Geology topics

Daniel R. MacNulty

Publications and source records attributed to Daniel R. MacNulty.

16 recordsLinked to original sources

USGS2024

Intrinsic and environmental drivers of pairwise cohesion in wild Canis social groups

Animals within social groups respond to costs and benefits of sociality by adjusting the proportion of time they spend in close proximity to other individuals in the group (cohesion). Variation in cohesion between individuals, in turn, shapes important group-level processes such as subgroup formation and fission–fusion dynamics. Although critical to animal sociality, a comprehensive understanding of the factors influencing cohesion remains a gap in our knowledge of cooperative behavior in animals. We tracked 574 individuals from six species within the genus Canis in 15 countries on four continents with GPS telemetry to estimate the time that pairs of individuals within social groups spent in close proximity and test hypotheses regarding drivers of cohesion. Pairs of social canids ( Canis spp.) varied widely in the proportion of time they spent together (5%–100%) during seasonal monitoring periods relative to both intrinsic characteristics and environmental conditions. The majority of our data came from three species of wolves (gray wolves, eastern wolves, and red wolves) and coyotes. For these species, cohesion within social groups was greatest between breeding pairs and varied seasonally as the nature of cooperative activities changed relative to annual life history patterns. Across species, wolves were more cohesive than coyotes. For wolves, pairs were less cohesive in larger groups, and when suitable, small prey was present reflecting the constraints of food resources and intragroup competition on social associations. Pair cohesion in wolves declined with increased anthropogenic modification of the landscape and greater climatic variability, underscoring challenges for conserving social top predators in a changing world. We show that pairwise cohesion in social groups varies strongly both within and across Canis species, as individuals respond to changing ecological context defined by resources, competition, and anthropogenic disturbance. Our work highlights that cohesion is a highly plastic component of animal sociality that holds significant promise for elucidating ecological and evolutionary mechanisms underlying cooperative behavior.

Ecology↗

USGS2023

Forage senescence and disease influence elk pregnancy across the Greater Yellowstone Ecosystem

For various temperate ungulate species, recent research has highlighted the potential for spring vegetation phenology (“green-up”) to influence individual condition, with purported benefits to population productivity. However, few studies have been able to measure the benefit on vital rates directly, and fewer still have investigated the comparative influence of other phenological periods on ungulate vital rates. In this study, we tracked phenological changes throughout the duration of the growing season and examined how their timing affected the probability of pregnancy in an ungulate population. We did this for elk ( Cervus canadensis ) across the Greater Yellowstone Ecosystem (GYE) by sampling 1106 adult females in winter at 25 sites over a 13-year period and assessing sources of variation in pregnancy using a Bayesian hierarchical model. Pregnancy rates were generally high across the GYE (82.4%), and the primary influences on probability of pregnancy were the timing of vegetation senescence (“brown-down”) in autumn and exposure to the reproductive disease brucellosis. Earlier forage brown-down in fall negatively influenced the probability of pregnancy of elk aged 6–9 years by an estimated 17.2% within the range (ca. 32 days) of observed brown-down end dates. While summer habitat quality has been inferred to influence elk pregnancy previously, our findings specify the key influence of foraging conditions later in the seasonal cycle, immediately before the breeding season. The reproductive disease brucellosis was also an important factor, reducing the probability of pregnancy by 12.4% in elk in the 6- to 9-year age class. Because pregnancy was tested before most disease-induced abortions occur, the apparent mechanism for this effect is a prolonged reduction in fertility beyond the period of initial exposure in which fetal mortality is typically expected. Our results prompt greater scrutiny of the combined effects of late-season phenology and disease on reproductive rates and population productivity in temperate ungulates.

Idaho, Montana, Wyoming↗

USGS2023

Diverse migratory portfolios drive inter-annual switching behavior of elk across the Greater Yellowstone Ecosystem

A growing body of evidence shows that some ungulates alternate between migratory and nonmigratory behaviors over time. Yet it remains unclear whether such short-term behavioral changes can help explain reported declines in ungulate migration worldwide, as opposed to long-term demographic changes. Furthermore, advances in tracking technology reveal that a simple distinction between migration and nonmigration may not sufficiently describe all individual behaviors. To better understand the dynamics and drivers of ungulate switching behavior, we investigated 14 years of movement data from 361 elk in 20 herds across the Greater Yellowstone Ecosystem (GYE). First, we categorized yearly individual behaviors using a clustering algorithm that identified similar migratory tactics across a continuum of behaviors. Then, we tested seven hypotheses to explain why some ungulates switch behaviors, and we evaluated how behavioral changes affected the proportions of different behaviors across the system. We identified four distinct behavioral tactics: residents (4.8% of elk-years), short-distance migrants (53.7%), elevational migrants (21.9%) and long-distance migrants (19.6%). Of the 20 herds, 18 were partially migratory, and 5 had all four movement tactics present. We observed switches between migratory tactics in all sets of consecutive years during our study period, with an average of 22.5% of individual elk changing movement tactics from one year to the next. Elk in herds with higher movement tactic diversity were significantly more likely to switch tactics and often responded more effectively to adverse environmental conditions, compared to those in herds with low movement tactic diversity. During our study period, switching increased the prevalence of both short- and long-distance migrants, decreased the prevalence of elevational migrants, and had no effect on the prevalence of residents. Our findings suggest that rather than contributing to the declining migratory behavior found in the GYE, switching behavior may enable greater resiliency to continuously changing environmental and anthropogenic conditions.

Wyoming↗

USGS2023

Multi-level thresholds of residential and agricultural land use for elk avoidance across the Greater Yellowstone Ecosystem

1. Conversion of land for settlements and agriculture is increasing globally and can influence wildlife space use. However, there is limited research to identify the thresholds of land-use change that incur wildlife avoidance and how these thresh-olds might vary across levels of selection. 2. We evaluated multi-level avoidance thresholds of elk Cervus canadensis impacted by residential development and irrigated agriculture across the Greater Yellowstone Ecosystem in Idaho, Montana and Wyoming. Using GPS data from765 elk in 21 herds, we estimated habitat selection in relation to development and agriculture at three levels (home range selection, within home range selection and movement path selection). Next, using individual selection covariates and as-sociated measures of land-use availability, we used functional-response models to evaluate how selection varied based on availability, and in turn, to estimate avoidance thresholds. 3. We found individual and level-specific variation in elk responses to environmental factors. Elk exhibited stronger responses (either selection or avoidance) when selecting home range locations (i.e. second-order selection) than when selecting areas within home ranges (i.e. third-order selection) or selecting movement paths (i.e. fourth-order selection). Importantly, elk avoidance of development and agriculture changed as the amount of land in these categories changed. Across all levels of selection elk exhibited neutral selection for human development at low levels of availability (<1.1%–2.2% developed) but avoided areas that were >1.1%–2.2% developed. Conversely, elk selected positively for irrigated agriculture at low to moderate levels of availability (<52.0%–66.2% agriculture) but exhibited neutral selection in areas that were >52.0%–66.2% agriculture. 4. Synthesis and applications . Elk avoidance of low levels of human development suggests conservation efforts such as restrictions on future development or conservation easements could focus on areas that are still below 2% developed. Additionally, because elk selection was strongest at the landscape scale, conservation actions that are based on information about the overall landscape structure may be most impactful. Our results highlight the importance of under-standing variability in wildlife habitat selection at multiple levels, particularly in relation to land-use change, and highlight how functional response modelling can help inform landscape conservation.

Montana, Wyoming↗

USGS2022

Examination of the interaction between age-specific predation and chronic disease in the Greater Yellowstone Ecosystem

Predators may create healthier prey populations by selectively removing diseased individuals. Predators typically prefer some ages of prey over others, which may, or may not, align with those prey ages that are most likely to be diseased. The interaction of age-specific infection and predation has not been previously explored and likely has sizable effects on disease dynamics. We hypothesize that predator cleansing effects will be greater when the disease and predation occur in the same prey age groups. We examine the predator cleansing effect using a model where both vulnerability to predators and pathogen prevalence vary with age. We tailor this model to chronic wasting disease (CWD) in mule deer and elk populations in the Greater Yellowstone Ecosystem, with empirical data from Yellowstone grey wolves and cougars. Model results suggest that under moderate, yet realistic, predation pressure from cougars and wolves independently, predators may decrease CWD outbreak size substantially and delay the accumulation of symptomatic deer and elk. The magnitude of this effect is driven by the ability of predators to selectively remove late-stage CWD infections that are likely the most responsible for transmission, but this may not be the age class they typically select. Thus, predators that select for infected young adults over uninfected juveniles have a stronger cleansing effect, and these effects are strengthened when transmission rates increase with increasing prey morbidity. There are also trade-offs from a management perspective—that is, increasing predator kill rates can result in opposing forces on prey abundance and CWD prevalence. Our modelling exploration shows that predators have the potential to reduce prevalence in prey populations when prey age and disease severity are considered, yet the strength of this effect is influenced by predators' selection for demography or body condition. Current CWD management focuses on increasing cervid hunting as the primary management tool, and our results suggest predators may also be a useful tool under certain conditions, but not necessarily without additional impacts on host abundance and demography. Protected areas with predator populations will play a large role in informing the debate over predator impacts on disease.

Wyoming↗

USGS2022

Disease outbreaks select for mate choice and coat color in wolves

We know much about pathogen evolution and the emergence of new disease strains, but less about host resistance and how it is signaled to other individuals and subsequently maintained. The cline in frequency of black-coated wolves ( Canis lupus ) across North America is hypothesized to result from a relationship with canine distemper virus (CDV) outbreaks. We tested this hypothesis using cross-sectional data from wolf populations across North America that vary in the prevalence of CDV and the allele that makes coats black, longitudinal data from Yellowstone National Park, and modeling. We found that the frequency of CDV outbreaks generates fluctuating selection that results in heterozygote advantage that in turn affects the frequency of the black allele, optimal mating behavior, and black wolf cline across the continent.

Science↗

USGS2021

Patterns and processes of pathogen exposure in gray wolves across North America

The presence of many pathogens varies in a predictable manner with latitude, with infections decreasing from the equator towards the poles. We investigated the geographic trends of pathogens infecting a widely distributed carnivore: the gray wolf ( Canis lupus ). Specifically, we investigated which variables best explain and predict geographic trends in seroprevalence across North American wolf populations and the implications of the underlying mechanisms. We compiled a large serological dataset of nearly 2000 wolves from 17 study areas, spanning 80° longitude and 50° latitude. Generalized linear mixed models were constructed to predict the probability of seropositivity of four important pathogens: canine adenovirus, herpesvirus, parvovirus, and distemper virus—and two parasites: Neospora caninum and Toxoplasma gondii . Canine adenovirus and herpesvirus were the most widely distributed pathogens, whereas N. caninum was relatively uncommon. Canine parvovirus and distemper had high annual variation, with western populations experiencing more frequent outbreaks than eastern populations. Seroprevalence of all infections increased as wolves aged, and denser wolf populations had a greater risk of exposure. Probability of exposure was positively correlated with human density, suggesting that dogs and synanthropic animals may be important pathogen reservoirs. Pathogen exposure did not appear to follow a latitudinal gradient, with the exception of N. caninum . Instead, clustered study areas were more similar: wolves from the Great Lakes region had lower odds of exposure to the viruses, but higher odds of exposure to N. caninum and T. gondii ; the opposite was true for wolves from the central Rocky Mountains. Overall, mechanistic predictors were more informative of seroprevalence trends than latitude and longitude. Individual host characteristics as well as inherent features of ecosystems determined pathogen exposure risk on a large scale. This work emphasizes the importance of biogeographic wildlife surveillance, and we expound upon avenues of future research of cross-species transmission, spillover, and spatial variation in pathogen infection.

Alaska, Alberta, Arizona, British Columbia, Michig↗

USGS2018

Diel predator activity drives a dynamic landscape of fear

A “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‐km 2 of 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.

Yellowstone National Park↗

USGS2018

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.

Wyoming↗

USGS2017

Sexually dimorphic aggression indicates male gray wolves specialize in pack defense against conspecific groups

Aggression directed at conspecific groups is common among gregarious, territorial species, and for some species such as gray wolves ( Canis lupus ) intraspecific strife is the leading cause of natural mortality. Each individual in a group likely has different measures of the costs and benefits associated with a group task, such as an aggressive attack on another group, which can alter motivation and behavior. We observed 292 inter-pack aggressive interactions in Yellowstone National Park between 1 April 1995 and 1 April 2011 (>5300 days of observation) in order to determine the role of both sexes, and the influence of pack, age, and other traits on aggression. We recorded the behaviors and characteristics of all individuals present during the interactions (n = 534 individuals) and which individuals participated in each step (i.e. chase, attack, kill, flight) of the interaction. Overall, all wolves were more likely to chase rivals if they outnumbered their opponent, suggesting packs accurately assess their opponent’s size during encounters and individuals adjust their behavior based on relative pack size. Males were more likely than females to chase rival packs and gray-colored wolves were more aggressive than black-colored wolves. Male wolves and gray-colored wolves also recorded higher cortisol levels than females and black-colored wolves, indicating hormonal support for more intense aggressive behavior. Further, we found a positive correlation between male age and probability of chasing, while age-specific participation for females remained constant. Chasing behavior was influenced by the sex of lone intruders, with males more likely to chase male intruders. This difference in behavior suggests male and female wolves may have different strategies and motivations during inter-pack aggressive interactions related to gray wolf mating systems. A division of labor between pack members concerning resource and territory defense suggests selection for specific traits related to aggression is an adaptive response to intense competition between groups of conspecifics.

Behavioural Processes↗

USGS2016

Reproduction in moose at their southern range limit

Reproduction is a critical fitness component in large herbivores. Biogeographic models predict that populations occurring at the edges of the range may have compromised reproductive rates because of inferior habitat at range peripheries. When reproductive rates are chronically low, ungulate populations may lack the resiliency to rebound quickly after periods of environmental stress, and this effect may be greatest for heat-sensitive organisms at their southern range limit. To assess the demographic vulnerability of moose ( Alces alces ), we studied relationships between reproductive rates, maternal age, and rump fat in the southernmost naturally occurring moose population in North America. For prime-aged moose in our study, pregnancy rates were high (92%), but moose aged < 3 or > 9 years had low pregnancy rates (32% and 38%, respectively). The relationship between rump fat and pregnancy was nonlinear such that a threshold of at least 2mm of rump fat yielded a high probability of being pregnant midwinter. In contrast, among pregnant moose, the probability of both producing a calf and recruiting it until spring increased linearly with rump fat. We also conducted a meta-analysis of pregnancy and twinning rates for adult (≥ 2 years) moose across a latitudinal gradient to compare reproductive rates from our study to other populations in North America. Moose living at southern latitudes tended to have lower reproductive rates than those living in the core of moose range, implying that southern moose populations may be demographically more vulnerable than northern moose populations.

Utah↗

USGS2016

Territoriality and inter-pack aggression in gray wolves: shaping a social carnivore's life history

When Rudyard Kipling wrote The Jungle Book in 1894 and included the famous line "For the strength of the Wolf is the Pack, and the strength of the Pack is the Wolf," he would have had no idea that over a century later, scientific research would back up his poetic phrase. Recent studies in Yellowstone have found that both the individual wolf and the collective pack rely on each other and play important roles in territoriality. At a time when most fairy tales and fables were portraying wolves as demonic killers or, at best, slapstick gluttons, Kipling seemed to have a respect or even reverence for the wolf. Wolves in The Jungle Book raise and mentor the main character Mowgli, with the pack's leader eventually dying to save the "man-cub" from a pack of wolves. Kipling may have extended intra- pack benevolence to a human boy for literary sake, but he was clearly enthralled with how pack members treat each other. As wolf packs are almost always family units, most commonly comprised of a breeding pair and their offspring from several years, amiable behavior within the pack is unsurprising. By contrast, wolf packs are fiercely intolerant of their neighbors, their rivals. And this competition is proving to be an important facet in the life of a wolf and its pack.

Yellowstone Science↗

USGS2015

Group composition effects on aggressive interpack interactions of gray wolves in Yellowstone National Park

Knowledge of characteristics that promote group success during intraspecific encounters is key to understanding the adaptive advantages of sociality for many group-living species. In addition, some individuals in a group may be more likely than others to influence intergroup conflicts, a relatively neglected idea in research on social animals. Here we use observations of aggressive interactions between wolf ( Canis lupus ) packs over an extended period and use pack characteristics to determine which groups had an advantage over their opponents. During 16 years of observation in Yellowstone National Park from 1995 to 2010, we documented 121 interpack aggressive interactions. We recorded pack sizes, compositions, and spatial orientation related to residency to determine their effects on the outcomes of interactions between packs. Relative pack size (RPS) improved the odds of a pack displacing its opponent. However, pack composition moderated the effect of RPS as packs with relatively more old members (>6.0 years old) or adult males had higher odds of winning despite a numerical disadvantage. The location of the interaction with respect to pack territories had no effect on the outcome of interpack interactions. Although the importance of RPS in successful territorial defense suggests the evolution and maintenance of group living may be at least partly due to larger packs’ success during interpack interactions, group composition is also an important factor, highlighting that some individuals are more valuable than others during interpack conflicts.

Wyoming↗

USGS2015

Wolves on the hunt: The behavior of wolves hunting wild prey

The interactions between apex predators and their prey are some of the most awesome and meaningful in nature—displays of strength, endurance, and a deep coevolutionary history. And there is perhaps no apex predator more impressive and important in its hunting—or more infamous, more misjudged—than the wolf. Because of wolves’ habitat, speed, and general success at evading humans, researchers have faced great obstacles in studying their natural hunting behaviors. The first book to focus explicitly on wolf hunting of wild prey, Wolves on the Hunt seeks to fill these gaps in our knowledge and understanding. Combining behavioral data, thousands of hours of original field observations, research in the literature, a wealth of illustrations, and—in the e-book edition and online—video segments from cinematographer Robert K. Landis, the authors create a compelling and complex picture of these hunters. The wolf is indeed an adept killer, able to take down prey much larger than itself. While adapted to hunt primarily hoofed animals, a wolf—or especially a pack of wolves—can kill individuals of just about any species. But even as wolves help drive the underlying rhythms of the ecosystems they inhabit, their evolutionary prowess comes at a cost: wolves spend one-third of their time hunting—the most time consuming of all wolf activities—and success at the hunt only comes through traveling long distances, persisting in the face of regular failure, detecting and taking advantage of deficiencies in the physical condition of individual prey, and through ceaseless trial and error, all while risking injury or death. By describing and analyzing the behaviors wolves use to hunt and kill various wild prey—including deer, moose, caribou, elk, Dall sheep, mountain goats, bison, musk oxen, arctic hares, beavers, and others—Wolves on the Hunt provides a revelatory portrait of one of nature’s greatest hunters.

Book↗

USGS2011

Nonlinear effects of group size on the success of wolves hunting elk

Despite the popular view that social predators live in groups because group hunting facilitates prey capture, the apparent tendency for hunting success to peak at small group sizes suggests that the formation of large groups is unrelated to prey capture. Few empirical studies, however, have tested for nonlinear relationships between hunting success and group size, and none have demonstrated why success trails off after peaking. Here, we use a unique dataset of observations of individually known wolves ( Canis lupus ) hunting elk ( Cervus elaphus ) in Yellowstone National Park to show that the relationship between success and group size is indeed nonlinear and that individuals withholding effort (free riding) is why success does not increase across large group sizes. Beyond 4 wolves, hunting success leveled off, and individual performance (a measure of effort) decreased for reasons unrelated to interference from inept hunters, individual age, or size. But performance did drop faster among wolves with an incentive to hold back, i.e., nonbreeders with no dependent offspring, those performing dangerous predatory tasks, i.e., grabbing and restraining prey, and those in groups of proficient hunters. These results suggest that decreasing performance was free riding and that was why success leveled off in groups with >4 wolves that had superficially appeared to be cooperating. This is the first direct evidence that nonlinear trends in group hunting success reflect a switch from cooperation to free riding. It also highlights how hunting success per se is unlikely to promote formation and maintenance of large groups.

Behavioral Ecology↗

USGS2001

Winter severity and wolf predation on a formerly wolf-free elk herd

We studied wolf (Canis lupus) predation on elk (Cervus elaphus) in Yellowstone National Park from 17 March to 15 April 1997 (severe winter conditions) and from 2 to 31 March 1998 (mild winter conditions) 2-3 years after wolves were reintroduced to the park. Elk composed 91 % of 117 kills. Data comparisons for 1997 versus 1998 were: hunting success rate, 26% versus 15%; kill rate, 17.1 kg/wolf/day versus 6.1; percent of kill consumed in first day, 7 versus 86; percent femur marrow fat of adult kills, 27 versus 70; calf:adult ratios of kills, 2:33 versus 17:23; sex ratio of kills, 14M:19F versus 17M:6F; mean age of elk killed, males 6.1 years, females 15.2 versus males, 4.8, females 13.0. Winter severity influenced the wolf-elk relationship more than the naivete of the elk herd to predation by wolves.

Journal of Wildlife Management↗