able to match birth with spring green-up. Shorter distance migrants completed migration sooner and gave birth earlier, seemingly trading off more time for offspring to grow and develop over greater access to resources. Thus, movement tactic had profound downstream effects on birth timing. These findings highlight a need to reconsider classical theory on optimal birth timing, which has focused solely on conditions at the birth site.publishe

Drivers of site fidelity in ungulates

1. While the tendency to return to previously visited locations - termed 'site fidelity' - is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals' recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity.2. We compared inter-year site fidelity in 669 individuals across eight ungulate species fitted with GPS-collars and occupying a range of environmental conditions in North America and Africa. We used a distance-based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size.3. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren-ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a 'win-stay, lose-switch' strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested.4. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter-annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species-specific differences in attraction to known sites, contribute to variation in the expression of this behaviour.5. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change.publishe

Biomarkers of Animal Nutrition: From Seasonal to Lifetime Indicators of Environmental Conditions

Nutrition underpins survival and reproduction in animal populations; reliable nutritional biomarkers are therefore requisites to understanding environmental drivers of population dynamics. Biomarkers vary in scope of inference and sensitivity, making it important to know what and when to measure to properly quantify biological responses. We evaluated the repeatability of three nutritional biomarkers in a large, iteroparous mammal to evaluate the level of intrinsic and extrinsic contributions to those traits. During a long-term, individual-based study in a highly variable environment, we measured body fat, body mass, and lean mass of mule deer (Odocoileus hemionus) each autumn and spring. Lean mass was the most repeatable biomarker (0.72 autumn; 0.61 spring), followed by body mass (0.64 autumn; 0.53 spring), and then body fat (0.22 autumn; 0.01 spring). High repeatability in body and lean mass likely reflects primary structural composition, which is conserved across seasons. Low repeatability of body fat supports that it is the primary labile source of energy that is largely a product of environmental contributions of the previous season. Based on the disparate levels in repeatability among nutritional biomarkers, we contend that body and lean mass are better indicators of nutritional legacies (e.g., maternal effects), whereas body fat is a direct and sensitive reflection of recent nutritional gains and losses