Northern Yellowstone National Park Research Articles

A 2000-year record of fecal biomarkers reveals past herbivore presence and impacts in a catchment in northern Yellowstone National Park, USA.

Molecular biomarkers preserved in lake sediments are increasingly used to develop records of past organism occurrence. When linked with traditional paleoecological methods, analysis of molecular biomarkers can yield new insights into the roles of herbivores and other animals in long-term ecosystem dynamics. We sought to determine whether fecal steroids in lake sediments could be used to reconstruct past ungulate use and dominant taxa in a small catchment in northern Yellowstone National Park. To do so, we characterized the fecal steroid profiles of a selection of North American ungulates historically present in the Yellowstone region (bison, elk, moose, mule deer, and pronghorn) and compared them with those of sediments from a small lake in the Yellowstone Northern Range. Analysis of a set of fecal steroids from herbivore dung (Δ5-sterols, 5α-stanols, 5β-stanols, epi5β-stanols, stanones, and bile acids) differentiated moose, pronghorn, and mule deer, whereas bison and elk were partially differentiated. Our results show that bison and/or elk were the primary ungulates in the watershed over the past c. 2300 years. Fecal steroid influxes reached historically unprecedented levels during the early and middle 20th century, possibly indicating high local use by ungulates. Comparison of fecal steroid influxes with pollen and diatom data suggests that elevated ungulate presence may have contributed to decreased forage taxa (Poaceae, Artemisia, and Salix), relative to long-term averages, and possibly increased lake production. Our results reflect past change within a single watershed, and extending this approach to a network of sites could provide much-needed information on past herbivore communities, use, and environmental influences in Yellowstone National Park and elsewhere.

The primacy of density‐mediated indirect effects in a community of wolves, elk, and aspen

AbstractThe removal or addition of a predator in an ecosystem can trigger a trophic cascade, whereby the predator indirectly influences plants and/or abiotic processes via direct effects on its herbivore prey. A trophic cascade can operate through a density‐mediated indirect effect (DMIE), where the predator reduces herbivore density via predation, and/or through a trait‐mediated indirect effect (TMIE), where the predator induces an herbivore trait response that modifies the herbivore's effect on plants. Manipulative experiments suggest that TMIEs are an equivalent or more important driver of trophic cascades than are DMIEs. Whether this applies generally in nature is uncertain because few studies have directly compared the magnitudes of TMIEs and DMIEs on natural unmanipulated field patterns. A TMIE is often invoked to explain the textbook trophic cascade involving wolves (Canis lupus), elk (Cervus canadensis), and aspen (Populus tremuloides) in northern Yellowstone National Park. This hypothesis posits that wolves indirectly increase recruitment of young aspen into the overstory primarily through reduced elk browsing in response to spatial variation in wolf predation risk rather than through reduced elk population density. To test this hypothesis, we compared the effects of spatiotemporal variation in wolf predation risk and temporal variation in elk population density on unmanipulated patterns of browsing and recruitment of young aspen across 113 aspen stands over a 21‐year period (1999–2019) in northern Yellowstone National Park. Only 2 of 10 indices of wolf predation risk had statistically meaningful effects on browsing and recruitment of young aspen, and these effects were 8–28 times weaker than the effect of elk density. To the extent that temporal variation in elk density was attributable to wolf predation, our results suggest that the wolf–elk–aspen trophic cascade was primarily density‐mediated rather than trait‐mediated. This aligns with the alternative hypothesis that wolves and other actively hunting predators with broad habitat domains cause DMIEs to dominate whenever prey, such as elk, also have a broad habitat domain. For at least this type of predator–prey community, our study suggests that risk‐induced trait responses can be abstracted or ignored while still achieving an accurate understanding of trophic cascades.

Spatial variation in density of American black bears in northern Yellowstone National Park

AbstractThe quality and availability of resources are known to influence spatial patterns of animal density. In Yellowstone National Park, relationships between the availability of resources and the distribution of grizzly bears (Ursus arctos) have been explored but have yet to be examined in American black bears (Ursus americanus). We conducted non‐invasive genetic sampling during 2017–2018 (mid‐May to mid‐July) and applied spatially explicit capture‐recapture models to estimate density of black bears and examine associations with landscape features. In both years, density estimates were higher in forested vegetation communities, which provide food resources and thermal and security cover preferred by black bears, compared with non‐forested areas. In 2017, density also varied by sex, with female densities being higher than males. Based on our estimates, the northern range of Yellowstone National Park supports one of the highest densities of black bears (20 black bears/100 km2) in the northern Rocky Mountains (6–12 black bears/100 km2 in other regions). Given these high densities, black bears could influence other wildlife populations more than previously thought, such as through displacement of sympatric predators from kills. Our study provides the first spatially explicit estimates of density for black bears within an ecosystem that contains the majority of North America's large mammal species. Our density estimates provide a baseline that can be used for future research and management decisions of black bears, including efforts to reduce human–bear conflicts.

Bison alter the northern Yellowstone ecosystem by breaking aspen saplings.

The American bison (Bison bison) is a species that strongly interacts with its environment, yet the effects of this large herbivore on quaking aspen (Populus tremuloides) have received little study. We documented bison breaking the stems of aspen saplings (young aspen >2 m tall and ≤5 cm in diameter at breast height) and examined the extent of this effect in northern Yellowstone National Park (YNP). Low densities of Rocky Mountain elk (Cervus canadensis) after about 2004 created conditions conducive for new aspen recruitment in YNP's northern ungulate winter range (northern range). We sampled aspen saplings at local and landscape scales, using random sampling plots in 87 randomly selected aspen stands. Across the YNP northern range, we found that 18% of sapling stems had been broken. The causal attribution to bison was supported by multiple lines of evidence: (1) most broken saplings were in areas of high bison and low elk density; (2) saplings were broken in summer when elk were not foraging on them; (3) we directly observed bison breaking aspen saplings; and (4) mixed-effects modeling showed a positive association between scat density of bison and the proportion of saplings broken. In a stand heavily used by bison, most aspen saplings had been broken, and portions of the stand were cleared of saplings that were present in previous sampling in 2012. Bison numbers increased more than fourfold between 2004 and 2015, and their ecosystem effects have similarly increased, limiting and in some places reversing the nascent aspen recovery. This situation is further complicated by political constraints that prevent bison from dispersing to areas outside the park. Thus, one important conservation goal, the preservation of bison, is affecting another long-term conservation goal, the recovery of aspen and other deciduous woody species in northern Yellowstone.

Rub tree use and selection by American black bears and grizzly bears in northern Yellowstone National Park

Several of the world's bear species exhibit tree-rubbing behavior, which is thought to be a form of scent-marking communication. Many aspects of this behavior remain unexplored, including differences in rub tree selection between sympatric bear species. We compiled rub tree data collected on Yellowstone National Park's Northern Range (USA) and compared rub tree selection of sympatric American black bears (Ursus americanus) and grizzly bears (U. arctos) at local and landscape scales. During 2017 and 2018, we identified 217 rub trees and detected black bears at 117 rub trees and grizzly bears at 18 rub trees, based on genetic analysis of collected hair samples. Rub trees generally were located in areas with gentle slopes and close to existing animal trails. Trees selected by black bears were typically in forested areas, whereas trees selected by grizzly bears were in forested and more open areas. Use of rub trees varied seasonally and between sexes for black bears, but seasonal data were inconclusive for grizzly bears. Black bears showed preferences for certain tree species for rubbing, but we did not find evidence that rub tree selection by grizzly bears differed among tree species. Both bear species selected trees that lacked branches on the lower portions of tree trunks and the maximum rub height was consistent with the body length of the bear species that used the tree. Although the sample size for grizzly bears was small, identifying the species and sex of bears based on genetic analysis enhanced interpretation of rub tree use and selection by bears. Scent-marking by black bears and grizzly bears on similar rub objects in well-traversed areas likely serves to enhance communication within and between the 2 species.

A noninvasive and integrative approach for improving density and abundance estimates of moose

AbstractAcquiring demographic data for moose (Alces alces) can be difficult because they are solitary in nature, they prefer densely vegetated and mountainous habitats, and they often occur at low density. Such data, however, are essential for long‐term population monitoring, evaluating management practices, and effective conservation. Winter aerial surveys are the standard method for estimating moose population parameters, but they can be logistically challenging, expensive, and subject to sightability correction, which necessitates the capture of study animals for initial model development. Herein, we demonstrate a noninvasive alternative approach for estimating population parameters of moose in northern Yellowstone National Park, where aerial surveys were attempted but proved ineffective. We determined individual moose genotype and sex using microsatellite polymerase chain reaction amplification of DNA extracted from fecal pellets, integrated ancillary pellet sample data (i.e., metadata) in genotype analysis to aid in the identification of matching genotypes, and used spatially explicit capture‐recapture (SECR) modeling to estimate sex‐specific density and abundance. We collected 616 samples over 3 consecutive winters (Dec 2013–Apr 2016) and within 2 sampling occasions each winter. We recorded 514 captures of 142 individual moose (69 males, 73 females). Overall density ranged between 0.062 moose/km2 and 0.076 moose/km2 and averaged 0.034/km2 for females and 0.033/km2 for males. Abundance estimates were 150 moose in 2013 (female = 76, 95% CI = 55–105; male = 74, 95% CI = 54–103), 186 in 2014 (female = 95, 95% CI = 63–142; male = 91, 95% CI = 60–138), and 160 in 2015 (female = 79, 95% CI = 58–108; male = 81, 95% CI = 59–110). Average population sex ratio was 0.99 males/female. We demonstrate that SECR analysis of fecal DNA genotypes, using metadata in genotype analysis to help identify matching moose genotypes, is a promising alternative method for estimating sex‐specific density and abundance of a low‐density moose population in a mountainous and forested landscape.

Accounting for imperfect detection in observational studies: modeling wolf sightability in Yellowstone National Park

AbstractImperfect detection is ubiquitous among wildlife research and is therefore commonly included in abundance estimation. Yet, the factors that affect observation success are largely unknown for rare and elusive species, such as large carnivores. Here, we took advantage of intensive ground‐based monitoring effort and an extensive GPS data set (2000–2018) and developed a winter sightability model for gray wolves (Canis lupus) in northern Yellowstone National Park, Wyoming, USA. Our resulting sightability model indicated that observation success was positively affected by the topographic nature of where wolves were in relation to observer locations (viewshed), areas being less forested (openness), and wolf group size, and negatively affected by distance from observer locations. Of these, viewshed had the strongest effect on the probability of observing a wolf, with the odds of observing a wolf being four times more likely when wolves were in the predicted viewshed. Openness was the next most influential covariate, and group size was the least influential. We also tested whether a wolf being harvested from a pack when they were outside of Yellowstone National Park had an effect on wolf sightability. We did not, however, find support for human‐induced mortality affecting wolf sightability inside of Yellowstone National Park. Our results indicate that the ability to observe wolves was greatly affected by ecological and landscape‐level factors, a finding that is likely to generally extend to other large carnivores. As such, our sightability model highlights the importance of considering landscape structure and variation in large carnivore use of the landscape when conducting observational‐based studies.

How climate impacts the composition of wolf-killed elk in northern Yellowstone National Park.

While the functional response of predators is commonly measured, recent work has revealed that the age and sex composition of prey killed is often a better predictor of prey population dynamics because the reproductive value of adult females is usually higher than that of males or juveniles.Climate is often an important mediating factor in determining the composition of predator kills, but we currently lack a mechanistic understanding of how the multiple facets of climate interact with prey abundance and demography to influence the composition of predator kills.Over 20 winters, we monitored 17 wolf packs in Yellowstone National Park and recorded the sex, age and nutritional condition of kills of their dominant prey—elk—in both early and late winter periods when elk are in relatively good and relatively poor condition, respectively.Nutritional condition (as indicated by per cent marrow fat) of wolf‐killed elk varied markedly with summer plant productivity, snow water equivalent (SWE) and winter period. Moreover, marrow was poorer for wolf‐killed bulls and especially for calves than it was for cows.Wolf prey composition was influenced by a complex set of climatic and endogenous variables. In early winter, poor plant growth in either year t or t − 1, or relatively low elk abundance, increased the odds of wolves killing bulls relative to cows. Calves were most likely to get killed when elk abundance was high and when the forage productivity they experienced in utero was poor. In late winter, low SWE and a relatively large elk population increased the odds of wolves killing calves relative to cows, whereas low SWE and poor vegetation productivity 1 year prior together increased the likelihood of wolves killing a bull instead of a cow.Since climate has a strong influence on whether wolves prey on cows (who, depending on their age, are the key reproductive components of the population) or lower reproductive value of calves and bulls, our results suggest that climate can drive wolf predation to be more or less additive from year to year.

Bison limit ecosystem recovery in northern Yellowstone

American bison (Bison bison) numbers in northern Yellowstone National Park increased during the last two decades, while those of elk (Cervus canadensis) decreased. We undertook this study to assess the potential effects of bison on woody vegetation and channel morphology in the park's northern ungulate winter range. Based on differences in the number of elk and bison, body mass, and the amount of time each utilizes the northern range, foraging pressure from bison began to exceed that of elk in 2007 and is currently ~10 times greater than that of elk. In the Lamar Valley study area, stand structure data (i.e., tree frequency by diameter class) for aspen (Populus tremuloides) and cottonwood (P. spp.) indicated that the growth of seedling/sprouts of these species into tall saplings and trees has been extensively suppressed by bison herbivory. For streams that crossed the valley floor, woody riparian vegetation was absent along their banks and channels were relatively wide and deep, conditions sustained by high levels of bison use. Overall, our findings indicated that the elevated numbers of bison, via herbivory, trampling, and tree bark effects, are limiting the structure, composition, and distribution of woody plant communities in the Lamar Valley, affecting the character of the valley's stream and river channels, and, in turn, influencing habitat and food-web support for terrestrial and aquatic wildlife species. The conservation success of bison recovery now may be adversely affecting another conservation goal, the restoration and maintenance of woody riparian vegetation and riparian ecosystems.

Predation shapes the evolutionary traits of cervid weapons.

Sexually selected weapons evolved to maximize the individual reproductive success of males in many polygynous breeding species. Many weapons are also retained outside of reproductive periods for secondary reasons, but the importance of these secondary functions is poorly understood. Here we leveraged a unique opportunity from the predator-prey system in northern Yellowstone National Park, WY, USA to evaluate whether predation by a widespread, coursing predator (wolves) has influenced a specific weapon trait (antler retention time) in their primary cervid prey (elk). Male elk face a trade-off: individuals casting antlers early begin regrowth before other males, resulting in relatively larger antlers the following year, and thus greater reproductive success, as indicated by research with red deer. We show, however, that male elk that cast their antlers early are preferentially hunted and killed by wolves, despite early casters being in better nutritional condition than antlered individuals. Our results run counter to classic expectations of coursing predators preferring poorer-conditioned individuals, and in so doing, reveal an important secondary function for an exaggerated sexually selected weapon-predatory deterrence. We suggest this secondary function played a key evolutionary role in elk; uniquely among North American cervids, they retain their antlers long after they fulfil their primary role in reproduction.

Diel predator activity drives a dynamic landscape of fear

AbstractA “landscape of fear” (LOF) is a map that describes continuous spatial variation in an animal's perception of predation risk. The relief on this map reflects, for example, places that an animal avoids to minimize risk. Although the LOF concept is a potentially unifying theme in ecology that is often invoked to explain the ecological and conservation significance of fear, little is known about the daily dynamics of an LOF. Despite theory and data to the contrary, investigators often assume, implicitly or explicitly, that an LOF is a static consequence of a predator's mere presence within an ecosystem. We tested the prediction that an LOF in a large‐scale, free‐living system is a highly dynamic map with “peaks” and “valleys” that alternate across the diel (24‐h) cycle in response to daily lulls in predator activity. We did so with extensive data from the case study of Yellowstone elk (Cervus elaphus) and wolves (Canis lupus) that was the original basis for the LOF concept. We quantified the elk LOF, defined here as spatial allocation of time away from risky places and times, across nearly 1,000‐km2of northern Yellowstone National Park and found that it fluctuated with the crepuscular activity pattern of wolves, enabling elk to use risky places during wolf downtimes. This may help explain evidence that wolf predation risk has no effect on elk stress levels, body condition, pregnancy, or herbivory. The ability of free‐living animals to adaptively allocate habitat use across periods of high and low predator activity within the diel cycle is an underappreciated aspect of animal behavior that helps explain why strong antipredator responses may trigger weak ecological effects, and why an LOF may have less conceptual and practical importance than direct killing.

Factors affecting gray wolf (Canis lupus) encounter rate with elk (Cervus elaphus) in Yellowstone National Park

Despite encounter rates being a key component of kill rate, few studies of large carnivore predation have quantified encounter rates with prey, the factors that influence them, and the relationship between encounter rate and kill rate. The study’s primary motivation was to determine the relationship between prey density and encounter rate in understanding the mechanism behind the functional response. Elk (Cervus elaphus Linnaeus, 1758) population decline and variable weather in northern Yellowstone National Park provided an opportunity to examine how these factors influenced wolf (Canis lupus Linnaeus, 1758) encounter rates with elk. We explored how factors associated with wolf kill rate and encounter rate in other systems (season, elk density, elk group density, average elk group size, snow depth, wolf pack size, and territory size) influenced wolf–elk encounter rate in Yellowstone National Park. Elk density was the only factor significantly correlated with wolf–elk encounter rate, and we found a nonlinear density-dependent relationship that may be a mechanism for a functional response in this system. Encounter rate was correlated with number of elk killed during early winter but not late winter. Weak effects of snow depth and elk group size on encounter rate suggest that these factors influence kill rate via hunting success because kill rate is the product of hunting success and encounter rate.

Predator foraging response to a resurgent dangerous prey

SummaryPrey switching occurs when a generalist predator kills disproportionately more of an abundant prey species and correspondingly spares a rarer species. Although this behaviour is a classic stabilizing mechanism in food web models, little is known about its operation in free‐living systems which often include dangerous prey species that resist predation.We used long‐term (1995–2015) data from a large mammal system in northern Yellowstone National Park,USA, to understand how prey preference of a wild, generalist predator (Canis lupus) responds to a shift in prey species evenness involving rising numbers of dangerous prey (Bison bison) and dropping numbers of relatively safer prey (Cervus elaphus).Contrary to the prey switching hypothesis, wolves attacked and killed disproportionately more of the rarer, but safer, species. Wolves maintained a strong preference against bison even when this species was nearly twice as abundant as elk. [Correction added after online publication on 26 April 2017: ‘more than’ changed to ‘nearly’]. There was also evidence that wolves were increasingly averse to hunting bison as relative bison abundance increased.Wolves seldom hunted bison because capture success was limited to a narrow set of conditions: larger packs (>11 wolves) chasing smaller herds (10–20 bison) with calves. Wolves scavenged bison carrion instead and did so more frequently as bison abundance increased.Our study demonstrates the overarching importance of prey vulnerability to understanding the prey preferences of generalist predators in ecological communities with dangerous prey. The formidable defences of such prey diminish the potential for switching and its stabilizing influence on population dynamics. In these communities, shifts from hunting to scavenging are perhaps more likely than shifts in prey preference. The assumption of switching may therefore overestimate the stability of multi‐prey systems that include dangerous prey species.Alay summaryis available for this article.

Ungulate control of grassland production: grazing intensity and ungulate species composition in Yellowstone Park

AbstractKnowledge of the envelope of grazing conditions within which grassland plant and soil processes are sustainable is important for the ecologically sound management of grazed grassland. Here we examined how a recent shift from elk to bison dominance of the northern ungulate community in Yellowstone National Park (YNP), and an associated increase in the duration that grassland was grazed, affected grazing intensity and aboveground net primary production (ANPP). Mean grazing intensity (GI, percentage ANPP removed) and stimulation (grazed ANPP minus ungrazed [exclosed] ANPP) were compared among three studies, two when elk (1988–1989, 1999–2001) and one when bison (2012–2014) dominated the northern YNP ungulate community. We also manipulated GI with small exclosures established for different lengths of time after the start of the growing season to directly determine the effect of the combination of grazing duration (GD) and GI on stimulation at a dry grassland and a mesic grassland. GI was greater under a bison‐dominant grazing regime (49%) compared with that measured during the two earlier periods when elk were the dominant ungulate species (31%, 13%). Plotting stimulation on GI for sites sampled across all three studies revealed a unimodal relationship, with peak stimulation of 34% occurring at a GI of 40%. Manipulating GI indicated that the greater GI and longer GD of grazing under a bison‐dominant regime reduced stimulation at a mesic grassland and negated it completely at a dry grassland. These findings revealed that a shift in the grazing ungulate community composition and an associated change in the migratory behavior of the dominant species impacted YNP grassland processes. Results also showed that grassland ANPP was resilient to the relatively high rates of prolonged grazing by the bison‐dominant community and did not reduce ANPP below paired, ungrazed conditions. However, YNP grassland should be continued to be monitored if such high rates of herbivory continue.

Long-term aspen dynamics, trophic cascades, and climate in northern Yellowstone National Park

We report long-term patterns of quaking aspen (Populus tremuloides Michx.) recruitment for five ungulate exclosures in the northern ungulate winter range of Yellowstone National Park. Aspen recruitment was low (<3 aspen·ha−1·year−1) in the mid-1900s prior to exclosure construction due to herbivory by Rocky Mountain elk (Cervus elaphus Linnaeus, 1758) but increased more than 60-fold within 25 years after exclosure construction despite a drying climatic trend since 1940. Results support the hypothesis that long-term aspen decline in Yellowstone’s northern range during the latter half of the 20th century was caused by high levels of ungulate herbivory and not a drying climate. Gray wolves (Canis lupus Linnaeus, 1758) were reintroduced during 1995–1996. For the period 1995–2012, we summarized annual predator–prey ratios, ungulate biomass, and drought severity. The average density of young aspen increased from 4350 aspen·ha−1 in 1997–1998 to 8960 aspen·ha−1 in 2012; during the same time period, those >1 m in height increased over 30-fold (from 105 to 3194 aspen·ha−1). Increased heights of young aspen occurred primarily from 2007 to 2012, a period with relatively high predator–prey ratios, declining elk numbers, and decreasing browsing rates. Consistent with a re-established trophic cascade, aspen stands in Yellowstone’s northern range have increasingly begun to recover.

Trophic cascades from wolves to alders in Yellowstone

We explored possible interactions among gray wolves (Canis lupus), Rocky Mountain elk (Cervus elaphus), and thinleaf alder (Alnus incana spp. tenuifoli) in northern Yellowstone National Park. We developed an alder age structure based on annual growth rings for plants growing along six streams in areas accessible to ungulates on the northern range. Alder stems (n=412) along the six streams originated only after wolf reintroduction. By 2013, 80% of the sampled alders along these streams were taller than 2m, in contrast with a historical pattern of height suppression by ungulate herbivory. This pattern of alder recruitment is consistent with a trophic cascade whereby new alder growth occurred across all study streams within several years after wolf reintroduction. Although declines in elk density since wolf reintroduction likely contributed to the release of alder from herbivory, the immediate onset of new alder recruitment following wolf reintroduction indicates that behavioral responses to predation may also have been an important component in the resulting trophic cascade. These results suggest that predator conservation could play a role in the management and ecological restoration of riparian areas.

Recovering aspen follow changing elk dynamics in Yellowstone: evidence of a trophic cascade?

To investigate the extent and causes of recent quaking aspen (Populus tremuloides) recruitment in northern Yellowstone National Park, we measured browsing intensity and height of young aspen in 87 randomly selected aspen stands in 2012, and compared our results to similar data collected in 1997-1998. We also examined the relationship between aspen recovery and the distribution of Rocky Mountain elk (Cervus elaphus) and bison (Bison bison) on the Yellowstone northern ungulate winter range, using ungulate fecal pile densities and annual elk count data. In 1998, 90% of young aspen were browsed and none were taller-than 200 cm, the height at which aspen begin to escape from elk browsing. In 2012, only 37% in the east and 63% in the west portions of the winter range were browsed, and 65% of stands in the east had young aspen taller than 200 cm. Heights of young aspen were inversely related to browsing intensity, with the least browsing and greatest heights in the eastern portion of the range, corresponding with recent changes in elk density and distribution. In contrast with historical elk distribution (1930s-1990s), the greatest densities of elk recently (2006-2012) have been north of the park boundary (approximately 5 elk/km2), and in the western part of the range (2-4 elk/km2), with relatively few elk in the eastern portion of the range (<2 elk/km2), even in mild winters. This redistribution of elk and decrease in density inside the park, and overall reduction in elk numbers, explain why many aspen stands have begun to recover. Increased predation pressure following the reintroduction of gray wolves (Canis lupius) in 1995-1996 played a role in these changing elk population dynamics, interacting with other influences including increased predation by bears (Ursus spp.), competition with an expanding bison population, and shifting patterns of human land use and hunting outside the park. The resulting new aspen recruitment is evidence of a landscape-scale trophic cascade in which a resurgent large carnivore community, combined with other ecological changes, has benefited aspen through effects on ungulate prey.

After long-term decline, are aspen recovering in northern Yellowstone?

In northern Yellowstone National Park, quaking aspen (Populus tremuloides) stands were dying out in the late 20th century following decades of intensive browsing by Rocky Mountain elk (Cervus elaphus). In 1995–1996 gray wolves (Canis lupus) were reintroduced, joining bears (Ursus spp.) and cougars (Puma concolor) to complete the guild of large carnivores that prey on elk. This was followed by a marked decline in elk density and change in elk distribution during the years 1997–2012, due in part to increased predation. We hypothesized that these changes would result in less browsing and an increase in height of young aspen. In 2012, we sampled 87 randomly selected stands in northern Yellowstone, and compared our data to baseline measurements from 1997 and 1998. Browsing rates (the percentage of leaders browsed annually) in 1997–1998 were consistently high, averaging 88%, and only 1% of young aspen in sample plots were taller than 100cm; none were taller than 200cm. In 2012, browsing rates were much lower at 44%, and young aspen were taller on average with 34% >100cm and 5% >200cm. Most (62%) of the variation in height of young aspen in 2012 was explained by browsing intensity. Furthermore, in 2012, 25% of stands had at least five aspen saplings tall enough to escape elk browsing (⩾200cm spring height), a condition that has not occurred for decades and happened despite a recent drought. Sapling recruitment did not increase until browsing decreased, following substantial changes in elk density and distribution, and was not significantly related to stand productivity or climate fluctuations. These results suggest that large carnivore restoration, through effects on prey, may aid aspen recovery where aspen have been suppressed by elk.

A Non-Invasive Population Study of Moose in Northern Yellowstone National Park

North American moose (Alces alces) populations are declining across much of their southern distribution from the Canadian Maritimes to the Rocky Mountain range of the Shiras moose subspecies (Alces alces shirasi). Shiras moose population declines have been documented in Montana and Wyoming with reduced productivity reported from Utah and Colorado. These declines are due to a combination of factors including the natural succession, loss, and degradation of habitat; predation by wolves and bears; disease caused by infection from artery worm; and parasitism by moose ticks. The effects of heat stress may also contribute to chronic malnutrition and a reduction in female fertility. Significant reductions in Montana and Wyoming moose populations adjacent to Yellowstone National Park (YNP) are indicative of regional moose population declines and suggest that moose numbers may be decreasing in YNP as well. In the northern portion of YNP, also known as the Northern Range (NR), significant loss of riparian willow browse due to overgrazing by elk for decades and by bison more recently, the reduction of mature and old-growth conifer forests from the fires of 1988 and from disease more recently have reduced both winter habitat quality and quantity for moose. Several moose with cropped ears, an external sign of the disease Elaeophorosis, or artery worm, have been observed over the past few years on the NR suggesting that the disease may also be present in YNP. Northern Range estimates of moose have decreased from almost 400 in 1970 to possibly fewer than 100 today. Despite evidence suggesting moose decline in YNP, no current population data exists. Knowledge of population demographics serves as a critical baseline for evaluating and understanding factors leading to population declines. However, because moose are solitary, prefer densely vegetated habitats, and are present at low densities, collecting population data is challenging. Traditional methods of studying moose that require capture, radio-collaring, and aerial surveys are costly, sometimes produce unreliable results, can be harmful to the study animal, and are discouraged in some jurisdictions such as national parks. Non-invasive sampling, the collection of data without having to capture, handle, or in any manner physically restrain study animals, has proven to be a valuable tool for acquiring accurate population data from free-ranging ungulates when using traditional methods is neither feasible nor practical. In December 2013, we initiated a three-year non-invasive moose population study in YNP with the main objective to estimate population demographics of NR moose. For three consecutive winters we will be systematically collecting fecal pellets from the extent of NR moose wintering habitat. We are extracting DNA from epithelial cells on the pellet surface and through genetic testing will be able to identify individual moose and their genders. Female pellet samples will be analyzed for pregnancy hormone concentrations to make inferences on pregnancy rates. Because fecal pellet size is directly related to moose size, and therefore to moose age, we will use various pellet measurements to differentiate between age classes. These data will be used in capture-recapture modeling to estimate population abundance and vital rates.

Holocene seasonal variability inferred from multiple proxy records from Crevice Lake, Yellowstone National Park, USA

A 9400-yr-old record from Crevice Lake, a semi-closed alkaline lake in northern Yellowstone National Park, was analyzed for pollen, charcoal, geochemistry, mineralogy, diatoms, and stable isotopes to develop a nuanced understanding of Holocene environmental history in a region of northern Rocky Mountains that receives both summer and winter precipitation. The limited surface area, conical bathymetry, and deep water (>31m) of Crevice Lake create oxygen-deficient conditions in the hypolimnion and preserve annually laminated sediment (varves) for much of the record. Pollen data indicate that the watershed supported a closed Pinus-dominated forest and low fire frequency prior to 8200calyr BP, followed by open parkland until 2600calyr BP, and open mixed-conifer forest thereafter. Fire activity shifted from infrequent stand-replacing fires initially to frequent surface fires in the middle Holocene and stand-replacing events in recent centuries. Low values of δ18O suggest high winter precipitation in the early Holocene, followed by steadily drier conditions after 8500calyr BP. Carbonate-rich sediments before 5000calyr BP imply warmer summer conditions than after 5000calyr BP. High values of molybdenum (Mo), uranium (U), and sulfur (S) indicate anoxic bottom-waters before 8000calyr BP, between 4400 and 3900calyr BP, and after 2400calyr BP. The diatom record indicates extensive water-column mixing in spring and early summer through much of the Holocene, but a period between 2200 and 800calyr BP had strong summer stratification, phosphate limitation, and oxygen-deficient bottom waters. Together, the proxy data suggest wet winters, protracted springs, and warm effectively wet summers in the early Holocene and less snowpack, cool springs, warm dry summers in the middle Holocene. In the late Holocene, the region and lake experienced extreme changes in winter, spring, and summer conditions, with particularly short springs and dry summers and winters during the Roman Warm Period (~2000calyr BP) and Medieval Climate Anomaly (1200–800calyr BP). Long springs and mild summers occurred during the Little Ice Age, and these conditions persist to the present. Although the proxy data indicate effectively wet summer conditions in the early Holocene and drier conditions in the middle and late Holocene, none point specifically to changes in summer precipitation as the cause. Instead, summer conditions were governed by multi-seasonal controls on effective moisture that operated over multiple time scales.