Human disturbance compresses the spatiotemporal niche.

Human disturbance may fundamentally alter the way that species interact, a prospect that remains poorly understood. We investigated whether anthropogenic landscape modification increases or decreases co-occurrence-a prerequisite for species interactions-within wildlife communities. Using 4 y of data from >2,000 camera traps across a human disturbance gradient in Wisconsin, USA, we considered 74 species pairs (classifying pairs as low, medium, or high antagonism to account for different interaction types) and used the time between successive detections of pairs as a measure of their co-occurrence probability and to define co-occurrence networks. Pairs averaged 6.1 [95% CI: 5.3, 6.8] d between detections in low-disturbance landscapes (e.g., national forests) but 4.1 [3.5, 4.7] d between detections in high-disturbance landscapes, such as those dominated by urbanization or intensive agriculture. Co-occurrence networks showed higher connectance (i.e., a larger proportion of the possible co-occurrences) and greater proportions of low-antagonism pairs in disturbed landscapes. Human-mediated increases in species abundance (possibly via resource subsidies) appeared more important than behavioral mechanisms (e.g., changes in daily activity timing) in driving these patterns of compressed co-occurrence in disturbed landscapes. The spatiotemporal compression of species co-occurrences in disturbed landscapes likely strengthens interactions like competition, predation, and infection unless species can avoid each other at fine spatiotemporal scales. Regardless, human-mediated increases in co-occurrence with-and hence increased exposure to-predators or competitors might elevate stress levels in individual animals, with possible cascading effects across populations, communities, and ecosystems.

Missing History of a Modern Domesticate: Historical Demographics and Genetic Diversity in Farm-bred Red Fox Populations.

The first record of captive bred red foxes (Vulpes vulpes) dates to 1896, when a breeding enterprise emerged in the provinces of Atlantic Canada. Because its domestication happened during recent history, the red fox offers a unique opportunity to examine the genetic diversity of an emerging domesticated species in the context of documented historical and economic influences. In particular, the historical record suggests that North American and Eurasian farm-bred populations likely experienced different demographic trajectories. Here, we focus on the likely impacts of founder effects and genetic drift given historical trends in fox farming on North American and Eurasian farms. A total of 15 mitochondrial haplotypes were identified in 369 foxes from 10 farm populations that we genotyped (n=161) or that were previously published. All haplotypes are endemic to North America. Although most haplotypes were consistent with eastern Canadian ancestry, a small number of foxes carried haplotypes typically found in Alaska and other regions of western North America. The presence of these haplotypes supports historical reports of wild foxes outside of Atlantic Canada being introduced into the breeding stock. These putative Alaskan and Western haplotypes were more frequently identified in Eurasian farms compared to North American farms, consistent with historical documentation suggesting that Eurasian economic and breeding practices were likely to maintain low-frequency haplotypes more effectively than in North America. Contextualizing inter- versus intra-farm genetic diversity alongside the historical record is critical to understanding of the origins of this emerging domesticate and the relationships between wild and farm-bred fox populations.

Landscape heterogeneity provides co-benefits to predator and prey.

Predator populations are imperiled globally, due in part to changing habitat and trophic interactions. Theoretical and laboratory studies suggest that heterogeneous landscapes containing prey refuges acting as source habitats can benefit both predator and prey populations, although the importance of heterogeneity in natural systems is uncertain. Here, we tested the hypothesis that landscape heterogeneity mediates predator-prey interactions between the California spotted owl (Strix occidentalis occidentalis)-a mature forest species-and one of its principal prey, the dusky-footed woodrat (Neotoma fuscipes)-a younger forest species-to the benefit of both. We did so by combining estimates of woodrat density and survival from live trapping and very high frequency tracking with direct observations of prey deliveries to dependent young by owls in both heterogeneous and homogeneous home ranges. Woodrat abundance was ~2.5 times higher in owl home ranges (14.12 km2 ) featuring greater heterogeneity in vegetation types (1805.0 ± 50.2 SE) compared to those dominated by mature forest (727.3 ± 51.9 SE), in large part because of high densities in young forests appearing to act as sources promoting woodrat densities in nearby mature forests. Woodrat mortality rates were low across vegetation types and did not differ between heterogeneous and homogeneous home ranges, yet all observed predation by owls occurred within mature forests, suggesting young forests may act as woodrat refuges. Owls exhibited a type 1 functional response, consuming ~2.5 times more woodrats in heterogeneous (31.1/month ± 5.2 SE) versus homogeneous (12.7/month ± 3.7 SE) home ranges. While consumption of smaller-bodied alternative prey partially compensated for lower woodrat consumption in homogeneous home ranges, owls nevertheless consumed 30% more biomass in heterogeneous home ranges-approximately equivalent to the energetic needs of producing one additional offspring. Thus, a mosaic of vegetation types including young forest patches increased woodrat abundance and availability that, in turn, provided energetic and potentially reproductive benefits to mature forest-associated spotted owls. More broadly, our findings provide strong empirical evidence that heterogeneous landscapes containing prey refuges can benefit both predator and prey populations. As anthropogenic activities continue to homogenize landscapes globally, promoting heterogeneous systems with prey refuges may benefit imperiled predators.

Reciprocated competition between two forest carnivores drives dietary specialization.

Competition shapes animal communities, but the strength of the interaction varies spatially depending on the availability and aggregation of resources and competitors. Among carnivores, competition is particularly pronounced with the strongest interactions between similar species with intermediate differences in body size. While ecologists have emphasized interference competition among carnivores based on dominance hierarchies from body size (smaller = subordinate; larger = dominant), the reciprocity of exploitative competition from subordinate species has been overlooked even though efficient exploitation can limit resource availability and influence foraging. Across North America, fishers Pekania pennanti and martens (Martes spp.) are two phylogenetically related forest carnivores that exhibit a high degree of overlap in habitat use and diet and differ in body size by a factor of 2-5×, eliciting particularly strong interspecific competition. In the Great Lakes region, fishers and martens occur both allopatrically and sympatrically; where they co-occur, the numerically dominant species varies spatially. This natural variation in competitors and environmental conditions enables comparisons to understand how interference and exploitative competition alter dietary niche overlap and foraging strategies. We analysed stable isotopes (δ13 C and δ15 N) from 317 martens and 132 fishers, as well as dietary items (n = 629) from 20 different genera, to compare niche size and overlap. We then quantified individual diet specialization and modelled the response to environmental conditions that were hypothesized to influence individual foraging. Martens and fishers exhibited high overlap in both available and core isotopic δ-space, but no overlap of core dietary proportions. When the competitor was absent or rare, both martens and fishers consumed more smaller-bodied prey. Notably, the dominant fisher switched from being a specialist of larger to smaller prey in the absence of the subordinate marten. Environmental context also influenced dietary specialization: increasing land cover diversity and prey abundance reduced specialization in martens whereas vegetation productivity increased specialization for both martens and fishers. Despite an important dominance hierarchy, fishers adjusted their niche in the face of a subordinate, but superior, exploitative competitor. These findings highlight the underappreciated role of the subordinate competitor in shaping the dietary niche of a dominant competitor.

Phenotypic variation in the molt characteristics of a seasonal coat color-changing species reveals limited resilience to climate change.

The snowshoe hare (Lepus americanus) possesses a broad suite of adaptations to winter, including a seasonal coat color molt. Recently, climate change has been implicated in the range contraction of snowshoe hares along the southern range boundary. With shortening snow season duration, snowshoe hares are experiencing increased camouflage mismatch with their environment reducing survival. Phenological variation of hare molt at regional scales could facilitate local adaptation in the face of climate change, but the level of variation, especially along the southern range boundary, is unknown. Using a network of trail cameras and historical museum specimens, we (1) developed contemporary and historical molt phenology curves in the Upper Great Lakes region, USA, (2) calculated molt rate and variability in and among populations, and (3) quantified the relationship of molt characteristics to environmental conditions for snowshoe hares across North America. We found that snowshoe hares across the region exhibited similar fall and spring molt phenologies, rates and variation. Yet, an insular island population of hares on Isle Royale National Park, MI, completed their molt a week earlier in the fall and initiated molt almost 2 weeks later in the spring as well as exhibited slower rates of molting in the fall season compared to the mainland. Over the last 100 years, snowshoe hares across the region have not shifted in fall molt timing; though contemporary spring molt appears to have advanced by 17 days (~ 4 days per decade) compared to historical molt phenology. Our research indicates that some variation in molt phenology exists for snowshoe hares in the Upper Great Lakes region, but whether this variation is enough to offset the consequences of climate change remains to be seen.

Older forests function as energetic and demographic refugia for a climate-sensitive species.

More frequent and extreme heat waves threaten climate-sensitive species. Structurally complex, older forests can buffer these effects by creating cool microclimates, although the mechanisms by which forest refugia mitigate physiological responses to heat exposure and subsequent population-level consequences remain relatively unexplored. We leveraged fine-scale movement data, doubly labeled water, and two decades of demographic data for the California spotted owl (Strix occidentalis occidentalis) to (1) assess the role of older forest characteristics as potential energetic buffers for individuals and (2) examine the subsequent value of older forests as refugia for a core population in the Sierra Nevada and a periphery population in the San Bernardino Mountains. Individuals spent less energy moving during warmer sampling periods and the presence of tall canopies facilitated energetic conservation during daytime roosting activities. In the core population, where tall-canopied forest was prevalent, temperature anomalies did not affect territory occupancy dynamics as warmer sites were both less likely to go extinct and less likely to become colonized, suggesting a trade-off between foraging opportunities and temperature exposure. In the peripheral population, sites were more likely to become unoccupied following warm summers, presumably because of less prevalent older forest conditions. While individuals avoided elevated energetic expenditure associated with temperature exposure, behavioral strategies to conserve energy may have diverted time and energy from reproduction or territory defense. Conserving older forests, which are threatened due to fire and drought, may benefit individuals from energetic consequences of exposure to stressful thermal conditions.

Human disturbance increases trophic niche overlap in terrestrial carnivore communities.

Animal foraging and competition are defined by the partitioning of three primary niche axes: space, time, and resources. Human disturbance is rapidly altering the spatial and temporal niches of animals, but the impact of humans on resource consumption and partitioning-arguably the most important niche axis-is poorly understood. We assessed resource consumption and trophic niche partitioning as a function of human disturbance at the individual, population, and community levels using stable isotope analysis of 684 carnivores from seven communities in North America. We detected significant responses to human disturbance at all three levels of biological organization: individual carnivores consumed more human food subsidies in disturbed landscapes, leading to significant increases in trophic niche width and trophic niche overlap among species ranging from mesocarnivores to apex predators. Trophic niche partitioning is the primary mechanism regulating coexistence in many communities, and our results indicate that humans fundamentally alter resource niches and competitive interactions among terrestrial consumers. Among carnivores, niche overlap can trigger interspecific competition and intraguild predation, while the consumption of human foods significantly increases human-carnivore conflict. Our results suggest that human disturbances, especially in the form of food subsidies, may threaten carnivores by increasing the probability of both interspecific competition and human-carnivore conflict. Ultimately, these findings illustrate a potential decoupling of predator-prey dynamics, with impacts likely cascading to populations, communities, and ecosystems.

Cascading effects of a disease outbreak in a remote protected area.

Disease outbreaks induced by humans increasingly threaten wildlife communities worldwide. Like predators, pathogens can be key top-down forces in ecosystems, initiating trophic cascades that may alter food webs. An outbreak of mange in a remote Andean protected area caused a dramatic population decline in a mammalian herbivore (the vicuña), creating conditions to test the cascading effects of disease on the ecological community. By comparing a suite of ecological measurements to pre-disease baseline records, we demonstrate that mange restructured tightly linked trophic interactions previously driven by a mammalian predator (the puma). Following the mange outbreak, scavenger (Andean condor) occurrence in the ecosystem declined sharply and plant biomass and cover increased dramatically in predation refuges where herbivory was historically concentrated. The evidence shows that a disease-induced trophic cascade, mediated by vicuña density, could supplant the predator-induced trophic cascade, mediated by vicuña behaviour, thereby transforming the Andean ecosystem.

Carnivore Niche Partitioning in a Human Landscape.

AbstractTo minimize competitive overlap, carnivores modify one of their critical niche axes: space, time, or resources. However, we currently lack rules for how carnivore communities operate in human-dominated landscapes. We simultaneously quantified overlap in the critical niche axes of a simple carnivore community-an apex carnivore (Puma concolor), a dominant mesocarnivore (Lycalopex culpaeus), and a subordinate small carnivore (Lycalopex griseus)-in a human landscape featuring pastoralists and semidomestic carnivores (i.e., dogs, Canis familiaris). We found that dominant species had strong negative effects on the space use of subordinate ones, which ultimately created space for subordinate small carnivores. Humans and dogs were strictly diurnal, whereas the native carnivore community was nocturnal and exhibited high temporal overlap. Dietary overlap was high among the native carnivores, but dogs were trophically decoupled, largely because of human food subsidies. Our results show that in landscapes with evident human presence, temporal and dietary partitioning among native carnivores can be limited, leaving space as the most important axis to be partitioned among carnivores. We believe that these findings-the first to simultaneously assess all three critical niche axes among competing carnivores and humans and their associated species (i.e., domesticated carnivores)-are transferable to other carnivore communities in human-modified landscapes.

Seasonality drives the survival landscape of a recovering forest carnivore in a changing world.

Ecological heterogeneity promotes species persistence and diversity. Environmental change has, however, eroded patterns of heterogeneity globally, stifling species recovery. To test the effects of seasonal heterogeneity on a reintroduced carnivore, American martens (Martes americana), we compared metrics of local and season-specific heterogeneity to traditional forest metrics on the survival of 242 individuals across 8 years and predicted a survival landscape for 13 reintroduction sites. We found that heterogeneity-created by forest structure in the growing season and snow in the winter-improved survival and outperformed traditional forest metrics. Spatial variation in heterogeneity created a distinct survival landscape, but seasonal change in heterogeneity generated temporal discordance. All translocation sites possessed high forest heterogeneity but there were greater differences in winter heterogeneity; recovery sites with the poorest snow conditions had the lowest viability. Our work links heterogeneity across seasons to fitness and suggests that management strategies that increase seasonal aspects of heterogeneity may help to recover other sensitive species to continuing environmental change.

A recovery network leads to the natural recolonization of an archipelago and a potential trailing edge refuge.

Rapid environmental change is reshaping ecosystems and driving species loss globally. Carnivore populations have declined and retracted rapidly and have been the target of numerous translocation projects. Success, however, is complicated when these efforts occur in novel ecosystems. Identifying refuges, locations that are resistant to environmental change, within a translocation framework should improve population recovery and persistence. American martens (Martes americana) are the most frequently translocated carnivore in North America. As elsewhere, martens were extirpated across much of the Great Lakes region by the 1930s and, despite multiple translocations beginning in the 1950s, martens remain of regional conservation concern. Surprisingly, martens were rediscovered in 2014 on the Apostle Islands of Lake Superior after a putative absence of >40 yr. To identify the source of martens to the islands and understand connectivity of the reintroduction network, we collected genetic data on martens from the archipelago and from all regional reintroduction sites. In total, we genotyped 483 individual martens, 43 of which inhabited the Apostle Islands (densities 0.42-1.46 km-2 ). Coalescent analyses supported the contemporary recolonization of the Apostle Islands with progenitors likely originating from Michigan, which were sourced from Ontario. We also identified movements by a first-order relative between the Apostle Islands and the recovery network. We detected some regional gene flow, but in an unexpected direction: individuals moving from the islands to the mainland. Our findings suggest that the Apostle Islands were naturally recolonized by progeny of translocated individuals and now act as a source back to the reintroduction sites on the mainland. We suggest that the Apostle Islands, given its protection from disturbance, complex forest structure, and reduced carnivore competition, will act as a potential refuge for marten along their trailing range boundary and a central node for regional recovery. Our work reveals that translocations, even those occurring along southern range boundaries, can create recovery networks that function like natural metapopulations. Identifying refuges, locations that are resistant to environmental change, within these recovery networks can further improve species recovery, even within novel environments. Future translocation planning should a priori identify potential refuges and sources to improve short-term recovery and long-term persistence.

Limited sexual segregation in a dimorphic avian scavenger, the Andean condor.

Sexual segregation is widely reported among sexually dimorphic species and generally attributed to intraspecific competition. Prey diversity and human activities can reinforce niche segregation by increasing resource heterogeneity. Here, we explored trophic and spatial sexual segregation in the only avian scavenger that exhibits pronounced sexual size dimorphism (up to 50% difference in body mass) and a highly despotic social system, the Andean condor (Vultur gryphus). We predicted that larger and dominant males would exclude smaller and subordinate females from high-quality resources, leading to sexual segregation particularly in human-dominated landscapes showing increased prey diversity. We compared resource use between females and males across six sites in Argentina featuring a range of prey diversity via stable isotopes analysis of molted feathers (n = 141 individuals). We then focused on two sites featuring contrasting levels of prey diversity and quantified assimilated diet via stable isotopes and space use via GPS monitoring (n = 23 and 12 tagged individuals). We found no clear differences in isotopic niche space, individual variation in isotopic signature, or assimilated diet between females and males. However, there were differences in foraging locations between sexes, with females apparently using areas of fewer food resources more frequently than males. Local conditions defined the dynamics of fine-scale sexual differences in foraging sites; yet, unpredictable and ephemeral carrion resources likely prevent segregation by sexes at the landscape scale. Our study highlights complex dynamics of sexual segregation in vultures and the relevancy of analyses under multiple spatial-temporal scales to explore segregation in social species.

Future winters present a complex energetic landscape of decreased costs and reduced risk for a freeze-tolerant amphibian, the Wood Frog (Lithobates sylvaticus).

Winter climate warming is rapidly leading to changes in snow depth and soil temperatures across mid- and high-latitude ecosystems, with important implications for survival and distribution of species that overwinter beneath the snow. Amphibians are a particularly vulnerable group to winter climate change because of the tight coupling between their body temperature and metabolic rate. Here, we used a mechanistic microclimate model coupled to an animal biophysics model to predict the spatially explicit effects of future climate change on the wintering energetics of a freeze-tolerant amphibian, the Wood Frog (Lithobates sylvaticus), across its distributional range in the eastern United States. Our below-the-snow microclimate simulations were driven by dynamically downscaled climate projections from a regional climate model coupled to a one-dimensional model of the Laurentian Great Lakes. We found that warming soil temperatures and decreasing winter length have opposing effects on Wood Frog winter energy requirements, leading to geographically heterogeneous implications for Wood Frogs. While energy expenditures and peak body ice content were predicted to decline in Wood Frogs across most of our study region, we identified an area of heightened energetic risk in the northwestern part of the Great Lakes region where energy requirements were predicted to increase. Because Wood Frogs rely on body stores acquired in fall to fuel winter survival and spring breeding, increased winter energy requirements have the potential to impact local survival and reproduction. Given the geographically variable and intertwined drivers of future under-snow conditions (e.g., declining snow depths, rising air temperatures, shortening winters), spatially explicit assessments of species energetics and risk will be important to understanding the vulnerability of subnivium-adapted species.

The demography of a resource specialist in the tropics: Cecropia trees and the fitness of three-toed sloths.

Resource specialists persist in a narrow range of resources. Consequently, the abundance of key resources should drive vital rates, individual fitness, and population viability. While Neotropical forests feature both high levels of biodiversity and numbers of specialist species, no studies have directly evaluated how the variation of key resources affects the fitness of a tropical specialist. Here, we quantified the effect of key tree species density and forest cover on the fitness of three-toed sloths ( Bradypus variegatus), an arboreal folivore strongly associated with Cecropia trees in Costa Rica, using a multi-year demographic, genetic, and space-use dataset. We found that the density of Cecropia trees was strongly and positively related to both adult survival and reproductive output. A matrix model parametrized with Cecropia-demography relationships suggested positive growth of sloth populations, even at low densities of Cecropia (0.7 trees ha-1). Our study shows the first direct link between the density of a key resource to demographic consequences of a tropical specialist, underscoring the sensitivity of tropical specialists to the loss of a single key resource, but also point to targeted conservation measures to increase that resource. Finally, our study reveals that previously disturbed and regenerating environments can support viable populations of tropical specialists.

Snow roosting reduces temperature-associated stress in a wintering bird.

Animals in temperate northern regions employ a variety of strategies to cope with the energetic demands of winter. Behavioral plasticity may be important, as winter weather conditions are increasingly variable as a result of modern climate change. If behavioral strategies for thermoregulation are no longer effective in a changing environment, animals may experience physiological stress, which can have fitness consequences. We monitored winter roosting behavior of radio-tagged ruffed grouse (Bonasa umbellus), recorded snow depth and temperature, and assayed droppings for fecal corticosterone metabolites (FCM). Grouse FCM levels increased with declining temperatures. FCM levels were high when snow was shallow, but decreased rapidly as snow depth increased beyond 20 cm. When grouse used snow burrows, there was no effect of temperature on FCM levels. Snow burrowing is an important strategy that appears to allow grouse to mediate the possibly stressful effects of cold temperatures. This is one of the first studies to explore how variable winter weather conditions influence stress in a free-living cold-adapted vertebrate and its ability to mediate this relationship behaviorally. Animals that depend on the snowpack as a winter refuge will likely experience increased stress and possible fitness costs resulting from the loss of snow cover due to climate change.

Integrating temporal refugia into landscapes of fear: prey exploit predator downtimes to forage in risky places.

The landscape of fear is an important driver of prey space use. However, prey can navigate the landscape of fear by exploiting temporal refuges from predation risk. We hypothesized that diel patterns of predator and prey movement and space use would be inversely correlated due to temporal constraints on predator habitat domain. Specifically, we evaluated habitat selection and activity of the vicuña and its only predator, the puma, during three diel periods: day, dawn/dusk, and night. Pumas selected the same habitats regardless of diel period-vegetated and rugged areas that feature stalking cover for pumas-but increased their activity levels during dawn/dusk and night when they benefit from reduced detection by prey. Vicuñas avoided areas selected by pumas and reduced activity at night, but selected vegetated areas and increased activity by day and dawn/dusk. Vicuña habitat selection and movement strategies appeared to reduce the risk of encountering pumas; movement rates of pumas and vicuñas were negatively correlated across the diel cycle, and habitat selection was negatively correlated during dawn/dusk and night. Our study shows that an ambush predator's temporal activity and space use patterns interact to create diel refugia and shape the antipredator behaviors of its prey. Importantly, it is likely the very nature of ambush predators' static habitat specificity that makes predator activity important to temporally varying perceptions of risk. Prey which depend on risky habitats for foraging appear to mitigate risk by feeding when they can more easily detect predators and when predators are least active.

Genomics meets applied ecology: Characterizing habitat quality for sloths in a tropical agroecosystem.

Understanding how habitat quality in heterogeneous landscapes governs the distribution and fitness of individuals is a fundamental aspect of ecology. While mean individual fitness is generally considered a key to assessing habitat quality, a comprehensive understanding of habitat quality in heterogeneous landscapes requires estimates of dispersal rates among habitat types. The increasing accessibility of genomic approaches, combined with field-based demographic methods, provides novel opportunities for incorporating dispersal estimation into assessments of habitat quality. In this study, we integrated genomic kinship approaches with field-based estimates of fitness components and approximate Bayesian computation (ABC) procedures to estimate habitat-specific dispersal rates and characterize habitat quality in two-toed sloths (Choloepus hoffmanni) occurring in a Costa Rican agricultural ecosystem. Field-based observations indicated that birth and survival rates were similar in a sparsely shaded cacao farm and adjacent cattle pasture-forest mosaic. Sloth density was threefold higher in pasture compared with cacao, whereas home range size and overlap were greater in cacao compared with pasture. Dispersal rates were similar between the two habitats, as estimated using ABC procedures applied to the spatial distribution of pairs of related individuals identified using 3,431 single nucleotide polymorphism and 11 microsatellite locus genotypes. Our results indicate that crops produced under a sparse overstorey can, in some cases, constitute lower-quality habitat than pasture-forest mosaics for sloths, perhaps because of differences in food resources or predator communities. Finally, our study demonstrates that integrating field-based demographic approaches with genomic methods can provide a powerful means for characterizing habitat quality for animal populations occurring in heterogeneous landscapes.

Conserving and managing the subnivium.

In regions where snowfall historically has been a defining seasonal characteristic of the landscape, warming winters have reduced the depth, duration, and extent of snowpack. However, most management and conservation has focused on how aboveground wildlife will be affected by altered snow conditions, even though the majority of species that persist through the winter do so under the snowpack in a thermally stable refugium: the subnivium. Shortened winters, forest management practices, and winter recreation can alter subnivium conditions by increasing snow compaction and compromising thermal stability at the soil-snow interface. To help slow the loss of the subnivium in the face of rapidly changing winter conditions, we suggest managers adopt regional conservation plans for identifying threatened snow-covered environments; measure and predict the effects land cover and habitat management has on local subnivium conditions; and control the timing and distribution of activities that disturb and compact snow cover (e.g., silvicultural practices, snow recreation, and road and trail maintenance). As a case study, we developed a spatially explicit model of subnivium presence in a working landscape of the Chequamegon National Forest, Wisconsin. We identified landscapes where winter recreation and management practices could threaten potentially important areas for subnivium persistence. Similar modeling approaches could inform management decisions related to subnivium conservation. Current climate projections predict that snow seasons will change rapidly in many regions, and as result, we advocate for the immediate recognition, conservation, and management of the subnivium and its dependent species.

Microbes are trophic analogs of animals.

In most ecosystems, microbes are the dominant consumers, commandeering much of the heterotrophic biomass circulating through food webs. Characterizing functional diversity within the microbiome, therefore, is critical to understanding ecosystem functioning, particularly in an era of global biodiversity loss. Using isotopic fingerprinting, we investigated the trophic positions of a broad diversity of heterotrophic organisms. Specifically, we examined the naturally occurring stable isotopes of nitrogen ((15)N:(14)N) within amino acids extracted from proteobacteria, actinomycetes, ascomycetes, and basidiomycetes, as well as from vertebrate and invertebrate macrofauna (crustaceans, fish, insects, and mammals). Here, we report that patterns of intertrophic (15)N-discrimination were remarkably similar among bacteria, fungi, and animals, which permitted unambiguous measurement of consumer trophic position, independent of phylogeny or ecosystem type. The observed similarities among bacterial, fungal, and animal consumers suggest that within a trophic hierarchy, microbiota are equivalent to, and can be interdigitated with, macrobiota. To further test the universality of this finding, we examined Neotropical fungus gardens, communities in which bacteria, fungi, and animals are entwined in an ancient, quadripartite symbiosis. We reveal that this symbiosis is a discrete four-level food chain, wherein bacteria function as the apex carnivores, animals and fungi are meso-consumers, and the sole herbivores are fungi. Together, our findings demonstrate that bacteria, fungi, and animals can be integrated within a food chain, effectively uniting the macro- and microbiome in food web ecology and facilitating greater inclusion of the microbiome in studies of functional diversity.

Cophylogenetics and biogeography reveal a coevolved relationship between sloths and their symbiont algae.

Specialized species, like arboreal folivores, often develop beneficial relationships with symbionts to exploit ecologically constrained lifestyles. Although coevolution can drive speciation by specialization of a symbiont to a host, a symbiotic relationship is not indicative of coevolution between host and symbiont. We tested for coevolved relationships between highly specialized two- and three-toed sloths (Choloepus spp. and Bradypus spp., respectively) and their symbiotic algae using cophylogenies and phylogeography. Our phylogeographic analysis showed a biogeographic pattern for the sloth distribution that was not found in the algal phylogeny. We found support for congruence between the sloth and algae phylogenies, implying cospeciation, only in the Bradypus lineage. Algae host-switching occurred from Bradypus spp. to Choloepus spp. Our results support a previously hypothesized symbiotic relationship between sloths and the algae in their fur and indicate that coevolution may have played a role in algae diversification. More broadly, convergent evolution may facilitate host switching between deeply diverged host lineages.