Genomic Analyses Capture the Human-Induced Demographic Collapse and Recovery in a Wide-Ranging Cervid.

The glacial cycles of the Quaternary heavily impacted species through successions of population contractions and expansions. Similarly, populations have been intensely shaped by human pressures such as unregulated hunting and land use changes. White-tailed and mule deer survived in different refugia through the Last Glacial Maximum, and their populations were severely reduced after the European colonization. Here, we analyzed 73 resequenced deer genomes from across their North American range to understand the consequences of climatic and anthropogenic pressures on deer demographic and adaptive history. We found strong signals of climate-induced vicariance and demographic decline; notably, multiple sequentially Markovian coalescent recovers a severe decline in mainland white-tailed deer effective population size (Ne) at the end of the Last Glacial Maximum. We found robust evidence for colonial overharvest in the form of a recent and dramatic drop in Ne in all analyzed populations. Historical census size and restocking data show a clear parallel to historical Ne estimates, and temporal Ne/Nc ratio shows patterns of conservation concern for mule deer. Signatures of selection highlight genes related to temperature, including a cold receptor previously highlighted in woolly mammoth. We also detected immune genes that we surmise reflect the changing land use patterns in North America. Our study provides a detailed picture of anthropogenic and climatic-induced decline in deer diversity and clues to understanding the conservation concerns of mule deer and the successful demographic recovery of white-tailed deer.

Experimental SARS-CoV-2 Infection of Elk and Mule Deer.

To assess the susceptibility of elk (Cervus canadensis) and mule deer (Odocoileus hemionus) to SARS-CoV-2, we performed experimental infections in both species. Elk did not shed infectious virus but mounted low-level serologic responses. Mule deer shed and transmitted virus and mounted pronounced serologic responses and thus could play a role in SARS-CoV-2 epidemiology.

Structural basis of white-tailed deer, Odocoileus virginianus, ACE2 recognizing all the SARS-CoV-2 variants of concern with high affinity.

SARS-CoV-2 has been expanding its host range, among which the white-tailed deer (WTD), Odocoileus virginianus, became the first wildlife species infected on a large scale and might serve as a host reservoir for variants of concern (VOCs) in case no longer circulating in humans. In this study, we comprehensively assessed the binding of the WTD angiotensin-converting enzyme 2 (ACE2) receptor to the spike (S) receptor-binding domains (RBDs) from the SARS-CoV-2 prototype (PT) strain and multiple variants. We found that WTD ACE2 could be broadly recognized by all of the tested RBDs. We further determined the complex structures of WTD ACE2 with PT, Omicron BA.1, and BA.4/5 S trimer. Detailed structural comparison revealed the important roles of RBD residues on 486, 498, and 501 sites for WTD ACE2 binding. This study deepens our understanding of the interspecies transmission mechanisms of SARS-CoV-2 and further addresses the importance of constant monitoring on SARS-CoV-2 infections in wild animals. IMPORTANCE Even if we manage to eliminate the virus among humans, it will still circulate among wildlife and continuously be transmitted back to humans. A recent study indicated that WTD may serve as reservoir for nearly extinct SARS-CoV-2 strains. Therefore, it is critical to evaluate the binding abilities of SARS-CoV-2 variants to the WTD ACE2 receptor and elucidate the molecular mechanisms of binding of the RBDs to assess the risk of spillback events.

Habitat alteration or climate: What drives the densities of an invading ungulate?

Anthropogenic habitat alteration and climate change are two well-known contributors to biodiversity loss through changes to species distribution and abundance; yet, disentangling the effects of these two factors is often hindered by their inherent confound across both space and time. We leveraged a contrast in habitat alteration associated with the jurisdictional boundary between two Canadian provinces to evaluate the relative effects of spatial variation in habitat alteration and climate on white-tailed deer (Odocoileus virginianus) densities. White-tailed deer are an invading ungulate across much of North America, whose expansion into Canada's boreal forest is implicated in the decline of boreal caribou (Rangifer tarandus caribou), a species listed as Threatened in Canada. We estimated white-tailed deer densities using 300 remote cameras across 12 replicated 50 km2 landscapes over 5 years. White-tailed deer densities were significantly lower in areas where winter severity was higher. For example, predicted deer densities declined from 1.83 to 0.35 deer/km2 when winter severity increased from the lowest value to the median value. There was a tendency for densities to increase with increasing habitat alteration; however, the magnitude of this effect was approximately half that of climate. Our findings suggest that climate is the primary driver of white-tailed deer populations; however, understanding the mechanisms underpinning this relationship requires further study of over-winter survival and fecundity. Long-term monitoring at the invasion front is needed to evaluate the drivers of abundance over time, particularly given the unpredictability of climate change and increasing prevalence of extreme weather events.

White-tailed deer population dynamics in a multipredator landscape shaped by humans.

Large terrestrial mammals increasingly rely on human-modified landscapes as anthropogenic footprints expand. Land management activities such as timber harvest, agriculture, and roads can influence prey population dynamics by altering forage resources and predation risk via changes in habitat, but these effects are not well understood in regions with diverse and changing predator guilds. In northeastern Washington state, USA, white-tailed deer (Odocoileus virginianus) are vulnerable to multiple carnivores, including recently returned gray wolves (Canis lupus), within a highly human-modified landscape. To understand the factors governing predator-prey dynamics in a human context, we radio-collared 280 white-tailed deer, 33 bobcats (Lynx rufus), 50 cougars (Puma concolor), 28 coyotes (C. latrans), and 14 wolves between 2016 and 2021. We first estimated deer vital rates and used a stage-structured matrix model to estimate their population growth rate. During the study, we observed a stable to declining deer population (lambda = 0.97, 95% confidence interval: 0.88, 1.05), with 74% of Monte Carlo simulations indicating population decrease and 26% of simulations indicating population increase. We then fit Cox proportional hazard models to evaluate how predator exposure, use of human-modified landscapes, and winter severity influenced deer survival and used these relationships to evaluate impacts on overall population growth. We found that the population growth rate was dually influenced by a negative direct effect of apex predators and a positive effect of timber harvest and agricultural areas. Cougars had a stronger effect on deer population dynamics than wolves, and mesopredators had little influence on the deer population growth rate. Areas of recent timber harvest had 55% more forage biomass than older forests, but horizontal visibility did not differ, suggesting that timber harvest did not influence predation risk. Although proximity to roads did not affect the overall population growth rate, vehicle collisions caused a substantial proportion of deer mortalities, and reducing these collisions could be a win-win for deer and humans. The influence of apex predators and forage indicates a dual limitation by top-down and bottom-up factors in this highly human-modified system, suggesting that a reduction in apex predators would intensify density-dependent regulation of the deer population owing to limited forage availability.

Spatial prey availability and pulsed reproductive tactics: Encounter risk in a canid-ungulate system.

Predation risk is a function of spatiotemporal overlap between predator and prey, as well as behavioural responses during encounters. Dynamic factors (e.g. group size, prey availability and animal movement or state) affect risk, but rarely are integrated in risk assessments. Our work targets a system where predation risk is fundamentally linked to temporal patterns in prey abundance and behaviour. For neonatal ungulate prey, risk is defined within a short temporal window during which the pulse in parturition, increasing movement capacity with age and antipredation tactics have the potential to mediate risk. In our coyote-mule deer (Canis latrans-Odocoileus hemionus) system, leveraging GPS data collected from both predator and prey, we tested expectations of shared enemy and reproductive risk hypotheses. We asked two questions regarding risk: (A) How does primary and alternative prey habitat, predator and prey activity, and reproductive tactics (e.g. birth synchrony and maternal defence) influence the vulnerability of a neonate encountering a predator? (B) How do the same factors affect behaviour by predators relative to the time before and after an encounter? Despite increased selection for mule deer and intensified search behaviour by coyotes during the peak in mule deer parturition, mule deer were afforded protection from predation via predator swamping, experiencing reduced per-capita encounter risk when most neonates were born. Mule deer occupying rabbit habitat (Sylvilagus spp.; coyote's primary prey) experienced the greatest risk of encounter but the availability of rabbit habitat did not affect predator behaviour during encounters. Encounter risk increased in areas with greater availability of mule deer habitat: coyotes shifted their behaviour relative to deer habitat, and the pulse in mule deer parturition and movement of neonatal deer during encounters elicited increased speed and tortuosity by coyotes. In addition to the spatial distribution of prey, temporal patterns in prey availability and animal behavioural state were fundamental in defining risk. Our work reveals the nuanced consequences of pulsed availability on predation risk for alternative prey, whereby responses by predators to sudden resource availability, the lasting effects of diversionary prey and inherent antipredation tactics ultimately dictate risk.

Wolves make roadways safer, generating large economic returns to predator conservation.

Recent studies uncover cascading ecological effects resulting from removing and reintroducing predators into a landscape, but little is known about effects on human lives and property. We quantify the effects of restoring wolf populations by evaluating their influence on deer-vehicle collisions (DVCs) in Wisconsin. We show that, for the average county, wolf entry reduced DVCs by 24%, yielding an economic benefit that is 63 times greater than the costs of verified wolf predation on livestock. Most of the reduction is due to a behavioral response of deer to wolves rather than through a deer population decline from wolf predation. This finding supports ecological research emphasizing the role of predators in creating a "landscape of fear." It suggests wolves control economic damages from overabundant deer in ways that human deer hunters cannot.

Resource selection by New York City deer reveals the effective interface between wildlife, zoonotic hazards and humans.

Although the role of host movement in shaping infectious disease dynamics is widely acknowledged, methodological separation between animal movement and disease ecology has prevented researchers from leveraging empirical insights from movement data to advance landscape scale understanding of infectious disease risk. To address this knowledge gap, we examine how movement behaviour and resource utilization by white-tailed deer (Odocoileus virginianus) determines blacklegged tick (Ixodes scapularis) distribution, which depend on deer for dispersal in a highly fragmented New York City borough. Multi-scale hierarchical resource selection analysis and movement modelling provide insight into how deer's movements contribute to the risk landscape for human exposure to the Lyme disease vector-I. scapularis. We find deer select highly vegetated and accessible residential properties which support blacklegged tick survival. We conclude the distribution of tick-borne disease risk results from the individual resource selection by deer across spatial scales in response to habitat fragmentation and anthropogenic disturbances.

Speciation without gene-flow in hybridizing deer.

Under the ecological speciation model, divergent selection acts on ecological differences between populations, gradually creating barriers to gene flow and ultimately leading to reproductive isolation. Hybridisation is part of this continuum and can both promote and inhibit the speciation process. Here, we used white-tailed (Odocoileus virginianus) and mule deer (O. hemionus) to investigate patterns of speciation in hybridizing sister species. We quantified genome-wide historical introgression and performed genome scans to look for signatures of four different selection scenarios. Despite ample modern evidence of hybridisation, we found negligible patterns of ancestral introgression and no signatures of divergence with gene flow, rather localized patterns of allopatric and balancing selection were detected across the genome. Genes under balancing selection were related to immunity, MHC and sensory perception of smell, the latter of which is consistent with deer biology. The deficiency of historical gene-flow suggests that white-tailed and mule deer were spatially separated during the glaciation cycles of the Pleistocene and genome wide differentiation accrued via genetic drift. Dobzhansky-Muller incompatibilities and selection against hybrids are hypothesised to be acting, and diversity correlations to recombination rates suggests these sister species are far along the speciation continuum.

Deer dietary responses to wildfire: Optimal foraging, individual specialization, or opportunism?

Increasing impacts of wildfire on arid regions of the world fuelled by climate change highlight the need to better understand how natural communities respond to fire. We took advantage of a large (1660-km2 ) wildfire that erupted in northern California during an in-progress study of black-tailed deer (Odocoileus hemionus columbianus) to investigate deer use of and diets within burned and unburned habitats before and after the fire. We compared deer diet breadth to predictions of optimal foraging theory, the niche variation hypothesis, and opportunistic (i.e., generalist) foraging expectations under the assumption that overall availability and diversity of forage in burned areas declined immediately after the fire and increased as the plant community recovered in the next 3 years after the fire. We used faecal pellet counts to document space use and metabarcoding to study diet during pre-fire, post-fire, and recovery periods. Pellet counts supported predictions that deer increased use of unburned sites and reduced use of burn sites after the fire and began to return to burned sites in subsequent sampling years. Diet diversity did not differ significantly between control and burn sites before the fire, but was lower in burn than control sites post-fire (p < .001), when and where diet was dominated by oak (Quercus spp). In contrast, during subsequent years, diet diversity was higher (including more herbaceous plants) in burn than control sites (p < .05). In contrast to predictions of optimal foraging and niche variation hypotheses, individual deer foraged as generalists for which changes in dietary niche breadth paralleled fire-induced changes in diversity of the plant community.

Integration of aerial surveys and resource selection analysis indicates human land use supports boreal deer expansion.

Landscape change is a driver of global biodiversity loss. In the western Nearctic, petroleum exploration and extraction is a major contributor to landscape change, with concomitant effects on large mammal populations. One of those effects is the continued expansion of invasive white-tailed deer populations into the boreal forest, with ramifications for the whole ecosystem. We explored deer resource selection within the oil sands region of the boreal forest using a novel application of aerial ungulate survey (AUS) data. Deer locations from AUS were "used" points and together with randomly allocated "available" points informed deer resource selection in relation to landscape variables in the boreal forest. We created a candidate set of generalized linear models representing competing hypotheses about the role of natural landscape features, forest harvesting, cultivation, roads, and petroleum features. We ranked these in an information-theoretic framework. A combination of natural and anthropogenic landscape features best explained deer resource selection. Deer strongly selected seismic lines and other linear features associated with petroleum exploration and extraction, likely as movement corridors and resource subsidies. Forest harvesting and cultivation, important contributors to expansion in other parts of the white-tailed deer range, were not as important here. Stemming deer expansion to conserve native ungulates and maintain key predator-prey processes will likely require landscape management to restore the widespread linear features crossing the vast oil sands region.

Logging, linear features, and human infrastructure shape the spatial dynamics of wolf predation on an ungulate neonate.

Humans are increasingly recognized as important players in predator-prey dynamics by modifying landscapes. This trend has been well-documented for large mammal communities in North American boreal forests: logging creates early seral forests that benefit ungulates such as white-tailed deer (Odocoileus virginianus), while the combination of infrastructure development and resource extraction practices generate linear features that allow predators such as wolves (Canis lupus) to travel and forage more efficiently throughout the landscape. Disturbances from recreational activities and residential development are other major sources of human activity in boreal ecosystems that may further alter wolf-ungulate dynamics. Here, we evaluate the influence that several major types of anthropogenic landscape modifications (timber harvest, linear features, and residential infrastructure) have on where and how wolves hunt ungulate neonates in a southern boreal forest ecosystem in Minnesota, USA. We demonstrate that each major anthropogenic disturbance significantly influences wolf predation of white-tailed deer fawns (n = 427 kill sites). In contrast with the "human shield hypothesis" that posits prey use human-modified areas as refuge, wolves killed fawns closer to residential buildings than expected based on spatial availability. Fawns were also killed within recently-logged areas more than expected. Concealment cover was higher at kill sites than random sites, suggesting wolves use senses other than vision, probably olfaction, to detect hidden fawns. Wolves showed strong selection for hunting along linear features, and kill sites were also closer to linear features than expected. We hypothesize that linear features facilitated wolf predation on fawns by allowing wolves to travel efficiently among high-quality prey patches (recently logged areas, near buildings), and also increase encounter rates with olfactory cues that allow them to detect hidden fawns. These findings provide novel insight into the strategies predators use to hunt ungulate neonates and the many ways human activity alters wolf-ungulate neonate predator-prey dynamics, which have remained elusive due to the challenges of locating sites where predators kill small prey. Our research has important management and conservation implications for wolf-ungulate systems subjected to anthropogenic pressures, particularly as the range of overlap between wolves and deer expands and appears to be altering food web dynamics in boreal ecosystems.

First description of the mitochondrial genomes of the Central American brocket deer Mazama temama (Kerr, 1792) and the Yucatán Peninsula brocket deer Odocoileus pandora Merriam, 1901.

BACKGROUND: The Central American (Mazama temama) and the Yucatán Peninsula brocket deer (Odocoileus pandora) are deer species with cryptic habits, and little is known about their biology. Odocoileus pandora is listed as Vulnerable on the 2015 IUCN Red List of Threatened Species, while M. temama is considered Data Deficient; however, it currently faces a decreasing population trend. METHODS AND RESULTS: We assembled the complete mitochondrial genome for two M. temama specimens and one complete and one partial for O. pandora from Illumina 150 bp paired-end reads. The mitogenomes of M. temama and O. pandora have a length of 16,479-16,480 and 16,419 bp, respectively, AT-biased; they consist of 13 protein-coding genes, two ribosomal RNA genes, 22 transfer RNA genes, and one non-coding control region, most of them follow a transcription direction in the heavy strand of the molecule. The mitochondrial genome of O. pandora shows some particularities compared to other deer species, like a shorter control region of 987-990 bp and a cytochrome b gene with a length of 1,143 bp. Our phylogenetic analyses confirm the close affinity of M. temama to South American M. americana and the nested position of the genus Odocoileus, including O. pandora, into the genus Mazama. CONCLUSIONS: Here, we described for the first time the complete mitochondrial genome for these two species. While our study provides additional information about the taxonomic status of the northern neotropical brocket deer, further research is needed to solve the complicated taxonomy of neotropical deer.

Limited detection of shared zoonotic pathogens in deer keds and blacklegged ticks co-parasitizing white-tailed deer in the eastern United States.

Deer keds, such as Lipoptena cervi Linnaeus (Diptera: Hippoboscidae), are blood-feeding flies from which several human and animal pathogens have been detected, including Borrelia burgdorferi sensu lato Johnson (Spirochaetales: Borreliaceae), the causative agent of Lyme disease. Cervids (Artiodactyla: Cervidae), which are the primary hosts of deer keds, are not natural reservoirs of B. burgdorferi sl, and it has been suggested that deer keds may acquire bacterial pathogens via co-feeding near infected ticks. We screened L. cervi (n = 306) and Ixodes scapularis Say (Ixodida: Ixodidae) (n = 315) collected from 38 white-tailed deer in Pennsylvania for the family Anaplasmataceae, Bartonella spp. (Hyphomicrobiales: Bartonellaceae), Borrelia spp., and Rickettsia spp. (Rickettsiales: Rickettsiaceae). Limited similarity in the bacterial DNA detected between these ectoparasites per host suggested that co-feeding may not be a mechanism by which deer keds acquire these bacteria. The feeding biology and life history of deer keds may impact the observed results, as could the season when specimens were collected. We separately screened L. cervi (n = 410), L. mazamae Róndani (n = 13), L. depressa Say (n = 10), and Neolipoptena ferrisi Bequaert (n = 14) collections from locations within the United States and Canada for the same pathogens. These results highlight the need to further study deer ked-host and deer ked-tick relationships.

Species and physiographic factors drive Indian cucumber root and Canada mayflower plant chemistry: Implications for white-tailed deer forage quality.

Nutrition is fundamental to white-tailed deer (Odocoileus virginianus) management given its relationship to habitat carrying capacity and population productivity. Ecological Sites (ESs) are a United States federal landscape management unit of specific land potential due to unique soils, topography, climate, parent material, and perhaps deer forage nutritional value. We present results of a study that extends the use of ESs to inform white-tailed deer management by evaluating indicator plant chemistry in two spring forb species, Indian cucumber root (Medeola virginiana) and Canada mayflower (Maianthemum canadense), across the northcentral Appalachians. We sampled spring forbs and underlying soils across two ESs: Dry, upland, oak-maple-hemlock hardwood forest (OMH) and Deep soil, high slope, northern hardwood forests (NHF). Plant elemental content, soil pH, and site aspect, slope and elevation were measured. Our results show that forb chemistry differs between species and within a species geographically. Indian cucumber root, as compared to Canada mayflower, has significantly higher Mg, Na, Cu, Fe, and Zn, and lower Mn. Canada mayflower in the NHF ES, versus OMH ES, was found to have significantly higher K, Mn, and B. Indian cucumber root in the NHF ES, versus the OMH ES, was found to have significantly higher Mg, Al, Fe, and Ca:P ratio but lower K. Linear discriminant analysis shows that plant tissue Mn was the best discriminator between species, and between ESs, Canada mayflower plant tissue Mn and Indian cucumber plant tissue P, K, Ca, Mg and Mn were best discriminators. Given that nutrition determines habitat carrying capacity, differences in forage nutrition between ESs may have different potentials to support deer. Forage nutrition is an important aspect of deer habitat conditions and carrying capacity, thus ESs are likely to support deer populations with different growth potential, which means that even if the same plant species occur in different ESs their nutritional value to deer may differ.

Intentions of Landowners towards Active Management of Ecosystem for Deer Habitat.

Active management such as prescribed fire and thinning can restore savanna and prairie ecosystem to maintain a full suite of ecosystem services and create suitable habitat for wildlife species such as white-tailed deer (Odocoileus virginianus). Active management comes with the cost of management and acceptance of management tools. The south-central transitional ecoregion of the USA, which otherwise was a mixture of forest, savanna, and tallgrass prairie, is increasing in woody plant dominance due to the exclusion of fire and other anthropogenic factors. Deer hunting is a vital source of revenue generation to offset the landowner's management cost in the region. We studied Oklahoma landowners' perceptions regarding active and sustainable management of forest and rangeland for deer habitat using two established theories of reasoned action and planned behavior as well as expanded theories adding moral norms. We analyzed mailed survey data using structural equation modeling. We found that subjective norms and perceived behavior control significantly affected deer hunting intention when moral norms were introduced into the model. Attitudes independently significantly affected intentions of deer hunting but have negative relations with the intentions. The study suggested that landowners have positive social pressure and were interested in active management but associated financial burden and risk could be shaping negative attitudes.

Responses to natural gas development differ by season for two migratory ungulates.

While migrating, animals make directionally persistent movements and may only respond to human-induced rapid environmental change (HIREC), such as climate and land-use change, once a threshold of HIREC is surpassed. In contrast, animals on other seasonal ranges (e.g., winter range) make more localized and tortuous movements while foraging and may have the flexibility to adjust the location of their range and the intensity of use within it to minimize interactions with HIREC. Because of these seasonal differences in movement, animals on seasonal ranges should avoid areas that contain any level of HIREC, however, during migration, animals should use areas that contain low levels of HIREC, avoiding it only once a threshold of HIREC has been surpassed. We tested this hypothesis using a decade of GPS collar data collected from migratory mule deer (Odocoileus hemionus; n = 56 migration, 143 winter) and pronghorn (Antilocapra americana; n = 70 migration, 89 winter) that winter on and migrate through a natural gas field in western Wyoming. Using surface disturbance caused by well pads and roads as an index of HIREC, we evaluated behavioral responses across three spatial scales during winter and migration seasons. During migration, both species tolerated low levels of disturbance. Once a disturbance threshold was surpassed, however, they avoided HIREC. For mule deer, thresholds were consistently ~3%, whereas thresholds for pronghorn ranged from 1% to 9.25% surface disturbance. In contrast to migration, both species generally avoided all levels of HIREC while on winter range. Our study suggests that animal responses to HIREC are mediated by season-specific movement patterns. Our results provide further evidence of ungulates avoiding human disturbance on winter range and reveal disturbance thresholds that trigger mule deer and pronghorn responses during migration: information that managers can use to maintain the ecological function of migration routes and winter ranges.

Stasis in forest regeneration following deer exclusion and understory gap creation: A 10-year experiment.

Chronically elevated ungulate browse pressure in temperate forests worldwide often generates ecological legacies characterized by low plant diversity and contributes to the formation of dense, nearly monodominant, and highly recalcitrant layers of understory vegetation. Once established, these recalcitrant layers combined with continued browsing may jointly constrain tree establishment and diversity so completely that understory recovery may be unattainable without mitigating browse pressure, the recalcitrant layer, or both. Here, we investigate the independent and synergistic effects of both white-tailed deer (Odocoileus virginianus) browsing and hay-scented fern (Dennstaedtia punctilobula) competition on tree regeneration in a 10-year experiment. Specifically, we examine how tree seedling establishment, growth, and composition are filtered by fern cover versus fern removal (gaps), browser presence versus absence (exclosures), and their combined effects during 10 years at three hardwood forest sites in Pennsylvania, USA. Fern gaps enhanced establishment for multiple tree species, increasing seedling density and diversity, particularly in the first 3 years post-treatment, and enhancing richness (≤1 species) over the course of the experiment. Excluding deer for a decade increased the height growth of other regeneration and altered species composition, but had no effect on diversity, richness, and density. Notably, we observed higher Prunus serotina seedling densities outside exclosures, possibly due to greater secondary dispersal. We argue that browsing legacies in second growth forests established at the turn of the last century created two conditions inimical to diverse forest regeneration: an overstory dominated by two species, P. serotina and Acer rubrum (86% of basal area), and a dense recalcitrant understory layer dominated by a native fern. The first condition limits propagule supply, the second strongly filters seedling establishment, and both create impoverished forest understories composed of few individuals and species. In undisturbed forest understories, the inertia toward impoverishment was sustained across the decade even where browsing was eliminated. Consequently, stand replacing disturbances (whether natural or anthropogenic) that disrupt the understory layer and reinitiate succession may be necessary to propel forests out of their current stasis and down a pathway leading to greater diversity.

Interactive effects of wildfires, season and predator activity shape mule deer movements.

Wildfires are increasing in size, frequency and severity due to climate change and fire suppression, but the direct and indirect effects on wildlife remain largely unresolved. Fire removes forest canopy, which can improve forage for ungulates but also reduce snow interception, leading to a deeper snowpack and potentially increased vulnerability to predation in winter. If ungulates exhibit predator-mediated foraging, burns should generally be selected for in summer to access high-quality forage and avoided in winter to reduce predation risk in deep snow. Fires also typically increase the amount of deadfall and initiate the growth of dense understory vegetation, creating obstacles that may confer a hunting advantage to stalking predators and a disadvantage to coursing predators. To minimize risk, ungulates may therefore avoid burns when and where stalking predators are most active, and use burns when and where coursing predators are most active. We used telemetry data from GPS-collared mule deer (Odocoileus hemionus), cougars (Puma concolor) and wolves (Canis lupus) to develop step selection functions to examine how mule deer navigated species-specific predation risk across a landscape in northern Washington, USA, that has experienced substantial wildfire activity during the past several decades. We considered a diverse array of wildfire impacts, accounting for both the severity of the fire and time since the burn (1-35 years) in our analyses. We observed support for the predator mediating foraging hypothesis: mule deer generally selected for burned areas in summer and avoided burns in winter. In addition, deer increased use of burned areas when and where wolf activity was high and avoided burns when and where cougar use was high in winter, suggesting the hunting mode of resident predators mediated the seasonal response of deer to burns. Deer were not more likely to die by predation in burned than in unburned areas, indicating that they adequately manage fire-induced changes to predation risk. As fire activity increases with climate change, our findings indicate the impact on ungulates will depend on trade-offs between enhanced summer forage and functionally reduced winter range, mediated by characteristics of the predator community.

Prey tells, large herbivores fear the human 'super predator'.

Fear of the human 'super predator' has been demonstrated to so alter the feeding behavior of large carnivores as to cause trophic cascades. It has yet to be experimentally tested if fear of humans has comparably large effects on the feeding behavior of large herbivores. We conducted a predator playback experiment exposing white-tailed deer to the vocalizations of humans, extant or locally extirpated non-human predators (coyotes, cougars, dogs, wolves), or non-predator controls (birds), at supplemental food patches to measure the relative impacts on deer feeding behavior. Deer were more than twice as likely to flee upon hearing humans than other predators, and hearing humans was matched only by hearing wolves in reducing overall feeding time gaged by visits to the food patch in the following hour. Combined with previous, site-specific research linking deer fecundity to predator abundance, this study reveals that fear of humans has the potential to induce a larger effect on ungulate reproduction than has ever been reported. By demonstrating that deer most fear the human 'super predator', our results point to the fear humans induce in large ungulates having population- and community-level impacts comparable to those caused by the fear humans induce in large carnivores.