Kevin L. Monteith

Timing of seasonal migration in mule deer: effects of climate, plant phenology, and life‐history characteristics

Phenological events of plants and animals are sensitive to climatic processes. Migration is a life-history event exhibited by most large herbivores living in seasonal environments, and is thought to occur in response to dynamics of forage and weather. Decisions regarding when to migrate, however, may be affected by differences in life-history characteristics of individuals. Long-term and intensive study of a population of mule deer (Odocoileus hemionus) in the Sierra Nevada, California, USA, allowed us to document patterns of migration during 11 years that encompassed a wide array of environmental conditions. We used two new techniques to properly account for interval-censored data and disentangle effects of broad-scale climate, local weather patterns, and plant phenology on seasonal patterns of migration, while incorporating effects of individual life-history characteristics. Timing of autumn migration varied substantially among individual deer, but was associated with the severity of winter weather, and in particular, snow depth and cold temperatures. Migratory responses to winter weather, however, were affected by age, nutritional condition, and summer residency of individual females. Old females and those in good nutritional condition risked encountering severe weather by delaying autumn migration, and were thus risk-prone with respect to the potential loss of foraging opportunities in deep snow compared with young females and those in poor nutritional condition. Females that summered on the west side of the crest of the Sierra Nevada delayed autumn migration relative to east-side females, which supports the influence of the local environment on timing of migration. In contrast, timing of spring migration was unrelated to individual life-history characteristics, was nearly twice as synchronous as autumn migration, differed among years, was related to the southern oscillation index, and was influenced by absolute snow depth and advancing phenology of plants. Plasticity in timing of migration in response to climatic conditions and plant phenology may be an adaptive behavioral strategy, which should reduce the detrimental effects of trophic mismatches between resources and other life-history events of large herbivores. Failure to consider effects of nutrition and other life-history traits may cloud interpretation of phenological patterns of mammals and conceal relationships associated with climate change.

GROWTH OF MALE WHITE-TAILED DEER: CONSEQUENCES OF MATERNAL EFFECTS

Abstract Identifying maternal effects on offspring is critical to interpreting population dynamics, but the duration of maternal effects and which life-history traits they influence is not well understood. We quantified growth and development of male white-tailed deer (Odocoileus virginianus) originating from the Black Hills in southwestern South Dakota and from eastern South Dakota in a controlled environment with high-quality nutrition. Despite being in good nutritional condition, males from the Black Hills ceased rapid growth 41 days earlier, were 29% smaller at asymptotic body mass, and grew significantly smaller antlers than males from eastern South Dakota. Females from eastern South Dakota were 14.9 kg larger than females from the Black Hills, yet birth mass of male offspring was similar for females from the 2 regions. Male offspring of 1st-generation deer from the Black Hills attained a 30% larger asymptotic body mass and grew significantly larger antlers than their sires. Body mass and antler size of 2nd-generation males of Black Hills origin approached that of 1st-generation males from eastern South Dakota at maturity. Suppression in growth of 1st-generation males of the Black Hills and increased growth by their offspring supported an influence of maternal and grandmaternal condition during gestation on subsequent growth of offspring and highlighted the significance of nutrition during gestation. These intergenerational effects indicate that measures of animal condition and population performance might reflect past rather than current conditions, and illustrate the potential for time lags in responses of populations to improved environmental conditions.

Behavior and nutritional condition buffer a large‐bodied endotherm against direct and indirect effects of climate

Temporal changes in net energy balance of animals strongly influence fitness; consequently, natural selection should favor behaviors that increase net energy balance by buffering individuals against negative effects of environmental variation. The relative importance of behavioral responses to climate-induced variation in costs vs. supplies of energy, however, is uncertain, as is the degree to which such responses are mediated by current stores of energy. We evaluated relationships among behavior, nutritional condition (i.e., energy state), and spatiotemporal variation in costs vs. supplies of energy available to a large- bodied endotherm, the North American elk (Cervus elaphus), occupying two ecosystems with contrasting climates and energy landscapes: a temperate, montane forest and an arid, high- elevation desert. We hypothesized that during spring through autumn, behavioral responses to the energy landscape would be both context dependent (i.e., would vary as a function of the contrasting environmental conditions experienced by elk in the forest vs. the desert), and state dependent (i.e., would vary as a function of the energy balance of an individual). We tested several predictions derived from that hypothesis by combining output from a biophysical model of the thermal environment with data on forage quality, animal locations, and nutritional condition of individuals. At the population level, elk in the desert selected areas that reduced costs of thermoregulation over those that provided the highest-quality forage. In the forest, however, costs imposed by the thermal environment were less pronounced, and elk selected areas that increased access to high-quality forage over those that reduced costs of thermoregulation. At the individual level, nutritional condition did not influence strength of selection for low-cost areas or high-quality forage among elk in the forest. In the desert, however, strength of selection for low-cost areas (but not forage quality) was state dependent; individuals in the poorest condition at the end of winter showed the strongest selection for areas that reduced costs of thermoregulation during spring and summer, and also expended the least amount of energy on locomotion. Our results highlight the importance of understanding the roles of behavior and nutritional condition in buffering endotherms against direct and indirect effects of climate on fitness.

Migrating Mule Deer: Effects of Anthropogenically Altered Landscapes

Background Migration is an adaptive strategy that enables animals to enhance resource availability and reduce risk of predation at a broad geographic scale. Ungulate migrations generally occur along traditional routes, many of which have been disrupted by anthropogenic disturbances. Spring migration in ungulates is of particular importance for conservation planning, because it is closely coupled with timing of parturition. The degree to which oil and gas development affects migratory patterns, and whether ungulate migration is sufficiently plastic to compensate for such changes, warrants additional study to better understand this critical conservation issue. Methodology/Principal Findings We studied timing and synchrony of departure from winter range and arrival to summer range of female mule deer (Odocoileus hemionus) in northwestern Colorado, USA, which has one of the largest natural-gas reserves currently under development in North America. We hypothesized that in addition to local weather, plant phenology, and individual life-history characteristics, patterns of spring migration would be modified by disturbances associated with natural-gas extraction. We captured 205 adult female mule deer, equipped them with GPS collars, and observed patterns of spring migration during 2008–2010. Conclusions/Significance Timing of spring migration was related to winter weather (particularly snow depth) and access to emerging vegetation, which varied among years, but was highly synchronous across study areas within years. Additionally, timing of migration was influenced by the collective effects of anthropogenic disturbance, rate of travel, distance traveled, and body condition of adult females. Rates of travel were more rapid over shorter migration distances in areas of high natural-gas development resulting in the delayed departure, but early arrival for females migrating in areas with high development compared with less-developed areas. Such shifts in behavior could have consequences for timing of arrival on birthing areas, especially where mule deer migrate over longer distances or for greater durations.

Large herbivores surf waves of green-up during spring

The green wave hypothesis (GWH) states that migrating animals should track or ‘surf’ high-quality forage at the leading edge of spring green-up. To index such high-quality forage, recent work proposed the instantaneous rate of green-up (IRG), i.e. rate of change in the normalized difference vegetation index over time. Despite this important advancement, no study has tested the assumption that herbivores select habitat patches at peak IRG. We evaluated this assumption using step selection functions parametrized with movement data during the green-up period from two populations each of bighorn sheep, mule deer, elk, moose and bison, totalling 463 individuals monitored 1–3 years from 2004 to 2014. Accounting for variables that typically influence habitat selection for each species, we found seven of 10 populations selected patches exhibiting high IRG—supporting the GWH. Nonetheless, large herbivores selected for the leading edge, trailing edge and crest of the IRG wave, indicating that other mechanisms (e.g. ruminant physiology) or measurement error inherent with satellite data affect selection for IRG. Our evaluation indicates that IRG is a useful tool for linking herbivore movement with plant phenology, paving the way for significant advancements in understanding how animals track resource quality that varies both spatially and temporally.

Supplemental feeding alters migration of a temperate ungulate

Conservation of migration requires information on behavior and environmental determinants. The spatial distribution of forage resources, which migration exploits, often are altered and may have subtle, unintended consequences. Supplemental feeding is a common management practice, particularly for ungulates in North America and Europe, and carryover effects on behavior of this anthropogenic manipulation of forage are expected in theory, but have received limited empirical evaluation, particularly regarding effects on migration. We used global positioning system (GPS) data to evaluate the influence of winter feeding on migration behavior of 219 adult female elk (Cervus elaphus) from 18 fed ranges and 4 unfed ranges in western Wyoming. Principal component analysis revealed that the migratory behavior of fed and unfed elk differed in distance migrated, and the timing of arrival to, duration on, and departure from summer range. Fed elk migrated 19.2 km less, spent 11 more days on stopover sites, arrived to summer range 5 days later, resided on summer range 26 fewer days, and departed in the autumn 10 days earlier than unfed elk. Time-to-event models indicated that differences in migratory behavior between fed and unfed elk were caused by altered sensitivity to the environmental drivers of migration. In spring, unfed elk migrated following plant green-up closely, whereas fed elk departed the feedground but lingered on transitional range, thereby delaying their arrival to summer range. In autumn, fed elk were more responsive to low temperatures and precipitation events, causing earlier departure from summer range than unfed elk. Overall, supplemental feeding disconnected migration by fed elk from spring green-up and decreased time spent on summer range, thereby reducing access to quality forage. Our findings suggest that ungulate migration can be substantially altered by changes to the spatial distribution of resources, including those of anthropogenic origin, and that management practices applied in one season may have unintended behavioral consequences in subsequent seasons.

Top-down versus bottom-up forcing: evidence from mountain lions and mule deer

Abstract We studied mountain lions (Puma concolor) and mule deer (Odocoileus hemionus) inhabiting a Great Basin ecosystem in Round Valley, California, to make inferences concerning predator–prey dynamics. Our purpose was to evaluate the relative role of top-down and bottom-up forcing on mule deer in this multiple-predator, multiple-prey system. We identified a period of decline (by 83%) of mule deer (1984–1990), and then a period of slow but steady increase (1991–1998). For mule deer, bitterbrush (Purshia tridentata) in diets, per capita availability of bitterbrush, kidney fat indexes, fetal rates (young per adult female), fetal weights, and survivorship of adults and young indicated that the period of decline was typical of a deer population near or above the carrying capacity (K) of its environment. Numbers of mountain lions also declined, but with a long time lag. The period of increase was typified by deer displaying life-history characteristics of a population below K, yet the finite rate of growth (λ = 1.10) remained below what would be expected for a population rebounding rapidly toward K (λ = 1.15–1.21) in the absence of limiting factors. Life-history characteristics were consistent with the mule deer population being regulated by bottom-up forcing through environmental effects on forage availability relative to population density; however, predation, mostly by mountain lions, was likely additive during the period of increase and thus, top-down forcing slowed but did not prevent population growth of mule deer. These outcomes indicate that resource availability (bottom-up processes) has an ever-present effect on dynamics of herbivore populations, but that the relationship can be altered by top-down effects. Indeed, top-down and bottom-up forces can act on populations simultaneously and, thus, should not be viewed as a stark dichotomy.

Effects of climate and plant phenology on recruitment of moose at the southern extent of their range

Climate plays a fundamental role in limiting the range of a species, is a key factor in the dynamics of large herbivores, and is thought to be involved in declines of moose populations in recent decades. We examined effects of climate and growing-season phenology on recruitment (8–9 months old) of young Shiras moose (Alces alces shirasi) over three decades, from 18 herds, across a large geographic area encompassing much of the southern extent of their range. Recruitment declined in 8 of 18 herds during 1980–2009, whereas others did not exhibit a temporal trend (none showed a positive trend). During those three decades, seasonal temperatures increased, spring–summer precipitation decreased, and spring occurred earlier, became shorter in duration, and green-up occurred faster. Recruitment was influenced negatively by warm temperatures during the year before young were born, but only for herds with declining recruitment. Dry spring–summers of the previous year and rapid rates of spring green-up in the year of birth had similar negative influences across declining and stable herds. Those patterns indicate both direct (yeart) and delayed (yeart−1) effects of weather and plant phenology on recruitment of young, which we hypothesize was mediated through effects on maternal nutritional condition. Suppressed nutrition could have been induced by (1) increased thermoregulatory costs associated with warming temperatures and (2) shortened duration of availability of high-quality forage in spring. Progressive reductions in net energetic gain for species that are sensitive to climate may continue to hamper individual fitness and population dynamics.

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.

Evaluation of Techniques for Categorizing Group Membership of White-Tailed Deer

Abstract We studied sexual segregation, particularly patterns of group membership for white-tailed deer (Odocoileus virginianus), in Lincoln County, Minnesota, USA, to evaluate current techniques used to categorize animals when studying sexual segregation. We categorized group membership according to Hirth (1977) and grouped individuals using our solitary categorization method. Our solitary method was most sensitive to changes in behavior and physiology exhibited by reproductively active females and their association with other deer during sexual segregation.